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Systema Porifera. A Guide To The Classification of Sponges. Hooper, John N. A. y Van Soest, Rob W.M.-2002

Q: Explain the significance of Acanthochaetetes' survival strategy in its long evolutionary history.

Acanthochaetetes' survival strategy involves its 'chaetetid' basal skeleton serving as a protective and resting body for special omnipotent cells like thesocytes and archaeocytes, enabling it to survive environmental crises. This strategy allows the species to initiate new growth on existing calicles, signaling its importance in its evolutionary history, contributing to a continuous lineage from the Lower Cretaceous to present .

Q: Describe the role of sponge-related microorganisms in the mineralization of sponge tissue based on the Devonian fossil record.

Sponge-related microorganisms play a crucial role in the mineralization of sponge tissue through processes such as sulfate reduction and ammonification. These processes contribute to the rapid preservation of sponge tissue by facilitating mineralization, which is essential for maintaining the integrity of sponge fossils over time. This mechanism of preservation is often captured in chemotaxonomic examinations of Devonian sponge fossils .

Q: What are the key factors that contributed to the rapid development of polyphyletic coralline sponges during the Devonian period?

The key factors contributing to the rapid development of polyphyletic coralline sponges during the Devonian period include medium-term fundamental oceanic changes that resulted in new niches, possibly due to increased nutrient input and alkalinity, along with a massive increase in Ca2+ dissolved in ocean water. These changes created new environmental conditions like deeper water and protected areas, which prompted the diversification of these sponges .

Q: What are the main arguments for the inclusion of non-skeletal features in the taxonomic classification of sponges?

Main arguments for including non-skeletal features in sponge taxonomy include the limited preservation of skeletal features, variability in spicule formation, and the need for a taxonomic framework that accommodates organisms with minimal or no skeletal records. Factors like cellular structures, reproductive strategies, and genetic markers provide important complementary data to enhance accuracy in classification .

Q: How do unique structural features of certain fossilized sponges provide insight into their biological functions and environmental adaptations?

Unique structural features, such as the 'thalarnid' and aragonitic spherulites in Subascosymplegma oussifensis, suggest specific biological functions like structural support and protection in high-energy environments. These structures, along with inhalant pore arrangements, also indicate adaptations to nutrient fluxes and protection against environmental stressors. Such features offer insights into the evolutionary pressures these sponges faced .

Q: Discuss the relationship between sponge symbionts and environmental adaptation as seen in recent and fossilized sponges.

Sponge symbionts play a critical role in environmental adaptation by facilitating nutrient acquisition, contributing to chemical defense, and supporting structural integrity against abiotic stress. In both recent and fossilized sponges, symbiotic relationships often indicate specialized adaptations to specific ecological niches, influencing the evolutionary success of some sponge taxa .

Q: What evidence suggests that some Archaeocyatha may be classified as demosponges?

Evidence suggesting some Archaeocyatha may be demosponges includes the occurrence of tetractinellid spicules within the basal skeletons of some specimens, reflecting characteristics common in demosponges. However, this hypothesis requires more substantial corroborative evidence for a definitive classification .

Q: How does the poor preservation potential of non-lithistid demosponges affect our understanding of their evolutionary trends?

The poor preservation potential of non-lithistid demosponges, due to their soft tissue and discrete siliceous spicules losing clarity or being replaced by other minerals, severely limits our knowledge of their evolutionary trends. This poor record makes it challenging to identify evolutionary progressions within this group, which impacts understanding both phylogeny and taxonomy .

Q: Assess the impact of geological preservation conditions on understanding the diversity of Lower Cambrian sponges.

The diversity of Lower Cambrian sponges is often underestimated due to unfavorable geological preservation conditions, which lead to poor representation of non-lithistid demosponge fossils. Preservation typically favors hard-bodied organisms, thus biasing our understanding and limiting insights into the diverse soft-bodied sponge taxa that existed during this period .

Q: How do the choanosomal and cortical structures of Porifera contribute to their physiological functions?

In Porifera, the choanosomal structures, often composed of oxyasters and radial arrangements, are crucial for maintaining water flow through the sponge, optimizing nutrient absorption and waste expulsion. Cortical layers, including sterraster and microxead arrangements, provide structural integrity and protect internal cells, influencing both defense mechanisms and environmental adaptations .

This document provides an introduction to a two-volume guide to the classification of sponges. Volume 1 will cover the class Demospongiae and introductions. It was edited by John Hooper and Rob Van Soest with contributions from several sponge researchers and taxonomists. The guide aims to revisit and accommodate new sponge taxa, correct past mistakes, and provide a phylogenetic framework for sponge systematics. It describes approximately 680 living genera in 127 families and 25 orders, as well as 1000 fossil genera. The collaborative effort involved over 45 researchers from around the world over six years.

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Systema Porifera. A Guide To The Classification of Sponges. Hooper, John N. A. y Van Soest, Rob W.M.-2002

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SYSTEMA PORIFERA

A Guide to the
Classification of Sponges
VOLUME 1
SYSTEMA PORIFERA
A Guide to the
Classification of Sponges
VOLUME 1

Edited by

John N. A. Hooper
Queensland Museum
South Brisbane, Queensland, Australia

and

Rob W. M. Van Soest

Zoological Museum
University ofAmsterdam
Amsterdam, Netherlands

Bibliographic Editor for Recent Taxa

Philippe Willenz
Royal Institute of Natural Sciences of Belgium
Brussels, Belgium

K1uwer Academic I Plenum Publishers

New York, Boston, Dordrecht, London, Moscow
Library of Congress Cataloging-in-Publication Data

Systema Porifera: a guide to the classification of sponges/scientific editors, John N.A.

Hooper and Rob W.M. Van Soest.
p.cm.
Includes bibliographical references (p. ).
ISBN 0-306-47260-0
1. Sponges-Classification. I. Hooper, John N. A. II. Soest, R. W. M. van
QL371 .8962002
693.4'012--dc21

2002072781

ISBN 0-306-47260-0
Published in two volumes:
Volume 1: Introductions and Demospongiae
Volume 2: Calcarea, Hexactinellida, Sphinctozoa, Archaeocyatha, unrecognizable taxa, and Index of higher taxa
© 2002 Kluwer Academic / Plenum Publishers, New York
233 Spring Street, New York, New York 10013
http://www.wkap.nll
10 9 8 7 6 5 4 3
A C.I.P. record for this book is available from the Library of Congress
All rights reserved
No part of this book may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical,
photocopying, microfilming, recording, or otherwise, without written permission from the Publisher, with the exception of any material supplied
specifically for the purpose of being entered and executed on a computer system, for exclusive use by the purchaser of the work.
Preface

Sponges are among the most ancient of multicellular these many new taxa, to correct the many nomenclat-
animals alive on the planet today. Ecologically and eco- ural and taxonomic mistakes of the past, and to provide
nomically, they are among the most highly successful a sound platform for the future development of sponge
life forms that have ever existed. First known from the systematics. For many groups the Systema is a major
Precambrian, and well established by the Cambrian, revision, and for all groups it provides a sound frame-
they were the major reef builders during the Palaeozoic work of sponge systematics based on phylogenetic
and Mesozoic Eras. Today they are highly diverse, with principles. Perhaps more importantly, this project also
an estimated 15,000 species living in all Recent marine provides a practical tool to allocate taxa within this
and freshwater habitats worldwide, but these undoubt- framework, including keys, comprehensive illustra-
edly represent only a small proportion of those that have tions and descriptions of the major characters used to
ever lived. Sponge body plans do not appear to have classify sponges - something that has not previously
changed significantly over the many millennia, such been attempted.
that some species alive today appear to be almost identi- The Systema is published in two volumes.
cal to those that once lived in Earth's ancient seas. Volume 1 includes a general introduction and the Class
Consequently, sponges are often regarded as ultra-con- Demospongiae. Volume 2 includes Classes Calcarea
servative in their morphology, presumably as successful and Hexactinellida, the fossil Class Archaeocyatha,
models of early evolutionary experiments. In contrast, fossil 'Sphinctozoa', and an annotated list of unrecog-
the living fauna is widely diverse in many other biologi- nisable taxa and unavailable names. We describe
cal features, especially in their biochemical con- approximately 680 genera of sponges in 127 families,
stituency that is currently of major interest to the 25 orders and three classes of the living fauna. In addi-
pharmaceutical industry. It is this latter feature, and also tion, approximately 1000 fossil genera in 245 families,
the renewed interest in marine biodiversity worldwide, 30 orders and six 'classes' are also covered. The living
that has attracted a m;:yor resurgence of studies on fauna is dealt with far more comprehensively than the
sponges with the consequence that over the past two fossil fauna, with overviews of the fossil classification
decades researchers have discovered a living fauna included mainly in an attempt to relate these two classi-
about twice the size of that published in the literature. fications more closely. A far more comprehensive treat-
Sponge systematics is generally perceived as ment of fossil sponges is currently in preparation for the
appallingly difficult. These difficulties stem partly revised edition of the Treatise on Invertebrate
from the sponges themselves, being highly plastic and Palaeontology (J.K. Rigby et al., editors).
easily susceptible to environmental modification and The Systema project was only possible to achieve
frequent character losses. They also stem, perhaps through an intensively collaborative effort amongst 45
more so, from the inadequacies and inconsistencies researchers from 17 countries, spanning about six years of
of the taxonomic framework. This includes a previ- research effort. We pride ourselves to have among our
ously fragmented and uncoordinated research effort, contributors the major contemporary spongologists,
whereby only a few higher groups have had contempo- including Patricia R. Bergquist, Nicole Boury-Esnault,
rary revisions applied to them, and with the remainder Franr;oise Debrenne, Claude Levi, Henry Reiswig,
largely unworked since the late 19th century. In tum, Joachim Reitner, Klaus Rtitzler, Michele Sara, Maria J.
this has led to increasing difficulties faced by new Uriz and Jean Vacelet, as well as several members of the
workers - among them a growing number from newly new generation of sponge researchers. The Systema
developing countries - to access the vast and mostly Porifera project is not an end - but a sound beginning for
ancient literature. this new generation to build on what we propose here.
The Systema Porifera project was devised to
revisit the poriferan higher taxa (genus level and John N.A. Hooper, Brisbane
above, based on type species data), to accommodate Rob W.M. Van Soest, Amsterdam

v
Contents of Volumes 1 and 2
VOLUME 1

Key to Symbols, Institutional Acronyms, Abbreviations .. .............. ......... ............ .......... ... .............. .... .... ......... xv
Acknowledgements ........................................................................................................................................ xvii
Introduction
John N.A. Hooper & Rob W.M. Van Soest .................................................................................................. 1
Bibliography of Introductions
Editor Philippe Willenz ............. .... .... .......................... ......... ..... ......... ......... ... .......... .................................... 4
Phylum Porifera Grant
John N.A. Hooper, Rob W.M. Van Soest & Fran~oise Debrenne ................................................................ 9
Class Demospongiae Sollas
John N.A. Hooper & Rob W.M. Van Soest ................................................................................................ 15
Bibliography of Class Demospongiae
Editor Philippe Willenz ........ ..... ......... ... ...... ... ..... ............. ........................... ............. ........ ......... ......... ..... ... 19
Non-Lithistid Fossil Demospongiae - Origins of their Palaeobiodiversity and Highlights in
History of Preservation
Joachim Reitner & Gert Worheide ............................................................................................................. 52
t'Class Stromatoporoidea' Nicholson & Murie, Stromatoporoids
Alex Cook ............................................................................. ..................... ........................... ......... ............ 69
Order Homosclerophorida Dendy. Family Plakinidae Schulze
Guilherme Muricy & Maria Christina Diaz ............................................................................................... 71
Order Spirophorida Bergquist & Hogg
Rob W.M. Van Soest & John N.A. Hooper ............................................................................................... 83
Family Tetillidae Sollas
Rob W.M. Van Soest & Klaus Rtitzler ... .............. .... ......... ................................... ........ ....................... ....... 85
Family Samidae Sollas
Rob W.M. Van Soest & John N.A. Hooper ................................................................................................ 99
Family Spirasigmidae Hallmann
John N.A. Hooper & Rob W.M. Van Soest .............................................................................................. 102
Order Astrophorida Sollas
John N.A. Hooper & Rob W.M. Van Soest .............................................................................................. 105
Family Ancorinidae Schmidt
Maria J. Driz ............................................................................................................................................ 108
Family Calthropellidae Lendenfeld
Rob W.M. Van Soest & John N.A. Hooper .............................................................................................. 127
Family Geodiidae Gray
Maria J. Driz ............................................................................................................................................ 134
Family Pachastrellidae Carter
Manuel Maldonado ........... ......... ...................... ............ ........................... ................. .................. .............. 141
Family Thrombidae Sollas
Maria J. Driz ............................................................................................................................................ 163
Astrophorida Incertae Sedis
John N.A. Hooper & Manuel Maldonado ................................................................................................ 165
Order Hadromerida Topsent
John N.A. Hooper & Rob W.M. Van Soest .............................................................................................. 169
Family Clionaidae D'Orbigny
Klaus Rtitzler ........................................................................................................................................... 173

vii
viii Contents of Volumes 1 and 2

Family Hemiasterellidae Lendenfeld

John N.A. Hooper .................................................................................................................................... 186
Family Placospongiidae Gray
Klaus Riitzler ............... ... ...................... ............................... ........................... ............. ........ ..... ......... ...... 196
Family Polymastiidae Gray
Nicole Boury-Esnault ............................................................................................................................... 201
Family Spirastrellidae Ridley & Dendy
Klaus Riitzler ........................... ..... ......... ....................................... ..... ............ ................................ .......... 220
Family Stylocordylidae Topsent
Rob W.M. Van Soest ................................................................................................................................ 224
Family Suberitidae Schmidt
Rob W.M. Van Soest ................................................................................................................................ 227
Family Tethyidae Gray
Michele Sara ... ......... ......................... ............................... ......... ..... ... .............. ................. ..... ........ ........... 245
Family Timeidae Topsent
Klaus Riitzler .............................................................................................................................. ............. 266
Family Trachycladidae Hallmann
John N.A. Hooper & Rob W.M. Van Soest .............................................................................................. 268
Family Acanthochaetetidae Fischer
Klaus Riitzler & Jean Vacelet ................................................................................................................... 275
Family Sollasellidae Lendenfeld
Rob W.M. Van Soest .............................................................................................................................. 279
Family Alectonidae Rosell
Klaus Riitzler ........................ ........ .................. ..... ........ ..... ..................... ..... ............................................. 281
Order Chondrosida Boury-Esnault & Lopes. Family Chondrillidae Gray
Nicole Boury-Esnault ............................................................................................................................... 291
'Lithistid'Demospongiae
Andrzej Pisera & Claude Levi ................ ......................... ......... ..... ......... ................................... .............. 299
Family Scleritodermidae Sollas
Andrzej Pisera & Claude Levi .. ..... ......... ......................... ..... ............. ..... ......... ........ ............. ................... 302
Family Corallistidae Sollas
Andrzej Pisera & Claude Levi ................................................................................................................. 312
Family Pleromidae Sollas
Andrzej Pisera & Claude Levi ................................................................................................................. 321
Family Theonellidae Lendenfeld
Andrzej Pisera & Claude Levi ................................................................................................................. 327
Family Siphonidiidae Lendenfeld
Andrzej Pisera & Claude Levi ................................................................................................................. 338
Family Neopeltidae Sollas
Andrzej Pisera & Claude Levi .. ......... .................. ................................... ....................................... .......... 344
Family Azoricidae Sollas
Andrzej Pisera & Claude Levi ................................................................................................................. 352
Family Desmanthidae Topsent
Andrzej Pisera & Claude Levi ............ ... .......... ............. ........ .......................................................... ......... 356
Family Vetulinidae Lendenfeld
Andrzej Pisera & Claude Levi ........... ..................................................... ............. ..... ............................... 363
Family Phymatellidae Schrarnmen
Andrzej Pisera & Claude Levi ................ .................. ................. ........ .......... ......... ... ...................... .......... 366
Family Isoraphiniidae Schrarnmen
Andrzej Pisera & Claude Levi ................................................................................................................. 374
Family Macandrewiidae Schrarnmen
Andrzej Pisera & Claude Levi ................................................................................................................. 377
Contents of Volumes 1 and 2 ix

Family Phymaraphiniidae Schrarnmen

Andrzej Pisera & Claude Levi ................................................................................................................. 380
Lithistids Incertae Sedis
Andrzej Pisera & Claude Levi ............................ ................. ........................... .......................... ............... 384
tFossil Lithistids: An Overview
Andrzej Pisera .......................................................................................................................................... 388
Order Poecilosclerida Topsent
John N.A. Hooper & Rob W.M. Van Soest .............................................................................................. 403
Suborder Microcionina Hajdu, Van Soest & Hooper
John N.A. Hooper .................................................................................................................................... 409
Family Acarnidae Dendy
John N.A. Hooper .................................................................................................................................... 412
Family Microcionidae Carter
John N.A. Hooper .................................................................................................................................... 432
Family Raspailiidae Hentschel
John N.A. Hooper .................................................................................................................................... 469
Family Rhabderemiidae Topsent
John N.A. Hooper .................................................................................................................................... 511
Suborder Myxillina Hajdu, Van Soest & Hooper
Rob W.M. Van Soest ................................................................................................................................ 515
Family Chondropsidae Carter
Rob W.M. Van Soest ................................................................................................................................ 521
Family Coelosphaeridae Dendy
Rob W.M. Van Soest ................................................................................................................................ 528
Family Crambeidae Levi
Rob W.M. Van Soest 547
Family Crellidae Dendy
Rob W.M. Van Soest 556
Family Dendoricellidae Hentschel
Rob W.M. Van Soest ................................................................................................................................ 567
Family Desmacididae Schmidt
Rob W.M. Van Soest ................................................................................................................................ 572
Family Hymedesmiidae Topsent
Rob W.M. Van Soest ................................................................................................................................ 575
Family Iotrochotidae Dendy
Rob W.M. Van Soest '" ..... ..... ....................................................................... ............................................ 594
Family Myxillidae Dendy
Rob W.M. Van Soest ................................................................................................................................ 602
Family Phellodermidae fam. nov.
Rob W.M. Van Soest & Eduardo Hajdu ................................................................................................... 621
Family Tedaniidae Ridley & Dendy
Rob W.M. Van Soest ......................................................................................................... ....................... 625
Suborder Mycalina Hajdu, Van Soest & Hooper
Rob W.M. Van Soest & Eduardo Hajdu.. ................................................ ................................................. 633
Family Cladorhizidae Dendy
Eduardo Hajdu & Jean Vacelet ................................................................................................................ 636
Family Desmacellidae Ridley & Dendy
Eduardo Hajdu & Rob W.M. Van Soest ................................. ................. ................................................. 642
Family Guitarridae Dendy
Eduardo Hajdu & Clea Lerner ........ ................................................ ...................... ................................... 651
Family Esperiopsidae Hentschel
Rob W.M. Van Soest & Eduardo Hajdu ................................................................................................... 656
x Contents of Volumes 1 and 2

Family Hamacanthidae Gray

Eduardo Hajdu ............ ....................... ......... ................................................. ............. ................................ 665
Family Mycalidae Lundbeck
Rob W.M. Van Soest & Eduardo Hajdu ................................................................................................... 669
Family Merliidae Kirkpatrick
Eduardo Hajdu & Rob W.M. Van Soest ................................................................................................... 691
Family Podospongiidae De Laubenfels
Michelle Kelly & Toufiek Samaai ........................................................................................................... 694
Family Isodictyidae Dendy
Eduardo Hajdu & Gisele Lobo-Hajdu ..................................................................................................... 703
Suborder Latrunculina subord. nov. incertae sedis
Michelle Kelly & Toufiek Samaai ........................................................................................................... 707
Family Latrunculiidae Topsent
Toufiek Samaai & Michelle Kelly ........................................................................................................... 708
Order Halichondrida Gray
Rob W.M. Van Soest & John N.A. Hooper ............................................................................................. 721
Family Axinellidae Carter
Belinda Alvarez & John N.A. Hooper ..................................................................................................... 724
Family Bubaridae Topsent
Belinda Alvarez & Rob W.M. Van Soest .................................................................................................. 748
Family Desmoxyidae Hallmann
John N.A. Hooper .................................................................................................................................... 755
Family Dictyonellidae Van Soest, Diaz & Pomponi
Rob W.M. Van Soest, Dirk Erpenbeck & Belinda Alvarez ...................................................................... 773
Family Halichondriidae Gray
Dirk Erpenbeck & Rob W.M. Van Soest .................................................................................................. 787
Order Agelasida Hartman
Rob W.M. Van Soest & John N.A. Hooper ............................................................................................. 817
Family Agelasiidae Verrill
Rob W.M. Van Soest ................................................................................................................................ 819
Family Astrosc1eridae Lister
Jean Vacelet .............................................................................................................................................. 824
Order Haplosclerida Topsent
Rob W.M. Van Soest & John N.A. Hooper .............................................................................................. 831
Suborder Haplosclerina Topsent
Rob W.M. Van Soest & John N.A. Hooper .............................................................................................. 833
Family Callyspongiidae de Laubenfels
R. Desqueyroux-Faundez & C. Valentine ................................................................................................ 835
Family Chalinidae Gray
Wallie H. De Weerdt ................................................................................................................................. 852
Family Niphatidae Van Soest
Ruth Desqueyroux-Faundez & C. Valentine ............................................................................................ 874
Suborder Petrosina Boury-Esnault & Van Beveren
Rob W.M. Van Soest & John N.A. Hooper .............................................................................................. 891
Family Phloeodictyidae Carter
R. Desqueyroux-Faundez & C. Valentine ................................................................................................ 893
Family Petrosiidae Van Soest
R. Desqueyroux-Faundez & Clare Valentine ............................................................................................. 906
Family Calcifibrospongiidae Hartman
Rob W.M. Van Soest ................................................................................................................................ 918
Suborder Spongillina subord. nov. : Freshwater Sponges
Renata Manconi & Roberto Pronzato ...................................................................................................... 921
Contents of Volumes 1 and 2 xi

Order Dictyoceratida Minchin

Steve de C. Cook & Patricia R. Bergquist 1021
Family Irciniidae Gray
Steve de C. Cook & Patricia R. Bergquist 1022
Family Thorectidae Bergquist
Steve de C. Cook & Patricia R. Bergquist 1028
Family Spongiidae Gray
Steve de C. Cook & Patricia R. Bergquist 1051
Family Dysideidae Gray
Steve de C. Cook & Patricia R. Bergquist 1061
Order Dendroceratida Minchin
Patricia R. Bergquist & Steve de C. Cook ............................................................................................. 1067
Family Darwinellidae Merejkowsky
Patricia R. Bergquist & Steve de C. Cook ............................................................................................. 1068
Family Dictyodendrillidae Bergquist
Patricia R. Bergquist & Steve de C. Cook .................................... .................. ..... ................. .............. ... 1072
Order Halisarcida Bergquist
Patricia R. Bergquist & Steve de C. Cook ............................................................................................. 1077
Family Halisarcidae Schmidt
Patricia R. Bergquist & Steve de C. Cook ............................................................................................. 1078
Order Verongida Bergquist
Patricia R. Bergquist & Steve de C. Cook ............................................................................................. 1081
Family Aplysinidae Carter
Patricia R. Bergquist & Steve de C. Cook ............................................................................................. 1082
Family Pseudoceratinidae Carter
Patricia R. Bergquist & Steve de C. Cook ............................................................................................. 1086
Family Ianthellidae Hyatt
Patricia R. Bergquist & Steve de C. Cook .......... ..................................................... .............. ................ 1089
Family Aplysinellidae Bergquist
Patricia R. Bergquist & Steve de C. Cook ............................................................................................. 1094
Recent 'Sphinctozoa', Order Verticillitida, Family Verticillitidae Steinmann
Jean Vacelet ............................................................................................................................................ 1097
Demospongiae Incertae sedis: Myceliospongia Vacelet & Perez
Jean Vacelet, Thierry Perez & John N.A. Hooper .................................................................................. 1099
Index of Higher Taxa to Volumes 1 and 2 ............................... ..... ................. .......................... ........................ xxi

VOLUME 2

Key to Symbols, Institutional Acronyms, Abbreviations ................................................................................. xv

Acknowledgements ........................................................................................................................................ xvii
Class Calcarea Bowerbank
Michael Manuel, Radovan Borojevic, Nicole Boury-Esnau1t & Jean Vace1et ....................................... 1103
Bibliography of Class Calcarea
Editor Philippe Willenz .......... ... .......... .... ................ ....................... ............. .............. ......... ................. ... 1111
tFossil Calcarea. An Overview
John Pickett ............................................................................................................................................ 1117
tOrder Heteractinida Hinde
John Pickett ............................................................................................................................................ 1121
xii Contents of Volumes 1 and 2

Order Clathrinida Hartman

Radovan Borojevic, Nicole Boury-Esnault, Michael Manuel & Jean Vacelet 1141
Order Murrayonida Vacelet
Jean Vacelet, Radovan Borojevic, Nicole Boury-Esnault & Michael Manuel 1153
Order Leucosolenida Hartman
Radovan Borojevic, Nicole Boury-Esnault, Michael Manuel & Jean Vacelet 1157
Order Lithonida Vacelet, Recent
Jean Vacelet, Radovan Borojevic, Nicole Boury-Esnault & Michael Manuel 1185
Order Baerida Borojevic, Boury-Esnault & Vacelet
Radovan Borojevic, Nicole Boury-Esnault, Michael Manuel & Jean Vacelet 1193
Class Hexactinellida Schmidt
Henry M. Reiswig .................................................................................................................................. 1201
Bibliography of Class Hexactinellida
Editor Philippe Willenz .......................................................................................................................... 1203
tFossil Hexactinellida: An Overview
Manfred Krautter .................................................................................................................................... 1211
Dictionary of Hexactinellida
Konstantin R. Tabachnick & Henry M. Reiswig .................................................................................... 1224
Order Amphidiscosida Schrammen
Henry M. Reiswig .................................................................................................................................. 1231
Family Hyalonematidae Gray
Konstantin R. Tabachnick & Larisa L. Menshenina ............................................................................... 1232
Family Monorhaphididae Ijima
Konstantin R. Tabachnick ...................................................................................................................... 1264
Family Pheronematidae Gray
Konstantin R. Tabachnick & Larisa L. Menshenina .............................................................................. 1267
Order Hexactinosida Schrammen
Henry M. Reiswig .................................................................................................................................. 1281
Family Aphrocallistidae Gray
Henry M. Reiswig .................................................................................................................................. 1282
Family Craticulariidae Rauff (Recent)
Henry M. Reiswig .................................................................................................................................. 1287
Family Cribrospongiidae Roemer (Recent)
Henry M. Reiswig .................................................................................................................................. 1290
Family Dactylocalycidae Gray
Henry M. Reiswig .................................................................................................................................. 1293
Family Euretidae Zittel
Henry M. Reiswig & Benjamin Wheeler .............................................................................................. 1301
Family Farreidae Gray
Henry M. Reiswig .................................................................................................................................... 1332
Family Tretodictyidae Schulze
Henry M. Reiswig .................................................................................................................................. 1341
Hexactinosida incertae sedis
Henry M. Reiswig .................................................................................................................................. 1355
Order Aulocalycoida Tabachnick & Reiswig
Henry M. Reiswig .................................................................................................................................. 1361
Family Aulocalycidae Ijima
Henry M. Reiswig ............. ..... .............................. ............. ............................................................. ........ 1362
Family Uncinateridae fam. nov.
Henry M. Reiswig .................................................................................................................................. 1372
Order Lychniscosida Schrammen
Henry M. Reiswig .................................................................................................................................. 1377
Contents of Volumes 1 and 2 xiii

Family Aulocystidae Sollas

Henry M. Reiswig .................................................................................................................................. 1378
Family Diapleuridae Ijima
Henry M. Reiswig .................................................................................................................................. 1383
Order Lyssacinosida Zittel
Henry M. Reiswig .................................................................................................................................. 1387
Family Euplectellidae Gray
Konstantin R. Tabachnick ...................................................................................................................... 1388
Family Leucopsacidae Ijima
Konstantin R. Tabachnick 1435
Family Rossellidae Schulze
Konstantin R. Tabachnick 1441
Lyssacinosida incertae sedis
Konstantin R. Tabachnick ...................................................................................................................... 1506
tFossii 'Sphinctozoa': Chambered Sponges (Polyphyletic)
Baba Senowbari-Daryan & Diego C. Garcfa-Bellido ............................................................................ 1511
Bibliography of Fossil 'Sphinctozoans'
Baba Senowbari-Daryan & Diego C. Garcfa-Bellido ............................................................................ 1534
tClass Archaeocyatha Bornemann
Frans:oise Debrenne, A. Yu. Zhuravlev & Pierre D. Kruse .................................................................... 1539
Bibliography of Class Archaeocyatha
Frans:oise Debrenne, A. Yu. Zhuravlev & Pierre D. Kruse .................................................................... 1693
Annotated List of Unrecognisable Sponge Taxa and Unavailable Names
John N.A. Hooper & Rob W.M. Van Soest ............................................................................................ 1701
Bibliography of Unrecognisable Sponge Taxa
Editor Philippe Willenz .......................................................................................................................... 1707
Index of Higher Taxa to Volumes 1 and 2 ......... .............. ............. ...................... ................................... .......... xxi
Key to Symbols, Institutional Acronyms, Abbreviations

t, Denotes an extinct taxon (is used primarily for fossil taxa IGNS, Institute of Geological and Nuclear Sciences (fonnerly
mentioned in chapters chiefly dealing with Recent sponges). New Zealand Geological Survey), Lower Hutt, New Zealand.
?, placed in front of taxonomic name denotes that there ICZN, International Code of Zoological Nomenclature (see
remains uncertain amongst the contemporary scientific community Anon., 1999).
about the validity or taxonomic placement of a species, genus, IgiG, United Institute of Geology, Geophysics and Mineralogy
family or order. (collections at CSGM), Siberian Branch of the Russian Academy
AM (AMS), Australian Museum, Sydney, Australia (includ- of Sciences; Novosibirsk, Russian Federation.
ing SUP collection, fonnerly of Department of Geology and 1M (ZEV), Indian Museum (including, Zoological Survey of
Geophysics, University of Sydney, Australia). India), Calcutta, India.
ANSP, Academy Natural Sciences of Philadelphia, USA. IMGP, Geowissenschaftliches Zentrum der Universitat
BGU, Buryatan Territorial Geological Survey, Ulan-Ude, G6ttingen, Section Geobiology [fonnely Institut und Museum fUr
Russian Federation. Geologie und Palaontologie, G6ttingen, Gennany.
BMAG, Bristol Museums and Art Gallery, Bristol, United INALI, Instituto Nacional de Limnologia, Santo Tome, Santa
Kingdom. Fe, Argentina.
BMNH, The Natural History Museum, London, United INM, National Museum of Ireland, Dublin, Ireland.
Kingdom [fonnerly British Museum (Natural History)]. IORAS, Institute of Oceanology of the Russian Academy of
BMS, Buffalo Museum of Science, Buffalo, USA. Sciences, Moscow, Russian Federation.
BPBM, Bernice P. Bishop Museum, Honolulu, Hawaii, USA. IRS(c)NB, Institut royal des Sciences naturelles de Belgique,
CAS, California Academy of Sciences, San Francisco, Bruxelles, Belgium (see also KBIN).
California, USA. ISM, Imperial Science Museum, Tokyo, Japan (see TID).
CE, Departemento de Paleontologia, Universidad Complutense KBIN, Koninklijk Belgisch Instituut voor
de Madrid, Madrid, Spain. Natuurwetenschappen, Brussel, Belgium (see also IRScNB).
CEAB, Centro de Estudios Avanzados de Blanes Collection, KGU, Krasnoyarsk Territorial Geological Survey,
Blanes, Spain. Krasnoyarsk State University, Russian Federation.
CMN, Canadian Museum of Nature, Ottawa, Ontario, KMMA, Koninklijk Museum voor Midden-Afrika, Tervuren,
Canada (fonnerly NMC, National Museums of Canada). Belgium (see MRAC).
CNIGRm, Central Scientific-Researching Geological- LFM, Merseyside County Museums, Liverpool, United
Exploration Museum, St Petersburg; Federal Republic of Russia. Kingdom (fonnerly Liverpool Free Museum).
(new transliteration TsNIGRm). LMJG, Landesmuseum Joanneum Graz (Abteilung fUr
CNRS, Centre National de la Recherche Scientifique, Paris, Zoologie), Graz, Gennany.
France. LSLH, Linnean Society of London Holdings (Linnean
CSGM, Central Siberian Geological Museum (see TsSGM= Herbarium), United Kingdom.
IGiG) (now holding collections of IgiG), Novosibirsk, Russian MABA, Museo Argentino de Ciencias Naturales 'Bernardino
Federation. Rivadavia', Buenos Aires, Argentina.
DTRG, Dipartimento per 10 studio del Territorio e delle MANSP, Museum of the Academy of Natural Sciences of
sue Risorse, Universita di Genova (Dip. Te. Ris.; ex-IZUG), Philadelphia, Pennsylvania, USA.
Italy. MCN, Museo de Ciencias Naturais da Fundarr1io Zoobotiinica,
DVGI, Far East Geological Institute, Far East Institute of Porto Alegre, RS, Brasil.
Mineral Resources, Khabarovsk, Russian Federation. MCNM (MNCN-CSIC), Museo Nacional de Ciencias
DVIMS, Far-East Institut of Mineral Resources, Khabarovsk, Naturales-Consejo Superior de Investigaciones Cientfficas, Madrid,
Russian Federation. Spain.
DVTGU, Far-East Territorial Geological Survey, Far-East MCZ, Museum of Comparative Zoology, Harvard University,
branch of the Russian Academy of Sciences, Khabarovsk, Russian Cambridge, Massachusetts, USA.
Federation. MCSN, Museo Civico di Storia Naturale, Verona, Italy.
FMNH, Field Museum of Natural History, Chicago, USA. MFSN, Museo Friulano di Storia Naturale, Udine, Italy.
FZRG, Fundacao Zoobotanica Rio Grande do SuI, Museu de MG, Geological Museum, New Zealand (see VUMG).
Ciencias Naturais, Porto Alegre, Brazil. MHNG, Museum d'Histoire naturelle, Geneve, Switzerland.
GIN, Geological Institute of the Russian Academy of MNHN, Museum national d'Histoire naturelle, Paris, France.
Sciences, Moscow, Russian Federation. MNRJ, Museu Nacional, Rio de Janeiro, Brazil.
GML, Geiseltalmuseum Martin Luther, University of Halle, MOM, Musee Oceanographique de Monaco, Monaco.
Halle, Gennany. MRAC, Musee Royal de l' Afrique Centrale de Tervuren,
GSC, Geological Survey of Canada, Ottawa, Canada. Belgium (see KMMA).
HBOI/M, Harbor Branch Oceanographic Institution! MRCN, Museu Rio-Grandense de Ciencias Naturais, Porto
Museum, Fort Pierce, Florida, USA. Alegre, Rio Grande do SuI, Brasil.
HMN, Hancock Museum, Newcastle-upon-Tyne, United MSNG, Museo Civico di Storia Naturale 'Giacomo Doria',
Kingdom. Genoa, Italy.

xv
xvi Key to Symbols, Institutional Acronyms, Abbreviations

MT Por, Museo e Istituto di Zoologfa Sistematica SNIIGGIMS, Siberian Scientific Research Institute of
dell 'Universita di Torino, Italy (Porifera collection). Geology, Geophysics and Mineral Resources, Novosibirsk, Federal
MY, Million years. Republic of Russia.
MYA, Million years ago. SUP, Department of Geology and Geophysics, The University
MZUS, Musee de Zoologie de l'Universite de Strasbourg, of Sydney, Australia (now housed in The Australian Museum;
France. see AM).
NHRM, Naturhistoriska Riksmuseet, Stockholm, Sweden. TEM, Transmission electron microscopy.
NIGP, Nanjing Institute of Geology and Paleontology of the TIU, Imperial University Museum, Tokyo, Japan.
Chinese Academy of Sciences, Nanjing, China. TsNIGRm, new transliteration for CNIGRm.
NMB, Naturhistorisches Museum zu Basel, Basel, TsSGM, new transliteration for CSGM.
Switzerland. UA, Department of Geology, University of Alberta, Edmonton,
NMNZ, Te Papa, Museum of New Zealand, Wellington, New Alberta, Canada.
Zealand (formerly National Museum of New Zealand, and for- UA, University of Adelaide (see SAM).
merly Dominion Musuem). UAM, University of Alaska Museum, Alaska, USA.
NMS, National Museum of Scotland, Edinburgh, Scotland. UBC, University of British Columbia, Vancouver, Canada.
NMV, Museums of Victoria, Melbourne, Australia (formerly UCLZ, University College of London, Zoology.
National Museum of Victoria). UCPM, University of California Paleontology Museum,
NSM, National Science Museum, Tokyo, Japan. Berkeley, California, USA.
NTM, Northern Territory Museum of Arts and Sciences, UFRJPOR, Universidade Federal do Rio de Janeiro, Brazil.
Museums and Art Galleries of the Northern Territory, Darwin, UKL, University of Kansas, Lawrence, USA.
Australia. UMML, Marine Invertebrate Museum, University of Miami,
NYSM, New York State Museum, Albany, New York State, Rosenstiel School of Marine and Atmospheric Sciences.
USA. USNM, National Museum of Natural History, Smithsonian
NZGS, New Zealand Geological Survey, Wellington, New Institution, Washington D.C., USA (formerly United States
Zealand. National Museum).
NZMW, National Zoological Museum Warsaw, Poland. UTBEG, Texas Memorial Museum, Bureau of Economic
PGO, Zabsibgeologiya, Novokuznetsk, Russian Federation. Geology, Austin, Texas, USA.
PGU, Primor'ye Territorial Geological Survey; Khabarovsk, UWU, University of West Indies, Mona, Jamaica, West Indies
Russian Federation. (Zoology Department) [MSUJ).
PIN, Paleontological Institute of the Russian Academy of VSEGEI, All-Russian Geological Institute, St Petersburg,
Sciences, Moscow, Russian Federation. Russian Federation (transfer of collections to TsNIGRm in
PMJ, Phyletisches Museum, Jena, Germany. progress).
PU, Princeton University Bedfords' Collections (now housed VUMG, Victoria University of Wellington, Museum of
in USNM Washington). Geology, Wellington; New Zealand.
QM, Queensland Museum, Brisbane and Townsville (includ- WAM, Western Australian Museum, Perth, Australia.
ing MTQ, Museum of Tropical Queensland). YPM, Yale University, Peabody Museum of Natural History,
RASMAS, Rosenstiel School of Marine and Atmospheric Yale University, New Haven, Connecticut, USA.
Sciences (see UMML). ZEV, (see IM).
RIB, (see IRScNB). ZIL, Zoological Institute of the Russian Academy of
RMM, Redpath Museum, McGill University, Montreal, Sciences, St Petersburg, Russian Federation.
Quebec, Canada. ZMA, ZoOiogisch Museum, Universiteit van Amsterdam,
RMNH, Nationaal Natuurhistorisch Museum, Leiden, The Amsterdam, The Netherlands.
Netherlands (formerly Rijksmuseum van Natuurlijke Historie). 2MB, Zoologisches Museum fiir Naturkunde an der
ROM, Royal Ontario Museum, Toronto, Canada. Universitat HumboldVzu Berlin, Berlin, Germany.
RSME, Royal Scottish Museum Edinburgh, Scotland. ZMH, Zoologdches Museum flir Hamburg, Hamburg,
SAMP, South African Museum, Pretoria, Republic of South Germany. .
Africa. ZMUB, Zoological Museum, University of Bergen, Norway.
SAM, South Australian Museum, Adelaide, Australia (includ- ZMUC, Zoologisk Museum Universitet Copenhagen,
ing collections of UA - The University of Adelaide). Copenhagen, Denmark.
SAMC, South African Museum, Capetown, Republic of ZMUZ, Zoologisches Museum der Universitat, Ziirich,
South Africa. Switzerland.
SBR, Station Biologique de Roscoff, France. ZPAL, Institute of Paleobiology of the Polish Academy of
SDCC, Private collections of Steve de C. Cook (destined for Sciences, Warsaw, Poland.
theNMNZ). ZRS, Zoologiska Rijkmuseum, Stockolm, Sweden.
SEM, Scanning electron microscopy. ZSGGU, West Siberian State Geological Trust (=WSSGE;
SME, Station Marine d'Endoume, Marseille, France. =WSSGT, =ZSTGU) Novokuznetsk, Russian Federation.
SMF, Natur-Museum und Forschunginstitut Senckenberg,
Frankfurt-am-Main, Germany.
Acknowledgements

The Systema Porifera project involved an extensive collabora- Rob Van Soest similarly acknowledges the technical assistance of
tion amongst many experts over many years. Without this coopera- Elly Beglinger, Henk de Haas, Mario de Kluijver, Raquel Gomez,
tion this book would never have been attempted. The scientific and Nicole de Voogd (all ZMA Amsterdam), and Bert Hoeksema
bibliographic editors, John Hooper, Rob Van Soest and Philippe (RMNH Leiden). Philippe Willenz thanks Laetitia Despontin for
Willenz, are most grateful to all the authors for their perseverance persevering assistance while amending the bibliographic refer-
and cooperation over the long gestation of this project. We would ences database from original documents.
particularly like to thank our editor at Plenum, Joanna Lawrence, The editors also acknowledge the continued support from
for her continued support and patience during the many and varied their respective institutions: the Queensland Museum, Brisbane,
challenges we overcame to get this work to press, and for main- the Zoi:ilogisch Museum, Universiteit van Amsterdam, and the
taining faith in this collaborative project. The existence and contin- Institut royal des Sciences naturelles de Belgique, Bruxelles.
uance of this sponge network is a positive and essential tool for the Financial support during the course of this project was received
future - to take the basic infrastructure and the taxonomic hypothe- from the following: John Hooper - from AstraZeneca R&D
ses we present here into further realms. Griffith University, Brisbane, and the Australian Biological
During the development of the Systema three workshops were Resources Study, Canberra; Rob Van Soest and colleagues - from
held (London, Paris and Brisbane) to promote collaboration EU-MAS3-CT97-0l44 (to Rob Van Soest), LSF NHM London
between the authors and to resolve both theoretical and practical (Vanessa Pike, to Rob Van Soest, LSF MNHN Paris (to Dirk
difficulties associated with the project. For these meetings the edi- Erpenbeck), LSF ZMUC Copenhagen (to Dirk Erpenbeck), and
tors are most grateful to the host institutions and sponsors: The NWO-ALW 8009.34.003 (to Rob Van Soest).
Natural History Museum, London, including the Linnean Society The authors and editors gratefully acknowledge the following
of London; the Museum National d'Histoire Naturelle, Paris; and publishers for pennission to reproduce copyrighted illustrations in
the Queensland Museum, Brisbane, including AstraZeneca R&D this work. Full citations of literature are provided in the various
Griffith University, Brisbane, and the Ian Potter Foundation, bibliographies of this work.
Melbourne. We are particularly grateful to Clare Valentine for her
continued support in accessing the vast BMNH resources over AA Balkema, Rotterdam, The Netherlands - Soest, R.W.M. Van, Kempen,
recent years. Th.M.G. Van & Braekman, J.C. (Eds), 1994, 'Sponges in Time and
Space' -Sara, M. & Burlando, B. Pp 111-116 (figs 1,4); Reiswig, H.M. &
The curators and collection managers of the major museum Mehl, D. Pp 151-165 (figs 1-3,9).
collections throughout the world are gratefully acknowledged for American Museum of Natural History, New York, USA - Bulletin of the
providing many loans of sponge types and other material over the American Museum of Natural History, Vol. 120, Finks, R.M., 1960
many years of this project. We are especially grateful to Penny (fig. 63, p. 108).
Association Argentina de Ciencias Naturales, Buenos Aires, Argentina -
Berents (AM Sydney), Clare Valentine and Shirley Stone (BMNH
Physis, Vol. 26(71): 129-140, Bonetto, A.A. & Ezcurra, I.D., 1966 (all
London), Beatrice Burch (BPBM Honolulu), A.K. MandaI (1M figures). Vol. 28(77): 351-357, Bonetto, A.A. & Ezcurra de Drago, I.D.,
Calcutta), C. O'Riordan (INM Dublin), Ruth Desqueyroux- 1969 (all figures).
Faundez (MNHG Geneva and Johanneum Museum Graz), Association of Australasian Palaeontologists, Canberra, Australia -
Takomura Hoshino [deceased] (MMBS Hiroshima), Claude Levi Alcheringa, Vol. 2: 27--47, Kruse, P.D. 1978 (figs 2D, 4A, 7B, l1A,
lIB). Vol. 10: 235-278, Debrenne, F. & Kruse, P.D. 1986 (figs 6A, 6B,
(MNHN Paris), Ms. Stock (NHM Basel), Helmut Sattmann (NHM 15A, 15B, 19A, 21A, 2IB, 22A, 22B). Vol. 17: 179-183, Debrenne, F.
Wien), Urs Rahm (NMB Basel), B.R. Stuckenberg (NM Natal), Zhuravlev, A.Yu. & Gravestock, D.l. 1993 (fig. 3A-B). Vol. 18: 1-54,
Frank Climo (NMNZ Wellington), Susan Boyd (NMV Zhuravlev, AYu. & Gravestock, D.I. 1994 (fig. 12B, E). Memoirs of the
Melbourne), Frank von Knorre (PMJ Jena), Bert Hoeksema and Association of Australasian Palaeontologists Vol. 1: 93-120, Pickett, J.
1983 (figs 4D-E). Vol. 2: 1-139, Gravestock, D.1. 1984 (figs 32L, 50E,
J.C. Den Hartog (deceased) (RMNH Leiden), Susan Chambers
55H, 60A, 62A, 62D, 62E, 62F).
(RSME Edinburgh), Shane Parker [deceased], Wolfgang Zeidler Australian Geological Survey Organisation, Canberra, Australia -
and (SAM Adelaide), Nicole Boury-Esnault and Jean Vacelet Memoirs of the Association of Australian Palaeontologists, Vol. 1:
(SME Marseille), Manfred Grasshoff (SMF Frankfurt), Klaus 85-92, Pickett, J.w. & Jell, P.A., 1983 (fig. 4A-C).
RUtzler and Kathleen Smith (USNM Washington), Loisette Marsh Birkhiiuser Verlag AG, Basel, Switzerland - Experentia Suppiementum,
No. 28, Wiedenmayer, F. 1977, 'Shallow water sponges of the western
and Jane Fromont (WAM Perth), Deiter KUhlmann (ZMB), and Bahamas' (pis 4.3, 9.2).
Ole Tendal (ZMUC Copenhagen). Board of the Museums and Art Galleries of the Northern Territory, Darwin,
The scientific editors are also grateful to the very many refer- Australia - The Beagle, Occasional Papers of the Northern Territory
ees for providing positive comments and suggestions to improve Museum of Arts and Sciences, Vol. 3(1): 167-189, Hooper, J.N.A 1986
(fig. 40). The Beagle, Records of the Northern Territory Museum of Arts
the various chapters in this book, ensuring that these were both sci-
and Sciences, Vol. 6(1): 133-140, Hooper, J.N.A. & Krasochin, V. 1989
entifically accurate and provided a practical working tool for (fig. 1). Vol. 8(1): 27-36, Hooper, J.N.A. et at. 1991 (figs 1-9). Vol. 12(1):
sponge systematics, wherever possible. 151-176, Bergquist, P.R. & Kelly-Borges, M. 1995 (figs 2a, e).
John Hooper thanks the following for technical assistance Board of the Museum of Victoria, Melbourne, Australia - Memoirs of the
during the production of this work: Stephen Cook, John Kennedy, Museum of Victoria, Vol. 50(1): 1-242, Wiedenmayer, F., 1989 (pI. 8,
fig. 11, pI. 9, fig. 4, pI. 28, figs 3--4).
Gert Worheide, Daniel Edson, Sue List-Armitage, Paula Board of the Queensland Museum, Brisbane, Australia - Memoirs of the
Thomkins, Brett Krueger and Monika Schlacher, and Justin Gough Queensland Museum, Vol. 38(1): Bergquist, P.R. 1995 (figs 17c, 20b, 22b,
for support developing an accessible web site (all QM Brisbane). 22c, 22d, 24a, 25c, 23a, 23b). Vol. 40: 1-626, Hooper, J.N.A. 1996 (figs 1,

xvii
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Geologisch-Paliiontologische Mitteilungen, Sonderband, Vol. 4: 1-55, et al., 2000 (figs 1,4,6-8,10, 14, 15, 17,23-25,27-29,31,33-40,46).
Mehl, D., 1996 (figs 2, 3, 4, 8, pI. 3, figs 1, 3, pI. 5, fig. 2, pI. 6, fig. 1). National Institute of Water and Atmospheric Research (NIWA) Ltd.,
Institut flir Geologie und Palaontologie der Universitat der Stuttgart, Wellington, New Zealand - Department of Scientific and Industrial
Germany - Profil Stuttgart, 1997, Krautter, M. 1997 (fig. 63). Research Bulletin, New Zealand Oceanographic Institute Memoir, No.
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Germany - Paliiontologische Zeitschrift, Vol. 42(1-2): 13-32, Rietschel, 1-85, Bergquist, P.R. 1970 (pI. 5, fig. B). No. 100: 1-98, Dawson, E.W
1968 (pI. 1, fig. 1). 1993 (cover photograph). No. 107: 1-53, Bergquist, P.R 1996 (cover
Institut ftir Paillontologie der Universitat Erlangen-Niimberg, Germany - photograph, pI. 5D, pI. 4F, pI. 2C, pI. IB, fig. 4, pI. 7D).
Facies, Vol. 38: 1-88, Worheide, G. 1998 (pI. 20, figs 6-7, pI. 21, fig. 16). National and Kapodistrian University of Athens, Department of Biology -
Institut de Ciencies del Mar, CSIC, Barcelona, Spain - Scientia Marina, 'Kalymnos and the secrets of the sea' - Castritsi-Catharius, J., 1998 (fig.
Vol. 57(4): 283-306, Diaz, M.e. et aI., 1993 (fig. 34). Vol. 57(4): 109).
381-393, Rtitzler, K. & Smith, K.P., 1993 (fig. 5). Northern Territory Geological Survey, Darwin, Australia - Report of the
Institut royal des Sciences naturelles de Belgique, Bruxelles, Belgium - Northern Teritory Geological Survey, Vol. 7: 1-58, Kruse, P.D., 1990 (pis
Cercle hydrobiologique de Bruxelles, Vol. 11(1): 1-56, Brien, P., 1967 25E,26A).
(all figures). Revue de Zoologiye et de Botanique Africaines, Vol. 88(3): Palaeontologia Polonica, Warszawa, Poland - No. 55: 9-36, Wrona, R &
585-624, Brien, P., 1974 (all figures). Zhuravlev, A Yu. 1996 (pI. 7, figs 2, 5).
Instituto Espanol Oceanografia - Boletin Instituto Espanol de Senckenbergiana Maritima, Frankfurt am Main, Germany - Vol. 30 (3/6):
Oceanografia, Vol. 6(4), No. 324: 103-154, Biblioni, M.A., 1981 (fig. 7). 161-221, Schonberg, C.H.L. 2000 (pI. 7, fig. 40).
Izdatelsvo Nauka, Moscow, Russia - Akademia Nauk SSSR, (1959): 1-236, SIR Publishing, Royal Society of New Zealand, Wellington, New Zealand-
Koltun, V.M. 1959 (fig. 78). Vol. 92: 6-116, Koltun, V.M. 1964 (fig. 19). New Zealand Journal of Zoology, Vol. 7, Bergquist, P.R. 1980 (figs la,
Koeltz Scientific Books, Koenigstein, Germany - Theses Zoologicae, Vol. 4a-d, 5c-d, 6a-c, 7a-b, 8e, lOb, d, 11 a, 12a, c-d, 13a-c, e, 14a-b, 15a,
23, Barthel, D. & Tendal, O.S. 1994 (fig. 14). 16a-b, 17d-e, 19c, 19f, 22f, 23a, 23b-c, 23f, 24b-c, 25b).
Acknowledgements xix

Smithsonian Insitution, Washington, USA - Smithsonian Institution Press, University of Amsterdam, The Netherlands - Bijdragen tot de Dierkunde,
Vos, L. De et al., 1991 'Atlas of sponge morphology' (p. 106, pI. 54). Vol. 64(3): 163-192, Van Soest, RW.M. et al., 1994 (figs 3, 6).
Smithsonian Contributions to Zoology, Vol. 598: 1-47, Alvarez, B. et aI., University of Amsterdam, Institute of Taxonomic Zoology (Zoological
1998 (fig. 11). Museum), The Netherlands - Beaufortia, Vol. 36 (6): 81-165, Weerdt,
Societe geologique de France, Paris, France - Bulletin de la Societe W.H. De, 1986 (figs 5b, 19b). Vol. 40(2): 15-62, Soest R.W.M. Van etal.,
Geologique de France, (8), Vol. 4, Debrenne, F. & Zhi-Wen Jiang, 1989 1990 (figs 22, 60, 69, 73).
(pI. II, fig. 6). University of Hawaii Press, Honolulu, USA - Pacific Science, Vol. 19 (2):
Societe zoologique de France, Paris, France - Bulletin de la Societe 123-204, Bergquist, P.R. 1965 (figs la, Ib, 7a, 8b).
zoologique de France, Vol. 103: 443-448, Levi, C. & Levi, P., 1979 University of Iowa, USA - Journal of Paleontology, Vol. 41(2): 511-515,
(figs 1-5). Rigby, J.K. 1967 (figs 3-4). Vol. 49(2): 329-339, Rigby, J.K. & Nitecki,
Springer-Verlag GmbH & Co. KG, Germany - Reitner, J. & Keupp, H. M.H. 1975 (pI. 1, figs 1-2). Vol. 52(3): 705-716, Rigby, lK. 1978 (pI. 1,
(Eds), 1991 'Fossil and Recent Sponges' - Reiswig, H.M., Pp 7-20 figs 1,3,5, fig. 3). Vol. 53: 475-493, Rigby J.K. eta/. 1979 (pI. I, figs 1-3,
(fig. lA-D); Volmer-Ribeiro, C. & Reitner, J., pp. 121-133 (ali figures); pI. 2, fig. 1). Vol. 65(1): 38-44, Rigby, J.K. 1991 [fig. 3 (1,3,4), fig. 4).
Vacelet, J., Pp. 252-265 (figs Ib, d, 2b). Marine Biology, Vol. 103: University of Kansas Paleontological Institute, USA - University of Kansas
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Morphologie und Okologie der Tiere (now Oecologia), Vol. 55(1): 1-82, (fig. 11(1-3)).
Riitzler, K., 1965 (fig. 11). University of Miami, USA - Bulletin of Marine Science, Vol. 59(2):
Staringia, Ordovicische zwetjsteensponzen, Grondboor & Hammer, 370-392, Sara, M. & Bavestrelio, G. 1996 (figs 2, 3, 4, 8, 11). Vol.
Vol. 9, Jaargang 55(1), Rhebergen, Eggink, Koops, Rhebergen, 2001 58(3): 764-774, Reiswig, H.M. & Tsurumi, M. 1996 (figs 3, 4, 5,
(figs 1, 13,46). 10,20,21).
Station Biologique de Roscoff, France - Cahiers de Biologie Marine, University of Tennessee, Department of Geological Sciences, USA -
(1967), Borojevic, R. 1967 (figs 3, 4, 11). Broadhead, T.W. (ed.) 'Sponges and spongiomorphs; notes for a short
Suid-Afrikaanse Museum, Cape Town, South Africa - Annals of the South course' Studies in Geology (Knoxville) Vol. 7: 70-89, Rigby, 1.K., 1983
African Museum, Vol. 67(8): 331-361, Debrenne, F. (figs 3a,b, 7a). (figs lB-D, 6A-D, 7B-C).
The Palaeontological Association, Aberystwyth, Wales, UK University of Wales Swansea, School of Biological Sciences - Journal of
Palaeontology, Vol. 24 (2), Debrenne, F. & James, N.P. 1981 (pI. 50, Natural History, Vol. 31: 315-328, Sandford, F. & Keliy-Borges, M.,
fig. 5, pI. 52, figs 1,5, pI. 53, figs 1,4). 1997 (figs 2d, 3a).
The Natural History Museum, London, UK - Burton, M., 1963, 'A revision Verlag Friedrich Pfeil - Miinchner Geowissenschaftliche Abhandlungen,
of the classification of the calcareous sponges' (fig. 163). No. 21: 1-326, Senowbari-Daryan, B., 1990 (various figures).
Transactions of the Royal Society of South Africa, Vol. 37 (1): 1-72, Yale University, Peabody Museum of Natural History, New Haven,
Levi, C. 1963 (pI. 4, fig. G). USA - Postilla, 137: 1-39, Hartman, W.D. 1969 (pI. 1, figs 1-2, text-figs I,
University of Alberta, Department of Earth & Atmospheric Sciences, 2,7,12).
Canada - Canadian Journal of Earth Sciences, Vol. 13(1): 120-125, Zoological Society of London, UK - Journal of Zoology, Vol. 149:
Rigby, J.K. 1976 (figs 4-6, 8A). Vol. 23(3): 344-349, Rigby, J.K. & 415-531, Bakus, J., 1966 (text-figs 16, 23). Vol. 234: 301-323, Kelly-
Goodbody, Q.H. 1986 (fig. 2A, J). Borges, M. & Bergquist, P.R., 1994 (pI. la).
SYSTEMA PORIFERA
A Guide to the
Classification of Sponges
VOLUME 2
SYSTEMA PORIFERA
A Guide to the
Classification of Sponges
VOLUME 2

Edited by

John N. A. Hooper
Queensland Museum
South Brisbane, Queensland, Australia

and

Rob W. M. Van Soest

Zoological Museum
University ofAmsterdam
Amsterdam, Netherlands

Bibliographic Editor for Recent Taxa

Philippe Willenz
Royal Institute of Natural Sciences of Belgium
Brussels, Belgium

K1uwer Academic / Plenum Publishers

New York, Boston, Dordrecht, London, Moscow
Library of Congress Cataloging-in-Publication Data

Systema Porifera: a guide to the classification of sponges/scientific editors, John N,A.

2002072781

ISBN 0-306-47260-0

Published in two volumes:

Volume 1: Introductions and Demospongiae
Volume 2: Calcarea, Hexactinellida, Sphinctozoa, Archaeocyatha, unrecognizable taxa, and Index of higher taxa

© 2002 Kluwer Academic / Plenum Publishers, New York

233 Spring Street, New York, New York 10013

http://www.wkap.nl/

10 9 8 7 6 5 4 3
A c.I.P. record for this book is available from the Library of Congress

All rights reserved

No part of this book may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical,
photocopying, microfilming, recording, or otherwise, without written permission from the Publisher, with the exception of any material supplied
specifically for the purpose of being entered and executed on a computer system, for exclusive use by the purchaser of the work.
Preface

Sponges are among the most ancient of multicellular these many new taxa, to correct the many nomenclat-
animals alive on the planet today. Ecologically and eco- ural and taxonomic mistakes of the past, and to provide
nomically, they are among the most highly successful a sound platform for the future development of sponge
life forms that have ever existed. First known from the systematics. For many groups the Systema is a major
Precambrian, and well established by the Cambrian, revision, and for all groups it provides a sound frame-
they were the major reef builders during the Palaeozoic work of sponge systematics based on phylogenetic
and Mesozoic Eras. Today they are highly diverse, with principles. Perhaps more importantly, this project also
an estimated 15,000 species living in all Recent marine provides a practical tool to allocate taxa within this
and freshwater habitats worldwide, but these undoubt- framework, including keys, comprehensive illustra-
edly represent only a small proportion of those that have tions and descriptions of the major characters used to
ever lived. Sponge body plans do not appear to have classify sponges - something that has not previously
changed significantly over the many millennia, such been attempted.
that some species alive today appear to be almost identi- The Systema is published in two volumes.
cal to those that once lived in Earth's ancient seas. Volume 1 includes a general introduction and the Class
Consequently, sponges are often regarded as ultra-con- Demospongiae. Volume 2 includes Classes Calcarea
servative in their morphology, presumably as successful and Hexactinellida, the fossil Class Archaeocyatha,
models of early evolutionary experiments. In contrast, fossil 'Sphinctozoa', and an annotated list of unrecog-
the living fauna is widely diverse in many other biologi- nisable taxa and unavailable names. We describe
cal features, especially in their biochemical con- approximately 680 genera of sponges in 127 families,
stituency that is currently of major interest to the 25 orders and three classes of the living fauna. In addi-
pharmaceutical industry. It is this latter feature, and also tion, approximately 1000 fossil genera in 245 families,
the renewed interest in marine biodiversity worldwide, 30 orders and six 'classes' are also covered. The living
that has attracted a major resurgence of studies on fauna is dealt with far more comprehensively than the
sponges with the consequence that over the past two fossil fauna, with overviews of the fossil classification
decades researchers have discovered a living fauna included mainly in an attempt to relate these two classi-
about twice the size of that published in the literature. fications more closely. A far more comprehensive treat-
Sponge systematics is generally perceived as ment of fossil sponges is currently in preparation for the
appallingly difficult. These difficulties stem partly revised edition of the Treatise on Invertebrate
from the sponges themselves, being highly plastic and Palaeontology (J.K. Rigby et al., editors).
easily susceptible to environmental modification and The Systema project was only possible to achieve
frequent character losses. They also stem, perhaps through an intensively collaborative effort amongst 45
more so, from the inadequacies and inconsistencies researchers from 17 countries, spanning about six years of
of the taxonomic framework. This includes a previ- research effort. We pride ourselves to have among our
ously fragmented and uncoordinated research effort, contributors the major contemporary spongologists,
whereby only a few higher groups have had contempo- including Patricia R. Bergquist, Nicole Boury-Esnault,
rary revisions applied to them, and with the remainder Fran90ise Debrenne, Claude Levi, Henry Reiswig,
largely unworked since the late 19 th century. In tum, Joachim Reitner, Klaus Rtitzler, Michele Sara, Maria J.
this has led to increasing difficulties faced by new Uriz and Jean Vacelet, as well as several members of the
workers - among them a growing number from newly new generation of sponge researchers. The Systema
developing countries - to access the vast and mostly Porifera project is not an end - but a sound beginning for
ancient literature. this new generation to build on what we propose here.
The Systema Porifera project was devised to
revisit the poriferan higher taxa (genus level and John N.A. Hooper, Brisbane
above, based on type species data), to accommodate Rob W.M. Van Soest, Amsterdam

v
Contents of Volumes 1 and 2
VOLUME 1

Key to Symbols, Institutional Acronyms, Abbreviations ................................................................................. xv

Acknowledgements ........................................................................................................................................ xvii
Introduction
John N.A. Hooper & Rob W.M. Van Soest .................................................................................................. 1
Bibliography of Introductions
Editor Philippe Willenz ...................... ..................... ................................... .................................................. 4
Phylum Porifera Grant
John N.A. Hooper, Rob W.M. Van Soest & Fran~oise Debrenne ................................................................ 9
Class Demospongiae Sollas
John N.A. Hooper & Rob W.M. Van Soest ................................................................................................ 15
Bibliography of Class Demospongiae
Editor Philippe Willenz ............................................ ........ ................................................. ........ ................. 19
Non-Lithistid Fossil Demospongiae - Origins of their Palaeobiodiversity and Highlights in
History of Preservation
Joachim Reitner & Gert Worheide ............................................................................................................. 52
t'Class Stromatoporoidea' Nicholson & Murie, Stromatoporoids
Alex Cook .. ..... ..................................................... ...................................................................................... 69
Order Homosclerophorida Dendy. Family Plakinidae Schulze
Guilherrne Muricy & Maria Christina Diaz ..................... ......... ............. .............. ................. ..................... 71
Order Spirophorida Bergquist & Hogg
Rob W.M. Van Soest & John N.A. Hooper ............................................................................................... 83
Family Tetillidae Sollas
Rob W.M. Van Soest & Klaus Riitzler ............................................................. ................. .................. ....... 85
Family Samidae Sollas
Rob W.M. Van Soest & John N.A. Hooper ................................................................................................ 99
Family Spirasigmidae Hallmann
John N.A. Hooper & Rob W.M. Van Soest .............................................................................................. 102
Order Astrophorida Sollas
John N.A. Hooper & Rob W.M. Van Soest .............................................................................................. 105
Family Ancorinidae Schmidt
Maria J. Uriz ............................................................................................................................................ 108
Family Calthropellidae Lendenfeld
Rob W.M. Van Soest & John N.A. Hooper .............................................................................................. 127
Family Geodiidae Gray
Marfa J. Uriz ............................................................................................................................................ 134
Family Pachastrellidae Carter
Manuel Maldonado ....................................................... ................................................................. .......... 141
Family Thrombidae Sollas
Marfa J. Uriz ............................................................................................................................................ 163
Astrophorida Incertae Sedis
John N.A. Hooper & Manuel Maldonado ................................................................................................ 165
Order Hadromerida Topsent
John N.A. Hooper & Rob W.M. Van Soest .............................................................................................. 169
Family Clionaidae D'Orbigny
Klaus Riitzler ........................................................................................................................................... 173

vii
viii Contents of Volumes 1 and 2

Family Hemiasterellidae Lendenfeld

John N.A. Hooper .................................................................................................................................... 186
Family Placospongiidae Gray
Klaus Riitzler .............................................................................................................................. ............. 196
Family Polymastiidae Gray
Nicole Boury-Esnault ............................................................................................................................... 201
Family Spirastrellidae Ridley & Dendy
Klaus Riitzler ........... ..................... ......... ..... ..................... ................................................ ..... ..... ... ........... 220
Family Stylocordylidae Topsent
Rob W.M. Van Soest ................................................................................................................................ 224
Family Suberitidae Schmidt
Rob W.M. Van Soest ................................................................................................................................ 227
Family Tethyidae Gray
Michele Sara ....... ..... ............. ...................... .......................... ........... ....... ... ..... .............. ... ..... .... ............... 245
Family Timeidae Topsent
Klaus Riitzler .. .......... ............................. ......... ............................... ........ ............. .......... ... ........................ 266
Family Trachycladidae Hallmann
John N.A. Hooper & Rob W.M. Van Soest .............................................................................................. 268
Family Acanthochaetetidae Fischer
Klaus Riitzler & Jean Vacelet ................................................................................................................... 275
Family Sollasellidae Lendenfeld
Rob W.M. Van Soest .............................................................................................................................. 279
Family Alectonidae Rosell
Klaus Riitzler ....... ............. ...................... ... ............................... ..... .......................... ................................ 281
Order Chondrosida Boury-Esnault & Lopes. Family Chondrillidae Gray
Nicole Boury-Esnault ............................................................................................................................... 291
'Lithistid'Demospongiae
Andrzej Pisera & Claude Levi ..................................................... ...... ..... ..................... ..... ....................... 299
Family Scleritodermidae Sollas
Andrzej Pisera & Claude Levi ............... .............. ............ ..... ................................................ ......... .......... 302
Family Corallistidae Sollas
Andrzej Pisera & Claude Levi ............... ......... ..... .... ........ .............. .......................... ............. ................... 312
Family Pleromidae Sollas
Andrzej Pisera & Claude Levi ................................................................................................................. 321
Family Theonellidae Lendenfeld
Andrzej Pisera & Claude Levi ................................................................................................................. 327
Family Siphonidiidae Lendenfeld
Andrzej Pisera & Claude Levi ................................................................................................................. 338
Family Neopeltidae Sollas
Andrzej Pisera & Claude Levi .. ......... ..... .................................. ......... ..... ................. ............. ................... 344
Family Azoricidae Sollas
Andrzej Pisera & Claude Levi ................................................................................................................. 352
Family Desmanthidae Topsent
Andrzej Pisera & Claude Levi ................................................................................................................. 356
Family Vetulinidae Lendenfeld
Andrzej Pisera & Claude Levi .......................................................................................... ....................... 363
Family Phymatellidae Schrammen
Andrzej Pisera & Claude Levi ... ......................... ......... .............. .............................................................. 366
Family Isoraphiniidae Schrammen
Andrzej Pisera & Claude Levi ................................................................................................................. 374
Family Macandrewiidae Schrammen
Andrzej Pisera & Claude Levi ................................................................................................................. 377
Contents of Volumes 1 and 2 ix

Family Phymaraphiniidae Schrammen

Andrzej Pisera & Claude Levi ................................................................................. ............ .................... 380
Lithistids Incertae Sedis
Andrzej Pisera & Claude Levi ................................................................................................................. 384
tFossil Lithistids: An Overview
Andrzej Pisera .......................................................................................................................................... 388
Order Poecilosclerida Topsent
John N.A. Hooper & Rob W.M. Van Soest .............................................................................................. 403
Suborder Microcionina Hajdu, Van Soest & Hooper
John N.A. Hooper .................................................................................................................................... 409
Family Acarnidae Dendy
John N.A. Hooper .................................................................................................................................... 412
Family Microcionidae Carter
John N.A. Hooper .................................................................................................................................... 432
Family Raspailiidae Hentschel
John N.A. Hooper .................................................................................................................................... 469
Family Rhabderemiidae Topsent
John N.A. Hooper .................................................................................................................................... 511
Suborder Myxillina Hajdu, Van Soest & Hooper
Rob W.M. Van Soest ................................................................................................................................ 515
Family Chondropsidae Carter
Rob W.M. Van Soest ................................................................................................................................ 521
Family Coelosphaeridae Dendy
Rob W.M. Van Soest ................................................................................................................................ 528
Family Crambeidae Levi
Rob W.M. Van Soest 547
Family Crellidae Dendy
Rob W.M. Van Soest 556
Family Dendoricellidae Hentschel
Rob W.M. Van Soest ............................................................................................ .................................... 567
Family Desmacididae Schmidt
Rob W.M. Van Soest ................................................................................................................................ 572
Family Hymedesmiidae Topsent
Rob W.M. Van Soest ................................................................................................................................ 575
Family Iotrochotidae Dendy
Rob W.M. Van Soest ................................................................................................................................ 594
Family Myxillidae Dendy
Rob W.M. Van Soest ................................................................................................................................ 602
Family Phellodermidae fam. nov.
Rob W.M. Van Soest & Eduardo Hajdu ................................................................................................... 621
Family Tedaniidae Ridley & Dendy
Rob W.M. Van Soest ................................................................................................................................ 625
Suborder Mycalina Hajdu, Van Soest & Hooper
Rob W.M. Van Soest & Eduardo Hajdu ................................................................................................... 633
Family Cladorhizidae Dendy
Eduardo Hajdu & Jean Vacelet ................................................................................................................ 636
Family Desmacellidae Ridley & Dendy
Eduardo Hajdu & Rob W.M. Van Soest .... ........................................... ...... .............................................. 642
Family Guitarridae Dendy
Eduardo Hajdu & Clea Lerner ................................................................................................................. 651
Family Esperiopsidae Hentschel
Rob W.M. Van Soest & Eduardo Hajdu ............ ............................................... .......................... .............. 656
x Contents of Volumes 1 and 2

Family Hamacanthidae Gray

Eduardo Hajdu ........ .............. ............ .......... ........ .............. ... ....................... ........ ..... ........ ............. ........... 665
Family Mycalidae Lundbeck
Rob W.M. Van Soest & Eduardo Hajdu ................................................................................................... 669
Family Merliidae Kirkpatrick
Eduardo Hajdu & Rob W.M. Van Soest ................................................................................................... 691
Family Podospongiidae De Laubenfels
Michelle Kelly & Toufiek Samaai ........................................................................................................... 694
Family Isodictyidae Dendy
Eduardo Hajdu & Gise1e Lobo-Hajdu ..................................................................................................... 703
Suborder Latrunculina subord. nov. incertae sedis
Michelle Kelly & Toufiek Samaai ........................................................................................................... 707
Family Latruncu1iidae Topsent
Toufiek Samaai & Michelle Kelly ........................................................................................................... 708
Order Halichondrida Gray
Rob W.M. Van Soest & John N.A. Hooper ............................................................................................. 721
Family Axinellidae Carter
Belinda Alvarez & John N.A. Hooper ..................................................................................................... 724
Family Bubaridae Topsent
Belinda Alvarez & Rob W.M. Van Soest .................................................................................................. 748
Family Desmoxyidae Hallmann
John N.A. Hooper .................................................................................................................................... 755
Family Dictyonellidae Van Soest, Diaz & Pomponi
Rob W.M. Van Soest, Dirk Erpenbeck & Belinda Alvarez ...................................................................... 773
Family Halichondriidae Gray
Dirk Erpenbeck & Rob W.M. Van Soest .................................................................................................. 787
Order Agelasida Hartman
Rob W.M. Van Soest & John N.A. Hooper ............................................................................................. 817
Family Age1asiidae Verrill
Rob W.M. Van Soest ................................................................................................................................ 819
Family Astrosc1eridae Lister
Jean Vacelet .............................................................................................................................................. 824
Order Haplosclerida Topsent
Rob W.M. Van Soest & John N.A. Hooper .............................................................................................. 831
Suborder Haplosclerina Topsent
Rob W.M. Van Soest & John N.A. Hooper .............................................................................................. 833
Family Cal1yspongiidae de Laubenfels
R. Desqueyroux-Faundez & C. Valentine ................................................................................................ 835
Family Chalinidae Gray
Wallie H. De Weerdt ................................................................................................................................ 852
Family Niphatidae Van Soest
Ruth Desqueyroux-Faundez & C. Valentine ............................................................................................ 874
Suborder Petrosina Boury-Esnault & Van Beveren
Rob W.M. Van Soest & John N.A. Hooper .............................................................................................. 891
Family Phloeodictyidae Carter
R. Desqueyroux-Faundez & C. Valentine ................................................................................................ 893
Family Petrosiidae Van Soest
R. Desqueyroux-Faundez & Clare Valentine ............................................................................................. 906
Family Calcifibrospongiidae Hartman
Rob W.M. Van Soest ................................................................................................................................ 918
Suborder Spongillina subord. nov. : Freshwater Sponges
Renata Manconi & Roberto Pronzato ...................................................................................................... 921
Contents of Volumes 1 and 2 xi

Order Dictyoceratida Minchin

Steve de C. Cook & Patricia R. Bergquist ............................................................................................. 1021
Family Irciniidae Gray
Steve de C. Cook & Patricia R. Bergquist ...... .................................................... ...................... ......... .... l022
Family Thorectidae Bergquist
Steve de C. Cook & Patricia R. Bergquist ............................................................................................. 1028
Family Spongiidae Gray
Steve de C. Cook & Patricia R. Bergquist ............................................................................................. 1051
Family Dysideidae Gray
Steve de C. Cook & Patricia R. Bergquist .............. .......... ....................................... ........ ..... ................. 1061
Order Dendroceratida Minchin
Patricia R. Bergquist & Steve de C. Cook ............................................................................................. 1067
Family Darwinellidae Merejkowsky
Patricia R. Bergquist & Steve de C. Cook ............................................................................................. 1068
Family Dictyodendrillidae Bergquist
Patricia R. Bergquist & Steve de C. Cook .......... .............. ............. ........ ......... ..... ...... ......................... ... 1072
Order Halisarcida Bergquist
Patricia R. Bergquist & Steve de C. Cook ........... ........ ............................... ............. ............. ......... ........ 1077
Family Halisarcidae Schmidt
Patricia R. Bergquist & Steve de C. Cook ............................................................................................. 1078
Order Verongida Bergquist
Patricia R. Bergquist & Steve de C. Cook ...... ...................... ..... ........ ............................... ..... ............ .... 1081
Family Aplysinidae Carter
Patricia R. Bergquist & Steve de C. Cook ............................................................................................. 1082
Family Pseudoceratinidae Carter
Patricia R. Bergquist & Steve de C. Cook ............................................................................................. 1086
Family Ianthellidae Hyatt
Patricia R. Bergquist & Steve de C. Cook ...... ..... ....................................... ..... ........ .............. ................ 1089
Family Aplysinellidae Bergquist
Patricia R. Bergquist & Steve de C. Cook ....... ........ ..... ...................... ............ .......... .............. ... ........ .... 1094
Recent 'Sphinctozoa', Order Verticillitida, Family Verticillitidae Steinmann
Jean Vacelet ............................................................................................................................................ 1097
Demospongiae Incertae sedis: Myceliospongia Vacelet & Perez
Jean Vace1et, Thierry Perez & John N.A. Hooper .................................................................................. 1099
Index of Higher Taxa to Volumes 1 and 2 .............. .............................. ..... ......... ... ..... .............. ........ ......... ...... xxi

VOLUME 2

Order Clathrinida Hartman

Radovan Borojevic, Nicole Boury-Esnault, Michael Manuel & Jean Vacelet 1141
Order Murrayonida Vacelet
Jean Vacelet, Radovan Borojevic, Nicole Boury-Esnault & Michael Manuel 1153
Order Leucosolenida Hartman
Radovan Borojevic, Nicole Boury-Esnault, Michael Manuel & Jean Vacelet 1157
Order Lithonida Vacelet, Recent
Jean Vacelet, Radovan Borojevic, Nicole Boury-Esnault & Michael Manuel 1185
Order Baerida Borojevic, Boury-Esnault & Vacelet
Radovan Borojevic, Nicole Boury-Esnault, Michael Manuel & Jean Vacelet 1193
Class Hexactinellida Schmidt
Henry M. Reiswig .................................................................................................................................. 1201
Bibliography of Class Hexactinellida
Editor Philippe Willenz .......................................................................................................................... 1203
tFossil Hexactinellida: An Overview
Manfred Krautter .................................................................................................................................... 1211
Dictionary of Hexactinellida
Konstantin R. Tabachnick & Henry M. Reiswig .................................................................................... 1224
Order Amphidiscosida Schrammen
Henry M. Reiswig .................................................................................................................................. 1231
Family Hyalonematidae Gray
Konstantin R. Tabachnick & Larisa L. Menshenina ............................................................................... 1232
Family Monorhaphididae Ijima
Konstantin R. Tabachnick ...................................................................................................................... 1264
Family Pheronematidae Gray
Konstantin R. Tabachnick & Larisa L. Menshenina .............................................................................. 1267
Order Hexactinosida Schrammen
Henry M. Reiswig .................................................................................................................................. 1281
Family Aphrocallistidae Gray
Henry M. Reiswig .................................................................................................................................. 1282
Family Craticulariidae Rauff (Recent)
Henry M. Reiswig ............ ..... ... ...... ........ ..... ...................... ........ .............. ........... ...... ... .............. ....... ...... 1287
Family Cribrospongiidae Roemer (Recent)
Henry M. Reiswig .......................................................... ,....................................................................... 1290
Family Dactylocalycidae Gray
Henry M. Reiswig .................................................................................................................................. 1293
Family Euretidae Zittel
Henry M. Reiswig & Benjamin Wheeler .............................................................................................. 1301
Family Farreidae Gray
Henry M. Reiswig .................................................................................................................................... 1332
Family Tretodictyidae Schulze
Henry M. Reiswig ................. ..... ........ ..... ......... ... .......... ............. .... ..... ............. ..................... ......... ..... ... 1341
Hexactinosida incertae sedis
Henry M. Reiswig .................................................................................................................................. 1355
Order Aulocalycoida Tabachnick & Reiswig
Henry M. Reiswig .... ........ .............. ............. ..... ...... ................... ......... ............................... ......... ......... ... 1361
Family Aulocalycidae Ijima
Henry M. Reiswig .................................................................................................................................. 1362
Family Uncinateridae fam. nov.
Henry M. Reiswig .................................................................................................................................. 1372
Order Lychniscosida Schrammen
Henry M. Reiswig .................................................................................................................................. 1377
Contents of Volumes 1 and 2 xiii

Family Aulocystidae Sollas

Henry M. Reiswig .................................................................................................................................. 1378
Family Diap1euridae Ijima
Henry M. Reiswig .................................................................................................................................. 1383
Order Lyssacinosida Zittel
Henry M. Reiswig .................................................................................................................................. 1387
Family Euplectellidae Gray
Konstantin R. Tabachnick ...................................................................................................................... 1388
Family Leucopsacidae Ijima
Konstantin R. Tabachnick 1435
Family Rossellidae Schulze
Konstantin R. Tabachnick 1441
Lyssacinosida incertae sedis
Konstantin R. Tabachnick ...................................................................................................................... 1506
tFossil 'Sphinctozoa': Chambered Sponges (Polyphyletic)
Baba Senowbari-Daryan & Diego C. Garcia-Bellido ...... .............. ...................... ...................... ............ 1511
Bibliography of Fossil 'Sphinctozoans'
Baba Senowbari-Daryan & Diego C. Garcia-Bellido ............................................................................ 1534
tClass Archaeocyatha Bornemann
Franc;oise Debrenne, A. Yu. Zhuravlev & Pierre D. Kruse .................................................................... 1539
Bibliography of Class Archaeocyatha
Franc;oise Debrenne, A. Yu. Zhuravlev & Pierre D. Kruse .................................................................... 1693
Annotated List of Unrecognisable Sponge Taxa and Unavailable Names
John N.A. Hooper & Rob W.M. Van Soest ............................................................................................ 1701
Bibliography of Unrecognisable Sponge Taxa
Editor Philippe Willenz .......................................................................................................................... 1707
Index of Higher Taxa to Volumes 1 and 2 .............. ............................ ................ .............................. ............... xxi
Key to Symbols, Institutional Acronyms, Abbreviations

t, Denotes an extinct taxon (is used primarily for fossil taxa IGNS, Institute of Geological and Nuclear Sciences (formerly
mentioned in chapters chiefly dealing with Recent sponges). New Zealand Geological Survey), Lower Hutt, New Zealand.
?, placed in front of taxonomic name denotes that there ICZN, International Code of Zoological Nomenclature (see
remains uncertain amongst the contemporary scientific community Anon., 1999).
about the validity or taxonomic placement of a species, genus, 19iG, United Institute of Geology, Geophysics and Mineralogy
family or order. (collections at CSGM), Siberian Branch of the Russian Academy
AM (AMS), Australian Museum, Sydney, Australia (includ- of Sciences; Novosibirsk, Russian Federation.
ing SUP collection, formerly of Department of Geology and 1M (ZEV), Indian Museum (including, Zoological Survey of
Geophysics, University of Sydney, Australia). India), Calcutta, India.
ANSP, Academy Natural Sciences of Philadelphia, USA. IMGP, Geowissenschaftliches Zentrum der Universitat
BGU, Buryatan Territorial Geological Survey, Ulan-Ude, G6ttingen, Section Geobiology [formely Institut und Museum flir
Russian Federation. Geologie und Palaontologie, G6ttingen, Germany.
BMAG, Bristol Museums and Art Gallery, Bristol, United INALI, Instituto Nacional de Limnologia, Santo Tome, Santa
Kingdom. Fe, Argentina.
BMNH, The Natural History Museum, London, United INM, National Museum of Ireland, Dublin, Ireland.
Kingdom [formerly British Museum (Natural History)]. IORAS, Institute of Oceanology of the Russian Academy of
BMS, Buffalo Museum of Science, Buffalo, USA. Sciences, Moscow, Russian Federation.
BPBM, Bernice P. Bishop Museum, Honolulu, Hawaii, USA. IRS(c)NB, Institut royal des Sciences naturelles de Belgique,
CAS, California Academy of Sciences, San Francisco, Bruxelles, Belgium (see also KBIN).
California, USA. ISM, Imperial Science Museum, Tokyo, Japan (see TIU).
CE, Departemento de Paleontologia, Universidad Complutense KBIN, Koninklijk Belgisch Instituut voor
de Madrid, Madrid, Spain. Natuurwetenschappen, Brussel, Belgium (see also IRScNB).
CEAB, Centro de Estudios Avanzados de Blanes Collection, KGU, Krasnoyarsk Territorial Geological Survey,
Blanes, Spain. Krasnoyarsk State University, Russian Federation.
CMN, Canadian Museum of Nature, Ottawa, Ontario, KMMA, Koninklijk Museum voor Midden-Afrika, Tervuren,
Canada (formerly NMC, National Museums of Canada). Belgium (see MRAC).
CNIGRm, Central Scientific-Researching Geological- LFM, Merseyside County Museums, Liverpool, United
Exploration Museum, St Petersburg; Federal Republic of Russia. Kingdom (formerly Liverpool Free Museum).
(new transliteration TsNIGRm). LMJG, Landesmuseum Joanneum Graz (Abteilung flir
CNRS, Centre National de la Recherche Scientifique, Paris, Zoologie), Graz, Germany.
France. LSLH, Linnean Society of London Holdings (Linnean
CSGM, Central Siberian Geological Museum (see TsSGM= Herbarium), United Kingdom.
IGiG) (now holding collections of IgiG), Novosibirsk, Russian MABA, Museo Argentino de Ciencias Naturales 'Bernardino
Federation. Rivadavia', Buenos Aires, Argentina.
DTRG, Dipartimento per 10 studio del Territorio e delle MANSP, Museum of the Academy of Natural Sciences of
sue Risorse, Universita di Genova (Dip. Te. Ris.; ex-IZUG), Philadelphia, Pennsylvania, USA.
Italy. MCN, Museo de CH~ncias Naturais da Fundac;ao Zoobotanica,
DVGI, Far East Geological Institute, Far East Institute of Porto Alegre, RS, Brasil.
Mineral Resources, Khabarovsk, Russian Federation. MCNM (MNCN-CSIC), Museo Nacional de Ciencias
DVIMS, Far-East Institut of Mineral Resources, Khabarovsk, Naturales-Consejo Superior de Investigaciones Cientificas, Madrid,
Russian Federation. Spain.
DVTGU, Far-East Territorial Geological Survey, Far-East MCZ, Museum of Comparative Zoology, Harvard University,
branch of the Russian Academy of Sciences, Khabarovsk, Russian Cambridge, Massachusetts, USA.
Federation. MCSN, Museo Civico di Storia Naturale, Verona, Italy.
FMNH. Field Museum of Natural History, Chicago, USA. MFSN, Museo Friulano di Storia Naturale, Udine, Italy.
FZRG, Fundacao Zoobotanica Rio Grande do Sui, Museu de MG, Geological Museum, New Zealand (see VUMG).
Ciencias Naturais, Porto Alegre, Brazil. MHNG, Museum d'Histoire naturelle, Geneve, Switzerland.
GIN, Geological Institute of the Russian Academy of MNHN, Museum national d'Histoire naturelle, Paris, France.
Sciences, Moscow, Russian Federation. MNRJ, Museu Nacional, Rio de Janeiro, Brazil.
GML, Geiseltalmuseum Martin Luther, University of Halle, MOM, Musee Oceanographique de Monaco, Monaco.
Halle, Germany. MRAC, Musee Royal de l' Afrique Centrale de Tervuren,
GSC, Geological Survey of Canada, Ottawa, Canada. Belgium (see KMMA).
HBOJIM, Harbor Branch Oceanographic Institutionl MRCN, Museu Rio-Grandense de Ciencias Naturais, Porto
Museum, Fort Pierce, Florida, USA. Alegre, Rio Grande do SuI, Brasil.
HMN, Hancock Museum, Newcastle-upon-Tyne, United MSNG, Museo Civico di Storia Naturale 'Giacomo Doria',
Kingdom. Genoa, Italy.

xv
xvi Key to Symbols, Institutional Acronyms, Abbreviations

MT Por, Museo e Istituto di Zoologfa Sistematica SNIIGGIMS, Siberian Scientific Research Institute of
dell'Universita di Torino, Italy (Porifera collection). Geology, Geophysics and Mineral Resources, Novosibirsk, Federal
MY, Million years. Republic of Russia.
MYA, Million years ago. SUP, Department of Geology and Geophysics, The University
MZUS, Musee de Zoologie de I'Universite de Strasbourg, of Sydney, Australia (now housed in The Australian Museum;
France. see AM).
NHRM, Naturhistoriska Riksmuseet, Stockholm, Sweden. TEM, Transmission electron microscopy.
NIGP, Nanjing Institute of Geology and Paleontology of the TIU, Imperial University Museum, Tokyo, Japan.
Chinese Academy of Sciences, Nanjing, China. TsNIGRm, new transliteration for CNIGRm.
NMB, Naturhistorisches Museum zu Basel, Basel, TsSGM, new transliteration for CSGM.
Switzerland. UA, Department of Geology, University of Alberta, Edmonton,
NMNZ, Te Papa, Museum of New Zealand, Wellington, New Alberta, Canada.
Zealand (formerly National Museum of New Zealand, and for- UA, University of Adelaide (see SAM).
merly Dominion Musuem). UAM, University of Alaska Museum, Alaska, USA.
NMS, National Museum of Scotland, Edinburgh, Scotland. UBC, University of British Columbia, Vancouver, Canada.
NMV, Museums of Victoria, Melbourne, Australia (formerly UCLZ, University College of London, Zoology.
National Museum of Victoria). UCPM, University of California Paleontology Museum,
NSM, National Science Museum, Tokyo, Japan. Berkeley, California, USA.
NTM, Northern Territory Museum of Arts and Sciences, UFRJPOR, Universidade Federal do Rio de Janeiro, Brazil.
Museums and Art Galleries of the Northern Territory, Darwin, UKL, University of Kansas, Lawrence, USA.
Australia. UMML, Marine Invertebrate Museum, University of Miami,
NYSM, New York State Museum, Albany, New York State, Rosenstiel School of Marine and Atmospheric Sciences.
USA. USNM, National Museum of Natural History, Smithsonian
NZGS, New Zealand Geological Survey, Wellington, New Institution, Washington D.C., USA (formerly United States
Zealand. National Museum).
NZMW, National Zoological Museum Warsaw, Poland. UTBEG, Texas Memorial Museum, Bureau of Economic
PGO, Zabsibgeologiya, Novokuznetsk, Russian Federation. Geology, Austin, Texas, USA.
PGU, Primor'ye Territorial Geological Survey; Khabarovsk, UWU, University of West Indies, Mona, Jamaica, West Indies
Russian Federation. (Zoology Department) [MSUJj.
PIN, Paleontological Institute of the Russian Academy of VSEGEI, All-Russian Geological Institute, St Petersburg,
Sciences, Moscow, Russian Federation. Russian Federation (transfer of collections to TsNIGRm in
PMJ, Phyletisches Museum, Jena, Germany. progress).
PU, Princeton University Bedfords' Collections (now housed VUMG, Victoria University of Wellington, Museum of
in USNM Washington). Geology, Wellington; New Zealand.
QM, Queensland Museum, Brisbane and Townsville (includ- WAM, Western Australian Museum, Perth, Australia.
ing MTQ, Museum of Tropical Queensland). YPM, Yale University, Peabody Museum of Natural History,
RASMAS, Rosenstiel School of Marine and Atmospheric Yale University, New Haven, Connecticut, USA.
Sciences (see UMML). ZEV, (see 1M).
RIB, (see IRScNB). ZIL, Zoological Institute of the Russian Academy of
RMM, Redpath Museum, McGill University, Montreal, Sciences, St Petersburg, Russian Federation.
Quebec, Canada. ZMA, ZoOiogisch Museum, Universiteit van Amsterdam,
RMNH, Nationaal Natuurhistorisch Museum, Leiden, The Amsterdam, The Netherlands.
Netherlands (formerly Rijksmuseum van Natuurlijke Historie). 2MB, Zoologisches Museum fur Naturkunde an der
ROM, Royal Ontario Museum, Toronto, Canada. Universitat Humboldt zu Berlin, Berlin, Germany.
RSME, Royal Scottish Museum Edinburgh, Scotland. ZMH, Zoologisches Museum flir Hamburg, Hamburg,
SAMP, South African Museum, Pretoria, Republic of South Germany.
Africa. ZMUB, Zoological Museum, University of Bergen, Norway.
SAM, South Australian Museum, Adelaide, Australia (includ- ZMUC, Zoologisk Museum Universitet Copenhagen,
ing collections of UA - The University of Adelaide). Copenhagen, Denmark.
SAMC, South African Museum, Capetown, Republic of ZMUZ, Zoologisches Museum der Universitat, Zurich,
South Africa. Switzerland.
SBR, Station Biologique de Roscoff, France. ZPAL, Institute of Paleobiology of the Polish Academy of
SDCC, Private collections of Steve de C. Cook (destined for Sciences, Warsaw, Poland.
theNMNZ). ZRS, Zoologiska Rijkmuseum, Stockolm, Sweden.
SEM, Scanning electron microscopy. ZSGGU, West Siberian State Geological Trust (= WSSGE;
SME, Station Marine d'Endoume, Marseille, France. =WSSGT, =ZSTGU) Novokuznetsk, Russian Federation.
SMF, Natur-Museum und Forschunginstitut Senckenberg,
Frankfurt-am-Main, Germany.
Acknowledgements

The Systema Porifera project involved an extensive collabora- Rob Van Soest similarly acknowledges the technical assistance of
tion amongst many experts over many years. Without this coopera- Elly Beglinger, Henk de Haas, Mario de Kluijver, Raquel Gomez,
tion this book would never have been attempted. The scientific and Nicole de Voogd (all ZMA Amsterdam), and Bert Hoeksema
bibliographic editors, John Hooper, Rob Van Soest and Philippe (RMNH Leiden). Philippe Willenz thanks Laetitia Despontin for
Willenz, are most grateful to all the authors for their perseverance persevering assistance while amending the bibliographic refer-
and cooperation over the long gestation of this project. We would ences database from original documents.
particularly like to thank our editor at Plenum, Joanna Lawrence, The editors also acknowledge the continued support from
for her continued support and patience during the many and varied their respective institutions: the Queensland Museum, Brisbane,
challenges we overcame to get this work to press, and for main- the Zo61ogisch Museum, Universiteit van Amsterdam, and the
taining faith in this collaborative project. The existence and contin- Institut royal des Sciences naturelles de Belgique, Bruxelles.
uance of this sponge network is a positive and essential tool for the Financial support during the course of this project was received
future - to take the basic infrastructure and the taxonomic hypothe- from the following: John Hooper - from AstraZeneca R&D
ses we present here into further realms. Griffith University, Brisbane, and the Australian Biological
During the development of the Systema three workshops were Resources Study, Canberra; Rob Van Soest and colleagues - from
held (London, Paris and Brisbane) to promote collaboration EU-MAS3-CT97-0144 (to Rob Van Soest), LSF NHM London
between the authors and to resolve both theoretical and practical (Vanessa Pike, to Rob Van Soest, LSF MNHN Paris (to Dirk
difficulties associated with the project. For these meetings the edi- Erpenbeck), LSF ZMUC Copenhagen (to Dirk Erpenbeck), and
tors are most grateful to the host institutions and sponsors: The NWO-ALW 8009.34.003 (to Rob Van Soest).
Natural History Museum, London, including the Linnean Society The authors and editors gratefully acknowledge the following
of London; the Museum National d'Histoire Naturelle, Paris; and publishers for permission to reproduce copyrighted illustrations in
the Queensland Museum, Brisbane, including AstraZeneca R&D this work. Full citations of literature are provided in the various
Griffith University, Brisbane, and the Ian Potter Foundation, bibliographies of this work.
Melbourne. We are particularly grateful to Clare Valentine for her
continued support in accessing the vast BMNH resources over A.A. Balkema, Rotterdam, The Netherlands - Soest, R.W.M. Van, Kempen,
recent years. Th.M.G. Van & Braekman, J.e. (Eds), 1994, 'Sponges in Time and
Space' - Sara, M. & Burlando, B. Pp 111-116 (figs 1,4); Reiswig, H.M. &
The curators and collection managers of the major museum Mehl, D. Pp 151-165 (figs 1-3,9).
collections throughout the world are gratefully acknowledged for American Museum of Natural History, New York, USA - Bulletin of the
providing many loans of sponge types and other material over the American Museum of Natural History, Vol. 120, Finks, R.M., 1960
many years of this project. We are especially grateful to Penny (fig. 63, p. 108).
Association Argentina de Ciencias Naturales, Buenos Aires, Argentina -
Berents (AM Sydney), Clare Valentine and Shirley Stone (BMNH
Physis, Vol. 26(71): 129-140, Bonelto, A.A. & Ezcurra, 1.0., 1966 (aU
London), Beatrice Burch (BPBM Honolulu), A.K. Mandai (1M figures). Vol. 28(77): 351-357, Bonetto, A.A. & Ezcurra de Drago, 1.0.,
Calcutta), e. O'Riordan (INM Dublin), Ruth Desqueyroux- 1969 (aU figures).
Faundez (MNHG Geneva and Johanneum Museum Graz), Association of Australasian Palaeontologists, Canberra, Australia -
Takomura Hoshino [deceased] (MMBS Hiroshima), Claude Levi Alcheringa, Vol. 2: 27-47, Kruse, P.O. 1978 (figs 20, 4A, 7B, l1A,
lIB). Vol. 10: 235-278, Debrenne, F. & Kruse, P.O. 1986 (figs 6A, 6B,
(MNHN Paris), Ms. Stock (NHM Basel), Helmut Sattmann (NHM 15A, 15B, 19A, 21 A, 2IB, 22A, 22B). Vol. 17: 179-183, Debrenne, F.
Wien), Urs Rahm (NMB Basel), B.R. Stuckenberg (NM Natal), Zhurav1ev, A.Yu. & Gravestock, 0.1. 1993 (fig. 3A-B). Vol. 18: 1-54,
Frank Climo (NMNZ Wellington), Susan Boyd (NMV Zhuravlcv, A.Yu. & Gravestock, 0.1.1994 (fig. 12B, E). Memoirs of the
Melbourne), Frank von Knorre (PMJ Jena), Bert Hoeksema and Association of Australasian Palaeontologists Vol. 1: 93-120, Pickett, J.
J.e. Den Hartog (deceased) (RMNH Leiden), Susan Chambers 1983 (figs 4D-E). Vol. 2: 1-139, Gravestock, 0.1. 1984 (figs 32L, 50E,
55H, 60A, 62A, 620, 62E, 62F).
(RSME Edinburgh), Shane Parker [deceased], Wolfgang Zeidler Australian Geological Survey Organisation, Canberra, Australia -
and (SAM Adelaide), Nicole Boury-Esnault and Jean Vacelet Memoirs of the Association of Australian Palaeontologists, Vol. 1:
(SME Marseille), Manfred Grasshoff (SMF Frankfurt), Klaus 85-92, Pickett, J.W. & JeU, P.A., 1983 (fig. 4A-C).
Riitzler and Kathleen Smith (USNM Washington), Loisette Marsh Birkhauser Verlag AG, Basel, Switzerland - Experentia Supplementum,
No. 28, Wiedenmayer, F. 1977, 'Shallow water sponges of the western
and Jane Fromont (WAM Perth), Deiter Kiihlmann (ZMB), and Bahamas' (pis 4.3, 9.2).
Ole Tendal (ZMUC Copenhagen). Board of the Museums and Art Galleries of the Northern Territory, Darwin,
The scientific editors are also grateful to the very many refer- Australia - The Beagle, Occasional Papers of the Northern Territory
ees for providing positive comments and suggestions to improve Museum of Arts and Sciences, Vol. 3(1): lfi7-189, Hooper, J.N.A. 1986
the various chapters in this book, ensuring that these were both sci- (fig. 40). The Beagle, Records of the Northern Territory Museum of Arts
and Sciences, Vol. 6(1): 133-140, Hooper, J.N.A. & Krasochin, V. 1989
entifically accurate and provided a practical working tool for (fig. 1). Vol. 8(1): 27-36, Hooper, J.N.A. et al. 1991 (figs 1-9). Vol. 12(1):
sponge systematics, wherever possible. 151-176, Bergquist, P.R. & Kelly-Borges, M. 1995 (figs 2a, e).
John Hooper thanks the following for technical assistance Board of the Museum of Victoria, Melbourne, Australia - Memoirs of the
during the production of this work: Stephen Cook, John Kennedy, Museum of Victoria, Vol. 50(1): 1-242, Wiedenmayer, F., 1989 (pI. 8,
fig. 11, pI. 9, fig. 4, pI. 28, figs 3-4).
Gert Worheide, Daniel Edson, Sue List-Armitage, Paula Board of the Queensland Museum, Brisbane, Australia - Memoirs of the
Thomkins, Brett Krueger and Monika Schlacher, and Justin Gough Queensland Museum, Vol. 38(1): Bergquist, P.R. 1995 (figs 17c, 20b, 22b,
for support developing an accessible web site (all QM Brisbane). 22c, 22d, 24a, 25c, 23a, 23b). Vol. 40: 1-626, Hooper, J.N.A. 1996 (figs 1,

xvii
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F,0,4C). 101-168, Levi, C. 1983 (various figs). Vol. 8 (A, Zoologie): 437-442,
Geological Society of America and the University of Kansas, Boulder City, Levi, C. 1986 (pI. 1, figs 1-4, pI. 2, fig. 4). Vol. 8 (A, Zoologie): 443-455,
Colorado, and Lawrence, Kansas, USA - Treatise on Invertebrate Borojevic, R & Boury-Esnault, N. 1986 (figs la, 230 2d).
Paleontology, Part E, Vol. 1, 1972. Archaeocyatha (D. Hill, ed.) (figs 87, Vol. 10 (A, 2): 241-263, Levi, C. 1988 (various figs). Vol. 12 (A, 2):
3a,78, 1acd, 60, 1b). 243-276, Borojevic, Ret al., 1990 (figs 1,4, 12, 19,21,23,25,31). Vol.
Geological Survey of Canada, Canada - Palaeontographica Canadiana, 12 (A, 2): 277-290, Levi, C. 1990 (figs 1-2, pI. 1, figs 1-2). Memoiresdu
No.2: 1-105, Rigby, J.K., 1986 (figs 24, 25, pI. 19, figs 1,4,9). Bulletin Museum national d'Histoire naturelle, 184: 39-52, Tabachnick, K. &
of the Geological Survey of Canada, Vol. 269: 121-129, Rigby, J.K. 1977 Levi, C. 2000 (figs 2, 11, 13 (1,5,6,11». Zoosystema, Vol. 9(1): 7-14,
(pI. 1, fig. 3). Vol. 269: 131-137, Rigby, J.K. 1977 (fig. 2, pI. 1, fig. 3). Tabachnick, K. & Levi, C. 1997 (figs 1-11). Vol. 20 (1): 5-22, Vace1et, J.
Geologisch-Paliiontologisches Institut der Universitat Innsbruck, Austria. & Perez, T. 1998 (figs 4a, 5a-b, 6a-b). Vol. 22: 203-263, Borojevic R
Geologisch-Palaontologische Mitteilungen, Sonderband, Vol. 4: 1-55, etal., 2000 (figs 1,4,6-8,10,14,15,17,23-25,27-29,31,33-40,46).
Mehl, D., 1996 (figs 2, 3,4,8, pI. 3, figs 1,3, pI. 5, fig. 2, pI. 6, fig. 1). National Institute of Water and Atmospheric Research (NIWA) Ltd.,
Institut fur Geologie und Paliiontologie der Universitat der Stuttgart, Wellington, New Zealand - Department of Scientific and Industrial
Germany - Profil Stuttgart, 1997, Krautter, M. 1997 (fig. 63). Research Bulletin, New Zealand Oceanographic Institute Memoir, No.
Institut fur Geologie und Paliiontologie der Philipps-Universitiit Marburg, 37: 1-105, Bergquist, P.R. 1968 (text-fig. 10, pI. 4a, pI. lIe-f). No. 197:
Germany - Paliiontologische ZeitschriJt, Vol. 42(1-2): 13-32, Rietschel, 1-85, Bergquist, P.R 1970 (pI. 5, fig. B). No. 100: 1-98, Dawson, E.W.
1968 (pI. 1, fig. 1). 1993 (cover photograph). No. 107: 1-53, Bergquist, P.R. 1996 (cover
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Facies, Vol. 38: 1-88, Worheide, G. 1998 (pI. 20, figs 6-7, pI. 21, fig. 16). National and Kapodistrian University of Athens, Department of Biology -
Institut de Ciencies del Mar, CSIC, Barcelona, Spain - Scientia Marina, 'Kalymnos and the secrets of the sea' - Castritsi-Catharius, J., 1998 (fig.
Vol. 57(4): 283-306, Diaz, M.C. et al., 1993 (fig. 34). Vol. 57(4): 109).
381-393, Riitzler, K. & Smith, K.P., 1993 (fig. 5). Northern Territory Geological Survey, Darwin, Australia - Report of the
Institut royal des Sciences naturelles de Belgique, Bruxelles, Belgium - Northern Teritory Geological Survey, Vol. 7: 1-58, Kruse, P.D., 1990 (pis
Cercle hydrobiologique de Bruxelles, Vol. 11(1): 1-56, Brien, P., 1967 25E,26A).
(all figures). Revue de Zoologiye et de Botanique Africaines, Vol. 88(3): Palaeontologia Polonica, Warszawa, Poland - No. 55: 9-36, Wrona, R &
585-624, Brien, P., 1974 (all figures). Zhuravlev, A. Yu. 1996 (pI. 7, figs 2, 5).
Instituto Espanol Oceanografia - Boletin Instituto Espanol de Senckenbergiana Maritima, Frankfurt am Main, Germany - Vol. 30 (3/6):
Oceanografia, Vol. 6(4), No. 324: 103-154, Biblioni, M.A., 1981 (fig. 7). 161-221, Schonberg, C.H.L. 2000 (pI. 7, fig. 40).
Izdatelsvo Nauka, Moscow, Russia -Akademia Nauk SSSR, (1959): 1-236, SIR Publishing, Royal Society of New Zealand, Wellington, New Zealand -
Koltun, V.M. 1959 (fig. 78). Vol. 92: 6-116, Koltun, V.M. 1964 (fig. 19). New Zealand Journal of Zoology, Vol. 7, Bergquist, P.R. 1980 (figs la,
Koeltz Scientific Books, Koenigstein, Germany - Theses Zoologicae, Vol. 4a-d, 5c-d, 6a-c, 7a-b, 8e, lOb, d, lla, 12a, cod, 13a-c, e, 14a-b, 15a,
23, Barthel, D. & Tendal, O.S. 1994 (fig. 14). 16a-b, 17d-e, 19c, 19f, 22f, 23a, 23b-c, 23f, 24b-c, 25b).
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Vos, L. De et aI., 1991 'Atlas of sponge morphology' (p. 106, pI. 54). Vol. 64(3): 163-192, Van Soest, RW.M. et aI., 1994 (figs 3, 6).
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Societe geologique de France, Paris, France - Bulletin de la Societe WH. De, 1986 (figs 5b, 19b). Vol. 40(2): 15-62, Soest R.WM. Van eta!.,
Geologique de France, (8), Vol. 4, Debrenne, F. & Zhi-Wen Jiang, 1989 1990 (figs 22, 60, 69, 73).
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Societe zoologique de France, Paris, France - Bulletin de la Societe 123-204, Bergquist, P.R. 1965 (figs la, Ib, 7a, 8b).
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(figs 1-5). Rigby, J.K. 1967 (figs 3-4). Vol. 49(2): 329-339, Rigby, J.K. & Nitecki,
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(figs 1, 13, 46). 10,20,21).
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(1967), Borojevic, R. 1967 (figs 3, 4, II). Broadhead, T.W. (ed.) 'Sponges and spongiomorphs; notes for a short
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Goodbody, Q.H. 1986 (fig. 2A, J). Borges, M. & Bergquist, P.R., 1994 (pI. la).
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Systema Porifera. A Guide to the Classification of Sponges

John N.A. Hooperl & Rob W.M. Van Soest2

"If I have seen further it is by standing on the shoulders a/Giants" (Sir Isaac Newton, 1676 [Turnbull et al., 1959])

INTRODUCTION mechanisms that have evolved to survive over Earth's changing

chemical history; their role in the cycles of life and death in the
Why write a book on sponge taxonomy; who would read such oceans; and their co-evolution with other symbiotic life forms.
a book; what is the relevance of sponges; and what are sponges This knowledge has also provided practical answers to funda-
anyway? mental biological questions such as understanding the biosynthesis
The first two questions are not easily answered, except to of chemicals, the evolution of eukaryotic immunology, cellular
offer the explanation that over the past few centuries there have theory and totipotency, understanding physiological adaptive
been many 'books' written on various aspects of sponges, and pre- strategies to cope with extreme enviromnents, gene function and
sumably over the next few centuries there will be many more. evolution of the Metazoa, and some realistic assessments of
During the late 19th to the mid 20th centuries a great many the magnitude of marine biodiversity, including impacts by
monographic publications on sponges were produced (e.g., humans on the marine environment. It is significant that many of
Johnston, 1842; Duchassaing & Michelotti, 1864; Schmidt, 1862, these contemporary studies have been collaborative, multidiscipli-
1864, 1868, 1870, 1879, 1880b; Bowerbank, 1864, 1866, 1874b, nary efforts that seek a broader understanding of the role of
1882; Haeckel, 1872; Pomel, 1872; Marshall & Meyer, 1877; sponges in time and space.
Zittell, 1877, 1879; Quenstedt, 1878b; Hinde, 1887-1893; It is also obvious from the existing sponge literature that this
Saville Kent, 1880--1882; Lendenfeld, 1888, 1889a, 1897a, 1907, present book is unique. Apart from the pioneering attempt by
191Oa,b, 1915; Ridley, 1884a; Polejaeff, 1883, 1884; Ridley & de Laubenfels (1936a) to make the sponge systematics 'widelyavail-
Dendy, 1887; Sollas, 1888; Rauff, 1893-1894; Topsent, 1892a, able' at the practical level of classification (families and genera), no
1900, 1901d, 1904b, 1913a, 1928c; Hall & Clarke, 1898; other publication has attempted to achieve what we propose here.
Lundbeck, 1902, 1905, 1910; Schulze, 1904; Schrammen, The latter questions are perhaps easier to answer. Sponges are
1910--1912, 1937; Dendy, 1922b; Wilson, 1925; Ijima, 1926; de among the most highly diverse and successful of the 28 aquatic
Laubenfels, 1936a, 1948, 1954; Reid, 1958a; Hill, 1965; Burton, invertebrate phyla, both in terms of the numbers of species and
1963; Penney & Racek, 1968; and others). This trend is now sadly their range of morphological characters. About 7,000 extant
in decline, although there are some notable exceptions among the species are currently described (but these undoubtedly represent
contemporary authors, especially the palaeontological literature only a small fraction of those that have ever lived). Sponges have
(e.g., Levi, 1956a; Hartman, 1958b; Koltun, 1959, 1964b, 1967; survived largely unchanged in their fundamental bauplans since the
Finks, 1960; Vacelet, 1964; Wiedenmayer, 1977b, 1994; Rigby, Late Cambrian (509MYA). They were important reef builders dur-
1986c; Rigby & Webby, 1988; Senowbari-Daryan, 1990; Boika ing the Phanerozoic - a role subsequently usurped by the faster
etal., 1991; Reitner, 1992; Rigby & Senowbari-Daryan, 1996; Pisera, growing scleractinian corals during the late Mesozoic. Although
1997, and others). There are also several general texts, mono- most of the older groups became extinct during the Late Devonian
graphic taxonomic overviews, and a few popular books published crisis (373MYA), sponges have radiated and diversified in Recent
on sponges (e.g., Delage & Herouard, 1899; Minchin et aI., 1900; seas, with representatives found in all aquatic habitats - from
Hentschel, 1923; Arndt, 1935; Hyman, 1940; Koltun, 1955; ephemeral (quasi-terrestrial) ones to the more stable marine
Moore, 1955; Teichert, 1972; Brien et al., 1973; Bergquist, 1978; abyssal zones. Consequently, sponges have demonstrated a huge
Hartman, 1982; Hartman et at., 1980; Simpson, 1984; Zea, 1987; capacity to adapt and survive in marginal life situations - and it has
Ackers et al., 1992; Rigby et at., 1993; Hooper & Wiedemnayer, been suggested that they are clearly set to outsurvive those who
1994; Levi et aI., 1998). Since 1970 there has also been a growing study them (Vacelet, 1999).
literature from the international sponge symposia, together repre- Sponges continue to have an economic potential beyond their
senting a comprehensive treatment of sponge biology (Fry, 1970; fundamental roles in the marine ecological processes. These
Harrison & Cowden, 1976; Levi & Boury-Esnault, 1979; Jones, include commercial 'bath sponges', important since early Greek
1987; Vacelet & Boury-Esnault, 1987; Rutzler, 1990; Reitner & civilization, and more recently as potential sources of therapeutic
Keupp, 1991; Uriz & Rutzler, 1993; Van Soest et al., 1994; drugs. This renewed interest in sponges has accelerated the discov-
Willenz, 1996; Watanabe & Fusetani, 1998; Hooper, 1999). ery and documentation of species in all the oceans, with recent
Together with the many thousands of other existing publications on estimates from the various museum collections that the extant
sponges this literature has contributed substantially to advancing fauna may be twice as diverse as that currently described (e.g.,
the knowledge of 'Life On Earth' - past and present; the diverse Hooper & Levi, 1994). As a consequence, it was necessary
sponge morphologies; their structural, physiological and biochemical to revisit the existing systematics in order to accommodate these

1
2 Systema Porifera. A Guide to the Classification of Sponges

substantial new collections, which was the initial impetus to Despite a number of important publications during the late 18th
produce this book. and early 19th centuries (Linnaeus, Cuvier, Lamarck, Lamouroux,
Pallas, Esper, Nardo etc.; see bibliography of Vosmaer, 1928),
Lendenfeld (1888) suggested that our earliest reliable taxonomic
HISTORY OF SPONGE TAXONOMY knowledge of sponges stemmed from the mid-19th century works
of Bowerbank (e.g., 1841) and Schmidt (e.g., 1862) on the British
A long and tortuous taxonomic history and Adriatic faunas, respectively. While it is true that these authors
introduced new suites of characters to sponge taxonomy, raising the
Johnston (1842), an 'amateur' sponge taxonomist, remarked standard of contemporary systematic debates, there were certainly
long ago that sponges were a "comparatively limited and isolated many other significant contributions to the growing body of knowl-
class of organized beings, obscure in character and possessed of edge at that time, including works of Carter, Duchassaing,
less interest than attaches every other. The class may be said to Gray, Hyatt, Johnston, Lieberkuhn, Norman, Thompson, Wyville
occupy at present the confines of debateable land, lying between Thompson and Verrill, to name only the more prominent, and many
the confines of the two organic Kingdoms, - too poor and barren to of whose descriptions are still recognizable today. Towards the end
be an object of contest with the subjects of either ... This is not an the 19th and beginning of the 20th centuries there was a phenome-
easy task, for there is so much that is in common to them, and each nal proliferation of sponge research that impacted on the systemat-
adapts itself so readily to circumstances and assumes a new mask, ics of Porifera, with the most prominent contributions by:
that it requires a tact, to be gained only by some experience, to rec- Annandale, Arndt, Arnesen, Bidder, Brpndsted, Dendy, DOderlein,
ognize them under their guises; while we labour; perhaps in vain, Fristedt, Haeckel, Hallmann, Hanitsch, Hentschel, Hinde, Ijima,
to devise phrases which shall aptly pourtray to others the charac- Jenkin, Keller, Kieschnick, Kirk, Kirkpatrick, Lambe, Lendenfeld,
teristics of objects that have no fixed shape, and whose distinctive Lindgren, Lundbeck, Maas, Marshall, Minchin, Moret, Okada,
peculiarities almost cheat the eye." Polejaeff, Rauff, Ridley, Row, Saville Kent, Schluter, Schrammen,
Over recorded human history the taxonomy of sponges has Schulze, Sollas, Thiele, Topsent, Vosmaer, Weltner, Whitelegge,
been long and tortuous, gaining an unenviable reputation among Wilson and Zittel. It was also during this period that most of the
zoologists as presenting appalling difficulties (Vacelet, 2000). 7,000 currently known species were documented, and the frame-
Homer made numerous references to them (Lendenfeld, 1888) as work of the existing classification developed (e.g., see statistics on
regular household commodities in early Greek society (referring to history of Australian sponge collections on the web at http://
the so-called 'bath sponges', which apparently served in diverse www.environment.gov.auJabrs/abif-faunalvolswww.htm).
roles, from cushioning body armour to cleansing bodies). He also In contrast with this earlier halcyon era of scientific endeavour
coined the name 'sponges', "to squeeze": "Then with a sponge he only a few authors studied sponges during the middle of the
drest, his face all over; necke and hands, and all his hairie breast" 20th century (e.g., Brien, Burton, de Laubenfels, Tanita, Topsent,
(translation by Chapman, quoted in Johnston, 1842). Aristotle was Tuzet), followed by several others over the next few decades
apparently the first to describe sponges as objects of scientific (e.g., Bergquist, Borojevic, Cuif, Gautret, Hartman, Jones, Koltun,
inquiry (Johnston, 1842), differentiating three 'species' within the Levi, Pulitzer-Finali, Rasmont, Sara, Simpson, Termier, Vacelet,
Mediterranean fauna - although it is highly likely that recognition Wendt, Wiedenmayer), with expertise gradually escalating to number
of more than one species of 'sponge' substantially predates his writ- several hundreds of researchers by the end of the 20th century, includ-
ings: Tragos, Manon, Achilleum. Pliny (translation by Holland, ing the 30 or so contributors to this present book. In comparison to
1634) also mentioned a fourth species: Aplysiae. More significantly, this escalating number of neontological authors our palaeontological
however, Aristotle was apparently also the first author to recognize capabilities continue to decline, with many of the more prominent
that sponges might be animals (Johnston, 1842), although Pliny palaeontological authors now retired or approaching the end of their
provided conflicting statements that they were "neither living crea- careers (Debrenne, Finks, Pickett, Reid, Rigby, Zhuravlev).
tures nor yet plants ... intermediate between vegetable and animal
kingdoms", and again, "sponges have a life ... yea and a sensible "Standing on the shoulders of giants"
life ... for there is found of their bloud settled within them".
The ensuing scientific void during the 'dark ages' contributed In 1928 Geo P. Bidder wrote "Some younger biologists incline
little to advancing biological or taxonomic knowledge of the phy- now to cut themselves loose from the lengthening chain of literature,
lum. Occasionally 'bestialities' included reference to sponges (e.g., and to read nothing that has appeared more than twenty years ago.
Entzel, 1551), and their medicinal properties (e.g., Pomet, 1694), Whatever they gain in time for research, they lose much ... They lose
sometimes reiterating Aristotle's assertion of their animality (e.g., knowledge because forgotten observations of great value lie hidden
Wotton, 1552), although with little comprehension of their origin in papers which excited little interest among contemporaries, and
and development - "fermentation of the sea's scum or its sponta- consequently have not been passed through the generations on the
neous pullulations ... certaine matter wrought together; of the fome stream of second-hand quotation. ... Traditional beliefs without rea-
and froth of the sea, which we call spunges" (Gerarde, 1597), and sonable foundation ... are at their strongest if we believe that our
sometimes asserting their vegetable nature - "cryptogamous veg- own generation has made all its own thought, and that no brave
etables and nearly allied to the Fungi" (Imperato, 1599). Indeed, man lived before Agamemnon" (Bidder, in Vosmaer, 1928).
Linnaeus (1759) initially arranged Spongia amongst the cryptoga- Unfortunately, the sponge literature is now so diverse, and some
mous algae, but later (1767) included it in Animalia. This debate works are nearly impossible to access in libraries, that the older
continued for well over a century (see Johnston, 1842; Levi, 1999), literature continues to rapidly recede into the past. Levi (1999)
with Pallas (1766) and Ellis (1755, 1786) being the first to provide provides some pertinent examples of knowledge that has been lost
an informed scientific opinion as to their animal nature, which was over time, or distorted during its transmission to the present, such
finally settled under the influential gaze of Grant (1826b). that sometimes we seem to repeatedly 'reinvent the wheel' .
Systema Porifera. A Guide to the Classification of Sponges 3

We take heed of Bidder's warning, and consequently have classes, subclasses, orders, suborders, families and genera - and
endeavored here to build on the accumulated knowledge of our within each genus, listing the pertinent literature (where this exists) to
forebears to provide (within the rules of the ICZN; Anon, 1999): a potentially identify species. This structure, however, is fundamentally
stable nomenclature; a sound contemporary classification; a rem- at odds with how most sponge taxonomists operate - which is from
edy for the nomenclatural mistakes of the past; and incorporating the species level upwards (a 'gestalt' approach). Anyone familiar with
more recent non-morphometric data into the contemporary classifi- the appalling difficulties associated with sponge systematics would
cation. We hope that these efforts will provide a solid platform for already know that keys usually do not work, except perhaps for the
the future, and that any shortcomings of this Systema Porifera will very few well-known small regional faunas, or for small, well-
mostly concern the phylogenetic controversies and unanswered characterised taxonomic groups (such as some genera and families).
biological questions, which we anticipate will be solved by The message is: beware! These difficulties are mostly due to the fre-
advances in our biological knowledge and our abilities to better quent losses of 'pivotal characters', or the modification of these char-
sample the sponge genome. acters beyond recognition as being 'typical' for a particular taxonomic
group, or the difficulty in interpreting whether characters are the same
or different between groups of species. This latter problem concerns
AIMS OF THE 'SYSTEMA PORIFERA' the interpretation of 'homology', whereby some prominent characters
that may appear to be the same across several taxonomic groups actu-
A Taxonomic Revision ally represent analogous features that do not necessarily reflect
phylogeny, or translate into a convenient morphological classification.
The primary aim of Systema Porifera is to summarise, and Consequently, the secondary structure of this book - and one
revise where necessary the supraspecific classification of the Phylum that is more important to its future as a sound platform to develop
Porifera (including spongiomorphs such as 'Sphinctozoans', the sponge systematics - concerns extensive analyses of sponge
Archaeocyatha and Heteractinida), based firstly on re-evaluation of characters and their relative importance (homology vs. analogy), to
type material for each genus when possible, and subsequently incor- arrive at both a practical classification and a theoretical reconstruc-
porating any recent biological evidence. The Systema focuses on the tion of sponge phylogenies. For those who are mostly interested in
living fauna, and in most cases provides only a cursory treatment of simply identifying sponges these phylogenetic analyses and
the fossil fauna and palaeontological literature. It is anticipated that lengthy discussions may appear as 'noise', although we contend
the forthcoming revision of the Treatise on Invertebrate Paleontology that this 'noise' is an essential quality control. Accurate interpreta-
will balance this bias (Rigby et al., in prep.). tion of characters is essential for the accurate identification of taxa.
Illustrated glossaries are useful to define characters and to assist in
A Sponge Guide recognising homologous from analogous characters. However, we
only provide a few of these here because many have already been
It is intended that the Systema Porifera will serve as a published in the contemporary literature (see below).
practical guide, providing accessibility to the supraspecific Thirdly, the Systema represents a concerted attempt to both
classification of sponges and spongiomorphs; providing a clear stabilize the sponge nomenclature (through our collective compre-
explanation of morphological characters, their importance to the hensive re-evaluations of the entire literature), and to revise the
taxonomy and systematics of each higher taxon, and to illustrate Poriferan classification (incorporating recent evidence held by
the major characters used to classify each group. experts in the field - who have an estimated combined taxonomic
expertise of more than 600 person-years!). Thus, many of these
Solving Nomenclatural Problems discussions and analyses are necessary to justify the new
taxonomic hypotheses presented here.
The Systema Porifera provides a sound baseline for future
debate on sponge taxonomy and to address many long-outstanding References to other works that may assist using this book
nomenclatural problems (such as those inadequately treated or
overlooked completely by de Laubenfels, 1936a). There are several illustrated glossaries and other useful publi-
cations that define sponge morphology. Ideally, these should have
been reproduced here, to make this book complete in itself, but this
ABOUT THIS BOOK was seen as a lUXury and a redundancy. Already the size of this vol-
ume had doubled from our original assessment of the Phylum at the
Firstly, an apology outset of this project, as we continued to uncover numerous forgot-
ten taxa. Consequently, there are two publications in particular that
It is obvious that we have committed a fundamentalJaux pas should be used in conjunction with this volume to identify sponges:
in the title of this book, which of course should be Systema the Thesaurus oj sponge morphology (Boury-Esnault & Riitzler,
Poriferorum. However, from the outset it was recognised that this 1997) and Atlas oj sponge morphology (De Vos et al., 1991).
classic title was both ponderous and ambiguous in identifying the Several other works may also be useful, providing additional glos-
Phylum Porifera as the target taxon, and consequently we chose to saries, relevant illustrations and interpretations of sponge characters
ignore classic nomenclature for pragmatic reasons. Our other (Brien et al., 1973; Bergquist, 1978; Hooper & Wiedenmayer, 1994
endeavours in this book, however, have been more stringent. [available through bookshop@isb.csiro.au, with an online version
at http://www.environment.gov.aulabrs/abif-faunalvolswww.htrn];
Structure of the 'Systema' Van Soest et al., 2000 [available through orders@ springer. de;
The structure of this book is aimed primarily at those faced with http://www.eti.uva.n1!Products/CD-catalogue.html]; Levi et al.,
the daunting task of identifying sponges - providing keys to diagnose 1998 [available through diffusion@bondy.orstom.fr].
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

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provenant des croisieres du Prince Albert ler de Monaco. Resultats des record, ecology, and global distribution of intermediate and higher taxa.
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from the N.E. Atlantic and Mediterranean Sea. NATO ASI Series G13. (Palaozoologie: 1876-1880). (Miinchen - Oldenbourg).
(Springer-Verlag: Berlin, Heidelberg): 1-332. ZITTEL, K.A 1915. 1. Unterstamm. Porifera. Pp. 52-84. In: Broili, F.
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VON LENDENFELD, R.: see LENDENFELD, R. VON. ZRZAVY, J., MIHULKA, S., KEPKA, P., BEZDEK, A. & TIETZ, D. 1998.
VOSMAER, G.C.J. 1928. In: Bidder, G.P. & Vosmaer-Roell, C.S. (Eds), Phylogeny of the Metazoa Based on Morphological and 18S Ribosomal
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Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Phylum Porifera Grant, 1836

John N.A. Hooper,! Rob W.M. Van Soest2 & Fran~oise Debrenne3

1 Queensland Museum, P.O. Box 3300, South Brisbane, Qld, 4101, Australia. (JohnH@qm.qld.gov.au)
2 Zoological Museum, University of Amsterdam, P.O. Box 94766,1090 GT, Amsterdam, Netherlands. (soest@science.uva.nl)
3 Paleontology, Museum National d'Histoire Naturelle, 8 Rue Buffon, 75005 Paris, France. (debrenne@club-internet.fr)

Phylum Porifera (sponges) are metazoans united by the unique possession of choanocyte chambers, a system of afferent and efferent
canals with external pores, lacking a tissue grade of construction but having a'highly mobile population of cells capable of totipotency,
and possessing siliceous or calcitic spicules in many (but not all) species. Four sponge classes are currently recognised, although the
monophyly (and indeed the existence of the 'Phylum Porifera') has been challenged by recent molecular data and chemical evidence.
Three classes have Recent species: Hexactinellida (syncytial choanoderm, discrete cells and pinacoderm, siliceous triaxone spicules,
ubiquitous fibrillar collagen, mostly deeper water), Demospongiae (discrete cells, siliceous monaxone or tetraxone spicules, ubiquitous
fibrillar collagen), and Calcarea (discrete cells, calcareous spicules, ubiquitous fibrillar collagen). Archaeocyatha was an important group
of sessile marine organisms during the Cambrian but is now presumed extinct. It was characterized in having double-walled inverted
conular growth forms with spaces between outer and inner walls filled by various skeletal structures, overall architecture of interlocking
polyhedral microgranular calcite, and lacking free spicules. Within this systematics there is scope for recognition of subphyla (e.g.,
Demospongiae + Calcarea vs. Hexactinellida, based on cytological evidence; or Demospongiae + Hexactinellida vs. Calcarea, based on
some molecular data), but this level is not applied here as the debate is still embryonic and there are still conflicting molecular data. Fossil
sponges are not always easy to classify with respect to Recent taxa, with some fossilised groups such as the 'lithistids' and 'sphincto-
zoans' distributed within a Recent classification, but with others treated independently. Keys to the classes of Recent and fossil sponges
are constructed independently to avoid presumptions of alleged phylogenetic affinities. This work describes 682 valid genera of the extant
fauna (of about 1600 nominal genera) in three classes, 25 orders and 127 families; and mentions over 1000 'valid' genera amongst the fos-
sil fauna in six 'classes', 30 orders and 245 families (although the fossil demosponge fauna is not substantially delineated in this work).
Fossil 'Classes' Sphinctozoa and Stromatoporoidea are clearly polyphyletic and represent grades of construction and not phylogenetic
clades.
Keywords: Porifera; Demospongiae; Calcarea; Hexactinellida; Archaeocyatha; 'Sphinctozoa'; 'Stromatoporoidea'.

DEFINITION, DIAGNOSIS, PURPOSE, SCOPE addition to other functions. Cells are highly mobile, such as those
that move freely within an extracellular matrix made of fibrils of
Definition collagen (the mesohyl): archaeocytes, collencytes, spiculocytes,
spongocytes, glycocytes, cells with inclusions, etc. (see definitions
Sessile metazoans with a differentiated inhalant and exhalant of terms in Boury-Esnault & Riitzler, 1997). Most of the cells, espe-
aquiferous system with external pores, in which a single layer of cially the archaeocytes, have an ability to continually evolve into
flagellated cells (choanocytes) pump a unidirectional water current several other cell types as required by the individual organism
through the body, containing a highly mobile population of cells (totipotency), which provide the sponge with a plasticity for its
capable of differentiating into other cell types (totipotency) and organisation. Firmness of the sponge body is provided by (1) colla-
conferring a plasticity to growth form, and with siliceous or calcitic gen fibrils of the mesohyl, (2) spongin fibres, and (3) an inorganic
spicules present in many species. skeleton consisting of various supporting mineral elements composed
of either calcium carbonate (CaC0 3) or silica (Si0 2) material
Diagnosis (including discrete spicules, articulated or fused spicules and/or
hypercalcified mineralised basal skeleton). Articulated and hyper-
Sessile metazoans possessing inhalant and exhalant pores con- calcified skeletons are absent in most Recent taxa, but were much
nected by chambers lined by choanocytes. The outside and the more prevalent in fossil faunas (,lithistid' and 'sclerosponge' bau-
canals are lined by pinacocytes (exo- and endopinacocytes respec- plans). These organic and inorganic materials are manufactured or
tively). Water is inhaled through small pores (10--100 fLm in diame- otherwise engineered by various types of cells. Sponges have
ter), traversing the afferent canals towards the choanocyte chamber, free-swimming or creeping larvae, although most groups have con-
and is expelled through efferent canals and the larger exhalant oscu- siderable means of asexual propagation, and all have extensive
lum. Water currents are unidirectional, maintained by an active regenerative powers that appear to be vital for sustaining local pop-
beating of a single layer of flagellated cells (choanocytes) usually ulations. There are three distinct classes (Hexactinellida,
contained within chambers. Food particles and oxygen are removed Demospongiae and Calcarea), with the extinct class Archaeocyatha
from the water by various cells, including the choanocytes. Other having suspected affinities with Demospongiae. The fossil
cells, including archaeocytes, are instrumental in transporting these 'classes' Sphinctozoa and Stromatoporoidea are obvious grades of
respiratory and dietary products throughout the sponge body, in construction and not phylogenetic clades (e.g., Wood, 1991a),

9
10 Phylum Porifera

although distribution of the many lower taxa amongst the four well- otherwise subjective taxonomic decisions based solely on morpho-
established classes remains largely unresolved. metric data (although these molecular datasets are sometimes
increasingly confounding, and in some cases clearly misleading).
Poriferan apomorphies Nevertheless, systematics derived from genotypic data must have
some corroboratory morphological support to be of any practical
Sponges were first recognised unequivocally as being animals value. This is the purpose of this book.
by Ellis (1755), Pallas (1766) and Ellis & Solander (1786), although
an acrimonious debate on this topic continued for at least a century Remarks
(see Levi, 1999). They were first acknowledged as a distinct taxon
within the Linnean classification, as 'Spongida', 'Spongiida' or Defining taxa of Porifera in morphological terms continues to
'Spongiae', but generally incorporated within the Protozoa (early be problematical. Fry (1970), when evaluating taxonomic contribu-
volumes of the Zoological Record). It was not until the 1872 volume tions presented at the first symposium of international researchers
of the Zoological Record that Liitken suggested 'Spongozoa' held in London, remarked that sponges "continue to enjoy a position
"should probably form a class by themselves", and subsequently in of doubtful parentage and relationship". Indeed, despite significant
1876 were they elevated to their own section of the Zoological technological and conceptual advances since 1970 this uncertainty
Record. Over the two centuries of earnest study the perception of persists within the poriferan phylogeny. The definition provided
sponges has evolved from suspected colonial representatives of uni- here covers most but not every putative 'sponge'. For example,
cellular protozoans (Clark, 1867; Saville Kent, 1880-1882), to a Cladorhizidae (Demospongiae, Poecilosclerida), a family of exclu-
member of the coelenterates (Haeckel, 1872), to an intermediate sively deep-water sponges, contains many species that lack both
taxon (Subkingdom Parazoa) (e.g., Hentschel, 1923), to a phylum choanocyte chambers and a canal system, with dietary requirements
lying at the base of the Metazoa (Hyman, 1940; Brien, 1967), and provided by a strategy of carnivory evolved for living in the deep sea
currently to several potential phyla of multicellular animals (e.g., (Vacelet, 1999a). These species have spicule and cellular character-
Bergquist, 1985; Zrzavy et al., 1998; Borchiellini et al., 2001). istics similar to other sponges - thus enabling us to recognise them
Their true affinities will certainly become better resolved as as sponges in the first place - and consequently it can be said confi-
our technical abilities to sample the genome increase, unclouded by dently that most sponges have most if not all apomorphies, and
the myriad of phenotypic problems that currently plague sponge where one or more of these are absent they are subsequent evolu-
systematics. In the meantime, however, we continue to treat sponges tionary losses. The definition of Porifera is also not practical when
(rightly or wrongly) as a monophyletic taxon, with the current applied to many fossil sponges or sponge-like organisms, where
definition based fundamentally on the common possession of choanocyte chambers, the smaller afferent and efferent canals, and
(1) choanocytes, (2) a differentiated inhalant and exhalant aquiferous the morphology and structure of cells and the mesohyl are not pre-
system with external pores, (3) a high cellular motility with cells served through the processes of fossilization - with the exception of
capable of totipotency, and (4) presence of spicules in many species. the remarkable Precambrian fossils from Guizhou mentioned above
Recent data have already contested some of these alleged (Li et al., 1998), that have apparent affinities to Haplosclerida
synapomorphies, with the discovery of carnivorous sponges (Vacelet (Demospongiae) (Vacelet, 1999a). Consequently, the 'sponge
& Boury-Esnault, 1995a) and some remarkable Precambrian fossils nature' of fossil groups that are no longer extant in Recent faunas is
from Guizhou in southern China (Li et al., 1998), both of which a matter of circumstantial evidence and thus continually open to
lack the first two synapomorphies (Vacelet, 1999a), and the last speculation. Our assignment of certain fossil groups to the Porifera,
one, spicules, not universal amongst the phylum. In a theoretical such as the Archaeocyatha and Heteractinida, is based on contempo-
context these modifications or losses to the fundamental poriferan rary expert opinion, but it is also accepted that these opinions may
bauplan (described as probable apomorphies by Vacelet, 1999a) are change with discovery of new evidence and also the reinterpretation
no problem, whereas they present a practical problem in clearly dif- of existing data using new technologies.
ferentiating all putative sponges from other multicellular animals. Finally, on a cautionary note, the practical identification of
In light of this new evidence, and in reviewing the concept of 'ani- sponges is fraught with pitfalls and traps due to the many losses or
mality' of the Porifera, Levi (1999) regarded the most important transformations of certain characters, especially spicule types, that
feature of the phylum to be their highly mobile cells and plasticity are otherwise pivotal to the definition of a particular taxon. These
in growth form, reiterating his previous assertions (e.g., Levi, 1970) reductions (or modifications) may confound even the best attempts
that sponges retain a plasticity of their cellular organisation unique to assign specimens to any reliable taxon. Fortunately, in most
amongst the Metazoa. Together with their theoretical capabilities cases, other key characters provide clues for correct diagnoses,
for cellular totipotency (Vacelet, 1990) these diverse cell types can although sometimes these clues are very subtle. Similarly, some
be found in all organisms that we presently assign to 'sponges'. sponge taxa are infamous for their ability to incorporate foreign
This definition has good empirical support from the cellular diver- spicules and/or detritus into their skeletons, challenging the abili-
sity observed in Recent species (e.g., pinacocytes, porocytes, ties of even the more experienced taxonomists to diagnose them
archaeocytes, sclerocytes) also allegedly present in the Precambrian correctly. Consequently, considerable effort has been devoted in
fossils from Guizhou; the cellular diversity and extremely high cel- this book to diagnose, describe and illustrate the variety of body
lular mobility present in carnivorous species; and the incidental, but plans and other diagnostic features used to define generic and
nevertheless graphic evidence that high cellular mobility is pivotal family level taxa. Nevertheless, proceed with caution!
to morphogenesis, homeostasis, locomotion, and ultimately to sur-
vivorship of sponges (e.g., Fry, 1970; Bond, 1992). Perceived prob- Current systematics
lems in defining Porifera may stem from our over-reliance on
morphometric datasets to make taxonomic decisions, with molecu- Monophyly of the Porifera. Several attempts have been
lar data providing increasingly important evidence to resolve made over the past 150 years to subdivide the Porifera into subphyla
Phylum Porifera 11

and other taxa. Gray (1867a) was the first to propose such a subdivi- paraphyly within the Porifera altogether, and thus sidestep any pro-
sion, to differentiate 'Porifera Silicea' (later to be dubbed 'Porifera posal to rearrange the classification of its established classes until
Incalcarea') from 'Porifera Calcarea' - a distinction that was main- the matter has been more satisfactorily resolved. Nevertheless, it is
tained for many years. More recently, Reiswig & Mackie (1983), predictable that escalating molecular evidence based on multiple
after their discovery of the syncytial nature of the choanoderm and gene sequences will soon approach a satisfactory resolution to
pinacoderm of the Hexactinellida, proposed a subdivision of the answer the question whether 'sponges' are monophyletic or para-
Phylum into two taxa: the 'Symplasma' (containing only the class phyletic. For the present we propose that these data are still con-
Hexactinellida) and 'Cellularia' (containing classes Demospongiae + tentious phylogenetic hypotheses, lacking substantial corroboratory
Calcarea). This proposal was subsequently supported by Bergquist support from other genes, whereby only a few taxa have been
(1985), who further suggested that the Symplasma should be ele- analysed thus far, and only incomplete (single gene sequences)
vated to a separate phylum. Mehl & Reitner (1996) also supported have contributed to potential phylogenies. These divergences of
the view of Reiswig & Mackie (1983), but pointed out the likely opinion have, and probably always will occur throughout sponge
primitive nature of discrete cells, and accordingly they proposed a systematics, and it is important for the young reader to understand
different name - 'Pinacophora' for the clade Demospongiae + that science is always progressing, and that there is NO absolute
Calcarea - but retained the clade Hexactinellida. truth - even in a huge volume like this.
More recent molecular data using 28S ribosomal DNA (e.g., Phylogenetic 'Clades' versus 'Grades' of construction.
Lafay et al., 1992) and 18S rDNA (e.g., Borchiellini et al., 2001) Hartman (1969, 1979), after his re-discovery of the sponge nature of
were interpreted as empirical support for paraphyly within the the so-called coralline sponges (sponges with solid limestone
Porifera, whereby calcareous sponges (Calcarea) appeared to be 'hypercalcified' basal skeletons), proposed a fourth class of Porifera,
more closely related to other metazoans than to siliceous sponges 'Sclerospongiae'. However, subsequent investigations (e.g., Vacelet,
(Demospongiae + Hexactinellida), showing deep radiations 1985) clearly showed that solid limestone skeletons have been devel-
between these two groups. Ironically, these current state-of-the-art oped independently in several unrelated lines of demosponges.
technical data corroborated the earliest proposal for a subdivision 'Sclerosponges', or coralline sponges, or hypercalcified sponges, are
of the Phylum Porifera by Gray (1867a), as noted above. Siddall here treated at the level of families included in various orders of
et al. (1995) and Cavalier-Smith et al. (1996) provided further data which the majority of families do not possess the solid limestone
to support a closer relationship between calcareans and the skeleton. Similarly, possession of basal skeletons composed of des-
ctenophorans than with the siliceous sponges, and Zrzavy et al. mas ('lithistids', previously assigned to order Lithistida), or different
(1998) listed some possible synapomorphies to define a clade grades of skeletal construction (e.g., 'sphinctozoans' in class
(Calcarea + Ctenophora + Cnidaria). These potential synapormor- Sphinctozoa, 'stromatoporoids' in class Stromatoporoidea) also
phies included the possible non-homology of choanocyte flagellae remain contentious (e.g., Wood, 1991a), with present indications sug-
throughout the Porifera (with calcareans having cross-striated fla- gesting that these features are homeoplastic and their indicated taxa
gellar rootlets found in some triploblasts but not in the diploblastic are polyphyletic - with the similar consequence that, where possible
siliceous sponges), and the animal-like mode of sponge embryoge- from other corroboratory evidence (e.g., geometry of free spicules),
nesis in calcareans but not in other poriferans. They also suggested these taxa are distributed amongst the established classes and orders
that the common possession of calcitic spicules in calcareans and of Porifera. Achieving this task completely, however, remains elusive
anthozoans was a potential synapomorphy, but this latter hypothe- and hence the systematics of Porifera is still largely unresolved at
sis is here rejected given that these characters are non-homologous, higher levels of classification. For this reason the Systema Porifera
whereby calcarean spicules are secreted extracellularly. Zrzavy project has deliberately focussed on the intermediate taxa (families,
et al. (1998), supported by Borchiellini et al. (2001), proposed to res- genera), and includes the higher taxa (suborders, orders, and above)
urrect Johnston's (1842) taxa 'Silicispongiae' (for Demospongiae + only to provide an 'indicative context' to these more practical units
Hexactinellida), with a potential apomorphy being the method of of classification. Resolving the higher systematics of sponges is
secretion of spicules and the ultrastructure of the sclerocytes, and clearly beyond the capabilities of this present book or morphometric
'Calcispongiae' (for Calcarea) as subphyla, or potential phyla, to data alone.
reflect the alleged deep molecular divergence between these
clades.
These data conflict with earlier phylogenetic hypotheses that Scope
support the monophyly of Porifera (e.g., Reitner & Mehl, 1996).
Furthermore, a recent investigation of new full-length 28S and 18S For the purpose of this project the Phylum Porifera is subdi-
rDNA sequences (Medina et al., 2001), including re-examination of vided into three (unchallenged) Recent classes, with a fourth,
some previously published sequences by these authors, found very apparently exclusive fossil class (Archaeocyatha) included - the
strong support for the clade (Demospongiae + Hexactinellida), for latter showing possible affinities to the Demospongiae based on
which they used the later name of Silicea Gray, 1867a. They did studies of immune responses and peculiar budding types
not, however, find conclusive or statistically significant support for (Debrenne & Zhuravlev, 1994). Due to diagenetic changes in fossil
poriferan paraphyly, or resolve the position of the Calcarea within sponges and the frequent loss of vital spicule components -
the phylum, suggesting that earlier conclusions about 'Phylum particularly free spicule types that are often useful as 'systematic
Calcispongiae' must be interpreted cautiously for the time being, indicators' - it is impossible to key out fossil and Recent taxa using
including any inferred relationships of the Calcarea with the a single key. Thus, we treat fossil and Recent specimens
Eumetazoa. These findings also suggest that these genetic markers separately here.
(18S and 28S rDNA) might not be the most appropriate to resolve In the present work Recent sponges are distributed amongst
this specific question of calcarean relationships. Consequently, we three classes, seven subclasses, 25 orders (one herein abandoned),
prefer, at this juncture of uncertainty, to avoid the issue of potential 127 families and 682 valid genera (with over 1600 nominal genera
12 Phylum Porifera

('available names', but here considered to be junior synonyms), plus treatment in the forthcoming reVlSlon of the Treatise on
approximately 500 other invalid (,unavailable') names). Treatment Invertebrate Paleontology (J.K. Rigby et al., editors).
of the fossil fauna is far less comprehensive and less critical, with
six 'classes', 30 orders, 245 families and 998 'valid' genera Further reading
mentioned, although fossil demosponges in particular are not sub-
stantially delineated in this work, awaiting a more comprehensive Brien et at., 1973; Bergquist, 1978; Hartman, 1982.

KEY TO RECENT PORIFERA

(1) Mineral skeleton absent ................................................................................................................................................ Demospongiae

Mineral skeleton includes a 'hypercalcified' basal skeleton of solid limestone .................................................................................. 2
Mineral skeleton consisting of discrete spicules .... ...... ... ........ .... ...... ........... .......... ........... ................ ...... ........................... ................. 3
(2) Soft parts contain siliceous spicules ............................................................................................................................. Demospongiae
Soft parts contain calcareous spicules (test with acid) ........................................................................................................... Calcarea
(3) Spicules siliceous, the larger ones are triaxonelhexactine (six-rayed), occurring both individually and
fused together .......... ..... ................ ................... ............. ........... ...... ............... ................. ................ ............... ...... ..... ...... HexactineUida
Spicules siliceous, the larger ones are tetraxone or monaxone ..................................................................................... Demospongiae
Spicules calcareous (test with acid), usually triactine or tetractine ........................................................................................ Calcarea

KEY TO FOSSIL PORIFERA AND 'SPONGIOMORPHS'

Key to the classes of fossil Porifera and sponge· like fossils

Preservation of sponges depends upon processes of fossilisation (generally restricted to the hard body parts), and taphonomy (com-
paction, dislocation of skeleton, diagenetic modifications). Consequently, a key to the fossil fauna differs in its approach and format to
that of the living fauna.

Isolated spicules in sediments

Siliceous spicules: stauractines, pentactines, hexactines ............................................................................................. Hexactinellida

Monaxones ................................................................................................................................. Demospongiae [since Precambrian]
Monaxones, tetractines, rare dermal spicules ................................................................................. Demospongiae [since Cambrian]
Calcareous: polyactines ........................................................................... Calcarea (Heteractinida, extinct group) [since Cambrian]

Complete (or almost complete) bodies

By early phosphatisation: cellular sponge structures and embryos .......................................... ? Demospongiae [Late Sinian, China]
In lagerstatten
(1) Cast of ....................................................................................................................... ? Hexactinellida [Precambrian of Ediacara]
(2) With siliceous hexactine spicules in bundles or distinguishable bodies ............................. Hexactinellida [Lower Cambrian of
Chengjiang, Middle Cambrian of Burgess Shales]
(3) With siliceous tetractine spicules ...................................................... Demospongiae [Lower Cambrian of Shansha, Greenland,
Middle Cambrian of Burgess Shales]
(4) With polyactine calcareous spicules ..................................................................... Calcarea [Middle Cambrian Burgess Shales]

Massive spicular skeleton

With six-rayed siliceous spicules ................................................................................................................................ Hexactinellida

With tetraxone/monaxone siliceous spicules .............................................................................................................. Demospongiae
With trimeral symmetric calcareous spicules ....................................................................................................................... Calcarea

Perforate calcareous skeleton consisting of uniform interlocked polyhedral microgranular calcite ............. ...... .... Archaeocyatha

'Hypercalci6ed' basal skeleton of solid limestone (aragonitic or calcitic)

With or without embedded siliceous spicules ............................................................................................................. Demospongiae

With or without embedded calcareous spicules ......... ............ .... ........ ... ................ ...... ..... ................ .......... ...... ...... ........... ... Calcarea
Phylum Porifera 13

'Spongiomorphs'

The systematic position of problematic organisms with calcified skeletons, Recent as well as ancient groups, has long been disputed.
Poriferan affinities are now generally admitted. But the discovery of hypercalcified 'living fossils' and the re-evaluation ofthe systematic
value of architectural patterns demonstrate that 'Sphinctozoa' (successive chambers), 'Stromatoporoidea' (laminae and pillar mesh-like
coenosteum) and 'Chaetetidae' ('Tabulata') (calicles) represent grades of organisation rather than systematic clades and are present in the
different classes of sponges. Systematic position within classes depends upon presence or absence of 'normal' sponge systematic charac-
ters. (Refer to chapters on: Class Archaeocyatha, Fossil Demospongiae, 'Class Stromatoporoidea', Fossil 'Sphinctozoa', Order
Heteractinida).
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademicIPlenum Publishers, New York, 2002

Class Demospongiae Sollas, 1885

John N.A. Hooper! & Rob W.M. Van Soest2

Demospongiae (Porifera) have discrete cellular elements, parenchymella or blastula larvae, with either viviparous or oviparous reproduc-
tive strategies. The skeleton is composed of monaxonic or tetraxonic siliceous spicules (never triaxonic) bound together with spongin in
discrete fibres or loosely aggregated, and Ubiquitous collagenous filaments forming the ground substance of the intercellular matrix.
Spicules andlor fibres, or both, may be absent in some taxa, and several other groups with solid calcitic or siliceous skeletons are also
included ('sclerosponges', 'sphinctozoans', 'lithistids') rendering the class morphologically heterogeneous. Three subclasses are recog-
nised based on larval morphology, reproductive strategy, tetraxonid versus monaxonic megascleres, and microsclere geometries, although
these divisions require further refinement, and as such they are often ignored by contemporary authors. Demosponges include about 85%
of all described Recent species, some are freshwater but predominantly they are marine species living from the intertidal to the deepest
seas, with around 15 orders (the exact number still in contention), 88 families, and about 500 valid genera.
Keywords: Porifera; Demospongiae; Homoscleromorpha; Tetractinomorpha; Ceractinomorpha.

DEFINITION, DIAGNOSIS, SCOPE flow (archaeocyte/pinacocyte). Choanocyte chambers may be

eurypylous, diplodal or aphodal. Larvae are mostly parenchymella,
Synonymy but in some groups there are cinctoblastula or blastula forms pro-
duced. Both oviparous and viviparous reproductive strategies occur.
Demospongiae Sollas, 1885c.
Remarks
Definition
Demospongiae contains about 85% of all living sponges, with
Porifera with siliceous spicules andlor a fibrous skeleton, or about 6000 'valid' living species of demosponges already
occasionally without a skeleton. Spicules are either monaxonic described in the literature. There are potentially double this number
(either monactine or diactine) or tetraxonic (tetractine), never triax- of species, with the extant poriferan fauna estimated to comprise at
onic. The axial filament is embedded in a triangular or hexagonal least 15,000 species worldwide (Hooper & Levi, 1994; based on
cavity. surveys of unpublished museum collections). Furthermore, this
estimate is probably conservative as it largely neglects the grossly
Diagnosis under-sampled and under-studied encrusting, cryptic, sciaphilic
and other small taxa that pervade the many crowded marine com-
Encrusting, massive, lobate, tubular, branching, flabellate, cup- munities, such as the coral reefs (Hooper et ai., 1998). Most demo-
shaped or excavating sponges. Skeleton composed of spongin fibres sponges are marine, but several dozens of species occur in
alone or together with siliceous spicules which are usually divided freshwater habitats allover the world (so far excluding Antarctica).
into megascleres and microscleres. Megascleres are basically monax- Within Demospongiae three subclasses are recognized
onic or tetraxonic; microscleres are diverse, polyaxial or monaxone, (Homoscleromorpha, Tetractinomorpha, Ceractinomorpha), although
often quite elaborate in shape and ornamentation. Spongin is almost there is an increasing number of anomalies and exceptions between
universally present, forming discrete fibres or binding the skeletal otherwise closely allied family groups - based on putative morpho-
elements. In most cases the spicular skeleton and fibrous skeleton logical similarities and their differing reproductive strategies - sig-
form a combined reinforcement. Fibrils of collagen are ubiquitous. naling that some subclass taxa require tighter definition based on
Some groups lack spicular skeletons, but compensate that by building new data. Nevertheless, a rough division is possible based on the
an elaborate fibre skeleton. A few (unrelated) groups have no skeletal mutually exclusive presence of aster microscleres and reticulate
elements other than diffuse fibrillar collagen. Other minor groups skeletal elements. These might be equated with Tetractinomorpha
have developed a hypercalcified basal skeleton in addition to other and Ceractinomorpha, respectively (although the original contents
skeletal elements, or have a solid aragonitic structure lacking free of these subclasses were different, e.g., based on reproductive
spicules. In total these skeletal variants contribute to a heterogenous strategies, larval morphology). Homoscleromorpha is homoge-
morphological concept of Demospongiae, although there are other neous, with a single order and family (Homosclerophorida,
(non-morphometric) characters that provide more valuable clues as to Plakinidae). Thus, although some taxa do not appear to fit this
their phylogenetic affinities (e.g., possession of viviparity, model this subclass system offers an hypothesis to evaluate the
parenchymella larvae). The cellular elements are discrete, never syn- diverse demosponge orders and families.
cytial, and cellular diversity may be considerable. The aquiferous A nominal supraordinal group, 'Keratosa', was proposed by
system is of the leucon-type, although one family of deep-water early authors for sponges lacking a siliceous skeleton. Such
Poecilosclerida (Cladorhizidae) has lost its aquiferous system and a group, if it had any plausibility today, would contain the orders
has assumed a carnivorous lifestyle, and dependent on fibroblastic Dictyoceratida, Dendroceratida, Verongida and Halisarcida, but

15
16 Porifera· Demospongiae

these groups in reality are quite disparate, and hence the taxon has Tetractinellidae Sollas, 1880. Tetraxonia Dendy, 1905. Astro-
no real basis for support (Bergquist et al., 1998). tetraxonida Hentschel, 1909.
Several species of 'living fossils' previously assigned to Definition. Demospongiae with parenchymella or blastula
'Sphinctozoa' (now included in the order Verticillitida) and larvae, predominantly oviparous reproduction (although in some
'Sclerospongiae' are also undoubted Demospongiae, possessing a genera young sponges are apparently incubated within the parent
viviparous reproductive strategy and producing parenchymella and set free as small adults; Bergquist, 1978). Megascleres are
larvae, in Recent species at least (Vacelet, 1979b). These spec- tetraxonic and monaxonic, occurring together or separately;
ies presently sit uneasily within a homogeneous concept of microscleres are asterose forms and derivatives; skeletal structure
Demospongiae, based on a poriferan bauplan. is usually radial or axially compressed.
Levi (1953a, 1956a, 1957b, 1973) provides an outline and dis- Remarks. Tetractinomorpha was recognised as a poly-
cussion of the various proposals subdividing the Demospongiae at phyletic taxon several decades ago (Bergquist, 1978), but it has
suprafamily levels, and he was also the first to provide a compre- nevertheless persisted in the contemporary classification in a
hensive synthesis of the 'modern' sponge classification. slightly modified form to the present. Four 'well established'
orders are traditionally included within the concept of
Scope Tetractinomorpha: Astrophorida (also known as Choristida) and
Spirophorida - both of which continue to be treated as a single
We recognize three subclasses of demosponges with extant order Tetractinellida by a few authors (with a shared character of
representatives, distributed amongst 15 orders, 88 families and tetractines; Chombard et at., 1998) - Hadromerida, and Chondrosida.
ca. 1000 nominal genera (although only about 500 genera are A fifth order Lithistida is clearly polyphyletic, with many taxa
presently considered valid). Most of these genera are marine but showing major affinities to Astrophorida and some to
there are also about 40 genera confined to freshwater. Orders Hadromerida, to which they may be eventually allocated. Despite
included at this time are: (1) Homoscleromorpha: Homos- recent attempts (e.g., Kelly, 2000), resolving the systematic affini-
clerophorida; (2) Tetractinomorpha: Astrophorida, Chondrosida, ties of most 'lithistids' remains elusive, especially those that lack
Hadromerida, most 'lithistids' (polyphyletic), Spirophorida; free spicules that may provide phylogenetic clues (see also
(3) Ceractinomorpha: Agelasida, Dendroceratida, Dictyoceratida, Remarks for Ceractinomorpha). The order is herein abandoned
Halichondrida, Halisarcida, Haplosclerida, Poecilosclerida (which (Pisera & Levi, this volume).
includes some 'lithistids'), Verongida, and Verticillitida (the latter a Allocation of Latrunculiidae has also oscillated between
fossil order to which a single Recent genus is currently assigned). Tetractinomorpha (close to Hadromerida) and Ceractinomorpha
Several other widely employed ordinal taxa are allocated to exist- (close to Poecilosclerida) throughout its long and tortuous taxo-
ing orders, following contemporary revisions of these groups, nomic history. The family is now recognised as two separate taxa
although not presently universally accepted (e.g., Axinellida (see (with the resurrection of Podospongiidae), and currently referred
Halichondrida), Ceratoporellida (see Agelasida), Choristida (see to Ceractinomorpha (with some support from molecular and
Astrophorida), Petrosida, also known previously as Nephelio- biochemical evidence).
spongida (see Haplosclerida». The tetractinomorph order Axinellida Bergquist is also con-
sidered polyphyletic by most contemporary authors (although not
Recent reviews universally accepted by all), with families distributed amongst the
predominantly viviparous Ceractinomorpha and the predominantly
Levi, 1973; Bergquist, 1978; Hartman, 1982; Hooper & oviparous Tetractinomorpha. This treatment is followed in the
Wiedenmayer, 1994. present volume because it has a more sound morphometric base
(e.g., asterose vs. non-asterose taxa), but there is currently no
SUBCLASSES OF DEMOSPONGIAE published corroboratory molecular support one way or the other.
Hemiasterellidae and Trachycladidae are the only 'axinellid' fami-
Subclass Homoscleromorpha Levi, 1973 lies that now remain in this subclass, with suggested affinities to
the Hadromerida.
Other names. Microsclerophora Sollas, 1887. Carnosa
Carter, 1875c. Subclass Ceractinomorpha Levi, 1953a
Definition. Demospongiae with cinctoblastula larvae and
viviparous reproduction; skeleton composed of tetraxonic siliceous Other names. Cornacuspongiae Vosmaer, 1887, Monaxonidae
spicules and derivatives with equal rays (diods, triods, lophate Sollas, 1882b, Monaxonida Ridley & Dendy, 1887.
spicules), arranged around choanocyte chambers reflecting the canal Definition. Demospongiae with parenchymella larvae and
structure; no differentiation between megascleres and microscleres predominantly viviparous sexual reproduction; generally with both
although size differences do occur between types of spicules; a spicule skeleton and well-developed spongin fibres forming a
spicules usually small (100!-Lm or less), not localised to any particu- diversity of skeletal structures (although siliceous spicules are lost
lar region; choanocyte chambers with large numbers of choanocytes. altogether in three orders and in several other genera, and spongin
Remarks. The subclass presently contains a single order fibres are lost or greatly reduced in several genera scattered
and family. throughout the subclass). Spicules are monaxonic (either monacti-
nal (styles) or diactinal (oxeas-strongyles», never tetractinal
Subclass Tetractinomorpha Levi, 1953a (although modifications to the ends of some monaxonic spicules
occur and may appear to be superficially tetractinal); microscleres
Other names. Chondrospongiae Lendenfeld, 1886. are diverse (meniscoid (chelae), oxeote, toxote, spheres) but never
Spiculispongiae Gray, 1867. Tetractinellida Marshall, 1876. asterose.
Porifera' Demospongiae 17

Remarks. Acceptance of the concept of subclass (Alvarez et al., 2000a). Further literature citations are provided in
Ceractinomorpha has diminished in the contemporary literature, the respective family chapters below as evidence in support of these
largely due to the declining acknowledgement that viviparity is a allocations, but nonetheless, they remain contentious by some
pivotal phylogenetic character to differentiate the taxon from the authors.
oviparous Tetractinomorpha. An increasing number of oviparous Nine orders are potentially allocated to Ceractinomorpha based
taxa have been found in Ceractinomorpha (e.g., Petrosiidae, on the predominance of viviparity, and/or the production of
Verongida), with sound biological, biochemical or other evidence parenchymella larvae, and/or they possess close morphological
to support this scheme (e.g., Bergquist, 1980a). In contrast, the similarities with other VIVIparous ceractinomorphs. These
allocation of some other oviparous (or suspected oviparous) taxa to include: Agelasida (oviparous), Dendroceratida, Dictyoceratida,
Ceractinomorpha remains controversial. Halichondrida, Halisarcida, Haplosclerida, Poecilosclerida (with one
Of the families formerly included in the polyphyletic order (Raspailiidae) or possibly two oviparous families (Rhabderemiidae)),
Axinellida six are now included in Ceractinomorpha. Axinellidae Verongida (oviparous), Verticillitida, with another (Petrosiidae,
and Desmoxyidae are included by some authors in the oviparous) family included within the predominantly viviparous
Halichondrida (e.g., Van Soest et al., 1990), even though they may Haplosclerida. Axinellidae is also included within the viviparous
be exclusively oviparous (although data are still rudimentary for Halichondrida, and despite recent molecular support for this alloca-
most genera). In the case ofAxinellidae there is molecular support tion it is still not universally accepted by all contemporary authors.
from 28S rDNA analyses that shows the family is (a) polyphyletic Several genera with exclusively or predominantly 'lithistid' or
but (b) indeed close to Halichondriidae (Alvarez et al., 2000a). 'sublithistid' skeletons are also included in various ceractinomorph
In the case of Desmoxyidae this relationship is far more speculative. families (Le., Crambe, Desmatiderma, Desmanthus, Lithochela,
Desmacellidae demonstrate both modes of reproduction. For exam- Petromica), and a few 'sublithistid' species are found in genera
ple, viviparity has been recorded for several species of Biemna with a predominantly 'non-lithistid' grade of construction (e.g.,
(with some also producing gemmule-like asexual bodies, similar to Esperiopsis). This issue is still controversial. One school of
asexual resting bodies seen in some Mycale), whereas Neofibularia thought suggests that the 'lithistid' grade of construction may have
nolitangere is clearly oviparous (e.g., see reviews by Fell, 1993; 'persisted' from a once more-widespread fauna (e.g., Ordovician
Boury-Esnault & Jamieson, 1999, and literature contained therein). and Devonian sponge reefs), with free spicules indicative of phylo-
The likelihood that the family is polyphyletic (i.e., their phylogeny genetic affinities (e.g., Van Soest & Zea, 1986; Hooper & Levi,
is erroneously based on morphological convergence) is negligible as 1989; Van Soest et al., 1990). Another school suggests that desmas
similarities between Desmacella, Biemna and Neofibularia are con- are indicative of phylogenetic affinities and taxa possessing them
vincing, and it can only lead to the conclusion that there is a dis- are recognisable at least at the family level (e.g., Kelly, 2000;
crepancy in the interpretation of mode of reproduction as Pisera, pers. comm.; Pisera & Levi, this volume). These arguments
phylogenetic markers for otherwise very closely related sponges remain unresolved without other supporting evidence, except in
(Le., concepts of 'vivipary' and 'ovipary' are non-homologous cases like Esperiopsis and Crambe which have clear affinities to
within the sponge phylogeny). Raspailiidae (oviparous, where Poecilosclerida based on the possession of microscleres that are
known) is also assigned to Poecilosclerida based on morphological unique to this order.
similarities to the viviparous family Microcionidae. While this rela-
tionship is supported by character analysis and chemotaxonomic Conclusions
data (Hooper et al., 1992), no sequence data is yet available to sup-
port this hypothesis. Rhabderemiidae (reproduction unknown) is Given all of these anomalies it is questionable whether repro-
less confidently included in Poecilosclerida. It possesses diverse ductive strategy is a suitable biological criterion to formulate
microscleres that are assumed homologues of poecilosclerid a phylogenetic hypothesis at the level of subclass, or indeed if the
microscleres (with this assumption supported by possession of true subclass classification is necessary at all. For this reason many con-
toxas in three species of Rhabderemia), but most of these microscle- temporary authors prefer not to group the different orders into sub-
res are unique and therefore potentially analogues of typical poe- classes, but we suggest that these higher taxa have value as
cilosclerid morphologies. Agelasidae also theoretically belong to working hypotheses to reconstruct evolutionary relationships
Tetractinomorpha (oviparous), but its allocation has oscillated between the diverse orders and families of demo sponges. It there-
between Ceractinomorpha and Tetractinomorpha. The family is fore remains a challenge for the future to re-evaluate and perhaps
now assigned to its own monophyletic order (Agelasida), and its expand this subclass classification for the Demospongiae,
inclusion within Ceractinomorpha has some molecular support incorporating new datasets.

KEY TO ORDERS OF RECENT DEMOSPONGIAE

This key may not always result in the assignment of each individual sponge specimen to its proper order due to the imperfectness of
juvenile or growth stages, phenomena like reduced spiculation in carbonate environments, or deviating species associated with orders
only through circumstantial similarity with species showing a full complement of ordinal characters. Consequently, several redundancies
are deliberately included in the key, but it is necessary to use it with care.

(1) Skeleton absent .................................................................................................................................................................................. 2

Skeleton present ................................................................ ,........... .......... ................ ................. .......................... ...... .......... ................. 7
(2) Firm sponges with cartilaginous consistency ...................................................................................................................................... 3
Soft sponges ....... .......... ................ ................. .......................... ........................................... ...... ............... ........ ... ................ ................. 4
18 Porifera· Demospongiae

(3) With leuconoid aquiferous system and diplodal choanocyte chambers; ectosome thick ... Homosclerophorida (Pseudocorticium)
With a well developed cortex made of thick fascicles of fibrillar collagen, numerous spherulous cells, and inhalant apertures
localised in special structures .................................................................................................................. Chondrosida (Chondrosia)
(4) With fibrillar collagen only ............................................................................................................................................................... 5
With a nodular spongin fibre skeleton ....................................................................................................... Chondrosida (Thymosia)
(5) Choanocyte chamber eurypylous, simple .......................................................................................................................................... 6
Choanocyte chambers tubular and branched, size about 100 fLm; ectosomal and subectosomal collagen highly organised and struc-
turally diversified ............................................................................................................................................................. Halisarcida
(6) Ectosome thin, with sylleibid-like aquiferous system; choanocyte chambers eurypylous, rounded, less than
60 fLm diameter ............................................................................................................................... Homosclerophorida (Oscarella)
With a thin cortex enriched with fibrillar collagen parallel to the surface, a superficial cuticle and pore-sieves
may be present ..................................................................................................................................... Chondrosida (Thymosiopsis)
Ectosome strongly collagen-reinforced and bounded by a distinct skin, with spherulous cells -10 fLm in diameter are common
throughout the mesohyl but particularly concentrated in the ectosome; sponge attaining a thickness of only about 5 mm; choanocyte
chambers large and sac-shaped ...................................................................................................................... Verongida (Hexadella)
(7) Megascleres present ....... ....... ................ .......... ....... ..... ... ... .... ...... ...... .......... ...... ................ ............... ...... ........... .......... ...... ......... ........ 8
Only asterose microscleres present .......................................................................................................... Chondrosida (Chondrilla)
Siliceous spicules absent (or secondarily lost) ................................................................................................................................ 17
(8) Spicules exclusively verticillate-spined styles or oxeas ..................................................................................................... Agelasida
Spicules may be spined or smooth but are not exclusively verticillate- spined ......... ...... ......... ...... .... ....... ... ... .......... ...... ....... ... ....... 9
Megascleres always include articulated siliceous desmas, with or without free
spicules ............................................................................................................................ Demospongiae 'lithistids' (polyphyletic)
(9) Megascleres are all monaxones ....................................................................................................................................................... 10
Megascleres include diods and/or triods, megascleres and microscleres undifferentiated, sometimes spicules are lost completely and
sponge may be superficially confused with compound ascidians ........ ................ ...................... ...... .......... .... ... Homosclerophorida
Megascleres include triaenes ........................................................................................................................................................... 16
(10) Megascleres exclusively diactines (oxeas and/or strongyles) ......................................................................................................... 11
Megascleres diverse or exclusively monactinal (tylostyles, styles, strongyloxeas) ........................................................................ 15
(11) Asterose microscleres ...... .... .... ... ...... ......... ....... ............... ................ ... ... ....... ... ...... ...... .... ...... ..... ................. .......... .... ... Astrophorida
No asterose microscleres ................................................................................................................................................................. 12
(12) Megascleres arranged in an isodictyal or anisodictyal reticulation ................................................................................................. 13
Megascleres arranged in a confused manner or plumose or plumo-reticulate .... ....... ...... ............... .......... .............. .... ..... .......... ..... 14
(13) Microscleres include chelae, megascleres often localized to distinct regions (e.g., inside fibres), sand/detritus may replace megascle-
res completely ............................................................................................................................................................ Poecilosclerida
No chelae; microscleres absent or restricted to sigmas, toxas, raphides, amphidiscs or microspined oxeas, megascleres diactinal
usually producing well-formed structures such as triangular, rectangular or polygonal meshes ................................ Haplosclerida
(14) Microscleres include chelae and or sigmas or toxas .................................................................................................. Poecilosclerida
No chelae, sigmas or toxas ......................................................................................................................................... Halichondrida
(15) Microscleres may be absent or may include asterose and monaxonic forms (microxeas, spirasters); skeleton peripherally radiate
forming palisades of spicules at the surface .... ................ ...... ...... .... ...... ...... ....... ... ................ ...... ............... ........... ....... Hadromerida
Microscleres include chelae and/or sigmas, occasionally microscleres are absent ..................................................... Poecilosclerida
No asters, and no other microscleres other than trichodragmas (or raphides); skeleton peripherally tangential or undifferentiated,
main skeleton composed of a criss-cross of spicules, or compressed into a distinct axis, or with plumose, plumo-reticulate or den-
dritic mineral skeleton, fibre system poorly developed or absent ........ ........................... ... ... ......... ...... ................. ..... Halichondrida
(16) Microscleres sigmaspires (rugose c- or s-shaped), spherical growth form usual, radial pattern of
triaenes and oxeas ..... .... ............. ................ .......... ............ ................ ...... ... ....... ...... .......... ................ ...... ....... ... ............. Spirophorida
Microscleres rugose sigmaspires, no oxeas, no radial skeleton, no spherical growth form .......................... Spirophorida (Samidae)
Microscleres asters or streptoscleres, large oxeas always present, sometimes with triaenes, skeleton only obviously radial at the sur-
face .. ...... ......... ....... ..... .......... ................ ...................... ...... ................ ....... ... ...... .......... ...... .......... ...... .......... .................. Astrophorida
(17) Solid carbonate skeleton, lacking free spicules, with a solid cortex producing a series of chambers on top of each other, the youngest
(uppermost) chambers lined with living tissue .............................................................................................................. Verticillitida
Skeleton of discrete spongin fibres .................................................................................................................................................. 18
(18) Fibres generally well laminated, containing a cellular mass visible as a dark pith in transmitted light, without differentiation of
primary or secondary elements, many taxa aerophobic (darken in contact with air) ........................................................ Verongida
Fibres contain a core of sand or spicule fragments or are entirely free of inclusions ..................................................................... 19
(19) Skeleton an anastomosing system of interconnected fibres, often well developed and relatively homogeneous fibre construction with
2-3 different sized networks, consistency not collagenous ........................................................................................ Dictyoceratida
Skeleton consists of dendritic fibres arising from basal attachment, with fibres strongly laminated ....................... Dendroceratida
Skeleton with reticulate, plumoreticulate or plumose fibres containing sand or spicule fragments, with vestigial spicules (check for
microscleres or echinating spicules) or occasionally no spicules at all ..................................................................... Poecilosclerida
Fibre skeleton well-developed, more-or-Iess regularly reticulate, and also with a tangential ectosomal (tertiary) network of fine
aspicular fibres and foreign material, whereas choanosomal fibres are aspicular and with only foreign material (or sometimes
extremely vestigial oxeas) .......................................................................................................................... Haplosclerida (Dactylia)
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

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Compilation: Philippe WilIenz

Institut royal des Sciences naturelles de Belgique, Rue Vautier 29, B-1000 Bruxelles, Belgium. (philippe.willenz@naturalsciences.be)

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19
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CABIOCH, L. 1968a. Contribution it la connaissance des peuplements ben- CARTER, H.J. 1872a. On two new Sponges from the Antarctic Sea, and on
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CALCINAI, B., CERRANO, e. & BAVESTRELLO, G. (in press). A Philippine Islands. Annals and Magazine of Natural History (4) 10(56):
new species of Scantiletta (Demospongiae, Clionidae) from the 110-113.
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CAPON, R.J. & MACLEOD, J.K. 1987a. Structural and Stereochemical 12(67): 17-30, pI. I.
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CAPON, R.I. & MACLEOD, J.K. 1987b. 5-Thio-D-mannose from the together with Facts elicited from their Deciduous Structures and
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Chemical Communication 15: 1200-1201. CARTER, H.J. 1874a. Descriptions and Figures of Deep-sea Sponges and
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Trikentrins: novel indoles from the sponge Trikentrion flabelliforme. 'Porcupine', chiefly in 1869; with Figures and Descriptions of some
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© Kluwer AcademiclPlenum Publishers, New York, 2002

Non-Lithistid Fossil Demospongiae - Origins of their Palaeobiodiversity and

Highlights in History of Preservation

Joachim Reitnerl & Gert Worheide2

lGeowissenschaftliches Zentrum Giittingen GZG, Abt. Geobiologie, Universitiit Giittingen, Goldschmidtstr. 3, D-37077 Giittingen,
Germany. Greitne@gwdg.de)
2 Queensland Museum, P.O. Box 3300, South Brisbane, Qld, 4101, Australia. (GertW@qm.qld.gov.au)

Available evidence suggests that the first demosponges occur in the Late Proterozoic, with forms characterized by bundles of long mon-
axonic spicules. In the Middle Devonian the first modem forms of Dendroceratida, 'axinellids' (mostly halichondrids), and first hap-
losclerids appeared. An important boundary for the demosponges is the Late Devonian extinction event, which caused a complete
overhaul of demosponge communities. The Late Permian and the Triassic, especially the Late Triassic, are the main eras for coralline
demosponge radiation and dominance, in which some modem taxa occur for the first time (Ceratoporella, Astrosclera, Vaceletia). In the
Late Jurassic the freshwater environments were occupied by certain (marine) demosponges, mostly Haplosclerida. The importance of
coralline demosponges as primary reef-builders decreases up to the Late Cretaceous.
Keywords: Porifera; Demospongiae; fossil taxa; coralline demosponges.

INTRODUCTION most significantly, the discrete siliceous spicules that are often cru-
cial to taxonomic identification. The silica of spicules is replaced
Demospongiae are an important group of sessile benthic organ- by other minerals, mostly calcite or secondary silica or pyrite.
isms showing a special potential for fossil preservation. Up to Isolated spicules are common in certain deposits and some of them
60% of their biomass consists of microorganisms, predominantly are of taxonomic and phylogenetic importance, but only few stud-
bacteria, in some cases Archaea, and partly of anaerobic taxa ies have yet been made, especially of the phylogenetically impor-
(sulfate-reducing bacteria (SRB) and Archaea) (Reitner & tant Early Palaeozoic strata. In special fossil lagerstiitten, like the
Schumann- Kindel, 1997). The SRB playa central role in the forma- Burgess shale deposits, entirely preserved sponges are present.
tion of Porifera-rich mud mounds and certain microbialites. To a Similarly, one of us (JR) has recently discovered entirely preserved
great extent the micrite in these structures was formed by various non-lithistid demosponges in Cambrian and Middle Devonian
types of sponge related microorganisms, (and/or) degraded and re- microbialites. Due to the large numbers of sponge-related bacteria
organized organic material originated from sponges (Reitner et al., in many demosponges the sponge tissue may mineralize rapidly,
1995), but some portions of these build-ups are also influenced by controlled by sulfate reduction and/or ammonification. In this par-
seeps and seepage. In the Upper Devonian a fundamental push in the ticular case certain sponge biomarkers (chemofossils) are pre-
development of sponges took place, from the long-existing, stem- served and thus allow chemotaxonomic examination.
group Palaeozoic taxa to modem groups of which the monophyla are This chapter does not provide a complete overview on fossil
still present. The driving forces for this change are so far unknown, demosponges - which could occupy a volume in itself, and will be
but most likely include medium-term fundamental oceanic changes more comprehensively addressed in the forthcoming revision of
(water chemistry, nutrient situation) producing new niches for new the Treatise of Invertebrate Paleontology (J.K. Rigby et ai., edi-
taxa, especially in deeper water and other protected areas. Porifera- tors). The purpose of this chapter is to demonstrate some highlights
rich build-ups and black shale environments provide a window to in the history of palaeontological preservation, which provides a
answer this question, which concerns reef-like structures, rich in general overview on the early development of the main mono-
micrites, without a metazoan framework. An increase in nutrient phyletic groups of non-lithistid demosponges. Some coralline
input could have impacted significantly on oceanic environments, sponge taxa are also included in this treatment, but these taxa are
possibly through a significant increase of alkalinity as well as restricted to those with modem representatives and those that rep-
by increased input of hydrothermal calcium and other metal resent important phylogenetic lines.
cations. Porifera react very rapidly to such changes, whereas sponge-
associated bacteria are even more sensitive to changes within the
general nutrient environment (eutrophism of organic and inorganic PHYLOGENETIC ASPECTS
nutrients), as we know from field experiments on modem Porifera.
In this way, the rapid development during the Devonian of the poly- New data suggest that 'sponges' are probably not mono-
phyletic coralline sponges (the so-called 'sclerosponges') may have phyletic (Borchiellini et ai., 2001), with Calcarea indicated to have
been a reaction to the global oceanic change during this critical inter- greater similarities with Cnidaria (perhaps the Ctenophores) than
val, and linked with increasing alkalinity and lor massive increase of to sponges with siliceous skeletons. These data include nucleic
the overall Ca2 + dissolved in ocean water (Arp et ai., 2001). acid analysis (18s rDNA), the spectra of fatty acids and the occur-
Little is known about the fossil record of non-lithistid demo- rence of sponge-related bacteria. In these latter characters the
sponges, in contrast to the 'lithistid' demosponges, and taxa with a Calcarea do not exhibit the very characteristic long-chained demo-
secondary calcareous basal skeleton, due to the poor preservation spongic acids, which are very good biomarkers for all demo-
potential of the soft tissue, the collagenous spongin skeleton and, sponges and hexactinellids (Thiel et al., 2002), and in most

52
Porifera· Demospongiae· Non-Lithistid Demosponges 53

Calcarea mesohyle bacteria are missing. This evidence certainly A

suggests a more distant phylogenetic position for the Calcarea,
although it does not necessarily imply sponge paraphyly.
Demosponges demonstrate the following clades based on
morphological characters:

(1) Homosclerophorida.
(2) Tetractinomophora, including the Astrophorida, Spirophorida
and Hadromerida.
(3) Ceractinomorpha, including the aspicular Dendroceratida,
Dictyoceratida and Verongida (so-called 'Keratosa') and spicu-
lose taxa like the Poecilosclerida, Haplosclerida, Halichondrida
('Axinellida') and Agelasida.

THE CONCEPT OF SPONGE-BIOFILMS

The origin of the sponge bauplan is probably related to the

B
development of special stromatolite-forming biofilms during the
Early Proterozoic - at the same time as endosymbioses of eukary-
otic cells evolved. This hypothesis is supported by direct evidence
from 1.8 billion year old stromatolites containing sponge-specific
C30 steranes and 24-isopropylcholestanes (Moldowan et al.,
1994a,b), which are extremely abundant in demosponges and
therefore good biomarkers for these animals (Thiel, 1997). A fur-
ther indirect argument for a phylogenetic relationship between the
modem morphology of sponges and ancestral biofilms is the pres-
ence of mid-chain branched carboxylic acids (MBCA) in modem
demosponges, which are probably related to anaerobic sulfate-
reducing bacteria within the sponge mesohyl (Thiel et al., 1999).
This type of bacterial biomarker was previously unknown in any
other marine environment but was recently discovered in het-
erotrophic biofilms of the modem highly alkaline Walker Lake in
Nevada. We hypothesise that Early Proterozoic oceanic environ-
ments had a higher alkalinity than Recent seas (Kempe &
Kazmierzcak, 1994), and thus the occurrence of certain species of
bacteria adapted to alkaline environments found in modem demo-
sponges - especially in bacteria-rich demosponges like the possi-
ble ancestral astrophorid taxon Geadia - supports this assumption.
Our working hypothesis - that sponges are highly developed
biofilms with a close relationship to choanoflagellate eukaryotic
cells - is based on the analysis of the sponge-related microorgan-
isms and biofilm analyses from extreme environments.
Demosponges exhibit a mesohyle community of Eubacteria,
mainly gamma and alpha proteobacteria, few gram-positive
bacteria and only in a few cases the Archaea (Schumann-Kindel
et al., 1997). In contrast to the demosponges all taxa of the
Hexactinellida demonstrate a community of microorganisms
dominated by Archaea (Thiele et al., 2002).

PALAEONTOLOGICAL HISTORY OF DEMOSPONGES

Late Proterozoic Demosponge Remains

Fig. 1. A, Late Proterozoic (ca. 555my) sponge with demosponge affini-
The spicule record of sponges starts in the Late Proterozoic. In ties from the Claudina-reefs of the Kuibis Formation (Zebra River,
most cases these are simple monaxonic types with uncertain affini- Southern Namibia). The small sponges with tylostyle-type spicules are
ties. Some of them show hexactinellid affinities with simple hexa- located between Claudina-tubes. Claudina has probably pogonophoran
ctines and stauractines. The oldest spicules with demosponge affinites and represents the first organism in earth history with a calcified
skeleton - onset of biomineralisation. B. Late Proterozoic (Ediacaran)
affinities were found by the senior author in ca. 750my old sponge with demosponge affinities from the White Sea coast. Northern
Noon Day Dolomite in Nevada, and in the Neoproterozoic Russia. This organism is constructed of long monaxonic spicules
(ca. 555my) Claudina-Reefs of southern Namibia. These latter reefs (scale 7 em).
54 Porifera· Demospongiae • Non-Lithistid Demosponges

demonstrate bundles of styles to tylostyles, typical demosponge (Reitner, 1992, Reitner & Mehl, 1995), which suggest a close rela-
spicules. Cloudina may be related to pogonophoran worms and the tionship to the Tetractinellida (Fig. 2). It is questionable, however,
Cloudina-mound environment may be related to hydrothermal seep- whether these spicules are part of the sponge skeleton or of
age. Small sponges are common between Cloudina-rich thrombolitic allochtonous origin. The Archaeocyatha possess a Mg-calcite basal
pillars (Fig. IA). Sponge remains with demosponge affinities from skeleton with distinctive microstructural characteristics which are
the Ediacara-type environment of the White Sea coast (Russia) are also know from the modem demosponge aspicular 'sphinctozoan'
currently under investigation (Fig. IB). These consist of sponges taxon Vaceletia, which occur first in the Middle Triassic (Reitner,
with radiating monaxonic spicule bundles characteristic of early 1992; Reitner et al., 1997). This taxon has some affinities to the
demosponges. This type is also known from the Early Cambrian ceractinomorph demosponges (see Vacelet, and Senowbari-Daryan
(Atdabanian) Sansha formation of China (Saetaspongia densa) & Garcia-Bellido chapters in this volume). In the older parts of the
(Steiner et al., 1993). The small 'sponges' and so-called 'sponge lar- archaeocyath skeleton a calcification phenomenon of a lens-shaped
vae' of the Wengan phosporites are highly questionable as to their microstructure is present, which is known from the deep skeletal
affinities to Porifera (Li et al., 1998). Some of these phosphatized structure of Vaceietia - called CWD (Ca-Waste-Deposits) (Reitner,
larvae exhibit some morphological similarities to simple demo- 1992). This phenomenon is often observed in similarly formed fossil
sponge parenchymella larve (Tethya-type), but this interpretation of sphinctozoans, like the middle Triassic Stylothalamia Ott, 1967 and
the fossil evidence is still equivocal. Late Triassic Cassianothalamia Reitner, 1987b and Uvanella
(Reitner, 1987b). This type of a basic biomineralization process is
Early Cambrian Non-Lithistid Demosponges based on Ca-detoxification and could be a model for all irregular,
micritic-granular basal skeletons of 'stromatoporoid' and 'thalarnid'
Archaeocyatha - demosponges with uncertain affinities. grades of organisation. Of special interest is that all types of
The first sponges in the fossil record with a calcareous basal skele- Archaeocyatha, the oldest known coralline sponges, exhibit this very
ton occur in the Early Tommotian - the Archaeocyatha (see specific type of calcification mode. The modem Vaceletia Pickett,
Debrenne et ai. chapter in this volume). Archaeocyatha are classi- 1982, thus, may be a modem 'archaeocyath' sponge (Reitner et al. ,
fied in two main taxa: the Regulares and the Irregulares (Debrenne & 1997,200lb).
Zhuravlev, 1992, 1994; Debrenne & Reitner, 2000). The phyloge- Early Cambrian Tetractinellida. The first record of
netic position of the entire Archaeocyatha is still unclear. Spicular tetractinellid spicules are four-rayed calthroPs from the base of the
skeletons are not known from most of these sponges, except in a Cambrian of the Flinders Ranges (South Australia) (Bengston
few cases where tetractinellid spicules have been occasionally et al., 1990).
incorporated into rapidly calcified buds of some archaeocyaths The Geodiidae are probably one of the most ancestral of
demosponge groups as suggested by their fossil record. They
exhibit the ancestral character of radially arranged larger spicules,
a plesiomorphy of the stem group of demo sponges (Reitner &
Mehl, 1996). The oldest remains of geodiid spicules are known
from the Early Cambrian deep water archaeocyath reefs of the
Mount Scott Range near the Flinders Ranges (South Australia)
(Reitner & Mehl, 1995) (Fig. 3). The observed spicules are various
types of large triaene dermal spicules, phyllotriaenes, and peculiar
kidney-shaped sterrasters with the characteristic impression of the
spicule forming cell (Gruber & Reitner, 1991). These spicules are
exclusively preserved in a polycrystalline calcite, in contrast to the
associated spicules of calcareous sponges which exhibit a charac-
teristic monocrystalline structure. However, these fossil astrophorid

Fig. 2. A, Coscinocyath archaeocyath from the Flinders Ranges (South Fig.3. Geodiid spicules remains from the archaeocyath reefs of the Mount
Australia) (scale 5 mm). B, Intramural, mostly tetractinellid spicules are Scott Range (Flinders Ranges, South Australia) (Reitner & Mehl, 1995).
common within the endothecal buds (Reitner, 1992; Reitner & Mehl, 1995) A, Megascleres resemble large triaenes (scale 200 fLm). B, The kidney-
(scale 1 mm). shaped structures are interpreted as remains of sterrasters (scale 1()() fLm) .
Porifera· Demospongiae • Non-Lithistid Demosponges 55

spicules differ in many aspects from Recent ones. Most of the Early Cambrian Demosponges with 'Axinellitl'-Type Spicule
observed spicules are larger (600/-lm - 3 cm) and thicker (more Arrangement. In the Lower Cambrian deep water archaeocyath
than 50/-lm) than those of modem species. reefs of the Mount Scott Range (Australia) entirely preserved small
Early Cambrian Hadromerid-Type Spicules. In additional sponges were observed, constructed of radially arranged monaxonic
to the astrophorid spicule remains in the Early Cambrian deep spicules. These demosponges (about 1 cm in diameter and smaller)
water archaeocyath reefs of the Mount Scott Range there are also are extremely abundant within the deep water archaeocyath facies
large tylostyle megascleres, which may have hadromerid affinities. and they are part of a spiculitic matrix. This occurrence is the first
Unfortunately, no hadromerid spicule arrangements were observed record in Earth's history of an autochtonous spiculite which are oth-
within these strata, and therefore their classification in the erwise common in modem temperate and polar seas (Henrich et al.,
Hadromerida is equivocal (Reitner & Mehl, 1995, 1996). 1992). The preservation of these small demosponges is not so good
Cambrian Demosponge Taxa with Uncertain Phylogenetic as to allow their accurate classification. The monaxonic spicule
Affinity. From the Chinese locality Sansha a very diverse hexa- arrangement is definitely not of the Tetractinellida model, but is
ctinellid fauna is known with only one spherical sponge, reminiscent of the 'Axinellida' bauplan.
Saetospongia densa Steiner et ai., 1993, which has alleged Choiidae de Laubenfels, 1955. From the Lower and Middle
demosponge affinities (Steiner et al., 1993). Saetospongia densa Cambrian strata relatively large demosponges are known which are
has bundles of small diactine spicules, an arrangement not seen called Choia Walcott, 1920 and Choiella Rigby & Hou, 1995. Both
previously in hexactinellids. Similar types of sponges from the taxa are very similar and probably synonyms (Mehl, 1999). The
lower Cambrian Sirrius Pas set of northern Greenland were investi- Choiidae exhibit radially arranged, several cm long large styles,
gated by the authors; they exhibit the same spicule arrangement. In diactines and small styles (1 cm and less). They are typical ances-
this particular case the spicules are plumose in arrangement (Fig. 4). tral demosponges with radially arranged simple spicules, with
postulated affinities to the Halichondriidae. These sponges
exhibit some similarities to modem hymeniacidonids (family
Halichondriidae). In any case, the Choiidae are members of the
stem group of the Demospongiae (Fig. 5).
Halichondritidae Rigby, 1986c. These sponges exhibit a
strong 'axinellid' spicule arrangement and probably developed
from the Choiidae. They have lost the entirely radially arranged
spicule architecture and the sponges are more cup-shaped. Oxeas
and styles are arranged axially, and in this regard differ from the
Choiidae. Both types occur in the same environments and are
known from the lower Cambrian Sirrius Passet from Greenland
and from the middle Cambrian Burgess Pass (Canada). Two taxa
are known, Halichondrites Dawson, 1889 and Pirania Walcott,
1920. The latter is characterized by very long styles (Pirania-style)
which protrude from radially arranged small oxea. The
Halichondritidae are very close to the Ordovician taxon

Fig. S. Choia sp. from the Lower Cambrian Sirius Passet from Northern
Fig. 4. Saetospongia densa, a demosponge from the Sansha fossil lager- Greenland (coIl. J. Peel). These spherical demosponges are good represen-
statten, China (Steiner et ai., 1993) (scales: A, I cm; B, 5 mm). tatives of the demosponge stem line (scale I cm).
56 Porifera· Demospongiae • Non-Lithistid Demosponges

Saccospongia Bassler, 1889 which also contains a certain type of Highlights of the Cambrian Demosponges. In the
desma (heloclones) (Mehl, 1999). The occurrence of this latter Cambrian the diversification of the main demosponge clades was
body plan can be seen in the enigmatic Esperiopsis desmophora realised. It is possible to distinguish between tetractinellid taxa
Hooper & Levi, 1989 (Recent deep-water Poecilosclerida from the (Geodiiidae), 'axinellid'lhalichondrid taxa (Choiidae, Pirania), and
continental slope off the Great Barrier Reef). Thus, the 'axinellid' ceractinomorph 'keratose' types (Vauxiidae). In the Middle
type of architecture is highly convergent within Demospongiae, Cambrian a new character evolved within both the main clades of
with some of the Recent taxa assigned to Halichondrida (Van the demosponges: desma spicules, which form rigid choanosomal
Soest, 1991; Reitner, 1992) (Fig. 6), Poecilosclerida (e.g., skeletons and increased their preservation potential. Desma-bearing
Raspailiidae) and Hadromerida (e.g., Hemiasterellidae) (see rele-
vant chapters in this volume).
Choiidae and Halichondritidae are a well-defined mono-
phyletic group and may be the stem group of the Saccospongiidae.
In any case, these demosponges exhibit many typical 'axinellid'
characters and they may be ancestors of the modem 'Axinellida' /
Halichondrita, which were first documented from the Middle
Devonian of the Boulonnais (France).
Early 'Keratose'Demosponges. Ceractinomorph sponges
without spicules were summarized in the taxon 'Keratosa' Grant,
1861. 'Keratosa' s.s. excludes the Halisarcidae Vosmaer 1887,
where the latter does not have spongin fibres common to all others.
In the modem scheme the term 'Keratosa' has virtually disap-
peared as a clade, although they may represent a grade of construc-
tion, with Recent species assigned to the Dendroceratida,
Dictyoceratida or Verongida. The fossil record of these sponges is
poor due to the fact that they have no spicules. There are some
Fig. 7. 'Sandsponges' from the late Proterozic Ediacaran of the Ukraine.
exceptions, however, when the entire tissue has been preserved due These structures may be remains of so-called 'Psammosponges'. Many
to rapid calcification via sulfate reduction (see section on Devonian 'keratose' demosponges agglutinate sandgrains to support their collagenous
'keratose' sponges). Some of these taxa agglutinate detritic mate- skeletons (scale I cm).
rial to support their spongin fibre skeleton. The amount of aggluti-
nated material may reach more than 50% of the entire sponge
volume. There are some Late Proterozoic small sand-structures
which resemble possible fossil remains of 'sand-sponges' (Fig. 7).
The best known taxa are the middle Cambrian Vauxiidae
Walcott, 1920 from the Burgess Shale. These sponges exhibit a
basket-shape fibrous, aspiculate skeleton, probably made by strong
spongin fibres. Based on this feature. Rigby (1986c) discussed a
strong affinity to modem 'Keratosa'. It is difficult to decide, based
on this material, whether or not this assumption is correct. In any
case, the Vauxiidae are representatives of the stem-group of the
'keratose' demosponges (which excludes the Halisarcidae) (Fig. 8).

Fig. 6. Halichondrites sp. from the Cheniang fossil lagersUitten (China). Fig. 8. Vauxia gracilenta from the middle Cambrian Burgess Shale. These
This demosponge is probably evolved form Choia and exhibits some simi- sponges are 'keratose' demosponges with affinities to the Verongida
larities to axinellid demosponges (scale 1 cm). (scales: A, 4cm; B, 1 mm).
Porifera· Demospongiae • Non-Lithistid Demosponges 57

demosponges (see chapters on 'Lithistida') have dominated the the Boulonnais displays a much better preservation of authochtho-
fossil record of demosponges since this time. Important is the fact nous compounds among the organic sediment fraction. This may be
that sigma-type microscleres were recorded (Mehl, 1999) in a close due to the tight sealing of the small carbonate deposits by fine-
relationship with the first middle Cambrian 'lithistid' demosponges grained hemipelagic sediments and a low degree of alteration during
(Rankenella mars Gatehouse, 1968) from the Ranken Limestone carbonate diagenesis. A remarkable feature of the Boulonnais sam-
(Northern Territory, Australia) and the Georgina Basin, which illus- ples is the occurrence of a pronounced series of bicyclic sesquiter-
trate the Cambrian diversification of the demosponges into the main penoid hydrocarbons, some of which displayed mass spectrometrical
clades Tetractinomorpha and Ceractinomorpha. Within the Middle characteristics consistent with a drimane-type carbon skeleton
Cambrian Australian Georgina Basin and Daly Basin characteristic (Reitner et al., 200 la). The occurrence of fossil sesquiterpenoids has
tetractinomorph spicules are commonly documented as various been demonstrated in several studies, but the significance of these
types of triaenes (orthotriane, trichotriane) and aster-microscleres, compounds has not yet been fully unravelled (Peters & Moldowan,
such as oxyasters (Van Kempen, 1978, 1990a; Kruse, 1990; Mehl, 1993). Some structures have been related to inputs from terrestrial
1999). However, most of these spicules are related to the 'lithistid' sources, whereas others, including the drimanes, have been sug-
demosponges (e.g., Anthaspidellidae-Orchocladina). It is difficult gested as microbial biomarkers. Notably, sesquiterpenoid natural
to decide whether these free tetractinellid and ceractinomorph products occur as main compounds in numerous recent demo-
spicules are related to 'lithistids' or to desma-free demosponges. In sponges, and are particularly prominent among the Halichondrida,
any case, the main steps of demosponge phylogeny had been and notably in the Axinellidae (e.g., Bergquist, 1979). An interpreta-
achieved in the Cambrian. tion of the sesquiterpenoids found as diagenetic derivatives of
sponge natural products would be in full accordance with the
palaeontological evidence found in these deposits.
Middle Devonian Non-Lithistid Demosponges from the This result is spectacular, since it gives evidence of the direct
Boulonnais (Northern France): the Beginning of connection with the 'axinellid' plumose spicule bundles ! In terms
Modern-Type Demosponge Communities of Earth's history this shows the earliest appearance of this modem
sponge taxon.
The Middle Devonian microbialites from Boulonnais, Northern The palaeobiodiversity of non-lithisitid demosponges within
France (Mistiaen & Poncet, 1983) show a characteristic ecological these cryptic environments looks very modem, and was never
sequence of rugose corals, cyanobacteria/algae (Rothpletzella), observed in comparable Early Palaeozic environments. It is the
microbialite crusts, and Porifera (Demospongiae) documenting a first time in Earth's history that cryptic demosponges playa signif-
facies poor in light (Reitner et al., 2001a). Within cryptic space hali- icant role in reef mound environments. The community is domi-
chondridl'axinellid' demosponges are common (Fig. 9), as well as nated by halichondrid demosponges, followed by 'keratose'/
relicts of dendroceratid demosponges (Fig. 10), which show dendroceratid sponges, and few haplosclerid demosponges.
an excellent preservation of the spicule skeletal frame due to automi- Hexactinellids have never been observed within this facies. The
crite formation. This result is supported by chemofossils, like history of modem demosponges started within these cryptic facies.
sesquiterpenes, which are characteristic of (and potentially good
biomarkers for) Recent halichondriidsl'axinellids'. Biomarker inves- Late Devonian to Early Carboniferous
tigations were performed on the decalcified residue of samples from
the Porifera-Rothpletzella sequence from the small microbialite The interval between the middle-Late Devonian to the Early
reefs. Compared to other Palaeozoic mud mounds, the material from Carboniferous is one of the most critical times for the development

Fig. 9. Within the small microbialite reefs in the surroundings of Ferque

(Boulonnais, Northern France) Axinella-type cryptic sponges are extremely Fig. 10. Within the small microbialite reefs in the surroundings of Ferque
abundant. Biomarker analysis of these rock have shown sesquiterpene bio- (Boulonnais, Northern France) dendroceratid ceractinomorph sponges are
markers, characteristic for the Halichondrida/'Axinellida' (Reitner et al., common. Diagenetically calcified spongin fibres are preserved (Reitner
2001a) (scale 200 /Lm). et al., 2001a) (scale 200 /Lm).
58 Porifera· Demospongiae· Non-Lithistid Demosponges

of modem sponge taxa, especially the demosponge communities, poecilosclerid spicules from middle Triassic sediments are espe-
due to fundamental changes in the oceanic systems and reorganisa- cially common (Mostler, 1976, 1986, 1990; Reitner, 1992;
tion and recovery of shallow shelf systems and reef environments. Wiedenmayer, 1994). The reorganisation of the modem taxon
This critical interval is called the 'Late Devonian' crisis and Demospongiae had been completed by the Early Triassic.
involved one of the biggest worldwide extinction events during Only one fundamental new development happened, probably
Phanerozoic Earth history (summarized in Walliser, 1996). In par- in the Early Jurassic - the invasion of some marine taxa,
ticular, a fundamental change in the 'lithistid' communities predominantly Haplosclerida, in freshwater environments. The
occurred in the Late Devonian (Famennian). The first tetracladine oldest known remains of freshwater sponges are from the continen-
'lithistid' demosponges were found in small bacterial reef mounds tal Late Jurassic of Portland (southern England). These are spi-
located directly on submarine hydrothermal spring/vent systems. culites of spiny spongillid oxeas, Spongilla purbeckenis (Hinde,
These sponges are part of a true vent-fauna with auloporid corals, 1883). The fossil record of Mesozoic freshwater sponges is
certain small rugose corals, tube worms and stalked crinoids extremely poor and only one further locality from the Lower
(Reitner et ai, 200la). The reef-type Famennian strata are Cretaceous is known, the Chubut-Formation in Argentina, how-
extremely rich in demosponges, although most of them are ever, with excellent well-preserved modem-type freshwater
currently of unknown taxonomic position. Within the Lower sponges (Volkmer Ribeiro & Reitner, 1991b) (Fig. 11).
Carboniferous most of the modem clades of demosponges
existed, and in the Permian all modern major taxa were present. CORALLINE DEMOPONGES WITH PRESERVED
Remarkable is that the first record of the Homosclerophora, a SPICULE SKELETONS
simply-organized group of demosponges, occurred late in the Early
Carboniferous with plakinid spicules and probably evolved from Characterization of Coralline Demosponges
astrophorid tetractinellid ancestors. The first remains of poe-
cilosclerid spicules are known from Permian strata. However, Coralline sponges or 'sclerosponges' are a polyphyletic group
with affinities to both Demospongiae and Calcarea (e.g., Hartman &
Goreau, 1969, 1972; Vacelet, 1985; Wood, 1987; Reitner, 1987c,
1989, 1992; Reitner & Mehl, 1996), which construct a secondary
basal skeleton of Mg calcite or aragonite (Reitner, 1992). Five basic
types of basal skeletons can be distinguished: (1) simple thin crusts;
(2) differentiation oftubes!calicles ('Ceratoporella-type'); (3) tubes/
calicles separated by tabulae ('chaetetid-type'); (4) more-or-less
irregular network of the primary organic skeleton with horizontal
(laminae) and vertical elements (pillars) ('stromatoporoid basal
skeleton'); and (5) chambered basa! skeleton ('sphinctozoid' or
'thalamid basal skeletons'). The first three types of basal skeletons
are related to sponges with thin organic tissue, the latter two are
characterized by a thicker living tissue (Reitner, 1992).
The first Recent specimens of coralline demosponges were
collected at the turn of the last century (DOderlein, 1897;
Kirkpatrick, 1908d, 191Oa,c, 1911, 1912a,b; Lister, 1900; Hickson,
1911), but from that time until relatively recently the knowledge of
these organisms had been nearly forgotten - no doubt due to the
diminished interest in the phylum during the mid-1900s (see gen-
eral Introductory chapter and chapter on Phylum Porifera). During
that time palaeontological investigations were also made on fossil
sponges with basal coralline skeletons, as stromatoporoids and
'Pharetronida', and Calcarea with a rigid dense network of calcitic
spicules (Rauff, 1913; Steinmann, 1882; Welter, 1910). However,
from those 'halcyon days of sponge discovery' until relatively
recently most of the fossil coralline sponges, especially the stro-
matoporoids, were misinterpreted as being hydrozoans, corals,
foraminifera, cyanobacteria etc. (e.g., Lecompte, 1951; Fliigel &
Fliigel-Kahler, 1968; Kazmierczak, 1981).
During the second half of the 1900s up to the present two
groups of biologists have independently recovered Recent coralline
sponges, including many new species, living especially in cryptic
marine reef environments (Vacelet & Levi, 1958; Vacelet, 1964;
Hartman & Goreau, 1969, 1972, 1975; Hartman 1979), and other
Fig. 11. Paiaeospongilla chubutensis from the Lower Cretaceous continen- new species with different types of basal coralline skeletons were
tal Chubut-Formation in Argentina. This specimen is the best known pre- found in the late 1900s (Reitner & Worheide, 1996; Willenz &
served fossil spongillid (Volkmer-Ribeiro & Reitner, 199Ib). The specimen Pomponi, 1996). All these discoveries were important for the inter-
shows lots of gemmulae embedded within the silicified tissue. The gemmu-
lae are surrounded by reticulate tracts of haploscierid oxea (scales: pretation of many fossil sponge remains, which are now classified
A, I mm; B, 300 /-Lm). as 'coralline demosponges' or 'pharetronids'. Hartman & Goreau
Porifera· Demospongiae • Non-Lithistid Demosponges 59

(1972) created a new class to accommodate sponges with a second- coralline algae in the Jurassic, hermatypic corals became more
ary basal skeleton, the Sclerospongiae Hartman & Goreau, dominant as reef-frame builders. Today, coralline sponges are virtu-
1970, but this taxon has clearly been shown to be invalid based on ally restricted to cryptic niches and deeper fore-reef areas, with less
its polyphyletic origin (e.g., Van Soest, 1984a; Vacelet, 1985; or no light and strict oligotrophic conditions.
Reitner, 1991, 1992; Wood, 1987). Sponges with coralline basal After the Cretaceous-Tertiary (KT-) boundary the fossil
skeleton are now assigned to widely distributed clades within record of coralline sponges is extremely poor.
demosponges, and this feature is no longer recognised as a phylo-
genetic autapomorphy; significantly, hexactinellids with a calcare-
ous basal skeleton have never been observed. The various basal Palaeozoic Record of Spicule-Bearing Coralline Demosponges
skeletons are highly convergent, and represent a grade of construc-
tion (sharing functional similarities) rather than any common Hadromerida. Only few taxa of coralline sponges from the
ancestry. Observations of spicule remains in fossil representatives Palaeozoic bear spicules and allow an informed evaluation of their
of coralline sponges have shown that a lot of coral-like fossils are taxonomic affinities. The largest group of these in the Palaeozoic
sponges, and that they were important reef dwellers and reef-form- are the classical Stromatoporoidea Nicholson & Murie, 1878, which
ing organisms in the past up until the end of the Cretaceous (Dieci do not bear spicules. However, due to their characteristic exhalant
et al., 1974, 1977; Kazmierczak, 1979; Gray, 1980; Reitner & canal systems (astrorhizae), most of the main taxa (Stromatopora
Engeser, 1983, 1985, 1987; Wood & Reitner, 1986, 1988; Reitner, Goldfuss, 1826, Actinostromaria, etc.) are now classified as
1987a,b,c, 1989, 1991, 1992; Wood, 1987; Wood et al., 1989). demo sponges. A more detailed classification and determination of
Calcified sponges were the dominant reef-building organisms the phylogenetic position has, up until now, been impossible.
since the beginning of the Phanerozoic. They were replaced Only one group of Palaeozoic stromatoporoids, the Early
by scleractinian corals in modem reefs as primary structural Devonian taxon Syringostroma cf. borealis (Nicholson, 1875), has
organisms, but they have living relatives ('coralline sponges') in a distinctive wall structure of densely packed spherical structures
cryptic niches of almost all Recent coral reefs. They were the first (Fig. 12). These spherical bodies are interpreted as aster microscle-
metazoans to produce a carbonate skeleton and their microstruc- res, comparable to those known from the modem hadromerid taxon
tural features have remained completely unchanged over this very Chondrilla, and found in the Cretaceous chondrillid coralline
long period of time. Their biomineralization processes are sponges Calcichondrilla crustans Reitner, 1991 and Calcistella
extremely conservative and these still exist in extant calcified tabulata Reitner, 1991 (Fig. 13) (Reitner, 1992).
sponges (Reitner et al., 1997, 2001b). If this taxonomic assumption is correct, the diversification of
Coralline demosponges first occur along with the the Hadromerida happened very early in the Phanerozoic. This
Archaeocyatha in the Lower Cambrian (Tommotian). At this time assumption is supported by the spicule record in the 'chaetetid'
archaeocyaths were very diverse in their morphotypes, dominated early- to middle Devonian taxon Pachytheca cf. stellimicans
the Cambrian reefs, and have strong poriferan affinities (Debrenne & (Schliiter, 1885). Within the basal skeleton of this taxon
Zhuravlev, 1993; Debrenne et al., 1990), now ascribed as potential hadromerid-like tylostyle spicules are preserved (Reitner, 1992).
demosponges (refer to chapter on Archaeocyatha by Debrenne et Both major clades of the hadromerid construction were realised in
al., this volume). Although demosponge spicules have been discov- the Middle Devonian: on the one hand the SuberiteslPolymastia-
ered within the basal skeleton of few Archaeocyatha 'Irregulares' like taxon with reduction of aster microscleres (Pachytheca), and
(Reitner, 1992; Reitner & Mehl, 1995), their allochtonous origin is on the other hand the Chondrilla-like taxon with a reduction of
possible. Coralline demosponges with a stromatoporoid basal skele- megascleres (Syringostroma Nicholson, 1875).
ton occur in the Middle Ordovician, except a few enigmatic ones
from archaeocyath-mounds. They playa major role as reef-building
organisms in the Silurian and especially in the Devonian. After the
Frasnian! Fammenian boundary coralline demosponges with
'chaetetid' basal skeletal (Chaetitidae, Tabulata) construction playa
major reef-building role, as well as chambered coralline sponges
('sphinctozoans'), which occur first in the Ordovician (Rigby &
Potter, 1986). The first (problematic) taxa of 'sphinctozoan'
sponges were known from the Lower Cambrian. During the
Permian and Middle Triassic 'sphinctozoans' were one of the most
important groups of reef builders, whereas modem forms of
coralline sponges originated in the Late Permian and had a first
maximum in the Late Triassic (Carnian-Norian). Special forms of
stromatoporoids (e.g., Dehomella) were important reef dwellers
during the Jurassic and Lower Cretaceous. Most of them are cryptic
benthic dwellers and had adapted to living in nutrient-poor deeper
shelf areas. In the Late Albian, modem cryptic coralline sponge
communities of the Tethys realm developed (Reitner, 1989, 1990,
1991, 1992, 1993). Since this time there have been no fundamental
changes observed in the evolution of coralline sponges, with Recent
Fig. 12. Syringostroma cf. borealis from the middle Devonian of northern
taxa morphologically very similar to the Cretaceous faunas . Up Spain with abundant spherical stuctures within the stromatoporoid basal
until the Lower Cretaceous coralline sponges were important reef- skeleton. These structures are interpreted as densely packed aster
building and reef-dwelling organisms, but since the development of microscJeres with hadromerid affinities (scale 200 fLm).
60 Porifera' Demospongiae • Non-Lithistid Demosponges

The 'chaetetid' basal skeleton bauplan is very important in the in silicified specimens (Fig. 14). The opaline silica of sponge
Lower Carboniferous, and some 'chaetetids' bear bundles of spicules is dissolved very fast and spicule remains, mostly calcitic
hadromerid tylostyles but no rnicroscleres. An important represen- pseudomorphs, are extremely rare. Therefore, it appears that the
tative of the Lower Carboniferous hadromerid 'chaetetids' is Hadromerida were well developed in the Palaeozoic.
Boswellia mortoni Gray, 1980. However, most of the Carboniferous Halichondridl'Axinellid' Palaeozoic Coralline Demosponges.
'chaetetids' lack characteristic spicules and this may be due to dia- There is only one record of coralline demosponge with 'axinellid'
genetic factors. The spicules in Boswellia are exclusively preserved affinities from the Late Carboniferous (Pennsylvanian) from
Kansas. This sponge was firstly described by Newell (1935), as
Parallelopora mira. This taxon was revisited by Wood et al. (1989)
and this was the first time that the spicular skeleton was described
in detail. In that paper the authors favoured a classification within
the Haplosclerida, due to the reticulate arrangement of the spicular
skeleton. However, the arrangement of the subtylostyle spicules in
the main pillars exhibit an axial condensation, which is more char-
acteristic of an 'axinellid' bauplan (Reitner, 1992) (Fig. 15).
Comparable skeletal features were seen in the Lower Cretaceous
taxon Euzkadiella erenoensis Reitner, 1987b.
A very intriguing halichondrid coralline demosponge is the
'thalarnid' Subascosymplegma oussifensis (Termier et al., 1977)
from the Late Permian of Djebel Tebaga in Tunisia. This locality is
famous for of its masses of coralline sponges and aragonite pre-
servation of most of the basal skeletons. The basal skeleton of
Subascosymplegma oussifensis is formed by aragonitic spherulites
and the inhalant pores of the 'thalarnid' skeleton are surrounded by
bundles of long styles (Reitner, 1992). The spicular types and
arrangements are comparable with the modem taxa Hymeniacidon,
Scopalina and the coralline demosponge Hispidopetra miniana
Hartman, 1969.
Agelasid Coralline Demosponges.
(1) Ceratoporellidae Hartman & Goreau, 1972 - agelasid
coralline demosponges with a heavy aragonitic basal skeleton
crust. The type genus of the Ceratoporellidae is Ceratoporella
Fig. 13. Calcichondrilla crus tans , a Lower Cretaceous hadromerid with (Hickson, 1911), which is restricted to the Caribbean region. Only
affinities to the extant Chondrilla. The distribution of microscleres are sim- one species of Ceratoporellidae, Stromatospongia micronesica, is
ilar to those seen in the Devonian Sysringostroma (scale 200 fLm). known from Pacific reefs. Ceratoporella is one of the extant

Fig. 14. Boswellia mortoni from the Lower Carboniferous from Wales. These chaetetid sponges are classified as hadromerids with strong affinities to
Suberites. This specimen exhibits a clear regular chaetetid growth pattern. The specimen is partly silicified and within the silicified portions of the chaetetid
the hadromerid-type tylostyles arranged in bundles are well preserved. All vertical elements of these chaetetids are primeriy constructed of tylostyles
spicules (scales: A, 1 cm; B, 150 fLm).
Porifera' Demospongiae • Non-Lithistid Demosponges 61

coralline demosponges ('living fossils') which first occurred in the The main problem of this taxon is that it occurs first in the
Late Permian of the Djebel Tebaga (Reitner, 1992; Wood, 1987) Late Permian, has a good fossil record in Triassic reefs, and then
(Fig. 16). The Ceratoporellidae are very common in the Late disappears at the end of the Triassic (Reitner, 1992) but reappears
Triassic, especially from the Carnian Cassian Beds (Northern next in the Pleistocene. Palaeontologists refer to these taxa as
Italy). From one species, Ceratoporella breviacanthostyla Reitner, 'Lazarus-taxa'. There is a time gap of about 21Omy. Modern repre-
1992, acanthostyle spicules are known (Fig. 17). The spicular sentatives of Ceratoporellidae differ little from Triassic faunas,
skeleton of this taxon is made of verticillitid acanthostyles of the either in the basal skeleton geochemistry or in the spicular skele-
Agelas type. They form a heavy aragonitic basal skeleton on the ton. The reason for this peculiar fossil record is unknown and could
top with small calicles in which most parts of the soft tissue of the be related with varying ocean chemistry, early diagenetic arago-
sponge is located. The soft tissue is also characterized by high nitic dissolution, or may be a facultative basal skeleton formation
amounts of symbiontic bacteria (ca. 60% of the entire biomass) by one taxon dependent on the Ca2 + content of the ambient seawa-
(Willenz & Hartman, 1989). In contrast to the 'sphinctozoan' ter. Permian and Triassic oceans were probably low in Ca2+, simi-
Vaceletia, the aragonite is orientated in clinogonal arranged fibres lar to Recent oceans, whereas Jurassic, Cretaceous and Tertiary
('water jet' structure). Shortly after the removal of the soft tissue oceans were enriched in Ca2+ (Arp et al., 2001). However, the
the calicles are closed by rapid epitactical growth of the aragonitic detailed physiological influences on varying Ca-concentrations are
fibres. The calcification fronts are easy to stain with calcein and there- still unknown and currently under investigation.
fore ideal for in situ growth measurements (Willenz & Hartman, (2) Astrosclera willeyana Lister, 1900 - a stromatoporoid
1985). These sponges are extremely slow growing, exhibiting only coralline sponge with a spherulitic basal skeleton. The main features
200-500 /-Lm yearly growth rates (Bohm et al., 2000). We have inves- of soft tissue and the processes of formation of the basal skeleton in
tigated specimens with an age of more than 600 years. The entire Astrosclera have recently been described in detail, and therefore only
basal skeleton is a thick aragonitic crust with high amounts of Sr a summary is provided here (refer to Worheide, 1998).
(10,000 ppm) and additional extraordinary high amounts of U The living tissue of Astrosclera penetrates the basal skeleton
(7-8 ppm) which allows excellent age determination using the UfTh to a maximum depth of 50%, depending on specimen size. The soft
method. The carbon used for skeletal formation is heavy and in equi-
librium with the ambient seawater (8 13C +5 - + 3,8).

Fig. 16. A, this specimen is the oldest know Ceratoporellidae from the Late
Permian Djebel Tebaga (Tunesia) (scale I cm). B, the aragonitic basal
Fig. 15. Newellia mira, an 'axinellid' coralline demosponge from the Late skeleton is comparable with Recent species, however, unfortunately no
Carboniferous of Kansas (USA) (Wood et al., 1987) (scale 250 J-Lm). spicules are preserved (scale 1 mm).
62 Porifera· Demospongiae • Non-Lithistid Demosponges

Fig. 17. A, this ceratoporellid specimen (Ceratoporella breviacanthostyla) from the Carnian of the southern Alps (Cassian Beds, Northern Italy) is the
only known material with acanthostyle megascleres (B, C) (Reitner, 1992) (scales: A, 2.5 mm; B--{,:, 20 fJ.m) .

tissue shows a stromatoporoid grade of organization (e.g., Wood, aragonite c-axis. When they attain a size of::!: 15fLm spherulites are
1987) and can be divided into three major zones: released from the cell and enveloped by basopinacocytes. In the
(A) The ectosome is the area directly beneath the exopinaco- second process, basopinacocytes transport the spherulites to the tips
denn, reaching a thickness of about 100-300/-lm. This zone is of skeletal pillars where they fuse by epitaxial growth. This epitax-
characterized by the absence of choanocyte chambers, an enrich- ial growth is controlled by acidic mucous substances in the extra-
ment of storage and supporting cells (SSC's), and archaeocyte-like cellular space (ECS) between the growing aragonitic fibers and the
large vesicle cells (LYC's), which are responsible for the initial basopinacoderm. The mucus is produced by basopinacocytes and
formation of aragonitic spherulites. (B) The choanosome, contigu- acts as a buffer for Ca2+ ions. The ECS-mucus is thought to have a
ous with the ectosome but with a more-or-less sharp transition, different composition from the mucus inside vesicles of the LYC's.
comprises the major part of the living tissue in Astrosclera, charac- The third process involves withdrawal of soft tissue during upward
terized by small choanocyte chambers (::!: 15 fLm diameter) and a growth, when it is pulled upwards from the lowermost parts of
high density of bacterial symbionts. Other cellular components of skeletal cavities. The space remaining is subsequently filled by epi-
the choanosomal mesohyle are archaeocytes, typical pluripotent, taxial growth of aragonite fibers. The zone of epitaxial backfill is
phagocytic demosponge cells, SSC's, and rare fiber cells. (C) The characterized by an absence of choanocyte chambers and a reduced
zone of epitaxial backfill (ZEB) is considered as a subzone of the number of bacteria, but sometimes by an increased number of
choanosome due to its important role for the syn vivo cementation SSC's. The ECS in the ZEB is also filled with acidic mucus, which
of the lowest parts of the basal skeleton. It is characterized by a controls the speed and direction of epitaxial growth. The number of
reduced number of (or absence of) choanocytes and bacteria and SSC's is sometimes increased, indicating their importance in nutri-
sometimes an enrichment of SSC's. ent supply during skeletal elaboration.
The aragonitic calcareous skeleton of Astrosclera is formed by Astrosclera is a very slow growing species, the mean growth
the combination of three processes. In the first, spherulites are rate being 230 fLm per year, with an average growth rate of 0.63 fLm
formed in LYC's in the ectosome. The LYC's posses a large vesicle per day, as determined by in vivo staining with Calcein-Na2,
which is filled with a three dimensional network of sheets and AMS, 14C and UlTh data (Worheide, 1998). The oldest known
fibers and acidic mucus. Sheets and fibers act as the insoluble modem specimen has an individual age of more then 500 years
organic matrix (10M) and mucus as soluble organic matrix (SOM) (Worheide, 1998).
for seed crystal nucleation. In the first stage, seed crystals are Taxonomic Value of the Spherulitic Basal Skeleton. The
randomly oriented, later they are oriented in the direction of the spherulitic basal skeleton-type was much more widespread at the
Porifera· Demospongiae • Non-Lithistid Demosponges 63

end of the Palaeozoic (upper Pennian, Djebel Tebaga, Tunisia), and 1992). Reitner (1992) also stated that different skeletal morpho-
in the Late Triassic (Carnian, St. Cassian, Dolomites, Italy; Norian, types appeared independently in various phylogenetic lineages,
Antalya, Lycien Taurus, Turkey), than in Recent seas. Generally, and suggested that intracellular formation of the aragonitic
this body plan can be elaborated by two distinct processes. The first spherulites in general is not an apomorphy of Astrosclera. This
is extracellular, where an organic skeleton (e.g., organic spongin) is grade of construction may be interpreted as a polyphyletic develop-
calcified extracellularily, producing a 'pseudo-spherulitic' skele- ment of several distinct sponge taxa, with no synchronous develop-
ton. This mode is seen in extant Calcifibrospongia actinostromari- ment (i.e., a convergent character), or a plesiomorphic character,
oides (Haplosclerida) from the Bahamas, and in the Aptian evolved in a common ancestor during the Precambrian! Cambrian.
axinellid Euzkadiella erenoensis from northern Spain (see Reitner, Therefore, an 'intracellularily formed spherulitic skeleton' repre-
1987a, 1992). The second, an intracellular mode of spherulite for- sents a character with no taxonomic value in the fossil record.
mation is where no primary skeleton is calcified. This mode was Worheide (1998) screened and studied thin sections of sam-
described in detail by Worheide (1998), and is summarized above, ples from St. Cassian and from Antalya (the latter by courtesy of
outlining early stages of skeletal formation in extant Astrosclera I.-P. Cuif) , to detennine the affinities of calcified sponges with
willeyana. Whether formed extra- or intracellularily, the spherulitic 'astrosclerid-like' spherulitic microstructures, and to find relatives
type of skeleton appears to be predominant amongst sponges from of the extant Astrosclera (if possible, spicule-bearing ones). A large
the Triassic deposits of St. Cassian and Antalya (Cuif & Gautret, variety of spherulitic microstructures was observed in thin sections
1991a), whereas the intracellular mode of formation is now only of this collection, belonging to a large variety of skeletal architec-
found in one extant taxon, A. willeyana. tures and taxa (e.g., Worheide 1998: pIs. 29/3, 29/4: the chaetetid
The first occurrence of sponges with a spherulitic grade of con- Cassianochaetetes cf. gnemidius (Klipstein); PIs. 29/5-7: an
struction was reported from the upper Pennian of Tunisia (e.g., unidentified sphinctozoan). Most of the different spherulitic skele-
Tennier et al., 1977). One of the reported taxa, Subascosymplegma tal morphologies of sponges from St. Cassian and Antalya were
oussifensis, was described by Reitner (1992: 210), as possessing a formed by an intracellular process in the early stage, and later by
basal skeleton made of intracellularily-formed aragonitic spherulites. epitaxial growth as in the Recent Astrosclera, and therefore this
Free spherulites, as found in the extant Astrosclera but with a larger mode was named 'astrosclerid-like' (e.g., Cuif & Gautret 1991b). It
diameter of 400--600 /-Lm, were also found in the biogene pore space has been demonstrated in detail by Worheide (1998) that the skele-
of that species. Subascosymplegma oussifensis had a spicular skeleton of Astrosclera is formed by three distinct processes. Only in the
ton of irregularly arranged long thin styles, a plumose ectosomal very early, initial stages are spherulites elaborated by an intracellu-
skeleton also composed of long thin styles, and Reitner (1992) sug- lar process, but the greater part of skeletal accretion happens by
gested affinities to the taxon 'Axinellida'/Halichondrida. Although two different extracellular epitaxial growth processes. In some of
the deposits of Djebel Tebaga contain a large variety of spherulitic- the Triassic thin sections screened, there were signs of (A) initial
type basal skeletons, only some of them appear to be formed intra- intracellular growth of the spherulite (non-mineralized dark centre
cellularily. Intracellular formation of spherulites is only recognizable of spherulite), (B) epitaxial fusion of spherulites (elongated
in spherulites with a dark centre, as found in extant Astrosclera. spherulites), and (C) epitaxial backfill. These specimens certainly
Calcified coralline sponges with a spherulitic skeleton are belong to a different taxon than Astrosclera (Worheide, 1998: PIs
present in the exceptionally well preserved Late Triassic deposits 29/3-7). Even if all these processes were found in one specimen, a
of Antalya and St. Cassian, previously described by Cuif (1974) definite decision could not be made about the taxonomic affinities
and Cuif (1983), Engeser & Taylor (1989), Reitner (1992), and given that a combination of the three processes appears to have no
others. These sponges have well-preserved original aragonitic min- taxonomic value. As noted, this grade of construction has probably
eralogy and microstructure, and were used to compare the taxo- appeared independently in different phylogenetic lineages of
nomical value of spherulitic skeletons based on comparisons with coralline sponges, representing a convergent character, whereas
Recent Astrosclera (Gautret, 1986; Cuif & Gautret, 1991b; Wood, taxonomic affinities can only be deduced if spicules are present.
1991b; Reitner, 1992) (Table 1). These authors showed that the Hypercalcified spherulitic basal skeletons are clearly a grade of
spherulitic grade of construction appeared independently in a large construction, as suggested by Wood (1991b), and no apomorphic
variety of skeletal grades of architecture in Triassic sponges character of particular taxa, as suggested by Hartman & Goreau
(i.e., sphinctozoan, chaetetid, stromatoporoid; Worheide, 1998: (1970) in general, and by Cuif & Gautret (1991b) for Astrosclera.
Plate 29), and in different taxa (e.g., Sestostromella robusta, Clearly, morphological characters are inadequate clues to the
Haplosclerida; Chaetosclera clipsteini, Halichondrida; Reitner affinities of extant Astrosclera with Triassic species. Because the

Table 1. Examples of selected coralline sponges with a basal skeleton made of intracellularily formed spherulites. This type of skeleton occurs in distinct
sponge taxa through time and represents a character with no taxonomic value (from Worheide, 1998).

Taxon Age Diameter of Spicular skeleton Affinities to

intracellular
spherulites
----_._---_.,,-- ..
Subascosymplegma oussifensis Upper Permian 400-600 fllIl Long, thin, styles, plumose Axinellidal Halichondrida
Sestrostomella robusta Lower Carnian 20-50 fllIl Long, thin, curved diactines Haplosclerida
Chaetosclera klipsteini Lower Carnian 10-20 fllIl Long styles Halichondrida
Astrosclera willeyana Recent 15-60 flm Thick, short, verticillate acanthostyles Agelasida
(basic form)
64 Porifera· Demospongiae • Non-Lithistid Demosponges

Fig. 18. A, holotype of Astroclera cuifi Worheide 1998 from the Norian of Antalya (Turkey); horizontal section of specimen in thin sec-
tion (scale 1 mm). B, one mamelon of A. cuifi (arrow) (scale 200 fLm). C, detailed view of A. cuifi showing spherulitic basal skeleton,
made out of intracellularily formed spherulities. Note dark centers of spherulites and free spherulites in biogene pore space, similar to
extent Astrosclera (scale 150 fLm). D, sub-acanthostyle megascaleres (arrows) in the basal skeleton of A. cUifi (scale 50 fLm). All figures
from Worheide (1998, Plates 29, 30).

Triassic material lacks spicules (Reitner 1992: 230), sponges, prob- similarities to one of the 'groups with similar spicule morphology'
ably Astrosclera, have to be identified initially from a combination (GSSMs) of extant Astrosclera. Worheide (1998) distinguished six
of morphological criteria based on biocalcification. A definitive different populations in Astrosclera. One of these populations, from
classification of Triassic 'relatives' is only possible based on the western Indian Ocean, showed a medium intra-population varia-
spicule morphologies, but in most cases spicules are not present in tion in spicule morphology and normally contains acanthostyle
the fossil record, leaving only the morphological criteria of megascleres, but also a few thick spicules without spines, or with
the basal skeleton (i.e., central Pacific populations of extant reduced spines, which are comparable to the ones observed from the
Astrosclera also lack spicules). Antalya species. Affinities of the Triassic sponge A. cuifi to
Thus, all of the microstructural features present in Astrosclera Astrosclera were therefore obvious. The Triassic species was
must be present in fossil taxa to infer affinities, not only the char- clearly new due to the slightly larger spherulite size, the rarer
acter 'intracellularily formed spherulitic' basal skeleton, since the spicules, and the different spicule morphology (Fig. 18D).
intracellular formation of spherulites appeared in the Triassic However, although the first occurrence of Astrosclera has
in different taxa. But even so, confirmation of phylogenetic affinity been confirmed in the Late Triassic of southern Turkey
with Astrosclera is at most only probable, and still uncertain if no (Astrosclera cuifi Worheide 1998), an important problem still
spicules are present. remains: the lack of a record during younger times until the
However, applying stringent criteria, Worheide (1998) was Holocene (thus, a 'L azarus' -taxon, see above).
able to discover and describe the first true relative of the taxon Vaceletia Pickett, 1982: A 'Thalamid' Coralline Demosponge
Astrosclera. Only one sponge from Antalya (Turkey) complied with with Archaeocyathan Affinities. The fossil record of this taxon
all the criteria, of all the sponges screened from both St Cassian and is, in contrast to other aragonitic coralline sponges, very good and
Antalya. Therefore, affinities to the taxon Astrosclera were clear, continuous from its first reported occurrence in the Middle Triassic
and moreover, spicules were also found in this specimen (Fig. 18). to Recent seas. They are 'thalamid' or 'sphinctozoid' sponges that
Astrosclera cuifi Worheide (1998) showed signs of all three of the exhibit a soft tissue organisation and structure already known from
distinct biocalcification processes of A. willeyana, although the size sponges with a stromatoporoid skeleton. Normally they possess a
of spherulites was larger than in extant Astrosclera. These biocalci- central cavity (spongocoel) and a chambered basal skeleton.
fication characters indicate initial and subsequent biocalcification The primary organic skeleton of Vaceletia is non-spicular
took place similar in a manner similar to the processes in extant (Vacelet, 1979). The choanocyte chambers are relatively large
A. willeyana (see above and Worheide, 1998 for details). The spicu- (50 fL m) and the mesohyl is enriched with bacteria (ca. 50% of the
lar skeleton, although not 'typical' acanthostyles, generally showed entire biomass). Large rounded cells with large inclusions are very
Porifera· Demospongiae • Non-Lithistid Demosponges 65

B
-
Fig. 19. Cenomanian Vaceletia from Liencres, northern Spain. These coralline sphinctozoan sponges are very common in the middle Cretaceous (Reitner,
1989. Reitner et aI., 1995) (scales: A, 5 mm; B, 2mm).

characteristic of this sponge, comparable with the LCG cells of the central part and the border of the pillars, in between the calcifi-
Acanthochaetetes Fischer, 1970 (Reitner & Gautret, 1996). It has a cation is slower.
trabecular organisation and is overlaid by a hemi-spherical top- (2) Calcification of the inactive parts of the skeleton. In the
layer (,dermal-layer'). The trabecules consist of irregular, organic ontogenetic development of the older parts of the skeleton a second
filaments with a very thick central filament. This centi'al filament calcification phenomenon is observed. The upwardly moving soft-
has a supporting function and could be seen as an 'organic spicule'. tissue is able to form organic phragmas via the basopinacoderm.
A network of very thin fibers surrounds this central filament. The These phragmas separate chambers which are filled with acidic
calcification of the secondary aragonitic skeleton starts between glycoproteic mucus ('soluble matrix', SOM). This SOM shows
these organic fibers. This secondary skeleton consists of irregular high concentrations of Asp (14.13 mol%) and Glu (11.42mol%) in
aragonitic micrite. The central filament would not be calcified. The combination with high values of sugars ARA (14mol%), XYL
formation of the secondary skeleton is not a continuous process, it (16.5mol%), and GLC (22mol%) (Reitner, 1992). Within these
happens step by step in the following order (cf. Reitner, 1992; chambers the toxic physiological surplus of Ca2+ is bonded to
Reitner et al., 1997, 2001b). acidic macromolecules and deposited as an aragonitic waste prod-
(1) Formation of skeletal-pillars. Formation of a new, non- uct. The SOM is interspersed by polymerised mucus fibres which
calcified chamber with a hemispherical dermal top-layer and a tra- act as the insoluble organic matrix (10M). The steps of mineraliza-
becular organisation and organic skeletal-pillars containing a thick tion are the same as seen in the pillars (Reitner, 1992; Reitner et al.,
central filament. These skeletal-pillars are filled with thin fibres. 2001b). This calcification phenomenon is often observed in many
The entire space inside the pillar is filled with acidic glycoproteicl fossil sphinctozoans of the Vaceletia-type, and could be a model
proteoglycanic mucous. This space is filled successively during for all irregular, micritic-granular basal skletons of stromatoporoid
ontogenesis by aragonite crystals. The mineralization starts and 'thaI amid' organisation. Most important is the mode of bio-
from the inside of the organic pillars. Further on the whole fibrous mineralization because the Archeocyatha of the Lower Cambrian
insoluble matrix of a newly formed pillar is substituted succes- exhibit the same type. It is suggested that Vaceletia may be
sively by aragonite crystals. The acidic mucous substances extremely ultraconservative and therefore comparable with certain
are reduced correspondingly. The thick central filament cannot be taxa of the Archaeocyatha (Reitner, 1992).
mineralized. This central filament has only a primary initial sup- Highlights of the Mesozoic Coralline Demosponges. The ear-
porting function, because the irregular fibres inside the pillars are liest fossil taxon is Stylothalamia dehmi Ott, 1977, recorded from the
not able to support the choanosome on their own. Newly formed Ladinian (Middle Triassic) (Ott, 1967) (Fig. 19). This type of 'sphinc-
chambers never show the complete structure and size of the later tozoan' is restricted to small caves and dark reef environments.
calcified ones. Such a chamber in statu nascendi is increasing The Stylothalarnidae were extant in the Late Triassic and apparently
slowly in size. The crystallization seems to start from the borders survived until the lower Jurassic (Domerian). The Stylothalmidae are
of the uncalcified skeletal elements. An initial, prismatic layer of characterized by vesicular skeletal structures, and sometimes oxeas
aragonite crystals is observed. Larger crystals overlay this layer are found incorporated into the micritic, aragonitic basal skeleton.
forming a loose network. The density of calcification is higher in However, most vaceletids do not bear any spicules. The spicule types
66 Porifera· Demospongiae· Non-Lithistid Demosponges

Fig. 20. Most of the Mesozoic coralline sponges are related to the Halichondrida! 'Axinellida', with axial spicule condensation. One taxon of the
Milleporellidae, the genus Murania, has a basal skeleton of aragonite in the Triassic and calcitic ones in the Lower Cretaceous. Murania kazmierczaki from
the Carnian Cassian beds has an aragonitic basal skeleton with excellent preserved subtylostyle spicules (Reitner, 1992) (scales: A, vertical section, Imm;
B, horizontal section, lmm; C, SEM micrograph - aragonite dissolved with EDTA (method described in Reitner & Engeser 1987),100 fLm).

and arrangement resembles the Haplosclerida, with some affinities

to the Callyspongiidae. In the Cretaceous Vertic illites Defrance,
1829 is very common and forms large specimens of up to 10 cm in
size. Vaceletia is known from various Tertiary localities (e.g.,
Vaceletia progenitor Pickett, 1982), and the deeper water form from
the Danian of Faxe (Denmark), Vaceletiafaxensis.
Recent Vaceletia are mostly non-branched solitary taxa and are
restricted to dark reef caves. However, common frame building
colonial forms were recently discovered in shallow water reef caves
at Osprey Reef (Coral Sea) and at the Astrolabe Reef, Fiji
(Worheide & Reitner, 1996). According to a recent genetic analysis
(Worheide et aI., unpublished) both forms are clearly different
species. There is also one deep-water (200-400 m) species from New
Caledonia realm (Vacelet, 1988, Vacelet et al., 1992). The genetic rela-
tionships of all these forms are currently under investigation.
Based on the ceractinomorph parenchymella larva and simple
monaxonic spicules in some fossil taxa, with a callyspongid
Fig.21. A, Murania merbeleri from the Barremian Schrattenkalk (Bavaria,
arrangement the fossil and Recent vaceletids may be classified
Gemany) exhibits a calcitic basal skeleton (scale lmm). B, This sponge
close to the Haplosclerida. shows the same spicule types and arrangements as M. kazmierczaki (scale
100 fLm).
Mesozoic Coralline Demosponges with 'Axinellid' Spicule
Arrangement the species Murania kazmierczaki Reitner, 1992. The basal skele-
ton is constructed of a fibrous aragonite and the spicules are subty-
In the Mesozoic, coralline demosponges with an 'axinellid' lostyles (Fig. 20). A good example of a Cretaceous representative
skeletal structure are major components of the benthos (Wood, is Murania merbeleri Scholz, 1984 from the Barremian
1987; Reitner, 1992). Most of the Jurassic stromatoporoids Schrattenkalk reef facies (Fig. 21). The Milleporellida became
(Milleporellidae), like Dehornella Lecompte, 1952, exhibit the extinct at the Cretaceous-Tertiary boundary.
typical axial concentration of monaxonic spicules (oxeas, styles). Acanthochaetetes Fischer, 1970, a Lower Cretaceous
They are important reef and shallow water dwellers, and exhibit in Spirastrellid Coralline Demosponge. The modem representative
the Jurassic and Cretaceous a low Mg calcite basal skeleton. In the of this taxon is the chaetetid Acanthochaetetes wellsi Hartman &
Triassic all of the coralline sponge with 'axinellid' affinities bear Goreau, 1975, which has been associated with the Recent genus
an aragonitic basal skeleton. Surprisingly, one taxon is able to do Spirastrella by Reitner (1991) but requires further molecular cor-
both, aragonite basal skeletons in the Triassic and calcitic ones in roboration. The basal skeleton is made of high-Mg Calcite (15-19
the Cretaceous. This taxon is a crust-forming coralline sponge mole% MgC0 3) with a generally tangential lamellar growth pat-
called Murania Kazmierczak, 1974. The spicule-trees (axially con- terns (Reitner & Gautret, 1996). Only the 0.5-1 mm thick youngest
densed spicules) become calcified probably during the upward generation of the calicles of the 'chaetetid' -type basal skeleton is
movement of the soft tissue. The taxon Murania occurs first in the occupied by the living soft tissue. Acanthochaetetes wellsi has an
Late Triassic (Carnian) of the Cassian Beds (northern Italy), with extremely slow growth rate of the basal skeleton of 50 ,.un/yr, The
Porifera· Demospongiae • Non-Lithistid Demosponges 67

Fig. 22. Acanthochaetetes is an ultra-conservative taxon, which occurs first in the Lower Cretaceous of northern Spain, A, taxa from the Albian (Lower
Cretaceous) incorporate parts of their spicular skeleton in the high Mg-calcite basal skeleton (scale 100/-Lm). B, the spicular skeleton is composed of
hadromerid tylostyles and spiraster microscieres (scale 20 /-Lm). C, Cretaceous specimen of Acanthochaetetes (scale 1 cm) (from Reitner & Engeser, 1997;
Reitner, \992).

soft tissue and basal skeleton exhibit a vertical anatomy divided in highly organized microstructure, both characteristic of the
six major zones. The formation of the basal skeleton can be sum- Acanthochaetetes skeleton, were never observed in these places.
marized as below (Reitner & Gautret, 1996). Skeletal formation starts inside the uppermost fibre template in the
(1) At the uppermost position there is a thick crust-like layer form of a soft structure in which elements have the shape and size
of spiraster microscleres (dermal area, zone I), and tylostyle of the future characteristic Acanthochaetetes crystals, but these are
megascleres which are arranged in clearly plumose bundles, not rigid, and they look like 'cooked spaghetti' (Reitner & Gautret,
demonstrating its alleged close relationship to the bauplan of 1996). This random structure becomes calcified and organized
Recent Spirastrella. when a mucus is secreted in the narrow space between the
(2) Below the outer dermal area, the internal dermal area (zone basopinacoderm and the calcified skeleton, by the pinacocytes
II) is formed by mesohyle tissue, devoid of choanocyte chambers, which are forming the most basal continuous cell layer. This mucus
and enriched in mobile cells. Large inhalant chambers (lacunae) and is highly soluble, making direct observations difficult to perform
distributing canals cross this zone, serving the choanosome with with electron microscopy. It is not preserved in TEM preparations
water filtered through the ostiae. The choanosome is characterized and at best, it can be recognized with the SEM through the col-
by very large choanocyte chambers (80-100 jl.m). The mesohyle is lapsed clumps which are closely related spatially to growing crys-
characterized by large cells (ca. IOjl.m) containing numerous inclu- tals in very well fixed specimens.
sions (LCG: large cells with granules), and lying directly on the cal- (3-5) The central part of the tubes (zone III) is characterized
careous skeleton (Reitner, 1992; Reitner & Gautret, 1996). Mesohyl by the choanosome which exhibits large choanocyte chambers (80-
bacteria are rare (ca. 5% of the mesohyl biomass) and they are very 100 jl.m) leading to large oscular channels. Few tylostyles are nor-
small (500nm). These highly mobile cells are undoubtedly responsi- mally present. Typical for a 'chaetetid' skeletal type is the
ble for the secretion of collagen fibres and they probably derive from occurrence of tabulae stopping the tubes (zone IV). These are
a special type of lophocyte. Collagen fibres form strong bundles formed by the basopinacoderm, first as a thin organic phragma or
which traverse the basal pinacocyte layer, and anchor into the rigid sheet. Below the choanosomal zone, LCG cells become enriched and
skeleton (Vacelet & Garrone, 1985). Thin collagenous fibres pro- produce the mineralization of the organic sheet. Continuous
duced by non-modified lophocytes are widely dispersed within the upwardly moving basopinacoderm form a space filled with Ca2+-
intercellular mesohyl. They condense and become organized into a binding and mineralizing organic mucus. This mineralization
frame-building matrix at the top of the walls and they stay process happens only when LCG cells are present (Reitner, 1992).
entrapped inside skeletal structures after calcification. Calcite Within the closed spaces between tabulae, they contain accumula-
formation occurs as soon as these two types of fibres are present, tions of modified archaeocytes with numerous storage granules
supporting the interpretation that they have an ability to attract (thesocyte-like cells) and few spiraster microscleres (zone V).
bivalent cations. However, the acicular shape of crystals and the These cells may play a role in regeneration processes (Vacelet,
68 Porifera· Demospongiae· Non-Lithistid Demosponges

1985, 1990; Reitner, 1992), making the sponge able to start grow- 'axinellid' sponges also radiated. In the Middle Devonian the first
ing again when it has been drastically damaged. modem forms of Dendroceratida, 'axinellids' (mostly halichon-
The 'chaetetid' basal skeleton has a very specific function as drids) , and the first haplosclerids occurred. Most of the Lower
a protective and resting body for special omnipotent cell types Palaeozoic stromatoporoids are probably demo sponges although
(thesocytes, archaeocytes), which are enclosed between two tabu- this also requires further corroboratory evidence given that in all
lae in a calicle (internal 'gemmulae'). This strategy allows the except one form spicules are missing. An important boundary for
sponge to survive environmental crises. Due to this survival strat- the demosponges is the Late Devonian extinction event which
egy the sponges may have many buds, representing a starting point caused a complete renovation of demo sponge communities. In the
for new growth on the top of one calicle. Early Carboniferous most of the main modem demo sponge taxa
The oldest known Recent specimen from a deep submarine were present. The first Poecilosclerida are known from Permian
cave of Cebu (Philippines) has individual age of more than 600 years. sediments. Notably, most of the stromatoporoids became extinct
The taxon Acanthochaetetes is very conservative and the first during the Late Devonian extinction event and were replaced by
fossil record is known from the Lower Cretaceous (Fischer, 1970; the 'chaetetid' body plan. Most Carboniferous 'chaetetids' are rep-
Reitner, 1982, 1990; Reitner & Engeser, 1983). In contrast to the resentatives of the Hadromerida. In the Permian, coralline demo-
most aragonitic species of coralline sponges, this taxon has a con- sponges became very important and this development continued
tinuous record since the Lower Cretaceous. During the Cretaceous into the Triassic. Most of the spicule-bearing coralline sponges are
three ecological niches were occupied, a shallow marine open 'axinellids', and most of these can be associated with halichondrid
water environment (Acanthochaetetes ramulosus), deep fore reef taxa, with only one appearing to be more closely related to the
environment (cf. A. seunesi), and cryptic niches (A. seunesi). In modem Poecilosclerida and some more closely related to modem
contrast to modem Acanthochaetetes the Cretaceous species incor- Haplosclerida. The Late Permian and the Triassic, especially the
porate parts of their spicular skeleton within the basal skeleton Late Triassic, are the main eras for coralline demosponge radiation
(Reitner & Engeser, 1986) (Fig. 22). and dominance, in which some modem taxa occur first
(Ceratoporella, Astrosclera, Vaceletia). At the Triassic-Jurassic
boundary another fundamental change occurred to demosponge
CONCLUSIONS: MAJOR TRENDS IN THE communities. Remarkable is the disappearance of the ceratoporel-
EVOLUTION OF NON-LITHISTID DEMOSPONGES lids and astrosclerids, which did not reappear again until the
Pleistocene, 200 my later. In the Late Jurassic the freshwater envi-
The main problem in interpreting trends in evolution of non- ronments were occupied by particular marine demosponges,
lithistid demosponges is that their fossil record is generally poor and mostly Haplosclerida. This was the last innovation in demosponge
discontiguous for most taxa, and most palaeontologists are not able phylogeny until Recent seas. The importance of coralline
to recognize sponge spicules within sediments. Therefore, our demosponges as primary reef-builders decreased up to the Late
knowledge of non-lithistid demosponges, and also of the coralline Cretaceous. Today, nearly all coralline sponges are restricted to
sponges, is poor, because only few bear spicules that allow a more- cryptic or deep marine environments and are considered to be
or-less unequivocal classification within the modem taxonomic relicts or 'living fossil' taxa. Their further study certainly holds
framework. However, some trends are recognisable. Available evi- clues to fossil sea environmental conditions and evolutionary
dence suggests that the first demosponges occur in the Late trends within Porifera.
Proterozoic, with forms characterized by bundles oflong monaxonic
spicules. In the Early Cambrian spherical forms with long styles
occur (Choia) and represent probably the stem line of the ACKNOWLEDGEMENTS
Phanerozoic demosponges. It is possible, or even likely, that some
Archaeocyatha are demosponges, based on the occurrence of occa- JR acknowledges financial support from the German Research
sional tetractinellid spicules incorporated in the basal skeleton, but Council (DFG) Re 665/8, 12 (Leibniz Award), 14, 16. GW
this interpretation requires further corroboration. In the Early acknowledges grants from the Australian Biological Resources
Cambrian the tetractinellids diversified with the taxon Astrophorida Study and Postdoctoral fellowships from the German Academic
(Geodiidae). At this time the first 'keratose', hadromerid and Exchange Service (DAAD) and the Queensland Museum.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

t 'Class Stromatoporoidea' Nicholson & Murie, 1878: Stromatoporoids

Alex Cook

Queensland Museum, P.O. Box 3300, South Brisbane, Qld, 4101, Australia. (AlexC@qm.qld.gov.au)

Stromatoporoidea Nicholson & Murie (? Porifera, ? Demospongiae) are calcareous skeletal remains of organisms present during the
Palaeozoic to Mesozoic, comprising sponge-like taxa with hypercalcified layered skeletons that were important to reef-building. Earlier
(Palaeozoic) forms lack spicules but some of the later (Mesozoic) taxa have siliceous monaxonomic or tetraxonic spicules in addition to
calcitic basal skeletons, inferring affinities to Demospongiae. 'Stromatoporoids' has been used at various levels (ordinal, class and sub-
phylum) within the phylum Porifera, but also within other 'lower' marine invertebrate phyla at various times. The contemporary hypoth-
esis is that these life forms may represent a grade (and not a clade) of poriferan construction, although it is also recognised that there may
be fundamental differences between the Palaeozoic and Mesozoic stromatoporoid faunas, such that this grade of skeletal construction
may not be homologous, and the taxon may be a completely artificial construct. Some stromatoporoids have also been associated with
the Calcarea and also the 'sphinctozoan' grade of construction. Only a brief overview is provided although stromatoporoids include about
110 genera in seven orders.
Keywords: ? Porifera; ? Demospongiae; 'Stromatoporoidea'; stromatoporoidomorphs.

HISTORY AND BIOLOGY skeleton, which open at the surface in the vicinity or summit of
astrorhizae. These astrorhizae (stellate grooves or canals on the
Concept of 'Stromatoporoidea' surface), are among the most prominent features of stromato-
poroids. The astrorhizid pattern is also visible in the interior of the
The term 'stromatoporoid' represents different taxonomic skeleton (coenosteum), spread out on distinct growth interruptions
concepts to many workers, just as views on their phylogenetic rela- (latilaminae). Horizontal tabulae separate major parts of the skele-
tionships are varied. Stromatoporoidea Nicholson & Murie, 1878- ton, with living tissue probably only covering the skeleton as a rel-
whatever that means - have been historically classified with inter atively thin veneer. Astrorhizae are also prominent features of the
alia the cnidarians, hydrozoans, foraminiferans, cyanobacteria, and Recent 'sclerosponges', such as Astrosclera (refer to chapter on
most recently the PoIifera. A review of the historical systematic Family Astrosclerida (Vacelet, this volume», with suggested affini-
assessment of the Palaeozoic stromatoporoids was provided by ties partly based on this inferred homology. The affinities between
Stearn (1982). Mesozoic stromatoporoids were reviewed by Wood Recent 'sclerosponges' having stromatoporoid skeletal structure
(1987). To paraphrase Reitner & Keupp (1991) "these represent and the Palaeozoic and Mesozoic stromatoporoids, however, are
a characteristic skeletal type". equivocal. Whereas most contemporary authors accept that these
Recent 'sclerosponges' represent surviving remnants of a once-
Morphological characteristics diverse hypercalcified sponge fauna, with affinities to Demo-
spongiae (e.g., Hartman & Goreau, 1970; Wood, 1990), the fossil
Stromatoporoids were once widespread in their distribution stromatoporoid fauna is likely highly polyphyletic whereby similar
and abundance. The fossil record suggests they rose to prominence structures occurring in the two groups are not all necessarily
during the Middle Ordovician (see Figure 22 in the Fossil Lithistida homologous.
chapter (pisera & Uvi, this volume) for geological time scales).
They peaked in diversity during the Upper Silurian and Middle Palaeozoic 'Stromatoporoidea'
Devonian when they were most successful reef builders. Their
apparent diversity waned during the Upper Carboniferous to Middle Despite a previous history of significant debate (e.g., Dong
Jurassic, and then again with an increase in diversity during the De-Yuan, 1988), it is currently generally agreed by most students
Upper Jurassic where they were important reef builders. They subse- of the group that Palaeozoic stromatoporoids represent a poriferan
quently declined during the Cretaceous-Tertiary and became extinct grade organism, although there are alternative views to this place-
during the Kff boundary (refer to detailed history in Lecompte, ment (Kazmierczak, 1976, 1981). There is recognition that the
1957). It has been hypothesised that some modem day demo- Palaeozoic stromatoporoids have not revealed any spicules. What
sponges, such as Astrosclera, Ceratoporella and Stromatospongia, is not agreed is: whether they are polyphyletic; their original skele-
represent living stromatoporoids, or alternatively, these few Recent tal mineralogy; their relationship to Mesozoic stromatoporoids
taxa retain a stromatoporoid body plan (see chapter on Non-Lithistid (stromatoporoidomorphs); and their putative placement within the
fossil Demospongiae (Reitner & Worheide, this volume». demosponge clade.
Stromatoporoid growth forms range from massive to laminar, The most recent review of the Palaeozoic stromatoporoids
with sizes also ranging from several centimetres to over 1 m in was that of Stearn et al. (1999), who regarded them as a Poriferan
diameter. The skeleton is a marked layered structure with vertical Class ranging from Ordovician to latest Devonian in the fossil
pillars and horizontal laminae, together forming a more-or-Iess record. These authors proposed the following classification:
irregular meshwork. The aquiferous system is composed of vertical Class Stromatoporoidea Nicholson & Murie, 1878. Order
and transverse canals, associated with the open framework of the Labechiida KUhn, 1939; Order Clathrodictyida Bogoyavlenskaya,

69
70 Porifera· Demospongiae • 'Stromatoporoidea'

1969; Order Actinostromatida Bogoyavlenskaya, 1969; Order Much of the evidence for original skeletal chemistry is cir-
Stromatoporellida Lecompte, 1951; Order Stromatoporida Stearn, cumstantial, at best. Stearn (1972, 1975), and Stearn & Mah (1987)
1980; Order Syringostromatida Bogoyavlenskaya, 1969; Order assessed that the original skeletal chemistry was aragonite,
Amphiporida Rukhin, 1938. whereas Galloway (1957), Kershaw (1990) and Rush & Chafetz
Within these higher taxa Stearn et al. (1999) placed 109 gen- (1991) concluded the skeleton was high-Mg calcite. The implica-
era and added 14 of dubious placement. Expounding this systemat- tions of these opinions for their phylogenetic affinities are critical,
ics further here is inappropriate within the largely Recent with Mesozoic stromatoporoids displaying uncontested aragonitic
biological context of the Systema Porifera, although a more skeletal structures, either primarily or secondarily (Wood, 1987).
detailed systematics of stromatoporoids will soon appear in the Intemalistically the diagenetic state of the skeleton also effects the
forthcoming revision of the Treatise on Invertebrate Palaeontology classification as a function of the interpretation of critical
(Rigby et al., Editors). microstructures. In an extreme example of this Sleumer (1969)
interpreted many fossil remains into a few taxa, and rejected many
Polyphyletic classification of the specific, generic, familial taxa previously established.

The Stromatoporoidea, be it subphylum (Stearn, 1993), or Relationship to Mesozoic Stromatoporoidea

another poriferan grade, has been a flag of convenience since the
erection of the genus Stromatopora Goldfuss, 1826. Whilst most The earliest stromatoporoids are Mid Ordovician, and the last
palaeontologists are happy to accept portmanteaus for groups of of the Palaeozoic groups disappeared close to the Devonian-
uncertain affinity, this belief does not sit well alongside biological Carboniferous boundary, following the Frasnian-Famennian crisis.
works such as others in this volume. The intragroup classification The problem for assessing the direct relationship between the two
has been developed since Nicholson (1886-92), and major works groups is that there is a considerable gap between the demise of
are that of Lecompte (1956), Galloway (1957), Stearn (1980) and Palaeozoic forms and the appearance in the Late Palaeozoic -
Stearn et al. (1999). The seven 'orders' of Palaeozoic stromato- Early Mesozoic of the stromatoporoidomorphs. Some hope is
poroids given by Stearn et al. (1999) show considerable variation given by the genus Newellia Wood et al., 1989, but its full relation-
in skeletal microstructure, in addition to macrostructure, and may ships require further investigation. This relationship is also
represent polyphyly within poriferan grades. However, there seems obscured by the possible differing skeletal mineralogy, presence of
to be a clear line of development from labechiid stock to clathrodic- spicules in some of the Mesozoic groups (giving them undoubted
tyids, actinostromatids and perhaps amphiporids. Stromatoporids, demo sponge affinity), and the paucity of completed studies on the
syringostromaellids and stromatoporellids are a natural grouping latter.
which mayor may not have direct descendents in the labechiids.
The work of Stearn & Pickett (1994) indicated that stromatoporel- Mesozoic Stromatoporoidea
lids and clathrodictyids have homologous relationships with the
sphinctozoan grade of construction (refer to chapter on Non- A much clearer understanding of the Mesozoic stromato-
Lithistid fossil Demospongiae (Reitner & Worheide, this volume)). poroids has arisen through the thorough study of microstructures
and other features by Wood (1987, et seq.). It is clear that Wood
Skeletal chemistry (1991b) regards Mesozoic stromatoporoids as demosponges, citing
several Recent taxa as examples of extant stromatoporoids (e.g.,
The nature of the basal calcareous skeleton, combined with see summary by Wood, 1990), but her interpretation of demo-
the vicissitudes of diagenesis in the geological settings in which sponge affinities between these and Palaeozoic stromatoporoid
they are found, and the instability of aragonite all conspire to cloud faunas has not been universally accepted by the palaeontological
the evidence for the type of original carbonate phase deposited by community. Further discussion of the stromatoporoid body plan in
the living organism. Many studies of diagenesis have failed to relation to other 'sclerosponge' skeletons is provided in the chapter
establish whether the skeleton was calcite, aragonite or some com- on Non-Lithistid fossil Demospongiae (Reitner & Worheide, this
bination thereof. Stromatoporoids are preserved in diagenetic envi- volume)).
ronments commonly hostile to pristine chemical preservation.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Order Homosclerophorida Dendy, 1905, Family Plakinidae Schulze, 1880

Guilherme Murley· & Maria Cristina Diaz2

1 Departamento de Invertebrados, Museu Nacional, Universidade Federal do Rio de Janeiro.

Quinta da Boa Vista, sino., Slio Crist6vlio. 20940-040 Rio de Janeiro, Brazil. (muricy@acd.ufrj.br)
2 Institute of Marine Sciences, University of California Santa Cruz, CA 94038, USA. (Diaz@cats.ucsc.edu)

Homosclerophorida Dendy (Demospongiae, Homoscleromorpha) contains a single family Plakinidae Schulze (including Oscarellidae
Lendenfeld and Corticiidae Vosmaer), with seven valid genera and about 60 valid species worldwide. Species live mainly in shallow
waters but a few have been recorded from abyssal depths (up to 2460m). Species are often encrusting, lobate, but massive species are
common in some genera (Plakortis, Plakinastrella); surface is usually smooth or microhispid and consistency varies from soft to carti-
laginous. All genera possess flagellated exo- and endopinacocytes, a basement membrane lining both choanoderm and pinacoderm, oval
to spherical choanocyte chambers with a sylleibid-like or leuconoid organization, and a unique incubated cinctoblastula-type larvae;
spicules, when present, are peculiar tetractines (calthrops) and derivatives. Genera are distinguished mainly by four morphological char-
acters: presence of a siliceous skeleton; presence of a cortex associated with a leuconoid aquiferous system and well-developed mesohyl
or a sylleibid aquiferous system with poorly developed mesohyl and ectosome; number of spicule size classes; and presence and type of
ramifications in the actines of calthrops (tetractinal spicules), with three distinct general morphologies recognized.
Keywords: Homosclerophorida; Plakinidae; Corticium; Oscarella; Placinolopha; Plakina; Plakinastrella; Plakortis; Pseudocorticium.

DEFINITION, DIAGNOSIS, SCOPE calthrop, Fig. 2F), or four rays (tetralophose calthrop, Fig. 2G-H).
Furthermore, two types of tetralophose calthrops are distinguished,
Synonymy those with all rays presenting a similar branching pattern
(homolophose calthrops, typical but not exclusive of Plakina
Order: Homosclerophorida Dendy, 1905: 64. Microscle- spp; Fig. 2G) and those in which one ray's branching pattern is dif-
rophora Sollas, 1887: 423 (in part). ferent from the other three (heterolophose calthrops or candelabra,
Family: Plakinidae Schulze, 1880: 447. Corticiidae Vosmaer,
1887: 324. Oscarellidae Lendenfeld, 1887a: 582. Placinidae Topsent,
1890d: 231.

Definition

Demospongiae with flagellated exo- and endopinacocytes, a

basement membrane lining both choanoderm and pinacoderm, oval
to spherical choanocyte chambers with a sylleibid-like or leu-
conoid organization, and a unique incubated cinctoblastula type
larvae; spicules, when present, are peculiar tetractines (calthrops)
and derivatives through reduction (diods and triods) or through
ramification of one to all four actines (lophose calthrops).

Diagnosis

Thin to massive-encrusting Demospongiae, with few species

presenting tubular growth forms (Fig. 1); surface usually smooth to
the touch, but sometimes wrinkled or with highly convoluted
appearance. Species are usually compressible and very dense, with
thinner species tending towards a soft or cartilaginous consistency.
A few encrusting species are fixed on the substrate by thin filaments
only (some species of Plakina and Corticium). Skeleton, when
present, is usually formed by a combination of small calthrops
andlor derivatives through reduction (diods and triods) (Fig. 2). The
rays of calthrops, diods, or triods may present multiple, sometimes Fig. 1. External morphology of plakinids (underwater close-ups). A,
complex branching, in which case the prefix 'lopho-' is used to Conicium candelabrum Schmidt (Marseille, France), specimen in top center
denominate the branching forms (lophodiods, lophotriods, and (scale 2em). B, Oscarella lobularis (Schmidt) (Marseille, France) (scale 2
em). C, Plakina jani Muricy et al. (Marseille, France) (scale Scm). D,
lophocalthrops). Among lophocalthrops the numbers of actines
Plakinastrella onkodes Uliczka (Carrie Bow Cay, Belize) (scale Scm). E,
ramifying may involve one ray (monolophose calthrop, Fig. 2D), Plakortis angulospiculatus (Carter) (Carrie Bow Cay, Belize) (scale 5 cm). F,
two rays (dilophose calthrop, Fig. 2E), three rays (trilophose Pseudocorticiumjarrei Boury-Esnault et al. (Marseille, France) (scale 2cm).

71
72 Porifera· Demospongiae • Homoscleropborida • Plakinidae

exclusive of Corticium spp; Fig. 2H). The pattern of ramification of into account the number of rounds of ramification in lophose
the lophose actines in lophocalthrops can be quite distinct among actines (1-3), their location along the actine ('p' for proximal, 'm'
species, and are valuable characters for both generic and specific for medial, 'd' for distal), and the presence of terminal spines (ts)
diagnoses. The patterns of ramification of lophose actines can (Fig. 3). In this system, the ramification pattern of a spicule with
be described by a simple alpha-numerical code, which takes only one round of ramifications close to the base of the lophose
actines and bearing tiny terminal spines at their extremities is coded
'lp, ts' (Fig. 3D). A spicule with one round of ramification at mid-
length of the actine, a second round of ramification close to the
extremity of the secondary ray, and bearing terminal spines is coded
'1m, 2d, ts', and so on (Fig. 3G; Muricy et al., 1998). Spicules are
generally found in one size class; however in Plakinastrella and
Placinolopha the spicules are present in two or more size classes.
Spicules are generally arranged uniformly in the sponge body,
surrounding the aquiferous system in a regular "alveolar" way or
more confusedly dispersed (Fig. 4). Minute diactinal microscleres
(Plakortis), and smalllophocalthrops (Corticium, Plakina), may be
found either dispersed on the sponge body, or concentrated at the
surface. The aquiferous system is either sylleibid-like (eurypylous
chambers uniformly arranged around large, parallel. descending
inhalant and exhalant canals) or leuconoid (mostly diplodal or

Fig. 2. Spicule types of plakinids. A-C, calthrops and derivates. A, diod

(25-750 fLm long). B, triod (actines 8-300 fLm long). C, calthrop (actines
8-320 fLm long). D-G, homolophose calthrops (actines 7-35 fLm long).
D, monolophose calthrop. E, dilophose calthrop. F, trilophose calthrop.
G, tetralophose caltbrop. H, heterolophose calthrop (=candelabrum). I, dis-
tally lophate calthrop (Placinolopha) (actines 275-300 fLm long).

Fig. 3. Ramification patterns of actines in plakinid lophose caltbrops. A,

'lp' (e.g., basal actines of Corticium candelabrum). B, '1m' (e.g., Plakina
monolopha and P. trilopha). C, 'Id'. D, 'lp, ts' (e.g., Plakina endoumensis). Fig. 4. Spicule arrangement in plakinids (photomicrographs of cross-
E, '1m, ts' (e.g., Plakina monolopha and P. trilopha). F, 'Id, 2d, ts' (e.g., sections). A-B, Corticium candelabrum (scale A, 100 fLm, B, 50 fLm). C,
Plakina crypta). G, '1m, 2d, ts' (e.g., Plakina janusi). H, 'Id, 2m, 3m, ts' Plakina monolopha (scale 100 fLm). D, P. trilopha (scale 200 fLm). E, Plakina
(e.g., Placinolopha bedoti). I, 'Ip, spined' (e.g., Corticium candelabrum). corticioides (scale 100 fLm). F, Plakortis angulospiculatus (scale 200 fLm).
Porifera· Demospongiae • Homosclerophorida • Plakinidae 73

aphodal chambers confusedly arranged in a complicated system of (Placinolopha, Fig. 21); and small heterolophose calthrops
small, sinuous inhalant and exhalant canals), often with a large (Corticium, Fig. 2H).
basal exhalant cavity, and sometimes also with ectosomal inhalant
cavities (Figs 4, 5). Choanocyte chambers, 30--90/Lm diameter, are Scope
usually eurypylous in Plakortis, Plakinastrella, Plakina, and
Oscarella, but are aphodal or diplodal in Corticium and The order Homosclerophorida (= Microsclerophora Sollas,
Pseudocorticium. A basement membrane underlines the choano- 1887) was created with three families, Plakinidae, Corticidae, and
derm and pinacoderm in genera that have been studied histologi- Thrombidae (Sollas, 1888; Dendy, 1905), of which only Plakinidae
cally (Corticium, Oscarella, Plakina, and Pseudocorticium). Both still remains. Until recently two families Plakinidae and
exo- and endopinacocytes are flagellated (Fig. 6A). Cytological Oscarellidae were recognized, but they were eventually merged
traits such as apopylar cell morphology and type of cell inclusions under Plakinidae and the order is currently monofamilial (Sole-
may be important generic and specific diagnostic characters of Cava et aI., 1992; Diaz & Van Soest, 1994; Boury-Esnauly et al.,
aspiculate plakinids (Fig. 6B-F). Larvae are incubated, of a unique 1995; Muricy et al., 1996a; Muricy, 1999). Placinidae Topsent,
cinctoblastula type, which is a hollow, ovoid larvae, with a single 1890d is a misspelling of Plakinidae.
external layer of flagellated cells and an equatorial belt of distinctly Over twelve nominal genera have been included in this family,
pigmented cells (Fig. 7; see also Boury-Esnault & Riitzler, 1997). but only seven are considered to be valid and now included:
Plakortis Schulze, 1880, Plakinastrella Schulze, 1880, Plakina
Remarks Schulze, 1880, Placinolopha Topsent, 1897a, Corticium Schmidt,
1862, Oscarella Vosmaer, 1887 and Pseudocorticium Boury-
A generic classification of the Plakinidae is centered around Esnault et al., 1995. The two incertae sedis genera, Corticellopsis
four principal diagnostic features: presence/absence of siliceous and Astroplakina, are excluded based on the clear astrophorid
skeleton; presence/absence of a cortex associated with the archi-
tecture of the aquiferous system and type of choanocyte chambers;
number of spicule size classes; and presence and type of ramifica-
tions in the actines of calthrops, with at least three distinct general
morphologies recognized: small homolophose calthrops (Plakina,
Fig. 2D-G); large lophodiods, lophotriods and lophocalthrops

Fig. 5. Aquiferous system of plakinids (photomicrographs of cross-

sections). A, Corricium candelabrum: micrograph showing the thick cortex Fig.6. Some cell types of plakinids. A, flagellated pinacocyte of Plakina jani
(Co) in the upper portion of the photograph, and a leuconoid system with (scale 1 f,Lm). B, apopylar cells of Oscarella imperialis, leading towards an
aphodal chambers (ac) towards the lower portion (scale 50 f,Lm). B, excurrent canal (scale 2 f,Lm). C-F, cells with inclusions that characterise
Oscarella lobularis: micrograph showing a sylleibid aquiferous system with genera or species among aspiculate plakinids. C, cells with paracrystalline
eurypilous chambers (ec) (scale lOOf,Lm). C, Plakina monolopha: micro- inclusions, cell type I, of Pseudocorricium jarrei (scale 2 f,Lm). D, 'crescent-
graph showing a sylleibid aquiferous system with eurypilous chambers (ec), shaped' cell with inclusions, cell with inclusions type I of Oscarella viridis
and canals (Ca) (scale 100 f,Lm); D, Pseudocorticium jarrei: micrograph (scale 2 f,Lm). E, spherulous cell with granular inclusions, cell type II of
showing the cortex (Co) in the upper portion of the photograph, and a leu- Oscarella microlobara (scale 2 f,Lm). F, vacuolar cell with paracrystalline
conoid system with aphodal chambers (ac) towards the lower portion; apho- inclusions, cell with inclusions type I of Oscarella imperialis (scale 1 f,Lm)
dus (ap) can be seen leaving some choanocyte chambers (scale 50 f,Lm). (from Boury-Esnault er aI., 1995, Muricy er al., 1996a).
74 Porifera· Demospongiae • Homosclerophorida • Plakinidae

biradiate forms, the triradiate and biradiate forms being derived

from the quadriradiate forms by reduction of one or two rays. The
biradiates occur in the form of weakly bent oxea, with an irregular,
knobby-knotty, curved and in general crooked-looking center".

. Subsequently, Sollas (1888) grouped the Plakinidae together with

Corticiidae Vosmaer and Thrombidae Sollas in the suborder
Microsclerophora, order Choristida, next to the suborder
Astrophora (which included Pachastrellidae, Ancorinidae, and
Geodiidae). The Choristida together with the Lithistida were
assigned to the subclass Tetractinellida. Topsent (1895) considered
the Microsclerophora as a suborder of the order Camosa Carter,
• which was separated from his order Tetractinellida on the basis of
lack of large diactinal megascleres and a trend towards reduction
in size of tetractinal spicules. Dendy (1905) changed the name

s··. ':
,
. '. .
,"
,

..
Microsclerophora to Homosclerophora based on the common belief
that the small plakinid spicules did not represent real microscleres.
De Laubenfels (1936a) viewed the Camosa Carter as composed of
the families Halinidae de Laubenfels, Plakinastrellidae de
Laubenfels, Chondrillidae Gray, Chondrosiidae Schulze and
Dedalopeltidae de Laubenfels. This arrangement was clearly artifi-
cial, and subsequent authors abandoned the use of the name
Camosa. The family Plakinidae was maintained in the order
Choristida, subclass Tetractinellida, until the introduction of repro-
ductive characteristics into sponge systematics (Levi, 1957b,
1973). Plakinids were then relocated in a separate subclass
.. Homoscleromorpha, order Homosc1erophorida, based on the
possession of a distinct morphology combined with a unique
.. amphiblastula-type larva which is non-homologous to the cal-
caronean amphiblastula. The order consisted of two families, the
Fig. 7. Embryos and larvae of plakinids. A, Oscarella lobularis embryos Plakinidae which included the original Schulze's plakinid genera
and larvae in different stages of development (scale 200 fLm). B, (Plakina, Plalwrtis, and Plakinastrella) and a few of the
Pseudocorticium jarrei (scale 50 fLm). Corticiidae genera (Corticium Vosmaer, Rachella Sollas), and
the family Oscarellidae Lendenfeld, which included sponges with-
nature of their spiculation. Until further studies clarify the supra- out any mineral or fibre skeleton (Oscarella Vosmaer). Until
generic relationships among homosclerophorid genera, we must recently, comprehensive systematic sponge literature (Levi, 1973;
consider both skeletal and non-skeletal genera as belonging Bergquist, 1978; Hartman, 1982) included twelve genera in the
to the former family Plakinidae Schulze (1880). Approximately family, which were recently reduced to six valid genera, based
70 species of sponges from these genera have been published mostly on the study of skeletal characters (Diaz & Van Soest,
worldwide but only 60 are considered valid (Table 1). The number 1994). Another recent modification of this sponge group was the
of species described in the literature is clearly geographically biased merging of both families (Plakinidae and Oscarellidae) under the
towards areas that have been better studied such as the former family Plakinidae (Sole-Cava et al., 1992; Boury-Esnault
Mediterranean, the West Pacific and the Caribbean. It is expected et al., 1992b; Diaz & Van Soest, 1994). The genera recognized
that when the least studied biogeographic regions are surveyed for were: Plakortis Schulze, 1880 (incl. Roosa de Laubenfels, 1934),
this group the species numbers will increase considerably. Species Plakinastrella Schulze, 1880 (incl. Dercitopsis Dendy, 1905),
of various genera such as Oscarella, Plakina, Plalwrtis, and Plakina Schulze, 1880 (incl. Plalwosa de Laubenfels, 1936b),
Corticium with relatively simple morphologies are found in distinct Placinolopha Topsent, 1897a (incl. Acanthoplakina Burton, 1959a;
biogeographic regions (e.g., Plalwrtis spp. in the Caribbean and in Diactinolopha Sara, 1960b), Corticium Schmidt, 1862, and
the West Pacific), and several species are considered cosmopolitan Oscarella Vosmaer, 1887 (incl. Octavella Tuzet & Paris, 1964).
(e.g., Oscarella lobularis, Plalwrtis simplex, Plakina monolopha). Astroplakina Dendy & Burton, 1926, and Corticellopsis Sollas,
Such disjunct populations, although morphologically similar, are 1888 were excluded from the family, due to their possession of
expected to be reproductively isolated and to have diverged geneti- euasters, and placed as incertae sedis (Diaz & Van Soest, 1994). A
cally enough in time to deserve a status of distinct species. new genus of sponges without skeleton, Pseudocorticium, with well-
developed cortex, diplodal choanocyte chambers, and a leuconoid
Corticium-like aquiferous system has been recently described from
History and Biology Mediterranean waters (Boury-Esnault et al., 1995; Figs IF, 5D, 6C,
7B). This discovery supported the artificiality ofhomosclerophorid
Schulze (1880) erected this family to include three closely suprageneric classification (Boury-Esnault et aI., 1992b; Sole-Cava
related, newly described genera: Plakina, Plakortis, and et al., 1992; Diaz & Van Soest, 1994), which distinguished a sepa-
Plakinastrella. He described the species in the family to include rate family for the skeleton-lacking genera (the Oscarellidae).
"Tetractinellida with isolated needles (not bound by horny Pseudocorticium is an aspiculate genus which presents closer affin-
substance), which consist of a series of quadriradiate, triradiate, and ity in both biochemical (allozymes) and histological traits with the
Porifera' Demospongiae • Homoscleropborida • Plakinidae 75

Table 1. Plakinid species, with the distribution of the original material described.

Species Reference Locality Species Reference Locality

Corticium Thomas, 1968e India P. monolopha Schulze, 1880 NW Mediterranean

acanthastrum P. pacifica Desqueyroux-Faundez & Galapagos
C. bargibanti Levi & Levi, 1983b New Caledonia Van Soest, 1997
C. candelabrum Schmidt, 1862 NW Mediterranean P. reducta (Pulitzer-Finali, NW Mediterranean
C. niger Pulitzer-Finali, 1996 New Guinea 1983)
C. quadripartitum Topsent, 1923 Caribbean P. tetralopha (Hechtel, 1965) Jamaica
C. simplex Lendenfeld, 1907 N Australia P. tetralophoides * Muricy et al., 1998 Japan
Oscarella Muricy et aI., 1996a NW Mediterranean P. topsenti (Pouliquen, 1972) NW Mediterranean
imperiolis P. trilopha Schulze, 1880 NW Mediterranean
O. lobularis (Schmidt, 1862) NW Mediterranean P. versatile (Schmidt, 1879) Gulf of Mexico
O. microlobata Muricy et al., 1996a NW Mediterranean P. weinbergi Muricy et aI., 1998 E Mediterranean
O. tuberculata (Schmidt, 1868) NW Mediterranean Plakinastrella (Dendy, 1905) Sri Lanka
O. viridis Muricy et al., 1996a NW Mediterranean ceylonica
Placinolopha (Thomas, 1970) India P. clathrata (Kirkpatrick, 1900b) Funafuti
acantolopha P. copiosa Schulze, 1880 NW Mediterranean
P. bedoti Topsent, 1897 a Indonesia P. mammillaris (Lendenfeld, 1907) W Australia
P. europae Vacelet & Vasseur, 1971 Madagascar P. mixta Maldonado, 1992 W Mediterranean
P. moncharmonti (Sara, 1960b) NW Mediterranean P. onkodes Uliczka, 1929 Caribbean
P. sarai Levi & Levi, 1989 Philippines P. oxeata Topsent, 1904b Azores
P. spinosa Kirkpatrick, 1900b Funafuti P. polysclera Levi & Levi, 1989 Philippines
Plakina (Sara, 1960b) NW Mediterranean P. trunculifera Topsent, 1927b Azores
bowerbanki Plakortis (Carter, 1879b) Caribbean
P. australis (Gray, 1867a) W Australia angulospiculatus
P. brachylopha Topsent, 1928c Azores P. copiosa Pulitzer-Finali, 1993 Kenya
P. corticioides Vacelet et al., 1976 Madagascar P. erythraena Levi, 1958 Red Sea
P. corticolopha Levi & Levi, 1983b New Caledonia P. galapagensis Desqueyroux-Faundez & Galapagos
P. crypta Muricy et al., 1998 NW Mediterranean Van Soest, 1997
P'dilopha Schulze, 1880 NW Mediterranean P. halichondroides (Wilson, 1902) Caribbean
P. elisa (de Laubenfels, 1936b) Caribbean P. kenyensis Pulitzer-Finali, 1993 Kenya
P. endoumensis Muricy et al., 1998 NW Mediterranean P. lita de Laubenfels, 1954 Caroline lsI.
P. fragilis Desqueyroux-Faundez & Galapagos P. nigra Levi, 1953b Red Sea
Van Soest, 1997 P. quasiamphiaster Diaz & Van Soest, 1994 Vanuatu
P. jamaicensis Lehnert & Van Soest, 1998 Jamaica P. simplex Schulze, 1880 NW Mediterranean
P.jani Muricy et al., 1998 NW Mediterranean P. zygompha (de Laubenfels, 1934) Caribbean
P. microlobata Desqueyroux-Faundez & Galapagos Pseudocorticium Boury-Esnault et al., NW Mediterranean
Van Soest, 1997 jarrei 1995

* Plakina tetralophoides Muricy et al., 1998 was first described as P. tetralopha by Tanita & Hoshino (1989).
Further Notes. Plakinastrella schulzei and P. intermedia are not included as they belong to the astrophorid genus Penares. Dercitopsis ceylonica and
D. minor were assigned to Plakinastrella (the latter considered a junior synonym of P. clathrata; Diaz & Van Soest, 1994). Placinolopha europae was described
as P. spinosa europae VaceIet & Vasseur, 1971; P. mirabilis de Laubenfels, 1954 probably belongs to Theonella (Theonellidae, 'Lithistida'); Oscarella cruenta
(Carter, 1881b) sensu Keller (1889) is probably a Chondrosia (Chondrosida: Chondrillidae). Oscarella membranacea Hentschel, 1909 and O. tenuis Hentschel,
1909 are considered insufficiently described (Muricy et ai., 1996a). Hooper & Wiedenmayer (1994) suggested that Achinoe australis Gray, 1867a might belong
to Corticium, whereas it more probably belongs to Plakina (see text). Species incertae sedis are Astroplakina stelligera Dendy & Burton, 1926, Corticellopsis
stelligera Schmidt, 1862, and Corticellopsis novaezealandiae Bergquist, 1968 - these are not included as they are probably astrophorids.

skeletal genus, Corticium, than with the non-skeletal genus Mediterranean is essential to understand the true diversity of the
Oscarella (Solt~-Cava et al., 1992; Boury-Esnault et al., 1995). family and the relationships among its members.
However, Pseudocorticium seems closer to Oscarella than to Biochemical characteristics of species from three genera
Corticium or Plakina in cell composition, and the possibility remains (Oscarella, Plakina, and Plakortis) have been described by several
that the aspiculate plakinids are indeed monophyletic (Muricy et aI., authors (Bergquist & Hartman, 1969; Cimino et al., 1975; Higgs &
1996a; Muricy, 1999). Histological, cytological and biochemical Faulkner, 1978; Ravi et al., 1979; Faulkner et al., 1979; Stierle &
studies were conducted on Oscarella, Corticium, Plakina, and Faulkner, 1980; Faulkner & Ravi, 1980; Rosser & Faulkner, 1984;
Pseudocorticium (Sole-Cava et al., 1992; Boury-Esnault et al., Bergquist et aI., 1984; Sakemi et al., 1987; Aiello et al., 1990;
1992b, 1995; Muricy, 1999; Muricy et al., 1996a,b, 1998, 1999). Inman et aI., 1990; West et al., 1990; Davidson, 1991). Among the
These studies demonstrated the importance of these traits to the char- most interesting chemical components of plakinid species are rare,
acterization of genera and species, and to the understanding of the cytotoxic epoxy-sterols in Oscarella (Aiello et al., 1990), and
phylogenetic relationships among plakinid genera. Similar studies antimicrobial steroidal alkaloids in Plakina (Rosser & Faulkner,
on the poorly known genera Plakortis, Plakinastrella, and 1984). Species of Plakortis are known to produce both monoter-
Placinolopha would be of particular interest to achieve a more pene derivatives unknown from other sponges (Faulkner & Ravi,
detailed picture of the phylogeny of the Plakinidae. Also, a complete 1980), cytotoxic alkaloid pigments (West et al., 1990), cytotoxic
characterization of the plakinid fauna in regions other than the peroxydes (Sakemi et aI., 1987; Davidson, 1991), and pentacyclic
76 Porifera· Demospongiae • Homoscleropborida • Plakinidae

aromatic alkaloids with anti-helmintic and anti-reverse transcrip- more definitive phylogenetic picture of the family. Further biological
tase activities (Inman et al., 1990). characterization of all plakinid genera (cytological, histological,
Plakinid species comprise mostly small, thinly to massively and genetic) such as in previous studies with species of Oscarelia,
encrusting species, widely distributed geographically and bathy- Corticium, Pseudocorticium, and Plakina will aid in understanding
metrically, from tidal depths up to at least 2460m (Plakina brachy- intergeneric affinities. Secondly, until the present, these sponges,
lopha; Topsent, 1928c). They grow only on hard substrates. Softer with their unique cinctoblastula larvae, a basement membrane lin-
and thinner forms predominate in sheltered habitats (caves, ing both choanoderm and pinacoderm, flagellated pinacocytes, and
crevices, etc.), whereas in exposed areas (rock outcrops, coral the distinctive morphology of their spicules, aquiferous system and
colonies, etc.) the species tend to be more massive and rigid larvae, remain isolated from other extant Demosponges and its out-
(Fig. 1). Species descriptions and surveys are found dispersed in a group relationships are still unclear (e.g., see Van Soest, 1984a;
series of sponge monographs from all the world's oceans (Table 1). Grothe, 1989; Muricy, 1999). The answer to this problem seems to
Definitions of the constituent taxa and important taxonomic char- depend on molecular studies (e.g., sequencing of 18S ribosomal
acters are given by Diaz & Van Soest (1994), Boury-Esnault et al. genes) to discern the relationships of this family with other sponge
(1984, 1992b, 1995), Muricy et al. (1996a, 1998, 1999). groups (e.g., Astrophorida, Ca1caronea). A first approach to the
phylogeny of the Plakinidae, including species of four genera
Remarks (Corticium, Oscarella, Plakina, and Pseudocorticium) was
attempted by Muricy (1999), using morphological and cytological
Most plakinid genera are clearly associated by the common characters and Discodermia (Lithistida) as the outgroup. Muricy
nature of their skeleton, aquiferous system, cytological features (1999) found little support for a monophyletic family Oscarellidae
and larval morphology. However two major problems still haunt clustering Oscarella and Pseudocorticium, and both aspiculate
our understanding of this group. Firstly, important biological genera were kept in the family Plakinidae. These analyses must
aspects of approximately half of plakinid genera remain poorly however be complemented by inclusion of other genera (Plakortis,
studied (Plakortis Schulze, 1880, Plakinastrella Schulze, 1880, Plakinastrella, and Placinolopha) and study of other characters
Placinolopha Topsent, 1897a). This prevents us from drawing a (e.g., DNA sequences).

KEY TO GENERA

(1) With inorganic (spicular) skeletal complement ................................................................................................................................... 2

Without inorganic (spicular) skeletal complement .............................................................................................................................. 6
(2) Skeleton mainly composed of diods, triods, and/or calthrops in one size class .................................................................................. 3
Skeleton mainly composed of diods, triods and/or calthrops with a large size variation .................................................................... 5
(3) Lophose diods, triods, or calthrops complement the main skeleton of non-lophose spicules ............................................................. 4
Lophose spicules absent, diactinal "microscleres" (microrhabs) present in some species .........................................................Plakortis
(4) Heterolophose calthrops (candelabra) complement the main skeleton of non-lophose spicules, which might be absent orrare; choanocyte
chambers usually aphodal .. .................. .......... .................. ... ............. .................. ..... ... .............. ..... ..... ..................................... ......... Corticium
Lophocalthrops with one to four homogeneously ramified actines complement the main skeleton of non-Iophose spicules; candelabra
absent; choanocyte chambers usually eurypylous .............................................................................................................................. Plakina
(5) Skeleton composed of non-lophose diods, triads and/or calthrops in three size classes ........................................................ Plakinastrella
Skeleton formed by diods, triods, and/or calthrops with large size variation; the larger spicular category presents tenninally-branching rays
.. ...... .... ......... ......... ...... ..... ...... ................................... .... ..... .... ............... ...................................... .... ........ ... ..................... ......... ...... Placinolopha
(6) With sylleibid-like aquiferous system and eurypylous choanocyte chambers; ectosome thin, proportion of mesohyl
to chambers < 1: 1 .................................................................................................................................................................. Oscarella
With leuconoid aquiferous system and diplodal choanocyte chambers; ectosome thick, proportion of mesohyl
to chambers >2:1 ...................................................................................................................................................... Pseudocorticium

CORTICIUM SCHMIDT, 1862 Diagnosis

Synonymy Thinly encrusting to cushion-shaped Plakinidae with a spicu-

lation consisting almost exclusively of non-lophose calthrops in
Corticium Schmidt, 1862: 42. one size class and heterolophose calthrops ('candelabra'; Fig. 2H).
Homolophose calthrops may also be present (Fig. 2D), and non-
lophose calthrops are absent in some species. Aquiferous system
Type species leuconoid, with aphodal choanocyte chambers, and about 300
choanocytes per chamber (Boury-Esnault et al., 1984). The species
Corticium candelabrum Schmidt, 1862: 42 (by monotypy). of this genus present variable but usually thick cortex (100-
300/-Lm), and a proportion of mesohyl to chambers of about 1: 1
(Boury-Esnault et al., 1995).
Definition
Description of type species
Thin to thick encrusting plakinid, with a skeleton dominated by
non-lophose calthrops and heterolophose calthrops; homolophose Corticium candelabrum Schmidt, 1862 (Figs lA, 2H, 3A, I,
calthrops can be found in certain species. 4A-B,5A).
Porifera· Demospongiae • Homosclerophorida • Plakinidae 77

Synonymy. Corticium candelabrum Schmidt, 1862: 42. (Cruz & Bacallado, 1981), Caribbean (Topsent, 1923), Indian
Material examined. Lectotype: LMJG 15353/0 - Sibenik, Ocean (Thomas, 1968e, 1970), Australia (Gray, 1867a;
Croatia (here designated). Other material. LMJG 15508 - no desig- Lendenfeld, 1907; Burton, 1934a; Wiedenmayer, 1989; Hooper &
nated locality. USNM 23889 - Adriatic. USNM 39217 - Marseille, Wiedenmayer, 1994), New Guinea (Pulitzer-Finali, 1996), and
France. ZMA Por. 52 - Banyuls, France. New Caledonia (Levi & Levi, 1983b) (Table 1).
Description. Thinly encrusting to cushion-shaped, lobate, up
to 3 cm long and 1.5 cm thick, which contracts when taken out of the
water to such an extent that its surface takes a highly convoluted OSCARELIA VOSMAER, 1884
appearance. Body fixed on the substratum by thin filaments.
Surface uneven, slightly rough to touch. Round oscula, 1-5 mm in Synonymy
diameter, contractile. Colour alive light brown to tan, preserved in
spirit. Consistency firm, cartilaginous. Ectosome with a well [Oscaria] Vosmaer, 1881: 163 (preocc. by Oscaria Gray, 1873
defined cortex 100-300 j.lm thick, with abundant amoeboid cells. (Reptilia». Oscarella Vosmaer, 1884: pI. 8 (explanation); 1887: 326
Subectosomal cavities absent, basal cavity well developed. (nom. nov. for [Oscaria] Vosmaer). Oscarella Vosmaer, 1887: 326.
Aquiferous system leuconoid, with ovoid, aphodal choanocyte Octavella Tuzet & Paris, 1964: 88 (no type specimens designated).
chambers, 50-70 j.lm in diameter. Skeleton confused, with spicules Taxonomic decision after Vosmaer (1887: 326); Boury-Esnault
scattered between choanocyte chambers. Although also present in et al. (1984: 13, 1992b: 282); Diaz & Van Soest (1994: 102).
the choanosome, candelabra are concentrated at the surface and bor-
dering canals. Spicules are irregular, non-Iophose calthrops in one
size class (actines 23-35j.lm); monolophose calthrops (actines Type species
25-32j.lm) with a ramification pattern in which lophose actines
have only one, proximal ramification point which gives rise to 3-5 Halisarca lobularis Schmidt, 1862 (by monotypy).
conical, smooth rays (a pattern coded '1 p, conical'), and candelabra
23-35j.lm long. The 3 'basal' (equally ramified) actines of cande-
labra have a ramification pattern similar to that of monolophose Definition
calthrops (1 p, conical); the fourth actine ramifies basally in 4-10
longer and thinner microspined rays (Figs 2H, 3A, I). Plakinidae without spicules, with an aquiferous system made
Remarks. The best apomorphy of the genus Corticium is the up of spherical, eurypylous chanocyte chambers uniformly arranged
presence of heterolophose calthrops (candelabra), which are a spe- around large, regular exhalant canals.
cial kind of tetralophose calthrops in which three actines are rami-
fied in a simple pattern (one medial ramification point, 3-4 conical
rays, smooth), and the fourth actine has a unique pattern of ramifi- Diagnosis
cation: seven to ten thin, micro-spined rays diverge from a proxi-
mal ramification point. Several species described in the genus Plakinidae without skeleton, with thinly encrusting to lobate
Corticium lack candelabra, having instead homogeneously rami- shape. Thin ectosome « 100 j.lm), often nearly limited to the pina-
fied trilophose and/or tetralophose calthrops typical of Plakina coderm, true cortex absent. Mesohyl ill-developed, with a propor-
(Corticium versatile Schmidt, 1879; C. bowerbankii Sara, 1960b; tion of mesohyl to chambers varying from 0.5: 1 to 1.2: 1. The
C. tetralophum Hechtel, 1965; C. topsenti Pouliquen, 1972; and C. aquiferous system has a sylleibid-like organization, with spherical,
reductum Pulitzer-Finali, 1983). These species were accordingly eurypylous choanocyte chambers uniformly arranged around large,
transferred to Plakina (Muricy et al., 1998). The genus Corticium regular exhalant canals, and a large basal exhalant cavity (Fig. 5B)
shares with Plakina the presence of simple, monolophose calthrops (after Boury-Esnault et al., 1995).
and non-Iophose calthrops, as well as large apopylar cells with
osmiophilic inclusions (Murlcy et al., 1999). Schmidt (1862) did Description of type species
not designate a holotype for the type species. We found two speci-
mens from the type locality (Sibenik, Croatia), which apparently Oscarella lobularis (Schmidt, 1862) (Figs lB, 5B, 7A).
belong to the original series. Specimen LMJG 15353/0 is here des- Synonymy. Halisarca lobularis Schmidt, 1862: 80.
ignated as the lectotype, in accordance with the ICZN (Art. 74) Octavella galangaui Tuzet & Paris, 1964: 88.
(Anon., 1999). Specimen LMJG 15508 (no designated locality) is Material examined. Neotype: MNHN LBIM DNBE 1991-1 -
here designated as a paralectotype. Corticium candelabrum seems Marseille, France. Other material. UFRJPOR 4378, 4379, 4380 -
to be a well defined, easily identifiable species, and it would seem Marseille, France.
inappropriate to consider a priori that non-Mediterranean records Description. Shape thinly to massively encrusting, lobate.
were misidentifications. However, in the closely related genus Size up to 20 cm wide and 3 cm high, lobes 1 cm wide and high.
Plakina it was shown that allozyme patterns and very subtle differ- Colour variable (purple, violet, or blue, often with cream tinges at
ences in the ramification pattern of lophocalthrops were diagnostic the base). Surface smooth, with scattered inhalant ostia and circu-
at the species level and may reveal cryptic speciation (Muricy lar oscula (5-10mm in diameter) at the top of the lobes.
et al., 1996b, 1998). Corticium candelabrum awaits similar studies Consistency soft. Aquiferous system sylleibid-like (with a radial
to prove or disprove its cosmopolitan nature. arrangement of chambers around exhalant canals), with eurypylous
choanocyte chambers 35-90 j.lm in diameter. Ectosome thin (5-50
Distribution j.lm), unspecialized, without ectosomal cavities. A large exhalant
basal cavity extends through the center of the lobes to the oscula.
Six species are known from the Mediterranean (Schmidt, Proportion of mesohyl to chambers approximately 0.5:1. Spicule
1862; Topsent, 1895; Uriz & Bibiloni, 1984), Eastern Atlantic and fibre skeleton absent.
78 Porifera· Demospongiae • Homosclerophorida • Plakinidae

Remarks. Oscarella species are homogeneous in morpho- Type species

logical, anatomical and reproductive characters: all are thinly to
thickly encrusting, lobate, with reduced mesohyl and thin ectosome PZacinoZopha bedoti Topsent, 1897a: 429 (by original
(usually less then 100/-Lm thick). The aquiferous system superfi- designation).
cially resembles the sylleibid organization of some Calcarea: spher-
ical, eurypylous choanocyte chambers are arranged around inhalant Definition
and exhalant canals, which run perpendicularly from the surface
down to a basal exhalant cavity, ramifying and anastomosing in the Plakinidae with a spiculation of large lophodiods, sometimes
choanosome (Fig. 5B). Oscarella species are hard to characterize supplemented by lophotriods and/or lophocalthrops, and a comple-
morphologically except in a few cases by the use of field characters. ment of non-Iophose diods, triods, and/or calthrops.
Examination of cytological characters (at least in semithin sections
of properly fixed specimens) is essential for species discrimination Diagnosis
in Oscarella, especially when taken in connection with molecular
methods such as allozyme electrophoresis (Muricy et aZ., 1996a), or Thinly to massively encrusting Plakinidae with a spiculation
DNA analysis. Oscarella shares with Pseudocorticium the absence of large lophodiods, sometimes supplemented by lophotriods
of skeleton and cytological traits such as thin, triangular apopylar and/or lophocalthrops, and a complement of non-Iophose diods,
cells and two peculiar types of cells with inclusions (Boury-Esnault triods, and/or calthrops. Tetralophose calthrops are found in one
et aZ., 1995; Muricy et aZ., 1996a; Fig. 6B, D-E). They are distin- species (P. acanthoZopha).
guished by the presence in Pseudocorticium of well-developed
mesohyl, a relatively thick cortex, leuconoid aquiferous system and Description of type species
diplodal choanocyte chambers (Fig. 5D). Oscarella ZobuZaris is very
simple morphologically (Fig. IB), and it has often being con- PZacinoZopha bedoti Topsent, 1897a (Figs 21, 3H).
founded with its sibling species O. tubercuZata (Schmidt, 1868), Synonymy. PZacinoZopha bedoti Topsent, 1897a: 429.
which is abundant in the NW Mediterranean (Boury-Esnault et aZ., Material examined. Holotype: MHNG C-12/6 (slide
1992b). These two species can only be confidently distinguished by MNHN LBIM DT1814) - Amboine, Indonesia (Desqueyroux-
molecular or cytological methods, and all records of O. ZobuZaris Faundez, 1981: 756).
published before this distinction was made must have their specific Description. Topsent (1897a) described PZacinoZopha
status re-evaluated (especially those from outside the Mediterranean). bedoti as a thinly encrusting sponge, 30 X 23 mm wide and 1-3 mm
The synonymy of Octavella and Oscarella is justified since the only thick. Colour white in spirit. Surface smooth, without visible open-
difference between the type species of both genera is the a1ledged ings. Consistency firm, brittle. Spiculation dense, composed of
absence of flagella in endopinacocytes of O. gaZangaui, which calthrops and derivatives in three size classes: smaller spicules are
is probably due to poor preservation conditions (Boury-Esnault abundant diods (100-160 X 3-7 /-Lm), sinuous, acerate, centrotylote
et aZ., 1984). and variable in shape, less common triods (actines 60-70 X 5/-Lm),
and rare calthrops (actines 50/-Lm long); medium-size spicules are
Distribution diods (350-550 X 11-25 /-Lm), triods and calthrops (both with
actines 110-190 X 20/-Lm); larger spicules are abundant lophodi-
Cosmopolitan: Mediterranean (Schmidt, 1862; Topsent, ods (750 X 50 /-Lm), rare lophotriods and abundant lophocalthrops
1895; Driz & Bibiloni, 1984; Levi, I957b), E Atlantic (Levi, (both with actines 275-300 X 35-50 /-Lm), and sometimes also
1952; Cruz & Bacallado, 1981; Boury-Esnault & Lopes, 1985), lophopentactines. Lophocalthrops predominate at the surface, and
N Atlantic (Van Soest & Weinberg, 1980; Burton, 1959a; Koltun, lophodiods are more common in the choanosome. Lophose
1964b), S Atlantic (Burton, 1930e), Caribbean (G. Murlcy, unpub- spicules ramify dichotomously several times, forming straight or
lished data), Red Sea (Levi, 1958), Indian Ocean (Vacelet et aZ., curved conical rays arranged in a tree-like fashion. Our analysis of
1976), NW Pacific (Koltun, 1962b) and Antarctic (Topsent, 1917). the type species material conforms overall to Topsent's (1897a)
However, only five species of Oscarella are currently recognized, description, to which a few observations could be added. The
all from the Mediterranean: the type species O. ZobuZaris (Schmidt, choanosomal skeleton has spicules of all types randomly dispersed.
1862), O. tubercuZata (Schmidt, 1868), O. viridis Muricy et aZ., The ramification pattern of lophose spicules is peculiar, with one
1996a, O. microZobata Murlcy et aZ., 1996a, and O. imperialis distal ramification point from which depart 2-3 conical, curved
Muricy et aZ., 1996a. The correct specific assignment of the several secondary rays which often bear a second and a third medial points
records of O. ZobuZaris worldwide awaits a detailed anatomical and of ramification (with give rise to 2-4 secondary rays each) before
cytological revision, but it is already clear that most of these might ending in acerate or less often terminally spined extremities (a gen-
consist of cryptic sibling species representing new taxa. eral pattern coded 'ld, 2m, 3m'; Figs 21, 3H). The number of ram-
ifications varies among actines of the same spicule or rays of the
same actine, but they always show the typical conical, curved
PLACINOLOPHA TOPSENT, 1897 shape. Spicules of MNHN LBIM DT1814 are similar in both gen-
eral shape and ramification pattern to MNHG C-12/6, although
Synonymy slightly larger: abundant diods (60-550 X 2-8 /-Lm); rare triods
(actines 20-40 X 2-6/-Lm); rare calthrops (actines 40-210/-Lm);
PZacinoZopha Topsent, 1897a: 429. PZakinoZopha de common lophodiods (430-720 X 15-50/-Lm); lophotriods absent;
Laubenfels, 1954: 249. [AcanthopZakinal Burton, 1959a: 156 common lophocalthrops (actines 190-250 X 25-52 /-Lm).
[unavailable name, see remarks]. DiactinoZopha Sara, 1960b: 2 Remarks. PlacinoZopha was synonymized with Plakina by
(after Levi & Levi, 1989: 45; Diaz & Van Soest, 1994: 102). Lendenfeld (1903) but reinstated by Topsent (1928c) and de
Porifera· Demospongiae • Homosclerophorida • Plakinidae 79

Laubenfels (1936a) without apparent justification (Sara, 1960b). or aphodal, usually with a radial arrangement around incurrent
These two genera are consistently different in the possession of spicule and excurrent canals (called sylleibid-like arrangement, such as in
size classes and the much greater upper range in the size of spicules in P. monolopha, P. elisa).
Plncinolophn (which are shared with Plakinastrella), and in the rami-
fication pattem of their lophose spicules. Plakinolophn de Laubenfels,
1954 is a misspelling of Placinolophn Topsent. In Placinolophn bedoti Description of type species
the actines of long lophodiods, lophotriods and lophocalthrops bear a
terminal ramification point giving rise to secondary rays which in turn Plakina monolopha Schulze, 1880 (Figs 2D, 3B, E, 4C, 5C).
ramify medially in conical rays; a few actines of some spicules may Synonymy. Plakina monolophn Schulze, 1880: 407.
also show a third, medial ramification round (a pattern coded' Id, 2m, Material examined. Lectotype: BMNH 1883.12.4.29 -
3m, conical') (Fig. 3H). In Plakina, smalliophocalthrops have several Naples (Murlcy et al., 1998). Other material. ZMA POR 4391,
different branching patterns which are diagnostic at the species level 4424 - Ireland. ZMA POR 5123 - Brittany, France. ZMA POR
(Fig. 2D-G; Murlcy et al., 1998). Diactinolophn Sara (l960b) was 1821 - Curar,:ao. ZMA POR 7978, 8064, 8398 - Indonesia. UFRJ-
synonymized with Placinolophn based on the similarity of the spicula- POR 4350, 4351, 4352 - Grand Congloue Island, Rioux
tion of its type species, D. monchnrmonti Sara, 1960b, to that of Archipelago, Marseille, France.
P. bedoti. [Acanthoplakina] Burton, 1959a is here considered an Description. Sponges small, thinly encrusting, up to 2 X 2 cm
unavailable name since it was erected without a description or defini- wide and 1-5 mrn thick. Shape discoidal or irregular, with smooth,
tion, or a valid type species designation (ICZN Art. 13). The genus elevated borders forming a 'ring canal' around the sponge body.
Plncinolophn has been very poorly studied and there is no information Body attached to the substratum by thin filaments. Surface
on the choanocyte chambers type or organization. Species distinction microlobate, with rugose, irregular lobes 0.5-2.0 mrn in diameter
is based on the size and number of spicule types and size categories, and height, which may fuse together. Colour alive and in spirit
together with external morphological characteristics. white or cream. Consistency soft, fragile. Ectosome poorly devel-
oped, 15-30 j.Lm thick, without subectosomal cavities. Spaces
Distribution between lobes form open inter-digitations 50-100 j.Lm wide;
inhalant canals are absent, and ostia lead water almost directly to
Six species have been described from the Mediterranean the chambers. Aquiferous system sylleibid-like, with spherical,
(Sara, 1960b), Indo-West Pacific (Topsent, 1897a; Kirkpatrick, eurypylous choanocyte chambers. Exhalant canals lead to a system
1900b; Levi & Levi, 1989), and Indian Ocean (Thomas, 1970; of basal cavities and then to the oscula, located at the borders of the
Vacelet & Vasseur, 1971) (Table 1). sponge. Proportion of mesohyl to choanocyte chambers varies
from 0.5 : 1 to 1.2: 1 (mean 0.8: 1). Skeleton dense, confused reti-
culation of diods, triods and calthrops in a single size class around
PLAKINA SCHULZE, 1880 the aquiferous system. Monolophose calthrops concentrated at the
surface of the sponge, most with their lophose actines pointing out-
Synonymy ward, with a few dispersed in the choanosome or lining canals and
basal cavities. Lophose and non-ramified spicules may form a
[Achinoe] Gray, 1867a: 546 (unavailable name, see Remarks). dense palisade at the surface, 25-50 j.Lm thick. Diods slender,
Plakina Schulze, 1880: 448. Placina Topsent, 1890d: 231. irregular, sinuous, with actines gradually pointing to sharp ends
Plakoosa de Laubenfels, 1936b: 462 (after Topsent, 1937: 7). (52-93 j.Lm). Actines often irregularly spined, with variable number
and size of spines. The central swelling may be knotty-crooked,
Type species centrotylote or almost smooth. Triods often with one or two ill-
developed or malformed actines. The central swelling, actine size
Plakina monolophn Schulze, 1880: 407 (by original and angle between actines vary widely (actines 11-34 fLm).
designation). Calthrops show the same general shape and variations, and one
actine is usually reduced to a small button (actines 15-31j.Lm).
Definition Monolophose calthrops, 8-31j.Lm long, are irregular, and each
lophose actine have two to six rays. Lophose actines ramify once at
Plakinidae with a spiculation of diods, triods, and calthrops in the middle of their length in two to five slender, cylindrical rays,
a single size class, and with hornolophose calthrops with one, two, which usually show two or three tiny terminal spines (a pattern of
three, or four lophate rays. ramification coded '1m, ts') (Fig. 3E).
Remarks. [Achinoe] Gray, 1867a, with type species Achinoe
Diagnosis australis Gray, 1867a (by monotypy) was erected for two spicule
drawings of an unknown sponge from Western Australia
Thinly to massively encrusting Plakinidae with a spiculation (Freemantle) by Bowerbank (1864: Figs 235-236), showing
of diods, triods, and calthrops, and with homogeneously ramified typical spined calthrops of Homosclerophorida. De Laubenfels
lophocalthrops with one, two, three, or four lophate rays. (1936: 80) associated it with Trikentrion and Cyamon
Candelabra (heterolophose calthrops) absent. Lophocalthrops usu- (Raspailiidae); Hooper & Wiedenmayer (1994) suggested the type
ally concentrated at the sponge surface and bordering canals. species may belong to Corticium; conversely, it is far more likely
Development of the ectosorne is variable, and subectosomal cavi- that it belongs to Plakina in which there are species with profusely
ties may be present (e.g., P. trilophn Schulze, 1880). A large basal spined lophocalthrops. [Achinoe] Gray, 1867a has seniority over
cavity is present in most species. Proportion of mesohyl to cham- Plakina Schulze, 1880, but the name can be suppressed as a nomen
bers varies from 0.7 to 1.8:1. Choanocyte chambers are eurypylous oblitum since it has not been used since. '(ICZN Art. 23.9.1)'
80 Porifera· Demospongiae • Homoscleropborida • Plakinidae

Plakoosa de Laubenfels (type species Plakoosa elisa de Diagnosis

Laubenfels, 1936b: 463, by monotypy; holotype USNM 22237), is a
clear synonym of Plakina, since the spicules called ''microme- Thinly to massively encrusting, sometimes lobate or tubular
soorthotrichotriaenes" by de Laubenfels (1936b) are in fact dilophose Plakinidae, usually tough in consistency and with surface smooth
calthrops similar to those of Plakina dilopha Schulze, 1880 (Topsent, to the eye but rough to the touch. Subectosomal inhalant cavities
1937). Placina Topsent (189Od) is a misspelling of Plakina Schulze. present in some species; choanocyte chambers eurypylous or diplo-
Plakina shares with Corticium the possession of spicules with true dal. Skeleton composed of non-lophose diods, triods, and/or
lophose rays (rays which subdivide close to the base in two to several calthrops with wide size variation, usually in three size classes. The
branches which then might ramify again). Moreover, the type species of small diactines are accumulated on the surface, either forming a
both genera share the presence of monolophose calthrops (although palisade or disposed tangentially to the surface.
with slightly different ramification patterns). Therefore, the genus
Plakina does not have any good synapomorphy of its own, and it is Description of type species
likely to be a paraphyletic group (see also Murley, 1999; Murley et ai.,
1998). However, 10phocalthrops of Plakina are always homogeneously Plakinastrella copiosa Schulze, 1880: 433.
ramified, whereas Corticium also has heterolophose calthrops (autapo- Synonymy. Plakinastrella copiosa Schulze, 1880: 433.
morphic for that genus). Thus, we prefer to consider Plakina as a valid Material examined. Holotype: Not available. Slide
genus until further phylogenetic studies demonstrate more clearly its UFRJPOR 4856 (made by Topsent) from an unregistered specimen
paraphyly and how to divide it in monophyletic subgroups. Species dis- collected from Banyuls, France.
tinction is based on the number of lophose actines in lophocalthrops Description. Schulze (1880) described P. copiosa as cushion-
(1-4) and on details of their patterns of ramification, as well as on exter- shaped, 2cm wide, 5mm high, light yellow in alcohol, with a low
nal morphological features and the architecture of the aquiferous sys- oscular tube at the top of the sponge. Ectosome well developed, with
tem. Plakina tetralophoides Murlcy et ai., 1998 is a nomen novum for regular subdermal cavities. Basal cavities absent. Canal system richly
P. tetralopha Tanita & Hoshino, 1989, required because it became a developed, ramifying like a tree. Skeleton confused, made up of
junior homonym of Corticium tetralophum Hechtel, 1965 after the spicules in three size classes, with transitions: larger spicules diods,
transfer of the latter to Plakina (with the consequent change of its spe- triods and calthrops, of which the first two may appear in the cortex;
cific epithet from tetralophum to tetralopha (Murley et al., 1998). medium-size spicules diactines and triactines only; small spicules
The possible conspecificity of the disjunct, morphologically diods, triods and calthrops restricted to the cortex, with some malfor-
similar populations of the type species, P. monolopha, seems unlikely mations. Candelabra absent. Topsent's (1895) description of the
and awaits genetic, reproductive, and/or cytological studies to be cor- species conforms to that of Schulze (1880), adding certain aspects to
roborated. Studies on the shape and pattern of ramification of lopho- the species description: ''A smooth surface, even or marked by small
calthrops suggest that most records from outside the Mediterranean ridges, covered by microscopic ostia. Choanosome well developed,
basin have been incorrectly identified, with the possible exception of with a rich canal system and eurypylous choanocyte chambers.
Northeastern Atlantic specimens (Murlcy et al., 1998). Endopinacocytes flagellated. Spherulous cells abundant in the
ectosome but also present in the choanosome. Viviparous. Skeleton
Distribution confused, dense, composed by calthrops and derivatives in three size
classes: larger spicules are diods (600-700 X 30-35 fJ.m), triods, and
Twenty-two species of Pia/dna are known worldwide, from the calthrops (both with actines 320 X 30-35 fJ.m); medium-sized
Mediterranean (Schulze, 1880; Sara, 1960b; Pulitzer-Finali, 1983; spicules are diods and triods, but not calthrops; smaller spicules are
Murlcy et al. 1998), Atlantic (Topsent, 1928c), Caribbean (Schmidt, diods (25-40 X 1-2 fJ.m), triods and calthrops (both with actines
1879; de Laubenfels, 1936b; Lehnert & Van Soest, 1998), Indian 8-12fJ.fD long). Larger spicules are concentrated at the choanosome,
Ocean (Vacelet et al., 1976), Pacific (Levi & Levi, 1983b; Tanita & and smaller spicules concentrated at the ectosome and around subder-
Hoshino, 1989; Desqueyroux-Faundez & Van Soest, 1997) and mal cavities." In the specimen from Banyuls spicules are diods and
Antarctic (Topsent, 1901a, 1917; Lendenfeld, 1907) (Table 1). triods in three size classes, supplemented by calthrops in the largest
size class only. Diods I: 310-520 fJ.m; diods IT: 73-130 fJ.m; diods Ill:
11-45 fJ.m; triods I: actines 120-325 fJ.m; triods IT: actines 24-39 fJ.fD;
PLAKINASTRELlA SCHULZE, 1880 triods Ill: actines 12-18 fJ.m; calthrops: actines 80-250 fJ.fD.
Remarks. The synonymy of Dercitopsis and Plakinastrella is
Synonymy justified because the type species of Dercitopsis, D. ceylonica
Dendy, 1905, is very similar to Plakinastrella copiosa in spiculation
Plakinastrella Schulze, 1880: 449. Placinastrella Sollas, and skeletal arrangement (Topsent, 1928c). Placinastrella Sollas
1888: 103. Dercitopsis Dendy, 1905: 65 (after Topsent, 1928c: 33). (1888: 103) is a misspelling of Plakinastrella Schulze. Pachamphilla
Lendenfeld, 1906 was considered a junior synonym of Dercitopsis
Type species by de Laubenfels (1936a: 180), but reexamination of the holotype
of the type species P. allata Lendenfeld, 1907 (M. Maldonado,
Plakinastrella copiosa Schulze, 1880: 433 (by monotypy). pers. comm.) indicates instead a relationship to Penares Gray
(Astrophorida). Species of Plakinastrella possess large calthrops
Definition similar to those of astrophorids such as Calthropella, Pachastrella,
Penares, and Erylus. Species described by Dendy (1905) in
Plakinidae with a skeleton composed of non-lophose diods, Plakinastrella (P. schulzei and P. intermedia) have been found to
triods, and/or calthrops with wide size variation, usually in three belong to the Penares (Topsent, 1928c; Diaz & Van Soest, 1994).
size classes. Plakinastrella shares the presence of spicules in three size classes
Porifera' Demospongiae • Homosclerophorida • Plakinidae 81

with Placinolopha, and the possession of small smooth calthrops and low in frequency. Compressible in life. Aquiferous system
and derivatives with Plakortis, Plakina, Corticium, and intermediate between sylleibid-like and leuconoid, with eurypy-
Placinolopha. We have not studied the nature of the aquiferous sys- lous choanocyte chambers regularly distributed around exhalant
tem of this genus, however, Schulze (1880) described the canal canals. Both ectosomal inhalant cavities and basal exhalant cavities
system of P. copiosa as well-developed, ramifying as a tree, with are present, the latter also being bordered by choanocyte chambers.
subdermal inhalant lacunae but without basal exhalant cavities. Skeleton confused, dense, without ectosomal specialization or dif-
Subdermal cavities are also found in P. clathrata Kirkpatrick and ferentiallocation of spicules. Spicules mostly diods centrotylote or
P. mammilaris Lendenfeld. Dendy (1905) described eurypylous with knobby-knotty centers, somewhat sinuous, 60-150 X 3-6 fLm,
choanocyte chambers in P. ceylonica, and diplodal chambers in and less common, sometimes absent smaller triods 25-50 X
P. mammilaris. Comparisons of aquiferous systems should be 3-6 fLm. Spicules are regularly distributed and densely packed
made among the different species of Plakinastrella and other throughout the body.
plakinid genera to clarify their relationships. Remarks. Roosa de Laubenfels, 1934 (type species
R. zygompha de Laubenfels, 1934: 2; by original designation;
Distribution holotype USNM 22277), was synonymized with Plakortis because
the spiculation of its type species, with diods 50-140/-Lm long
Nine species are known from the Mediterranean (Schulze, and triods 25-50 /-Lm long, is typical of Plakortis species (Topsent,
1880; Maldonado, 1992), Indian Ocean (Dendy, 1905), W Pacific 1937). Placortis Topsent, 1895 is a misspelling of Plakortis
(Kirkpatrick, 1900b; Lendenfeld, 1907; Levi & Levi, 1989), Atlantic Schulze. The original definition of the genus was modified (Diaz &
(Topsent, 1904b, 1927b), and Caribbean (Uliczka, 1929) (Table 1). Van Soest, 1994) to include the possibility of finding calthrops and
diactinal 'microscleres', as complement to diods and triods. Due
to the simplicity of the spiculation of this genus, the species diag-
PLAKORTIS SCHULZE, 1880 nosis is quite difficult without the observation of anatomical char-
acters (e.g., architecture of the aquiferous system) and external
Synonymy characteristics.

Plakortis Schulze, 1880: 449. Placortis Topsent, 1895: 557. Distribution

Roosa de Laubenfels, 1934: 2 (after Topsent, 1937: 7).
Eleven species of Plakortis are known from the Mediterranean
Type species (Schulze, 1880), Atlantic (Pulitzer-Finali, 1993), Caribbean (Carter,
l879b; Wilson, 1902; de Laubenfels, 1934; Zea, 1987), Red Sea
Plakortis simplex Schulze, 1880: 430 (by original designation). and E Africa (Levi, 1953b, 1958; Diaz & Van Soest, 1994),
E Pacific (Desqueyroux-Faundez & Van Soest, 1997), and W Pacific
Definition (de Laubenfels, 1954; Diaz & Van Soest, 1994) (Table 1). From the
literature, it would seem that some Plakortis species have an
Plakinidae with a skeleton formed by small diods with apparent cosmopolitan distribution (i.e., Plakortis simplex,
triods in varying abundance. Diactine-derived 'microscleres' P. angulospiculatus). However biogeographical considerations (iso-
(microrhabds) may be present in some species. lation of the Pacific and Atlantic basins for 3-5 MY; low dispersal
potential of sponges) are at odds with such an hypothesis. The appar-
Diagnosis ent conspecificity of these specimens might be just a consequence of
the simple and non-diagnostic nature of their skeleton and external
Thinly to massively encrusting plakinids with a skeleton morphology (see also Levi, 1953b). The conspecificity of such
mainly formed by small (50-200/-Lm) diods with triods in varying morphologically similar but geographically disjunct populations
abundance. Deformed calthrops can be found in some specimens. awaits more detailed anatomical, reproductive and genetic studies.
Some species have microrhabds (5-20/-Lm) distributed regularly
in the sponge body. Aquiferous system intermediate between
sylleibid-like and leuconoid, with eurypylous choanocyte cham- PSEUDOCORTICIUM BOURY-ESNAULT ET AL., 1995
bers regularly distributed around exhalant canals. Both ectosomal
inhalant cavities and basal exhalant cavities are usually present. Synonymy
Skeleton confused, dense, without ectosomal specialization or
differential location of spicules. Pseudocorticium Boury-Esnault et al., 1995: 28.

Description of type species Type species

Plakortis simplex Schulze, 1880. Pseudocorticium jarrei Boury-Esnault et al., 1995 (by origi-
Synonymy. Plakortis simplex Schulze, 1880: 449. nal designation).
Material examined. Holotype: Not available. Other mate-
rial. USNM 8433, 9433 - Ireland. ZMA paR 7054, 7143, 7149- Definition
Cape Verde Islands, 1-15 m depth. SME 59 - Marseille, France.
Description. Thinly-encrusting sponges, generally 2-5 mm Plakinidae without mineral skeleton, with a well-developed
thick, usually light in color: brown, white, yellow, or tan. Surface ectosome, a leuconoid organization of the aquiferous system, and
smooth and regularly pierced by ostia; oscules < 1 mm diameter diplodal choanocyte chambers.
82 Porifera· Demospongiae • Homoscleropborida • Plakinidae

Diagnosis absent. NW Mediterranean (Marseilles; Boury-Esnault et al.,

1995) (Table 1).
Plakinidae without skeleton, with a well-developed ectosome, Remarks. Pseudocorticium shares the absence of skeleton,
a leuconoid organization of the aquiferous system, and diplodal thin, triangular apopylar cells, and two types of cells of inclusions
choanocyte chambers. Proportion of mesohyl to chambers greater with species of Oscarella, from which it differs by its well devel-
than 2:1. oped cortex, leuconoid aquiferous system and diplodal choanocyte
chambers. The recent proposal to synonymise the families
Description of type species Plakinidae and Oscarellidae (Sole-Cava et al., 1992; Diaz & Van
Soest, 1994; Boury-Esnault et al., 1995) was based on its greater
Pseudocorticiumjarrei Boury-Esnault et al., 1995. similarity to Corticium than to Oscarella in anatomical and exter-
Synonymy. Pseudocorticium jarrei Boury-Esnault et aI., nal morphological characters, supported by allozyme data.
1995: 28.
Material examined. Holotype: MNHN DNBE-94.1 - Jarre Distribution
Island, Marseille, in a cave. Other material. UFRJPOR T-5 and
T-6 - Jarre Island, Marseille, in a cave. This monotypic genus is so far known only from two semi-
Description. Thickly encrusting to lobate sponge, with obscure caves at Marseilles, France (NW Mediterranean) (Table 1;
lobes up to 12cm long by 2cm wide, hanging down from the Sole-Cava et al., 1992; Boury-Esnault et al., 1995; J. Vacelet,
sponge in ceilings of caves. Base irregular, 3-lOcm wide and pers. comm.).
5-30 mm thick. Colour alive cream; in spirit it varies from yellowish
white to brownish gray. Surface smooth and slippery but corru-
gated, folded with irregular depressions. Superficial exhalant ACKNOWLEDGEMENTS
canals (2-5 mm in diameter) lead to circular oscula (5-10 mm in
diameter). Oscula located at the top or on the sides of the lobes, The authors thank Dr. Klaus Rtitzler and Kate Smith from the
5 mm high, surrounded by a slightly transparent oscular rim. Smithonian Institution (Washington, D.C., USA), Dr. Claude Levi
Consistency firm, cartilaginous. Within the lobes there is usually a from the Museum National d'Histoire Naturelle (Paris, France),
central exhalant canal 1-3 mm in diameter. Ectosome with a dense and Claire Valentine from the Natural History Museum (London,
cortex, 50-350 f.1m thick. Aquiferous system leuconoid, with diplo- UK) for the kind loan of specimens. We also thank Dr. MarciaAttias
dal choanocyte chambers (25-60 f.1m diameter). Mesohyl granular, and Noemia Rodrigues from the Laboratorio de Ultraestrutura
with abundant bacteria, collencytes, archaeocytes and cells with Celular Hertha Meyer (Universidade Federal do Rio de Janeiro,
inclusions. Choanocyte chambers and canals surrounded by a clear, Brazil) and Chantal Bezac from the Station Marine d'Endoume
relatively bacteria-free layer of mesohyl (1-6 f.1m thick). A thin (Marseille, France) for their help with S.E.M. Critical reading by
basement membrane underlines both choanoderm and pinacoderm. J.N.A. Hooper and an anonymous reviewer greatly improved the
Proportion of mesohyl to choanocyte chambers in the choano- manuscript. This work was supported by grants and fellowships
some varies from 2.2: I to 2.8: 1. Both spicule and fibre skeleton from CNPQ and FAPERJ (Brazilian government).
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Order Spirophorida Bergquist & Hogg, 1969

Rob W.M. Van Soest1 & John N.A. Hooper2

1 Zoological Museum, University of Amsterdam, P.O. Box 94766, 1090 GT, Amsterdam, Netherlands. (soest@science.uva.nl)
2 Queensland Museum, P.O. Box 3300, South Brisbane, Qld, 4101, Australia. (JohnH@qm.qld.gov.au)

Spirophorida (Demospongiae) is a well-defined small group of non-lithistid tetractinomorph sponges characterized by sigmaspire
microscleres (spined c- or s-shaped spicules of 10-40 /-Lm length). Triaenes and tetractine spicules are shared with sponges of the order
Astrophorida, but are absent in some taxa. Sigmaspires are shared with the lithistid family Scleritodermidae, which points to a probable
close relationship with this family, once again demonstrating polyphyly of the 'order Lithistida'. Spirophorida comprise three families,
Samidae Sollas is monotypic, Spirasigmidae Hallmann with two genera, and Tetillidae Sollas has eight valid genera. Samidae are exca-
vating sponges with amphitriaenes and lacking long-shafted triaenes and lacking oxeas; Spirasigmidae are encrusting to small conical
sponges with only oxea megascleres, lacking triaenes, with a plumose skeleton fanning out towards the periphery, and one genus with a
secondary skeleton of roughened 'microxeas'; Tetillidae are globular sponges with protriaenes as the characteristic megascleres.
Keywords: Porifera; Demospongiae; Spirophorida; Samidae; Spirasigmidae; Tetillidae.
------------------------------------------

DEFINITION, DIAGNOSIS, SCOPE sharing characteristic sigmaspire microscleres. Aside from this
character two of the three families are quite divergent in their habit
Synonymy and further spiculation and they are not considered particularly
closely related. Spirasigmidae is incertae sedis, containing two
Sigmatophora Sollas, 1887, 1888. Spirosclerina Reid, 1963a. genera each with a single species; Tetillidae comprise about 150
Craniellida Reid, 1968d. Spirophoridae Brien, 1968. Spirophorida species, and Samidae only a few.
Bergquist & Hogg, 1969. Spirophorina Wiedenmayer, 1977: 183.
Remarks
Definition
Sollas (1886b, 1888) recognized spigmaspire microscleres as
Tetractinomorpha with triaene megascleres and sigmaspire an important discriminating character and erected a suborder
microscleres. Sigmatophora in his order Choristida for sponges possessing them.
He also erected two families, Tetillidae and Samidae, to acknowl-
Diagnosis edge the gap in structural diversity among the members of
Sigmatophora, with the 'golf-ball' sponges of the family Tetillidae
Globular, occasionally massive, rarely excavating or encrusting and the excavating sponges of the family Samidae. This classifica-
sponges. In non-excavating sponges the structure is strongly radiate, tion found general support (e.g., Wilson, 1925), but was challenged
with thickly developed spicule bundles originating from a focal and modified by Lendenfeld (1903, 1907) and de Laubenfels
point in the middle of the sponge and running to the surface where (1936a), who erected special families and genera for species lack-
they cause a conulose or hispid-bristly surface. At the periphery a ing the sigmaspires. De Laubenfels (1936a) went even so far as to
strongly collagenous cortical region strengthened by special cortical consider that sigmaspire-bearing and sigmaspire-Iacking represen-
oxeas may be developed in some genera. Megascleres include tatives were unrelated and distributed these over different parts of
protriaenes and anatriaenes, occasionally amphitriaenes and plagio- the Choristida, abandoning the (sub-)order Sigmatophora. A con-
triaenes, and large oxeas. Triaenes may be lost in some taxa. trary view was taken by Dendy (1922b), followed by Burton
Microscleres are c- or s-shaped finely spined sigmaspires. In exca- (1934a) and others. These authors assumed homology of sig-
vating sponges (one genus) oxeas, protriaenes and anatriaenes are maspires with the sigmata and derived microscleres of the
lacking, all megascleres are amphitriaenes of a single type. They Poecilosclerida and Haplosclerida. Dendy (1922b) erected a major
are small and insignificant bioeroders. In encrusting or small coni- order Sigmatotetraxonida Dendy, comprising all demosponges lack-
cal sponges (two genera), triaenes are absent, with only large ing asters; Burton (1934a) called this group the Sigmatosclerophora
strongyloxeas forming sequentially radial, plumose and eventually Burton. Subsequent authors eventually reinstated Sollas' group, but
tangential halichondroid skeletal tracts extending from the basal to the name was changed (first to Spirosclerina Reid, 1963a, then to
the peripheral skeleton, and one genus (Spirasigma) has a second- Craniellida Reid, 1968d and eventually to Spirophorida Bergquist &
ary disorganised skeleton of roughened 'microxeas' distributed Hogg, 1969) in order to avoid confusion with sponges possessing
throughout the entire skeleton, overlaying the main skeleton. smooth true sigmas (members of Poecilosclerida and
Haplosclerida). However, recently Gruber (1993), in the spirit of
Scope Dendy and Burton, proposed that Spirophorida, in the sense
employed here, and sigma-bearing Poecilosclerida were considered
The order is a small compact group consisting of three fami- to be a monophyletic group, thus again interpreting sigmaspires and
lies, Tetillidae Sollas, Samidae Sollas and Spirasigmidae Hallmann, true sigmas as homologous. We do not subscribe to this unrealistic

83
84 Porifera· Demospongiae • Spiropborida

point of view, which ignores the structural differences between the The position of Samidae in Spirophorida is controversial,
two orders. The name Spirophorida in the presently employed sense because some authors (e.g., Topsent, 1928c; Wiedenmayer, 1994)
has gained general support and for that reason it is preferred instead consider the amphitriaenes to be microscleres, interpreted as
of the earlier Sigmatophora (the ICZN rules of priority do not apply to amphiasters. In combination with the excavating habit this would
supra-family level taxa). A third family of sponges, Scleritodermidae, point to the family Thoosidae of the order Hadromerida. We prefer
possesses the characteristic sigmaspires, but these sponges have to stress the similarity of the sigmaspires of Tetillidae and
a desma-skeleton. Several recent authors (Gruber, 1993; Hooper & Samidae, and retain the two families within Spirophorida. The
Wiedenmayer, 1994) included Scleritodermidae within the inclusion of a third family, Spirasigmidae, in this order is proposed
Spirophorida, and in view of the likely polyphyletic nature of the here, albeit one of incertae sedis and possibly concerning highly
'order Lithistida' this is understandable. However, in the present aberrant Cinachyrellas.
volume we have taken the conservative approach to retain this
taxon within the 'Lithistida', although we expect that in the near
future most of the families of this order will be distributed over var- Recent reviews
ious orders of the Tetractinomorpha. A fully elaborated proposal
for this is still lacking. Rtitzler, 1987; Hooper & Wiedenmayer, 1994.

KEY TO FAMILIES

(1) Excavating or encrusting on corals ........ ............ .... ................ ................ ........... ..................... ...... ...... ......... ....... ................ ................. 2
Globular or semiglobular sponges, oxeas forming the main spicule type, with protriaenes and other
long-shafted triaenes ........................................................................................................................................................... ... TetiUidae
(2) Excavating in corals, lacking oxeas, possessing peculiar small amphitriaenes as the only megascleres ............................... Samidae
Encrusting or small conical sponges on coral, lacking triaenes, with large strongyloxeas as main megascleres, forming
mainly plumose skeletal structure, almost tangential at the periphery, and one genus with a secondary disorganised
skeleton of microxeas ..... ... ................ .......... ................. ... ............. ... ....... ... ............. ...... .......... ..... ................ ................. Spirasigmidae
Systema Porifera: A Guide to the Classification oj Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademicIPlenum Publishers, New York, 2002

Family Tetillidae Sollas, 1886

Rob W.M. Van Soest1 & Klans Rfitzler

1 Zoological Museum, University of Amsterdam, P.O. Box 94766, 1090 GT, Amsterdam, Netherlands. (soest@science.uva.nl)
2National Museum of Natural History, Smithsonian Institution, Washington D.C. 20560, U.S.A. (ruetzler.klaus@nmnh.si.edu)

Tetillidae Sollas (Demospongiae, Spirophorida) are for the most part globular sponges ('golf ball' sponges) belonging to a small order on
account of their possession of sigmaspire microscleres and triaene megascleres. The characteristic megasclere is the protriaene, which is
found in most members of the family with very few exceptions. In addition there are long oxeas and frequently anatriaenes, rarely also
calthrops or amphitriaenes. The skeletal architecture is strictly radial. Eight valid genera are recognized, differentiated by presence of cor-
tical structures, specialized pore-sieves, and composition of the spicule complement. Tetillids are common and ubiquitous sponges in all
oceans and at all depths.
Keywords: Porifera; Demospongiae; Spirophorida; Tetillidae; Acanthotetilla; Amphitethya; Cinachyra; Cinachyrella; Craniella;
Fangophilina; Paratetilla; Tetilla.

DEFINITION, DIAGNOSIS, SCOPE megascleres). The family comprises more than one hundred
species distributed over all oceans and habitats.
Synonymy
Biology
[Lophurellidae] Gray, 1872a. [Casuladae] Gray, 1872a.
[Tethyina] Carter, 187Sb (not Tethyadae Gray, 1848). Tetillidae Tetillids have a preference for sedimented habitats and some
Sollas, 1886a. Tethyopsillidae Lendenfeld, 1903. Tethydae species possess a root of long spicule bundles to attach them to the
Lendenfeld, 1907. Ectyonillidae Ferrer-Hernandez, 1914a. substrate. Reproductive patterns range from the extrusion of fertilized
Craniellidae de Laubenfels, 1936a. eggs, which fix to the substrate and develop directly, to incubation
of complete young sponges which are then expelled by localised
Definition breakdown of the pinacoderm. No free larvae have yet been
described.
Spirophorida with sigmaspire microscleres and a radiate
skeleton of triaenes and oxeas. Remarks

Diagnosis Gray (1867a) included sponges of this group under the genus
name Tethya in a larger Tethyadae comprising a wide range of
Typically with spherical growth form, often with characteristic tetractinomorph genera. Carter (187Sb) was the first to distinguish
pits called porocalices containing the inhalant and occasionally also a separate group (subfamily) for tetillids, but like Gray called the
exhalant orifices. Cortical region often strengthened by collagen group Tethyina. The explanation for this is that at that time there
fibers and special cortical megascleres, but this region may be thin was still a great confusion over the identity of Tethya. Lamarck
or absent in several genera. Skeleton with tetraxonic and monaxonic (1815) included in his genus Tethya next to T. lyncurium
megascleres (triaenes, huge oxeas) organized in a radiate pattern of (=T. aurantium) and others, Tethya cranium (MUller, 1789 as
spicule bundles, often spiralling outwards from the centre of the Alcyonium). Carter was aware of the great difference between the
body; oxeas, protriaenes and anatriaenes are most common and type of Tethya (Le., T. lyncurium) and T. cranium (see for example
these often protrude from the surface producing a conulose or hairy Carter, 1872a), but he ignored Schmidt's (1870) proposal to
surface. In sponges provided with porocalices they form a palisade employ a separate genus Craniella for Alcyonium cranium.
of long spicules, protruding far beyond the sponge surface, sur- Schmidt (1870) included Craniella in his family Ancorinidae along
rounding the porefields or oscular apertures, which lie in rounded with Stelletta and other astrophorid sponges, but Sollas (1886a)
depressions. Microscleres are unique contorted microspined sig- revived Carter's Tethyina under a different name by erecting the
maspires. Reproduction is oviparous without a larval stage, or family Tetillidae based on Schmidt's (1868) genus Tetilla. In many
viviparous with production of young adults within a parent. ways this was an unwise choice of type genus, inspired by some
notion that Tetilla would represent the most primitive of tetillid
Scope genera. The description of Tetilla euplocamos by Schmidt (1868:
40, pI. V fig. 10) is insufficient to conclude with certainty what its
Out of twenty six nominal genera eight are considered valid, properties were. Selenka (1880: 469) thought to recognize
Acanthotetilla, Amphitethya, Cinachyra, Cinachyrella, Craniella, Schmidt's species in an intertidal habitat in the Bay of Rio de
Fangophilina, Paratetilla, Tetilla. They are differentiated on gross Janeiro, and provided some additional data. Sollas (1888) was able
morphological structure (presence or absence of a cortical region, to verify from a Schmidt slide that at least protriaenes were pres-
presence or absence of specialized pore areas), and spicule comple- ent, but apparently this species has no sigmaspires (cf. below).
ment (possession of calthrops, amphitriaenes or other auxiliary Even though protriaenes are of common occurrence in tetillids,

85
86 Porifera· Demospongiae • Spiropborida • Tetillidae

they are by no means confined to that group and we have to rely and Tethyopsillidae was incorrect, since the type of Tetillidae and
on the subtle tetilliform protriaene-type as proof that Tetilla genus Tetilla (T. euplocamos) has no sigmaspires and thus would
euplocamos is a member of what is generally understood as Tetilla belong in Tethyopsillidae, making this family an objective synonym
and the Tetillidae. In summary: we have a type species of the type of Tetillidae. De Laubenfels (1936a) for the same reason refused to
genus of a family lacking the major synapomorphy of the order to accept Tetilla euplocamos as a close relative of Craniella, and
which the family belongs! Some confidence may be obtained from erected a separate family Craniellidae for the genera possessing sig-
the fact that Tetilla species lacking sigmaspires are not uncommon. maspires. Tetilla and Tetillidae were defined as 'Choristida lacking
Despite this dubious type species designation, Sollas' (1888) mono- microscleres' and an unrelated group of genera was assigned to it.
graphic treatment of Tetillidae was followed by most subsequent Needless to say that neither scheme gained general support.
authors with the exception of Lendenfeld (1888, 1903, 1907) and de
Laubenfels (1936a). Lendenfeld continued to employ Tethya for Previous reviews
sponges answering to the definition of Craniella, but at least kept
this in the family Tetillidae. He erected a family Tethyopsillidae for Sollas (1888), Lendenfeld (1903), Wilson (1925), Topsent
sponges without sigmaspires. Lendenfeld's division into Tetillidae (1928c), Levi (1973), Riitzler (1987).

KEY TO GENERA

(1) Specialized pore-bearing pits ('porocalices') present ......................................................................................................................... 2

No porocalices ..................................................................................................................................................................................... 6
(2) Spicules include medium-sized heavily spined oxeas ('megacanthoxeas') ................................................................... Acanthotetilla
No megacanthoxeas ............................................................................................................................................................................. 3
(3) Among the megascleres there are short-shafted triaenes with equal-sized rays (calthrops). N.B. these may have bifid cladi but are not
to be confused with long-shafted amphiclads ...................................................................................................................... Paratetilla
No calthrops, all triaenes - if present - are long-shafted .................................................................................................................... 4
(4) In cross section, there is a clear cortical region with special cortical megascleres (short oxeas) and collagenous
reinforcement ........ .... ... .... ...... ......... ........ .................... ....... ... .... ... ... ... ....... ... .... ...... ...... ...... ... ....... ... ... .... ... ... .... ............ ... ..... Cinachyra
In cross section there is no discernible peripheral layer different from that of the interior ........................................... ,... .......... ..... 5
(5) Porocalices all similar, undifferentiated morphologically ................................................................................................ Cinachyrella
Porocalices differentiated into a morphologically distinct inhalant and exhalant type ................................................... Fangophilina
(6) Stalked sponges with differently structured stalk and main body; among the megascleres of the stalk there are triaenes with cladi on
both ends (amphitriaenes, -diaenes, -monaenes, collectively called amphiclads) ............................................................ Amphitethya
No amphiclads, if stalked, then it is a thin root-like projection, not a proper stiff stalk ..................................................................... 7
(7) In cross section, there is a clear cortical region with collageneous reinforcement ................................................................ Craniella
In cross section there is no discernible peripheral layer different from that of the interior .. .... ... ...... .......... ................ ..... .......... Tetilla

ACANTHOTETILLA BURTON, 1959 Previous review

Synonymy Van Soest (1977b).

Acanthotetilla Burton, 1959a: 201. Acanthocinachyra Levi, Description of type species

1964b: 386.
Acanthotetilla hemisphaerica Burton, 1959a (Fig. lA-B)
Type species Synonymy. Acanthotetilla hemisphaerica Burton, 1959a: 201,
fig. 5; Van Soest, 1977b: 2, pI. I figs a-b, pI. II figs c-d, text-fig. 1.
Acanthotetilla hemisphaerica Burton, 1959a: 201 (by Material examined. Holotype: BMNH 1936:3:4:530 - John
monotypy). Murray Exped. stat. 45, coast of SArabia, litotharnnion bottom,
38m depth.
Definition Description. Semiglobular (Fig. 1A), 4-5 cm diameter. Colour
in alcohol yellowish brown. Surface hispid-bristly due to megascle-
Tetillidae with megacanthoxeas as auxiliary megascleres. res projecting 1 mm or more beyond the ectosome. Porocalices
numerous, 1-2mm diameter, scattered over the surface. No appar-
Diagnosis ent oscules. Consistency hard, incompressible. Skeleton radiate,
forming a tight palisade at the surface. The main megascleres are
Globular, hemispherical or irregularly massive sponges. smooth long oxeas and medium-sized megacanthoxeas, but protri-
Surface extremely hispid and provided with numerous small-sized aenes and rare anatriaenes are also present. Interiorly, the skeleton
porocalices. Skeleton radiating bundles of oxeas, protriaenes, anais less dense, making the choanosome cavernous, with megacan-
triaenes and megacanthoxeas, developed into an impenetrable pal- thoxeas and sigmas pires as the main spicule categories present.
isade at the periphery. Internally cavernous with megacanthoxeas Spicules. Protriaenes, simple, straight, relatively rare, shaft
and sigmas pires as the main spiCUle complement. Four species are 1000-2520 X 6--14/lm, cladi 30-56/lm; anatriaenes even rarer,
known from the Western Indian Ocean and the Caribbean, depth shaft 1250-1600 X 10 /lm, cladi 55-80/lm; oxeas, forming the
range 38-100 m. bulk of the radiating skeleton, straight, 3000-4400 X 25-40 /lm;
Porifera' Demospongiae • Spiropborida • Tetillidae 87

Fig.!. Acanthotetilla spp. A-B, Acanthotetilla hemisphaerica Burton, 1959a. A, holotype BMNH 1936.3.4.530 (scale 1 em). B, megacanthoxea ofholotype
(scale 50 ILm). C, Acanthotetilla enigmatica (Levi, 1964b as Acanthocinachyra), megacanthoxea (scale 50 ILm). D-H, Acanthotetilla gorgonosclera Van Soest,
1977b. D, oxea (scale 10 ILm). E, protriaene (scale 10 ILm). P, megacanthoxea (scale 50 ILm). G, detail of same (scale 10 ILm). H, sigmaspire (scale 1ILm).

megacanthoxeas (Fig. lB), thickly spined, spines arranged in irreg- 1977b: 7 was described from Barbados (see Fig. 2D-H). It has char-
ular 'whorls', with points directed towards the centre, 325-414 X acters in between A. seychellensis and A. hemisphaerica, and in
40-60/-Lm, with 20--25 whorls of spines. Juvenile growth stages of view of its Atlantic occurrence is also considered a valid species.
the megacanthoxeas are centrotylote smooth oxeas. Distribution
and ecology. Only recorded from the type locality.
Remarks. The structure of the skeleton is reminiscent of a AMPHITETHYA LENDENFELD, 1907
cortical specialization (impenetrable palisade of oxeas and mega-
canthoxeas at the surface), but no organic fibrous cortical region has Synonymy
been described so far, so this remains undecided. Burton (1959a)
gave a faulty description of A. hemisphaerica by expressly stating Amphitethya Lendenfeld, 1907: 126.
that sigmaspires were absent. This induced Levi (1964b) to erect a
genus Acanthocinachyra for material very similar to Burton's spec- Type species
imen, but possessing numerous microscleres. The type species (by
monotypy) Acanthocinachyra enigmatica Levi, 1964b: 386, fig. 2, Amphitethya microsigma Lendenfeld, 1907: 126 (by subsequent
from Inhaca island, Mozambique (material from MNHN re- designation; Riitzler, 1987, confirmed by Hooper & Wiedenmayer,
examined, megacanthoxea figured in Fig. 2C) shares most features 1994: 429).
with A. hemisphaerica and since Burton's species (even his own
preparations) contain numerous sigmaspires, the two genera are Definition
obvious synonyms. The two type species were kept as separate
species by Van Soest (1977b), but additional material may bridge Tetillidae with long-shafted amphiclad triaenes and
the few discrepancies and it is predicted that they both belong to the plagiotriaenes.
same species. A third species from the Western Indian Ocean,
Acanthocinachyra seychellensis Thomas, 1973: 80, has unusually Diagnosis
thin and curved megacanthoxeas as well as much smaller smooth
oxeas, and may indeed be a valid separate species of Acanthotetilla. Globular-stalked sponges with conulose surface, lacking
The fourth species, Acanthotetilla gorgonosclera Van Soest, porocalices. Small scattered oscules. Skeleton of the main body
88 Porifera' Demospongiae • Spiropborida • Tetillidae

'.1
TIJ
( It
:rr1'

Fig. 2. Amphitethya microsigma Lendenfeld, 1907. A-D, habit and spicules reproduced from Lendenfeld, pI. XV fig. 19-3. A, habit (scale 1 cm).
B, spicules. C, detail of amphitriaene. D, detail of sigmaspire (sizes see text). E-H, photos of spicules made from slides of one of the types kept in BMNH.
E, cross section of peripheral region of peduncle (scale 500 !Lm). F, plagiotriaene (scale 100 !Lm). G-H, amphitriaenes (scale 100 !Lm).

radiate, consisting of oxeas and normal triaenes, and that of the of Western Australia, 82-110m depth. BMNH 1908.2.9.33-35
stalk longitudinally arranged, consisting of all megascleres includ- (not seen) - 3 slides of one of the syntypes, labelled "Valdivia &
ing amphiclad triaenes. Cortical region present in stalk and main Gazelle Lendenfeld".
body. Megascleres oxeas, protriaenes, anatriaenes, plagiotriaenes Description (summary of Lendenfeld's 1907 extensive
and long-shafted amphiclad triaenes. Microscleres sigmaspires. description). Stipitate sponge (Fig. 2A). Main body globular, up to
1\vo species, both from Australian waters. 4.2 cm in diameter, surface conulose, optically smooth between
conules. Stalk up to 7 cm long, 1.2 cm in thickness, circular in cross
section, surface also slightly conulose. Colour in alcohol is coffee-
Recent review brown. In cross section, both the main body and the stalk show
a distinct cortical layer of about 250 J.Lm thick, in which only
RUtzler (1987). microscleres and auxiliary megascleres are found. The amphiclad
megascleres are confined to the stalk and are found in the subecto-
Description of type species somal layer (Fig. 2E). The main skeleton consists of oxeas, packed
longitudinally in the stalk and in radiating bundles in the main
Amphitethya microsigma Lendenfeld, 1907 (Fig. 2A-H). body, where they follow a spiral course as is found in many other
Synonymy. Amphitethya microsigma Lendenfeld, 1907: tetillids. Megascleres (Fig. 2B), protriaenes, cladome regular, with
126, pI. XV figs 19-39. fusiform shaft, 3000-5000 X 10 J.Lm; anatriaenes, confined to the
Material examined. Syntypes: 2MB (not seen) - collected peripheral region of the stalk, 5000-7000 X 10 J.Lm, cladi 15 J.Lm;
by the 'Gazelle' Exped., Dirk Hartog Island, off the NW coast plagiotriaenes (Fig. 2F), confined to the peripheral region of the
Porifera· Demospongiae • Spiropborida • Tetillidae 89

stalk (Fig. 2E), straight shaft, rounded at the end, 320-1100 X they are mixed with protriaenes. These protrude beyond the
13-30 f.Lm, cladi 50-250 f.Lm; amphiclad triaenes (Fig. 2C, G-H), layer of cortical oxeas and are the cause of the bristly surface.
confined to the peripheral region of the stalk, shaft 160-540 X Megascleres choanosomal large oxeas, cortical small oxeas,
14-30f.Lm, cladome usually irregular with cladi of l00-150f.Lm, protriaenes, and anatriaenes. Microscleres sigmaspires. So far, only
mostly with different lengths, or occasionally absent (diaene and a single valid species has been recognized with certainty to belong
monaene variations are not uncommon); oxeas, straight, or more to this genus, most records of Cinachyra species concern the genus
often curved, smooth, gradually but sharply pointed, 6000-8000 X Cinachyrella (cf. below).
20-60f.Lm. Microscleres sigmaspires (Fig. 2B,D), 10-12 X 0.5-0.8
f.Lm. Distribution and ecology. Dirk Hartog Island, off the coast of Previous reviews
NW Australia, 82-110m.
Remarks. In view of the similarity and the structural position Wilson (1925), Rutzler (1987).
they share, the amphiclad triaenes and the plagiotriaenes are very
likely to be considered as the same spicule type, in which the Description of type species
amphiclad condition mayor may not have developed. Lendenfeld
assigned to his genus Amphitethya a second tetillid species with Cinachyra barbata Sollas, 1886a (Fig. 3A-C).
amphiclad triaenes, Tethya stipitata Carter, 1886c: 460 (redescribed Synonymy. Cinochyra barbata Sollas, 1886a: 183; Cinachyra
by Sollas (1888: 49) as Tetilla?). This differs in details of spicule barbata Sollas, 1888: 23, pIs ill, XXXIX.
sizes from A. microsigma, but is otherwise similar and is an Material examined. Syntypes (not seen): BMNH
obvious Amphitethya. The fact that both species assigned to 1889.1.1.14-19, 107-108 (slides 1894.11.16.37-51) - the type
Amphitethya are prominently stalked sponges is not emphasized series consists of more than 60 syntypes all dredged from the
here, as root-organs of other tetillids (e.g., Tetilla euplocamos or shores of Balfour Bay, Kerguelen.
Cinachyra barbata) may be considered homologous to a stalk. A Description (from Sollas, 1888). Globular sponges (Fig. 3A),
further species assigned by Lendenfeld, viz., Tetilla bacca Selenka, seated on a dense spicular basal mass; ash-grey in alcohol. Oscules
1867, is considered a member of Paratetilla. Its auxiliary spicules inconspicous, scattered. Porocalices large and numerous, up to
are calthrops-like, not amphiclad, although the cladi may be occa- 5mm in diameter, scattered over the lateral surfaces, but relatively
sionally bifid. Likewise, assignments of Tetilla merguiensis sensu rare on top, and lacking from the basal mass. The porocalices are
Topsent, 1897a and Paratetilla aruensis Hentschel, 1912 to flask-shaped, penetrating deep into the interior. Internally, they are
Amphitethya by Wilson (1925) are incorrect. The differences are smooth-walled and -rimmed, and have a poresieve at the bottom.
perceived such that the Amphitethya amphitriaene is a derivate of a Surface hispid-bristly, with megascleres protruding far beyond the
long-shafted triaene (plagiotriaene), whereas the occasional bifid ectosome. Size of main body up to 7 X 8.5 X 7 cm, size of basal
Paratetilla auxiliary megasclere derives from equal-rayed mass up to 5 X 10 X7 cm. Basal mass apparently a fused root sys-
calthrops-like spicules. tem which gradually expands during growth of an individual, it is
absent in the smallest collected specimen. In cross section (Fig. 3B)
the cortical region is visible as a dense peripheral layer, up to
CINACHYRA SOLLAS, 1886 1.75 mm in thickness, its whitish colour showing off from the yel-
lowish choanosomal interior. The cortex consists of a distinct
Synonymy organic collagenous tissue, reinforced by special cortical oxeas
strewn at all angles and microscleres. The choanosomal skeleton
[Cinochyra] Sollas, 1886a: 183 (nomen corrigendum); consists of bundles of oxeas issuing from an eccentrically located
Cinachyra Sollas, 1888: 23. focus and following a spiral course towards the periphery. The
peripheral bundles contain a mixture of oxeas and triaenes, with the
Type species latter protruding far beyond the surface. Megascleres (Fig. 3C).
Protriaenes, occasionally prodiaenes, in two distinct size categories,
Cinochyra barbata Sollas, 1886a: 183 (by monotypy). large structural ones, fusiform, shaft up to 13,000 X 30 f.Lm, cladi up
to 180X 16f.Lm, and small, sinuous, hair-like, up to 130X4f.Lm,
Definition with cladi 16-30 f.Lm; anatriaenes, occurring only in the basal mass,
where they function as anchoring spicules, up to 40,000 X 24 f.Lm,
Tetillidae with cortex reinforced by auxiliary oxeas, with cladi up to 215 X 28 f.Lm; oxeas, in two distinct size categories, large
flask-shaped porocalices. choanosomal ones, sharply pointed, straight or curved, up to
8000 X 70 f.Lm; small cortical oxeas, fusiform, bluntly pointed, up to
Diagnosis 900 X 36 f.Lm. Micmscleres sigmaspires, 12-16 f.Lm. Distribution.
Kerguelen, Patagonia, Antarctica, sediment-rich bottoms, 45-549 m.
Globular sponges with surface covered by numerous porocal- Remarks. The genus name has been widely used by many
ices. Full-grown specimens develop an irregular basal mass as authors for sponges now recognized to belong in a separate genus
wide as the globular upper part, equivalent perhaps to a stalk. Cinachyrella Wilson, 1925 (cf. below). This shares the porocalices,
Porocalices large and deep, flask-shaped, surrounded by a fringe of but lacks any cortical specialization. SoUas (1888) maintains that
long megascleres. In cross section there is a prominent cortex some of the porocalices are exhalant, whereas others are inhalant.
strengthened by special cortical oxeas. Skeleton of the globular However, Kirkpatrick (1905) found small, simple oscules separate
part radiate, consisting of bundles of oxeas originating in a central from the complicated porocalices, and similar findings are reported
nucleus and spirally curving outwards toward the periphery, where by Boury-Esnault & Van Beveren (1982) and Rutzler (1987).
90 Porifera· Demospongiae • Spirophorida • Tetillidae

c
Fig. 3. Cinachyra barbata Sollas, 1888, drawing of habit (A), peripheral skeleton in cross section (B), and spicules (C), reproduced from Sollas' pI. III)
(scales see text).

CINACHYRELLA WILSON, 1925 Synonymy. Tetilla hirsuta Dendy, 1889: 75; Cinachyra
hirsuta; Lendenfeld, 1903: 28; Tetilla (Cinachyrella) hirsuta;
Synonymy Wilson, 1925: 363.
Material examined. Lectotype (here designated) (not seen):
[Psetalia] Gray, 1873c: 234 (nomen oblitum). [Labaria] Gray, BMNH 1889.1.21.22 - (the larger of the two specimens),
1873c: 235 (nomen oblitum). Cinachyrella Wilson, 1925: 363. Rameswaram Island, Gulf of Manaar, Sri Lanka, Thurston collec-
Raphidotethya Burton, 1934a: 526. Uliczka de Laubenfels, 1936a: tion. Paralectotype (not seen): BMNH 1925.11.1.769 - the smaller
174. specimen from the same locality.
Description (from Dendy, 1889). Globular sponge, size up
Type species to 5 cm in diameter, colour dark grey in alcohol. Surface 'hirsute'
due to projecting spicules. Porocalices scattered irregularly over
Tetilla hirsuta Dendy, 1889: 75 (by subsequent designation; the body. Some porocalices are hemispherical and shallow, others
Riitz1er, 1987). deep and tubular. The floor of the porocalices is alternatively perfo-
rated by a number of small pores, or some oscu1ar tubes, or is
Dermition devoid of openings. Dendy assumed the porocalices were either
inhalant or exhalant, but these are not morphologically differenti-
Tetillidae with undifferentiated porocalices, without cortex, ated. In cross section there is no special cortical skeleton. The
without auxiliary megascleres. choanosomal skeleton consists of stout radiating fibers issuing
from a dense central nucleus. Megascleres. Protriaenes, often
Diagnosis of hair-like dimensions, i.e., comparatively thin and short, up to
460 x 14 fLm, cladi 50 X 7 fLm; anatriaenes similar in dimension,
Globular sponges with hispid-bristly surface provided with slightly shorter; oxeas, fusiform, sharply pointed, up to 3500 X
numerous scattered porocalices; oscu1es inconspicuous. Skeleton 42 fLm. Microscleres the usual sigmaspires, up to 22 fLm.
radiate, consisting of bundles of oxeas issuing from the centre Choanocyte chambers 20 fLm in diameter. Distribution. Known
and running spirally to the surface. No cortical specialization. only from the type locality.
Megasc1eres protriaenes, anatriaenes, occasionally plagiotriaenes, Remarks. No images of the type species are currently
and dominant large oxeas. Microscleres sigmas pires, which may be available, but it is likely that it is closely similar to e.g., Cina-
lacking occasionally, finely spined oxeas and/or raphides occur. chyrella apion (Uliczka, 1929 as Cinachyra), as described and
More than one hundred species distributed in shallow water of illustrated by Riitzler & Smith (1992). Their illustrations are
tropical and subtropical seas. here reproduced in Fig. 4 (habit and surface characteristics) and
Fig. 5A (spicules).
Recent review Burton (1934a) attempted to revise a large amount of
Cinachyrella species (as Cinachyra), arriving at the remarkable con-
Riitzler (1987). clusion that most tropical representatives are members of a single
intertropical species to be called Cinachyra australiensis (Carter,
Description of type species 1886b). Needless to say that such a conclusion is unacceptable.
The genus Psetalia Gray, 1873c: 234 was erected for type
Cinachyrella hirsuta (Dendy, 1889). species P. globulosa Gray, 1873c: 234 (by monotypy). There are
Porifera· Demospongiae • Spiropborida • Tetillidae 91

Fig.4. Cinachyrella apion (Uliczka, 1929), habit and surface characteristics, reproduced from Riitzler & Smith (1992: fig. 6). A, specimen in situ attached
to alga, showing numerous buds. B, alcohol-preserved specimen. C, close-up of buds. D, close-up of porocalyx. E, close-up of oscule.

apparently four specimens of this species from Singapore. These are recognizable. The main genus character is the absence of triaenes,
globular sponges of up to 6 cm diameter, covered by prominent instead of which there are only oxeas as megascleres, arranged in
"tubercles each containing a tuft of filamentous spicules". A deep the usual radiating bundles. At the surface and throughout the
concavity is found at the upper surface. It seems clear that this is choanosome there are oxeas of 730 X 12 f.Lm about half the length
a description of porocalyces, and as its type locality is in the tropics, of the choanosomal main oxeas which measure 1400 X 24 f.Lm.
it seems a safe conclusion that it concerns a species of Cinachyrella. Microscleres are sigmaspires apparently in two size categories,
The name Psetalia is an unused name in the sense of ICZN Article 24 and 9 /Lm, and raphides also in two size categories, 30 and
23.9 (not used after 1899) and accordingly it is proposed to suppress 15 f.Lm. Despite the fact that no porocalices were described, this
it in favour of the prevailing junior name Cinachyrella. genus is considered a junior synonym of Cinachyrelia, as it has no
The genus Labaria Gray, 1873c: 235 was erected for type conulose surface, is hispid, and lacks a cortex. The absence of tri-
species Labaria hemisphaerica Gray, 1873c: 235 (by monotypy). aenes and the possession of spicule size categories is not of generic
Like Psetalia, this hemispherical sponge of 5 cm diameter appears value, as there is a large diversity in these features among the many
to be a Cinachyrella, as Gray described structures on the surface - Cinachyrella species. Moreover, Hooper (pers. comm.) has
" ... cylindrical perforations, from the centre of which are emitted observed specimens possessing triaenes with their cladomes stick-
tufts of elongated filiform spicules ... " - which can only be ing some distance beyond the surface. This may also have been the
porocalices. The name Labaria is an unused name in the sense of case for the type specimen.
ICZN Article 23.9 (not used after 1899) and accordingly it is pro- The genus Uliczka de Laubenfels, 1936a: 174 was erected
posed to suppress it in favour of the prevailing junior name for type species Cinachyra schistospiculosa Uliczka, 1929: 45,
Cinachyrella. The name is in any case preoccupied by Labaria figs 27-30 from Barbados (by original designation). This species
Carter, 1873c (Hexactinellida, Pheronematidae). is a commonplace Cinachyrella, assigned to the synonymy of
The genus Raphidotethya Burton, 1934a was erected for type Cinachyrella kuekenthali (Uliczka, 1929 as Cinachyra) by Riitzler &
species (by monotypy) Raphidotethya enigmatica Burton, 1934a: Smith, 1992: 154. It possesses occasional 'split' oxeas, but these are
526, fig. 2 (here reproduced as Fig. 5B). The holotype (BMNH obvious artefacts, and certainly not a specific or generic character
1930.8.3.27; not examined) is described as a ficiform sponge of (see also Van Soest & Sass, 1981 and Riitzler & Smith, 1992).
4.5 X 2 cm, with a thick stalk and a uniformly hispid surface in Wilson (1925) erected Cinachyrella as a subgenus of Tetilla,
which no pores or oscules were visible. No cortical specialization rather than of Cinachyra, thus apparently assuming the lack of
92 Porifera' Demospongiae • Spirophorida • Tetillidae

.~
. :1

c .." y ,

" rf\f
a

IMr_ 600 IJm

--;0;;;:;;-
a.
c -9
b
vc;; " t
~l

' dI
h
100 ~m

A B
Fig. 5. A, Cinachyrella apion (Uliczka, 1929), drawing of spicules, reproduced from Rtitzler & Smith (1992: fig. 7). B, Cinachyrella enigmatica (Burton,
1934a as Raphidotethya), drawing of cross section and spicules reproduced from Burton's fig. 2 (scales see text).

a cortex is a more important feature than the presence of porocalices. CRAN/ELLA SCHMIDT, 1870
The presence or absence of a cortical specialization is of dubious
phylogenetic significance, as it occurs both in tetillids with poro- Synonymy
calices (Cinachyra and Cinachyrella) and in those without
(Craniella and Tetilla). Moreover, the cortical specialization is also Craniella Schmidt, 1870: 66. Tethya of authors (e.g., Carter,
used as a discriminatory character in other tetractinomorph groups. 1872a; Lendenfeld, 1903) (not Tethya Lamarck, 1815). Polyurella
Cortical specialization is rather easily detected when the cortex is Gray, 1870b: 312. [Lophurella] Gray, 1872a: 471 (invalid name).
thick and strengthened by special cortical megascleres, but remains [Lophiurella] Gray, 1873c: 234 (lapsus). Tethyopsilla Lendenfeld,
difficult to demonstrate when it is thin and cortical spicules are 1888: 45. Craniellopsis Topsent, 1913a: 14.
lacking. In the case of Cinachyra, the cortex is well-developed, but
so far only a single species is known, whereas the bulk of the Type species
species described as Cinachyra are devoid of a cortex apparently,
and need to be assigned to Cinachyrella. We maintain here the dis- Craniella tethyoides Schmidt, 1870: 66 (by subsequent desig-
tinction between the two, because Cinachyra barbata differs also nation; de Laubenfels, 1936a: 175).
quite strongly from Cinachyrella hirsuta in spicule sizes and cate-
gories. A further complication for the use of Cinachyrella is the Definition
relationship with Fangophilina Schmidt (cf. below). This is main-
tained as a separate genus of Tetillidae on the strength of the occur- Tetillidae without porocalices, with a distinct cortex strength-
rence of clearly morphologically differentiated oscular and ostial ened by special cortical oxeas.
porocalices. Still, Dendy (1889) examined the bottom of several
porocalices in C. hirsuta and distinguished oscular and poral types. Diagnosis
Wilson (1925) described special simple oscules from the
Philippine specimen he identified as C. hirsuta, and thus casted Globular sponges with conulose but optically smooth surface
some doubt even though he acknowledged the possibility that over most of the upper body; at the base there are bundles of
Dendy may have been right. It is also not quite certain that Dendy's spicules acting as a root. Oscules few, usually on the top. Ostia in
and Wilson's material was conspecific, because there is a large size sieve-like groups overlying subdermal cavities. In cross section
discrepancy in the protriaenes recorded from the Indian and there is a distinctly visible cortical layer, which is perhaps divisible
Philippine specimens. Later on, Riitzler (1987) expressed strong in a tightly collagenous layer strengthened by radially or confus-
doubts about the reality of the specialization of differentiated oscu- edly arranged special cortical megascleres, and a less dense outer
lar and ostial porocalices, maintaining that all porocalices are layer in which there are many subdermal cavities. Choanosomal
inhalant and that separate simple oscules remain usually unde- skeleton radiating in spiral fashion, with bundles of oxeas originat-
tected in most Cinachyra and Cinachyrella. If Riitzler is right and ing from a central focus and spiralling outwards towards the sur-
the differentiation will be demonstrated to be an artifact, then face where they are mixed with protriaenes to protrude in groups
Fangophilina would be a synonym of Cinachyrella. and push up the ectosome into conical elevations. Megascleres,
Porifera· Demospongiae • Spirophorida • Tetillidae 93

A B

Fig. 6. Craniella tethyoides Schmidt, 1870. A, BMNH slide of lectotype. B-D, photos of spicules from lectotype (scales 50j..Lm). B, sigmaspires.
C, protriaene. D, anatriaenes.

protriaenes, anatriaenes, choanosomal oxeas, shorter cortical is sufficient to recognize the genus as valid and of common occur-
oxeas. Microscleres, sigmaspires (may be lost not infrequently). rence. The genus is similar in many aspects to Tetilla, but this
About 25 species are recorded from most seas and oceans, often genus lacks a cortical specialization.
from somewhat deeper localities. The genus Polyurella Gray, 1870b: 312 was erected for type
species Polyurella schmidtii Gray, 1870b: 312 (by monotypy).
Previous reviews Gray (1870b) admits himself that this is the same (Le., a junior
synonym) as Tetilla (spelled as Tetella (sic» polyura Schmidt,
Sollas, 1888; Topsent, 1894d; Wilson, 1925. 1870: 66, pI. VI fig. 8. This species is here assigned to Craniella on
account of the presence of a cortical skeleton of smaller oxeas
Description of type species (based a.o. on subsequently assigned specimens in the collection of
ZMA). Gray's (1870b) article was published after the appearance
Craniella tethyoides Schmidt, 1870 (Fig. 6A-D). of Schmidt's monograph, so Polyurella is a clear junior synonym
Synonymy. Craniella tethyoides Schmidt, 1870: 66, of Craniella Schmidt, 1870.
pI. VI fig. 9; Sollas, 1888: 54; Tetilla tethyoides; Lendenfeld, The genus [Lophurella] Gray, 1872a: 461 was erected for type
1903: 25. species Tetilla lophura "Schmidt, table" (Gray's notation). There is
Material examined. Lectotype (here designated): BMNH no record of Tetilla lophura in any of Schmidt's papers; and it is
1939.2.10.30 (not seen), slide examined BMNH 1870.5.3.54 - also not mentioned in Desqueyroux-Faundez & Stone (1992). We
labeled in Schmidt's handwriting "Craniella tethyoides n.g. (116), must assume this is a misspelling, most likely of Tetilla polyura,
Florida 100-230 fathoms" (here reproduced in Fig. 6A), presum- since the definition of the family [Lophurellidae] Gray, 1872a: 460
able made from the lectotype which was apparently obtained in an to which this genus is assigned mentions the characteristic numer-
exchange with MCZ. The latter museum is presumably holding ous rooting spicules, used by Schmidt to characterize Tetilla
a further specimen (here designated paralectotype). Other material. polyura. This is confirmed also by Gray (1873c: 234), where he
BMNH 1870.5.3.55 (slide) - Iceland, mentioned in Schmidt's indicates subsequently that Tetilla polyura is the type of his genus
report, and not considered part of the type series as it appears to be [Lophiurella] (sic!). Even so, [Lophurella] is not an available genus
from a different species. Two further slides from Florida in BMNH name as it cannot be unambiguously assigned to a nominal species-
bearing this name on the labels are likewise not part of the type group taxon (ICZN Art. 12.2.5), which rules out type species
material. assignment by indication (pers. comm. by Dr Alice Wells of the
Description (from Schmidt, 1870, and Sollas, 1888). Australian Biological Resources Survey, Canberra). Likewise,
Globular, with conulose surface (where surface membrane is still [Lophurellidae] Gray, 1872a is unavailable as a family name as it is
intact). Distinct fibrous cortex strengthened by cortical oxeas based on an unavailable genus name.
arranged radially. Choanosomal oxeas robust, forming the bulk of The genus Tethyopsilla Lendenfeld, 1888 was erected (by
the megascleres and protruding beyond the surface. Megascleres. monotypy) for type species Tethyopsilla stewarti Lendenfe1d, 1888:
Protriaenes (Fig. 6C) present (no size recorded), anatriaenes 45, to accomodate species lacking sigmaspire microscleres. The
(Fig. 6D) with cladi of 70-90 fLm, cladome 80-110fLm in diameter type (which was recently found to be missing from the
(no length recorded); oxeas in two distinct size categories (no sizes 2MB, and also not located elsewhere, cf. Hooper & Wiedenmayer,
recorded). Microscleres (Fig. 6B), sigmaspires up to 35 fLm. 1994: 431) is described as a small (2cm) spherical sponge, with
Distribution. Florida, down to 400 m. smooth surface and slit-like oscules. There are abundant protri-
Remarks. The type species remains ill-known and awaits aenes, prodiaenes and promonaenes (up to 2000 X lOfLm), anatri-
proper redescription from type and fresh material. Nevertheless, the aenes (no size quoted) and oxeas (1500X lOfLm), no microscleres.
combination of the lack of porocalices and a cortical specialization The genus was made the type of a family Tethyopsillidae apparently
94 Porifera· Demospongiae • Spiropborida • Tetillidae

originally founded on the possession of irregular protriaenes, but Description of type species
later (Lendenfeld, 1903) on the absence of sigmaspires. The poly-
mastiid genus Proteleia Dendy & Ridley, 1886 was also included Fangophilina submersa Schmidt, 1880b (Fig. 7A).
in this nominal family. The absence or rarity of sigmaspires is Synonymy. Fangophilina submersa Schmidt, 1880b: 73,
rather a common phenomenon in unrelated tetillids and not worthy pI. X fig. 3; Topsent, 1920a: 2; Cinachyra submersa; Lendenfeld,
of genus or family distinction. Wilson (1925) assigned Tethyopsilla 1903: 28.
to Tetilla, but Lendenfeld (1903) expressly mentions a cortex of Material examined. Lectotype (here designated) (not seen):
1.2mm thickness (admittedly without special cortical oxeas), so it ZMUS P0160 - dry, Caribbean. Paralectotypes: ZMUS P.0166 -
seems more appropriate to assign the genus to Craniella. Hooper & dry fragments. Fragments of types: 2MB 6650 (6 slides) - fide
Wiedenmayer (1994) remarkably assigned T. stewarti to Cinachyra, Desqueyroux-Faundez & Stone, 1992.
but this seems unlikely in view of the smooth surface and slit-like Description (from Schmidt, 1880b and Topsent, 1920a).
oscules. The type redescribed by Topsent represents half of the original
The genus Craniellopsis Topsent, 1913a was likewise erected specimen. It is 3.4 cm high and has a diameter of 3.8 cm. It has no
for tetillids possessing a cortex and lacking sigmaspires. Topsent trace left of the spicular root mentioned in Schmidt's description.
(1913a) named three species, of which de Laubenfels (1936a: 171) The surface is hispid and many foreign objects such as
subsequently designated Tetilla zetlandica (Carter, 1872a: 417) as foraminifera and sand grains have attached to it. At opposite sides
the type species. Four syntypes (BMNH 1983.6.14.1-4), as well as there are two large porocalices, both with palisade of very long
several slides made from these (fide Hooper & Wiedenmayer, spicules protruding up to 1.5 cm above the surrounding sponge sur-
1994: 432), are extant but have not been examined for this study. face. The porocalices are 1-1.5 cm deep and 0.8-1 cm in diameter.
This is a common species in the North Atlantic, sharing the two The left one has a perforated bottomplate, the other shows a few
size categories of oxeas with C. tethyoides, the smaller category of slit-like apertures in the bottom and the sides, but is generally with-
which is concentrated in the cortex. The lack of sigmas pires is con- out visible apertures. Schmidt apparently misinterpreted the func-
sidered of low importance. Lendenfeld (1903) and de Laubenfels tions of both types, but this was corrected by Topsent: the one
(1936a) maintained a separation at the genus and family level depicted left in Schmidt's drawing (here reproduced as Fig. 7A)
for species with (Craniellidae de Laubenfels) and without sig- is the inhalant porocalyx, the other the exhalant porocalyx. The
maspires (Tetillidae sensu de Laubenfels, Tethyopsillidae spicule palisade is made up of oxeas and protriaenes/prodiaenes. In
Lendenfeld) (cf. above). cross section the structure of the sponge - apart from the porocal-
ices - is radiate, with bundles of oxeas originating from a focal
point traversing towards the surface. There is no cortical special-
ization. Megascleres. Protriaenes and prodiaene modifications,
FANGOPHIUNA SCHMIDT, 1880 length not recorded, thickness of shaft up to 20 fLm, c1adi up to 400
fLm long, c1adome 130 fLm wide; anatriaenes, length not recorded,
Synonymy thickness up to 8 fLm, cladi up to 47 fLm, cladome 27 fLm; plagio/
orthodiaenes and -monaenes, length not recorded, thickness 25 fLm,
Fangophilina Schmidt, 1880b: 73. Spongocardium Kirkpatrick, c1adi, sometimes flexuous, up to 420 fLm; oxeas in two size cate-
1902: 224. gories, largest 25000 X 75 fLm, smallest 900-1500 X 15-30 fLm (the
latter are found in the choanosome intercrossing the bundles of
Type species oxeas). Microscleres sigmaspires 15-35/-Lm, larger ones often
elongated, wavy, toxiform.
Fangophilina submersa Schmidt, 1880b: 73 (by subsequent Remarks. Although he did not define it, Schmidt (1880b)
designation; RUtzler, 1987: 189). apparently considered Fangophilina to comprise a wider group of
sponges than only Fangophilina submersa, since he assigned
Definition Tetilla polyura Schmidt, 1870, Tetilla euplocamos Schmidt, 1868,
and Tetilla radiata Selenka, 1880 to it in the discussion of the new
Tetillidae with morphologically distinct exhalant and inhalant genus. He apparently forgot that Tetilla had been erected earlier by
porocalices positioned laterally, without cortical specialization, himself with Tetilla euplocamos as the type. Thus, we may surmise
without auxiliary megascleres. that Fangophilina was intended for at least several tetillids that did
not share the peculiar porocalyx-specialization of F. submersa. The
Diagnosis genus is of rather dubious validity, since several authors (e.g.,
RUtzler, 1987) have expressed doubts about the nature ofthe exha-
Globular sponges with hispid surface provided with two large lant porocalyx. Possibly, true oscules have become obscured
porocalices situated laterally. These are differentiated into an exha- through contraction in preserved state. Still, the fact that several
lant and an inhalant porocalyx. Skeleton radiate, with strongly pro- species have been described with similar structure is considered
truding megascleres. Cortical specialization absent. Megascleres sufficient to maintain the genus as valid for the time being (there is
protriaenes, anatriaenes, oxeas. Microscleres sigmaspires. Three also the consideration, that Fangophilina is senior to Cinachyrella,
species have been assigned to this genus, distributed in the the genus most similar to it).
Caribbean and Indian Ocean. Kirkpatrick, 1902, erected Spongocardium for type species
(by monotypy) Spongocardium gilchristi Kirkpatrick, 1902: 224,
Previous reviews pI. II fig. I, pI. ill fig. 1. This sponge (Fig. 7B) is very similar to
Schmidt's Fangophilina submersa, with the poral and oscular
Lendenfeld (1907); Topsent (1920a); Wilson (1925). porocalyx in a lateral position. There are a few spicular differences
Porifera· Demospongiae • Spiropborida • Tetillidae 95

.... ,

Fig. 7. Fangophilina species. A, Fangophilina submersa Schmidt, 1880b, drawing of cross section of ho1otype reproduced from Schmidt's pI. X fig. 3
(scale 1cm). B-C, Fangophilina gilchristi (Kirkpatrick, 1902 as Spongocardium). B, lectotype and cross section, reproduced from Kirkpatrick's pI. II fig. 1
(scale 1cm). C, drawing of spicules from Kirkpatrick's pI. III fig. I (scale see text).

(Fig. 7C) (e.g., there are orthotriaenes, 4900 X 35jLm and a second Diagnosis
smaller category of protriaenes, 5300 X 31 and 690 X 2jLm). Other
measurements are similar, oxeas up to 10mm X 80-100 jLm, anatri- Globular sponges with hairy-bristly surface, provided with
aenes ditto, sigmaspires 35-45jLm. However, Topsent's conclusion porocalices. Skeleton radiate, with bundles of oxeas originating
that Spongocardium gilchristi is a junior synonym of Fangophilina from a central focus and running radially to the surface. Spicules
submersa appears incorrect in view of the considerable differences. protriaenes, anatriaenes, calthrops-like triaenes, oxeas, sig-
Later, Kirkpatrick (1905) realized that Schmidt's genus was maspires. So far only a single wide-spread Indo-West Pacific
defined on the same characters, and pronounced the two genera species is reliably recorded.
synonyms. This was followed by Lendenfeld (1907: 157), who
erected a third species of Fangophilina, F. hirsuta. Previous reviews

Riitzler (1987).

PARATETILLA DENDY, 1905 Description of type species

Synonymy Paratetilla bacca (Selenka, 1867) (Fig. 8A-C).

Synonymy. Stelletta bacca Selenka, 1867: 569, figs 14-15;
Paratetilla Dendy, 1905: 97. Paratetilla bacca; Hooper & Wiedenmayer, 1994: 433, with addi-
tional synonyms. Tethya merguiensis Carter, 1883a: 366, pl. XV
Type species figs 6-8; Tetilla merguiensis; Sollas, 1888: 14. Tetilla tematensis
Kieschnick, 1896: 527. Paratetilla cineriformis Dendy, 1905: 97,
Tethya merguiensis Carter, 1883a: 366 (by subsequent desig- pl. III fig. 7.
nation; de Laubenfels, 1936a: 175). This is generally considered a Material examined. Type material: not located - Samoa. Type
junior synonym of Stelletta bacca Selenka, 1867. material of T. merguiensis. 1M (presumed) - Burma (=Myanmar).
Other material. ZMA specimens - Indonesia, 'Siboga' and 'Snellius
Definition II' collections (subjectively assigned to this species).
Description (based on Carter, 1883a and Selenka, 1867).
Tetillidae with porocalices, with megascleres including Globular (Fig. 8B), size 2-3 cm diameter (larger in subsequently
calthrops-like short-shafted triaenes. recorded specimens), surface uniformly hispid due to projecting
96 Porifera· Demospongiae • Spirophorida • Tetillidae

Fig. 8. Paratetilla haeca (Selenka, 1880) (scales see text). A, drawing of spicules reproduced from Selenka, 1867: fig. 15, as Stelletta). B-C, drawing of
habit (B) and spicules (C) of Tetilla merguiensis Carter, 1883a: pI. XV figs 6-8, considered a junior synonym.

megascleres, with numerous larger apertures, 0.5-1 cm diameter, Cinaehyrella. The calthrop-like short-shafted triaenes are probably
scattered evenly over the surface in Selenka's specimen, concen- derived from plagiotriaenes, which are occasionally recorded in
trated in the median region in Carter's specimen. Both authors Cinaehyrella (cf. Rtitzler, 1987; Rtitzler & Smith, 1993), and thus
interpret these as exhalant, but it is obvious they are porocalices. could be interpreted as a species rather than a genus character.
Colour: black-brown (according to Carter). In cross section there is
no cortex (Selenka mentions a thin cortical layer containing a layer
of calthrops, but this is simply the peripheral region), and the skele- TETILIA SCHMIDT, 1868
ton is radiate, with bundles of oxeas issuing from a central focus.
Megascleres (Fig. 8A, C; sizes combined from data and drawings Synonymy
of Selenka, Carter, Sollas and Dendy). Protriaenes, up to 3500 X 15
f.Lm, cladi up to 90 f.Lm; anatriaenes 3500 X 15 f.Lm, cladi 60 f.Lm; [Daetylella] Gray, 1872a: 461 (nomen oblitum). [Casula]
calthrop-like short-shafted triaenes (Selenka: 'vierzackige Sterne'; Gray, 1872a: 461 (nomen oblitum). Tetilla Schmidt, 1868: 40.
Carter: 'zone-spicule', Sollas: 'orthotriaenes'), cladi 100-200 f.Lm Chrotella Sollas, 1886a: 181. Spiretta Lendenfeld, 1888: 42.
(Carter gives I156th of an inch, which would be 450 f.Lm, clearly Traehygellius Topsent, 1894c: 8. Ectyonilla Ferrer-Hernandez,
in excess of the sizes found in other recorded specimens); oxeas, 1914a: 452. Kaira de Laubenfels, 1936a: 175.
up to 7000 X 42 f.Lm. A smaller category of oxeas is mentioned by
Carter (1883a) and Dendy (1905 as Paratetilla cinerijormis). Type species
Microscleres sigmaspires (not recorded by Selenka), 11-17 f.Lm.
Distribution and ecology. Widespread Indo-West Pacific (Samoa, Tetilla euploeamos Schmidt, 1868: 40 (by monotypy).
Myanmar, Indonesia, NE Australia, Sri Lanka), reefs, and deeper
water, down to + 200 m. Definition
Remarks. Rtitzler (1987) nominated Paratetilla cineri-
formis Dendy, 1905 as the type species, but this was superseded by Tetillidae without porocalices, without cortical specialization,
de Laubenfels' (1936a) previous nomination of Tethya merguiensis without auxiliary megascleres.
as the type. Since this species was mentioned by Dendy as a
species of Paratetilla, de Laubenfels' designation is valid. All Diagnosis
described tetillid species with calthrop-like short-shafted triaenes
appear to be referable to a single widespread species, Paratetilla Globular sponges with conulose or uniformly hispid surface,
baeca (cf. synonymy in Hooper & Wiedenmayer, 1994). If this is with few oscules, usually situated at the top. Usually with spicule
confirmed in comparative studies, the validity of this genus is dubi- strands at the base, acting as root system. Surface without porocal-
ous, because the possession of the genus character is then limited ices. In cross section, there is no visible cortex. The skeleton is spi-
to a single species, which in all other aspects conforms to rally radiate, with bundles of oxeas originating from a central focus
Porifera· Demospongiae • Spirophorida • Tetillidae 97

/
Fig.9. Tetilla euplocamos Schmidt, 1868. A, drawing of type and protriaene reproduced from Schmidt, 1868: pI. V fig. 10 (sizes see text). B, BMNH slide
of type. C-D, photos of spicules from BMNH slide. C, oxeas and bundle of small? protriaenes (scale lOOjJ.m). D, protriaene (arrow) (scale lOOjJ.m).

and radiating in a spiral manner to the surface. At the periphery, (Fig. 9C, D) in photos made from the BMNH slide. Distribution
these bundles of oxeas become mixed with triaenes, predominantly and ecology. Known only from type locality.
protriaenes. Microscleres sigmaspires (absent in the type species). Remarks. For no apparent reason, Sollas (1888) changed
About 50 species have been described from all parts of the world euplocamos into euplocamus. The species remains ill-known.
oceans, mostly from deeper localities. Since many authors did not Schmidt's description is casual and short and subsequent speci-
distinguish between Tetilla and Craniella, the actual number of mens from the same area identified under this name (Selenka,
species may be less. 1880) are only presumably conspecific. Nevertheless, there are
many species of tetillids sharing the characters described by
Previous reviews Schmidt, and the genus is considered clear and unequivocal.
The genus Dactylella Gray, 1872a: 461 was erected for type
Sollas (1888); RUtzler (1987). species Tethya dactyloidea Carter, 1869a: 15, figs 1-4 from the SE
coast of Arabia (by original designation). This is undoubtedly a
Description of type species member of Tetillidae, because Carter described a radiate structure
and oxeas and protriaenes as megascleres. No certainty exists about
Tetilla euplocamos Schmidt, 1868 (Fig. 9A-D) the presence of sigmaspires, because Carter had "given away the
Synonymy. Tetilla euplocamos Schmidt, 1868: 40, pl. V specimen" (to Dr Bowerbank, cf. Carter, 1871e: 103). In view of
fig. 10; Selenka, 1880: 469, pl. 27 fig. 5. the absence of clear porocalyces and apparent lack of a cortical
Material examined. Holotype (not seen): MZUS P0206 - skeleton, this is likely to be a Tetilla. Since the name Dactylella
Desterro, Bay of Rio de Janeiro, intertidal, colI. F. MUller. Slide of predates Tetilla it needs to be suppressed as an unused name in the
the holotype (examined; Fig. 9B): BMNH 1868.3.2.32 - bearing sense of ICZN Article 23.9. The name Tetilla has been frequently
Schmidt's handwriting, labeled "Desterrorretilla euplocamos Sch used in the past 50 years by many authors, and thus fulfils the
1868". Fragment 2MB 7165 - (see Desqueyroux-Faundez & demands for continued usage under Article 23.9. The name
Stone, 1992). Dactylella has not been used for a tetillid sponge after its original
Description (from Schmidt, 1868). Pear-shaped (Fig. 9A), proposal, although subsequently, [Dactylella] was also used by
with long root ('Nadelschopf'). No cortex. Surface finely hispid. Thiele, 1898 for type species Dactylella hilgendorfi Thiele, 1898:
In cross section the choanosomal skeleton consists of bundles of 56, pI. 4 fig. 8, pI. 5 fig. 25, pI. 8 fig. 41 a-b. This is a sponge of the
oxeas originating from a core spiralling outwards toward the sur- family Dictyonellidae, and the name is now considered preoccu-
face, where they mix with triaenes. The latter protrude beyond the pied, and replaced by Lipastrotethya de Laubenfels, 1954.
surface and cause a fine downy pelt of spicules. Megascleres. The genus Casula Gray, 1872a: 461 was erected, in Casuladae
Protriaenes (Fig. 9A, no size recorded), apparently no anatriaenes; Gray, 1872a: 461, for Tethya casula Carter, 1871e: 99, pI. IV from
oxeas 2300 X 22 }Lm. No sigmaspires were detected in the slide, nor South Africa (by monotypy). The description of Carter leaves no
recorded by Schmidt or others. Some spicules are here illustrated doubt that this is a Tetilla, probably considerably damaged during
98 Porifera· Demospongiae • Spiropborida • Tetillidae

collection giving rise to the rather strange schematic drawing of the The genus Trachygellius Topsent, 1894c: 8 was erected for
habit in his fig. 1. In its turn this induced Gray (1872a) to empha- type species Trachya globosa Carter, 1886a: 121 from shallow
size in his definition of the family Casuladae, that the sponge is water off Port Phillip Head, Victoria (by original designation). This
free-living and has a "funnel-shaped expansion or disc formed of is a yellow globular sponge with a short stalk and uneven surface.
elongated spicules united together". Spiculation include protri- Oscu1es in a group around the top. The species is described by
aenes up to 10 mm, oxeas up to 5 mm, and sigmaspires of 14 f.Lm. Carter (1886a) as having a skeleton that is radiate with a "thick der-
Since the family group name Casuladae Gray, 1872a is an unused mis". Spicules are reported to be oxeas of 4250 X 36 f.Lm, and sig-
name in the sense of ICZN Article 23.9 (not used after 1899), it is maspires of 8 f.Lm. Topsent (1894c) erected this genus in a footnote
here suppressed in favour of Tetillidae (used many times by many discussing the genus Gellius (Haplosclerida), assuming apparently
authors in the past 50 years). that the microscleres of T. globosa were true sigmas. However, it is
The genus Chrotella Sollas, 1886a was erected for type clear from Carter's description that they are sigmaspires and that
species Chrotella simplex Sollas, 1886a: 181; Sollas, 1888: 17, T. globosa belongs to Tetilla or possibly Craniella, and in view of
pI. II figs. 1-4 (by subsequent designation by Sollas, 1888: cxxv), the fact that special cortical oxeas were not reported, it is here con-
South Australia, 274m. The type series (BMNH 1889.1.1.12, with sidered a junior synonym of Tetilla. This assumption has been
slides BMNH 1894.11.16.27-32, not examined) consists of four recently verified by re-examination of the type material (slides AM
cream-coloured subspherical specimens of up to 2 cm in diameter. G2751-2) by J.N.A. Hooper (courtesy of Dr Penny Berents, AMS),
Oscules small, pores in sieve-like areas overlying subdermal confirming its allocation to Tetilla and not to Craniella.
spaces, surface uniformly pilose. Sollas records a cortex but neither The genus Ectyonilla Ferrer-Hernandez, 1914a: 452 was
his detailed description nor his drawings show a clear cortical spe- erected for type species Tetilla truncata Topsent, 1892a: 36,
cialization. There are many subdermal cavities but no peripheral pI. VIII fig. 7 from the NW coast of Spain (by original designation).
collagenous region, nor cortical spicules. Megascleres are protri- The genus was assigned to a new family Ectyonillidae defined as
aenes, with rhabds of up to 3400 X 20 f.Lm, cladi up to 158 X 16 f.Lm; "Sigmatophora with a skeleton ofAxinellidae". The holotype is
anatriaenes up to 5300 X 16 f.Lm, cladi up to 47 X 12 f.Lm; oxeas up housed in MOM (not examined). According to Topsent's descrip-
to 3000 X 24 f.Lm. Microscleres sigmaspires 12 f.Lm. The characters tion, this is a fragment of a definite Tetilla, with a radiate skeleton
of C. simplex appear to overlap with those of Tetilla and accord- of styles of 1200 f.Lm instead of oxeas (probably oxea-derived), and
ingly it is considered a junior synonym. anatriaenes and protriaenes. No microscleres. The styles are
The genus Spiretta Lendenfeld, 1888: 42 was erected for type unusual, but have nothing in common with those ofAxinellidae.
species Spiretta raphidiophora Lendenfeld, 1888: 43 (subsequent It is clear that it is a commonplace tetillid, and we assign it to the
designation herein, lectotype in AM G9076, designation herein, with synonymy of Tetilla on account of the absence of special cortical
a small fragment in BMNH 1886.8.27.634, here designated paralec- megascleres.
totype). The description of Lendenfeld makes it likely that this is a The genus Kaira de Laubenfels, 1936a: 175 was erected for
species of Tetilla, and Lendenfeld admitted as much himself at a later type species (by original designation) Tethya hebes Lendenfeld,
date (Lendenfe1d, 1903). Although he mentions a 'cortex' this was 1907: 98, pI. XVI figs 19-38 from NW Australia (type 2MB 768
apparently used to indicate the peripheral skeleton, rather than a real (not examined), with fragment BMNH 1908.9.24.66). This species
cortex. The species has an unusual spiculation by its complement of was assigned into synonymy with Cinachyra australiensis (Carter,
styles of 600 X 70 f.Lm, small choanosomal oxeas of 240 X 20 f.Lm, 1886b: 127 as Tethya cranium var. australiensis) by Burton
and ectosomal strongyles of 400 X 3 f.Lm. Furthermore there are radi- (1934a: 524), followed by Hooper & Wiedenmayer (1994: 430),
ating oxeas 3000-4000 X 50 /Lm, anatriaenes 1000 X 10 /Lm, sig- because both possessed rugose small oxeas. However, several dis-
maspires 14 f.Lm; apparently no protriaenes. Spiretta was relegated crepancies between the descriptions of australiensis and hebes are
into synonymy with Cinachyra by Burton (1934a: 524), as he apparent (presence of plagiotriaenes and anatriaenes in the latter).
assumed that the apertures of the type specimen represented 'col- Moreover, Lendenfeld's specimen possessed only a single lateral
lapsed porocalices'. This synonymy assignment was followed by oscule and no other apertures, whereas C. australiensis, which is
Hooper & Wiedenmayer (1994: 430). Conversely, we prefer here to an obvious representative of Cinachyrella, has abundant porocal-
consider Spiretta raphidiophora a member of the genus Tetilla on ices. Until fresh material has been found, we prefer to consider
account of the absence of clear porocalices and lack of cortex. Tethya hebes a member of Tetilla.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademicIPlenum Publishers, New York, 2002

Family Samidae Sollas, 1888

Rob W.M. Van Soest1 & John N.A. Hooper

Samidae Sollas (Demospongiae, ? Spirophorida) is an enigmatic monotypical family of excavating sponges. The affinities of the genus
Samus Gray are obscure, but the possession of sigmaspire microscleres is considered evidence for membership of the order Spirophorida.
The typical family of Spirophorida, Tetillidae Sollas, differs considerably in habit and additional spiculation. Although several species
have been described in the genus Samus, only one species name remains recognized in the genus, the others having been transferred to
other genera of different families. The single species Sam us anonymus Gray is reported to be almost cosmopolitan (Brazil, Caribbean,
Indian Ocean, Mediterranean, Australia), but is probably a complex of closely related species. It forms insignificant excavations in corals
and coralline algae and was long thought to be a facultative dweller of empty holes excavated by clionids.
Keywords: Porifera; Demospongiae; Spirophorida; Samidae; Samus.

DEFINITION, DIAGNOSIS, SCOPE controversy until the present. Major authors like Topsent (1928c:
28), Levi (1973: 599) and Wiedenmayer (1994: 29) considered the
Synonymy combination of excavating habit, smallness of the amphitriaenes
and their resemblance to amphiasters of Thoosa and Alectona, for
Samidae Sollas, 1888: 244. example, compelling evidence for membership of Hadromerida,
whereas many contemporary taxonomists attach more weight to
Definition the presence of the sigmaspires as a synapomorphy with Tetillidae.
De Laubenfels (1936a: 180) assigned Samus to Homosclerophorida
Spirophorida (?) with amphitrichotriaenes, without long- near Corticium, but that has found few followers. In the present
shafted triaenes or oxeas. volume we choose for inclusion of Samidae in Spirophorida, based
on the sigmaspires, which are shared with Tetillidae, and based on
Diagnosis the argument that the excavation habit is not exclusive to
Hadromerida. We realize that there is an inconsistency in this
Small sized excavating sponges with scanty skeletons consist- emphasis on shared possession of sigmaspires, because the family
ing of confusedly arranged small amphitriaenes and sigmaspires. Scleritodermidae is assigned to 'Lithistida' in spite of its posses-
No further spicule types. sion of sigmaspires. However, because at this moment in time,
redistribution of the polyphyletic 'Lithistida' over the various
Scope orders and families of Demospongiae cannot be confidently exe-
cuted, we prefer to keep 'lithistids', including Scleritodermidae,
A single genus, Samus, is recognized. Its only valid species as a sponge group defined pragmatically on the presence of an
has been reported from many parts of the world. articulated desma skeleton.

History Previous reviews

Bowerbank (1864: pI. II figs 41-42) figured the megascleres Sollas (1888); Lendenfeld (1903); Hooper & Wiedenmayer
of an unknown sponge from Bahia. Gray (1867a: 526) subse- (1994, as member of Tetillidae).
quently named this Samus anonyma, assigning it to his family
Clioniadae. He indicated relationship with Axos cliftoni and Cliona
species (as Vioa and Euryphylle), but gave little comment or SAMUS GRAY, 1867
descriptions. We may conclude however, that Gray tended to regard
the amphitriaenes as microscleres (the true microscleres were not Synonymy
known to him then). Carter (1879b) gave a full description of
S. anonyma based on material from Australia, and discussed the Sam us Gray, 1867a: 526.
affinities. He concluded that Samus was not related to either Axos
or Cliona, but appeared to belong to a group of excavating sponges Definition
like Dotona and Alectona, later to be united in the family
Thoosidae. Sollas (1888: 244) described the original material from See definition of family.
Bahia and erected a separate family Samidae in his suborder
Sigmatophora (=Spirohorida) based on the amphitriaenes. This Diagnosis
fundamental difference - alliance with either Thoosidae in
Hadromerida or a separate family in Spirophorida - remains a See diagnosis of family.

99
100 Porifera' Demospongiae • Spiropborida • Samidae

Fig.t. Samus anonymus Gray, 1867a. A, Bowerbank's (1864) drawings of the spicules of an unnamed sponge (copied from his plate II figs. 41-42) (scale
see text). B-D, SEM images of spicules of a Cura~ao specimen, made by Ms EUy Beglinger (ZMA). B, lateral view of amphitriaene (scale 10 ILm).
C, amphitriaene from the head (scale 10 ILm). D, sigmaspire (scale 1 ILm).

Description of type species may vary from 20--80 fLm in length by up to 38 fLm in thickness, total
length of amphitriaene including cladi may be 150 fLm; cladome may
Samus anonymus Gray, 1867a: 526 (Fig. 1). be up to 160 fLm, with protocladi and deuterocladi each about 15-75
Synonymy. Unnamed sponge of Bowerbank, 1864: 234, pI. fLm in length. Microscleres (Fig. ID) sigmaspires with relatively few
II figs. 41-42. Samus anonyma Gray, 1867a: 526; Carter, 1879b: and long spines, 10--14 fLm. Distribution and ecology. Reported from
350, pI. XXIX figs 1-4; Samus anonymus; Carter, 1880b: 59. NE Brazil, 'Australia', Sri Lanka, Singapore, Florida, Palau Islands,
Material examined. Holotype (not seen): BMNH West Africa, Mediterranean, Colombia, Cura«;ao, from shallow
1877.5.21.221 (slide) - Bahia (see remarks below). Other material. depths, excavating limestone substrates, often in caves.
ZMA 13348, 14063, 14069 - reef cavities Cura«;ao, colI. Remarks. Ms Clare Valentine kindly provided the following
S. Wiegman, E. Wierenga and I. Wunsch. information on the type material. The type, a dry specimen, is sup-
Description. Excavating sponge making tiny holes and corri- posed to be in BMNH but a recent search did not turn it up.
dors in limestone of corals and coralline algae. Colour variously However, there is a slide (registered as BMNH 1877.5.21.221)
recorded as yellowish, purple or blueish, but that may have been with Bowerbank number Bk. 221 on it and the locality Bahia.
influenced by other boring sponges which are invariably present in It also reads "type". In the BMNH register the entry reads:
samples containing this species. The blue colour especially may be "Sponge spicules - illustration Fig. 42 Vol. I. Mon. Brit.
derived from e.g., Cliona schmidti. The surface character and pres- Spongidae". This is evidently the material upon which Gray
ence of papillae or similar extralithic parts of the sponge remain founded the species. Hooper & Wiedenmayer (1994: 433) erro-
unknown, as the species so far is detected only in preserved samples, neously indicated another specimen in BMNH from Australia as
collected mostly because of the presence of more obvious excavat- the holotype. This is the material mentioned in Carter's (1879b:
ing sponges. Skeleton a ? confused arrangement of megascleres, 350) subsequent record of the species, in which he described a
with numerous microscleres. Megascleres (Fig. lA-C) short-shafted hitherto undescribed Bowerbank specimen (bearing Carter's num-
amphipro- or amphiplagiotriaenes, basically symmetrical amphitri- ber 699) from Australia.
chotriaenes. However, frequently cladi are only dichotomous, and In addition to the type material from Bahia and the Australian
then symmetry is usually lacking. An often smaller variation is dis- material of Carter (l879b), the species is reported from Sri Lanka
tinctly asymmetrical in having only three undivided cladi at one end (Carter, 1880b), Singapore (Sollas, 1902), Florida (de Laubenfels,
of the shaft and three trichotomously divided cladi at the other end. 1936a), Palau Islands (de Laubenfels, 1954), West Africa (Levi,
Such spicules may appear to be calthrop-like, but they are basically 1959), Seychelles (Thomas, 1973), the Mediterranean (Pulitzer-
the same amphitriaene-type as the fully developed larger ones, and Finali, 1983), Colombia (Wintermann-Kilian & Kilian, 1984), and
only Sollas proposed to distinguish them as a distinct spicule type Cura«;ao (this volume). The various records from all over the world
(normal amphitriaenes and 'heteropolar' amphitriaenes). Rhabds are suspect from a genetic point of view. It is likely that widely
Porifera· Demospongiae • Spiropborida • Samidae 101

separated populations comprise different species, but the absence Pachastrellidae) and S. complicatus (=Rhachella, Theonellidae).
of spicular differentiation precludes definite conclusions in this Diaz & Van Soest (1994) suggested the latter to be a true Samus,
respect. All reported spicule sizes and forms are similar. Of the fur- but the isolated spicule upon which the genus Rhachella was
ther species assigned to Sam us by Carter (1880b) not one appears founded is not an amphitriaene, but a cladocalthrops, i.e.,
to share the characters of Samus anonymus. These are S. simplex a calthrops with proliferated cladi such as is found in some
( = Stoeba, Pachastrellidae), S. parasiticus ( = Triptolemma, theonellid 'lithistids'.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Spirasigmidae Ballmann, 1912

John N.A. Hooper l & Rob W.M. Van Soest2

Spirasigmidae Hallmann (Demospongiae, Spirophorida) contains two genera, both known only from single specimens from remote
islands of the Pacific and Indian Oceans. They are thinly encrusting to small conical shapes, with radial skeletons that fan out and become
plumose in the periphery where they become nearly tangential at the surface, composed of large strongyloxeas. One genus (Spirasigma)
also has a secondary disorganised skeleton of roughened 'microxeas'. Both lack triaenes, and their allocation to Spirophorida is based on
possession of true sigmaspires, although it is included in Spirophorida as a family incertae sedis.
Keywords: Porifera; Demospongiae; Spirophorida; Spirasigmidae; Spirasigma; Tentorina.

DEFINITION, DIAGNOSIS, SCOPE and Trachycladus Carter, and was supposedly characterized by the
presence of sigmaspires and oxeas arranged radially (reminiscent
Synonymy of Tetillidae). Later, Hallmann (1916a) did not mention
Sigmaspiridae when discussing Trachycladus, and we can only
Spirasigmidae Hallmann, 1912: 131; Hooper & Wiedenmayer, assume that he had changed his mind. De Laubenfels (l936a)
1994: 440. referred both the former genera to his subfamily Rhaphidistiinae
(in Epipolasida, roughly corresponding to Coppatiidae, now in
Definition Ancorinidae). Hooper & Wiedenmayer (1994) tentatively included
Trachygellius and Spirasigma in the family Thrombidae
Spirophorida without long-shafted triaenes, with strongylox- (Astrophorida), as genera incertae sedis, but this allocation is now
eas forming a primary ascending skeleton that fans out towards the rejected (Uriz, Family Thrombidae, this volume) as Thrombidae
periphery and becomes nearly tangential at the surface. are defined by their possession of amphiaster microscleres and
acanthotriaene megascleres, lacking any oxeas, which clearly dif-
Diagnosis fer from these two former taxa.
Re-examination of the type species of Trachygellius (Trachya
Thickly encrusting to conical sponges with main skeleton globosa Carter), showed that it was a clear synonym of Tetilla
composed of large strongyloxeas forming radial bundles initially in (Tetillidae), whereas allocation of Spirasigma is more problematic.
the basal skeleton, plumose tracts towards the periphery of the No Tetillidae have an encrusting growth form, and the possession
skeleton, and nearly tangential tracts at the surface; microscleres are of a secondary skeleton of roughened 'microxeas' is also unique to
sigmaspires, and roughened microxeas in one genus (Spirasigma). the family. Similarly, although several tetillids lack triaenes (pre-
sumed secondary losses, such as in Tetilla and Cinachyrella), the
Scope same logic does not necessarily apply to Spirasigma. One clue
which does confer possible affinities is the possession of true sig-
Two poorly known and monospecific genera, Spirasigma maspires (roughened, c- and s-shaped or multiple twists, abrupt
Hallmann and Tentorina Burton. terminations differing substantially from sigmas), and on this basis
Spirasigrnidae is allocated to Spirophorida as a family incertae
History sedis. Tentorina Burton is also here assigned to this family, being
very similar to Spirasigma but lacking the secondary skeleton of
Spirasigrnidae, as proposed by Hallmann (1912), initially microxeas.
included three genera, Trachygellius Topsent, Spirasigma Hallmann

KEY TO GENERA

(1) With roughened microxeas forming a secondary skeleton ................................................................................................. Spirasigma

Lacking microxeas .... ........ ............ .... ........... ........................... ........... .............. ............. ...... ....... ... .......... .... ....... .... .... ... ... ...... Tentorina

SPIRA SIGMA HALLMANN, 1912

Synonymy Type species

Spirasigma Hallmann, 1912: 131; de Laubenfels, 1936a: 159; Gellius aculeatus Whitelegge, 1897: 326 (by original
Hooper & Wiedenmayer, 1994: 440. designation).

102
Porifera· Demospongiae • Spiropborida • Spirasigmidae 103

Definition long, 1 mm wide. Texture soft, compressible, tough. Ectosome

partially transparent, oscules and ostia not visible but reported as
Spirasigmidae which has a secondary skeleton of roughened "occurring between the aculeate processes, subcircular ... 1.2-1.5 mm
microxeas overlaying the primary ascending, plumose skeleton of in diameter". The choanosomal skeleton consists of thick multispicu-
strongyloxeas. lar tracts of larger strongyloxeas, arising from the basal skeleton,
attached to a coral base. In the basal region these tracts form discrete
Diagnosis bundles, without any bifurcation or anastomoses. Approximately
mid-way through the choanosomal region tracts begin to bifurcate,
Thickly encrusting sponge with main skeleton composed of and continue increasingly towards the surface where they become
large strongyloxeas forming radial bundles initially in the basal plumose, fanning out to touch neighboring tracts. In the peripheral
skeleton, plumose tracts towards the centre of the skeleton, and skeleton these tracts become almost disorganised, with unispicular
halichondroid tracts that are nearly tangential at the surface; or paucispicular tracts protruding through the surface, and also
a dense secondary skeleton of roughened microxeas overlays the similar sized tracts (often bispicular) nearly tangential to the sur-
primary ascending skeleton, and microscleres also include sig- face, lying just below the surface. In the region of surface conules
maspires concentrated mainly around choanocyte chambers. these tracts may form anastomoses with adjacent tracts and reform
to produce multispicular bundles supporting the conules. The over-
Description of type species all impression of the choanosomal structure is radial at the base and
nearly halichondroid at the surface. Throughout the skeleton are
Spirasigma aculeata (Whitelegge, 1897) (Fig. 1). dispersed numerous large 'microxeas', forming a dense secondary
Synonymy. Gellius aculeatus Whitelegge, 1897: 326-327, skeleton in between the more or less ascending larger oxeas. These
pI. 13, fig. 3; Hallmann, 1912: 131-132. microxeas are slightly less common (but equally disorganised) at
Material examined. Holotype: AMS G1659 (slides AM the surface, and most dense (and disorganised) in the basal skele-
G 1659, Z5385) - West Funafuti (Tuvalu), central South Pacific, ton. There is no real basis to support Hallmann's (1912) contention
deep water in the lagoon, colI. C. Hedley. that this secondary skeleton of microxeas is isotropic, allegedly
Description. Thickly encrusting, 45 X 20 mm long, up to resembling that of Gellius, but it is true that individual microxeas
12 mm thick. Surface shaggy, with small pointed conules up to 8 mm do cross in all directions, and depending on the plane of focus
through thick sections they may be construed of having some sort
of reticulate organisation. It is suggested that any interpretation of
isotropic structure is an artefact. Collagen is abundant throughout,
heavier in the basal skeleton than at the periphery. No fibres were
observed surrounding tracts of megascleres. Sigmaspires appear
to be concentrated several layers deep around oval-elongate
choanocyte chambers (up to 100 I-\-m diameter). Megascleres are
strongyloxeas, or more accurately, large thick oxeas with long
tapering rounded ends, with straight or slightly curved smooth
shafts. Generally, the larger megascleres have rounded tips and the
smaller, thinner ones are sharply pointed, with a range of termina-
tions in between (740--1670 X 6-18I-\-m). 'Microxeas' were
described as megascleres by Whitelegge (1897), but these are
roughened and almost resemble very thick onychaetes (except they
are consistently symmetrical and sharply pointed), and as such they
are considered here to be microscleres. These microxeas are pre-
dominantly straight (112-184 X 2-6l-\-m). 'Sigmas' are actually
sigmaspires, with c- and s-shapes, and sometimes with double
twists, with their surfaces minutely roughened (11 X241-\-m chord
length X 1.5-31-\-m maximum width).
Remarks. De Laubenfels (1936a: 159) suggested that
Spirasigma was defined to contain two types of megascleres,
smaller oxeas and larger strongyles, and that microscleres were 'spi-
rals' something like those in Trachygellius and Spiroxya Topsent.
This is not an accurate description, however, with the larger spicules
being strongyloxeas and smaller oxeas are roughened and could
legitimately be classed as microscleres (and for this reason they are
described here as 'microxeas'). Similarly, the comparison with the
other two genera is not entirely accurate either. In the case of
Trachygellius (now assigned to Tetillidae), these 'spirals' are sig-
maspires, c- and s-shaped, with roughened surfaces. This latter
Fig. 1. Spirasigma aculeata (Whitelegge). Holotype AMS G1659. A-B, genus also has a completely radial arrangement of huge oxeas in
larger and smaller strongyloxeas (oxeas) (scale 250 fLm). C-D, termina-
tions of the same (scale 10 fLm). E, roughened microxea (scale 20 fLm).
bundles, and is clearly a species of Tetilla. In the case of Spiroxya
F, sigmaspires (scale 10 fLm). G, plumose radial skeleton near periphery (now assigned to Alectonidae) these 'spirals' consist of curled or
(scale 500 fLm). H, secondary skeleton of microxeas (scale 500 fLm). curved rhabds, bearing no resemblance to sigmaspires whatsoever.
104 Porifera· Demospongiae • Spiropborida • Spirasigmidae

Hallmann (1912) also made a misleading comparison between parsimonious allocation of this taxon, but for the fact that the spec-
Spirasigma and Trachycladus. He was correct in assuming that the imen is truly encrusting on a coralline base (i.e., it is not a fragment
sigmaspirae of Spirasigrna were not "ordinary sigmata", but incor- of a tom specimen), its main skeleton is plumose (and eventually
rect in stating that they were more similar to "sigmaspires of halichondroid) in the peripheral region, unlike typical radial struc-
Trachycladus" (in the latter they are curled rhabds, not sigmaspirae). tures seen in Cinachyrella, and its possession of a secondary skele-
The allocation of Sigmaspira to a higher taxon is not straight- ton of microxeas. The genus remains incertae sedis within
forward. Its larger megascleres are more-or-Iess radially arranged Spirophorida, with true sigmaspires the only definite apomorphy.
in bundles, vaguely reminiscent of Tetillidae as suggested by
Hallmann (1912), but these bundles fan out near the surface and as
such they are clearly plumose in the peripheral skeleton. Hallmann TENTORlNA BURTON, 1959
(1912) also suggested that if "the fibre-forming strongyla disap-
peared, the species would, without doubt, owing to the mode of Synonymy
distribution of its smaller oxea, be classed as a Gellius"
(Chalinidae), but this comparison is purely superficial. Microxeas Tentorina Burton, 1959a: 205.
of Sigmaspira do not form a true isotropic or isodictyal reticulation
but are dispersed in such heavy numbers that they appear at all Type species
angles to the main skeletal tracts, depending on the plane of view in
thick sections. Similar microxeas with roughened surfaces also Tentorina sigrnatophora Burton, 1959a: 205 (by monotypy).
occur in several Cinachyrella (e.g., c. australiensis, C. kuekenthali;
see also Rtitzler & Smith, 1992), so they are not unique to Definition
Spirasigmidae, although their distribution in a secondary skeleton
in Sigrnaspira is most unusual. It could also be argued that if Spirasigmidae lacking microscleres other than sigmaspires.
not for the encrusting shape Sigrnaspira could be assigned to
Cinachyrella, as an atypical species or a very young or badly dam- Diagnosis
aged specimen (the latter supposition supported by its shaggy sur-
face, which is often a sign of a maltreated Cinachyrella half-ripped Conical sponge with radiating tracts of strongyloxeas, which
off the substrate). This may indeed be the situation, and the most level off at the surface to form a tangential layer. Microscleres
sigmaspires. Maldives. Monotypical.

Description of type species

Tentorina sigrnatophora Burton, 1959a (Fig. 2).

Synonymy. Tentorina sigmatophora Burton, 1959a: 205,
fig. 7.
Material examined. Holotype: BMNH 1936.3.4.516 (slide)-
Maldives, John Murray Expedition, Stn 157, 04°44'N, n055'E,
229 m depth, coral rock.
Description (after Burton, 1959a). Small, conical sponge
with smooth, even surface. Apparently there are one or more apical
oscules. The consistency is described as soft and delicate. Colour
pale yellow (in alcohol). The skeleton consists of radiating curving
tracts of megascleres, thickness 50--220 J.Lm, consisting of 12-25
spicules in cross section, spaced apart 100--300 J.Lm, fanning out
and levelling off at the surface to form apparently a more-or-Iess
tangential layer at the surface. Between the tracts there is a dense
homogeneous presence of sigmaspires. Megascleres are perfectly
formed fusiform strongyloxeas with a tapering but rounded-off end
and a sharply pointed end. Size variable, 400--900 X 10--24 J.Lm, no

s distinct size categories are apparent and the various sizes are mixed
in the tracts without localization.
Remarks. This is a monotypical genus with rather perplex-
ing skeletal and spicular characters. With Spirasigma aculeata it
shares the strongyloxeas in a relatively small size, the sigmaspires,
the soft consistency, and the irregular non-globular shape. It differs
from that species in lacking the spined microxeas and in having a
smooth rather than a shaggy surface and lacking conules. It
Fig.2. Tentorina sigmatophora Burton. Strongyloxea and sigmaspires (for remains to be determined whether these differences are sufficiently
sizes see text) (from Burton, 1959a). different to keep the two as separate genera.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob w.M. Van Soest
© Kluwer AcademicIPlenum Publishers, New York, 2002

Order Astrophorida Sollas, 1888

John N.A. Hooper! & Rob W.M. Van Soest2

I Queensland Museum, P.O. Box 3300, South Brisbane, Qld, 4101, Australia. (JohnH@qm.qld.gov.au)
2Zoological Museum, University of Amsterdam, P.O. Box 94766, 1090 GT, Amsterdam, Netherlands. (soest@science.uva.nl)

Astrophorida Sollas (Demospongiae), also sometimes known as Choristida Sollas, include sponges with asterose microscleres and
tetractinal megascleres (either sometimes lost), together with microxeas, microrhabds and oxeas, and skeletal architecture always radial,
at least at the surface, but more confused towards the centre of the body. Five families are currently included (with 38 genera and two sub-
genera), and species are known from all oceans and at all depths.
Keywords: Porifera; Demospongiae; Astrophorida; Ancorinidae; Calthropellidae; Geodiidae; Pachastrellidae; Thrombidae.
-------- ._. _ .._ ... __. _ - - - _ ..__ .._ ..- ... _ . __ .._ .. _ - - - _ . _ -

DEFINITION, DIAGNOSIS, SCOPE Brachiaster, Characella, Cladothenea, Halina, Pachastrella,

Poecillastra, Stoeba, Thenea, Triptolemma, Vulcanella, Vulcanella
Synonymy (Annulastrella), Vulcanella (Vulcanella), and Thrombidae Sollas,
1888 (monotypic: Thrombus).
[Asterophora] Sollas, 1887 (nomen nudum). Astrophora Sollas,
1888. Astrophoridae Brien, 1968. Astrophorina Wiedenmayer, Remarks
1977b. Choristidae Sollas, 1880 (part). Choristida Sollas, 1885b
(part). Tetractina Vosmaer, 1885b (part). Not Tetractinellida Sollas's (1888) early concept of suborder Astrophora was
Marshall, 1876. Epipolasida sensu de Laubenfels, 1936a (in part). defined as "Choristida in which one or more of the microscleres is
an aster", with order Choristida defined as Tetractinellida lacking
Definition 'lithistid' desmas, and in turn the Tribe Tetractinellida defined to
include Demospongiae with triaene or tetraxon megascleres, or
Demospongiae with asterose microscleres, sometimes with desmas. Disassembling this hierarchy of taxa into an inclusive def-
microxeas and microrhabds, and with tetractinal megascleres and inition for the order Astrophorida we arrive exactly at the definition
oxeas radially arranged at least peripherally. presented by Hartman (1982), repeated above. That this concept
has survived over a century of revisions is perhaps testimony to its
Diagnosis robustness amongst the Porifera. Early histories of the order are
provided by Sollas (1888: cii) and Topsent (1928c: 25).
Sponges usually have a coarse texture emphasizing silica con- This order is sometimes referred to as Choristida, a taxon
tent over spongin in the skeleton. Microscleres are asters in one or which is still occasionally used by some contemporary authors.
more categories (sometimes lost), sometimes accompanied by However, 'Choristida', as originally proposed by Sollas (1880) and
microxeas and microrhabds. Megascleres are tetractinal, usually subsequently refined by Vosmaer (1882b) and Sollas (1886a), dif-
triaenes, calthrops, or short-shafted triaenes, together with oxeas, fers from Astrophorida in both its content and definition.
always with some radial skeletal architecture obvious at least in the Conversely, usage of 'Choristida' by contemporary authors (e.g.,
peripheral skeleton, and with architecture often more confused Bergquist, 1978) generally refers to Astrophorida s.s. and not to
towards the centre of the body. In genera that have lost tetractinal Choristida s.s., because under Sollas's (1886a) definition the latter
megascleres, leaving only oxeas, there is always some radial skele- taxon contained families currently assigned to Astrophorida
tal organisation remaining. Reproduction is oviparous although (Pachastrellidae, Theneidae, Stellettidae and Geodiidae) +
gametes are so far described for very few species. Larval stages are Homosclerophorida (Corticiidae) + Spirophorida (Tetillidae).
not yet known. Tetractinellida sensu Zittel (1879), which explicitly excluded
'Lithistida', essentially mirrors that of Choristida, and for this
Scope reason the taxon was renamed Tetractina by Vosmaer (1885b).
In the 'Challenger' report on the Tetractinellida Sollas (1888)
Five families, 38 genera and two subgenera are currently again revised Choristida and proposed three suborders: (1)
included in this order: Ancorinidae Schmidt, 1870 (with 15 genera: Sigmatophora Sollas, 1887 (containing Tetillidae and Samidae),
Stelletta, Tethyopsis, Cryptosyringa, Rhabdastrella, Jaspis, which we now know as order Spirophorida; (2) Microsclerophora
Ancorina, Disyringa, Stryphnus, Ecionemia, Psammastra, Penares, Sollas, 1887 (containing Plakinidae, Corticiidae and Thrombidae),
Melophlus, Asteropus, Tribrachium, Holoxea), Calthropellidae which is clearly polyphyletic; and (3) Astrophora Sollas, 1888
Lendenfeld, 1907 (with four genera: Calthropella, Chelotropella, (with three 'Demi' containing Theneidae, Pachastrellidae,
Pachastrissa, Pachataxa), Geodiidae Gray, 1867a (with six genera: Stellettidae, Epipolasidae (with question), Geodiidae and
Erylus, Caminus, Geodia, /sops, Pachymatisma, Sidonops), Placospongiidae) (see below). Bergquist (1968) employed
Pachastrellidae Carter, 1875b, including Theneidae Carter, 1883 'Choristida' in the sense of Lendenfeld (1903), Hentschel (1923),
(with 12 genera and two subgenera: Acanthotriaena, Ancorella, Topsent (1928c), de Laubenfels (1936a) and others, all of who

105
106 Porifera· Demospongiae • Astrophorida

contributed to refining the taxon to a point where it became nearly (1) Euastrophorida (with euasters - including sterrasters, oxyasters,
synonymous with our present understanding of Astrophorida aspidasters, strongylasters etc.), contained Geodiidae, Ancorinidae
(although Bergquist retained Tetillidae in the order but relegated and Caltbropellidae. (2) Streptosclerophorida (with streptosclere
Pachastrella to the Homosclerophorida). microscleres - including sanidasters, metasters, amphiasters,
Thus, the concept of the taxon Choristida has now nearly plesiasters etc.) (refer to Chombard et al., 1998, for illustrations of
merged with that of Astrophorida, and the choice of one name over microsclere morphology), contained Pachastrellidae (including
another might be decided by the Rules of Priority (Choristida has Theneidae) and Thrombidae, in addition to many desma-bearing
seniority over Astrophorida). However, suprafamilial taxa are not 'lithistid' taxa with obvious astrophorid affinities (Le., those bear-
bound by the rules of the ICZN (Anon., 1999), and consequently we ing triaenes: Tetracladina Zittel, l878c; Megamorina Zittel, 1878c;
follow Levi (1973) and Hartman's (1982) usage of Astrophorida. Dicranocladina Schrarnmen, 1924a and Streptosclerina Reid,
This decision is justified on the basis that its present usage is nearly 1963a (also known as Asterostreptidae Topsent, 1902». This pro-
identical to the original concept of Sollas (1887), whereas posal has potential merit, and as such the Key to Families, provided
Choristida, as used by contemporary authors, bears little resem- below, was constructed based on this fundamental subdivision of
blance to the original taxon. Furthermore, as far as we can ascertain the order Astrophorida. However, it requires further morphological
there has never been a genus 'Choristes', so in effect Choristidae and molecular resolution, particularly with regard to the question
Sollas, 1880 could also be construed as a nomen nudum. of incorporating 'Lithistida' into the present concept of
This present understanding of Astrophorida follows Levi Astrophorida (see below), and as such it is not been progressed fur-
(1973: 595), who elevated it to ordinal status. He excluded ther here. Furthermore, there appears to be an overlap between the
[Epipolasidae] Sollas, 1888 (which he placed in an order incerlae groups Euastrophorida and Streptosclerophorida. Genera such as
sedis) and Placospongiidae (which he referred to Hadromerida), and Stryphnus and Asteropus (and to a lesser extent Ancorina and
added the families Thrombidae and Caltbropellidae. The most recent Ecionemia), have both euasters and streptasters. It may be signifi-
comprehensive treatment of Astrophorida, that of Hartman (1982), cant that sanidasters and amphiasters of these genera appear to be
included seven families (Stellettidae, Geodiidae, Caltbropellidae, based on a straight rod, and appear at first glance not to be spiral, as
Pachastrellidae, Theneidae, Thrombidae and Jaspidae (also known are the spirasters and metasters of Pachastrellidae. But this
as Coppatiidae», and some of these are merged here to form problem of homology is indeed contentious and needs further study.
a revised ordinal taxon that contains only five families. Discussion of the concepts of Astrophorida and Choristida
Other families included in Astrophorida at one time or another also requires consideration of the nominal order Tetractinellida,
are now allocated elsewhere as follows: Stellettidae Carter, proposed by Marshall (1876) and further developed by Sollas
Coppatiidae Topsent and Jaspidae de Laubenfels are included here (1888) and Lendenfeld (1907) in particular. Sollas (1888: xcix)
in Ancorinidae; Erylidae Lendenfeld is included in Geodiidae; defined the order as "Demospongiae in which some or all of the
[Epipolasidae] Sollas, 1888, is also a nomen nudum with the genus scleres are tetraxons, triaenes, or desmas". Tetractinellida contains
Epipolasis de Laubenfels, 1936a (not erected until half a century what we now know as Astrophorida + Spirophorida + 'Lithistida',
after the family was established), now included in Halichondriidae sharing the possession of tetraxon spicules as its only apomorphy.
while the other genera included in [Epipo1asidae] by Levi (1973) Inclusion of 'Lithistida' into this clade, based on this apomorphy at
(Jaspis, Stelletinopsis, Asteropus) are referred in Ancorinidae (see least, is questionable given that not all 'lithistids' contain triaene
chapter by Uriz, this volume). Theneidae Carter is included here in megascleres, but this observation merely supports the already well-
Pachastrellidae, although this allocation remains controversial (see espoused hypothesis that 'Lithistida' are polyphyletic (e.g., Kelly
chapter by Maldonado, this volume). Corticiidae is a junior syn- Borges & Pomponi, 1994), whereas the majority of 'lithistid' taxa
onym of P1akinidae, and now allocated to order Homosclerophorida, are probably true astrophorids (or tetractinellids). In this regard,
and Tetillidae belongs to the order Spirophorida. There have been Reid's (1963a) proposal to subdivide the 'Lithistida' into several
several proposals to subdivide the order Astrophorida, based on orders and suborders (based on both the desma and free spicule
aster morphology. The earliest of these schemes was that of Sollas morphologies) may have some merit, although his allocation
(1888: cxiii) who proposed three clades. (1) Demus Streptastrosa of other non-lithistid taxa to this scheme is highly contentious and
Sollas, 1888 ("in which one of the microscleres is a spiraster; or not accepted by most contemporary authors. Interestingly, Sollas
when this is not the case one of the megascleres is a calthrops"), (1888: c) adopted a contrary position, arguing that 'Lithistida'
containing Theneidae and Pachastrellidae. (2) Demus Euastrosa lacking tetraxons (or triaxons) had probably lost them, and he
Sollas, l886a ("in which euasters are always present, but never strongly supported the possession of desmas as a phylogenetically
spirasters nor sterrasters ... Triaenes are present but not informative character to unite the clade 'Lithistida'. This position
calthrops"; Sollas, 1887), containing the Stellettidae (and is one to which we have returned to in this volume (see chapters on
[Epipolasidae]). (3) Demus Sterrastrosa Sollas, 1887 ("in which 'Lithistida' by Pisera & Levi, this volume) - but one that is perhaps
the characteristic microsclere is a sterraster"), containing the based more on ignorance rather than on any substantial new knowl-
Geodiidae (and Placospongiidae). These three 'Demi' were united edge of this taxon since the early 20th century. Resolving the
by the common possession of some form of aster, and also 'lithistid' dilemma remains a future challenge that might best be
frequently possessing a second distinctive category of aster resolved from molecular datasets.
(forming the clade Astrophorida). This scheme was 'unnatural' This concept of Tetractinellida, including Reid's (1963a) pro-
(polyphyletic) and has long since been abandoned. posed classification, was recently revisited using both morphologi-
By comparison, the most recent of these schemes was sug- cal and 28s rRNA molecular data (Chombard et al., 1998), lending
gested by Reid (1963a, 1968d), and reiterated by Levi (1991) and support to the recognition of a monophyletic clade, Tetractinellida,
Chombard et al. (1998), who recognised two orders, Euasterophorida containing [(Astrophorida + 'Lithistida') and Spirophorida]. These
Reid, 1963a and Streptosclerophorida Dendy, 1924, to remove authors noted that some pivotal astrophorid characters, such as
the traditional barrier between the astrophorids and lithistids. sanidaster and amphiaster microscleres, were not necessarily
Porifera' Demospongiae • Astrophorida 107

homologous amongst putative astrophorid taxa, and their use in other genes, but for the present these hypotheses are contentious. In
phylogenetic analysis required more careful evaluation. They also this work we define Astrophorida to contain only non-lithistid
suggested that the family Ancorinidae was polyphyletic, with two aster-bearing taxa, with tetraxon megascleres and oxeas, and a
genera (Penares and Stelletta) indicated as being closer to skeleton that retains at least a peripheral radial arrangement.
Geodiidae and Calthropellidae, respectively, implying that the Astrophorids lacking asters or tetraxons are presumed secondary
genus Penares should be reallocated from Ancorinidae to developments.
Geodiidae (even though there was little obvious morphological
support for this allocation). However, as noted by Uriz (see family Previous reviews
Ancorinidae, this volume), future phylogenetic analyses might bet-
ter evaluate these discrepancies (and the validity of an order Levi, 1973; Bergquist, 1978; Hartman, 1982; Hooper &
Tetractinellida), with the addition of further astrophorid taxa and Wiedenmayer, 1994; Chombard et ai., 1998.

KEY TO FAMILIES

(1) Microscleres include euasters ............................................................................................................................................................. 2

Microscleres are never euasters ... .............. ....... .... ... ....... ... ........... .... .... ....... .... ... .... .... ... .................... ........... .......... .... ...................... ... 3
(2) Microscleres are euasters, sanidasters or rnicrorhabds; megascleres are long-rhabdome triaenes
(sometimes absent) and oxeas .......................................................................................................................................... Ancorinidae
Microscleres are euasters, usually spherasters, sometimes also with rnicrorhabds; megascleres are irregularly arranged calthrops or
short-rayed mesotriaenes forming the deeper choanosomal skeleton, and radially orientated mesotriaenes or dichocalthrops forming
a peripheral skeleton, mostly short-rayed; with or without oxeas .............................................................................. Calthropellidae
Microscleres are sterrasters, together with euasters, spherules or rnicrorhabds; megascleres large oxeas, protriaenes, anatriaenes, and
plagiotriaenes/orthotriaenes, some of which may be almost calthrops ................................................................................ Geodiidae
(3) Microscleres are streptasters and (in most cases) rnicroxeas, microrhabds; megascleres include a variety of tetraxons (i.e., calthrops,
short-shafted triaenes, mesotriaene, mesotriaenes-derived desmas, or long-shafted triaenes) ................................... Pachastrellidae
Microscleres are metasters; sponge with special aquiferous openings; megascleres include long-shafted
triaenes ................................................................................................................................. Pachastrellidae (Thenea + Ciadothenea)
Microscleres are amphiaster streptasters; megascleres are small spiny triaenes (acanthotriaenes) with simple, bifurcate and trifurcate
clads .............. .......................... ....... .......... .......... ................. ............... ...................................................................... ......... Thrombidae
Microscleres only microrhabds ....................................................................................................................... Ancorinidae (Hoioxea)
Microscleres only sanidasters ................................................................................................................... Ancorinidae (Tribrachium)
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Ancorinidae Schmidt, 1870

Maria J. Uriz

Centre d'Estudis Avacan<;ats de Blanes (CEAB), CSIC, Carni de Sta Barbara sIn. 17300 Blanes (Girona), Spain. (iosune@ceab.csic.es)

Ancorinidae Schmidt (Demospongiae, Astrophorida) is revised to contain 15 valid genera (of 37 nominal genera): Stelletta,
Cryptosyringa, Tethyopsis, Rhabdastrella, Jaspis, Ancorina, Disyringa, Stryphnus, Ecionemia, Psammastra, Penares, Asteropus,
Melophlus, Tribrachium, and Holoxea. Some of them (e.g., Stelletta, Ecionemia) have a large representation of species widespread around
the world while some others are monotypic (e.g., Cryptosyringa, Disyringa). The main microsclere, which characterizes this family is the
euaster, but it can be absent from several species (e.g., Penares candidata, Holoxeafurtiva). The diagnostic characters traditionally used
for separating genera within Ancorinidae are the type of microscleres (euasters versus microrhabds or sanidasters), the presence or
absence (assumed by reduction) of triaenes and the presence/absence of particular inhalant and/or exhalant structures (tubes). Any possi-
ble combinations of these three types of characters outline the diagnosis of the different genera. Several of these genera are polyphyletic
in a cladistic analysis based on the structural and morphological characters at hand but are maintained here with a practical purpose until
new characters (e.g., genetic sequences) will allow a more reliable phylogenetic analysis. Ancorinid sponges live on soft, detritic, and hard
(rocky) bottoms from shallow waters to bathyal depths.
Keywords: Porifera; Demospongiae; Astrophorida; Ancorinidae; Stelletta; Tethyopsis; Cryptosyringa; Rhabdastrella; Jaspis; Ancorina;
Disyringa; Stryphnus; Ecionemia; Psammastra; Penares; Melophlus; Asteropus; Tribrachium; Holoxea.

DEF1NITION, DIAGNOSIS, SCOPE History and biology

Synonymy Ancorinidae is a large complicated family, and several

attempts have been made in the past to subdivide it into subfamilies
Ancorinidae Schmidt, 1870. Stellettidae Carter, 1875b: 67 (as (e.g., Sollas, 1888; Topsent, 1894d) using microsclere geometry
Stellettina). Euastrosa Sollas, 1886a. Coppatiidae Topsent, 1898c. and the presence/absence of a cortex as the main diagnostic charac-
[Epipolasidae] Sollas, 1888 (nomen nudum). [Streptasteridae] ters. These proposed schemes are as follows:
Topsent, 1928c: 35 (nomen nudum). Jaspidae de Laubenfels,
- Subfamily Homasterina (with one aster category). Myriastra
1936a. Seiriolidae Hanitsch, 1889.
Sollas (without cortex); Pilochrota Sollas (with cortex).
- Subfamily Euasterina (with two euaster categories). Anthastra
Definition Sollas (without cortex, without tubes); Stelletta Schmidt,
Dragmastra Sollas (with cortex, without tubes); Tethyopsis Stewart
Astrophorida with long-rhabdome triaenes, which may be (with tubes).
reduced or even absent, and oxeas. Microscleres are euasters, - Subfamily Rhabdasterina (with a euaster and a microrhabd).
sanidasters or microrhabds. Ecionemia Bowerbank, Penares Gray (without cortex), Psammastra
Sollas (with cortex).
- Subfamily Sanidasterina (with a euaster and a streptaster
Diagnosis
(sanidaster or amphiaster». Ancorina Schmidt, Tribrachium
Weltner, Disyringa Sollas (with radial skeleton); Stryphnus
Encrusting, irregularly massive or clearly spherical sponges,
(without radial skeleton).
in some cases with long inhalant and/or exhalant tubes. The megas-
cleres are triaenes with long rhabdome and oxeas. Microscleres However, not all the currently accepted genera of Ancorinidae
are euasters (oxyasters, spherasters, chiasters, tylasters), strep- can be unambiguously placed in a given subfamily, and conse-
tasters (sanidasters, amphiasters), and spiny or smooth quently this subfamily classification has not been followed here.
microrhabds. Several members of Ancorinidae display characteristic growth
habits, with particular aquiferous structures, which also involve a
Scope specific and distinct distribution of the different spicule types.
Inhalant or exhalant orifices may be clustered in some areas and or
This family contains a large number (37) of nominal genera, be placed on long, more or less complex tubes.
of which 15 are recognized as valid here. The presence of triaenes The separation of genera based on the presence or absence of
is not a constant character in these sponges, and as such I include in cortex, followed by some early authors (e.g., Sollas, 1888; Topsent,
Ancorinidae those genera previously referred to Coppatiidae bear- 1894d), has proven to be impractical given that the cortex may dif-
ing euasters, sanidasters or spiny, sanidaster-derived microrhabds fer substantially in thickness in representatives of the same genus.
such as Jaspis, Holoxea, Melophlus and Asteropus. Similarly, among the skeletal characters, the presence, abundance

108
Porifera· Demospongiae • Astrophorida • Ancorinidae 109

and type of triaenes is particularly variable, even within a given (Chombard, 1998; Chombard et al., 1998) indicated a close rela-
genus or species. The same can be said of the confused! radiate tion of Penares with Erylus and Pachymatisma which would sug-
skeleton. Variability in all these characters provides no support to gest moving the genus Penares from Ancorinidae to Geodiidae.
maintain those genera without triaenes and with a more-or-Iess However, additional examples of 'true' Ancorinidae must be exam-
confused skeleton separate from Ancorinidae, which have asters ined to allow a more complete phylogenetic assessment of this
and spiny microrhabds as microscleres (usually included in the hypothesis before a decision can be taken
family Coppatiidae). Coppatiidae has been frequently maintained
for practical reasons, although its polyphyletic nature has been rec-
ognized (Hajdu & Soest, 1992; Kennedy, 2000), and several rea- Remarks
sons suggest they be included in Ancorinidae. These include the
absence of triaenes in several genera otherwise considered 'true' The concept of Ancorinidae has been expanded here, accord-
Ancorinidae, with the presence of triaenes not a synapomorphic ing to Hooper et al. (2000) and Kennedy (2000), and as was previ-
character. Consequently, we are including in Ancorinidae those ously suggested by Hajdu & Van Soest (1992), to include several
genera of Coppatidae bearing euasters, sanidasters or spiny, genera traditionally allocated to the separate (although recognized
sanidaster-derived microrhabds. Other coppatid genera have more as artifIcial) family Coppatiidae. Different members of this family,
dubious affinities and cannot be included at present in Ancorinidae which lack triaenes as a presumed synapomorphy, appear to be
(e.g., Jasplakina de Laubenfels, 1954 and Lamellomorpha related to members of Astrophorida due to the presence of large
Bergquist, 1968). Conversely, microsclere geometry appears to be monaxons and some form of aster or aster-derived microsclere.
a more consistent character to differentiate genera and species in Some genera of Ancorinidae (e.g., Rhabdastrella) have vestigial
this family, although differences between sanidasters and spiny triaenes and support the contention that triaene-free genera may be
microrhabds are not always clear. Our current knowledge on the members of Ancorinidae.
phylogeny of these taxa has not progressed much since the early
classic authors (e.g., Lendenfeld, Sollas, Dendy, Topsent etc.) Previous reviews
because no new diagnostic characters are available since the time
when the fIrst descriptions were made. The only studies addressed Sollas, 1888; Lendenfeld, 1903, 1907; Dendy, 1916c; Levi,
to assess the homology of morphological characters in Astrophorid 1973; Bergquist, 1978; Hartman, 1982; Wiedenmayer, 1989;
sponges based on molecular data (i.e., the 28S rNA gene) Hooper & Wiedenmayer, 1994; Kennedy, 2000.

KEY TO GENERA

(1) Only euasters (and occasional trichodragmata) as microscleres ....................................................................................................... 2

With other microscleres apart from trichodragmata (with or without euasters) ............................ ................ .......... ...... .................... 3
(2) With an ectosomallayer of paratangential oxeas (triaenes absent) ........................................................................................... Jaspis
Without an ectosomallayer of paratangential oxeas .... ...................... ................................ .................................................... ........... 4
(3) Euasters and streptasters or microrhabds ........................................................................................................................................... 5
Only sanidaster-like microrhabds and (sometimes) trichodragmata ............................................................................................... 14
(4) Oxyasters, chiasters or tylasters (these two last types may have a marked centrum (small spherochiasters or spherotylasters) ...... 6
Large oxyspherasters or sterrospherasters among the euasters (triaenes may be rare or absent) ................................. Rhabdastrella
(5) Euasters and streptasters (sanidasters or amphiasters) ...................................................................................................................... 8
Euasters and microrhabds .................................................................................................................................................................. 9
(6) With a tubular aquiferous structure with several ducts ....................................................................................................... Tethyopsis
Without tubular structures .................................................................................................................................................................. 7
(7) Minute (less than 5 mm long) ovoid sponge with the exhalant (sieve) and inhalant areas at the opposite sides ......... Cryptosyringa
Normal size, massive sponge, without polarisation of the inhalant and exhalant areas ......................................................... Stelletta
(8) Without triaenes .................................................................................................................................................................. Asteropus
With triaenes .................................................................................................................................................................................... 12
(9) Smooth microrhabds ............................................................................................................................................................... Penares
Spiny microrhabds ........................................................................................................................................................................... 10
(10) Without triaenes ................................................................................................................................................................. Melophlus
With triaenes .................................................................................................................................................................................... 11
(11) Thick cortex ..................................................................................................................................................................... Psammastra
Cortex thin or indistinguishable ......................................................................................................................................... Ecionemia
(12) With short-shafted triaenes ................................................................................................................................................. Stryphnus
With long-shafted triaenes ............................................................................................................................................................... 13
(13) With tubular structures ........................................................................................................................................................ Disyringa
Without tubular structures .................................................................................................................................................... Ancorina
(14) With triaenes; only sanidasters as microscleres; simple tube ......................................................................................... Tribrachium
Without triaenes, sanidaster-like microrhabds; without tubes ............................................................................................... Holoxea
110 Porifera· Demospongiae • Astrophorida • Ancorinidae

STELLEITA SCHMIDT, 1862 pointed, 6-25 /-Lm in length. Skeletal arrangement. Orthotriaenes
and oxeas radiate at the sponge periphery, with the clads tangential
Synonymy to the sponge surface. Oxeas and occasional protriaenes protrude
the sponge surface producing hispid areas. Oxeas densely placed
Stelletta Schmidt, 1862. Myriastm Sollas 1886a: 188. Pilochrota but disarranged in the central zone of the sponge. Oxyasters and
Sollas, 1886a: 189. Anthastm Sollas, 1886a: 191. Dmgmastm Sollas, chi asters spread thorough the sponge.
1888: 187. Dorypleres Sollas, 1888: 426. Astrella Sollas, 1886a: 193. Remarks. The tenn 'somal' is used here in the sense of the
Cryptotethya Dendy, 1905: 110. Not [Cryptotethya] de Laubenfels, older literature. This tenn refers to those spicules which are not
1949. ? Monotria de Laubenfels, 1936a: 179 (incertae sedis). exclusively restricted to either the ectosome or choanosome, but
are present throughout the sponge.
Type species Some variation in spicule size has been observed in specimens
of the type species recorded from different localities. Thus, speci-
Stelletta grubei Schmidt, 1862 (by subsequent designation; mens from the Mediterranean (Topsent, 1894d; Uriz, 1981) have
Burton & Rao, 1932). smaller spicules than those from the North Atlantic (Arndt, 1935).
Although the genus Dorypleres Sollas, 1888 has been consid-
Definition ered to be synonymous with Jaspis by previous authors (e.g.,
Bergquist, 1968, Sanders et al., 1999), based on similarities in their
Ancorinidae with euasters without a marked centrum spicule complement, it has clear affinities with Stelletta, as stated
(oxyasters, chi asters and tylasters) as the main microscleres. by Burton and Rao (1932) and Kennedy (2000). The type species
(D. dendyi Sollas 1888), D. affinis (Carter 1879b, as Hemiasterella)
Diagnosis

Massive sponges with a more-or-Iess collagenous rich cortex;

triaenes often abundant, more rarely absent, oxeas, and from one to 100 ~m s.b 20 ~m c.d
three types of euasters, one of them confined to the choanosome,
the other(s) sparse thorough the sponge. Occasional accessory
ortho- or trichodragmata.

Previous reviews
A
Sollas, 1888; Topsent, 1894d; Lendenfeld, 1903; Burton & Rao,
1932; Uriz, 1981; Hooper & Wiedenmayer, 1994; Kennedy, 2000.

Description of type species

Stelletta grubei Schmidt, 1862 (Fig. 1).

Synonymy. Stelletta grubii Schmidt, 1862: 46. Stelletta
boglicii Schmidt, 1862: 47. Tethya collingsii Bowerbank 1866: 87;
Stelletta collingsii; Sollas, 1888: 181. Tethya schmidtii Bowerbank,
1866: 89; Collingsia schmidtii Gray, 1867a: 541. Collingsia
samiensis Gray, 1867a: 544. Stelletta anceps Schmidt, 1868: 31;
Astrella anceps; Sollas, 1888: 181. Ecionemia coactum
Bowerbank, 1874: 269; Stelletta coactum; Sollas, 1888: 185.
Material examined. Syntypes (holotype not fixed):
LMJG15702 - Quarnero; LMJG15272 - Zlarin, Adriatic. Other
material. Specimens of Stelletta grubei: CEAB.POR.BIO.013a,
13b, 013c, 013d, 013e, 013f. Carter's material of Pilochrota
lactea: BMNH 468-EK3. Stelletta normani: BMNH 10:1:1:571 -
Nonnan Collection.
Description. Sponge subspherical or irregularly massive.
Surface even or hispid according to the zones, with encrusted for-
eign debris. Color from white to gray. Cortex coriaceous,
15000-3000/-Lm thick. Small oscula sparse. Ostia cribriporal.
Megascleres: oxeas straight, fusifonn, sharply pointed, 1600-2700 X
40-50/-Lm in size; orthotriaenes with straight or slightly curved
rhabdome, 1000-2000 X 40-60 /-Lm in size; clads rather thin,
30-40/-Lm in thickness, abruptly curved at a distance of about 70
/-Lm from the origin and with a total length up to 150/-Lm.
Microscleres: somal chi asters with 8-10 actines, 6-8 /-Lm in length, Fig. 1. Stelletta grubei Schmidt, 1862. Spicules from specimen CEAB.
cylindrical, ending in a swelling, and a total diameter of 11-18 /-Lm; POR.BIO.013a. A, oxeas. B, orthotriaenes. C, choanosomal oxyasters.
choanosomal oxyasters with 6-12 actines smooth, conical, sharply D, somal chiasters.
Porifera· Demospongiae • Astrophorida • Ancorinidae 111

and D. splendens de Laubenfels, 1954 lack the paratangential ecto- Description of type species
somal layer of oxeas typical of Jaspis. In contrast, as in Stelletta,
the oxyasters form an ectosomal layer. The reduction or total loss Cryptosyringa membranophila Vacelet, 1979c (Figs 2 and 3).
of triaenes is a recurrent character in species of different genera of Synonymy. Cryptosyringa membranophila Vacelet,
Ancorinidae and, consequently, does not seem to be synapomor- 1979c: 33.
phic, as Hajdu & Van Soest (1992) suggested. De Laubenfels' Material examined. Holotype, paratypes (3): MNHN.D.
description (1954) of the species, Dorypleres splendens, highlights JV78.1 - with single registration number (the holotype is labeled);
another character making the genus more distant from Jaspis, viz., central Atlantic, Jamaica, coral reef cave, 20 m in depth.
the surface protuberances in Dorypleres (Sanders et al. , 1999), Description. Typically pyriform, small sponge, 2.3-3.5 mm
which are the products of the radial multispiculate tracts of oxeas, high, 1.8-2.1 mm wide, with a globular base and a narrower,
in contrast to the confused arrangement of oxeas in Jaspis . rounded apical zone. The inhalant orifices are located at the apical
The genus Monotria de Laubenfels, 1936a: 179 (with type zone, while the oscula are clustered together in a well-delimited
species Coppatias solidissirrw Wilson, 1902) is also a possible sieve at the bottom of the globular part. The sponge is inserted in a
ancorinid, with some triaenes mentioned by the original author, and foreign membranous bag with just the exhalant sieve piercing the
thus inferring possible synonymy with Stelletta. De Laubenfels membrane and the rest of the body inside the bag. The surface is
(1936a) states that the taxon is characterised by oxeas and triods, even, without foreign debris. The color is white both in living
with the latter plausibly regarded as calthrops that have lost one of specimens (Vacelet, 1979c) and after preservation in alcohol. The
the four rays. This assumption suggests that the genus should be
placed in Calthropellidae, due to possession of reduced calthops.
However, re-examination of the type specimen slide (in the USNM,
courtesy of Klaus Riitzler, pers. comm.), found neither triaenes nor
calthrops, suggesting that the taxon was inaccurately described and
that it has more probable affinites with Ancorinidae than with
Calthropellidae. Consequently, the nominal genus is included here
as Ancorinidae incertae sedis within Stelletta, but requires more
detailed study to confirm this hypothesised synonymy.

Distribution

Cosmopolitan.

CRYPTOSYRINGA VACELET, 1979

Synonymy

Cryptosyringa Vacelet, 1979c: 34.

Type species

Cryptosyringa membranophila Vacelet, 1979c (by monotypy).

Definition

Ancorinidae with oscula and ostia clustered at the opposite

sides of the sponge's longest axis. Megascleres are strongyles,
oxeas, and dichotriaenes. Microscleres are euasters (tylasters
and/or spherasters).

Diagnosis

Sponge, small, pyriform, with inhalant orifices at the narrow

apical part, and oscula clustered in a well-delimited area at the oppo-
site zone. Megascleres are mainly strongyles, which form a conden-
sation following the longitudinal axis, and are also arranged radially
in the choanosome, dichotriaenes with the cladome tangential to the
sponge surface and the rhabdome inwards. The somal microsclere is
a tylaster. Spherasters are confined to the basal exhalant zone.

Fig. 2. Cryptosyringa membranophila Vacelet, 1979c. A, lateral view of a

Previous reviews paratype with the posterior zone inserted (bottom on the left) in the foreign
membrane. B, paratype showing the sieve area on the bottom (arrow) where
Vacelet, 1979c. the exhalant canals flow.
112 Porifera· Demospongiae • Astrophorida • Ancorinidae

consistency is fleshy. The spicule complement is weak. Remarks. This is a monotypic genus. The very odd and
Megascleres: subtylotes to strongyles straight or slightly bent, with cryptic habitat of the type species, if this is a generic adaptation,
an axial vesicle clearly discernible in the tyles, 500---ti50 X 5-7 /Lm makes it potentially difficult to find more species of this genus.
in size; thin oxeas, probably corresponding to immature subtylotes,
can only be rarely found; fragile dichotriaenes, with clads and
rhabdome ending in rounded points; deuteroclads (150--160 X TETHYOPSIS STEWART, 1870
4.4.5/Lm) much longer than protoclads (20 X 4-5 /Lm) and rhab-
dome straight, 425-550 X 4-7 /Lm in size; occasional protriaenes Synonymy
with one or two undivided clads can also be found. Microscleres:
tylasters, 5-6/Lm in diameter, with actines ending in a flat knob; Tethyopsis Stewart, 1870: 281. Monosyringa Brl'lndsted,
oxyspherasters of the basal sieve, 18-26/Lm in diameter, with con- 1924: 441.
ical actines, and a centrum of about 5 /Lm (the occasional large
oxyasters with few actines, 12.5-30 /Lm long, described by Vacelet Type species
(1979c) in the holotype may be foreign spicules since they are not
present in the paratype). Skeletal arrangement: axial condensation Tethyopsis columnifer Stewart, 1870 (by monotypy).
of subtylotes, which also are spread in a radial manner; dichotri-
aenes at the sponge periphery with the clads tangential to the sponge Definition
surface and the rhabdome directed radially inwards, together with a
couple of subtylostrongyles. The oxyspherasters are exclusively Ancorinidae with a four- to six ducted tube. Choanosomal
found at the inner face of the exhalant, sieve area. The tylasters megascleres are oxeas and orthotriaenes to plagiotriaenes.
accumulate at the periphery forming a dense layer although they can Particularly modified orhotriaenes form the tube skeleton.
also be found thorough the sponge (somal microsclere). Microscleres are euasters, to which orthodragmata may be added.
Distribution and ecology. Known only from the type locality.
The sponge grows in very particular microhabitats consisting of Diagnosis
small (10--25mm in diameter) crevices of the rock covered by a
foreign membrane. From 3-10 small individuals live together in Globular sponge with a four- to six ducts tube, with exhalant and
each cavity. inhalant (?) functions. Sponge body with a radiately arranged skeleton.
Megascleres are oxeas and orthotriaenes to plagiotriaenes in the
sponge body and modified orthotriaenes with a clad much longer than
the other two, or orthodiaenes (also with a clad much longer than the
other one). Microscleres are chiasters to spherochiasters and oxyasters.
Orthodragmata may be present. Sanidasters are absent.

Previous reviews

~A' 8
Stewart, 1870; Sollas, 1888.

Description of type species

Tethyopsis columnifer Stewart, 1870 (Figs 4 and 5).

Synonymy. Tethyopsis columnifer Steward, 1870: 281.
Material examined. Holotype: BMNH 1870.11.2.1 -
Philippines.
A Description. Globular sponge, 3.5 cm in diameter, with
the basal part tom away, covered by foreign debris. Cortex 350--420
/Lm thick. From the sponge apex arises a cylindrical tube, free of
foreign debris, rounded at the distal end. Four main conducts run
along of the tube. The tube surface protrudes into conical projec-
tions. Inhalant orifices are present on the main sponge body, on the
rare areas free of foreign bodies and, probably, on the tube walls.
E The tube, as in other Ancorinidae with complex tubes, seems to

*
serve as both inhalant and exhalant functions. Megascleres of the
main body: plagiotriaenes with a conical, straight or curved rhab-
~ dome, 5000--6654 X 52-118 /Lm in size, which ends in a very thin

~
D ~ ~ 20l1 m A'.C.D,E
point; clads relatively short and stout, 190--386 X 79-85/Lffi in size;
oxeas 5400--7000 X 60--90 /Lm rare according to Sollas (1888).
f1.
o I'm Megascleres of the tube: orthotriaenes, 5800---ti800 X 85-110 /Lm in
~
A,S
A' size, with one clad much more developed (370-400 /Lm long) than
Fig. 3. Cryptosyringa membranophila Vacelet, 1979c. Spicules from the other two. Microscleres: spherasters (spherochiasters) with
the paratype. A-A', subtylotes. B-C, dichotriaenes. D, tylasters. E, short rounded actines, 7-9/Lm in diameter; oxyasters with numer-
oxyspherasters. ous conical, pointed actines, 13.6-25/Lm in diameter (the smallest
Porifera· Demospongiae • Astrophorida • Ancorinidae 113

Fig. 4. Tethyopsis columnifer Stewart, 1870 (holotype). A, transverse section showing the radial arrangement of megascleres. B, external surface of the
spherical body completely covered by foreign debris.

walls. Plagiotriaenes and oxeas, arranged in radiate tracts, form

the skeleton of the spherical sponge body. Chiasters are spread
through the sponge; oxyasters are mainly concentrated in the
choanosome.
Remarks. Wilson (1925) expanded the genus diagnosis of
Sollas (1888) by stating that "euaster forms may be slightly modi-
fied in the direction of a streptaster" in order to accommodate his
new species Tethyopsis dubia, in which (according to his figs

A A' c D
45a-c), sanidasters are clearly present whereas in the diagnosis of
this genus Sollas (1888) specifies that sanidasters are absent.
Furthermore, plagiotriaenes and orthodiaenes were described in
T. dubia. As a whole, the spicule complement of T. dubia resembles
more that of a Disyringa (with sanidasters) than to that of
Tethyopsis. Sollas (1888) also reports on the presence of choanoso-
mal chiasters in the holotype. However, he describes these chiasters
with actines slender, "hair-like, reminding one of a young ster-
raster". This description does not match that of a chi aster but that of
the oxyasters I found from re-examination of the holotype. As for
the choanosomal orthotriaenes reported by Sollas (1888), they are
clearly plagiotriaenes with the clads directed obliquely upwards.
The genus Monosyringa BrjlJndsted (1924) (type species
M. mortenseni BrjlJndsted, 1924: 442) is a junior synonym of
Tethyopsis. The external shape, tube structure, and spicule comple-
ment are similar in the type species of both genera. The only differ-
ence is the presence of modified triaenes in the tube of Tethyopsis
500 11m a,b.c,d 20llm a,f,g
versus the modified diaenes in Monosyringa. Amazingly,
100 11m a' BrjlJndsted (1924) did not mention Tethyopsis when he erected the
genus Monosyringa, whereas he compared his new genus with
Tribrachium and Disyringa, which have sanidasters instead of
Fig,S, Tethyopsis columnifer Stewart, 1870. Spicules from the holotype. chiasters. Subsequently, Bergquist (1968) suggested that the
A- B, plagiotriaenes from the spherical body. C, orthotriaene of the tube
species Agilardiella radiata Marshall, 1883, which Sollas (1888)
with a clad longer than the other two. D, oxea. E, trichodragmata.
F, oxyasters. G, spherochiasters. placed into Tethyopsis, may be conspecific with Monosyringa
mortenseni but, according to de Laubenfels (1936a), she consid-
ered A. radiata unrecognizable and did not include T. columnifer in
her discussion.
have the highest number of actines); trichodragmata, 29.5-34 X
9-10 /Lm in size. Skeletal arrangement. The main skeleton of the Distribution
tube is made of a central spicular axis formed by oxeas and modi-
fied orthotriaenes with the longer clad directed toward the tube Pacific and Antarctic Oceans, between 54-117 m depth.
114 Porifera· Demospongiae • Astrophorida • Ancorinidae

RHABDASTRELLA THIELE, 1903 ]ASPIS GRAY, 1867

Synonymy Synonymy

[Aurora] Sollas, 1888: 187 (preoccupied; Lepidoptera Astropeplus Sollas, 1888: 416, 422. Coppatias Sollas, 1888:
(Ragonot 1887». Rhabdastrella Thiele, 1903a: 934. Diastra Row, 206. Astroplakina Dendy & Burton, 1926. Zaplethea de
1911: 300. [Aurorella] de Laubenfels, 1957: 245 (nomen nudum Laubenfels, 1950b: 32.
for Aurora Sollas, 1888).
Type species
Type species
Vioajohnstonii Schmidt, 1862 (by original designation).
Coppatias distinctus Thiele, 1900 (by original designation).
Definition
Definition
Ancorinidae without triaenes, spicules are oxeas and euasters
Ancorinidae with euasters, among which, large spherasters or without a marked centrum.
sterrospherasters are abundant.
Diagnosis
Diagnosis
Encrusting or massive sponges without triaenes; choanosomal
Ancorinidae with large spherasters or sterrospherasters, skeleton composed of oxeas irregularly interlaced, ectosomal
mainly concentrated in the cortex. Triaenes may be reduced or skeleton formed by a layer of paratangential oxeas, generally
absent in some species. smaller than those in the choanosome; microscleres are euasters
without a centrum (never spherasters).
Previous reviews
Previous reviews
Dendy, 1916c; Bergquist, 1968; Kennedy, 2000.
Topsent, 1900; Dendy, 1916c; Hadju & Van Soest, 1992;
Description of type species Sanders et al., 1999; Kennedy, 2000.

Rhabdastrella distincta (Thiele, 1900) (Fig. 6).

Synonymy. Coppatias distinctus Thiele, 1900: 56;
Rhabdastrella distincta (Thiele 1900: 56). IUOllm a
Material examined. None. Holotype is missing - Temate, 20)1m b,c.d
Moluccas, Indonesia.
Description (from Thiele, 1900). Fragment of a massive
sponge, 1 cm thick, 3.5 cm in area, blackish in color in alcohol. A
Cortex 200/LID thick. Megascleres: oxeas, fusiform, with short
points, about 850 X 25IJ.m in size: Microscleres: spherasters up to
40 IJ.m in diameter, with 15-16 conical, smooth actines; oxyasters
up to 80 IJ.m in diameter, without a centrum, with few long actines;
oxyspherasters about 15IJ.m in diameter, with a centrum of 51J.m,
with conical actines. Skeletal arrangement. Oxeas radiately
arranged from the sponge periphery toward the innermost zones
#c
where they are confusedly arranged. Spherasters form a crust at the
external zone of the cortex. Oxyasters and oxyspherasters are
dispersed thorough out the choanosome.
Remarks. [Aurora] was erected by Sollas (1888) for species
of Stelletta having large oxyspherasters. Lendenfeld (1903) merged
it with Stelletta but Dendy (1916c) convincingly considered it a good
genus and provided arguments to consider the genera [Aurora],
Diastra and Coppatias synonyms. Dendy's opinion has been fol-
D
lowed by modern authors (e.g., Bergquist, 1968). RhabdastreIla,
including its several synonyms, is characterised by a frequent reduc-
tion of triaenes, which may be completely absent in species such as
R. sterrastrosa (Row, 1911) and R. cribriporosa (Dendy, 1916c), and
the presence of a cortex formed by large spherasters.

Distribution Fig. 6. Rhabdastrella distinctus (Thiele, 1900). Spicules from the holotype
(redrawn from Thiele, 19(0). A, oxea. B, choanosomal oxyaster. C, choanoso-
Indian and Pacific Oceans. mal oxyspheraster. D, large spherasters from the cortex.
Porifera· Demospongiae • Astrophorida • Ancorinidae 115

contrast to Dorypleres, is maintained separately from Stelletta due to

the possession of a paratengential layer of oxeas at the sponge
periphery.
One species of Jaspis, J. stellifera (Carter), has been copi-
ously described or cited in the contemporary literature, particularly
that pertaining to marine natural products chemistry, but Kennedy
(2000) demonstrated that virtually all contemporary records of this
B species refer to Rhabdastrella globostellata (Carter).

c Distribution

)~ I
East and West Atlantic, Indian and Pacific Oceans,
Mediterranean and Red Sea.

ANCORlNA SCHMIDT, 1862

~ a,b Synonymy
250 Ilm c
Fig. 7. Jaspis johnstoni (Gray, 1867a). Spicules (redrawn from Dendy,
Ancorina Schmidt, 1862: 51. Sanidastrella Topsent, 1892b: 18.
1916c). A, oxyasters. B, centrotylote small oxeas. C,large oxeas. Seiriola Hanitsch, 1889: 170.

Type species
Description of type species
Ancorina cerebrum Schmidt, 1862 (by subsequent designation).
Jaspis johnstonii (Schmidt, 1862) (Fig. 7).
Synonymy. Vioa johnstonii Schmidt, 1862: 78. Definition
Material examined. Syntypes (not seen): LMJG 15256,
15258, 15268, 15649 - Adriatic Sea, Sebenico; see Desqueyroux- Ancorinidae with triaenes, oxeas, sanidasters and euasters.
Faundez & Stone (1992). Other material. Specimens of J. john-
stoni: MA:9(4); MA:ll; MA:31; MA:E(P3) - from caves at the Diagnosis
Balearic Islands (Spain, western Mediterranean).
Description. Thinly encrusting sponge inhabiting rock Ancorinidae with a conspicuous cortex, with triaenes and
crevices or covering bivalve shells. Surface smooth, perforated by oxeas as megascleres and sanidasters and euasters (chiasters,
sparse oscules, 0.4-0.5 mm in diameter. Cortex undifferentiated. tylasters or oxyasters) as microscleres.
Ectosome reinforced by paratangential small oxeas and asters.
Megascleres: oxeas fusiform and slightly curved, of variable size Previous reviews
from 70 X 1.2 f.Lm up to 1000 X 15 f.Lm, the larger ones predomi-
nantly located in the choanosome and the smaller in the ectosome; Sollas 1888; Hooper & Wiedenmayer 1994.
the smallest ones may be centrotylote. Microscleres: oxyasters
without a marked centrum, with 6-10 conical, sharp-pointed Description of type species
actines, 4-13 f.Lm long (total diameter 10--28 f.Lm). Skeletal
arrangement: the large oxeas are radial at the sponge periphery and Ancorina cerebrum Schmidt, 1862 (Fig. 8).
confused toward the interior. The small oxeas are arranged tangen- Synonymy. Ancorina cerebrum Schmidt, 1862: 51.
tially to the sponge surface. Oxyasters dispersed thoroughout the Ancorina verruca Schmidt, 1862: 52.
sponge but more abundant in the peripheral region. Distribution. Material examined. Syntypes: 2MB2414 (slides), 2MB
Adriatic; Mediterranean, (coasts of France and Spain); Atlantic 6441 (1 slide), BMNH 1867.3.11.12 - Adriatic Sea (holotype not
(Azores and Cape Verde). designated).
Remarks. The monotypic genus Zaplethea de Laubenfels, Description. Large globose, lobate sponge, with a folded
1950b has a similar spicule complement to Jaspis. According to surface like that of a human brain. Cortex up to 3 mm thick with
Sanders et al. (1999), it only differs from Jaspis in possessing double- a collagen rich outer layer and a fibrous inner layer. Megascleres:
bent small oxeas, which does not seem to justify a different genus. oxeas up to 3000 X 51.6-58 f.Lm; orthotriaenes to dichotriaenes
Moreover, these double-bent spicules have also been described in the with a rhabdome 3250--3400 X 39--60 fLm in size, protoclads
type species of Jaspis (Topsent, 1900). The type species of Zaplethea 60--65 f.Lm, deuteroclads 75-80 f.Lm; anatriaenes with rhabdome
(Z digonoxea de Laubenfels, 1950b, holotype USNM 22746) has 2800--3400 X 26-29 f.Lm in size, clads 100 f.Lm long, 150 f.Lm chord
been considered synonymous with the type species of Jaspis in previ- length. Microscleres: ectosomal sanidasters, 7-8 f.Lm long; somal
ous works (see Sanders et al., 1999), although these authors did not chi asters with actines slender, ending in a swelling (tylasters),
take into account the very disjunct distribution of the two species 8-9 f.Lm in diameter. Skeletal arrangement. Triaenes with the
(Mediterranean-Adriatic versus Indo-Pacific, respectively), which cladome in the cortex and the rhabdome inwards; oxeas radial at
alone casts some doubt on this synonymy. The genus Jaspis, in the sponge periphery and confusedly arranged toward the interior.
116 Porifera· Demospongiae • Astrophorida • Ancorinidae

Sanidasters concentrated in the cortex; chiasters dispersed throughout Description of type species
the sponge. Distribution. Adriatic, Zara and Quamero, in a wide
bathymetrical range. Disyringa dissimilis (Ridley, 1884a) (Fig. 9).
Remarks. Following Sollas (1888), Lendenfeld (1903), Synonymy. Tethyopsis dissimilis Ridley, 1884a: 447.
Wilson (1925), Levi (1973) and Hooper et al. (2000), among Material examined. None. Holotype and paratype (not
others, I consider Ecionemia as a genus different from Ancorina. seen): BMHH 1882.2.23.220 - consisting of fragments (tubes);
However, the main spicule difference between species of both gen- according to Fry & Fry (1979) these and some of Sollas' (1888)
era (presence of sanidasters in Ancorina versus spiny microrhabds specimens correspond to different species. Type locality - Port
in Ecionemia) is not always clear because it is not always possible Darwin, 13-22 m in mixed sand and mud bottom.
to determine whether or not the spiny microrhabds have a Description (from Fry & Fry, 1979). Free sponge with a
sanidaster origin by reduction of the actine length. spherical body, which produces a long cylindrical whole tube
(exhalant tube) ending in a single opening, and at the opposite side
Distribution of the sponge another long, cylindrical tube containing four ducts,
which are arranged symmetrically around a central spicule axis.
Mediterranean, Atlantic, Indian, and Pacific Oceans. The latter tube ends blindy at a solid pad of packed oxea, which
extend radially to give the appearance of a fringed funnel. Surface
even, rough to touch. Ostia forming clusters, 0.57 mm in diameter,
DISYRINGA SOLLAS, 1888 along the walls of the inhalant (four-ducted) tube. Ostia inconspic-
uous. Megascleres: somal oxeas fusiform, straight or gently
Synonymy curved, variously pointed, 4.641 X 59.3 fLm in size. Orthotriaenes
with conical, sharply pointed clads, 470 fLm long, and straight,
Disyringa Sollas, 1888: 161. sharply pointed rhabdome, 3600 X 50 fLm in size; they derive into
asymmetrical orthodiaenes (with a clad longer than the other one)
Type species and orthomonaenes by reduction of one or two clads. The orthodi-
aenes in the tube have the long clad much longer (700-2142 fLm)
Tethyopsis dissimilis Ridley, 1884a (by original designation). than those spicules in the spherical body. In some cases, the

Dermition

Ancorinidae of subspherica1 growth form, with inhalant

and exhalant tubes placed at opposite sides of the sponge. Mega-
scleres: oxeas, orthotriaenes, orthodiaenes and orthomonaenes.
Microscleres: oxyasters, sanidasters and orthodragmata.

Previous reviews
A B c D
Sollas, 1888; Ridley, 1884a; Hentchel, 1912; Fry & Fry, 1979.

1 t
E
A B D

500 11m a.b.c,d

20 11m e ,f

250 11m a,b,c 100 11m d

Fig. 9. Disyringa dissimilis (Ridley, 1884a). Spicules (redrawn from
Fig. 8. Ancorina cerebrum Schmidt, 1862: spicules from the holotype Sollas, 1888). A, oxea. B, orthodiaene from the spherical body. C,
(redrawn from Schmidt, 1862). A, orthotriaene. B, D, anatriaene. C, cladome orthomonoaene from the spherical body. D, orthodiaene with an aborted
of a dichotriaene. clad, from the tube; E, sanidasters. F, oxyaster.
Porifera· Demospongiae • Astrophorida • Ancorinidae 117

monodiaenes have the only clad divided but one of the deuteroclads Description of type species
is aborted. Fry & Fry (1979) reported the presence of numerous
styles at the inhalant tube, which were not mentioned by Sollas Stryphnus niger Sollas, 1886a (Fig. 10).
(1888). Microscleres. Oxyasters with comparatively large centrum Synonymy. Stryphnus niger Sollas, 1886a: 193. Stelletta
and numerous, conical, sharp-pointed actines, 8-11.8 IJ-m in diam- mucronata Schmidt, 1868: 19; Stryphnus mucronatus; Sollas, 1888:
eter, sanidasters of normal shape, 11.8-15.8 IJ-m long; orthodrag- 193; Marenzeller, 1889: 16. Stelletta carbona ria Schmidt, 1880:
mata 23.7-27.6 X 11.8IJ-m in size, confined to the inhalant tube 280; Wehner, 1882: 52; Stryphnus carbonarius; Sollas, 1888: 192.
(called exhalant by Sollas, 1888). Skeletal arrangement. The skele- Material examined. Holotype: BMNH 1889.1.1.79 - Port
ton of the exhalant tube consists of an external layer of tangential Jackson, New South Wales, Australia, 55.5-64.7 m depth.
oxeas and an inner layer of orthodiaenes, besides numerous Comparative material. S. fortis Vosmaer, 1883: MNHN CP98-E45,
sanidasters lining the duct and tube surfaces. The skeleton of the CP98-E43. S. ponderosus Sollas, 1886a: CEAB.POR.BIOL.146a,
inhalant (four-ducted) tube is made of a central axis of oxeas and 146b,169b.
asymmetrical orthodiaenes with the long clad arising radially; Description. Large, massive, irregularly lobate sponge,
sanidasters and small styles are also abundant. The body skeleton 18 X 12 X 8 cm in size. Colour black outside, gray inside. Surface
contains oxeas, triaenes, diaenes, sanidasters, oxyasters, and generally even but rough to touch, with some hispid zones. Cortex
trichodragmata. 1750-3000 f.Lm thick. Ostia uniporal. Oscula simple, 3 mm in
Remarks. This genus is monotypic. The holotype and diameter, clustered on the top of lobes. Megascleres: oxeas large,
paratype of this species fixed by Ridley (1984) only consisted of stout, fusiform, usually curved obtusely, sharply or round pointed,
fragments of two tubes in which there were not represented all the 1563-4720 X 43-60 IJ-m; dichotriaenes small, with a rhabdome
spicule types present in the sponge. Furthermore, according to Fry conical, obtusely pointed, 430-450 X 29-36 f.Lm in size, proto-
& Fry (1979), these tubes appear not to belong to the same species. clads, 50-558 f.Lm long, projecting chiefly outwards, and deutero-
Thus, we describe here the morphology of the type species on the clads horizontal, 72-80 IJ-m long; cladome chord 254 IJ-m.
basis of the reconstruction provided by Fry & Fry (1979) who Microscleres: oxyasters with numerous, conical, sharply pointed
revised Ridley's and Sollas's material. Spicule data are taken from actines, 11.5-29.5 IJ-m in diameter; amphiasters with a straight,
the redescription of the type material by Sollas (1888). short axis and long rounded or conical actines, 13.6-19.8 IJ-m long
(included actines) and 2.3-4.5 IJ-m wide (without actines). Skeletal
Distribution arrangement. Oxeas lying in various directions, densely crowded in
the choanosome; they are tangential, oblique-or perpendicularly to
North Australian coasts: Great Barrier Reef, Arafura Sea, the sponge surface at the ectosome. The dichotriaenes, confined to
Timor Sea and Indonesia, 5.5-50m depth. the cortex, with the cladome at the sponge periphery and the rhab-
dome inwards. Amphiasters mainly concentrated in the cortex

STRYPHNUS SOLLAS, 1886

Synonymy

Stryphnus Sollas, 1886a: 193. A B

Type species

Stryphnus niger Sollas, 1886a: 193 (by subsequent designation).

Definition

Ancorinidae with large oxeas and ortho-, plagio- or dichotri-

aenes as megascleres, and euasters and streptasters (amphiasters or
sanidasters) as microscleres.

Diagnosis

Ancorinidae with large oxeas as the somal megascleres,

densely packed in the choanosome, and tangential, oblique or
pependicular to the surface at the peripheral zone. The ectosomal
megascleres include ortho-, plagio- or dichotriaenes. The micro-
scleres are a euaster and an irregular amphiaster or sanidaster.
The cortex is collenchymatous.
IOOllm a,b ~
Previous reviews 20l1m c,d
Fig. 10. Stryphnus niger Sollas, 1886a. Spicules from the holotype. A,
Sollas, 1888; Topsent, 1894d; Lendenfeld, 1903. oxea. B, dichotriaene. C, choanosomal oxyasters. D, somal sanidasters.
118 Porifera· Demospongiae • Astrophorida • Ancorinidae

although they are also present in the choanosome. The oxyasters Holotype of Hezekia demera de Laubenfels, 1934: USNM 22286-
chiefly located in the choanosome. Distribution. E. coast of dry specimen.
Australia, and Adriatic and Mediterranean Seas (the latter as Description. Ovoid, massive sponge, 5.6cm high, 4.8cm
S. mucronatus). wide, with the surface minutely hispid and the oscula dispersed.
Remarks. The disjunct distribution of this species
(Australia and Mediterranean/Adriatic) casts some doubt over the
alleged synonymy between S. niger and S. mucronatus (Schmidt).
However, there are no current spicular and morhological charac-
ters that allow us to consistently differentiate both species. The
spicules described for S. mucronatus (Topsent, 1894d) are: stout
oxeas, fusiform, straight or bent, sharply pointed, 2000-2500 X
55-60 J.Lm, dichotriaenes with deuteroclads 40-60 J.Lm long,
shorter than the protoclads, 70 J.Lm long, rhabdome straight,
400-450 X 20-25 J.Lm; amphiasters, 10-l3 J.Lm long; oxyasters
12-20 J.Lm in diameter.

Distribution

East coast of Australia, Mediterranean, Arctic, NE Atlantic

and Indian Ocean.

ECIONEMIA BOWERBANK, 1864

Synonymy

Ecionemia Bowerbank, 1862: 1101. Stellettinopsis Carter,

1879b: 348 (in part). Algol Sollas, 1888: 200. Thalassomora Fig. 11. Ecionemia acervus Bowerbank, 1864: specimen BMNH
Lendenfeld, 1888: 40. Hezekia de Laubenfels, 1934: 4. 1882.10.17.53.

Type species

Ecionemia acervus Bowerbank, 1864 (by monotypy).

A
Definition

Ancorinidae with oxeas and triaenes as megasc1eres, euasters

(but never oxyasters) and spiny microrhabds as microsderes.

Diagnosis

Massive or thickly encrusting sponges without a distinct cor-

tex. Triaenes of different types, and large oxeas as megascleres.
Microscleres include spiny microrhabds in addition to euasters.
D C'
tc- '*
Microrhabds usually form a dermal layer.

Previous reviews

Bowerbank, 1864; Sollas, 1888; Wilson, 1925.

~ *
Description of type species C
Ecionemia acervus Bowerbank, 1864 (Figs 11 and 12).
E
a. c::.
Synonymy. Ecionemia acervus Bowerbank, 1864: 173;
Stelletta acervus Ridley, 1884a: 627. Ecionemia rotundum Sollas,
= =>
£:> am
1888: 198. Ancorina nova-zealandiae Dendy, 1924: 301.
Material examined. Holotype: Lost, previously deposited at
the Museum of the Royal College of Surgeons, London (Stone, 500 w;n c 20. U\D...• ·.d.e.
1986) - Pacific, Fiji. Other material. BMNH 1882.10.17.53 -labeled Fig. 12. Ecionemia acervus Bowerbank, 1864: spicules from specimen
as "E. rotundum Sollas". Specimen of Stellettinopsis corticata BMNH 1882.10.17.53. A-A', anatriaene. B, plagiotriaenes. C-C', oxeas.
Carter (holotype of Algol and Stellettinopsis): BMNH 1885.3.14.8. D, somal chiasters. E, cortical rough microrhabds.
Porifera· Demospongiae • Astrophorida • Ancorinidae 119

Brown in color when dried. Megascleres: oxeas fusiform, abruptly holotype I also had occasion to see these formations and can state
pointed, 17~OOO X 43-72.5 ~m; orthotriaenes with rhabdome that they are certainly not spicules. In contrast, I found true small
strongylate, 1890--3030 X 43.5-84 ~m in size and clads 130-- euasters in the choanosome (Fig. 14). These spicules are very small
270 X 36.5-{)5 f..Lm; anatriaenes with rhabdome 1200--4000 X and scarce but they certainly belong to the sponge. Thus, this genus
19.5-29 ~m in size, cladome 110--118 ~m in chord length, and clads contains microrhabds and euasters, and consequently it cannot be
83-97.5 f..Lm long; Microscleres: microstrongyles rough, 9-13.6 X separated from Ecionemia, although the microrhabds in both gen-
2.6-3.4 ~m; chiasters with terminal tylote actines, 8-15 ~m in era (at least in the type species of both genera), appear to be typical
diameter, some of them with a conspicuous centrum (called rhabds and sanidaster-like, respectively.
spheraster by Sollas, 1888). Skeletal arrangement: central conden- Hooper & Wiedenmayer (1994) already considered H. walk-
sation of oxeas from which spicular bundles arise radially toward eri de Laubenfels, 1954 synonymous with Ecionemia acervus.
the sponge surface. Triaenes with the cladome placed immediately However, examination of the respecticve holotypes allowed me to
beneath the sponge surface and the rhabdome directed inwards. verify that E. acervus has abundant, characteristic chi asters, up to
A layer of micro-strongyles beneath the ectosome. Chi asters 16 ~m in diameter, together with the spiny microrhabds. The latter
dispersed through the sponge. Distribution. W Pacific, Indian are relatively thick (11.3-13.6 X 2.7-3.4 ~m) and most often cen-
Ocean, Indo-Pacific, Australia and New Zealand, to 180m depth. trotylote. In contrast, in H. walkeri the microscleres are only spiny
Remarks. The differences between Ecionemia and microrhabds, slender (only 1-1.5 ~m thick) and not centrotylote.
Ancorina rely mainly on the type of microsclere which accompa- Consequently, E. acervus and H. walkeri are not con specific.
nies the euaster. It is basically a sanidaster in Ancorina and a spiny After revising the type species of Algol (Stellettinopsis corti-
microrhabd in Ecionemia. However, some spiny microrhabds cata Carter, BMNH 1855.3.14, Fig. 13), I found only two triaenes
might have a sanidaster-origin, and both genera likely display in the several slides ofthe holotype. Carter (1879b) did not find tri-
strong relationship. aenes, and Sollas (188) remarked on their rarity. This is probably
The genus Hezekia de Laubenfels is also a junior synonym of why de Laubenfels (1954) erroneously placed into synonymy
Ecionemia. The type species of Hezekia (H. demera de Laubenfels, Stellettinopsis corticata and Stellettinopsis simplex (type species of
1934) was described as possessing microrhabds as the only certain Asteropus). This is another example of the reduction of triaenes in
microsclere. De Laubenfels (1934), however, reported the presence a species of a genus such as Ecionemia, which typically bears tri-
of some other kind of aster, 18 ~m in diameter, which were not aenes, and supports the relatively low diagnostic value of this
present in boiled-out spicule mounts but which were observed in spicule morphology as synapomorphic for the family.
sponge sections. He stated that these forms might be the result of
a particular arrangement of microrhabds (rosette-like) or crystal
precipitation due to specimen preservation. When revising the

D
c

4;-
E ~

F
-
100 11m a.b.c.d 20j1m &.1

Fig. 14. Hezekia demera de Laubenfels, 1934. Spicules from the holotype.
Fig. 13. Stellettinopsis corticata Carter, 1879b (type species of Algol A, oxea. B, plagiotriaenes. C, aborted triaenes which resemble tylostyles.
Sollas and Stellettinopsis Carter): holotype. D, anatriaene. E, choanosomal chiasters. F, cortical rough microrhabds.
120 Porifera· Demospongiae • Astrophorida • Ancorinidae

Distribution

Indian, Pacific and Atlantic Oceans.

PSAMMASTRA SOLLAS, 1886

Synonymy

Psammastra Sollas, 1886a: 195. Rhabdodragma Dendy,

1916c: 239.

Type species

Psammastra murrayi Sollas, 1886a (by monotypy).

Definition

Ancorinidae with a thick, collagen-rich, conulose cortex, and

spiny microrhabds (microstrongyles) and euasters as microscleres.

Diagnosis

Sponges characterized by a conulose surface, thick collagen-

rich, cortex often completely encrusted with thin sediment, oxeas
and triaenes with short clads as megascleres, and spiny
microrhabds and from one to several types of euasters as
Fig. 15. Psammastra murrayi Sollas, 1886a: holotype.
microscleres. Trichodragmata may be present.

Previous review

Sollas, 1888.

Description of type species

Psammastra murrayi Sollas, 1886a (Figs 15 and 16). A

Synonymy. Psammastra murrayi Sollas, 1886a: 195.
Material examined. Holotype: BMNH 1889.1.1.80 (dry) -
East Moncoeur Island, Bass Strait, Victoria, Australia, 70 m depth,
sandy bottom with shells.
Description. Subspherical sponge, 4 X 5 mm in diameter, 0
with a conulose surface completely covered by a thin sand cortex. <®
The conules, irregular in size (1-5 mm high) and unevenly distrib- B Q
uted on the surface, are the result of fibro-spicular processes. Three
small oscules, 2-5 mm in diameter, lie between the conules.
Ostia, 8-15JLm in diameter, are clustered in oval sieve-plates. .'!f
C* "*~
Conspicuous cortex, 2-3 mm thick, fibrous, prolonged at the
sponge base into stout processus for attachment to the substratum.
Color brown on the sides and upper surface, pale gray on the lower
surface. Megascleres: oxeas fusiform, straight, sometimes slightly
bent along the first third of its length, mostly with very sharp
ends, occasionally with blunt ends, 1090-4290 X 20--80 JLm;
l'
Plagiotriaenes with rhabdome conical, sharply pointed, 1815- D t;l ~
3090 X 36.5-65.6JLm in size, clads very short 45-109 X 29-36.5
JLm. Cladoxeas with the cladal end rounded or strongylate, and f>
100 I'm 8
from one to four aborted clads, directed at right angles or projecting
p
forwards and/or backwards; rhabdome straight, ending in a sharp _Ol'm b.c.d
point; most of them are smaller (1800--2200 X 43-43.5JLm) than the
normal triaenes. Microstrongyles minutely spiny, sometimes with
Fig. 16. Psammastra murrayi Sollas, 1886a. Spicules from the holotype. A,
a central constriction, relatively wide, 11.5-16 X 4.5-8JLm in size. plagiotriaenes (normal and modified into cladoxea). B, rough microrhabds.
Oxyasters with a centrum more or less developed giving rise to C, oxyasters with variable centrum and actines shape. D, spherasters.
Porifera· Demospongiae • Astrophorida • Ancorinidae 121

spherasters (11.35-20.5lJ.m) and actines variable in shape Synonymy. Stelletta helleri Schmidt, 1864: 32; Penares hel-
(rounded, spiny at the ends, conical); spherasters generally smaller leri; Gray, 1867a: 542; Vosmaer, 1890: 37; Stelletta helleri; Weltner,
than the oxyasters and with a higher number of actines (I 1.3-16 IJ.m). 1882: 46; Papyrula helleri; Sollas, 1888:199; Marenzeller, 1889: 17.
Skeletal arrangement. Megascleres radiate from the surface Material examined. Holotype: Unknown - Adriatic, Lissa,
conules to the sponge inwards. Cladoxeas and plagiotriaenes 64.7 m depth. Other material. MNHN DR49.75-162 - Balgim N.O.
immediately below the cortex, and in the flbrospicular tracks, 'Cryos' Expedition; CEAB.POR. Me:56(2) and Ma:9(3) - western
which form the conules. Occasional plagiotriaenes protrude from Mediterranean.
the conules. Microstrongyles form a layer at the cortex and are also Description. Irregularly massive sponge with smooth surface,
spread thorough the choanosome. Asters are mingled together both uniporal ostia and simple oscula. Choanosomal skeleton radiate at
in the inner part of the cortex and in the choanosome. the sponge periphery. Cortex very thin and coriaceous. Color white
Remarks. TIlis genus has been considered a synonym of inside, white or dark brown outside. Megascleres: oxeas straight or
Ecionemia Bowerbank (Lendenfeld, 1903) or Ancorina Schmidt slightly bent, fusiform, 1430 X 391J.m; dichotriaenes with proto-
(Hooper & Wiedenmayer, 1994). It shares with Ecionemia the pres- clads 60-90!-Lm long, deuteroclads of 190-240!-Lm and relatively
ence of rough microrhabds, but in contrast to that genus it possesses short rhabdome, 400 X 35 IJ.m in size, fusiform, ending in a
a thick cortex. Furthermore, the cortex in Psammastra may be rounded point. Microxeas smooth, fusiform and centrotylote,
densely encrusted with finely calibrated sand. Dendy (l916c) 32-150 X 6!-Lm in size; oxyasters, with a small centrum and rough
erected the genus Rhabdodragma for Psammastra conulosa conical actines 20 IJ.m in length. Skeletal arrangement: dichotri-
Kieschnick, 1896, due to the presence of trichodragmata. As stated aenes with the cladome placed at the sponge surface and the rhab-
previously, the presence of trichodragmata appears to be a recurrent dome inwards. Centrotylote microxeas tangential to the sponge
character in species of several genera of Ancorinidae (e.g., Stelletta) surface, forming a crust at the external part of the cortex; they are
and, thus it is not considered here to be a diagnostic character at the also spread through the choanosome. Oxeas are tangential to the
genus level. sponge surface at the inner part of the cortex and arranged more-
or-less radially in the choanosome. Oxyasters in the choanosome.
Distribution Remarks. The type species is a common sublittoral
Mediterranean sponge, which can also be found at greater depths in
Pacific Ocean. the adjacent Atlantic, living in zones under the direct influence of
the Mediterranean water flow through Gibraltar Strait.
The genus Pachamphilla Lendenfeld, 1906 (type species
PENARES GRAY, 1867 P. alata Lendenfeld, 1906, by original designation), appears to be
close to Penares due to the characteristic, curved, centrotylote
Synonymy smooth microrhabds, as stated by Levi (1967b) who maintained the
genus in its original allocation within Pachastrellidae. I have not
Penares Gray, 1867a; Topsent, 1894d: 357, 1896a: 124; sufficient information to certainly assess whether or not this genus
1925c: 629; 1934a: 5; Lendenfeld, 1903: 61; Babic, 1921: 16; is a synonym of Penares or even if it is valid. Penares and
Uvi, 1957a: 204; Sara, 1961: 32; Riitzler, 1965a: 311, 1965b: 11; Pachamphilla share the presence of smooth microrhabds and the
Pulitzer-Finali, 1979: 15; Boury-Esnault, 1971: 297; Uriz, 1981: absence of oxyasters in some species (e.g., Penares sphaera
35. Papyrula Schmidt, 1868: 18. ? Pachamphilla Lendenfeld,
1906.

Type species A
Stelletta helleri Schmidt, 1864 (by monotypy). 100 ~m a.b

Definition c 50~m c.d

Ancorinidae with dichotriaenes, oxeas, smooth microrhabds

and euasters.

Diagnosis

Sponges irregularly massive with a very thin cortex, smooth

microrhabds, centrotylote or not, forming a crust in the ectosome;
euasters may be absent.

Previous reviews

Sollas, 1888; Topsent, 1894d.

Description of type species Fig. 17. Penares helleri (Schmidt, 1864). Spicules from specimen MNHN
DR49-75-162). A, oxea. B, dichotriaenes. C, centrotylote microxeas.
Penares helleri (Schmidt, 1864) (Fig. 17). D, choanosomal oxyasters.
122 Porifera· Demospongiae • Astrophorida • Ancorinidae

Lendenfeld and Pachamphilla alata Lendenfeld, respectively). two whorls of actines at each end, leaving free the central zone.
However, both genera differ in the presence of calthrops or tri- Skeletal arrangement. Skeleton dense, formed by oxeas confusedly
aenes, respectively. The presence of calthrops has been considered arranged, sometimes forming ill-defined tracks directed toward the
a distinctive character in Pachastrellidae although their diagnostic sponge surface. Oxeas densely packed tangential to the surface at
value is not clearly established the sponge periphery. Oxyasters in the choanosome. Sanidasters
form an ectosomal crust but are also present in the choanosome.
Distribution Remarks. Sollas (1888) defined this genus as "resembling
Stryphnus from which it differs only in the absence of triaenes". The
Mediterranean, Atlantic, Indo-Pacific and Antarctic Oceans. inclusion (or exclusion) of Asteropus and other related genera lack-
ing triaenes but with large oxeas and asters in (or from) Ancorinidae
has been a recurrent issue in the literature. Some authors suggested
ASTEROPUS SOLLAS, 1888 that Asteropus may belong in Ancorinidae (e.g., Van Soest &
Stentofi, 1988; Hajdu & Van Soest, 1992). Other authors placed
Synonymy them in the polyphyletic families Coppatiidae Topsent,
Epipolasidae Sollas (e.g., Dendy, 1905) or Jaspidae de Laubenfels
Asteropus Sollas, 1888: 205. (e.g., Bergquist, 1968) although for obvious practical reasons.
However, triaenes range from abundant to rare in ancorinids and,
Type species thus, it seems reasonable to think that they have been definitely lost
in some genera such as Asteropus. Asteropus shares the presence of
Stellettinopsis simplex Carter, 1879b (by monotypy). a paratangential external skeleton of oxeas with other triaene-free
genera. The interpretation of this tangential layer as an adaptive
Definition answer to strengthen a peripheral skeleton, which has lost the tri-
aene clads as well as the loss of triaenes in different ancorinid line-
Ancorinidae without triaenes, with oxeas, oxyasters and ages, as suggested by Hajdu & Van Soest (1992), seems reliable.
sanidasters. De Laubenfels (1954) wrongly considered the type species of
Asteropus (Stellettinopsis simplex) synonymous with S. corticata
Diagnosis Carter (a species of Ecionemia) due to the rarity of the triaenes in
the latter. This is another example on the variation of triaene abun-
Massive sponges, occasionally bearing long hollow fistulae. dance in an ancorinid genus, which supports de decision to allocate
Main skeleton is a dense feltwork of large oxeas, which are placed coppatiid genera in Ancorinidae.
tangential to the surface at the sponge periphery. Triaenes are
absent. Microscleres are oxyasters and sanidasters, to which tri- Distribution
chodragmata may be added.
Indian and Pacific Oceans.
Previous reviews

Dendy, 1916c; Dendy, 1924; Bergquist, 1968; Hajdu & Van

Soest, 1992; Kennedy, 2000.

Description of type species 250 Ilm a

~b,c
Asteropus simplex (Carter, 1879b) (Fig. 18).
Synonymy. Stellettinopsis simplex Carter, 1879b: 349;
de Laubenfels, 1936a: 160; Asteropus simplex Hentschel, 1909:
369; Dendy, 1916c: 251; 1924: 306; Dendy & Frederick, 1924:
494; Wilson, 1925: 327; Vacelet et a1., 1976: 23; Bergquist, 1968:
32; Van Soest & Stentoft, 1988: 31. Asteropus haeckeli Dendy,
1905: 109.
Material examined. Holotype: BMNH 1886.12.15.362
(dry) - Fremantle, Western Australia.
Description. Massive convex sponge of about 6.5 X 2.5 cm
in diameter. Surface even, irregularly undulating. Oscules clustered
in a shallow cavity on the lower surface. Consistency firm and
compact. Color brownish both after drying and in alcohol. Cortex
about 1 mm thick. Megascleres: oxeas fusiform, stout, slightly
curved, 1320-1900 X 28-35/Lm in size, sometimes suddenly bent
near one end, resembling reduced triaenes (promonaenes);
Microscleres: oxyasters 11-15 /Lm in diameter, with a small cen-
trum or without centrum, and with a variable number of rough,
conical actines. Sanidasters 14-17.5/Lm long with a slender axis, Fig. 18. Asteropus simplex (Carter, 1879b). Spicules from the holotype.
and actines either concentrated at both ends and sometimes with A, oxeas and style. B, rough oxyasters. C, sanidasters.
Porifera· Demospongiae • Astrophorida • Ancorinidae 123

MELOPHLUS TmELE, 1899 TRlBRACHIUM WELTNER, 1882

Synonymy Synonymy

Melophlus Thiele, 1899: 8. [Tribrachion] Weltner, 1882: 50 (imperfect name); de

Laubenfels, 1936a: 168. Tribrachium; Ridley, 1884a: 479; Sollas,
Type ·species 1886a: 194. Van Soest & Stentoft, 1988: 34. Kapnesolenia de
Laubenfels, 1934: 6.
Melophlus sarasinorum Thiele, 1889 (by monotypy).
Type species
Definition
Tribrachium schmidtii Weltner, 1882 (by monotypy).
Ancorinidae without triaenes, with large oxeas, oxyasters and
rough microrhabds. Definition

Diagnosis Ancorinidae spherical with a simple hollow long tube. Somal

oxeas, triaenes and diaenes as megascleres; sanidaster-like
Thickly encrusting or massive sponge with a dense feltwork of microrhabds as microscleres.
large oxeas, which are placed tangential to the surface at the sponge
periphery but in disarray within the choanosome. Microscleres are Diagnosis
oxyasters and rough microrhabds. Triaenes are absent.
Ancorinidae spherical with a simple hollow exhalant tube
Previous reviews supported by a skeleton of orthodiaenes with long clads, which ter-
minates in a simple osculum. Ostia are evenly spread over the
Thiele, 1903a; Hadju & Van Soest, 1992. spherical body. Skeletal arrangement of the main body is clearly
radial. The skeleton of the tube walls consists of vertically overlap-
Description of type species ping orthodiaenes with the rhabdome directed towards the sponge
body.
Melophlus sarasinorum Thiele, 1899 (Fig. 19).
Synonymy. Melophlus sarasinorum Thiele, 1899: 8.
Material examined. None. Holotype: presumably in 2MB -
Celebes Islands (Sulawesi), Indonesia.
Description (from Thiele, 1899). Irregularly massive sponge,
20cm high, 14cm wide (at the widest zone), with basal rhizome-like 20 lim b,c,d
prolongations up to 10 cm long. Brownish in color. Surface covered
by papillae, variable in size and form. On the upper zone there is an A
atrial cavity, 2 cm wide, 9 cm deep, with smooth walls. The cortex is
4mm thick. Megascleres: oxeas fusiform with short points
Microscleres: spiny microrhabds of three categories: (I) small,
rough, centrotylote, 18-20 X 3 fLm in size, (II) stout, fusiform, 60 X

B~
6 fLm, with round points, and (Ill) long and slender, 175 X 4 fLm in
size; oxyasters with 15-20 cylindrical actines, 15-18 fLm in diame-
ter, without a conspicuous centrum. Skeletal arrangement. The skele-
ton of the cortex consists of abundant oxeas confusedly arranged,
tangential to the surface at the sponge periphery. Oxeas are scattered
in disarray within the choanosome. Asters are located in the cortex.
Microrhabds are located both within the cortex and the choanosome.
Remarks. To be consistent with the criterion followed to
differentiate Ancorina from Ecionemia, we consider the genus
Melophus (with spiny microrhabds) is different from Asteropus
(with sanidasters among the microscleres), following the proposal
of Hajdu & Van Soest (1992). These authors also highlighted the
absence of trichodragmata in the two known species of Melophlus
(Le., M. sarasinarum Thiele, 1899 and M. cherbonnieri Levi,
1961a) versus their presence in Asteropus. However, the presence
D ~
of trichodragmata appears to have a poor diagnostic value at a
=-
genus level in Ancorinidae.

Distribution Fig. 19. Melophlus sarasinorum Thiele, 1889. Spicules (redrawn from
Thiele, 1899). A, oxea. B. medium and large microrhabds. C, oxyasters.
Pacific Ocean, Carribbean. D, small, rough microrhabds.
124 Porifera· Demospongiae • Astrophorida • Ancorinidae

Previous reviews in size, straight, fusiform, with sharp points; protriaenes to orthotri-
aenes with the rhabdome (1800-4000 X 20.5-52.2 J..Lm) straight,
Sollas, 1888; Weltner, 1882; de Laubenfels, 1934. ending in a sharp point and two types of clads: short (63-80 X
16-18.2 J..Lm), directed upwards, and long (114-295 X 34-41 J..Lm),
Description of type species first slightly upwards and then brusquely doubled perpendicularly
to the rhabdome (occasional divided clads can also be observed);
Tribrachium schmidtii Weltner, 1882 (Figs 20-21). anatriaenes, small, with short rounded clads, 13-35 J..Lm long and
Synonymy. Tribrachion schmidtii Wehner, 1882: 50; rhabdome straight or curved, sharply narrowing below the cladome
Tribrachium schmidtii; Ridley, 1884a: 479. and then enlarged at the point, which is typically rounded or even
Material examined. Holotype: missing - only consisted of in the shape of a water droplet (821-1816 X 9-13.5 J..Lm in size).
the atrial tube. Neotype (here designated): BMNH 1889.1.171D- Sanidaster-like microrhabds of two types: those from the tube (type
Bahia, 'Challenger' Expedition, 12.6-36m depth. Other material. I) are straight and thinner (9-11 X 0.5-1 J..Lm); those from the
Holotype of Kapneso/enia fisheri de Laubenfels, 1954: USNM choanosome (type II) are wider and irregularly twisted (8-11.5 X
22370 - specimen in alcohol and five slides, three of them from 2.3-2.7 J..Lm). Skeletal arrangement. Oxeas and ortho- and dichotri-
Johnson's collection. aenes, with a radial arrangement, form the main skeleton of the
Description. Sponge spherical, 0.8 cm in diameter with a spherical body. The monodiaenes, densely placed, with the rhab-
hollow tube, 4 cm long, 4 mm wide at the base, tapering toward the dome downwards, and the clads strongly entangled, form the tube
apex and ending in a single opening. Cortex 200-300 J..Lm thick. skeleton. Microrhabds of type I are extraordinarily abundant along
Inhalant orifices distributed over the spherical sponge body. Small the tube walls and also present in the globular body. Microrhabds
orifices (inhalant ?) are also located on the tube walls. The color is of type II are dispersed thorough the choanosome.
whitish outside and brownish inside, in alcohol. Megascleres of Remarks. The oxyasters described by Sollas (1888) in the
the tube: orthodiaenes with long symmetrical clads, 360-500 X specimens from Bahia are foreign spicules, as he already suspected.
22-26.5 J..Lm, and a long, completely straight rhabdome, Furthermore, Sollas (1888) did not mention the occasional division
2800-4900 X 26-95 J..Lm in size, tapering toward a very thin, sharp of clads in orthotriaenes, and thus the presence of dichotriaenes.
point. Spicules of the sponge body: oxeas 1971-6438 X 4-97 J..Lm The occasional presence of dichotriaenes and dichomonaenes has

.!!!!..1!!!2... a .b.c.c'

20 m d.e.!

Fig. 21. Tribrachium schmidtii Weltner, 1882. SpicUles from the neotype.
A-A', oxeas. B, orthodiaene from the tube skeleton. C-C', orthotriaenes to
Fig. 20. Tribrachium schmidtii Weltner, 1882. A, specimen BMNH protriaenes from the body skeleton. D, anatriaenes from the body skeleton.
1889.1.171D (neotype). Detail of the tube wall showing small (inhalant?) E, sanidaster-1ike microrhabds from the choanosome. F, rough microrhabds
orifices. from the tube.
Porifera· Demospongiae • Astrophorida • Ancorinidae 125

also been reported in other tubular species, such as Tethyopsis schmidtii, which according to Van Soest & Stentoft (1988) is con-
mortenseni (type of Monosyringa) (see Bergquist, 1968) and specific with K. fisheri, is in fact T. fisheri (but clearly different
Disyringa (Sollas, 1888), respectively. from T. schmidtii). It is possible that these two species have been
The biological function of the tube in this species is somewhat confounded in the literature. The bathymetrical distribution
controversial. Sollas (1888) considers the tube to play an exhalant reported for T. schmidtii (shallow waters (less than 12 m depth) and
function (the atrial cavity). Fry & Fry (1979) state that the small deep waters (more than 700m depth), respectively), is suspect and
openings clearly visible on the lateral sides of the tube are ostia and the specimens of T. schmidti reported from deep zones might
thus the tube would have both exhalant and inhalant functions. belong to T. fisheri.
However, the presence of these small orifices does not necessarily
guarantee their inhalant function. Similar small orifices are fre- Distribution
quently observed along the large exhalant canals of several encrust-
ing sponges (author, unpublished data) and may contribute to Central Atlantic.
decrease water pressure over the canal walls by releasing exhalant
water, and thus favoring flow speed. On the other hand, it is diffi-
cult to imagine how these small orifices, which in Tribrachium run HOLOXEA TOPSENT, 1892
into a hollow tube along which a main exhalant current flows,
could play an inhalant function. Synonymy
Kapnesolenia (Fig. 22) is a junior synonym of Tribrachium,
according to Van Soest & Stentoft (1988). However, the type Holoxea Topsent, 1892b: 26; 1900: 272; 1928c: 144; Hallmann,
species of these two genera, although they show a similar growth 1917c: 654.
habit and inhabit the same geographical region, have clear differ-
ences in their spicule complements and thus cannot be considered Type species
conspecific. Tribrachium schmidti differs from T. fisheri in the rar-
ity of the dichotriaenes and the presence of two types of sanidaster- Holoxeafurtiva Topsent, 1892b (by monotypy).
like microrhabds: those in the choanosome wider and with the axis
curved (type II), and those in the tube straight and thinner (type I), Definition
while T. fisheri has stout dichotriaenes and spiny microrhabds only
of the latter type although longer (11.5-20.4 X 0.6-2.2/-1m). Ancorinidae with oxeas, spiny (sanidaster-like) microrhabds
The specimen described by Wilson (1902) as Tribrachium and trichodragmata.

Diagnosis

Massive or encrusting growth forms with oxeas of two sizes

as megascleres and minute sanidaster-like microxeas, and raphides
as microscleres, the latter occurring singly or in trichodragmata.
Choanosomal skeleton irregularly halichondroid without a clear
axial and extra-axial differentiation. Subectosomal megascleres
present singly or in loose tracts protruding through the surface;
ectosome with smaller category of oxeas erect on surface.

Description of type species

Holoxeafurtiva Topsent, 1892b (Fig. 23).

Synonymy. Holoxea furtiva Topsent, 1892b: 26; Pulitzer-
Finali, 1983: 477.
Material examined. Schizotype: MNHN LBIM DT2414-
Bandol, Banyuls, Mediterranean. Other material. MNHN - speci-
men from La Calle (Topsent, 1898c).
Description. Thinly encrusting and endolithic sponge which
occupies small crevices between calcareous conglomerates; surface
hispid, even; consistency coriaceous; color white. Megascleres:
oxeas (I) fusiform 1155-1660 X 25-55/-1m; oxeas (II) 450-785 X
8-13/-1m. Microscleres: sanidaster-like, spiny microxeas, 18-26 X
1-2/-1m in size, covered by irregular tubercles, fusiform, straight or
slightly bent; trichodragmata small, 8-12 X 5-8 /-1m in size. Skeletal
arrangement. Ectosomal skeleton with erect or paratangentiallayer
of thinner ectosomal oxeas occurring singly or in loose bundles;
choanosomal skeleton with disorganized, nearly halichondroid
Fig. 22. Kapneso/enia fisheri de Laubenfels, 1954 (type species of
Kapneso/enia, junior synonym of Tribrachium). A, holotype. B, orthodi- criss-cross of long smooth oxeas, mostly in tracts tangential
aenes arrangement at the tube walls. C, surface of the spherical body show- to the substratum, some erect and protruding through the surface;
ing the cladome of dichotriaenes. ectosome with the smaller category of oxeas erect on surface;
126 Porifera· Demospongiae • Astrophorida • Ancorinidae

sanidaster-like spined microxeas, scattered predominantly in the

A ectosomal skeleton, less so throughout the choanosome; raphides
occur singly or in trichodragmata abundant in the choanosome.
RelTUlrks. Hallmann (1917c) suggested that Holoxea was
related to Desmoxya in spicule geometry, whereas the presence of
an ectosomal skeleton in H. furtiva may be of little taxonomic sig-
nificance since it was absent in other members of the genus. In con-
trast, Topsent (l928c) suggested that Holoxea was more closely
related to the Astrophorida (e.g., Stelletta, Dercitus), having lost
tetractinal megascleres and having developed a spicule morphol-
ogy analogous to the Desmoxyidae. Topsent's decision was
adopted by several authors (de Laubenfels, 1936a; Pulitzer-Finali,
1983) and is followed here.

Distribution
50).l.m
Mediterranean (H. furtiva Topsent), Indonesia (H. collectrix
Thiele and H. valida Thiele).

B ACKNOWLEDGEMENTS

I thank G. Agell and G. Carreras from the CEAB for their pre-
cious help in drawings and processing photographs. M. Bardaji
helped in slide preparation and spicule measurements. J. Hooper
200).l.m
provided most information and spicule photographs for Holoxea.
C. Valentine (BMNH London), J. Vacelet (SME Marseille),
Fig. 23. Holoxeafurtiva Topsent, 1892b. A, light photograph of spicUles C. Levi (MNHM Paris), K. Rtitzler (USNM Washington) and
complement from the schyzotype (MNHN LBIM DT2414) with oxeas of R. Desqueyroux-Faundez (MHN Geneva) kindly loaned holotypes
two sizes and abundant sanidaster-like microrhabds. B, large oxea. and other museum material.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Calthropellidae Lendenfeld, 1907

Rob W.M. Van Soest1 & John N.A. Hooper

'Zoological Museum, University of Amsterdam, P.O. Box 94766, 1090 GT, Amsterdam, Netherlands. (soest@science.uva.nl)
2 Queensland Museum,
P.O. Box 3300, South Brisbane, Qld, 4101, Australia. (JohnH@qm.qld.gov.au)

Calthropellidae Lendenfeld (Demospongiae, Astrophorida) is a small family of Astrophorida positioned between the major families
Ancorinidae and Pachastrellidae. With Ancorinidae (+ Geodiidae) they share the possession of euasters, and with Pachastrellidae they
share calthrops-like megascleres. There are no unique features, and it is suspected that the family may turn out to be artificial. All genera
have calthrops and euaster microscleres, and some have modified triaenes and tetractines, mesotriaenes (mesocalthrops or mesodichotri-
aenes) and oxeas. Genera are differentiated by the presence or absence of oxea megascleres, organisation of triaenes in the choanosomal
and subectosomal skeletons, and euaster morphology and diversity. Four valid genera are recognised and only about two dozen species
worldwide, with records from the North Atlantic, Mediterranean, South Africa, western Indian Ocean, Indonesia and New Zealand, found
mainly in warmer waters, moderately deep (about 600m) but also recorded occasionally from the littoral.
Keywords: Porifera; Demospongiae; Astrophorida; Calthropellidae; Calthropella; Chelotropella; Pachastrissa; Pachataxa.

DEFINITION, DIAGNOSIS, SCOPE to Calthropellidae (Pachastrissa pathologica (Schmidt, 1868)

sensu Lendenfeld, 1903, and Pachataxa enigmatica Levi & Levi,
Synonymy 1983b) shared the possession of short-shafted mesotriaenes (meso-
calthrops or mesodichotriaenes) with some representatives of
Calthropellidae Lendenfeld, 1907: 301. Pachastrellidae (Yodomia Lebwohl, Triptolemus Sollas) and
'Lithistida' (Brachiaster Wilson). He concluded that these spicules
Definition were non-homologous, and the presence of mesotriaenes in
Calthropellidae and Pachastrellidae was a product of convergent
Astrophorida with irregularly arranged calthrops or short- evolution (with Brachiaster now also included in Pachastrellidae;
rayed mesotriaenes forming the deeper choanosomal skeleton, and Maldonado, this volume).
radially orientated mesotriaenes or dichocalthrops forming a
peripheral skeleton, mostly short-rayed; with or without oxeas; Scope
microscleres are euasters, usually spherasters, never sterrasters,
sometimes also with microrhabds. Seven nominal genera are included in Calthropellidae, four of
which are probably valid although two of these are monotypic.
Diagnosis Only about two dozen species have been described, distributed in
warmer parts of the ocean, in moderately deep water (about 600 m
Encrusting, massive to subspherical globular sponges with a depth) occasionally from the littoral zone, with records from the
rough surface and hard consistency. The main skeleton tends to be north Atlantic, Mediterranean, South Africa, East Africa, Indonesia
confused, but may be radially arranged if triaenes and/or oxeas are and New Zealand.
present. Megascleres include calthrops (with rays ranging from
two to four, some of which may be bifurcate (dichocalthrops», or Taxonomic history
short-shafted triaenes, and sometimes long-shafted triaenes and/or
large oxeas; one genus (Chelotropella) has radially oriented dicho- Although Sollas (1888) described or revised most of the
triaenes and oxeas throughout the skeleton, whereas most species species now considered to belong to Calthropellidae, he still
have radial organisation only apparent in the subectosomal region, assigned them to Pachastrellidae. Likewise, Lendenfeld (1903) in
if at all, with calthrops (and other megascleres if present) irregu- his revision of the tetraxonid sponges kept them in Pachastrellidae.
larly dispersed through the choanosome. Microscleres are euasters, As late as 1907, the family was erected by Lendenfeld with the
usually spherasters, sometimes modified, and microrhabds or above given definition (here modified slightly), and three genera
microxeas. Rugose microrhabds occur in one genus (Pachataxa). assigned to it. Since then the family was retained in its original for-
mat by most authors. De Laubenfels (1936a: 179) erected a genus
Remarks Pachataxa for Schmidt's species Pachastrella lithistina, which was
later assigned to Calthropellidae by Pulitzer-Finali, 1978. De
Lendenfeld (1907) originally included three genera, Laubenfels (1936a) was the only author to reject Lendenfeld's family.
Calthropella, Pachastrissa, and one new genus ChelotropeUa in his He completely mixed up members of Calthropellidae, Pachastrellidae
family Calthropellidae for sponges similar to Pachastrellidae but (Astrophorida) and Plakinidae (Homosclerophorida), and distributed
with euaster microscleres. One additional genus, Pachataxa de these over two families, Halinidae and Plakinastrellidae, whose
Laubenfels (1936a) is now included here (Maldonado, 1993). contents are obviously artificial. Not surprisingly, none of the
Maldonado (1993) also noted that a few species currently assigned major authors of the previous century followed this classification.

127
128 Porifera· Demospongiae • Astrophorida • Calthropellidae

Although there is an apparently wide acceptance among recent equiangular equal-length rays) and 'short-shafted' triaenes, of
authors of the validity of the family Calthropellidae in its present which one of the rays is longer and considered homologous to the
concept, there is ample reason to reexamine it critically, e.g., by rhabd of a long-shafted triaene.
sequencing appropriate genes and including various pachastrellid
and ancorinid genera in the analysis. There are no morphological
markers for the family, it can only be defined on the unique combi- Reviews
nation of calthrops and euasters. And even that combination is
threatened by the vague separation of genuine calthrops (with four Lendenfeld (1907), Maldonado (1993).

KEY TO GENERA

(1) Microscleres include rugose microrhabds (ataxasters) in addition to regular euasters; calthrops are irregularly arranged throughout the
skeleton; short-shafted mesotriaenes occur in some species ................................................................................................ Pachataxa
No rugose microrhabds ........................................................................................................................................................................ 2
(2) Megascleres include oxeas .................................................................................................................................................................. 3
No oxea megascleres; with calthrops irregularly arranged throughout skeleton; microscleres regular euasters .............. Calthropellll
(3) Skeleton radial throughout; calthrops (dichotriaenes) and oxeas radially orientated in the peripheral skeleton and form two distinct
subectosomal layers; long-shafted triaenes present; microscleres diverse euasters (strongylasters, acanthotylasters, spherasters,
oxyspherasters) ... ....... ........ ... ....... .... ... .......... ... ....... ............... ............. .......... ........ ... ........... ...... ............... ............ ............ Chelotropellll
Skeleton irregular except at surface; microscleres regular euasters (spherasters to spheroxyasters); short-shafted (dicho-) triaenes may
be present ........................... ........... ............................... ................................. ........... .......... ................. ................ .............. Pachastrissa

CALTHROPELLA SOLLAS, 1888 of Ancorinidae and Pachastrellidae, but are also shared with
Geodiidae, Theneidae and Spirophorida, together forming the nom-
Synonymy inal order Tetractinellida (i.e., Astrophorida + Spirophorida) (e.g.,
Chombard et al., 1998). In some genera in each of these families
Calthropella Sollas, 1888: 107. [Corlicella] Sollas, 1888: 281 long-shafted triaenes are independently lost, including most
(preocc. by COrlicella Ehrenberg, 1872, Protoctista). Corlicellopsis Calthropellidae except some species of Chelotropella, with the
Bergquist, 1968: 62. consequence that this feature may not be diagnostic above the
species or genus level.
Type species [Corlicella] Sollas, 1888: 281 was erected for type species
Corlicium stelligerum Schmidt, 1868: 25, pI. III fig. 6. This is a
Calthropella simplex Sollas, 1888: cxxxiii, 107 (by original white encrusting Mediterranean sponge, represented by type speci-
designation). mens in LMJG 15351, 15352 and 15507 (Desqueyroux-Faundez &
Stone, 1992), of which the characters overlap entirely with those of
Definition Calthropella. Sollas (1888: 281) defines the genus as having 'poly-
actinose as well as tetractinose asters', but the sizes quoted for the
Calthropellidae with megascleres only calthrops irregularly 'tetractinose' aster (rays of 330 j.lm) make it clear that these are
arranged throughout the skeleton; without long-shafted triaenes or nothing but calthrops. The true 'polyactinose' asters have 6-12 rays
oxeas; and microscleres are only regular euasters. and measure 20 j.lm in diameter. Topsent (1904b) employed
COrlicella (sensu Sollas) for what he considered a combined
Diagnosis species Calthropella geodioides (Carter, 1876) and Calthropella
simplex. However, C. geodioides in Carter's sense has oxeas, and is
Massive to thickly encrusting or globular sponges of hard here considered the type of a genus Pachastrissa (see below).
consistency, with irregular conulose, tuberculate and microhispid [Corlicella] is a synonym of Calthropella, and because the lat-
surface. Ectosomal region with a dense organic cortex and a layer ter has page priority, it is considered the senior name. Moreover,
of predominantly smaller euasters. Choanosomal skeleton with a Bergquist (1968: 62) discovered that [Corlicella] Sollas, 1888 is
confused mass of irregularly dispersed calthrops or short-shafted preoccupied by the protoctist genus COrlicella Ehrenberg, 1872.
triaenes, typically simple, with sharply pointed single rays, or For that reason, she erected Corlicellopsis (type species not fixed,
bifurcate (dichocalthrops, dichotriaenes) in some species, and here designated as COrlicium stelligerum Schmidt, 1868). The differ-
larger euasters intermingled. ence she brought up to differentiate Corlicellopsis from Calthropellll,
viz., the occurrence in the latter of triactines and bent diactines, are
Remarks generally considered of minor significance, and does not justify main-
taining two genera. Accordingly, Corlicellopsis, like Corlicellll sensu
The characters on which Calthropella is distinguished from Sollas, is considered a junior synonym of Calthropella.
the other genera of the family are particularly negative. On the one Sollas (1888) suggested Calthropella was similar to
hand, if we assume that Pachastrellidae and Ancorinidae are Pachastrellidae (at that time included in suborder Astrophorida,
closely related families, the absence of oxeas and long-shafted tri- demus Streptastrosa), and intermediate between 'typical' pachas-
aenes may be interpreted as losses, and thus as synapomorphies. trellids, like Pachastrella, and Theneidae (both the latter now
Alternatively, long-shafted triaenes are not only a synapomorphy included in a single family Pachastrellidae; see Maldonado, this
Porifera· Demospongiae • Astrophorida • Calthropellidae 129

volume). In the same monograph Sollas (1888) included the name Halinidae de Laubenfels (1934), over Pachastrellidae
[Corticella] Sollas based on its possession of polyactinal as well as (which they erroneously attributed to Hentschel, 1923), arguing for
tetractinal aster morphologies and in this regard differing from its recognition of seniority of their respective type genera, with Halina
alleged sibling taxa Corticium and Plakortis. All three were allo- Bowerbank having priority over Pachastrella Schmidt. This action
cated to a family Corticidae Vosmaer. Sollas (1888) gave no indica- is now recognised as both contrary to the Rule of Priority (ICZN,
tion that he considered either Calthropella or [Corticella] Sollas Anon., 1999), and erroneous (i.e., Pachastrellidae was erected by
were remotely closely related. Lendenfeld (1903) subsequently syn- Carter, 1875c and has clear seniority over Halinidae). Bergquist
onymised the two taxa, referring C. stelligera to Calthropella, not- (1968) assigned the family to the order Homosclerophorida and also
ing that the possession of modified (polyactinal) asters in the former including in it the genus Pachastrella.
was not significantly different from the uniform (regular) euasters
in Calthropella. Pulitzer-Finali (1970) followed this action. Description of type species
Conversely, de Laubenfels (1936a) rejected this interpretation and
stated that Calthropella had calthrops, triads and bent diacts Calthropella simplex Sollas, 1888 (Figs lA-C).
whereas [Corticella] Sollas had only calthrops and spiny euasters. Synonymy. Calthropella simplex Sollas, 1888: 107-108,
Bergquist (1968) followed de Laubenfels' (1936a) system, adopting pI. 10, figs 13-14,21-29.

B -

Fig. 1. Calthropella Sollas, 1888. A-C, Calthropella simplex Sollas, 1888. A-B, habit and cross section (from Sollas, 1888: pI. 10, figs 13-14). C,
calthrops and spherasters (from ditto pI. 10, figs 21-29) (scales: A-B, I cm; C, 100 J.Lm (megascleres), 25 J.Lm (microscleres». D-E, 'Corticella' geodioides
sensu Topsent, 1892a, 1904b (= ? Calthropella spec.). D, habit (scale 1 em) (from Sollas, 1888 pI. 4, fig. 14). E, calthrops and spherasters (scales: 100 J.Lm
(megasc1eres), 10 J.Lm (microscleres» (from Topsent, 1904: pUO fig. 12).
130 Porifera· Demospongiae • Astrophorida • Calthropellidae

Material examined. None. Holotype: BMNH (not seen) - CHELOTROPELLA LENDENFELD, 1907
Praia, Sao Tiago, Cape Verde Islands. The type species was
recorded subsequently from the Azores, at depths of 400-800m, Synonymy
'Challenger' Expedition.
Description (from Sollas, 1888). Massive, irregular ridges Chelotropella Lendenfeld, 1907: 302.
with flattened base and rounded sides, overgrowing a species of
Vulcanella (as Sphinctrella) (Figs lA-B), together they are Type species
9X3.5X3.8cm. Surface smooth, raised in places to form small
rounded tubercles, each with a crater-like depression at the summit, Chelotropella sphaerica Lendenfeld, 1907 (by original
0.15--0.5 mm diameter and a centrally perforated floor (a sieve). designation).
Both oscules and inhalant pores are similar in having a sieve cover-
ing them. This central sieve on the tubercles leads into compara- Definition
tively large subdermal cavities. Cortex collagenous, about 0.2 mm
thick, containing numerous granular, elongated fusiform cells, with Calthropellidae with calthrops, oxeas and peripheral dichotri-
a subectosomal layer of spheraster euasters immediately below, aenes forming a radial skeleton throughout, and in the peripheral
mostly small with occasional larger ones dispersed in between. region forming two subdermal layers, and with the addition of
Choanosomal skeleton highly collagenous mesohyl with granular long-shafted triaenes; microscleres diverse forms of euasters
cells scattered throughout, a well-developed canal system traversed (strongylasters, acanthotylasters, spherasters, oxyspherasters).
by numerous prosopyles surrounded by concentrically arranged
myocytes and fusiform co11encytes radiating inwards from the mar- Diagnosis
gin; choanocyte chambers about 20 X 16 fJ.m. Spicule skeleton is a
confused mass of calthrops (Fig. 1C) divisible into two size cate- Irregular spherical globular growth form with granular surface.
gories, both with 1-4, mostly 3 conical actines, smooth, simple, Subectosomal skeleton characterized by large subdermal cavities
oxeote, tornote or strongylote, the larger with rays 800 X 90 fJ.m and with long-rayed dichotriaenes radially oriented in the upper part of
the smaller with rays 150 X 20 fJ.m long. Microscleres (Fig. 1C) are the sponge, their cladomes forming two layers parallel to the sur-
spherasters, more-or-Iess in 2 sizes, the larger with a relatively face. Skeleton comp9Sed of radial bundles of oxeas and dichotri-
large centrum, numerous actines reduced to rounded tubercles, aenes extending from the proximal part of the choanosomal skeleton
24 fJ.m in diameter, and the smaller with fewer tubercles, 12 fJ.m in to the periphery, with a more disorganized skeleton of calthrops in
diameter, with actines more slender, conical and oxeote, and most deeper parts of the skeleton. Megascleres consist of oxeas, dicho-
commonly found in the subectosomal region. triaenes, calthrops of two sizes. Microscleres consist of diverse
Remarks. Sollas (1888) originally included two species in the euaster morphologies including strongylasters, acanthotylasters,
genus, C. simplex and C. (Pachastrella) geodioides (Carter) (the lat- spherasters and oxyasters.
ter with dichocladose actines on calthrops). Following the revisions
of various calthropellid genera by Lendenfeld (1903, 1907) and Remarks
others, five species are currently included in this genus, three from the
Atlantic-Mediterranean region (c. recondita Pulitzer-Finali, 1970, This genus was erected by Lendenfeld (1907) for a single
C. simplex So11as, 1888, and C. stelligera (Schmidt, 1868», and two species which he considered was intermediate between other calthro-
from the Indo-Pacific (C. novaezealandiae (Bergquist, 1961c), pellids and members of the Stellettidae. It differs from other nominal
C. digitata Pulitzer-Finali, 1993). A possible further species is the calthropellids in possessing two distinct layers of dichotriaene
sponge described by Topsent (1904b) as Corticella geodioides, (see cladomes, parallel to the surface, one in the peripheral part of the
Figs 1D-E), which does not appear to belong to Pachastrissa geodi- subectosome and the other at the junction of the cavernous subecto-
oides (cf. below) as Topsent did not describe the presence of oxeas. some and choanosome, and possibly also by its diverse euaster mor-
Other species included in Calthropella at one time or another, phologies not seen in other species. The genus remains monotypic.
C. inopinata Pulitzer-Finali, 1993, Stelletta pathologica Schmidt,
1868, and Pachastrella geodioides Carter, 1876 were subsequently Description of type species
referred to Pachastrissa. So11as (1888: 111) also referred
Pachastrella exostitus Schmidt, 1868 to Calthropella (with question Chelotropella sphaerica Lendenfeld (Figs 2A-C).
mark), based on its supposed similarities in spherasters with those of Synonymy. Chelotropella sphaerica Lendenfeld, 1907:
C. geodioides and C. simplex, but he also noted that the former had 302-304, pI. 34, figs 1-7; Pulitzer-Finali, 1993: 254; Levi, 1969:
tuberculated microrhabds, and in this respect he indicated that it 954; Levi & Levi, 1983b: 150.
might have closer affinities to Derr:itus (pachastrellidae). Pulitzer- Material examined. None. Holotype (not seen): 2MB -
Finali (1983) suggested further that C. pathologica, C. geodioides Agulhas Banks, South Africa coast, 'Valdivia' Expedition.
and C. simplex may constitute a single, variable species given simi- Description (from Lenden/eM, 1907). Growth form is
larities in their spiculation and apparently only minor differences in irregular, globular spherical with a granular surface. Small oscules
other morphometric characters, but no decision is possible to con- 1-2mm diameter arranged in groups. Ectosomal cortex about 700
firm or refute this hypothesis, which requires a far more detailed fJ.m thick covered by a thin dermal membrane and overlying a cav-
study of the respective type material than is possible here. ernous subectosome free of choanocyte chambers. Choanosomal
region with spherical choanocyte chambers, 15-20 fJ.m diameter.
Distribution Choanosomal skeleton (Fig. 2C) composed of radial bundles of
longitudinal long-rayed dichotriaenes with calthrops ('chelotrope')
Mediterranean, NE Atlantic, SW Pacific. and euasters. Radial bundles of slender oxeas and dichotriaenes
Porifera' Demospoogiae • Astropborida • Caltbropellidae 131

Fig. 2. Chelotropella, Pachastrissa, Pachataxa. A~, ChelotropeUa sphaerica Lendenfeld, 1907. A, dichotriaenes, calthrops, oxeas (scales: left 500 j.l.m, right
1000 j.l.m). B, strongylasters, acnthotylasters, spherasters and oxyaster euasters (scale 25 j.l.m). C, cross section of peripheral skeleton (scale 1000 j.l.m) (from
Lendenfeld, 1907, pI. 34 figs 1-7). D, Pachastrissa geodioides (Carter, 1876 as Pachastrella), habit (scale 1cm), oxeas, calthrops, short-shafted triaenes
and dichotriaenes (scale 500j.l.m) and microscleres (scale 25j.l.m) (from Carter, 1876: pI. 14 fig. 23). E, Pachataxa lithistina (Schmidt, 1880b as
Pachastrella), microrhabds (scale 10 j.l.m) (from Schmidt, 1880b: pI. 9 fig. 3).

extend from the proximal portion of the centre of the sponge to the between calthrops and triaenes. Calthrops are mostly regular in
distal portion of the choanosome, just below the cortex. Cladomes shape but differentiated into two size classes, the large ones are
of dichotriaenes are arranged in two layers above each other, one at regular but less numerous and have four congruent straight conical
the external surface carrying a crust of euasters and the other mark- pointed rays 700-1050 X 85-120 11m. More numerous smaller
ing the boundary between the subectosomal and choanosomal calthrops have rays 170-700 X 20-85 j.l.m, and all their rays have a
regions. Dichotriaene cladomes are more numerous in the upper similar conical shape. Euasters (Fig. 2C) include diverse mor-
layer than in the deeper subectosomal cavernous layer. Calthrops phologies: large thick-rayed stongylasters with 6-7 tapering rays,
are very abundant and irregularly dispersed throughout the smooth towards the centrum and spined elsewhere, and with hook-
choanosome, including smaller, possibly juvenile forms, forming a like spines, 21-2711m diameter; 2-5 rayed euasters 26-4511m
confused mass. Microscleres are variable in morphology. In the diameter. Acanthotylasters with 8-12 rays tapering only slightly
subectosomal region are numerous, mostly large, thick-rayed towards the end, cylindrical conically pointed, with only few spines
strongylasters, whereas in the deeper parts of the choanosome there except for larger spines at the end of actines, 8-16 11m diameter.
are oxyasters, acanthotylasters and very small spherasters with Spherasters divisible into smaller sphere-like asters with 14-20
acanthose actines (and many transitional forms). Megascleres conical rays each bearing a crown of spines at its end and appear-
(Fig. 2A) are oxeas, mostly symmetrical, slightly curved, 3600- ing acanthose, 5-9 11m diameter, and oxyspherasters with 10-14
5600 x 50-80 11m. Triaenes are exclusively dichotriaenes, conically rays, conically pointed, with the proximal half and tip smooth and
pointed rays, straight or slightly bent and constricted at the the remainder covered with spines, 20-32 11m diameter.
acladome end, rhabdome 2800-4400 X 100-440 11m, cladome Remorks. Monotypic. The species has been recorded from
650-1300 11m in diameter, clades 130-170 11m long and the paired EastAfrica, South Africa and South West Africa. Pulitzer-Finali, 1993:
clades 300-550 j.l.m long. There are apparently intermediate forms 254 mentions the presence of anatriaenes in his material from Kenya
132 Porifera' Demospongiae • Astrophorida • CaithropeUidae

assigned to this species; possibly this pertains then to a second species Synonymy. Pachastrella geodioides Carter, 1876: 407, pI. 14,
separate from C. sphaerica. Anatriaenes were also reported in a fur- fig. 23. Calthropella geodioides; Sollas, 1888: 111. Calthropella
ther record of Chelotropella from New Caledonia, viz., Chelotropella geodioides var. Topsent, 1897a: 434. Pachastrissa geodioides;
neocaledonica Levi & Levi, 1983b: 148, fig. 27. This species has Lendenfeld, 1903: 81. ? Corticella geodioides; Topsent, 1892a: 42;
oxeas, long-shafted dichotriaenes, anatriaenes and calthrops as mega- 1904b: 77, pI. 4, fig. 14, pI. 10, fig. 12. ? Calthropella geodioides;
scleres and the combination of strongylasters, spherasters and centro- Desqueyroux-Faundez, 1981: 730--732, figs 13,15, 106.
tylote microrhabds. It is possible that this species belongs to Material examined. None. Holotype (not seen): BMNH -
Ancorinidae, close to Ecionemia, rather than to Chelotropella. The Cape St Vincent, off the Portugese south coast, North Atlantic,
difference of Chelotropella and Calthropella is the presence of long- 525m depth.
shafted triaenes and oxeas, features which may be interpreted as Description (from Carter, 1876). Growth fonn globular
ancestral. The presence of these spicules have a profound influence on (Fig. 2D) to thickly encrusting. Surface even or slightly undulating,
the structure of the skeleton, which becomes radiate because of paral- correlated with size of specimen, slightly hispid, with oscules scat-
lel arrangement of the long shafts. This genus also marks the thin tered over surface. Ectosomal region without cortex but with for-
dividing line between Calthropellidae and Ancorinidae. But for the eign detritus and euasters interdispersed and fonning a confused
presence of the calthrops, this genus could easily fit in Stelletta. mass, although not markedly offset from choanosomal skeleton.
Choanosomal skeleton with heavy mesohyl containing foreign
Distribution detritus and two size classes of calthrops and oxeas dispersed with-
out apparent order. Larger calthrops (Fig. 2D) have clads lying par-
Known only from South Africa, SW Africa and Kenya. allel to the surface, intenningled with and largely covered by
spherasters. Calthrops morphology is variable, with the most
numerous fonns with simple actines commonly four in number,
PACHASTRISSA LENDENFELD, 1903 sometimes three and occasionally reduced to two. Less common
are dichocalthrops, with three rays bifurcated and extended in a
Synonymy plane tangential to the surface and with the fourth ray perpendicu-
lar to the others, consisting only of a short extension of the central
Pachastrissa Lendenfeld, 1903: 80. Jasplakina de Laubenfels, shaft, and producing a dichotriaene-like spicule with short rhab-
1954: 228. dome. Smaller calthrops ('subquadriradiates' of Carter) have the
fourth ray more-or-Iess atrophied which is extended into a short
Type species round prominence more or less prolonged, arms of equal length,
smooth, round, sharply pointed, and slightly curved. Dichotriaenes
Pachastrella geodioides Carter, 1876 (by original designation). (Fig. 2D) are equally numerous and have the rhabdome much
shorter than the rays (rays 700X 85 fLm). Oxeas (Fig. 2D) fusifonn,
Definition smooth, sharply pointed and slightly curved, 736X9.3 fLm.
Microscleres (Fig. 2D) vary from spherasters to spheroxyasters.
Calthropellidae with oxeas and calthrops irregularly arranged The fonner are most abundant at the surface, consisting of a thick
except at surface, lacking long-shafted triaenes, with short-shafted centrum and numerous cylindrical, short round tubercles (rays),
(dicho-)triaenes present in some species; microscleres only regular smooth or crowned with very fine spines, 7-25 fLm diameter. The
euasters. latter are more sparsely dispersed, with a poorly developed cen-
trum and numerous rays, possibly representing younger fonns of
Diagnosis the fonner, with blunt or sharply pointed rays, 12-15 fLm diameter.
Remarks. Topsent's (1892a, 1904b) records of this species
Massive, globular to thickly encrusting sponges with rough (as Corticella) are uncertain because he fails to mention the pres-
surface. Few small oscules. Ectosomal skeleton with a layer of con- ence of oxeas in these specimens (see Figs ID-E). He also
fused euasters covering the clades of larger calthrops lying parallel expressed his opinion that geodioides and Calthropella simplex
to the surface. Choanosomal skeleton largely confused, with oxeas, comprise a single species. It is possible that his material concerns a
calthrops and/or short-shafted triaenes and dichotriaenes, and some- species of Calthropella. A second species assigned to Pachastrissa
times mesotriaenes dispersed without apparent order. Microscleres by Lendenfeld is Stelletta pathologica Schmidt, 1868. 19, pI. III
are euasters ranging from spherasters to spheroxyasters. figs 3-4 (see redescription in Sollas, 1888: 202). This is not a
straightforward Pachastrissa because true calthrops are apparently
Remarks lacking, instead of which there are mesotriaenes. Still, it is con-
ceivable that the mesotriaenes represent modified calthrops.
Short-shafted mesotriaenes are known for several species, e.g., Lendenfeld also assigned Pachastrella connectens Schmidt, 1870:
P. pathologica (Schmidt, 1868) and P. inopinata (Pulitzer-Finali, 65, pI. VI fig. 5 to Pachastrissa, but the affinities of that sponge
1983), but are considered convergent and non-homologous with remain uncertain. Sollas (1888: 111) apparently had a slide of
those of Pachastrellidae, such as Triptolemus (Maldonado, 1993). Schmidt's material and from this it may be concluded that it could
Mesotriaenes are reportedly not present in the type species of perhaps be a Penares. The apparent possession of both microxeas
Pachastrissa. and centrotylote strongyles make membership of Calthropellidae
less likely. In any case, Sollas (1888: 112) synonymised P. con-
Description of type species nectens with Characella agassizi, but this action was apparently
not accepted by Lendenfeld (1903). Uliczka (1929: 50, figs 41-45)
Pachastrissa geodioides (Carter) (Fig. 2D). described Pachastrissa hartmeyeri from deep water off Barbados.
Porifera • Demospongiae • Astrophorida • CaltbropeUidae 133

Subsequently, Pulitzer-Finali (1986: 73) reported this species from calthrops. Microscleres irregularly shaped rugose microrhabds
shallow water off the Dominican Republic. It has much smaller (called ataxasters) and spherasters. Long-shafted triaenes and
oxeas and calthrops than P. geodioides. Pachastrella connectens oxeas are absent, although mesotriaenes occur in some species.
was previously synonymised with Characella agassizi by Sollas Remarks. Short-shafted mesotriaenes are not considered to
(1888: 112), but this action was apparently not accepted by be typical of this genus, found in some species (e.g., P. enigmatica
Lendenfeld (1903). Pulitzer-Finali (1983) described a fourth Levi & Levi, 1983b) but not in others (including the type species,
species, P. inopinata, as Calthropella, from the Mediterranean P. lithistina (Schmidt, 1880b) and P. lutea Pulitzer-Finali, 1987).
which was subsequently referred to Pachastrissa by Maldonado Calthrops and derivatives display interspecific variation in
(1993). Finally, Topsent's (1897a) record of a variety of the type Pachataxa, absent from P. enigmatica and present in P. lithistina
species from Ambon, Indonesia, is also questionable as to its con- and P. lutea.
specificity with the North Atlantic population, and it too possibly
represents a distinct species. Redescription of this material by Description of type species
Desqueyroux-Faundez (1981) suggests that there are several slight
morphometric differences between Indonesian and North Atlantic Pachataxa lithistina (Schmidt, l880b) (Fig. 2E).
populations. In fact Topsent himself (1904b: 79) commented on Synonymy. Pachastrella lithistina Schmidt, 1880b: 68,
several differences between these popUlations, although this pI. 9, fig. 3; Topsent, 1923: 6, fig. 1. Pachastrellidae incertae sedis,
alleged 'cosmopolitanism' still remains unresolved. Sollas, 1888: 112 Dercitus lithistina; Lendenfeld, 1903: 82.
The genus Jasplakina de Laubenfels, 1954: 228 was erected Pachataxa lithistina; de Laubenfels, 1936a: 179.
for type species Jasplakina nux de Laubenfels, 1954: 228, fig. 157 Material examined. None. Lectotype (here designated):
(by original designation). The holotype USNM 23120 (not exam- MZUS - dry specimen without locality data other than
ined) is described as a jet black rounded mass of 6 X 8 X 5 cm "Mexic.busen" (redescribed by Topsent, 1923: 6) Caribbean. The
(internally much lighter coloured), with undulate surface showing MZUS specimen conforms closely to Schmidt's description and is
a few oscules of 6mm diameter. The ectosome is packed with here indicated as the lectotype (following information from
microxeas of 25-100 X 0.5-2 11m and euasters with reduced num- Desqueyroux-Faundez & Stone, 1992: 52). Schmidt's original
ber of rays (four or more) up to 36 11m in diameter. Megascleres description gives no locality data. Paralectotypes: MCZ - an
arranged vaguely radiate, comprised of oxeas of up to 830 X 12 11m assumed 'holotype' from Grenada. 2MB 6870 - 2 slides without
and relatively rare reduced triactine calthrops, rays up to 250 X locality data.
10 11m. The microxeas are unique in the family, and indicate a pos- Description (from Schmidt, 1880b, and Topsent, 1923).
sible relationship with Penares in the Ancorinidae. The combined Thickly encrusting to cushion-shaped 10-20mm thick. Upper sur-
characters conform closely to Pachastrissa and accordingly face with numerous low conules each with a terminal osculum 0.5
Jasplakina is here considered a junior synonym of this genus. mm diameter. Under surface is even but with similar oscules and
pores dispersed. Subectosomal skeleton with large cavities forming
Distribution a labyrinthine network under the thin ectosomal cortex and perme-
ating the whole sponge. Skeleton a dense mass of calthrops under-
Three species are currently described from the Atlantic, a pos- neath a surface layer of euasters; the latter are also frequent in the
sible valid fourth species from Indonesia, and a tentatively fifth interior. Spicules. Calthrops in a large size range (no sizes given).
member occurs in the Central Pacific. Microscleres spherasters to oxyspherasters with numerous rays,
10-1611m diameter, and irregularly shaped rugose microrhabds
(Fig. 2E), often somewhat ellipsoid, but some appear to have short
PACHATAXA DE LAUBENFELS, 1936 rounded lobes and these are possibly derived from asters.
Remarks. De Laubenfels (1936a) erected this genus on the
Synonymy basis that its megasc1ere spiculation apparently consisted almost
exclusively of calthrops and that microscleres were ataxasters,
Pachataxa de Laubenfels, 1936a: 179. whereas Topsent (1923) regarded these features as merely modifi-
cations to characters typical of Pachastrella monilifera. Lendenfeld
Type species (1903: 82) transferred the type species to Dercitus, but de
Laubenfels suggested that it was so "radically different from other
Pachastrella lithistina Schmidt, 1880b (by original forms" as to warrant a new genus. Two other species have been
designation) described, one Pachataxa lutea Pulitzer-Finali, 1987, also from the
Caribbean (Jamaica). It differs a.o. in having some of the calthrops
Definition with two rays reduced resulting in diactines. Levi & Levi (1983b)
recorded Pachataxa enigmatica from deep water off New
Calthropellidae with calthrops irregularly arranged throughout Caledonia. This is closely similar in spiCUlation to P. lithistina,
the skeleton; short-shafted mesotriaenes occur in some species but the asters are only 4-5 11m in diameter. The microrhabds
although true triaenes absent; microscleres are regular euasters of Pachataxa are peculiar and constitute an easy character for
(spherasters) and characteristic rugose microrhabs (ataxasters). the genus. In other aspects, Pachataxa is closely similar to
Calthropella.
Diagnosis
Distribution
Massively encrusting, hard sponges. Ectosomal crust of
euaster microscleres. Choanosomal skeleton a dense mass of Caribbean and New Caledonia.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Geodiidae Gray, 1867

Maria J. Uriz

Centre d'Estudis Avamrats de Blanes, Cami de Sta Barbara sIn 17300. Blanes (Girona) Spain. (iosune@ceab.csic.es)

Geodiidae Gray (Demospongiae, Astrophorida) includes astrophorid sponges with sterrasters as the main cortical microsclere and various
forms of triaenes among the megascleres. Other microscleres present are euasters and microrhabds. Sponges are generally massive or
thickly encrusting and often develop into globular forms. The inhalant or exhalant orifices (or both) may be organized in a sieve. The
skeletal arrangement is radiate at the periphery and confused in the inner region of the skeleton. These sponges are usually cream, gray or
white in color inside, and gray, brown to black outside, often dependant on their exposure to light. They inhabit bathyal, soft bottoms and,
less often, are found in caves and overhangs of the sublittoral zone. The family contains twelve nominal genera of which six are valid:
Erylus, Caminus, Pachymatisma, Geodia, /sops, and Sidonops, with species distributed worldwide.
Keywords: Porifera; Demospongiae; Astrophorida; Geodiidae; Erylus; Caminus; Geodia; /sops; Pachymatisma; Sidonops.

DEFINITION, DIAGNOSIS, SCOPE recognizing the presence of true sterrasters in the former and flat
sterrasters (aspidasters) in the latter. However, the somal microscle-
Synonymy res in both families can be either a microrhabd or an aster; protri-
aenes and anatriaenes may be absent from some genera; and
Geodiadae Gray, 1867a: 504. Geodinidae Schmidt, 1870: 68. more-or-Iess subglobular sterrasters are present in genera allocated
Sterrastrosa Sollas, 1887, 1888: 209. Erylidae Lendenfeld, 191Oa: to both subfamilies. Thus, there is no consistent character to divide
11, 191Ob: 267. genera bearing sterrasters into two different families, and this clas-
sification is rejected here. The term 'somal' microsclere is used
Definition here, although an antiquated terminology, because it does not
imply that these microscleres are restricted to any particular region
Astrophorida with large oxeas and triaenes as megascleres. of the sponge, where the terms 'ectosomal' or 'choanosomal'
Microscleres are sterrasters, together with euasters, spherules or microscleres presume their strict localization.
microrhabds. However, recent phylogenetic molecular studies (Chombard
et al., 1998; Chombard, 1998) have suggested the paraphyly of the
Diagnosis family Geodiidae, and that the concept of Erylidae-Geodidae fami-
lies have to be reevaluated in a general cladistic revision.
Thickly encrusting, massive to globular growth forms with Several genera assigned to Geodiidae have a similar spicule
a well-developed cortex. The cortex is made of two differentiated complement and consequently earlier authors considered other
layers: the extemallayer ('ectochrote') is fleshy and collagenous characters to differentiate higher taxa, including the aquiferous
("collenchymatose") with one of the various specialized microscle-
res (euasters, spherules or microrhabds); the inner layer is filled
with sterrasters. Euasters or microrhabds may be also present in the
choanosome. Megascleres are regular, long shafted triaenes, and
oxeas.

Scope

This family has twelve nominal genera, six of which are rec-
ognized here as valid: Erylus, Caminus, Pachymatisma, Geodia,
/sops, and Sidonops.

History and Biology

The family Geodiidae was named Sterrastrosa by Sollas

(1888), referring to the consistent presence of sterrasters. Sollas
(1888), followed by Topsent (1894d), divided genera into two sub-
families: Erylina and Geodina, and Lendenfeld (191Oa), followed
by Dendy (1916c) and Wilson (1925), elevated these to family sta-
Fig. 1. Schematic representation of the arrangement of incurrent and
tus (Erylidae and Geodiidae) based on alleged fundamental differ- excurrent canals in the different genera of Geodiidae. A, Erylus and [sops.
ences between Erylus and Geodia in their microsclere spiculation. B, Caminus, Pachymatisma, and Sidonops. C-D, Geodia (Geodia) and
Dendy's diagnosis of Geodiidae differs from that of Erylidae in Geodia (Cydonium), respectively.

134
Porifera • Demospongiae • Astrophorida • Geodiidae 135

system, and particularly the structure of the inhalant and exhalant by means of a formal cladistic analysis, it is nevertheless preferable
orifices and canals as a diagnostic character, which combined with to maintain them as separate genera until other types of characters
skeleton features, allowed them to define genera. These criteria (e.g., genetic sequences) become available to perform a reliable
were not followed by some contemporary authors, such as Hajdu phylogenetic reconstructions. The types of inhalant and exhalant
et al. (1992) and Hooper & Wiedenmayer (1994), who considered structures found in members of this family are depicted in Fig. 1.
that spicules were more relevant characters indicative of phyloge- Most species of Geodiidae have a bathyal distribution, living
netic affinities and regarded that differences in aquiferous system predominantly on soft bottoms, but some representatives inhabit
structures were simple adaptations to living in particular habitats. dark habitats in the littoral and shallow sublittoral zones, such as in
This opinion is rejected here because there are some morphological caves, crevices, and overhangs.
characters, particularly several features of the cortex, that corrobo-
rate differences in aquiferous systems between these genera. It is Previous Reviews
difficult to contemplate that these complex structures are simply
ecological adaptations to survive in certain environmental condi- Sollas, 1888; Topsent, 1894d; Lendenfeld, 1903, 1907, 191Oa,
tions, and not genetically determined. However, as no assessment 191Ob; Dendy, 1905, 1916c; Wilson, 1925; de Laubenfels, 1936a;
of the monophyletic status of these genera has yet been attempted Hooper & Wiedenmayer, 1994; Chombard et al., 1998.

KEY TO GENERA

(1) Both inhalant and exhalant orifices are cribriporal .................................................................................................................... Geodia
Inhalant, exhalant or both orifices are uniporal ......... ........................................ ............................ ............... .... .... ............................... 2
(2) Both inhalant and exhalant orifices are uniporal .............. .... ................. .... ...... ............... .............................. .... ................................... 3
Inhalant orifices cribriporal, exhalant uniporal .... .... ...... ... ........................... .... .... .................................................... .... ................ .... ... 4
(3) Somal microscleres are euasters of different types ....................................................................................................................... lsops
Microrhabds among the somal microscleres .... .... .... .......................... .............. .... ....................................... .... ...... ..................... Erylus
(4) Microstrongyles among the somal microscleres ........................................................................................................... Pachymotisma
The somal microsclere is an aster or aster-derived form ..................................................................................................................... 5
(5) The somal microsclere is an euaster ....................................................................................................................................... Sidonops
The somal microsclere is a spherule .................. .... ..................................... ....... .... .... .......... ......................... ...... .... .... ............ Caminus

ERYLUS GRAY, 1867 Description of type species

Synonymy Erylus mammillaris (Schmidt, 1862) (Fig. 2).

Synonymy. Stelletta mammillaris Schmidt, 1862: 46. Erylus
Erylus Gray, 1867a: 549; Stelletta (in part) Schmidt 1862: 46; mammilaris; Gray, 1867a: 549.
Scutastra Ferrer-Hernandez, 1912: 582. Material examined. Holotype: BMNH 1867.3.11.32
(slide), LBIM DT2208 (61) (slide) - Adriatic. Comparative
Type species material. Specimens of E. euastrum (Schmidt, 1868): MNHN
CP95-107, CEAB.POR.BIOL Mal(3,6); CEAB.POR.BIOL Ca43.
Stelletta mammillaris Schmidt, 1862 (by monotypy). Scutastra cantabrica Ferrer-Hernandez, 1912: MNVN (Madrid)
holotype and specimen no. 54).
Definition Description. Sponge massive, ovoid or thickly encrusting
with rounded lobes, several cm in size, brownish in color in alcohol.
Geodiidae with short-shafted triaenes (ortho- or plagiotri- Cortex about 500 /Lm thick. Surface perforated by multiple uniporal
aenes); sterrasters usually more-or-Iess flattened (aspidasters). The orifices; a unique oscule, 2 mm in diameter at the top of each lobe.
somal microsclere is a centrotylote microrhabd. Uniporal, inhalant Megascleres: oxeas curved or sinuous, 750-1500 X 18-32 /JJll;
and exhalant orifices. dichotriaenes with rhabdome conical, straight, 532-716 X 25-44 /Lm
in size, ending in a sharp point; protoclads 90 X 32-35 /JJll; deutero-
Diagnosis clads 60-90 /Lm long. Microscleres: sterrasters, flattened, ellipsoidal
or more-or-Iess elongated, 62-106 X 29-52 /Lm in size, with clearly
Thickly encrusting to massive globular sponges with simple differentiated actines, rough at the end; choanosomal oxyasters,
(uniporal) afferent and efferent orifices. Megascleres are triaenes of 14-28 /Lm in diameter, with actines conical, pointed and rough at the
different types according to different species (plagiotriaenes, ortho- end. Somal microrhabds, extraordinarily abundant, 13-24 X 2-4/Lm
triaenes, etc.) and oxeas. Microscleres are microrhabds (ectosomal in size, cylindrical, very rough, occasionally centrotylote. Skeletal
microscleres) and different types of euasters (in the choanosome). arrangement: oxeas slightly radiate at the periphery, confused
towards the interior; the triaenes have their cladome below the cortex
Previous reviews with the rhabdome directed inwards; cortical layer of sterrasters cov-
ered by a crust of somal microrhabds; rough microrhabds and
Topsent, 1984; Lendenfeld, 1903, 191Ob; Dendy, 1916c; oxyasters throughout the choanosome. Distribution. Adriatic,
Wilson, 1925; Mothes et al., 1999; Adams & Hooper, 2001. Mediterranean (Algiers coasts), North Atlantic (Azores).
136 Porifera· Demospongiae • Astrophorida • Geodiidae

Remarks. Certain differences in spicule sizes of specimens Previous reviews

from the Adriatic and Atlantic have been reported: the dichotriaenes
are stronger and the oxyasters are weaker in Atlantic specimens Sollas, 1888; Topsent, 1894d.
(Topsent, 1928c) than in those from the Adriatic (Sollas, 1888).
Distribution. Mediterranean, Atlantic and Pacific Oceans. Description of type species

Caminus vulcani Schmidt, 1862 (Fig. 3).

CAMINUS SCHMIDT, 1862 Materiol examined. Holotype: missing - Adriatic Sea,
Sebenico. Neotype (here designated): BMNH 1867.7.26.48 - dry
Synonymy specimen from the type locality, Sebenico. CEAB.PORBIO.OO8a,
CEAB.PORBIO.OO8b, and CEAB.PORALB.Ollb - Mediterranean.
Caminus Schmidt, 1862: 48. Geodia (in part) SOllas, 1888: 241. Description. Globular sponge with a single atrial oscule
at the top. Surface even, perforated by a network of minuscule
Type species orifices (cribriporal ostia). Cortex hard, 1.5-2mm thick (thicker
closer to the sponge base). Megascleres: Strongyles straight or
Caminus vulcani Schmidt, 1862 (by monotypy). slightly curved, 850-880 X 15-17 /-Lm; orthotriaenes with rhab-
dome straight, 480-572 X 15-16.5 /-Lm, and cladome 320-360/-Lm
Definition in chord length, with long and straight clads. Microscleres:
sterrasters, 100-115 X 87-90 /-Lm in diameter, subspherical,
Geodiidae with globular sterrasters. The somal microsclere is depressed, with short actines; oxyasters with few, conical actines
a spherule (spheraster with reduced actines). Cribriporal inhalant (35-42/-Lm in diameter); spherules, 3-4/-Lm in diameter, without
orifices and simple atrial oscules. conspicuous actines or with actines barely differentiated. Skeletal
arrangement: radiate at the periphery, confused towards the inte-
Diagnosis rior; the triaenes have their cladome lying in the cortex or just
below it, with the rhabdome directed inwards; cortical layer of ster-
Hemispherical to lobate sponges with a single atrial osculum rasters covered by a subectosomal crust of spherules; spherules and
at the top of each lobe, and cribriporal inhalant ostia. The megas- oxyasters dispersed throughout the choanosome. Distribution.
cleres are oxeas, slightly curved, with blunt points, often trans- Adriatic, western Mediterranean, 15-40m depth.
formed into strongyles, and orthotriaenes. Sterrasters subspherical. Remarks. As the holotype of this species is missing, we
The somal microsclere is a spherule. The choanosomal spicule is designate here the specimen BMNH 1867.7.26.48 from the type
an oxyaster. locality (Sebenico) as the neotype.

+'
A

c ~+
E
c?oOo
= 00° E
D ~ D
=
A B

20~m

100 11m
a,b

c,d,e
./ I()()~m

20 11m
l .b

c.d e

Fig.2. Erylus mammillaris (Schmidt). Spicules from the holotype (BMNH Fig. 3. Caminus vulcani Schmidt. Specimen from the Blanes littoral
1867.3.11.32). A, oxea. B, dichotriaenes. C, oxyasters. D, spiny (Mediterranean). A, strongyles. B, orthotriaenes. C, oxyasters. D,
microrhabds. E, flattened sterrasters (aspidasters). spherules. E, sterraster.
Porifera· Demospongiae • Astrophorida • Geodiidae 137

Distribution Diagnosis

Atlantic Ocean and Mediterranean. Thickly encrusting or irregularly massive growth forms.
Surface even. Oscules sparse or arranged in clusters or rows.
Cortex conspicuous. Sterrasters located throughout the cortex.
PACHYMATISMA BOWERBANK, 1842 Inhalant orifices flow into single inhalant canals, which cross the
cortex, and end in subcortical ostia. Microstrongyles (somal
Synonymy microscleres) located at the external part of the cortex and along
the walls of canals. Oxyasters and microrhabds sparse within the
Pachymatisma Bowerbank in Johnston, 1842: 244. choanosome.

Type species Previous reviews

Halichondria johnstoni Bowerbank, 1842 (by subsequent Sollas, 1888; Topsent, 1894d; Lendenfeld, 1903.
designation).
Description of type species
Definition
Pachymatismajohnstonia (Bowerbank, 1842) (Figs 4 and 5).
Geodiidae with subspherical sterrasters. The somal micro- Synonymy. Halichondria johnstonia Bowerbank, 1842
sclere is a microrhabd. Cribriporal inhalant orifices and simple (in Johnston, 1842: 198); Pachymatisma johnstonia; Bowerbank,
oscules. in Johnston, 1842: 244; Pachymatisma johnstonia; Bowerbank,
1866: 4, 51. Pachymatisma normani Sollas, 1888: 243. Caminus
osculosus Grube, 1872 (see Koehler, 1986: 53).
Material examined. ? Holotype: BMNH BK 839, No.
R2.3.7.9 (l77h) (slides) - dry specimens from the type locality,
North Atlantic, Ireland.
Description. Massive sponge, with rounded borders, up to
15 cm thick. Surface smooth, bright, extensively perforated by
cribriporal inhalant orifices. Oscula simple, 2-3 mm in diameter,
surrounded by a whitish ring, arranged in clusters. Color bluish
gray, violet, brown pinkish or reddish in the ectosome, whitish in
the choanosome. Cortex up to 1 mm thick. Megascleres:

B'
~

~ C

100 11m a.b.c E

-::::::>
=
=
F
o11m b' =::::>

20l1m d ,e ,l

Fig. 5. Pachymatisma johnstoni (Bowerbank). Spicules from specimen

Fig. 4. Pachymatisma johnstoni (Bowerbank): dry specimens (BMNH BK BMNH RI.3.7.9 (l77h). A, B, B', strongyles. C, orthotriaene. D, rough
839) showing two different arrangements of oscula. oxyasters. E, sterraster. F, microrhabds.
138 Porifera· Demospongiae • Astrophorida • Geodiidae

strongyles, straight or irregularly curved, 600-1100 X 13-20 j..Lm; Synonymy. Geodia gibberosa Larrnarck, 1815: 333;
orthotriaenes with a straight rhabdome 440-700 X 13-26 j..Lm Topsent, 1918: 611; 1931: 3; Hechtel, 1965: 68; Wiedenmayer,
in size, and clades straight or slightly curved, up to 300 j..Lm long. 1977b: 178; Van Soest & Stentoft, 1988: 12; Pyxitis gibberosa;
Microscleres: sterrasters ellipsoid in shape, 90-120 X 71-93 j..Lm; Schmidt, 1870: 70.
oxyasters 22--63 j..Lm in diameter; microrhabds, 18-32 X 2.7-4.5 j..Lm Material examined. Holotype: MNHN (not examined) -
long. Skeletal arrangement: strongyles, radiately arranged at the tropical Atlantic, Guiana. Other material. Pyxitis gibberosa
periphery, confused towards the interior; the triaenes have their Schmidt: BMNH 1870.5.3.167 - dry fragment. Comparative mate-
cladome in the cortex with the rhabdome directed inwards; cortical rial. CEAB.POR.BIOL.l46 ter, and 212 - specimens of Geodia
layer of sterrasters covered by a subectosomal crust of microrhabds; cydonium Jameson. MNHN CP62-Ilb, CP63-E5, and CP92-88 -
microrhabds and oxyasters dispersed within the choanosome. specimens of Geodia barretti Bowerbank. MMNHN CP63-E2 and
Distribution. North Atlantic, Ireland, Norway, England, coasts of CP95.110 - specimens of G. nodastrella Carter.
France, Portugal, and Spain, intertidal and subtidal. Description. Massive, globular lobate sponge, several cm in
Remarks. This genus has a spicule complement similar diameter, with a wide zone where oscula are located. Color whitish
to that of Caminus, the main difference being the presence of an outside in dark habitats, dark brown in zones exposed to light,
ectosomal layer of spiny microrhabds in Pachymatisma instead of cream inside, both in life and in alcohol. Surface smooth, with
spherules found in Caminus. The cortex also is notably thinner in several sieve areas bearing ostia 50-100 j..Lm in diameter. Cortex
Pachymatisma. 0.5-1 mm thick. Oscula 0.3-1.2 mm in diameter, clustered in circu-
lar or ovoid, well-delimited areas, 1-8 cm wide. Megascleres:
Distribution oxeas straight or slightly curved, 780-1900 X 20-40 j..Lm; cortical,
fusiform oxeas, 125-250 X 2-4 j..Lm in size; orthotriaenes derived
Atlantic and Pacific Oceans. from plagiotriaenes (or vice-versa), 420-900 X 15-22 j..Lm in size,
with clads of 192-235 X 13-16 j..Lm. Sterrasters spherical, 40-
100 j..Lm in diameter; somal spherasters with a large centrum and
GEODIA LAMARCK, 1815 reduced strongylote rays, 4-10 j..Lm in diameter; choanosomal
oxyasters, 10-30 j..Lm in diameter, with rough, conical actines, vari-
Synonymy able in number, and a centrum up to 5 j..Lm in diameter. Skeletal
arrangement: radiate at the periphery, confused towards the inte-
Cydonium Fleming, 1828: 516. Pyxitis Schmidt, 1870: 70. rior; the triaenes have their cladome in the cortex with the rhab-
Geodinella Lendenfeld, 1903: 117; 191Oa: 205. ? Stellogeodia dome directed inwards; cortical layer of sterrasters covered by a
Czerniavsky, 1880 [1879]: 115 [280]. Geodistrongyla Hoshino, subectosomal crust of somal spherasters (the cortex at the pore
1981: 252. sieves is devoid of sterrasters); spherasters and oxyasters dispersed
within the choanosome. Distribution. Tropical W coast of Africa,
Type species

Geodia gibberosa Larrnarck, 1815 (by monotypy).

Definition

Geodiidae with large oxeas and regular triaenes arranged radi-

c A B
ally at or near the surface. Globular sterrasters and euasters as
microscleres. Inhalant and exhalant conducts are cribriporal.

Diagnosis D
Thickly encrusting, massive, or globular sponges. Surface
irregularly hispid. Cortex conspicuous, formed by a crust of ster-
rasters with a layer of euasters in the outer zone. Cladome of the tri-
aenes located at the cortex. Megascleres (oxeas and triaenes)
radially arranged at the peripheral zone of the sponge, oxeas more
disorganized in arrangement in the interior. Inhalant and exhalant
orifices arranged in clusters under a sieve of the cortex (cribriporal).

Previous reviews

Sollas, 1888; Topsent, 1894d, 1918, 1931; Wilson, 1925;

F 50J.lm a,b.c
Lendenfeld, 1903, 1910a; Bergquist, 1968; Wiedenmayer, 1977b.
20~m d.e.f
Description of type species Fig. 6. Geodia gibberosa Lamarck, specimen BMNH 1870.5.3.167. A,
oxea. B, orthotriaene. C, cortical oxea. D, sterraster. E, choanosoma1
Geodia gibberosa Lamarck, 1815 (Fig. 6). oxyasters. F, somal spherochiasters.
Porifera· Demospongiae • Astrophorida • Geodiidae 139

Horida, Cuba, Antilles, Bermudas, Brazil, St. Thome, common in Plagiotriaenes with rhabdome up to 4000 X 40 fJ-m, and clads
shallow waters. 500--600 fJ-m long. Protriaenes with rhabdome up to 7500 X 72 fJ-m,
Remarks. The genus Geodia has been traditionally and anatriaenes of very different sizes, with a straight rhabdome up to
distinguished from Cydonium (type species C. muelleri Heming) by 2550 X 13-20 fJ-m and clads of 62-146 X 4-18 fJ-m. Sterrasters, sub-
the absence of an exhalant sieve. However, the type species of spherical, 90-111 X 88-110 fJ-m in diameter with the actines spiny at
Geodia (G. gibberosa) also has a typical exhalant sieve, negating this their point. Spherasters (somal rnicroscleres) with a marked centrum
alleged difference. Furthermore, closer examination of some species and conical, pointed actines, in two size categories of 5-7 fJ-m and
of Geodia such as G. barretti showed that the exhalant cavity corre- 11-20 fJ-m in diameter, respectively; choanosomal oxyasters (also
sponds to an invagination of the sponge, which inside contains a called spherasters by Sollas) with a small centrum, 20-70 fJ-m in
cribriporal structure (Boury-Esnault et al., 1994b). Consequently, diameter. Skeletal arrangement: layer of sterrasters in the cortex
both genera are clearly synonymous. Geodistrongyla Hoshino (type under a subectosomal crust of spherasters. Spherasters are also con-
species G. strongyla Hoshino, 1981) is also clearly a Geodia with centrated along the canal walls; chiasters are dispersed within the
strongyloid ends of actines, and is transferred into synonymy here. choanosome; triaenes extend their cladome within the inner layer of
the cortex directing their rhabdome inwards; oxeas radiately arranged
Distribution at the sponge periphery but more confusedly arranged in the interior.
Remarks. Sollas (1888) put into synonymy with l. phlegraei
Cosmopolitan. two species previously described by Vosmaer (1882a) as [sops pal-
lida and l. sphaeroides. Both species, however, seem to be charac-
terized by the presence of dichotriaenes and orthotriaenes (instead
1S0PS SOLLAS, 1880 of protriaenes and plagiotriaenes in l. phlegraei), and by oxyasters
(instead of the spherasters). Thus, I agree with Lendenfeld (1903)
Synonymy that these two species are not synonyms of l. phlegraei.

[sops Sollas, 1880: 396. Synops Vosmaer, 1882a: 20. Not Distribution
Synops sensu Sollas, 1886a: 198, 1888: 227, 265. Caminella
Lendenfeld, 1894: 150. North Atlantic, Indian and Pacific Oceans, and Mediterranean
and Red Sea.
Type species

[sops phlegraei Sollas, 1880 (by monotypy).

Definition

Geodiidae with uniporal inhalant and exhalant orifices.

Sterrasters are globular. The somal rnicrosclere is a euaster.

Diagnosis
A
J()() j.lm
Globular sponges with oscules and ostia consisting of simple
apertures (uniporal). Cortex relatively thin. Megascleres are oxeas
and triaenes with a long rhabdome. Sterrasters subspherical. Somal
rnicroscleres are euasters.

Previous reviews

Sollas, 1888; Topsent, 1894d; Lendenfeld, 1903, 191Oa.

*
Description of type species

[sops phlegraei Sollas, 1880 (Fig. 7).

F~
Synonymy. [sops phlegraei Sollas, 1880: 396.
Material examined. Holotype: not seen - North Atlantic,
Norway, Kors Fjord, 329 m depth. Other material. BMNH
1910.1.1.913 - Norman Collection. Comparative material. MNHN
CP63-E1 and CP95-E17 - specimens of l. pachydermata Sollas.
MNHN CP63-185 - specimen of l. intuta Topsent.
Description. Small, subspherical sponge, 2.5 cm in diame- IOOj.lm b,c 40 j.lm d,e,!
ter. Brown color in alcohol. Surface hispid due to protruding Fig. 7. [sops phlegraei Sollas, specimen BMNH 1910.1.1,913, Norman
megascleres. Oscula and ostia simple (uniporal). Cortex about Collection. A, anatriaene. B, protriaene and plagiotriaene. C, dichotriaene.
600 fJ-m thick. Long oxeas, up to 6000 X 60 fJ-m in size. D, somal spherasters. E, choanosomal oxyasters. F, sterraster.
140 Porifera· Demospongiae • Astrophorida • Geodiidae

SIDONOPS SOLLAS, 1889 Description. Cup-shaped sponge, 40cm high and from 12cm
(at the base) to 31 cm (at the broadest part) wide, with thick walls,
Synonymy brown in color (dried). Surface corrugated and smooth. Cortex 800
fLm thick. Cribriporal, inhalant orifices on the external face of
Synops Vosmaer, sensu Sollas, 1886a: 198, 1888: 227, 265 the cup. Oscules simple, spread over the inner side of the cup.
(in part). Not Synops Vosmaer, 1882a: 20. Sidonops Sollas, Megascleres: oxeas, fusiform, straight or curved with blunt,
1889: 276. sharp or rounded ends (strongyles and styles), 1250--1500 X
17-23 fLm; triaenes are mainly orthotriaenes, with a rhabdome
Type species 900--1050 X 15:....20 fLm in size, tapering to a sharp point, and a
cladome curved outwards, 300--362 fLm in chord length, with clads
Synops neptuni Sollas, 1886a (by original designation). 180 fLm long; anatriaenes very scarce (only one was reported from
the holotype by Sollas, 1888),270--380 X 11-14 fLm in size (rhab-
Definition dome) and 17-21X8-lOfLm (clads). Microscleres: sterrasters,
spherical, about 45-55 fLm in diameter; chi asters (ectosomal
Geodiidae with triaenes arranged radially at the sponge microscleres) 5-8 fLm in diameter, with short, abruptly truncated,
periphery. Somal microscleres are euasters. Inhalant orifices are rod-like actines and with a more-or-less marked centrum (sphe-
cribriporal; exhalant ones are uniporal. rochiasters); choanosomal oxyasters, 21-28 fLm in diameter, with
numerous, rough actines, 11-15 fLm long. Skeletal arrangement:
Previous reviews the layer of sterrasters in the cortex is covered by a subectosomal
crust of chiasters. Spherasters and chi asters dispersed within the
Lendenfeld, 1903, 191Oa; Wilson, 1925. choanosome; the triaenes extend their cladome within the inner
layer of the cortex and direct the rhabdome toward the
Description of type species choanosome; oxeas are radially arranged at the sponge periphery
but more disarranged in the interior; occasionally they protrude
Sidonops neptuni (Sollas, 1886a) (Fig. 8). through the sponge surface, rendering the surface slightly hispid.
Synonymy. Synops neptuni Sollas, 1886a: 198. Sidonops Remarks. The genus Sidonops Sollas, 1889 was maintained
neptuni; Sollas, 1889: 276. by Lendenfeld (1903) and Wilson (1925) despite the fact that no
Material examined. Holotype: not seen - tropical Atlantic skeletal characters differentiate it from Geodia. The only difference
(pernambuco), 58 m depth, mud. Other material. BMNH is the arrangement of the exhalant orifices: cribriporal in Geodia
1894.11.16.327. and uniporal in Sidonops.
The diagnosis initially provided by Lendenfeld (1903) for the
genus Geodinella (type species G. cylindrica): "Geodiidae with
reduced triaenes arranged radially in the periphery of the sponge
and occurring in longitudinal bundles forming an axial skeleton
in the interior" was subsequently modified by the same author
(191Oa) to include other species which did not have an axial skele-
ton of triaenes but only triaenes distributed in the choanosome
(e.g., G. robusta). Other authors (e.g., Wilson, 1925; Bergquist,
1968) included in Geodinella other species such as G. spherastrosa
(inhalant uniporal, exhalant cribriporal) and G. vestigifera Dendy,
by ignoring the character "triaenes in the choanosome". Currently,
as a result the only diagnostic character for Geodinella is the pres-
ence of modified triaenes. Aborted triaenes as those in Geodinella,
have also been described in some species of [sops, Geodia, and
Sidonops and, consequently, this character has no diagnostic value.
As the inhalant orifices of Geodinella are uniporal and the oscula
are cribriporal, this genus becomes synonymous of Sidonops.

Distribution

Arctic, Antarctic, E and W Atlantic, Indo-Malayan region and

E Pacific.

100 11m B.b.c.d ACKNOWLEDGEMENTS

20 11m e,f,g.h I thank M. Bardaji from the CEAB for preparing the microscope
slides, G. Carreras, and G. Agell for helping in processing drawings
Fig. 8. Sidonops neptuni (Sollas), specimen BMNH 1894.11.16.327. A,
oxea. B, strongyle. C, style. D, orthotriaene. E, microxea. F, rough oxyaster. and photographs. Clare Valentine, C. Uvi, and R. Desqueyroux-
G, spherochiasters. H, sterraster. FaUndez kindly loaned holotypes and other museum material.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Pachastrellidae Carter, 1875

Manuel Maldonado

Centro de Estudios Avanzados de Blanes (CSIC), Centre d'Estudis Avan"ats de Blanes, Acceso a Cala San Francesc 14, Blanes 17300,
Girona, Spain. (maldonado@ceab.csic.es)

Pachastrellidae Carter (Demospongiae: Astrophorida), including Theneidae Carter, contains 12 valid genera and 2 subgenera. Most gen-
era contain cup-like, massive, and submassive species and are distributed from tropical to high latitudes, mostly in epibathyal and bathyal
habitats. A few genera also include encrusting or cavity-filling forms found predominantly in tropical or temperate shallow-water habitats.
This astrophorid family is defined by the possession of tetraxons (calthrops, short-shafted triaenes, mesotriaene-derived desmas, or long-
shafted triaenes) in combination with streptasters (never euasters) and in most cases by monaxonic microscleres (Le., microxeas,
microstyles and microstrongyles). The microscleres are densely packed at the ectosome forming a feltwork. The relationship between
genera possessing a typical pachastrellid skeleton and some genera bearing tetraxial desmas other than mesotriaene-derived desmas
(mesotriders), which have traditional been included in Lithistida, is still a matter of contention.
Keywords: Porifera; Demospongiae; Astrophorida; Pachastrellidae; Acanthotriaena; Ancorella; Brachiaster; Characella; Cladothenea;
Dercitus; Pachastrella; Poecillastra; Stoeba; Thenea; Triptolemma; Vulcanella; Vulcanella (Annulastrella) sUbgen.nov., Vulcanella
(Vulcanella).

DEFINITION, DIAGNOSIS, SCOPE (2) Vulcanella species also have special aquiferous openings
(Fig. 3D-E); and (3) a wide variety of streptasters, apart from
Synonymy metasters, can be found in many species traditionally allocated
to both Theneidae and Pachastrellidae. Pachastrellidae and
Pachastrellidae Carter, 1875c. Theneidae Carter, 1883b. Theneidae also shared the absence of euasters.
Streptastrosa Sollas, 1888. [Asterostreptidae] Topsent, 1902. Given that skeletal characters do not allow inequivocal discrim-
[Astrostreptidae] Auct. (lapsus). Metastrosa Lendenfeld, 1906. ination between Theneidae and Pachastrellidae, both families are for-
Halinidae de Laubenfels, 1934. mally merged here. This proposal is not a new one, as it has been
previously suggested by several earlier authors. Sollas (1888) sug-
Definition gested the name Streptastrosa (with Demus level) for a general
astrophorid line including Pachastrellidae and Theneidae, and charac-
Astrophorida having a variety of tetraxons (Le., calthrops, terized by the presence of streptasters in combination with tetraxons.
short-shafted triaenes, mesotriaene, mesotriaene-derived desmas, Sollas also stated (1888: CN): "indeed, there is a most evident transi-
or long-shafted triaenes) in combination with streptasters (never tion from Thenea to Pachastrella through the intermediate genus
euasters) and, in most cases, monaxonic rnicroscleres (usually Poecillastra". The Sollas 'Demus' taxon was also used by Lendenfeld
rnicroxeas and/or rnicrostrongyles). (1906) and Lebwohl (1914), but under the name Metastrosa.
Combining the families was first proposed by Topsent (1902), who
Diagnosis maintained Theneidae and Pachastrellidae as subfamilies of his new
family [Asterostreptidae] (which is a nomen nudum because there is
Astrophorida whose megascleres are calthrops, short-shafted no genus Asterostrepta, -us). Ferrer-Hernandez (1914b) and Wilson
triaenes, mesotrider desmas, or long-shafted triaenes (Fig. 1), usu- (1925) also supported this combination but did not differentiate the
ally, but not always, in combination with monaxonic megascleres subfamilies. The former author proposed the name Pachastrellidae
(typically oxeas, rarely styles or strongyles). Microscleres are while the latter erroneously considered that Theneidae had priority
streptasters and/or monaxonic microscleres (Fig. 2), but never over Pachastrellidae. More recently, the combined family, under the
euasters. The ectosomal skeleton is constituted by a layer of name Pachastrellidae, was also claimed by Wiedenmayer (1994).
microscleres, forming a feltwork that has occassionally been Twelve nominal genera are considered here to be valid within
referred to as "pseudocortex". Pachastrellidae: Acanthotriaena Vacelet, Vasseur & Levi, 1976;
Ancorella Lendenfeld, 1906; Brachiaster Wilson, 1925; Characella
Scope Sollas, 1886a; Cladothenea Koltun, 1964b; Dercitus Gray, 1867a;
Pachastrella Schmidt, 1868; Poecillastra Sollas, 1888; Stoeba
Prior to this review, Pachastrellidae and Theneidae were con- Sollas, 1888; Thenea Gray, 1867a; Triptolemma de Laubenfels,
sidered to be separate and valid families (e.g., Levi, 1973). 1955b; Vulcanella Sollas, 1886a.
Theneidae was diagnosed as having special aquiferous openings, Several other nominal genera previously assigned to
long-shafted triaenes and metasters, while Pachastrellidae was Pachastrellidae (including Theneidae) are herein abandoned, syn-
defined as having calthrop-like tetraxons and a large variety of onymised into other genera, or transferred to other families. For
streptasters. However, it has been shown that: (1) several species instance, Dercitancorina Topsent, 1902, formerly erected for
of typical pachastrellid genera (such as Characella, Poecillastra Pachastrella lesinensis Lendenfeld, 1894, is now regarded as a
and Vulcanella (=Sphinctrella» have long-shafted triaenes; junior synonym of Stoeba, being S. lesinensis a valid species.

141
142 Porifera· Demospongiae • Astrophorida • Pachastrellidae

Fig. 1. Pachastrellidae megascleres. A, anatriaene of Thenea. B, anatriaene of Characella. C, dichotriaene of Cladothenea. D, regular calthrops and has-
tate oxeas of Pachastrella. E, triactinal tetraxon of Characella. F-G, mesocalthrop and mesodichotriaene, respectively, of an undescribed Pachastrella from
!be Galapagos Islands. H-I, end of multibranched clad in mesotriaene of Triptolemma and !be entire spicUle, respectively. J-M, several branching stages of
mesotriaenes in transition to mesotrider desmas in Brachiaster. N-O, general view and detail of a piece of the choanosomal network made by fused
mesotriders and pierced by fascicles of strongyloxeas in Brachiaster. P-Q, details of the end of cladotyles in Cladothenea. R, hastate oxea of Pachastrella.

Pachamphilla Lendenfeld, 1906 (type species P. alata Lendenfeld, which was formerly considered as a theneid (e.g., Lendenfeld,
1906) is a junior synonym of the ancorinid genus Penares Gray, 1906), is now regarded as a junior synonym of the ancorinid genus
1867a (see also Levi, 1967b). The monotypic genus Chelotropaena Penares (see chapter on Ancorinidae). The genus Plakinastrella
Lendenfeld, 1906, with type species C. tenuirhabda Lendenfeld, Schulze, 1880, which was included (as Placinastrella) in
1906, is a synonym of Poecillastra. The genus Yodomia Lebwohl, Theneidade by Sollas (1888) and in Pachastrellidae by Lendenfeld
1914, with type species Y. ijimai Lebwohl, 1914, is a synonym of (1894), is now regarded as a valid plakinid genus (see chapter on
Characella, with its other known nominal species - Y. perfecta Plakinidae). The genus Nethea Sollas, 1888 was invalidated fol-
Dendy, 1916c - being a Pachastrella species. The genus Halinastra lowing Levi (1973) and Maldonado (1993), as it was an artificial
de Laubenfels, 1936a, formerly erected for Pachastrella exostotica taxon erected to contain species whose tetraxons have underdevel-
Schmidt, 1868, is herein considered to be a junior synomyn of oped actines; its type species Nethea nana (Carter, 1880a) and
Stoeba, with S. exostotica a valid species. another species N. amygdaloides (Carter, 1876) clearly belong to
Several other genera had previously been discarded for differ- Poecillastra, while Nethea dissimilis Sara, 1959a probably belongs
ent reasons. For instance, the genus Papyrula Schmidt, 1868, to the genus Stoeba. The genera Calthropella Sollas, 1888 and
Porifera· Demospongiae • Astrophorida • Pachastrellidae 143

Fig. 2. Pachastrellidae microscleres. A, large and small, spiny microxeas of Characella. B, large annulate microxea and small, uniformly spiny microxea
of Vulcanella. C, microxea of Triptolemma. D, oval microstrongyles, spiraster-like microstrongyles and curved, oval microstrongyles of Pachastrella.
E, detail of ornamentation of spiraster-like microstrongyle, suggesting that this spicule type is a streptaster with reduced actines. F, sanidasters of Stoeba.
G, detail of a sanidaster-ataxaster of Stoeba, showing a spheraster-like morphology, likely derived from an evolutionary shortening of the central axis.
H, metaster of Pachastrella. I, large plesiaster and small spiraster of Poecillastra. J, large plesiaster and small amphiaster of Cladothenea. K, spiraster of
Vulcanella. L, amphiaster of Characella on two large, spiny microxeas.

[Corticella] Sollas, 1888, (preocc. by Corticella Ehrenberg, 1872, anatomical findings, being difficult to predict the direction of such
Protoctista), regarded as pachastrellids by Sollas (1888) and changes. Future research might result in a family split, reinstating
Lendenfeld (1894) respectively, are now considered to be syn- the former theneids and pachastrellids, as suggested by an anony-
onyms, with the senior genus Calthropella being the type of the mous reviewer. However, it is more likely that the present family
family Calthropellidae (see chapter on Calthropellidae). Similarly, concept will be expanded by the addition of a variety of genera
the genus Pachastrissa Lendenfeld, 1903, formerly erected in from the polyphyletic 'order Lithistida', which is gradually being
Pachastrellidae, is currently considered a valid genus of dismembered. Indeed, a potential relationship between 'lithistids'
Calthropellidae. The monotypic genus Neothenea de Laubenfels, and pachastrellids was suggested by some earlier workers, such as
1934 was invalidated by Van Soest & Stentoft (1988), who con- Carter (1875c), who even included several 'lithistids' in his family
firmed that the type species, Neothenea enae de Laubenfels, 1934, Pachastrellidae. This relationship between pachastrellids and
was likely conspecific with Characella aspera Sollas, 1886a desma-bearing genera also appears to be supported by the Late-
(the type species of Characella). Finally, the name [Picraster] Cretaceous genus Propachastrella Schrammen, 1910 (type species
attributed to Sollas (1888) by de Laubenfels (1936a) for a putative Pachastrella primaeva Zittel, 1878b t), which is characterized by
pachastrellid is apparently an error, as no data has been found in the calthrop-like spicules with irregularly curved actines ended in
literature to validate de Laubenfels' information (Le., nomen dubium). syzygial plates, showing a morphology transitional between regu-
Undoubtedly the present scope of Pachastrellidae, as defined lar calthrops and tetraxonic desmas (e.g., Wiedenmayer, 1994).
here, will change on the basis of future genetic, biochemical, and The affinity between Pachastrellidae and some 'lithistid' genera is
144 Porifera· Demospongiae • Astrophorida • PachastreUidae

Fig. 3. Pachastrellids with specialized aquiferous systems. A-C, Thenea muricata. A, specimen showing basal roots and a transversal recess of the body,
in which the inhalant areas are located. B-C, details of the inhalant areas and ostioles. D-E, Vulcanella tricomis. D, specimen showing the atrial areas of
the exhalant side. E,detail of the exhalant areas, which are surrounded by a largely protruding palisade of flexuous spicules.

also supported by the spiny tetraxons of Acanthotriaena and the should be kept in mind when reallocating some of the genera cur-
multibranched mesotriaenes of Triptolemma and Brachiaster, rently in the 'order Lithistida' .
which even become tetraxonic desmas (mesotriders) in this latter The diagnoses and definitions of genera revised here are
genus. The stout, short-shafted dichotriaenes of some species of the expanded to include both the traits of the type species and the most
genus Stoeba also resemble the tetraxons of some 'lithistids'. On distinctive traits that consistently occur in all or most species in
the basis of morphological similarities some authors (e.g., Reid, each genus. Lists of all valid species in each genus are not provided
1970) have suggested the hypothesis of a common ancestor for although special attention has been given to the remarks on each
streptaster-bearing 'lithistids' and pachastrellids. Indeed, the idea genus to clarify the taxonomic status of controversial species
that a variety of tetraxon-bearing 'lithistids' may be pachastrellids assigned to them.

KEY TO GENERA

(1) Tetraxons being only a variety of long-shafted forms (i.e., triaenes) ................................................................................................ 2
Tetraxons being calthrops/short-shafted tetraxons alone or in combination with long-shafted forms .............................................. 3
(2) With cladotyles (styles with a crown of tubercles at its rounded end) ....................................... .. .................................. Cladothenea
Without cladotylotes; monaxonic megascleres being regular oxeas or styles ........................................................................ Thenea
(3) With a number of short-shafted tetraxons becoming mesotriaena-derived desmas (mesotriders) ................................... Brachiaster
Tetraxons never becoming mesotrider desmas .... ....... .. .............. .......... ................ ................. ... ........ ... .... .. ...... ............. .. ...... ........ .... 4
(4) Tetraxons with spiny rhabdome ................................................................................................................................. Acanthotriaena
Tetraxons being entirely smooth ........................................................................................... ................................. ............................ 5
(5) All tetraxons being pentactinal (i.e., mesotriaenes) ....................................................................................................... Triptolemma
Tetraxons being triactinal and tetractinal forms, alone or in combination with mesotriaenes ... ........... ...................... ................ ...... 6
(6) Without streptasters ...................................................................................................................................................... .. ..... Ancorella
With streptasters ...... ........... ... .... ...... ... ... .............. ..... ..... ............. .......... ........ ... ...... ... ........ ... ...... ....... .... .... ................................ ... ...... 7
(7) Without microxeas ............................................................................................................................................................................. 8
With microxeas ... ....... .......... .. ............ ............................................................................ ................................ ......... .. .................... 10
(8) With toxa ................. ... .... ........... ...... ...... ....... ... ..... ......... ........ ................ .............................. .... ... ........... ........... ........ ............. Dercitus
Without toxa .. ....................................... ................... ........ ................ ....... .... ....... ....... ............. .. ........................ .... ........................... 9
(9) With oval microstrongyles .............................................................................................................................................. Pachastrella
Without oval microstrongyles; with microrhabd-like sanidasters ............................................................................................ Stoeba
(l0) Cribiporal oscules surrounded by a palisade of protruding spicules; annulate microxeas and/or annulate
plesiasters ......... ............... ..... .... ....... ... .......................... ............... ... .... ....... ...... ... .... ... .................... ....... ....... .... .......... ....... Vulcanella
Simple oscules; uniformly spiny microxeas or, more rarely, smooth microxeas ............................................................................ 11
(11) Microxeas in at least two categories; streptasters never being plesiasters ............................ ................. ................. .......... Characella
Microxeas in one category; streptasters always including plesiasters ............................................................................. Poecillastra
Porifera· Demospongiae • Astrophorida • Pachastrellidae 145

ACANTHOTRIAENA VACELET, VASSEUR & LEVI, 1976 also been reported in the late-Triassic genus Costamorpha Mostler,
1986 (see also Wiedenmayer, 1994).
Synonymy
Description of type species
Acanthotriaena Vacelet, Vasseur & Levi, 1976: 21.
Acanthotriaena crypta Vacelet, Vasseur & Levi, 1976 (Fig. 4).
Type species Material examined. None. The description is taken from
Vacelet et al. (1976).
Acanthotriaena crypta Vacelet, Vasseur & Levi, 1976: 21 (by Description. Small encrusting sponge filling cavities on the
monotypy). underside of a boulder, from which only small fragments could
be collected. The material did not allow description of the skeletal
Definition structure and casts doubts about the exogenous or endogenous ori-
gin for some of the spicule types found. The skeleton consists of
Pachastrellidae having megascleres of which are triaenes with four spicule types: oxeas, acanthotriaenes, spirasters and raphides.
spiny rhabdomes (acanthotriaenes). Oxeas are smooth, regular in shape, curved at the middle, and
measuring 115-130 X 5-6 /-Lm. Acanthotriaenes are dichotriaenes
Diagnosis with smooth protoclads and deuteroclads measuring 15-35/-Lm X
5-15/-Lm and 15-55 X 5-15/-Lm, respectively, and a spiny rhab-
Encrusting Pachastrellidae, the megascleres of which are dome measuring 155-500 X 5-20/-Lm. The rhabdome, although it
spiny tetraxons (acanthotriaenes) and small oxeas; its microscleres may occasionally be smooth, is often provided with 1-6 conical
are spirasters and raphides. spines, each 40/-Lm long, that contain a branch of axial canal in
their bases. Spirasters have short actines relative to the central axis
Remarks and measure 16-18 X l/-Lm in total length. Raphides are flexuous
and measure 50--250 X 0.5-1 /-Lm.
The family allocation for Acanthotriaena, a monotypic genus Remarks. The genus and species is known only from the
known only by fragments of its type species, is arguable. This holotype. Although the oxeas described above were intimately
genus is apparently related to other genera with spiny tetraxons, intermingled with the remaining spicules, Vacelet et al. (1976) sug-
such as the Jurassic-Cretaceous Acanthastrella Schrammen, 1924a gested the possibility that they may actually belong to a Cliona.
and the Recent Thrombus Sollas, 1888. Furthermore, the skeleton
of two species ofthe latter genus, Thrombus abyssi (Carter, 1873b)
and Thrombus jancai Lehnert, 1998, consists of a combination of ANCORELLA LENDENFELD, 1906
spiny short-shafted tetraxons and peculiar amphiasters. Based on
their skeletal similarities it is possible that Acanthotriaena is not a Synonymy
pachastrellid but a second member of the unconnected, monotypic
family Thrombidae. Conversely, it is also possible that Thrombus is Ancorella Lendenfeld, 1906: 248.
a reduced pachastrellid and thus Thrombidae an empty family (see
chapter on Thrombidae for alternative views). Dichotriaenes and Type species
anatriaenes with acanthose decoration on clads and rhabdome have
Ancorella paulini Lendenfeld, 1906: 248 (by monotypy).

Definition

Pachastrellidae without streptasters.

Diagnosis

= Pachastrellidae without streptasters. Megascleres are strongy-

loxeas transitional to strongyles, calthrops reduced to three actines,
and anatriaenes. Microscleres are uniformly and finely spiny
microxeas.

Remarks

The absence of asters complicate the taxonomic interpretation of

this monotypic genus. Although Ancorella has been traditionally con-
sidered as a pachastrellid that has lost its streptasters (Lendenfeld,
1906), it might also be regarded as an ancorinid that had lost its
euasters. However, several of its features, which also occur in typical
Fig. 4. Acanthotriaena. A. crypta, holotype. A, sIDooth oxeas (scale pachastrellid genera, suggest that Ancorella is a skeletally-reduced
35 !LID). B, acanthodichotriaenes (scale 75 !LID). C, spirasters (scale pachastrellid, as follows: the general appearance and consistence of
17 !LID). D, flexuous raphides (scale 40 !LID). the sponge, the ectosomal feltwork of microxeas, the arrangement,
146 Porifera· Demospongiae • Astrophorida • Pachastrellidae

shape and size of the reductional stages of calthrops, the presence whole cladome is reduced to a knob, yielding a spicule morphol-
of anatriaenes with a blunt rhabdome (see Maldonado 1996), and ogy similar to that of a tylostyle; clads measure 45-60 j.l.m in length
the decoration of the microxeas. Indeed, the body form and skeletal and rhabdomes measure 750-1100 X 5-11j.l.m. Microxeas are in a
features of the type species, Ancorella paulini Lendenfeld, 1906, single size category (67.5-210 X 2.5-4j.l.m), being slightly curved,
strongly resemble those of the Atlantic-Mediterranean Characella uniformly and finely spiny, and with acerate to hastate points.
tripodaria (Schmidt, 1868), which is characterized by the presence The ectosomal skeleton consists of a feltwork of microxeas. It is
of anatriaenes with blunt rhabdomes and malformed clads internally reinforced by abundant reduced calthrops and few
(Figs lB, 7M), as well as malformed and reduced calthrops, and, strongyloxeas, both placed tangentially to the sponge surface.
what is more important, an extremely low density of streptasters in Anatriaenes, arranged perpendicular to the surface, pierce the ecto-
its tissue (Maldonado, 1996). some projecting the cladome out of the sponge. Microxeas,
strongyloxeas and calthrop-derived spicules also occur abundantly
in the choanosome.
Description of type species Remarks. A category of smooth microxeas mentioned in the
original description and omitted herein is likely to be exogenous to
Ancorella paulini Lendenfeld, 1906 (Fig. 5). the sponge, as also suggested by Lendenfeld (1906). The type
Material examined. Holotype: BMNH 1908.2.9.199-122 species is known only from type material collected off the Pacific
(slides). coast of Chile (38°34'S, 77°38'66"W) at 672m depth.
Description. 1\vo specimens collected. One specimen is a
triangular, plate-like sponge, 75mm long, 55mm wide and 30mm
thick, attached to the substratum through one of the sides. The other BRACH/ASTER WILSON, 1925
is a 115 mm long encrustation growing on a coral. Subspherical pro-
tuberances give a lumpy, sulcate appearance to the surface. Synonymy
Aquiferous orifices are not visible on the sponge surface, but abun-
dant aquiferous canals, up to 5 mm wide, occur in the choanosome. Brachiaster Wilson, 1925: 471.
Color in alcohol is light brown (from Lendenfeld, 1906). Spicules
consist of strongyles transitional to strongyloxeas, triactinal and Type species
diactinal spicules derived from calthrops and short-shafted tri-
aenes, anatriaenes and microxeas in a single size category. Brachiaster simplex Wilson, 1925: 471 (by monotypy).
Strongyloxeas transitional to strongyles are slightly curved,
centrotylote, measuring 1300-2300 X 18-26j.l.m. Triactinal and
Definition
diactinal spicules are likely derived from calthrops and have
straight, slender actines measuring 200-990 X 20-70 j.l.m and
Pachastrellidae, whose tetraxons are short-shafted triaenes
600-1100 X 40-70 j.l.m, respectively. Anatriaenes have blunt rhab-
and/or mesotriaenes that become mesotrider desmas.
domes and frequent malformations in the clads; sometimes the

A======--
~========-- --
- Diagnosis

Megascleres are styloxeas, hastate oxeas, short-shafted tri-

aenes and mesotriaenes with clads diversely branched transitional
to tetracrepidal and mesotrider desmas, respectively. Desmas can
be articulated through subcircular syzygial plates, but, at the inner-
most parts of the sponge, they fuse to each other, making a solid
network. Microscleres are spiny microxeas, oval microstrongyles,
and amphiasters transitional to metasters.

Remarks

The diagnosis of this monotypic genus given above is based on

the skeletal traits reported from the two available records of its type
species (Wilson, 1925; Levi & Levi, 1989). Nevertheless, the skele-
tal differences between the material collected in both cases suggests
the existence of high levels of intraspecific skeletal variability and
the possibility of further diagnosis readjustments when additional
material is collected.

Description of type species

Fig. 5. Ancorella. A. paulini, holotype. A, fragments of long strongyloxeas
(scale 250 fJ.-m). B, anatriaene with blunt rhabdomes (scale 50 j.l.m). Brachiaster simplex Wilson, 1925 (Figs lJ-O, 6).
C, triactinal tetraxons (scale 250 fJ.-m). D, spiny microxeas (scale 40 fJ.-m). Material examined. Holotype (not seen): USNM. Specimen
E, detail of microxea end (scale 6 fJ.-m). of Levi & Levi (1989): MNHN.
Porifera· Demospongiae • Astrophorida • Pachastrellidae 147

~
F

Fig. 6. Brachiaster. B. simplex. A, styloxea (scale 150 f1m). B, hastate oxeas (scale 40 f1m). C, short-shafted mesotriaenes with undivided clads (scale 75
f1m). D, short-shafted mesotriaenes with diversely branched clads (scale 75 f1m). E, desma mesotrider (scale 200 f1m). F, spiny microxeas (scale 40 f1m) .
G, oval microstrongyles(scale 20 f1m). H, amphiasters transitional to metasters (scale 20 f1m).

Description. Cup-like sponge, 145 mm tall, 120 mm wide externally covered by a region of isolated mesotriaenes, which in
and with a 25 mm thick body wall. Color is grayish ochre with pink- tum is externally covered by a feltwork of oval microstrongyles.
ish tones. 1\vo different regions are externally distinguished on the Fascicles of styles and styloxeas, passing through the holes of the
concave side of the sponge: a central region, where the surface is desma network, run from the innermost choanosome to the internal
rugose and hispid, and a peripheral region, where the surface is side of the sponge, which is slightly hispid. Towards the external
smooth and glabrous, with a radial striation pattern caused by the side of the cup, the internal layer of desmas is covered by an outer-
internal aquiferous canals. The spicule set in Levi's material most layer rich in cells, oval microstrongy!es and metasters.
consists of styloxeas, has tate oxeas, a variety of mesotriaenes Microxeas are scattered in the interstitial tissue.
transitional to mesotrider desmas, microspiny microxeas, micro- ReTTUlrks. The description above is based on Wilson's
strongyles and streptasters. Styloxeas are long, thin, and very flexu- (1925) initial description and re-examination of Levi & Levi's
ous, measuring up to 4 mm in length and 6 mm in thickness. Hastate (1989) subsequent material. By comparing Wilson's and Levi's
oxeas are smooth and slightly curved, measuring 100-225 X 6-11 descriptions it appears that the presence or absence of dichotri-
f.Lm. Mesotriaenes have nearly symmetrical rhabdomes, being aenes is a subject of intraspecific variability. Dichotriaenes were
clearly shorter than the clads; clads may be undivided, dichoto- reported by Wilson (1925) in addition to the mesotriaenes, both
mous, trifurcated and even multibranched. Isolated mesotriaenes, types becoming desmas. Nevertheless, both observations are com-
the clads of which measure 150-350 X 15~9 f.Lm, grow irregularly patible as it is likely that the mesotriaenes may grow initially
in thickness and also in length to become mesotriders (300-500 X through a dichotriaene stage before developing the epirhabdome.
80-250 f.Lm), which often occur fused to each other forming a solid The type species is known from Philippine waters at 198-188 m
network. Microxeas are slightly curved, entirely and unifonnly depth (Wilson, 1925; Levi & Levi, 1989).
microspiny, measuring 100-160 X 3.5-5 f.Lm. Oval microstrongyles
measure ~ X 2-3 f.Lm, being neither centrotylote nor curved.
Streptasters, which measure 12-15 f.Lm in total length, are a variety CHARACELLA SOLLAS, 1886
of amphiasters transitional to metasters, with a ~ f.Lm thick axis
and 4-8 f.Lm-long actines. The skeleton of the innermost regions of Synonymy
the sponge body wall consists of a rigid network of mostly fused
des mas, although articulate spicules can be found towards the Characella Sollas, 1886a: 186. Yodomia Lebwohl, 1914.
periphery. Towards the internal side of the cup, the desma layer is Neothenea de Laubenfels, 1934.
148 Porifera· Demospongiae • Astrophorida • Pachastrellidae

Type species Description of type species

Characella aspera Sollas, 1886a: 186 (by original designation). Characella aspera Sollas, 1886a (Fig. 7A-F).
Synonymy. Normania goliath Sollas, 1886a: 187.
Definition Material examined. Holotype: BMNH 1894. 11.16.149-150
(slides).
Pachastrellidae the microscleres of which consist of streptasters Description. Massive sponge, with an irregular, hispid sur-
of straight axis (never spirasters) and at least two categories of curved face that shows ridges and folds. Numerous ostia are scattered on
monaxonic spicules (microoxea, microstyles, microstrongyloxeas). the surface, sometimes grouped in depressed areas. Oscules, 1-2 mm
in diameter, are also visible on the sides and summits of the ridges.
Diagnosis The spicule set consists of oxeas, short-shafted orthotriaenes and
dichotriaenes, microxeas in two size categories, and amphiasters.
Pachastrellidae whose megascleres consist of abundant oxeas Oxeas are slightly curved and fusiform, measuring 1100-2700 X
and scarce calthrops (andlor short-shafted triaenes), mostly restricted 25-75 f.Lm. Orthotriaenes have clads measuring 200-630 X 225-50
to subectosomal regions. Microscleres are spiny or smooth microxeas- f.Lm; dichotriaenes have protoclads and deuteroclads measuring up
microstrongyles in at least two size categories and streptasters with a to 143 f.Lm and 270 f.Lm, respectively; rhabdomes are 200-400 X
straight central axis (amphiasters or sanidasters); streptasters may be 20-45 f.Lm in both spiCUle types. Microxeas are slightly curved,
very scarce in some species. Anatriaenes with cladomes that pro- fusiform, smooth, never centroangulated and rarely centrotylote.
trude the sponge surface and blunt rhabdomes embedded in the There are two size categories, 50-80 X 2.5-3 f.Lm and 150-300 X
choanosome occur in some species. 4-5 f.Lm, although a few microxeas of intermediate sizes occur.
Amphiasters have two whorls of four to six, relatively long, thin, and
Remarks sharply-pointed actines, and a relatively short and irregularly thick
axis, the ends of which become sharp actines. Amphiasters measure
Streptasters are scattered in very low density in the tissue of sev- 14-25 f.Lm in total length along the axis and actines 18 to 23 f.Lm. The
eral species, such as in C. aspera Sollas, 1888 from the North Atlantic ectosomal skeleton consists of a feltwork of microxeas supported
(BMNH 1894.11.16.149-152 (slides» and C. tripodaria (Schmidt, internally by large oxeas and triaenes that pierce largely the sponge
1868) from the Atlantic-Mediterranean region (Fig. 2L; BMNH surface. The triaenes are mostly restricted to subectosomal regions
1868.3.2.36 (slide); MNCN-36; CEAB Alb-8, Alb-31, Alb-6Op). and the large oxeas are irregularly scattered through the whole
This makes Characella close to the monotypic genus Ancorella in its sponge. Microxeas are also abundant in the choanosome, particularly
skeletal structure, which is characterized by the absence of strep- reinforcing the internal epithelia along with the amphiasters.
tasters and the presence of anatriaenes. It is noteworthy that anatri- Remarks. Characella aspera (Fig. 7 A-F), known from
aenes also occur in some species of Characella, such as C. tripodaria 640 m deep, muddy bottoms off Brazil, is clearly differentiated
(Fig. 1B, 7M; see also Maldonado, 1996). They were also observed from the close North-Atlantic C. pachastrelloides (Fig. 7G-L), with
in re-examination of the holotype of C. pachastrelloides (Carter, the former lacking anatriaenes and spines on the microxeas.
1876) (BMNH 1882.7.28.125 (dry» and in many other specimens of Although the holotype of C. aspera has short-shafted dichotriaenes
this species collected around Cape Saint Vincent (Fig. 7G-H) by the in addition to the short-shafted orthotriaenes, this spicule combina-
'Porcupine' Expeditions (e.g., BMNH 1910.1.1.1680 (slide) Norman tion should not be considered as a species specific trait. It is known
ColI.), as well as in specimens described by Stephens (1915a) that the presence/absence of dichotriaenes is subject to population
(BMNH 1953.II.II.30) from Irish waters, and eastern-Atlantic speci- variability in other species of the genus (Topsent, 1902; Maldonado,
mens described by Levi & Vacelet (1958). Anatriaenes have also been 1996). The occasional siliceous globules (40-160 f.Lm in diameter)
found in specimens of an undescribed Characella collected from the described in the holotype of C. aspera (Fig. 7F) are probably the
Galapagos Island (Maldonado & Pomponi, unpublished). result of failure in the silicification process and they should not be
Re-examination of diverse material has also led to the conclu- considered as a true spicule type. I also found such globules during
sion that several species originally described in other astrophorid gen- the re-examination of the holotype of C. stellettodes.
era may belong to Characella. For instance, Pachastrella connectens
Schmidt, 1870 (BMNH 1870.5.3.45 (slide), also 1870.5.3.170 (dry,
fragment from type) from Florida is a valid species of Characella CLADOTHENEA KOLTUN, 1964
(Figs lE, 7T-X), while Ancorina pachastrelloides Schmidt, 1870
(Fig. 7R-S; BMNH 1870.5.3.48 (slide» from Florida (but not Synonymy
Characella pachastrelloides (Carter, 1876) from the northeastern
Atlantic) is a junior synonym of C. connectens (Schmidt). Contrary to Cladothenea Koltun, 1964b: 16.
the suggestion of Topsent (1923, 1928c), C. connectens can be clearly
distinguished from both C. pachastrelloides (Fig. 7G-L; BMNH Type species
1895.1.23.2 (wet» and all other species in the genus because the large
microxeas have been replaced by strongyles. Pachastrella stellettodes Cladothenea andriashevi Koltun, 1964b: 16 (by monotypy).
Carter, 1885e (BMNH 1882.12.31.8 (dry» from the Japan Sea is a
valid species of Characella, the skeleton of which consists of oxeas, Definition
calthrops, short-shafted orthotriaenes, two size categories of microx-
eas, and scarce amphiasters (Figs 2A, 7N-Q). Yodomia ijimai Pachastrellidae with distinctive cladotyles, which are style-
Lebwohl, 1914 (BMNH 1938.7.20.4 (dry) 1938.4.20.4a (slide», also like spicules characterized by having a crown of tubercles at the
from the Japan Sea, is also likely a junior synonym of C. stellettodes. rounded spicule end.
Porifera· Demospoogiae • Astrophorida • Pachastrellidae 149

o
E_
F

J .

p
==-

T
-----=
W
x--==-
Fig. 7. Characel/a . A-F, C. aspera, holotype. A, oxea (scale 250 fLm). B, orthotriaene (scale 150 fLm) . C, amphiasters (scale 25 fLm) . D, large microxeas
(scale 80 fLm) . E, small microxeas (scale 80 fLm). F, siliceous globule (scale 80 fLm). G-L, C. pachastrel/oides, holotype. G-H, anatriaenes (scale
100 fLm). I, short-shafted orthotriaene and calthrop (scale 300 fLm). J, sanidaster (scale 30 fLm) . K, large microxea (scale 100 fLm). L, small microxea,
microstrongyle-microstyle (scale 50 ILm). M, C. tripodaria , blunt anatriaene (scale 75 fLm) . N-Q, C. ijimai, holotype. N, short-shafted piagiotriaenes, with
some occasional malformations (scale 350 fLm) . 0 , amphiaster (scale 25 ILm). P, large microxeas (scale 50 fLm). Q, small microxeas (scale 50 ILm). R- X,
C. connectens. R-S, calthrop and dichotriaene (holotype of Ancorina pachastrelloides Schmidt) (scale 200 fLm) . T, triactinal calthrop (holotype of
Pachastrella connectens Schmidt) (scale 200 fLm) . U, metasters (scale 25 fLm) . Y, large microstrongyles (scale 80 fLm). W, large microxeas (scale 80 fLm).
X, small microxeas (scale 50 fLm) .

Diagnosis reduction may have occurred in other theneids, also leading to

monaxon-like morphologies. For instance, the oxytylote - a
Pachastrellidae with radial, protruding fascicles of oxeas and monaxon-like spicule characterized by a terminal swelling and
cladotyles. Long shafted tetraxons (i.e., dichotriaenes) occur at the found in the roots of Thenea delicata Sollas (BMNH MC.5; 1.13)-
periphery of the choanosome, with their rhabdome pointing toward appears to have evolved from anatriaenes (Sollas, 1888). The fact
the innermost part of the pseudospherical sponge body and the that the axial canal branches incipiently within the terminal
cladome reinforcing the ectosomal skeleton. Microscleres are strep- swelling in the case of T. delicata, and that a canal branch goes into
tasters in more than one category (i.e., plesiasters and amphiasters). each tubercle in the case of Cladothenea (Figs lQ, 8B), is evidence
to support the tetraxon origin of this spicule. Nevertheless, it
Remarks remains unexplained why the number of axial branches and tuber-
cles ranges from 4 to 8 or even more in cladotyles. Note that
This is a monotypic pachastrellid genus characterized by Koltun's (1964b) concept of a cladotyle differs from that of Boury-
the presence of cladotyles. As inferred from the spicule name, Esnault & Rtitzler (1997), in which they are considered to be
cladotyles were assumed by Koltun to be monaxons evolved from spicules with a cladome at one end and a tyle at the other.
tetraxons by a process of cladome reduction. If so, such a cladome Cladotyle-like spicules, called cladotylostyles, also occur in the
150 Porifera· Demospongiae • Astrophorida • Pachastrellidae

Fig. 8. Cladothenea. C. andriashevi, holotype. A, cladotyles (scale 40/Lm). B, cladotyle ends in detail, showing axial canals (scale 200 /Lm). C, oxeas
(scale 200 /Lm). D, dichotriaene and detail of dichotriaene clads (scale 200 /Lm). E, tetractinal, triactinal, and diactinal plesiasters (scale 70/Lm). F, amphi-
asters (scale 70/Lm). G, detail of amphiaster (scale 10 /Lm).

hadromerid genera Tylexocladus Topsent, 1898b and Sphaerotylus (oxeas and cladotyles) that protrude 1-2mm beyond the sponge
Topsent, 1898b, which are apparently unrelated to pachastrellidae. surface. Spicules are oxeas, cladotyles, long-shafted dichotriaene,
The resemblance in skeletal structure and external morphol- plesiasters, and amphiasters. Oxeas are curved, slender, slightly
ogy between Cladothenea and Thenea is striking, the two only fusiform, in a wide size range (800 to more than 4000 X 20-55 /Lm);
differentiated significantly by the presence or absence of a poorly long fragments of isodiametric, 8 to 10 /Lm-thick oxeas also occur
understood spicule, the cladotyle. This, along with the fact that in the examined slides. Cladotyles are stout, nearly straight, some-
Cladothenea is a monotypic genus and known only from a single, what fusiform, with a crown of 4 to 8 tubercles, and measuring
fragmented individual, suggests further re-assessment of the genus 1200-2000 X 40-65 /Lm. Dichotriaenes with conical, slender rhab-
if additional material comes to hand. domes (1500-3400 X 20-70 /Lm); protoclads and deuteroclads
measuring 55-180 X 20-65 /Lm and 450-1100 X 20-60/Lm,
Description of type species respectively. Plesiasters with 2 to 6, finely spiny actines, measuring
60-120/Lm in total length. Amphiasters with finely spiny actines,
Cladothenea andriashevi Koltun, 1964b (Figs 1C, 1P-Q, 21, 8). measuring 10-22 /Lm in total length. The ectosomal skeleton consists
Material examined. Holotype: ZIT... 6300 (skeletal slides). of a feltwork of oxeas and cladotyles; it is also reinforced by amphi-
Description. Holotype fragmented in several, 5-10mm asters and the cladomes of the dichotriaenes. Oxeas and cladotyles
pieces, precluding a proper description of the external sponge mor- form hispid tufts at the sponge surface, but also radial tracts that pen-
phology. Sponge originally described as a gray, apparently sub- etrate the choanosome along with the rhabdomes of dichotriaenes.
spherical individual, with a deep transverse furrow in the upper Abundant plesiasters reinforce the internal pinacoderms.
part of the body. Oscula were not appreciable, but pore sieves were Remarks. The type species is known from a single speci-
identifiable in some fragments with ectosome. There is a 2 mm- men collected from Balleny Islands (Antarctic shores) at 3000m
thick, bristly ectosomal skeleton provided with spicules tufts deep.
Porifera· Demospongiae • Astrophorida • Pachastrellidae 151

DERCITUS GRAY, 1867 also converged to microrhabdose morphology (e.g., Topsent,

1897a; Levi & Levi, 1983a; Maldonado, 1993).
Synonymy
Description of type species
[Halina] Bowerbank, 1858: 288 (preocc.). Dercitus Gray,
1867a: 542. Battersbyia Bowerbank, 1874b: 343. Dercitus bucklandi (Bowerbank, 1858) (Fig. 9).
Synonymy. Halina bucklandi Bowerbank, 1858: 288; 1861:
Type species 235; Hymeniacidon bucklandi, Bowerbank, 1866: 226; Dercitus
bucklandi, Gray, 1868a: 542; Pachastrella bucklandi, Schmidt,
Halina bucklandi Bowerbank, 1858: 288 (by subsequent des- 1870: 76; Dercitus niger Carter, 1871a: 3; Battersbyia bucklandi,
ignation; de Laubenfels, 1936a: 179). Bowerbank, 1874b: 343.
Material examined. Holotype: Not seen. Slide of
Definition Battersbyia bucklandi: BMNH 1974, BK-542. Slide from Norman
collection: BMNH 10.1.1.1705. Other material. Specimen of
Pachastrellidae with microscleres including toxas. D. bucklandi; Burton: BMNH 1947.7.1.5a - Tombay Survey.
Holotype of Dercitus niger Carter, 1871a : BMNH (no registration
Diagnosis number found).
Description. Massive or cushion-shaped, sometimes filling
Pachastrellidae having toxas as microscleres in addition to cavities in hard substrata. Sponge surface is finely hispid. Oscules
characteristic sanidasters with a thick central axis relative to the are 1-2mm in diameter, isolated or in small groups, and with a dis-
actines. Megascleres are calthrops or short-shafted triaenes; oxeas tinctive sphincter-like structure. Ostioles are punctiform and irregu-
are absent. larly scattered over the entire surface. The ectosome of the living
sponge is black with gray or brown tinges, while the choanosome is
Remarks beige to cream. Spicules are calthrops to short-shafted p1agiotri-
aenes, sanidasters and toxa. Calthrops and short-shafted p1agiotri-
Halina Blainville, 1830: 497 predates [Halina] Bowerbank, aenes have clads measuring 170-450 X 30-50 f.Lm; clads are often
1858: 288 (according to Neave), with the former a junior synonym blunt, and incipiently divided in some of the plagiotriaenes.
of Halichondria (see chapter on Halichondriidae). Consequently, Sanidasters measure 27-35 X 2-6 f.Lm and have a relatively thick,
although [Halina] Bowerbank, 1858 and Dercitus are objective straight or curved central axis entirely covered by abundant minute
synonyms (with the same type species), the former name is actines, which make the spicule to look like a spiny microstrongyle.
unavailable and the latter is here adopted. Dercitus is considered Toxa measure 50-110 X 1-1.5 f.Lm, being slender and isodiametric,
here to be a monotypic genus although it has been traditionally with pointed, smooth ends. The skeleton consists of a feltwork of
merged with Stoeba Sollas, 1888 - type species Stoeba simplex sanidasters, the internal side of which is reinforced by the cladomes
(Carter, 1880a) - following the concept of Topsent (1902) and of short-shafted plagiotriaenes. The choanosomal skeleton consists
Lendenfeld (1903). Following recommendations of Dendy (1905), of great abundance of ca1throps and toxa, with scarce sanidasters.
Burton (1926), Vacelet & Vasseur (1971) and Maldonado (1993), Remarks. The type species occurs typically in sublittoral
and given that Halina bucklandi has toxas while Stoeba does not, habitats of the northeastern Atlantic and the western Mediterranean
the two genera are retained as distinct for the time being. The pres- (Templado et al., 1986), growing preferably in sites with low expo-
ence of toxas is of no value in providing information on the phylo- sure to sun light.
genetic relationships of Dercitus, because no other astrophorid has
them. A few toxa were described in the second record of the
pachastrellid Poecillastra rickettsi de Laubenfels, 1930, but were PACHASTRELIA SCHMIDT, 1868
subsequently regarded as being foreign (de Laubenfels 1932).
Topsent's hypothesis (1902) that Dercitus' toxas are reduced Synonymy
euasters is unlikely because there is no remnant of a centrum.
Conversely, vestiges of a centrum can be found in the reduced toxa- Pachastrella Schmidt, 1868: 64.
like asters of other astrophorids, such as Erylus (e.g., Topsent,
1927b; Pu1itzer-Finali, 1983; Maldonado, 1992).
The microrhabdose form of the sanidasters, which has been
the principal trait used to support the synonymy between Dercitus
and Stoeba may be also deceiving when used to infer taxonomic
relationships. The evolutionary paths through which the micro-
rhabdose spicules of Dercitus (Fig. 9C) and Stoeba (Fig. 12B, D, E,
H) have evolved (which are currently assumed to be sanidasters
with reduced actines) are hard to retrace. It is probable that they
have both evolved from a streptaster stage by actinal reduction.
Nevertheless, we cannot rule out the hypothesis that they are
derived from a microdiactinal stage, similar to the oval micro-
strongyles of Pachastrella or the small microxeas of Characella.
Furthermore, reduced euasters, such as the ataxasters of some Fig. 9. Dercitus. D. bucklandi, holotype. A, calthrops (scale 175/-Lm).
calthropellid genera (e.g., Pachataxa de Laubenfels, 1936a), have B, toxa (scale 10 /-Lm). C, sanidasters (scale 10 /-Lm).
152 Porifera· Demospongiae • Astrophorida • PachastreUidae

Type species Kirkpatrick, 1902: BMNH 02.2.13.4 to 6 - Natal Coast. Holotype

of Pachastrella caliculata Kirkpatrick, 1902: BMNH 02.2.13.18-
Pachastrella monilifera Schmidt, 1868: 64 (by monotypy). South Africa. Specimens of P. caliculata; Lendenfeld, 1906:
BMNH 08.2.9.107 to 110 - South Africa. Specimens of P. monilif-
Definition era; de Laubenfels, 1935b: BMNH (no registration number) - Baja
California. Holotype of Pachastrella chuni Lendenfeld, 1906:
Pachastrellidae with microscleres including oval BMNH 08.2.9.111 to 114 - Cape Verde.
microstrongyles, but never microxeas. Its tetraxons do not become Description. Encrusting, massive or cup-like sponges, with
desmas. a rough surface to the touch, the holotype being an irregular nodu-
lar fragment. Ostioles are punctiform, widely scattered through
Diagnosis the body, sometimes grouped in small depressions of the surface.
Oscules are irregularly scattered in most specimens, but concen-
Pachastrellidae having oxea megascleres in one or more cate- trated on the internal side in cup-like forms, and over the lateral
gories along with calthrops and/or short-shafted triaenes that can sides and the top of massive individuals. Oscules are 0.5 to 2 mm in
be mesotriaenes. Microscleres consist of oval microstrongyles that diameter and may be fringed in white in alive specimens. Spicules
make an ectosomal feltwork and one or several kinds of streptasters are oxeas, calthrops, microstrongyles, and several types of strep-
that may include forms with reduced actines; they never are tasters. There are two categories of oxeas: 1) large, robust fusiform
microxeas. oxeas (950-1900/-Lm X 18-30/-Lm) similar to the choanosomal
oxeas of other pachastrellids and (2) small, curved hastate oxeas
Remarks (150-350/-Lm X 5-6/-Lm) similar to those found in the haplosclerid
genus Haliclona. Calthrops occur in a large size variety, having
It is worth noting that, unlike in the related genus Brachiaster; conical, straight or curved clads that measure 150-800/-Lm X
mesocalthrops never become tetracrepidal desmas. 8-lOO/-Lm and show occasional malformations. Microstrongyles
After re-examining diverse material, it is concluded that several are typically oval, microspiny, and slightly centrotylote spicules
species originally described in Pachastrella belong to other genera, (8.5-15/-Lm X 4-6/-Lm). Small variations in microstrongyle mor-
and vice versa. The holotype of P. stylifera Lendenfeld, 1897d phology include oval and curved forms (Figs 2D, 1OF), depending
(BMNH 27.IIl.MC3) and P. tenuipilosa Lendenfeld, 1906 (BMNH upon specimens. There are also elongated, spiny worm-like
no registration number) are actually conspecific with Poecillastra spicules (25-50/-Lm X 2-3/-Lm) that have been occasionally
compressa. The holotype of P. amygdaloides Carter, 1876 (BMNH regarded to be microstrongyle-derived. Nevertheless, SEM obser-
00047; I. 1.2) also belongs to Poecillastra cf. compressa. Pachastrella vations (Fig. 2E) revealed that they are spiraster-like streptasters
connectens Schmidt, 1870 (BMNH I. 1.1; 70.5.3.4.5) and P. stellet- with a spiral shaft and reduced actines (Maldonado, 1993). Apart
todes Carter, 1885e (BMNH 1.1.1.1) are two valid species of from these microrhabdose streptasters, typical amphiasters transi-
Characella. Pachastrella lesinensis Lendenfeld, 1894 (BMNH tional to metasters occur, showing a 11-14/-Lm-long, relatively
96.11.5.32-34) and P. exostitus Schmidt, 1868 (BMNH 08.9.24.80) thick axis and short actines. The ectosomal skeleton consists of a
are two valid species of Stoeba (Fig. 12C-H). The holotype of feltwork of microstrongyles, irregularly reinforced by calthrops.
Yodomia peifecta Dendy, 1916c (BMNH MC5; RN.I to RN IX) and The choanosomal skeleton is made of oxeas and calthrops scattered
subsequent material collected by Burton (1956; BMNH 1936.3.4. in high density and with no preferential arrangement through the
184) are conspecific with Pachastrella ovisternata (Fig. 10Q-N). body; oxeas never protrude the ectosome. Streptasters occur rein-
forcing the internal side of the ectosomal skeleton and the
Description of type species endopinacoderms.
Remarks. The type species occurs throughout the Atlantic
Pachastrella monilifer Schmidt, 1868: 15 (Figs ID, lR, 2D, and western Mediterranean, in sublittoral and bathyal depths grow-
2H, lOA-N, 15R, 16D). ing on hard substrata in rocky and soft-bottom communities and
Synonymy. Pachastrella abyssi Schmidt 1870: 64; caves (Pouliquen, 1972; Bibiloni, 1990). There are also records of
Pachastrella caliculata Kirkpatrick, 1902: 227; Pachastrella isor- the species from the Pacific (de Laubenfels, 1935b), the Antarctic
rhopa Kirkpatrick, 1902: 228. (e.g., Koltun, 1964b), the Indian Ocean (Burton, 1959a), and the
Material examined. Holotype (fragment): BMNH China Sea (Levi & Levi, 1989). However, a global revision of
10.1.1.854 - Algeria. Slides of holotype: BMNH 1868.3.3.2. material should be undertaken before concluding that P. monilifera
Holotype of Pachastrella abyssi Schmidt, 1870: BMNH 00036; is a single cosmopolitan species. For example, some of the Indian
1870.5.3.59 (slides) - Florida. Other material. Slides of P. abyssi; ocean material revised here was found to belong to the related
Sollas, 1888: BMNH (no registration number) - South Atlantic. species P. ovistemata.
Slides of P. abyssi; Carter, 1876: BMNH Norman ColI. 10.1.1. Some authors have regarded P. monilifera as a species with
702-704: Cape St. Vincent. Slides of P. monilifera; Kirkpatrick, a variable spiculation, with dichotriaenes and mesodichotriaenes
1902: BMNH 24.5.1.74-75 - South Africa. Specimens of P. monil- occasionally occurring in addition to calthrops (e.g., Topsent,
ifera: BMNH 1936.3.4.144, 148, 149, 180 and 493 - Indian Ocean, 1902, 1904b). Nevertheless, a great deal of this reported variability
John Murray Expedition. BMNH 33.8.12.12ba, 131;132; is artificial and a consequence of lumping P. ovistemata into
33.8.13.133; 1947.2.15.3 - Siboga Expedition. BMNH 28.2.14.24 synonymy with P. monilifera. Indeed, P. ovistemata is a valid
to 26 - 'Discovery' Expedition. Specimens of P. monilifera; species characterized by the presence of long isodiametric, hispi-
Maldonado, 1993: CEAB-ALB-7-lOa, -13, -18, -30, -32 and -35- dating oxeas in addition to the typical, robust, fusiform choanoso-
western Mediterranean. Holotype of Pachastrella isorrhopa mal oxeas, as well as by the presence of dichotriaenes and
Porifera' Demospongiae • Astrophorida • Pachastrellidae 153

M
a
o

U
T ~
.....,; • 0 v

Fig. 10. Pachastrella. A-N, P. monilijera, holotype. A, fragment of oxea (scale 250 f-Lm). B, hastate oxeas and detail of their end (scale 75 f-Lm). C, calthrops
(scale 250 f-Lm). D, oval and lengthened microstrongyles (scale 25 f-Lm). E, amphiaster (scale 25 f-Lm). F-H, curved oval microstrongyles, spiraster-
like microstrongyles, and a sanidaster (from Indo-Pacific specimens collected by Burton, 1959a) (scale 25 f-Lm). I-N, oxeas, hastate oxeas, calthrops,
spiraster-like microstrongyles, oval microstrongyles, and metaster-amphiasters (from holotype of P. abyssi (now P. monilijera)) (scales: I, 250/-Lm, J, 50/-Lm,
K, 250 /-Lm, L-N, 25 /-Lm). O-W, Pachastrella ovisternata, from material formerly described as P. abyssi by Carter. 0, oxea and detail of oxea end (scale 250
/-Lm). P, hastate oxea (scale 75 /-Lm). Q, calthrops and detail of malformation (scale 250 /-Lm). R, dichotriaene (scale 150/-Lm). S, mesotriaenes (scale 150 /-Lm).
T, spiraster-like microstrongyles (scale 25 /-Lm). U, oval microstrongyles (scale 25 /-Lm). V,amphiaster (scale 25 /-Lm). W, plesiaster (scale 25 /-Lm).

mesodichotriaenes in addition to calthrops (Maldonado, 1993). Re- In the material re-examined here there are nominal species,
examination of material revealed that several BMNH specimens such as P. abyssi, P. caliculata, and P. isorrhopa, that are junior
catalogued under diverse names belong to P. ovisternata: Norman synonyms of P. monilifera. The presence of small dichotriaenes but
collection from Cape St. Vincent (Fig. IOQ-W) labeled as P. abyssi no mesotriaene in the holotype of P. chuni from Madeira suggests
O.S. by Carter (10.1.1. 1702-1704); Stephens's (1915a) material that this may be a valid species, although close to P. monilifera.
from the Irish coasts labeled as P. monilifera (S.R 151: 1953. The hastate oxeas described here in P. monilifera (Figs lR,
11.11.24); material from Madeira (1954.29.12); diverse material lOB, J) have traditionally been regarded as exogenous spicules.
from the Indian ocean, such as the holotype of Yodomia perfecta However, they have been found in the choanosome of all material
Dendy, I916c, and specimens of Y. perfecta and P. monilifera (Fig. of P. monilifera examined, including specimens originally
IOF-H) collected by Burton (1959a). Mesotriaenes also occur in an described under different names. Similar hastate oxeas also occur
undescribed Pachastrella from the Galapagos Islands (Fig. IF-G). in individuals of the close species P. ovistemata (Fig. 6P) as well
154 Porifera· Demospongiae • Astrophorida • PachastreUidae

as in the closely related genus Brachiaster (Fig. 6B), and con- 258; Nonnania crassiuscula Sollas, 1886a: 185; Poecillastra
sequently these oxeas are considered to be characteristic of both incrustans Sollas, 1888: 105; Pachastrella stylifera Lendenfeld,
genera. 1897d: 82; Pachastrella tenuipilosa Lendenfeld, 1906: 234.
Material examined. Holotype: BMNH BK.353, BK.354 -
slides from Ecinonemia compressa Bowerbank, 1866. Holotype of
POEClUASTRA SOLLAS, 1888 Nonnania crassa Bowerbank, 1874b: BMNH BK-831 to 832;
R.1631, R.1635. Specimens of Ecionemia compressa: BMNH
Synonymy 10.1.1.1686, 10.1.1.1688 - Norman coIL, Shetland. Specimens of
Nonnania compressa: BMNH 10.1.1.1129 to 1131- Norman colI.,
[Nonnania] Bowerbank, 1869a: 328 (preocc.). Poecillastra Norway. Specimens of Hymeniacidon placentula: BMNH
Sollas, 1888: 105. Nethea Sollas, 1888: 103. Chelotropaena 10.1.1.1684 - Norman colI., Shetland. Specimens of P. compressa:
Lendenfeld, 1906: 231. BMNH 1887.9.30.8 - NE Atlantic, colI. H.M.S. 'Triton'. BMNH
(no registration number) - Icelandic Ingolf Exp. Specimens of
Type species Stephens, 1915a: BMNH R.S. 483, 1953.11.11.19. Specimens of
P. compressa; Maldonado, 1992: CEAB-ALB-6-5, ALB-6-22,
Poecillastra compressa (Bowerbank, 1866: 55) (by original ALB-7-21b - Mediterranean, Alboran Sea. Holotype of
designation). Poecillastra incrustans Sollas, 1888: BMNH 94.11.16.134 to
135 - North Atlantic. Holotype of Pachastrella tenuipilosa
Definition Lendenfeld 1906: BMNH (no registration number) - Central
Atlantic. Holotype of Stelleta scabra Schmidt, 1868: BMNH
Pachastrellidae, the microscleres of which include always 68.3.2.41 - Algeria.
streptasters in the form of spirasters and plesiasters, as well as Description. Polymorphous sponge often showing a plate-
microxeas in a single category. like (as the holotype) or cup-like body form, with punctiform osti-
oles distributed on one side of the sponge and 1-3 mm oscules
Diagnosis usually provided with a contractile diaphragm and placed on the
opposite sponge side. More rarely, the species is encrusting or
Pachastrellidae whose megascleres are calthrops and/or short- massive. Alive specimens are whitish or pale yellowish. The
shafted triaenes scattered through the whole choanosome. spicules of the holotype consist of oxeas, calthrops and short-
Microscleres consist of microxeas, always in a single size category, shafted orthotriaenes, microxeas in a single size category, large
and several streptaster types, always including small spirasters and plesiasters transitional to amphiasters, and spirasters. Oxeas are
large plesiasters (transitional to metasters or amphiasters) with fusiform, curved in the middle, with sharp ends, measuring
scarce, long actines. 960-2080 X 15-35 /Lm. Calthrops and short-shafted orthotriaenes
are relatively scarce, with actines measuring 125-400 X
Remarks 10-20/Lm. Tetraxons are frequently malformed and one or more
actines can be missing in many spicules; the rhabdome is typically
[Nonnania] Bowerbank, 1869a: 328 is preoccupied by missing in those spicules that reinforce the ectosome. Microxeas
Nonnania Brady, 1866, Crustacea, and therefore Poecillastra is the are finely spiny, slightly curved, sometimes centrotylote, usually
next available name. Re-examination of diverse museum collections measuring 125-200 X 3-5 /Lm. Plesiasters transitional to amphi-
discovered several species of Poecillastra that were originally asters have three to five long actines relative to the central shaft;
assigned to other genera, and vice versa. The holotype of they are 35-55 /Lm in their greater diameter, with actines measur-
Chelotropaena tenuirhabda Lendenfeld, 1906 (BMNH 02.2.9. ing 12-15 X 1 /Lm. Spirasters have a relatively thin axis of several
104-106), from the Magellan Strait, appears to be a valid species of revolutions and numerous, relatively large actines, measur-
Poecillastra with short-shafted plagio- and dichotriaenes ing15-25 /Lm in total length. The ectosomal skeleton consists of a
(Fig. 11 G-N). Stelleta scabra Schmidt, 1868 (Fig. II00R; BMNH layer of tangential microxeas and streptasters. It is internally rein-
68.3.2.41) from Algeria and Pachastrella tenuipilosa Lendenfeld, forced by calthrops, and bundles of oxeas that emerge from the
1906 (BMNH no registration number) from the Central Atlantic are choanosome and become brushes just under the ectosome. Oxeas
conspecific with Poecillastra compressa (Bowerbank, 1866). It is very rarely hispidate the sponge surface. Microxeas and strep-
also worth mentioning that the skeleton of the holotype of tasters also occur in the choanosome, with streptasters preferen-
Sphinctrella theneides Burton, 1959a (BMNH 1936.3.4.313a tially reinforcing the endopinacoderms.
to 314a) looks very similar to that of Poecillastra, despite the fact Remarks. The type species occurs in the northeastern
that the sponge is subspherical, with a single osculum on the top and Atlantic and the western Mediterranean over a wide depth range
a single areolated, depressed area on the underside, both fringed by that goes from intertidal habitats (Sara, 1964b) to depths of 3500 m
long, hispidating spicules, as typically found in Vulcanella. (Koltun, 1970).
It has been suggested several times (e.g., Topsent, 1894d;
Burton, 1930c) that the Indo-Pacific P. laminaris (Sollas, 1886a)
Description of type species (BMNH 94.11.16.120-130), the Southern Ocean P. schulzii Sollas,
1886a (BMNH 27.MC4, no number), and the Japanese P. tenuilam-
Poecillastra compressa (Bowerbank, 1866) (Figs 21, lIA-F). inaris Sollas, 1886a (BMNH 27.MC4, no number) are conspecific
Synonymy. Ecionemia compressa Bowerbank, 1866: 55; with the Atlantic-Mediterranean P. compressa. Re-examination of
Stelleta scabra Schmidt, 1868: 19; Hymeniacidon placentula the respective holotypes confirmed that P schulzii is a valid
Bowerbank, 1874b: 189; Nonnania crassa Bowerbank, 1874b: species. Unlike P. compressa, it possesses two categories of oxea
Porifera • Demospongiae • Astropborida • Pacbastrellidae 155

-
E

- -

dz5PL!;' ~'~

- =--
)(~ *P:M-«3:::."~_
Fig. 11. Poecillastra. A-F, P. compressa. holotype. A, oxea (scale 125 fLm). B, calthrop, short-shafted plagiotriaene and some malfonnations (scale 125 fLm).
C, triactinal calthrop tangential to the ectosome (scale 125 fLm). D, plesiaster (scale 25 fLm) . E, spiraster (scale 25 fLm). F, microxeas (scale 40 fLm) . G-N,
P. tenuirhabda. G, oxea (scale 250 fLm) . H, short-shafted plagiotriaene (scale 250 fLm) . I, short-shafted dichotriaene (scale 250 fLm). J, diactinal, tetractinal
and pentactinal plesiasters (scale 40 fLm). K, detail of plesiaster actine (scale 10 fLm). L, spirasters (scale 40 fLm). M, detail of spiraster actine (scale
10 fLm) . N, mature and thinner, developing microxeas (scale 50 fLm) . O-R, Microscleres of Stelleta scabra (now Poecillastra compressa), holotype.
O. plesiasters (scale 20 fLm). P, plesiasters transitional to amphiasters (scale 20 fLm). Q, spirasters (scale 20 fLm). R, microxea (scale 20 fLm) .

megascleres: fusiform, robust, choanosomal oxea and isodiametric, as valid species, the former occurring in the Indo-Pacific and the
slender ectosomal oxeas that hispidate the sponge surface. Southern Ocean and the latter occurring in the North Pacific from
Poecillastra lamifUlris from the Indo-Pacific is possibly a junior the Japan Sea to the American coasts.
synonym of P. schulzii, as well as P. eccentrica Dendy & Burton,
1926 (BMNH R.N.V.I. 26.10.1.95) from the same area. In contrast,
P. tenuilaminaris, which lacks the slender ectosomal oxeas, is STOEBA SOLLAS, 1888
probably a valid species. It resembles P. compressa in its skeleton
but its plesiasters are transitional to metasters and are much smaller Synonymy
(only up to 22 f.Lm in largest diameter). Diverse material from the
Pacific coast of North America all appear to be conspecific with Stoeba Sollas, 1888: 102. CalcabrifUl Sollas, 1888: 280.
P. tenuilaminaris, including material of P. tenuilaminaris, de Dercitancorina Topsent, 1902: 13. HalifUlstra de Laubenfels,
Laubenfels (1932) and Dickinson (1945), the holotype of 1936a: 179. [Dercitus pars]
Poecillastra rickettsi de Laubenfels, 1930 and subsequent
Californian specimens (BMNH 19.8.22.7a, 29.8.22.7a) described Type species
by de Laubenfels (1932), which are contaminated with Vulcanella
spicules. Thus, P. schulzii and P. tenuilamifUlris should be regarded Samus simplex Carter, 1880a: 60 (by original designation).
156 Porifera· Demospongiae • Astrophorida • Pachastrellidae

Definition Synonymy. Samus simplex Carter, 1880a: 60; Dercitus

simplex, Thiele, 1900: 20.
Pachastrellidae having as microscleres only one or more cate- Material examined. Specimen of Dercitus plicatus var.
gories of peculiar microrhabd-like sanidasters characterized by a simplex (Carter): BMNH (Ind. Mus. Coll. 31.1.1.21a) - Indian
relatively thick axis and reduced actines. Ocean, Adaman Sea, Invisible Bank.
Description. Encrusting sponge, filling excavated cavities
Diagnosis in calcareous structures. The skeleton consists of two spicule types,
short-shafted dichotriaenes and sanidasters. Dichotriaenes with
Pachastrellidae in which the only megascleres, in most protoclads being clearly shorter than deuteroclads. In BMNH spec-
cases, are calthrops or short-shafted dichotriaenes; oxea mega- imens, protoclads are 30-50 X 12-60/-Lm and deuteroclads are up
scleres are absent in most species but are present in two. to 150 X 35-40 /-Lm; rhabdome is 40-225 X 60-75/-Lm, and 210 X
Microscleres are sanidasters with a relatively thick axis and 42/Lm in the holotype. Sanidasters have a thick, slightly twisted
reduced actines, resembling spiny microrhabds. All known axis and reduced actines, measuring 22-28 X 4--6 /Lm in total
species are encrusting and often filling cavities in calcareous length. The sanidasters are extremely abundant in the ectosome,
substrata. making a feltwork that is reinforced at their inner side by the
cladome of the triaenes; the rhabdomes point inwards.
Remarks ReTlUlrks. The type species is known from the Indian Ocean
(Gulf of Manaar, Adaman Sea), excavating Melobesian nodules
Sollas (1888) proposed Stoeba for Samus simplex Carter, and other calcareous structures, from uncertain depth.
1880a from the Gulf of Manaar, but Lendenfeld (1903) followed
by Topsent (1904b) and most modem authors merged it in
Dercitus, whose type species possesses toxa. Following Dendy THENEA GRAY, 1867
(1905), it is preferred here to retain the distinction between these
two taxa for species with or without toxas given that the evolution- Synonymy
ary origin of toxas within the order Astrophorida remains enig-
matic and intriguing. Thenea Gray, 1867a: 541. Tisiphonia Thomson, 1869a: 712.
Apart from the type species, other apparently valid species of Dorvillia Kent, 1870c: 293. Wyvillethomsonia Wright, 1870: 80.
the genus are S. plicatus (Schmidt, 1868) from the Mediterranean [Dorvilia] Pocta, 1883: 382 (lapsus). Clavellomorpha Hansen,
(BMNH 1.2.4), S. occultus (Hentschel, 1909) from Shark Bay, 1885: 19. Thisiphonia Lendenfeld, 1887b: 563. Theuca Lundbeck,
Western Australia, S. extensa Dendy, 1905 from Mutwal Island, Sri 1907: 559.
Lanka (BMNH 07.2.1.5a; 338.4.7a), S. natalensis Burton, 1926
from South Africa, and S. syrmatitus (de Laubenfels, 1930) from Type species
California (BMNH 1. 21). In addition, Pachastrella lesinensis
Lendenfeld, 1894 from the Adriatic (BMNH 96.11.5.32-34), Tethea muricata Bowerbank, 1858: 308 (by original designation).
which is characterized by having oxeas, calthrop-like tetraxons and
sanidasters with rounded ends, is also a valid species of Stoeba Definition
(Fig. 12F-H). Although its sanidasters were interpreted as
microstrongyles by Lendenfeld, they are not like those in Pachastrellidae with specialized poriferous areas and root-like
Pachastrella but are irregular and slightly twisted. Pachastrella processes for attachment to the substratum, which are partially
lesinensis Lendenfeld is also the type species of Dercitancorina made of flexuous anatriaenes. Its microscleres always include
Topsent (1902: 13) (by original designation), which makes streptasters in the form of plesiasters; microxeas are absent.
Dercitancorina a junior synonym of Stoeba. Apart from S. lesinen-
sis, monaxonic megascleres are only known from material Diagnosis
described as Stoeba spp Vacelet & Vasseur, 1971, in which thin
strongyloxeas along with calthrops and sanidasters were found. Distinctive ovate or mushroom-shaped sponges, with radial or
Pachastrella exostitus Schmidt, 1868 from the Red Sea bilateral symmetry, with one to few oscula at upper portion of the
(BMNH 08.9.24.80), characterized by calthrops, thick sanidasters body, and with at least part of the ostia grouped in a one or more
and tiny spheraster-like spicules with reduced actines is also a valid inhalant areas located near the equator of the body, in a transversal
species of Stoeba (Fig. 12C-E). SEM observations have revealed recess. Because most species live in deep, soft bottoms, the
that the spheraster-like spicules are slightly elongated, thus being a sponges are provided with basal, root-like processes for anchoring
kind of compressed sanidasters or ataxasters (Fig. 2F-G). to the sediment or buried substrata. The transversal recess, some-
Consequently, because Pachastrella exostitus Schmidt, 1868 was times provided with a protruding palisade of spicules, is thought to
also designated the type species of Halinastra de Laubenfels, prevent the inhalant areas from clogging by the fine silt that char-
1936a, this latter genus becomes now a junior synonym of Stoeba. acterizes deep-sea habitats (Fig. 3A-C). The skeleton consists of at
It is also concluded here that Stoeba loricatus (Lebwhol, 1914) is least a type of monaxons (oxeas and/or oxeotylotes), one or more
not a valid species, its holotype (BMNH 1942.6.12.11) being con- types of long-shafted tetraxons (dichotriaenes, anatriaenes,
specific with Pachastrella c.f. monilifera: mesoanatrienes, protriaenes, mesoprotriaenes, and/or orthotri-
aenes), and one or more categories of streptasters (plesiasters,
Description of type species amphiasters, metasters, spirasters); true microxeas are always
absent, although plesiasters reduced to two-ray, microxea-like
Stoeba simplex (Carter, 1880a) (Fig. 12A-B). forms may occur in some species.
Porifera· Demospongiae • Astrophorida • Pachastrellidae 157

F
.-::;;:::::;:;:~~--- -- -- -.::=:::::::::::::::~

Fig. 12. Stoeba. A-B, S. simp/ex. A, dichotriaenes (scale 100 .... m). B, sanidasters (scale 25 .... m). C-E, S. exostitus, holotype. C, calthrops (scale 100 .... m).
D, sanidasters (scale 15 ....m). E, ataxaster (scale 15 ....m). F-H, S. /esinensis , holotype. F, oxea (scale 150 .... m). G, calthrops (scale 100 ....m). H, sanidasters
(scale 15 .... m).

Remarks of oxeas, dichotriaenes, protriaenes and anatriaenes as megascleres;

microscleres are plesiasters and spirasters. Oxeas are isodiametric,
Owing to the large size that the plesiasters attain in some curved or flexuous, measuring in the holotype 2000-5000 X
species (e.g., T. novazelandiae Bergquist, 1961c) they have occa- 10-15 J.Lm, but up to 15 mm in giant individuals. Dichotriaenes
sionally been misinterpreted as megascleres and described as spiny have conical, long, somewhat flexuous rhabdomes, measuring
calthrops. However, true calthrops have never been found in Thenea. up to 5000-6000 X 150 J.Lm; protoclads are shorter than deutero-
clads, measuring 100-330 X 30-100 J.Lm and 300-1100 X 30-80
Description of type species J.Lm, respectively. Protriaenes have a straight rhabdome measuring up
to 5000 X 30 J.Lm, and curved or straight clads measuring 150-400 X
Thenea muricata (Bowerbank, 1858) (Figs lA, 3A-C, 13). 12-20 J.Lm. Anatriaenes have thin, flexuous rhabdomes measuring up
Synonymy. Tethea muricata Bowerbank, 1858: 308; Thenea to 300 X 25 J.Lm, and straight, conical clads measuring 40-140 X
muricata, Gray 1867 a: 541; Wyvillethomsonia wallichii Wright, 15-20 J.Lm. The form of the asters is affected by some inter-individual
1870: 8; Dorvillia agariciformis Kent, 1870c: 293; Tisiphonia variability; in the holotype, the plesiasters have 2 to 6, finely spiny,
agariciformis, Thomson, 1873: 167; Clavellomorpha minima 40-100 J.Lm long actines. The spirasters, which measures 25-35 J.Lm
Hansen, 1885: 19; Thenea schmidtii Sollas, 1886a: 183; Thenea in total length, occur in two morphologies: (1) with relatively few,
intermedia Sollas, 1888: 97. long actines and a thin central shaft; (2) with many short actines and
Material examined. Holotype: BMNH 1.1.7. Other mate- a relatively thick, central shaft. The latter category occurs exclusively
rial. Specimens of Thenea muricata: CEAB collections - Gibraltar at the ectosome of the inhalant areas. The skeletal arrangement fol-
Straits (Maldonado, unpublished). Comparative material. BMNH lows a clearly radial pattern. The oxeas, along with the rhabdomes of
types of T. delicata Sollas, 1886a, T. megastrella Lendenfeld, 1903, the tetraxons, form fascicles that penetrate radially from the sponge
and T. grayi Sollas, 1886a; Sollas's (1888) material of T. Jenestrata surface to the innermost choanosome. The oxeas also protrude
(Schmidt, 1880b), ZIL material of T. delicata Koltun (1964b). beyond the sponge surface, giving support to the budding processes
DescriptWn, Ovate to mushroom-shaped sponges, 0.5 to 2cm for asexual reproduction and forming the hispid, equatorial palisade
in diameter, white to gray in color. Giant individuals - up to 12cm that protects the inhalant areas. The cladomes of the tetraxons, along
in diameter - have been described from the Norwegian fjords with the spirasters, reinforce the ectosome, and may protrude the
(Steenstrup & Tendal, 1982). One or few open oscules (2-5 mm in sponge surface here and there. Spirasters and plesiasters occur every-
diameter) placed at the upper part of the body. One or more areolated where in the body, the former being more abundant in the ectosome
areas located within a transversal recess that may also be provided and the latter ones in the endosome. The rootlet processes are mainly
with a protruding palisade of megascleres. This spicule palisade may made by entangled anatriaenes.
form a complete rim at the equator of the sponge body, occur at some Remarks. The abundant literature on this species indicates
points only, or, more rarely, be absent, depending upon individuals. that T. muricata shows a significant intraspecific variability in
Most individuals develop a variable number of flexible root-like external morphology, dimensions and abundance of the various
processes made of naked, entangled spicules that allow the anchoring spicule types (Boury-Esnault et al., 1994b). The type species
of sponges in soft bottoms. At the upper part of the body, budding occurs in the northeastern Atlantic and the Mediterranean, mostly
processes for asexual reproduction may occur. The skeleton consists in deep waters (120-4020m depth) and preferably in soft-bottom
158 Porifera' Demospongiae • Astrophorida • Pachastrellidae

Fig. 13. Thenea. T. muricata, holotype. A, oxea (scale 300 !Lm). B, protriaenes (scale 150 !Lm). C-D, dichotriaenes (scale 250 !Lm). E, choanosomal anatri-
aene (scale 150 !Lm). F, anatriaene from the root-like processes (scale 150 !Lm). G-H, plesiasters in a wide size range (scale 50 !Lm). I, spirasters (scale 50 !Lm).

communities. There are also records of the species in sublittoral Remarks

caves (Russ & Riitzler, 1959; Sara, 1964b; Riitzler, 1966).
There are four known species in the genus: T. intextus (Carter,
1876) from Cape St. Vincent, northeastern Atlantic (Fig. 14B-E;
TRIPTOLEMMA DE LAUBENFELS, 1955 BMNH 10.1.1. 1693-97), growing on and in Corallistes bower-
bankii; T. incertus (Kirkpatrick, 1903a), from South Africa
Synonymy (Figs lH-I, 2C, 14F-I; BMNH 02.5.2b-5b), growing on and in
Discodermia natalensis; T. simplex (Sara, 1959a), from the
[Triptolemus] Sollas, 1888: 93 (preocc.). Triptolemma de Mediterranean, growing within various demosponges; T. cladosus
Laubenfels, 1955b: E43. (Sollas, 1888) from Ki Island, Indian Ocean. This latter species
possibly also occurs in the Gulf of Manaar (Carter, 1880a), where
Type species it has been regarded as an Indian population of the North-Atlantic
Samus parasiticus Carter, 1876. Indeed the holotype of S. parasiticus
Triptolemma cladosus (Sollas, 1888: 93) (by original from Cape St. Vincent appears to be conspecific with T. intextus
designation). and was proposed by Levi & Levi (1983a) to be a junior synonym
of T. intextus.
Definition Monaxonic megascleres are absent in all species, since,
according to Sara (1959a), those reported in the literature of
Cryptic Pachastrellidae having megascleres exclusively short- T. cladosus and T. intextus (Fig. 14B) presumably belonged to the
shafted mesotriaenes with diversely branched clads, but never host sponges.
becoming mesotrider desmas.
Description of type species
Diagnosis
Triptolemma cladosus (Sollas, 1888) (Fig. 14A).
Pachastrellidae that penetrates the tissue (parasitises?) of Synonymy. Samus quadripartita Carter (unpublished
other sponges. It is characterized by megascleres that are exclu- manuscript).
sively short-shafted mesotriaenes with diversely branched clads. Material examined. Three slides containing also spicules of
Microscleres are streptasters in one or two categories, with microx- the host sponge Corallistes thomasi (BMNH 1891.5.4.12) were
eas occurring in some species. examined, but only a few microscleres potentially belonging to the
Porifera· Demospongiae • Astrophorida • Pachastrellidae 159

Fig. 14. Triptolemma. A, T. cladosus, holotype, mesotriaene with irregular, tetrafurcated clads (redrawn from Sollas, 1888) (scale 35 11m). B-E, T. intexta,
holotype (redrawn from Carter, 1876). B, oxeas, the exogenous nature of which has been suggested (scale 50 11m). C, short-shafted mesotriaene with tetra-
furcate clads (scale 50 11m). D, sanidasters (scale 10 11m). E, metasters transitional to amphiasters; note that microxeas occur, but are not illustrated (scale
10 11m). F-I, T. incertus. F, short-shafted mesotriaenes with diversely branched clads (scale 100 11m). G, spiny microxeas (scale 20 11m). H , sanidasters
(scale 10 11m). I, metasters (scale 10 11m).

holotype of T. cladosus were found. No other type material appears most evidence suggests that all four known species of Triptolemma
to be available. Therefore, the holotype description below follows lack oxeas.
Sollas (1888: 93).
Description. Encrusting specimen growing on and penetrat-
ing the tissue of a Corallistes. The skeleton consists of short-shafted VULCANELLA SOLLAS, 1886
mesotriaenes, microxeas and sanidasters (formerly described as tri-
chose microxeas) and spirasters. Mesotriaenes have symmetrical, Synonymy
short and blunt rhabdomes that measure up to 60 X 21 fLm each;
clads are irregularly trifurcated or tetrafurcated, with desmoid [Sphinctrella] Schmidt, 1870: 65 (preocc.). Vulcanella Sollas,
appearance. The dichotomy of the cladome is not confined to a sin- 1886a: 186. Sphincterella de Laubenfels, 1936a: 180.
gle plane and the level of division also varies within a spicule, co-
occurring dichotomous and tetrafurcated clads; protoclads are up to Type species
52 X 21 fLm, deuteroclads up to 55 fLm long, tritoclads up to
27.6 fLm long, and tetraclads up to 27.6 fLm long. Microxeas were Vulcanella horrida (Schmidt, 1870: 65) (by original
formerly described as being smooth - although this is unsupported designation).
by the current knowledge on other species in the genus - and more
or less fusiform, measuring up to 118 X 4 fLm. Sanidasters have a Definition
relatively thin axis covered with minute, erect spines and measure
up to 27.6 fLm in total length. Spirasters have a thin axis provided Pachastrellidae with special cribriporal oscula surrounded by
with relatively long and thin actines and measure up to 11.8 fLm in a palisade of protruding oxeas (Fig 3D-E) and with one or more
total length. The skeletal arrangement consists of abundant microsclere types showing an annulate decoration (Fig. 2B).
sanidasters forming an ectosomal feltwork in which the cladomes of
the mesotriaenes are embedded tangentially to the sponge surface. Diagnosis
ReTfUlrks. Sollas (1888) was unsure about whether oxea that
he observed belonged to Triptolemma or to its host Corallistes. Encrusting, thick lamellate or submassive Pachastrellidae in
Although the issue remains unsolved, Sara (1959a) pointed that which oscula are grouped in specialized exhalant areas - called
160 Porifera· Demospongiae • Astrophorida • Pachastrellidae

'fenestrate cloacas' sensu Sollas (1888) - and fringed by a protrud- Definition

ing palisade of flexuous, atrial oxeas. Oxeas are in two categories,
slender atrial oxeas and stout choanosomal oxeas. Tetraxonic Vulcanella lacking tetraxonic megascleres and possessing
megascleres are absent in some species (subgenus Annulastrella n. large plesiasters with two to six annulate actines.
subg.), but calthrops transitional to short-shafted triaenes, occa-
sionally dichotriaenes, are present in most species (nominal sub- Diagnosis
genus Vulcanella). Microscleres consist of one or two categories of
microxeas, which may be absent in some species, and one to sev- Vulcanella lacking tetraxonic megascleres. Oxeas may be
eral types of streptasters. Streptasters are usually metasters and spi- either absent or present in one or two categories (atrial and
rasters (Fig. 2K), more rarely amphiasters; triactinal and choanosomal oxeas), depending on species. Microscleres are large
calthrop-like plesiasters with annulate actines also occur in the plesiasters with two to six annulate actines (traditionally called
subgenus Vulcanella. microtriods and microcalthrops), along with one or two categories
of smaller streptasters (spirasters, metasters or amphiasters).
Remarks Although annulate microxeas have been described in some species,
it is likely that they are plesiasters reduced to a diactinal stage in
Some species of Vulcanella have small both calthrop-like and most cases, except those described by Topsent (1923) in material of
triactinal spicules. These microscleres have been misinterpreted by Stellettinopsis annulata Schmidt, 1880b.
some authors, who regarded them as true megascleres, called
microtriods and microcalthrops. However, accumulation of obser- Remarks
vations over time has made it clear that the number of actines in
these spicules may range from 2 to 6, depending on the species. Apart from the type, three species are included in this sub-
This indicates that they are derived from asters, possibly plesiasters genus. Sphinctrella verrucolosa Pulitzer-Finali, 1983, from the
with an extremely short central axis. Similarly, the large plesiasters western Mediterranean, is characterized by atrial and choanosomal
with four actines that occur in some species of Thenea have occa- oxeas, spirasters (ranging to metasters and amphiasters), annulate
sionally been misinterpreted as true megascleres. It is interesting plesiasters with 2-6 actines, and a category of annulate, centroty-
that asters converging on calthrop-like morphologies also occur in lote microxeas, which probably are plesiasters reduced to a diacti-
other unrelated sponge lines, such as, for instance, in the nal stage. Sphinctrella ornatus Sollas, 1888 (Sollas, 1888; Topsent,
hadromerids Timea cumana and T. tetractis (e.g., Maldonado, 1992). 1892a, 1904b, 1928c) from the Azores and Cape Verde (north-
In most species of Vulcanella, one or more types of microscle- eastern Atlantic Ocean), is characterized by atrial and choano-
res (i.e., the largest microxeas and plesiasters) show an annulate somal oxeas, spirasters, metasters (to amphiasters), and annulate
ornamentation which originates from the coalescence of plesiasters, the number of actines of which varies from 2-5.
microspines to make verticils or spires (Fig. 2B). This ringed orna- Stellettinopsis annulata Schmidt, 1880b, from the Gulf of Mexico,
mentation may be a useful and reliable diagnostic tool at the is characterized by atrial and choanosomal oxeas, spirasters,
generic level, although there are some species where this feature metasters, plesiasters with three to four finely annulate actines, and
can only be seen under SEM (e.g., V. cribrifera, V. aberrans, spiny, non-annulate microxeas (Topsent, 1923). Note that
V. cribiporosa). Furthermore, there are many species described Stellettinopsis annulata Schmidt, when transferred to the genus
prior to the use of SEM techniques for which we have no data on Vulcanella (Annulastrella), becomes a junior secondary homonym
ringed ornamentation, requiring comprehensive re-examination of the type species Vulcanella (Annulastrella) annulata Carter, and
using new techniques to assess the utility of spiny verticils in is renamed here as Vulcanella (Annulastrella) schmidti nom. nov.,
microxeas and/or streptasters as a diagnostic trait. after its former author.
In an attempt to organise the skeletal variability in Vulcanella, Topsent (1923, 1928c) considered the species mentioned
two subgenera are proposed here. Nominal subgenus Vulcanella is above, excluding the recently described V. verrucolosa, to be junior
erected for Vulcanella species that always have tetraxonic mega- synonyms of V. annulata (Carter). However, small differences in
scleres and microxeas, but lack triactinal and tetractinal plesiasters their skeletons and the enormous geographical distance between
with annulate actines. Annulastrella subgen. nov. is created for populations make it sensible to maintain the Mediterranean, the
those species that lack tetraxonic megascleres, but possess triacti- Indian, the north-eastern and north-western Atlantic material as
nal and calthrop-like plesiasters with annulate actines, with or separate species.
without microxeas as microscleres. Triassic and Cretaceous species of the genus Monilites Carter,
1871b are apparently related to Vulcanella, as they have similar,
annulate microxeas and plesiasters (see Wiedenmayer, 1994). In my
VULCANELLA (ANNULASTRELLA) SUBGEN.NOV. opinion, no relationship can be supported between the annulate
microscleres of Vulcanella and the cricostyles and ringed protriaenes
Type species found in Jurassic-Cretaceous deposits (see Wiedenmayer, 1994).

Vulcanella (Annulastrella) annulata (Carter, 1880a: 140) (by Description of type species
monotypy).
Vulcanella (Annulastrella) annulata (Carter, 1880a) (Fig. 15).
Synonymy Synonymy. Tisiphonia annulata Carter, 1880a: 140.
Material examined. None. The description is taken from
Tisiphonia annulata Carter, 1880a: 140; Sphinctrella annulata, Carter (1880a) from specimens dredged up from the Gulf of
Sollas 1888: 100. Manaar (Indian Ocean).
Porifera· Demospongiae • Astrophorida • Pachastrellidae 161

A
-
--------::==-- ~-
~~
Fig. 15. Vulcanella (Annulastrella). V. (A) ornata, holotype. A, cloacal oxea (scale 250 fLm). B, choanosomal oxea (scale 250 fLm). C, triactinal and
tetractinal, annulate, plesiasters (scale 50 fLm). D, spirasters (scale 30 fLm). Spicules illustrated are very similar to those of the type species, V. (A) annulatus,
see Carter (l880a).

Description. Massive, white sponge measuring 2.12mm in that covers an osculum remains as the only extant type material for
diameter and growing among the detritus attached to a large, mas- this species. BMNH BK 1251, BK 1252a-f - slides from two spec-
sive specimen of another astrophorid sponge (from Carter, 1880a). imens labeled as S. horrida by Bowerbank. Comparative material.
The megascleres consist of fusiform, slightly curved, oxeas meas- Holotype of Sphinctrella horrida Schmidt, 1870: BMNH
uring 885 X 20/-Lm. The microscleres consist of four-rayed plesi- 70.5.3.47. BKI2524a-f, BK1251 - material from America and
asters, the actines of which measure up to 240 X 20 /-Lm and are Florida labeled by Bowerbank. Holotype of Sphinctrella gracilis
covered by minute micros pines arranged in marked rings; there are Sollas, 1888: BMNH 94.11.16.144-146 (slides) - Cape Verde.
also spirasters with a relatively thin axis and numerous long MSNG NIS.70.2, PF.455, CEAB-ALB-7-11cx - Mediterranean
actines, measuring above 12.7 /-Lm in total length. specimens described by Pulitzer-Finali (1983) and Maldonado
Remarks. The type species is known from the Indian Ocean (1993), respectively. Syntypes of Sphinctrella aberrans
(Gulf of Manaar and Ambon; Carter, 1880a; Topsent, 1897a; Maldonado & Uriz, 1996a: CEAB-POR.BIO 173a-c - Alboran
Desqueyroux-Faundez, 1981) Sea. Specimens of Vulcanella tricomis (Wilson, 1904): HBOI,
unpublished - Galapagos Islands. Holotype of Sphinctrella
cribrifera Sollas, 1886a: BMNH 89.1.1.39 - Cape Verde. MSNG
VULCANELLA (VULCANELLA) SOLLAS, 1886 PF-268: - Mediterranean specimens erroneously reported by
Pulitzer-Finali (1983) as S. horridn. Holotype of Sphinctrella
Type species theneides Burton, 1959a: BMNH 1936.3.4.313a-314a - Maldives
Island. Holotype of Sphictrella omatus Sollas, 1888: BMNH
Vulcanella (Vulcanella) horrida (Schmidt, 1870): 65. 94.11.16.146-149 - Cape Verde. BMNH SR.353, 1953.1l.II.27 -
material described Stephens (1915a) from Irish waters.
Definition Description. A 2-3 cm thick plate, with rounded edges,
rough surface, and oval elongated atriums lined by a fenestrated
Vulcanella species that have tetraxonic megascleres and membrane and fringed by a palisade of long spicules (Sollas,
microxeas but lack triactinal and tetractinal plesiasters with annulate 1888). The skeleton consists of atrial and choanosomal oxeas,
actines. calthrops, microxeas in two categories, and spirasters. Only a bro-
ken, 12 /-Lm-thick, atrial oxea occurs in the type slide, but fusiform,
Diagnosis slightly curved choanosomal oxeas are up to 3000 X 85 /-Lm; oxeas
measuring up to 5000 X 142 /-Lm were reported in a redescription of
Nominative subgenus containing Vulcanella species that have the type (Sollas, 1888). Calthrops are mostly well-formed, with
tetraxonic megascleres and microxeas but lack triactinal and clads measuring 400-1100 X 40-100 /-Lm. Microxeas in the large
tetractinal plesiasters with annulate actines. Microxeas are spiny, category are fusiform, stout, with a very patent ringed ornamenta-
with spines arranged in either a marked annulate pattern or a nearly tion, and measure 200-500 X 8-13 /-Lm. Microxeas in the small cat-
uniform distribution, depending upon species. egory measure 100-180 X 3-4.5 /-Lm, being comparatively slender,
finely spiny, and with a ringed pattern only perceptible around the
Description of type species middle of the spicule. Spirasters are 12-18/-Lm in length, with
numerous, short actines and a relatively thick axis with two or three
Vulcanella (Vulcanella) horridn (Schmidt, 1870) (Fig. 16B-F). revolutions.
Synonymy. Sphinctrella horrida Schmidt, 1870: 65. Remarks. The type species is recorded from the North
Material examined. Holotype (slide): BMNH-70.5.3.4a - Atlantic, from Florida (e.g., Schmidt, 1870; unpublished material
containing only a piece of peripheral choanosome and the ectosome labeled by Bowerbank in BMNH) to the Azores (e.g., Topsent,
162 Porifera· Demospongiae • Astrophorida • PachastrelUdae

E
--=

~ F

Fig. 16. Vulcanella (Vulcanella). V. (V.) horrida, holotype. A, atrial oxea (scale 300 j.Lm). B, choanosomal oxea (scale 300 j.Lm). C, caltrop (scale 200 j.Lm).
D, large, annulate microxeas (scale 50 j.Lm). E, small, unifonnly spiny microxeas (scale 50 j.Lm). P, spirasters (scale 15 j.Lm).

1904b, 1928c), at depths of 60-2500m. It has erroneously been re-examination, Gustavo Carreras for help with line drawings,
reported in the Mediterranean (Pulitzer-Finali, 1983) as result of a Carmen Carmona and Ram6n Coma for help with processing of
misidentification of fragmentary material probably belonging to digital figures, and Esther Cuiiado and Xus Cartes for help with
Vulcanella cribrifera (see Maldonado & Uriz, 1996a). bibliographic compilation. I am indebted to Vanessa Pike and Clare
Valentine for supporting my application to the Large-Scale
ACKNOWLEDGMENTS Facilities (LSF) Program to visit the BMNH facilities. This
research has been supported by European Community funds pro-
I thank Prof. Claude Levi, Prof. Jean Vacelet, and Ms. Clare vided by the LSF Program (1998) of the Natural History Museum
Valentine for help in locating and providing material for of London and a grant of the Spanish Government (PB 98-0485).
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Thrombidae Sollas, 1888

Mana J. Uriz

Centre d'Estudis Avan~ats de Blanes, Cami de Sta Barbara sin 17300. Blanes (Girona), Spain. (iosune@ceab.csic.es)

Thrombidae Sollas (Demospongiae, Astrophorida) contains only a single genus and five species to date, characterized by peculiar spiny
triaenes (acanthotriaenes) of a size intermediate between that of megascleres and microscleres, often with clads divided dichotomously or
trichotomously. Microscleres, when present, are streptasters (amphiasters). No clear relationships of this family with any other sponge
group have been established.
Keywords: Porifera; Demospongiae; Astrophorida; Thrombidae; Thrombus.

DEFINITION, DIAGNOSIS, SCOPE from Halinidae de Laubenfels (with streptasters), whereas

Thrombus lacks euasters but has a form of streptasters (amphi-
Synonymy asters). More recently, Levi (1973), followed by contemporary
authors (e.g., Hooper & Wiedenmayer, 1994), revived Thrombidae
Thrombidae Sollas, 1888. for Thrombus in recognition of its very peculiar characteristics.
Hooper & Wiedenmayer (1994), however, also included in the fam-
Definition ily two genera incertae sedis, previously included in Spirasigmidae
(Hallmann, 1912), with the consequence that the latter family
Astrophorida with diplodal aquiferous system. The spicule became ajunior synonym ofThrombidae. However, Spirasigmidae,
complement consists of small spiny triaenes (acanthotriaenes) with as proposed by Hallmann (1912) to contain the genera
simple, bifurcate and trifurcate clads. Microscleres, if present, are Trachygellius Topsent and Spirasigma Hallmann, is characterized
amphiaster streptasters. by the presence of sigmaspires and oxeas arranged radially (such as
in Tetillidae). In contrast, in Thrombidae the microscleres are
Diagnosis amphiasters, the 'megascleres' are small acanthotriaenes, and oxeas
are absent. Consequently, this proposed synonymy is rejected here.
Irregular, massive or encrusting, creeping-stoloniferous Members of Thrombidae are found in the Atlantic,
sponges, with diplodal aquiferous system and cortex barely dis- Mediterranean and Pacific. Most species inhabit bathyal depths to
cernible. Without monaxonid spicules. The megascleres are small 1378 m but some have also been found in shallow waters, in caves
spiny triaenes of different types, with simple, bifurcate or trifurcate of the Mediterranean Sea, and more recently in the Caribbean
clads (i.e., plagio-, dicho-, and trichotriaenes), one or more may (Lehnert, 1998).
be absent. Microscleres absent or amphiasters. The triaenes are
arranged in the ectosomal region with the clads tangential to the Taxonomic remarks
surface, and the rhabdome directed inwards. Similar spicules
are disarranged within the choanosome. The family's sole genus contains five nominal species:
T. abyssi (Carter), T. challengeri Sollas, T. kittoni (Carter), T. orna-
Scope tus Sollas and T. jancai Lehnert, although some of these have been
considered to be dubious by earlier authors. However, differences
Monogeneric (Thrombus). in sizes, morphology, and types of spicules between these species
probably correspond more to species-specific differences than to
History and biology individual variability.

This family was erected by Sollas (1888) to include only Previous reviews
Thrombus, which still has obscure phylogenetic relationships. It
was initially allocated to the suborder Microsclerophora (termed Sollas, 1888; Topsent, 1895; Lendenfeld, 1903.
Homosclerophora by Dendy, 1905) within the order Choristida
Sollas. Topsent (1895) subsequently allocated the genus to the sub-
order Microtriaenosa within the order Camosa Carter, together THROMBUS SOLLAS, 1886
with the genera Triptolemus Sollas, Rhachella Sollas, Samus Gray,
Dercitus Gray and Corticella Sollas, thus leaving the family Synonymy
Thrombidae empty.
Lendenfeld (1903) placed Thrombus in the Family Plakinidae Thrombus Sollas, 1886.
Schulze together with Plakina Schulze, Plakinastrella Schulze,
Plakortis Schulze and Corticium Schmidt. De Laubenfels (1936a) Type species
then transferred it to Corticiinae Vosmaer even though this subfam-
ily was defined by the presence of euasters, and thus differentiated Corticium abyssi (Carter, 1873b) (here designated).

163
164 Porifera· Demospongiae • Astrophorida • Thrombidae

Definition Previous reviews

Thrombidae with spiny small triaenes (acanthose Sollas, 1888; Topsent, 189S; Van Soest & Stentoft, 1988;
pseudotriaenes). Boury-Esnault et al., 1994b.

Diagnosis Description of type species

Massive, thickly encrusting, irregular or branching sponges Thrombus abyssi (Carter, 1873b) (Figs 1 and 2).
with non-hispid but rough surface. The main spicules are small Synonymy. Corticium abyssi Carter 1873b: 18.
acanthotriaenes (plagiotriaenes, dichotriaenes, or trichotriaenes), Material examined. Holotype: not seen - North Atlantic,
sometimes accompanied by amphiasters. English Channel, 900 m depth. Other material. MNHN CPS-80 -
North Atlantic.
Description. Thickly encrusting, irregular sponge, which
incorporates shells and other foreign debris of organic origin.
Smooth (non-hispid) but rough surface. Consistency hard. Cortex
not differentiated. Cream color in alcohol. The main spicules are
spiny dichotriaenes and trichotriaenes with the cladome 4S-SS f.Lm
in cord length, protoclads 8-12 f.Lm long and deuteroclads 10001S
f.Lm long ending in a round point; the rhabdome is up to 60 f.Lm long
with a more-or-Iess sharp point and the spines directed toward the
cladome. Microscleres are amphiasters, S f.Lm in length. Skeletal
arrangement. The trichotriaenes form a layer at the superficial zone
with the cladome tangential to the sponge surface and the rhab-
dome inwards. The same spicule type is placed in disarray within
the choanosome. The amphiasters are scattered thorough the
sponge.
Remarks. The genus Thrombus has no clear relationships
with any other sponge genus known so far, and thus Family
Thrombidae remains monotypic. The small size of the acanthotri-
aenes of Thrombus, closer to that of microscleres than to megascle-
res, precludes any comparison with other Astrophorida genera
bearing normal (megasclere) triaenes with some spines, such as
Fig. 1. Thrombus abyssi (Carter, 1873b), fragment of specimen MNHN Acanthotriaena crypta Vacelet et al., 1976, or other fossil genera
CPS-80. such as Acanthastrella Schrammen, 1924a. The small size of the
acanthotriaenes of Thrombus, and the presence of amphiasters are
somewhat reminiscent of spicules in some species of Corallistidae
(with tuberculated dichotriaenes (occasionally also trichotriaenes),
such as Corallistes tubulatus (Van Soest & Stentfort, 1988), and
even those of Callipelta (Macandrewia) cavemicola Vacelet &
Vasseur (1964) from Madagascar (with phyllotriaenes and tubercu-
lated dichotriaenes, 30--S0 X 9 f.Lm in size, and amphiasters 9-12
f.Lm long). Despite these vague similarities the phyllogenetic affini-
ties of this family remain obscure. The absence of true megascleres
induced early authors to consider this genus within the
Homoscleromorpha. Lendenfeld (1903) placed it in Plakinidae,
,• I Ilm whereas de Laubenfels (1936a) placed it together with Corticium,
Placinolopha, etc. in the family Corticiidae (subfamily Corticiinae).

Distribution

Atlantic and Indo-Pacific oceans.

ACKNOWLEDGEMENTS

I thank G. Carreras and G. Agell from the CEAB for their help
in drawings, scanning and treatment of figures, respectively.
C. Levi (MNHM) kindly loaned Museum material.

Fig.2. Thrombus abyssi (Carter, 1873b), specimen MNHN CPS-80, lateral

and frontal views of two trichotriaenes.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademicIPlenum Publishers, New York, 2002

Astrophorida Incertae Sedis

John N.A. Hooper l & Manuel Maldonado2

I Queensland Museum, P.O. Box 3300, South Brisbane, Qld, 4101, Australia. (JohnH@qm.qld.gov.au)
2 Centro de Estudios Avanzados de Elanes (CSIC), Camino de Santa Barbara sin Blanes 17300, Girona, Spain. (maldonado@ceab.csic.es)

Lamellomorpha Bergquist (Demospongiae, ? Astrophorida) is a monotypic genus from cool temperate and subantarctic New Zealand. It
has contort oxeas, strongyles and strongyloxeas as megascleres forming a lax confused choanosomal skeleton with only slight traces of
radial structure near the surface, and a skin-like membranous ectosomal skeleton packed with microstrongyles, and with streptasters
(amphiaster- , metaster- and spiraster-like) scattered throughout. Its precise taxonomic allocation cannot be presently resolved, with dis-
cussion provided on possible affinities with Astrophorida (e.g., Ancorinidae, Pachastrellidae), some 'lithistid' demosponges (e.g.,
Corallistidae, Isoraphinidae, Phymaraphiniidae), Hadromerida (e.g., Alectonidae) and Halichondrida (e.g., Axinellidae, Bubaridae), and
is left incertae sedis within Astrophorida.
Keywords: Porifera; Demospongiae; Astrophorida; incertae sedis; Lamellomorpha.

LAMELLOMORPHA BERGQUIST, 1968 172°30'E, 6O-120m depth, colI. MV 'Tui', 22 X .58, NZOI
stn. B93.
Synonymy Description. Massive thickly folded lamellate growth fonn,
13-16 cm high, with incurved lamellae 2-6 cm thick, supported by
Lamellomorpha Bergquist, 1968: 30. stout stalk about 3 cm in diameter. Texture finn, compressible, eas-
ily broken. Surface smooth, oscules scattered over the convex sur-
Type species face of lamellae, flush with the surface. Ectosomal skeleton with
a distinct skin-like dennal membrane densely packed with
Lamellomorpha strongylata Bergquist, 1968 (by monotypy). microstrongyles. Choanosomal skeleton with only slight traces of
radial construction, where in the subectosomal region of both sur-
Definition faces of lamellae tracts of megascleres, of variable thickness, curve
outwards and intersect the surface at acute angles. These tracts are
? Astrophorida with only contort monaxons (oxeas, strongyles absent from deeper regions of the choanosomal skeleton which is
and strongyloxeas) as megascleres, and microscleres being strept- lax and confused. Microstrongyles and streptasters scattered
asters and microstrongyles. throughout the choanosomal skeleton. Megascleres are basically
oxeote with strongylote and strongyloxeote modifications,
Diagnosis predominantly contort/sinuous, sometimes merely curved,
1000-(1980)-2808/-Lm long, 14-(26)-33/-Lm wide. Microscleres
Massive folded lamellate sponge with smooth surface. are squat microstrongyles, evenly rounded, slightly roughened,
Ectosomal skeleton a skin-like membrane packed with micro- occasionally centrotylote (23-(24.8)-30,...,m long, 2.3-(2.9)-4.3
strongyles. Choanosomal skeletal architecture lax and confused /-Lm wide), and small streptasters with 3-12 smooth, sharply
with only slight traces of radial structure, with subectosomal tracts pointed rays and mostly a straight shaft (amphiaster- transitional
of contorted oxeas or modified fonns (strongyles, strongyloxeas) - to metaster- and spiraster-like, but not plesiasters as stated by
curving outwards and supporting the ectosomal skeleton. Micro- Bergquist, 1968) (8.0-(9.8)-11.0 /-Lm long).
strongyles and streptasters (amphiaster-, metaster- and spiraster- Remarks. The original description of this species is rela-
like) scattered throughout the body. tively comprehensive and well-illustrated, yet the affinities of the
genus are still highly problematic.
Remarks Bergquist (1968) made comparisons between Lamellomorpha
and Jaspis (Coppatiidae, now Ancorinidae) based on skeletal struc-
The genus contains one or possibly two species (see remarks ture and megasclere geometry, in particular J. serpentina Wilson,
below), and is known only from southern temperate and subantarctic but differing significantly in their microsclere composition, with
waters (Three Kings Island, Campbell Plateau, and South Africa). putative differences being the presence of streptasters in the for-
mer, tenned plesiasters by Bergquist but clearly more similar to
Description of type species amphiasters-metasters (with a straight shaft) and spirasters (with
a curved shaft), and the presence of euasters in the latter. Bergquist
Lamellomorpha strongylata Bergquist (Fig. 1). (1968) also suggested that Coppatias baculifer Kirkpatrick from
Synonymy. Lamellomorpha strongylata Bergquist, 1968: South Africa may also belong to Lamellomorpha, being similar to
31-32, pI. 4a, 1Ie-f, fig. 10. L. strongylata in most respects, also having microstrongyles, but
Material examined. None. Holotype: NMNZ Por.33 lacking streptasters. She hypothesised further that microstrongyles
(not seen): South of Three Kings Island, New Zealand, 34°00'S, of C. baculifer were derived from streptasters through loss of rays,

165
166 Porifera· Demospongiae • Astrophorida • Incertae Sedis

c~
~_::-;:'::;.:-.:':::;;:'y
ct£:£:.:: '-':""<" ::~" ." :'.':~

G/ :-<:'. :':~

.
~ ~~
krk~~~h
Fig. 1. Lamellomorpha strongylata Bergquist. A, holotype (size 13 cm high). B, light micrograph of contort oxeas, strongyles and strongyloxeas. C,
microrhabds. D, streptasters (refer to text for sizes) (all modified from Bergquist, 1968).

and L. strongylata presented an intermediate condition between that pachastrellids - and astrophorids in general- are unlikely to be
Jaspis and C. baculifer (having both sorts of microscleres). Indeed bright green. Astrophorids are usually whiteish, with only some
microstrongyles in L. strongylata do appear to be derived from shallow-water species (which is not the case of Lamellomorpha,
streptasters, through loss of rays (Bergquist, 1968: pI. 11, collected from 60 to 120m depth) showing melanine-derived
Fig. F2), but it is unlikely that these are related to euasters, and the blackish or grayish colours for UV protection. Green colouration in
only obvious homology between Lamellomorpha and Coppatiidae sponges usually results from either subectosomal development of
(=Ancorinidae) is the lack (loss) oftriaene megascleres. It is there- photoautotrophic symbionts or translocation of undegraded phyto-
fore unlikely that these species belong to Ancorinidae as they lack plankton pigments, likely for UV protection. Both possibilities are
euasters (a relatively strong synapomorphy for the family, although unlikely in sponges with a thick cortex of microscleres or tangen-
some ancorinid genera do lack them; see chapter by Uriz, this tial felts, which diffract incident light. Thus, despite similarities in
volume). several features between Lamellomorpha and some Pachastrellidae
A second alternative, and possibly more plausible allocation, it is unsure whether these are homologous features or analogous
is with the Pachastrellidae, but this hypothesis is also speculative convergences, and the genus is not included here either.
given the absence of triaene megascleres or their derivatives with A third alternative is that Lamellomorpha may be a 'lithistid'
which to judge affinities (see chapter by Maldonado, this volume). sponge that has lost its desmas. There are several 'lithistids' that
Ignoring for the moment the absence of triaenes (andlor calthrops) have contorted strongyles and streptasters, in addition to monocre-
in Lamellomorpha (which could be hypothesised as a secondary pidial or tetracrepidial desmas (such as Corallistidae and
loss), the genus shows some superficial similarities to Pachastrella Isoraphinidae, and Phymaraphiniidae, respectively; refer to chap-
which also has an ectosomal skeleton composed of a felt of ters by Pisera & Levi, this volume). Maldonado et al. (1999)
microstrongyles, and amphiaster- to metaster-like streptasters. demonstrated that production of desmas may be inhibited in
However, the shape of oxeas and strongyloxeas in Lamellomorpha shallow-water 'lithistids' because the levels of dissolved silicon in
is not the same as those seen in various genera of Pachastrellidae. shallow waters are lower than those required for secretion of des-
Microstrongyles in the former are not necessarily homologous to mas and other hypersilicified spiCUles. This might also be the case
the oval microstrongyles of Pachastrella, for example (measuring for Lamellomorpha (with a known depth range of 60-120m). To
8.5-15 f.Lm X 4--6 f.Lm), but are longer and thinner (23-30 f.Lm X test this idea would require searching for a significant resemblance
2.3-4.3 f.Lm). Bergquist (1968) described for Lamellomorpha "In between the free (non-lithistid) skeletons of Lamellomorpha and
the subdermal region, on each surface of the sponge, tracts of various 'lithistids'. However, given that the concept of 'order
megascleres curve outward and intersect the surface at an acute Lithistida' is now formally abandoned (Pisera & Levi, this vol-
angle". This feature is so far unknown in typical pachastrellids. ume), such a relationship would need to be postulated at the family
Furthermore, she adds "A distinct skin-like membrane is present or genus level to have any taxonomic meaning. While a number of
and is densely packed with microstrongyles", whereas a skin-like 'lithistids' have obvious affinities with various families in the
membrane is so far unknown from typical pachastrellids, and orders Poecilosclerida, Halichondrida and Spirophorida, most are
indeed none have an ectosomal felt that can be easily detached. not yet resolved but many are astrophorids and it is plausible that
Lamellomorpha is described as bright green in life, and although Lamellomorpha belongs to one of these.
sponge colouration often may be taxonomically unimportant, there Finally, it is possible that Lamellomorpha may not belong
are some physiological pathways and ecological concerns about to Ancorinidae at all, but may have affinities to Hadromerida
colour in sponges (see Maldonado & Young, 1996c) suggesting or Halichondrida. For example, some genera of Alectonidae
Porifera' Demospongiae • Astrophorida • lncertae Sedis 167

(Hadromerida) have oxeas and strongyles (and they are known to strongyles vaguely reminiscent of those found in Lamellomorpha
lose their megascleres), and streptasters (amphiasters) and strongy- (see chapters by Alvarez et al., this volume), with the supporting
lote microscleres. In addition, the latter ones usually show thick, feature being possession of a stalked growth form (common to
tubercule-like microspines similar to those illustrated by Bergquist Axinellidae), and a lax choanosomal skeleton (characteristic of
(1968: pI. 11, Fig. F2) in some spicules of Lamellomorpha, which many halichondrids). Nevertheless, this evidence is very weak
she called "abnormal microrhabds". Nevertheless, alectonid and highly speculative. All these above hypotheses require further
strongylote microscleres are usually bent, spiral and/or with spi- testing through comparative morphological studies based on
rally arranged microspines (see chapter by Riitzler, this volume). type material (including a redescription of the holotype of
Similarly, some species ofAxinellidae and Bubaridae Lamellomorpha), and until such a study is completed this genus
(Halichondrida) have contorted oxeas, strongyloxeas and remains incertae sedis within Astrophorida.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© KIuwer AcademiclPlenum Publishers, New York, 2002

Order Hadromerida Topsent, 1894

John N.A. Hooperi & Rob W.M. Van Soest2

Hadromerida Topsent (Demospongiae), including Clavulina Vosmaer and Astromonaxonellida Dendy, contains 23 nominal families or
subfamilies of which 13 families are recognised here (two allocated as incertae sedis). The order includes sponges with monaxonic
megascleres (tylostyles, subtylostyles, oxeas or derivatives) producing radiate or subradiate skeletal arrangement, although this arrange-
ment may be obvious only in the peripheral skeleton in some, often with scarce spongin and consequently firm, non-elastic consistency.
Ectosomal spicules are often smaller than choanosomal ones, and where present these may form a prominent cortical skeleton.
Microscleres may include various forms of euasters, spirasters, rhabds, microxeas and/or raphides, although many hadromerids lack
microscleres altogether. Hadromerid sponges are known from all oceans at all depths.
Keywords: Porifera; Demospongiae; Hadromerida; Acanthochaetetidae; Alectonidae; Clionaidae; Hemiasterellidae; Placospongiidae;
Polymastiidae; Sollasellidae; Spirastrellidae; Stylocordylidae; Suberitidae; Tethyidae; Timeidae; Trachycladidae.

DEFINITION, DIAGNOSIS, SCOPE elastic. Bright colours are common (yellows, oranges, reds) but
generally not diagnostic, and in some cases (e.g., encrusting
Synonymy forms), colouration may be produced by symbionts. Where known
choanocytes have a periflagellar sleeve. Microscleres may con-
[suborder] Hadromerina Topsent, 1894. [order] Hadromerina sist of euasters, sterrasters, streptasters and derivatives, spherasters,
Topsent, 1928c. Hadromerida de Laubenfels, 1936a. Clavulina micrasters, anthasters, lophasters, pseudasters (amphiasters), spi-
Vosmaer, 1885b (part). Astromonaxonellida Dendy, 1905. rasters, spiraster-like spirules, microrhabds, microxeas or
microstrongyles, but are absent in several families. Most groups are
Definition oviparous (where known), with eggs extruded and development of
parenchymella larvae (in one case a blastula larva) directly in
Demospongiae with monaxonic megascleres (tylostyles, seawater. Budding is common in Tethyidae.
subtylostyles, oxeas or derivatives) forming radiate or subradiate
skeletal arrangement, sometimes only obvious in the peripheral Scope
skeleton; ectosomal spicules usually smaller than choanosomal
ones, and where present they may produce a cortical skeleton; Twenty three family or subfamily taxa have been included in
spongin often sparse producing firm non-elastic consistency; Hadromerida at one time or another, although only 13 families are
microscleres may include various forms of euasters, spirasters, recognised in this work (two incertae sedis): Acanthochaetetidae
rhabds, microxeas and/or raphides in trichodragmata, or absent in Fischer, 1970; Alectonidae Rosell, 1996b; Clionaidae d'Orbigny,
many taxa. 1851; Hemiasterellidae Lendenfeld, 1889b; Placospongiidae
Gray, 1867a; Polymastiidae Gray, 1867a; Sollasellidae Lendenfeld,
Diagnosis 1887a (incertae sedis); Spirastrellidae Ridley & Dendy, 1886;
Stylocordylidae Topsent, 1928c; Suberitidae Schmidt, 1870;
Sponges with uniform spiculation of monaxonic megascleres Tethyidae Gray, 1848; Timeidae Topsent, 1928c; Trachycladidae
(monactinal or diactinal, most often tylostyles or subtylostyles, but Hallmann, 1917c (incertae sedis). Other nominal families are syn-
also including exotyles, strongyloxeas, oxeas or modified forms of onymised as follows: [Astraxinellidae] Dendy, 1905 and
these). Sponges are frequently massive with megascleres radially Leptosastrinae Topsent, 1928c are synonyms of Hemiasterellidae;
arranged throughout the entire skeleton (e.g., Tethyidae), or with [Spirophorellinae] Lendenfeld, 1889b (nomen nudum) and
this arrangement only obvious at surface (e.g., Polymastiidae), but Rhaphidistiinae de Laubenfels, 1936a are synonyms of
typical growth forms also include excavating forms (Clionaidae), Trachycladidae; Thoosidae Rosell & Uriz, 1997 is a synonym
thinly encrusting (e.g., Timeidae), spherical (Tethyidae), stipitate of Alectonidae; Choanitidae de Laubenfels, 1936a is a synonym of
(Tethyidae, Polymastiidae, Stylocordylidae, Suberitidae), vasiform Spirastrellidae; Tabulospongiidae Mori, 1976 is a synonym of
and arborescent branching forms (e.g., Hemiasterellidae, Acanthochaetetidae; Donatiadae Carter, 1875c and Xenospongiidae
Trachycladiidae). Megascleres usually consist of more than one Carter, 1882b are synonyms of Tethyidae. Chondrosiidae Schulze,
size class, with ectosomal spicules often smaller than choanosomal 1877 (emend. Chondrosiidae Wiedenmayer, 1977; =Chondrillidae
spicules, usually standing perpendicular to the surface, protruding Gray, 1872a) is allocated to its own order.
through the ectosome and forming a hispid palisade or cortical
skeleton. Spongin fibres are generally poorly developed (if at all Remarks
present), frequently producing a firm, non-elastic, friable consis-
tency, although in several genera (e.g., Cauiospongia, Suberites) Topsent (1898a) initially included nine families into his sub-
spongin is abundant and these taxa are more compressible and order Hadromerina, and further proposed to subdivide the taxon

169
170 Porifera· Demospongiae • Hadromerida

into two Sections, Aciculida (as Aciculides) with diactinal megas- (including Polymastiidae); Placospongiidae; Clionidae (now
cleres (containing Coppatiidae, Strepasteridae, Tethyidae, Clionaidae); and Gastrophanellidae (for a small group of 'lithis-
Stylocordylidae) and Clavulida (as Clavulides) with monactinal tids' with tylostyles or exotyles). Other taxa that we now include in
megascleres (containing Clionaidae, Spirastrellidae, Polymastiidae, Hadromerida were mostly assigned to his order Epipolasida, such as
Suberitidae, Mesapidae). In his 1904 monograph Topsent also Tethyidae, Stylocordylidae (which he included in Podospongiidae),
included some genera now allocated to Desmoxyidae (e.g., and Trachycladidae.
Heteroxya) in his group Aciculides. Hentschel (1923) and Wilson Major contemporary syntheses of the sponge classification (e.g.,
(1925) adopted Topsent's scheme, and added the families Levi, 1973; Bergquist, 1978; Hartman, 1982; and others) agree on a
Chondrosiidae and some other genera of [Astraxinellidae] core component of Hadromerida (containing eight families:
(=Hemiasterellidae). Topsent (1928c: 33) subsequently elevated Tethyidae, Polymastiidae, Suberitidae, Spirastrellidae, Clionidae
Hadromerina to ordinal status, and also abandoned his earlier (=Clionaidae), Placospongiidae, Timeidae and Stylocordylidae).
subdivision of the order given that Tethyidae had monactinal Conversely, there are several other families disputed by contempo-
megacleres (but otherwise conformed with Aciculida, which was rary authors as having Hadromerida affinities. Some of these have
characterized by diactinal megascleres), and that Chondrosiidae been recently settled based on additional non-morphological evi-
'suppressed' its megascleres making its allocation to one or the dence. Chondrosiidae (= Chondrillidae) was elevated to full ordinal
other group equivocal. In this revision Topsent (1928c) included status (Boury-Esnault & Lopes, 1985), and is now allocated to its
Spongosoritidae (now in Halichondrida), Coppatiidae (now in own order Chondrosida. Latrunculiidae was found to be poly-
Astrophorida) and Chondrosiidae (now in its own order phyletic, now subdivided into two families, both allocated to the
Chondrosida), together with seven 'typical' families (Clionidae Poecilosclerida (see chapters by Kelly & Samaai and Samaai &
[now Clionaidae], Polymastiidae, Spirastrellidae, Stylocordylidae, Kelly, this work). Acanthochaetetidae are hypercalcified sponges
Suberitidae, Tethyidae, Timeidae). He admitted (p. 34) that ('sclerosponge' grade of construction) which show clear morpho-
Chondrosiidae, in particular, was not a straightforward allocation logical (spicular) affinities to the Spirastrellidae (e.g., Van Soest,
to Hadromerida, and he also reallocated [Astraxinellidae] 1984a; Wood, 1990), with additional recent molecular support from
( = Hemiasterellidae) to the Halichondrida, although again with 28S rRNA sequences to support this hypothesis (Chombard et al.,
some reservations (pp. 38-39) about the validity and affinities of 1997). Sollasellidae is incertae sedis within Hadromerida, showing
this family. (perhaps) superficial similarities to some 'true' hadromerids. It is
Some 19th and 20th century authors did not follow Topsent's similar to Polymastiidae in having a cortical skeleton, especially to
(1898a, 1904, 1928c) system (e.g., Dendy, 1922b; Burton, 1932b, Pseudotrachya with which it shares the combination of choanoso-
1959a), preferring instead the earlier classifications, such as those mal styles and ectosomal oxeas. Similarly, it shows similarities with
proposed by Ridley & Dendy (1887) and Sollas (1888), in which some Suberitidae, such as Homaxinella, and Stylocordylidae, shar-
hadromerids were often assigned to a suborder Clavulina Vosmaer, ing a stalked habit and axially condensed skeleton (see chapter by
1885b, or sometimes to a family Clavulidae. In this latter scheme, Van Soest, this volume). However, Sollasellidae remains poorly
as modified by Ridley & Dendy (1887) two families were recog- known and these affinities are relatively speculative.
nised: Suberitidae (which included genera now assigned to The allocation of Hemiasterellidae and Trachycladidae to
Stylocordylidae, Polymastiidae, Clionaidae) and Spirastrellidae Hadromerida is more controversial, and to date lacks any definitive
(which also included Latrunculiidae). Ridley & Dendy (1887) also molecular or other non-morphological support. These families are
remarked on the probable relationship between these taxa and remnants of the now dismantled order Axinellida, with both
the Halichondrida, notably the common possession of radially assigned to Hadromerida based on megasclere and microsclere
arranged tylostyles; cork-like granular 'ground substance'; pres- morphologies and skeletal structure, whereas other families once
ence of a distinct fibrous cortex; and absence of spongin in the assigned to the axinellids are allocated to Poecilosclerida and
skeleton. Halichondrida (see respective chapters in this volume). Assigning
De Laubenfels (1936a) was probably the first to formally Hemiasterellidae to Hadromerida is fairly straightforward using
adopt Topsent's scheme, and was followed by most contemporary morphological criteria, with the common possession of euasters,
authors working on hadromerids (e.g., Bergquist, 1968 et seq.). De monactinal megascleres and radial skeletal architecture, at least
Laubenfels' (1936a) treatment of the order is a useful precis of at the surface (Voultsiadou-Koukoura & Van Soest, 1991a).
hadromerid apomorphies, noting that the possession of a radiate or Chemotaxonomic support for this hypothesis (Hooper et al., 1992),
subradiate architecture, the total, or almost total, lack of spongin, however, is equivocal. Hemiasterellids differ from other
and a pronounced cortical specialization in the ectosome were hadromerids, such as Polymastiidae and Spirastrellidae, in having
indicative of affinities with the 'tetraxon sponges' (now assigned axially compressed choanosomal skeletons (sometimes loosely
to the orders Astrophorida, Spirophorida and Chondrosida, and constructed or hymedesmoid, e.g., Paratimea) and an ectosomal
including many 'Lithistida'), and he suggested that they differ from palisade which appears to be homologous to the 'typical'
these 'tetraxon' orders principally in having tylostyles as their chief hadromerid cortical skeleton. Hemiasterellids also show similari-
megascleres. Conversely, his speculation that this latter feature, ties to Aaptos (Suberitidae) and Tethya (Tethyidae) in having pre-
together with the possession of spongin in some taxa, indicated that dominantly style megascleres, rather than tylostyles found in many
they may be more closely related to Poecilosclerida, is now dis- hadromerid taxa, although some hemiasterellids retain long oxeas
counted and supported by non-morphological evidence (e.g., com- in the extra-axial skeleton (see chapter by Hooper, this volume).
mon possession of oviparous reproductive strategy), and to a large Conversely, the affinities of Trachycladidae remain unclear.
degree validates a 'tetraxon sponge' clade (e.g., Bergquist, 1978). There are no reproductive, molecular or chemical data to support
De Laubenfels (1936a) recognised only five families, differentiated any morphological hypotheses. Br~ndsted (1924b) suggested a
primarily by their microsclere composition: Spirastrellidae (as close relationship between Trachycladus and Spirastrellidae, given
Choanitidae), which included Timeidae (as a subfamily); Suberitidae fundamental similarities in their microsclere geometries, although
Porifera· Demospongiae • Hadromerida 171

with equally fundamental differences in patterns of spination of these 1999); (5) the phylogenetic significance of an excavating habit
microscleres. Peripheral skeletal architecture of trachycladids is plumo- amongst hadromerids (Rossell & Uriz, 1997); (6) an analysis of
reticulate; the axial skeleton is compressed (both composed of oxeas or the phylogenetic significance of skeletal architecture (including
modified oxeote megascleres); and the ectosomal skeleton is composed presence or absence of axial compression), megasclere geometry
of a cortical skeleton of microscleres (spined vermiform spinispirae), (presence, absence and secondary loss of triaenes), microsclere
together with smooth microrhabds scattered throughout the skeleton geometry (including euaster geometries and their derivatives),
(see chapter by Hooper and Van Soest this volume). These characteris- reproductive strategy (ovipary versus vivipary), and other fea-
tic spinispirae resemble spirasters, common to some hadromerids, but tures used to assess the relationships of Hadromerida within
the homology of these two forms of spicules remains to be determined. a tetractinellid clade (amongst others) (Van Soest, 1991); (7) an
Trachycladidae is incerlae sedis within Hadromerida. evaluation of chemistry used as chemotaxonomic markers (Van
Several more recent publications provide new insights on the Soest & Braekman, 1999) - who evaluated the approximately 185
affinities of Hadromerida, although none of these treatments secondary metabolites described from Hadromerida and concluded
include all the families assigned here, with most concerning only that there were no distinct compounds that could be used reliably
the 'typical' families ofhadromerids. These works include: (1) new as taxonomic markers for the order, although several appeared
alkaloids as taxonomic markers for the order (Bergquist et at., to be useful for family-level taxa.
1991a); (2) structure of the aquiferous system (Bavestrello et al.,
1998b); (3) 28S rRNA sequence data (Chombard et al., 1999) - Distribution
which hypothesised the inclusion of family Halichondriidae in the
Hadromerida (supporting Topsent's (1928c) earlier proposal to Recorded from all oceans at all depths.
include Spongosoritidae in this order), and proposed further to
recognise four suborders: Spirastrellina, Timeina, Polymastiina Previous reviews
and Suberitina; (4) 18S rRNA sequence data (Kelly-Borges et al.,
1991), which apparently were insufficiently informative to resolve Levi, 1973; Bergquist, 1978; Hartman, 1982; Chombard et al.,
the phylogeny at the higher taxonomic level (Chombard et al., 1999.

KEY TO FAMILIES

(1) Microscleres euasters ................... ................ ........... ........... .......... .... ....... ........... ........... ................... ............. ................................. ...... 2
Microscleres sterrasters ..... ........... .......... ...... .... ....... .... ........... ........... ........... ...... .... ........ ... .... ....... ........... ...... ............... ....................... 3
Microscleres streptasters and/or microstrongyles or microrhabds ...................................................................................................... 4
Microscleres spined 'corkscrew shaped' spinispirae ........................................................................................................................... 6
Asterose microscleres absent ... .... ................ ........... ........... ...................... .......... ........... ........... ........... .......... ................................ ....... 7
(2) Megascleres styles, oxeas or both, forming compressed axial (or basal) skeletons and plumo-reticulate or radial peripheral skeletons;
growth forms commonly arborescent, massive and vasiform, occasionally encrusting; microscleres euasters with smooth and/or
partially microspined rays ......................................................................................................................................... Hemiasterellidae
Megascleres stylote, mainly strongyloxeas (but sometimes subtylostyles, tylostyles or anisostrongyles) forming radial bundles or
sometimes parallel tracts, and frequently with a well-developed ectosomal cortical skeleton; growth forms commonly spherical, glob-
ular or massive; microscleres often include two categories of euasters (micrasters and megasters), sometimes also with microrhabds,
microstrongyles, exotyles and spheres .................................................................................................................................. Tethyidae
Megascleres single or tracts of tylostyles running to the surface; growth forms predominantly encrusting; microscleres are euasters
(including anthasters and lophasters) or pseudasters (amphiasters) forming a densely packed cortical skeleton ................. Timeidae
(3) Microscleres sterrasters (selenasters or amphiaster-like (amphinolasters» forming a strong cortical skeleton of polygonal plates, and
with accessory microscleres being spirasters, spherasters and spherules; megascleres are tylostyles in tracts radiating from the base
toward the surface and supporting the margins of the cortical plates ..... ........... ...................... ........... ................ ....... Placospongiidae
(4) Megascleres tylostyles ......................................................................................................................................................................... 5
Megascleres styles, oxeas or strongy1es (occasionally absent completely), being smooth or spiny and kinked or sometimes branched;
microscleres amphiasters (streptasters), usually microspined with spines arranged uniformly, clustered, or spiraled, and/or
microrhabds (straight, bent, or spiral); growth forms mostly excavating, forming chambers .......................................... Alectonidae
(5) Megascleres sparse tylostyles forming ascending radial tracts and protruding through the surface; microscleres streptasters (spirasters,
diplasters, spiny microstrongyles) forming a dense cortical skeleton and also a basal crust; early stages may be excavating but adults
are only encrusting or submassive, not endolithic, with the aquiferous system confined to the external surface and consisting of mean-
dering vein-like structures converging on oscula ......................................................................................................... Spirastrellidae
Megascleres tylostyles, styles or oxeas; skeletal arrangement varies depending on stage of development, confused in excavating
(alpha) growth forms, or confused and with ill-defined tracts in encrusting (beta) and massive (gamma) forms; ectosomal skeleton
a cortical skeleton of megascleres in a palisade; microscleres streptasters and/or microrhabds of different types ............ Clionaidae
Megascleres erect tylostyles and microscleres relatively large streptasters both reinforcing a thin veneer of soft tissue covering
a hypercalcified ('sclerosponge') basal skeleton ................................................................................................. Acanthochaetetidae
(6) Megascleres oxeas, strongyles and/or (tylo-)styles, forming a compressed axial region and a plumose or plumo-reticulate extra-axial
region; microscleres spined 'corkscrew shaped' spinispirae and smooth microrhabds .............................................. Trachycladidae
172 Porifera· Demospongiae • Hadromerida

(7) Cortical skeleton well-developed ....................................................................................................................................................... 8

No well-developed cortical skeleton .................................................................................................................................................. 9
(8) Ectosome with a more-or-less complicated cortical skeleton, always with an outer palisade of ectosomal spicules (tylostyles, or
oxeas and/or exotyles); megascleres tylostyles, subtylostyles, strongyloxeas, styles or oxeas forming a radiating choanosomal
skeleton; rnicroscleres absent apart from occasional centrotylote rnicroxeas, acanthose microxeas or raphides in
trichodragmata ..... .... ....... .... ....................... ................. .... ........... .............. ........ .......... ........... ...... .... .... ..... ....... ....... ... ... Polymastiidae
(9) Ectosomal skeleton with bouquets or a palisade of spicules ........................................................................................................... 10
Ectosomal skeleton composed of oxeas or stylotes ......................................................................................................................... 11
(10) No ectosomal cortical skeleton present but megascleres are arranged in bouquets or a palisade in the peripheral skeleton; megascleres
usually tylostyles, often with a pronounced or modified tyle, but occasionally styles, strongyloxeas or centrotylote oxeas; choanoso-
mal structure may be strictly radial, or show a strong axial orientation; more massive species become progressively more confusedly
arranged towards the interior; microscleres absent apart from occasional microrhabds or trichodragmas ..................... Suberitidae
(11) Megascleres centrotylote oxeas or strongyles forming radial tracts in the sponge 'body', also often with smaller oxeas or stylotes
at the surface, partly arranged in a tangential layer both on the main body and lining the stalk; stipitate habit, with a thin stalk and
globular-oval 'body' flattened at the top (confined to deep water) ........................................................................... Stylocordylidae
Megascleres long styles and short oxeas forming three distinct zones: a confused mass in the centre of the body; a subcortical region
with low spicular density traversed by bundles of a single style surrounded by a mass of oxeas bound by spongin; and an external
cortical skeleton strengthened by short oxeas; growth form is stalked, ramose with polygonal ornamentation of lines of inhalant
openings on the surface ............................................................................................................................................... '" Sollasellidae
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademicIPlenum Publishers, New York, 2002

Family Clionaidae D'Orbigny, 1851

Klaus Riitzier

Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, DC, 20560--0163, USA.
(ruetzler.klaus@nmnh.si.edu)

Clionaidae (Demospongiae, Hadromerida) receives a new definition by accepting Spheciospongia and massive species previously
assigned to Spirastrella (Spirastrellidae). A recently described genus, Cervicomia (for Alcyonium cuspidiferum Lamarck) is included here
and distinguished by its specialized incurrent fistules and endopsarnrnic choanosomal pulp. Clionaopsis new name is suggested to replace
the preoccupied [Clionopsis] Thiele. Spheciospongia remains distinct for its pore sieves. All other genera are separated by their spicule
complements which include tylostyles (with some oxeote or stylote modifications or additions) as megascleres, and delicate (compared to
Spirastrellidae) spirasters, amphiasters, smooth and spiny microxeas, and bent or spiral, smooth or spiny microstrongyles and microrhabds
as microscleres. Eight genera are considered to be valid.
Keywords: Porifera; Demospongiae; Hadromerida; Clionaidae; Cervicomia; Cliona; Clionaopsis; Cliothosa; Pione; Spheciospongia;
Thoosa; Volzia.

DEFINITION, DIAGNOSIS, SCOPE that many members had spicule complements that were radically
different from that of the type genus Cliona. For instance, de
Synonymy Laubenfels (1936a: 154) diagnosed the Clionaidae as sponges
with spiculation of Spirastrella in the family Choanitidae
[Clionidae] d'Orbigny, 1851: 209 (preocc. by Clionidae (=Spirastrellidae), with tylostyles as megascleres and spirasters as
Rafinesque, 1815 (Gastropoda». [Clioniadae] Gray, 1867a: 524 microscleres, but set off from the latter "on the basis [that they] bore
(lapsus). Spirastrellidae sensu Ridley & Dendy, 1886: 490. holes in calcareous material and live in the resulting cavities". Yet in
Thoosidae Rosell & Uriz, 1997: 350. Clionaidae d'Orbigny, 1851 discussing the scope of the family, de Laubenfels (1936a: 155) lists
(emended by Bouchet and Rtitzler, 2002). genera such as Aka (for Acca Johnson), which has only oxeas and is
now widely considered a haplosclerid (cf. Aka and Siphonodictyon,
Definition Phloeodictyidae) (Rutzler & Stone, 1986). Today we know that
species of Aka (and Siphonodictyon) are potent destructors of reef
Excavating Hadromerida with tylostyles and a great variety of coral (Rutzler, 1971), but also that members of other sponge groups
microscleres; microscleres, including spirasters, amphiasters, microx- that are clearly not related to Cliona, for instance, the
eas, spiral microstrongyles, and derivatives may be lacking altogether. Poecilosclerida Paracomulum and Zyzzya (Rutzler & Stone, 1986;
Schonberg, 2000) are able to efficiently excavate limestone.
Diagnosis Boring into the calcareous substratum results in a three-
dimensional network of interconnected chambers or galleries
Hadromerida with limestone-excavating capability, having below the substrate surface. The substrate chambers are filled with
tylostyles as principal megascleres, in some genera accompanied by sponge 'tissue' and spicules without orientation. Papillae are often
oxeas or styloid modifications. Microscleres may be absent entirely fortified by contractile cells and specially arranged megascleres
or in some specimens or popUlations. If present, they include spi- (e.g., tylostyles in bouquets or in a palisade), but not by prominent
rasters, amphiasters, microxeas, microrhabds, or raphides; some layers of microscleres (as in the Spirastrellidae). In gamma-stage
spirasters display secondary branching of spines, microrhabds may sponges, some megascleres are organized in loose tracts radiating
be smooth or microspined, straight, bent, or spiral. toward the surface; microscleres are dispersed throughout or occur
near the surface or along major canals.
Scope Limestone excavation is accomplished by chemical etching
compounds that are secreted by pseudopodial processes of special
Eight genera are here assigned: Cervicomia, Cliona, etching cells and help liberate minute (up to about 60 f.1m) chips
Clionaopsis, Cliothosa, Pione, Spheciospongia, Thoosa and Volzia. (Rutzler & Rieger, 1973; Pomponi, 1980). These chips are expelled
Two (Cervicomia, Spheciospongia) were previously placed in the with the excurrent water, only 2-3% of the calcium carbonate is
Spirastrellidae. Several thickly encrusting and massive species for- removed in dissolved form. The remaining cavities or series of cham-
merly assigned to the genera Anthosigmella Topsent, 1918, and bers (galleries) have characteristically pitted walls and are occupied
Spirastrella sensu lato (Ridley & Dendy, 1886; 1987), fall to the clion- by the sponge. Communication with the water column is maintained
aid genera Cliona or Cliothosa (Rutzler, 1990; Rosell & Uriz, 1997). through ostia- and oscula-bearing papillae that protrude slightly from
the substrate surface but can fuse to form a continuos sponge crust
History and biology (beta stage). Systematists may get confused by non-boring sponge
species that live cryptically in abandoned clionaid cavities.
Historically, the family Clionaidae was applied to sponges Most clionaids have simple ostia and oscula for water
with capacity for excavating limestone substrata, despite the fact exchange but there are some modifications to accommodate special

173
174 Porifera· Demospongiae • Hadromerida • Clionaidae

living habits. Sponges in alpha-stage of excavating (Vosmaer, massive with age or after the original substrate piece has been out-
1932) develop separate ostial and oscular papillae which puncture grown (gamma stage). The original incurrent papillae may still
the substratum surface, the former bearing many ostia, the latter show as wart-like structures in gamma-stage sponges although the
only one osculum. Sponges in the genus Spheciospongia, which calcareous substrate is no longer visible.
excavate rock substrata initially and live on sandy bottoms with
high sedimentation rate, are distinguished by pore sieves. These Remarks
sieves, already apparent in the papillate alpha stage, may help
avoid clogging of the actual ostia which are located in the In this treatment, emphasis is put on spicular and structural
vestibules below. Cervicomia was introduced for sponges with an features and not the excavating capacity alone to assign genera.
uncommon morphology and life style, also an adaptation to living The presence of tylostyles is the governing character and not the
on sediment bottoms. Large, epibenthic incurrent fistulas serve a multitude of microscleres. For other genera that excavate lime-
pulp-like choanosome that is completely buried in sand or shell and stone, see Alectonidae (e.g., Alectona), Phloeodictyidae in the
sends root-like exhalant tubes deep into the substrate (Rutzler, Haplosclerida (AkaiSiphonodictyon), and Acarnidae in the
1997). Poecilosclerida (Paracomulum, Zyzzya).
Clionaid sponges typically live in their burrows and connect The distinction between Spirastrellidae, Clionaidae, and
to the outside world through the papillae. Papillae may fuse and Placospongiidae in this revision is further supported by 28Sr RNA-
extend to encrusting habit (beta stage). Some species become genetic analysis (Chombard, 1998).

KEY TO GENERA

(1) Sponges excavating chambers in limestone substrates, with ostial and oscular papillae remaining at the surface (alpha stage), or form-
ing continuous encrustations (beta stage), or masses (gamma stage) that grow beyond the original substrate piece; without specialized
aquiferous structures except, in some species, the original papillae ...... .......... .... .... ............... ........ ....... .... ........ .......... ....................... 2
Sponges massive, without endolithic chambering in the adults, with specialized aquiferous morphology ..... ............. ...... ................ 7
(2) Spicules include oxeas ......................................................................................................................................................................... 3
Spicules do not include oxeas .... .... .......................... .... ........................ ...... ............. ............. ....... ............ ............ .... ........ ........... .......... 5
(3) Oxeas are spiny microxeas, located in the chambers (criss-cross fashion), and are accompanied by straight or sinuous spiny
microrhabds ............. ...... .... .... .... .................... ........ .... ........ ...... ......................................... .... .............. .... .... .......... ................ ........ Pione
Oxeas are smooth and form a palisade in the papillae ....... ...... ............................................................ ....... ...... .... .............. .......... ....... 4
(4) Spiculation is restricted to tylostyles and oxeas, no microscleres ............................................................................................... Volzia
Spiculation includes spirasters in addition to tylostyles and oxeas .................................................................................... Clio1UlOpsis
(5) Microscleres are raphides or spirasters, or missing altogether; spirasters may include diplastrose modifications or have lost spines
entirely (sinuous microstrongyles) ............................................................................................................................................. Cliona
Microscleres are amphiasters or euasters ...... .... .... .... ............ ........ ... ................. ...... ...... ...... .... .... .......... .......... ...... .............. .... ...... ....... 6
(6) Microscleres are two types of amphiasters: stout, smooth, nodulose, and thin-rayed with terminal branching ................... Cliothosa
Microscleres are microspined amphiasters and smooth, thin-rayed oxyasters ........ ....... ...... ....................... ...... ...... .... .............. Thoosa
(7) Sponges irregularly massive, with cribiporal chones and simple but often multiple oscula ... .... ............ .................... Spheciospongia
Sponges form large, branched, antler-like inhalant fistules and amorphous endopsammic masses with
exhalant stolons ....................................................... .... .... ...... .............. ... ........... .... ...... ..................... .................................. Cervicornia

CERVICORNIA RUTZLER & HOOPER, 2000 pulp. Exhalant stolons end underground. No alpha or beta stages
are known. With large tylostyles arranged in dense tracts in the
Synonymy fistulas, without orientation in the choanosome. Small and rare
spirasters and amphiasters located in ectosome and canal linings.
Cervicomia Rutzler & Hooper, 2000: 342.
Previous review
Type species
Rutzler & Hooper, 2000.
Alcyonium cuspidiferum Lamarck (by original designation).
Description of type species
Definition
Cervicomia cuspidifera (Lamarck) (Figs 1,2).
Clionaidae with specialized, ectosomal incurrent fistulas, Synonymy. Alcyonium cuspidiferum Lamarck, 1815: 168.
abundant robust tylostyles, and minute uncommon spirasters and Spirastrella cuspidifera; Topsent, 1933: 41, fig. 4, pI. 2, fig. 4.
derivatives. Xestospongia tiemeyi (de Laubenfels); Wiedenmayer 1977B: 117,
fig. 130, pI. 15, fig. 3, pI. 16, fig. 1. (Not Prianos tiemey de
Diagnosis Laubenfels, 1953a = Spheciospongia vesparium). Spheciospongia
cuspidifera; Pulitzer-Finali, 1986: 94, figs 22, 23; Vicente et at.,
Clionaid sponges with large, smooth, inhalant fistulas protrud- 1991: 217, figs 3d, 4; Rutzler, 1997: 1392, figs 1-2. Cervicomia
ing from the substratum and buried (endopsammic) choanosomal cuspidifera; Rutzler & Hooper, 2000: 342.
Porifera· Demospongiae • Hadromerida • Clionaidae 175

';',

., ....
~';

.
.... .

• ".
.
•
(.' ..
Ii

A B
Fig. 1. Cervicomia cuspidifera, habitus and anatomy. A, schematic of habitus in situ (scale 5 em). B, tylostyles (scale 50 fLm).

Fig. 2. Cervicomia cuspidifera, morphology and microscleres. A,habit of two specimens from Puerto Rico, West Indies (scale 5 cm). B, SEM images of
microscleres (scale 5 11m). (From Vicente et aI., 1991.)

Material examined. Holotype: MNHN LBIM DT 652 - stalactites. Tylostyles, some modified to tylostrongyles, average
labeled "151. Alcyonium cuspidiferum Lmk"; specimens examined 330-400 X 8-11 /Lm. Spirasters are small (4--13/Lm long), many
by Rtitzler in Vicente et al. (1991 : 217). smaller ones (4--6/Lm) are modified amphiasters.
Description. The dry type at MNHN is a cluster of about 8 The type species was recently described and illustrated in
slender conical tubes, fused at the bottom, free but blind-ending detail (Vicente et al., 1991: 217f, figs 3c, d, 4; as Spheciospongia).
distally, 44 cm tall, 20-25 mm in diameter at about midway to the These large, hollow and blind-ending fistulas were considered by
top. The original author compared them to a bundle of inverted previous authors to be the entire sponge until in situ observations
176 Porifera· Demospongiae • Hadromerida • Clionaidae

and fine-structure study showed their true nature and function Previous reviews
(Riitzler, 1997). The latter work also showed that the choanosome
is burrowing and concealed in unconsolidated calcareous substrate Topsent (1900); Hartman (l958b); Rosell & Uriz (1997).
into which exhalant water is discharged.
Remarks. Misidentification of this sponge as Xestospongia Description of type species
(Wiedenmayer, 1977B) is explained by the appearance, in some
populations, of the tylostyles as strongyles (only few subterminal Cliona celata Grant, 1826a (Figs 3-7).
swelling appear here and there) and the rarity and small size of the Synonymy. Cliona celata Grant, 1826a: 78; Hymeniacidon
spirasters. Prianos tierney de Laubenfels, 1953a turns out to be a celata Bowerbank, 1866: 212; Vioa celata Schmidt, 1866a: 40 (see
similar modification of Spheciospongia vesparium (Vicente et al., also Topsent, 1900). Spongia sulfurea Desor, 1848. Papillina
1991). suberea Schmidt, 1862: 69.
Material examined. Ho10type (not seen): believed to be in
the Grant Museum of Zoology and Comparative Anatomy at the
CLIONA GRANT, 1826 University College, London, where R.E. Grant worked (Clare
Valentine, personal communication; this information could not be
Synonymy confirmed). Description is based on literature records deemed reli-
able and specimens at BMNH and USNM. Northeast Atlantic,
Ctiona Grant, 1826a. Vioa Nardo, 1833. Poterion Schlegel, Celtic Sea - BMNH 1910.1.1.411; Tenby, Wales. Mediterranean-
1858. Raphyrus Bowerbank, 1862b. [papi/tina] Schmidt, 1862 USNM uncatalogued slide: alpha-stage, Tunisia. USNM 31374:
(preocc. by Papillina Conrad, 1855, Mollusca). Raphiophora Gray, Croatia. East coast ofthe United States (alpha- and gamma-stages)
1867a. Taguilla Boweberbank, 1874: xi (nom. nov. for [Papillina] - USNM 5349: off Massachusetts. USNM 7283: Rhode Island.
Schmidt). Tapiliata Bowerbank, 1874: xi (nom. nov. for [papi/tina] USNM 7492: New York. USNM 22705: Virginia. USNM 23648:
Schmidt). Euryphylla Duchassaing & Michelotti, 1864. Euryphylle North Carolina. USNM 33391: South Carolina. USNM 31065:
Duchassaing & Michelotti, 1864. Hymeniacidon sensu Bowerbank, Georgia. USNM 39608: Florida Gulf Coast.
1866 [not Hymeniacidon Bowerbank, 1859]. /damon Gray, 1867a. Distribution. Considered 'worldwide' but that statement
Myle Gray, 1867a. Pronax Gray, 1867a. Saptine Gray, 1867a. would have to be confirmed with improved taxonomic tools, such
Oscutina Schmidt, 1868. Spirastrella sensu Ridley, 1884a (non as molecular techniques. Reliable records indicate occurrence on
Schmidt, 1868). Papillella Vosmaer, 1885b. Papillissa Lendenfeld, both sides of the North Atlantic Ocean, in the Mediterranean, and
1888. Anthosigmella Topsent, 1918. [Rhaphiophora] de on the Great Barrier Reef, Australia (for instance, Topsent, 1900;
Laubenfels, 1936a (Iapsus for Raphiophora Gray). Delaubenjelsia Volz, 1939; Hartman, 1958b; Riitzler, 1973; Hoeksema, 1983;
Dickinson, 1945. Bernatia Rosell & Uriz, 1997 (see also Topsent, Wesche et al., 1997; Schonberg, 2000). Alpha-stages are particu-
1900; Vosmaer, 1932 (but beware of excessive lumping); de larly common (and obvious) in oyster shells ('oyster pest')
Laubenfels, 1936a). between the intertidal and 10m depth, gamma-stages occur free on
sediment bottoms to 40 m depth (New England fishermen call them
Type species 'bay pumpkins' and 'porpoise dung').

Ctiona celata Grant, 1826a (by monotypy).

Definition

Excavating, mostly cryptic Clionaidae without elaborate

aquiferous morphology; with microscleres composed of raphides
or spirasters, including amphiastrose modifications of spirasters or
entirely smooth forms (sinuous microrhabds).

Diagnosis

Sponges primarily in alpha growth form (excavating cham-

bers, communicating through papillae), some species developing
beta stage by merging of papillae, very few regularly outgrowing
their substratum and occurring in gamma stage. Some gamma-
stage species attain large, irregular massive or cup shape but do not
develop specialized incurrent or excurrent features other than,
in some forms, the original pori- and oscula-bearing papillae.
Spicules are tylostyles as megascleres, very thin oxeas (raphides)
as accessory scleres but never of structural importance, and
spiraster-like microscleres. Spirasters are straight, bent, kinked,
spiraled, or undulate spiny rhabds, including amphiastrose forms
clearly derived from true spirasters; extreme forms may be entirely Fig. 3. Cliona celata, underwater photograph of gamma stage, eastern
smooth. North Atlantic, Ireland (ca. 0.5 X natural size) (photo courtesy T. Crabtree).
Porifera· Demospongiae • Hadromerida • Clionaidae 177

A o

B
c __ E
Fig. 4. Cliona celata, habitus and spiculation. A, alpha stage in oyster shell, specimen USNM 23360, Tangier Sound, Chesapeake Bay, western Atlantic
(scale 5mm). B, gamma stage, specimen USNM 8514, off Woods Hole Massachusetts, USA (scale 5mm). C, tylostyles and raphides (from a juvenile),
USNM 23360 (scale 50 /-Lm) . D, tylostyle heads magnified (scale 5 /-Lm). E, spirasters, reported for some juvenile specimens of the species (Topsent, 19(0),
but possibly belonging to C. viridis (afterVosmaer, 1932) (scale 5 /-Lm).

Fig.5. Cliona caribbaea anatomy and excavations. A, longitudinal section through a single papilla (flanked by rock substratum), showing incurrent canals
and perpendicular spicule orientation (scale 200 /-Lm). B, SEM view of fractured ectosome with tylostyles in bouquet arrangement (scale 100 /-Lm). C, exca-
vated chambers in (fractured) oyster shell, sponge tissue removed (scale 5 mm). D, excavated papillar openings in oyster shell (viewed from above) , sponge
tissue removed (scale 5 mm) . (A, C, D from Rtitzler, 1974.)

Description. The color of live specimens is rich yellow to older specimens with much of the substratum removed, the sponge
golden and orange yellow. The consistency of the ectosome ectosome becomes continuous at the surface, between the papillae
(including papillae) is rather tough (from spicule reinforcement), which retain their structure. After additional growth periods the
the choanosomal chamber tissue is soft. In the alpha-stage, papillae sponges become massive, hemispherical or irregular, depending on
are well separated, 0.5-3 .0mm in diameter. They connect directly environmental conditions and the nature of the original substratum.
to large chambers, 2.5-4.5 mm and more. The walls of the cham- Spicules in the papillae are arranged in palisade-fashion, points
bers (after sponge material is removed) often appear angular and outward and protruding beyond the ectosome. In the chambers
are connected by numerous small (up to 0.7 mm) circular pores. In spicules are scattered, particularly in early stages, later become
178 Porifera· Demospongiae • Hadromerida • Clionaidae

partly organized in tracts. Spicules consist mainly of tylostyles

which may be accompanied by raphides as the only microscleres
(forming trichodragmas in the chambers). For young specimens,
small spirasters have been reported to occur mainly in the papillae
(Topsent, 1900). Tylostyles are long and slim and have inconspicu-
ous inflations for heads which are usually subterminal.
Their dimensions range with specimen and location from about
200-400 X 3-12 /-Lm, the average is about 300 X 8 /-Lm and tends to
be smaller when located in the choanosome. Raphides are 100-
150 X < 1 /-Lm. Spirasters are reported to be short and stout,
20-25 X 2-3 /-Lm.
Remarks. It has been well established that clionaids may
lose one or the other spicule type during the course of one life
cycle, for instance, only early stages of Cliona celata are with
microscleres (Topsent, 1900), making systematic judgement based
on museum specimens difficult or impossible. However, cladistical
analysis of a large number of species (Rosell & Uriz, 1997) sup-
ports the grouping as here defined and the inclusion of the genus
Anthosigmella Topsent whose characteristic 'anthosigmas' are rare
or absent in many specimens but are really derived from typical
Fig. 6. Cliona spiculation. A, tylostyles, C. amplicavata (scale 20 j.Lm).
B, raphide, C. amplicavata (scale 20 j.Lm). C, raphide tip enlarged, C. spirasters. I do, however, disagree with the decision by Rosell &
amplicavata (scale I j.Lm). D, tylostyle head and raphides, C. amplicavata Uriz (1997) to establish a separate genus Bematia (for Cliona ver-
(scale 10 j.Lm). E, tylostyle head, C. paucispina (scale 5 j.Lm). F, G, tylostyle mifera Hancock; with smooth, vermiform microstrongyles). The
heads, C. caribbaea (scale 5 j.Lm). (From Riitzler, 1974.) lack of spines on the microscleres of this sponge are considered a
minor modification and does not justify generic separation; a tran-
sitional form with strongly reduced spines is exemplified by Cliona
paucispina Riitzler (1974). Delaubenfelsia Dickinson is massive
without known aquiferous specialization (the original material
exists only in fragments), and its microscleres are very similar to
those of Anthosigmella. Two other generic synonyms are included
here, following recent rediscoveries of subsequent names to replace
[Papillina] Schmidt, 1862 (preoccupied by Papillina Conrad, 1855,
Mollusca): Taguilla Boweberbank, 1874, Tapiliata Bowerbank,
1874, both curiously introduced in the same work and on the same
page. Raphiophora Gray, 1867a is an objective synonym of
Poterion Schlegel, 1858, both having Spongia patera Hardwicke,
1822 as their type species.

CUONAOPSIS NOM NOV.

Synonymy

[Clionopsis] Thiele, 1905: 412 (preocc. for Clionopsis (lapsus

pro Cliopsis) Bronn, 1862: 645, Mollusca) (according to Neave,
1940).

Type species

Clionopsis platei Thiele, 1905 (by original designation) (Fig. 8).

Definition

Clionaidae with oxeas and spirasters.

Diagnosis
Fig. 7. Cliona spiculation, spiraster-type microscleres at the same scale. A,
spirasters, C. flavifodina. B, spirasters and diplasters, C. dioryssa. C, spi-
raster and diplaster, C. caribbaea. D, spirasters, C. paucispina. E, sinuous
Excavating sponges with ty10sty1es and oxeas as megascleres
microstrongyles, C. vermifera. F, anthosigma and spiraster, C. varians and spirasters as microscleres. Only beta- to gamma-stage (thickly
(scale 10 j.Lm). (From Riitzler, 1974.) encrusting) specimens are known (type material).
Porifera' Demospongiae • Hadromerida • Clionaidae 179

CUOTHOSA TOPSENT, 1905

Synonymy

Cliothosa Topsent, 1905a: 95. Thoosa sensu Topsent, 1888:

81 (not Thoosa Hancock, 1849).

Type species

Thoosa hancocki Topsent, 1888 (misspelled as T. hancocci;

by original designation).

Definition

Clionaidae with amphiasters as microscleres.

Diagnosis

Excavating sponges of alpha growth form with large

tylostyles and generally two types of amphiasters as microscleres:
stout, nodulose in the papillae and delicate thin-rayed with termi-
nal branching of the slender rays, in the chambers. The nodulous
amphiasters may be absent in some specimens or species (Topsent,
1928b; Riitzler, 1973). On the other hand, a third kind, a very small
diplaster has been reported to occur in the papillae (Volz, 1939).

Previous review
Fig. 8. Clionaopsis platei, syntype (ZMB 3270). A, oxeas and spiraster I
(scale 50 fLm). B, tylostyles (scale 50 fLm). C, spirasters I (scale 10 fLm). Topsent (1928b); Rosell & Uriz (1997).
D, spirasters II (scale 10 fLm) .
Description of type species
Description of type species
Thoosa hancocki Topsent, 1888 (Figs 9, 10).
Clionaopsis platei (Thiele, 1905) (Fig. 8). Synonymy. Thoosa hancocki (as T. Hancocci) Topsent,
Synonymy. Clionopsis platei Thiele, 1905: 412, fig. 37a-d. 1888: 81, pI. 7, fig. 12. Thoosa hancocki; Topsent, 1928b: I, fig. I;
Material examined. Syntype: 2MB 3270 - Calbuco, Chile. Volz, 1939: 25, fig. 8, pI. 4, figs. 2~; Riitzler, 1973: 634, fig . 7;
Description. The examined type is a flat, 1 cm thick cushion Rosell & Uriz, 1997: 363, fig. 2B. Vioa ramosa Lendenfeld, 1897a:
of about 7 cm2 coverage. According to the original describer 81. Cliona seurati Topsent, 1905a: 96.
(Thiele, 1905: 412), specimens are flat (3~ cm diameter, 1 cm Material examined. Holotype: MNHN DT 2559 (two slides)-
thick, tapering toward the margin) to cone-shaped (5 cm tall) and "in Tridacna gigas"; no further location given. Specimens from
incorporate calcareous particles including balanid shells. Colour in Tunisia (deposited at INSTOP, Salammbo) are described by
alcohol is grayish brown. Inhalant and exhalant areas (2-3 mm Ri.itzler (1973). The species is common in the Mediterranean and
papillar regions) are evident on the surface as they are darker than the Indopacific region; 0.5-lOm depth.
the surrounding surface and slightly depressed. The cortex is gen- Description. The sponge develops only the alpha-stage.
erally formed by a dense arrangement of oxeas accompanied by Papillae are bright yellow but may have an orange tinge; they are
fewer tylostyles and short spirasters. The papillar regions are circular, well separated, 1-6 mm in diameter. Chamber tissue is also
mainly supported by tylostyles which are oriented perpendicularly yellow, but duller. Chambers may be minute in young specimens
to the surface, with points outward, and also joined by short spi- (0.3-1.2mm; Riitzler, 1973) but tend to merge and fill larger spaces
rasters. The choanosome is packed with oxeas and tylostyles in advanced stages (5-20mm; Topsent, 1905a; Schonberg, 2000).
without particular orientation and contains long, thin spirasters. The chambers connect to the papillae through a narrow canal of 10
Choanocyte chambers are oval in cross-section, 20 X 30 f.Lm in mm length, or more. Spicules consist of large, robust tylostyles, and
diameter. Oxeas are smooth, bent in the center, sharply pointed on two types of amphiasters. Tylostyles are straight, with prominent
both ends but in places modified to styloid forms; they measure circular or oval head, usually subterminal; in the type they measure
440-550 X 17-23 .....m. Tylostyles, 300-400 X 11-15 .....m, have 30~ lOX 8-12 .....m. Ramose amphiasters occur in the choano-
rounded heads, rarely irregular or subterminal. Microscleres some and have long, delicate rays with hooks at their distal ends;
consist of two classes spirasters: short, stout with strong spines they measure 26 X 21 .....m, overall. Nodulose amphiasters, if pres-
(15-35 X 7-23 .....m, including spines); and long, slim, sinuous, ent, are in the papillae; they have bulbous swellings instead of rays;
with delicate spines (53-86 X 5-8 .....m, with spines). 20 X 15 .....m. Amphiasters are rare in preparation MNHN DT
Remarks. The specimen described by Desqueyroux (1972) 2559(1) but common in MNHN DT 2559(2).
from Montemar (Valparaiso), Chile, is actually Cliona chilensis Remarks. The validity of this genus was doubted by
(R. Desqueyroux-Faundez, pers. comm.). Annandale (1915a: 22) on the basis of the fact that the spicule
180 Porifera· Demospongiae • Hadromerida • Clionaidae

A __
B c
Fig. 9. Cliothosa hancocki, spiculation. A, tylostyles and enlarged tylostyle heads (scale 50 jJ-m; 10 jJ-m for heads). B, nodulose amphiasters (scale 10 jJ-m).
C, ramose amphiasters (scale 10 jJ-m).

Type species

Cliona northumbrica Hancock, 1849: 336, pi. 14, fig. 5

(=Cliona vastifica Hancock, 1849: 342, pi. 15, fig. 2) (by subse-
quent designation, Rosell & Driz, 1997).

Definition

Clionaidae with tylostyles and rnicrospined oxeas and

microrhabds.

Diagnosis

Sponges in alpha- and beta-forms. Delicate tylostyes as megas-

cleres, accompanied by microspined (rarely smooth) rnicroxea and
rnicrospined rnicrorhabds as rnicroscleres. Microrhabds are usually
straight or bent but may also be centrotylote and sinuose.
Microxeas do not have a particular location or orientation in the
tissue.

Previous review
Fig. 10. Cliothosa hancocki, SEM views of spicules. A, thin-rayed amphi- Rosell & Driz (1997).
aster (scale 10 jJ-m). B, nodulose amphiaster (scale 10 jJ-m). C, tylostyle
(scale 100 jJ-m). D, tylostyle head (scale 10 jJ-m). (A, C, D, specimen ZMA
POR 8661, Indonesia, photo courtesy R.W.M. Van Soest; B, from Description of type species
Schonberg, 2000, photo courtesy C.H.L. Schonberg.)
Pione vastifica (Hancock) (Figs 11-13).
Synonymy. Cliona northumbrica Hancock, 1849: 336, pi. 14,
complement of a species may shift during different growth phases. On fig. 5; Hancock, 1867: 237, pi. 7, fig. 1; RUtzler & Stone, 1986: 665,
the other hand, the possible importance of the presence of two size fig. 5e, f. Cliona vastifica Hancock, 1849: 342, pi. 15, fig. 12;
classes of tylostyles in this genus has been pointed out (Rosell & Driz, Hancock, 1867: 237, pi. 7, fig. 2; Topsent, 1900: 56, pi. 2, figs 3-9;
1979). Two species (in addition to the type) were recently transferred to Topsent, 1932c: 558, fig. 3; Topsent, 1934c: 99; Volz, 1939: 8, fig. 2,
this genus: Cliona quadrata Hancock (by RUtzler & Stone, 1986: 666) pi. 1, fig 3, pi. 2, fig. 3, pi. 3, fig. 1; RUtzier, 1973:633, fig. 6; RUtzler
and Spirastrella aurivillii Lindgren (by Rosell & Driz, 1979: 363). & Stone, 1986: 667, fig. 6c, d. Cliona lampa de Laubenfels forma
occulta RUtzler, 1974: 23, fig 17b, d; 20d. Pione vastifica (Hancock),
PlONE GRAY, 1867 Rosell & Driz, 1997: 362, fig. 3C. (See also Topsent, 1900: 56.)
Material examined. Syntypes (Cliona northumbrica): HMN
Synonymy 4.16.15-4.16.17 (3 slides) - northeast coast of England. Syntypes
(Cliona vastifica): HMN 4.16.36-4.16.39 (4 slides) - Firth of
Pione Gray, 1867a: 525. Archaeocliona Czemiavsky, 1878: Forth, Scotland. (For further information see RUtzler & Stone,
396; 1879: 244, pi. iv, figs 17a-f; 1880: 7. Gapoda de Laubenfels, 1986: 665, 667.) Raphyrus hixonii von Lendenfeld: BMNH
1936a: 144. 86.8.27.605 - Port Jackson, N.S.W, Australia.
Porifera· Demospongiae • Hadromerida • Clionaidae 181

Distribution. The species has been reported from many loca- because their size is at the borderline of light-microscope resolution.
tions in the boreal and tropical Atlantic, the Mediterranean, and Still, some of the microxeas may indeed be smooth. Tylostyles
Indopacific; it has tolerance for a wide range of salinity. are straight and slender with well pronounced rounded head.
Description. Only microscope preparations (cleared tissue AVeraging several populations and localities, tylostyles measure
squashes and acid-cleaned spicule spreads) are left of the types. about 250-300 X 4-5 /-Lm. Microspined microxeas are fusiform,
Material of Pione northumbrica shows tylostyles (330 X 7 /-Lm), slightly bent in the center, 60-100 X 2-4 /-Lm. Spiny microstrongyles
microspined oxeas, more or less bent in the center (110 X 5 /-Lm), and are straight or bent, or sinuous, depending on the locality; 8-16 X
microspined microrhabds (14 X 3/-Lm) that are bent or wavy (2-5 1-3 /-Lm. The species tolerates fairly low levels of salinity (Old, 1941),
bends); slides of Pione vastifica contain tylostyles (300 X 5 /-Lm), and presence of gemmules has been reported (Riitzler, 1974).
microspined oxes, slightly bent in the center (100 X 3/-Lm), and Remarks. Rosell & Uriz (1997) resurrected this genus based
microspined microrhabds (12 X 3/-Lm) that range from straight and on their cladistic analysis. These authors also pointed out that
spindle-shaped to s- and w-shape (Riitzler & Stone, 1986: 665,667; Topsent (1888) synonymized the type species of Pione, Cliona
figs 5e, f, 6c, d). The living sponge is usually red to orange-red, in northumbrica Hancock with Cliona vastifica Hancock and favored
some locations light yellow, with duller-colored interior. Papillae are the latter name although C. northumbrica was described several
very small, under 1 mm in diameter. Chambers are equally small and pages before C. vastifica (Hancock, 1849: 336, 342). Because the
connect directly to the papillae, without canal. As a rule there are name 'northumbrica' remained unused for more than 50 years, the
three types of spicUles, tylostyles, microxeas and microrhabds. Some type species is correctly called Pione vastifica (ICZN Art. 23.9.1.).
authors report one or the other category of microscleres missing but Riitzler (1974) introduced a new papillate 'variety' of Pione (as
thorough search, possibly of several specimens in a population, usu- Cliona) lampa, forma occulta, which was considered an alpha-
ally reveals them. The microscleres are covered by minute spines form of the encrusting P. lampa (de Laubenfels, 1950a), but subse-
that are easily overlooked by a low-quality compound microscope quently was shown to be a separate species, P. vastifica (Riitzler,
2002). Archaeocliona Czemiavsky is considered to be a brackish-
water form of Pione, with smooth microxeas and spiny
microstrongyles lost or overlooked by the author. Gapoda de
Laubenfels (established for Raphyrus hixonii von Lendenfeld,
1886) has the typical spiculation of Pione, but with a third, spi-
raster-like microsclere added.

SPHECIOSPONGIA MARSHALL, 1892

Synonymy

Spheciospongia Marshall, 1892. Heterocliona Verrill, 1907.

Type species

Fig. 11. Pione iampa, beta-stage habitus, thinly coating a recently dead Alcyonium vesparium Lamarck, 1815: 78 (by original
coral; from the type locality, Bermuda (0.5 X). designation).

A
B __ c
Fig. 12. Pione vastifica, spicules. A, tylostyles (scale 50 j.Lm). E, microxeas (scale 10 j.Lm). C, spiny microrhabds (scale 2 j.Lm).
182 Porifera· Demospongiae • Hadromerida • Clionaidae

Fig. 13. Pione, SEM views of microscleres. A, microspined microxea, P. lampa. B, enlarged view of spination on microxea, P. lampa. C, spiny microrhabds,
P. lampa. D, spiny microxeas, P. hixonii. E, spirasters, P. hixonii. F, spiny microrhabds, P. hixonii. (Scales lO!-Lm for A, D, E; 5!-Lm for B, C, E)

Definition Description. Topsent (1933: 30) states that the Lamarck

collection at the Paris Museum contains four specimens, two
Massive Clionadae with complex, incurrent pore sieves, abun- relatively small ones having a label by Lamarck, and two larger
dant large tylostyles in the choanosome and rare, minute spirasters ones, labeled simply "Suberites, Antilles, M. Mauge, 1799".
and their amphiastrose derivatives in the ectosome. The specimen piece with the label cited above is about 43 cm tall
and 30 cm in diameter. has a large oscular cavity with an 11 em
Diagnosis opening on top and several large (5-20 mm) pore sieves along the
sides. Pores are 0.5-1.5 mm in diameter and open in a large
Large-growing sponges with cribiporal chones (pore sieves) canal that branches into the interior of the body. Dry specimens are
and simple, multiple, or septate oscula. In the ectosome, numerous grayish brown to purple, the live animal is nearly black, with a pur-
robust tylostyles forming a dense tangental network and ple tinge, but there is also a yellow variety (forma pallida, Vicente
rare, minute spirasters and spiraster-derived amphiasters; the et aZ., 1991). The shape may be that of a large cake with many
latter are also located in the canal linings. In the choanosome, small oscula in a group on top, some resemble volcanoes with just
tylostyles occur densely scattered and in strands. TYlostyles may a few large oscula close together. The spicules are large, densely
be modified to styles, tylostrongyles, and strongyles. Sponges packed tylostyles and minute, rare spirasters and spiraster-derived
excavate limestone in early growth stage, forming large, tissue- amphiasters, located mainly in ectosome and canal linings.
filled cavities, not a series of distinct chambers as do other Tylostyles are curved or double-bent, with small rounded heads
clionaids. and uniform thickness until near the tip where they taper to a point;
in the type they are 250-470 X 8-9 J.Lm, population means range
Previous review 280-410 X 6--9 J.Lm. Spirasters have 1-5 bends and measure 8-26 X
4 J.Lm, including spine clusters; the shorter forms are straight
Topsent, 1933; Vicente et aZ., 1991. amphiasters.
Remarks. The original author of the genus was not aware
Description of type species of the small spirasters and considered this sponge a simple suberitid
(Marshall, 1892: 32). However, Topsent (1933: 32), recognizing
Spheciospongia vesparium (Lamarck) (Figs 14, 15). the importance of the pore sieves of these sponges, left the genus
Synonymy. AZcyonium vesparium Lamarck, 1815: 78; intact and transferred these sponges to Spirastrellidae. A canal
Spheciospongia vesparium Marshall, 1892: 32. (See also Topsent, system of substantial extent and bore diameter adds to the
1933; Vicente et aZ., 1991). unique character of these sponges and supports a large and diverse
Material examined. Holotype: MNHN LBIM DT 3419 - fauna of endozoic invertebrates (Pearse, 1934; Westinga &
"labelled 142. Alcyonium vesparium Lamarck". Hoetjes, 1981).
Porifera· Demospongiae • Hadromerida • Clionaidae 183

c
Fig. 14. Spheciospongia vesparium, habitus and spiculation. A, early excavating stage, Bermuda (scale 10 mm). B, tylostyles (scale 50 /Lm). C, spirasters
(scale 5/Lm).

Fig. 15. Spheciospongia vesparium, adult habitus and SEM views of spicules. A, specimen from Puerto Rico showing pore sieve (arrow; O.17X).
B, spirasters and amphiastrose derivatives (scale 2/Lm). (From Vicente et al., 1991.)

THOOSA HANCOCK, 1849 Diagnosis

Synonymy Sponges are always papillate. Tylostyles may be missing in

some specimens, populations, or species. There are many types
Thoosa Hancock, 1849 (not Thoosa sensu Topsent, 1888: 81). and variations of microscleres but most commonly there are amphi-
[Annandaleal Topsent, 1928b: 6. Annandalena Topsent, 1932c: asters and oxyasters in the spiculation. Amphiasters are mostly
573. Thooce de Laubenfels, 1936a. tylote, with microspination on the terminal bulbs but variations in
different species include smooth oxeote rays. Oxyasters have a very
Type species small center with 1-{) long, slender rays projecting. The common
reduction of rays can lead to centrotylote sigmoid or tylostylote
Thoosa cactoides Hancock, 1849 (by subsequent designation; fonns. Because rays tend to be curved, biradiate fonns often appear
de Laubenfels, 1936a: 157). like birdwings. Accessory microscleres in some species may
include microxeas and pseudosterrasters (microspined spheres or
Definition rhabds).

Clionaidae with alpha-fonn morphology and microscleres Previous review

composed of small, nodular amphiasters with microspined radii
and large, thin-rayed euasters. Riitzler & Stone (1986).
184 Porifera· Demospongiae • Hadromerida • Clionaidae

-,
.
".
~;.~;
.~.

:
0: "
•
~
, ,
-:

A---- B c
Fig. 16. Thoosa, spiculation. A, tylostyles in T. mollis Volz (1939) (scale 50 fLm). B,amphiasters of T. cactoides (scale 5 fLm). C, oxyasters (after Topsent,
1905a; Volz, 1939) (scale 20 fLm).

Description of type species

Thoosa cactoides Hancock (Figs 16, 17).

Synonymy Thoosa cactoides Hancock, 1849: 445, pI. 13,
figs 1,2; Riitzler & Stone, 1986: 667, fig. 7b.
Material examined. 'Type': HMN 4.17.05, 4.17.06 -
2 slides (acid-cleaned spicule mounts), no locality stated; substrate:
'Meleagrina margaritifera', a mollusk shell.
Description. The type slides show only the characteristic
nodular amphiasters. They are stout, with tylote rays ornamented
by microspination, 23 X 15 fLm in average. The original author
illustrated the excavation pattern (Hancock, 1849, pI. 13, fig. 1),
which extended 15-18 cm through the substrate shell and gave the
diameter of the elliptical chambers as 5.1 mm. From better known
species (Thoosa annata, Topsent, 1904b; T. mollis, Volz, 1939) it
appears that the papillae are always small (up to 1 mm), at least in
comparison to the relatively large chamber, inconspicuous in color
(whitish), and contain the nodulose amphiasters (and pseudoster-
rasters, in T. annata). The chamber tissue is supported by tylostyles
and the large euasters. Reproduction through gemmulae has been
observed. Members of the genus are known from a depth range of
2m to almost 600m.
Remarks. The lack of certain spicule types in a population
or in some species is not unusual and does not justify establishing a
separate genus, as was done in the case of Thooce de Laubenfels,
1936a (for Thoosa socia lis Carter, 1880b: 56). Topsent (1920b)
Fig. 17. Thoosa armata, SEM views of spicules, specimen ZMA POR
already arrived at that conclusion when he compared species or 2390, Cura~ao, West Indies. A, amphiaster (scale 10 fLm). B, C, calthrop-
forms of Thoosa missing either tylostyles or oxyasters. like oxyasters (scale 10 fLm). D, enlarged center of oxyaster (scale I fLm).
The genus [Annandalea] Topsent, 1928b: 6 (preocc. by (Photo courtesy R.W.M. Van Soest).
Annandalea Needham, 1909, Neuroptera, Insecta) was erected
for type species Thoosa laeviaster Annandale, 1915a: 22, fig. 4 size. When Topsent became aware that the name Annandalea was
(by original designation), a Thoosa-like species with angular already taken, he changed the name in a footnote to Annandalena
microxeas of 80 X 3 fLm in addition to the usual knobby amphi- Topsent, 1932c: 573, which is here designated a junior synonym of
asters of 40-80 X 6-13 fLm and reduced oxyasters of similar Thoosa.
Porifera· Demospongiae • Hadromerida • Clionaidae 185

VOLZIA ROSELL & URlZ, 1997 Previous review

Synonymy Rosell & Uriz (1997).

Volzia Rosell & Uriz, 1997: 363. Description of type species

Type species Cliona albicans Volz, 1939 (Fig. 18).

Synonymy. Cliona albicans Volz, 1939: 21, fig. 6,
Cliona albicans Volz, 1939 (by original designation). pI. 3 fig. 3; Riitzler, 1965: 21; Volzia albicans; Rosell & Uriz,
1997: 363.
Definition Material examined. Type material: presumed lost during
WW II (but the description by the original author is precise). Other
Clionaidae with accessory oxeas arranged in the papillae in material. A few specimens were found near the type locality off
palisade fashion. Rovinj, Croatia, 1.2-10 m depth (Riitzler, 1965).
Description. Papillae of the live sponge are whitish to
Diagnosis brownish and very small (0.5-0.8 mm). The chambers, too, are
small (2 mm), spherical, and extend to 20 mm below the substratum
Sponges display only the papillate (alpha-) growth form. surface; they are filled with whitish tissue. Spicules consist of rare
Tylostyles are rare and restricted to the chambers. Oxeas are small tylostyles and an abundance of oxeas. The tylostyles are fairly
but common. Oxeas are mixed-in with tylostyles in the chambers; straight and slender, with gradually narrowing sharp points,
in the papillae they are arranged in palisade and in peripheral 300-360 X 5.00-7.5 fLm. Tylostyle heads are barely wider than the
strands. Microscleres absent. shaft and subterminal. The numerous oxeas are slightly bent,
shorter when located in the chambers (88-95 fLm), longer in the
papillae (90-110 fLm).
Remarks. Only two species (the type, and Volzia rovignen-
sis) from one locality (Rovinj, Croatia) are known. It is significant
that the oxeas, although rather small, take the place of tylostyles in
forming a palisade in the ectosomal papillae.

ACKNOWLEDGEMENTS

I am most grateful to the following friends and colleagues for

helping to improve my contributions to the Systema Porifera proj-
ect, for this and other chapters in this volume. Specimens from
B various collections and related data and illustrations were provided
by Clare Valentine (BMMH London), Rob Van Soest and Elly
Beglinger (ZMA Amsterdam), Claude Levi (MNHN Paris), Nicole
Boury-Esnault and Jean Vacelet (SME Marseille), Maria Uriz
(Blanes, Spain), Ruth Desqueyroux-Faundez (MNHN Geneva),
Barbara Calcinai, Maurizio Pansini, Roberto Pronzato and Michele
Sara (DTRG Genova), Gorgio Bavastrello (Ancona, Italy),
Giuseppe Corriero (Bari, Italy), Peter Dworschak and Helmut
Sattman (Vienna), Cristine Schonberg (Oldenburg, Germany), and
Penny Berents (AM Sydney). Scanning electron micrographs
(SEMs) were taken and processed by Susann Braden, Kathleen
McGinley, and Michelle Nestlerode. Literature was researched by
Kathleen McGinley, Michelle NestIerode, and Martha Robbart.
For drawing, digital figure processing, and illustration design
I thank Patricia Condit, Kathleen McGinley, Ana Maria Mardones,

A
c __ and Molly Kelly Ryan. Travel funds for meetings relating to
Systema Porifera were provided by the Smithsonian Research
Fig. 18. Volzia albicans, spiculation. A, tylostyles (scale 50 jJ.m).
Opportunities Fund (administrated by Ross Simons and Wendy
B, tylostyle head, enlarged (scale 5 11m). C, oxeas (scale 20l1m). (After Wiswall) and the Caribbean Coral Reef Ecosystems Program
Volz, 1939.) (Contribution number 622).
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob w.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Hemiasterellidae Lendenfeld, 1889

John N.A. Hooper

Queensland Museum, P.O. Box 3300, South Brisbane, Qld, 4101, Australia. (JohnH@qm.qld.gov.au)

Hemiasterellidae Lendenfeld (Demospongiae, Hadromerida) contains 11 nominal genera, of which six are valid, and approximately 20
described species worldwide. Species live in all oceans, predominantly in shallow waters, are often dichotomously branching or flabel-
late, with hispid surface or conules and stiff texture. The family has a unique combination of euasters mainly at the surface, and monax-
onic megascleres (styles and/or oxeas) with distinctly structured arrangement of axial and extra-axial skeletons. Genera are differentiated
predominantly on the basis of whether megascleres are diactinal or monactinal, development of the axial and extra-axial skeletons,
presence or absence of ectosomal specialisation, and euaster morphology.
Keywords: Porifera; Demospongiae; Hadromerida; Hemiasterellidae; Adreus; Axos; Hemiasterella; Leptosastra; Paratimea; Stelligera.

DEFINITION, DIAGNOSIS, SCOPE However, it is more likely that the concept of 'Axinellida' is poly-
phyletic and Hemiasterellidae belong to Hadromerida, with the
Synonymy implication that the characters compression of the axial skeleton
and differentiated axial and extra-axial skeletons are homoplasious
Hemiasterellidae Lendenfeld, 1889b. Leptosastrinae Topsent, throughout the Demospongiae. Topsent (1928c) established
1928c: 48, 56. [Astraxinellidae] Dendy, 1905 (nomen nudum). Leptosastrinae for two of his own genera, Leptosastra Topsent,
1904b, and Spirorhabdia Topsent, 1918 (with type species
Definition Spirastrella vidua Schmidt, by monotypy), based on their common
possession of ectosomal rhabds - smooth in the former and spined
Hadromerida with compressed axial plumose - plumo- in the latter. The latter genus is now included in Crellidae whereas
reticulate extra-axial skeleton composed of styles, oxeas or both, Leptosastra, and hence the subfamily name, belongs to
and smooth or partially microspined euasters. Hemiasterellidae based on possession of euasters (but see below
under Remarks for genus Leptosastra).
Diagnosis
Differences with similar families
Encrusting, cup-shaped, arborescent or branching growth
forms; megascleres are styles (in one genus acanthostyles), oxeas Hemiasterellids share their monactinal megascleres with the
or both enclosed within axially compressed spongin fibres, or Hadromerida and for this reason they were formally transferred
basally compressed in encrusting taxa, and plumose to plumo- there by Voultsiadou-Koukoura & Van Soest (1991a). They differ
reticulate extra-axial branches, or sometimes without a definite from other hadromerids such as Polymastiidae and Spirastrellidae
axis; bouquets of smaller styles and/or oxeas sometimes protrude in having axially compressed choanosomal skeletons (sometimes
through surface; microscleres are euasters, smooth or partially loosely constructed or hymedesmoid, e.g., Paratimea, Leptosastra)
microspined, often confined to an ectosomal crust. and bouquets, tracts or single extra-axial spicules near the periph-
ery, but they show similarities to Aaptos (Suberitidae) and Tethya
Scope (Tethyidae) in having predominantly style megascleres rather than
tylostyles although some hemiasterellids retain long oxeas in the
Eleven genera have been included, or are potentially included extra-axial skeleton. Hemiasterellidae can be further differentiated
in this family, although only six of these are now recognised, one from Tethyidae in lacking an organic cortex and a substantially
(Leptosastra) monotypic, poorly known, and included here specu- poorer diversity of euaster morphologies, and from Jaspis
latively. Species are found in all seas but only about 25 species (Ancorinidae, Astrophorida), the latter having megascleres that are
have so far been described. diactinal and confused in arrangement in the choanosomal skele-
ton, with a subectosomal tangential arrangement of megascleres
History and biology supporting a crust of euasters. Voultsiadou-Koukoura & Van Soest
(1991a) suggested further that similarities in euaster morphology
Hemiasterellidae was established for Hemiasterella Carter between some Jaspis and some Hemiasterella may show in future
and Epallax Sollas (not Gray, cf. de Laubenfels, 1936a; that these genera are more closely related than presently
Wiedenmayer, 1977b) with brief diagnosis "Axinellidae with stel- acknowledged by their ordinal classification (see below).
late microsc/era" (Lendenfeld, 1889b). Reid (1970) hypothesised
that the common possession of asters in Axinellida (Le., Previous reviews
Hemiasterellidae), Astrophorida and Hadromerida formed a 'natu-
ral' taxon (Astrotetraxonida) having common ancestral stock. Hooper (1986), Voultsiadou-Koukoura & Van Soest (1991a).

186
Porifera· Demospongiae • Hadromerida • Hemiasterellidae 187

KEY TO GENERA

(1) Axial skeleton poorly differentiated, plumo-reticulate skeleton .................................................................................... Hemiosterella

Axial skeleton well developed, compressed reticulate ...... .......... ................ ................ ...... .......... .......... .................. ... ............ ............. 2
Axial skeleton lacking, but with a basal hymedesmioid skeleton ....................................................................................................... 4
(2) Cortex poorly developed, with only a light crust of euasters (with smooth or spined rays, with oxeote, strongylote or
tylote terminations) ............................................................................................................................................................................. 3
Cortex well developed, with a thick crust of hexaradiate to cruciform euasters (anthasters) with thick heavily spined rays, confined
mainly to the peripheral region .... .......................... ................ ................. ............... ................. .......... ................. .......... ..... ............ Axos
(3) Microscleres euasters with thin strongylote or tylote rays, usually curved or sinuous, often branched, smooth or spined rays with
small centrum ......... .......... ...... .......... ........... ........... ........... .......... ................ ...... ....... ... ................. ............. ............. ........... ........ Adreus
Microscleres short rayed oxyspheraster euasters with entirely smooth rays and thick centrum ............ ............................... Stelligera
(4) Choanosomal megascleres tylostyles erect on substratum ................................................................................................... Paratimea
Choanosomal megascleres acanthostyles erect on substratum .............. ................ .......... ................. ................. .......... ...... Leptosastra

ADREUS GRAY, 1867 thick, slightly curved at centre, evenly rounded smooth bases,
tapering long points often telescoped (520-780 X 6-10 fLm),
Synonymy smaller extra-axial styles ranging from straight to sinuous, with
smooth rounded bases and sharp fusiform points (180-350 X
Adreus Gray, 1867a: 545. 3-5 fLm); euasters with thin strongylote or tylote rays, usually
curved or sinuous, often branched, smooth or spined rays with
Type species small centrum (entire spicule 8-12 fLm diameter).
Remarks. Adreus is arborescent with monactinal megascle-
Dictyocylindrus fascicularis Bowerbank, 1866, 1874b (by res occurring in bundles, ascending towards the periphery and with
monotypy). apparent size differences between choanosomal and ectosomal
megascleres (Topsent, 1928c). Euasters are not abundant, found
Definition mainly in the ectosomal region, but they are unusual in often hav-
ing branching sinuous rays. Sexual reproduction with oviparous
Hemiasterellidae with strongly developed axial skeleton larvae has been reported by Brien (1973a). Voultsiadou-Koukoura &
composed of long styles, poorly developed extra-axial skeleton Van Soest (l991a) included only three species in the genus
composed of plumose brushes of smaller styles, and euasters with (Dictyocylindrus fascicularis Bowerbank, 1866 from the British
smooth or spined, thin strongylote or tylote sinuous rays which Isles, Vibulinus micraster Burton, 1956 from W Africa and
may be branched. Mauritania, and Timea stylifera Arndt, 1927 from the Caribbean).
The type species has usually been attributed by authors to the first
Diagnosis volume of Bowerbank's monograph series (1864), whereas I could
only find it mentioned in the second volume (1866) and figured in
Arborescent growth form; choanosomal skeleton with the third volume (1874b).
strongly compressed axis composed of long smooth tylostyles in
bundles running longitudinally through branches, and poorly Distribution
developed extra-axial skeleton composed of sparse plumose
brushes of smaller smooth styles ascending to periphery; euasters East Atlantic, Caribbean.
with curved or sinuous, smooth or spined, strongylote or tylote rays
often branched, mainly confined to the ectosomal region.
AXOS GRAY, 1867
Description of type species
Synonymy
Adreusfascicularis (Bowerbank, 1866) (Fig. lA-H).
Synonymy. Dictyocylindrus fascicularis Bowerbank, 1866: Axos Gray, 1867a. Echinospongia Gray, 1870a. Taxonomic
110; 1874b: 45. Adreusfascicularis; Gray, 1867a: 545. decision for synonymy: de Laubenfels (1936a).
Material examined. Syntypes: BMNH 1910.1.38,39,43,44
(slides BMNH 1910.1.1.2461-2) - Vajon Bay, Guernsey. Type species
Description. Thin, stiffly branched, arborescent growth
form; axial skeleton strongly developed occupying majority of Axos cliftoni Gray, 1867a (by monotypy).
branch diameter, composed of bundles of long styles running lon-
gitudinally through branches; extra-axial skeleton more poorly Definition
developed with sparse plumose brushes of smaller styles, some
vestigial or vermiform, ascending to ectosome but rarely protrud- Hemiasterellidae with compressed axial skeleton and radial
ing through surface; peripheral skeleton with light crust of extra-axial skeleton composed of a single category of smooth
euasters; fibres well developed and collagen abundant throughout styles; hexaradiate and cruciform euasters (anthasters) with thick
mesohyl; megascleres styles of 2 sizes: larger axial styles long, heavily spined rays form a thick surface crust.
188 Porifera· Demospongiae • Hadromerida • Hemiasterellidae

E
G

A
~

Fig. 1. Adreus fascicularis (Bowerbank) (syntypes). A, choanosomal styles (scale 150 I-l-m). B, choanosomal style base and point (scale 10 I-l-m).
C, ectosomal styles (scale 50 I-l-m). D, ectosomal style base and point (scale 10 I-l-m). E, euasters (scale 10 I-l-m). F, section through branch (scale 250 I-l-m).
G, reconstruction of idealised skeletal structure. H, specimen figured by Bowerbank, 1874b (from his plate 18, fig. I).

Diagnosis Material examined. Holotype: BMNH 1870.8.26.31 - NW

Australia.
Cylindrical branching and flabelliform growth forms with Description. Dichotomously branching, thickly cylindrical,
prominent surface conules; well developed compressed axial skele- whip-like, rubbery, flexible growth form with stipitate, rhizomous
ton composed of styles; extra-axial skeleton composed of radial basal attachment, and prominent surface conules; ectosomal skele-
spicule tracts of the same styles perpendicular to the axis and ton detachable with heavy cortex of euasters pierced at regular
protruding into surface conules; asters hexaradiate to cruciform intervals by tracts of extra-axial styles forming light, continuous
with thick heavily spined rays (anthasters), confined mainly to the palisade; choanosomal axial skeleton massive, heavily com-
peripheral region forming a thick crust. pressed, irregular in cross· section (corresponding to distribution
of surface conules), consisting of tight bundles of styles packed
Description of type species together running longitudinally through branches; extra-axial
skeleton plumose, non-anastomosing, composed of styles radiating
Axos cliftoni Gray, 1867a (Fig. 2A-G). from axis to surface conules, with scattered euasters becoming
Synonymy. Unidentified sponge - Bowerbank, 1864: 260. more dense towards the periphery; megascleres consist of a single
Axos cliftoni Gray, 1867a: 546 (Not Axos cliftoni; Carter, 1879d: category of smooth fusiform styles (178-912 X 1.5-22 ILm);
284); Dictyocylindrus dentatus Bowerbank, 1873b: 321 ; microscleres hexaradiate or cruciform euasters with spined rays
Echinospongia australis Gray, 1870a: 272. (anthasters) (8-37 ILm).
Porifera· Demospongiae • Hadromerida • HemiastereUidae 189

Fig. 2. Axos cliftoni Gray (holotype). A, styles (scale 100 j.Lm). B. style base and point (scale 10 j.Lm). C, smaller euasters (scale 10 j.Lm). D, larger euasters
(scale \0 j.Lm). E, peripheral skeleton (scale 250 j.Lm). F, axial skeleton (scale 250 j.Lm). G, reconstruction of idealised skeletal structure. H, specimen of
A. flabelliformis in situ (photo the author).

Remarks. Axos was established for A. cliftoni, named from (type species E. australis Gray, 1870a (by monotypy» was syn-
a single spicule depicted by Bowerbank (1864). De Laubenfels onymous with Axos, although the holotype is unknown and this
(1936a) suggested that Axos and Adreus were closely related, cannot be corroborated.
with similar growth form and spiculation, with only euaster mor-
phology (branching rays) and a greater proportion of styles than Distribution
oxeas in Axos differentiating the two genera. This observation is
erroneous, whereby Axos has only monactinal megasderes Endemic to northern Australia: WA, NT and Far N Qld.
(styles) and euasters have spined (not branching) rays (in fact
euasters of Adreus are characterised by possession of sinuous
branching rays, not those ofAxos as supposed by de Laubenfels, HEMIASTERELLA CARTER, 1879
1936a). Only two species of Axos are known, endemic to northern
Australia. Burton (1934a) synonymised both species, presumably Synonymy
based on their nearly identical gross morphology, but Hooper
(1986) showed that they differed substantially in their euaster Hemiasterella Carter, 1879d. Epallax Sollas, 1888. Kalastrella
morphology, both of which are classed as 'anthasters' as defined Kirkpatrick, 1903b. Tylaspis Levi & Levi, 1983b. Taxonomic deci-
by Wiedenmayer (1994), for euasters with forked rays. De sion for synonymy: Voultsiadou-Koukoura & Van Soest (1991a)
Laubenfels (1936a) also suggested that Echinospongia Gray and present work.
190 Porifera· Demospongiae • Hadromerida • Hemiasterellidae

Fig. 3. Hemiasterella typus Carter (holotype). A, styles (scale 250 fLm). B, style base and point (scale 10 fLm). C, small euasters (scale 10 fLm). D, large
euasters (scale 10 fLm). E, peripheral skeleton (scale 250). F, axial skeleton (scale 250 fLm). G, reconstruction of idealised skeletal structure.
H, Hemiasterella (Epallax) callocyathus Sollas, 1888 (from his plate 10, fig. 1).

Type species regions, composed of styles and/or oxeas without functional

arrangement to any particular part of skeleton; ectosomal euasters
Hemiasterella typus Carter, 1879d (by original designation). typically with thick, acanthose, strongylote rays, sometimes
calthrops-like (rays reduced to 2-4, curved, asymmetrical or
Definition branching rays), predominant in peripheral region but not forming
a surface crust.
Hemiasterellidae with vasiform, cup-shaped or massive
growth forms lacking axial condensation, with plumo-reticulate Description of type species
choanosomal skeleton; euaster with thick acanthose strongylote
rays, sometimes calthrops-like. Hemiasterella typus Carter, 1879d (Fig. 3A-G).
Synonymy. Hemiasterella typus Carter, 1879d: 146.
Diagnosis Materilll examined. Holotype: BMNH unknown. Other
Material. BMNH 1936.3.4.406, slide BMNH 1936.3.4.406a -
Exclusively vasiform, plate-like, flattened branching or massive 'John Murray' Expedition.
growth forms; choanosomal and peripheral skeletons are loosely Description. Thinly flabellate, convoluted margins, short
arranged more-or-less plumo-reticulate, without apparent axial stalk, firm texture; ectosomal skeleton with thick cortex composed
compression or differentiation between axial and extra-axial of euasters, not detachable, with few thick styles protruding a short
Porifera· Demospongiae • Hadromerida • Hemiasterellidae 191

distance through surface; extra-axial skeleton not well developed De Laubenfels (1936a) included several species of
with few radial megascleres embedded near periphery or forming Hemiasterella in the Coppatiidae (Astrophorida), although
sparse radial tracts ascending from axis; extra-axial region packed Wiedenmayer (1977b) suggested that skeletal architecture is quite
with euasters; axial region loosely reticulate, slightly compressed different between these groups. Nevertheless, there are several
tracts of megascleres without obvious fibre component; megascleres species of Jaspis without obvious radial architecture and with mod-
long, thick, straight or slightly curved, fusiform styles, sometimes ified oxeote megascleres (strongyles, strongyloxeas and quasi-mon-
subtylostyles, usually with rounded bases and sharp points, some- actinal spicules) which may indeed belong to the Hemiasterellidae
times points slightly telescoped (550--1450 X 22-35 11m); microscle- (e.g., Coppatias carteri (Ridley, 1884a» (Hooper, 1986).
res euasters of more than one variety, most commonly with thick, Voultsiadou-Koukoura & Van Soest (1991a) included 10 species
acanthose, strongylote, terminally microspined rays (25--{)0I1m in the genus (H. typus Carter, 1879d from Australia, H. ajfinis Carter,
diameter), smaller euasters with terminal subtylote swellings usually 1879d from unknown locality, H. callocyathus (Sollas, 1888) from
microspined (12-20I1m diameter), and less commonly oxyasters Northern Australia and PNG, H. vasiformis (Kirkpatrick, 1903b)
with small centrum and pointed, smooth rays (8-20 11m diameter). from SE Africa, H. complicata Topsent, 1919 from Madagascar and
Re1TUlrks. Hemiasterella typus has exclusively stylote mega- Indonesia, H. intermedia Dendy, 1922b from W. Indian Ocean and
scleres, whereas the genus currently contains a diverse group of Indonesia, H. elongata Topsent, 1928c from Morocco and
species in which megascleres include monactinal, diactinal spicules, Cape Verde Is, H. digitata Burton, 1929a from Antarctica, H. strongy-
or a combination of both. The architecture is commonly more-or-less lophora Levi, 1956c from Madagascar, and H. aristoteliana
plumo-reticulate, and growth form is exclusively vasiform or cup- Voultsiadou-Koukoura & Van Soest (1991a) from the Aegean Sea).
shaped (Topsent, 1919; Dendy, 1922b). Species, like the type species The inclusion of Tylaspis topsenti Levi & Levi, 1983b, from deeper
with monactinal megascleres, have apparent affinities with Axos and waters off New Caledonia into synonymy with Hemiasterella brings
Timea (Timeidae, Hadromerida), whereas other species with only the total number of described species to 11.
oxeas (e.g., H. ajfinis Carter, 1879d), or both oxeas and styles (e.g.,
H. complicata Topsent, 1919) show superficial similarities to Jaspis Distribution
(Coppatiidae, Astrophorida). For this reason the higher taxonomic
placement of Hemiasterella, and hence Hemiasterellidae, is still Indian and West Pacific Ocean, Antarctica, E Atlantic,
equivocal (e.g., Bergquist, 1968; Wiedenmayer, 1977b; Hooper, E Mediterranean.
1986; Voultsiadou-Koukoura & Van Soest, 1991a).
Epallax and Kalastrella are clearly synonyms of
Hemiasterella. Epallax (type species E. callocyathus Sollas, 1888 PARATIMEA HALLMANN, 1917
(by monotypy) schizotypes BMNH 1894.11.16.480--3) (Fig. 3H)
is vasiform, has diactinal megascleres occassionally modified to Synonymy
quasi-monactinal styles or tylostyles, and an architecture consisting
of longitudinal bundles of megascleres in a more-or-less radial Paratimea Hallmann, 1917c. Halistella Sara, 1958.
plumose arrangement. The type species has, in addition to simple
oxyasters common to other hemiasterellids, branching calthrops- Type species
like asters for which Sollas (1888) inferred parallels with the genus
Thenea Gray (Theneidae, Astrophorida). Kalastrella (type species Bubaris constellata Topsent, 1893c (by original designation).
K. vasiformis Kirkpatrick, 1903b (by monotypy) schizotype BMNH
1902.11.16.6) is vasiform, has both monactinal and diactinallquasi- Definition
diactinal megascleres, and microspined rays on euasters. Skeletal
architecture is more-or-less compressed with interconnected lacu- Hemiasterellidae lacking an axially compressed skeleton,
nae which bear extra-axial small bundles of oxeas and styles pro- having instead a (sub-)hymedesmioid choanosomal skeleton with
jecting at right angles to the ectosome (Kirkpatrick, 1903b), and it long tylostyles erect on the substratum, and bundles of smaller cen-
too was merged with Hemiasterella by Topsent (1919) and Dendy trotylote tomotes or oxeas in both the peripheral and choanosomal
(1921), most closely related to H. complicata Topsent. skeletons; microscleres are smaller thick-centred euasters and
Tylaspis (type species Tylaspis topsenti Levi & Levi, 1983b larger short-rayed oxyspherasters.
(by original designation), holotype MNHNDCL2920 (not seen», is
less confidently included into synonymy with Hemiasterella given Diagnosis
that it has an amorphous massive growth form (unlike the well-
defined vasiform, plate-like or branching growth forms of most of Hemiasterellidae with thickly encrusting growth form and
the described species), and on this basis there may be some justifi- essentially hymedesmioid skeleton arrangement composed of erect
cation for recognising it as a distinct genus. However, it shares with choanosomal tylostyles and paratangential tracts of centrotylote
these 'typical' Hemiasterella species the lack of any axial condensa- ectosomal tomotes, the latter also scattered throughout the skeleton
tion of the choanosomal skeleton, unlike other hemiasterellids, and in dragmata. Microscleres are oxyasters without a prominent
moreover Hemiasterella species do commonly demonstrate centrum and oxyaster euasters with slightly swollen centrum.
intraspecific variability in growth forms. Levi & Levi (1983b) sug-
gested T. topsenti had closest affinities to Jaspis-like taxa (now in Description of type species
Ancorinidae) and Topsent's (1928c) concept of Halicnemia, now
dispersed amongst the hemiasterellids including Stelligera (see Paratimea constellata (Topsent, 1893c) (Fig. 4A-H).
chapter on Desmoxyidae), but Tylaspis appears to be most closely Synonymy. Bubaris constellata Topsent, 1893c: 33;
related to Hemiasterella to which it is here assigned. Paratimea constellata; Hallmann, 1917c: 675.
192 Porifera' Demospongiae • Hadromerida • Hemiasterellidae

IA BI DI

J
Fig. 4. A-H, Paratimea constellata (Topsent) (Mauritania specimen). A, tylostyle base and point (scale 50 ....m). B, tylostyle base (scale SO ....m). C, cen-
trotylote tomote centra (scale SO .... m). D, centratylote tomate (scale 100 .... m). E, oxyaster euasters (scale 10 ....m). F, ectosomal crust of oxyasters (scale
40 ....m). G, peripheral skeleton (scale 2S0 ....m). H, reconstruction of idealised skeletal structure. J-K, Leptosastra constellata Topsent (reproduced from
Topsent, 1904b, pI. 15, fig. 15). I, choanosomal acanthostyle. J, ectosomal 'rhabd'. K, oxyspheraster euasters (for sizes refer to the text).

Material examined. Holotype: MOM (not seen). tylostyles, very long and relatively robust, with swollen terminal
Other material. ZMA POR7348 - Maio I., Cape Verde Is., colI. or subtylote basal swellings (,trilobate' heads according to Topsent),
R.W.M. Van Soest, 40m depth, dredge, 27.viii.86. ZMA and long tapering sharp points (greater than 1 mm-3.5 mm X
POR15292 - Mauritania, dredge. 12-18ILm); tomotes with centrotylote swellings, variable in mor-
Description. Thinly encrusting, conulose and highly hispid phology (Le., multiple tylotes), straight or slightly bent at centre,
surface; ectosomal skeleton with long choanosomal tylostyles occasionally sinuous and in curved dragmata within the skeleton,
protruding more than 1 mm through the surface, often in bundles, with more-or-less symmetrical, sharply-pointed ends (750-1100 X
occasionally singly, and long tomotes in bundles perpendicular, 6-11ILm). Microscleres are oxyasters without a prominent centrum
paratangential or occasionally tangential to the surface; choanosomal (18-251Lm diameter), and oxyaster euasters with slightly swollen cen-
skeleton with erect, paratangential or tangential bundles of trum (14-l9lLm diameter), both with numerous sharply pointed rays.
centrotylote tomotes scattered throughout the skeleton forming Remarks. The original description of B. constellata was
irregular criss-crossing tracts; single or sparse bundles of tylostyles only very brief and lacked illustrations, whereas Topsent (1897b:
are erect or paratangential to the substrate, forming an essentially 245) redescribed this material in more detail and also provided
hymedesmioid skeleton (although this is difficult to observe in (rudimentary) spicule drawings. He again mentioned the species
either available material or published descriptions); the mesohyl is based on new material (Topsent, 1928c: 181), but added little to
abundantly invested with collagen throughout; asters are scattered the original description, whereas the present material (based on
throughout the skeleton although most dense in both peripheral and collections of R.W.M. Van Soest from two disparate localities:
basal parts of the skeleton. Megascleres consist of choanosomal Cape Verde Islands and Mauritania), represents the first record of the
Porifera· Demospongiae • Hadromerida • Hemiasterellidae 193

species from outside the Mediterranean, and subsequent to LEPTOSASTRA TOPSENT, 1904
Topsent's original records. This present material shows some vari-
ability from the holotype having slightly larger asters with a more Synonymy
prominent centrum (Mauritania specimen), and a reduced number
of megascleres (Cape Verde Islands specimen). Nevertheless, Leptosastra Topsent, 1904b: 194; 1928c: 56; de Laubenfels,
megasclere morphology is consistent between all populations, and 1936a: 111.
it is considered here that populations are conspecific. The
redescription above, and the reconstruction of skeletal architecture Type species
(Fig. 4H) is based on both this new material and Topsent's (1897b)
extensive redescription of the holotype given that the ZMA mate- Leptosastra constellata Topsent, 1904b (by monotypy).
rial lacks some detail due to its thinly encrusting nature.
Paratimea differs from most other hemiasterellids in Definition
lacking an axially compressed skeleton, having instead a sub-
hymedesmioid basal skeleton (with predominantly paratangential Hemiasterellidae lacking an axially compressed skeleton,
spicule tracts, and only few true erect spicules on the substrate), having instead a hymedesmioid choanosomal skeleton composed
and in this respect is most similar to, and potentially congeneric of erect acanthostyles, with smooth, very thin subtylote or
with, Leptosastra (coincidentally with the same type species name centrotylote monactinal (or diactinal) 'rhabds' (tomotes), and an
although both clearly different taxa). The two genera are retained ectosomal cortex of oxyspheraster euasters.
separately for the time being given substantially different geome-
tries of their respective choanosomal megascleres: tylostyles Diagnosis
versus acanthostyles (see Leptosastra).
Hallmann (1917c) suggested that P. constellata was similar Very thinly encrusting plaque; ectosomal skeleton with a dense
to Halicnemia in possessing tylostyles and centrotylote diactinal crust of spheraster euasters and reinforced with sparse bundles of
spicules forming the ectosomal skeleton, and were it not for the pres- very thin monactinal (or occasionally diactinal) 'rhabds' (tomotes)
ence of these ectosomal spicules he would have assigned the taxon below this crust; choanosomal skeleton hymedesmioid with acan-
to Timea (Timeidae). He tentatively included it in the Spirastrellidae. thostyles embedded in basal spongin and erect on the substratum.
Hooper (1986) suggested that it might be an encrusting form of
Stelligera, as judged from its relatively poor published description, Description of type species
given that their respective type species were superficially similar in
aster morphology, whereas Voultsiadou-Koukoura & Van Soest Leptosastra constellata Topsent, 1904b (Fig. 4I-K).
(1991a) indicated that it may be a valid taxon. These latter authors Synonymy. Leptosastra constellata Topsent, 1904b:
assigned five species to the genus: Bubaris constellata Topsent 194-195, pI. 15, fig. 15; Stephens, 1921: 63.
(from the Mediterranean (plus the two new locality records noted Material examined. None - Azores.
here», P. galaxea de Laubenfels (1936a) (from Florida), Halicnemia Description (from Topsent, 1904b). Thinly encrusting form-
duplex (Topsent, 1927b) (from Morocco), P. pierantonii Sara, 1958 ing an extensive plaque, colour white, smooth surface; choanosomal
and P. oxeata Pulitzer-Finali, 1977 (both from Naples). To this list skeleton composed of acanthostyles erect on (echinating) the substra-
should be added Halistella pierantonii Sara, 1958, the type species tum, with their bases embedded in basal spongin and points supporting
of the genus Halistella, which is clearly also a Paratimea. the ectosomal skeleton; subectosomal and ectosomal regions rein-
The higher taxonomic placement of Paratimea has been in forced with tomotes (described as 'rhabds'); ectosome with a thick
question for some time. Hallmann (1917c) proposed to combine crust of euasters confined entirely to the outer surface, through which
the aster-bearing axinellids, including Halicnemia Bowerbank, sparse bundles of tomotes occasionally protrude. Choanosomal acan-
Laonoenia Hallmann and Paratimea Hallmann (all possessing thostyles heavily and evenly spined, in a single size class, with straight
choanosomal tylostyles and ectosomal centrotylote oxeas) within shaft, swollen tyle and sharply pointed (80-100 X 7 f,Lm). Ectosomal
the Desmoxyidae (which are otherwise defined by their acanthose and subectosomal 'rhabds' occur in restricted numbers, embedded in
microxeote ectosomal spicules). Topsent (1928c) rejected this sys- small groups in the mass of euasters, having a straight smooth shaft,
tem, as well as Hallmann's genera (including Paratimea), retaining asymmetrical ends (defined as either monactinal or diactinal, although
these taxa within a single genus (Halicnemia). However, in the strict only the former are figured by Topsent), and a subtylote swelling
sense Halicnemia (viz., H. patera Bowerbank, H. gallica (Topsent) closer to the basal end than to the point of the spicule, occasionally poly-
and H. geniculata Sara) has tylostyles, ectosomal oxeas (often tylote (135 X 2f,Lm). Euasters are spherasters with numerous conical,
polytylote), and acanthose microxeas (Bowerbank, 1864, 1866; pointed actines and a thick centrum (20-23 f,Lm diameter).
Topsent, 1897b; Descatoire, 1966), and clearly belongs with the Remarks. Leptosastra is monotypic. Despite having type
Desmoxyidae (Levi, 1973), closely related to Higginsia. By compar- species with identical names Leptosastra and Paratimea are clearly
ison, Halicnemia sensu Hallmann is polyphyletic, with one group not identical, differing substantially in the geometry of their principal
belonging to the Astrophorida, and a second group (viz., Paratimea (choanosomal) structural megascleres - acanthostyles and tylostyles,
s.s., and including H. duplex Topsent, H. constellata (Topsent), and respectively. Whether these differences can sustain these genera sepa-
Paratimea galaxea de Laubenfels) having euasters, centrotylote rately remains uncertain, but re-examination of the holotype of the
oxeas and choanosomal tylotes, belonging to the Hemiasterellidae. former is essential to such a decision (presumably located in the
MOM, but not confirmed). Nevertheless, the current decision to not
Distribution amalgamate both genera is supported by the nature (and possibly ori-
gin) of the subectosomall ectosomal megascleres, which are defined
Mediterranean, NE and central E Atlantic, Caribbean. as 'rhabds' and tomotes, respectively. Both can be potentially defined
194 Porifera' Demospongiae • Hadromerida • Hemiasterellidae

ABC

Fig. 5. A-G. Stelligera stuposa (Montagu) (schizotype). A. choanosomal styles (scale 150 !Lm). B, ectosomal styles (scale 150 !Lm). C, extra-axial styles
(scale 150 !-Lm). D, euasters (scale 10 !-Lm). E, peripheral skeleton (scale 250 !-Lm). F, axial skeleton (scale 250 !-Lm). G, reconstruction of idealised skeletal
structure. H, Spongia stuposa Montagu, 1818 (from his plate 3, fig. I).

as tomotes, whereas those of Leptosastra are significantly thinner and STELUGERA GRAY, 1867
shorter than those of Paratimea that they may indeed be microscleres,
and hence warranting description as 'rhabds' by Topsent (1904b). Synonymy
The type species of Leptosastra was initially allocated close to
the hymedesmiids, such as Leptosia, based on its possession of Stelligera Gray, 1867a. Vibulinus Gray, 1867a. Taxonomic
a hymedesmioid skeleton composed of acanthostyles, with geometry decision for synonymy: Voultsiadou-Koukoura & Van Soest
resembling echinating acanthostyles found in many poecilosclerid (1991a).
families. However, Topsent (1940b) reported that the species had
euasters forming a distinct, thick cortex. If this is true then it negates Type species
any such phylogenetic interpretation, with the genus more appropri-
ately allocated to the present concept of Hemiasterellidae. It is possi- Raspailia stelligera Schmidt, 1862 (by original designation)
ble, but not yet proven, that L constellata belongs to Hymedesmiidae, (junior synonym of Spongia stuposa Montagu, 1818; Voultsiadou-
with its reported euasters potentially representing modified acan- Koukoura & Van Soest, 1991a).
thostyles (as suggested by Dendy, 1922b) or chelae (with potential
affinities to Spirorhabdia or AcantJuzncora). Correct allocation of this Definition
genus requires re-examination of the presently elusive holotype.'
Hemiasterellidae with several distinct structural regions:
Distribution a slightly compressed axial skeleton of longer styles and oxeas;
a reticulate extra-axial skeleton of smaller styles and/or oxeas;
Known only from the North Atlantic. longer extra-axial styles radiating from the axis to periphery; and
Porifera· Demospongiae • Hadromerida • Hemiasterellidae 195

an ectosomal skeleton of small styles surrounding the longer hispid surface; choanosomal skeleton with slightly compressed axis
projecting extra-axial styles; microscleres are short-rayed composed of thinner styles and oxeas, and with extra-axial skeleton
oxyspheraster euasters with a thick centrum and forming a thin divided into two components, one composed of shorter axial
ectosomal crust. styles/oxeas forming irregular reticulation throughout mesohyl, the
other composed of longer extra-axial styles radiating from axis to
Diagnosis periphery; megascleres are long, thick, straight or slightly curved
styles of the extra-axial skeleton, with rounded or slightly telescoped
Thinly branching growth form and prominently hispid sur- points and evenly rounded bases (1200-2850X 15-25 fLm), shorter,
face; choanosomal skeleton composed of long styles and oxeas thick styles, stongylote styles, or sometimes oxeas, slightly curved,
forming a slightly compressed axis; extra-axial skeleton with radial with rounded bases and strongylote or fusiform points (550-950 X
and reticulate components, the former composed of long projecting 8-13 fLm), ectosomal megascleres are short, thin styles or anisoxeas,
styles perpendicular to axis, the latter consisting of shorter curved, often sinuous, occasionally rhaphidiform (380-720 X 3-8
styles/oxeas forming an irregular reticulation; ectosomal skeleton fLm); microscleres short-rayed oxyspheraster euasters with entirely
composed of brushes of small styles surrounding protruding smooth rays and thick centrum (8-22 fLm).
extra-axial styles (analogous to the Raspailiidae ectosomal condi- Remarks. Stelligera is arborescent to lobodigitate, occasion-
tion), and also with a thin ectosomal crust of thick-centred, short- ally slightly flabelliform, and has both oxeas and styles (or
rayed oxyspheraster euasters. tylostyles) as megascleres and smooth euasters as microscleres.
Re-examination of type material supports the proposal by
Description of type species Voultsiadou-Koukoura & Van Soest (1991a) that S. stelligera and
S. stuposa are synonyms (possibly together with S. rigida (Montagu,
Stelligera stuposa (montagu, 1818) (Fig. 5A-H). 1818) from the British Isles and S. nux Lendenfeld, 1897a from the
Synonymy. Raspailia stelligera Schmidt, 1862: 60; Spongia Mediterranean, although these types have not yet been redescribed),
stuposa Montagu, 1818: 79. making Vibulinus (with type species Spongia stuposa Montagu,
Material examined. Holotype: SM not seen, schizotype 1818 (by monotypy) schizotypes BMNH 1910.1.1.2463-4) an
MNHN DCL1239L - Quamero, Adriatic. Other material. objective synonym of Stelligera. This leaves only two valid species
Schizotypes of S. stuposa: BMNH 1910.1.1.2463-5 - British Isles. in the genus (the type species from the British Isles, Mediterranean
Description. Thinly cylindrical, dichotomously branching and Adriatic, and S. columnata Levi, 1959 from W Africa).
growth form with prominently hispid surface; ectosomal skeleton with
thin crust of euasters, through which protrude long extra-axial styles Distribution
surrounded by bouquets of thin ectosomal styles tangential to the sur-
face; megascleres protrude up to 400 fLm from surface, producing East Atlantic, Mediterranean.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John NA. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Placospongiidae Gray, 1867

Klaus Riitzler

Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, DC, 20560-0163, USA.
(ruetzler.klaus@nmnh.si.edu)

Placospongiidae Gray (Demospongiae, Hadromerida) has the typical radial skeleton structure of Hadromerida and Spirastrella-like
tylostyle-spiraster spicule complement but stands out by the possession of a rigidly cemented cortex of large microscleres. The cortical
spicules are ovoid or bean-shaped (selenasters) or dumbbell-like (amphinolasters). The cortex is broken up into distinctive plates that are
separated by contractible grooves containing oscula and ostia. Species are encrusting or branching massive and most occur in shallow
tropical and subtropical waters, from the intertidal to almost 200 m. The family, until recently considered monotypic, includes two genera,
Placospongia (including Physcaphora) and Onotoa (including Amphinolana).
Keywords: Porifera; Demospongiae; Hadromerida; Placospongiidae; Onotoa; Placospongia.

DEFINITION, DIAGNOSIS, SCOPE as Schmidt (1870), Sollas (1888), Hanitsch (1895), Minchin
(1900), and Topsent (1933) believed in the close relationship of
Synonymy Placospongia then the only genus known in the family, and
Geodia, although, as Keller (1891) demonstrated, the
Placospongiidae Gray, 1867b: 127 (Gray also applied the Placospongia 'sterraster' actually develops from a spiraster, origi-
common name 'Stony Sponges'). nating from a rod-shaped center (Vosmaer & Vernhout, 1902),
whereas Geodia sterraster starts as an euaster (from one center
Definition point).
Species of Placospongia may be encrusting or branching; in
Hadromerida with cortex and selenasters or amphinolasters the latter case, the selenasters of the surface crust form also a dense
(amphispherasters) as cortical spicules. axial tract. The color is generally brown to tan (tan to white when
preserved or dry) but purplish-red specimens are known where the
Diagnosis (from Rutzler & Hooper, 2000) pigment resides in the selenasters and is not dissolved in preserva-
tion alcohol (Vosmaer & Vemhout, 1902).
Hadromerida with tylostyles as megascleres and selenasters Species in this family occur in tropical or subtropical shallow
or amphiaster-like sterrasters (amphinolasters) as primary water (from intertidal to 117 m depth), particularly on coral reefs.
microscleres forming polygonal cortical crusts. Cortical plates sep- Small crusts occur on the lower surfaces of intertidal rock and coral
arated by contractile ectosomal pore grooves bearing ostia and rubble where they can survive periods offalling dry by contracting
oscula. Tylostyles in tracts radiating from the base toward the sur- the unprotected surface grooves; epizoic algae and invertebrates
face and supporting the margins of the cortical plates. Accessory are common.
microscleres include spirasters, spherasters, and spherules.
Remarks
Scope
The distinction between Spirastrellidae, Clionaidae, and
1\\'0 valid genera, Onotoa de Laubenfels and Placospongia Placospongiidae in this revision is further supported by 28S-
Gray. genetic analysis (Chombard, 1998).
Developmental stages of selenasters and amphinolasters,
History and biology spicules with fused rays similar to the sterraster of Geodiidae,
show their relationship to both spirasters and amphiasters.
Because fully formed selenasters are similar to sterrasters in Associated typical spirasters indicate kinship with Spirastrellidae
shape and and also form a strong cortex, several older authors, such and Clionidae.

KEY TO GENERA

(1) Microscleres ofthe cortex are ovoid or bean-shaped selenasters ................ ...... .................................. ...... .......... ...... ..... Placospongia
Microscleres of the cortex are dumbbell-like amphispherasters ............................................................................................... Onotoa

196
Porifera· Demospongiae • Hadromerida • PJacospongiidae 197

o __
,,
,

I
B c_ E F_
Fig. 1. Onotoa amphiastra de Laubenfels, holotype USNM 35385. A, specimen detail showing the cortical plates bounded by ridges (closed grooves)
(scale 10 mm). B, cross-section demonstrating skeleton structure: Cortex of cemented amphinolasters and tylostyle bundles supporting cortical plates along
the aquiferous groves (scale 50 !Lm). C, tylostyles (scale 50 !Lm). D, large spirasters and amphiasters (scale 10 !Lm). E, microspined spirasters (scale
10 !Lm). F, amphinolasters, stages of development (scale 2 !Lm).

ONOTOA DE LAUBENFELS, 1955 Description of type species

Synonymy Onotoa amphiastra de Laubenfels, 1955a (Figs 1-2).

Synonymy. Onotoa amphiastra de Laubenfels, 1955a: 140,
Onotoa de Laubenfels, 1955a: 140. Amphinolana Riitzler & fig. 1.
Hooper, 2000: 339. Material examined. Holotype: USNM 35385 (de
Laubenfels' original number: 52003) - Onotoa Atoll, Gilbert
Type species Islands (ca. 1 50'S, 176°30'E).
0

Description. Small, hard and tough crust (37 x 20 mm),

Onotoa amphiastra de Laubenfels, 1955a (by original pale tan (alcohol), with polygonal cortical plates separated by
designation). ridges (contracted grooves). Fascicular tracts of tylostyles running
perpendicularly from substrate to a cortex composed of rigidly
Definition cemented amphinolasters. The choanosome contains scattered
microscleres of three types: Large spirasters and amphiasters with
Placospongiidae with a cortex of dumbbell-like thin axis and long, slender spines, some branched into secondary
amphispherasters (amphinolasters). spines near the end (called streptasters by de Laubenfels, 1955a);
tiny micro-spined spirasters with 1-2 bends (not mentioned by the
Diagnosis original author), and developmental stages of amphinolasters.
Spicule dimensions (range and means of 10 measurements, unless
Crustose sponges with cortical plates separated by grooves. otherwise indicated): Tylostyles, straight with spherical head that is
Tracts of megasc1eres (tylostyles) radiating from the base toward only slightly thicker than the shaft, and most with rounded point
the surface. Stony hard cortex and basal spicule layer formed by (length X maximum width, head width), 570-820 X 8-13, 10-15
unusual, cemented microsc1eres, barbell-shaped amphispherasters j.l.m (716.0 X 10.5, 13.5j.l.m), smaller sizes occur in the cortical
(or amphisterrasters). This spicule type was termed amphinolaster zone but do not form a separate size class; amphinolasters, dumb-
(Riitzler & Hooper, 2000) to contrast the selenaster of bell-shaped and covered by low, broad-based spines (length X
Placospongia, a spiraster derivative (Vosmaer & Vernhout, 1902). maximum diameter, shaft diameter), 58-70 X 25-38, 10-20 f.1m
Accessory microspined spirasters and long-rayed amphiasters (63.7 X 31.2, 14.2 f.1m); immature amphinolasters with thin spines,
(only in the type species) are microscleres found mainly in the 30-50 X 5-15, 1-4 f.1m (40.6 X 10.5, 2.8 f.1m); large amphiasters
choanosome. Two species from intertidal habitats in the Pacific and spirasters with long slender spines, 24-38 X 22-25, 3-4 f.1m
Ocean. (30.8 X 24.0, 3.8 f.1m); and microspined spirasters (length X width,
n=7), 6-14 X 2-3 f.1m (11.7 X 2.7 f.1m). Distribution: Tropical
Previous review Pacific Ocean.
Remarks. Only one specimen of Onotoa amphiastra was
None; but detailed description of a representative of the ever found and the species was all but forgotten until R. W.M. Van
genus, Amphinolana (=Onotoa) claudelevii, in Riitzler & Hooper Soest (pers. comm.) pointed out its description by de Laubenfels
(2000). (1955a: 137) who stated that the holotype was sent to USNM. After
198 Porifera· Demospongiae • Hadromerida • Placospongiidae

Fig.2. Onatoa spicules, SEM images. A, C-F, O. amphiastra de Laubenfels.; B, O. claudelevii (Riltzler & Hooper). A, tylostyle heads and amphinolasters
(scale SO !Lm). B, amphinolasters and (adhering to tylostyie shaft) microspined spirasters. C-D, developing amphinolasters. E, large spiraster (streptaster).
F, microspined spiraster (scales B-F, 10 !Lm).

lengthy search, the specimen with de Laubenfels' original number Diagnosis

was indeed found among unidentified Pacific sponges.
The early stages of amphinolasters of Onotoa, which are Encrusting or lobate and branching sponges covered by
formed in the choanosome, are very similar to undeveloped smooth cortical plates separated by contractible grooves. Size from
Placospongia selenasters. Both start as spiny microrhabds, spines small crusts (average 25cm2 , 2mm thick) to erect branches (36cm
longer toward the distal parts, but the shafts of the former stay tall, 1.5 cm and more in diameter). Live color of known species red-
straight whereas those of the latter bend gradually until fusion of dish brown to almost black. Skeleton structure radiate, tylostyle
the two heads. Both genera have small accessory spirasters. bundles running from a basal crust or a central axis (branching
specimens) of selenasters toward the cortex and supporting the mar-
gins of cortical plates. Megascleres are tylostyles of two size classes
PLACOSPONGIA GRAY, 1867 (the shorter ones in the ectosome), microscleres include selenasters
of the cortex and base layer (or central axis) and choanosomal and
Synonymy ectosomal spirasters, spherasters, and spherules; developmental
stages of selenasters occur throughout the choanosome.
Placospongia Gray, 1867b: 128. Physcaphora Hanitsch, 1895.
Previous review
Type species
Vosmaer & Vemhout, 1902.
Placospongia melobesioides Gray, 1867b (by original
designation). Description of type species

Definition Placospongia melobesioides Gray, 1867b (Figs 3, 4).

Synonymy. Placospongia melobesioides Gray, 1867b: 128,
Placospongiidae with selenasters as cortical spicules. figs 1-4.
Porifera· Demospongiae • Hadromerida • PJacospongiidae 199

A B c

1
F

.
~.
. 'i7....

G __ II

H __
E
Fig. 3. A-G, Placospongia melobesioides Gray, holotype BMNH 52.4.1.14. H, P. carinata (Bowerbank) (B, C, H, after Vosmaer & Vernhout, 1902).
A, habitus of fragment, USNM 39649, Gulf of Mexico, 76m (scale 20mm). B, cortical plates and grooves (scale 5mm). C, cross-section of branch show-
ing anatomy: selenasters forming central core and exterior cortex, and tylostyle tracts (scale I mm). D, tylostyles (scales 100 J,J.m, 200 J,J.m for magnified
ends). E, selenasters and stages of development (scale 10 J,J.m). F, spheraster (scale 5 J,J.m). G, spheres (scale 2 J,J.m). H, large spirasters and amphiasters
characteristic of P. carinata (scale 10 J,J.m).

Material examined. Holotype: BMNH 52.4.1.14 (2 dry (963.0 X 14.1, 17.0fLm), (IT) 350--560 X 8-10, 9-13fLm (438.8 X
fragments, 1 slide) - Borneo. 9.1, 10.5 fLm), smaller sizes occur in the cortical zone; selenasters,
Description. Chalky white, stone-hard (dry) fragments (60 X bean-shaped to oval, 38-48 X 25-38 fLm, 42.3 X 30.1 fLm); imma-
22 mm, 54 X 37 mm) of a branching specimen. Cortical plates make ture selenasters, 25-33 X 13-20 fLm (28.4 X 16.4 fLm); spherasters
up the outermost layer. Tylostyle bundles radiate from an axis of (diameter, n=5), 14-16 fLm (15.2 fLm); and spherules (diameter,
packed selenasters to the cortical plates that are also made up by the n=5), 1-3 fLm (1.7/-Lm). Distribution: Tropical Atlantic and
characteristic microscleres. Spicules (dimensions are ranges and Indopacific.
means of 10 measurements unless otherwise indicated): Tylostyles, Remarks. Placospongia melobesioides lacks spirasters
straight with spherical head and most with rounded point (length X (slender-spined streptasters as well as microspined rhabds) that are
maximum width, head width), (I) 720--1200 X 13-15, 15-18 fLm common in P. carinata (Bowerbank). According to Vosmaer &
200 Porifera· Demospongiae • Hadromerida • Placospongiidae

Fig. 4. Placospongia melobesioides Gray, SEM images of spicules, holotype BMNH 52.4.1.14. A, tylostyle and developing stages of selenasters (scale
50 j.Lm). B-C, selenasters (scales: B, 50 j.Lm; C, 10 j.Lm). D, spheraster (scale \0 j.Lm). E, spheres (scale 5 j.Lm).

Vernhout (1902: 9), only three of the six species of Placospongia 205; including synonym P. graeffei Lendenfeld, 1897b: 48).
described up to that year could be recognized as valid: the type Subsequently, one more species was described, P. cristata Boury-
species Placospongia melobesioides Gray, P. carinata (Bowerbank) Esnault, 1973: 276, which was distinguished by the possession of
(Geodia carinata Bowerbank, 1858: 308; including synonyms P. small spherasters and the absence of spirasters, both characteristics of
intermedia Sollas, 1888: 272 and P. mixta Thiele, 1900: 72), and P. P. melobesioides; hence, it may be a junior synonym of the latter
decorticans (Hanitsch) (Physcaphora decorticans Hanitsch 1895: species.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademicIPlenum Publishers, New York, 2002

Family Polymastiidae Gray, 1867

Nicole Boury-Esnault

Universite de la Mediterranee, Centre d'Oceanologie de Marseille, Station marine d'Endoume, UMR-CNRS 6540, rue de la Batterie des
Lions, 13007-Marseille, France. (esnault@com.univ-mrs.fr)

Polymastiidae Gray (Demospongiae: Hadromerida) contains 14 genera and approximately 100 species. Species live in all oceans, but
some genera are found predominantly in shallow waters at high latitude and in deep waters elsewhere. They are often globular, cushion
shape or discoid. The genera of this family share a radiating choanoskeleton and a more or less complicated cortex, the outer layer being
always a palisade of ectosomal spicules.
Keywords: Porifera; Demospongiae; Hadromerida; Polymastiidae; Polymastia; Atergia; Acanthopolymastia; Proteleia; Pseudotrachya;
Quasillina; Radiella; Ridleia; Sphaerotylus; Spinularia; Trachyteleia; Tentorium; Tylexocladus; Weberella.

DIAGNOSIS & SCOPE valid. More than one hundred species have been published world-
wide although many of them may be synonyms. The faunas of
Synonymy the Atlantic and deep polar regions are relatively well known,
including northern New Zealand, the North Sea, and the
Polymastiidae Gray, 1867a: 527; Topsent, 1900: 131; Wilson, Mediterranean in particular, but other faunas are relatively poorly
1925: 347; Boury-Esnault, 1987: 29; Kelly-Borges & Bergquist, known.
1997: 400.

Definition History and biology

Hadromerida with a radiating choanoskeleton and a more or The anatomy and cytology have been described only in some
less complicated cortex, the outer layer being always a palisade of species of Polymastia (Boury-Esnault, 1974, 1976; Boury-Esnault
ectosomal spicules (tylostyles, or oxeas and/or exotyles). Mega- et al., 1994a). The volume of choanocyte chambers is between
scleres are tylostyles, subtylostyles, strongyloxeas, styles or oxeas; 3400-7800/-lm3 which is within the range observed for other
microscleres may include centrotylote microxeas, acanthose Hadromerida, and the choanocytes show a periflagellar sleeve as in
microxeas, or raphides in trichodragmata. many other hadromerids (Boury-Esnault et al., 1990). Different
types of cells with inclusions are present which differ between
Diagnosis species and can help to discriminate them. However, too few
cytological studies have been conducted on Polymastiidae to use
Hadromerida of massive, encrusting, globular, discate or cytological characters at the family level of taxonomy.
pedunculate growth form. Surface slightly velvety or very hispid. Reproduction is known only in the genus Polymastia (Fig. 1). They
The choanoskeleton is constituted by radial megasclere tracts. are oviparous and the eggs stay in a thick mucus, where the cleav-
A complex specialised ectosomal skeleton is developed to a greater age occurs, after leaving the parent body. The mature larva is a flat-
or lesser degree in different genera. It is composed of at least a pal- tened blastula which is benthic and slithers along the substrata
isade of tylostyles, or oxeas and/or exotyles (cladotylostyles, spher- (Borojevic, 1967c). A case of asexual reproduction through bud-
ostyles, etc.). Spicules are two or three size categories of tylostyles, ding of papillae has been also reported for Polymastia arctica
subtylostyles, strongyloxeas, styles or oxeas. Free spiCUles are (Merejkowsky, 1878). This phenomenon occurs quite frequently in
always present in the choanosome which may be intermediary or this species and has been redescribed recently by Plotkin &
ectosomal tylostyles as well as smooth centrotylote microxeas, Ereskovsky (1997).
acanthose microxeas, or raphides in trichodragmata. A fringe of
long spicules is often present at the edge of the body at the contact
with the substrata. When known, reproduction is oviparous, extru- Remarks
sion of oocytes and sperm occurs in the sea, and after cleavage of
the egg a flattened blastula larvae is developed. Asexual reproduc- This family was erected by Gray, (1867a: 527) mostly for the
tion by the production of stalked bead-like buds has been observed. genus Polymastia such that the definition given by Gray corre-
Several genera are only found in very deep and or polar waters. sponds to the definition of Polymastia. Subsequently there has
been a widespread consensus about the content of Polymastiidae
Scope such that since Wilson (1925: 347) generic composition has not
changed very much, even if some authors like Dendy (1922b: 147)
Twenty-six nominal genera have been included in the family and de Laubenfels (1936a: 151) combined Suberitidae with
at one time or another but only 14 are now considered to be Polymastiidae.

201
202 Porifera' Demospongiae • Hadromerida • Polymastiidae

A - B -

c D
Fig. 1. Polymastia robusta, reproduction (R. Borojevic). A, egg (scale 25.3 /-Lm). B, two blastomere embryo (scale 28.3 /-Lm). C, four blastomere embryo
(scale 26.4 /-Lm) . D, free larva (scale 30 /-Lm).
-
KEY TO GENERA

(1) Polymastiidae with exotyles .............................................................................................................................................................. 2

Polymastiidae without exotyles ......................................................................................................................................................... 3
(2) Polymastiidae with grapnel-like spicules ............................................................................................................................. Proteleia
Polymastiidae with spherotylostyles .............................................................................................................................. Sphaerotylus
Polymastiidae with cladotylostyles ................................................................................................................................ Tylexocladus
Polymastiidae with acanthose tylostyles ......................................................................................................................... Trachyteleia
(3) Polymastiidae with the outer layer of the ectosomal skeleton constituted by a palisade of tylostyles ............................................. 4
Polymastiidae with the outer layer of the ectosomal skeleton constituted by a palisade of oxeas .............................. Pseudotrachya
(4) Free spicules of the choanoskeleton are smooth centroty lote oxeas, acanthose rnicroxeas or raphides in trichodragmata ....... ....... 5
Free spicules of the choanoskeleton are ectosomal or intermediary tylostyles or subtylostyles ....................................................... 6
(5) Free spicules are smooth centrotylote rnicroxeas .................................................................................................................... Atergia
Free spicules are acanthose rnicroxeas ................................................................................................................. Acanthopolymastia
Free spicules are raphides in trichodragmata ..................................................................................................................... Spinularia
(6) Polymastiidae with reduced choanoskeleton ..................................................................................................................................... 7
Polymastiidae with developed choanoskeleton .................................................................................................................................. 8
(7) Choanoskeleton is reduced to bundles of free tylostyles ................................................................................................... Quasillina
Choanoskeleton is reduced to a layer of tangential tylostyles in its outer part ........................................................................ Ridleia
(8) Choanoskeleton is radial .................................................................................................................................................................... 9
Choanoskeleton is reticulate ................................................................................................................................................ Weberella
(9) The ectosomal skeleton is constituted by a single layer of tylostyles perpendicular to the surface ................................... Tentorium
The ectosomal skeleton is constituted by at least two layers ........................................................................................................... 10
(10) The ectosomal skeleton from the upper part is different from the one of the lateral or lower part .......... .................... ......... Radiella
The ectosomal skeleton is identical on the whole surface ................................................................................................. Polymastia
Porifera· Demospongiae • Hadromerida • Polymastiidae 203

Fig.2. Polymastia. A, thick longitudinal section through a papilla of a Swedish specimen of Polymastia mamillaris (abbreviations: P, papilla; E, ectosomal
skeleton) (scale 340 )J.m). B, thick perpendicular section through the body of type specimen of Polymastia mamillaris (abbreviations: E, ectosomal skele-
ton; I, intermediary layer of the ectosomal skeleton; F, tracts of principal spicules; the arrow indicates the collagenous layer) (scale 340 )J.m).

POLYMASTIA BOWERBANK, 1864 MUller, 1806 with S. penicillus Montagu, 1818. Bowerbank (1864)
followed this view and seems to have ignored the description of
Synonymy MUller as there is no mention of this author in any text of
Bowerbank (1864, 1866, 1872b) concerning Polymastia mamil-
Polymastia Bowerbank, 1864: 177. Rinalda Schmidt, 1870: 51; laris. Vosmaer (1882a), followed by Levinsen (1887) and Fristedt
Morrow & Boury-Esnault, 2000: 329.Pencillaria Gray, 1867a: 527. (1887), considered both species to be distinct mainly because of
the presence of very long spiCUles around the margin of specimens
Type species of 'penicillus'. In fact, as stressed by Topsent (1900), they had con-
fused this species with Trichostemma hemisphaericum Sars, 1872
Spongia mamillaris Miiller, 1806 (by monotypy). while the description of 'penicillus' by Montagu is clearly not a
Trichostemma.
Diagnosis
Distribution
Thickly encrusting Polymastiidae, spherical or cushion-
shaped, always with papillae. Skeleton composed of radial tracts of About 50 species of Polymastia have been described in the lit-
principal spicules between which free spicules are scattered. erature. The Atlantic (Boury-Esnault, 1987; Boury-Esnault et al.,
Ectosomal skeleton composed of at least two layers, the superficial 1994b; Morrow & Boury-Esnault, 2000) and New-Zealand regions
one is a palisade of small tylostyles, the lower layer is made of (Kelly-Borges & Bergquist, 1997) are the best studied and each
intermediary spicules, tangential to the surface. The principal containing about 18 species. Several species such as Polymastia
spicules can be tylostyles, subtylostyles, styles, and strongyloxeas, mamillaris have been cited worldwide but it has been demonstrated
intermediary spicules are most often tylostyles, and ectosomal that this cosmopolitanism is based on weak taxonomy (Boury-
spicules are always tylostyles. Esnault et al., 1994a).

Remarks Description of type species

Polymastia was erected by Bowerbank in 1864. He designated Polymastia mamillaris (Miiller, 1806) (Fig. 2).
Halichondria mamillaris 'Johnston, 1842' as the type species con- Synonymy. Spongia mamillaris MUller, 1806: 44;
sidering it to be 'the best type of Polymastia'. In fact 'mamillaris' Bowerbank, 1864: 177; Morrow & Boury-Esnault, 2000:
was not described by Johnston but by MUller (1806) under the name 329-330.
Spongia mamillaris. Johnston (1842) transferred 'mamillaris' from Material examined. Holotype: ZMUC '2.1.93' - BohnsHin,
Spongia to Halichondria and also synonymised S. mamillaris North-Atlantic, Swedish west coast, 58°15'N, llo50'E. Other
204 Porifera· Demospongiae • Hadromerida • Polymastiidae

material. ZMUC - various material, Swedish west coast, colI. PROTELEIA DENDY & RIDLEY, 1886
Ole Tendal, Bjorms Rev., l20-200m, 11.07.1975; Ulvillarna,
150-225m, 21.03.1977; East of Rams6, 99-108m, 16.05.1974; Synonymy
Sacken, 76m, 13.05.1974; Sacken, 80-90m, 16.10.1974.
Description. The type specimen is a fragment of a cushion- Proteleia Dendy & Ridley, 1886: 152; Ridley & Dendy, 1887:
shaped, attached sponge approximately 35 X 18 X 7 mm thick. The 214; Lendenfeld, 1903: 29.
upper surface is hispid. The surface of the sponge traps silt and the
colour is only discernible on the papillae. The color is cream in Type species
alcohol. The specimen has 26 inhalant papillae and one exhalant.
The mean length of the inhalant papillae is 8 X 2 mm in diameter. Proteleia sollasi Dendy & Ridley, 1886 (by monotypy).
The exhalant papillae is 11 mm long and approximately 4 mm in
diameter. The ectosomal skeleton (Fig. 2) is about 400!Lm thick Diagnosis
and composed of three layers: the upper layer is a dense palisade
(-300 !Lm thick) of fusiform tylostyles, the middle layer is collage- Disc-shaped Polymastiidae with aquiferous papillae. The
nous (-20!Lm thick) and the lower layer is a tangential layer ectosomal skeleton consists of a palisade of two layers of tylostyles
(-80 !Lm thick) made of intermediary spicules. Choanosomal tracts which lie on a tangential layer of intermediary tylostyles. The ecto-
of principal spicules are 53-1 06!Lm thick. These tracts are divided somal skeleton is echinated by grapnel-like spicules (exotyles).
into two to three smaller ones below the ectosome. They cross the The choanosomal skeleton is composed of stout fibres of principal
ectosome and echinate the surface by approximately 875!Lm. fusiform styles which end in a palisade. Between the fibres groups
Groups from two to five ectosomal spicules are scattered between of grapnel-like spicules are visible. Besides the grapnel-like
the choanosomal tracts. They are particularly abundant below the spicules three types of tylostyles and styles are present.
tangential layer of intermediary spicules in a layer approximately
500!Lm thick. The skeleton of the inhalant papillae consists of Remarks
ascending multispicular megasclere tracts that run the length of the
papillae. These are supported by a network of intermediary Three valid species have been described in this genus: Proteleia
tylostyles arranged perpendicularly to the megasclere tracts. sollasi, P. borealis Swartschewsky, 1906, and P. burtoni Koltun,
Towards the periphery there is a layer of tangentially arranged 1964b: 28. Tetilla truncata Topsent, 1890b: 13 is here excluded as it
intermediary tylostyles and external to this a palisade composed was allocated erroneously to Proteleia by Lendenfeld, 1903: 29.
of ectosomal tylostyles (Fig. 2A). Ectosomal tylostyles are
fusiform, straight or slightly bent with a well marked head: Distribution
143-196 X 5.3-16!Lm (mean l69X 1O.6!Lm, N>50). Intermedi-
ary styles or subtylostyles, straight 243-561 X 8-15.9!Lm (mean The genus has a bi-polar distribution and wide bathymetric
445 X 13.2!Lm, N > 50). Principal spicules are straight, fusiform distribution from 18-3000m.
strongyloxeas, 742-1378 X 8-32!Lm (mean 1052 X 24.5 !Lm
N>50). Description of type species
Remarks. Specimens of Polymastia from the Swedish west
coast collected between 76 and 225 m deep correspond to the form Proteleia sollasi Dendy & Ridley, 1886 (Fig. 3).
described by MUller. These show the following characters: ectoso- Synonymy. Proteleia sollasi Dendy & Ridley, 1886: 152;
mal skeleton composed of three layers, groups from 2-5 ectosomal Ridley & Dendy, 1887: 214.
spicules scattered between the choanosomal tracts and particularly Material examined. Holotype: BMNH-1887.5:2.62 -
abundant below the tangential layer of the ectosome, shape of the Simon's Bay, near Cape of Good Hope, South Atlantic, 18-37 m.
ectosomal tylostyles (101-182X5.2-11.7, mean 148X8.4!Lm, Description. The holotype is 63 mm long, 31 mm wide and
N > 50) and of the principal choanosomal spicules (strongyloxeas about 13 mm in the thicker part. It is disc-shaped and had aquiferous
461-1320 X 10.6-26.5, mean 853 X 20 !Lm, N > 50) (Morrow & papillae which can reach 8 mm long and 4 mm in diameter at the base.
Boury-Esnault, 2000). Colour in alcohol is yellowish-grey. Surface is minutely hispid and
Conversely, the holotype of Polymastia mamillaris does not considerable amount of foreign matter is trapped in the palisade. The
correspond with what it is commonly called P. mamillaris in the papillae are less hispid and are clean. The ectosomal skeleton is about
Northeast Atlantic (Channel, North Sea, Irish Sea). The common 800-1000 !Lm thick and is composed of a complex palisade and a tan-
NE Atlantic Polymastia has a two-layered cortex, free intermediary gentiallayer of spicules. The palisade is composed of two layers: an
tylostyles in the choanosome and the shape of spicules is not outer layer of thin tylostyles disposed vertically about 150!Lm thick.
fusiform (see Boury-Esnault, 1974, 1987). This species is actually An inner layer of fusiform tylostyles (-350!Lm thick) intermingled
P. penicillus Montagu, 1818 (Morrow & Boury-Esnault, 2000). with the first one and also vertically arranged (Fig. 3A). Grapnel-like
Redescription of characters of Spongia mamillaris Muller, spicules are present in this palisade and project freely for a certain
1806 does not modify the definition ofthe genus Polymastia as it is distance from the surface (Fig. 3B). The palisade lies on a tangential
presently understood and accepted (Boury-Esnault, 1987; Boury- layer of fusiform tylostyles similar to those of the inner part of the
Esnault et al., 1994a; Kelly-Borges & Bergquist, 1997). palisade. The choanosomal skeleton is made of stout fibres of about
Consequently, the type species of Polymastia is Spongia 250 !Lm. These primary fibres expand into divergent brushes which
mamillaris Muller, 1806, misidentified as Halichondria mamillaris penetrate right into the cortex (Fig. 3C). The skeleton of the papillae
'Johnston, 1842' in the original designation by Bowerbank, 1864. is similar to the one of the main body (Fig. 3D). However the ectoso-
The species is known only from the Swedish west coast between mal skeleton of the papilla is thinner and the exotyles are absent.
75 and 225 m depth. In particular the tangential layer is not well separated from the inner
Porifera' Demospongiae • Hadromerida • Polymastiidae 205

Fig. 3. Proteleia. A, micrograph of a slide of the ectosomal skeleton of Proteleia sollasi made by Dendy & Ridley (BMNH 87.5.2.62) (abbreviations:
E, ectosomal skeleton; P, tangential layer of intermediary tylostyles) (scale 40 jJ-m). B, details of grapnel-like spicules echinating the palisade (scale 80 jJ-m).
C, micrograph of a slide of the choanosomal skeleton of Proteleia sollasi made by Dendy & Ridley (BMNH 87.5.2.62) (abbreviation: F, tracts of principal
spicules) (scale 40 jJ-m). D, transverse section of a papilla through the specimen type (BMNH 87.5.2.62) (abbreviations: E, ectosomal skeleton; F, tracts of
principal spicules) (scale 20 jJ-m). E, ectosomal, intermediary and principal tylostyles (scale 60 jJ-m).

layer of the palisade. Primary megascleres are fusiform styles sharply Topsent, 1913a: 23 (under the name Sphaerotylus schoenus); Boury-
and gradually pointed: 530-1200 X 15-30 /-Lm; intermediary megas- Esnault & Van Beveren, 1982: 39; Levi, 1993: 25; Uriz, 1988: 43.
cleres are fusiform tylostyles which could be slightly curved:
190-222 X 11-19/-Lm; ectosomal tylostyles slender and slightly Type species
curved occurring in the outermost layer of the ectosomal skeleton:
131-175 X 4-5 /-Lm; exotyles which are slender tylostyles with the Polymastia capitata Vosmaer, 1885a (by original designation).
point transformed in what has been called by Dendy & Ridley a grap-
nel and which is constituted by a small knob with three to four small Diagnosis
teeth: 350-520 X 5-6 fl-m.
Remarks. This very beautiful sponge has not been observed Polymastiidae with aquiferous papillae. Primary skeleton com-
since it was originally described by Dendy & Ridley (1886). The posed of tracts of principal spicules and an ectosomal skeleton made
observation of slides from the type-specimen has shown that the up of an inner tangential layer of intermediary tylostyles and an outer
skeleton has a very characteristic organisation without equivalent layer of small tylostyles reinforced by exotyles, called spheroty-
in the other genera. The type species is known only from the lostyles by Topsent. The principal skeleton of papillae is constituted
sublittoral near the Cape of Good Hope. by longitudinal tracts of principal spicules and an ectosomal skeleton
constituted by a palisade of small tylostyles only. The inner layer of
SPHAEROTYLUS TOPSENT, 1898 tangential spicules and the exotyles are absent in the papillae.

Synonymy Remarks

Sphaerotylus Topsent, 1898b: 244; Radiella schoen us Sollas, Sphaerotylus was erected by Topsent (1898b) for the species
1882a: 162 considered as nomen nudum by Kirkpatrick, 1908c: 18; Polymastia capitata Vosmaer, 1885a. However in 1913 Topsent
206 Porifera· Demospongiae • Hadromerida • Polymastiidae

Fig. 4. Sphaerotylus. A, perpendicular section through the body of Sphaerotylus capitatus (abbreviations: E, ectosomal skeleton; I, tangential layer of
intermediary tylostyles; F, tracts of principal spicules) (scale 40/-Lm). B, dissociated spicules of Sphaerotylus capitatus, portion of type (Norman collection
BMNH 10.1.1.1199) (abbreviations: E, ectosomal tylostyles of the palisade; T, principal spicules, polytylote fusiform style; S, exotyles) (scale 30/-Lm).
C, detail of the head of a principal style (scale 8 /-Lm). D, detail of the head of an exotyle (scale 8/-Lm).

placed into synonymy the species P. capitata with Radiella Description of type species
schoenus, taking into account the description and the figure of
Dendy & Ridley (1886: 156). Kirkpatrick (1908c: 18) considered Sphaerotylus capitatus (Vosmaer, 1885a) (Fig. 4).
that R. schoenus, which has no type specimen, type-locality or Synonymy. Polymastia capitata Vosmaer, 1885a: 16;
original description, should be declared a nomen nudum. About Topsent, 1913a: 23; not Boury-Esnault & Van Beveren, 39; not
seven species or sub-species have been described. The type species Kirkpatrick, 1908c: 18;notlJriz, 1988: 43.
Sphaerotylus capitatus has been reported from both Arctic and Material examined. Paralectotype: BMNH 10.1.1.672 -
Antarctic regions. This discontinuous distribution seems to be an Willents Barentz Expedition 1880-1881, Barents Sea, 72°14'8N,
indication that we are faced with a new case of two sibling species 22°30'9E. Lectotype (not seen): RMNH 704. Paralectotype (not
(morphospecies), as already suggested by Levi (1993). seen): ZMA 1841. Other material. MOM - stn 1052 (Topsent,
1913a, as Sphaerotylus schoenus).
Distribution Description. Globular specimen of about 2 cm in diameter
and short papillae. The choanosomal skeleton is constituted by
North Atlantic and Arctic region and sub-antarctic and tracts of principal spicules of about 135 /-Lm in diameter which do
Antarctic region, SW Pacific. not go beyond the surface; between the tracts free intermediary
Porifera· Demospongiae • Hadromerida • Polymastiidae 207

spicules are found. The ectosomal skeleton is composed of an inner most taxonomists with the exception of Dickinson (1945) who
layer of tangential intermediary spicules about 400 J.Lm thick, and described Atergia corona, and Koltun (l964b) who described
a palisade of ectosomal spicules (400 J.Lm thick) which is rein- Atergia acanthoxea Koltun, 1964b. More recently Levi (1993)
forced by a more or less dense layer of exotyles which can go described a new species of Tylexocladus from New Caledonia with
350 J.Lm over the palisade. The skeleton of the papillae is consti- cladotylostyles but without microxeas, and an Atergia species with
tuted of longitudinal tracts of principal spicules between which acanthose microxeas and of course without cladotylostyles. After
free intermediary spicules are present. The ectosomal skeleton in redescribing the type specimens of T. joubini and A. corticata
the region of the papilla is composed only by a palisade of ectoso- Kelly-Borges & Bergquist (1997) considered that even if both gen-
mal spicules. Principal spicules (Fig. 4B, C) are slightly fusiform era were synonymous both species were distinct, and that T. joubini
polytylote styles: 753-950 X 19-26 J.Lm; intermediary spicules and T. corticata both occurred in the NE Atlantic. At the same time
are tylostyles: 314--656 X 12-16 J.Lm; ectosomal spicules are they described a new polymastiid genus Acanthopolymastia with
fusiform tylostyles with a well-marked head: 109-141 X 5.8-6.4; acanthose microxeas similar to Atergia acanthoxa. Reconsidering
exotyles are spherotylostyles (Fig. 4B, D), these spicules being all the species and the type-species described for Tylexocladus,
tylostyles the tip of which is transformed in a slightly spiny sphere: Atergia and Acanthopolymastia, and to avoid confusion until a pre-
650-950 X 18-32 J.Lm, the thickest part of the shaft is close to the cise cladistic approach can be achieved, all three genera are
sphere. retained here for the time being. There are three species of Atergia
Remarks. Specimens of Sphaerotylus capitatus described [A. corticata Stephens, 1915a, A. corona Dickinson, 1945 and
from the South Atlantic, sub-Antarctic or Antarctic region A. villosus (Kelly-Borges & Bergquist, 1997 as Tylexocladus)], three
(Kirkpatrick, 1908c; Boury-Esnault & Van Beveren, 1982; Driz, species of Acanthopolymastia [A. acanthoxa (Koltun, 1964b),
1988) are very likely a new species. All the specimens identified as A. pisiformis Kelly-Borges & Bergquist, 1997 and A. bathamae
Sphaerotylus capitatus or S. schoenus from the sub-Antarctic or Kelly-Borges & Bergquist, 1997] and two species of Tylexocladus
Antarctic regions have to be revised. The type species is known [Tylexocladus jouhini Topsent, 1898b and Tylexocladus hispidus
from the North Atlantic and Arctic Oceans. Levi, 1993].

TYLEXOCLADUS TOPSENT, 1898 Distribution

Synonymy NE Atlantic, from 600-1360 m depth for T. joubini, and SW

Pacific 150 m depth for T. hispidus.
Tylexocladus Topsent, 1898a: 102; 1898b: 242; 1904b:
122-123. Not Atergia Stephens, 1915a: 32; Topsent, 1928c: 151;
Levi, 1993: 23-24; Kelly-Borges & Bergquist, 1997: 393. Not Description of type species
Boury-Esnault et al., 1994b: 75.
Tylexocladus jouhini Topsent, 1898b (Fig. 5).
Type species Synonymy. Tylexocladus joubini Topsent, 1898b: 242;
1904b: 122-123; not Atergia corticata Stephens, 1915a: 32-33.
Tylexocladus jouhini Topsent, 1898a (by original designation). Not Boury-Esnault et ai, 1994b: 75-76.
Material examined. Holotype: MOM Azores,
Diagnosis 39°21'20"N, 33°26'8'W, depth 1360m.
Description. The type specimen (Fig. 5A) resembles the
Massive spherical to hemispherical Polymastiidae without illustration of Topsent (1904b, plate I, fig. 9). It is disc-shaped,
inhalant papillae. The surface is hispid. Oscule at the top of a small attached to a branch of coral, about 2 cm in diameter and only
papilla. Choanosomal skeleton with robust primary tracts com- 2 mm thick. It has a single oscule located on a top of a small papilla.
posed of tapering tylostyles linked by groups of one to three small The surface is finely hispid except on the oscular papilla which is
tylostyles. Ectosomal skeleton consists of a palisade of small, smooth. The choanosomal skeleton is composed of primary tracts
robust tylostyles and cladotylosty1es. Centrotylote microxeas which stretch out from the base to the surface and consist of long
scattered within the choanosome may be present. tapering tylostyles. Between these tracts, groups of small and
robust tylostyles are scattered and intermingled with abundant
Remarks centrotylote microxeas. The ectosomal skeleton is formed by
a dense palisade of small and robust tylostyles which is pierced at
The genus was described by Topsent, 1898a,b for regular intervals by solitary, long and stout cladotylostyles
Polymastiidae with centrotylote microxeas and cladotylostyles. In (Fig. 5B). These spicules are set up below the palisade which they
1915 Jane Stephens described Atergia corticata for a species which pierce and produce the hispid surface. Principal tylostyles: straight,
contained centrotylote microxeas, with long tylostyles mainly hisp- with a well marked head and a very long and slender extremity; up
idating the periphery, thick spicules tracts and a palisade of robust to 1 mm long and 15 J.Lm at the thickest. Ectosomal tylostyles:
tylostyles but without cladotylostyles. Topsent (l928c: 151) fusiform and slightly curved they have a globular head and a short
described new specimens of T. jouhini two of which lacked extremity (175-300 X 10-12 J.Lm). Cladotylostyles: these exotyles
cladotylostyles and had very long tylostyles. Consequently, he con- have a very characteristic denticulate distal extremity (Fig. 5C, D)
sidered that these six specimens, collected from the same locality, where the axial canal of the spicule divided in several ramifica-
all belonged to the same species and thus synonymised Tylexocladus tions. The head is well-marked (550-650 X 20-25 J.Lm). Centrotylote
joubini and Atergia corticata. This synonymy was accepted by microxeas (50-120 X 3-5 J.Lm) (Fig. 5C, D).
208 Porifera· Demospongiae • Hadromerida • Polymastiidae

A -
B -

-
l
T
c D
Fig. 5. Tylexocladus . A, type specimen of Tylexocladus joubini (from Topsent 1904b, pI. I, fig. 9) (scale 26 mm). B, schema of the palisade composed by
small and robust tylostyles which is pierced at regular intervals by c1adotylostyles (from Topsent, 1904b, pI. XII, fig. 10) (scale 115IJ.m). C, detail of the
denticulate extremity of a c1adotylostyle and two centrotylote microxeas (scale 29IJ.m). D, spicules (from Topsent 1904b, pI. XII, fig. II) (abbreviations:
M, centrotylote microxeas; E, ectosomal tylostyles; C, c1adotylostyles; T, principal tylostyles; D, details of the extremities of the c1adotylostyles and the
head of the principal tylostyles) (scales T, 183 fLm; C, E, 60 fLm; D, M, 32 fLm).

Remarks. In other specimens described by Topsent (1928c : Diagnosis

152) dimensions of cladotylostyles may reach 1700 X 351J.m.
Tylexocladus joubini was found only by the 1896 and 1905 Prince Spherical to hemispherical Polymastiidae without inhalant
Albert I de Monaco expeditions to the N Atlantic, West of the papillae. Choanosomal skeleton composed by radial tracts of prin-
Azores Islands from 650--1360 m depth, in areas between cipal tylostyles which fan out very slightly beneath the cortex
39°21'-39°26'N to 31 °05' -31 °22'W. which they pierce. A hispid fringe is often visible around the
margin. The ectosomal skeleton is composed of a palisade of small
tylostyles. Scattered in the choanosome are small tylostyles and
centrotylote microxeas.
ATERGIA STEPHENS, 1915
Remarks
Synonymy
Refer to discussion for Tylexocladus. Three species are
Atergia Stephens, 1915a: 32-33; Topsent, 1928c: 152 allocated here to Atergia: A. corticata, A. villosus (Kelly-Borges &
(as Tylexocladus joubini for two specimens from station 2210); Bergquist, 1997), and A. corona Dickinson, 1945, but not
Dickinson, 1945: 36; Levi, 1993: 26; Boury-Esnault et al., 1994b: A. purpurea Laubenfels, 1954: 204.
75 (as Tylexocladusjoubini); Kelly-Borges & Bergquist, 1997: 396
(as Tylexocladus corticata). Distribution

Type species NE Atlantic, from the West of Ireland to the Gibraltar strait;
500--1200 m depth. Pacific Chatham islands and St James Bank off
Atergia corticata Stephens, 1915a (by original designation). Cape San Lucas (22°50'30"N. 1l0015'W (150m depth).
Porifera· Demospongiae • Hadromerida • Polymastiidae 209

head and a short extremity (175-340 X 12/-Lm). Hispidating

tylostyles may reach more than two millimetres in length, with
very long and thin extremities, but are quite similar in shape to the
principal tylostyles. Microxeas with a slight swelling at the centre
of the spicule (45-100 X 3 /-Lm) (Fig. 6B).
Remarks. The hispidating spicules vary in abundance and
length between different specimens, mostly however because they
are usually broken. This kind of spicule is very frequent in species
living in mud bottoms and their abundance may be linked to envi-
ronmental conditions. The type species is distributed in the NE
M Atlantic, from the West of Ireland to the Gibraltar Strait and around
the Azores Islands; 500-1200 m depth.

ACANTHOPOLYMASTIA KELLY-BORGES &

BERGQUIST, 1997

Synonymy

Acanthopolymastia Kelly-Borges & Bergquist, 1997.

Type species

Atergia acanthoxa Koltun, 1964b (by original designation).

Diagnosis

Polymastiidae with a cushion-shaped or discoid shape and a

single low oscular papilla. The edge of the sponge body is very hispid
whereas the other part of the surface is only minutely hispid. The
choanoskeleton is composed by tracts of tylostyles or subtylostyles
which project well beyond the surface at the edge. The ectosomal
skeleton is composed of a palisade of small tylostyles. Acanthose
microxeas are very abundant both in the ectosome and choanosome.

Remarks
B
T This genus was erected by Kelly-Borges & Bergquist (1997)
Fig. 6. Atergia. A, type specimen of Atergia corticata (from Stephens, for the species previously placed in Atergia but having acanthose
1945, pI. III, figs 2-3) (scale 4.5mm). B, spicules (from Stephens, 1915a, microxeas densely packed in the choanosome and the ectosome.
pI. V, fig. 4) (abbreviations: T, principal tylostyles; E, ectosomal tylostyles; Three species are included in this genus: A. acanthoxa (Koltun,
Ta, detail of the head of a principal tylostyles; M, centrotylote microxeas)
(scales T, 80 !Lm; E, Ta, 34 !Lm; M, 18 !Lm).
1964b), A. pisijonnis Kelly-Borges & Bergquist, 1997 (described
as Atergia acanthoxa by Levi, 1993), and A. bathamae Kelly-
Borges & Bergquist, 1997.
Description of type-species
Distribution
Atergia corticata Stephens, 1915a (Fig, 6).
Synonymy. Atergia corticata Stephens, 1915a: 32-33. This genus is so far known only from the Southwest Pacific,
Material examined. Holotype: BMNH 1953.11.11.12 - from deeper waters 500-3300m depth.
54°17'N, llo33'W, 700m depth. Other material. Gibraltar strait,
35°56'N, 5°35'E, 534-604m depth (Boury-Esnault et al., 1994b). Description of type species
Description. Disc-shaped specimen, 15 mm in diameter and
5 mm in thickness in the centre (Fig. 6A). The surface is hispid par- Acanthopolymastia acanthoxa (Koltun, 1964b) (Fig. 7).
ticularly at the periphery. The choanosomal skeleton is composed Synonymy. Atergia acanthoxa Koltun, 1964b: 27. Not
of strongly radiating tracts, which emanate from the base of the Atergia acanthoxa Levi, 1993: 26; Acanthopolymastia acanthoxa
sponge and end beyond the surface. Centrotylote microxeas and Kelly-Borges & Bergquist, 1997: 396.
small tylostyles criss-cross the primary tracts. The ectosomal Material examined. Holotype (fragment, courtesy of
skeleton is a very dense palisade of small tylostyles. The principal Vladimir Koltun): SME-Po57 - Antarctica, Ross Sea, 68°00'S,
tylostyles often pierce the surface making it very hispid. Principal l60 07'W.
0

tylostyles are straight with a well marked head and a very long, Description. Body-cushion shaped, about 4cm diameter
slender extremity (780-1350 X 12-18 /-Lm). Ectosomal tylostyles and 1 cm thick (Fig. 7A). A very long fringe is present at the edge.
are slightly curved and have a fusiform shaft, with a well-marked The choanoskeleton is formed by radial tracts of principal spicules
210 Porifera· Demospongiae • Hadromerida • Polymastiidae

c Tf E Tp
Fig. 7. Acanthopolymastia. A, type specimen of Acanthopolymastia acanthoxa (from Koltun, 1964b, as Atergia, fig. 3). Note: in the center the oscular
papilla and at the periphery the fringe of long spicules (scale 7.5 mm). B, part of the ectosomal skeleton where the dense layer of acanthoxeas is clearly
visible (abbreviation: A, acanthoxea) (scale 30 !Lm). C, spicules from the type specimen (from Koltun, 1964b, fig. 3) (abbreviations: Tf, spicules of the
fringe; E, ectosomal tylostyles; A. acanthoxea; Tp. principal tylostyles) (scales Tf. E. Tp. 77 IJ.-m; A. 22IJ.-m).

between which there is numerous acanthoxeas. The ectosomal Diagnosis

skeleton, 500-600 fLm thick, is composed of a dense layer of irregu-
larly distributed acanthoxeas (Fig. 7B) and an outer palisade of ecto- Polymastiidae with a short peduncle and an ovoid body, with
somal spicules. Principal tylostyles or subtylostyles 1650-2350 X a single apical oscule at the summit. From the inside to the outside
16-21 fLm; styles or subtylostyles of the fringe up to 7 mm long and the ectosomal skeleton is composed of ascending primary tracts,
14-34 fLm wide; ectosomal tylostyles 460-630 X 8-10 fLm; acan- a tangential layer of spicules perpendicular to the primary tracts
those centrotylote microxeas 98-185 X 5-8 fLm (Fig. 7C). and an extemallayer made of tufts of small styles perpendicular to
Remarks. This species is known only from the Antarctic the surface. The choanoskeleton is reduced to consist only of dis-
coast, Ross Sea, from considerable depth (3200-3400m). persed bundles of small styles. Megascleres are styles, subty-
lostyles, strongyloxeas or tylostyles of two or three sizes.
QUASILLINA NORMAN, 1869
Remarks
Synonymy
Four species are included in the genus: Q. brevis, Q. richardi
Quasi/Una Norman, 1869b: 329. Vosmaer, 1885a: 20; Ridley Topsent, 1913a and Q. intermedia Boury-Esnault, Pansini & Uriz,
& Dendy, 1887: 226; Dendy, 1888: 520; Topsent, 1898a: 102; 1994b, Q. translucida Desqueyroux-Faundez & Van Soest, 1997.
Topsent, 1900: 157; Boury-Esnault et al., 1994b: 70. ? BursaUna Quasillina quiza described by de Laubenfels (1954: 211) from the
Schmidt, 1875: 116. Also Bowerbank, 1862a: 71. central West Pacific is certainly not a Polymastiidae. The most
important character of this genus is the fact that the tracts of princi-
Type species pal megascleres are localized within the inner part of the ectosome
and do not constitute the choanosomal skeleton as in the other
Euplectella brevis Bowerbank, 1862a (by original designation). Polymastiidae. The choanoskeleton is reduced to bundles of small
Porifera· Demospongiae • Hadromerida • Polymastiidae 211

Fig. 8. Quasillina. A, histological section ofthe ectosome of Quasi/tina brevis from the 'Challenger' collection (abbreviations: E, ectosomal skeleton, see
tufts of small styles; T, tangential layer of principal styles) (scale 50 !Lm). B, histological section of the choanosome of Quasiltina brevis from the
'Challenger' collection (abbreviation: F, small styles in free bundles) (scale 50 !Lm).

styles which correspond to the free spicules often found between 5 to 12 (Fig. 8B). The ectosomal skeleton is composed from inside
the principal tracts in most of the polymastid genera. to outside by a layer of primary tracts which are closely related to a
second layer of tangential spicules, exactly perpendicular to the
Distribution tracts, and superficially by tufts of small styles perpendicular to the
surface (Fig. 8A) and between which the ostia open. Principal
This genus is known predominantly from the Northern hemi- styles are fusiform and resemble strongyloxeas: 600-990 X
sphere, from the Arctic, NE Atlantic and Mediterranean seas. The 10-26 fLm; small ectosomal styles: 135-210 X 2.6-5.3 fLm.
bathymetric distribution varies between 15-700m depth with the Reproduction. Spermatogenesis was observed in Arctic
most littoral stations located in the Arctic. material in June (Vosmaer, 1885a).
Remarks. This species was perfectly described by Topsent
(1900: 158). Its seems to be relatively abundant in all the NEAtlantic
Description of type species
and Mediterranean, with one record from the tropical E Pacific.
Euplectella brevis (Bowerbank, 1862a) (Fig. 8).
Synonymy. Euplectella brevis Bowerbank, 1862a: 71. RlDLEIA DENDY, 1888
Polymastia brevis; Bowerbank, 1866: 64; Bowerbank & Norman,
1882: 16 & 31; Lendenfeld, 1897a: 222; Quasillina brevis; Norman, Synonymy
1869b: 329; Vosmaer, 1885a: 20; Vosmaer, 1887: 330; Ridley &
Dendy, 1887: 226; Dendy, 1888: 520; Hanitsch, 1894a: 175 & 203; Ridleia Dendy, 1888: 515; Topsent, 1898a: 102; Boury-
Topsent, 1900: 158-164; Lundbeck, 1909: 452; Br0ndsted, 1914: Esnault et aI., 1994b: 72-73. Not Ridleia dendiia Laubenfels,
522; Stephens, 1915a: 30; Rezvoj, 1928; 81; Hentschel, 1929: 869; 1934: 10-11. Not Ridleia peleia Laubenfels, 1954: 207. Not
Br0ndsted, 1933: 14; Burton, 1959b: 13; Koltun, 1964b: 150; 1966: Ridleia sensu de Laubenfels, 1936a: 151.
89; Boury-Esnault et al., 1994b: 71.
Material examined. Holotype: BMNH 10.1.5 & 6 - North Type species
Atlantic, Shetland. Other material. MOM - stn 247 (Topsent,
1904b: 132); BALGIM collection (Boury-Esnault et al., 1994b: 71). Ridleia oviformis Dendy, 1888 (by monotypy).
Description. Whitish pedunculate sponge with an ovoid
body about 2-3 cm high and 5-10 mm in diameter. The oscule is Diagnosis
located at the upper part of the sponge. The surface is velvety. The
ectosome is tough whereas the choanosome is fleshy. The Pedunculate egg-shaped sponge with an oscular papilla.
choanoskeleton is composed only of small styles in free bundles of Skeleton mainly confined to the ectosome. From outside to inside
212 Porifera· Demospongiae • Hadromerida • Polymastiidae

Fig. 9. Ridleia. A, histological slide from the Dendy collection of Ridleia oviformis (abbreviations: P, palisade of the ectosomal skeleton; I, tangential layer
of intermediary spicules; F, tracts of principal spicules; C, collagenous layer at the limit between ectosome and choanosome; T, tangential layer of ectoso-
mal tylostyles in the outermost region of the choanosome; Ch, choanosome) (scale 32j.Lm). B, different kinds of tylostyles present in Ridleia oviformis
(scale 36j.Lm).

there is a palisade of ectosomal tylostyles, a layer of intermediary material. From the ibero-moroccan Gulf, 35°54'I"N, 6°14'5W,
tylostyles and longitudinal tracts of principal tylostyles which lie 150m depth (Boury-Esnault et al., 1994b: 72-73); MOM - Prince
on a collagenous layer. In the outermost region of the choanosome Albert I de Monaco expedition, station 2214, West of Flores,
there is a tangential layer of ectosomal tylostyles. The spicules are 39°16'lO"N, 31 °2I'20"W (identified by Topsent, 1928c: 150-151).
tylostyles in three size categories. Choanocyte chambers are Description. The holotype has an egg-shaped body,
diplodal. 15 mm high and 7 mm wide, ending in an oscular papilla and fixed
by a smaJl peduncle to pebbles. The colour in spirit is pale yellow
Remarks and the consistency is cork-like. The skeleton is almost entirely
confined to the ectosome. From outside to inside, a palisade of thin
In addition to the type species two other species were allo- ectosomal tylostyles lies on a tangential layer of intermediary
cated to Ridleia by de Laubenfels (1 936a): Ridleia dendiia tylostyles. In the inner part of the ectosome, longitudinal tracts of
Laubenfels, 1934 and Ridleia peleia Laubenfels, 1954. These principal tylostyles followed by a collagenous layer which indicate
species do not belong to Polymastiidae, indeed if they belong to the limit between ectosome and choanosome. In the outer-most
Hadromerida at all, the skeleton being confused and without region of choanosome a layer of tangential ectosomal tylostyles is
a spicule palisade. De Laubenfels (1936a) referred them to Ridleia present, representing the only remnant of the choanosomal skele-
based on the shape of their spicules but did not take into account ton (Fig. 9A). Principal tylostyles are straight tylostyles with a
the organization of the skeleton. well-marked head: 400-900 Il-m X l2ll-m. Intermediary tylostyles
are slightly fusiform, with a well-marked head: 250-350 X 7 Il-m.
Distribution Ectosomal tylostyles are very slender, frequently curved and they
taper gradually to the apex: 130-200 X 21l-m (Fig. 9B).
Northeast Atlantic, from 60°0' to 35°54'I"N and 5°13' to Reproduction. Small oocytes were observed in the
31°21'20"W, from 150-950m depth. histological slides of the Dendy collection.
Remarks. Although this species has been collected only on
Description of type species four occasions Dendy's (1888) description was so accurate that it
makes it perfectly recognisable. It shares with all Polymastiidae
Ridleia oviformis Dendy, 1888 (Fig. 9). longitudinal tracts of megascleres and palisade of ectosomal
Synonymy. Ridleia oViformis Dendy, 1888: 515; Topsent, tylostyles but the skeleton is almost exclusively confined to the
1904b: 124; 1928c: 150-151; Boury-Esnault et al., 1994b: 72-73. ectosomal region. The type species has been recorded from various
Material examined. Holotype: BMNH 83.12.13.69 - NE sites in the NE Atlantic - Northwest of Scotland (60 0 0'N, 5°13'W,
Atlantic, NW of Scotland, 60 0'N, 5°13'W, 571 m depth. Other
0
571 m depth), close to the Azores (Banc Princesse Alice, 37°57'N,
Porifera' Demospongiae • Hadromerida • Polymastiidae 213

29°14'45'W, 200m depth, W of Flores, 39°16'1O''N, 31°21'20'W, Polymastia: Trichostemma grimaldi Topsent, 1913a (by Topsent,
650-914m depth), and more recently from the ibero-morrocan 1927c: 257) and Radiella tissieri Vacelet, 1961 b (by Vriz & Rosell,
Gulf (35°54'I"N, 6°14'5W, 150m depth). 1990: 378).

Distribution
RADIELLA SCHMIDT, 1870
Three species have been described from the Atlantic: R. sol
Synonymy Schmidt, 1870 (West central Atlantic), R. hemisphaericum (Sars,
1872) (Arctic and North Atlantic), R. sarsi (Ridley & Dendy, 1886)
[Trichostemma] Sars, 1869: 250 (nomen nudum). Radiella (NE Atlantic and Mediterranean sea), and two species from the
Schmidt, 1870: 48; Schmidt, 1880b: 77; de Laubenfels, 1936a: Pacific ocean R. irregularis (Ridley & Dendy, 1886) and
150. Trichostemma Sars, 1872: 62-63; Ridley & Dendy, 1887: R. straticularis (Wilson, 1925). All belong to the deep water fauna.
216; Topsent, 1898a: 101; Dendy, 1922b:151; Wilson, 1925: 347;
Boury-Esnault et al., 1994b: 73. Description of type species

Type species Radiella sol Schmidt, 1870 (Fig. 10).

Synonymy. Radiella sol Schmidt, 1870: 48; Sars, 1872:
Radiella sol Schmidt, 1870 (by subsequent designation). 62-63; Topsent, 1913a: 20-21; Ridley and Dendy, 1886: 216;
Lundbeck, 1909: 451-452; Polymastia hemisphaericum; Vosmaer,
Diagnosis 1885a; Koltun, 1964a: 149; Koltun, 1966: 78.
Material examined. ? Holotype: VSNM 990 - 17°28'N,
Circular lenticular or hemispherical Polymastiidae. The skele- 77°30'W, 1116m depth, 'Blake' expedition 1878-1879 (identified
ton of the lower face of the sponge is composed of an envelope of by Schmidt). Other material. Trichostemma hemisphaericum:
principal subtylostyles or styles covered by a thin palisade of ecto- MOM - Prince Albert I de Monaco expedition to the Atlantic, stn
somal tylostyles. The upper face is composed of one or two layers 1052; Norwegian coast 65°41'N, 9°30'E, 440m; stn 960 between
of tylostyles. The choanosomal skeleton is composed of diverging Norway and Bear Island, 72°37'N, 20 00'E, 394m depth.
0

tracts emanating from a central nucleus at the base of the sponge Comparative material. MOM - misidentified as T. hemisphaericum
and by groups of fusiform tylostyles. The inhalant and exhalant Topsent, 1892a, stn. 16146°4'40''N, 49°2'33"W, 1267m depth.
apertures are grouped on papillae. Description. The holotype is hemispherical, 2 cm in diame-
ter, 7 mm thick (Fig. lOA). On the upper side 15 papillae about
Remarks 1 mID high were observed. The edge of the body is covered by a
very long fringe. The surface is velvety. The ectosomal skeleton of
The status of the genus Radiella has been controversial. the upper face is about 1500 f-Lm thick, composed of a palisade
Schmidt (1870: 48) described briefly a new genus Radiella without of ectosomal tylostyles (-850 f-Lm thick) and a paratangentiallayer of
designation of a type species or a type specimen. He allocated two criss-crossed fusiform tylostyles (-650 f-Lm thick) (Fig. lOB). The
species to this genus: R. spinuiaria (Bowerbank, 1866) and his new choanosomal skeleton is composed of diverging tracts of principal
species from the Gulf of Mexico R. sol. Tethea spinularia tylostyles (250-340/-Lm in diameter) emanating from a central
Bowerbank, 1866 is in fact the type species of the genus Spinularia nucleus at the base of the sponge, and by numerous rosettes of
Gray, 1867a, which was considered to be valid by Stephens fusiform tylostyles. The skeleton of the lower face of the sponge is
(1915a). Previously Sars, 1869 alluded to a new genus and species, composed of an envelope of principal subtylostyles covered by
Trichostemma hemisphaericum, in a species list detailing little a thin palisade of ectosomal tylostyles (Fig. 1OC). Ectosomal
more than only geographical information (Lofoten islands, Arctic- tylostyles straight or slightly curved with a well marked head
Atlantic area, between 200-300m depth). The species and genus 290-390 X 11 /-Lm; intermediary fusiform tylostyles 460-620 X
were only later carefully described by Sars in 1872. Thus, Schmidt 26 f-Lm; principal subtylostyles straight and with an acerate extremity:
(1880b: 77) claimed that Trichostemma hemisphaericum described 795-2290 X 11-32 f-Lm and subtylostyles of the fringe which are
by Sars in 1872 was synonym of Radiella sol, described in 1870. >5 mID long and about 50 /-Lm thick.
Since this time both genera have been used equally by authors, or Remarks. Presently it is impossible to ascertain if this
following Ridley & Dendy (1887) who used Trichostemma in pref- specimen from the West central Atlantic is a synonym of
erence because it was well described, or following the 'ruling' of 'hemisphaericum' from the Arctic and North Atlantic. A detailed
Schmidt (I 880b ) and the rules of zoological nomenclature. A spec- revision of specimens from the Arctic with other specimens from
imen identified by Schmidt as Radiella sol was discovered in the West central Atlantic is necessary before making such a decision.
VSNM Washington, although it does not correspond to the illustra- The type species is known from the West central Pacific, from
tion of Schmidt (1870, plate 4, fig. 6). Nevertheless, Trichostemma deep waters.
is considered here to be a synonym of Radiella following Article
12 of the International Code of Zoological Nomenclature (Anon.,
2000). WEBERELLA VOSMAER, 1885
Five species are presently considered to belong to this genus:
Radiella sol Schmidt, 1870, R. hemisphaericum (Sars, 1872), Synonymy
R. sarsi (Ridley & Dendy, 1886), R. irregularis (Ridley & Dendy,
1886) and R. straticularis (Wilson, 1925). Two other species Weberella Vosmaer 1885a: 16; Topsent, 1928c: 149; Vacelet,
have been assigned to this genus but are reallocated to 1960a: 261; Vriz, 1975: 429-442; Boury-Esnault et aI., 1994b: 76.
214 Porifera· Demospongiae • Hadromerida • Polymastiidae

Fig. 10. Radiella. A, type specimen of Radiella sol (from USNM 990) (scale 4 mm). B, detail of the ectosomal basal skeleton through a thick section of
Radiella sol (scale 170 11m). C, transverse section of a specimen of Radiella sol (abbreviations: E, ectosomal skeleton; F, fringe) (scale 1 mm).

Type species Description of type species

Alcyonium bursa Muller, 1806 (by original designation). Weberella bursa (Muller, 1806) (Fig. 11).
Synonymy. Alcyonium bursa Muller 1806: 43; Weberella
Diagnosis bursa Vosmaer, 1885a: 16; Topsent, 1928c: 149; Uriz, 1975: 430;
Boury-Esnault et al., 1994b: 76. Polymastia bursa Koltun, 1964a:
Polymastiidae with small aquiferous papillae, reticulate 149; Koltun, 1966: 76. Not Suberites bursa Schmidt 1862: 68. Not
choanosomal skeleton, and an ectosomal skeleton composed of a Polymastia bursa Lendenfeld, 1897a: 117.
palisade of tylostyles which lie on a paratangential layer of criss- Material examined. Holotype: Unknown - Arctic Ocean,
crossed tylostyles. Only two category of tylostyles present. n036'N, 24°57'E, depth 250m. Other material. BMNH
63.71.29.87 - from Koltun Collection registered in Leningrad
Remarks Museum under the number 7827; BALGIM collection - stn DR-42
(Boury-Esnault et al., 1994b); MOM, stn 1463 (Topsent, 1928c).
Two species have been described for this genus: Weberella Description. Globular or ovoid sponge (Fig. l1A) with a finn
bursa (Muller, 1806) from the NE Atlantic and W. verrucosa consistency and a finely hispid surface. The largest specimens
Vace1et, 1960a from Mediterranean Sea and the Gibraltar strait. In described are about 10cm in diameter. Small oscular papillae
the literature there is a confusion between Suberites bursa Schmidt, (1-5mrn high) are located on the upper part. The choanoskeleton is
1862: 68 which has been allocated to Polymastia by Lendenfeld composed of ascending primary tracts of tylostyles of about 50 fLm
(1897a: 117) and the species of Muller (1806) which has also been in diameter, which are intersected by secondary tracts. This reticula-
allocated to the genus Polymastia by Koltun (1964a: 149). tion is easily visible at the periphery. Within the mesh of this net free
groups of ectosomal tylostyles are present. The ectosomal skeleton
Distribution (-350 fLm thick) is composed of an outer palisade of small tylostyles
(-250fLm thick) (Fig. 11B) which lie on an inner paratangential
NE Atlantic and Mediterranean seas. layer of criss-crossed tylostyles (-90 fLm thick). The skeleton of the
Porifera· Demospongiae • Hadromerida • Polymastiidae 215

A
Fig.n. Weberella. A. type specimen of Weberella bursa (from Vosmaer, 1885a, pI. I, fig. 12) (scale 1 cm). B, schematic drawing of the organization of the
skeleton of Weberella bursa (from Vosmaer, 1885a, pI. 17, fig. 6) (abbreviation: E, ectosoma1 skeleton) (scale 500 f,Lm).

papillae is similar. Principal tylostyles are slightly fusiform with an genus is clearly recognizable given the precise description of
ovoid often mucronated head: 34()....{j50 X 6-12j.Lm; ectosomal Vosmaer (1885a: 18-19) the allocation of other species, such as
tylostyles are slightly curved with a well-rounded head: 90-160 X papillatus (Kirkpatrick, 1907a) by Topsent (1917: 36) or caminatus
3-6j.Lm. (Ridley & Dendy, 1887) by Koltun (1976: 168) require a very care-
Remarks. This species is easily recognised although rarely ful revision of all specimens before any decision could be made as
collected, found in the NE Atlantic from the Arctic to the ibero- to their correct allocation to this genus. For the present we consider
moroccan Gulf; from 130-960m depth. there are two species in this genus: the type species and Tentorium
papillatum. The type species has been reported from both the
Arctic and Antarctic regions, although this discontiguous distribu-
TENTORIUM VOSMAER, 1887 tion may indicate the possibility that there are two sibling species
hiding under this name.
Synonymy
Distribution
[Thecophora] Schmidt 1870: 50 (preocc.; Vosmaer, 1885a:
18). Tentorium Vosmaer, 1887: 329; Ridley & Dendy, 1887: 221; NW Atlantic, South Atlantic, Arctic and Antarctic regions.
Topsent, 1917: 36; Topsent, 1928c: 151; Boury-Esnault & Van
Beveren, 1982: 37. Description of type species

Type species Tentorium semisuberites (Schmidt, 1870) (Fig. 12).

Synonymy. Thecophora semisuberites Schmidt 1870: 50;
Thecophora semisuberites Schmidt, 1870 (by original desig- Vosmaer, 1885a: 18; Brs<Sndsted, 1914: 522-523; Ferrer-
nation). Hernandez, 1914b: 19; Stephens, 1915a: 29-30; Hentschel, 1929:
868-869; Koltun, 1959: 163; 1964a: 148. Not T. semisuberites
Diagnosis Boury-Esnault & Van Beveren, 1982: 37 and all material from the
South Atlantic and Antarctic.
Polymastiidae with colunmar or globular growth form pro- Material examined. Holotype: ZMUC - Ubekendl, Arctic
tected by a dense cylindrical sheath of longitudinally placed spicules Ocean "det. Schmidt".
which form a solid imperforate layer. The ectosomal skeleton con- Description. The body is cylindrical with a semi-spherical
tains bundles of smaller spicules arranged in a palisade and is limited upper part and has about 1-3 cm high and 0.5-1.5 cm in diameter
to the upper surface of the cylinder. Ostia are present on the upper (Fig. 12A). The surface of the cylinder is smooth whereas the
surface. The oscule at the top of a small papilla is on centre of the upper part is rough. There are 1-3 oscules located at the summit of
upper surface. Megascleres are subtylostyles and tylostyles. small papillae. Ostia are located on the upper part of the sponge.
The principal skeleton is composed of tracts of tylostyles oriented
Remarks according to the axis of the cylinder (Fig. 12B). The ectosomal
skeleton is composed of a palisade of ectosomal fusiform tylostyles
Thecophora Schmidt was preoccupied by a species of Diptera, on the upper surface and by a tangential layer of intermediary
Insecta (Thecophora Rondani, 1845), and hence the creation of fusiform tylostyles on the side of the cylinder. Principal tylostyles
a new name by Vosmaer (1887). Although the type specimen of the or subtylostyles with a tapering tip: 950-2400 X 13-24 j.Lm;
216 Porifera· Demospongiae • Hadromerida • Polymastiidae

A - B -
Fig. 12. Tentorium. A, type specimen of Tentorium semisuberites (from Vosmaer, 1885a, pI. I, fig. 23) (abbreviations: 0, oscule; U, upper part of the
sponge; C. cylindrical body) (scale 0.6cm). B, schematic drawing of a longitudinal section through the body of a specimen of Tentorium semisuberites
(from Vosmaer, 188Sa, p. 19, fig . 9) (abbreviations: E, ectosomal skeleton; F, tracts of principal spicules) (scale 0.5 cm).

intermediary fusiform tylostyles: 800-1150 X 20-33 f.Lm; ectoso- Distribution

mal fusiform tylostyles: 270-670 X 13-20 f.Lm.
Remarks. The southern specimens from South Atlantic NE Atlantic and SW Pacific in the bathyo-abyssal zone
(Inaccessible Island close to Tristan da Cunha, Ridley & Dendy, (650-4000m deep). In the Northern part of the Atlantic (Norway,
1887) and from Kerguelen (Boury-Esnault & Van Beveren, 1982) Greenland and Sweden coasts) it can be collected as shallow as
probably represent a new species of Tentorium as they are not con- 50-250 m deep.
specific with the type species. The type species is found in the
Artic and North Atlantic, from Greenland to the Azores, relatively Description of type species
deep water except in high latitudes where it may be collected in the
littoral zone. Spinularia spinularia (Bowerbank, 1866) (Fig. 13).
Synonymy. Tethea spinularia Bowerbank, 1866: 94.
Spinularia tetheoides Gray, 1867a: 524. Radiella spinularia
SPINULARlA GRAY 1867 Schmidt, 1870: 48. Polymastia spinularia Hanitsch, 1894a: 202.
Rhaphidorus setosus Topsent, 1898b: 245; Alander, 1942: 76.
Synonymy Material examined. Holotype: BMNH: 10.1.1.1183 - Norman
collection (slides), Shetland. Other material. MOM - stns 683 and
Spinularia Gray 1867a: 524. Rhaphidorus Topsent, 1898b: 3293 (Topsent, 1904b, 1928c). Not BMNH 1O.1.1.594A (Norman
244; 1904b: 120; Stephens, 1915a: 31; Topsent, 1928c: 150; Levi, collection) [=Atergia corticata).
1993: 25. Description. (Morphology based on MOM material, skele-
ton based on BMNH slides) Small discoid sponge with a diameter
Type species of about 12mm and a thickness of 3-4mm. Specimens are often
fixed to pebbles or shells (Fig. 13A). The surface is especially
Tethea spinularia Bowerbank, 1866 (by original designation). hispid towards the edge of the sponge where the spicules project
a considerable distance. No aquiferous papillae are present and
Diagnosis the oscule is slightly raised above the surface. Choanosomal
skeleton is composed of tracts of tylostyles between which
Small, more or less discoid Polymastiidae. Surface minutely raphides in trichodragmata are present. The ectosomal skeleton
hispid except at the edge where there is a strong hispidation pres- is composed of a simple palisade of ectosomal tylostyles
ent. Choanosomal skeleton composed of primary tracts of (Fig. 13B). Principal tylostyles are slightly fusiform, with an acer-
tylostyles between which trichodragmata are found. Ectosomal ate tip and an ovoid head: 900-1450 X 18-22 f.Lm; echinating
skeleton is composed of a palisade of ectosomal tylostyles. There tylostyles of the edge can reach up to 2000-2500 f.Lm; ectosomal
are no true aquiferous papilla and the oscule is located at the upper tylostyles are straight, fusiform and with a well-marked head:
part of the sponge and is slightly raised above the general surface. 180-310 X 10-13 f.Lm; raphides in trichodragmata (Fig. 13C):
70-120 f.Lm.
Remarks Remarks. All descriptions of this species are congruent
although the length of trichodragmata is the most variable element
This genus has a relatively complicated story which was even- and can reach 260 f.Lm in some Azorean and Irish specimens
tually solved by Stephens (1915a).A second species of this genus was (Topsent, 1928c; Stephens, 1915a). The type species is known only
recently found in the bathyal zone of New Caledonia (Levi, 1993). from the NE Atlantic.
Porifera· Demospongiae • Hadromerida • Polymastiidae 217

Fig. 13. Spinuiaria. A, specimen of Spinuiaria spinuiaria (from Topsent, 1904b, as Rhaphidorus setosus, pI. I, fig. 6) (scale I cm). B, thick section through
the body of Spinuiaria spinuiaria, slide from the Norman collection (BMNH 10.1.1.1183) (abbreviations: E, ectosomal skeleton; F, tracts of the principal
spicules) (scale 100 f1.m). C, detail of raphides in trichodragmata (scale 14 f1.m).

PSEUDOTRACHYA HALLMANN, 1914 Diagnosis

Synonymy Polymastiidae with radial skeleton of subtylostyles or styles

which project through the surface rendering it very hispid.
Pseudotrachya Hallmann, 1914: 286; Vacelet, 1969: 178; Ectosomal skeleton composed of a palisade of small oxeas. Small
Boury-Esnault et al., 1994b: 69. Anomolissa (partim) de Laubenfels, oxeas of the same type are scattered in the choanosome.
1934: 17; Van Soest & Stentoft, 1988: 79 (for A. amaza).
Remarks
Type species
Pseudotrachya was erected by Hallmann (1914) for Solasella
Trachya hystrix Topsent, 1892a (by monotypy). hystrix (Topsent, 1892a) and has been previously assigned to the
218 Porifera· Demospongiae • Hadromerida • Polymastiidae

T'
I
B
Fig. 14. Pseudotrachya. A, type specimen of Pseudotrachya hystrix (from Topsent, 1892a, as Trachya hystrix, pI. I, fig. 10) (scale 0.6 em). B, schema of
a style (T) echinating the palisade of oxeas (E) (from Topsent, 1892a, pI. XI, fig. 13) (scale 180 /Lm).

Axinellidae, by Topsent (l904b, 1928c) and Hallmann (1914), but is composed of an ectosomal palisade of small oxeas (Fig. 14B) and
always with some doubt. It was recently considered to be a choanosomal radial tracts of megascleres which protrude through
hadromerid by Van Soest & Stentoft (1988) and included in the palisade. Ectosomal oxeas are also dispersed within the tracts of
Tethyidae, but again with some doubt. However, Topsent (1904b) choanosomal megascleres. Oxeas are slightly fusiform with acerate
stated it was not a Tethyidae and Boury-Esnault et al. (1994b) sug- extremities (185-253 X 6/J..m) and tylostyles or subtylostyles which
gested its closest affinities were with Polymastiidae. can reach 7 mm length in the holotype. In other material these
tylostyles were shorter reaching only about 2000-4000 X 44 /-Lm. In
Distribution some Mediterranean specimens the megascleres can vary from
styles to oxeas (Sara, 1959b: 3-6; Vacelet, 1969: 178).
Atlantic ocean (from the Western Mediterranean to the Puerto Remarks. Pseudotrachya is presently considered to be
Rican Deep in the east, and to South Africa in the south), 100-954 m monospecific. However, the type species has a very wide alleged
depth although specimens have been recorded from 25 m depth in distribution, from Mediterranean to the Caribbean and to South
obscure caves on the Provence coast (Western Mediterranean basin). Africa, and it is highly suspected that it comprises two or more
cryptic sibling species. In fact we can hypothesise that these dis-
Description of type species contiguous populations comprise at least three or four species:
P. hystrix from Azores and perhaps also extending into the
Pseudotrachya hystrix (Topsent, 1892a) (Fig. 14). Mediterranean; a Caribbean species which could be called
Synonymy. Trachya hystrix Topsent, 1892a: 75. SoIasella P. amaza as already suggested by Lehnert & Van Soest (1999), and
hystrix Topsent, 1898a: 111; Kirkpatrick, 1903b: 247; Topsent, a South African species as yet unnamed. To date, however, there
1904b: 142. Pseudotrachya hystrix Topsent, 1928c: 180; Vacelet has been no formal revision of this species complex and the
1969: 178; Van Soest & Stentoft, 1988: 79; Boury-Esnault et aI., conspecificity between these regional populations remains
1994b: 70. Pseudotrachya oxystyla Sara, 1959b: 3-6. Anomolissa speculative.
amaza de Laubenfels, 1934: 17. ? Suberitechinus de Laubenfels,
1949a: 21.
Material examined. Holotype: MNHN-DT. 896 TRACHYTELEIA TOPSENT, 1928
'L'Hirondelle' expedition, 1888, StD. 234, Azores, between Pico and
Sao Jorge islands, 38°30'35''N, 28°16'20''W. Other material. Synonymy
Mediterranean, stn 153 and 135 from the BALGIM expedition
(personal collection) (Boury-Esnault et aI., 1994b). Anomolissa Trachyteleia Topsent 1928c: 152. ? Suberitechinus de
amaza: UNSM.22348 (de Laubenfels, 1934: 17). Laubenfels, 1949a: 21.
Description. The holotype is attached to pebbles, with
a cushion shape and a maximum length of 4 cm and 2 cm width Type species
(Fig. 14A). The colour in alcohol is whitish outside and pale yellow
inside. The surface is echinated by very long spicules. The skeleton Trachyteleia stephensi Topsent, 1928c (by monotypy).
Porifera· Demospongiae • Hadromerida • Polymastiidae 219

Diagnosis

Polymastiidae without papillae, with radiating tracts of

tylostyles and tracts of small tylostyles in between. The ectosomal
skeleton is composed by a palisade of small tylostyles. The surface
is echinated by distally acanthose tylostyles which are consider-
ably thicker than the primary spicules.

Remarks

The type species has never been collected since the Prince
Albert I de Moncao expedition in 1911. Unfortunately the entire
specimen was used by Topsent for his spicule preparation although
in this preparation the different types of spicules are very clear in
addition to some non-dissociated parts of the skeleton.

Distribution

NE Atlantic, Azores.

Description of type species

...
Trachyteleia stephensi Topsent, 1928c (Fig. 15). E
Synonymy. Trachyteleia stephensi Topsent, 1928c,
152-154, pI. 6, fig. 11.
Material examined. Holotype (slide only): MNHN DT
1285 - 38°01'N, 25°21'W, Azores, 1740m depth.
Ti
Description. The holotype consists of a very small hispid
crust and was grey in alcohol (now destroyed and only extant as
a spicule preparation). There are neither visible papillae nor visible Ex
o Tp
apertures. The choanosomal skeleton is composed of radial poly- Fig. 15. Trachyteleia. Spicule types present in Trachyteleia stephensi (from
spicular tracts between which intermediary tylostyles are scattered; Topsent, 1928c, pI. VI, fig. 11) (abbreviations: Ex, exotyle; Tp, principal
the ectosomal skeleton is composed of a tangential layer of interme- tylostyle; n, intermediary tylostyle; E, ectosomal tylostyle; D, details of
ectosomal tylostyle and principal tylostyle) (scales E, Tp, Ti, 100 j.Lm;
diary tylostyles on which lies a palisade (300 fLm thick) of ectoso- D, Ex, 25 j.Lm).
mal tylostyles reinforced by exotyles which hispidate the surface.
Principal tylostyles are straight with a mucronate head and with a
very thin and acerate extremity: 490-570 X 11 fLm; the intermedi-
ary tylostyles are straight or slightly curved with the same general
shape that the principals: 310-395 X 8 fLm; the ectosomal tylostyles tylostyles and apparently lacks papillae. This allocation requires
are slightly curved: 170-275 X 5 fLm. The exotyles are straight, the verification through checking type material in the National
axial canal is well visible and can be seen to pass through the central Museum of Canada, Ottawa, if it still exists.
cavity of the head; the sharp extremity of the exotyle is rough and
has a length of about 80 fLm: 590-735 X 23 fLm.
Remarks. It is with some hesitation that we decide to retain ACKNOWLEDGEMENTS
this monospecific genus. However, the type preparation and the
description of Topsent are excellent, and we prefer to keep it sepa- I thank the Natural History Museum, London (Shirley Stone
rate from other Polymastiidae until new data become available. & Clare Valentine), the Musee Oceanographique de Monaco (Dr
Suberitechinus de Laubenfels, 1949a (type species Tethea hispida Christian Carpine), the Museum national d'Histoire naturelle, Paris
Bowerbank, 1864b, from off Portland, Maine, NW Atlantic) is cur- (Prof. Claude Levi), the Zoological Museum in Copenhagen (Dr
rently uncertain in its allocation. From its published description Ole Tendal), the Smithsonian Museum (Dr Klaus Rtitzler) and the
and recommendations of Hentschel (1929: 924), it appears to have Zoological Institute of the Academy of Sciences, St Petersbourg
affinities to Polymastiidae rather than Suberitidae, possibly related (Prof. Vladimir Koltun) for having made available slides and type
to Trachyteleia, being strongly hispid, with a cortex, three sizes of specimens for this revision of Polymastiidae.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademicIPlenum Publishers, New York, 2002

Family Spirastrellidae Ridley & Dendy, 1886

Klaus Rutzler

Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington DC, 20560-0163, USA.
(ruetzler.klaus@nmnh.si.edu)

Spirastrellidae Ridley & Dendy (Demospongiae, Hadromerida) is reduced from its traditional definition by excluding massive sponges,
such as Spheciospongia and part of Spirastrella sensu lato, with rare and minute spirasters or amphiasters which are now placed in the
Family Clionaidae. The new definition applies to primarily encrusting sponges with prominent layers of relatively large spirasters, amphi-
asters or diplasters at the surface and the base or throughout the entire body. In contrast to clionaids, tylostyles in spirastrellids are quan-
titatively unimportant and occur in bouquets or simple tracts perpendicular to the substrate, without forming dense networks or tangential
layers. Two genera represent the family, Diplastrella and Spirastrella, with several species that are common in shaded tropical and
subtropical shallow-water habitats including coral reefs.
Keywords: Porifera; Demospongiae; Hadromerida; Spirastrellidae; Diplastrella; Spirastrella.

DEFINITION, DIAGNOSIS, SCOPE History and biology

Synonymy Based solely on the spicule combination of tylostyles and spi-

rasters and ignoring other morphological characteristics, Ridley &
Spirastrellidae Ridley & Dendy, 1886: 490 (part). Choanitidae Dendy (1886, 1887) in their 'Challenger' Expedition monograph
de Laubenfels, 1936a: 140. lumped together incrusting and massive sponges under
Spirastrellidae. This concept prevailed in the literature, particularly
after Vosmaer (1911) reduced more than 30 species (including the
Definition genus type) under the name of the massive species Spirastrella
purpurea (Lamarck) and despite Topsent's (1918) argument
Encrusting Hadromerida with few tylostyles in fascicular against such radical measure. Also de Laubenfels (1950a: 96)
tracts and very abundant and fairly large Streptasters forming pointed out that although sponges such as Spirastrella sensu stricto
cortical and basal crusts. and Spheciospongia had the same spicule combination (tylostyles
and spirasters), the former had incrusting instead of massive
growth form and large and stout instead of delicate and small spi-
Diagnosis rasters. A recent cladistical analysis of clionaids (Rosell & Uriz,
1997) confirmed this separation.
Encrusting sponges with limestone-excavating capability in Members of Spirastrellidae are generally colorful
early stages but without adult endolithic habit. Exhalant canal sys- (orange, red) and dominate many shaded habitats such as caves,
tem apparent at the surface (particularly obvious in live sponges, overhangs, and the lower surfaces of rocks. They are prominent in
contracted in preserved specimens) as meandering, vein-like struc- tropical and SUbtropical shallow-water habitats including coral
tures converging on oscula. Skeleton formed by relatively uncom- reefs.
mon megascleres in ascending radial tracts and dense layers of
microscleres in the ectosomal region and as base layer. Megascleres
mainly tylostyles, tracts starting at the sponge base and ending in Taxonomic Remarks
bouquet-like fashion (spicule points outward) in the ectosomal
region or protruding beyond the sponge surface; some megascleres The family name Choanitidae established by de Laubenfels
oriented at random in the choanosome. Microscleres composed of (1936a: 140) was again rejected (de Laubenfels, 1955a: 140)
large and stout streptasters (spirasters, diplasters), very common because it was shown (Topsent, 1933: 27) that the type of the genus
throughout the sponge but forming extra-dense cortex-like layers in Choanites is a compound ascidian.
the ectosome and at the base (substrate attachment). Massive species among the Spirastrellidae sensu lato, with an
abundance of large megascleres, with rare, minute spirasters and
derivatives, and special incurrent or excurrent features (pore sieves,
Scope papillae) are now assigned to the Clionaidae (Cervicomia, Cliona,
Poterion, Spheciospongia) and are further discussed in that chapter.
As a result of the present revision, four nominal genera are The distinction between Spirastrellidae, Clionaidae, and
included of which two are recognized here as valid: Diplastrella Placospongiidae in this revision is further supported by 28S-rRNA
and Spirastrella. genetic analysis (Chombard, 1998).

220
Porifera· Demospongiae • Hadromerida • Spirastrellidae 221

The genus Diplastrella (formerly in the Timeidae) is included microscleres (diplasters) which, as has already been suggested by
among the Spirastrellidae because of its structure and robust Rosell & Uriz (1997), are obviously derived from spirasters.

KEY TO GENERA

(1) Microscleres composed primarily of spirasters .................................................................................................................. Spirastrella

Microscleres composed primarily of diplasters ................................................................................................................. Diplastrella

DIPIASTRELLA TOPSENT, 1918 Synonymy. Tethya bistellata Schmidt, 1862: 45, pI. VII,
fig. 1 [pI. IV, fig. 1 as stated by Schmidt is incorrect]. Suberites
Synonymy bistellatus; Schmidt, 1864: 36. Hymedesmia bistellata; Topsent,
1900: 125, pI. ill, figs 13 & 16. Diplastrella bistellata; Topsent,
Diplastrella Topsent, 1918: 549. 1918: 549; Vosmaer, 1933: 416. Not Spirastrella bistellata;
Lendenfeld, 1897a: 55, pI. VI, fig. 59, pI. VII, fig. 65, pI. IX,
Type species figs. 120-122; Vosmaer, 1911: 8, pI. 12, fig. 9; Vosmaer, 1933: 416,
pI. 32, figs 12-14, pI. 41, fig. 11, pI. 54, figs 8-9 (=Spirastrella
Tethya bistellata Schmidt, 1862: 45 (by original designation). cunctatrix Schmidt, as was pointed out by Topsent 1918: 549 and
verified by our study of Lendenfeld's slide preparations deposited
Definition at the Naturhistorisches Museum Wien, Austria, catalog number
9099, and USNM, uncatalogued).
Spirastrellidae with diplasters and spherasters as principal Material examined. Holotype: LMJG - not found. BMNH
microscleres. 1867.3.11.16A - slide labelled "Suberites bistellatus Schmidt"
(see Desqueyroux-Faundez & Stone, 1992), re-examined in May,
Diagnosis 2001 (courtesy Ms Clare Valentine), was found to be another
preparation of Spirastrella cunctatrix, thus making it necessary to
Thin incrustations of typical spirastrellid structure, with erect establish a neotype. Neotype: USNM 2022309 (accession number)-
tylostyles in bouquet-like arrangement and a basal layer of diplas- Tremiti Islands, Adriatic, Italy, Grotta Bue Marino, 1.5 m depth,
ters and spherasters, and an ectosomal crust of diplasters. coli. G. Corriero, 30.v.2001 (courtesy of G. Corriero); collected
Modification of these spicules include spirasters, but they occur in from very near the type locality which is off the southern coast of
small numbers, relative to the other microscleres. the Adriatic island Lesina (now, Hvar, Croatia).
Description. The original author described the sponge as
Previous review a red incrustation (which he considered ajuvenile) with tylostyles,
spirasters, and diplasters (Schmidt, 1862: 45; pI. VII, fig. 1, shows
Topsent (1918); for more illustrations refer to Riitzler & Sara a tylostyle head portion and one of the characteristic diplasters).
(1962). The neotype was reported to be pale yellow in life (G. Corriero
pers. comm.). Schmidt may have confused the color with that of
Description of type species co-occurring Spirastrella cunctratrix because only yellowish,
whitish pink, and whitish brown forms are known from underwater
Diplastrella bistellata (Schmidt, 1862) (Figs 1-3). photographs of cave walls (for instance, in Harmelin et al., 1987:

A __
I
,I

~ ij ~ i$*~
c_ E

Fig. 1. Diplastrella, habitus and anatomy. A-B, after Riitzler & Sara (1962). C-E, D. bistellata Schmidt (neotype USNM acc. no. 2022309). A, detail of
surface of live specimen of D. ornata Riitzler & Sara (scale 0.5 mm). B, cross-section of D. ornata illustrating skeleton structure (scale 200 /-Lm).
C, tylostyles (scale 10 /-Lm). D, large diplasters (scale 10 /-Lm). E, small diplasters/spirasters (scale 10 /-Lm).
222 Porifera· Demospongiae • Hadromerida • Spirastrellidae

129; M. Panzini & R. Pronzato, pers. comm.). Surface appearance or oval head outlines and taper to a rounded (most common),
and anatomy of this species is similar to that of Spirastrella (see stepped, or sharp point. They measure 450-630 (515) fLm X 10-13
below). The neotype is a very thin (0.5 mm) crust covering an area (4.7) fLm, with 13-18 (15.2)fLm heads. Diplasters fall into two size
of about 20 X 30 mm; collapsed subsurface canals are faintly classes. The larger ones are stout and appear like large spherasters
visible. The color in alcohol is a pale drab. Tylostyles have circular except that the spines originate from two closely spaced centers.
Many of the smaller size are more distinctly amphiastrose, with
a short but smooth shaft and distally divergent spines, but there
are also spiraster-like modifications. Diplasters: (I) 25-45 X
22-42 (34.8 X 40.3)/-1m; (II) 11-20 X 8-15 (13.6 X 11.3)/-1m.
In other specimens described before, tylostyles were noted to
have rounded points (approaching tylostrongyles) and a few of the
larger diplaster were reported with rays split near the distal end
(Sara, 1961). Several authors confirmed the abundance of this
characteristic species in the Mediterranean, particularly in shaded
habitats, such as shallow caves and under rocks (Topsent, 1918;
Sara, 1961).
Remarks. Topsent (1918: 549) points out that a misplaced
label at the Graz Museum has led to Lendenfeld's (1897a) and
Vosmaer's (1911) misinterpretation of this species.

SPIRASTRELIA SCHMIDT, 1868

Synonymy

Spirastrella Schmidt, 1868: 17. Chondrillina de Laubenfels,

1936a (for Chondrilla phyllodes Schmidt, 1870). Taxonomic
decision for synonymy: Wiedenmayer, 1977b: 162.

Type species

Spirastrella cunctatrix Schmidt, 1868: 17 (by monotypy).

Fig. 2. Diplastrella bistellata Schmidt (neotype USNM 2022309), SEM Definition

images of spicules. A, tylostyles and diplasters. B, tylostyle point enlarged.
C, large diplasters. D, small diplasters/spirasters (all scales 10 !Lm). Spirastrellidae with spirasters as principal microscleres.

Fig.3. Diplastrella megastellata Hechtel (paratype USNM 24503), SEM images of spicules to show extreme development of diplasters. A, tylostyle (scale
20 !Lm). B-C, diplasters (scales: B, 10 !Lm; C, 2!Lm).
Porifera' Demospongiae • Hadromerida • Spirastrellidae 223

B c_ E
Fig. 4. Spirastrella cunctatrix Schmidt, habitus and anatomy (USNM 23944). A, habitus from an underwater photograph (scale lOmm). B, cross section
showing skeleton arrangement (scale 1 mm). C, typostyles (scale 10 j.Lm). D, large spirasters (scale 10 j.Lm). E, small spirasters (scale 20 j.Lm).

Previous review

Vosmaer (1911), Topsent (1918, 1938b).

Description of type species

Spirastrella cunctatrix Schmidt, 1868 (Figs 4,5).

Synonymy. Spirastrella cunctatrix Schmidt, 1868: 17, pI. 3
fig. 8. NotWiedenmayer, 1977b: 162 (=Spirastrella mollisVerrill).
Material examined. Holotype (fragments): MNHN DT 735
(1 slide, labeled "type, Algerie"); BMNH 1868.3.2.4 (1 slide,
labeled "Algier", original balsam spicule mount by Schmidt) -
Coast of Algeria. Other material. USNM 23944 - specimen pho-
tographed and collected in a shallow cave near Rovinj, Croatia
(Riitzler, 1965).
Description. Live specimens are dull pinkish-red crusts, 5-10
cm or more in diameter, 1-5 mm thick, depending on the state of con-
traction. They are soft and show vein-like, branching surface canals
converging toward elevated oscula, as it is common for encrusting
species, with incurrent pores dispersed in between. The ectosomal
membranes that cover the canals collapse upon fixation, leaving a
pattern of radiating grooves; preserved sponges are leathery tough,
with rough surface texture, and about 1 mm thick. Topsent (1938b)
describes the type specimen as a faintly pinkish (in alcohol) crust,
5 X 8 cm, 1 mm thick. In cross section, bundles of a few tylostyles
each extend from the base to the surface of the crust, spicule points
directed outward. The smaller spirasters are condensed near the sur-
face, the larger ones near the base, but there are many of both types
Fig. 5. Spirastrella cunctatrix Schmidt, SEM views of spicules (USNM strewn in between. Tylostyles have rounded to oval, pronounced
23944). A, tylostyle and spirasters (scale 20 j.Lm). B, large spiraster (scale heads from which they taper gradually to a sharp point; they measure
5 j.Lm). C-E, small spirasters (scales 2 j.Lm). 480 X 8 /-lm on average. Small spirasters vary 10-25/-lm in length,
5-23/-lm in width (including spines), large spirasters are 28-50 X
Diagnosis 23-35/-lm.
Remarks. Vosmaer (1911) considered this species a junior
Spirastrellidae with dense layers of spirasters in both ectosomal synonym of Spirastrella purpurea (Lamarck), and later (Vosmaer,
and basal choanosomal regions of the sponge. Spirasters are very 1933) synonymized it with Diplastrella bistellata (Schmidt).
common and of considerable size range; tylostyles comparatively Topsent (1918) argued strongly and correctly against this unjusti-
small and rare. fied lumping of distinctive species.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Stylocordylidae Topsent, 1892

Rob W.M. Van Soest

Zoological Museum, University of Amsterdam, P.O. Box 94766,1090 GT Amsterdam, The Netherlands. (soest@science.uva.nl)

Styolocordylidae Topsent (Demospongiae, Hadromerida) is a monotypical family with characteristic club-shaped habit. They are consid-
ered closely related to Suberitidae from which they differ in the possession of long centrotylote megascleres building the stalk and form-
ing the central core of the main body. The single genus Stylocordyla occurs in deeper water in all three oceans.
Keywords: Porifera; Demospongiae; Hadromerida; Stylocordylidae; Stylocordyla.
- - - - _ .. __ . - - - _..__ ... _ - - - - - _ . - _ . _ - _ - . - - - - -. . _ - -
... -'-- ---_._--

DEFINITION, DIAGNOSIS, SCOPE latrunculiids, and they are also obviously unrelated to
Stylocordylidae. Podospongiidae shows superficial similarities to
Synonymy Suberitidae, particularly with the stipitate genus Rhizaxinella
Keller, 1881, but is clearly a poecilosclerid with suspected affini-
Stylocordylidae Topsent, 1892: 58. ties to Mycalina families (see chapter on Podospongiidae). Both
Vosmaer (1887: 331) and Ridley & Dendy (1887: 222) assigned
Definition Stylocordyla to Suberitidae. However, from this family
Stylocordyla differs in the absence of tylostyles and the possession
Hadromerida with stipitate habit, and centrotylote oxeas or of tangential arrangement of small oxeas. With Aaptos Vosmaer,
strongyles in radial arrangement; usually also with smaller partly 1887 it shares the possession of oxeote spicules and peripheral
tangential oxeas or stylotes at the surface. smaller megascleres intermingled with the radiating megascleres,
however the megascleres of Aaptos are strongyloxeas without any
Diagnosis centrotylote swellings, and it has no tangential spicules. Hooper &
Wiedenmayer (1994: 402) suggested the family is incertae sedis
Deep-water sponges with thin stalk and globular-oval 'body', and possibly belongs in Ancorinidae having lost their triaenes. In
which is flattened at the top. Skeleton of the main body radially view of the centrotylote spicules and axial orientation of the skele-
arranged. Megascleres include centrotylote oxeas, with strongy- ton, this is considered unlikely.
lote, or occasionally stylote modifications; in addition many
species have smaller oxeas or styles at the surface, partly arranged Previous reviews
in a tangential layer both on the main body and lining the stalk.
Reproduction is viviparous, with the added peculiarity that larvae Burton (1934c), Bergquist (1972), Hooper & Wiedenmayer
are kept in the body until they have developed into young adults (1994).
(Bergquist, 1972). Distribution is cosmopolitan, with a predomi-
nance of records from cold water.
STYLOCORDYLA THOMSON, 1873
Scope
Synonymy
Four nominal genera, only one of which is considered valid.
Stylocordyla Thomson, 1873: 113. Stylorhiza Schmidt, 1880:
Taxonomic history 79 (Not Haeckel, 1880: 612). Oxycordyla Topsent, 1904b: 135. ?
Microcordyla Zirpolo, 1927: 290.
The family was erected by Topsent (1892: 58) for Stylocordyla
with the simple definition 'Aciculida without microscleres', because Type species
he considered the centrotylote oxeas related to those of Dorypleres
and other triaene-less Astrophorida. However, he noted the parallel Hyalonema boreale Loven, 1868: 105 (by original designation).
with Suberitidae in his order Clavulida (now Hadromerida).
The genus Tethycordyla de Laubenfels, 1934 was assigned to Definition
this family by Levi (1973), who also considered Podospongiidae
de Laubenfels, 1936 a junior synonym of Stylocordylidae. As for the family. Stalked Stylocordylidae with centrotylote
However, Tethycordyla possesses asters and is obviously closely megascleres arranged poly serially in the stalk and in an axial-radial
related to Tethya. It is a member of Tethyidae (see chapter on fashion in the main-body. At the surface and scattered in the body
Tethyidae). Podospongia and Podospongiidae have streptaster-like are smaller oxeas andlor styles. About 12 nominal species and
microscleres (spinose microrhabds, or spinorhabds) superficially 'varieties' have been described from all over the world oceans from
resembling acanthodiscorhabds, or 'chessman' spicules of the depths between 13 and 3000m.

224
Porifera· Demospongiae • Hadromerida • StyJocordyJidae 225

rio'

r..,

)~"
ro.

\'
f
r...

" . (((~. ~" rff'1J

I",.J

Fig. I. Stylocordyla borealis (Loven, 1868). A, reproduction of Loven's (1868) plate I, with drawings of habit, cross section and spicules of the type spec-
imens (sizes see text). B, schematic cross sections of the main body, reproduced from Ridley & Dendy's (1887) text-fig. 9. C, Schmidt's drawing of
Stylorhiza stipitata considered a junior synonym (size see text). D-G, specimen ZMA POR. 2224 from the Arctic Ocean. D, photo of habit (scale I cm).
E, SEM photo of middle sized centrotylote megasdere from the choanosomal region (scale 10 !Lm). F, small oxea from the ectosomal region (scale 10 !Lm).
G, styloid end of a small oxeote from the ectosomal region (scale I !Lm).

Description of type species Material examined. Type material (not examined) is stated
to be in the Stockholm Museum, consisting of two specimens, the
Stylocordyla borealis (Loven, 1868) (Fig. lA-F). specimen illustrated by Loven, 1868 from 360 m off the N coast of
Synonymy. Hyalonema boreale Loven, 1868: 105, pI. 2; Norway (Finmarken), is here designated as lectotype, and a second
Stylocordyla borealis; Thomson, 1873: 113; Topsent, 1896c: 286, specimen from the coast of Norway collected by G. von Yhlen
pI. vm, figs 11-15; Stylocordyla borealis var. typica Burton, 1934c: is the paralectotype. Other material. ZMA 2224 - 'Varna'
13 (with further synonyms). Polymastia stipitata Carter, 1876: 393. Exped. 1883, Arctic Ocean, (det. E. Arnesen). No further data.
226 Porifera' Demospongiae • Hadromerida • Stylocordylidae

Description. Club-shaped on long thin stalk, provided with conspecific with the North Atlantic material of Carter. The genus
a proliferated root system. The lectotype is 5.2cm long, with a name Stylorhiza was also used for a cnidarian by Haeckel (1880).
body of 1.3 cm and a stalk of 3.9 cm. The oval body is somewhat Oxycordyla Topsent, 1904b: 135 was erected for O. pellita Topsent,
flattened at the top and has an optically smooth surface. A single 1904b: 135 (by monotypy), and subsequently it was synonymized
oscule is apparent, to the side of the body near the flattened top. with Stylocordyla by Bergquist (1972: 128). The type (in MOM, not
Skeleton of the stalk made up of a tight mass of aligned larger reexamined) has the shape and structure of Stylocordyla, but was
megascleres, somewhat spirally arranged. The stalk penetrates distinguished by the absence of small monactinal or diactinal sur-
macroscopically far into the main body and from it radiating bun- face spicules. The only spicules noted by Topsent were long thin
dles of larger megascleres fan out towards the surface. Small oxeas of 1500--2500 X23 /-Lm. It is proposed here to follow
spicules form a palisade at the surface in between the radiating Bergquist's opinion and assign this species to Stylocordyla. The
bundles of megascieres. Small spicules also form a dense tangen- genus Microcordyla Zirpolo (1927: 290, fig. unnumbered) was
tial crust. Spicules, oxeas, often with rounded or strongylote end- erected (by monotypy) for Microcordyla asteriae Zirpolo, 1927. The
ings, often centrotylote, in three size categories, 790--2930 X description and very crude drawing of the habit are inadequate for
10--47/-Lm, 400--1140 X 11-13 /-Lm, and 340--450 X 4-4.5 /-Lm. definite classification. The stalked specimen of only 7 mm length
Ectosomal microxeas, 70--100 X 1.5-2m. Distribution and apparently was attached to one of the arms of an asteroid of the genus
ecology. North Atlantic, deep water. Asterias, with three peculiar-shaped uncinate attachment organs. The
Remarks. Judging from the records of the type species its vague reference to oxeas, the description of the cup-shaped oscule
distribution is bipolar and includes large sections of the North and with a circle of spine-like 'spicules', and the apparent 'muscular'
South Atlantic and Pacific Oceans. Hooper & Wiedenmayer (1994) nature of the attachment organs all contribute to the impression that
considered Southern Ocean 'varieties' described by several authors this is not a stylocordylid sponge, but for a firm conclusion the type
(var. globosa Ridley & Dendy, 1887: 224; var. acuata Kirkpatrick, material, presumably lodged in the collections of the Stazione
1908c: 22; var. irregularis Hentschel, 1914: 55) conspecific with the Zoologica at Naples, will have to be studied.
nominal populations from the North Atlantic, but this is here left
undecided in view of the apparent discontinuous distribution.
Stylorhiza Schmidt, 1880: 79 was erected for type species ACKNOWLEDGEMENTS
Polymastia stipitata Carter, 1876: 393 (by original designation).
This is here considered a junior synonym of S. borealis, although Professor Roberto Pronzato is thanked for making a copy of
it is uncertain whether Schmidt's Gulf of Mexico material was Zirpolo's paper available to me.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Suberitidae Schmidt, 1870

Rob W.M. Van Soest

Zoological Museum, University of Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, The Netherlands. (soest@science.uva.nl)

Suberitidae Schmidt (Demospongiae, Hadromerida) is characterized by the absence of a cortex and microscleres other than microrhabds
or trichodragmas. Spicules are for the most part tylostyles, often in several size categories; in exceptional cases, oxeas or strongyloxeas
are the main megascleres. Usually Suberitidae have an ectosomal specialization in the form of a palisade or brushes of spicules making
the surface velvety or microhispid in texture (one genus excepted). The interior skeleton tends to be irregular, confusedly reticulate, or
condensed, and in general lacks a clear radial arrangement (one genus excepted). There are massive-globular, arborescent-stalked,
or thinly encrusting growth forms. Many species and genera live in sedimented environment, often at some depth, either by rooted stalks
or by living on the empty shells of gastropods occupied by hermit crabs which carry the sponges around. The family in its present concept
includes 26 nominal genera, eleven of which are considered valid. One commonly used genus name, Laxosuberites, was found to be
untenable, and species assigned to it are for the most part transferred to a revived genus name Protosuberites.
Keywords: Porifera; Demospongiae; Hadromerida; Suberitidae; Aaptos; Caulospongia; Homaxinella; Plicatellopsis; Prosuberites;
Protosuberites; Pseudospongosorites; Pseudosuberites; Rhizaxinella; Suberites; Terpios.
-----

DEFINITION, DIAGNOSIS, SCOPE History and biology

Synonymy Taxonomic history. Schmidt (1870) included in his concept of

Suberitidae sponges without cortical structures, tylostyles in tracts or
Suberitidinae Schmidt, 1870: 46. Suberitidae Vosmaer, in confusion and at the periphery directed outwards. He included next
1887: 330. to Suberites, the genus Papillina (now in Clionaidae), and furthermore
a number of genera with cortical structures: Radiella, Polymastia in
part, Thecophora, Rinalda (all four now Polymastiidae), and
Definition Cometella and Tethya (now Tethyidae), thus violating in typical
Schmidt manner his own definition. Vosmaer (1887) separated the
Hadromerida without cortex and without microscleres other polymastiid genera from the Suberitidae, and added Rhizaxinella,
than microrhabds; spicules tylostyles or styles, exceptionally Quasillina (now Polymastiidae), Poterion (now Clionaidae) and
oxeas, arranged in brushes or palisade in the peripheral region. Stylocordyla (now Stylocordylidae). Subsequently, Lendenfeld
(1896) united the Polymastiidae and Suberitidae again, but split off
Stylocordylidae. Topsent's (1900) revision of the 'Clavulina' of
Diagnosis France was the first to define the family in its present outline. He
defined Suberitinae as Clavulina without microscleres other than
Globular, ramose, StIpltate, massive or encrusting habit. microrhabds and rhaphides, without a cortex, and a skeleton having
Megascleres usually tylostyles, occasionally styles, strongyloxeas no strictly radiate arrangement of the spicules. He included Ficulina
or centrotylote oxeas; microscleres usually absent, when present Gray, I 867a, Laxosuberites Topsent, I 896a, Pseudosuberites
confined to microrhabds and trichodragmas. In cross section, Topsent, 1896a, Prosuberites Topsent, 1893c, Rhizaxinella Keller,
megascleres are usually arranged in bouquets at the surface, in 1880a, Suberites Nardo, 1833 and Terpios Duchassaing & Michelotti,
more massive species becoming progressively confusedly arranged 1864. He assigned Aaptos (as Tuberella) to Tethyidae. Burton
towards the interior, but overall structure may also be strictly radial (1930d) more-or-Iess followed Topsent's classification, adding
or show a strong axial orientation. In one genus the spicules at the Caulospongia Saville Kent, 1871, Protosuberites Swartschewsky,
surface are arranged tangentially. There is no recognizable cortex. 1905, and erected two additional genera Laxosuberella (here consid-
In thinly encrusting species spicule orientation is either parallel or ered Clionaidae) and Terpiosella. He also included Tentorium
perpendicular to the substratum. Modifications of shape and posi- Vosmaer, 1885a (here considered Polymastiidae) and Axosuberites
tion of the tylostyle heads are common; they can be lobate, pear- Topsent, 1893a (here considered Microcionidae). Burton (1930d)
shaped, drop-shaped or subterminal. also indicated that in his view the differences between Polymastiidae
and Suberitidae were questionable. Perhaps inspired by this remark,
de Laubenfels (1936a: 147) expanded the family to include amongst
Scope others Polymastiidae by maintaining a simple definition of "compris-
ing genera having a spiculation of tylostyles as megascleres, and no
Eleven genera (out of 26 nominal genera) are considered valid: microscleres at all". This is followed by an unprecedented and partly
Aaptos, Caulospongia, Homaxinella, Plicatellopsis, Prosuberites, unexplained plethora of 40 genera, several of which did not even con-
Protosuberites, Pseudospongosorites, Pseudosuberites, Rhizaxinella, form to his simple definition. It is beyond the scope of these present
Suberites and Terpios. brief comments to list and discuss each of his assignments.

227
228 Porifera· Demospongiae • Hadromerida • Suberitidae

Remarkably, de Laubenfels (1936a: 143) assigned Ficulina (as stalked or arborescent forms with root systems for attachment, or by
Choanites Mantell, 1822) to a family Choanitidae, along with encrusting empty gastropod shells with symbiont hermit-crabs, carry-
spiraster-bearing genera Spirastrella and Spheciospongia. ing the sponge around. In addition, several genera are thinly encrusting
Understandably, his classification found few followers, and subse- on shells, rocks and corals. Colder, deeper waters are favored by many
quent authors (e.g., Levi, 1973; Bergquist, 1978; Hartman, 1982; suberitids, although most genera have representatives in shallow tropi-
Voultsiadou-Koukoura & Van Soest, 1993; Riitzler & Smith, 1993) cal environments. Several species and genera have a means of asexual
followed basically Topsent's (1900) scheme, with later additions reproduction by gemmule-type of bodies, formed at the contact area of
mentioned above. The present concept in its turn deviates from that the substratum, especially in forms living on gastropod shells. These
scheme only on details of inclusion of individual genera. For example, gemmules are not structurally comparable with the elaborate gem-
the genus Poterion is often included in Suberitidae (e.g., Hooper & mules of freshwater sponges, but may have similar functions.
Wiedenmayer, 1994: 404), but is here referred to Clionaidae. Chemistry. A chemical marker for at least two genera of the
Biology. Suberitid sponges include many forms capable of liv- family is the group of compounds dubbed 'aaptamines' (Bergquist
ing in sedimented environments, either through the development of et al., 1991a; Van Soest & Braekman, 1999).

KEY TO GENERA

(1) Sponge stipitate or arborescent, i.e. having a stem of some length separate from further body parts (ovate bodies, branches, lobes,
leaf-like structures) .... ... ....... ...... .... .................... ... ........... ...... .......... ........ ....... ...... ...... .......... ..... ....... ....... ... ...... .......... ................. ...... 2
Sponge globular, massive or encrusting; if erect, there is no distinct stem ..... ... ... ...... ... .... ....... ............ ... .................. ................... .... 5
(2) Sponge arborescent, with branches similar to the stem .................................................................................................................... 3
Sponge stipitate with ovate body, or provided with lobes or leaf-like structures .............................................................................. 4
(3) Spicules predominantly styles, extra-axial skeleton predominantly with single spicules arranged in brushes at the
surface ............................................................................................................................................................................ Homaxinella
Spicules predominantly tylostyles; extra-axial skeleton with strong spicule bundles fanning out at the
surface to form brushes .............. ................ ...... .... ................. ........... ... .... .... ... ... .... ...... ...... .... .... ....... .... .... ... ...... ............. Plicatellopsis
(4) Sponges with simple stem and ovate body, which may be pearshaped or otherwise elongated; brushes of smaller
spicules over radiate bundles of larger spicules . ..... ............. ........... ... .... ... .... ... .......... ... ... ............... .... ... .... ............... ..... Rhizaxinella
Sponges with elaborate shape, either pagoda-like with leafshaped lobes, or flabellate or a system of reticulate
branches; tylostyles in a single size category, with peculiarly flattened tyles .............................................................. Caulospongia
(5) Sponge globular, massively lobate or digitate, or massively encrusting ........................................................................................... 6
Sponge thinly encrusting, usually not exceeding a cm in thickness ...... ........... ................ ........... .... ........... ... ............. ................. ...... 9
(6) Spicules exclusively tylostyles .......................................................................................................................................................... 7
Spicules include oxeas or strongyloxeas ........................................................................................................................................... 8
(7) Surface skeleton is a tangential and usually detachable crust carried by subectosomal spicule brushes ................. Pseudosuberites
No tangential spicules, surface velvety due to palisade of small tylostyles ......................................................................... Suberites
(8) Oxeas symmetrical relatively small and sharply pointed; skeletal arrangement more or less
confused .......... ................. ..... ... ....... ...... ...... .... ........ ... ................. ....... .............. ...... ............. ... .... ........... ............ Pseudospongosorites
Strongyloxeas in a large size range, usually accompanied by small styles and/or tylostyles in a low quantity;
skeletal arrangement strictly radiate ..... ...... ................ ................ ........... .................................. .............. ................. ............... ... Aaptos
(9) Tylostyles single erect on the substrate ........................................................................................................................................... 10
Tylostyles in bundles or confused ................................................................................................................................................... 11
(10) Macroscopically hispid, no ectosomal specialization ..................................................................................................... Prosuberites
Macroscopically smooth, at the surface tylostyles in brushes ..................................................................................... Protosuberites
(11) Low spicular density, gelatinous consistency, tylostyles predominantly with peculiarly flattened, lobate tyles; size
uniformly small ....................................................................................................................................................................... Terpios
Tylostyles in a dense palisade at the surface and in thicker specimens arranged in parallel bundles; usually
a large size range; lobate tyles not common ................................................................................................................ Protosuberites

AAPTOS GRAY, 1867 Definition

Synonymy Suberitidae with radiate skeleton of strongyloxeas in three size

categories, the smaller of which form a dense ectosomal palisade.
Aaptos Gray, 1867a: 519. Trachya Carter, 1870a: No microscleres.
176. Tuberella Keller, 1880a: 277. Tethyophaena Schmidt,
1880a: 281. Diagnosis

Type species Lobate or spherical sponges with a radial skeleton, often con-
sisting of confluent globular or lobate units. Surface smooth or
Ancorina adriatica Gray, 1867a: 519 (by original designa- tuberculate-papillate, usually rough to the touch. Some species
tion) [=Aaptos aaptos (Schmidt, 1864: 33)]. show a distinct colour change when taken out of the water. In cross
Porifera· Demospongiae • Hadromerida • Suberitidae 229

c
Fig. 1. Aaptos aaptos (Schmidt, 1864). A, paralectotype LMJG 15705 from unknown locality (scale 1 em). B, cross section of peripheral skeleton of
lectotype BMNH 1910.1.1.866, Lagoste, Algeria (reproduced from Kelly-Borges & Berquist, 1994: pI. Ia) (scale 1000 !Lm). C, spicules (reproduced from
Lendenfeld, 1896: pI. VII fig. 87) (scale 100 !Lm).

section, the outer region is often fibrous and may be considered as a Material examined. Lectotype (designation by Kelly-
cortex which grades into the choanosome. Skeleton strictly radiate, Borges & Bergquist, 1994: 304): BMNH 1910:1:1:866 - Lagoste,
with tracts and single spicules issuing from the centre of the lobe or Algeria. Paralectotype (designated herein): LMJG 15705 -
globular body. At the surface the tracts fan out and form a dense pal- Schmidt specimen of unknown locality in the Graz Museum
isade consisting of smaller spicules intermingled between the ends of (cf. Desqueyroux-Faundez & Stone, 1992: 7, pI. ill fig. 22, here
the larger spicules. Spicules are strongyloxeas, in three overlapping reproduced as Fig. lA). Other material. ZMA POR. 6164 -
size categories; the intermediate and smaller spicules are occasion- Prainha, Madeira, 2m, colI. R.W.M. van Soest, 3 April 1986.
ally oxeas, styles or tylostyles. The genus is cosmopolitan and cur- Description. Massive, lumpy, bluntly lobate (Fig. lA), typi-
rently about 15-20 species have been described, but several more cally 3-5 cm in diameter, occasionally larger, fist-size. Surface irreg-
may hide among the widespread records of Aaptos aaptos. Several ular, hispid, occasionally smooth, seldom elevated in distinct
species produce a distinctive compound aaptarnine (Van Soest & papillae. Oscules up to 3 mm in diameter, few in number, contractile,
Braekman, 1999), which appears to be a good marker for the genus. occasionally elevated. Consistency compact, firm, hard. Colour
brown-yellow, occasionally pale yellow or greyish, or blackish
Previous reviews brown. Interior yellow. Skeleton architecture (Fig. IB) is radiate with
spicule tracts of 1.5-2 cm length, 10--20 spicules in thickness, radiat-
Kelly-Borges & Bergquist, 1994: 303. ing from a focal point towards the surface. At the surface these
bundles are strengthened and linked by brushes of small styles, mak-
Description of type species ing a continuous ectosomal palisade of about 1 mm in thickness,
which is however not macroscopically distinct and not fibrous.
Aaptos aaptos (Schmidt, 1864) (Fig. lA-C). Inhalant canals traverse the peripheral region in a sinuous way.
Synonymy. Ancorina aaptos Schmidt, 1864: 33, pI. IV Below this the choanosomal tracts are separated by subectosomal
fig. 11; Aaptos aaptos; Vosmaer, 1887: 327; Kelly-Borges & cavities. Spicules (Fig. lC) strongyloxeas and styles. The strongylox-
Bergquist, 1994: 304, pI. Ia figs 2a-b; Suberites aaptos; eas are characteristically thickest in the middle, thinning out towards
Lendenfeld, 1896: 140, pI. vn XII; Tuberella aaptos; Topsent, the blunt apex, and are divisible in two size categories, 1500--1900 X
1900: 285, pI. Vill figs 12, 13. Aaptos adriatica Gray, 1867a: 519. 12--45,....m and 750--1000 X 10--25 ,....m. Ectosomal small styles,
Tuberella tethyoides Keller, 1880a: 277. Tethyophaena silifica occasionally subtylostyles: 135-230 X 1-5,....m (data from the
Schmidt, 1880a: 281. Suberites spissus Topsent, 1892b: xxvii. holotype, taken from Kelly-Borges & Bergquist, 1994: 306).
230 Porifera' Demospongiae • Hadromerida • Suberitidae

Cell biology. The choanosome is rich in spherulous cells. The distinct flattened tyles in a single size category. No microscleres.
choanocyte chambers are small, eurypylous, 20-25 fLm in diame- The genus is exclusively Australian in distribution and contains
ter. Distribution and ecology. Mediterranean and also occurring off eight species.
the coasts of NW Spain and Portugal, Azores, Madeira and Canary
Islands. Mostly known from deeper water, but occasionally occur- Previous review
ring in the shallow sublittoral.
Remarks. Gray (1867a) erected a species of his own Fromont, 1998: 67.
(Aaptos adriatica) for Schmidt's Ancorina aaptos for no apparent
reason and without indicating material other than Schmidt's; con- Description of type species
sequently this is an objective junior synonym. The type species is
not well-known. It has been reported from many areas of the world, Caulospongia perfoliata (Lamarck, 1814) (Fig. 2A-D).
but it is almost certain that these conceru other species of Aaptos. Synonymy. Spongia perfoliata Lamarck. 1814: 439;
The genus Trachya Carter (1870a: 176) was erected for the Caulospongia perfoliata; Hallmann, 1914: 306, pI. 18 fig. 1.
West Indian species (by monotypy) Trachya pemucleata Carter Caulospongia verticillata Saville Kent, 1871: 616, pI. XLVIII
(1870a: 176). It was re-examined by Kelly-Borges & Bergquist fig. 1; Chalina verticillata; Bowerbank, 1876b: 769, pI. LXXIX.
(1994: 316) and assigned to Aaptos, which is confirmed here. Material examined. None. Ho10type: MNHN DT 3368 -
The genus Tuberella Keller, 1880a was erected for type extensively redescribed by Topsent, 1932: 85 and Fromont, 1998:
species (by monotypy) Tuberella tethyoides Keller, 1880a: 277, 68. The type of C. verticillata, from North Australia, was stated to
pI. XIV figs 7-9). The type material, supposedly in 2MB, was not be in the Liverpool Free Museum and is now considered lost; it is
reexamined, but the species has been discussed by Vosmaer (1887: described and figured in Saville Kent, 1871.
327, pI. XXVI fig. 9) and Topsent (1900: 287), making it clear that Description (from Fromont, 1998: 68). Erect pagoda-like
it is an Aaptos species closely similar to and probably conspecific sponge (Fig. 2A, B) with a solid stem and flattened irregular leaf-
with A. aaptos. Tuberella is thus a clear synonym of Aaptos. like lobes which are hollow inside and are sloping upwards towards
The genus Tethyophaena Schmidt, 1880a was erected for type the apex of the sponge. Length 30-36 cm, with a stem of 5-13 cm,
species (by monotypy) Tethyophaena silifica Schmidt, 1880a: 281. lobes 6-9 cm wide around the stem, 0.3-0.5 cm in thickness.
Although Desqueyroux-Faundez & Stone (1992) do not list any Surface smooth, porous, oscules not apparent. Texture very firm
extant type specimens, there is an unnumbered specimen in the but compressible. Colour cream or yellow-orange. Ectosomal
Leiden Museum from Naples with label in Schmidt's handwriting skeleton (Fig. 2C) as defined for the genus with both brushes of
('type'). This is here designated the lectotype. It is an obvious spicules and tangentially arranged spicules together forming
specimen of Aaptos aaptos (Schmidt himself likened his new a detachable surface crust of 90-150 fLm in thickness. Choanosomal
species to Tuberella tethyoides). skeleton (Fig. 2C) is a reticulation of spongin-encased spicule tracts
consisting of primary tracts of about 60 fLm diameter with a core of
five or more spicules and interconnecting thinner tracts of 30 fLm
CAULOSPONGIA KENT, 1871 diameter and a core of 2-4 spicules. In the stem, spicule tracts may
be as thick as 320 fLm. Spicules (Fig. 2D) tylostyles with character-
Synonymy istic flattened tyles which frequently are lobate, in a single size cat-
egory, 118-215 X 2.5-10 fLm. Distribution and ecology. SW
Caulospongia Kent, 1871: 616. Plectodendron Lendenfeld, Australia, on rocks among kelp, II-30m.
1888: 66. Remarks. This is a clearly demarcated species and genus
through the combination of elaborate shape, reticulate skeleton
Type species and flattened tyles of the spicules. The genus Plectodendron
Lendenfeld, 1888: 66 was erected for Plectodendron elegans
Caulospongia vertic illata Kent, 1871: 616 (by subsequent Lendenfeld, 1888: 66 (by monotypy). Hallmann (1914: 306, pI. 18
designation; Hallmann, 1914: 306) (=junior synonym of Spongia fig. 1) assigned the species to Caulospongia, and this was followed
perfoliata Lamarck, 1814: 439). by Fromont (1998: 78). The lectotype (designated by Fromont, 1998:
79), AM G9186, from Broughton Island, New South Wales, and para-
Definition lectotypes AM Z5221 and AM Z5271 , both from Port Jackson, NSW,
were redescribed by Fromont, 1998. Although the gross morphology
Suberitidae with erect pagoda-like or lobed-lamellated habit deviates quite substantially from the type species of Caulospongia,
and reticulate skeleton of spicule tracts. Surface skeleton with tan- the skeletal and spicule characters are the same. Moreover, a further
gential spiCUles carried by ectosomal brushes. Spicules tylostyles species of Caulospongia, C. reticulata Fromont, 1998: 80, bridges
with flattened-lobate tyles. the gap between the habits of C. perfoliata and Plectodendron
elegans. The two genera are here confirmed as synonyms.
Diagnosis

Erect sponges with solid stem and either a lobed or interlock- HOMAXINELLA TOPSENT, 1916
ing, reticulate branching habit or forming erect fans. Choanosomal
skeleton is a p1umo-reticulate or reticulate system of spicule tracts; Synonymy
interstitial spicules are frequent. Ectosomal skeleton consists of
brushes of spicules covered by a narrow or diffuse layer of spicules Homaxinella Topsent, 1916: 4. Pachaxinella Burton,
parallel to the surface. Spicules relatively small tylostyles with 1930b: 503.
Porifera· Demospongiae • Hadromerida • Suberitidae 231

Fig.2. Caulospongia perfoliata (Lamarck, 1814). A, drawing of holotype of Caulospongia verticillata Kent, 1871 = jun. syn. of C. perfoliata, reproduced
from Kent, 1871: pI. XLVIII, fig. 1 (scale Scm). B, photo of holotype of Spongia perfoliata, MNHN DT 3368 (reproduced from Topsent, 1932: pI. III
fig. 4) (scale 1 cm). C, cross section of peripheral skeleton with marked difference between ectosomal and choanosomal arrangement (reproduced from
Fromont's 1998: fig. 2c) (scale 100ll-m). D, spicules (reproduced from Fromont's 1998: fig . 3a) (scale IOll-m).

Type species Collection, 36-108 m. Paralectotype of Axinelia supratumescens:

MNHN 760 - 'Charcot' Expedition 1905, labeled 'type'.
Axinella supratumescens Topsent, 1907: 74 (by original des- Description. Ramose (Fig. 3A), with branches up to 20 cm
ignation), (=generally considered a junior synonym of Axinella long and max. 0.8 cm diameter, at branching points up to 1.5 cm,
balfourensis Ridley & Dendy, 1887: 480). they may appear somewhat flattened. Branches divide, but seldom
anastomose. Stalk is attached to the substrate by a system of roots
Detinition and/or creeping stolons. Colour various shades of yellow. Surface
smooth, microscopically hispid. Skeleton (Fig. 3C, D) divided in
Suberitidae with ramose habit and axial choanosomal skele- an axial and extra-axial region. The extra-axial region is rather soft
ton; extra-axial skeleton of brushes of spicules and spicules in con- and consists of somewhat loose brushes of styles (Fig. 3C) with
fusion. Megascleres exclusively styles in a large size variation; no points directed outwards, but many are strewn in confusion. Axial
microscleres. skeleton consisting of a dozen or more unidirectionally or spirally
aligned bundles of 80-120 jJ.m diameter separated by less densely
Diagnosis spiculated areas. The outer tracts of the central core tend to give off
thinner bundles (Fig. 3D) protruding through the surface mem-
Persistently thinly ramose sponges with pointed branches and a brane beyond the extra-axial brushes. At the apices of the branches
system of roots, stolons or branches creeping over the substrate. this structure blurs into a predominantly confused arrangement of
Surface optically smooth although may be somewhat furrowed or spicules. Megascleres (Fig. 3B) exclusively thin styles, mostly
annular, with small oscules on the branches. Consistency wiry, due to faintly subtylostylote (i.e., with an almost imperceptible constric-
axial skeleton, outer parts soft. Skeleton of the axis a mass of aligned tion at some distance from the blunt ending), in a large size range
tracts or more commonly confusedly arranged larger megascleres, with perhaps a distinguishable smaller and larger category,
extra-axial skeleton consisting of brushes of the smaller spicules 100-350 X 1-8jJ.m (Alvarez, 1998). Distribution and ecology.
'echinating' the axial skeleton. Styles are the only spicules. About Subantarctic and Antarctic waters, 35-200 m.
20 described species, recorded from all parts of the three oceans. Remarks. The genus was previously assigned to Axinellidae,
but the structure of its skeleton, lacking a reticulation and possessing
Description of type species brushes of megascleres at the surface, fits much better in Suberitidae.
The structure is in fact quite similar to that of Rhizaxinella
Homaxinella balfourensis (Ridley & Dendy, 1887) (Fig. 3A-D). (cf. below), but that genus has club-shaped habit and tylostyles.
Synonymy. Axinella balfourensis Ridley & Dendy, 1886: The genus Pachaxinella Burton, 1930b: 503 was erected for
480; Ridley & Dendy, 1887: 179, pI. XXIII fig. 1. Axinella supra- type species (by original designation) Halichondria subdola
tumescens Topsent, 1907: 74; Topsent, 1908: 32, pI. II fig. 4, pI. III Bowerbank, 1866: 247. This is a ramose species with axial and
fig. 6, pI. V fig. 2; Homaxinella supratumescens; Topsent, 1916: 4. extra-axial skeletons and exclusively styles, 220-510 X 3-6jJ.m.
Materml examined. Holotype of Axinelia balfourensis: The shape of the sponge (Fig. 4A, B, D) with pointed branches and
BMNH 1887.5.2.18 - Balfour Bay, Kerguelen, 'Challenger' optically smooth surface is similar to Homaxinella balfourensis.
232 Porifera· Demospongiae • Hadromerida • Suberitidae

Fig. 3. Homaxinella balfourensis (Ridley !& Dendy, 1886). A, drawing of habit of lectotype of Axinella balfourensis, BMNH 1887.5.2.18 (reproduced
from Ridley & Dendy, 1887: pI. XXIII fig. 1) (scale I cm). B, SEM images of apices of a representative spicule of ditto (scale 10 11m). C, SEM image of
ectosomal spicule brushes of ditto (reproduced from Alvarez, 1996) (scale 100 11m). D,SEM image of cross section of periphery of a branch of ditto (from
unpublished Alvarez, 1996) (scale 100 !-Lm).

Burton's reasons for erecting a separate genus are the apparent Type species
inversion of spicule density between the axis (rather loosely
arranged skeleton) and the peripheral regions with dense mass of Plicatellopsis arborescens Burton, 1932b: 332 (by original
spicules in confusion (Fig. 4C). However, these sponges contract designation).
notably in preservation, and through this the brushes of smaller
styles, present in the fresh condition, are distorted. Bowerbank's Definition
holotype, BMNH 1910.1.1.228 (and not BMNH 1929.11.27.1 as
Burton: 1930b: 503 stated, Ms Clare Valentine in litteris), was re- Suberitidae with ramose habit and axially condensed skeleton
examined and in many places the extra-axial skeleton is completely of tylostyles; extra-axial skeleton of bundles of spicules issuing
gone leaving only a macerated axis, showing the vague spicule from the central axis to the surface; surface skeleton a dense
bundles characteristic of Homaxinella. A recently collected speci- palisade of smaller tylostyles.
men of this species, ZMA POR. 14107, from Roscoff, France
(SYMBIOSPONGE 98IFR1SEPIIIMKl21), still shows the extra-
axial brushes. It is proposed here to consider Pachaxinella a junior Diagnosis
synonym of Homaxinella.
Thickly branched ramose or irregularly flabellate sponges.
Branches with rounded tips, surface microhispid. Surface skeleton
PUCATELLOPSIS BURTON, 1932 consists of the usual brushes of smaller megasc\eres forming a pal-
isade together with the endings of megasc\ere bundles of the extra-
Synonymy axial skeleton. The latter originate from the axial skeleton and
traverse the extra-axial region towards the surface. Choanosomal
Plicatellopsis Burton, 1932b: 332. skeleton a concentration of aligned or irregularly reticulated
Porifera· Demospongiae • Hadromerida • Suberitidae 233

Fig.4. Homaxinella subdola (Bowerbank, 1866), type of Pachaxinella (Burton, 1930b), considered jun. syn. of Homaxinella . A, habit and spicules (repro-
duced from Bowerbank, 1874: pI. XLIII fig. 14) (scale 1 em). B, photo ofBMNH holotype (courtesy Ms C. Valentine) (scale 1 em). C, in situ photo of liv-
ing specimen, St. Mary's, Scilly Isles (courtesy Dr B. Picton). D, drawing of cross section of branch to show axial differentiation of the skeleton
(reproduced from Burton, 1930b: fig. 8) (scale 1 mm).

spicule bundles. Megascleres tylostyles (Burton, 1932b: 333 states strong bundles. Rhizaxinella australis Hentschel, 1909 may be an
that the type species has styles - and also figures them as such in Australian representative of the genus, which is otherwise known
text-fig. 37b, cf. fig. 5C - but subsequent examination of the type from colder southern ocean waters.
material revealed only tylostyles, be it with small tyles). The genus
is so far only recorded from southern oceans: Falkland Islands,
Patagonia, Antarctica, possibly South Australia, approximately 6-7
described species. PROSUBERITES TOPSENT, 1893

Description of type species Synonymy

Plicatellopsis arborescens Burton, 1932b (Fig. 5A-E). Prosuberites Topsent, 1893c: xlii.
Synonymy. Plicatellopsis arborescens Burton, 1932b: 332,
pI. LVI fig. 5, text-figs 36b, 37b. Type species
Material examined. Holotype: BMNH 1928.2.15.388 -
Falkland Islands. Prosuberites iongispina Topsent, 1893c: xlii (by monotypy).
Description. Stalked-arborescent (Fig. 5A), more or less
dichotomously divided into a three-dimensional bush, 9 cm high, with Definition
thick branches of 0.6-1 cm diameter, rounded at the ends. Surface
velvety, microhispid, slightly irregular in places, with small rounded Suberitidae with a basal plate of spongin from which rise sin-
depressions on the branches which may represent contracted oscules. gle tylostyles. No ectosomal specialization, no localized distinct
Consistency firm, compressible. Colour brown (in alcohol). Ectosomal spicule size categories.
skeleton (Fig. 5B) a dense palisade of tylostyles which is more or less
continuous in preserved specimens. Extra-axial skeleton (Fig. 5B, D) Diagnosis
formed by strong, diverging bundles of up to 2.5 mm length, and 200
(at their origin near the axis) to 400 J.1m wide (at the surface). Axial Thinly encrusting sponges with hispid surface. Skeleton con-
skeleton a dense mass of aligned spiCUle bundles, intertwined and sisting of single, long, tylostyles oriented perpendicularly to the
anastomosed, 1-2 mm in diameter. Spicules tylostyles (Fig. 5E) with substrate, heads embedded in a basal plate of spongin. Spicules
tyle only slightly swollen, in a large size variation, probably divisible tylostyles of variable size, but no distinctly localized size cate-
in two categories, smaller tylostyles150-320 X 1-8 J.1m concentrated gories. A dozen or more species, distributed over most ocean
at the surface, and larger tylostyles, 850-1200 X 10-15 J.1m (meas- regions.
urements from Alvarez, 1998). Distribution and ecology. Falkland
Islands, 130-140m, on sandy shelly bottom. Previous reviews
Remarks. The genus is very closely related to Homaxinella
but is distinguished by its possession of tylostyles rather than Topsent, 1900: 174; Riltzler, 1965: 26; Riltzler & Smith,
styles, and also by the more elaborate extra-axial skeleton with its 1993: 383.
234 Porifera· Demospongiae • Hadromerida • Suberitidae

c
E

I I
Fig. 5. Plicatellopsis arborescens (Burton, 1932b). A, photo of habit ofholotype (reproduced from Burton, 1932b: pI. LVI fig. 5) (scale 1 cm). B, drawings
of cross sections of various branches of Plicatellopsis (reproduced from Burton's text·fig. 36b) (scale see text). C, drawing of spicules of P. arborescens
reproduced from Burton's 1932b: fig. 37b (subsequent examination of type specimen demonstrated that the styles are in reality all tylostyles) (scale see
text). D, SEM image of cross section of a branch of the holotype to show strong axial and extra-axial specialization (reproduced from Alvarez, 1996) (scale
1000 j..Lm). E, drawing of spicule of holotype specialization (reproduced from Alvarez, 1996) (scales: left, 50 j..Lm; right, 100 j..Lm).

Description of type species juvenile growth stages of more elaborate adult specimens may be
confused with specimens which do not become more elaborate later
Prosuberites longispina Topsent, 1893c (Fig. 6A, B). in their life. Prosuberites is here limited to specimens which: (1) have
Synonymy. Prosuberites longispina Topsent, 1893c: xlii; their tylostyles singly embedded with their heads in a basal spongin
Topsent, 1900: 174, pI. VI fig. 14. plate - a feature which is otherwise shared with poecilosclerid genera
Material examined. Holotype (slide): MNHN DT. 2467 - like Hymedesmia and Clathria (Microciona); (2) are strongly hispid
Banyuls, Western Mediterranean. due to projection of long tylostyles through the surface membrane;
Description (based on Topsent, 1900). Thinly encrusting, and (3) do not have a surface specialization of brushes of tylostyles or
with strongly hispid surface due to projecting spicules. Lateral size styles. This definition excludes several well-established Prosuberites
several cm2 . Punctiform oscules. Consistency fleshy, rather soft, species from the genus, the most prominent being P. epiphytum,
rather low spicular density. The skeleton (Fig. 6B) consists of sin- which is here transferred to Protosuberites (cf. below).
gle long spicules standing erect on the substrate with heads
attached to it by a spongin layer. Spicules (Fig. 6A). smooth, long
tylostyles, all sizes, from 170 X 4 f.1m up to 2000 X 17 f.1m, with PROTOSUBER1TES SWARTSCHEWSKY, 1905
tyle 25 f.1m in diameter. Occasionally, the spicule is polytylote.
Smaller tylostyles may occur in groups at the substratum. Synonymy
Distribution and ecology. Western Mediterranean and Adriatic,
Canary Islands, Azores. Common in the Banyuls area (Boury- Protosuberites Swartschewsky, 1905: 36. Laxosuberites sensu
Esnault, 1971a). Topsent, 1938a: 20 (not Topsent, 1896a: 126). ? [Gilchristia]
Remarks. The distinction of the encrusting suberitid genera Burton, 1928a: 129 (preocc.) (not Jordan, 1907). TerpioseUa
remains a difficult problem on account of the possibility that Burton, 1930d: 675. Keratylum de Laubenfels, 1936a: 125.
Porifera' Demospongiae • Hadromerida • Suberitidae 235

--=====-- -
Fig. 6. A, B, Prosuberites longispina Topsent, 1893c. A, drawing of spicules (reproduced from Topsent, 1900: pI. VI fig. 14) (scale 10 /-Lm). B, photo of
cross section of skeleton (reproduced from Riitzler, 1965: fig. 11) (scale 100/-Lm). C, Protosuberites prototipus Swartschewsky, 1905, drawings of cross
section of skeleton and tylostyles (reproduced from Swartschewsky, 1905: pI. VI fig. 5) (scale 100/-Lm).

Type species of spicules arranged in all directions but probably originally

consisting of loosely arranged brushes. Choanosomal skeleton
Protosuberites prototipus Swartschewsky, 1905: 36 (by (Fig. 6C) consists of single tylostyles arranged in arranged, heads
monotypy). on the substrate. Spicules (Fig. 6C) tylostyles in a large size range,
162-417 X 5-7 fJ-m. Smaller tylostyles are drawn in the surface
Definition brushes and strewn in the interior among the larger erect tylostyles.
Distribution and ecology. Black Sea (Liaspi), 60m.
Suberitidae with encrusting habit and a skeleton differentiated Remarks This genus name is here revived to take the place of
in a choanosomal arrangement of single tylostyles or bundles of Laxosuberites sensu Topsent, 1938a: 20. That genus name can no
tylostyles, and ectosomal brushes of tylostyles. longer be employed because Laxosuberites Topsent, 1896a: 126 is a
junior synonym of Hymeniacidon. Its type species (by original desig-
Diagnosis nation) Suberites rugosus Schmidt, 1868, was found to be a specimen
of Hymeniacidon by Topsent (1938a: 20), and his subsequent
Thinly to more massively encrusting sponges with a velvety attempts to convey the properties of Laxosuberites to the Banyu1s
or microhispid surface. Oscular veins often prominently visible. specimen described as Laxosuberites rugosus by Topsent, 1900
The surface skeleton consists of brushes of tylostyles, which are (cf. Fig. 7A, B) are inadmissible because a genus cannot have two
often somewhat smaller than those of the choanosome. Choanosomal different type species. Several species traditionally assigned to
skeleton consists variably of single spicules erect on the substrate Prosuberites, e.g., P. epiphytum (Lamarck, 1814), show skeletal
or bundles running from the substrate to the surface, usually paral- structures intermediate between Prosuberites longispina (Le.,
lel to each other, without any form of anastomosing. The genus in single spicules erect on the substrate) and 'Laxosuberites' spp.
its present concept includes dozens of species from all parts of the (bundles of aligned spicules), even by Topsent's own admission
world oceans. (see Topsent, 1900: 183). Swartschewsky's Protosuberites prototipus
is likewise somewhat intermediate between Prosuberites and
Description of type species 'Laxosuberites' by combining single spicules erect on the substrate
with ectosomal specialization of brushes of spicules. Protosuberites
Protosuberites prototipus Swartschewsky, 1905 (Fig. 6C). is here extended to include thinly encrusting sponges with choanoso-
Synonymy. Protosuberites prototipus Swartschewsky, mal bundles of spicules, as an acknowledgement of the occurrence in
1905: 36, pI. VI fig. 5 (in Russian, with a German summary of the a single species of specimens with single spicules erect on the sub-
species description on p. 52). strate and specimens with bundles of spicules. In Swartschewsky's
Material examined. None. The whereabouts of the type own words 'The genus described here has properties of the above
material is not recently confirmed but the material was deposited in described genera' - he refers to Topsent's genera Suberites,
the collections of the Academy of St Petersburg and the Zoological Prosuberites, Laxosuberites and Pseudosuberites - 'and this gives me
Museum of the University of Kharkov. reason to consider it a genus older than the others, and even older per-
Description (from Swartschewsky, 1905). Very thin encrus- haps than their ancestors. Hence I propose the name Protosuberites'
tation of a rust-brown color. Ectosomal skeleton (Fig. 6C) a mass (translation courtesy ofMs T. Das, University of Amsterdam).
236 Porifera • Demospongiae • Hadromerida • Suberitidae

-
Fig. 7. Protosuberites spp. A, B, Protosuberites rugosus (Topsent, 1938a). A, habit of Banyuls specimen (scale I cm). B, spicules from ditto (scale
100 !-Lm) (both reproduced from Topsent, 1900: pl. V). C-F, Protosuberites epiphytum (Lamarck, 1814). C, drawing of habit of OphlitaspongiaJucoides
Bowerbank, 1876b (jun. syn. of P. epiphytum) (scale 5 cm). D, drawing of skeleton of ditto (scale 500 !-Lm). E, spicules of ditto (scale 100 !-Lm) (all repro-
duced from Bowerbank, 1876b: pl. LXXX). G, SEM image of spicule and detail made from the holotype of Alcyonium epiphytum (scale: left, 10 !-Lm; right,
I!-Lm). F, habit of Lamarck's type specimen (reproduced from Topsent, 1933: pl. III fig . 2) (scale I cm).

The genus [Gilchristia] Burton, 1928a: 129 was erected for fusiform styles, but this is interpreted as a protection against the
type species (by original designation) Bubaris reptans Kirkpatrick host sponge, a common enough phenomenon in epizoic sponges.
(1903b: 248, pI. V fig. 14, pI. VI fig. 15a-c). The name is preoccu- The genus Terpiosella Burton, 1930d: 675 was erected for
pied by Gilchristia Jordan, 1907 (Pisces, Proc. U.S . Nat. Mus. 32: type species (by original designation) Ophlitaspongia fucoides
236). Burton (l928a: 129) gave no definition other than stating that Bowerbank (1876b: 771, pI. LXXX) from Shark Bay, Western
Kirkpatrick's species is not a Bubaris and that the new genus was Australia. This is a hispid, encrusting sponge on an alga (Fig. 7C-E),
incertae sedis in Suberitidae. The description of Kirkpatrick is with spicules erect on the substrate and some aligning the algal thal-
peculiar: it concerns an epizoic sponge on Halichondria pachas- lus strands (Fig. 7D); in places the skeleton is confused. Spicules
trelloides, apparently forming a meandering crust, leaving free (Fig. 7E) are tylostyles of 100-400/-Lm in length. Burton (1930d)
polygonal 'islands' of surface of Halichondria. The crust had a emphasized the symbiosis with the alga as a generic character, and
skeleton of erect bundles of large fusiform styles (which are here also assigned Terpios symbiotica Hentschel, 1909 to Terpiosella. The
interpreted as tylostyles), up to 530 X 15 /-Lm in size, and ectosomal epiphytic habit is not considered a generic feature, as many unrelated
brushes of small tylostyles, size given as 16 X 6/-Lm, which cannot sponges exhibit such behaviour. The structure of O. fucoides con-
be other than a misprint, assumed to be 160 X 6 /-Lm. The properties forms to the definition of Protosuberites and thus Terpiosella is a
of this sponge, if interpreted correctly, conform to Protosuberites. junior synonym of this genus. Topsent (1933: 35) in his redescription
Kirkpatrick (1903b) also described a basal tangential layer of the of what he called Prosuberites epiphytum (Lamarck, 1814 as
Porifera· Demospongiae • Hadromerida • Suberitidae 237

Alcyonium, type specimen MNHN DT3421) pointed out the similar- Material examined. Holotype: USNM 32441 - North
ities with this species, including the extensive overgrowing of algae Carolina, 32 m. Paratype: ZMA POR. 5762 - Margarita Island,
(cf. Fig. 7F), and suggested they were synonyms. Prosuberites epi- Venezuela, 56 m.
phytum is considered a cosmopolitan species with characteristic Description. 'Mobile' sponge living as a rule as a massive
spicules (Fig. 7G); it is reported especially from Europe (e.g., encrustation on empty gastropod shell of the genera Cantharus or
Mediterranean: Topsent, 1900: 178; Netherlands: Van Soest, 1977a: Epitonium, inhabited by a pagurid hermit crab. Shape (Fig. 8A)
266), but there are strong indications that the type originated from irregularly spherical to ovoid, with lobose outgrowths in larger
Australia (collected by Peron and Lesueur). The con specificity of specimens, typically 2-5 cm. Usually a single oscule with raised
Ophlitaspongia fucoides and Alcyonium epiphytum is here accepted rims, about 3 mm in diameter. Surface smooth, velvety.
and thus its ultimate designation is Protosuberites epiphytum. Consistency firm. Colour greyish green, tan or orange, sometimes
The genus Keratylum de Laubenfels, 1936a: 125 was erected mottled. Ectosomal skeleton (Fig. 8B, C) arranged as dense erect
for type species (by original designation) Suberites capillitium tufts. Below this there is an area with many subdermal spaces
Topsent (1892a: 130, pI. II fig. 10, pI. IX figs 19-10). The species is (Fig. 8C) and generally less dense skeleton. Further into the
an encrustation on brachiopod shells collected at greater depths in choanosome, the skeleton consists of irregular tracts of spicules
the North Atlantic; its skeleton consists of bundles oflarge tylostyles, making a confused reticulation with many loose spicules. At the
up to 1000 X 27 f.1m, ending in brushes of small tylostyles as small as contact with the gastropod shell, the skeleton is a dense confused
250 X 7 f.1m. These characters conform to Protosuberites in the pres- layer of spicules, and in many specimen this region also contains
ent concept and thus Keratylum is a junior synonym. round flattened spongin encased 'gemmules' of 150-500 f.1m in
diameter. Spicules (Fig. 8D) exclusively smooth sharp pointed
oxeas, occasionally centrotylote and bent, occasionally stylote or
PSEUDOSPONGOSORITES MCCORMACK & strongylote, probably divisible in two overlapping size categories,
KELLY, 2002 176-295X2-12f.1m and 125-185 X 2-10f.1m. Distribution and
ecology. Common in the Gulf of Mexico and the Atlantic coasts of
Synonymy Florida and North and South Carolina; also recorded from
Venezuela, shoreline down to 60 m.
Pseudospongosorites: McCormack & Kelly, 2002: 1019. Remarks. Some discussion about the phylogenetic affinities
has been going on in the recent literature. Although the similarities in
Type species habit and ecology with Suberites domuncula are compelling, the
exclusive occurrence of oxeas as spicules in this species was consid-
Spongosorites suberitoides Diaz et al., 1993: 299 (by original ered prohibitive for inclusion in Suberitidae until recently. Initially, this
designation). species was assigned to Wilson's (1902) Petrosia halichondrioides by
Wells et al., 1960; however, Wilson's species was discovered to be a
Dermition Plakortis and unrelated to the hermit crab sponge. Diaz et al. (1993)
noted the similarity with Suberites domuncula but considered it to be a
Suberitidae with exclusively small oxeas as spicules, arranged convergent association and assigned the species to the halichondrid
in brushes at the surface and confusedly in the interior. genus Spongosorites. Sandford & Kelly-Borges (1997) described the
association in more detail, expressed their dissatisfaction with the
Diagnosis assignment to Spongosorites, but felt there was not enough evidence
to approach the species to Suberitidae. Subsequently, McCormack &
Sponge massive, irregular in outline, initially encrusting Kelly (2002) produced molecular evidence for close relationship of
gastropod shells and developing a symbiosis with hermit crabs. Suberites and S. suberitoides, and proceded to assign it to a new
Surface smooth, velvety. Skeleton of the surface the usual brushes genus in the family Suberitidae. This is here accepted.
of megascleres. Choanosomal skeleton predominantly confused
with a vague reticulation of spicule tracts. Spicules exclusively
small oxeas, in two overlapping size categories. The genus is mono- PSEUDOSUBERITES TOPSENT, 1896
typical, recorded from shallow water in the Central West Atlantic.
Synonymy

Previous reviews Pseudosuberites Topsent, 1896a: 127.

Diaz et al. (1993); Sandford & Kelly-Borges (1997). Type species

Description of type species Hymeniacidon hyalina Ridley & Dendy, 1887: 168 (by subse-
quent designation; Burton, 1930d: 674).
Pseudospongosorites suberitoides (Diaz et al., 1993)
(Fig. 8A-D). Definition
Synonymy. Xestospongia halichondrioides sensu Wells et al.,
1960: 210 (not Petrosia halichondrioides Wilson, 1902). Suberitidae with tangential ectosomal skeleton of tylostyles
Spongosorites suberitoides Diaz et al., 1993: 299, figs 28, 34; over a confused choanosomal skeleton of tylostyle bundles and
Sandford & Kelly-Borges, 1997: 317, figs 1-5; Pseudospon- individual spicules. Tylostyles variable in length but no localized
gosorites suberitoides; McCormack & Kelly, 2002: 1019. size category.
238 Porifera· Demospongiae • Hadromerida • Suberitidae

Fig. 8. Pseudospongosorites suberitoides (Dfaz et ai., 1993). A, habit of paratype ZMA 5762 (scale I cm). B, drawing of skeleton and spicules (repro-
duced from Dfaz et aI., 1993: fig. 34) (scales: 200 fJ-m, 100 fJ-m). C, photo of peripheral skeleton (reproduced from Sandford & Kelly-Borges, 1997: fig. 3a)
(scale 500 fJ-m). D, photo of spicules (reproduced from Sandford & Kelly-Borges, 1997: fig. 2d) (scale 100 fJ-m).

Diagnosis specimen: ZMA POR. 14929 - Chile, Isla San Ambrosio, 26°20'S
80 0'W, coll. SEPBOP Exped. stat. 17-676B, 160-170m, 12 July
0

Massively or thinly encrusting sponges, structured like 1966 (donated by USNM-SOSC). European specimen: ZMA POR.
Suberites, but with a smooth surface due to a tangential ectosomal 5013 - Norway, Bergen area, Vaagegrund, 100m, dredged, colI.
skeleton. Consistency fleshy, due to an irregular arrangement of R.W.M. Van Soest, c.s. 19 July 1982.
spicule tracts. Spicules tylostyles in a large size variation but with- Description. The type consists of a few shapeless fragments
out distinct size categories. Distribution cosmopolitan, with about (Fig. 9B), encrusted by foreign organisms. The Chilean SEPBOP
20 species described, some of which are reported to have unusually specimens (Fig. 9D) thought to be this species are whitish lobate
disjunct occurrence. fragments with indeed some encrusting hydroids. The species is
'hyaline' in appearance due to large subdermal cavities roofed over
Previous reviews by the surface skeleton (Fig. 9C). The type is rather soft and spongy,
but the recent specimens, both the Chilean and the putative repre-
Topsent, 1900: 164; Burton, 1930d: 674; Riitzler & Smith, sentatives from Norway, are rather firm. Oscules were stated to be
1993: 383. large but a question mark was added; in the recent material no
oscules are apparent. The ectosomal skeleton is unusual for a suber-
Description of type species itid in containing a distinct tangential arrangement of spicules car-
ried by thick bouquets of subdermal spicules spaced rather far apart
Pseudosuberites hyalina (Ridley & Dendy, 1887) and leaving large subdermal cavities in between (Fig. 9C). The
(Fig. 9A-D). choanosomal skeleton is largely confused, with few spicule bun-
Synonymy (restricted). Hymeniacidon? hyalina Ridley & dles. Tylostyles (Fig. 9A) robust, with prominent globular heads
Dendy, 1887: 168, pI. XLV fig. 6; Pseudosuberites hyalina; ('pin-shaped'), fusiform; those at the surface and in the subdermal
Topsent, 1896a: 127; Topsent, 1900: 164; Burton, 1930d: 674. bouquets often smaller, but probably not clearly divisible in two size
Material examined. Holotype: BMNH 1887.5.21.41 - as categories: 220-1200 X 2-25 fLm. Distribution and ecology. SE
'Hymeniacidon (7) hyalina' (wet), off SW coast of Patagonia, Pacific, and recorded disjunctly from the North Atlantic, in deeper
'Challenger' colI. (not examined). Other material. Topotypical water. Many other records, including tropical localities, are suspect.
Porifera· Demospongiae • Hadromerida • Suberitidae 239

Fig.9. Pseudosuberites hyalina (Ridley & Dendy, 1886). A,drawing of spicules (reproduced from Ridley & Dendy, 1887: pI. XLV fig. 6) (scale 100 !Lm) .
B, habit of type material BMNH 1887.5.21.41 (photo courtesy Ms C. Valentine) (scale see text) . C,cross section of skeleton (reproduced from Kirkpatrick,
1908c: pI. XXVI fig . 7) (scale 500 !Lm). D, photo of topotypical fragments , ZMA 14929 (scale 1 em).

Remarks. The original habitat of this species is not recorded, been right in his assumption that the two are synonyms then
other than that it came from off SW Patagonia. The type material is Suberanthus would have had priority, since the publication date is in
meagre and the authors even state that they were at loss about its fact January 9, 1896. Fortunately, Suberanthus is a synonym of
affinity and had doubts describing it as a new species. It is surprising, Suberites (cf. below).
that it subsequently was recorded from all over the world. The iden-
tity of those records remains in most cases dubious, because of the
paucity of data available on the original specimen. Pseudosuberites RHIZAXINELLA KELLER, 1880
hyalina was recorded from such extremely different habitats as
deep water (500-600m) in the Mediterranean (Topsent, 19OO: 171), Synonymy
tropical mudflats in the Suez area (Row, 1911), and the freezing
cold water at McMurdo Sound and other Antarctic locations at Rhizaxinella Keller, 1880a: 272.
depths of 300m or more (Kirkpatrick, 1908c: 21 ; Burton, 1929a:
445), as well as from the Arctic Ocean between Bear Island and Type species
Norway (Topsent, 1913a: 26). It is obvious that not all these records
concern the same species. From Topsent's (l896a, 1900) treatment Rhizaxinella clavigera Keller, 1880a: 272 (by monotypy)
of Pseudosuberites, it is clear that he intended the well-established (=junior synonym of Alcyonium pyrifera delle Chiaje, 1828: lSI).
species P. sulphurea (Bowerbank, 1866: 208, as Hymeniacidon) as
the type of the genus. Unfortunately, he mentioned Hymeniacidon ?
hyalina Ridley & Dendy (1887) as a second species of Definition
Pseudosuberites, and omitted to state expressly that P. sulphurea was
the type. Equally unfortunately, Burton (1930d) chose hyalina as the Suberitidae with spherical or ovoid body carried on a long
'genotype', thus making the status and affinities of Pseudosuberites thin stalk. Skeleton radially arranged with ectosomal spicule
less clear than it would have been with P. sulphurea as the type, brushes at the surface. Spicules tylostyles, and if present rhaphides
because apart from all the dubious records from various localities arranged in trichodragmas .
around the world there have been no further published records from
the Chilean Patagonian coast. The nearest subsequent record is from Diagnosis
the Falkland Islands (Burton, 1932b), but no further descriptive data
were provided. P. sulphurea is a yellow-orange encrusting species Pedunculate sponges, with spherical, ovoid or cylindrical body
fairly common along the coasts of Europe and well-established by carried on a simple or ramified stalk, normally attached to the sub-
type and additional material. Burton (1930d: 674) mentioned strate by a root system; usually an apical oscule. Body with a more-
Suberanthus Lendenfeld as a junior synonym of Pseudosuberites, or-less radial skeleton and with brushes of spiCUles at the surface,
giving Lendenfeld's publication as 1897 (1897a). If he would have skeleton of the stalk a tightly packed mass of aligned megascleres,
240 Porifera· Demospongiae • Hadromerida • Suberitidae

I
I
C

Fig. 10. Rhizaxinella pyrifera (Delle Chiaje, 1822). A, drawing of habit (scale 1 em). B, cross section of main body (scale 1 mm). C, tylostyles (scale
100 J.1m) (all reproduced from Vosmaer, 1935: pis 38-40).

bound by spongin. Megascleres are tylostyles of different sizes witb Remarks. This is a clearly demarcated genus. It shows some
raphides in trichodragmas. About 20 species have been described superficial resemblance to Stylocordyla (family Stylocordylidae)
from all three ocean basins, usually from deeper or colder waters. but differs in having only tylostyles as megascleres.

Previous reviews
SUBERITES NARDO, 1833
Keller (1880a), Vosmaer (1887), Topsent (1900), Vosmaer
(1935). Restricted synonymy

Description of the species Suberites Nardo, 1833: 523. Ficulina Gray, 1867a: 523.
Suberanthus Lendenfeld, 1896: 144. Suberella Thiele, 1905: 416.
Rhizaxinella pyrifera (delle Chiaje, 1828) (Fig. lOA-C). [Suberella] Burton, 1929a: 446. Carnleia Burton, 193Oc: 519;
Synonymy. Alcyonium pyriferum delle Chiaje, 1828: 151; Laxosuberella Burton, 1930d: 675.
Rhizaxinella pyrifera; Vosmaer, 1887: 331; Topsent, 1900: 243.
Rhizaxinella clavigera Keller, 1880a: 272, pI. xm figs 1-3. Type species
Material examined. Holotype: Delle Chiaje's material is
not located. Type material of Keller may be in 2MB (not seen). Alcyonium domuncula Olivi, 1792: 241 (by original
Other material. Aegean Sea specimens reported in Voultsiadou- designation).
Koukoura & Van Soest, 1993.
Description. Stalked sponges with pyriform or cylindrical Definition
main body (Fig. lOA), lengtb of total sponge 3-23 cm, main body
0.5-19 cm. Stalk usually gradually broadening into tbe main body. Suberitidae with ectosomal skeleton consisting of bouquets of
Stalk fixed in or to substratum by a fine-threaded root system. smaller tylostyles carried by subradiate bundles of larger tylo-
Color whitish or greyish. Surface velvety. Usually a single apical styles; choanosomal skeleton confused or alveolar arrangement of
oscule. Skeleton (Fig. lOB) axially condensed, in the stalk the sur- larger tylostyles. If present, microscleres are spined centrotylote
face brushes are immediately perpendicular on the aligned and microstrongyles.
spongin-enforced axial spicules, but in the main body the axis
diverges into thinner secondary axes and finally end at the surface Diagnosis
in the brushes. In between numerous trichodragmas are found.
On tbe whole tbe spicules at the periphery are smaller tban tbose of Massive, compact, usually witb velvety smooth surface, caused
tbe axial parts, but no sharp distinction can be made. Spicules by dense ectosomal arrangement tylostyles oriented perpendicularly
(Fig. lOC) tylostyles, 300-2500 X 3-20 fLm, possibly divisible in to tbe sponge surface, points outward; peripheral choanosomal
smaller tylostyles 300-1200 fLm at the surface, and larger, often skeleton consists of closely packed strands of tylostyles distinctly
flexuous tylostyles 800-2500 fLm of the axial bundles. Raphides, larger tban ectosomal ones, with interior skeleton of densely
usually in trichodragmas, 80-120 fLm. Distribution and ecology. packed tylostyles in confusion. Centrotylote, minutely spined micro-
Mediterranean, deep water, 70-600 m. strongyles may be present in a few species and if so are concentrated
Porifera· Demospongiae • Hadromerida • Suberitidae 241

Fig. 11. Suberites domuncula (Olivi, 1792). A, habit of neotype ZMA 2237 (scale I cm). B, SEM image of cross section of peripheral skeleton (scale
100 /-Lm). C, SEM image of cross section of choanosome (scale 100 /-Lm). D, SEM image of spicules (scale 10/-Lm). E, SEM images of details of tylostyle
heads (scale I /-Lm).

at the surface. Approximately 80 species have been described an oscule 1 cm in diameter. Other specimens may have two oscules,
(obvious synonyms and invalid genus assignments omitted). arranged occasionally also nearer to the hermit crab aperture.
The genus is cosmopolitan in distribution but is most common in Many specimens have a layer of asexual rounded bodies on the sur-
cold temperate waters. face adhering to the mollusc shell or the hermit crab cavity, often
referred to as 'gemmules'. The colour of the neotype is not given,
Previous reviews but most specimens recorded alive are bright orange, some are mot-
tled with brown, red or blueish spots (so called 'marmorated' form).
Lendenfeld (1896), Topsent (1900), Vosmaer (1935), Burton Skeleton (Fig. liB, C) perpendicularly arranged in the peripheral
(1953), Riitzler & Smith (1993). 10 mm, becoming progressively confused towards the interior,
where the spicules are packed around interior holes and canals. The
Description of type species ectosomal spicules are arranged in bouquets of smaller spicules
carried by radiating larger spicules. Spicules tylostyles
Suberites domuncula (Olivi, 1792) (Fig. l1A-E). (Fig. lID, E) with a slightly subterminal (drop-shaped) but mostly
Restricted synonymy. Alcyonium domuncula Olivi, 1792: well-formed tyle, except for rare annular swellings in the neck
241; Suberites domuncula; Nardo, 1833: 523; Topsent, 1900: 225, region. Small surface tylostyles, 100-350 X 4-8 f.Lm, larger
pI. VI figs 1-9; Burton, 1953: 353. choanosomal tylostyles 250--480 X 5-8 f.Lm, the length of small sur-
Material. Type material: considered lost. Neotype (desig- face tylostyles averaging 53% of the length of choanosomal
nated herein): ZMA POR. 2237 - Triest, no further data. Other mate- tylostyles. Distribution and ecology. Originally described from the
rial. Numerous ZMA specimens from the Mediterranean-Atlantic. Adriatic, but subsequently widely recorded from all over the world.
Description. The neotype (Fig. l1A) has an ellipsoid-oval It is likely that this widespread occurrence is an artefact of low
shape, size 7 X 7 X 6 cm. Surface velvety smooth. Consistency amount of discriminatory morphological characters, and in reality
firmly compressible. Two apertures are apparent. One leads to the represents a complex of species living in symbiosis with hermit
cavity in which a hermit crab is lodged. In smaller specimens an crabs. The depth preference of the species is between 10 and 100 m.
empty gastropod shell or its remains are still to be found and then Remarks. Suberites domuncula occurs in a uniform orange
the hermit crab that lives in it; however, larger specimens usually and a marmorated form. The latter is extremely common along the
have no trace of the shell, and the crab lives inside the sponge. On sandy coasts of West Africa (Van Soest, 1993). So far it is unclear
the opposite side of the hermit crab aperture, in the neotype, there is what the cause is of the difference in surface markings; in alcohol
242 Porifera' Demospongiae • Hadromerida • Suberitidae

the distinction between the colour morphs disappears. The species was erected, was Suberanthus flavus (see Lendenfeld, 1896: pI. V
is reliably recorded from the Mediterranean, and the adjacent figs 45, 46, pI. VII fig. 71, pI. XII figs 215, 216, 222, 223, 226),
Atlantic coasts. Records from boreal Atlantic waters may have so this is firmly established as type species. However, Lendenfeld's
been confused with Suberites suberia (Montagu, 1818), which has definition and the synonymy he gave of S. flavus implied that he
a duller colour and possesses spined microstrongyles, but other- intended the genus to be what is now generally considered as the
wise is similar in habit and hermit crab symbiosis. This species genus Pseudosuberites (cf. below). He defined Suberanthus as hav-
may also occur in the Aegean Sea (Voultsiadou-Koukoura & Van ing a reticulation of tangential megascleres at the surface and
Soest, 1993: 179, as Suberites ficus) and in the Black Sea included Hymeniacidon sulphurea Bowerbank, 1866 in the syn-
(Swartchevsky, 1905). Lithumena Nardo, 1833 is associated with onymy of S. flavus. Topsent (1900: 236) rejected this synonymy
Suberites domuncula by Lieberkiihn, 1869: 520, but this was not and considered S. flavus a form of Suberites camosus Johnston
verified. If this is indeed the case, then Lithumena is an objective (1842), thereby implying that Suberanthus is a junior synonym of
synonym of Suberites because both share the same type species. Suberites.
The genus Ficulina Gray, 1867a: 523 was erected for The genus Suberella Thiele, 1905: 416 was erected as a sub-
Suberitidae with spined microstrongyles. Its type species (by genus of Suberites for type species (by original designation)
monotypy) is Halichondriaficus Johnston, 1842. The synonymy of Suberites heros Schmidt, 1870: 46 from the West Indies. The
this genus with Suberites has been the subject of a long debate description of Schmidt is somewhat ambiguous, mentioning both
among taxonomists (see review by Burton, 1953: 353). With the tylostyles and oxeas. A specimen in ZMUC labeled "type,
species Ficulina ficus - as with many other Suberites species - ?Vestindien" is here designated as lectotype (cf. Fig. 12F). It con-
there are taxonomic as well as nomenclatorial problems: the name forms in all respects to Suberites domuncula, including the usual
ficus goes back to Pallas (1766, as Alcyonium), but that is generally hole of a hermit crab and exclusively tylostyles in two size cate-
assumed to have been a compound ascidian. Subsequent uses of the gories. The original locality of this specimen is dubious because so
name ficus by Linnaeus (1767), Esper (1794) and Lamarck (1814), far no recent records of such sponges are reported from the West
all as Alcyonium, were likewise uncertain. Only when British Indian region, but in any case Suberella Thiele is a clear junior
authors (Johnston, 1842; Bowerbank, 1866) used this name in synonym of Suberites.
combination with descriptions of the spicules, the name became The genus [Suberella] Burton, 1929a: 446 was erected for
more profiled. It is proposed here to retain Suberites ficus authored type species (by original designation) Suberites topsenti Burton
by Johnston, 1842: 144, pI. XV figs 4,5 as the proper name for the (1929a: 446, pI. N fig. 5). The holotype is BMNH 1926.10.26.186
type species of Ficulina (as originally stated by Gray, although this from McMurdo Sound, Antarctica. However, belatedly, Burton
may not have been intentional), and BMNH 1847.9.7.51 is here (1930d) discovered that Suberella was preoccupied and erected the
chosen as the lectotype. Suberites ficus sensu Johnston, 1842 genus Laxosuberella Burton (1930d: 675) with the same type
(Fig. 12A-E), is a big massive lobate, occasionally cylindrical, species as a replacement. The genus was intended for sponges
orange sponge with one or more conspicuous, large oscules. It has intermediate between Pseudosuberites and Suberites, showing
a velvety smooth appearance. Growth forms may include speci- a confused choanosome and organised 'cortical layer' of spicules,
mens enveloping gastropod shells and encrusting scallops. Size which are arranged with the apices pointing outwards. The name
variable, 10-40cm in length. Contraction noticeable: when Suberella topsenti was intended to replace Suberites montiniger
removed from the water the sponge contracts to about 3/4 of its size sensu Topsent, 1915b: 39, a globular-massive species with papillae
when undisturbed underwater. Internally the skeleton is confused, at the surface resembling those of Cliona or Polymastia. Spicules
almost halichondroid. Tylostyles (Fig. 12C) divided into two size are tylosty1es in a single but variable size, no tangential or cortical
groups (100-250 j.l-m and 350-500 X 5-10 j.l-m), the larger growing structures are present. (The original Suberites montiniger Carter,
in a wide size range. Microscleres (Fig. 12D) are microrhabds 1880d: 256, described from the Arctic, was considered a different
(microspined microstrongyles, usually centrotylote), often rare and species by Burton (1929a), but later authors (e.g., Koltun, 1964b)
needing careful searching of the ectosome, amongst the tips of the disagreed and reverted to Suberites montiniger again). It is
palisade spicules (Fig. 12B, arrow). Size 15-50 j.l-m. Considered as here acknowledged that Suberites montiniger Carter is probably
cosmopolite this species is very likely a complex of sibling species. different from Suberites montiniger sensu Topsent (1915b),
It has been demonstrated that among Suberites ficus from the Irish but the characters on which to distinguish a separate genus
Sea three genetically isolated species are present (Sole-Cava & Laxosuberella appear to be insufficiently different from Suberites
Thorpe, 1986). De Laubenfels (1936a: 143) considered Choanites species, and accordingly Laxosuberella is considered a junior
Mantell, 1822: 178 a senior synonym of Ficulina, but this has not synonym.
been further verified. Subsequently, the same author (de Laubenfels, The genus Camleia Burton, 1930c: 519 was erected for type
1955b: E108) considered Choanites a genus of Tunicata based on species Biemna rhaphidiophora Brfilndsted, 1923: 136, fig. 17 from
the technicality that the type species was named Alcyonium ficus New Zealand (original designation). The type material (in ZMUC,
sensu Pallas (probably a tunicate) and not sensu Johnston. In view not examined) is described as a series of lumpy, club-shaped
of the fact that Ficulina is here considered a junior synonym of sponges, attached to shells, of which the biggest is 4.5 cm in diam-
Suberites, we propose to follow de Laubenfels' technical solution; eter. Surface smooth. Skeleton of the surface a partly perpendicular
if not, Choanites would have to be suppressed because it threatens arrangement of megascleres, becoming less densely arranged and
the junior name Suberites. more confused towards the interior. Megascleres are two size cate-
The genus Suberanthus Lendenfeld, 1896: 144 was erected gories of tylostyles, 275-460 X 5j.l-m and 143-208 X 5-6j.l-m,
for type species (by monotypy) Halichondria flava Lieberkiihn, apparently not localized in the skeleton. Microscleres reported as
1859: 521, pI. XI fig. 5), now generally considered a junior syn- sigmas, probably in two size categories, 10-65j.l-m, but they are
onym of Suberites massa Nardo, 1847a. Suberanthus is a case of quoted to be infrequent, probably foreign (Bergquist, 1970: 26) and
thorough confusion. The only species described when the genus microxeas 46-52 X 1.5-2.5j.l-m (likewise foreign according to
Porifera· Demospongiae • Hadromerida • Suberitidae 243

Fig. 12. Suberites spp. A-E, Suberites ficus sensu Johnston, 1842. A, habit in situ (photo B. Picton). B, SEM view of surface showing palisade of spicules
and (arrow) microrhabd (scale 100 fLm). C, SEM images of tylostyles (scale 10 jJ.m). D, SEM image of spined microrhabd (scalelO fJ-m). E, photo of lecto-
type of Halichondria ficus Johnston, 1842, BMNH 1847.9.7.51 (scale 1 cm). F, ZMUC lectotype of Suberites heros Schmidt, 1870, type of the genus
Suberella Thiele, 1905, considered jun. syn. of S. domuncula (scale 1 cm).

Bergquist, I.c.). Both Bergquist (1970) and Hooper (1984: 34) Definition
proposed to synonymize Camleia with Suberites, and this is
followed here. Suberitidae with encrusting habit and with a single category
of tylostyles with flattened lobate tyles.

TERPIOS DUCHASSAING & MICHELOTTI, 1864 Diagnosis

Synonymy Thinly encrusting sponges with optically smooth surface, but

microscopically hispid due to projecting spicules. Consistency
Terpios Duchassaing & Michelotti, 1864: 102. gelatinous, soft. No special ectosomal skeleton other than brush-like
endings of the weakly developed choanosomal spicule strands.
Type species Choanosome with relatively low spicular density and many loose
spicules in confusion. Bacterial symbionts of common occurrence
Terpios fugax Duchassaing & Michelotti, 1864: 102 (by sub- and often causing blue-pigmented or other bright coloration.
sequent designation; Topsent, 1900: 192). Spicules thin relatively short tylostyles in a single size category and
244 Porifera· Demospongiae • Hadromerida • Suberitidae

Fig. 13. Terpios fugax Duchassaing & Michelotti. 1864. A, water color image of type (reproduced from Duchassaing & Michelotti, 1864: pI. XXIV
fig. 6) (size see text). B, SEM images of ty10styles (scale 10 fLm).

possessing characteristic flattened-lobate or lumpy, wrinkled tyles. dark green or blue. Surface smooth, consistency soft, easily dam-
Distributed over all three oceans with a predominance in warm or aged. Skeleton feebly developed, with thin strands of tylostyles
warm-temperate waters. About 10--15 species have been described. running from substrate to surface, with single spicules scattered in
between. Spicules (Fig. 13B) tylostyles in a single size category,
Previous review 150-460 X 2.5-5 J.Lm, with peculiarly lobate, semiglobular or flat-
tened tyles. The tyles contain a branched axial canal. Symbiotic
Riitzler & Smith (1993). bacteria blue-pigmented, filamentous (multicellular). Distribution
and ecology. Originally described from the West Indies, in shallow
Description of type species reefs. Subsequently recorded from many different parts of world,
including Europe.
Terpiosjugax Duchassaing & Michelotti, 1864 (Fig. 13A, B). Remarks. The European representatives of T. jugax show
Synonymy. Terpios jugax Duchassaing & Michelotti, 1864: some discrepancies in spicule size (cf. Voultsiadou-Koukoura &
102, pI. XXN figs 6, 7. Van Soest, 1993: 178) and probably belong to a species separate
Material examined. Lectotype (designated by Van Soest from the West Indian population. They may conveniently be called
et al., 1983: 201): BMNH 1928.11.12.11 - spicule slide from Terpios gelatinosa (Bowerbank, 1866: 222, as Hymeniacidon). The
St Thomas, West Indies. Other material. ZMA POR. 5155 - Cura9ao, genus is well-established by the combination of the lobate tylostyle
reef, 15 m, coIl. G. van Moorsei. ZMA POR. 8355 - St Croix, heads (which appear similar to those of Caulospongia), encrusting
Virgin Islands, coIl. B. Gladfelter. habit and low spicular density.
Description (mostly from Riitz),er & Smith, 1993). Thin
crusts (Fig. 13A) on and under dead and overhanging corals, colour
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademicIPlenum Publishers, New York, 2002

Family Tethyidae Gray, 1848

Michele Sara

Department for the Study of the Territory and its Resources Corso Europa 26, 16132 Genoa, Italy (sara@dipteris.unige.it).

Tethyidae Gray (Demospongiae, Hadromerida) includes sponges with stylote megascleres and two types of asters: micrasters and
megasters. The body shape is typically globose but also irregularly massive or encrusting, and in some genera is supported by a stalk. The
surface is verrucose with tubercles. A cortex is frequently more-or-less developed around a softer choanosomal core. The skeleton struc-
ture is usually made by radiate bundles of stylote megascleres (strongyloxeas). The type genus Tethya contains the greatest number of
species but the family contains 14 valid genera and approximately 80 described species distributed worldwide, in tropical, temperate seas
and one species in the arctic. They are mainly found in shallow waters but also occasionally in the deep-sea. Genera are differentiated
essentially on the basis of their skeletal structure, which may detennine changes in the body shape and spicular types.
Keywords: Porifera; Demospongiae; Hadromerida; Tethyidae; Tethya; Anthotethya; Burtonitethya; Columnitis; Halicometes; Laxotethya;
Nucleotethya; Oxytethya; Stellitethya; Tectitethya; Tethyastra; Tethycometes; Tethytimea; Xenospongia.

DEFINITION, DIAGNOSIS, SCOPE Scope

Synonymy Fourteen valid genera.

Tethyadae Gray, 1848, 1867. Donatiadae Gray, 1872. History and biology
Xenospongiidae Carter, 1882. Tethyidae Sollas, 1888. Donatiidae
Baer, 1906. Type genus: Tethya Lamarck, 1814. The family Tethyidae is widely attributed to Gray (1867a), as
Tethyadae, but this name was used twenty years earlier by Gray
Definition (1848: 1) to defme a family that included a genus Tethya, with two
species Tethya cranium and T. aurantium, and also embracing all
Hadromerida usually with stylote megascleres, mainly existing eight genera of 'Silicea'. Although Tethya cranium was
strongyloxeas, generally in radiate tracts and two categories of euas- subsequently assigned to another genus (Craniella), and the other
trose microscleres, micrasters and megasters, sometimes rhabds. genera to other families and orders, and despite that it was also
originally considered the only family of the 'order Silicea', the
Diagnosis name Tethyadae (corrected as Tethyidae) must be considered to
date from 1848 because it includes the type species of the genus
Body shape typically globose (spherical, hemispherical, Tethya Lamarck, T. aurantium. Gray (1867a) provided a diagnosis
ovoid), but also sometimes irregularly massive, lobate or encrust- of the family that is now considered obsolete as it included many
ing. Sometimes with a basal stalk. The surface is verrucose with tetractinellids and hadromerids (25 genera). Tethya aurantium was
tubercles, typically hispid and frequently flattened. Oscules rarely referred to Donatia Nardo 1833, because, erroneously, the name
visible, generally apical, and the pores are in grooves between Tethya was conserved for C. cranium. In a second system Gray
tubercles. A cortex may be well-developed and sharply distinct (1872) provides a narrower concept of the family including only
from the choanosome, but sometimes poorly developed or indis- Donatia (= Tethya), Collingsia, according to Schmidt Stelletta, and
tinct (pseudocortex). The skeleton structure is constructed by sty- Tethyopsis, another Ancorinidae. Vosmaer (1887) assigned to the
lote (or modified stylote) megascleres, usually in radiate tracts (but family, now correctly named Tethyidae, the genera Tethya,
in some genera in parallel tracts running from the substrate to the Tuberella and Aaptos (although Tuberella was later placed into
tubercles). The main bundles may branch and sometimes are loose synonymy with Aaptos).
in confused tracts, or crossed by thinner transverse bundles. Sollas (1888) provided a clearer diagnosis of Tethyidae,
Interstitial megascleres may also be scattered between the tracts. placing in it Tethya, Columnitis and Xenospongia (three genera
Spicules generally include stylote megascleres and two types of included in this present revision), plus Magog which is now recog-
asters: megasters and micrasters. Stylote megascleres are nised as a synonym of Chondrilla. For Topsent (1898a) the family
frequently strongyloxeas (fusiform styles), but may be also included in addition to Tethya the genera Tethyorrhaphis
styles, subtylostyles, tylostyles or anisostrongyles. In one case Lendenfeld, Tuberella Keller (=Aaptos), Trachya Carter (=Aaptos)
(Oxytethya) the megascleres are oxeas. Megasters are spherasters, and Heteroxya Topsent. None of these genera, with the exception
oxyspherasters and oxyasters. Micrasters are euasters of the of Tethya, is presently included in the family. Tethyorrhaphis is a
tylaster, strongylaster and oxyaster type. Exceptionally one cate- synonym of Tethya (Bergquist & Kelly-Borges, 1991; Sara,
gory of aster (megasters or micrasters) may be lacking, a presumed 1994a); Aaptos has been removed from Tethyidae and assigned
secondary loss. Other spicules occasionally found are polyrhabds, to Suberitidae (see Van Soest, this volume); Heteroxya is assigned
microstrongyles, exotyles and spheres. Reproduction may be to Desmoxyidae (see Hooper, this volume). Baer (1906), following
oviparous with a parenchymella larvae, or asexual by external (and Gray (1867a, 1872) and Thiele (1903a), placed Tethya aurantium
sometimes internal) buds. into synonymy with Donatia lyncurium (=aurantium) because, in

245
246 Porifera' Demospongiae • Hadromerida • Tethyidae

his opinion, the name Tethya should be assigned, following the Tectitethya and established the new genus Nucleotethya. More
nomenclature rules to Craniella. The family name became recently, three new genera from Australia (Anthotethya,
Donatiidae and this name was used by all authors for the following Laxotethya, Oxytethya) were described (Sara & Sara, 2002). In this
two decades. In general reviews of the family made by Burton present chapter I recognize the validity of the generic name
(1924) and Wilson (1925) the name Donatiidae was conserved. Tethytimea de Laubenfels, 1936a for Donatia tylota Hentschel,
For Hentschel (1909, 1912) and Dendy (1916c) Donatiidae 1912, and Tethyorrhaphis oxyaster Burton, 1934a as type species
included three genera: Donatia Nardo, Tuberella Keller and of the new genus Tethyastra.
Xenospongia Gray. The large revision by Burton (1924) included The family Tethyidae presently contains fourteen genera. A
four genera in Donatiidae: Donatia Nardo, 1833, Halicometes cladistic analysis of the family (Sara & Burlando, 1994) included
Topsent, 1898a, Tethyorrhaphis Lendenfeld, 1888 andXenospongia eight genera, and showed that Tethyidae are divided into three
Gray, 1858. The genus Halicometes refers to Cometella stellata clades. One clade, the closest to the family root, is represented by
Schmidt, 1870, which Topsent (1898a) had previously assigned Tethya, a cosmopolitan and speciose genus, with species living
to Halicometes but included the taxon in the family Stylocordylidae. mainly in shallow tropical and warm-temperate areas, but with
However, Topsent (1920a), re-describing Cometella stellata as some species recorded from deep waters and cold-temperate areas.
Halicometes stellata, referred Halicometes to the family Tethyidae. One species (T. norvegica) lives also in arctic waters. The other
Significantly, in the revision of Burton, Tuberella (=Aaptos) two clades give rise to two adaptive radiations: (1) stalked forms,
was not included in Donatiidae. By the end of 1920s nearly developed in deep-water, soft bottom habitats (Burtonitethya,
all authors returned to the names Tethya and Tethyidae on the Halicometes, Tethycometes); (2) massive and encrusting forms spe-
basis of the criticisms made by Topsent (1920d, 1928c) and cialized for shallow-water environments (Columnitis, Xenospongia,
Vosmaer (1933). Stellitethya, Tectitethya). A subsequent cladistic analysis including
De Laubenfels (1936a) included Tethyidae in the order three other genera showed that Nucleotethya and Tethytimea were
Epipolasida Sollas instead of in Hadromerina Topsent, defining placed in two successive bifurcations at the basis of the massive-
Epipolasida as having as principal spicules oxeas and occasionally encrusting clade, while Tethyastra, although not stalked, lies at the
styles instead of tylostyles. This assignment was not followed and basis of the stalked clade in a position near Tethya. The suggested
cannot be applied to Tethyidae in which oxeas are normally lacking phylogenetic position of Anthotethya and Laxotethya is near the
and tylostyles can sometimes be present, and Tethyidae is cur- massive shallow water clade, while that of Oxytethya is near Tethya
rently assigned to Hadromerina (emended to Hadromerida). De (Sara & Sara, 2002).
Laubenfels (1936a) included in the family only Tethya, Tethycordyla
(a new genus erected for T. thyris de Laubenfels, 1934 but later Distribution
synonymised with Halicometes (Sara, 1994», and Tethytimea
(a new genus established for Donatia tylota Hentschel, 1912). The family is widely distributed in the tropical and warm-
Aaptos, Tethyorrhaphis, Halicometes and Xenospongia were temperate regions but the genus Tethya has some cold temperate
referred to other families. and one arctic representative. The depth ranges from intertidal
Levi (1973) included in Tethyidae the genera Tethya, Aaptos habitats to 2100m (Halicometes hooperi Levi, 1993).
and Xenospongia, whereas Bergquist (1978) included only
Tethya and Aaptos. Bergquist & Kelly-Borges (1991) placed
Tethyorrhaphis into synonymy with Tethya. More recently, Hooper DIAGNOSTIC FEATURES OF THE FAMILY
& Wiedenmayer (1994) included Tethya and Xenospongia in the
family, with Aaptos referred to Polymastiidae based on evidence Main diagnostic features
from morphology (e.g., Boury-Esnault, 1987) and biochemistry
(Kelly-Borges et al., 1991). Tethyorrhaphis Lendenfeld, Tethycordyla Tethyidae is distinctive for its spicular pattern with stylote
de Laubenfels and Tethytimea de Laubenfels are considered megascleres and two categories of euasters (megasters and micras-
synonyms of Tethya. ters). Tethyidae is placed in the Hadromerida based on the absence
In the first revision of Tethyidae sincer Burton (1924), Sara of triaenes and of any other tetractinal form of spicules, but species
(1994) recognized eight valid genera in the family, based on a show a clear relationship with the Astrophorida based on the
study of types and description of new material: Tethya Lamarck, common occurrence of euasters. On the other hand, the relation-
Burtonitethya Sara, Columnitis Schmidt, Halicometes Topsent, ship between Tethya and other Hadromerids (Suberitidae and
Stellitethya Sara, Tectitethya Sara, Tethycometes Sara and Clionaidae) has been demonstrated by Kelly-Borges et al. (1991)
Xenospongia Gray. Three of the four genera included in the revi- using RNA sequence analysis. Aaptos, previously included in the
sion of Burton (Tethya for Donatia, Halicometes andXenospongia) Tethyidae, is suggested, on the basis of a cladistic analysis of the
were maintained, while Tethyorrhaphis was considered as a syn- family (Sara & Burlando, 1994) to represent the sister group of
onym of Tethya. Burtonitethya and Tethycometes were erected for the Tethyidae and Aaptos is now arranged among the Suberitidae
new material discovered in the BMNH. Columnitis Schmidt was (Van Soest, this volume). The generic classification of the
kept as valid, while Stellitethya and Tectitethya referred to species Tethyidae is chiefly centered on some general features of the skele-
previously included in Tethya. However, T. crypta was originally ton structure. Among these are the radiate, parallel or branched
described by de Laubenfels (1949b) as belonging to Cryptotethya megasclere tracts, the megasc1ere shape, the megaster shape and
Dendy (whose type species is C. agglutinans, a non-Tethyidae distribution, the occurrence of a central spicular nucleus and of
species). Tethycordyla de Laubenfels was placed in synonymy with transversal bundles of secondary megascleres, and the inclusion of
Halicometes, and Aaptos was not included in the family Tethyidae sediment grains. However, genera are usually recognizable also by
at all. Successively Sara & Bavestrello (1996), on the basis of their body shape that may be globular, ovoid and stalked, massive,
new material from the Caribbean, redescribed Columnitis and or encrusting. In globose and ovoid stalked genera the megasc1eres
Porifera • Demospongiae • Hadromerida • Tetbyidae 247

fonn radiate bundles. Radiate bundles occur also in some massive Australia and New Zealand. (Sara, 1998). The number of Indo-
genera as Tectitethya, Nucleotethya and Columnitis. In Tectitethya, Pacific species is significantly greater than Atlantic species. One
Anthotethya, Oxytethya, and especially Laxotethya, the radiate species of Tethya occurs in arctic seas and scattered spicules have
bundles show branching and anastomosing tracts, and in been recorded on the shores of Antarctica. Of the 14 genera, two
Columnitis and Nucleotethya are crossed by transverse bundles of are cosmopolitan, nine Indopacific and three Atlantic. Bathymetric
auxiliary megascleres. In Stellitethya and in the encrusting genera, distribution is from intertidal to 2100m, although the majority of
Tethytimea and Xenospongia the megasclere skeleton shows paral- known species live in shallow waters between 0-50m depth.
lel tracts coming from the substrate to the upper surface of the
sponge. The megasclere tracts end always with diverging fans that Terminology
make the surface tubercles bristly. Megascleres may be strongylox-
eas (globose and stalked genera), styles, anisostrongyles, subty- Spicular terminology for Tethyidae differs among recent
lostyles and tylostyles (massive and encrusting genera). In the authors as, for example, in the aster classification and tenns
globose Oxytethya they are, exceptionally, oxeas. Megasters are reported by Bergquist & Kelly Borges (1991). Some of these dif-
generally spherasters or oxyspherasters but may be represented by ferences are not trivial but concern basic assumptions that require
oxyasters in genera such as Burtonitethya, Halicometes, further clarification. In this chapter I follow the indications of Sara
Tethyastra, Tethycometes and Xenospongia, and by anthasters in (1994), which agree with the common use of tenns in the majority
Anthotethya. Micrasters in the genus Tethya may be unifonn, gen- of papers on Tethyidae. Accordingly, I have adopted the following
erally tylasters, or polymorph, oxyasters, strongylasters, tylasters, terminology for asters. Asters are separated into two categories,
mingled or split in two categories, different in size and shape, for megasters and micrasters. Megasters are named spherasters, when
the cortex (smaller tylasters) and the choanosome (larger the RIC ratio (ray length compared to center diameter) is less than
oxyasters). In several other genera micrasters are unifonn, gener- 1; oxyspherasters (=spheroxyasters) when RIC is 1 or between
ally strongylasters, but in Columnitis, Anthotethya, Laxotethya and 1-2; oxyasters when this exceeds 2. Micrasters have spiny rays and
Oxytethya they are split into two different categories. Other types are differentiated into tylasters with tylote ray tips, chiasters with
of microscleres occur in some genera: polyrhabds in two species of truncated, strongylasters with rounded, and oxyasters with pointed
Tethya and in Tethyastra, exotyles in Halicometes, spheres in ray tips. This difference between the micraster types, using optical
Tethytimea, microstrongyles in Tethycometes. microscopy, is based essentially on the spinosity of the ray tips
The number of genera has gradually increased over recent (and confinned by electron micrographs), which is maximal in
years. The revision of Burton (1924) listed four genera; Sara tylasters and minimal in strongylasters and oxyasters. Chi asters
(1994) included eight genera; Sara & Bavestrello (1996) nine represent an intennediate situation. The difference between
genera, Sara & Sara (2002) 12, and now 14 are recognised in the strongylasters and oxyasters is related not only to the tip, rounded
present work. Described valid species of Tethyidae number 81 or pointed, but also to the ray shape that is cylindrical in strongy-
(56 alone for the genus Tethya), with about 40 recognizable but still lasters and more-or-Iess conical in oxyasters. Incorporated sedi-
undescribed new species remaining. The family is practically cos- ment is important in the genera Columnitis, Tectitethya,
mopolitan, particularly rich in tropical seas, but there appears to be Anthotethya, Laxotethya and Xenospongia and may represent a
a special zone of diffusion in the temperate shores of south-eastern skeleton support.

KEY TO GENERA

(1) Without basal stalk (or with stalk not exceeding half of the body diameter) ...................................................................................... 2
With basal stalk longer than half of the body diameter .... ........ ..................................................................................... .... .................. 8
(2) With a well developed and distinct cortex, body spherical or subspherical ..... .......... .... ........ .... .... ............. ...... .... .... ....... ............. ...... 3
With a little developed or indistinct cortex, irregularly massive or encrusting body.......... .................................. ........................ ...... 4
(3) Megasters spherasters or oxyspherasters, rather homogenous in size not exceeding 180 J.Lm in diameter ................................ Tethya
Megasters oxyasters, very heterogeneous in size even exceeding 180 J.Lm in diameter ....................................................... Tethyastra
Without megasters, megascleres oxeas ..... .................................. ...... .... .... ........ .... ............................. .... ...... ........ ................. Oxytethya
(4) Megascleres styles, subtylostyles or strongyles ................................................................................................................................... 5
Megascleres tylostyles ..... .... ...... ................. ...... ...... ...... ......... ................................. ...... ....... ............... .... ...... .... .... ............... Tethytimea
(5) Megascleres in radiate tracts proximally crossed by thinner megascleres ....................................................................... Nucleotethya
Megascleres in radiate ascending tracts, branching and anastomosing .. ...... ................ ........................... .......... .... .... .......................... 6
Megascleres in parallel ascending tracts .... ........................... .... ....... ................ ...... .... ................... ....................................................... 7
(6) Megasters anthasters ............................................ ......... ...................... ........................................................ ....................... Anthotethya
Megasters spherasters or oxyspherasters .. ...... ...... ............... ........................... ...... .... ........ .... ....... ........................ ................ Tectitethya
Without megasters, megasclere tracts partially loosened or confused ........................................... ............. .... ........... ......... lAxotethya
(7) With megasclere tracts arising from a central sediment core, proximally crossed by thinner megascleres, with
a winding cylindrical body .................................................................................................................................................. Columnitis
With megasclere tracts arising from a sediment basis, discoid body........... ...... ...... .... ........................... ...... .... ............... Xenospongia
With giant megasters, thick encrusting or massive body........ ..................................... ...... ........ ......................................... Stellitethya
(8) With stalk about the same length of body diameter ........... ........ ..................................................................................... Burtonitethya
With stalk about two to four times the body diameter .................................. .......... .............................. ................. ........... Halicometes
With stalk about eight times the body diameter ................................ .... .... ...... .......... ........ .......... .... ....... ...... .... .... ........... Tethycometes
248 Porifera' Demospongiae • Hadromerida • Tethyidae

Fig. 1. Tethya aurantium. A, neotype. B, skeleton structure. C, strongyloxeas. D, spherasters. E, cortical (above) and choanosomal (below) micrasters
(scales: A, I cm; B, 1 mm; C, D, 50 11m; E, 12ILm).

TETHYA LAMARCK, 1814 Micrasters are tylasters, strongylasters or oxyasters, normally with
spined rays, with polyrhabds in some species.
Synonymy
Description of type species
Tethya Lamarck, 1814: 71; Selenka, 1880: 472; Lendenfeld,
1888: 48; Sollas, 1888: 427; Thiele, 1898: 11. Donatia Nardo, 1833: Tethya aurantium (Pallas) (Fig. lA-E).
522. Lyncuria Nardo, 1833: 715. Amniscos Gray, 1867a: 549. AZemo Synonymy. Alcyonium aurantium Pallas, 1766: 357.
Wright, 1881: 16. Tethyorrhaphis Lendenfeld, 1888: 51. Taboga de Alcyonium Zyncurium Linne, 1767: 1295; Tethya Zyncurium
Laubenfels, 1936a: 452. Lamarck, 1814: 71; Donatia Zyncurium Nardo, 1833: 522. Tethya
morum Schmidt, 1862: 44. Tethya aurantium Topsent, 1920d: 640;
Type species Sara & Melone, 1965: 123. Tethya limski MUller & Zahn, 1968:
473. Taxonomic decision for synonymy: Topsent, 1920d, and
AZcyonium aurantium Pallas, 1766: 357 (subsequent designa- Sara & Melone, 1965.
tion by Topsent, 1920d: 643). Material examined. Holotype: unknown, "seas of Europe".
Neotype: MSNG 49670 - Bay of Naples. Other material. Many
Diagnosis specimens from different Mediterranean localities: Azores, western
North African coast; spicular slides from the Caribbean.
Tethyidae with a spherical, sometimes hemispherical body Electrophoretic allozyme patterns of Mediterranean populations
with a well-developed cortex, distinct from the choanosome (Sara et aZ., 1989; Sara, 1990; Sara & Manara, 1991; Sara et aZ.,
(medulla), dense or rich in lacunae. Main skeleton formed by 1992; Bavestrello & Sara, 1994).
strongyloxea bundles radiating from the center of the sponge and Description. Shape spherical, 1-5 cm in diameter, surface
bristling, generally flattened, sometimes conical, tubercles on the orange with rounded and flattened tubercles, consistency moder-
surface. The whole choanosome or its periphery may be filled by ately hard, with a thick and dense cortex without large subdermal
thinner auxiliary megascleres that accompany the distal fans of lacunes. One oscule is sometimes visible, as in the neotype.
megasclere bundles in the tubercles. Main megascleres are usually Megascleres represented by main, thicker, and auxiliary, thinner,
strongyloxeas, interstitial (auxiliary) megascleres are often styles. strongyloxeas but the two categories overlap in their lengths and
Megasters and micrasters are variously distributed in the cortex widths. Length 400-2500 fLm, width 5-30 fLm. Fans of radiating
and choanosome. Megasters are spherasters or oxyspherasters. megasclere bundles not forked. Interstitial megascleres fill the
Porifera· Demospongiae • Hadromerida • Tethyidae 249

space among the main megasclere bundles. Megasters represented mainly hydrodynamic conditions. In coral reefs Tethya may grow
by spherasters 18-105 fLm in diameter, generally 50--80 fLm under coral or in small crevices in an irregular, often cushion-like
(mean 62.04 ± 11.34) with RJC = 0.4-0.6 (mean 0.49 ± 0.14) shape. Some species, also in temperate seas, have anchoring rhi-
(Bavestrello & Sara, 1992). Megasters densely packed in many zoids. External color is often a specific trait and ranges from red,
layers in the endocortex. Some meg asters also in rows or scattered pink, orange, yellow, green, gray, brown to black. The surface may
in the mesocortex, especially along the canals and in the outer be covered by sediment. Consistency may be hard or soft, related to
choanosome. Micrasters are different between the cortex and the different development of lacunes and the distribution and den-
choanosome. In the cortex they are represented by a heterogeneous sity of the megasters. Oscules are rarely visible, and when they are
set of tylasters, strongylasters and slender oxyasters, 10--15 fLm in there is only a single apical osculum. Cyanobacterial filamentous
diameter (mean 12.5 fLm), with 9-14 apically spined rays. They are symbionts occur frequently in the cortex, influencing its color. Aster
distributed both as a dense layer on the surface of the sponge and shape, size and distribution represent the main diagnostic features
around the canals and lacunae of the cortex. In the choanosome for distinguishing species. Megasters may be different in shape and
they are represented by an uniform set of oxyasters, 15-25 fLm in size, and their distribution in the cortex and choanosome follows
diameter (mean 18.5 fLm) with 12-14 rays bearing few scattered specific patterns. Micrasters may be uniform in the cortex and
microspines, and ray tips are more spined and slightly knobbed choanosome, and are generally tylasters, or there may be different
under light microscopy. Both types of micrasters occur jointly in shapes mingled in the same part of the sponge, or they may also dif-
the inner cortex. Reproduction is sexual (in summer) and asexual, fer in shape and size between the cortex and the choanosome. Buds
by external buds (in winter). frequently occur, generally external, sometimes internal.
Geographic distribution. Mediterranean, Atlantic North-
African coasts, Azores. This restricted distribution depends from
the distinction between T. aurantium and T. citrina, and from a crit- ANTHOTETHYA SARA. & SARA., 2002
ical evaluation of the existing data. In the Mediterranean both
aurantium and cifrina occur, but in North European seas only Synonymy
T. cifrina is present, eventually flanked by T. norvegica. All the
museum specimen labelled 'aurantium' or 'lyncurium' examined Anthotethya Sara & Sara, 2002.
by myself, and the newly collected material of Tethya coming from
the Channel and northern seas, belong to T. cifrina Sara & Melone Type species
or T. norvegica Bowerbank. The attribution of Caribbean popula-
tions to this species, as well as its reported occurrence along the Anthotethya Jromontae Sara & Sara, 2002 (by original
South-European Atlantic coasts, should be confirmed. Californian designation).
(Monterey Bay) specimens of T. aurantia var. califomiana de
Laubenfels, 1932 have been assigned to the new species T. califor- Diagnosis
niana (Sara & Corriero, 1993). The material labelled T. aurantium
coming from Australia and New Zealand and deposited in BMNH, Tethyidae with irregular conical shape and thin cortex. The
partially described as T. aurantium by Bergquist & Kelly-Borges megasclere skeleton is made by ascending tracts of styles (some-
(1991), does not belong to T. aurantium but to a different species times anisostrongyles or subtylostyles) and interstitial styles. Main
(based on its described traits and from my personal observations on tracts branch at different body levels and also may anastomose.
this material). Depth range: 0.3-40m (Pulitzer-Finali, 1970; Megasters are represented by anthasters, and micrasters by spiny
Bavestrello & Sara, 1994). strongyl asters and oxyasters.
Remarks. As noted by Topsent (1920d), the first figures of
Tethya are by Aldrovandi (1642) and Marsilli (1725), and the first Description of type species
description and assigned name is Tethya sphaerica by Donati
(1750). But, as this name is prior to 1758, the first valid name for AnthotethyaJromontae Sara & Sara, 2002 (Fig. 2A-F).
the type species is Alcyonium aurantium Pallas, 1766. The taxo- Synonymy. Anthotethya Jromontae Sara & Sara, 2002.
nomic identity of T. aurantium has been well defined by Sara & Material examined. Holotype: WAM Z5793. Paratype:
Melone (1965), with indication that two different species T. auran- WAM Z5410 - Dampier Archipelago, Western Australia. Depth:
tium (Pallas, 1766) and T. citrina (Sara & Melone, 1965) have intertidal to 14 m.
often been confused. Pallas' description refers clearly to T. auran- Description. The body is irregularly conical, 5-6 cm high
tium, but under the name 'aurantium' or 'lyncurium' the successive and broad, with some little distinct rounded lobes 2 cm high. The
authors have also described T. cifrina. surface is covered by small flattened tubercles 1-2 mm wide, often
Tethya contains the most species of the family, with 56 species transformed in narrow irregular ridges with a winding pattern.
presently described and about 40 others recognized but hitherto not The cortex is about 0.5 mm thick and distinct from the choanosome
yet described. Its distribution (Sara, 1998) is nearly cosmopolitan, due to a dense concentration of anthasters and micrasters. Lacunae
mainly tropical, with a peculiar biodiversity in the temperate and canals are abundant throughout the body. Foreign particles
Australia-New Zealand area. Indo-pacific species represent about occur on the surface and inside the sponge. Consistency elastic.
85% of the total diversity. The occurrence in Arctic seas is evi- Color (in ethanol) externally reddish-orange, internally ochre-
denced by only one species, and in Antarctica only by scattered brown. The skeleton shows ascending megasclere tracts that
spicules in the sediment. Depth distribution is between 0--805 m. generally run in parallel and end at the surface in small fans.
Most species have been found in shallow waters (Burton, 1958). Megasclere tracts may often branch and anastomose at different
External features are important in distinguishing species but the body levels. Auxiliary megascleres placed in the interstices
growth form of Tethya may be influenced by environmental, between the tracts are very abundant throughout the sponge.
250 Porifera· Demospongiae • Hadromerida • Tethyidae

Fig. 2. AnthotethyaJromontae. A, holotype. B, skeleton structure. C, stylote and subtylote bases of megascieres. D, anthasters. E, oxyasters. F, strongy-
lasters (scales: A, 2cm; B, 1 mm; C, F, 5 /-Lm; D, E, 10/-Lm).

Megascleres are generally styles but also anisostrongyles and Type species
subtylostyles: 300-1750 X 5-35 /-Lm. Megasters are anthasters with
4, or more frequently, 6 strong rays that fork in 3-5 (generally 4), Tethya gemmijormis Burton & Rao, 1957 (by original
slightly curved strong spines, 40-80 /-Lm in diameter. Micrasters are designation).
strongylasters, 10-20/-Lm with 8-10 microspined rays, and
oxyasters, 10-50 /-Lm with 3-8 smooth rays and bifid tips. Diagnosis
Remarks. This monotypic genus from Australia was erected
primarily for its unique shape of megasters, resembling anthasters Tethyidae with subspherical body and a basal stalk, approxi-
as in the Hemiasterellidae genus Axos (Hooper, 1986). Other fea- mately as long as the body diameter. Cortex rich in lacunae, poorly
tures unique amongst Tethyidae are the cortex and the skeleton developed; with a large central spicular nucleus placed at the inser-
structure, and the micrasters are also very peculiar. On the whole, tion of the stalk skeleton in the radiate body skeleton. The nucleus is
the massive shape, the poorly developed cortex, the branching and formed by clumps of thin strongyles and styles around the bases of
anastomosing skeleton with little fusiform styles as megascleres, the radiating bundles of megascleres, represented by strongyloxeas.
and the abundance of sediment particles in the body place this genus Surface with peculiar button-like tubercles, fringed by megascleres
near the Australian Laxotethya and the Caribbean Tectitethya. projecting nearly horizontally to, or at very acute angles with, the
surface. Megasters represented by oxyasters filling the whole
sponge, very heterogeneous in size, and reaching giant dimensions,
BURTONITETHYA SARA, 1994 up to 300 /-Lm in diameter. Micrasters represented by strongylasters.

Synonymy Description of type species

Burtonitethya Sara, 1994: 358. Burtonitethya gemmijormis Sara, 1994 (Fig. 3A-F).
Porifera· Demospongiae • Hadromerida • Tethyidae 251

CJ {J oD

B C-

Fig. 3. Burtonitethya gemmiformis. A, reconstruction of the sponge structure. A' , reconstruction of the whole body. B, main strongyloxea. C, accessory
style. D, slrongyles. E, micrasters. F, megasters (oxyasters) (scales: A, 4mm; A', IOmm; B-D, 20J.l.m; E, 10 J.l.m; F, 100 J.l.m).

Synonymy. Burtonitethya gemmiformis Sara, 1994: 358. megasters, represented by oxyasters (which fill the sponge with a
Material examined. Holotype: BMNH 1957.7.15.1 (section heterogeneous set of sizes and include giant spicules), a reduced
slide) - Andaman Islands, NE. Indian Ocean, depth unknown. Other cortex, a stalk and a nucleus which surrounds the bases of megas-
material. BMNH 1934.11.24.392 (section slide) - same locality. cleres and formed by peculiar spicules.
Description. Body shape a depressed sphere 8 mm high and
10 mm broad. Overall height of body and stalk about 18 mm. Stalk
10 mm long formed by bundles of megascleres coated by a thin layer COLUMN/TIS SCHMIDT, 1870
of micrasters. Cortex is less than 0.5 mm thick, with lacunae. Surface
rough, with button-like tubercles, 0.5 mm high. Color pinkish-brown Synonymy
in ethanol. The body center is occupied by a large spicular nucleus
formed by transverse and concentric rows of thin styles and Columnitis Schmidt, 1870: 25; Sollas, 1888: 441; Lendenfeld,
strongyles, frequently slightly bent, around the bases of the strongy- 1890a: 397; Sara, 1994: 361; Sara & Bavestrello, 1996: 374.
loxeas bundles. Main strongyloxeas of the radiating bundles very Donatia sensu Burton, 1924: 1034. TImea sensu de Laubenfels,
long, 2400 X 20 ILm. Accessory strongyloxeas, styles and strongyles 1932: 47. Tethya sensu Wiedenmayer, 1977b: 171. Taxonomic
200--600 X 3-10 ILm. Auxiliary megascleres, besides the nucleus, decision for synonymy: Sara, 1994: 361.
are found along the main bundles and stop just below the surface.
Megasters are oxyasters, 60-300 ILm in diameter, with a small cen- Type species
trum about 10% of the ray length (RIC = 10). Rays 8-12, often irreg-
ularly bent in their apical and middle tract, sometimes forked or with Columnitis squamata Schmidt (by monotypy).
rounded tips. Micrasters are slightly spined strongylasters, 10012lLm
in diameter, with the ray tips little enlarged, without centrum and Diagnosis
with, on the average, 12 slender rays. The micrasters are uniform
throughout the whole sponge and form a dense layer on the surface. Tethyidae with a thickly encrusting or massive body, horizon-
Remarks. This monotypic genus from the Indian Ocean tally developed into winding branches and lobes. Surface with
was erected for a species mentioned in a manuscript of Burton and flattened, roughly hexagonal tubercles, contiguous above and
attributed to Tethya. But this species is sharply differentiated from separated below by grooves. Cortex indistinct but a thickening of
the other Tethya species based on the distribution and shape of their asters under the surface is present (pseudocortex), with large
252 Porifera' Demospongiae • Hadromerida • Tethyidae

ill
ill
ill c

~ $*~
**** E

. Fig. 4. Columnitis squamata. A, holotype. B, skeleton structure. C, subtylostyles. D, megasters. E, micrasters (scales: A, 1cm; B, I mm; C, 30 fLm;
D, 50 fLm; E, 10 fLm).

subdermal lacunes. Skeleton made by parallel bundles of megas- Description. Irregular massive body horizontally developed
cleresending in the tubercles and crossed in the inner part of the with winding branches and rounded lobes. The erroneously nomi-
sponge by thinner megascleres isolated or in bundles. Megascleres nated neotype measures 7.5 X 5 X 2.5 cm. Surface smooth, covered
are subtylostyles, sometimes styles. Megasters are oxyspherasters. by flattened, roughly hexagonal tubercles. Parallel bundles of
Micrasters are of two sizes: the larger represented by oxyasters and megascleres arise from the inner sediment core and end with fans in
strongylasters, with rays almost completely covered by spines; the the tubercles. Thinner megascleres, generally in smaller transverse
smaller particularly abundant on the sponge surface, and repre- bundles or scattered, cross the main bundles in the inner part of the
sented by tylasters. A large amount of sediment, forming a massive sponge, around the sediment core. Megascleres are sUbtylostyles,
central core, is included inside the sponge. sometimes styles, 300-2000 X 3-30 f..Lm. The vertically disposed
main megascleres are generally 20-25 f..Lm thick while the second-
Description of type species ary transverse ones are only 5-10 f..Lm thick. Megasters and micras-
ters are densely distributed throughout the whole sponge but more
Columnitis squamata Schmidt, 1870 (Fig. 4A-E). heavily concentrated under the sponge surface in a pseudocortical
Synonymy. Columnitis squamata Schmidt, 1870: 25; layer. Megasters are oxyspherasters with rays often bent or swollen
Sollas, 1888: 441; Sara & Bavestrello, 1996: 374. at their tips, 50-80 f..Lm, RIC = 1-2. Micrasters in two categories:
Material examined. Fragment of holotype: 2MB 6560 strongylasters, sometimes oxyasters, with 12 spined rays, 15-25 f..Lm
(slide) - from Schmidt type (lost) - "Antilles?". Other material. and without center; and very small tylasters 3-5 f..Lm diameter.
USNM 34537 - Florida, Gulf of Mexico (erroneously nominated Remarks. This genus was erected by Schmidt for the
as Neotype by Sara & Bavestrello (1996», USNM 34548, species C. squamata, now represented only by a spicular slide
BA 831 - Florida, Gulf of Mexico. from the type stored in the 2MB . After checking this slide
Porifera· Demospongiae • Hadromerida • Tethyidae 253

Sara & Bavestrello (1996) identified three specimens from the Gulf Tethya stellata Sollas, 1888: 440. Halicametes stellata Topsent,
of Mexico as belonging to C. squamata, and one of these specimens 1898a: 112; Topsent, 1920a: 32.
was erroneously chosen as neotype of the species. The existence of Material examined. Lectotype: MZS.PO 114 A. Syntypes:
a surviving slide from the type in the 2MB, which is here recog- MZS.PO 114 A, B - two specimens labelled "Tethya cametes
nised as the holotype (ICZN Art. 72.5.5), allowed Sara & O. Schm. 24° 8N, 82° 51 W (329-339 Fd) Golf v. Mexico. Agassiz
Bavestrello (1996: 374) to identify three subsequent specimens of 1876". This material comes from the locality (North of Cuba)
this species (not from Schmidt's description but from the contents and from the depth of the original description by Schmidt. It
of the slide; ICZN Art. 75), negates the subsequent neotype desig- represents the two extremes of a range of variability from an
nation. On the basis of a more precise characterization of hispid and a smooth variety of the species as described by Schmidt
Columnitis from this new material it was also possible to attribute to (1879: 78) in an enlargement of the species diagnosis. MZS.PO
this genus the BMNH specimen 1938.b.30.38, labelled by Burton as 114C - labelled "Tethya cametes O. Schm. Golf v. Mexico.
"Tethya anomala" but not described. This specimen was described Agassiz". BMNH 1939.2.10.38, BMNH 1870.5.3.98 (teased
by Sara & Bavestrello (1996) as a second species of Columnitis, fragments).
C. anomala. Conversely, the BMNH specimen 1928.5.12.167 from Description. With an ovoid body, 7 mm high and a basal
the Crawshay collection (West Indies) labelled "c. squamata", does stalk 22 mm long. In the body, radiating megasclere bundles ending
not belong to this species but to Tectitethya crypta. in hispid cylindrical tubercles (in the more frequent variety). Less
Peculiar traits of Columnitis are the body shape, the inner core frequently, in another variety, tubercles are smooth. Megascleres of
of sediment, and the occurrence of two categories of micrasters, the choanosomal bundles are strongyloxeas, slightly flexuous (not
well-differentiated in size and shape, also in the absence of a dis- anisostrongyles as indicated by Topsent, 1920a): 1900-2250 X
tinction between cortex and choanosome. The larger category of 30-35 J..lm (15-20 J..lm at the base). In the stalk they are styles up to
micrasters occurs only in Columnitis. 3000 X 30 J..lm. Auxiliary megascleres, bristling also in the tuber-
Two West Indies species: C. squamata is known from the cles, are thinner styles, slightly flexuous, 800-1000 X 10-12 J..lm.
Gulf of Mexico, off the south-western Florida coast, 30 m depth; Megasters of the body and stalk are oxyspherasters, with 18-20
C. anomala is known from the Caribbean, off the Nicaragua coast rays, very heterogeneous in size, 15-180 J..lm, with RJC = 1.2-1.8.
(Sara & Bavestrello, 1996). The RJC is higher in the hispid variety. Micrasters are chiasters,
with 12 micros pined rays, covering a dense layer at the surface of
the body and stalk and also scattered inside the sponge. They have
HALICOMETES TOPSENT, 1898 a diameter of 10-15 J..lm, and a centrum, more marked in the
smooth variety. The stalk is bristled by numerous exotyles, often
Synonymy flexuous, 45-325 X 5-7 J..lm. In the stalk there are also anomalous
asters with a reduced number of rays and differing in the shape of
[Cometella], in part, Schmidt, 1870: 49 (nomen oblitum). tetractines, triactines or diactines, 50-70 J..lm in diameter.
Tethya sensu Schmidt, 1880b: 78; Sollas, 1888: 440. Halicometes Remarks. The synonymy of Tethycardyla (stalked
Topsent, 1898a: 112; Topsent, 1920a: 32; de Laubenfels, 1936a: Tethyidae without exotyles) with Halicometes (stalked Tethyidae
145; Levi, 1993: 27; Sara, 1994: 361; Sara & De Rosa Barbosa, with exotyles) was established by Sara (1994) on consideration
1995: 167; Sara & Bavestrello, 1996: 255. Tethycordyla de that, at the present state of knowledge, the lack of the stalk exotyles
Laubenfels, 1934: 8; Levi, 1964a: 72. Taxonomic decision for as a unique distinctive trait is not sufficient to differentiate these
synonymy: Sara, 1994: 362. nominal genera. In addition to the fact that a single spicular trait,
even if important, should be used more for discriminating species
Type species than genera, the exotyles if rare may be easily overlooked or lost in
the preservation of the very tiny specimens of Halicometes.
Halicometes stellata (Schmidt, 1870: 49) (by original [Cometellal Schmidt, 1870 originally included [Cametella]
designation). gracilior Schmidt, 1870 (=Asbestopluma) and [Cometella] stellata
Schmidt, 1870. De Laubenfels, 1936: 151 designated the former
Diagnosis as type species of [Cametella] and thus Cametella is a cladorhizid
(proposed to be suppressed under ICZN Art. 29.3., see Hajdu &
Tethyidae with an ovoid body and a basal stalk 2-4 times Vacelet, this volume).
longer than the body height. Cortex indistinct or poorly developed. Halicometes is sharply differentiated from Tethya, not only
Exotyles on the surface of the stalk, sometimes rare or even for the stalk and the deep-water habitat, but also for the indistinct
lacking. The megascleres occur in radiating bundles, bristling at or reduced cortex and the megaster shape, size and distribution.
the surface tubercles, and may be strongyloxeas, anisostrongyles Based on megaster shape and size, and the occurrence of giant
or styles. The megasters, distributed throughout the sponge, are spiCUles, Halicometes slides in museum collections (such as the
oxyasters or oxyspherasters of different sizes, including giant BMNH) are sometimes erroneously labelled as Tethya repens
spicules (more than 150 J..lm in diameter). Micrasters may be Schmidt (or Donatia repens). However, Tethya repens Schmidt
strongylasters, chiasters, tylasters, sometimes oxyasters. 1870 from Florida is a species inquirenda, likely a Stellitethya, but
this cannot be unequivocally identified at the species level because
Description of type species its description is insufficient and the type is represented only by a
spicule slide. Presently Halicometes contains six species from deep
Halicametes stellata (Schmidt, 1870) (Fig. SA-G). waters (150-2100 m depth) of the Atlantic and Indo-Pacific regions
Synonymy. [Cametella] stellata Schmidt, 1870: 49. Tethya Halicametes stellata (Schmidt 1870) has been found off Cuba
caudata Dezso, 1879: 648. Tethya cometes Schmidt, 1879: 78. (650m depth) and the Gulf of Mexico (619m depth); H. thyris
254 Porifera· Demospongiae • Hadromerida • Tethyidae

Fig.S. Halicometes stellata. A, the two syntypes (on the left the hispid syntype chosen as lectotype, on the right the smooth syntype). B, sponge structure.
C, megascleres (main strongyloxea, auxiliary style and stalk style). D, exotyles. E, megasters. F, anomalous stalk megasters. G, micrasters (scales:
A, 2mm; B, Imm; C, 60 ILm; D-F, 50 ILm; G, 10 ILm).

(de Laubenfels 1934) off Puerto Rico (350m); H. pediculata (Levi, Diagnosis
1964a) off Durban (South Africa, Indian Ocean, 495 m); H. hooperi
Levi, 1993, off New Caledonia (2100 m); H. minuta Sara & De Rosa Tethyidae with massive irregular body, ill-defined cortex and
Barbosa, 1995 off the Brazilian coast (190m); H. stonei Sara & surface covered by irregular tubercles and sinuous ridges.
Bavestrello, 1996, west of Andaman Islands (732 m). Megasclere skeleton formed by subtylostyles packed in winding,
more-or-Iess compact tracts and scattered in the interstices. The
tracts branch and anastomose, and generally ascend towards the sur-
LAXOTETHYA SARA & SARA, 2002 face, ending in fans. Microscleres are only micrasters, mainly
tylasters, but also smaller oxyasters, filling the whole body. Sediment
Synonymy particles are abundant on the surface and inside the sponge.

Laxotethya Sara & Sara, 2002. Description of type species

Type species Laxotethya dampierensis Sara & Sara, 2002 (Fig. 6A-G).
Synonymy. Laxotethya dampierensis Sara & Sara, 2002.
Laxotethya dampierensis Sara & Sara, 2002 (by original Material examined. Holotype: WAM Z3948 - Dampier
designation). Archipelago, Western Australia, depth 20 m.
Porifera· Demospongiae • Hadromerida • Tethyidae 255

Fig. 6. Laxotethya dampierensis. A, holotype. B, skeleton structure. C, tylote and subtylote bases of megascleres. D, megasclere. E, convergence and
forking of megasclere tracts. F, oxyasters. G, small and large tylasters (scales: A, 2 cm; B, I mm; C, G, 10 ILm; D, 100 ILm; E, 500 ILm; F, 5 ILm).

Description. Cushion-shaped compressed body of 5.5 X peculiarities are the body shape, the subtylostyle, and sometimes
3 X 3 cm. The scabrous and uneven surface is covered by irregular tylostyle megascleres, the lack of megasters and the abundance
tubercles, 1-3 mm broad, 1-2mm high, fused partly into winding of sediment. These traits put this genus near the Australian
ridges. The cortex is macroscopically indistinct and may be Anthotethya and the Caribbean Tectitethya.
observed only as a 0.5 mm wide histologically peculiar zone. The
aquiferous system shows some small oscules, grooves harboring
cribriporal areas, and several lacunae throughout the body. NUCLEOTETHYA SARA & BAVESTRELLO, 1996
Consistency firm and color (in ethanol) whitish-brown externally
and internally. Megasclere tracts ascend towards the surface where Synonymy
they end in small fans, and may be compact or loosened, run in a
winding pattern, branching and anastomosing. The interstices Nucleotethya Sara & Bavestrello, 1996: 380.
between tracts are filled by abundant scattered megascleres.
Generally the skeleton pattern may be defined as dendritic reticu- Type species
late. Megascleres are short subtylostyles, sometimes tylostyles and
styles, 300--920 X 5-20 fLm. Megasters are absent. Micrasters are Nucleotethya bifida Sara & Bavestrello, 1996 (by original
generally tylasters, 10--20 fLm, smaller in the micraster crust of the designation).
surface zone, with 4-12 rays bearing spiny knobs, or more rarely
oxyasters, 5-15 fLm in diameter, with 4-10 thin rays and often with Diagnosis
a small apical knob.
Remarks. This monotypic genus from Australia was erected Tethyidae with a conical or ovoid massive body. Surface with
for a Tethyidae characterized by an extensive loosening, anastomos- flattened, rounded or polygonal tubercles. Cortex indistinct, marked
ing and branching of the ascending megasclere tracts, producing only by the density of asters. Skeleton composed of radiating bun-
a skeleton pattern that may be defined as dendritic-reticulate. Other dles of megascleres ending in the tubercles, and in the inner part of
256 Porifera· Demospongiae • Hadromerida • Tethyidae

D
c

Fig.7. Nucleotethya bifida. A, holotype. B, skeleton structure. C, styles. D, megasters. E, micrasters (scales: A, I cm; B, I mm; C, 30 !Lm; D, 100 !Lm; E, 5!Lm).

the sponge by transverse bundles of thinner megascleres crossing anisostrongyles, 800-2300 X 20-30 f.Lm, ending in tubercles with
the radiating bundles and surrounding a large spicular nucleus fan-like expansions. The transverse rows are composed of thinner
(about 0.5--0.66 of the sponge diameter). The nucleus is made of a styles, 350-1600 X 5-10 /-Lm. The styles in the nucleus measure
confused network of megascleres accompanied by meg asters, 300-2000 X 5-20 /-Lm. Megasters are oxyspherasters 40-160 f.Lm in
rnicrasters and sediment. Megascleres are styles and subtylostyles. diameter (in the nucleus 30-80/-Lm) with RIC = 1-1.5, and rays
Megasters are oxyspherasters and micrasters tylasters. Surface cov- generally truncated with forks and teeth. Micrasters are tylasters
ered by sediment, abundant also inside the sponge. with slender rays and swollen tips, in diameter 7-12, sometimes
15-18/-Lm.
Description of type species Remarks. The genus Nucleotethya was established prima-
rily for its nucleus, which strongly differs in its structure from other
Nucleotethya bifida Sara & Bavestrello, 1996 (Fig. 7A-E). nuclei occasionally found in Tethyidae. Other distinctive characters
Synonymy. Nucleotethya bifida Sara & Bavestrello, differentiating it from Tethya are the indistinct cortex, the large
1996: 380. size of the megasters, and the megascleres represented generally
Material examined. Holotype: USNM 34558 - SW Florida by styles or subtylostyles. Nucleotethya belongs to the same
coast, Gulf of Mexico, 85 m depth. Paratypes: USNM 34559, BA clade as Columnitis and Tectitethya based on its external appear-
477 - same locality. ance, the indistinct cortex, the megasc1ere shape and the abun-
Description. Body irregularly conical (6 X 3 X 5 cm for dance of sand. The only known species comes from SW Florida
holotype and one paratype) or ovoid (the other paratype). The color coast.
(in ethanol) is brownish gray at the sandy surface and pink-brown
in the interior. The consistency is hard, incompressible. One small
oscule located on the top of the sponge. Large subdermal lacunes OXYTETHYA SARA. & SARA., 2002
under the pseudocortex. The surface is finely hispid and covered by
sediment. The tubercles are flattened, 2-4 cm in diameter, rounded Synonymy
or with a roughly polygonal outline. The skeleton is composed
of radiating bundles of styles or subtylostyles, sometimes Oxytethya Sara & Sara, 2002.
Porifera' Demospongiae • Hadromerida • Tethyidae 257

Fig. 8. Oxytethya mirabilis. A, holotype, lateral view. B, holotype, view of the surface. C, skeletal structure. D, oxea in three sections. E, oxyspherasters.
F, oxyasters. G, oxyasterlstrongylaster. H, polyrhabds (scales: A, B, 2cm; C, 1 cm; D, 100 fLm; E, 2 fLm; F, G, 5 fLm; H, 10 fLm).

Type species Synonymy. Oxytethya mirabilis Sara & Sara, 2002.

Material examined. Holotype: AM Z5684 - Long Reef,
Oxytethya mirabilis Sara & Sara, 2002 (by original designation). New South Wales, SE Australia, depth 37.5 m.
Description. The holotype, represented by a large fragment
Diagnosis of a radial sector of the body, 9.5 X 8 cm, is reconstructed to proba-
bly be globose (subspherical or spherical), with a probable diame-
Tethyidae with globose, subspherical or spherical body and ter of about 12-16 cm. The cortical surface is covered by pinnacles,
radiate skeleton. Cortex well developed. Surface with high pinna- 2.5 cm high, connected by crests irregularly spaced and sometimes
cles and crests. Megascleres are oxeas with similar ends and widest fused. The surface texture is characterized by megasclere tracts,
at the middle of the spicule. Megasclere tracts branch and anasto- visible in transverse section, with the cortex under the pinnacles
mose in the choanosome. Microscleres are represented only by and crests having a thickness of 2-3 cm. The aquiferous system
micrasters, which are oxyasters, oxyspherasters, and polyrhabds. shows large lacunae in the cortex and choanosome. The consis-
tency is elastic, the color in ethanol is pale brownish. Megascleres
Description of type species radiate in partially winding tracts and may branch, but also anasto-
mose in the choanosome. They run tangential to the surface, ending
Oxytethya mirabilis Sara & Sara (Fig. 8A-H). without fan expansions in the pinnacles and crests. Main megascleres
258 Porifera· Demospongiae • Hadromerida • Tethyidae

are giant oxeas, 4000-4970 X 50-65 fJ.m, straight or moderately Material examined. Lectotype: BMNH 1925.11.1111 (slide
bent in the middle, with similar tips sometimes sharp but more from ZMH syntype). Hooper & Wiedenmayer (1994) report the
often a little blunt or tapered. Auxiliary megascleres are also oxeas, following type data. Syntypes: 2MB 6589 (2 slides), slide BMNH
180-210 X 3-5 fJ.m. Megasters are absent. Micrasters are repre- 25.11.1111 (slide from type material), 2MB 4267 (fragment of
sented by a varied set of oxyspherasters, oxyasters, oxyastersl type), BMNH (unregistered slide, ex ZMH 15302/54). Lectotype
strongylasters. Other microscleres are polyrhabds. Oxyspherasters locality: NW of Middle Bluff Shark Bay, Western Australia, 7-8 m
and oxyasters with a more-or-less developed center, measuring depth, rocky bottom with corals. The locality of the other syntype
8-20 fJ.m in diameter, with 12-16 short conical rays, with rays gen- is not recorded.
erally spiny. The rarer oxyasters/strongylasters measure 15-20 fJ.m Description. The two specimens described by Hentschel
in diameter, with 8-12 cylindrical spiny rays, and sometimes a differ in shape and size but are apparently similar in their spicular
poorly developed center. The polyrhabds are varied in size and traits. The lectotype is cushion shaped and thickly encrusting (9.5 X
shape, 10-16 X 3-10 fJ.m, with irregularly verticillated spines. 6 X 1.5 cm). The skeleton shows parallel megasclere bundles that
Remarks. Oxytethya is a very distinct genus of Tethyidae, so run from the sponge base to the tubercles, where they expand into
far known only from the Pacific coast of SE Australia characterized large fans. The other dry syntype, represented in Fig. 9 (reproduced
primarily by the extraordinary occurrence of oxeas as main and aux- from the original illustration of Hentschel, 1909), is irregularly
iliary megascleres but also by the surface structure and the lack of massive (19 X 11 X 6.5 cm). Cortex is indistinct. Surface with con-
megasters. In some traits, such as the occurrence of polyrhabds, there tiguous flattened tubercles, variable in breadth and height. In the
is a similarity to some Tethya, and consequently its phylogenetic lectotype they are 1-3mm broad, often low and only in some
position is likely to be near the Tethya clade. An association of oxeas places 3 mm high. In the syntype they are 3-10 mm broad and high.
with euasters occurs also in Asteropus Sollas, 1888 and Melophlus The surface is covered with sediment, which is also found inside
Thiele, 1899. Both these genera were transferred to Ancorinidae the sponge, and by a thick layer of tylasters. External color
(Hajdu & Van Soest, 1993), and Oxytethya differs sharply from them (in ethanol) is reddish-yellow on the free surface. Main and auxil-
in its skeleton (formed by well-defined radiating megasclere tracts) iary megascleres are styles, sometimes distally abbreviated. In the
and without their specialized tangential ectosomal skeleton. The lectotype the largest megascleres are generally anisostrongyles.
microrhabds of Melophlus are different from the polyrhabds of Their sizes (reported by Hentschel) for the main and auxiliary
Oxytethya and other Tethyidae. megascleres are respectively of 1056--2560 X 35-52 fJ.m and
701-1353 X 12-21 fJ.m (700-3250 X 5-60 fJ.m from re-examination
of the lectotype). Megasters are oxyspherasters with 20-24, some-
STELUTETHYA SARA, 1994 times dichotomous rays, 20-200 fJ.m in diameter, RIC = 1-1.5,
generally 1.25. Micrasters are tylasters with 10-12 slender rays,
Synonymy 10-13 fJ.m (10-19 fJ.m according to Hentschel).
Remarks. The two specimens described by Hentschel
Tethya sensu Schmidt, 1870: 51; Wiedenmayer, 1977b: 172. (1909), and considered to be the syntypes differ in shape and size but
Donatia sensu Hentschel, 1909: 374. Tethytimea sensu Burton, are similar in spicular traits. The lectotype designation is made for
1937: 12; Thomas, 1968a (thesis unpublished); Thomas, 1980: 14. 2MB 6589 on the basis that it has a known type locality and the
Stellitethya Sara, 1994: 362; Sara & Bavestrello, 1996: 257. skeleton structure is known and can be corroborated with Hentschel's
Taxonomic decision for synonymy: Sara, 1994: 362. description. The other syntype has not been designated as paralecto-
type because it has yet been personally examined by me.
Type species The first record of a species pertaining to the genus
Stellitethya may be Tethya repens Schmidt, 1870 from Florida.
Stellitethya extensa (Hentschel, 1909) (by original Unfortunately, Schmidt's description is incomplete and the only
designation). clue for this possible assignment is its thickly encrusting shape.
'TYpe material is represented only by the spicule slide BMNH
Diagnosis 1870.5.3.91, which shows some resemblance in its megasters with
those of S. stellagrandis, but this material is not sufficient to con-
Tethyidae with a body of large size and irregularly massive clusively decide on the precise taxonomic status of repens.
shape, sometimes subspherical, frequently thick encrusting. Without Consequently, Sara (1994) chose Donatia jissurata var. extensa as
a distinct cortex. Megascleres styles or strongyloxeas (fusiform type species (Donatia jissurata Lendenfeld, 1888: 48, should be
styles), vertically crossing the sponge in parallel bundles and ending assigned to Tethya according to the original description). However,
in large flattened tubercles at the surface. Megasters represented by Tethya repens has been also attributed to the genus Tethytimea de
oxyspherasters, filling the whole sponge, heterogeneous in size, and Laubenfels, 1936a because, erroneously, Burton (1924) considered
including giant spicules (more than 150 fJ.m in diameter and reach- Donatia tylota Hentschel, 1912, the type species of Tethytimea, as a
ing 500 fJ.m in two species). Micrasters usually represented by synonym of T. repens. Stellitethya is, however, differentiated from
tylasters. Tethytimea by many important structural and skeletal characters.
Stalked specimens from the Andaman Islands have also been
Description of type species assigned to Donatia repens by Dendy & Burton (1926), but these
likely belong to Halicometes. Indeed, I have found that under the
Stellitethya extensa (Hentschel, 1909) (Fig. 9A-F). name Tethya repens' or 'Donatia repens' there are labelled slides
Synonymy. Donatiajissurata var. extensa Hentschel, 1909: or specimens in the BMNH which belong to other species of
374. Stellitethya extensa Sara, 1994: 362. Tethya repens; Hooper & Stellitethya, such as S. murrayi, but also perhaps to the very different
Wiedenrnayer, 1994: 425. genus Halicometes (for the size and shape of the megasters).
Porifera • Demospongiae • Hadromerida • Tethyidae 259

000
D 0 0
QcO ~I

Fig. 9. Stellitethya extensa. A, the syntype figured by Hentschel. B, skeleton structure (lectotype). C, main styles and anisostrongyles. D, auxiliary styles.
E, megasters. F, micrasters (scales: A, 1 cm; B, 1 mm; C-E, 50/Lm; F, 10 /Lm).

The genus contains four species from the Indian Ocean, South Type species
Arabian coasts (Sarli & Bavestrello, 1996) and S Western Australia
(Hentschel, 1909), with depth range 4O--70m. The possible extension Tectitethya crypta (de Laubenfels, 1949b) (by original
of the geographic range of the genus to the West Indies depends on designation).
a more certain attribution of Tethya repens to Stellitethya.
Diagnosis

TECTITETHYA SARA, 1994 Tethyidae with a massive, amorphous, irregularly conical,

cylindrical or plurilobate body of large size. The cortex is ill
Synonymy defined, nearly indistinct. Tubercles flattened and irregularly
rounded, of different size. The surface is covered by sediment or
Cryptotethya sensu de Laubenfels, 1949b: 20 (not: Dendy, sea-weed, and sediment may also be found inside the sponge.
1905). Tethya sensu Reiswig, 1971 : 569; Wiedenmayer, 1977b: Megascleres are anisostrongyles or strongyles, frequently with
171. Tectitethya Sara, 1994: 362; Sara & Bavestrello, 1996: 381. subtylote heads, forming stout bundles that end in fans at the tuber-
Taxonomic decision for synonymy: Sara, 1994: 362. cles and frequently branch and anastomose along their course.
260 Porifera· Demospongiae • Hadromerida • Tethyidae

* * 1(1
*" + $}f

ill OJrnW(I)m~ * =0=

D
Fig. 10. Tectitethya crypta. A, paratype. B, skeleton structure. C, anisostrongyles. D, megasters. E, micrasters (scales: A, B, I em; C, D, 20 fLm; E, 10 fLm).

Scattered megascleres lie in the interstices among the bundles. gray-olive. Megascleres in radiating bundles frequently branched
Megasters, sometimes rare, are represented by oxyspherasters of and fused, described by de Laubenfels as strongyloxeas, but are
small or medium size, with rays frequently reduced or anomalous. generally anisostrongyles to strongyles, 500-1400 X 10-40 !-Lm.
Micrasters are represented by strongylasters or oxyasters. Megasters are represented by oxyspherasters occurring throughout
the sponge, but generally in reduced number, with forked rays and
Description of type species other anomalies, 10-50!-Lm in diameter, RIC = 1-2. Ray tips fre-
quently rounded with ray length twice their basal width. Micrasters
Tectitethya crypta (de Laubenfels) (Fig. lOA-E). are strongylasters, sometimes oxyasters, 8-12!-Lm in diameter.
Synonymy. Cryptotethya crypta de Laubenfels, 1949b: 20; Remarks. The name Tectitethya refers to the species
Tethya crypta Reiswig, 1971: 569; Wiedenmayer, 1977b: 171; described as Tectitethya crypta by de Laubenfels (1949b) and subse-
Tectitethya crypta Sara, 1994: 362; Sara & Bavestrello, 1996: 383. quently included in Tethya by Reiswig (1971). This species has been
Material examined. Holotype: AMNH 473, in two frag- carefully re-described by Wiedenmayer (1977b), but always attrib-
ments - Bimini, Bahamas. Paratype: AMNH 500 - same locality. uted to Tethya. Conversely, Sara (1994) and Sara & Bavestrello
Other material. Tethya crypta: USNM 30278 - Bimini. Pulitzer- (1996) showed clearly by the body shape and skeleton structure that
Finali colI. BW4 - Bimini. SDC 10 - San Domingo. Pansini coIl. T. crypta is not a Tethya but belongs to a separate genus. It cannot be
2206 - San Salvador (Little Bahamas). Columnitis squamata: called Cryptotethya because this name was previously used by
BMNH 1928.1.12.167 - Crawshay collection, West Indies. Dendy (1905) for a species (c. agglutinans) which is not a Tethyidae
Description. According to de Laubenfels (1948) the body and bears no affinities to C. crypta. Striking differences of
is amorphous, often of fist size or in slabs, 4 X 7 X 12cm, but it Tectitethya from Tethya are seen in the branching and anastomosing
may also be conical, hemispherical or stubby cylindrical megasclere bundles, the lack of a cortex, the megasclere type repre-
(Wiedenmayer, 1977b), covered on much of its surface by low, sented by strongyles or anisostrongyles, and the megaster and
irregularly rounded and flattened tubercles, 3-5 mm in diameter, micraster shape. The genus contains four shallow-water species, all
and by abundant sediment. Color of the surface, devoid of sediment, found in the Caribbean (Sara & Bavestrello, 1996).
Porifera· Demospongiae • Hadromerida • Tethyidae 261

c
I 8

Fig. 11. Tethyastra oxyaster. A, holotype. B, skeleton structure. C, strongyloxeas. D, megasters. E, polyrhabds (scales: A, Iem; B, Imm; C, 50 fLm;
D, \00 fLm; E. \0 fLm).

TETHYASTRA GEN.NOV. Synonymy. Tethyorrhaphis oxyaster Burton, 1934a: 568;

Tethya oxyaster Bergquist & Kelly-Borges, 1991: 60.
Synonymy Material examined. Holotype: BMNH 1930.8.13.30
(section slide BMNH 1930.8. 13.30a) - Great Barrier Reef,
Tethyorrhaphis sensu Burton, 1934a: 568. Tethya sensu Australia, 65 m depth, mud bottom.
Bergquist & Kelly-Borges, 1991: 60. Description. Body subspherical, 2 cm in diameter, with
rounded tubercles and an apical oscule. Cortex thin, about
Type species 0.5-1 mm thick, but distinct and with lacunes. Megascleres are
strongyloxeas, 1120 X 24 j..Lm, in distally bent bundles. Megasters
Tethyorrhaphis oxyaster Burton, 1934a: 568 (here designated). are oxyasters, very heterogeneous in size. Choanosomal oxyasters
20-200 j..Lm in diameter, cortical oxyasters, sometimes with a small
Diagnosis centrum, 20-120 j..Lm. Ray number: 8-12. Micrasters are spiny
polyrhabds sometimes with branches (according to the description
Tethyidae with a subspherical body with rounded tubercles and of Burton, spirasters with lateral branches). They measure 5-20 X
distinct but thin cortex. Megascleres are strongyloxeas arranged in 1-3 j..Lm (3-18 j..Lm according to Burton).
radiating bundles. Megasters are represented by oxyasters filling the Remarks. The genus Tethyorrhaphis, to which Burton
whole sponge, very heterogeneous in size, and with giant spicules. (1934a) attributed T. oxyaster, was established by Lendenfeld
Micrasters are spiny microrhabds, sometimes with branches. (1988) for Tethyidae species, such as T. laevis (Lendenfeld) with
polyrhabds in addition to micrasters. Bergquist & Kelly-Borges
Description of type species (1991), on the basis of a new species with polyrhabds (Tethya com-
munis), placed Tethyorrhaphis in synonymy with Tethya, and then
Tethyastra oxyaster (Burton, 1934a) (Fig. 11A-E). also Tethyorrhaphis oxyaster. I agree with this synonymy except for
262 Porifera· Demospongiae • Hadromerida • Tethyidae

**
E

* + F
t:;l .

C _D
~**
* 1(
Fig. 12. Tethycometes sibogae. A, holotype (on the right) and paratype. B, sponge structure. C, basal strongyloxea. D, distal strongyloxea. E, megasters
(oxyasters). F, rnicrasters. G, microstrongyles (scales: A, 5 mm; B, 1 mm; C, D, F, 10 !Lm; E, 100 !Lm; G, 50 !Lm).

the inclusion of T. oxyaster. Indeed, a single trait such as the occur- Type species
rence of polyrhabds in addition to micrasters, may be considered as
species descriptor but not for the establishment of a new genus. But Tethycometes sibogae Sara, 1994 (by original designation).
this is not the case for T. oxyaster. This species differs from other
species of Tethya not only for the polyrhabds, but more importantly, Diagnosis
for the role of the megasters in the skeleton structure accompanied
by a reduced cortex, and the megaster features (shape, size and dis- Tethyidae with an ovoid and very small body, and with a thin
tribution). The megasters are represented by oxyasters, with a wide basal stalk about eight times the length of the body. Without a
range of sizes and reaching giant dimensions, and are distributed distinct cortex. Surface with rounded and hispid tubercles. Body
uniformly and densely throughout the whole sponge. The distribu- skeleton formed by bundles of strongyloxeas, one-spicule long,
tion pattern, as well as the shape and size of the oxyasters, is more which radiate from the body centrum. Fans of diverging thinner
similar to the genus Stellitethya than to other species of Tethya. But strongyloxeas, one spicule-long, originate from the bundles and bris-
T. oxyaster cannot be attributed to Stellitethya given its globose reg- tle the tubercles. Megasters represented by oxyasters, heterogeneous
ular shape, accompanied by a radiating megasclere skeleton and a in size, reaching large dimensions (more than 150 fLm in diameter),
distinct cortex. It is therefore a representative of a new genus, some- and filling the whole sponge body. Two types of rnicrasters:
what intermediate for its reduced cortex and megaster geometry, oxyasters and strongylasters. In addition there are rnicrostrongyles
between Tethya and Stellitethya. The genus contains only one around the base of the main strongyloxeas.
species from the Great Barrier Reef (Burton, 1934a).
Description of type species

TETHYCOMETES SARA, 1994 Tethycometes sibogae Sara, 1998 (Fig. 12A-G).

Synonymy. Tethycometes sibogae Sara, 1998: 363.
Synonymy Material examined. Holotype and one paratype: BMNH
1933.8.12.245 - 'Siboga' Exped. stat. 321, Java sea, Indonesia, 82m
Tethycometes Sarl!, 1994: 363. depth.
Porifera' Demospongiae • Hadromerida • Tethyidae 263

c
00
F

Fig. 13. Tethytimea tylota. A, holotype (indicated by arrows) on a stone with other sponges. B, skeleton structure. C, tylostyles. D, megasters. E, micras-
ters. F spheres (scales: A, 5 mm; B, 1 mm; C-D, F, 50 fLm; E, 10 fLm).

Description. The ovoid body measures 3 X 2 mm, the long Type species
and thin stalk 25 X 0.3 mm. Main strongyloxeas in bundles,
700 X 10 fLm, thinner strongyloxeas in the distal fans, 600 X 5 fLm. Donatia tylota Hentschel, 1912 (by original designation).
The thin microstrongyles, grouped around the base of the main
strongyloxeas, range from 15 X 2 to 160 X 6 fLm, and are straight Diagnosis
or variously bent or twisted. Megasters are oxyasters with or with-
out a small centrum (RIC = 10) and 60-200 fLm in diameter. Tethyidae with thinly encrusting body and indistinct cortex.
Micrasters are oxyasters without centrum with 8-12 slightly spined Megasclere skeleton composed of parallel bundles of tylostyles
slender rays, 10-20 fLm in diameter. On the surface layer are sev- ending in tubercles. Megasters are oxyspherasters, heterogeneous
eral strongylasters, slightly spined and sometimes slightly tylote, in size and with giant spicules. Micrasters are tylasters. Other
also cruciform, 10 fLm in diameter. It was not possible to ascertain microscleres, occasionally present, are spheres.
if there are exotyles in the stalk.
Remarks. Tethycometes is differentiated from other stalked Description of type species
Tethyidae (Burtonitethya, Halicometes) by the length of the stalk in
proportion to the body size, the very small size of the body, its sim- Tethytimea tylota (Hentschel, 1912) (Fig. 13A-F).
ple skeletal structure formed by only two series of non-overlapping Synonymy. Donatia tylota Hentschel, 1912: 41; Tethytimea
strongyloxeas (a basal layer of bundles of larger strongyloxeas tylota de Laubenfels, 1936a: 164; Donatia tylota Sara, 1994: 367.
and a distal layer with fans of smaller strongyloxeas), the hetero- Tethya repens; Hooper & Wiedenmayer, 1994: 425.
geneity in shape of the micrasters, and the peculiar occurrence of Material examined. Holotype: SMF 992-996 (fragment
microstrongyles. The genus contains only one species from MNHN DCL 2208) - Sungi Manumbai, near Dosi, Am Islands,
Indonesia (Sara, 1994). Arafura Sea, Indonesia collected 29.v.1908, 20m depth. Paratype:
SMF (not seen) - Sungi Manumbai (Kapala Sungi), Arafura sea,
Indonesia, 5.v.1908, 20m depth.
TETHYTIMEA DE LAUBENFELS, 1936 Description. Encrusting body of 3 cm diameter, 3 mm thick.
Tubercles about 1-5 mm broad. Indistinct cortex (pseudocortex)
Synonymy 600-700 fLm thick. Tylostyles, with a more-or-1ess developed head,
of two sizes: 1192-1624 X 15-25fLm and 320--640X5-10fLm.
Donatia sensu Hentschel, 1912: 41. Tethytimea de Laubenfels, Megasters are oxyspherasters 40-250 fLm in diameter with about
1936a: 164. Not Tethytimea; Thomas, 1968a; 1980: 14. 14 rays, frequently bent, forked or trunked with spines
264 Porifera· Demospongiae • Hadromerida • Tethyidae

Fig. 14. Xenospongia patellijormis. A, holotype (upper face on the right). B, detail of the upper face. C, sponge structure. D, auxiliary styles. E, main style.
F, megasters. G, micrasters (scales: A, B, 1 em; C, 1 mm; D, 100 fLm; E, F, 50 fLm; G, 10 fLm).

(RIC = 1-1.5). Micrasters are tylasters, 10-13 /-Lm in diameter with to Tethytimea, and in this opinion he was followed by Thomas
6-14 rays. According to Hentschel spheres are abundant in (1968a, 1980). However, the diagnostic traits of T. tylota do not
the holotype but not recorded in the paratype and measure occur in T. repens, which as noted above, is likely to be
44-60 /-Lm in diameter, but I observed only few of these in the a Stellitethya. The genus contains only a single species from the
holotype. One anomalous spicule formed by two joined spheres Aru Islands, Indonesia (Hentschel, 1912).
measures 70 X 40 /-Lm.
Remarks. The holotype in the SMF is encrusting on a stone,
associated with other sponges. From the original description of XENOSPONGIA GRAY, 1858
Hentschel (1912) it clearly refers to the specimen from Station 7,
Sungi Manumbai. The other reported specimen is a fragment col- Synonymy
lected from Station 17 and should be considered a paratype.
Hooper & Wiedenmayer (1994) proposed the type material of this Xenospongia Gray, 1858: 230; Dendy, 1905: 115.
species included only 'Syntypes (2 specimens) SMF 993 wet', and
that Hentschel's species was a synonym of Donatia repens schmidt, Type species
but this is rejected here.
This genus, erected by de Laubenfels (1936a) for Donatia Xenospongia patelliformis Gray, 1858 (by monotypy).
tylota Hentschel 1912, is well characterized by the thinly encrust-
ing body and the megascleres represented by tylostyles in parallel Diagnosis
bundles. Another peculiarity is the occurrence of spheres. These
traits clearly distinguish Tethytimea from Tethya repens Schmidt, Tethyidae with a discoid body. Cortex is thin, poorly devel-
1870. De Laubenfels (1936a) suggested to attribute Tethya repens oped. Surface with relatively small rounded tubercles and oscules
Porifera' Demospongiae • Hadromerida • Tethyidae 265

on slight prominences in the central part of the disk. Much sedi- pores in two peripheral grooves. Texture firm, compact, leathery.
ment occurs in the deeper part of the choanosome. Megascleres are Color (in ethanol) yellowish-gray with a pinkish fringe. Cortex thin
represented by styles in parallel bundles ending in the tubercles, and represented essentially by a dense layer of asters (pseudocor-
crossed by thin bundles of styles in the lower part of the tex) about 130 f.Lm thick. Megascleres are flexuous styles
choanosome at the contact with the sand layer. Megasters are 1700-3500 X 12/-Lm and more. Megasters are represented by
oxyasters, heterogeneous in size and shape, reaching large dimen- oxyasters with small centrum and heterogeneous in size, with
sions (also 180 f.Lm in diameter): Micrasters are spiny oxyasters, diameters between 50-180 f.Lm with about 11 rays frequently irreg-
strongylasters, chi asters and tylasters. ularly curved and more-or-Iess branched. The larger and more
branched spicules are characteristic of the sandy layer. Micrasters
Description of type species are strongylasters (chiasters-tylasters), generally 8 f.Lm in diameter
with about 8 stout rays and sometimes a center, and oxyasters-
Xenospongia patelliformis Gray, 1858 (Fig. 14A-G). tylasters with small center, 11 rays sometimes branched with
Synonymy. Xenospongia patelliformis Gray, 1858: 230; oxeote or tylote tips. Sand grains in the lower two-thirds of the
Dendy, 1905: 115. disk, surrounding the bases of styles and asters.
Material examined. Holotype: BMNH 1883.1.25.9 - Torres Remarks. This genus belongs to the shallow-water clade of
Straits, N Australia. Tethyidae with massive or encrusting shape (Sara & Burlando,
Description. Body discoid, slightly concave above and 1994). The main peculiarities are the flattened discoid shape and
convex below, 3-12 cm or more in diameter and 6 mm thick. The the inner layer of sediment that surrounds the megasclere bases.
holotype is 12 cm in diameter. Surface minutely tuberculate or The genus contains only one species, found in the Indo-Pacific,
conulose. Oscules multilocular placed in the center of the disc, from Torres Straits to Sri Lanka, depth 20-35 m (Dendy, 1905).
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Timeidae Topsent, 1928

Klaus Riitzler

Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, DC, 20560-0163, USA.
(ruetzler.kiaus@nmnh.si.edu)

Timeidae Topsent (Demospongiae, Hadromerida) now contains only Timea because Diplastrella was moved to the Spirastrellidae. These
sponges are thin, colorful (mostly red) crusts with choanosomal tylostyles arranged in vague tracts, ectosomal tylostyles in bouquet
arrangement. Euasters occur throughout the body and form a dense layer at the surface. Species are cryptic and are found in shaded
habitats of warm-temperate to tropical shallow-water environments.
Keywords: Porifera; Demospongiae; Hadromerida; Timeidae; Timea.

DEFINITION, DIAGNOSIS, SCOPE of Diplastrella to the Spirastrellidae (see Rtitzler, this volume),
only one genus, Timea, remains in the family.
Synonymy

Timeidae Topsent, 1928c: 145. Choanitidae de Laubenfels TIMEA GRAY, 1867

(subfamily Timeinae Topsent), de Laubenfels (1936a, 1950a).
Spirastrellidae, in part, Wiedenmayer (1977b). Synonymy

Definition Hymedesmia, in part, Bowerbank, 1866: 149; 1874b: 71.

Timea Gray, 1867a: 544. Kotimea de Laubenfels, 1936a: 147.
Crustose Hadromerida with tylostyles and euasters. Halicometes sensu de Laubenfels, 1950a: 99. Not Halicometes
Topsent, 1898: 112. Timeopsis Levi, 1958: 22.
Diagnosis
Type species
Sponges form very thin crusts on shells or under rock.
Tylostyles are arranged in tracts ending as bouquets at the sponge Hymedesmia stellata Bowerbank, 1866 (by monotypy).
surface where they cause a hispid appearance. Additional tylostyles
occur in criss-cross fashion between the tracts. Microscleres are Definition
euasters, which are dispersed throughout the tissue, denser at the
substrate and toward the surface, and are forming an ectosomal crust. Same as family.

Scope Diagnosis

Of four nominal genera only one, Timea, is recognized. Same as family.

History and biology

All species known in this family are encrusting (1 mm thick,

or less), and therefore have a similar body plan as many spirastrel-
lids and clionaids. Tracts of tylostyles rise from the substrate
toward and beyond the surface (ectosome), causing a velvety feel
when touched. Microscleres form a protective crust at the surface.
The major exhalant canals leading toward the oscula bulge up and
give a radiating vein-like appearance. Color reported for Timeidae
is generally red or brown. The sponges cover only a few square
centimeters of substratum and are best found by examining the
lower surfaces of rock or coral plates in shallow water. Nothing is
known about reproduction or other aspects of biology.

Remarks
Fig. 1. Timea stellata (Bowerbank). Illustration original prepared by the
Recent authors treated Diplastrella as part of the Timeidae author of the holotype and kept with the specimen in the collection of
(Hartman, 1982; Hooper & Wiedenmayer, 1994). With the transfer BMNH. It was the base of Bowerbank's (1874b) plate XXVIII figs 6-8.

266
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Tracbycladidae Hallmann, 1917

John N.A. Hooper l & Rob W.M. Van Soest2

Trachycladidae Hallmann (Demospongiae, Hadromerida), contains two genera with at least nine species, all having an axial (or basally)
compressed skeleton, and an extra-axial plumo-reticulate skeleton, with megascleres predominantly oxeas, less often strongyles and/or
(tylo-)styles, in spongin-enforced multispicular tracts, or in the case of encrusting species, megascleres erect on the substrate (in
hymedesmioid arrangement). Ectosomal skeleton is a crust of spinispirae (spined vermiform spiraster-like spicules), with smooth
microrhabds and/or spinispirae dispersed throughout the mesohyl. The two genera are differentiated on the basis of fundamental differ-
ences in spinispirae morphology. Species are known from the shallow subtidal to about 220 m depth, distributed from temperate Australia,
New Zealand and New Caledonia, Indonesia, East Africa and South Africa, Mediterranean and the western North Atlantic.
Keywords: Porifera; Demospongiae; Hadromerida; Trachycladidae; Rhaphidhistia; Trachycladus.

DEFINITION, DIAGNOSIS, SCOPE convincingly) that some Spirastrellidae (e.g., S. dilatata

Kirkpatrick) have megascleres united into spongin-enforced tracts
Synonymy reminiscent of Trachycladus. Conversely, he noted that
Trachycladus had skeletal structure reminiscent of sigma-bearing
Trachycladidae Hallmann, 1917c: 673. [Spirophorellinae] poecilosclerids (which we now know as Desmacellidae), and thus
Lendenfeld, 1889b (nomen nudum). Rhaphidistiinae de supposedly represented a link between Spirastrellidae and
Laubenfels, 1936a. Poecilosclerida. Brlilndsted (1924b) also stressed the close relation-
ship between Trachycladus and Spirastrellidae and Latrunculia,
Definition although the latter taxon is no longer widely included in this
relationship (see chapters on Latrunculiidae and Podospongiidae,
Hadromerida with spined vermiform spinispirae and smooth this volume). Similarly, Trachycladidae was included in the
microrhabds, with a differentiated axial and extra-axial skeleton polyphyletic order Axinellida by Bergquist (1970, 1978), Levi
cored by oxeas, strongyles and/or (tylo-)styles. (1973), Hartman (1982) and others, as opposed to an allocation
in Hadromerida, implicitly based on possession of an axially
Diagnosis compressed and extra-axially plumo-reticulate skeletal structure,
spongin-enforced spicule tracts, bright colouration, arborescent
Encrusting, massive or branching growth forms. Oscules are growth form (Levi, 1973). Unfortunately there is no corroboratory
small (less than 1 mm diameter) and ostia are scattered singly or evidence to support a close relationship between typical axinellids
grouped. Skeleton is condensed in the axial region and plumo- (e.g. Axinella) and Trachycladus (e.g., oviparous mode ofreproduc-
reticulate in the extra-axial region, with ascending multispicular tracts tion, parenchymella larvae), and in the absence of this evidence
joined at infrequent intervals by single spicules. Encrusting species Trachycladidae is included in Hadromerida. Conceivably, however,
have a hymedesmioid architecture. Skeletal tracts are composed it could justifiably be merged in Spirastrellidae given the fundamen-
of spongin fibres enclosing oxeas, strongy1es and/or (tylo-)styles. tal similarities in their microscleres, although with equally funda-
Microscleres are smooth microstrongyles (microrhabds) and/or mental differences in patterns of spination, hypothesised to reflect
spined vermiform spiraster-like spiru1es (spinispirae), with either a ontogenetic differences in their respective development.
curled shaft bearing longitudinal rows of spines along its length, or
a straight shaft bearing concentric spiral rows of spines around its Scope and biology
circumference and length.
Four nominal genera have been included, or potentially
Remarks included in the family, of which two are considered here to be
valid, Trachycladus and Rhaphidhistia. There are 17 nominal
Trachycladus was allocated to Axinellidae as 'Axinellidae species or subspecies referred to the family at one time or another,
with microscleres', but nevertheless recognised as anomalous of which only nine are probably valid: seven allocated to
(Hallmann, 1916a: 454), and later (in postscript) allocated to a Trachycladus, and two to Rhaphidhistia. These are: (I) T. laevispir-
monogeneric subfamily Trachycladinae (Hallmann, 1917c: 673). ulifer Carter (the type species, from E, SE, Sand SW Australia,
Topsent (l928c) overlooked or ignored Hallmann's new subfamily including as synonyms: digitata Lendenfeld, 1888; scabrosus
in his revision of Demospongiae, and retained the genus in Hallmann, 1916a;fastigatus Hallmann, 1916a; gracilis Hallmann,
Axinellidae. Hallmann (1916a: 454) remarked on the obvious 1916a; clavatus Hallmann, 1916a, reteporosus Hallmann,
derivation of spinispirae from spirasters of Spirastrella, the fact 1916a; and pustulosus Hallmann, 1916a); (2) T. strongylatus
that microrhabds occur commonly in hadromerids, and (less Hallmann, 1916a (as T. digitatus strongylatus, from Victoria)

268
Porifera· Demospongiae • Hadromerida • Timeidae 267

Fig. 2. Timea stellata (Bowerbank). SEM images of spicules. A, tylostyles of two size classes, and euaster (scale 20 /-Lm). B, small euasters (scale 5 /-Lm).
C. large euasters (scale 5/-Lm).

Previous review tapering to a sharp point. Many are broken in the slide preparation
but there seem to be two size classes, 90-120 X 2.5-3.5 fLm and
Hooper & Wiedenmayer (1994). 190-230 X 2.5-4.5 fLm. The euasters have a small center and 6-8
(mostly 7) rays. The rays are either smooth or microspined along the
Description of type species shaft but are always spined near the distal end. Euasters occur in two
size categories which measure 8-10 fLm and 14-22 fLm in diameter.
Timea stellata (Bowerbank) (Figs 1-2). The species was recorded from the North Atlantic, the West
Synonymy. Hymedesmia stellata Bowerbank, 1866: 150; Indies, and the Southwestern Pacific off East Australia but the
1874b: 71, pI. xxvrn figs 5-8. Timea stellata; Gray, 1867a: 544. conspecifity of the latter populations is doubtful (Hooper &
Material examined. Holotype: BMNH 10.1.1.97 - Guernsey, Wiedenmayer. 1994).
Channel Islands, English Channel. Remarks. Kotimea was established by de Laubenfels
Description. The dry specimen is grayish drab and covers (1936a) (with type species Hymedesmia moo rei Carter, 1880b: 50,
the interior surface of a bivalve shell. The thin (> 1 mm) crust pI. 4 fig. 11, by original designation), for Timeidae with styles
spreads over less than 2.5 cm2• Tylostyles run in strands parallel to in addition to tylostyles. However, the type specimen is lost and
the substrate and are without orientation between the tracts; toward neither Carter's description nor his figure indicate the presence
the surface they fan out and protrude from the sponge in bouquet of styles. Timeopsis was established by Levi (1958) for T. intermedia,
fashion. Euasters occur throughout the body but form a dense crust differing from 'typical' Timea only in having a very compact skele-
in the ectosome. The tylostyles are straight or slightly curved, with tal structure composed of large masses of megascleres with distinct
inconspicuous subterminal head and the opposite end gradually subtylote constrictions.
Porifera· Demospongiae • Hadromerida • Tracbycladidae 269

including T. digitatus sensu Levi & Levi, 1983b (from New Trachycladus laevispirulijer from southern Australia was dis-
Caledonia); (3) T. stylijera Dendy, 1924 (from northern and south- covered to have organohalogen compounds (trachycladine) with
ern New Zealand, Bergquist, 1970); (4) T. cervicornis Burton, highly cytotoxic activity against the P388 leukemia cell line
1959a (from east Africa); (5) T. tethyoides Burton, 1959a (from (Searle & Molinksy, 1995; Gribble, 1998), but few other biological
east Africa and Indonesia, Van Soest, unpublished); (6) T. 'spin- or chemical data are yet known for any species of the family.
ispirulijer'sensu Carter, 1879b (as Suberites) from E, SE, Sand
SW Africa (Carter, 1886a; Dendy, 1897; Burton, 1959a; Levi, Reviews
1959); and Indonesia (as Spirastrella dilatata; Kieschnick, 1896;
Thiele, 1900); and T. 'spinispirulijer' sensu Bergquist, 1970, from Topsent (1928c: 37), Bergquist (1970: 22; 1978), Brien et al.
N New Zealand, and Driz (1987) (from SW Africa» - the two (1973), Wiedenmayer (1989: 53), Hartmann (1982), Hooper &
groups of 'spinispirulijer' potentially also not conspecific; Wiedenmayer (1994: 444).
(7) T. minax (Topsent, 1888) (as Hymeraphia) from the
Mediterranean, NW Africa and the coast of France (Topsent, 1900; Distribution
Burton, 1956; Sara, 1958, 1961; Cabioch, 1968b; Pulitzer-
Finali, 1977); (8) Rhaphidhistia spectabilis Carter, 1879b; and Species are known from the shallow subtidal to depths of at
(9) R. mirabilis (Dendy, 1924) from N New Zealand (as least 220m (Hartman, 1982), distributed in southern, western and
Dotonella). Hallmann (1916a, plate 24) mentions another species eastern Australia, New Zealand and New Caledonia, Indonesia,
(T. formosus), but this appears to be a manuscript name and thus East Africa and South Africa, Mediterranean and the western North
nomen nudum. Atlantic.

KEY TO GENERA

(1) Spinispirae with spines forming regular longitudinal rows running the length of the stem, with the spicule itself (not the spines) curled
or contorted into twisted shapes; only one size class of spinispirae ............................................................................... Trachycladus
Main spinispirae with a long and straight shaft, but with spines forming a complete full tum around the circumference of the
shaft, producing concentric spiral rows along its length; a second smaller, curly size class of spinispirae may also be
present ............... ...... .... ................ ............................ .......... ........... ..... ................ ............................ ........................... ...... Rhaphidhistill

TRACHYCIADUS CARTER, 1879 shaft, and the geometry of the shaft ranging from simply curved,
c-shaped, to twisted with up to two complete turns, and also with
Synonymy microrhabds, smooth or occasionally faintly centrotylote.

Trachycladus Carter, 1879b: 343. [Spirophora] Lendenfeld, Remarks

1887c: 794 (preocc.). Spirophorella Lendenfe1d, 1888: 236.
Taxonomic decision for synonymy Hallmann, (1914: 429). Spirophorella Lendenfeld (with type species Spirophora dig i-
tata Lendenfeld, I 887c, by subsequent designation, Wiedenmayer in
Type species Hooper & Wiedenmayer, 1994: 444), was undoubtedly proposed as a
replacement name for [Spirophora] Lendenfeld (a junior homonym
Trachycladus laevispirulifer Carter, 1879b (by monotypy). of Spirophora Milne-Edwards, 1836 (Bryozoa), and [Spirophora]
Zopf, 1884 (Protozoa», although Lendenfe1d (1888) was not
Definition explicit about this.

Trachycladidae with spines on the spinispirae forming regular Reviews

rows that run longitudinally along the stem of the spicule, and the
spicule itself (not the spines) contorted into curled shapes. Smooth The genus has been reviewed by Hallmann (1916a: 453),
or occasionally faintly centrotylote microrhabds are also typically Topsent (1928c: 37), Bergquist (1970: 22), Levi & Levi (1983b:
present. 942) and Wiedenmayer (1989: 53).

Diagnosis Description of type species

Tubulo-digitate, arborescent, whip-like to flabellate, rarely Trachycladus laevispirulijer Carter, 1879b (Fig. 1).
encrusting growth forms. Ectosome with a distinct cortex packed Synonymy. Trachycladus laevispirulijer Carter, 1879b: 343,
with spinispirae. Choanosomal skeleton compressed in the axial pI. 28, figs 1-5. Spirophora bacterium Lendenfeld, 1887c:
region, plumoreticulate in the extra-axial region, with paucispicu- 795. Spirophora digitata Lendenfeld, 1887c: 794. Trachycladus
lar connecting fibres. Fibres fully cored by oxeas, but also includ- digitata; Hallmann, 1916a, pI. 22 fig. 2. Trachycladus digitatus
ing styloid or strongyloid modifications resembling strongyloxeas, clavatus Hallmann, 1916a: 474, fig. 6, pI. 22 fig. 4, pI. 23 fig. 3,
anisoxeas, mucronated anisoxeas, true styles or strongyles, and in pI. 25 fig. 2, pI. 27 fig. 3, pI. 28 fig. 5, pI. 29 fig. 1. Trachycladus
one species tylostyles. Microscleres are 'spinispirae', smooth or digitatus gracilis Hallmann, 1916a: 472. Trachycladus fastigatus
minutely spined, with spines forming longitudinal rows along the Hallmann, 1916a: 462. Trachycladus pustulosus Hallmann,
270 Porifera· Demospongiae • Hadromerida • TrachycIadidae

Fig. 1. Trachycladus laevispirulifer Carter, 1879b. A, habit of Trachycladus digitatus Hallmann, 1916a Gun.syn. ofthe type species) (reproduced from his
plate XXII fig. 2; (scale 1 cm). B, SEM overview of spicules of holotype of T. laevispirulifer, BMNH unregistered (scale 100 j.Lm). C, details of stylote
megasclere of the same (scale 1 j.Lm). D, spinispirae (scale 1 j.Lm). E, microrhabd (scale 1 j.Lm).

1916a: 486, fig. 9, pI. 21 fig. 5, pI. 26 figs 5, 8, pI. 27 fig. 6, pI. 39 Additional material. QM collections and Fromont (1999b: 177).
figs 6, 7, pI. 39 figs 6, 7. Trachycladus reteporosus Hallmann, Comparative material. Holotype of T. digitatus strongylatus: AM
1916a: 479, pI. 21 figs 2,3, pI. 23 figs 5-8, pI. 24 figs 1-3, pI. 25 Z1760 - Port Phillip Bay, Victoria. Holotype of T. stylifer: BMNH
fig. 1, pI. 26 figs 1, 4, 7, pI. 27 fig. 5, pI. 28 figs 1-4, pI. 29 fig. 2. 1923.10.1.158 - Three Kings Is., New Zealand. Syntypes of
Trachycladus scabrosus Hallmann, 1916a: 459. Not Trachycladus T. spinispirulifera (Carter, 1879b): BMNH 1871.5.12.1, BMNH
digitatus strongylatus Hallmann, 1916a: 477. Not Trachycladus 1954.3.9.443, 2MB 3039 - Port Elizabeth, South Africa.
stylifer Dendy, 1924: 377. Refer to Wiedenmayer (1989: 53) for Description. Highly polymorphic growth form, ranging
more complete synonymy citations. from tubulo-digitate, shrub-like to simple arborescent, di- or poly-
Material examined. Lectotype: BMNH unregistered dry. chotomous branching sponge, rarely unbranched whip-like or fla-
Paralectotype: BMNH 1883.1.25.10 - South Australia. Other type bellate. Branches variable in length between specimens, ranging
material. Holotype of S. bacterium: BMNH 1887.4.27.31 (slide from: relatively short, club-like, thickened more towards their apex
AM G3476) - Westernport Bay, Victoria, Australia. Lectotype of S. than at their point of attachment to the stalk, with tapering rounded
digitata: AM Z488 - Port Jackson, NSW. Paralectotype AM Z1333 - or club-shaped terminations, often bifurcate at the extremities; to
Encounter Bay, South Australia. Holotype of T. digitatus clavatus: long thin tendrils tapering to fine points. Branches fleshy in life.
AM Z1270 - Port Phillip Bay, Victoria, Australia. Holotype of Stalk relatively firm and thin compared to branch diameter.
T. digitatus gracilis: AM Z963 - Port Jackson, NSW. Holotype Oscules small, up to 3 mm diameter (living material), or < 1 mm
of T. Jastigatus: AM E3710 - Great Australian Bight. Holotype of diameter when preserved (if visible at all), scattered either on lat-
T. pustulosus: AM Z1761 - Port Phillip Bay, Victoria. Holotype of eral (less often terminal) parts of branches, in vague longitudinal
T. reteporosus: AM Z1620 - Port Phillip Bay and Heads, Victoria. rows, or clustered into poriferous areas on the distal parts of
Holotype of T. scabrosus: AM Z1501 - off Port Jackson, NSW. branches. Texture compressible, rubbery in life, firm and harsh
Porifera· Demospongiae • Hadromerida • Tracbycladidae 271

Fig. 2. A-B, Trachycladus styiifer Dendy, 1924. A, SEM overview of microsc1eres of ZMA l3000, from 695-798 m off New Zealand (scale 10 f.Lm).
B, detail of microrhabd of the same (scale 1 f.Lm). C-E, Trachycladus spinispiruiifer (Carter, 1880 as Suberites), SEM images of the holotype BMNH
1871.5.12.1. C, detail of tylostyle (scale 10 f.Lm). D, spinispira (scale I f.Lm). E, overview of spicules (scale 100 f.Lm).

when preserved. Surface macroscopically smooth and even in life between fibres. Mesohyl heavily collagenous, with scattered spin-
but microscopically conulose due to ascending fibres terminating ispirae and microrhabds, the latter often clustered, and sarcode per-
at the surface. When preserved the surface is porous and highly meated with large aquiferous canals near the surface, becoming
conulose, with prominent sharp conules ('scopuliform process') increasingly smaller and converging as they approach the axial
corresponding to the lines of reticulate fibres below the surface, but region. Megascleres predominantly oxeas, symmetrically curved at
this is certainly not a feature of the living sponge. Subdermal centre with varying terminations ranging from slightly fusiform,
canals clearly visible below surface when detached. Colour red, gradually tapering to abruptly pointed with telescoped points;
red-orange, vermilion to pale pinkish alive, beige in ethanol with occasionally styloid resembling strongyloxeas, anisoxeas,
a distinctly white surface membrane that largely disintegrates after mucronated anisoxeas, true styles or strongyles, 230--315 X 5-7
preservation. Ectosome with a distinct and detachable cortical /-Lm. Microscleres 'spinispirae', resembling spirulae, toxaspires
region packed with spinispirae. Choanosomal skeleton axially and sigmaspires, smooth or minutely spined, with spines forming
compressed, with axis occupying at least half of branch diameter, regular rows that run longitudinally along the stem; the spicule
composed of longitudinally directed multi spicular tracts intercon- itself (not the spines) are curly, ranging from simply curved, c-
nected by short, oblique lateral tracts together forming a dense shaped, to contorted and forming up to two complete turns of the
reticulation, becoming more plumose towards the periphery of the shaft, 6-15 X 1-2 /-Lm; microrhabds smooth, occasionally faintly
axis. Ascending fibres arising from the central axis are radial, centrotylote, 6-16 X 1-4/-Lm.
plumose, becoming increasingly thinner towards the surface, with Remarks. Wiedenmayer (1989) notes that there are so many
primary multispicular ascending fibres and paucispicular lateral descriptions of the type species (and its synonyms) that it comprises
connecting fibres, the former extending to just below the cortex. one of the most prolifically and thoroughly described and illus-
Fibres fully cored by tracts of mostly oxeas, but also including trated of sponges in the Australian fauna (refer to literature citations
styloid or strongyloid spicules, with many also scattered irregularly in Wiedenmayer, 1989: 53-54). It has a confirmed distribution from
272 Porifera· Demospongiae • Hadromerida • TracbycIadidae

coastal waters of southern Queensland (J.N.A. Hooper, unpublished Description of Rhaphidhistia spectabilis (partly from Carter,
collections), NSW, Victoria, South Australian and southern Western 1879b and holotype slide; Fig. 3A-D). Lamelliform, extremely
Australia (to the Houtman Abrolhos) (refer to records cited by thinly encrusting, size up to 2.5 cm in lateral expansion, whitish
Wiedenmayer, 1989). Other records included in synonymy with the yellow in dry condition, texture delicate. Surface smooth, even,
type species by Wiedenmayer (1989) are excluded here and now with oscules raised on monticular processes. Choanosomal skele-
considered valid species: T. stylifer from New Zealand (Dendy, ton described as consisting of a delicate sarcode with 'loosely'
1924: 377; Bergquist, 1970: 21), and T. 'digitatus'sensu Levi & Levi arranged oxeas and large lacunae (Carter, 1879b). In fact the skele-
from deeper waters off New Caledonia (Levi & Levi, 1983b: 942). tal preparation consists of a tissue tease which contains plumose
Suberites spinispirulijer sensu Carter, 1879b from Africa has been bundles of oxeas packed with spinispirae, and it is possible that
traditionally assigned to Spirastrella but has curly 'spirasters' that these bundles may have been erect on the surface of the living
closely resemble those of Trachycladidae, in addition to having sponge. This feature is not possible to verify based on existing
tylostyles for megascleres and encrusting growth form, substantially museum material. Oxeas large, slightly curved, slightly asymmetri-
expanding the definition of the genus. cal, abruptly pointed or slightly tapering, occasionally with
rounded blunt tips, 440-648 X 11-22 /-Lm. Microscleres are long,
straight or irregularly curved spinispirae, up to 28-75 X 4-9 /-Lm,
RHAPHIDHISTIA CARTER, 1879 with up to 10 spiral twists of spines running around the circumfer-
ence of the spicule forming concentric spiral rows along its
Synonymy length. Spinispirae are not apparently restricted to any particular
region of skeleton but are heavily clustered throughout the entire
Rhaphidhistia Carter, 1879b: 300. Dotonella Dendy, 1924: 379. skeleton.
Description of Dotonella mirabilis Dendy (partly from
Type species Dendy, 1924 and holotype; Fig. 3E-l). Thinly encrusting
sponge, about 22 X 16mm in dimensions, cream-white in pre-
Rhaphidhistia spectabilis Carter, 1879b (by monotypy). served condition, with a strongly hispid surface (although now
abraded due to transportation and handling). Basal skeleton, in
Definition contact with substratum, has a dense layer oflarger spinispirae, and
likewise the ectosomal skeleton has a dense layer of smaller spin-
Trachycladidae with long straight spinispirae, with a straight ispirae (slides of the holotype suggest that the ectosome is a remov-
shaft and spines forming spiral rows along the length of the shaft, able cortex), and with a hymedesmioid skeleton of large tylostyles
each forming a complete full turn around the circumference of the embedded in the basal layer and points protruding a long way
shaft. A second category of smaller, curly spinispirae may also be through the ectosomallayer and beyond the surface, producing the
present, forming an ectosomal cortex. hispid appearance. Megascleres are tylostyles with well-developed
basal tyle, a somewhat curved shaft, and gradually tapering to
Diagnosis sharp or abrupt points, 800-1000 X 14-24 f..Lm (although the
exact length is difficult to measure since most spicules are broken
Thinly or thickly encrusting growth forms, with 1-2 size cate- in the slide preparations), with rounded heads 20-24 f..Lm diameter.
gories of spinispirae, and if both are present, the smaller forming Microscleres consist of two distinct size categories of spin-
an ectosomal cortex. Choanosomal skeletal architecture ranges ispirae, both with tight spiral concentric whorls of spines over the
from loosely dispersed megascleres in vaguely plumose arrange- spicule; the larger basal ones have a long, thick and straight shaft,
ment to hymedesmioid architecture, consisting of erect megascle- 90-135 X 10-19 f..Lm, with concentric spiral spination; and the
res embedded in the substratum and protruding through the smaller ones are tightly curled, varying from c-shapes to up to two
surface. Megascleres are oxeas or tylostyles. Microscleres are spin- complete twists, or sometimes recurved upon itself forming commas,
ispirae in one or two size categories, the larger with a long straight and resembling those of the type of T. laevispirulifer, 9-14 X 4-7 f..Lm,
shaft and bearing concentric spiral rows of spines forming a com- and are densely covered by tiny spines.
plete tum around the circumference of the shaft, producing a Remarks. Rhaphidhistia, including Dotonella Dendy (pre-
smooth inner curve, and the smaller with similar spination but with viously referred to Spirastrellidae), stands apart from Trachycladus
the shaft curled into C-, s- or contorted-shapes. in having long straight spinispirae (as opposed to curly ones), and
also with quite different patterns of spination on spinispirae. In
Remarks Rhaphidhistia spinispirae are even more distinct from typical spi-
rasters of Spirastrellidae than are those of Trachycladus, having a
See remarks for type species below. straight shaft with spines forming concentric spiral rows, with each
row forming a complete full turn around the circumference of the
Description of type species shaft, and producing a smooth inner curve (see Fig. 3). By compar-
ison, spines on the spinispirae of Trachycladus (see Figs 1-2) form
Rhaphidhistia spectabilis Carter (Fig. 3). regular rows that run longitudinally along the stem of the spicule,
Synonymy. Rhaphidhistia spectabilis Carter, 1879b: and the spicule itself (not the spines) forming a contorted curl or
300-301, pI. 26, figs 10, 13, 14 (by monotypy). twist. We consider these differences may represent fundamental
Material examined. Holotype: BMNH 1877.5.21.1877a ontogenetic processes in the formation of microscleres and hence
(slide) - Mauritius. Other material. Holotype of Dotonella mirabilis justify differentiating these two genera.
Dendy, 1924: BMNH 1923.10.1.159 (fragment and 3 slides) - Three Dotonella (type species Dotonella mirabilis Dendy, 1924: 379,
Kings Island, New Zealand, 'Terra Nova' Expedition, 180 m depth. pI. 15, figs 43-45; by monotypy), differs from Rhaphidhistia s.s. in
Porifera· Demospongiae • Hadromerida • Trachycladidae 273

Fig. 3. A-D, Rhaphidhistia spectabilis Carter, 1879b. A, oxeas (scale 100 j.l.m). B, large spinispirae (scale 25 j.l.m). C,ectosomal skeleton (scale 200 j.l.m).
D, drawing of encrusting specimen (from Carter, 1879b). E-I, Rhaphidhistia mirabilis (Dendy, 1924 as Dotonella). E, ends of tylostyles (scale 50 j.l.m).
F, larger spinispirae (scale 50 j.l.m). G, smaller spinispirae (scale 5 j.l.m). H, large microrhabd surrounded by spinispirae (scale 10 j.l.m). I, spinispirae
(two types or one developing stage), enlarged (scale 10 j.l.m). H-I, courtesy of Klaus Rtitzler.

having a differentiated deeper layer of large straight spinispirae and similar to the hymedesmioid arrangement seen in Dotonella.
a superficial layer of much smaller curly spinispirae, and with Dendy (1924) compared Dotonella to Trachycladus and Dotona
megascleres consisting of tylostyles (as opposed to oxeas). There (an excavating Clionaidae), suggesting that only the small spinispi-
are also reported differences in their respective choanosomal skele- rae of D. mirabilis resembled those of Trachycladus, and the larger
tal structures (hymedesmioid versus 'loosely arranged'), but these ones were more coarsely spined than corresponding spicules in
might plausibly be attributed to relative differences in the thick- Dotona, implying a close relationship between Trachycladidae,
nesses of their respective growth forms. Dendy (1924) also Clionaidae and Spirastrellidae - but perhaps also inferring it was
remarked on the similarities between its microscleres with those of one related to their respective growth formslhabits. We reject this
Rhaphidhistia spectabilis Carter and Spiroxya heteroclita Topsent, assumption, instead regarding that the characteristics of spinispirae
1900: 280, although he discounted any inferred relationship are more informative to this hypothesized phylogeny than are mod-
between these groups based on the fact that both the latter have ifications to megascleres (with Rhaphidhistia having either oxeas
oxeas for megascleres. However, it is not certain that the skeleton or tylostyles, and Trachycladus having megascleres that range from
of Rhaphidhistia spectabilis is 'loosely arranged', as quoted by oxeas, strongyloxeas, anisoxeas, mucronated anisoxeas, true styles,
Carter (1879b), where slide preparations reveal distinct plumose strongyles or tylostyles). Dendy (1924) included Dotonella in
bundles of oxeas, which are obviously flattened in slide prepara- the family Clavulidae, subfamily Spirastrellinae, and considered
tions, but potentially these bundles might have formed erect its type species an equivalent to Dotona pulchella Carter in the
hymedesmioid or microcionid architectures in the living sponge, Clionaidae. It is feasible that Dotonella and Rhaphidhistia could be
274 Porifera· Demospongiae • Hadromerida • TracbycIadidae

assigned to Spirastrellidae (as redefined by Rtitzler. this volume). of relevant type material. He also notes that Hinde & Holmes
but this hypothesis is also rejected given the profound similarities (1892: pl.7 fig. 38) illustrated a spinispira resembling. but much
in microsclere geometries to those of Trachycladus. larger than that of Dotonella mirabilis. from the Tertiary deposits of
Although de Laubenfels (1936a) subsequently referred Oamaru, South Island, New Zealand. It is also theoretically feasible
Spirastrella aculeata Topsent (1890b: 69; 1892: 127) to Rhaphid- that Spirastrella-like encrusting species which all have tylostyles
histia. that species has strongyloxeas (or tomotes) and spirasters should be differentiated from Trachycladus proper (with oxeasl
typical of spirastrellids (and quite unlike those of the two strongyles/styles). except for the existence of species like Dotonella
Trachycladidae genera - clearly indicating that it should remain in mirabilis with both tylostyles and Rhaphidhistia-like long straight
Spirastrella). Dendy (1924) suggests that Spiroxya heteroclita spinispirae microscleres. Thus. there are potentially more species
Topsent. 1900 also belongs in Rhaphidhistia, having two kinds of of thinly encrusting 'spirastrellids' that may belong to this genus.
spinispirae. although quite different from those of the New Zealand but no decision is possible without careful re-examination of
sponge. but this assertion has not yet been tested from examination respective type material.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcaderniclPlenurn Publishers, New York, 2002

Family Acanthochaetetidae Fischer, 1970

Klaus Riitzlerl & Jean Vacelet2

IDepartment ofInvertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, DC, 20560-0163, USA.
(ruetzler.klaus@nmnh.si.edu)
2Centre d'Oceanologie de Marseille, Universite de la Mediterranee, Station Marine d'Endoume, 13007 Marseille, France.
(jvacelet@com.univ-mrsJr)

Acanthochaetetidae Fischer (Demospongiae, Hadromerida), is a largely extinct family of 'sclerosponges' represented by two extant
species, Acanthochaetetes wellsi and Willardia caicosensis, our knowledge of which indicates its relationship to the Porifera rather than
Cnidaria. The siliceous spicules of these sponges indicate close ties to the genera Spirastrella and Diplastrella but morphological peculi-
arities of spicules and the presence of an elaborately structured calcareous basal skeleton justifies the distinction of Acanthochaetetidae
from Spirastrellidae.
Keywords: Porifera; Demospongiae; Hadromerida; Acanthochaetetidae; Spirastrellidae; Acanthochaetetes; Willardia.

DEFINITION, DIAGNOSIS, SCOPE found by research submersible on vertical rock walls (100-119m)
near the Turks and Caicos Islands (tropical western Atlantic Ocean)
Synonymy (Willenz & Pomponi, 1996). The calcareous base in both species is
covered by a thin layer of soft tissue that is attached by a basopina-
Acanthochaetetidae Fischer, 1970: 199. Tabulospongiidae coderm and includes small, rounded choanocyte chambers,
Mori, 1976: 8. choanocytes with periflagellar sleeves, collagen strands, and cells
with various inclusions (Hartman & Goreau, 1975; Vacelet, 1981,
Definition 1990; Boury-Esnault et al., 1990; Reitner, 1991; Willenz &
Pomponi, 1996). Acanthochaetetes wellsi has two important
Encrusting Hadromerida with calcareous basal skeleton and features of the living tissue that are absent in W caicosensis:
siliceous spicules (tylostyles, streptasters) reinforcing the soft-tissue. (1) fascicles of collagen fibrils that extend through canaliculi
in the limestone base skeleton (Fig. 1) and anchor the soft tissue,
Diagnosis including masses of pseudogemmules; and (2) presence of these
pseudogemmules, apparently dormant bodies consisting of
'Sclerosponges' with a massive calcareous skeleton by which clusters of thesocyte-like cells, enclosed in the basal part of some
the animals are attached to the substratum. Living tissue coating pseudocalices (crypt tissue) (Vacelet, 1990).
the outermost layer of the basal skeleton and containing siliceous
spiculation of tylostyles as megascleres, points outward, and Taxonomic remarks
common, relatively large streptasters.
The genus Acanthochaetetes Fischer, based on the fossil species
Scope A. seunesi Fischer, is represented by the 'living fossil' species A. wellsi
Hartman & Goreau which shows strong affinities to ilie Spirastrellidae
Only two extant genera, Acanthochaetetes and Willardia. (Van Soest, 1984a; Wood, 1990; Chombard, 1997). It has been pro-
posed to assign A. wellsi to this family, even as a subgenus to the
History and biology genus Spirastrella (Reitner, 1991: 195), but scrutiny of morphological
differences supports the validity of Acanthochaetetidae. The genus
The first live specimens of Recent Acanthochaetetidae were Willardia, with its only species W caicosensis Willenz & Pomponi,
discovered by divers in submarine caves (i-30m depth) on reefs of was placed by its original authors in the family Timeidae (Willenz &
Guam in 1968, belonging to Acanthochaetetes wellsi. Soon there- Pomponi, 1996). However, comparing spicule shape and the parallel
after it was shown that the species is common in similar habitats morphology evident in Spirastrella and Diplastrella we decided to
throughout the western Pacific Ocean (Hartman & Goreau, 1975). place both genera in the Acanthochaetetidae (see also discussion in
The second genus, represented by Willardia caicosensis, was the chapter Spirastrellidae).

KEY TO GENERA

(1) Basal skeleton composed of calcite, with vertical tabulate pseudocalicles; siliceous microscleres include
spirasters ............ ...... ..................................................... ................................ ...... .................. ........ ....... .......... ........... Acanthochaetetes
Basal skeleton composed of aragonite, with perpendicular pillar-shaped processes; siliceous microscleres are
exclusively diplasters ..................... ................ ........................................... .... ............................ ........... .......... ......................... Willardia

275
276 Porifera· Demospongiae • Hadromerida • Acanthochaetetidae

Fig. 1. Acanthochaetetes wellsi Hartman & Goreau, 1975. A, schematic vertical section of skeleton and living tissue, with a superficial layer of diplasters,
and pseudogemmulae in crypts under horizontal tabulae (from Vacelet, 1990) (abbreviations: Ab, anchoring collagen bundles; Cc, choanocyte chambers;
Ct, crypt tissue; Ht, horizontal tabulae; S, spines; Sk, calcareous skeleton; Sp, spicules). B, SEM photo of the surface and fracture of the skeleton, with
tabulae (arrows) (scale 0.5 mm). C, SEM photo of calic1e, showing diplasters adhering to the skeleton, spines and micropores corresponding to anchoring
collagen bundles (scale 75 fLm). D-F, siliceous spicules (scale 10 fLm). G, diplaster (scale 1 fLm). (B, C, specimen from Philippines; D--G, specimen
USNM 51487, from Palau).

ACANTHOCHAETETES FISCHER, 1970 Definition

Synonymy Spirastrellid-like 'sclerosponge' with basal calcite skeleton of

lamellar microstructure. Calicles (vertical tubes) have shared walls
Acanthochaetetes Fischer, 1970: 199. Tabulospongia Mori, and are beset by vertical rows of spines. Basal part of the calicles
1976: 2 (type species T. horiguchii Mori 1976: 2). closed by horizontal tabulae.

Type species Diagnosis (combining information from fossil and

Recent material)
Acanthochaetetes seunesit Fischer, 1970: 199 (by original
designation). Sponge has a basal calcite skeleton composed of contiguous
Only extant representative. Acanthochaetetes wellsi vertical, tabulate calicles with lamellar microstructure and covered
Hartman & Goreau, 1975. at the lower surface by an epitheca with concentric growth lines.
Porifera· Demospongiae • Hadromerida • Acanthochaetetidae 277

The inner walls of the calicles are ornamented by vertical rows or 1983). Growth rate of A. wellsi is very slow; estimated from carbon
clumps of spines. Growth occurs by intramural budding. Living tis- isotope records it ranges from 50-450 fLmlyr (Bohm et al., 1996;
sue is restricted to a thin surface layer and to the lumen of the out- Reitner & Gautret, 1996).
ermost tabulae and contains a siliceous spiculation of megascleres Remarks. Based on Acanthochaetetes wellsi, the only
in the choanosome and common, relatively large microscleres known living representative of the genus, Hartman & Goreau
condensed in the ectosome. Exhalant canals converging upon (1975) and Hartman (1982) suggested to place the 'sclerosponge'
the slightly elevated oscula create a star-like pattern that is also family Acantho-chaetetidae in a separate order, Tabulospongida.
impressed into the surface of the calcitic calicles and is termed We disagree with this suggestion as it overemphasises the presence
astrorhizae. Spicules consist of tylostylote megascleres, points of a calcareous skeleton and prefer to position Acanthochaetetes in
directed outward, and spiraster-like microscleres. the Hadromerida.

Previous reviews
WILLARDIA WILLENZ & POMPONI, 1996
Hartman & Goreau (1975); Vacelet (1981, 1990); Hartman
(1982); Reitner & Engeser (1987); Hooper & Wiedenmayer (1994). Synonymy

Description of type species Willardia Willenz & Pomponi, 1996.

Acanthochaetetes seunesit Fischer, 1970 (Fig. 1). This Type species

description is based on the Jurassic type species and supple-
mented by description of the closely related 'living fossil' Willardia caicosensis Willenz & Pomponi, 1996 (by original
Acanthochaetetes wellsi Hartman & Goreau, 1975. designation).
Synonymy. Acanthochaetetes seunesit Fischer, 1970: 199,
201-202, fig. 32, pI. F3-5. Only extant representative: Definition
Acanthochaetetes wellsi Hartman & Goreau, 1975: 2-12,
figs 1-9, 11-14 = Tabulospongia horiguchii Mori, 1976. Spirastrellid sclerosponge with basal aragonite skeleton of
Material examined. Holotype (not seen): MNHN Institut de peniciliate spherulitic microstructure, ornamented by pillar-shaped
Paleontologie R05599 - type material consists of 1 specimen, 2 frag- processes at the surface.
ments, 4 sections; description is partly based on Fischer's (1970)
photographs (pI. F, figs 3-5). Holotype of Acanthochaetetes ramulo- Diagnosis
sust Fischer: USNM 32196. Paratype of A. wellsi Hartman &
Goreau: USNM 24410 - Puntan Madag, Saipan, Marianas Islands, Encrusting sponges secreting a basal calcareous skeleton of
colI. T. F. Goreau, 27 July 1969. Other material. USNM 51487 - aragonite that is attached to the substratum but may have free edges
South Point Cave (wall), Ngermeuangel, Koror, Palau, coIl. D. curling downward. The surface of the calcareous base, where it is
Williams & J. Bozanic, 18 April 1985. Numerous specimens from covered by cellular tissue, is densely ornamented by finger-like
New Caledonia and Philippine from the authors' collection. processes (Willenz & Pomponi, 1996: fig. 5), ca. 1 mm tall. Oscula
Description. The Mesozoic material consists of a frag- are slightly elevated and in the center of converging, vein-like sur-
mented but seemingly spheroid colony composed of radiating face canals. Siliceous megascleres (tylostyles) are arranged in
tubes, oval or circular in cross-section, ranging from 0.8 X 0.8 mm brushes at the sponge surface. Microscleres (diplasters) are con-
to 1.1 X 0.7mm in maximum perpendicular diameters. Recent centrated near the surface and along major aquiferous canals. In
material of Acanthochaetetes wellsi: Massive sclerosponge with places, spicules may become embedded in aragonite.
calcitic skeleton made up of adjoining vertical tubes (calicles) with
common walls. Basal parts of the calicles partitioned by vertical Previous review
tabulae. Walls ornamented by spines that are arranged in vertical
lines or clumped irregularly. Both walls and spines have Willenz & Pomponi, 1996.
a microstructure of stacked lamellae. The surface of the skeleton
shows starlike impressions (astrorhizae) from meandering exhalant Description of type species
canals converging upon single oscula. Epitheca with concentric
growth lines covering lower surface of sponge. Size ranging from Willardia caicosensis Willenz & Pomponi, 1996 (Fig. 2).
under 1 cm to over 18 cm (diameter of live tissue area). Living Synonymy. Willardia caicosensis Willenz & Pomponi,
tissue (as seen on only extant species, A. wellsi) cream-colored, 1996: 208, figs 1-22.
restricted to a coating of the calcareous skeleton, including the Materinl examined. Holotype (not examined): RIB-POR 49-
spaces in the calicles above the outermost tabulae. Siliceous Grand Turk Island, Turks and Caicos Islands, 21 °31'59"N,
spicules (observed in A. wellsi) occur in the living tissue and 71°07'97"W, 114m, collected 1994. Paratypes: BMNH, HBOI,
include erect tylostyles (points toward the surface, 286 X 3.4 fLm, YPM - same locality.
7.4 fLm head diameter) and spiraster-like and amphiaster-like Description (from Willenz & Pomponi, 1996). The sponge
microscleres (5 X 6-20 X 28 fLm) localized in a layer in the outer forms 5 mm thick, plate-like crusts, up to 20 cm in diameter, and is
tissue. Microsclere spines are often branched and closely spaced attached to the substratum by its aragonite basal skeleton; the
thus obscuring the axis. Some microscleres can be seen adhering to plate margin is free and curled downward, toward the substratum.
the calcareous skeleton (Fig. 1) and may thus become incorporated The live tissue is yellow to tan orange and has a velvety appear-
during fossilization, as described for A. seunesi (Reitner & Engeser, ance from perpendicular spicule brushes embedded in the surface.
278 Porifera· Demospongiae • Hadromerida • Acanthochaetetidae

Fig. 2. Willardia caicosensis Willenz & Pomponi, 1996. A, SEM photo of the surface of calcareous skeleton with pillar shaped processes (scale 500 ILm).
B, section perpendicular to the surface (abbreviations: Ac, aquiferous canal; Ar, aragonite skeleton; Ct, collagen tracts; S, surface; Sc, spherulous cells
accumulated at the surface (scale 200 ILm). C, tylostyles and amphiasters (scale 200 ILm). D, diplasters (scale 20 ILm). (From Willenz & Pomponi, 1996.)

Oscula are slightly raised, 3-4 mm in diameter, supplied by con- ca. 20 cells per chamber), spherical, and eurypylous. The
verging, anastomosed canals that form a star-like pattern. choanocytes have a periflagellar sleeve surrounding the base of the
Siliceous spicules consist of tylostyles arranged in bouquet fash- long flagellum, typical of hadromerids. There is also a central cell
ion at the surface, points protruding out beyond the ectosome, and controlling flow through the apopyle. The mesohyl is rich in colla-
diplasters, also in the surface and lining the canals. A few spicules gen fibrils which anchor the cellular tissue to the aragonitic base
are embedded in the calcareous base. Tylostyles have small and includes spherulous cells and glycocytes. There are a few inter-
rounded heads and mostly dull points; they measure 577 X 9.8 fLm cellular bacteria.
in mean dimensions. Diplasters (amphiasters) have short, blunt Remarks. The following important characteristics separate
spines and average 17.6 X 15.8 fLm. The basal skeleton plate is Willardia from Acanthochaetetes. Willardia has a base skeleton of
aragonite with finger-shaped processes 1 mm tall, 0.25 mm spaced aragonite mineralogy and finger-like ornamentation instead of cal-
protruding into the soft tissue and a microstructure of radiating citic pseudocalicles and tabulae; tylostyles are forming surface
needle like crystalline units. The soft-tissue sheet is 0.1-5 mm brushes instead of strands in the choanosome; and microscleres are
thick. Choanocyte chambers are small (16.8 fLm in diameter, only diplasters without spiraster-like variants.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Sollasellidae Lendenfeld, 1887

Rob W.M. Van Soest

Zoological Museum, University of Amsterdam P.O. Box 94766,1090 GT Amsterdam, Netherlands.

(soest@science.uva.nl)

Sollasellidae Lendenfeld (Demospongiae, Hadromerida) comprises a single Australian species, Sollasella digitata Lendenfeld, 1888. This
species is ramose with branches having in cross section a three-layered skeleton consisting of a cortical region of brushes of megascleres,
a subcortical region of low spicular density and an axial core of confused spicules and spicule bundles. Its status among Hadromerida is
incertae sedis, but closest relatives appear to be Polymastiidae, Suberitidae and Stylocordylidae.
Keywords: Porifera; Demospongiae; Hadromerida; Sollasellidae; Sollasella.

DEFINITION, DIAGNOSIS, SCOPE considered members of the family Halichondriidae, like

Epipoiasis, Axinyssa; these lack a cortical region and an axially
Synonymy condensed skeleton, and their spiculation is oxeas and may include
trichodragmas. Sollasella was assigned to Coppatiidae (a dustbin
Sollasellidae Lendenfeld, 1887a: 584. family for putative Epipolasida) by Hooper & Wiedenmayer (1994:
141), but these authors gave insufficient grounds for this. In its
Definition present concept, Sollasellidae is maintained solely for Sollasella.
With Polymastiidae, it shares the presence of a cortex and it is par-
Hadromerida with a cortex strengthened by short oxeas and ticularly similar to Pseudotrachya, with which it shares the combi-
an axial skeleton consisting of a mixture of long styles and short nation of choanosomal styles and ectosomal oxeas. With some
oxeas. No microscleres. Suberitidae (Homaxinella, Plicatellopsis, Rhizaxinella) it shares
the stalked habit and axially condensed skeleton. Some affinity
Diagnosis with Stylocordylidae is also apparent.

Stalked ramose sponges with polygonal ornamentation of

lines of inhalant openings. A cortical region is supported by a pal- SOLLASELLA LENDENFELD, 1888
isade of oxeas, and is separated from the axial core of confused
masses of styles and oxeas by a subcortical region with low spicu- Synonymy
lar density traversed by bundles consisting of a single style sur-
rounded by a mass of oxeas bound by spongin. Spicules oxeas and Sollasella Lendenfeld, 1888: 56.
styles, no microscleres.
Type species
Scope
Sollasella digitata Lendenfeld, 1888: 56 (by monotypy).
A single genus and species.
Definition
History and biology
Sollasellidae with stalked ramose habit and with polygonal orna-
The family was erected by Lendenfeld (1887a: 584), and later mentation of lines of inhalant openings on the surface. Choanosomal
clarified with the erection of the genus Sollasella with the definition skeleton consisting of an axial core of aligned spicule bundles and
, ... cortex of expanding tufts of radiating spicule-bundles ... interior extra-axial perpendicular bundles set at right angles to the axial core
skeleton irregular non-radial disposed spicule-bundles ... '. The and traversing a subcortical region. Ectosomal skeleton a distinct
family was subsequently maintained by Hallmann (1914: 286), pre- cortex of spicule brushes. Megascleres small oxeas and large styles.
dominantly because of the incompatibility of S. digitata with estab-
lished families. Topsent (l904b: 142) assigned Trachya hystrix to Previous review
Sollasella, but Hallmann (1914: 286) erected for that species the
genus Pseudotrachya and referred this to Axinellidae (now consid- Hallmann, 1914: 286.
ered to belong to Polymastiidae). De Laubenfels (1936a: 161)
assigned the family to his order Epipolasida (intended to receive Description of type species
families and genera with astrophorid affinities but lacking triaenes).
This order is no longer recognized and aster-bearing members are Sollasella digitata Lendenfeld, 1888 (Fig. lA-F).
assigned to Ancorinidae and Hemiasterellidae. De Laubenfels Synonymy. Sollasella digitata Lendenfeld, 1888: 56;
(1936a) assigned next to Sollasella also a series of genera here Hallmann, 1914; 287, pI. XV figs 1-2, text-fig. 1.

279
280 Porifera· Demospongiae • Hadromerida • SoUasellidae

A c

I
mm
mm
Fig. 1. Sollasella digitata Lendenfeld, 1888. A, habit of spirit specimen (scale I cm). B, BMNH lectotype (scale I cm). C, drawing of styles (scale
10 fLm) and details of heads (same scale I fLm). D, drawing of oxeas (scale 10 fLm) and details of apices (scale I fLm). E-F, SEM images of spicules (right
scales 1 fLm, left scale 10 fLm). E, oxea and detail of apex. F, style. A-D, reproduced from Hallmann, 1914: pI. XV figs 1-2 and text-fig. 1.

Material examined. Lectotype (per Hallmann, 1914): bundles bound by considerable amount of spongin, the bundles
BMNH 1886.8.27.639 (spirit specimen), including 2 slides. consist of a mixture of styles and oxeas. The subcortical region is
Paralectotype (here designated): AM G9I07 - 'syntype' mentioned traversed by spicule bundles with orientation perpendicular to the
by Hooper & Wiedenmayer (1994: 144). Other material. BMNH axial core, and these consist of a single central style and a mass of
unnumbered - dry specimen from 'Marley Beach, July 91931 surrounding oxeas. The styles protrude to some extent beyond the
No. 61, Dendy collection'. surface. The cortical skeleton is supported by the brushed endings
Description (mostly from Hallmann, 1914). Stalked ramose of the subcortical spicule bundles. Choanocyte chambers about
sponge (Fig. lA, B) of up to 14cm long, with rounded short 25 f.Lm in diameter, confined to the axial core. Cortex densely
branches. Consistency very firm, tough. Surface hispid due to pro- fibrous. Spicules (Fig. IC-F), oxeas with sharp points, often annu-
truding spicules, and provided with a characteristic polygonal pat- lated, 340-760 X 10-16 f.Lm; styles with often slightly swollen
tern of lines of round shallow depressions (Fig. IA), presumed to head and mucronate points (dubbed 'tylostrongyles' by Hallmann)
be inhalant openings. Oscules up to 2 mm in diameter, few in num- 2000--4000 X 10-35 f.Lm. Distribution and ecology. Port Jackson,
ber. In cross section the branches (Fig. IB) show three distinct E coast of Australia, no further data.
regions, an outer cortical region, an open subcortical region of low Remarks. The species has never been recorded since
spicular density, and an axially condensed region. The axial region Hallmann, 1914, and is apparently rare. Its true affinities remain to
consists of confusedly strewn spicules and anastomosing spicule be established.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Alectonidae Rosell, 1996

Klaus Riitzler

Department ofInvertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, DC, 20560-0163, USA.
(ruetzler.klaus@nmnh.si.edu)

Alectonidae Rosell (Demospongiae, Hadromerida), including Thoosidae Rosell & Uriz (part) and [Scolopidae] Sollas (a nomen oblitum),
contains species with limestone-excavating capability and habitus similar to many Clionaidae, including some massive (gamma) forms,
but lacking tylostyles. Instead, megascleres may be missing or include oxeas, styles, and strongyles, some with fanciful ornamentation.
Microscleres are generally large strongylote forms, bent or spiral and with simple or spirally arranged microspines, and small amphiasters
or diplasters. Because many of the species are rare and rather small and living cryptically inside calcareous substrates with other excavat-
ing sponges, contamination of spicules with foreign elements may be common. More material and reliable live observations are needed to
confidently separate genera and species. Six genera are presently recognized.
Keywords: Porifera; Demospongiae; Hadromerida; Alectonidae; Aleetona; Deleetona; Dotona; Neamphius; Seolopes; Spiroxya.
------- --- ---------

DEFINITION, DIAGNOSIS, SCOPE History and biology

Synonymy Despite the strong disagreement in spicule complement, species

in this family have traditionally been lumped with relatives of Cliona
[Scolopidae] Sollas, 1888: 432 (nomen oblitum). Alectonidae (possessing tylostyles and spirasters) under Clionaidae (or its junior
Rosell, 1996b: 53. Thoosidae Rosell & Uriz, 1997: 350 (part). homonym [Clionidae]), solely based on the common capability to
excavate limestone substrates. Most species known are small and live
in burrows made in calcareous algae, scleractinian corallites, or the
Definition axial skeleton of octocorals. Asexual reproduction through planc-
tonic, armoured propagules ('gemmules') has been reported for
Hadromerida with oxeas and their derivatives, microrhabds, some species (Topsent, 1920b; Tregouboff, 1942; Rosell, 1996a).
and amphiasters.
Remarks

Diagnosis The lack of tylostyles as principal megascleres in this family of

excavating sponges is considered a more important character than
Hadromerid excavating sponges, but including massive the large variety of microscleres that prompted some authors to
gamma forms, with oxeas, styles, or strongyles as megascleres separate a family Thoosidae from Clionaidae (Rosell & Uriz, 1997:
(or megascleres absent) and microrhabs (straight, bent, or spiral) or 350). Despite the similarities between amphiasters of Aleetona-like
amphiasters, or both, as microscleres. Megascleres may be smooth sponges, Thoosa and Cliothosa, I consider the latter two genera to
or spiny and kinked or branched forms may occur. Microscleres be closer to the Clionaidae than those discussed below.
are usually ornamented by microspines that are arranged uni- The massive species of Scolopes were previously allocated
formly, clustered, or spiraled. Excavations are chambered, as in to Coppatiidae (=Ancorinidae) but demonstrating Hadromerida
Clionaidae. Papillae, where known, are small and inconspicuous, structure, megasclere and microsclere geometry. They possess
massive sponges are rare. oxea megascleres and amphiaster and granular microstrongyle
microscleres, typical of other Alectonidae sponges, in particular
Neamphius. The inclusion of this genus here would theoretically
Scope require that the family name Scolopidae Sollas, 1888 takes prece-
dence over Alectonidae Rosell, 1996b. However, the name
Six genera are here assigned: Aleetona, Deleetona, Dotona, Scolopidae Sollas has not been used since 1899 and therefore can
Neamphius, Seolopes and Spiroxya. be suppressed under Article 23.9.1.1. of the ICZN (Anon., 1999).

KEY TO GENERA

(1) Megascleres present ............................................................................................................................................................................. 2

Megascleres absent .................................................................................................................................................................. ............ 6
(2) Robust tuberculate and smooth diactines (or derived polyactines) for megascleres, amphiasters for microscleres .............. Alectona
Megascleres are smooth monactines and diactines ............................................................................................................................. 3
(3) Megascleres are smooth oxeas ............................................................................................................................................................ 4
Megascleres are smooth styles or strongyles, microscleres include microspined rhabds and delicate, microspined
diplasters ........................ ............... ................................ ................. ........... .......... ................ ................ ................ ................. ...... Dotona

281
282 Porifera· Demospongiae • Hadromerida • Alectonidae

(4) Microscleres are only microspined rhabds ............................................................................................................................. Spiroxya

Microscleres include amphiasters; sponges grow massive .................................................................................................................. 5
(5) Microscleres are delicate, microspined amphiasters .......................................................................................................... Neamphius
Microscleres are minute diplaster-like amphiasters and granular microstrongyles ................................................................ Seolopes
(6) Microscleres are stout rhabds with annular swellings or with amphiastrose ornamentation, and delicate diplasters .......... Deleetona

ALECTONA CARTER, 1879 two symmetrical sets of perpendicular lumps (rays) near the center;
all microspined. Species are reported from the Indopacific and
Synonymy Atlantic region, and the Mediterranean Sea.

Corticium sensu Carter, 1879b: 353. Alectona Carter, l879c: Previous reviews
493 (not Alectona sensu Carter, 1880b: 58). [Nisella] Johnson,
1899: 463 (preocc.). Bavestrello et al. (1998a); Vacelet (1999b: 627).

Type species Description of type species

Alectona millari Carter, 1879c (by monotypy). Alectona millari Carter, 1879c (Figs 1-3).
Synonymy. Alectona millari Carter, 1879c: 495, pI. 17
Definition figs 1-7.
Material examined. Ho10type: BMNH 82.4.27.4 - spicule
Alectonidae with robust diactine and polyactine megascleres, preparation, 'from type', North Atlantic, 59°56'N, 6°27'W,
smooth or covered by spines or tubercles; with delicate amphiasters. between north of Scotland and Faroe Islands, 'Porcupine' Exped.,
1869, stn. 54, 202 m depth. Other material. BMNH - there is also a
Diagnosis specimen (fragment) labeled 'Alectona millari, pres. by Dr. Millar
(in Lophohelia), nr. 85.12.22.1, cf. slide in collection'; it is possible
Excavating sponges occupying chambers similar to clionaids that this is the holotype specimen from which the two available
in alpha stage. Most species have delicate papillae that are easily microscope slide preparations were made and that it was cata-
overlooked on the substrate surface, some exhibit massive growth logued at a later date under a different number.
outside the limestone-penetrating base. Spicules occur throughout Description (based on Carter, 1979a; with my own obser-
the sponge. Megasc1eres are stout oxeas, some kinked or with extra vations and measurements of spicules added). The sponge is
rays thus forming tri- or polyactines that are usually spined or cov- excavating a coral skeleton from which small papillae (ca. 1 mm in
ered by a few or many spines, tubercles, or mushroom-shaped diameter) are protruding. Spicules in the papillae are arranged in
knobs. Microscleres are delicate amphiasters with small center and radial fashion but they are dispersed throughout the choanosome
straight, slender or conical rays that are beset by minute spines or without particular orientation. There are three principal types of
hooks. In some species, amphiasters have a stout diactine axis and spicules: (1) Large spiny oxeas, bent, kinked, or sharply angled in

~r.' ~
F
..

B
G--

c ___ H_

..
~
.,
.'. " -~:.
, . ".
.
,;. . :
C - ' •• '

A __ E __ F __ 1 _ _• •
o
Fig. 1. Alectona, spicule characteristics. A-D, Alectona millari Carter, holotype BMNH 82.4.27.4. E-F, A. walichii Carter, drawn after SEM photomicro-
graphs in Bavestrello et al. (I 998a). G-I, A. triradiata Levi & Levi, drawn from SEM photomicrographs in Bavestrello et al. (1998a). A, large acanthoxea
(scale 50 !Lm). B, spiny triactines (scale 20 !Lm). C, small spiny oxeas (scale 20 !Lm). D, amphiasters (scale 20 !Lm). E, larger tuberculate oxea (scale 100 !Lm).
F, amphiaster (scale 5!Lm). G, spined triactine (one ray divided) (scale 50 !Lm). H, large amphiaster (scale 10 !Lm). I, small amphiaster (scale 5!Lm).
Porifera· Demospongiae • Hadromerida • Alectonidae 283

Fig. 2. Alectona millari Carter, holotype BMNH 82.4.27.4. SEM view of substrate excavation with large acanthoxea protruding (scale 100 11m).

Fig. 3. Alectona millari Carter, holotype BMNH 82.4.27.4. SEM view of spicules. A, small acanthoxeas. B, acanthotriactine. C, spiny micro strongyle,
transition to amphiasters (scale to 11m).

the center, with tubercle-like spines arranged in longitudinal rows; Remarks. De Laubenfels (1936a: 156) included Alectona
225-370 f.Lm long, 16-28 f.Lm in diameter. (2) Small, spiny or wallichii (Carter, 1874a) with A. millari, which would make
smooth oxeas, sharply kinked or bent in the center (a small number A. wallichii a senior synonym, but Bavestrello et al. (1998a)
with a third ray originating near the center, or polyactine), with demonstrated that the two species are distinct.
only few, sporadic, microspined tubercles and covered in patches The genus [Nisella] Johnson, 1899: 463 (preocc. by Nisella
by a fine spination; large size variation, 64-132 X 6-16 f.Lm. Heine & Reichenow erected in 1890, Aves; see Neave, 1940: 338)
(3) Amphiasters with fusiform, microspined shaft and two central was erected for type species [Nisella] verticillata Johnson, 1899:
circles of microspined tubercles (a few extra tubercles may occur 463, pI. VI fig. 6. No type material has been retrieved. Several
along the shaft); 35-60 X 9-14 f.Lm. specimens of this sponge were found boring in deep-water corals
284 Porifera· Demospongiae • Hadromerida • Alectonidae

off Madeira. The spicules include two types of amphiasters, one communicate with each other and with the outside through small
with two regular whorls of knobbed rays, the other with a short openings in the substratum. There are three types of spicules, all
shaft and unequal rays. One of the sponges contained in addition microscleres: Microstrongyles, slightly curved, with rounded ends
bent centrotylote oxeas. This description clearly fits the genus and annular constrictions alternating with microspined ridges
Alectona, and accordingly [Nisella] Johnson becomes a junior along the length of the rhabd; 21-85 f.Lm long, 6.4-10.6 f.Lm in
synonym. diameter. Hair-like, mostly toxiform oxea, ca. 50 f.Lm long.
Amphiasters with straight, thin shaft and equally delicate rays radi-
ating from two points near the center of the shaft; both shaft and
DELECTONA DE LAUBENFELS, 1936 rays are microspined, the amphiasters measure 21 X 20 f.Lm in
overall length by width.
Synonymy

Alectona sensu Carter, 1880b: 58 (not Alectona Carter, 1879c:

493). Delectona de Laubenfels, 1936a: 156.

Type species

Alectona higgini Carter, 1880b (by original designation).

Definition

Alectonidae without megascleres. Microscleres occur in haphaz-

ard arrangement and include rhabds and fusiform amphiasters
with tuberculate surface, toxiform oxeas, and amphiasters with long,
delicate rays.

Diagnosis

Sponges excavating and filling small chambers in limestone

(alpha stage). Papillae have not been observed. Spicules occur
throughout the choanosome, without particular orientation. There
A __ B __ c __
are no confirmed megascleres. The most common spicules are
Fig. 4. Delectona higgini (Carter), spicules (from Carter, 1880b; with
microstrongyles (rhabds) which are ornamented by microspined or
details from Bavestrello et aI., 1996). A, micro strongyle (scale 10 11m).
smooth annular swellings or tubercles and may be modified to B, toxiform oxea (scale 10 11m). C, amphiaster (scale 5 11m).
fusiform amphiasters or diplasters. Less common are toxiform
oxeas (some with central swelling) and delicate amphiasters with
long, thin, microspined rays (some may be reduced to oxyasters);
rugose discs occur in some species. All known species are boring in
skeletons of coralline algae (Melobesia) and cnidarians (Corallium,
Leptopsammia, Madracis). Distribution is Indian Ocean and
Mediterranean Sea.

Previous reviews

Bavestrello et al. (1996, 1997); Rosell (1996a).

Description of type species

Delectona higgini (Carter, 1880b) (Figs 4-5).

Synonymy. Alectona higgini Carter, 1880b: 58-59, pI. 5
fig. 25a-c.
Material examined. None. According to letters exchanged
between Shirley Stone (BMNH) and the curator at Liverpool
Museum (1979) the material (ZI.3033, syntypes and 2 spicule
slides) is missing since bombing during World War II (in lit.,
2000, Clare Valentine, BMNH). The type specimens occupied
coralline nodules dredged from the Gulf of Mannar (Manaar)
(Sri Lanka).
Description (from Carter, 1880b). The syntype specimens
are yellowish, in the dry state, and their tissue coats the honeycomb- Fig. 5. Delectona ciconiae Bavestrello, Calcinai, & Sara, SEM photomi-
like chambers they excavate in coralline (melobesian) nodules of crographs of microstrongyles (unpublished images, courtesy of the authors)
up to 30 mm diameter. The chambers reach 4 mm in diameter and (scale 10 11m).
Porifera' Demospongiae • Hadromerida • Alectonidae 285

c _

E _ G __
A_ 1:1_ 0_ F_

Fig. 6. Dotona, spicule characteristics. A-C, D. pulchella Carter, after Carter (l880b) and Topsent (1904b). E-G, D. davidi Kirkpatrick from SEM pho-
tomicrographs (E, F) and after Kirkpatrick (1900b) (E, adjusted to reflect the proportions of the measurements). A, styles and raphides (scale 10 fLm).
B, spiral-spined microstrongyles (scale 10 fLm). C, spiny microstrongyle (scale 10 fLm). D, diplasters (scale 5 fLm). E, strongyles (scale 20 fLm). F, raphides
(scale 10 fLm). G, spiral-spined microstrongyle (scale 20 fLm).

Remarks. The presence of rather large subtylostyles Synonymy. Dotona puZchella Carter, 1880b: 57, pi. 5
(345-355 X 10-27/-Lm) was signaled from one species in this fig. 24a-d.
group (Rosell, 1996a) but in view of other recent studies of related Material examined. None. According to letters exchanged
taxa (Bavestrello et aZ., 1996, 1997) it is likely that these spicules between Shirley Stone (BMNH) and the curator at Liverpool
were foreign. Future studies of new material should focus on this Museum (1979) the material (ZI.3032, holotype and 1 spicule slide)
discrepancy. is missing since bombing during World War II (in lit., 2000, Clare
Valentine, BMNH). The specimens occupied burrows in coralline
nodules dredged from the Gulf of Mannar (Sri Lanka).
DOTONA CARTER, 1880 Description (from Carler, 1880b). Specimens occurred in
small (3 mm), whitish masses filling cavities excavated in coralline
Synonymy (melobesian) nodules. Spicules of three kinds: curved strongyles
covered by a tightly wound spiral of minute spines; average size,
Dotona Carter, 1880b: 57. Dyscliona Kirkpatrick, 1900b: 352. 50.8 X 6.4 /-Lm. Very thin, hair-like spicules (raphides), about 85 /-Lm
long. Straight microstrongyles with diverging spines at both ends
Type species and a ring of spines around the center (diplasters), 8.5 X 2.1 /-Lm.
No megascleres are mentioned in the original description but
Dotona puZchella Carter, 1880b (by monotypy). Topsent (1904b), studying material from the Azores, emphasized
the presence of small styles (100 X 2 /-Lm), concentrated in bundles,
Dermition point outward, in the papillae. The same author provided additional
measurements and illustrations (Topsent, 1904b: 108, pi. 12, fig. 2)
Alectonidae with smooth styles or strongyles for principal of the spiral-spined strongyles (110-120 X 6/-Lm) and small spiny
megascleres and two types of microscleres, large, bent or undulate, 'spirasters' (diplasters), 6-8 X 2-3 /-Lm, which are located in the
microspined strongyles and small diplasters. papillae.
Remarks. Topsent (1904b) considered the small styles to be
Diagnosis megascleres and the (longer) spiral-spined strongyles to be very
large microscleres. We support this distinction because the styles
Alpha-stage excavating sponges with whitish tissue filling playa structural role in the papillae.
very small chambers and tiny papillae. Megascleres are styles or In contrast to the older authors, Rosell & Driz (1997) report
strongyles which may form perpendicular bundles in the papillae the presence of oxeas in material examined by them. If this obser-
or are scattered through the choanosome. The main microscleres vation is substantiated, a merger of Dotona with the genus
are bent or undulating strongyles ornamented by fine spination Spiroxya (see below) should be contemplated. Stylote and strongy-
which most commonly is patterned in a spiral around the rhabd. lote modifications of oxeas are not unusual, for instance, in Cliona
Less conspicuous microscleres, if present, are short, stout, spiny acus (Bavestrello, Ca1cinai, & Sara) and C. sarai (Melone), but
microstrongyles or diplasters and hair-like (raphide) spiCUles. Dotona, here, is understood to be without oxeas as principal
Specimens of the few species and forms so far reported are from megascleres.
Indopacific, Atlantic, and Mediterranean locations. Dotona davidi Kirkpatrick, 1900b: 352 (type of Dyscliona
Kirkpatrick, holotype BMNH 1900.10.19.12) clearly contains
Previous review strongyles as megascleres; unfortunately, a preparation made from
the small remaining holotype did not reveal the spiral-spined
Rosell & Driz (1997). microstrongyles illustrated by Kirkpatrick (1900b: pi. 14, fig. Id).
The strongyle size originally reported as 126-246 X 3-5.5 /-Lm was
Description of type species found to be larger (up to 268 X 11 /-Lm) and contrary to the original
description, they are never roughened on the ends (SEM obser-
Dotona pulchella Carter, 1880b (Fig. 6). vation). The microstrongyles measured by Kirkpatrick were
286 Porifera· Demospongiae • Hadromerida • Alectonidae

A B c
Fig. 7. Neamphius huxleyi (Sollas), holotype BMNH 1889.1.1.76. A, habitus with attachment base added to suggest boring capacity evident from
incorporated rock fragment (scale 10 mm). B, oxeas (scale 100 !Lm). C, amphiaster (scale 5 !Lm).

90 X 3 /-Lm and considered rather rare. Inclusion by Kirkpatrick Description. The original description (Sollas, 1888) could
(l900b: 353, pI. 14, fig. If, g) of Cliona purpurea Hancock in be confirmed. The specimen measures 11 X 6 cm, 7 cm high. There
Dyscliona was inappropriate, as it was shown to belong to are several fused mounds topped by oscula, 4-10 mm in diameter.
Paracomulum in the Poecilosclerida (see discussion in Riitzler & The ectosome ('skin') is brown and wrinkled, the texture is soft
Stone, 1986). elastic. Pores are clustered, sieve-like. Choanocyte chambers are
diplodal and measure about 24 /-Lm in diameter. There are only two
spicule types. The megascleres are oxeas, partly scattered but also
NEAMPHIUS DE LAUBENFELS, 1953 organized in radial strands. Oxeas are slender, straight or gently
and evenly bent, gradually tapering to sharp points; they measure
Synonymy 425-600 X 6-8 /-Lm. Microscleres are amphiasters and are dispersed
throughout the choanosome. They are delicate, with two whorls of
[Amphius] Sollas, 1888: 177 (preocc. by a Coleoptera). four slender, microspined rays, slightly bent outward toward an iden-
Neamphius de Laubenfels, 1953a: 545. tical fifth ray which is a continuation of the axial shaft. Amphiasters
measure 16-20 X 12-16 /-Lm, rays are about 2 /-Lm in diameter.
Type species Remarks. De Laubenfels (1936a: 156) listed Amphius huxleyi
under Clionaidae but commented that it was 'not certainly boring'.
Amphius huxleyi Sollas, 1888 (by monotypy). His hunch seems well founded, however, because fragments of sub-
strate in close contact with the sponge, examined by SEM, show fresh
Definition etchings and dislodged chips characteristic of sponge burrowing.
We do not agree with Burton & Rao (1932) who emended
Alectonidae of massive (gamma stage), initially boring, with the original diagnosis to 'microscleres in the form of euasters or
oxeas in radial tracts and delicate, microspined amphiasters. some modification thereof' to accommodate their species Amphius
sollasi, which has euasters and does not belong here.
Diagnosis

Massive sponges with large oscula elevated on mounds. They SCOLOPES SOLLAS, 1888
excavate calcareous substrate at their base but alpha-stage borers
are hitherto unknown. Radial skeleton structure. Megascleres are Synonymy
oxeas, scattered throughout the choanosome and organized in
vague tracts radiating from the base toward the surface where they Scolopes Sollas, 1888: 432. Paracordyla Hallmann, 1912: 132.
are oriented in tangential fashion. Microscleres are abundant and
scattered throughout the choanosome. Only known from moderate Type species
depth (120m) in the southern Pacific Ocean.
Scolopes moseleyi Sollas, 1888 (by monotypy).
Previous review
Definition
Burton & Rao (1932).
Alectonidae with two size categories of oxeas. Microscleres
Description of type species include both minute amphiasters (diplaster-like) and granulated
centro-angulate microstrongyles.
Neamphius huxleyi (Sollas, 1888) (Figs 7-8).
Synonymy. Amphius huxleyi Sollas, 1888: 178, pI. 42. Diagnosis
figs 5-11.
Material examined. Holotype: BMNH 1889:1:1:76 - Massive (gamma) growth form, with oxea megascleres in two
'Challenger' Exped., off Api, New Hebrides, 108-126 m. size classes, the larger forming a compact choanosomal skeleton,
Porifera· Demospongiae • Hadromerida • Alectonidae 287

Fig. 8. Neamphius huxleyi (Sollas), holotype BMNH 1889.1.1.76, SEM photomicrographs. A-B, tylostyles and amphiasters (scales: A, 100 ....m; B, to ....m).
C, amphiaster enlarged (scale 5 .... m). D, substrate grain with characteristic pitting of sponge excavations, one chip freshly etched out but still attached
(scale 20 ....m).

the smaller forming an ectosomal palisade, and with an intermedi- Description of second species
ate category of oxeas (or occasionally modified styles) forming a
sparse paratangential subectosomal skeleton that protrudes through Seolopes lignea (Hallmann, 1912) (Fig. 9).
the surface. Microscleres are minute amphiasters (diplaster-like), Synonymy. Paraeordyla lignea Hallmann, 1912: 132.
and granulated centro-angulate microstrongyles. Seolopes lignea; de Laubenfels, 1936a: 160.
Material examined. Ho10type: AM Z195 (slide) - Off
Scope Sandon Bluffs, southern NSW, 70-80 m depth, RV 'Endeavour' .
Description (from type material and original description).
Three species: Seolopes moseleyi Sollas, Paraeordyla lignea Sponge massive, apparently with branch-like elongations (? blind
Hallmann, 1912, Seolopes megastra de Laubenfels, 1953. fistules), with dense and solid consistency. Surface even and pilose.
Oscules not observed. Choanosomal skeleton is composed nearly
Description of type species exclusively of densely packed larger oxeas in more-or-Iess parallel
arrangement, thicker towards the axis and at the core of the fistules.
Seolopes moseleyi Sollas, 1888 (Fig. 9). In the subectosomal region are bundles of larger oxeas criss-
Synonymy. Seolopes moseleyi Sollas, 1888: 432, pI. XLll crossing and projecting through the surface in vaguely plumose
figs 1-9. bundles. The ectosomal skeleton is a dense erect or paratangential
Material examined. Holotype: BMNH (not seen) - Bahia. palisade of small oxeas (described as microscleres by Hallmann,
Description (from Sotlas). Massive growth form, surface 1912, but more accurately a second category of smaller megascle-
even and uniformly pilose, oscules and pores not observed, with a res). Contrary to Hallmann's observations only very few of these
dense spicular cortex. Choanosomal skeleton with coarse spicular are modified to styles. Smaller oxeas are also dispersed throughout
fibres radiating to the surface, cored by large oxeas, with interme- the mesohyl, particularly surrounding lacunae of the aquiferous
diate sized oxeas projecting through the surface in loose plumose system. Amphiasters and granulated microstrongyles are rare
tracts. Ectosome with radially arranged smaller oxeas in an erect within the ectosomal or choanosomal skeletons. Megascleres large
palisade lying between the protruding medium sized oxeas. oxeas, symmetrical or asymmetrical, slightly bent towards one
Smaller oxeas also scattered within the mesohyl and around lacu- end, between 650-3500/Lm or more long, 15-55 /J-m wide. Some
nae formed by the aquiferous system. Megascleres oxeas of two smaller oxeas, modified styles or occasionally tylostyles present
clearly differentitated size classes. Largest (choanosomal oxeas) in surface brushes, and probably intermediate examples between
straight or slightly curved, symmetrical, sharply pointed the two size categories of megascleres (uncommon, reported as
(2250 X 60/Lm); medium sized (subectosomal oxeas) more cylin- 200-700/J-m long and maximum diameter 18/J-m). Small oxeas of
drical, abruptly pointed (556 X 13/Lm); smallest (ectosomal oxeas) the ectosomal skeleton usually symmetrical, occasionally asym-
cylindrical, straight, abruptly pointed (125 X 4/Lm). Microscleres metrical and bent slightly towards one end, but with symmetrical
amphiasters with spines at the ends in terminal whorls (7X3/Lm). points (80-135/J-m long, 3-6/J-m wide). Microscleres are minute
288 Porifera· Demospongiae • Hadromerida • Alectonidae

= -
Fig. 9. Seoiopes, spiculation and habitus. A-D, S. moseieyi Sollas. A, habitus. B, choanosomal skeletal structure. C, ectosomal palisade and hispidation.
D, spiculation. E-M, S. lignea (Hallmann). E, habitus (scale 40 mm). F, choanosomal skeletal structure (scale 300 !Lm). G, tip of large choanosomal
oxea (scale 60 !Lm). H, tip of medium sized subectosomal oxea (scale 30 !Lm). I, small oxea of ectosomal palisade (scale 20 !Lm). J, granular microrhabd
(scale 10 fLm). K, amphiasters (scale 5 fLm). L, large choanosomal oxeas (scale 500 fLm). M, small and medium sized ectosomal oxeas (scale 100 !Lm).
(A-D, after Sollas, 1888, pI. XLIII. E-F, after Hallmann, 1912, fig. 27, pI. xxii fig. 2.)

amphiasters, slightly resembling diplasters, with clusters of spines Both genera have been previously allocated to
at either end and a smooth rod in between, rare (3--6!Lm long, Rhaphidhistiinae (=Trachycladidae) (de Laubenfels, 1936a),
1-2 fLm wide at the centre). Microstrongyles are also present, gran- Coppatiidae (= Ancorinidae), or Spirastrellidae, and comparisons
ulated over the whole surface and with a slight centrotylote have also been made with Suberitidae (e.g., Trachya = Aaptos).
swelling, also rare (12-16 fLm long, 2-3 fLm wide). Very few spi- It is clear, however, that Scolopes belongs to Hadromerida based
rasters also seen (about 14 X 3 fLm) but most likely contaminants. on its skeletal structure and spiculation, whereas its supposed
Remarks. Paracordyla Hallmann is an obvious synonym of resemblance to Jaspis, Trachycladus, Rhaphidhistia or Trachya (=
Scolopes, despite Hallmann's (1912) arguments to the contrary. Aaptos) is superficial. Conversely, the definitive allocation of
These arguments concern the possession of fibres in the cortex Scolopes to a family is more difficult given that on the one hand
and the radial skeletal structure in Scolopes, allegedly different it has an organised ectosomal skeletal palisade that fits better with
from features of Paracordyla. However, illustrations provided by Polymastiidae (e.g., Pseudotrachya), but on the other hand its
both Sollas (1888) and Hallmann (1912) are nearly identical, and amphiaster and microstrongyle microscleres are the same as those
observed differences are probably due to different growth states of found in some Alectonidae. The possession of oxeas (not
preservation of their respective material. The type material of tylostyles) also precludes this genus from Clionaidae, and for these
Scolopes from the BMNH has not yet been examined, and conse- reasons the genus fits closest with Alectonidae. It is closest to
quently a description of P. lignea is also provided as this material Neamphius in having a massive (gamma) growth stage, and amphi-
has been re-examined. No new material of Scolopes has yet been aster microscleres. By comparison, Scolopes has two size categories
collected, with the descriptions given above based on antiquated of oxeas (Neamphius has only one), with the smaller organised into
museum specimens over a century old. Consequently, it is not pos- an ectosomal palisade, and in having both amphiasters and
sible at this time to confirm whether Scolopes has an excavating microstrongyles (versus only amphiasters in Neamphius). In the
growth stage. case of the type species, S. moseleyi, microscleres are apparently
Porifera· Demospongiae • Hadromerida • Alectonidae 289

A c E G

Fig. 10. Spiroxya spicules. A-C. S. heteroclita Topsent, after Topsent (1900), and from holotype, MNHN DT 2413. D-E, Cliona ievispira Topsent, after
Topsent (1898b), and from holotype, MNHN DT 873. G-F, after SEM photomicrographs of Scantiletta sp. (courtesy B. Calcinai). A, oxeas (scale 100 j..Lm).
B, spiral microstrongyles (scale 10 j..Lm). C, spiral-spined microstrongyles (scale 10 j..Lm). D, oxeas with styloid modifications (scale 100 j..Lm). E, spiral
microstrongyles and amphiasters (scale 10 j..Lm). F, spiral micro strongyle with microspination (scale 10 j..Lm). G, amphiaster (scale 2 j..Lm).

abundant and clearly native, albeit very small. In the case of Diagnosis
P. lignea, it is uncertain whether or not all the microscleres are
native to the sponge (notwithstanding re-examination of the type Excavating and thinly encrusting, small sponges inhabiting
material; courtesy of J.N.A. Hooper), as they are very rare. In both the scleraxis of octocorals and crevices in rock. Oxeas are present
species amphiasters are extremely small (about 5/-Lm long), with as megascleres and occur in two size classes: the larger in tracts
clusters of spines at either end reminiscent of diplasters or nodu- crossing the choanosome, smaller ones arranged in palisade-
lose amphiasters. Furthermore, a second form of microsclere was fashion in the papillae (Calcinai et al., in press). Styloid modifica-
also discovered in P. lignea, consisting of small granular tions or accessory spicules may occur. Microscleres are dispersed
microstrongylote rhabds, with slight centrotylote swelling (remi- throughout the sponge body. The microscleres are microspined,
niscent of acanthorhabds seen in some Theonellidae lithistids). spiral rhabds or short, straight spiny microstrongyles, or both.
These were not mentioned by Sollas (1888) for S. moseleyi
although he depicts something similar in his plate XLm, figure 4,
Previous reviews
and it can only be assumed that he considered these were growth
stages of amphiasters.
Rosell & Driz (1997); Calcinai et al. (in press).

SPIROXYA TOPSENT, 1896 Description of type species

Synonymy Spiroxya heteroclita Topsent, 1896a (Fig. 10).

Synonymy. Spiroxya heteroclita Topsent, 1896a: 122;
Spiroxya Topsent, 1896a: 121. [Scantilla] Johnson, 1899: 462 Topsent, 1900: 280, pl. 8 fig. 11.
(preocc. by an Insecta). Donotella de Laubenfels, 1936a: 156 (for Material examined. Holotype: MNHN DT 2413 (spicule
Cliona acustellaAnnandale, 1915a: 14). Scantiletta de Laubenfels, preparation) - Banyuls (Cap Abeille), France (Mediterranean).
1936a: 156 (replacement for [Scantilla] Johnson). Cliona Grant, Other material. Cliona levispira Topsent, 1898b: 235, fig. 2ii:
1826a, in part (e.g., c. levispira Topsent, 1898b: 235; C. pruvoti MNHN DT 873 (8 PA 1895), DT 874 (8 PA 1897) -Azores. Cliona
Topsent, 1900: 104; C. sarai Melone, 1965: 348; C. acus pruvoti Topsent, 1900: 104, pl. 3, fig. 6: DT 2424 - Banyuls.
Bavestrello et al., 1995: 380). Description. The original author describes a thin whitish
crust penetrating crevices in rock, no openings known. Nothing
Type species else is known about the live type species. Related taxa are whitish
or yellowish and clearly boring, with chambers ranging 1-6 mm in
Spiroxya heteroclita Topsent, 1896a: 122 (by monotypy). diameter and very small papillae (Melone, 1965; Calcinai et al., in
press, pers. comm.). Oxeas are bent, asymmetrical, and taper to
Definition sharp points. These megascleres have a great size range and can be
separated into a smaller category of about 76-110 X 1.5-2.5 /-Lm
Alectonidae with oxeas as principal megascleres and one or and a larger one of 200-900 X 15-20 /-Lm. The microscleres are of
two size classes of microspined rhabds as microscleres. two kinds: spiral strongylotes with 4-8 turns, smooth or minutely
290 Porifera· Demospongiae • Hadromerida • Alectonidae

spined, 60 X 3 flm in average dimension; curved (but not spiral) Because several combinations of oxeas with spiral rhabds, spiral-
strongyles, 70 X 4 flm, ornamented with small spines that run in a spined rhabds, and spiny rnicrostrongyles may occur in this group of
spiral around the rhabd. excavating species there is no justification of separating genera until
Remarks. Topsent (1896a) considered the spiral rhabds to be more material becomes available. In this context, Spiroxya is the
smooth but high magnifications show that many have rnicrospination. senior genus.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Order Chondrosida Boury-Esnault & Lopes, 1985.

Family Chondrillidae Gray, 1872

Nicole Boury-Esnault

Universite de la Mediterranee, Centre d'Oceanologie de Marseille, Station marine d'Endoume, UMR CNRS 6540, rue de la Batterie des
Lions, 13007-Marseille, France. (esnault@com.univ-mrs.fr)

Chondrosida Boury-Esnault & Lopes (Demospongiae) contains only a single family Chondrillidae Gray, including junior syno-
nym Chondrosiidae Schulze, with seven nominal genera of which four are valid: Chondrosia, Chondrilla, Thymosia and Thymosiopsis.
Chondrillidae was previously considered close to both the order Hadromerida and other tetractinellid orders, such as Astrophorida,
warranting recognition in a new order as there was no argument in favour of one solution over the other. This order was subsequently
supported by recent molecular data confirming its monophyly although there are still no clear phylogenetic hypotheses on relationships
with the other orders of Demospongiae. Chondrosids are oviparous, encrusting to massive with a marked cortex enriched with fibrillar
collagen, inhalant apertures localised in special structures and a skeleton (when present) composed of siliceous spicules or spongin fibres.
Chondrosid species are widely distributed mainly in shallow waters, exceptionally up to 750m depth. They are often present in semi-
obscure conditions, on vertical walls, at the entrance of submarine caves, or below rocks.
Keywords: Porifera; Demospongiae; Chondrosida; Chondrillidae; Chondrosia; Chondrilla; Thymosia; Thymosiopsis.

DEFINITION, DIAGNOSIS, SCOPE History and biology

Synonymy The concept of Chondrosida is relatively recent and the order

was erected in 1985 by Boury-Esnault & Lopes for the unique fam-
Order: Chondrosides Levi, 1956. Chondrosiina Wiedenmayer, ily Chondrillidae. Chondrosida includes oviparous species with
1977b. Chondrosida Boury-Esnault & Lopes, 1985: 172. Family: a well developed cortex reinforced by fascicles of fibrils of colla-
[Gummineae] Schmidt, 1862 (nomen oblitum). [Gumminidae] gen, with numerous spherulous cells, relatively small choanocyte
Schmidt, 1862 (nomen oblitum). Chondrillidae Gray, 1872a. chambers and a mesohyl with very abundant symbiotic bacteria.
Chondrosidae Schulze, 1877. Chondrissinae Lendenfeld, 1885: 14. Skeleton may be absent or present but is always reduced and made
Chondrosiidae Wiedenmayer, 1977b. of either siliceous microscleres of the aster type or irregular and
nodular spongin fibres. Megascleres are always absent. The inhalant
Diagnosis of the order and family apertures are always localised in pore-sieves or cribriporal chones.
There is some debate on the correct name for this family.
Demospongiae, encrusting to massive, with a marked cortex Gray (1872a) erected Chondrillidae for Chondrilla and Corticium
enriched with fibrillar collagen, with inhalant apertures localised Schmidt, 1862, and Schulze (1877) subsequently erected
in pore-sieves or cribriporal chones and a skeleton often absent, Chondrosiidae to include these two genera plus Chondrosia.
composed when present of nodular spongin fibres or aster micro- Corticium was later transferred to Homoscleromorpha. Although
scleres only (never megascleres). Collagen always very abundant. these two family names have been used together in recent literature
Oviparous. (e.g., Wiedenmayer, 1977b), most authors used Chondrosiidae in
the sense of Topsent (1895), including three genera Chondrosia,
Type genus Chondrilla and Thymosia, whereas Hooper & Wiedenmayer
(1994) emended this name to Chondrillidae based on the rule of
Chondrilla Schmidt, 1862 (by original designation). priority. Similarly, although Chondrillidae is a junior synonym of
Gummineae Schmidt, 1862, the prevailing usage of Chondrillidae
Scope must be maintained (ICZN, Art. 23.9) based on 'reversal of prece-
dence'. Gummineae has not been used subsequent to 1899 whereas
Single family with over seven generic names of which only Chondrillidae has been used as a valid name by at least 10 authors,
four are now considered valid and most appropriately referred here. in at least 25 works in the immediately preceding 50 years
Approximately 50 species have been published worldwide mostly (Colin & Arneson, 1995; Desqueyroux-Faundez & van Soest,
belonging to the genera Chondrosia Nardo, 1847b (10 valid 1997; Erhardt & Moosleiner, 1995; Fromont, 1999a; Hechtel,
species) and Chondrilla Schmidt, 1862 (24 valid species). 1965, 1969, 1976; Hooper, 1994, 1997; Hooper & Wiedenmayer,
Although some of the nominal chondrosid species are now known 1994; de Laubenfels, 1950a, 1954; Lazoski et al., 2001; Lehnert &
to be junior synonyms, others which are allegedly cosmopolitan van Soest, 1998, 1999b; Pattanayak, 1999; Pulitzer-Finali, 1993;
(Chondrosia reniformis Nardo, 1847b and Chondrilla nucula Reitner, 1991; Thomas, 1973, 1979; Vacelet et al., 2000; Vacelet &
Schmidt, 1862) are certainly complex of species (Klautau et al., Perez, 1998; Wiedenmayer, 1977b, 1989, 1994). The seniority of
1999; Lazoski et al., 2001). The genera Thymosia Topsent, 1895 Chondrillidae over Chondrosiidae has no bearing on the choice of
and Thymosiopsis Vacelet & Perez, 1998 contain two species. the ordinal name Chondrosida (ICZN Art. 40).

291
292 Porifera· Demospongiae • Chondrosida • Chondrillidae

Chondrillidae has been tossed back and forth within the have a worldwide distribution, and have been recorded from the NE
demo sponge ordinal classification, sometimes even by the same Atlantic and Mediterranean (Topsent, 1895; Pulitzer-Finali, 1983),
author, considered close to Hadromerida (e.g., Topsent, 1925c; West Atlantic and Caribbean (Wiedenmayer, 1977b; de Laubenfels,
Bergquist, 1978; Hooper, 1997), and Tetractinellida (e.g., Topsent, 1936a; Pulitzer-Finali, 1986), South Atlantic (Boury-Esnault, 1973)
1895; Levi, 1973), prompting Boury-Esnault & Lopes (1985) to and Indo-Pacific and Australia (Hooper & Wiedenmayer, 1994).
erect the order Chondrosida for the family, citing that there is no Thymosia has been recorded in the NE Atlantic and in SW Pacific
argument to favour one solution or the other. Recent molecular (N. Zealand) whereas Thymosiopsis has to be considered for the
analyses have confirmed that Chondrosida is a monophyletic clade moment as an endemic of the NW Mediterranean.
distinct from Hadromerida and Tetractinellida, although analyses Although the monophyly of the family is supported on molec-
failed to find clear phylogenetic relationships with other orders of ular grounds (Chombard, 1998; Vacelet et ai., 2000), it is less evi-
Demospongiae (Chombard, 1998). dent with morphological characters. The synapomorphies between
The most important debate concerns the placement of Chondrosia and Chondrilla are cribriporal chones, thick cortex
Thymosia, a genus with a spongin fibre skeleton. It has, in tum, with dense fascicles of collagen, disjunction between both cellular
been considered as Chondrosiidae (Topsent, 1895), Spongiidae (de and collagen elements and a granulo-fibrillar matrix (Vacelet &
Laubenfels, 1936a), Verongiinae (de Laubenfels, 1948), Dysideidae Donadey, 1987). The synapomorphies between Thymosia and
(Borojevic et ai., 1968) and finally again Chondrosiidae (Hiscock Thymosiopsis are presence of pore-sieves and an ectosome consti-
et ai., 1984; Boury-Esnault & Lopes, 1985; Rosell, 1988). tuting a weakly developed cortex with fibrillar collagen. The prob-
Recently Vacelet & Perez (1998) described a new genus with- able homology between pore-sieves and cribriporal chones has
out a skeleton from the Western Mediterranean Sea, Thymosiopsis, yet to be confirmed, as does the homology on the structure of the
also included in this family. A second species of this genus was dis- cortex. Relationships between the four genera also remain to be
covered recently and molecular analysis of both species, and of confirmed. The generic classification of the Chondrillidae centres
Thymosia guemei, confirmed that they are closely related and both around the presence/absence of a skeleton and if present the type of
genera made a monophyletic clade with Chondrosia reniformis the skeleton, i.e., spongin fibre versus siliceous spicules.
(Vacelet et ai., 2000).
Chondrosid sponges are widely distributed mainly in shallow Previous reviews
waters, exceptionally up to 750m depth. They are often present
in semi-obscure conditions, on vertical walls, at the entrance Schulze, 1877: 1-38; Topsent, 1895: 512; Wilson, 1925: 353;
of submarine caves, or below rocks. One species, Thymosiopsis Boury-Esnault & Lopes, 1985: 172; Hooper & Wiedenmayer,
cuticuiatus, is typical of obscure caves. Chondrosia and Chondrilla 1994: 122.

KEY TO GENERA

(1) Chondrosida with skeleton ................................................................................. 2

Chondrosida without skeleton ................................................................................ 3
(2) Chondrosida with siliceous skeleton ................................................................. Chondrilla
Chondrosida with spongin skeleton .................................................................... Thymosia
(3) Chondrosida with a well developed cortex ............................................................ Chondrosia
Chondrosida with a thin cortex and a cuticle .......................................................... Thymosiopsis

CHONDROSIA NARDO, 1847 of the genus Celluiophana Schmidt, 1862, which was also assigned
to Gumrninae is a colonial ascidian (Schulze, 1877: 35). Chondrosia
Synonymy tubercuiata Schmidt, 1868: 24, was synonymised by Schulze, 1877:
31, to Oscarella (Halisarca) iobuiaris and is considered a valid
Chondrosia Nardo, 1847b: 267; Schmidt, 1862: 40; Gummina species of Oscarella by Boury-Esnault et ai. (1992b).
Schmidt, 1862: 37; Schmidt, 1864; Topsent, 1895: 568; Topsent, Among the species described nine seem to be valid. However
1929c. the genus probably has many more species because several, such as
Chondrosia reniformis, C. piebeja Schmidt, 1868, C. collectrix
Type species Lendenfeld, 1888 are allegedly cosmopolitan and whereas they
most likely represent a complex of species, as demonstrated for
Chondrosia reniformis Nardo, 1847b (by monotypy). C. reniformis (Lazoski et al., 2001). There are also many nomenclat-
ural problems which have to be resolved through re-examination of
Diagnosis type specimens, if they are still available, one being Celluiophana
collectrix Schmidt, 1862, the type specimen of which is lost.
Chondrillidae without skeleton and with a well developed However the type specimens need to be sufficiently well preserved
cortex made of thick fascicles of collagen and with numerous to make histological slides, the only way to discriminate the
spherulous cells. species. Dry specimens are useless.

Remarks Distribution

Gummina gliricauda Schmidt, 1862 (type species of Gummina) Cosmopolitan, tropical, subtropical and temperate zones,
was synonymised by Schmidt himself with Chondrosia reniformis mostly in the littoral zone above 50m depth. Chondrosia piebeja
(Schmidt, 1864). Celluiophana pileata Schmidt, 1862, type species was dredged once at 760 m depth.
Porifera· Demospongiae • Chondrosida • Chondrillidae 293

Fig. 1. Chondrosia. A, in situ photograph of a Mediterranean specimen of Chondrosia reniformis (J. Vacelet) (scale 0.5 cm). B, microphotograph of histo-
logical slide from the Lendenfeld collection BMNH 96.11.5.109 (abbreviations: CO, cortex; CH, choanosome; sp, foreign spicule; arrow, cribriporal
chone) (scale 400 /-Lm).

Description of type species spicules are present in the choanosome and in the internal layer
of the cortex. The border cortexlchoanosome is underlined by a
Chondrosia renifonnis Nardo, 1847b (Fig. I). line of spherulous cells (Fig. lB). Choanocyte chambers ovoid to
Synonymy. Chondrosia renifonnis Nardo, 1847b: 267; spherical are about 40 fLm in diameter with a small aphodus at
Schmidt, 1862: 40; Gummina ecaudata Schmidt, 1862: 38; Gummina the apopy1e of the chamber. Extracellular symbiotic bacteria and
gliricauda Schmidt, 1862: 38; Schulze, 1877: 13; Topsent, 1895: bacteriocytes are present in the mesohyl (anatomical description
568; Topsent, 1925c. taken from neotype and other slides made by Lendenfeld in 1896
Material examined. Holotype: Unknown - North Adriatic from Adriatic specimens).
Sea. Neotype: MNHN DNBE 2000; BMNH: 96.11.5.112 (slides)- Reproduction. The type species is oviparous (Scalera-Liaci
coli. February in Trieste, Lendenfeld Adriatic sponges; BMNH: et aZ., 1971). Oogenesis occurs from May to August and spermato-
96.11.5.109 (radial serial section, thick) - Trieste, Collection Rao; genesis from July to August. The emission of the spermatozoids
BMNH: 33.3.1.45 - Napoli. Other material. Personal collection - and oocytes occurs between the first quarter and the full moon of
specimens from Western Mediterranean (Provence Coast). Gummina August (Levi & Levi, 1976). The oocytes remain trapped in follic-
ecaudata Schmidt - evidently from Schmidt species, Adriatic. Slides ular cells near the basis of the sponge. The fecundation is external.
from BMNH (labelled Chondrosia renifonnis) - doubtful or evi- The larva is a blastula. During embryogenesis, there is a transmis-
dently misinterpretations: BMNH 158.12.29.127 - Hebrew sion of bacteria and spherulous cells from the mother-sponge to the
University collection, Eylath Red Sea; BMNH 1954.2.23.37 - larva (Levi & Levi, 1976).
Herdman's Ceylon sponges; BMNH 1936.11.2b.8a - Reef bay, Remarks and distribution. We consider that only populations
Pt Elizabeth South Africa, 5.7.36, Presd. Prof. JA Stephenson; BMNH of Chondrosia renifonnis from the Mediterranean and the nearest
32.4.5.16a - Dry Tortugas, Dendy collection; BMNH 1939.5.8.56- Atlantic (Coasts of Spain, Portugal and Morocco) (Lazoski et ai.,
off Tampa Bay, USA, Belgian Museum ColI. 'Mercator'. 2001) belong to this species. Specimens from localities outside the
Description. Specimens are generally lobate and can reach Mediterranean vary principally in the localisation and abundance of
30 cm in greatest dimension and 3 cm thick. The colour in life is from foreign materials and spherulous cells and are certainly different
black on parts exposed to the light to white on parts unexposed to species (Lazoski et aZ., 2001).
light. The consistency in life is cartilaginous, firm, and tough. The
surface is smooth, shiny (Fig. lA) (features taken from living
Mediterranean specimens). The upper cortex is composed of two lay- CHONDRILLA SCHMIDT, 1862
ers: one superficial layer with numerous spherulous cells, one inter-
nallayer with few spherulous cells. The basal cortex shows a lower Synonymy
density of spherulous cells and the external zone is devoid of any
spherulous cells. Spherulous cells are abundant also around canals. Chondrilla Schmidt, 1862: 38. Magog Sollas, 1888: 442 (for
The density of the spherulous cells in the choanosome is similar to Chondrilla saccifonnis Carter, 1879b: 299); de Laubenfels, 1936a:
that of the internal layer of the upper cortex. The spherulous cells 182; Topsent, 1895: 512; Topsent, 1918: 601. Chondrillastra
contain about 20 spherules of about 3 fLm in diameter. Cortex is also Topsent, 1918: 603 (for C. australiensis Carter, 1873b); de
characterised by a net of large fascicles of collagen fibrils. Foreign Laubenfels, 1936a: 182.
294 Porifera· Demospongiae • Chondrosida • Chondrillidae

Fig. 2. Chondrilla. A. in situ photograph of a living specimen of Chondrilla nucula from the Provence coast (western Mediterranean) (J. Vacelet) (scale
1 cm). B, microphotograph of a slide from the Lendenfeld collection BMNH 96.11.5.108 (abbreviations: CO, cortex; CH, choanosome) (scale 500 !Lm).
C, SEM micrograph of a spheraster of the type specimen (courtesy of R. Desqueyroux-Faundez) (scale 7 !Lm).

Type species Distribution

Chondrilla nucula Schmidt, 1862 (subsequent designation by Cosmopolitan, tropical, subtropical and temperate zones,
de Laubenfels, 1936a). mostly in the littoral zone above 50m depth.

Diagnosis Description of type species

Chondrillidae with a skeleton only composed of siliceous Chondrilla nucula Schmidt, 1862 (Fig. 2).
spicules of the aster type localized mainly in the cortex and around Synonymy. Chondrilla nucula Schmidt, 1862: 38; Schulze,
the canals. 1877: 24; Topsent, 1918: 601.
Material examined. Holotype: LMJG 15018 - Quarnero,
Remarks North Adriatic, Mediterranean Sea. Paratype: LMJG 15687 (with
handwritten labels by Lendenfeld; Desqueyroux-Faundez & Stone,
Among the 26 nominal Chondrilla species described about 1992). Other material. Personal collection: Western Mediterranean
20 seem to be valid. However, the genus probably has many Sea (Ligurian and Provence coast). BMNH 96.11.5.108 (slide)-
more species because one (Chondrilla nucula) is considered as Lesina, Adriatic, colI. of Lendenfeld. BMNH 1938.5.26.109 -
cosmopolitan (Klautau et al., 1999), and others have a very Acca, Palestine, colI. of Dr K. Reich. Slides from BMNH (labelled
large distribution in the Indo-Pacific region like Chondrilla Chondrilla nucula) - doubtful or evidently misinterpretations.
australiensis. A complete revision is necessary with examination of BMNH 23.10.1.257 - Dendy, 1924 collection, 15m, R.N.VIII,
type specimens which are well preserved and of living popUlations. 'Terra Nova' sponges; BMNH 24.5.1.115 - Port Philip Heads,
Porifera· Demospongiae • Chondrosida • Chondrillidae 295

RN 490, Victorian sponges, J.B. Wilson coIl; BMNH 1938.4.26.15- Remarks

Richardson Cove, Bermuda, 74.13.16, collected by Dr J.F.Cr.
Wheeler; BMNH 1954.2.12.258 - Dendy collection, South coast of The nodular spongin fibres present in the type species were
Australia. misinterpreted by Bergquist (1980b) who suggested that they may
Description. The type specimen is small about 0.9 cm in have been a foreign hydroid skeleton, whereas they are so abundant
length and 0.6 cm in width and has a rounded shape. A paratype of in well preserved specimens that it is impossible to miss them.
unknown locality is lobate and about 4.5 cmll cm. The colour in Although the genus is currently monotypic, the only species
life of Mediterranean specimens is brown to brown violet and the described, Thymosia guemei, is distributed over a large area of the
general shape is rounded to lobate (Fig. 2A). The consistency is NE Atlantic. Originally described by Topsent (1895) from the
cartilaginous, firm, and tough. The surface is smooth. In the south coast of French Brittany, the species has been redescribed in
Lendenfeld's slides of Adriatic specimens, the cortex is about detail from diverse populations throughout the NE Atlantic (Boury-
175-280/-Lm thick with a cuticle of about 5/-Lm (Fig. 2B). It is Esnault & Lopes, 1985; Rosell, 1988).
composed of two layers; the upper one is a dense sheet of spheru-
lous cells and spherasters, whereas the internal one is made of scat- Distribution
tered spherulous cells and spherasters. The choanocyte chambers
are ovoid, aphodal and have a diameter of about 40-50 /-Lm. In the Northeast Atlantic from Lundy (Great Britain) to Lagos
choanosome the spherulous cells and the spherasters are scattered (South Portugal) and Azores (Portugal) - 51°11' -37"N,
in the mesohyl. Around the canals the spherasters are organized in 4°40' -28°39'W - in caves and overhanging walls. Depth: 0-36m.
an unispicular layer. Extracellular symbiotic cyanobacteria are Apparently absent from the Mediterranean. Recently a new species
present in the upper layer of the cortex and symbiotic bacteria and have been found at the entrance of a cave in New Zealand
bacteriocytes are present in the mesohyl. Spicules. The spicules are (Bergquist, pers. comm.).
spherasters. Average diameter varied along the year from 20-
27/-Lm (Bavestrello et ai., 1993). The number of rays varied Description of type species
between 20-25 in the holotype. The extremities of the rays are
most often rugose or slightly spinose (Fig. 2C). Thymosia guemei Topsent, 1895 (Fig. 3).
Reproduction. The species is oviparous as are all species of Synonymy. Thymosia guemei Topsent, 1895: 574; Hiscock
the order. Oogenesis and spermatogenesis occurs mostly during et al., 1984: 20; Boury-Esnault & Lopes, 1985: 172; Rosell,
August although oogenesis may last from July to October. The 1988: 353.
oocyte is surrounded by follicular cells (Scalera-Liaci et al., 1971; Material examined. Holotype: MNHN DT 1895 -
Gaino, 1980). The release of the gametes, the embryogenesis and Concameau, south coast of French Brittany, North East Atlantic,
the larva have never been observed. Topsent collection (dry). Other material. Personal collection: alco-
Remarks and distribution. The morphology and the size of hol fixed specimens from Lagos Portugal, 37"N, 8°50'W; Sagres,
the spherasters can vary between populations of the same species Portinho da Arrabida, and Azores (llha de Faial and Ilha de Sao
and seem not to be a good character to discriminate between the Miguel: 38°31'N, 28°37'W to 37°45'N, 25°30'W); and Lundy,
species of Chondrilla (Bavestrello et al., 1993; Klautau et al., 51 °Il 'N, 4°40'W, Great Britain.
1999). However their distribution and abundance in the cortex and Description. Plate-like in shape with a maximum dimen-
the choanosome, as well as the distribution and abundance of the sions of about 2 cm X 10 cm. The surface is conulose with cone-
spherulous cells, seem to be better discriminant characters. As shape projections of about 0.5 mm high raised up by the termination
these characters have been used rarely a precise redescription of all of the fibres. The colour in life is light grey to cream. The pore
the species is necessary. sieves are located in some depressions and are about 5 mm in
We consider Chondrilla nucula to comprise only the popula- diameter. The oscules are at the top of cone-shaped projections and
tions from Mediterranean and the nearest Atlantic (Coasts of Spain, are 2 mm in diameter. At the basis a layer of foreign detritus is most
Portugal and Morocco) (Klautau et al., 1999). often present (taken from dry holotype). The cortex of about half
a millimetre thick is composed of two layers of collagen. From
outside to inside are present the exopinacocyte layer composed
THYMOSIA TOPSENT, 1895 of T-shaped cells, a layer of dense collagen up to 50 /-Lm thick, a
second layer of collagen up to 450 /-Lm filled up by numerous
Synonymy spherulous cells. The spongin fibres stop at the level of the first
layer and slightly raise the surface. The choanosome is composed of
Thymosia Topsent, 1895: 574; de Laubenfels, 1948: 103; spherical to ovoid choanocyte chambers of about 30 /-Lm in diame-
Bergquist, 1980b: 499; Boury-Esnault & Lopes, 1985: 172; Rosell, ter. A central cell is most often visible at the apopyle of the cham-
1988: 353. bers. In the mesohyl different cell types are present (lophocytes,
glycocytes, archaeocytes) and two types of spherulous cells. Type 1
Type species has a diameter up to 13 fLm and is filled by 3 to 5 spherules of about
3 fLm in diameter. They are particularly numerous in the layer 2 of the
Thymosia guemei Topsent, 1895 (by monotypy) cortex. Type 2 are cells of about 16 /-Lm in diameter and filled with 20
small spherules. They are less abundant than type 1. The skeleton is
Diagnosis composed of fibrillar collagen which give the particular texture of the
cortex and by dendritic heavily nodular fibres. The choanosome is
Chondrillidae with skeleton composed of nodular spongin densely packed by the fibres which are perpendicular to the surface
fibres. with a diameter varying from 100/-Lm at the basis to 40 /-Lm at the top.
296 Porifera· Demospongiae • Chondrosida • Chondrillidae

Fig. 3. Thymosia. A, specimen of Thymosia guernei from the south coast of Portugal (Portinho da Arrabida) (scale 35 mm). B, transverse section of
Thymosia guernei where spongin fibres (arrow) are visible (scale 35 mm).

The bark is laminar and the pith is granular. The fibres are dendritic,
although some anastomoses are present. They are easily breakable
(anatomical details taken from MNHN Azores specimen).
Remarks and distribution. The nodular fibres have been
interpreted as a hydroid skeleton (Bergquist, 1980b) whereas in
living specimens these fibres are clearly native to the species and
cannot be confused with a hydroid skeleton (Topsent, 1895; Boury-
Esnault et Lopes, 1985; Rosell, 1988). Found mostly on wave
exposed coasts on vertical to overhanging walls and entrance of
caves from 6-20m depth.

THYMOSIOPSIS VACELET & PEREZ, 1998

Synonymy

Thymosiopsis Vacelet & Perez, 1998: 7; Vacelet et al.,

2000: 316.

Type species
Fig. 4. Thymosiopsis cuticu/atus. In situ photograph of a specimen from the
Thymosiopsis cuticulatus Vacelet & Perez, 1998 (by original type locality (J. Vacelet) (scale 1 em).
designation).

Diagnosis discovered recently at the entrance of the same cave (Vacelet et aI.,
2000). The cortex of Thymosiopsis is considerably thinner than that
Chondrillidae with a general organization similar to that of of Chondrosia.
the genus Thymosia having a smooth surface, a marked cortex
(40-50 !Lm) enriched with fibrillar collagen parallel to the surface, Distribution
but lacking spongin fibres. A superficial cuticle and pore-sieves
may be present. The choanocyte chambers are eurypylous. Known only from caves 3PP, Fauconniere and Plane on the
Provence coast of Mediterranean (France).
Remarks
Description of type species
The genus has been recently described from a dark cave on
the coast of the French Mediterranean. A second species was also Thymosiopsis cuticulatus Vacelet & Perez, 1998 (Fig. 4).
Porifera· Demospongiae • Chondrosida • Chondrillidae 297

Synonymy. Thymosiopsis cuticulatus Vacelet & Perez, volume is made by cells with inclusions of three types often
1998: 7-13. grouped in clusters. Reproduction has not been observed.
Material examined. Holotype: MNHN-D JY.59 - '3PP Remarks. This sponge appears to have affinities with
cave', 1.2 km south-west of La Ciotat, coast of the French Thymosia. The two species share the presence of pore-sieves and
Mediterranean, 43°09' 47"N, 5°36'Ol"E, 16-22 m depth, on vertical a weakly collagenous developed cortex. A molecular study using
or overhanging walls, 30-80m from the cave opening. Paratypes: 28SrRNA has shown that Thymosia and Thymosiopsis made a
MNHN D JV 60 - same locality. monophyletic clade (Vacelet et al., 2000). The obvious morpholog-
Description. Encrusting up to 15-20cm, and 3-5mm thick ical differences between Thymosiopsis and Chondrosia are the
in the centre, thinner on the edges. The colour is white or yellowish dimension of the cortex, the abundance of collagen bundles as well
white in living specimens (Fig. 4). The surface is smooth, but as the type of choanocyte chamber and canal system organization,
irregular as the sponge closely follows the irregularities of the sub- which precludes its allocation to the latter genus.
stratum. The consistency is quite cartilaginous, although easy to
tear. There is neither spicules nor a spongin fibre skeleton. A small ACKNOWLEDGEMENTS
amount of foreign material is frequently included in the
choanosome. The ectosome is 40-50 fLm thick and is lined on the I thank the Natural History Museum, London (Clare
outer surface by a thin, non-cellular cuticle 2 fLm thick. The ecto- Valentine), and the Musee de Geneve (Ruth Desqueyroux-Faundez)
some contains bundles of collagen fibrils parallel to the surface, for having made available slides and type specimens of Chondrosia
and two types of cells with inclusions without special localisation. reniformis and Chondrilla nucula, Javier Cristobo and Pilar Rios for
Choanocyte chambers occurs in relatively low density, they are collection of specimens of Thymosia from Portugal, Jean Vacelet
eurypylous and 15-30fLm in diameter. Most of the choanosome and Thierry Perez for slides and type specimens of Thymosiopsis.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

'Lithistid'Demospongiae
Andrzej Pisera 1 & Claude Levi2

1 Instytut
Paleobiologii, PolskaAkademia Nauk, ul. Twarda 51155, 00-818 Warszawa, Poland. (apis@twarda.pan.pl)
2 Museum National d'Histoire Naturelle, Laboratoire de Biologie des Invertebres Marins et Malacologie, 57, Rue Cuvier, 75231 Paris
Cedex 05, France. (ievi@mnhnJr)

'Lithistid' demosponges are a polyphyletic group (formerly order Lithistida Schmidt, 1870) containing 13 extant families and 36 included
genera, plus five poorly known genera of uncertain status (most probably synonyms of other genera but their affinities are not yet certain).
The only shared character amongst the lithistids is the presence of more-or-less irregular articulated choanosomal spicules called desmas,
which form a rigid skeleton in most genera. The polyphyletic nature of 'lithistids' is revealed by the wide array of geometry (tetraxial, mon-
axial, polyaxial or anaxial) of its desmas and of ectosomal spicules (dichotriaenes, phyllotriaenes, discotriaenes, rhabds, oxeas, or pseudo-
discotriaenes and pseudophyllotriaenes), as well as microscleres (amphiasters, spirasters, sigmaspires, raphides). We propose here, for the
reason of their undoubted polyphyletic nature, to formally abandon the order Lithistida Schmidt, both for Recent and fossillithistids.
Keywords: Porifera; Demospongiae; Lithistida, 'lithistids'; Azoricidae; Corallistidae; Desmanthidae; Isoraphiniidae; Macandrewiidae;
Neopeltidae; Phymaraphiniidae; Phymatellidae; Pleromidae; Scleritodermidae; Siphonidiidae; Theonellidae; Vetulinidae.

DEFINITION, DIAGNOSIS, SCOPE 1910, Pleromidae Sollas, 1888, Scleritodermidae Sollas, 1888,
Siphonidiidae Lendenfeld, 1903, Theonellidae Lendenfeld, 1903,
Synonymy Vetulinidae Lendenfeld, 1903.

Lithistidae Schmidt, 1870: 21; Carter, 1873c: 437; Zittel, Remarks

1878a: 96. Lithistina Carter, 1875b: 185; 1880b: 496; Rauff,
1893: 194. Lithistida Sollas, 1888: 284; Lendenfeld, 1903: Originally the group name Lithistidae had been proposed by
125; de Laubenfels, 1955b: E44; Bergquist, 1978: 158; Hartman, Schmidt (1870) and later used as an ordinal taxon by Zittel
1982: 644. (1878a). The corrected spelling Lithistida was formally proposed
by Sollas (1888) who defined it as 'tetractinellida provided with a
Definition consistent skeleton by the zygosis of modified spicules or desmas' .
Earlier, the spelling Lithistina was used by Carter (1875b, 1880b)
'Lithistid' Demospongiae (polyphyletic) characterized by in an informal way to describe a group within his family
choanosomal articulated spicules called desmas forming a rigid Pachastrellida; he also used the spelling Lithistidae in his earlier
skeleton in most genera (informal group). work (Carter 1873c: 437). Sollas (1888) divided Lithistida in two
suborders: Hoplophora (with ectosomal spicules and microscleres)
Diagnosis and Anoplia (without ectosomal spicules and microscleres).
Anoplia contained families Azoricidae and Anomocladidae Zittel
Polymorphic, polyphyletic (informal) group of demosponges (for Vetulina). On the other hand, Hoplophora were divided by
which may be encrusting, massive irregular, cup or ear-shaped, fla- Sollas in Triaenosa, having ectosomal spicules in the form of tri-
bellate cylindrical, branched, globular, with or without pedicel, aenes and microscleres as spirasters, amphiasters and/or rhabds,
with or without atrial cavity. Ectosomal spicules when present may and Rhabdosa, having ectosomal spicules as microstrongyles or
be discotriaenes, pseudodiscotriaenes, phyllotriaenes, pseudophyl- their derivatives and monocrepid desmas. Triaenosa included fami-
lotriaenes, dichotriaenes, rhabds and oxeas. Choanosomal desmas lies Tetracladidae Zittel (corresponding to our Theonellidae and
equally diversified and may be either tetraxial (tetraclones), Phymatellidae), Corallistidae (corresponding to our Corallistidae
monaxial (rhizoclones, megaclones, dicranoclones, heloclones, or and Neopeltidae) and Pleromidae. Rhabdosa included Scleritoder-
various complex branched forms), polyaxial or anaxial (sphaero- midae and Cladopeltidae (=Siphonidiidae). This subdivision is not
clones). Microscleres may be represented by various rhabds, oxeas, followed here because polyphyly of lithistids is generally accepted,
spirasters, amphiasters, sigmaspires and raphides (or various and at least some families used by Sollas are also polyphyletic.
combinations of these). Levi (1973) united lithistid families bearing some form of ectoso-
mal triaene into Desmophorida, leaving other forms as heteroge-
Scope neous assemblages classified only to family level.
The polyphyletic nature of Lithistida is revealed by the wide
The informal group 'lithistids' contains 13 extant families: array of desma geometries (tetraxial, monaxial, polyaxial or anax-
Azoricidae Sollas, 1888, Corallistidae Sollas, 1888, Desmanthi- ial), as well as of ectosomal spicules (dichotriaenes, phyllotriaenes,
dae Topsent, 1894d, Isoraphiniidae Schrammen, 1924a, rhabds, oxeas and others) and microscleres (amphiasters, spiras-
Macandrewiidae Schrammen, 1924a, Neopeltidae Sollas, 1888, ters, sigmaspires, raphides). Lithistids were recognised as being a
Phymaraphiniidae Schrammen, 1910, Phymatellidae Schrammen, polyphyletic taxon a long time ago (SchranImen, 1910, 1924a;

299
300 Porifera· Demospongiae· 'Lithistids'

Dendy, 1921; Burton, 1929b), based on differences in form and however, there is still no general proposition for distributing partic-
development of desmas and other spicules, and this tenet now is ular lithistid families among the various orders of Demospongiae,
generally accepted (Reid, 1963a, 1968d, 1970; Bergquist, 1978; and thus we have taken here a conservative but practical approach
Levi, 1973, 1991; Gruber, 1993), being also supported by molecu- to keep all such forms together in an informal group, proposing,
lar studies (Kelly-Borges & Pomponi, 1994). Unfortunately, however, to formally abandon the order Lithistida Schmidt.

KEY TO EXTANT FAMILIES OF LITHISTIDA

(1) Desmas as sphaeroclones ................................................................................................................................................... Vetulinidae

Desmas as megaclones ......... .... .......... ................. .... .... .... ... ................. ........... ................. .............. ........... ..... ...................... Pleromidae
Desmas as heloclones ................................................................................................................................................... Isoraphiniidae
Desmas as dicranoclones .. ........... ...... .......... ........... ..................... ................ ...................... ........................................... ... Corallistidae
Desmas as triders ...................................................................................................................................................... Phymaraphiniidae
Desmas with triaenose, rarely monaxial crepis, ectosomal phyllotriaenes to discotriaenes ..................................... Macandrewiidae
Desmas as rhizoclones ........................................................................................................................................................................ 2
Desmas as tetraclones .......................................................................................................................................................................... 3
Desmas monaxial or probably monaxial but not rhizoclones .............................................................................................................. 4
(2) Ectosomal spicules when present are various rhabds or oxeas, microscleres when present are
sigmaspires ........ .... ............ ...... .... ..... .... ... ............... ............................ ................................ ... ....... ........... ................. Scleritodermidae
No ectosomal spicules, microscleres absent or raphides .................................................................................................... Azoricidae
No ectosomal spicules, exotylostyles present ................................................................................................................ Siphonidiidae
(3) Ectosomal spicules as dichotriaenes ............................................................................................................................. Phymatellidae
Ectosomal spicules as phyllotriaenes or discotriaenes .................................................................................................... Theonellidae
(4) Desmas trider-like (probably monaxial), styles present ................................................................................................ Desmanthidae
Desmas of complex branching shape, ectosomal pseudophyllotriaenes or pseudodiscotriaenes ..................................... Neopeltidae

LITHISTID SPICULE TERMINOLOGY DendrocIone, monaxial desma with long central shaft and
terminal zygomes, sometimes may develop branches and then
There are several papers, including the most recent Thesaurus resembles tetraclone (only fossil) (see Figs 1-2 in Fossil Lithistids).
(Riitzler & Boury-Esnault, 1997) that present definitions of various Desmas, articulating choanosomal megascleres of various
skeletal features of lithistid sponges. No single source, however, geometry and usually complex morphology, often secondarily
includes all terms, and in some cases they are conflicting or con- modified and very irregular, in lithistids.
fusing. Consequently, we include a short glossary of spicular terms Dichotriaene, ectosomal spicule of triaenose symmetry with
particular to lithistid sponges as used in this revision. Most of the regularly developed cladome (may be smooth or tuberculated or
terms are based on, and may be found in Reid (1970). Some lithis- spinose) which is tangential to the sponge surface; rhabdome,
tid terms are given also in Kelly (2000) although we diverge from which is perpendicular to the cladome usually long, axial canals
her usage of some of these terms in several respects. extend along the whole length of the rhabdome and clads (see
Fig. 9A-D in Phymatellidae).
Recommended terms Dicranoclone, monaxial arched desma bearing well devel-
oped tubercles, it may be bi, tri or sometimes tetrapodial. Zygomes
Amphitrider, tetraclone resembling amphitriaene. terminal (articulating with upper tuberculated portion of other
Astroclone, desma with various number of arms radiating desma) (see Fig. lIA-B in Corallistidae and Fig. 7B-J in Fossil
from the center (may be polyaxial or anaxial) (e.g., fossil Lecanella Lithistids ).
Zittel, 1878a). DidymmocIone, monaxial desmas resembling two sphaero-
Brachyome, shortened (without zygome) arm of the trider clones connected by a horizontal beam (only fossil) (see Fig. 9B-I
(see definition). in Fossil Lithistids).
ChiastocIone, monaxial or anaxial desma resembling dendro- Discotriaene, ectosomal spicule with tangential part rounded
clone (see definition) but having short shaft, often with numerous to oval in shape (flat or slightly concave) and entire or incised mar-
radiating branches (only fossil) (see Fig. 3D in Fossil Lithistids). gin, with short rhabd, crepis tetraxial, very short in the disc (see
Cladome, see under definition of Phyllotriaene. Fig. 2A-B in Family Theonellidae).
Clones, ray like arms of the desmas which are partly (i.e., HelocIone, elongated monocrepidial desmas with sinuous
crepis extends only in the part of the desma close to its center) or shaft, no branches, and zygomes as lateral notches; axial canal usu-
entirely anaxial (crepis absent); number of clones is governed by ally present along the whole desma length (see Fig. 31-0 in Family
the crepis geometry. Isoraphinidae).
Clonome, in trider, three similar arms of the desma. MegacIone, arch-shaped, usually branched, smooth or rarely
Crepidome, see crepis (Rauff, 1893 term). mounded desma (with an uneven surface), with short monaxial
Crepis, the inceptional body of a desma (=axial filament) (when visible at all) crepis; zygomes terminal (articulating with
visible as canal (monocrepid desma) or canals (tetracrepid desma) lateral and or upper part of other desma) (see Fig. 6 in Family
(Sollas 1888 term). Pleromidae and Fig. lOB-J in Fossil Lithistids).
Porifera· Demospongiae· 'Lithistids' 301

Phyllotriaene, ectosomal spicule with one ray called rhab- Figs lA-B and 3A-B in Family Phymaraphiniidae, and Fig. 16A
dome, usually perpendicular to the sponge surface and penetrating in Fossil Lithistids).
it, and three others are more or less flat (tangential to the sponge Zygome, articulating part of the de sma.
surface) and irregular, branching, called cladome; crepis tetraxial,
very short in the cladome (see Figs 7A-C and lOC-F in Family Terms recommended to be abandoned
Theonellidae, and Fig. 21B in Fossil Lithistids).
Pseudodiscotriaene, ectosomal spicule resembling discotri- Lophotriaenes, to be abandoned (erroneus interpretation),
aene and analogous to it but monocrepid (monaxial) (see Fig. llA used by Hentschel for des mas of his Lophocanthus.
in Family Neopeltidae). Megarhizoclone, term used to described desmas of various
Pseudophyllotriaene, ectosomal spicule closely resembling geometry (modified dicranoclones, strongly branched astroclones
phyllotriaene and analogous to it, but monocrepid (crepis may be and even megaclones) and having nothing in common, except very
located in the rhabd or in the cladome) (see Fig. 9J-L in Family superficial resemblance, with real rhizoclones; for this reason it
Neopeltidae). should be abandoned.
Rhabdome, see under definition of Phyllotriaene.
Rhizoclone, monaxial (monocrepidial) desma, usually with
numerous spines and or/pointed tubercles which serve as zygomes ACKNOWLEDGEMENTS
(mostly lateral) (see Fig. 5C-D in Family Azoricidae and
Fig. 14B-J in Fossil Lithistids). The authors are indebted to Clare Valentine (NHM, London),
Sphaeroclone, anaxial desma in which several ray-like Ardis Jonston (MCZ, Harvard), Prof. Nancy Voss (RSMAS,
arms extend from a globular center, which may be spinose Miami, Florida), Dr. Shirley Pomponi and John Reed (HBOI, Fort
(see Fig. 12B-L in Fossil Lithistids). Pierce, Florida), Dr. Valter Raineri (Museo Civico di Storia
Tetraclone, tetraxial (tetracrepidial) desma, may be smooth Naturale 'Giacomo Doria', Genova) for their invaluable help to
or tuberculated; very regular, but often secondarily modified (some locate and organize the loan of original specimens. Royal Society
rays shorter, angles between clones different from 120 degrees, (London), DAAD (Bonn) and Museum National d'Histoire
clones branching, sometimes resembling rhizoclone etc.), its real Naturelle (Paris) supported AP's visits to the sponge collections in
nature may be recognized by crepis observation which is usually London, Bonn, Gottingen, Tiibingen and Paris, respectively.
very short; zygomes mostly terminal (tips of desma articulating Drs. Hans Jahnke (Universitiit Gottingen), Alexander Liebau
with other tips) (see Fig. 15E-F in Family Theonellidae and (Universitat Tiibingen) and Sarah Long (NHM, London) helped
Fig. 21A in Fossil Lithistids). during our studies of fossil lithistid collections in their care;
Trider, variant of tetraclone with triaenose symmetry; Mr. Freek Rhebergen (Emmen, Holland) and Mr. Ulrich von Hacht
zygomes terminal, one arm without articulation (brachyome); (Hamburg, Germany) send us some photographs; we would like to
crepis tetraxial present along the whole length of clones (see express our gratitude to all of them for their help.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Scleritodermidae Sollas, 1888

Andrzej Pisera1 & Claude Levi2

I Instytut Paleobiologii, Polska Akademia Nauk, ul. Twarda 51155, 00-818 Warszawa, Poland. (apis@twarda.pan.pl)
2 Museum National d'Histoire Naturelle, Laboratoire de Biologie des Invertebres Marins et Malacologie, 57, Rue Cuvier, 75231 Paris
Cedex 05, France. (levi@mnhnJr)

Scleritodermidae Sollas (Demospongiae, 'lithistids') consists of five valid genera and about 14 nominal species, usually deep water and
a few from shallow reef cavities, occurring worldwide in a warm climatic belt. The family is characterized by choanosomal rhizoclone
desmas, ectosomal spicules when present are acanthorhabds, smooth strongyloxeas, or strongyles/styles with spinose distal ends;
microscleres when present are spinose sigmas pires.
Keywords: Porifera; Demospongiae; 'lithistids'; Scleritodermidae; Aciculites; Amphibleptula; Microscleroderma; Scleritoderma; Setidium.

DEFINITION, DIAGNOSIS, SCOPE acanthorhabdslstrongyles, styles or smooth strongyloxeas; micro-

scleres when present are spinose sigmaspires.
Synonymy
Scope
Scleritodermidae Sollas, 1888: 346; Topsent, 1894d: 290;
Schrammen, 1903: 20; Wilson, 1925: 463; de Laubenfels, 1936: Five extant genera: Aciculites Schmidt, 1879; Amphibleptula
165; Scleritodermatidae Lendenfeld, 1903: 141. Taprobaneidae de Schmidt, 1879; Microscleroderma Kirkpatrick, 1903a; Scleritoderma
Laubenfels, 1936: 74 (partim). Amphibleptulidae de Laubenfels, Schmidt, 1879; Setidium Schmidt, 1879. Some fossil rhizomorine
1936: 102 (partim). lithistids may also belong here, as indicated by their structure.

Definition Remarks

'Lithistid' Demospongiae with rhizoclone desmas, ectosomal Further studies may finally lead to splitting this family, as some
spicules when present are acanthorhabds/strongyles, styles or smooth sponges included here have no microscleres, and are allocated based
strongyloxeas, microscleres when present are spinose sigmaspires. on their general structure, ectosomal spicules and type of rhizoclone
desmas, while others having microscleres display no ectosomal
Diagnosis spicules. On the other hand, acanthorhabds/strongyles, styles or
smooth strongy10xeas occurring in this family cannot, most probably,
Polymorphic: massive, encrusting, ear-shaped, foliated, be considered homologous to similar spicules occurring in non-
cup- to vase-shaped or flabellate; desmas are thorny or tubercu- lithistid demosponges, and thus used as characters enabling alloca-
lated rhizoclones; ectosomal spicules when present are various tion of particular genera to non-lithistid families or higher taxa.

KEY TO EXTANT GENERA

(1) Microscleres present ............................................................................................................................................................................ 2

Microscleres absent, ectosomal spicules are acanthorhabds/strongyles ............................................................................... Aciculites
(2) Ectosomal spicules are acanthorhabds ........................................................................................................................... Scleritoderma
Ectosomal spicules are smooth strongyloxeas ........................................................................................................................ Setidium
Ectosomal spicules absent, vase shaped or foliate in shape ........ ..... ............................ ........ ......... ...... .... .... ...... ...... Microscleroderma
Ectosomal spicules absent, massive conical or globular in form, with several raised complex oscular areas ..... ... ... .... Amphibleptula

ACICUUTES SCHMIDT, 1879 Definition

Synonymy Scleritodermidae with acanthose strongyles and/or styles as

ectosomal spicules; no microscleres.
Aciculites Schmidt, 1879: 29; Sollas, 1888: 347; Wilson,
1925: 463; Dendy 1905: 101; Van Soest & Stentoft, 1988: 63. Diagnosis
Sympyla Sollas, 1888: 353.
Polymorphic: massive, foliated or incrusting with short papil-
Type species lae. Rhizoclone desmas bear strong blunt spines; ectosomal
spicules are acanthose anisostrongyles and/or styles; microscleres
Aciculites higginsi Schmidt, 1879: 29 (by monotypy). of any type absent.

302
Porifera· Demospongiae· 'Lithistids'· Scleritodermidae 303

Remarks. The position of Aciculites within Scleritodermidae,

despite the absence of microscleres, is based on the presence of
tangential layer of acanthostrongyles and rhizoclone desmas, char-
acters shared with Scleritoderma, which has both ectosomal
spicules and microscleres. Support for such placement is also given
by molecular data (Kelly-Borges & Pomponi, 1994). About seven
species are attributed to this genus.

Description of type species

Aciculites higginsi Schmidt, 1879 (Figs 1-3).

Synonymy. Aciculites higginsi Schmidt, 1879: 29, pI. 2: 1,
4,13; Van Soest & Stentoft, 1988: 63, pI. 8: 6-7, fig. 29.
Material examined. Specimens: MZUS P0153; ZMA POR
5249, 5250. MNHN DeL 1158; ZPAL Pf.1512 (ex ZMA
POR5250).
Description. Pyriform (when small) to pillow-shaped sponge
having a flat top with strong and wide vertical ridges separating Fig. 1. Aciculites higginsi Schmidt, lateral view, ZPAL Pf.15/2 (ex ZMA
depressed areas which are inhalant and bear densely distributed pores. POR5250), (scale 10 mm).

Fig. 2. Aciculites higginsi Schmidt, A-C, E-F, ZPAL Pf.15/2 (ex ZMA POR52S0); D, type MZUS POlS3. A, view of the ectosome with tangential styles
(scale 200 /-Lm). B, surface of a choanosomal skeleton on a top of a sponge showing oscules and superficial canals in the choanosomal skeleton (scale 2 mm).
C, surface of the choanosomal skeleton from the lateral inhalating area (scale 1 mm). D, surface of the choanosomal skeleton (scale 200 /-Lm).
E, choanosomal rhizoc1one desmas (scale 200 /-Lm). F, details of rhizoclone desmas (scale 100/-Lm).
304 Porifera· Demospongiae· 'Lithistids'· ScIeritodermidae

Fig. 3. Aciculites higginsi Schmidt. A-D, tylostyles and strongyl (A-B, ZPAL Pf.15/2 ex ZMA POR5250; C-D, MZUS P0153; scale 50 j-Lm).
E-G, heads and tips of tylostyles and strongyle (E-F, ZPAL Pf.15/2 ex ZMA POR 5250, scale 20 j-Lm; G, MZUS PO 153, scale 20 j-Lm).

Oscules, one or several, slightly raised, 860-1180 f.Lm in diameter, Diagnosis

located on the top of the sponge being the center of radiating and sin-
uous subdermal canals about 560 f.Lm wide, visible on the surface of Massive conical to cylindrical sponges with several raised
the choanosomal skeleton. Ectosomal megascleres are styles and oscular areas in which open numerous exhalant canals; desmas are
tylostyles, usually slightly curved, with smooth or spinose head, vary- highly tuberculated rhizoclones. No special ectosoma1 spicules,
ing greatly in size, 188-350 f.Lm long, 8-11 f.Lm thick. They form a microsc1eres are sigmaspires. Other megascleres are long oxeas.
dense tangential layer on the sponge surface. Choanosomal skeleton
consists of massive rhizoclones about 200-350 f.Lm in size bearing Remarks
blunt spines (or tubercles when young and on canal surfaces); these
spines are usually divided into smaller spines that bear in tum very Van Soest & Stentoft (1988) considered the genera Micro-
small spines at the top. No microscleres present of any kind. This scleroderma Kirkpatrick, 1903a and Taprobane Dendy, 1905 as
species is known only from the Caribbean from a depth of 183 m. junior synonyms of Amphibleptula; here we prefer to retain
Remarks. AcicuUtes higginsi sensu Van Soest & Stentoft Amphibleptula in its original scope, based on differences in sponge
(1988) differs from the Schmidt material only in having longer structure and habitus between these nominal genera. Zittel (1878a)
strongyles, but this is clearly intraspecific variability as strongyles described a new rhizomorine lithistid Pomelia schmidti (see Zittel
generally show large variability even within one specimen. 1878a: 62, pI. 1: 4) from original Schmidt material (one speci-
men). Examination of the type material (MZUS) revealed that it is
Distribution a deciduous specimen, displaying des mas and a structure typical of
Amphibleptula Schmidt, but lacking any loose spicules, and for this
Caribbean, Madagascar, Eastern Africa (Kenya Banks), reason we leave it in synonymy with question mark. If any new
Ceylon, Philippines. material eventually comes to hand containing loose spicules show-
ing its identity with A. madrepora, as we suspect (see remarks
below), then Amphibleptula Schmidt should be considered as a
AMPHIBLEPTULA SCHMIDT, 1879 junior synonym of Pomelia Zittel, 1878a. Pisera (1997) reported
the genus Amphibleptula from the late Jurassic. This report was
Synonymy based, however, on the understanding of this genus as proposed by
Van Soest & Stentoft (1988). In the present sense it should
Amphibleptula Schmidt, 1879; 28; Sollas, 1888: 351; Van be regarded as a probable representative of Microscleroderma
Soest & Stentoft, 1988: 61 (partim); Gruber, 1993: 62 (partim). (see below). It is a monospecific genus.
PomeUa Zittel, 1878a: 62 (126).
Description of type species
Type species
Amphibleptula madrepora Schmidt, 1879 (Figs 4-7).
Amphibleptula madrepora Schmidt, 1879: 28 (by monotypy). Synonymy. Amphibleptula madrepora Schmidt, 1879: 28,
pI. 1: 6, pI. 3: 7; Sollas, 1888: 351; Van Soest & Stentoft, 1988: 61,
Definition pI. 8: 3-5, fig. 28. Gruber, 1993: 62, pI. 18: 3-6. Leiodermatium
madrepora Lendenfeld 1903: 146.
Massive conical to cylindrical Scleritodermidae bearing one Material examined. 'Cotype': MCZ 6009 (orig. no. 253).
or several raised oscular areas with numerous openings of exhalant Other material. MNHN DT 2188 slide (from Schmidt MZUS
canals. No ectosomal spicules, microscleres are sigmaspires. P0l73); ZPAL Pf.15/3 (ex ZMA POR without number).
Porifera· Demospongiae· 'Lithistids'· Scleritodermidae 305

Description. Holotype is a small conical sponge with widely 10-12 fLm thick in the case of choanosomal ones. The holotype
spread base, and flattened summit which bears a large oscular area displays no microscleres (deciduous specimens) but the specimens
and smaller elevated lateral oscula; other investigated specimens are from ZMA have strongly spinose sigmaspires 8-12 fLm long. This
irregularly conical with several elevated oscular areas of the same sponge is common around Barbados extending from 100 m depth,
importance, or even clavate with a summit single oscular area. The and has been also reported from Cuba.
type from MCZ is about 2 cm high and 2 cm wide at the base; the
specimens from ZMA are larger and may reach 3 cm high and 3 cm
wide. Oscular area in the holotype is divided into compartments by
numerous vertical septa-like skeletal lamellae (probably the effect
of erosion), which may be up to 14 in number. Upper part of the
cone below the summit is smooth, with the lower part bearing
numerous short processes, 1.5-2.0 mm wide, which open with one
or more vertical canals. Skeleton very dense, composed of massive,
tuberculatedlblunt spinose rhizoclones, 240-380 fLm in size; with
articulation general. Other megascleres are straight oxeas,
500-1200 fLm long, 1-3 fLm thick in the case of ectosomal ones, or

Fig. 4. Amphibleptula madrepora Schmidt, holotype MCZ 6009 (253) Fig. 5. Amphibleptula madrepora Schmidt, ZPAL Pf.15/3 (ex ZMA with-
(scale lOmm). out number), (scale lOmm).

Fig. 6. Amphibleptula madrepora Schmidt, 1879, ZPAL Pf.15/3 (ex ZMA). A, view of the oscular area of the choanosomal rhizoc1one skeleton (scale
I mm). B, details of the choanosomal rhizoc1one skeleton with protruding bundles of oxeas in the oscular area (scale 500 !Lm). C, surface of the choanoso-
mal rhizoclone skeleton of the lateral surface (scale 500 !Lm). D, details of rhizoclone desmas (scale 50 !Lm).
306 Porifera' Demospongiae • 'Lithistids' • ScIeritodermidae

Fig. 7. Amphibleptula madrepora Schmidt. 1879; ZPAL Pf.15/3 (ex ZMA). A-D, sigmaspires (scale 5 j.Lm). E, oxea tips (scale 10 j.Lm).

Remarks. The specimen from MCZ is partly deciduous,

which is probably the reason it lacks microscleres. It also represents
some extreme variability for the species having an atypical and
very regular habitus. There is no doubt that the lamellae observed
at the top of sponge (central oscular area), and influencing the
choice of species name (resembling septa of coral), are artifacts
caused by erosion of the sponge summit; whereas in well-preserved
specimens these are just round openings of the exhalant canals
gathered together in a bundle.

Distribution

Caribbean.

MICROSCLERODERMA KIRKPATRICK, 1903A

Synonymy

Microsclerodenna Kirkpatrick, 1903a: 173. Taprobane

Dendy, 1905: 103; Dendy, 1922b: 7; Wilson, 1925: 468.

Type species

Microsclerodenna hirsutum Kirkpatrick, 1903a: 173 (by

monotypy).

Definition Fig. 8. Microscleroderma hirsutum Kirkpatrick, 1903a, holotype BMNH

1902.5.26.1 (scale 4cm). A, upper surface view. B, lateral view.
Scleritodermidae without special ectosomal spicules;
microscleres are sigmaspires.
spicular components, but in our opinion they are retained as sepa-
Diagnosis rate taxa given their differences in morphology and structure as
well as spiculation. On the other hand, examination of the holotype
Cup-shaped or ear-shaped; ectosomal spicules absent; desmas of Taprobane herdmani Dendy, 1905 proved that there are no basic
are thorny rhizoclones fonning reticulate fibers; microscleres are differences between the genus Microsclerodenna Kirkpatrick,
spinose sigmaspires. 1903a and Taprobane Dendy, 1905, even in morphology - thus
they are synonimized here following the suggestions of Wilson
Remarks (1925), Levi & Levi (1983a) and Van Soest & Stentoft (1988).
Kirkpatrick (l903a) considered sigmaspires as ectosomal spicules
Van Soest & Stentoft (1988) considered this genus a syn- homologous to microstrongyles and rhabds in Scleritodenna and
onym of Amphibleptula Schmidt, 1879, based on similarity oftheir Aciculites, but this view is not supported here, as sigmaspires are
Porifera' Demospongiae • 'Lithistids' • Scleritodennidae 307

Fig. 9. Microsclerodenna hirsutum Kirkpatrick. 1903a. holotype, BMNH 1902.5.26.1. A, hispid outer surface of the sponge (scale I mm). B, transverse
cross section of the wall, outer surface at the top, showing reticulate skeleton and bundles of oxeas (scale I mm). C, choanosomal skeleton of rhizoclones,
upper sponge surface (scale I mm). D, choanosomal skeleton of rhizoc1ones showing clearly reticulate nature, outer sponge surface (scale I mm).
E, longitudinal section of the wall (growth edge at the top) showing regular reticulate choanosomal skeleton and choanosomal bundles of oxeas (scale
2 mm). F, details of desmas of the upper surface (scale 200 11m).

clearly choanosomal, not ectosomal spicules. DOderlein (1884)

described a new species of rhizomorine 'lithistid' Seliscothon
chonelloides from Japan that he attributed to the fossil genus
Seliscothon Zittel. Examination of the holotype of S. chonelloides
(MZUS), revealed that it is a deciduous specimen - displaying no
characters of the fossil genus Seliscothon at all - and resembles it
only in habitus. Details of the structure, type of rhizoclones and
probably presence of sigmaspires in S. chonelloides suggest its
affinities are with Microscleroderma, and this species is therefore
treated as belonging to this genus and not to Mesozoic Seliscothon
Zittel. Four species have been included in Microscleroderma and
all seem to belong here.

Description of type species

Microscleroderma hirsutum Kirkpatrick, 1903a (Figs 8-10). Fig. 10. Microscleroderma hirsutum Kirkpatrick, 1903a; sigmaspires;
Synonymy. Microscleroderma hirsutum Kirkpatrick, 1903a: holotype BMNH 1902.5.25.1 (scale 5 11m).
173, pI. 4: 1. Burton, 1929: 6. Levi & Levi 1989: 47, fig. 15, pI. 2: 3.
Material examined. Holotype: BMNH 1902.5.26.1; densely and uniformly distributed oscules, about 1.2 mm in diame-
BMNH 1902.5.26.1a (3 slides from holotype). ter, located on low conical expansions of the surface. This surface
Description. Cup-shaped sponge, about 17 cm high, 23 cm is also densely covered with oxeas protruding from the
in diameter at the top, with a short and massive pedicle; the wall is choanosome, but shorter than those on the outer surface and never
1.5-2.0 cm thick. Outer surface bears 1 cm wide and up to 1 cm forming bundles. Choanosomal skeleton consists of strongly tuber-
high irregular ridges, which run from the bottom to the top, and culated or thorny and arched rhizoclones organized in reticulate
branch more-or-Iess dichotomously. Entire outer surface densely fibers; both on the surface and in cross-section they form a reticu-
covered with bunches of long oxeas protruding from the late net, with meshes about 130--500 fJ-m in diameter and separated
choanosome. Inner surfaces generally smooth except for numerous by skeletal bridges 10--15 fJ-m wide. In longitudinal section there
308 Porifera· Demospongiae· 'Litbistids'· Scleritodermidae

is also some radial, or fan-like pattern of skeletal fibers visible. the Article 75.6 of the ICZN ('Conservation of Prevailing Usage';
Other choanosomal megascleres are radially arranged and bundled, Anon., 1999) to choose Scleritoderma flabelliforme Sollas as the
thin oxeas which may be 2000-5000 fLm long and up to 30 fLm type species of the genus Scleritoderma, to conserve prevailing
thick, with pointed ends. The bundles of oxeas close to the surface usage of Scleritoderma, and to leave Setidium obtectum as the type
protrude well above the sponge surface in a fan-like form. species of the genus Setidium Schmidt, as demonstrated by Pisera
Microscleres are spinose sigmaspires that are 15-7 fLm long and (1999b) (Application to ICZN, Case No. 3217). This genus resem-
2.2-2.8 fLm thick. This species is known from off the SE coast of bles Setidium Schmidt, 1879, with which it must be very closely
Africa, and South Arabian coast from depths between 38-100 m. related, as indicated by the type of desmas, ectosomal skeleton and
sigmaspire microscleres. It differs in general structure as well as in
Distribution acanthose ectosomal spicules. Five species have been attributed to
Scleritoderma, and all assignations seem to be justified.
Caribbean, Philippines, off SE of South Africa (Natal), South
Arabian coast, Ceylon (Sri Lanka) and New Caledonia. Description of type species

Scleritodermaflabelliforme Sollas, 1888 (Figs 11-13).

SCLERITODERMA SOLLAS, 1888 Synonymy. Scleritoderma flabelliformis Sollas 1888: 316,
pI. 35: 26-50; S.flabelliforme Levi & Levi, 1989: 47, fig. 15, pI. 2: 3.
Synonymy Material examined. Holotype: BMNH 1891.5.4.10 (dry).
Other material: MNHN DCL3207, 3208.
Scleritoderma Sollas, 1888: 315; Lendenfeld, 1903: 141. Van Description. Small, up to 42 mm high, 2-5 mm thick,
Soest & Stentoft, 1988: 59; Gruber, 1993: 63. Not Scleritoderma flabelliform or ear-shaped sponges, or digitate masses, with one or
Schmidt, 1879: 28 (=Setidium Schmidt). both surfaces covered with large oscules, 0.4-0.6 mm in diameter,
irregularly dispersed and raised above the surface. Ectosomal spicules
Type species are acanthorhabds that form a dense tangential layer, 60-80 fLm long,
8.0-9.3 fLm thick in the center, and usually curved. Choanosomal
Scleritodermaflabelliforme Sollas 1888 (here designated). Not skeleton very dense, composed of thorny rhizoclones. Large openings
Scleritoderma paccardi Schmidt, 1879: 28, pI. 2: 3 (=Setidium on the surface of the choanosomal skeleton, corresponding to oscules,
obtectum Schmidt, 1879). Not Scleritoderma paccardi Sollas, measure 470-570 fLm in diameter. Microscleres are spinose sig-
1888: 346 (=Setidium obtectum Schmidt, 1879). maspires 7.8-11 fLm long, 1.4-2.3 fLm thick. This species is known
from Indonesia and Philippines from depths between 96-256m.
Definition Remarks. Examination of Schmidt type material from the
MZUS, as well as a slide from BMNH prepared from the Schmidt
Scleritodermidae with ectosomal acanthostrongyles or material, clearly shows that Scleritoderma paccardi Schmidt is in
acanthorhabds and sigmaspires as microscleres. fact identical with, but only a young, small specimen of Setidium
obtectum Schmidt, as revised by Pisera (1999b).
Diagnosis
Distribution
Polymorphic: flabelliform, encrusting, and massive with
nodes, or cup- to vase-shaped. Desmas are spinose rhizoclones; Caribbean, Madagascar, Philippines, New Caledonia.
ectosomal spicules acanthostrongyles/rhabds forming a dense
tangential layer on the surface; microscleres are sigmaspires. SETIDIUM SCHMIDT, 1879

Remarks Synonymy

The type species of Scleritoderma was never formally desig- Setidium Schmidt, 1879: 30; Sollas, 1888: 350; Pisera,
nated. In theory it should be S. paccardi (by monotypy) as the first 1999b: 473.
described species of the genus. Studies of original Schmidt mate-
rial of Scleritoderma paccardi Schmidt (1879: 28) have shown that

,
it is identical with Setidium obtectum Schmidt (1879: 30). Thus, in ~"
theory, Setidium obtectum should be considered as a junior syn- ... '
:

onym of Scleritoderma paccardi. At the same time, since Sollas's

(1888) publication, the concept of the genus Scleritoderma was
based solely on his Scleritoderma flabelliformis (following from
.
the fact that Scleritoderma paccardi was initially very inadequately
described and illustrated), and in this sense it is clearly a genus
different from Setidium. If we follow strictly the ICZN Principle
of Priority, we should substitute the genus Scleritoderma for
Setidium, and find a new name for Scleritoderma sensu Sollas, as
used since Sollas's time. This action, however, would produce sub-
stantial instability of the nomenclature and subsequent numerous Fig. 11. Scleritoderrna flabelliforrne Sallas BMNH 1891.5.4,10 (after
misunderstandings. As a result, we propose here, in accordance with Sallas 1888, pI. 36: 26--28).
Porifera· Demospongiae· 'Lithistids'· Scleritodermidae 309

Fig. 12. Scleritodermaflabelliforme Sollas BMNH 1891.5.4.10. A. view of the ectosome with crust of curved acanthose microstrongyles (scale 100 j-Lm).
B. surface of choanosomal skeleton showing risen oscules (scale 2 mm). C, details of choanosomal skeleton of rhizoclones (scale 200 j-Lm). D, details of
bifurcating spines of rhizoclones (scale 100 j-Lm).

Fig. 13. Scleritodermaflabelliforme Sollas BMNH 1891.5.4.10. A-F, acanthorhabds (scale 20j-Lm). G, details of acanthorhabd sculpture (scale 5 j-Lm).
H-J, sigmaspires (scale 5 j-Lm).
310 Porifera· Demospongiae· 'Lithistids'· Scleritodermidae

Type species

Setidium obtectum Schmidt, 1879: 30 (by monotypy).

Definition

Scleritodermidae with smooth strongyloxeas as ectosomal

spicules and sigmaspires as microscleres.

Diagnosis

Ectosomal spicules are smooth strongyloxeas with rounded

ends, which show all transitional stages to desmas; choanosomal
spicules are strongly thorny rhizoclones forming a very dense con-
fused skeleton; with bundles of long oxeas protruding from the
choanosome.

Remarks

This genus is very close to Scleritoderma, differing mostly in

having relatively smooth strongyloxeas rather than acanthorhabds
as ectosomal spicules, and having a confused choanosomal skele-
ton. Setidium is so far a monospecific genus.

Description of type species

Setidium obtectum, Schmidt, 1879 (Figs 14--16). Fig. 14. Setidium obtectum Schmidt, holotype MCZ 6462 (scale IOmm).

Fig. 15. Setidium obtectum Schmidt UMML4.S41 MIM-RSMAS Pl14111 in Pisera, I 999b. A, upper surface showing bundles of oxeas protruding from
the choanosome (scale SOD fLm). B, detail of the upper surface showing oscules and tangential layer of ectosomal spicules (scale SOO fLm). C, lower (outer)
surface showing pores protected with oxea palisades (scale I mm). D, surface of the choanosomal skeleton composed of rhizoc1one desmas (scale I mm).
E, details of rhizoc1ones from lower surface of the choanosomal skeleton (scale 200 fLm). F, details of rhizoc1one desmas (scale 100 fLm).
Porifera· Demospongiae· 'Lithistids'· Scleritodermidae 311

Fig. 16. Setidium obtectum Schmidt, UMML4.S41 (old number MIM-RSMAS PI141). A-F, sigmaspires (scale 5 fLm). G-L, ectosomal strongyloxeas
(scale SO fLm).

Synonymy. Setidium obtectum Schmidt, 1879: 30, pI. 1: 9, on the upper surface measure 160-220 f.Lm in diameter (when ecto-
pI. 2: 14; Sollas, 1888: 350; Pisera, 1999b: 473, figs 2-4; somal spicules are present) and corresponding canal openings in
Leiodermatium obtectum Lendenfeld, 1903: 146. Scleritoderma the choanosomal skeleton are 230-300 f.Lm. Those on the lower
paccardi Schmidt 1879: 28, pI. 2: 3; Lendenfeld, 1903: 142; Sollas, surface are slightly larger and measure 200-280 f.Lm in diameter
1888: 346; Thiele, 1900: pI. 3: 11. (when ectosomal spicules present) and corresponding canal open-
Material examined. Holotype: MCZ 6462, MNHN DT ings in the choanosomal skeleton are 230-280 f.Lm. Choanosomal
2193 (slide from MZUS material). Other material: UMML4.541 skeleton consists of strongly branched and thorny rhizoclone des-
(old number MIM-RSMAS PI141), UMML4.539 (old number mas, which measure 200-400 f.Lm long. Generally the choanoso-
MIM-RSMAS G688) and UMMlA.540 (old number MIM- mal skeleton is very dense and hard. Ectosomal spicules as smooth,
RSMAS G1312). Scleritodermapaccardi Schmidt: MZUS P0l75. slightly curved strongyloxeas with rounded tips showing all transi-
Description (based on Pisera, 1999b). Irregular vase- tional types toward the rhizoclone desmas; they are arranged
shaped or turbinate with walls up to 1.5 cm thick, supplied with a tangentially to the sponge surface and may be concentrically
short peduncle; the holotype is about 5 cm wide, 3 cm high, while arranged around the pores, or perpendicular on the conulae surface.
the largest non-typical specimen from Great Inagua is slightly They measure 120-21Of.Lm long and 10-14f.Lm thick when still
larger. Both sponge surfaces are rugose as a result of numerous regular amphioxeas. Bunches of very long and thin, at least 1 cm
irregularly to slightly radially distributed conules, 265 mm high, long, and 15-23 f.Lm thick oxeas protrude from the choanosome;
from which protrude bundles of long oxeas, up to 20 spicules each they are invariably broken, thus their lengths cannot be accurately
(now all broken, but at least 1 cm long); because these conules are measured. Microscleres are thorny sigmaspires, 8-13 f.Lm long, up
slightly oblique to the surface they may form kind of short, radially to 1.7 f.Lm thick. This species is known from the Caribbean from
aligned ridges. Ostia of the lower surface are very densely distrib- depths between 234-516m.
uted, practically touching each other, and placed on top of small
elevations. Pores on the outer surface are surrounded and covered Distribution
by a teepee-like structure formed by delicate oxeas, up to 0.5 mm
long (length of visible part only) protruding from the surface. Pores Caribbean.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Corallistidae Sollas, 1888

Andrzej Pisera1 & Claude Levi 2

I Instytut Paleobiologii, Polska Akademia Nauk, ul. Twarda 51/55,00-818 Warszawa, Poland. (apis@twarda.pan.pl)
2Museum National d'Histoire Naturelle, Laboratoire de Biologie des Invertebres Marins et Malacologie, 57, Rue Cuvier, 75231 Paris
Cedex 05, France. (levi@rnnhn.fr)

Corallistidae (Demospongiae, 'lithistids') contains four extant genera (two described as new) with about 20 nominal (mostly poorly
known) species distributed worldwide. Corallistidae are known at least since the Late Jurassic and have numerous representatives in the
fossil record. Today they live, as do most 'lithistids' , in deeper waters, showing cup, vase, lamellar, club or massive shaped habits. Desmas
are monaxial, strongly tuberculated large dicranoclones which articulate with main body of other dicranoclones (non-terminal articula-
tion). Ectosomal megascleres are smooth or spinose dichotriaenes, sometimes also triaenes; other megascleres may be oxeas.
Microscleres are streptasters (often two types of spirasters), and/or microxeas, microrhabds or microstyles. Differentiation is based mostly
on the type of ectosomal dichotriaenes and microscleres, as well as habitus.
Keywords: Porifera; Demospongiae; 'lithistids'; Corallistidae; Corallistes; Herengeria; Neoschrammeniella gen. nov.; Neophrissospongia
gen. nov.
------------------------------

DEFINITION, DIAGNOSIS, SCOPE gen. nov., Neoschrammeniella gen. nov. Species occur in deep-
water worldwide. About 20 nominal species have been included,
Synonymy most very poorly known and badly in need of revision.

Corallistidae Sollas, 1888: 301 (partim); Topsent, 1894d: 290

(partim); Schrarnmen, 1910: 64 (partim); Hartman, 1982: 644 (par- Remarks
tim); Corallistinidae Schrammen, 1903: 14, 18. Corallistides
Moret, 1926: 103 (partim). Coscinospongiidae Lendenfeld, 1903: We restrict here the family to contain genera with dichotri-
135 (partim); Lendenfeld, 1907: 354; Wilson, 1925: 459. aenes as ectosomal spicules, excluding those having ectosomal
Gignouxiidae de Laubenfels, 1955b: E61. (For an extensive syn- phyllo- or discotriaenes (referred to Macandrewiidae Schrammen,
onymy of fossil corallistid families see chapter on fossillithistids.) 1924a, and which were earlier included in Corallistidae by
Sollas (1888), Lendenfeld (1903, 1907), Schrammen (1910) and
Definition subsequent authors). Lendenfeld (1903) erroneously
included Coscinospongia Bowerbank, 1869 (nomen nudum)
'Lithistid' Demospongiae with dicranoclone choanosomal (=Heterophymia Pomel, 1872) in this family, which is undoubt-
desmas and ectosomal dichotriaenes; among microscleres usually edly a rhizomorine sponge. In effect he also introduced
some kind of streptaster. the unfounded concept of Coscinospongiidae, considering
Corallistidae Sollas as a synonym. For reasons discussed above the
Diagnosis family Coscinospongiidae must be abandoned and Corallistidae
Sollas are considered valid. All except one corallistid (i.e.,
Polymorphic: massive ear, vase, cup or lamellar shaped; Corallistes microstylifer Levi & Levi, 1983, referred here to
choanosomal desmas as dicranoclones, ectosomal spicules as Neophrissospongia gen. nov.) have some kind of streptasters
dichotriaenes and sometimes additionally simple triaenes; among their microscleres, although this issue remains to be
microscleres can be spirasters (often two types), streptastersl clarified. In the fossil record there are several families of lithistids
amphiasters, microxeas, microstyles and microstrongyles. with dicranoclone desmas, corres-ponding to the concept of
Corallistidae Sollas, and which are classified as Dicranocladina
Scope (refer to chapter on fossillithistids). All these fossil families were
differentiated on the basis of their external morphology rather than
Four extant genera are included here in this family: Corallistes on spicule composition, and are therefore considered here to be
Schmidt, 1870, Herengeria Levi & Levi, 1988, Neophrissospongia synonymous with Corallistidae Sollas, 1888.

KEY TO EXTANT GENERA

(1) Ectosomal dichotriaenes smooth on the upper surface ...... .......................... .... ............ ............... ............ ...... .......... ...... ......... .............. 2
Ectosomal dichotriaenes with spines and tubercles on the upper surface, microscleres
streptasters/amphiasters .......................................................................................................................................... Neophrissospongia
(2) Microscleres two types of spirasters with pointed rays ........................................................................................................ Corallistes
Microscleres two types of spirasters (with short blunt and long pointed rays) and smooth anisoxeas ................. Neoschrammeniella
Microscleres spirasters with long and pointed rays, rhabd-like spirasters and spinose, often centrotylote microxeas ....... Herengeria

312
Porifera' Demospongiae· 'Lithistids" Corallistidae 313

CORALUSTES SCHMIDT, 1870 Moret, 1926, which contains similar ectosomal spicules.
De Laubenfels (1955b) erroneously considered Corallistes to be a
Synonymy rhizomorine sponge, and thus incorrectly assigned it to the family
KaJiapsidae de Laubenfels, 1936 ( = Phymaraphiniidae Schrammen,
Corallistes Schmidt, 1870: 22 (partim); Zittel, 1878a: 34,56; 1910). About 20 species have been assigned to this genus, most of
Sollas, 1888: 301 (partim); de Laubenfels, 1955b: E45; Gruber, them are valid but poorly known and badly in need of revision.
1993: 33 (partim); Coscinospongia Lendenfeld, 1903: 135 (par-
tim); Lendenfeld, 1907: 354; Wilson, 1925: 460 (partim). Description of type species

Type species Corallistes typus Schmidt, 1870 (Figs 1-3).

Corallistes typus Schmidt, 1870: 22, pI. 3: 3 (here designated).

Definition

Corallistidae with smooth upper surface of ectosomal dicho-

triaenes and spiraster microsc1eres.

Diagnosis

Cup to vase shaped, sometimes simple or folded plates;

desmas strongly tuberculated dicranoc1ones. Ectosomal spicules
smooth dichotriaenes; microsc1eres are spirasters, usually two
types. Large endosomal oxeas may be present.

Remarks

We propose here to restrict the genus Corallistes to contain

only species with a smooth upper surface of dichotriaenes, with
those having tuberculated and/or spinose upper surface of dichotri-
aenes assigned to a new genus, Neophrissopospongia gen. nov. Fig.t. Corallistes typus Schmidt, 1870. A, ZPAL Pf.B/1 (ex UMML4.543)
This condition is analogous to the fossil genus Phrissospongia (scale lOmm); B, UMML4.S43 (scale lOmm).

Fig.2. Corallistes typus Schmidt, 1870. A, Outer surface with ectosomal dichotriaenes, ZPAL Pf.13/l (ex UMML4.S43) (scale 200 fLm); B, Surface view
of the choanosomal skeleton, ZPAL Pf.13/1 (ex UMML4.S43) (scale bar I mm); C, choanosomal desmas in lateral view ZPAL Pf.13/1 (ex UMML4.S43)
(scale 200 fLm); D, details of the choanosomal desmas in top view, type MZUS PO 172 (scale 200 fLm).
314 Porifera' Demospongiae· 'Lithistids" Corallistidae

Fig. 3. Corallistes typus Schmidt, 1870. A-F subectosomal spirasters, ZPAL Pf.13f1 (ex UMML4.S43) (A-B scales SfLm, C-F scale lOfLm);
0-1 choanosomal spirasters (scale 10 fLm); J-L, ectosoma1 dichotriaenes in various views, holotype MZUS PO 172 (scale 100 fLm); M - ectosomal
dichotriaene, ZPAL Pf.13/1 (ex UMML4.S43) (scale 100 fLm).

Synonymy. Coralistes typus Schmidt, 1870: 22, pi. 3: 3; dichotriaenes which vary strongly in size in various parts of the
Zittel, 1878a: 120; (?) Sollas, 1888: 301, pI. 34: 14-18a; (?) sponge: the cladome is 219-350/-Lm in diameter, while conical
Burton, 1929b: 5; Van Soest & Stentoft, 1988: 54, fig. 24, pI. 7: rhabdome is 273-725 /-Lm long, and up to 67 /-Lm wide at the base,
2-4; Gruber, 1993: 34, pI. 5: 7-8, pI. 6: 1-2. with a blunt tip. Choanosomal desmas are massive dicranoclones
Material examined. 'Cotypes': MZUS PO 172, MZUS PO bearing numerous rather irregularly developed and distributed
209, MNHN DT 2187 (slide from the type). Other material. mushroom-shaped large tubercles. Dicranoclones are 200-300 /-Lm
BMNH 1886.12.31.6-7 (Not C. typus contrary to its label); ZMA in size. Other megascleres are amphioxeas which are over 700 /-Lm
POR 3814, 3828, 5239 - Barbados. UMML4.543, ZPAL Pf.13/1 long, 4 /-Lm thick. Microscleres consist of two types of spirasters
(ex UMML4.543). No lectotype designation is made due to the with pointed microspinose rays. Ones with more numerous, shorter
current uncertainty of 'acceptable' variability within the type series rays (subectosomal) are 19.6-25.7/-Lm long, 12.7-19.1/-Lm wide,
(see below). while the others (choanosomal) have less numerous and longer
Description. Ear or cup shaped sponges; the Caribbean rays, 28-29.4 /-Lm long, 22-24/-Lm wide. This species has been
specimen here investigated (Schmidt's material is known only from reported from 90-640 m depth from Florida, Barbados, coast of
fragments of the skeleton) is about 5 cm high, 4.5 cm wide, with Brazil, and the Natal Coast of South Africa.
walls about 6-7 mm thick, and a rounded margin. Specimens Remarks. The specimen from Pernambuco described by
described by Van Soest & Stentoft (1988) from Barbados are cup to Sollas (1888) may in fact be a different species as it bears relatively
vase shaped and larger, up to 20 cm in diameter and wall 1 cm larger oscules, 0.5-1 mm in diameter, whereas Caribbean material
thick, as well as often having a squarish margin. Specimens has oscules only 0.15-O.4mm in diameter. Specimens from Natal
described by Sollas (1888) are transversely oval plates and display, (Burton, 1929b) have spicules of similar size, although these were
according to Sollas (1888), the presence of large oscules up to not examined and their allocation to the type species is also tenta-
1 mm in diameter, whereas these are only 0.15-O.40mm in other tive. Specimens from Barbados described by Van Soest & Stentoft
material examined from the Caribbean. Thus, the assignment of (1988) have in general less massive spicules, especially dichotri-
Sollas's (1888) specimen to this species is tentative. The investi- aenes and spirasters. Their spicule morphometrics, however,
gated material displays numerous hair-like tiny oxeas protruding are the same as in the type series and material from UMML, thus
from both surfaces. Ectosomal spicules are massive smooth interpreted here as intra-specific variability only.
Porifera· Demospongiae· 'Lithistids'· Corallistidae 315

Distribution Description. Massive ovoid sponges; reaching about 15 mrn

high and 10 mrn wide. slightly flattened at the top. from which pro-
Mediterranean Sea. Florida. Caribbean. Mexican Gulf. northern trude dense bundles of oxeas about 10 mrn long. At the base of one of
to southern coast of Brazil. Southern Africa (Natal Coast). New these bundles short oscular chimneys were observed. On one side. at
Caledonia. Papua New Guinea. about 1/3 its height. there is a lateral flattened outgrowth reaching 7
mm long. 6mm wide and 2mrn thick. bearing inhalant pores.
HERENGERIA LEVI & LEVI, 1988 Ectosomal layer contains dichotriaenes and abundant rnicroscleres.
Dichotriaenes are smooth. often with tips bent downward. measuring
Synonymy 245-500 fLm in diameter (protoclads 50-80 fLm long. deuteroclads
90-200 fLm long) and the rhabds 300-475 fLm long. Large oxeas
Herengeria Levi & Levi. 1988: 250. occurring in bundles reach at least 10 mm in length and are 10-18 fLm
thick. Desmas as typically regular dicranoclones. slightly curved.
Type species usually tripodial. and bearing numerous mushroom-shaped tubercles
which are in turn covered with small. low. densely packed tubercles.
Herengeria auriculata Levi & Levi. 1988: 250. fig . 5. pI. 5: 5 Desmas are 450-800 fLm in size. and with clones 50-75 fLm thick.
(by monotypy). Zygoses are usually tenninal to lateral around the tubercles. Young
desmas are smooth or bear smooth tubercles. The skeleton around
Definition bundles of oxeas from the apical part of the sponge is less dense and
consists of more slender. more arched and less regular. less tubercu-
Massive Corallistidae with lateral inhalant auriculum and lated desmas. and often supplied with long spine-like outgrowths.
bundles of large oxeas protruding from the top. Microscleres are numerous and represented by three different types
of spicules: those on the surface include microspinose rnicroxeas
Diagnosis (often centrotylote). usually slightly curved. 70-100 fLm long.
5-8 fLm thick; the other ectosomal type is a rhabd-like long irregular
Massive in form with inhalant lateral auriculum; endosomal spiraster covered with low massive rnicrospines. measuring
spicules are dicranoclones; ectosomal spicules are smooth dichotri- 32-40 fLm long and 4-5 fLm thick. Choanosomal rnicroscleres are
aenes; rnicroscleres are microxeas and spirasters. Endosomal bun- typically rnicrospinose spirasters with long. more or less pointed
dles of large oxeas protrude from the upper surface. rays. 19-25 fLm long and 15-19 fLm wide. The species occurs at
depths from 505-515m off New Caledonia.
Remarks Remarks. So far monotypic.

Monotypic. Distribution

Description of type species New Caledonia.

Herengeria auriculata Levi & Levi. 1988 (Figs 4-8). NEOPHRISSOSPONGIA GEN. NOV.
Synonymy. Herengeria auriculata Levi & Levi. 1988: 250.
fig. 5. pI. 5: 5. Synonymy
Material examined. Holotype: MNHN DCL 3543.
Paratypes: MNHN DCL 3544. Corallistes Schmidt. 1870 (partim).

Fig. 5. Herengeria auriculata Levi & Levi. 1988. holotype MNHN

Fig. 4. Herengeria auriculata Levi & Levi. 1988. ho1otype. MNHN DCL3543 (top right) and paratypes MNHN DCL 3544 (scale lOmm)
DCL3543 (scale lOmm) (modified after Levi & Levi. 1988). (modified after Levi & Levi. 1988).
316 Porifera· Demospongiae • 'Lithistids' • Corallistidae

Fig. 6. Herengeria auriculata Levi & Levi, 1988, MNHN DCL3544. A-B, ectosomal dichotriaenes (scale 100 ....m); C, details of choanosomal desmas in
lateral view (scale 500 ....m); D, details of strongly modified choanosomal desmas in the central part of the sponge surrounding bunches of oxeas (scale 500 ....m);
E, details of choanosomal desmas in top view, young dicranoclone in the center (scale 200 ....m); F, details of the dicranoclone sculpture (scale 100 .... m).

Type species

Corallistes nolitangere Schmidt, 1870: 23, pI. 3: 6; Zittel,

1878a: 120: pI. 1: 2 (here designated).

Etymology

Referring to its resemblance to the Cretaceous genus

Phrissospongia Moret, 1926.

Definition

CoraIlistidae with spinose or tuberculated upper surface of

ectosomal dichotriaenes and streptasters/amphiasters among
microscleres.

Diagnosis

Shallow cup shaped or ear shaped growth forms. Ectosomal

spicules are dichotriaenes bearing on the upper surface strong spines
Fig. 7. Herengeria auriculata Levi & Levi, 1988, MNHN DCL3544; A-F, or tubercles; endosomal desmas as dicranoclones; microscleres are
subectosomal microxeas (scale 20 .... m); G, details of microxea (scale 5 ....m). streptasters/amphiasters.
Porifera· Demospongiae • 'Lithistids' • Corallistidae 317

Fig. 8. Herengeria auriculata Levi & Levi, 1988, microscleres (scale 10 J.Lm). A-F, choanosomal spirasters; G-J, subectosomal rhabd-like spirasters.

Remarks

This Recent genus closely resembles the Cretaceous genus

Phrissospongia Moret, 1926, which has dichotriaenes with vertical
spines on the upper surface of their clads. Lagneau-Herenger (1962)
considered this and morphologically similar species of dicran-
ocladine sponges to constitute a new family, Phrissospongiadae.
However, following Brimaud & Vachard (1986) we do not support
this view as spicular components of both Recent and fossil taxa are
similar and warrant their inclusion within the Corallistidae. Apart
from the type species we also tentatively include here one Recent
species from New Caledonia described by Levi & Levi (1983) as
C. microstylifer, based on its dichotriaenes. This species differs
from the type species in its microsclere geometry. Separation of
Corallistes species with spined dichotriaenes from those with
smooth dichotriaenes was first suggested by Levi (1991) and
supported by Kelly (2000).

Description of type species

Fig. 9. Neophrissospongia nolitangere (Schmidt, 1870), type MZUS
POlS7 (scale lOmm).
Neophrissospongia nolitangere (Schmidt) (Figs 9-11).
Synonymy. Corallistes nolitangere Schmidt, 1870: 23, pI. 3:
6: Carter, 1873c: 439; Zittel, 1878a: 103, 120, pI. 1: 2; Sollas, be branched. The tips of deuteroclads may be branched as well.
1888: 339; Topsent, 1904: 59, pI. 9: 11; Coscinospongia Choanosomal skeleton relatively loose and consists of slender dicra-
nolitangere Lendenfeld, 1903: 137. noclones bearing strong, mushroom-shaped tubercles with heads
Material examined. 'Types': MZUS P0139 (desmas), divided into low callosities. Desmas measure about 350-500 fLm
MZUS PO 157 (type?); MNHN DT 781 - Fayal, Portugal. MNHN in size. Microscleres are small acanthose streptasters/amphiasters,
DT 198 (slide from type in Strasbourg). sometimes resembling metasters, with short and blunt rays,
Description. Shallow cup or ear-shaped sponges with thick 10-13 fLm long, 8-10 fLm wide. Their morphology varies widely.
walls and rounded margins, attached by a short pedicel, and walls This species occurs off Portugal (depth unknown), Azores and
about 1 cm thick. Oscules numerous and distributed on inner Cape Verde Islands.
surface measuring about 0.5-1 mm in diameter; pores on the outer Remarks. Carter (1873c) and following him Sollas (1888)
surface. Ectosomal spicules are dichotriaenes with rhabdome about and Lendenfeld (1903), reported the presence of microxeas among
450-1200 fLm (usually 650-820 fLm) long and with blunt tip, and microscleres. These were not observed in either the original
cladome 230-310 fLm in diameter. The upper surface of the cladome Schmidt (1870) material or that of Topsent (1904) who also
is covered with numerous 10 fLm high spines or tubercles which may reported them in his specimen from Azores.
318 Porifera· Demospongiae • 'Lithistids' • Corallistidae

Fig.lO. Neophrissospongia nolitangere (Schmidt, 1870), type MZUS P0157. A, Surface view of the desmas of the choanosomal skeleton (scale 500 Il-m);
B, details of desma in lateral view (scale 100 Il-m); C, details of desma sculpture in top view (scale 50 Il-m); D, details of desma articulation (scale 100 Il-m).

Fig. 11. Neophrissospongia nolitangere (Schmidt, 1870), holotype MZUS P0157. A-E, streptasters/amphiasters (scale 5 fLm); F-I, ectosomal dichotri-
aenes in various views (scales F, 200 Il-m; G, 50 fLm; H-I, 100 fLm); J-K, points of ectosomal dichotriaenes (scale 20 Il-m).

Distribution Type species

Portugal, Azores, Cape Verde Islands, New Caledonia. Iouea moreti Levi & Levi, 1988: 248, pI. 1: 4, fig. 4 (by
monotypy).
NEOSCHRAMMENIEUA GEN. NOV.

Synonymy Etymology

Iouea sensu Levi & Levi, 1988: 248 (partim, Recent represen- Named for the close similarity to a related fossil
tative only). genus.
Porifera· Demospongiae • 'Lithistids' • Corallistidae 319

Definition Description. Cylindrical sponges with narrow (about 2 mm

wide) axial cavity, which extends to half the height and located at
Cylindrical Corallistidae with narrow and deep atrial cavity the opening of the conical sponge summit with the osculum, which
and two types of spirasters as microscleres. is closed in most specimens. Sponges are 20-55 mm high and 6-15
mm in diameter. Surface is even with only irregularly distributed
Diagnosis small ostia measuring 68-115 f..lm diameter. Ectosomal megascleres
are smooth dichotriaenes with long conical rhabdome 250-430 f..lm
Cylindrical sponges supplied with a narrow and deep atrial long and have a more or less pointed tip; cladome is 280-550 f..lm
cavity. Ectosomal spicules are smooth dichotriaenes; desmas are in diameter. Outer surface of the choanosomal skeleton with
dicranoclones with 2-4 clones. Outer surface of the choanosomal numerous simple pores leading to straight canals. Choanosomal
skeleton has numerous simple pores leading to straight canals. skeleton consists of strongly tuberculate and thick dicranoclones;
Microscleres consist of two types of spirasters, with short blunt and
long pointed rays.

Remarks

The type species of this genus has been attributed by Levi &
Levi (1988) to the fossil genus Iouea de Laubenfels, 1955b
(=[Phalangiuml Schrammen, 1910, preocc.), which is a junior
synonym of Schrammeniella Breistorfer, 1947 (see Lagneau-
Herenger, 1962: 168). Here we again follow Sollas's (1888)
approach and propose a new name, Neoschrammeniella gen. nov.,
for the Recent form attributed to Iouea, because of significant
differences in the composition of desmas and the uncertainty of
microsclere geometries in the fossil genus.

Description of type species

Neoschrammeniella moreti (Levi & Levi) (Figs 12-15).

Synonymy. Iouea moreti Levi & Levi, 1988: 248, pI. 4: 4,
fig. 4. Fig. 12. Neoschrammeniella moreti (Levi & Levi, 1988) (scale IOmm)
Material examined. Holotype: MNHN DCL 3541. (modified after Levi & Levi, 1988). A, C, paratypes MNHN DCL3542;
Paratypes: MNHN DCL 3542. B, holotype MNHN DCL3541.

Fig. 13. Neoschrammeniella moreti (Levi & Levi, 1988), MNHN DCL3542; A, outer surface with pores and ectosomal dichotriaenes (scale 200 jJ.m);
B, desmas of the choanosomal skeleton (scale 500 jJ.m); C, details of the choanosomal desmas (scale 100 jJ.m); D, details of the strongly modified desmas
on the surface of atrial cavity (scale 500 jJ.m).
320 Porifera' Demospongiae • 'Lithistids' • Corallistidae

Fig. 14. Neoschrammeniella moreti Levi & Levi, 1988, MNHN DCL3542 (scales A-D, 100 fLm); A, ectosomal triaene, B-D, ectosomal dichotriaenes
in various views; E-G, choanosomal microanisoxeas (scale 50 fLm).

Fig. 15. Neoschrammeniella moreti (Levi & Levi, 1988) MNHN DCL3542; A-D, ectosomal spirasters (scale 5 fLm); E-G, choanosomal spirasters
(scale 10 fLm).

tubercles are mushroom-shaped and slightly subdivided into Remarks. The type species was initially referred by Levi &
smaller very low callosities. The resulting skeleton is very dense. Levi (1988) to Iouea de Laubenfels, 1955b (=[Phalangium]
The dicranoclones are very irregular and form an even more dense Schrammen, 1903), based on similarities with this Cretaceous
skeleton on the surface of the axial cavity. Other choanosomal genus. However, these fossil sponges have no preserved microscleres
megascleres are long oxeas measuring 900-1400/-Lm long and and thus the type species of this new extant genus displays only par-
15-20/-Lm thick; there are also smaller anisoxeas which measure tial similarity to the fossil forms. Consequently, we have decided to
250-400 /-Lm long and 4-7 /-Lm thick. Microscleres are 2 types of follow Sollas (1888) in erecting a new taxon for Recent sponges,
spirasters; those occurring in great numbers in the ectosome are but we underline their close similarities to fossil genera by employ-
very irregular, have blunt short rays which are strongly spinose, ing the prefix Neo- to the stem of the related fossil genus name.
and measure 19-30/-Lm long and 8-14/-Lm wide. Within the
choanosome there are numerous spirasters with long, pointed and Distribution
spinose rays measuring 20-36/-Lm long and 16-20/-Lm wide.
Known only from the New Caledonia region at 435-570 m depth. New Caledonia.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Pleromidae Sollas, 1888

Andrzej Pisera1 & Claude Levi2

1 Instytut Paleobiologii, Polska Akademia Nauk, ul. Twarda 51155, 00-818 Warszawa, Poland. (apis@twarda.pan.pl)
2Museum National d'Histoire Naturelle, Laboratoire de Biologie des Invertebres Marins et Malacologie, 57, Rue Cuvier, 75231 Paris
Cedex 05, France. (levi@rnnhn.fr)

Pleromidae Sollas (Demospongiae, 'lithistids'), including P1eromatidae Lendenfeld and Megamorina Zittel, contains two valid extant gen-
era, with four species found in deep water, and known predominantly from the Pacific. These are polymorphic massive irregular to thick
auricular and conical sponges. Fossil pleromids are reported with certainty from the Late Jurassic, with numerous representatives in the
Late Cretaceous and also Jurassic. Desmas are arched megaclones, usually smooth. Other megascleres include large choanosomal and ecto-
somal oxeas, and ectosomal dichotriaenes and anatriaenes. Microscleres are spirasters or amphiasters, usually of two types, and
microspined microxeas and/or styles. Genera are differentiated on the basis of presence or absence of anatriaenes and type of microscleres.
Keywords: Porifera; Demospongiae; 'lithistids'; Pleromidae; Pleroma; Anaderma.

DEFINITION, DIAGNOSIS, SCOPE spirasters or amphiasters, microxeas and/or styles; large oxeas in
choanosome and protruding from the surface.
Synonymy
Scope
Pleromidae Sollas, 1888: 312; Topsent, 1894d: 290; Wilson,
1925: 461; de Laubenfels, 1955b: E50; Hartman, 1982: 644; Levi, Three Recent nominal genera were attributed to Pleromidae,
1991: 80. Pleromatidae Lendenfeld, 1903: 140. Pleromides Moret, but only two of these, Pleroma Sollas, 1888 and Anaderma Levi &
1926: 118. Megamorina Zittel, 1878a: 99, 130 (partim); de Levi, 1983a are considered here to be valid and represented by four
Laubenfels, 1955b: E50 (partim). species. All species are reportedly common in the Pacific except
Schmidt's (1870) deciduous specimen described from Cuba as
Definition Lyidium (and synonymized here with Pleroma).

'Lithistid' Demospongiae with megaclone choanosomal Remarks

desmas and dichotriaenes among ectosomal megascleres.
Hartman (1982) also included Costifer Wilson, 1925 in
Diagnosis Pleromidae, which has heloclones, not megaclones, as choanosomal
desmas. For this reason Costifer is here excluded from Pleromidae
Polymorphic; extant forms massive irregular or conical, thick and referred to Isoraphiniidae Schrarnmen, 1924, following Reid
lamellar to auriculate. Desmas are megac10nes (usually smooth); (1970) and Levi (1991). Pleromidae has numerous undoubted pre-
ectosomal spicules are dichotriaenes, sometimes additionally also served fossil representatives known from the late Jurassic, while the
anatriaenes protruding from subectosomal region; microscleres are first megaclones appear in the Middle Triassic (Wiedenmayer, 1994).

KEY TO GENERA

(1) Megascleres include ectosomal anatriaenes ......................................................................................................................... Anaderma

Without ectosomal anatriaenes ................................................................................................................................................ Pieroma

ANADERMA LEVI & LEVI, 1983 Definition

Synonymy P1eromidae with anatriaenes among ectosomal spicules;

microscleres include styles and amphiasters.
Anaderma Levi & Levi, 1983a: 107.
Diagnosis
Type species
Massive reversed conical sponges with shallow depression at the
Anaderma rancureli Levi & Levi, 1983a: 107 (by monotypy). top. Desmas as smooth, slender and poorly branched megaclones

321
322 Porifera· Demospongiae • 'Lithistids' • Pleromidae

displaying weak zygosis; outermost layer of megac10nes with long

spine directed outward. Ectosomal spicules are dichotriaenes and
anatriaenes. Other megasc1eres are oxeas of two size categories.
Microsc1eres are streptasters/amphiasters, microxeas and microstyles.

Description of type species

Anaderma rancureli Levi & Levi, 1983a (Figs 1-4).

Synonymy

Anaderma rancureli Levi & Levi, 1983a: 107, pI. 3: 4-5,

pI. 13: 2, 3, fig. 3.
Material examined. Holotype: MNHN DCL 2767.
Paratype: MNHN DCL 2768 - New Caledonia. Fig. 1. Anaderma rancureli Levi & Levi MNHN DeL 2767 (holotype).
Description. Massive sponge in the form of an inverted A, lateral view. B,upper surface view (scales 2 em) (modified after Levi &
Levi, 1983a).
cone, reaching 90 mm high, 90 mm in diameter at the top. Upper

Fig. 2. Anaderma rancureli Levi & Levi MNHN DeL 2767 (holotype). A, section perpendicular to the outer surface showing choanosome, subectosomal
spaces and bunches of anatriaenes and oxeas (scale 1 mm). B, details of the same section (note fan-like organization of anatriaenes and oxeas protruding
from the subectosome) (scale 500 f,Lm). e, outer surface of choanosomal skeleton (note canal openings) (scale 2 mm). D, details of the outer surface of the
choanosomal skeleton (note long spines of megaclones of the external layer) (scale I mm). E, interior of the choanosomal skeleton (surface to the right)
(note bunches of long oxeas) (scale 1 mm). P, details of the articulation of megaclones (scale 200 f,Lm).
Porifera· Demospongiae • 'Lithistids' • Pleromidae 323

Fig. 3. Anaderma rancureli Levi & Levi MNHN DCL 2767 (holotype). A-C, ectosomal dichotriaenes (scales: A, 200 f.Lm; B-C, 500 f.Lm). D, large
choanosomal oxea (scale 50 f.Lm). E, small ectosomal oxea accompanying anatriaenes (scale 20/-Lm). F-G, ectosomal anatriaenes (scales: F, 200 /-Lm,
G, 50 /-Lm). H-J, megacione desmas from the interior of the skeleton (scale 500 f.Lm).

Fig. 4. Anaderma rancureli Levi & Levi MNHN DCL 2767 (holotype). A-F, variability of streptasters/amphiasters (scale 5 f.Lm). G-I, variability of styles
(scale 20 f.Lm). K-L, variability of microxeas (scale 20 /-Lm).
324 Porifera· Demospongiae· 'Lithistids'· Pleromidae

surface with concavity about 25 mm deep, with dispersed oscules microspinose microxeas and two types of microspinose spirasters:
about 2 mm in diameter. Outer surface of the sponge is covered by ectosomal one which has short rays, and choanosomal one with
1-3 mm thick layer without desmas which is composed of densely long slender rays.
packed radial bunches of long oxeas and anatriaenes, which are
deeply rooted in the choanosome. These oxeas (often anisoxeas)
are 1.2-1.5 mm long, about 20--30 f.Lm thick and with pointed ends; Remarks
the anatriaenes are 700--900 f.Lm long and with cladome about
90--130 f.Lm in diameter. Dennal dichotriaenes are 1.2-2.4 mm Schmidt (1870) described Lyidium torquilla based on a dead
long, with cladome up to about 780 f.Lm in diameter (protoclad specimen displaying only megaclone desmas, making its exact
40--50 f.Lm long, deuteroclad 250--380 f.Lm long). The dennal mem- determination very difficult. However, megaclones in Lyidium are
brane occurring below is marked by sparse dichotriaenes and very similar (if not identical) with megaclones of Pleroma turbina-
numerous microscleres which occur also in great numbers in the tum Sollas, 1888. Based on these similarities, and because Lyidium
subdennal region, characterized by large interconnected spaces was never properly defined, it is treated here as a synonym of
and rare desmas. Choanosomal skeleton consists of megaclone des- Pleroma. Sollas (1888) himself considered the genus incertae
mas which display rather weak zygosis producing a delicate skele- sedis. Pleroma has a fossil representative known from the Tertiary
ton. At the growing edge and surface particular megaclones have of South Western Australia.
a long spine projecting outside. Megaclone desmas are slender and
poorly ramified with long zygomes at the tips; they are 1.0--1.5 mm
long, 0.07-0.1 mm thick. The choanosomal skeleton displays, apart Description of type species
from rather large meshes, slightly larger oval openings which rep-
resents canals entrances. Apart from desmas there are numerous Pleroma turbinatum Sollas, 1888 (Figs 5-7).
radially oriented, long oxeas in the choanosome, but their length is
difficult to measure; those seen in preparations are up to 3 mm long
and with blunt ends. Microscleres are curved microxeas, often cen- Synonymy
trotylote, 120--150f.Lm long, 8-11 f.Lm thick, with short spines.
There are also massive styles bearing the same sculpture, 105- Pleroma turbinatum Sollas, 1888: 312, pI. 33; Lendenfeld,
125 f.Lm long, 10--14 f.Lm thick. Streptasters/amphiasters have a 1903: 140. de Laubenfels, 1955b: E50, fig. 31: 3; Levi & Levi
wide size range, 12-18 f.Lm long, 9-13 f.Lm wide, with rays varying 1983a: 105, pI. 2: 5-8, textfig. 2. (Not pI. 1: 1-2 (=Corallistes.)
from pointed to blunt. The holotype was collected at a depth of Material examined. Holotype: BMNH 1899.1.1.98A- sub-
390-400 m near New Caledonia. sample in alcohol and slide, 'Challenger' Expedition, stn. 173, Fiji.
Remarks. Levi & Levi (1983a) described next to the Other material. MNHN DCL 2766 - Havannah, New Caledonia.
holotype two other specimens of this species which differ slightly Lyidium torquilla Schmidt: MNHN DT 2196, DCL 1243 - mega-
from it in the shape and details of spiculation. clone desmas in preparations from the Schmidt original type
(MZUS POl64, deciduous fragment).
Distribution Description. Small conical (holotype) to thick lamellate-
auricular sponge with walls up to 10 mm thick; in the conical fonn
Known only from the New Caledonia region. the upper surface is slightly convex, with rounded margins, oscula
small occurring only on the upper surface (in the 'Challenger' spec-
imen); pores simple, distributed on the sponge sides, 83-100 f.Lm in
PLEROMA SOLLAS, 1888 diameter. Desmas large, smooth megaclones up to 528-700 f.Lm
long, 50--100 f.Lm thick; articulation realized in such a fashion that
Synonymy the saddle-shaped tips of rays of one desma clasp the arched main
body of the other desma. Ectosomal spicules are dichotriaenes with
Pleroma Sollas 1888: 312; Lendenfeld, 1903: 140; Lendenfeld, long and conical and sharply pointed rhabdome, up to 1-2 mm long,
1955b: E50; Gruber, 1993: 30. Lyidium Schmidt, 1870: 84 (type
species L. torquilla Schmidt 1870: 84); Zittel, 1878a: 99, 132;
Lendenfeld, 1903: 144.

Type species

Pleroma turbinatum Sollas, 1888: 312 (by monotypy).

Definition

Pleromidae in which microscleres are microxeas and spirasters,

without ectosomal anatriaenes.

Diagnosis

Irregular massive or thick lamellate to auricular in shape.

Desmas are massive smooth megaclones. Microscleres are Fig.5. Pleroma turbinatum Sollas, MNHN DeL 2766 (scale 1em).
Porifera· Demospongiae • 'Lithistids' • Pleromidae 325

Fig. 6. Pleroma turbinatum Sollas, MNHN DeL 2766. A, surface showing ectosomal dichotriaenes (scale 500 J.Lm). B, outer surface of the choanosomal
skeleton (note numerous canal openings) (scale 100 J.Lm). e, details of the choanosomal skeleton (note numerous thin oxeas (broken) protruding from the
choanosome) (scale 500 J.Lm). D, details of desma articulation (scale 100 J.Lm).

Fig. 7. Pleroma turbinatum Sollas, MNHN DeL 2766. A-F, variability of choanosomal spirasters with long pointed rays (scale 5 J.Lm). G-J, variability of
spirasters with short rounded rays (scale 5 J.Lm). K-L, spinose microxeas (scale 5 J.Lm). M-N, ectosomal dichotriaenes (scale 200 J.Lm).
326 Porifera· Demospongiae· 'Lithistids'· Pleromidae

cladome reaching about 400---600 IJ-m in diameter; other megascle- described by Levi & Levi (1983a), notably in spirasters and oxeas,
res represented by somewhat sinuous oxeas (occurring chiefly in the the shape and size of sponges, may be due to the fact that the holo-
choanosome, but sometimes protruding from it), up to 1350-1700 type is young (very small). Only one type of spiraster was reported
IJ-m long, 3-81J-m thick. Microscleres are fusiform spinose rnicrox- by Sollas (1888) and Levi & Levi (1983a), with the other probably
eas, often centrotylote, sharply pointed and measuring 100-210 IJ-m overlooked by both authors. The type species was collected from
long, 3-51J-m thick. Other rnicrosclere types are spinose spiraster of 573-576 m depth from Fiji, on a coral mud bottom.
two types: one has slender, long and pointed rays, 14-251J-m long,
10-16IJ-m thick; the other is slightly smaller, 11-161J-m long, 5-9 Distribution
IJ-m thick, with short, massive and bluntly ended rays.
Remarks. Three species are attributed to this genus. Size Pleroma is known from Fiji and New Caledonia, with its
differences observed between the holotype and specimens synonym Lyidium Schmidt recorded from Cuba.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Theonellidae Lendenfeld, 1903

Andrzej Pisera1 & Claude Levi2

1 Instytut
Paleobiologii, Polska Akademia Nauk, ul. Twarda 51/55, 00-818 Warszawa, Poland. (apis@twarda.pan.pl)
2 Museum National d'Histoire Naturelle, Laboratoire de Biologie des Invertebres Marins et Malacologie, 57, Rue Cuvier, 75231 Paris
Cedex 05, France. (levi@mnhn.fr)

Theonellidae Lendenfeld (Demospongiae, 'lithistids') contains five genera and about 50 nominal species (most poorly known) occurring
both in shallow and deep waters worldwide. They are polymorphic, ranging from thin encrusting to massive irregular, cylindrical, erect-
branching, cup-shaped or clavate, with or without axial cavity. Theonellidae are characterized by ectosomal phyllotriaenes and/or disco-
triaenes, choanosomal tetraclones and microscleres as acanthorhabds to pseudospherasters and/or microxeas, and/or streptasters/
amphiasters. Genera are differentiated based on the type of ectosomal spicules and microscleres, as well as habitus. This family is basi-
cally Recent, but it corresponds to the family Discodermiidae Schrammen of paleontologists, which has numerous genera known since the
Cretaceous, and is here considered as a junior synonym of Theonellidae.
Keywords: Porifera; Demospongiae; 'lithsitids'; Theonellidae; Discodermia; Manihinea; Racodiscula; SiUquariaspongia; Theonella.

DEFINITION, DIAGNOSIS, SCOPE Scope

Synonymy There are five valid extant genera attributed here to

Theonellidae: Discodermia du Bocage, 1870, Manihinea Pulitzer-
Theonellidae Lendenfeld, 1903: 125 (partim); Lendenfeld, Finali, 1993, Racodiscula Zittel, 1878a, Siliquariaspongia Hoshino,
1907: 343 (partim); Wilson, 1925: 447; Hartman, 1982: 644 (par- 1981 b and Theonella Gray, 1868b.
tim); Hooper & Wiedenmayer, 1994: 438 (partim); Levi, 1991: 79.
Discoderrninae Schrantmen, 1910: 97. Discoderrniidae Schrantmen,
1924a: 37,48; de Laubenfels, 1955b: E58 (partim). Remarks

Lendenfeld (1903, 1907) included in this family such

Definition genera as: Kaliapsis Bowerbank, having triders as desmas, which
is here considered Phymaraphiniidae Schrammen, 1924a;
'Lithistid' Demospongiae with tetraclone choanosomal Sulcastrella Schmidt, 1870, which is here considered
desmas and phyllo- to discotriaenes as ectosomal spicules. Desmanthidae Topsent; and Jereopsis Schmidt (=Neosiphonia
Sollas, 1888) which has dichotriaenes as ectosomal spicules and is
here included in Phymatellidae Schrammen, 1910. Hooper &
Diagnosis Wiedenmayer (1994) considered Phymatellidae Schrammen,
1910 and Kaliapsidae de Laubenfels, 1936a as synonyms of
Polymorphic; choanosomal spicules are tetraclone desmas; Theonellidae Lendenfeld, but this view is not supported here
ectosomal spicules are phyllo- to discotriaenes; microscleres are because Phymatellidae have exclusively dichotriaenes as ectoso-
spinose microrhabds and/or microxeas, and streptasters, and/ mal desmas, while Theonellidae are characterized exclusively
or pseudospherasters which are believed to be derivatives of by having phyllo- to discotriaenes as ectosomal spicules.
acanthorhabds or amphiasters; large choanosomal oxeas may be Conversely, Kaliapsidae is a junior synonym of Phymaraphiniidae
present. Schrammen, 1910.

KEY TO EXTANT GENERA

(1) Ectosomal spicules exclusively discotriaenes, microscleres are acanthorhabds and acanthoxeas ...................................................... 2
Ectosomal spicules include phyllotriaenes .......................................................................................................................................... 3
(2) Ectosomal spicules discotriaenes, desmas are typical tetraclones, microscleres are acanthoxeas and
acanthorhabds ..... ............... ........... ...... .......... ...................... ...................... .... ........... ........... ...... ..... ...... .......................... ... Discodermia
Ectosomal spicules discotriaenes, desmas are tetralophs, microscleres are acanthostrongyles ............................... Siliquariaspongia
(3) Ectosomal spicules phyllotriaenes, microscleres are pseudospherasters ............................................................................ Manihinea
Ectosomal phyllotriaenes to discotriaenes, microscleres are acanthorhabds ........................................................................ Theonella
Ectosomal disco- to phyllotriaenes, microscleres are streptasters and acanthorhabds ...................................................... Racodiscula

327
328 Porifera· Demospongiae • 'Lithistids' • Theonellidae

DISCODERMIA DU BOCAGE, 1870

Synonymy

Discodermia du Bocage, 1870: pi. 11: 1; Zittel, 1878a: 37, 87;

Sollas, 1888: 292 (partim); Lendenfeld, 1903: 128; Dendy, 1905:
99; Dendy, 1922b: 6 (partim); Wilson, 1925: 455; de Laubenfels,
1955b: E58; Van Soest & Stentoft 1988: 50; Levi & Levi, 1989: 51;
Gruber, 1993: 41; Hooper & Wiedenmayer, 1994: 438 (partim).

Type species

Dactylocalyx polydiscus Bowerbank, 1869b: 96 (here

designated).

Definition
Fig. 1. Discodermia polydiscus (Bowerbank, 1869b), holotype BMNH
Theonellidae with exclusively discotriaenes as ectosomal 40.10.23.12 (scale 10 mm).
spicules; microscleres acanthoxeas and acanthorhabds.

Fig. 2. Discodermia polydiscus (Bowerbank, 1869b), holotype BMNH 40.10.23.12. A, view of the outer surface with ectosomal discotriaenes (scale 200
/-Lm). B, detached crust of ectosomal discotriaenes from below with numerous microscleres (scale 200 /-Lm). C, view of the choanosome showing
choanosomal oxeas (scale 200 /-Lm). D, details of choanosomal desma articulation (scale 100 /-Lm). E, choanosomal skeleton of tetraclone desmas (scale
400 /-Lm). F, details of tetraclones (scale 200 /-Lm).
Porifera· Demospongiae • 'Lithistids' • Theonellidae 329

Diagnosis Description of type species

Polymorphic: massive irregular, cup-shaped, branched, cylin- Discodermia polydiscus (Bowerbank, 1869b) (Figs 1-5).
drical. Ectosomal spicules exclusively discotriaenes; choanosomal Synonymy. Dactylocalyx polydiscus Bowerbank, 1869b: 96,
spicules are regular tetraclone desmas; microscleres are acanthoxeas pI. 6: 9-14. Discodermia polydiscus Zittel, 1878a: 87, pI. 1: 7 (par-
and acanthorhabds. tim); Sollas, 1888: 330 (partim); Van Soest & Stentoft, 1988: 50, pI. 7:
1, fig. 22 (partim). Non Discodermia polydiscus Barboza du Bocage,
Remarks 1870: 15, pI. 11: 1. Non Discodermia polydiscus Schmidt, 1879: 22.
Non Discodermia polydiscus Pouliquen, 1972: 749: pI. 9: 1, pI. 10: C.
Many authors have included species with a mixture of disco- Material examined. Holotype: BMNH 40.10.23.12 -
triaenes and phyllotriaenes in the genus Discodermia, while here St. Vincent Island, Caribbean. Other material. ZMA POR
we consider them as representatives of Theonella, following Levi 5256-5258 - Barbados.
& Levi (1989), while the genus Discodermia has only discotri- Description. Small irregular mushroom shaped, 25 X 20 mm
aenes and two sizes of microscleres. About 30 species have been large, 18mm high sponge with strongly concave upper side, very
attributed at various times to this genus, many however, belong to short stem and slightly expanded attachment base. Upper side with
other genera, especially Theonella. slightly raised, widely separated and irregularly distributed oscula,
which are oval and measure 0.5-O.8mm across. Lower side with
smaller round pores, 0.20-0.25 mm in diameter, widely separated and
irregularly distributed over the whole surface. Except for these open-
ings both surfaces are smooth and covered with a dense crust of ecto-
somal round to oval discotriaenes, 250-350 JLm in diameter, and with
short conical rhabd measuring 87-108JLm long. Holotype is
encrusted in places with bryozoans, serpulid tubes and thin crusts of
other demosponges. Choanosomal desmas are regular massive tetra-
clones with strongly branched and tuberculated zygomes and smooth
rays; articulation of the desmas is mostly terminal. Desmas are about
300-450 JLm in size, 100--110 JLm thick, and form a relatively dense
skeleton with meshes about 500--600 JLm wide. Other choanosomal
megascleres are long and thin oxeas (exact length unknown as always
broken). Microscleres are of two types: the larger ones are slender
fusiform and slightly curved or bent acanthoxeas (spines are hook-
like), about 38-59JLm long, 2.4-4JLm thick; the smaller ones are
fusiform, massive acanthorhabds, 15-22JLm long, 2--4.5 JLm thick.

Remarks

There are numerous reports of D. polydiscus, especially from

the Atlantic and Mediterranean, which are based on generic rather
than specific characters. As a consequence the present understand-
ing of D. polydiscus is that it represents either a different species
Fig. 3. Discodermia polydiscus (Bowerbank, 1869b), holotype BMNH
40.10.23.12, microscleres. A-F, microxeas (scale 10 fLm). G, details of (as in the case of the Mediterranean record of Pouliquen, 1972), or a
microxea sculpture (scale 5 fLm). mixture of various species (as is probably the case in D. polydiscus

Fig. 4, Discodermia polydiscus (Bowerbank, 1869b), holotype BMNH

40.10.23.12, microsc1eres. A-E, fusiform acanthorhabds (scale 5 fLm). Fig. 5. Manihinea conferta Pulitzer-Finali, 1993, holotype MSNG 48296
F, details of rhabd sculpture (scale 2 fLm). (scale 20 mm). A, lateral view. B, longitudinal section view.
330 Porifera· Demospongiae • 'Lithistids' • Theonellidae

from Barbados as reported by Van Soest & Stentoft, 1988). These Diagnosis
authors also synonymized several other species with D. polydiscus,
which we consider is unjustified. Clavate to hemispherical sponges with deep and narrow
axial cavity; ectosomal spicules are exclusively phyllotriaenes;
Distribution desmas as tetraclones; microscleres are pseudospherasters with
massive spinose rays (most probably modified amphiasters or
Worldwide, but especially common in the Caribbean and rhabds).
Atlantic, other occurrences are from the Philippines, Mediterranean,
Indian Ocean. Remarks

Kelly (2000) attributed this genus to Phymaraphiniidae, but

MANIHINEA PULITZER-FINAL!, 1993 this is clearly a mistake because it has typical tetraclones, not
triders, as choanosomal desmas. Pseudospheraster microscleres
Synonymy observed in this genus are most probably modified amphiasters (or
rhabds) in-filled with silica. Apart from the type species this genus
Manihinea Pulitzer-Finali, 1993: 262. probably contains only two species, both still undescribed: one
from the Caribbean (Pisera, in prep.) and one from Micronesia
Type species (Kelly, 2000).

Manihinea conferta Pulitzer-Finali, 1993: 263 (by original Description of type species
designation).
Manihinea conferta Pulitzer Finali, 1993 (Figs 6-7).
Definition Synonymy. Manihinea conferta Pulitzer-Finali, 1993: 263,
fig. 8.
Theonellidae with exclusively phyllotriaenes as ectosomal Material examined. Holotype: MSNG 48296 - North
spicules and pseudospherasters as the only microscleres. Kenya Banks, East Africa, Indian Ocean, 110-115 m depth.

Fig. 6. Manihinea conferta Pulitzer-Finali, 1993, holotype MSNG 48296. A, view of the outer surface with ectosomal phyllotriaenes (scale 200 fLm) .
B, details of A showing dense crust of ectosomal microscleres (scale 50 fLm) . C, view of the choanosome showing bundles of styles (scale 200 fLm) .
D, outer surface of choanosomal skeleton (scale I mm). E, details of choanosomal skeleton of tetraclone desmas (scale 200 fLm) . F, articulation of tetra-
clone desmas (scale 50 fLm).
Porifera· Demospongiae • 'Lithistids' • Theonellidae 331

Fig. 7. Manihinea conferta Pulitzer-Finali. A-C, ectosomal phyllotriaenes (scale 200 j-Lm). D-E, styles (scale 20 j-Lm). F-K, pseudospherasters (scale 2 j-Lm).

Description. Medium sized clavate sponge, about Wcm Distribution

high and 5.5 cm wide, with a short and narrow branch, about 3 cm
long, 2 cm wide, in the upper portion. Axial cavity about 6 cm deep, Western Indian Ocean (East Africa), Micronesia.
narrowing toward the interior, and about 1.5 cm wide at the oscu-
lum. The surface of the axial cavity is covered with large, 1 X 2 mm
long, densely spaced canal openings. These openings lead to the RACODISCULA ZITTEL, 1878
canal system cutting radially (which are arched toward the upper
surface), and the canal system running toward the outer surface. Synonymy
Other similar canal systems have openings at the outer surface and
run in the opposite direction, toward the axial cavity surface, and Racodiscula Zittel, 1878a: 37, 87 (151); Sollas, 1888: 331
canals in it are more-or-less perpendicular to the previous ones. (partim); Lendenfeld, 1903: 131; de Laubenfels, 1955b: E58; Van
They open on the axial cavity with large (approximately 1-2mm Soest & Stentoft, 1988: 52.
diameter) openings. Ectosomal spicules are smooth phyllotriaenes
with poorly branched and very narrow clads, 360-490 fA.m in diame- Type species
ter, rhabds about 170-220 fA.m long. Between them occur a dense
crust of microscleres. Choanosomal desmas are typical tetraclones Corallistes polydiscus Schmidt, 1870, pI. 3: 8 (by monotypy).
which measure 290-460 fA.m in size and which are sparsely
tuberculated on rays, and strongly tuberculated at their tips (articula- Diagnosis
tion). The surface of the choanosomal skeleton displays numerous
round to oval openings of the choanosomal canals, 0.5-1 mm in Massive pillow to subspherical 'lithistid' sponges with phyllo-
diameter. Other megascleres are choanosomal styles with rounded triaenes to discotriaenes as ectosomal spicules and microscleres are
lower tips, 700-1300 X 7.5-11 fA.m, which may occur in bundles. streptasters/amphiasters with long slender rays and two types of
Microscleres are irregular pseudospherasters with short, massive acanthorhabds which may vary to pseudospherasters.
spinose rays (most probably infilled amphiasters or rhabds),
8.25-9.30 fA.m in diameter. Remarks
Remarks. Microscleres in this species were interpreted by
Pulitzer-Finali (1993) as spherasters. Their irregularity clearly sug- This genus was proposed by Zittel (1878a) based on
gests, however, that these are pseudospherasters, for example amphi- Schmidt's original material, and the only skeletal character distin-
asters infilled with silica as suggested by Kelly (2000), or modified guishing it from other genera was the presence of phyllotriaenes -
rhabds. No styles were reported by Pulitzer-Finali (1993), but our Zittel (1878a) illustrated only a phyllotriaene of Racodiscula sp.
preparations leave no doubt about their attribution of this species. from Philippines. He referred to Carter (1873c) as the author of the
332 Porifera· Demospongiae • 'Lithistids' • Theonellidae

species R. asteroides, which is not true. Carter (1873c: 441) neither 1988: 52. Non Racodiscula asteroides Carter, 1885 (probably
described nor illustrated this species, but only referred to Schmidt's Discodermia).
slides in BMNH, misspelling their names at the same time (aster- Material examined. Neotype: ZPAL Pf.1511 (ex ZMA
oides not asterodiscus as they are labeled). Subsequently, Schmidt POR5264). Other material. BMNH 70.5.3.7 - Corallistes aster-
(1880) considered Carter (1873c) to be the author of the genus odiscus Horida (Racodiscula asterodiscus O. Schdt.; Corallistes
Racodiscula (which is clearly erroneous), and for this reason de asterodiscus Sdt. var. of Discodermia polydiscus Boc. Horida).
Laubenfels (1955b: E58) designated Corallistes asteroides Carter, Not BMNH 70.5.3.8 - Corallistes asterodiscus n. sp. var. Cuba
1873c as the type species of the genus Racodiscula Zittel. This des- (Corallistes asterodiscus Sdt. var. of Discodermia polydiscus Boc.,
ignation is incorrect, however, as it does not conform with the Cuba). ZMA POR 5262-64 - Racodiscula asteroides, Barbados.
facts, as the first formal description was given by Schmidt (1870) Description. Massive pillow to subspherical sponge with mul-
under the name Corallistes polydiscus, and the name Racodiscula tiple oscules located at the shallow depression on the top, supplied
was proposed by Zittel (1878a). The original material of this with an iris-like diaphragm. On the lateral surface occur slightly
species included two clearly different specimens: one from Cuba depressed inhalant areas. Ectosomal spicules are phyllo- to discotri-
and one from Horida, and according to Schmidt (1880) it was con- aenes; on the inhalant surfaces phyllotriaenes with slender rays are
sidered by him to be a synonym of Discodermia polydiscus du present, up to 420/Lm in diameter; in non-inhalant areas there are
Bocage. The slides from BMNH London are both designated in disco- to phyllotriaenes, with wide rays, which measure 250--380 /Lm
Schmidt's handwriting as Corallistes asterodiscus, (not asteroides), in diameter. Pores on lateral inhalant surfaces measure 500--600 /Lm
one of which is from the specimen from Cuba (mostly typical in diameter. Surface of the choanosomal skeleton reveals numerous
discotriaenes, and common streptasterlamphiaster microscleres, irregularly running furrows which are extensions of the choanosomal
strongly tuberculated desmas), and the other from Horida (much canal system, 0.5 mm wide. These canals run more-or-Iess radially
larger and strongly incised discotriaenes to phyllotriaenes, into the sponge; there is another system of canals in the center which
microscleres are acanthoxeas and acanthorhabds) - these clearly run more-or-Iess vertically and lead to oscular openings. In inhalant
belong to different species, and most probably to different genera. areas there are canal openings about 0.5-0.7 mm wide. Choanosomal
Racodiscula asteroides, as currently defined, corresponds more desmas are typical tetraclones with sparsely tuberculated rays, and
closely to Schmidt's illustration (1870, pI. 3: 8) and slide BMNH strongly tuberculated tips; articulation is terminal. Microscleres are of
1870.5.3.7 - but unfortunately no desmas are present in this prepa- three types: massive microrhabds, densely covered with complex
ration. Such a situation does not permit us to fully characterize the spines, 6.7-10.6/Lm long, 4.8-5.3/Lm thick, and which may vary to
species, or even genus, as very similar phyllotriaenes, and/or pseudospherasters, 6.53-7.41/Lm in diameter (with all intermediate
microscleres are known in other species (and even genera; see for forms present), found mostly in the ectosomal region; slender and
example Rimella Schmidt). On the other hand, the first complete curved, strongly spinose rhabds, 12-13/Lm long, 3.6-4/Lm thick,
description of this genus and its type species (although named found mostly in the subectosomal region; spinose streptasters/amphi-
Racodiscula polydiscus), was given by Sollas (1888) based on the asters with long slender spinose rays, 14.7-20.7/Lm long, 11.2-
specimen from Cuba (determined by Schmidt, but clearly not 19 .4/Lm wide, found in the choanosome.
the one described in his 1870 paper). This specimen is missing from Remarks. Lendenfeld (1903) also included here PomeUa
the collection of BMNH, but the original description of the sponge schmidti Zittel, which is clearly a rhizomorine lithistid, leaving the
morphology leaves no doubts that various unrelated forms have genus monotypic.
been included by Sollas (1888) into this species as well. Conversely, Distribution. The type species is known from Barbados ~
the specimens from Barbados attributed to Racodiscula asteroides 108-170m depth, and from Florida. Undoubted occurrences are
by Van Soest & Stentoft (1988) fit the diagnosis of the genus from the Caribbean, with other records from the Philippines
Racodiscula as given by Sollas (1888), but do not agree entirely needing to be verified.
with any of Schmidt's slides or Sollas's description of R. asteroides.
Thus, at present the situation is as follows : the described type
species is clearly an amalgam, with no original material except for SILIQUARIASPONGIA HOSIDNO, 1981
two slides which undoubtedly consist of different species (perhaps
even genera), but this material does not allow for a precise defini- Synonymy
tion of the species as not all features can be found in them.
The species name is a misspelling of the original slide designation. Siliquariaspongia Hoshino, 1981b: 255.
To avoid confusion in the future, and to objectively determine a
nominal species, we propose (in accordance with ICZN article 75.1
and 75 .5; Anon., 1999) to establish a neotype of the species
Racodiscula asteroides Zittel. We have chosen the specimen ZPAL
Pf.15/1 (ex ZMA POR 5264), from Barbados, 85 fathoms, as the
neotype.

Description of type species

Racodiscula asteroides Zittel (Figs 8-11).

Synonymy. Corallistes polydiscus Schmidt, 1870, pI. 3: 8-9
(partim). Corallistes asteroides Carter, 1873c: 441 ; Racodiscula
polydiscus Sollas 1888: 332 (partim). Racodiscula asteroides Zittel, Fig. 8. Racodiscula asteroides Zittel, 1878, neotype ZPAL Pf.l511
1878a: 151; Lendenfeld 1903: 132 (partim); Van Soest & Stentoft, (ex ZMA POR5264) (scale lOmm).
Porifera· Demospongiae· 'Lithistids'· Theonellidae 333

Fig. 9. Racodiscula asteroides Zittel, neotype ZPAL Pf.151l (ex ZMA POR5264). A, ectosomal phyllotriaenes in non-inhalant area (scale 200 j.Lm).
B, phyllotriaenes and dense crust of microscleres in lateral inhalant area (scale 200 J,Lm). C, ectosomal microscIeres in inhalant area (scale 20 j.Lm). D, sur-
face of the choanosomal skeleton from lateral inhalant area (scale 2mm). E, choanosomal skeleton of tetraclone desmas (scale 400 j.Lm). F, details of tetr-
ac10ne desmas (scale 100 J,Lm).

Fig. to. Racodiscula asteroides Zittel, neotype ZPAL Pf.15tl (ex ZMA POR5264), (scale lOmm). A-F, ectosomal disco- to phyllotriaenes (scale 200 j.Lm).
G, choanosomal styles (scale 10 j.Lm).
334 Porifera' Demospongiae • 'Lithistids' • Theonellidae

Fig. 11. Racodiscula asteroides Zittel, neotype ZPAL Pf.lS/I (ex ZMA PORS264). A-B, acanthorhabds type II (scale 10 f.Lm). C, details of A (scale
2 f.Lm). D-H, streptasters/amphiasters (scales 5 f.Lm). I-M, acanthorhabds to pseudospherasters (scale 5 f.Lm).

Fig. 12. Siliquariaspongia japonica Hoshino, 1981b, holotype SAT-078

(after Hoshino, 1981b) (scale lOmm).

Type species

SiUquariaspongiajaponica Hoshino, 1981b: 256 (by original

Fig. 13. Siliquariaspongia japonica Hoshino. A, microscleres (acan-
designation). thorhabds) (scale 20 f.Lm). B, tetraclone desmas at various growth stages
(scale 100 f.Lm). C, discotriaenes (scale 100 f.Lm) (modified from Hoshino,
Definition 198Ib).

Theonellidae with tetraloph desmas. it from the latter. Based on this diagnosis some other species
assigned earlier to Discodermia (e.g., D. dissoluta Schmidt), may
Diagnosis also belong here.

Endosomal tetraclone des mas in the form of nonarticulated Description of type species
tetralophs; ectosomal spicules are discotriaenes; microscleres are
acanthostrongyles. Siliquariaspongia japonica Hoshino (Figs 12-13).
ReTTUlrks. This genus is very close to Discodermia, but the Synonymy. Siliquariaspongia japonica Hoshino, 1981b:
peculiar form of desmas and the absence of microxeas differentiate 256, fig. 37, pI. 3: 6.
Porifera· Demospongiae • 'Lithistids' • Theonellidae 335

Material examined. Holotype (not examined): SAT 078 - spicules are considered here to be unrelated. About 16 species have
Uchinoura, Japan, 1975. been attributed to the genus, which is essentially Recent, although
Description (from Hoshino, 1981b). Irregular massive it was also reported by Pis era (2000) from the Tertiary of the
sponge with smooth and even surface; oscules and pores not visi- Ukraine, based only on similarities of tetraclones and phyllotri-
ble; ectosomal spicules are densely packed discotriaenes. Desmas aenes with Recent species, whereas no microscleres were found in
are smooth tetralophs about 100--200/-lm in size and with rays fossil specimens.
about 15-30 /-lm thick. Discotriaenes are 100--200/-lm in diameter,
rhabdome short, conical about 20--30 /-lm long and 8-10 /-lm wide. Description of type species
They have very characteristic saw-toothed margins. Microscleres
are acanthostrongyles which are straight to bent, bean-shaped, Theonella swinhoei Gray (Figs 14--16).
measuring 20-30 /-lm long and 4--5 /-lm thick. Synonymy. Theonella swinhoei Gray, 1868b: 566, fig. 1;
Remarks. The holotype, apparently common in the mid- Zittel, 1878a: 87, pI. 1: 9b; Lendenfeld, 1903: 126; Thiele, 1900:
tidal zone of Uchinora, Japan, is symbiotic with Siliquaria 53, pI. 3: 3-4; Wilson, 1925: 448; Levi, 1958: 5, fig. 1; Levi,
cumingi (Morch). We have not seen the type material of this 1961a: 4, fig. 1; Vacelet & Vasseur, 1971 : 58, fig. 1; Levi & Levi,
sponge, but it is described with two types of microscleres: one a 1989: 56, fig. 24; Pulitzer-Fin ali, 1993: 261; (7) Gruber, 1993: 44,
bean-shaped acanthorhabd and the other a straight to bent acan- pI. 10: 7-8, pI. 11: 1-2.
thorhabd. Material examined. Holotype: BMNH 65.12.15.2 - Taiwan
(slides BMNH 1865.12.l5.2a, 2 specimens). Other material.
Distribution MNHN DCL 3216 - Philippines, Musorstom 2.
Description. Massive, short, shallow cup-shaped sponge with
Known only from Japan. a broad base by which it is attached to the rocky substrate, supplied
with a deep and narrow axial cavity. The sponge measures 3.5 cm
high and about 4 cm wide; walls are about 6 mm thick. A single large
THEONELLA GRAY, 1868 osculum, about 1 X 0.7 cm wide, is located in the middle of the cup-
shaped depression leading to a rapidly narrowing axial cavity. Outer
Synonymy side of the sponge is supplied with irregular thick ridges. Ectosomal
spicules are strongly differentiated phyllotriaenes with a very short
Theonella Gray, 1868b: 438; Zittel, 1878a: 37, 87; Sollas, rhabd; they may have relatively long and very slender (narrow) clads,
1888: 284; Lendenfeld, 1903: 126; Lendenfeld 1907: 344; Dendy, or conversely, relatively massive flattened clads in various portions
1922b: 4 (partim); Wilson, 1925: 447; de Laubenfels, 1955b: E58; of the sponge. Those with narrow clads occur mostly in the inhalant
Van Soest & Stentoft, 1988: 48; Gruber, 1993: 44; Hooper & areas on the outer sponge surface, while those with massive clads
Wiedenmayer, 1994: 439. occur mostly on the inner surface of the cup. The clads may bear
sparse tubercles, especially in the central portion of the cladome.
Type species Phyllotriaenes are 460--560/-lm in diameter. There are numerous
microscleres occurring on the dermal membrane between the
Theonella swinhoei Gray, 1868b: 566 (by monotypy). phyllotriaenes, as well as in subdermal lacunae. Microscleres are
spinose rhabds which may be straight, but in most cases are bent in
Definition the middle, sometimes at 90°, and they are strongly spined. The
spines are complex and divided into several smaller spines at the top,
Theonellidae with phyllotriaenes to discotriaenes among ectoso- usually with tips bent outwards. The size of these rhabds vary
mal spicules; microscleres are acanthorhabds and/or acanthoxeas. between 14--23/-lm, with thickness from 3.4-4.7 /-lm. Choanosomal
skeleton is composed of regular tetraclone desmas, which may be
Diagnosis

Polymorphic habitus; ectosomal spicules are phyllotriaenes

(ranging to discotriaenes); microscleres are acanthorhabds and/or
acanthoxeas; large endosomal oxeas may also be present.

Remarks

The holotype is attached by a wide base to the ferrugineous

rocky substrate. Lendenfeld (1907) differentiated Theonella and
Discodermia based on the presence or absence of microxeas, and
not on the nature of ectosomal triaenes. This view is not supported
here, and we adopted the delimitation of both genera as proposed by
Levi & Levi (1989), based on the exclusive presence of dis-cotri-
aenes in Discodermia; Theonella displays a dominance of phyllotri-
aenes while discotriaenes may be present in certain areas only.
Dendy (1922b) defined the genus Theonella as having "phyllotri-
aenes or (and) dichotriaenes": this view is also not supported here, Fig. 14. Theonella swinhoei Gray, 1868b, holotype, BMNH 65 .12.15.2
as sponges with phyllotriaenes and dichotriaenes as ectosomal (scale IOmm).
336 Porifera' Demospongiae· 'Lithistids" TheonelIidae

Fig. 15. Theonella swinhoei Gray, 1868b, holotype BMNH 65.12.15.2. A, view of the ectosome with slender phyllotriaenes in inhalant areas (scale
500/Lm). B, view of the ectosome with massive phyllotriaenes in noninhalant areas (scale 500 /Lm). C, crust of ectosomal phyllotriaenes as seen from
below, note short rhabds of phyllotriaenes (scale 200 /Lm). D, details of desma articulation (scale 100 /Lm). E, choanosomal skeleton of tetraclone desmas
from the outer surface (scale 200 /Lm). E, details of choanosomal skeleton of tetraclones, inner (upper) surface (scale 200 /Lm).

Fig. 16. Theonella swinhoei Gray. A-G, acanthorhabds (scale 5 /Lm). H, details of acanthorhabd surface (scale 2/Lm). I, ectosomal phyllotriaene with
tuberculated surface (scale 200 /Lm). J, tips of strongyle (scale 20 /Lm).
Porifera· Demospongiae • 'Lithistids' • Theonellidae 337

smooth or poorly tuberculated (mostly towards the zygomes) close Sollas (1888), describe long cylindrical sponges. On the other
to the outer surface, or strongly tuberculated along the whole length hand, their spicules are very similar to those of the holotype, includ-
(as observed on the inner surface). They measure approximately ing size. Thus, it seems that the holotype represents one end of a
325-360/-Lm in size. Zygomes are branched and strongly tubercu- highly variable morphological spectrum demonstrated by the type
lated. Young desmas are entirely smooth, calthrop-like. Other species.
choanosomal megascleres are radially oriented, slender, slightly Distribution. The type species is known from the Western
curved strongy1es, 700-900 /-Lm long, 9-11 /-Lm thick. Pacific, Indian Ocean, Indonesia and Red Sea. The genus is cosmo-
Remarks. The above description is based on the holotype, politan and has been reported from the Indian Ocean, Red Sea,
whereas most subsequent descriptions, notably following after Pacific and Atlantic (rare), from both shallow and deep waters.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John NA. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Siphonidiidae Lendenfeld, 1903

Andrzej Piseral & Claude Levi2

1 Instytut
Paleobiologii, Polska Akademia Nauk, ul. Twarda 51155,00-818 Warszawa, Poland. (apis@twarda.pan.pl)
2 Museum National d'Histoire Naturelle, Laboratoire de Biologie des Invertebres Marins et Malacologie, 57, Rue Cuvier, 75231 Paris
Cedex 05, France. (levi@mnhn.fr)

Siphonidiidae Lendenfeld (Demospongiae, 'lithistids') contains three valid genera and about ten nominal species from deep water, occurring
worldwide in warm seas. They are polymorphic, massive to cylindrical-clavate, with or without an atrial cavity, often with long fistules, or
ear- to vase- and cup-shaped. Siphonidiidae are characterized by rhizoclone choanosomal desmas, and choanosomal exotylostyles and/or
styles, and absence of any ectosomal spicules or microscleres. Genera are differentiated based on habitus and structure of the skeleton.
Keywords: Porifera; Demospongiae; 'lithistids'; Siphonidiidae; Gastrophanella; Siphonidium; Lithobactrum.

DEFINITION, DIAGNOSIS, SCOPE Diagnosis

Synonymy Polymorphic: cylindrical with deep atrial cavity, vase-shaped

or massive to globular with fistules; no special ectosomal spicules;
[Cladopeltidae] Sollas, 1888: 317 (nomen nudum); choanosomal spicules are rhizoclone desmas; other choanosomal
Topsent, 1894d: 290. Siphonidiidae Lendenfeld, 1903: 140; megascleres are exotylostyles with ornamented spinose head or
Lendenfeld, 1907: 360; Wilson, 1925: 469; Van Soest & Stentoft, styles; no microscleres of any kind.
1988: 66; Muricy & Minervino, 2000: 600; Gastrophanellidae de
Laubenfels, 1936a: 157 (partim); Tremaulidiidae de Laubenfels, Remarks
1936a: 81.
Sollas (1888) proposed the new family Cladopeltidae for the
Scope genus Siphonidium, the name of which, unfortunately, was not
based on any valid genus name and hence contrary to the rules and
Three valid genera: Gastrophanella Schmidt, 1879, recommendations of the ICZN (Anon., 1999). Consequently, we
Lithobactrum Kirkpatrick, 1903a, Siphonidium Schmidt, 1879. accept here Lendenfeld's (1903) Siphonidiidae as the first available
name for this family group based on the genus Siphonidium, despite
Definition the priority of Sollas's name. Some fossil rhizomorine lithistids
may also belong here as suggested by the structure of their skele-
Rhizomorine 'lithistid' Demospongiae with exotylostyles tons. Exotylostyles occurring in this family are most probably not
with smooth to ornamented head, and/or styles, among choanoso- homologous to similar spicules known from non-lithistid sponges,
mal megascleres. and thus should not be used as a linking character between the two.

KEY TO EXTANT GENERA

(1) With long fistules, exotylostyles with ornamented heads ................................................................................................ Siphonidium
No fistules ...................... ,.................................................................................................................................................................... 2
(2) Deep and narrow atrial cavity, ear- or vase-shaped ..................................................................................................... Gastrophanella
No atrial cavity but a bundle of vertical canals in the center .......................................................................................... Lithobactrum

GASTROPHANELLA SCHMIDT, 1879 Definition

Synonymy Siphonidiidae which are cylindrical to globular with deep

atrial cavity, or vasiform to ear-shaped in growth form.
Gastrophanella Schmidt, 1879: 29; Sollas 1888: 349;
Lendenfeld, 1903: 148 (partim); Burton, 1929b: 7; Van Soest & Diagnosis
Stentoft, 1988: 68; Gomez, 1998: 774; Mothes & da Silva 1999:
129; Muricy & Minervino, 2000: 600. Clavate to cylindrical, or globular to massive lobate with deep
and narrow central cavity, or vase- to ear-shaped. No special ecto-
Type species somal spicules; desmas are rhizoclones forming reticulate fibers on
the surface; other choanosomal spicules are smooth styles, and/or
Gastrophanella implexa Schmidt, 1879: 29 (by monotypy). exotylostyles with ornamented to smooth heads.

338
Porifera' Demospongiae· 'Lithistids" Siphonidiidae 339

Remarks occur at any place but terminal zygoses are most common;
articulation very strong and complex. The rhizoclones reach 100-150
This genus was sometimes regarded as belonging to f..lm in size. Other choanosomal megascleres are long smooth styles
Azoricidae Sollas, but members of this family never have styles and exotylostyles with spinose heads, about 400-500 f..lm long,
among their megascleres. Five species have been attributed to the
genus at various times, of which probably only four are valid. For
more detailed discussion on the position of this genus within the
Demospongiae see recent review by Muricy & Minervino (2000).

Description of type species

Gastrophanella implexa Schmidt, 1879 (Figs 1-2).

Synonymy. Gastrophanella implexa Schmidt, 1879: 29, pI. 1:
7, pI. 3: 8; Sollas, 1888: 349; Lendenfeld, 1903: 149; Van Soest &
Stentoft, 1988: 68 (partim; only tube shaped forms). Gastrophanella
stylifera Mothes & da Silva, 1999: 129, figs 12-26, 37-41.
Material examined. Holotype: MCZ 6232 - between
Florida and Cuba, depth 232m. MNHN DT 2191 (slide of the
Schmidt specimen MZUS).
Description. Club-shaped sponge with single oscule located
at the top and leading to a narrow and very deep central cavity (run-
ning through entire sponge); sponge about 4.5 cm high, 2 cm wide at
the upper portion, osculum 3 mm diameter. Outer surface with very
dense round to meandering elongated pores, which are separated
by very narrow skeletal bridges formed by fibers composed of rhi-
zoclone desmas. Inner surface also reticulate but with rounded
pores, 100-200 f..lm in diameter, separated by narrow skeletal
bridges 25-50 f..lm wide. Incurrent canals more-or-Iess radially
arranged and very narrow; exhalant canals curved and following
the upper surface. Desmas are rhizoclones, often resembling tetra- Fig.l. Gastrophanella implexa Schmidt, holotype MCZ 6232 (scale 10 nun).
clones, bearing conical spines which bifurcate; articulation may A, lateral view. B, longitudinal section view.

Fig. 2. Gastrophanella implexa Schmidt, holotype MCZ 6232. A, inner part of choanosomal skeleton of rhizoclone desmas; note numerous vertically
arranged oxeas (scale 200 11m). B, details of choanosomal rhizoclone desmas (scale 100 11m). C, surface of the choanosomal skeleton from the atrial cav-
ity (scale 500 11m). D, details of rhizoclone articulation (scale 20 11m). E, exotylostyle head (scale 10 11m).
340 Porifera· Demospongiae • 'Lithistids' • Siphonidiidae

5 f.1m thick. Long oxeas (length not measured) are oriented more- Material examined. Holotype: BMNH 1902.5.2.67. BMNH
or-less vertically. This species is known from the Caribbean and 1902.5.2.67a (slide from holotype).
southern Brazil from depths between 150-228 m. Description. Massive club-shaped sponge which may reach
Remarks. Apart from typical tube-shaped forms Van 18 cm high, 8.5 cm wide, with the holotype about 7 cm high and ellip-
Soest & Stentoft (1988) also included vase-shaped to ear-shaped tical in cross-section at the top measuring 4 X 3 cm. The summit of the
specimens in this species, but in our opinion these latter specimens sponge is elliptical with numerous oscules about 0.5 mm diameter
belong to a different but very closely related (cryptic) species, which lead to the vertically arranged exhalant canals. These oscules
whose shape clearly exceeds the observed specific variability are surrounded by irregularly radiating subectosomal canals visible on
among rhizomorine lithistids. These authors also suggest that the surface of the choanosomal skeleton. Fine parallel incurrent canals
the genera Siphonidium and Gastrophanella should probably be radiate horizontally inwards from the pores. Pores in the choanosomal
united. We offer a different opinion on the basis that there are clear skeleton uniformly and densely distributed on the sides, 200-324 fLm
differences between both, not only in habitus but also in choanoso- in diameter. Choanosomal skeleton consists of rhizoclone desmas up
mal skeleton. Mothes & da Silva (1999) described a new species, to 450 f.1m in size; the skeleton is composed of very dense reticulate
G. stylifera, which differs from G. implexa only by the presence of complex fibers of rhizoclones, producing a very strong skeleton. The
smooth styles, supposedly absent in G. implexa. In fact these styles resulting meshes of the skeleton are rounded to elongate, 250-300 f.1m
also occur in this species, and thus G. stylifera is included in in diameter, while particular fibers are 140-180 f.1m thick. The desmas
synonymy with the type species. are very densely and covered with strong tubercles (or massive spines),
which are often slightly bifurcate, and which are supplied with 1-3
Distribution small spines at the top. No special ectosomal spicules. Other
choanosomal megascleres are exotylostyles with large, rounded and
Caribbean, NE and S Brazil, Belize, South Africa, Pacific strongly spinose heads, about 200-420 f.1m long and 6-7 fLm thick,
coast of Mexico, both from shallow (submarine caves) and deeper with head about 11 f.1m in diameter. This species has been collected so
waters. far only off the coast of SE Africa, at 100 m depth.

L1THOBACTRUM KIRKPATRICK, 1903

Synonymy

Lithobactrum Kirkpatrick, 1903a: 175.

Type species

Lithobactrumforte Kirkpatrick, 1903a: 175 (by monotypy).

Definition

Club-shaped Siphonidiidae with a bundle of vertical canals in

the center.

Diagnosis

Club-shaped, massive Siphonidiidae, with uniformly distrib-

uted pores at the sides, and with numerous small star-like oscules
on the summit; excurrent canals form bundles running vertically
upwards to the oscules.

Remarks

The structure, canalization and morphology of this genus are

very close, if not identical to the fossil genus Jereica Zittel.
Kirkpatrick (1903a) considered this species as an Azoricidae, but
the presence of typical tylostyles with spinose heads indicates its
position lies within the Siphonidiidae. Only one species has been
ascribed to this genus so far.

Description of type species

Lithobactrumforte Kirkpatrick, 1903a (Figs 3-5).

Synonymy. Lithobactrum forte Kirkpatrick, 1903a: 175, Fig. 3. Lithobactrum Jorte Kirkpatrick, holotype BMNH 1902.5.2.67
pI. 4: 5, 5a-g. (scale 10 mm). A, lateral view. B, top view with star-like oscules visible.
Porifera· Demospongiae • 'Lithistids' • Siphonidiidae 341

Fig. 4. Lithobactrum forte Kirkpatrick, holotype BMNH 1902.5.2.67. A, Surface of the choanosomal skeleton (scale 1 mm). B, interior of the choanoso-
mal skeleton composed of rhizoc1one desmas organized in reticulate fibers (scale 1 mm). C, details of rhizoc1one desmas from the surface (scale 100 fLm).
D, rhizoclone desma articulation and exotylostyles (scale 50 fLm).

reason why exotylostyles where not reported by Kirkpatrick

(1903a): i.e., they do not occur in spicule preparations. On the
other hand, Kirkpatrick (1903a) recorded the presence of
choanosomal amphityles, which may be variations of tylostyles.

Distribution

Known only from the coast of SE Africa.

SIPHONIDIUM SCHMIDT, 1879

Synonymy

Siphonidium Schmidt, 1879: 28; Sollas, 1888: 317, 348; Van

Soest & Stentofi, 1988: 66. Gruber, 1993: 50. Siphonidiella Burton,
1928a: 112. Tremaulidium Schmidt, 1879: 31, pI. 2: la-I, 2, pI. 4: 1-2.

Type species

Leiodermatium ramosum Schmidt, 1870: 23 (by original

designation).

Definition

Fig.5. Lithobactrumforte Kirkpatrick, holotype BMNH 1902.5.2.67. A, head Siphonidiidae bearing fistules, and choanosomal exotylostyles
of exotylostyle (scale 20 fLm). B, head and end of exotylostyle (scale 10 fLm). with heads directed outwards.

Remarks. There are other spicules, not mentioned above, Diagnosis

occurring in the superficial zone of the holotype, but they clearly
belong to some other encrusting sponge. The tylostyles, on the con- Encrusting, massive irregular, hemispherical or irregular
trary, occur deeply within the choanosome and are often strongly cylindrical to club-shaped with fistules; desmas are rhizoclones;
embedded into the desma skeleton. The latter was probably the no special ectosomal spicules but the most external flattened desmas
342 Porifera· Demospongiae· 'Lithistids'· Siphonidiidae

form a very dense armor-like layer; other megascleres are exoty-

lostyles with rugose heads directed outwards.

Remarks

Four or five species are included in this genus. Burton (l928a)

described a new genus Siphonidiella, which differs, according to
him, from Siphonidium only in having dichotriaenes dispersed
within the choanosomal skeleton. Examination of the original
preparations by Burton (BMNH 1926.10.1.15) revealed no dicho-
triaenes at all, but rhizoclones as in Siphonidium. Furthermore,
from the theoretical point of view, such an assemblage of spicules
seems artificial. As a result, we regard the dichotriaenes as contam-
inants and we synonymize Siphonidiella with Siphonidium.

Description of type species

Siphonidium ramosum (Schmidt, 1870) (Figs 6-8).

Synonymy. Leiodennatium ramosum Schmidt, 1870: 21,
pI. 3: 1; Siphonidium ramosum Schmidt, 1879: 28, pI. 1: 8; Sollas,
1888: 348; Topsent, 1892a: 52; Lendenfeld, 1903: 144; Topsent,
Fig. 6. Siphonidium ramosum Schmidt. A, MZUS P0162 (scale 1 cm). 1904b: 63, pI. 4: 9, pI. 8: 5; Van Soest & Stentoft, 1988: 66, fig. 31;
B, MCZ 6322. C, MCZ 6321 (scale 1 cm). Mothes & da Silva 1999: 131, figs 27-32, 42-46.

Fig. 7. Siphonidium ramosum Schmidt (A-D, original Schmidt specimen MZUS P0162; E-F, ZPAL Pf.l5/5 (ex. ZMA POR5253)). A, General view
of the fistules (scale 1 mm). B, details of dense desma armor of the choanosomal skeleton (scale 100 11m). C, interior of the choanosomal skeleton in the
fistule (scale 100 11m). D, surface of the choanosomal skeleton close to the edge of fistule (scale 100 11m). E, surface of the choanosomal skeleton on the
fistule (scale bar 200 11m). F, details of the surface of the choanosomal skeleton on the fistule, ZPAL Pf.I5/5 (ex ZMA POR 5253) (scale 50 11m).
Porifera· Demospongiae • 'Lithistids' • Siphonidiidae 343

Fig. 8. Siphonidium ramosum Schmidt, ZPAL Pf.1515 (ex ZMA POR 5253). A-B, exotylostyles (scale 50 ILm). C-E, heads and tips of exotylostyles
(scale 5ILm).

Material examined. Type material: MCZ 6321 and 6322- The heads of extotylostyles are directed outwards. This species is
Schmidt material, Florida. BMNH 1886.12.31.2-3 - Schmidt reported from Florida and the Caribbean, Azores, Brazil, and the
material, Cuba. MNHN DT 780 - Schmidt material from MZUS Mediterranean, from depths between 150-500m.
P0162. Other material. ZMA POR 5235 - Barbados. MZUS Remarks. Contrary to the observations of Van Soest &
P0162 - dry specimen (Schmidt material). Stentoft (1988), who did not observe the outermost dense layer of
Description. Small, about 2-5.5 cm high and 1 cm wide, desmas, we have found it in both the Schmidt type specimens from
irregularly massive to cylindrical sponges bearing numerous small MCZ as well as from MZUS, and ZMA collection from Barbados.
fistules; usually they are directed upward, but often may run in As a result, this feature can be used as a valid diagnostic character,
various directions. These fistules are about 1-2mm in diameter as originally proposed by Sollas (1888). On the other hand, we did
and open, although some may be entirely closed. Surface smooth not see desmas without zygosis in the subectosomal region. These
except wrinkled. No ectosomal spicules present of any kind. probably occur only at the top of fistules, and thus a new definition
Choanosomal desmas are massive rhizoclones, which are supplied of the genus Siphonidium is here proposed. Contrary to the opinion
with a long spine-like outgrowth in the interior and at the top of fis- of Lendenfeld (1903) we consider that the outermost layer of
tules, and at most of the surface are strongly tuberculated, tightly interlocking desmas is not homologous with specialized
measuring about 180-220 fLm in size. The desma skeleton is rela- ectosomal megascleres, but forms a part of choanosomal skeleton
tively dense, especially at the sponge surface. The most exterior represented by modified desmas.
desmas are flattened, more strongly branched and tightly interlock-
ing in one plane so there are no meshes or pores visible but only Distribution
a solid surface. The desmas are slightly less tightly interlocking
within fistules. Other choanosomal megascleres are exotylostyles Florida and the Caribbean, Azores, Western Africa, New
with spinose heads, measuring 160-220 fLm long, 2-3 fLm thick. Caledonia and Indonesia.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Neopeltidae Sollas, 1888

Andrzej Pisera1 & Claude Levi2

1 Instytut Paleobiologii, Polska Akademia Nauk, ul. Twarda 51/55, 00-818 Warszawa, Poland. (apis@twarda.pan.pl)
2Museum National d'Histoire Naturelle, Laboratoire de Biologie des Invertebres Marins et Malacologie, 57, Rue Cuvier, 75231 Paris
Cedex 05, France. (levi@mnhn.fr)

Neopeltidae Sollas (Demospongiae, 'lithistids') contains four genera and about 20 nominal species, some presently incorrectly allocated,
and known from all oceans. Species are polymorphic, ranging from thickly encrusting to massive irregular, massive clavate shapes with
shallow axial cavity, to subglobular with a stalk and thick lamellae consisting of several individuals. They are known both from deep
and very shallow waters, including coral reefs. Neopeltidae is characterized by monaxial ectosomal spicules (pseudophyllotriaenes or
pseudodiscotriaenes), monaxial complex shaped branching desrnas and microscleres which are amphiasters and/or spinose microxeas.
Genera are differentiated on the basis of the morphology of ectosomal pseudophyllotriaenes and pseudodiscotriaenes, and microscleres
geometries.
Keywords: Porifera; Demospongiae; 'lithistids'; Neopeltidae; Callipelta; Daedalopelta; Homophymia; Neopelta.

DEFINITION, DIAGNOSIS, SCOPE Homophymia Vacelet & Vasseur, 1971, and Neopelta Schmidt,
1880b.
Synonymy
Remarks
Neopeltidae Sollas, 1888: 344. Topsent, l894d: 290;
Schrammen, 1903: 20; de Laubenfels, 1955: E49 (partim); Kelly, The genera Callipelta Sollas and Homophymia Vacelet &
2000: 281 (partim). Daedalopeltidae de Laubenfels, 1936a: 184. Vasseur have usually been regarded as representatives of
Corallistidae. They have, however, ectosomal spicules of monaxial
Definition nature accompanied by monaxial desmas, similar to Neopelta
Schmidt, and for this reason they were transferred to Neopeltidae
'Lithistid' Demospongiae with monocrepid desmas of by Kelly (2000), which is here followed. She included Neopeltidae
complex morphology and monaxial ectosomal pseudophyllotriaenes together with Corallistidae in the Dicranocladina, but we refute this
and/or pseudodiscotriaenes, and amphiasters among microscleres. allocation based on fundamental differences in their ectosomal
spicules (in Corallistidae these are exclusively dichotriaenes, while
Diagnosis in Neopeltidae they are pseudophyllotriaenes/pseudodiscotri-
aenes). Consequently, we consider them to be unrelated groups.
Polymorphic; ectosomal spicules are monaxial pseudodisco- In the recent review of the family Kelly (2000) included in
triaenes to pseudophyllotriaenes, and/or siliceous plates; it Macandrewia, in addition to Callipelta, Daedalopelta,
choanosomal spicules are monocrepid des mas of complex branch- Homophymia and Neopelta. Again we do not support this alloca-
ing shape, often resembling tetraclones, sometimes accompanied tion based on the possession of true tetracrepidial phyllotriaenes
by strongyles; microscleres are amphiasters and/or spinose oxeas. in Macandrewia, as well as triaenose crepis in the majority of its
desmas. De Laubenfels (1936a) proposed a new family
Scope Daedelopeltidae for the genus Daedalopelta Sollas, and included
in it unrelated genera as Lyidium Schmidt (=Pleroma Sallas) and
Four Recent genera are here attributed to Neo- Vetulina Schmidt. We consider his family to be synonymous with
peltidae: Callipelta Sollas, 1888; Daedalopelta Sollas, 1888; Neopeltidae.

KEY TO GENERA

(1) Ectosomal spicules are pseudodiscotriaenes (monaxial discs), microscleres are oxeas ........................................................ Neopelta
Ectosomal spicules are pseudophyllotriaenes ..................................................................................................................................... 2
(2) Clads not incised and smooth, microscleres are one type of amphiasters ...................................................................... Homophymia
Ectosomal spicules are tuberculated pseudophyllotriaenes with dentate clads, microscleres are two types of
amphiasters ............... ... ................. .......... ........... ........... .......... ................. ... ............. .......... ................ ........... ..................... .... Callipelta
Ectosomal spicules are dentate pseudophyllotriaenes, microscleres are one type of amphiasters .................................. Daedalopelta

344
Porifera· Demospongiae· 'Lithistids'· Neopeltidae 345

CALLlPELTA SOLLAS, 1888 Synonymy. C. ornata Sollas, 1888: 309-312, pI. 37: 28-30
(as Corallistes callipelta); Vacelet & Vasseur, 1971: 62.
Synonymy Macandrewia ornata; Vacelet et al., 1976: 26; Vacelet & Vasseur,
1965: 84.
Callipelta Sollas, 1888: 309. Material examined. Holotype: BMNH 1877.5.21.1382,
1383 (both slides), MNHN DeL 2842 (slide) - Tulear, and
Type species E596bis (Tulear).
Description. Small pillow-like, 1-3 cm long, up to 1 cm thick,
Callipelta ornata Sollas, 1888: 309 (by monotypy). with several oscules, or hemispherical at early stages of growth with a
single oscule at the top. Ectosomal skeleton consists of pseudophyllo-
Definition triaenes to pseudodiscotriaenes (crepis in the rhabd) which are usually
strongly tuberculated on the upper surface and have very complex
Neopeltidae with dentate pseudodisco- to pseudophyllotri- shape with strongly incised or dentate margins; cladome 210-320 fLm
aenes bearing tubercles on the upper surface and with two types of in diameter, rhabdome 50-75 fLm long. Surface of the choanosomal
amphiasters as microscleres. skeleton displays groups of choanosomal canal openings about

Diagnosis

Polymorphic; desmas monocrepid, irregular but resembling

tetraclones; ectosomal spicules are pseudodiscotriaenes to pseudo-
phyllotriaenes (monocrepidial), usually with strongly incised mar-
gins or clads and a strongly tuberculated upper surface; microscleres
are two types of amphiasters.

Remarks

This genus presently contains 6 nominal species, some of

which, however, may belong to Neopelta and Daedalopelta, as
here defined.

Description of type species

Callipelta ornata Sollas (Figs 1-3). Fig. 1. Callipelta ornata Sollas. E596bis, Tulear (scale 5 mm).

Fig. 2. Callipelta ornata Sollas. E596bis, Tulear. A, Surface of the sponge showing ectosomal pseudophyllotriaenes and microscleres (scale 200 fLm);
B, surface of the choanosomal skeleton of desmas (scale 2 mm); C, desmas of the choanosomal skeleton (scale 200 fLm); D, details of articulation between
choanosomal desmas (scale 50 fLm).
346 Porifera· Demospongiae • 'Lithistids' • Neopeltidae

Fig. 3. Callipelta ornata Sollas, B596 bis. A-D, choanosomal amphiasters (scale 5 fLm); E-I, ectosomal amphiasters (scale 5 fLm); J-L, ectosomal pseudo-
phyllotriaenes (scale 100 fLm).

300 f.Lm in diameter (corresponding to oscular openings on the Definition

sponge surface) from which radiate irregularly subdermal canals of
similar width. Monaxial des mas irregular and usually smooth along Neopeltidae with dentate pseudophyllotriaenes as ectosomal
most of their surface, only at the tips strongly tuberculated and spicules and one type of amphiaster as microscleres, but occurring
slightly branched. Articulation between desmas terminal. Other in two size classes.
choanosomal megascleres include slender oxeas, 140-300 f.Lm long,
2 fLm thick. Microscleres include two types of amphiasters: subec- Diagnosis
tosomal ones are massive and short, 8.0-9.2 f.Lm wide, 9.4-
10.9 /-Lm long, with short rays which have microspinose blunt heads; See definition.
choanosomal ones are 9.7-11.6 /-Lm wide, 12.4-13.8 /-Lm long, with
spinose, pointed and long rays. This species was reported from a Remarks
depth of 236 m from the Indonesian region (Banda Sea), and from
Madagascar (tunnels in the Tulear Reef) at depths of 3--6 m. This poorly known monotypic genus is very close to
Remarks. Vacelet & Vasseur (1965) also recorded styles Callipelta, from which it differs in having only one kind of amphi-
among the choanosomal megascleres. aster with long and slender rays, but occurring in two size classes. It
is conceivable that the two genera may be merged in future studies.
Distribution
Description of type species
Philippines, Indonesia, Madagascar, New Caledonia.
Discodermia clavatella var. nodosa Schmidt (Figs 4--6).
Synonymy. Discodermia clavatella var. nodosa Schmidt,
DAEDAWPELTA SOLLAS, 1888 1879, 25, pI. 3: In, 0; Daedalopelta nodosa Sollas, 1888: 342.
Macandrewia nodosa Lendenfeld, 1903: 139.
Synonymy Material examined. Specimen: HBOI 003: 00949 -
Bahamas.
Daedalopelta Sollas, 1888: 342. Macandrewia Lendenfeld, Description. Small massive clavate or irregular cylindrical
1903 (partim). to cushion shaped sponge which may be up to 38 mm high and
24 mm wide. At the top is a shallow concavity or atrium with numer-
Type species ous canal openings leading to radial canals within the sponge and
on the surface of the choanosomal skeleton, where they form a
Discodermia clavatella var. nodosa Schmidt, 1879: 25 branching stellate pattern. Ectosomal megascleres are pseudophyl-
(=Daedalopelta nodosa Sollas, 1888: 342) (by monotypy). lotriaenes with irregular narrow cladi, with margins, with numerous
Porifera· Demospongiae • 'Lithistids' • Neopeltidae 347

indentations. Upper surface may also bear tubercles.

Pseudophyllotriaene cladomes measure 250-350 fLm in diameter,
with conical rhabdome 50-115 fLm long. Other choanosomal
megascleres may include slender cylindrical oxeas 200 fLm long,
3 fLm thick. Microscleres consist of a single type of amphiaster, but
occurring in two size classes: the smaller 10-11 fLm long, 12-
13 fLm wide, relatively massive and has shorter rays; the larger is less
massive, 17-26fLm long, 13-19fLm wide, with longer and more
slender rays. Occasionally metasters are found, about 24 fLm in
length. This species was reported by Schmidt (1879) from the Gulf
of Mexico and Kelly (2000) from the Bahamas from 451 m depth.
Remarks. This is very poorly known species having so
far been described only by Schmidt (1879) from Florida and
redescribed by Sollas (1888). No original material was found,
so this description is based on newly discovered specimens from
the HBOI collection.

Distribution
Fig. 4. Daedalopelta nodosa (Schmidt), HBOI 003: 00949, Bahamas,
(scale 10mm). Florida, Bahamas.

Fig. 5. Daedalopelta nodosa (Schmidt), HBOI 003: 00949, Bahamas. A, inhalant surface (scale 200 ILm); B, details of the surface with ectosomal pseudo-
phyllotriaenes (scale 200 ILm); C, surface of the choanosomal skeleton (scale I mm); D, details of choanosomal skeleton composed of pseudotetraclone
desmas (scale 200 ILm); E, details of desma terminal articulation (scale 100 ILm); P, modified desmas on choanosomal canal (scale 200 ILm).
348 Porifera· Demospongiae· 'Lithistids'· Neopeltidae

Fig. 6. Daedalopelta nodosa (Schmidt), HBOI 003: 00949. A-C, choanosomal amphiasters (scale 5 fAom); D-F, subectosomal amphiasters (scale 5 fAom);
O-J, ectosomal pseudophyllotriaenes (scale 100 fAom).

HOMOPHYMIA VACELET & VASSEUR, 1971

Synonymy

Homophymia Vacelet & Vasseur, 1971: 63; Kelly, 2000: 273.

Type species

Homophymia lamellosa Vacelet & Vasseur, 1971: 63 (by

monotypy).

Definition

Neopeltidae with smooth, not dentate pseudophyllotriaenes as

ectosomal spicules and one type of amphiaster as microscleres.
Fig. 7. Homophymia lamellosa Vacelet & Vasseur, holotype MNHN DJV
26, (scale 30mm).
Diagnosis

Thickly lamellar (consisting of several cylindrical individu-

als) or thickly globular with long stalk supplied with axial cavity. genus into Neopeltidae is based on its monaxial desmas and ecto-
Desmas are pseudotetraclones (monocrepid), ectosomal megascle- somal pseudophyllotriaenes (see also Kelly, 2000). TWo species
res are smooth pseudophyllotriaenes (monocrepid, crepis in the have so far been described.
rhabd); microscleres consist of a single type of amphiaster.
Description of type species
Remarks
Homophymia lamellosa Vacelet & Vasseur (Figs 7-9).
This genus was originally classified with some uncertainty Synonymy. Homophymia lamellosa Vacelet & Vasseur,
within the Corallistidae, based on its monaxial desmas 1971: 63, fig. 6, pI. 1: 4-7; Vace1et eta/., 1976: 27.
(Corallistidae, as here understood, has dichotriaenes as their ecto- Material examined. Holotype: MNHN DJV 26 - Tulear,
somal spicules, never pseudophyllotriaenes). The allocation of this Madagascar. MNHN DCL 2847 (slides from the holotype).
Porifera· Demospongiae· 'Lithistids'· Neopeltidae 349

Fig. 8. Homophymia lamellosa Vacelet & Vasseur, MNHN DJV 26. A, inhalant surface with ectosomal pseudophyllotriaenes (scale 500 ILm); B, Exhalant
surface with ectosomal pseudophyllotriaenes (scale 500 ILm); C, outer surface of choanosomal skeleton with subdermal canals (scale 2 mm); D, details of
choanosomal skeleton composed of pseudotetraclones (scale 200 ILm).

Fig. 9. Homophymia lamellosa Vacelet & Vasseur, MNHN DJV 26. A-E, amphiasters (scale 5 ILm); F, strongyle (scale 20 11m); G, style (scale bar 20 11m);
H-J, tylotes (scale 5 ILm); J-L, ectosomal pseudophyllotriaenes (scale 100 ILm).
350 Porifera· Demospongiae • 'Lithistids' • Neopeltidae

Description. Thickly lamellar sponge (8 em high, 10 em wide, Remarks

2cm thick) with a rounded outline consisting of several more-or-Iess
individualized (especially at the top) but fused, cylindrical sponges Desmas of Neopelta are very close in shape to the desmas of
with separate small oscules (1.5-2.0mm diameter) situated at the top, Theonellidae, especially some Discodermia species. They differ,
corresponding to the deep and narrow axial cavity. The two surfaces however, in being clearly monocrepidial, similar to their ectosomal
are different. One displays subectosomal canals (visible below the irregular discs (where the crepis is in the disc). De Laubenfels (1936)
ectosome) and has pseudophyllotriaenes (crepis in the rhabd) with stated that N. perfecta should be the type species while
slender clads, which fonn regular meshes (about 120-170 /.Lm across) de Laubenfels (1955) pointed out that N. imperfecta was established
in the center of which pores are placed. The other surface has less as the type species, but Schmidt (1880b) never designated a type
developed subectosomal canals and is covered with pseudophyllotri- species. Furthennore, the illustration of ectosomal discs of
aenes with thicker and more branched clads; pores are rare. The N. imperfecta by de Laubenfels (1955) corresponds to those
pseudophyllotriaenes with slender clads on the inhalant surface meas- in N. perfecta and not N. imperfecta. In our opinion, in this situation
ure 220-340 /.Lm across, while those with thick clads from the oppo- N. perfecta should be considered the type species by subsequent des-
site surface are slightly larger and measure 330-455/.Lm diameter. ignation by de Laubenfels (1936a). So far only 3 species have been
Desmas, about 250-350/.Lm in size, resemble tetraclones but are assigned to Neopelta although some of those currently described as
monocrepidial. They are slightly tuberculate, especially toward their Callipelta may be better placed here. The genus Lepidothenea de
zygomes which are terminal. Other choanosomal megascleres are Laubenfels, 1936a (type species Lepidospongia incrustans Dendy,
large styles to strongyles/tylotes measuring 268-500 /.Lm long and 1924: 316), is probably a junior synonym of Neopelta. It apparently
6-8/.Lm thick There are also small oxeas measuring 77-97/.Lm long lacks desmas but has pseudodiscotriaenes as ectosomal spicules and
and 0.7-2.2/.Lm thick Microscleres are microspinose amphiasters fusifonn acanthoxeas or acanthorhabds as microscleres.
with long slender rays and are very abundant at the surface of the
axial cavity and choanosomal canals; they measure 10-14/.Lm long Description of type species
and about 9-11/.Lm wide (rays about 3-6/.Lm long). This species was
found in 2-11 m depth in reef overhangs and cavities. Neopelta perfecta Schmidt, 1880b (Figs 10-12).
Remarks. Vacelet & Vasseur (1971) did not observe small Synonymy. Neopelta perfecta Schmidt, 1880b: 88, pI. 5: 3;
oxeas in their preparations, and they also reported large strongyles Sollas, 1888: 344.
rather than tylotes, which were slightly longer. These differences Material examined. Type material: two unnumbered slides
may be due to their preparations being made from different parts from MZUS - originally illustrated by Schmidt, 1880b (pI. 5: 3).
of the sponge than our preparations, illustrating a small level of Other material. HBOI 003: 00948 (2 specimens) - Grenada.
variability within this species. Description. Small massive cylindrical, short clavate or
irregular rounded sponges constricted above the base, measuring
Distribution about 3.8 cm high, 2 cm wide. Osculum, if present, at the top
(RBOI specimen) or several oscules are located on small erect
Madagascar, New Zealand, Reunion Island. processes (reported by Schmidt, 1880b and Sollas, 1888).
Ectosomal spicules are monocrepidial discs (pseudodiscotriaenes),
more-or-Iess oval in outline, often slightly concave, rarely with
NEOPELTA SCHMIDT, 1880 perpendicular process resembling rhabdomes, and very thin, about
143-232 /-Lm in diameter. Below the choanosome and visible on the
Synonymy surface of the choanosomal skeleton are subectosomal meandering
canals. Choanosomal desmas, forming a confused network, are of
Neopelta Schmidt, 1880b: 88; Sollas, 1888: 344; Lendenfeld,
1903: 140; de Laubenfels, 1955: E49; Levi & Levi, 1988: 252.
? Lepidothenea de Laubenfels 1936a: 176.

Type species

Neopelta perfecta Schmidt, 1880b: 88 (subsequent designa-

tion de Laubenfels, 1936a: 176).

Definition

Neopeltidae with monaxial discs (pseudodiscotriaenes) as

ectosomal spicules and microscleres as microxeas and amphiasters.

Diagnosis

Neopeltidae with ectosomal spicules as pseudodiscotriaenes

of oval to elongated shape (crepis in the disk); microscleres are
microxeas and amphiasters. Desmas are of complex shape, often Fig. 10. Neopelta peifecta Schmidt, HBOI 003: 00948, Grenada
resembling tetraclones but monocrepidial. (scale lOmm).
Porifera· Demospongiae • 'Lithistids' • Neopeltidae 351

Fig. 11. Neopelta perfecta Schmidt, HBOI 003: 00948, Grenada. A, outer surface with ectosomal pseudodiscotriaenes (scale 200 fJom); B, outer surface of
choanosomal skeleton (scale I mm); C, details of the outer surface of choanosomal skeleton composed of monaxial desmas (scale 200 fJom); D, details of
the interior of choanosomal desmas (scale 500 fJom).

Fig. 12. Neopelta perfecta Schmidt, HBOI 003: 00948, Grenada. A-C, microxeas (scale 10 fJom); D, points of microxeas (scale 5 fJom); E-H, choanosomal
amphiasters (scale 5 fJom).

complex shape and are monocrepidal, with smooth rays, slender on species is known only from the Caribbean, from the 188 m depth
the skeleton surface and irregular, 345-480 f.Lm in diameter, with off Barbados, and 116 m depth off Grenada.
terminal articulations. They are more regular and more massive Remarks. The morphology of the holotype as described by
inside the choanosome. Other megascleres are slender, slightly Schmidt (1888), and which has not yet been found (only slide
curved oxeas occurring singly or in bundles, and parallel to the preparations were investigated), differs from that of the investi-
main canals, measuring 420-540 f.Lm long, 5-8 f.Lm thick. gated specimens from HBOI, although their spicule compositions
Microscleres are fusifonn rnicroxeas with spinose surface, 55--68 f.Lm were identical.
long, 3-5 f.Lm thick, which occur both at the surface as a tangential
layer below the discs in the ectosome, and in the choanosome. Distribution
Other rnicroscleres are choanosomal amphiasters with spinose
rays, measuring about 10.4-14.4 f.Lm long, 9.5-12 f.Lm wide. This Caribbean, New Caledonia.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Azoricidae Sollas, 1888

Andrzej Pisera 1 & Claude Levi2

1 Instytut Paleobiologii, PolskaAkademia Nauk, ul. Twarda 51/55,00-818 Warszawa, Poland. (apis@twarda.pan.pl)
2Museum National d'Histoire Naturelle, Laboratoire de Biologie des Invertebres Marins et Malacologie, 57, Rue Cuvier, 75231 Paris
Cedex OS, France. (levi@mnhnJr)

Azoricidae (Demospongiae, 'lithistids') consists only of two genera, Leiodennatium and Jereicopsis, and at least six species, differenti-
ated on the basis of habitus and microscleres. They occur in the Atlantic and Pacific in deep water. The family is based on the presence of
strongly spinose rhizoclone desmas; microscleres when present are raphides; choanosomal and ectosomal oxeas may also be present.
Future studies may prove it to be polyphyletic and the family may even disappear.
Keywords: Porifera; Demospongiae; 'lithistids'; Azoricidae; Leiodennatium; Jereicopsis.

DEFINITION, DIAGNOSIS, SCOPE Remarks

Synonymy This family is based partly on very few characters, and as such
it may be polyphyletic or may disappear completely with genera
Azoricidae Sollas, 1888: 319; Topsent, I 894d: 291; allocated to other families - although the true relationships are
Schrammen, 1903: 20; Leiodermatidae Lendenfeld, 1903: 145; equivocal. The type of rhizoclones present in both genera suggests
Lendenfeld, 1907: 362; Wilson, 1925: 465 (partim). a relationship with Scleritodermiidae, whereas the absence of ecto-
somal rhabds and presence of raphides in Jereicopsis, and absence
Definition of microscleres and ectosomal rhabds in Leiodennatium, sets them
apart from scleritodermiids. Family Azoricidae was established by
Rhizomorine 'lithistid' Demospongiae with strongly spinose Sollas (1888) who maintained a distinction between the genera
rhizoclone desmas; microscleres if present are raphides. Azorica Carter, 1873c and Leiodermatium Schmidt, 1870.
Lendenfeld (1903) subsequently synonymized the genera, with
Diagnosis Leiodermatium being the most senior available name. As a
consequence of this synonymy and disappearance of the name
Polymorphic. Desmas strongly spinose rhizoclones; no special Azorica Lendenfeld (1903) changed the family name to the
ectosomal spicules; microscleres if present are raphides. Leiodermatidae, followed subsequently by Wilson (1925). This
action is in contravention to the ICZN, and thus Azoricidae is rein-
Scope stated here as the most senior available name. Wilson (1925) also
included Microsclerodenna Kirkpatrick in the family, considered
The family contains only two valid genera: Leiodennatium here synonymous with Taprobane Dendy and allocated to
Schmidt, 1870, and Jereicopsis Levi & Levi, 1983a. Scleritodermidae.

KEY TO GENERA

(1) Foliated of vase shaped in habit, no microscleres ........................................................................................................ Leiodermatium

Massive-conical in habit with bundles of oxeas protruding from the center, microscleres raphides .................................. ]ereicopsis

]EREICOPSIS LEVI & LEVI, 1983 Diagnosis

Synonymy Massive rhizomorine sponge with excurrent system similar

to iereica Zittel; oscules concentrated at the summit, with
Jereicopsis Levi & Levi, 1983a: 129. protruding oxeas; rhizoclones strongly spinose; microscleres are
raphides.
Type species
Remarks
iereicopsis graphidophora Levi & Levi, 1983a: 129 (by
monotypy). This is currently a monotypic genus. Its placement in
Azoricidae is tentative, as it has raphide microscleres, in contrast to
Definition the type genus Leiodermatium which has no microscleres at
all. However, both genera share characters such as the absence of
Massive Azoricidae with raphides as microscleres. ectosomal spicules and very similar spinose rhizoclones.

352
Porifera· Demospongiae • 'Lithistids' • Azoricidae 353

Description of type species sponge summit numerous (up to 30) oscu1ar openings are grouped;
each oscule surrounded by a peculiar modified (elongated) desma
Jereicopsis graphidophora Levi & Levi, 1983a (Figs 1-3). and contains a bundle of 2500-3000 /Lm long and 10-20/Lm thick
Synonymy. Jereicopsis graphidophora Levi & Levi, 1983a: oxeas; there are other, thinner oxeas protruding from the rest of the
129, pI. 3: 1-3; pI. 10: 15. sponge surface. Desmas are massive rhizoclones bearing numerous
Material examined. Holotype: MNHN DCL2789 - New complex spines, usually bifurcating several times. The des mas
Caledonia region, 410-415 m depth. Paratypes: MNHN DCL2790. measure up to 500 /Lm in size. Microscleres are extremely abun-
Description. Small massive sponge in the shape of a trun- dant raphides, often occurring in bundles, measuring 17-37/-Lm
cated cone, 8-12mm high, 15-25mm wide. In the center of the long, 0.5-1.0 /-Lm in thickness.
Remarks. In the canalization and shape this sponge resem-
bles the fossil genus Jereica Zittel, hence the name, but no bundles
of oxeas have been ever reported from the fossil material.

Distribution

Known only from the New Caledonia region.

LEIODERMATIUM SCHMIDT, 1870

Synonymy

Leiodermatium Schmidt, 1870: 22; Zittel, 1878a: 34, 58;

Sollas, 1888: 352; Lendenfeld, 1903: 145 (partim); Wilson, 1925:
465; Van Soest & Stentoft, 1988: 69; Gruber, 1993: 49. Azorica
Carter, 1873c: 439; Zittel, 1878a: 34 (98), 57(121); Sollas, 1888:
319; de Laubenfels, 1955: E49. Heterophymia Pomel, 1872: 143;
Zittel, 1878a: 57; Sollas, 1888: 343. ? Tretolophus Sollas, 1888: 325.

Fig. 1. Jereicopsis graphidophora Levi & Levi. A-D, holotype MNHN

Type species
DCL2789 and paratypes MNHN DCL2790 (scales 5 mm) (modified after
Levi & Levi, 1983a). Leiodermatium lynceus Schmidt, 1870: 22, pI. 3: 2.

Fig. 2. Jereicopsis graphidophora Levi & Levi, MNHN type series. A, oscular region of the choanosomal skeleton (scale 2mm). B, details of oscular
region of desmas skeleton (scale I mm). C, choanosomal skeleton of rhizoc1one desmas (scale 200 ILm). D, details of rhizoc1ones (scale 100 ILm).
354 Porifera· Demospongiae· 'Lithistids'· Azoricidae

Fig. 3. Jereicopsis graphidophora Levi & Levi. MNHN type series. A-B.
raphides; (A. scale 20 fLm; B. scale 5 fLm).
Fig. 4. LeioderTlUltium lynceus Schmidt. A-B, habitus of the holotype
MZUS P0145 (scale 1 em).

Definition Heterophymia heteroformis characters typical of Corallistidae: from

the illustrations of Bowerbank (1869b) it clearly follows that
Ear-shaped to foliated Azoricidae without ectosomal spicules H. heteroformis (Bowerbank) = Dactylocalyx heteroformis
or microscleres. Bowerbank is a rhizomorine sponge. The source of Sollas's (1888)
mistake remains unknown. Based on Sollas's (1888) diagnosis and
Diagnosis description Lendenfeld (1903) considered the species to be a coral-
listid, which is incorrect, choosing it as the typical genus of the
Foliated or vase to ear-shaped in habit; one surface with large family, and rejecting the generic name Corallistes Schmidt, 1870
elevated oscules the other with small pores. Desmas strongly spin- as a junior synonym of Coscinospongia Bowerbank, 1869b.
ose rhizoclones; no special ectosomal spiCUles; no microscleres. Further, he also changed the family name from Corallistidae
Accessory megascleres are large oxeas. Sollas, 1888 to Coscinospongiidae Lendenfeld, 1903. Eleven
species have been included into this genus at various times, but
Remarks probably only six of these belong here. The paucity of characters
makes species differentiation and determination difficult, and the
This genus has been included into various families: Sollas genus badly needs revision. Azorica (=Leiodermatium) has also
(1888) attributed it into his family Azoricidae; Lendenfeld (1903) to been reported from the Middle Jurassic (Moret, 1928) and
his family Leiodermatidae; Van Soest & Stentoft (1988) included it Cretaceous (Moret. 1926). but this attribution was based solely on
in Siphonidiidae Sollas, 1888 without justification. Lendenfeld morphological grounds and thus needs to be verified.
(1903) syonymised it with Azorica Carter, 1873c, followed by
Wilson (1925) and this view is accepted here. De Laubenfels (1955) Description of type species
for unknown reasons attributed Azorica (=Leiodermatium) to
Scleritodermidae. Most probably the genus Tretolophus Sollas, 1888 Leiodermatium lynceus Schmidt, 1870 (Figs 4-5).
is also a junior synonym of Leiodermatium, but the type material Synonymy. Leiodermatium lynceus Schmidt, 1870: 22,
consists only of a very small fragment of choanosomal skeleton pI. 3: 2; Zittel, 1878a, 34 (98), p. 58 (122), pI. 1: 5; Sollas, 1888:
composed of rhizoclone desmas, and this is currently impossible 352; Van Soest & Stentoft, 1988: 69, fig. 33. Not Boury-Esnault
to corroborate. Heterophymia Pomel, 1872 is also included in syn- et al., 1994b: 58.
onymy here. Its holotype (MNHN DCL 3675) is housed in the Material examined. Holotype: MZUS P0145 (slide
MNHN Paris, and has a complex history. Bowerbank (1869b: 85) MNHN DT2205; slides MNHN DCL from MZUS material). Other
described a new species Dactylocalyx heteroformis Bowerbank material. ZMA POR 5255 - Barbados.
which was based on an undescribed specimen from Shanghai, housed Description. Narrow ear-shaped or cylindrical sponge about
at present in MNHN under manuscript name Coscinospongia het- 6cm high, 3 cm wide, with wa1l3-4mm thick, with deeply incised
eroformis Valenciennes, 1872. Using the same specimen, and clearly rounded margin. Outer surface with evenly distributed oscules
unaware of Bowerbank publication, Pomel (1872) decided to change about 500-750 fLm in diameter, located on the top of small, flat eleva-
the manuscript name of Valenciennes, as not entirely appropriate tions. Inner surface even with densely distributed pores 156-188 fLm
[sic] to Heterophymia valenciennessi. On the other hand, Sollas in diameter. Choanosomal skeleton consists of rhizoclones, which
(1888: 343) confused the issue further by accepting Pomel's generic are strongly branched and supplied with numerous long branched
name Heterophymia but reinstating Bowerbank's specific name het- spines, tips of which are divided into small spine-like processes.
eroformis. It was Sollas (1888) who also first ascribed to the species No other megascleres or microscleres have been found in the
Porifera· Demospongiae· 'Lithistids'· Azoricidae 355

Fig. 5. Leiodennatium lynceus Schmidt, MZUS P0145. A, outer surface of the choanosomal skeleton (scale I mm). B, inner surface of the choanosomal
skeleton (scale I mm). C, choanosomal skeleton of rhizoc1one desmas (scale 200 j.Lm). D, details of rhizoc1ones (scale 20 j.Lm).

holotype, but Van Soest & Stentoft (1988) also reported accessory Distribution
oxeas in their material from Barbados.
Remarks. There are numerous reports of the species L lynceus, This genus has been widely reported from both the Atlantic
but usually these are specimens without large oscules on the outer side, and Pacific oceans, from deep water, as most other 'lithistids', but
suggesting it might be a different and most probably new species. This as it badly needs revision a synthesis of its published distribution
material is not included here in the concept of this species. would be misleading.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Desmanthidae Topsent, 1893

Andrzej Pisera1 & Claude Levi2

1 Instytut
Paleobiologii, Polska Akademia Nauk, ul. Twarda 51155,00-818 Warszawa, Poland. (apis@twarda.pan.pl)
2 Museum National d'Histoire Naturelle, Laboratoire de Biologie des Invertebres Marins et Malacologie, 57, Rue Cuvier, 75231 Paris
Cedex 05, France. (levi@mnhn.fr)

Desmanthidae Topsent (Demospongiae, 'lithistids') contains four genera, Desmanthus, Paradesmanthus, Sulcastrella, and provisionally
at least, Petromica, together with about eleven species world-wide, both from shallow and deep water. These are mostly encrusting, rarely
massive sponges, characterized by monaxial complex branching desmas, often loosely articulated, and styles and/or oxeas which are per-
pendicular to the sponge surface and may occur in bundles. No microscleres are present, except in one species. Genera are differentiated
on the base of habitus and desma shape, and presence of microscleres.
Keywords: Porifera; Demospongiae; 'lithistids'; Desmanthidae; Desmanthus; Paradesmanthus; Sulcastrella; Petromica .
. _ - - - _ .. __..- '--"-'--"'--' - _ .. _ - . _ .. _ - - - - - - - - - -

DEFINITION, DIAGNOSIS, SCOPE (1936a) was clearly unaware of the taxon Desmanthidae Topsent,
1893 when he independently proposed the same name for
Synonymy Desmanthus and some other unrelated demosponges. Van Soest &
Hajdu (2000), in their review of Desmanthidae, also include
Desmanthidae Topsent, 1894d: 290; Lendenfeld, 1903: 144; Lithobubaris Vacelet, 1969 as a valid genus, whereas we consider
Wilson, 1925: 470; Van Soest & Hajdu, 2000: 302. Desmanthidae here that it is a younger synonym of Sulcastrella Schmidt.
de Laubenfels, 1936a: 126 (partim). Petromicidae Topsent, 1928: We also include the genus Petromica in Desmanthidae, provi-
105. Monanthidae de Laubenfels, 1936a: 139 (partim). sionally at least, which is set slightly apart from other genera of
this family (its desmas are quite different from those of other gen-
Definition era of the family), but this placement is equivocal and dependent
on resolving clearer relationships between all families and genera
'Lithistid' Demospongiae encrusting, rarely massive in habit; attributed to Halichondrida, Axinellidae and also several groups
choanosomal skeleton composed of branching desmas supposedly within Hadromerida. The precise affinities of this genus require
monocrepidial or clearly monocrepidial; other megascleres are additional datasets, such as molecular sequences to resolve the
oxeas and/or styles. Microscleres in one genus only, slightly problem, but for the matter of convenience we prefer to include it
resembling sanidaster. in Desmanthidae until a final decision is reached. Petromica was
considered by Van Soest et al. (1990) and Muricy et al. (2001) to
Diagnosis belong to Halichondriidae on the basis that loose spicules are the
most important for determining sponge systematics, with des mas
Encrusting, rarely massive in habit; basal skeleton composed considered to be only accessory spicules - a view which is not sup-
of clearly monaxial (or supposedly monaxial) desmas of complex ported here. Their reasoning was also based on the skeletal organi-
branching pattern to trider-like morphology, accompanied by oxeas sation of Monanthus ciocalyptoides Van Soest & Zea, 1986, which
and styles/tylostyles perpendicular to the sponge surface. One has a clearly halichondriid structure. This species was later trans-
genus has large microscleres slightly resembling sanidasters. ferred to Petromica based on the proposed synonymy between
Monanthus and Petromica (Van Soest et al., 1990), but this pro-
Scope posal is also questioned here because neither Petromica grimaldi
nor Monanthus plumosus (their respective type species) have con-
Desmanthus Topsent, 1893; Pardesmanthus gen. nov.; fused ectosomal skeletons and/or confused-reticulate choanosomal
? Petromica Topsent, 1904; Sulcastrella Schmidt, 1879. skeletons of monaxons (otherwise indicating affinities with the
halichondriids). In support of our position we cite the existence of
Remarks plumose columns of oxeas and/or styles also occurring in sclerito-
dermids, for example, and more especially the heterogeneous
This family presently contains about eleven species although range of desma types observed in putative species of Petromica and
some of these may not truly belong here. For example, Topsent Monanthus. Thus, while 'Monanthus' ciocalyptoides may indeed
(1904) suggested that in Desmanthus desmas may be really appear to be closely related to the halichondriids we consider, at
tetracrepidial (i.e., not monocrepidial or acrepidial), and similarly, least provisionally, the type species and at least several other
Schmidt (1879) reported that desmas of Sulcastrella were tetraxial species of Petromica, and also the type species of Monanthus
(although we were not able to observe this), and consequently we (as its synonym), to belong to Desmanthidae (see chapter on
reserve our decision of this assessment at present. De Laubenfels Halichondriidae by Erpenbeck & Van Soest).

356
Porifera· Demospongiae • 'Lithistids' • Desmanthidae 357

KEY TO GENERA

(1) Massive in habit, desmas branching in various planes ......................................................................................................... Petromica

Encrusting in habit ............................................................................................................................................................................... 2
(2) With microscleres resembling sanidasters ................................................................................................................. Paradesmanthus
No microscleres ................................................................................................................................................................................... 3
(3) Desmas of the outer layer trider-like ................................................................................................................................ Desmanthus
Desmas hook-like, with pointed spines .............................................................................................................................. Sulcastrella

DESMANTHUS TOPSENT, 1894 revealed no crepis at all. He also mentioned the presence of lopho-
triaenes, but examination of the type material, again, revealed that
Synonymy he mistook these for typical Desmanthus-like desmas. Six
described species are included here. The species described by Uriz
Desmanthus Topsent, 1894d: 311; Lendenfeld, 1903: 144. (1988) is excluded as so-called desmas (II) are microscleres
Gruber, 1993: 51; Van Soest & Hajdu 2000: 302. Lophacanthus slightly resembling sanidaster, which are absent in Desmanthus;
Hentchel, 1912: 306. we propose for it a new genus Paradesmanthus (see below).

Type species Description of the type species

Aciculites incrustans Topsent, 1889: 32 (by subsequent Desmanthus incrustans (Topsent) (Figs 1-2).
designation; Topsent, 1894d) Synonymy. Aciculites incrustans Topsent 1889: 32, fig. 1;
Desmanthus incrustans Topsent: 1894d: 311; Lendenfeld, 1903:
Definition 144; Vacelet et al. 1976: 29, pI. 1: 1, fig. 11; Pulitzer-Finali, 1983:
481, fig. 18; Levi & Levi 1989: 50, fig. 19. Gruber, 1993: 51,
Thinly encrusting Desmanthidae with trider-like desmas pI. 15: 4-6; Van Soest & Hajdu, 2000: 304, fig. 2A-F, table. 1.
(but supposedly monocrepidial). Material examined. Holotype: MNHN DT 232 (slides DT
1853) - Campeche Bank, Caribbean. Other material. RMI
Diagnosis 74.7.23.3 - Barbados.
Description. Thinly encrusting sponge with a strongly
Encrusting in habit, desmas trider-like but without crepis visi- hispid outer surface. Basal skeleton of choanosomal desmas which
ble (monocrepidial) with long conical outer ray (usually sculptured); are of two types (all with intennediates): the uppennost are trider-
other megascleres are radially arranged styles; no special ectoso- like, tri- to tetrapodial, branched, and bearing a strong and long,
mal spicules, no microscleres. upwardly directed conical spine. The desma surface is tubercu-
lated; the more deeply located young desmas are less regular
Remarks rhizoclone-like, branched several times in one plane and also
tuberculated. Articulation of desmas is terminal or terminal to
Topsent (1889) described desmas of D. incrustans as shoulder. Perpendicular to the choanosomal skeleton are long
tetracrepid. Similarly, Van Soest & Hajdu (2000) suggested that the styles protruding from it, producing the hispid aspect of the
outennost layer of desmas in this species bears rhabdomes, but this sponge; they are 330-1000 f.Lm long, 13-26 f.Lm thick. No
interpretation is clearly erroneous. Examination of numerous microscleres of any kind.
desma preparations of D. incrustans revealed no crepis at all, and Remarks. According to Van Soest & Hajdu (2000) world-
although they have a relatively large spine this does not constitute a wide reports of D. incrustans most probably indicate that it is in
rhabdome. Trider-like morphology of the desmas in Desmanthus fact a complex of several closely related species.
are thus considered here to be superficial, and most probably they
are of the same type as in other genera of Desmanthidae (i.e., Distribution
monocrepidial). Interpretation of the large spine in the outennost
layer of desmas as rhabdome is in our opinion erroneous. The des- The type species has been reported from the Caribbean,
mas of Desmanthus incrustans seem very different, at first glance, Mediterranean, Indian Ocean, Philippines and Papua New Guinea.
from other desmas in the family by having the appearance of The type material comes from the Caribbean from depths of 12-
triders. Such desmas are characteristic, however, only of the 30 m from cryptic habitats, with other undoubted occurrences from
uppennost layer of the sponge, and its long conical outgrowth Jamaica and Barbados (depth about 90m). The genus has been
directed outward may be interpreted as a major spine and not a reported from Caribbean, Southern Atlantic, Mediterranean Sea,
real clone. The des mas occurring below this outer layer, as well Madagascar, Indian Ocean, Pacific coast of Panama, Philippines,
as younger desmas, resemble rhizoclones branched in one Papua New Guinea.
plane and are similar in geometry to the desmas of Sulcastrella
clausa.
Lophacanthus Hentschel, 1912 (type species L. rhabdophorus PARADESMANTHUS GEN. NOV.
Hentschel, 1912) has spiculation identical to Desmanthus and they
are clearly synonymous, following Van Soest & Hajdu (2000). Type species
Hentschel (1912) described the desmas of Lophacanthus as tetracre-
pidial, but re-examination of slides from the original material Desmanthus macphersoni Uriz, 1988 (here designated).
358 Porifera· Demospongiae· 'Lithistids'· Desmanthidae

Fig. 1. Desmanthus incrustans (Topsent), RMI 74.7.23.3. A, view of the outer surface with styles protruding from the ectosome (scale 200 jl.m). B, outer
surface of the choanosomal skeleton showing trider-like desmas (scale 100 jl.m). C, details of outer layer of choanosomal des mas (scale 50 jl.m). D, lower-
most layer of choanosomal desmas: view from below (scale 100 jl.m).

Remarks

Uriz (1988) considered her described microscleres as a second

type of desma, but this is not true. Considering their completely dif-
ferent morphology, the size of these spicules, their location in the
ectosomal region, and no traces of articulation suggest that they are
microscleres, but of exceptional size. The presence or absence of
microscleres, or their different types, are usually considered a generic
character, and for this reason we propose here a new genus for the
species described by Uriz (1988) as Desmanthus macphersoni.

Description of type species

Paradesmanthus macphersoni (Uriz) (Figs 3-4).

Synonymy. Desmanthus macphersoni Uriz 1988: 30, fig. 8,
pI. 4a; Desmanthus (?) macphersoni Van Soest & Hajdu, 2000: 308.
Material examined. Paratype: CEAB POR Bio182.
Fig. 2. Desmanthus incrustans (Topsent), RMI 74.7.23.3. A-B, styles Description. Thick (up to 8 mm) encrusting sponge with
(scale 100 jl.m). C-D, details of head and points of styles (scale 20 jl.m).
strongly hispid outer surface. Oscula 1-1.5 mm wide. Choanosomal
skeleton reticulate, displaying large, more-or-less oval openings,
Definition which are up to 3 mm long and 1 mm wide. Choanosomal desmas
irregular trider-like (but no crepis visible), tri- to tetrapodial, tubercu-
Encrusting Desmanthidae with trider-like (but monocrepidial) late, and bearing long irregular conical spine directed outward.
desmas, and large microscleres resembling sanidasters in the Articulation of desmas is terminal, or terminal to shoulder.
ectosomal region. Perpendicular to the choanosomal skeleton are long styles protruding
from it, producing the hispid aspect of the sponge; they are 545-700
Diagnosis f.Lm long and 27-42 f.Lm thick. Microscleres are exceptionally large,
occurring in great numbers in the ectosomal region. They are in the
Encrusting Desmanthidae with trider-like (but monocrepidial) form of rhabds, straight to bent in the middle, with numerous side
des mas and large microscleres in the form of straight to bent in the outgrowths that slightly resemble sanidasters; they bear low mas-
middle rhabds with numerous side outgrowths (resembling sive spines on their distal parts; they are 85-180 f.Lm long and
sanidaster); they bear low massive spines on their distal parts. 50-110 thick (including side outgrowths).
Porifera· Demospongiae • 'Lithistids' • Desmanthidae 359

Fig. 3. Paradesmanthus macphersonii (Uriz) , paratype CEAB POR Bio182. A, outer surface view (scale 1 mm). B, upper surface of the choanosomal
skeleton (scale 2mm). C, upper layer of choanosomal skeleton (scale 200 f,Lm). D, details of choanosomal desmas (scale 200 f,Lm).

Fig. 4. Paradesmanthus macphersonii (Uriz), paratype CEAB POR Bio182. A-B, styles (scale 100 f,Lm). C-I, microscleres (scale 50 f,Lm).

Remarks. The spicules described as microscleres and PETROMICA TOPSENT, 1898

occurring in great numbers in the ectosomal region of this sponge
are much larger than typical microscleres; they slightly resemble Synonymy
sanidasters although this interpretation is not certain.
Petromica Topsent, 1898: 226, 1904: 64. Dendy 1905:
Distribution 104. Dendy, 1921: 8; Topsent, 1928: 106; Van Soest et al.,
1990: 41; Gruber, 1993: 47; Boury-Esnault et al., 1994b:
This genus and species has been reported only from South 57; Muricy et ai., 2001: 105. Monanthus Kirkpatrick,
Atlantic (off Namibia) from the depth of 160 meters. 1903a: 176.
360 Porifera· Demospongiae· 'Lithistids'· Desmanthidae

Type species a very well developed skeleton composed of strongly fused rhizo-
clones. Conversely, these authors argued that the family
Petromica grimaldii Topsent, 1898: 226 (by monotypy). Desmanthidae Topsent was supposedly characterized by tetraxial
desmas in Desmanthus Topsent. We reject this proposal where
Definition careful re-examination of various Desmanthus material showed no
tetracrepidial desmas, with Petromica displaying clearly monaxial
Massive Desmanthidae with desmas branching in various desmas and therefore included here, provisionally at least, in
planes. Desmanthidae (see remarks for the family). Monanthus
Kirkpatrick, 1903a, as defined by its type species at least, is con-
Diagnosis sidered a junior synonym of Petromica Topsent because it has iden-
tical spiculation (including the type of desmas) and skeletal
Massive sponges with irregular monocrepid desmas (some structure. This genus has recently been revised by Muricy et al.
without a visible crepis) which are branched in all planes; smooth (2001) who considered it Halichondriidae. They present detailed
except at tips; and poorly articulated or non-articulated at all; other discussion of its scope, phylogeny and biogeography. Seven
megascleres are large, usually bent, oxeas, strongyloxeas and species are presently recognized.
anisorhabds. No microscleres.
Description of type species
Remarks
Petromica grimaldii Topsent (Figs 5-7).
This genus was originally included in Azoriciidae Synonymy. Petromica grimaldi Topsent, 1898: 226, fig. la;
(=Leiodermatidae) (Topsent, 1898, 1904), or even considered a Topsent, 1904: 64, pI. 5: 2-4, pI. 8: 6; Pulitzer- Finali, 1970: 334;
synonym of Leiodermatium (Lendenfeld, 1903), but this proposal Boury-Esnault, 1994b: 57; Murlcy et al., 2001: 105, Figs la, 2a--c, 3.
is unjustified because Azoricidae has no styles and usually displays Leiodermatium grimaldii; Lendenfeld, 1903: 145.
Material examined. Holotype (Topsent slide): MNHN DT
850 - Azores. Other material. MNHN unregistered specimen -
Azores, stn. 587, 'Le Prince de Monaco'.
Description. Massive sponge with conical form, up to 4 cm
high, 3 cm wide; surface very irregular covered with conules; pores
dispersed. Each conule is supported by bundles of oxeas, strongy-
loxeas and anisorhabds. No special ectosomal spicules.
Choanosomal desmas measure up to 570/-Lm in size, are very
loosely interwoven resulting in a very supple skeleton; desmas are
very irregular and branched in all planes, generally smooth except
at tips which are divided into massive low spines. The desmas lack
a visible crepis and display no zygomes, except those which encir-
cle oxeas/styles, no other desmas. Other choanosomal megascleres
are large, usually bent anisoxeas to styles (or strongyloxeas) or
even rhabds, occurring in bundles supporting the conules on the
surface and in the choanosome as well. Oxeas measure 956-1250 /-Lm
long, 23-30/-Lm thick; styles and rhabds are 700--1280/-Lm long,
23 /-Lm thick.
Remarks. In the slide preparations from the original mate-
Fig. 5. Petromica grimaldii Topsent, MNHN unregistered specimen of rial we also found some oxyasters, measuring 22-72 /-Lm in size,
Topsent (1904, pI. 5: 4) (scale lOmm). but these are relatively rare and likely to be contaminants.

Fig. 6. Petromica grimaldii Topsent, MNHN unregistered specimen of Topsent (1904, pI. 5: 4). A, outer surface view, note bundles of anisoxeas to
anisorhabds (scale I mm). B, view of the choanosome, with both desmas and anisoxeas visible (scale 500 fLm).
Porifera· Demospongiae • 'Lithistids' • Desmanthidae 361

Fig.7. Petromica grimaldii Topsent. MNHN unregistered specimen of Topsent (1904, pI. 5: 4). A, choanosornal anisoxea (scale 200 /lorn). B, points of the
same anisoxea (scale 20/lorn). C, ends of anisorhabd (scale 20/lorn). D, strongyloxea (scale 200/lorn). E, ends of the same strongyloxea (scale 20/lorn).
F-I, choanosornal desrnas (scale 200 /lorn). J, tips of desrna branch (scale 20/lorn).

Distribution Remarks

The type species is known from the Azores, NE Atlantic, from The morphology as well as organization of desmas and styles
200-599 m depth. The genus is distributed in the Azores, Sri Lanka is very similar in Sulcastrella and in Lithobubaris Vacelet 1969,
(Ceylon), South-West Africa, probably Caribbean. (with holotype MNHN DN7 re-examined), such that we consider
both genera to be synonymous. According to Schmidt (1879) des-
mas of Sulcastrella display triradiate crepis, but we were not able
SULCASTRELLA SCHMIDT, 1879 to verify this as only SEM studies have been carried out, while des-
mas in Lithobubaris display no crepis at all. Three species are
Synonymy attributable to the genus Sulcastrella, as here understood. The
genus Desmascula de Laubenfels, 1950a, (type species D. desde-
Sulcastrella Schmidt, 1879; Lendenfeld, 1903: 134. mona de Laubenfels, 1950a: 121) is possibly a junior synonym of
Lithobubaris Vacelet, 1969: 184; Van Soest & Stentoft, 1988: 72; Sulcastrella Schmidt, based on desmas and styles (reported oxeas
Gruber, 1993: 51; Van Soest & Hajdu, 2000: 308. may come from Spheciospongia), or less probably some rhi-
zomorine lithistid. Strange microsc1eres in D. desdemona are the
Type species problem because the only known specimen of Sulcastrella was
dead, thus nO microsc1eres have been ever found. However if One
Sulcastrella clausa Schmidt, 1879: 27 (by monotypy). can ascribe microsc1eres to Rhabderemia, than perhaps they are
contamination, or Sulcastrella has affinities with Rhabderemia.
Definition

Thickly encrusting Desmanthidae; choanosomal desmas Description of type species

hook-like.
Sulcastrella clausa Schmidt (Figs 8-9).
Diagnosis Synonymy. Sulcastrella clausa Schmidt, 1879: 27, pI. 1: 5,
pI. 2: 6, pI. 3: 7; Lendenfeld, 1903: 134.
Ectosomal spicules are radically arranged styles; choanoso- Material examined. Holotype: MCZ 6583 (orig. no. 289)-
mal spicules as peculiar spinose desmas with hook-like rays. Sand Key, Florida, 236 m depth.
362 Porifera' Demospongiae • 'Lithistids' • Desmanthidae

Fig. 8. Sulcastrella clausa Schmidt, holotype MCZ 6583(289). A, upper surface view of the holotype (scale 10mm). B-D, choanosomal desmas (scale
100 11m). E, style head and point (scale 50 11m).

Fig.9. Sulcastrella clausa Schmidt, holotype MCZ 6583(289). A, surface of the choanosomal skeleton (scale I mm). B, vertical section of choanosomal skele-
ton with perpendicular styles visible (scale 200 11m). C,details of choanosomal desma skeleton (scale 200 11m). D, articulation of desmas (scale 100 11m).

Description. Thick (about 2-2.5 mm) encrusting in habit, Remarks. The holotype is a deciduous specimen, but micro-
28 X 35 mm in size; surface raised into low convex areas, each of scope examination revealed the remnants of soft tissue with numer-
which is occupied by a stellate system of anastomosing subdermal ous styles, previously not reported from Sulcastrella, which are
canals radiating from the center. Choanosomal skeleton consists of still attached to the choanosomal skeleton.
strongly articulated massive (probably monaxial) desmas about
200-300 /-Lm in size. They are irregularly branched with clads which Distribution
are hook-like, and with strong and high conical spines; articulations
mostly terminal with the aid of spines. Other megascleres are slightly The type species is known only from Florida. The genus is
bent styles which are perpendicular to the sponge surface and pro- distributed in Florida, Barbados and the Mediterranean Sea.
truding from the choanosomal skeleton; their length is at least 300
/-Lm (entire length was not observed) and about 15 /-Lm thick.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Vetulinidae Lendenfeld, 1903

Andrzej Pisera1 & Claude Levi2

1 Instytut Paleobiologii, PolskaAkademia Nauk, ul. Twarda 51155,00-818 Warszawa, Poland. (apis@twarda.pan.pl)
2Museum National d'Histoire Naturelle, Laboratoire de Biologie des Invertebres Marins et Malacologie, 57, Rue Cuvier, 75231 Paris
Cedex 05, France. (levi@mnhnJr)

Vetulinidae Lendenfeld (Demospongiae, 'lithistids'), including Anomocladina Sollas and Anomocladidae Zittel, contains only one genus
and species, Vetulina stalactites Schmidt, 1879. Undoubted occurrences are only from the Caribbean (Barbados) from 126--601 m depth.
This species has neither ectosomal spicules nor microscleres, the only distinguishing character being anaxial and polyclad desmas
(astroclones!sphaeroclones). It is related to the common Mesozoic sphaerocladine lithistids, probably being the only extant species of this
lineage.
Keywords: Porifera; Demospongiae; 'lithistids'; sphaerocladine demosponges; Vetulinidae; Vetulina.

DEFINITION, DIAGNOSIS, SCOPE Definition

Synonymy As for the family.

Vetulinidae Lendenfeld, 1903: 149; Wilson, 1925: 470. Diagnosis

Anomocladina Sollas, 1885a: 492 (partim). Anomocladidae sensu
Sollas, 1888: 354 (partim). Flabellate or vase shaped; no special ectosomal spicules;
endosomal spicules as acrepid polyaxial desmas (sphaeroclones to
Definition astroclones); no microscleres.

'Lithistid' Demospongiae with acrepid polyaxial (astro- or Remarks

sphaeroclone) desmas. No other megascleres or microscleres.
So far only the type species is known. Both Van Soest &
Diagnosis Stentoft (1988) and Gruber (1993) considered that the genus was
close to Siphonidium and/or Leiodermatium; the present authors do
Monospecific (see type species description). not support this view because it has clearly unaxial or polyaxial
sphaeroclone or astroclone-like desmas. We consider Vetulina as
Scope close to the fossil Sphaerocladina Schrammen, supporting the view
presented already by Sollas (1885a, 1888). De Laubenfels (1955b)
Only one extant genus and species, Vetulina stalactites included this genus into his new family Chiastoclonellidae which
Schmidt, 1879 has been attributed to this family, but in the fossil encompasses both some Mesozoic sphaerocladine sponges and the
record there are numerous lithistids with sphaeroclone and/or astro- Paleozoic genus Chiastoclonella Rauff, 1894. Unfortunately, the
clone desmas known since Paleozoic (see chapter on fossillithistids). genus Chiastoclonella has desmas which are of a different and
unrelated type (chiastoclones - see Reid, 1970) than those (sphae-
Remarks roclones) occurring in other genera here included. Consequently,
the family Chiastoclonellidae de Laubenfels should be abandoned
The Recent family Vetulinidae Lendenfeld, 1903 corresponds as it contains various unrelated sponges. Moret (1926) reported this
to some Mesozoic representatives of the fossil suborder genus from the Upper Cretaceous of France, but this assignment
Sphaerocladina Schrammen. was made mostly on the basis of similarities in habitus, and thus
seems doubtful.

VETUUNA SCHMIDT, 1879 Description of type species

Synonymy Vetulina stalactites Schmidt, 1879 (Figs 1-2).

Synonymy. Vetulina stalactites Schmidt, 1879: 19,
Vetulina Schmidt, 1879: 19; Sollas, 1885a: 486; Sollas, 1888: pI. 1: 1, pI. 2: 9; Sollas, 1885a: 486: pis 3, 4; Sollas, 1888: 354.
354; de Laubenfels, 1955b: E63; Van Soest & Stentoft, 1988: 71. Lendenfeld, 1903: 150; de Laubenfels, 1955b: E63, fig. 46: 4;
Gruber, 1993: 49. Van Soest & Stentoft, 1988: 71, text-fig. 43; Gruber, 1993: 49,
pI. 14: 4-8.
lYPe species Material examined. Syntypes: MCZ 6640, MCZ 6642,
MCZ 6644 - Barbados 183-329m depth, 'Hassler' Expedition.
Vetulina stalactites Schmidt, 1879: 19 (by monotypy). MNHN DT 2204 (slide from Schmidt specimen). Other material.

363
364 Porifera· Demospongiae • 'Lithistids' • Vetulinidae

Fig. 1. Vetulina stalactites Schmidt (scale 10mm). A-B, 'co-type' , specimen MCZ 6640, Barbados, outer and inner surface view respectively.
C-D, 'co-type' , specimen MCZ 6642, Barbados, outer and inner surface view respectively.

Fig. 2. Vetulina stalactites Schmidt. A-B , Inner surface view of the choanosomal skeleton with exhalant pores, MCZ 6640, Barbados (scales: A, 500 !Lm;
B, 50 !Lm). C-D, outer surface view of the choanosomal skeleton, MCZ 6642 Barbados (scales: C, 500 !Lm; D, 200 !Lm). E, inner part of choanosomal skeleton,
MCZ 6642 Barbados (scale 100 !Lm). F, detail of articulation of the desma in the interior of the choanosomal skeleton, MCZ 6642 Barbados (scale 50 !Lm).
Porifera· Demospongiae • 'Lithistids' • Vetulinidae 365

ZMA POR 5254 - Barbados. MZUS P0154 and P0155 - dry skeleton: inside the wall they are more regular, as described above;
fragmented specimens. those occurring on the inner surface are less regular, having the
DescriptWn. Flabellate or sinuous plate, or vase shaped, globular center less clearly developed and bearing numerous very
sometimes with tubular portions; surfaces even, although outer sur- long processes; those occurring on the opposite outer surface are
face with well developed concentric growth lines; walls 0.7-1 cm also less regular than those from the center of the wall, and their
thick. Pores on the outer surface evenly distributed but without any centers are flattened in the form of a thick irregular, strongly tuber-
order, about 0.1 mm in diameter and irregular. Oscules (?) on the culated plate.
inner surface have the same pattern but larger, about 0.2-0.3 mm in Remarks. Undoubted reports of the type species are only
diameter, and less densely placed, more-or-Iess rounded. Desmas from Barbados, 153-601 m depth (Schmidt, 1879) and 153-216m
with irregularly globular center, 0.05-0.07 mm in diameter, from depth (Van Soest & Stentoft, 1988). The unlabelled specimen in the
which emanate 2-8 rays terminating in expanded zygomes; these Bristol Museum described by Sollas (1885a) should be, according
rays may be sparsely tuberculated and/or spinose. The rays origi- to him, from shallow water. Sollas (1885a, 1888) reported the pres-
nate usually only from the lower part of the globular center, while ence of cylindrical styles, which have not been found by the pres-
its upper portion displays numerous tubercles and long, branching, ent authors either in the specimens from Harvard or in those from
spine-like processes, which in tum may bear short and wide spines Strasbourg. We observed some fragments on the inner surface of
on their tips. In some regions of the skeleton some desmas have specimen MCZ 6640 containing numerous, irregularly distributed,
a hollow centre instead of being globular in this part. These desmas, finely spinose microxeas about 100---120/Lm long. Most probably
150---220/-lm in diameter, are arranged in such a way that the part all these spicules are contaminants as they were not found in other
devoid rays is directed toward the surface of the sponge. material examined.
Articulation is realized in such a fashion that the rays of several des-
mas meet the upper portion, devoid of rays, of the globose center, Distribution
articulating with its processes and with rays of other desmas.
There is often regional differentiation between desmas in the Known only from the Carribean.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcaderniclPlenum Publishers, New York, 2002

Family Phymatellidae Schrammen, 1910

Andrzej Pisera 1 & Claude Levi2

I Instytut Paleobiologii, Polska Akademia Nauk, ul. Twarda 51155,00-818 Warszawa, Poland. (apis@twarda.pan.pl)
2Museum National d'Histoire Naturelle, Laboratoire de Biologie des Invertebres Marins et Malacologie, 57, Rue Cuvier, 75231 Paris
Cedex 05, France. (levi@mnhn.fr)

Phymatellidae Schrammen (Demospongiae, 'lithistids') contains three extant genera, all valid, with four species occurring mostly in the
Pacific. It is known since the Late Jurassic with numerous representatives in the fossil record. Recent species live in deeper water and are
lamellar cup-shaped, club-shaped or spherical with a stem. Desmas are smooth to sparsely tuberculated, more or less irregular large tetra-
clones, which are usually poorly articulated. Ectosomal megascleres are smooth dichotriaenes; other megascleres may be oxeas.
Microscleres are ectosomal robust microspined streptasters/amphiasters and choanosomal microspined streptasters/amphiasters with long
slender rays. Genera are differentiated mostly by their habitus.
Keywords: Porifera; Demospongiae; 'lithistids'; Phymatellidae; Neoaulaxinia; Reidispongia; Neosiphonia.

DEFINITION, DIAGNOSIS, SCOPE Scope

Synonymy Four Recent nominal genera are attributed, but only three,
Neoaulaxinia nom. nov. (pro Recent Aulaxinia Levi & Levi, 1988),
Phymatellinae Schrammen, 1910: 72. Phymatellidae Reidispongia Levi & Levi, 1988 and Neosiphonia Sollas, 1888 are
Schrammen, 1924a: 37 (see also fossil Phymatellidae); Levi, considered valid and they are represented by five species, mostly
1991: 79. from the Pacific.

Definition Remarks

'Lithistid' Demospongiae with tetraclone desmas and This family has several fossil genera and is known mostly
ectosomal dichotriaenes. from the Upper Cretaceous, its representatives were reported
already from the Late Jurassic (Schrammen, 1937; Pisera, 1997). It
Diagnosis is based on the fossil genus Phymatella Zittel, 1878a (type species
Eudea intumescens Roemer, 1864, Upper Cretaceous), and was
Polymorphic; clavate, lamellar, spherical with a stem. Large proposed first by Schrammen (1910) as subfamily Phymatellinae,
regular tetraclone desmas (usually smooth) as choanosomal and finally raised to the family level by Schrammen (1924a).
spicules; ectosomal spicules smooth dichotriaenes; microscleres Phymatellidae were first applied for living genera from the New
are amphiasters (streptasters). Caledonia region by Levi & Levi (1988).

KEY TO EXTANT GENERA

(1) Lamellar or cup-shaped in habit ...................................................................................................................................... Reidispongia

Nonlamellar in habit ............................................................................................................................................................................ 2
(2) Clavate with shallow apical cavity .................................................................................................................................. Neoauloxinia
Spherical with stem and a bundle of vertical canals in the middle ................................................................................... Neosiphonia

NEOAUIAXINIA GEN. NOV. Definition

Synonymy Club-shaped Phymatellidae with numerous longitudinal canals

on the surface of choanosomal skeleton and shallow apical cavity.
Aulaxinia Levi & Levi 1988: 242 (partim, for Recent forms).
Diagnosis
Type species
Elongated cylindrical to apple-like in habit, with short stem and
Aulaxinia clavata Levi & Levi, 1988: 243. apical cavity of various depth; numerous subectosomallongitudinal

366
Porifera· Demospongiae • 'Lithistids' • Phymatellidae 367

canals converging toward the apical cavity visible on the surface of are also simple triaene-like spicules. The surface of the choanosomal
choanosomal skeleton. Desmas are smooth or sparsely tubercu- skeleton displays irregular longitudinal exhalant canals, which are
lated; microscleres as amphiasters/streptasters. partly open; in cross section one may see that there are more such
canals which run entirely within the choanosomal skeleton, but very
Remarks close to its surface. These canals converge at the sponge summit

The type species of the new genus here proposed was origi-
nally described by Levi & Levi (1988) as Recent representative of
the fossil genus Aulaxinia Zittel. Because of technical problems
with the proper description of genus characters based on fossil
material, where often desmas are not known from the holotype, and
microscleres always absent, we decided to adopt the Sollas' (1888)
approach to the problem: using separate generic names for fossil
and Recent forms, with the use of the prefix Neo- to stress the rela-
tionship of both taxa.

Description of type species

Neoaulaxinia clavata Levi & Levi, 1988 (Figs 1-5).

Synonymy. Aulaxinia clavata Levi & Levi, 1988: 243, pI. 1,
fig. 1.
Material examined. Holotype: MNHN DCL 3636.
Paratypes: MNHN DCL3537 - all from Norfolk Ridge, south of
New Caledonia.
Description. Club-shaped sponges, with the lower more
cylindrical part very long, and the upper clavate part shorter, 30-
60mm high, 10-30mm in diameter; surface is smooth except for
some protruding ectosomal dichotriaenes. The ectosomal dichotri-
aenes may occur both as groups clearly protruding above the sur-
face, but also embedded in the ectosome. They have smooth, long, Fig. 1. Neoaulaxinia clavata (Levi & Levi) (scale 20 mm) (modified after
slender and pointed clads: cladome 250-600 /-Lm in diameter; rhab- Levi & Levi, 1988). A, specimen MNHN DeL 3536 (holotype). B, speci-
dome is shorter, narrow conical, 200-480 /-Lm long. Sometimes there men MNHN DeL 3537 (paratype).

Fig. 2. Neoaulaxinia clavata (Levi & Levi) MNHN DeL 3537 (paratype). A, outer surface with protruding ectosomal dichotriaenes and thick crust of
robust amphiasters (scale 200 fJ,m). B, outer surface of choanosomal skeleton of desmas (scale bar 2mm). e, details of choanosomal skeleton oftetracJone
desmas (scale 200 fJ,m). D, tetracJone desmas articulating with each other (scale 100 fJ,m).
368 Porifera· Demospongiae • 'Lithistids' • Phymatellidae

Fig. 3. Neoaulaxinia clavata (Levi & Levi) MNHN DeL 3537 (paratype). A-F, variability of massive ectosomal amphiasters (scale 5 f1m).

Fig. 4. Neoaulaxinia clavata (Levi & Levi) MNHN DeL 3537 (paratype). A-F, variability of large choanosomal streptasters/amphiasters with long and
slender rays (scale 20 f1m).

in the shallow central cavity. Choanosomal skeleton consists of

tetraclone desmas which are sparsely covered with prominent,
somewhat mushroom-shaped tubercles. Articulation chiefly
terminal and rather weak. The size of des mas varies between
600--1000 /-Lm. Other choanosomal megascleres are oxeas
0.7-4.0mm long, 10--50/-Lm thick, perpendicular to the surface,
often in bundles in the upper part of the sponge. Microscleres are of
three types: ectosomal ones are robust amphiasters with rounded
tips, and microspined rays, 13-19 /-Lm long, 10--14 /-Lm wide;
choanosomal microscleres are streptasters to amphiasters with long
slender and microspined rays which have pointed tips, in two size
categories, with the larger ones 37-50/-Lm long, 36-46/-Lm wide
and having less numerous rays, and smaller ones 21-29 /-Lm long,
18-26/-Lm wide, and having more numerous rays. This species is
known only from Norfolk Ridge, New Caledonia region from the
depth of 515-505 m.
Remarks. So far this is a monospecific genus. It resembles
Fig. 5. Neoaulaxinia clavata (Levi & Levi) MNHN DeL 3537 (paratype).
the fossil genus Aulaxinia Zittel, 1878a in morphology and canal
A-D, variability of small choanosomal streptasters/amphiasters with long development, differing from fossil representatives in desma char-
and slender rays (scale lO f1m). acteristics which are very loosely articulated in Neoaulaxinia.
Porifera· Demospongiae • 'Lithistids' • Phymatellidae 369

Fig. 6. Neosiphonia superstes Sollas. BMNH 1891.5.4.11. holotype. (after Sollas, 1888, pI. 31: 7). A-B, lateral views. C, upper surface view (scale 10 mm).

Distribution

New Caledonia.

NEOSIPHONIA SOLLAS, 1888

Synonymy

Neosiphonia Sollas, 1888: 299; Levi & Levi 1988: 247.

?[Jereopsis] Schmidt, 1879: 20; Lendenfeld 1903: 133; Wilson,
1925: 457.

Type species

Neosiphonia superstes Sollas, 1888: 299 (by original

designation). Fig. 7. Neosiphonia superstes Sollas. MNHN DCL 3540 (scale lOmm)
(modified after Levi & Levi, 1988).

Definition
in fact it belongs to a separate genus. Thus, synonymy of [Jereopsis]
Phymatellidae with a rounded body supported on a stem; in Schmidt with Neosiphonia Sollas has to be verified. Sollas
the center a bundle of vertical canals. compared Neosiphonia with fossil genera Jerea and Siphonia but
used a new generic name because we do not know microscleres in
Diagnosis fossil forms. Contrary to the chosen name the genus Neosiphonia
resembles more the fossil genus Jerea, which lacks an atrial cavity
Massive sponges with a rounded body supported on a stem of and displays an identical bundle of vertical canals in the middle,
various length; ectosomal smooth dichotriaenes; tetraclones than Siphonia, which is supplied with an atrial cavity. More impor-
smooth, ramified; microscleres amphiasters/streptasters. No axial tant, the name [Jereopsis] Schmidt, 1879 is a junior homonym of
cavity, canal system a bundle of vertical canals in the center of the Jereopsis Pomel, 1872. In effect the name Jereopsis had to be aban-
sponge. doned for this sponge. Wilson (1925) considered Neosiphonia
Sollas, 1888 as a younger synonym of Jereopsis Schmidt, 1879, but
Remarks for the reasons explained above his approach is rejected here; he
also described a new species J. fruticosa, attributed to the sponge
The generic name Neosiphonia was proposed by Sollas (1888) described by Schmidt as Jereopsis gen. nov., on desma morphology
who used it also for the lithistid sponge described as Jereopsis by which is very similar in both sponges but very different from that in
Schmidt (1879). The name [Jereopsis] Schmidt is considered Neosiphonia. Both forms also differ in habitus.
invalid because Schmidt presented neither diagnosis nor species
name. For the material of Schmidt (1879) described as Jereopsis, Description of type species
Sollas (1888) proposed the new name Neosiphonia schmidti.
A small fragment, labelled by Schmidt as 'Jereopsis n.g, Havana', Neosiphonia superstes Sollas, 1888 (Figs 6-10).
and preserved in the collection of the Musee Zoologique, Synonymy. Neosiphonia superstes Sollas, 1888: 299, pi. 31:
Strasbourg, displays however, tetraclone desmas completely differ- 7-12; Levi & Levi, 1983a: 119, fig. 10; Levi & Levi, 1988: 47,
ent from those known in Neosiphonia, and one cannot exclude that pI. 3, fig. 3. Jereopsis superstes; Lendenfeld, 1903: 133.
370 Porifera· Demospongiae • 'Lithistids' • Phymatellidae

Fig. 8. Neosiphonia superstes Sollas. MNHN DeL 3540. A. Outer surface with inhalant pores, ectosomal dichotriaenes and thick crust of ectosomal
robust amphiasters (scale 200 /-Lm). B, Outer surface without pores and with ectosomal dichotriaenes and crust of massive amphiasters (scale 200/-Lm).

Fig. 9. Neosiphonia superstes Sollas. A-D, ectosomal dichotriaenes in various views, MNHN DeL 3540 (Scale 200/-Lm). E-H, choanosomal tetraclone
desmas MNHN DeL 3540 (scale 200 /-Lm). I-L, choanosomal tetraclone desmas of the holotype, BMNH 1891.5.4.11 (scale 200 /-Lm).

Material examined. Holotype: BMNH 1891.5.4.11 - trichotriaenes, but this seems to be a pathological development)
deciduous, dry specimen from Fiji. Other material. MNHN DCL with short conical rhabdome, about 100-150mm long, and
3540 - specimen with ectosomal spicules and microscleres, cladome about 0.5 mm in diameter. Microscleres are short spi-
Norfolk Ridge. MNHN DCL 2808 - New Caledonia. rasters with long rays, about 45 /Lm long. This species was reported
Description. Approximately spherical sponge supplied with by Sollas (1888) from Fiji (depth 576m) and by Levi & Levi
a short massive stem. The holotype is about 46 mm high, and (1983a, 1988) from the Norfolk Ridge (depth of 515-505 m) and
39 mm in diameter. Upper surface concave with numerous openings New Caledonia (depth 430--500m).
from the large bundle of vertical exhalant canals; near the sponge Remarks. The holotype is a deciduous specimen, with only
surface they run rather parallel to it and measure about 1.5 mm in traces of dermal membrane. Consequently the redescription of
diameter. Outer surface, including the stem displays numerous ectosomal spicules and microscleres is based on both the holotype
rounded openings of incurrent canals which are loosely spaced and (Sollas, 1888) and examination of Levi & Levi's (1983a, 1988)
smaller, about 1 mm in diameter, than the excurrent ones. They run very well preserved specimens from Norfolk Ridge and New
radially toward the sponge center. Choanosomal desmas are large, Caledonia. The desmas in the holotype are larger and more
strongly branched tetraclones reaching 600--700/Lm in size. branched than those described by Levi & Levi (1988), but we
Ectosomal megascleres are smooth dichotriaenes (sometime consider this to be merely intraspecific variability.
Porifera' Demospongiae • 'Lithistids' • Phymatellidae 371

Fig. 10. Neosiphonia superstes Sollas. MNHN DeL 3540. A-F, variability of ectosomal robust amphiasters (scale 5 j.Lm). G-L, variability of
choanosomal streptaster/amphiasters (scale 10 j.Lm).

Distribution

New Caledonia and Fiji Islands.

REIDISPONGIA LEVI & LEVI, 1988

Synonymy

Reidispongia Levi & Levi, 1988: 245.

Type species

Reidispongia coerulea Levi & Levi, 1988: 245 (by monotypy).

Definition

Lamellar or cup-shaped Phymatellidae.

Diagnosis

Lamellar or cup-shaped, tetraclone desmas large, smooth and

ramified several times, with poor or no zygosis; ectosomal smooth
Fig. 11. Reidispongia coerulea Levi & Levi (scale bar 20mm) (modified
dichotriaenes; microscleres are two types of amphiasters (Levi & after Levi & Levi, 1988). A, holotype MNHN DeL 3538. B, paratype
Levi, 1988: 245). MNHN DeL 3539.
372 Porifera' Demospongiae • 'Lithistids' • Phymatellidae

Fig. 12. Reidispongia coerulea Levi & Levi (paratype), MNHN DCL 3539. A, outer surface with numerous inhalant pores and thin oxeas (scale 500 /Lm).
B, details of the same surface showing thick crust of ectosomal streptasters/amphiasters. C, surface of the choanosomal skeleton showing numerous canal
openings (scale I mm). D, details of choanosomal skeleton composed of branched tetrac10ne desmas (scale 200/-Lm).

Fig. 13. Reidispongia coerulea Levi & Levi (paratype), MNHN DCL 3539 (scale 100 /-Lm). A, C-D, ectosomal dichotriaenes. B, ectosomal triaene.

Remarks 25X15mm. Wall is very thin, 2-3mm thick. Outer surface nearly
smooth, slightly hispid when enlarged and with fine pores; both sur-
This is a monotypic genus resembling the Mesozoic genus faces similar, with canal openings measuring 180--300 fJ.m in diam-
Kalpinella Hinde, 1883. eter and separated by a distance of 200--500 /Lm. Ectosomal smooth
dichotriaenes occur on both surfaces, 150--300 fJ.m in diameter
Description of type species (protoclades 40118 fJ.m, deuteroclades 100--150 fJ.m), and have rhab-
dome 102-190 fJ.m long. Other megascleres are represented by
Reidispongia coerulea Levi & Levi, 1988 (Figs 11-14). superficial oxeas which measure at least 500--1000 /Lm long,
Synonymy. Reidispongia coerulea Levi & Levi, 1988: 245, 7-8 fJ.m thick. Below the ectosomallayer there are openings of per-
pI. 2, fig. 2. pendicular canals separated from each other which traverse only
Materiill examined. Holotype: MNHN DCL 3538. Paratypes: part of the wall. Endosomal skeleton consists of tetraclone desmas
MNHN DCL 3539 - Norfolk Ridge, depth 515-505 m. which are smooth and branched several times; there is very poor or
Description. Cup shaped sponge with folded wall, about no terminal articulation resulting in a fragile sponge. Two types of
60--80 mm high, 90--100 wide at the top, attached by a simple base microscleres are present: one is an ectosomal amphiasterlstreptaster
Porifera' Demospongiae • 'Lithistids' • Phymatellidae 373

Fig. 14. Reidispongia coerulea Levi & Levi (paratype), MNHN DeL 3539. A-D, variability of ectosomal robust streptasters/amphiasters (scale 5 j,Lm).
E-H, variability of choanosomal streptasters/amphiasters with long slender rays (scale 5 j,Lm).

with short and robust rays covered by numerous low spines and Remarks. Malformed dichotriaenes, as well triaene-like
rounded tips, 10-13/-Lm long, 10-12/-Lm wide; the other type is a spicules were commonly observed in the ectosome of this species.
choanosomal amphiasterlstreptaster but it has long and slender
rays with pointed ends and is covered with hook-like spines, 14- Distribution
20/-Lm long, 17-20/-Lm wide. This species is known only from
Norfolk Ridge, from the depth of 515-505 m. New Caledonia.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AeademielPlenum Publishers, New York, 2002

Family Isoraphiniidae Schrammen, 1924

Andrzej Pisera l & Claude U;vi2

1 Instytut Paleobiologii, Polska Akademia Nauk, ui. Twarda 51155, 00-818 Warszawa, Poland. (apis@twarda.pan.pl)
2Museum National d'Histoire Naturelle, Laboratoire de Biologie des Invertebres Marins et Malacologie, 57, Rue Cuvier, 75231 Paris
Cedex 05, France. (levi@rnnhn.fr)

Isoraphiniidae Schrammen (Demospongiae, 'lithistids') contains only one extant genus Costifer Wilson, known only from two species
from deep water of the Pacific. There are, however, several fossil Mesozoic genera which belong to this family. The family is character-
ized by heloclone desmas and ectosomal triaenes. Microscleres in Costifer are of three types: acanthoxeas, spiraster-like acanthorhabds,
and streptasters.
Keywords: Porifera; Demospongiae; 'lithistids'; Isoraphiniidae; Costifer.

DEFINITION, DIAGNOSIS, SCOPE

Synonymy

Helomorinidae Schrammen 1910: 128; Helomorinides Moret

1926: 128; Isoraphiniidae Schrammen, 1924a: 38; de Laubenfels
1955b: E51; Lagneau-Herenger 1962: 157. Isoraphiniadae
Schrammen 1937: 75.

Definition

'Lithistid' Demospongiae with choanosomal desmas as

heloclones and triaene ectosomal spicules.

Diagnosis

Choanosomal desmas are heloclones, ectosomal spicules are

Fig, 1. Costifer vasiformis Wilson; USNM 21343, morphology of the
triaenes (pi agio- and/or dichotriaenes). Microscleres are strep- sponge (from Wilson, 1925, pI. 43: 6) (scale !Oem).
tasters and acanthoxeas, and spiraster-like acanthorhabds.

(based on the Cretaceous genus Isoraphinia Zittel), and proposed

Scope Helomorina as suborder of the order 'Lithistida'.

One Recent genus, Costifer Wilson, 1925, with 2 species;

there are also several fossil genera (Mesozoic and Tertiary). COSTlFER WILSON, 1925

Synonymy
Remarks
Costifer Wilson, 1925: 461.
The differentiating characters of Isoraphiniidae Schrammen,
1924a are heloclones, monocrepidial desmas with sinuous shaft Type species
displaying numerous zygoses in the form of lateral notches along
their length; the axial canal extends along the whole desma length Costifer vasiformis Wilson, 1925: 461, pi. 43: 6, pi. 50: 8, 13,
(Levi, 1991). This last feature makes the heloclones different from 14, pi. 51: 1 (by monotopy).
megaclones, with which they are often compared, and which
are also monocrepidial, but an axial canal if present is very Definition
short. Originally Schrammen (1910: 128) proposed the family
Helomorinidae for fossil lithistids with heloclones and triaenes. Isoraphiniidae with 3 types of microscleres: acanthoxeas,
Later (Schrammen 1924a: 38) he corrected it to Isoraphiniidae streptasters and spiraster-like acanthorhabds.

374
Porifera· Demospongiae • 'Lithistids' • Isoraphiniidae 375

Fig. 2. Costifer vasiformis Wilson. USNM 21343; A, view of the inner surface with oscula (scale I mm); B, crust of microscleres on the inner surface
(scale 50 !-Lm); C, crust of ectosomal plagiotriaenes on the outer surface (scale 500 !-Lm); D, microxeas in the choanosome (scale 100 !-Lm).

Fig.3. Costifer vasiformis Wilson, USNM 21343; A-H, ectosomal triaenes (scale 500 !-Lm); 1-0, choanosomal heloclones (scale 500 !-Lm).

Diagnosis spicules are heloclones with lateral zygomes; other megascleres

are large radially protruding oxeas and smaller endosomal
Vase shaped or lamellar; ectosomal spicules are plagiotriaenes, ones; microscleres are microxeas, streptasters and spiraster-like
which may be dichotriaene-like, rarely anatriaenes; endosomal acanthorhabds.
376 Porifera· Demospongiae • 'Lithistids' • Isoraphiniidae

Fig. 4. Costifer vasiformis Wilson, USNM 21343; A-C, microxeas (scale 20 !Lm); D, microxeas tips (scale 5 !Lm); E-H, streptasters (scale 5 !Lm);
I-N, spiraster-like acanthorhabds (scale 5 !Lm).

Remarks a result of dichotomous branching of some rays; also anatriaene-like

spicules have been observed. The plagiotriaenes are strongly pro-
This genus is represented by only two Recent species. There truding from the membrane forming an external crust on the outer
are, however, numerous fossil (Mesozoic) genera attributed to the surface, which was not observed on the inner surface. Choanosomal
fossil group Helomorina, which have identical desmas and triaenes desmas are typical heloclones with numerous lateral zygoses, with
among ectosomal spicules (Schrammen, 1924a; Levi, 1991). See rounded or tapering ends; axial canal extends through entire desma.
Levi (1991) for detailed discussion of relationship between fossil In some rare cases, there may exist small branches. The articulation
and Recent forms. of the desmas may occur at any point of the desma length; desmas
are 1.88-2.37 mm long, 220-280 fLm thick. Long oxeas are radially
Description of type species arranged in the choanosome, 2.0-2.5 mm long, 70 fLm thick.
Microscleres are of three types: evenly tapering acanthoxeas with
Costifer vasiformis Wilson (Figs 1-4). pointed tips, very abundant in the choanosome, 90-165 fLm long,
Synonymy. Costifer vasiformis Wilson, 1925: 461, pI. 43: 6, 4--5 fLm thick; spiraster-like acanthorhabds, small, very irregular and
pI. 50: 8, 13, 14, pI. 51: 1. of very variable shape, often with several large protuberances on the
Material examined. Holotype: USNM 21343 - Philippines, surface, very abundant in the ectosome and measure 18-25 fLm long,
Sulu Archipelago, Tawi Tawi Islands, 145m depth. MNHN DCL 5.4--7.7 fLm thick; streptaster microscleres very variable in shape,
2633 (two slides from the holotype). with long and pointed rays and a very short central axis or not visible
Description. Large (about 27 cm high and 28 cm wide) vase- at all, 15-19 fLm long, 12-17 fLm wide. This species was reported
shaped sponge, attached by a flattened base. The central cavity from the Philippines from a depth of 145 meters.
extends through the entire sponge. Walls are 20-25 mm thick at the Remarks. The second species of this genus described by
upper margin. Ectosomal spicules are projecting plagiotriaenes. Levi (1993) differs mostly in having only dichotriaenes as ectoso-
Closely spaced canal openings on the surface, partly covered by pla- mal spicules, and in the absence of articulation notches in the
giotriaenes, measuring 0.20-0.25 mm in diameter. The inner surface desmas; microscleres are very similar.
of the sponge displays closely spaced oscules 0.5-1 mm in diame-
ter. Plagiotriaenes vary in size, with a rhabdome about 850- Distribution
1310 fLm long, 45 fLm thick, and cladome 525-830 fLm in diameter;
sometimes there may be dichotriaene-like triaenes produced as Philippines, New Caledonia.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Macandrewiidae Schrammen, 1924

Andrzej Pisera1 & Claude Levi2

1 Instytut Paleobiologii, Polska Akademia Nauk, ul. Twarda 51155,00-818 Warszawa, Poland. (apis@twarda.pan.pl)
2Museum National d'Histoire Naturelle, Laboratoire de Biologie des Invertebres Marins et Malacologie, 57, Rue Cuvier, 75231 Paris
Cedex 05, France. (levi@mnhn.fr)

Macandrewiidae Schrammen (Demospongiae, 'litbistids') contains only one valid genus Macandrewia Gray, 1859, with about 6 species
known from the Atlantic and Pacific. They are polymorphic, cup shaped, branching or massive irregular, and occur in relatively deep
water as do most lithistids. The differentiating characters of the family are ectosomal dentate phyllo- to discotriaenes and smooth desmas
of complex morphology resembling tetraclones which have triaenose (rarely monaxial) crepis; microscleres are smooth microxeas.
Keywords: Porifera; Demospongiae; 'lithistids'; Macandrewiidae; Macandrewia.

DEFINITION, DIAGNOSIS, SCOPE long and otbers very short). This was already noted by Schmidt
(1870) and Sollas (1888) in M. clavatella (Schmidt) and by
Synonymy Topsent (l904b) in M. azarica, as well as by Levi & Levi (1989)
but its importance is disregarded. There are other basic differences
Macandrewiidae Schrammen, 1924a: 39. in desmas of Macandrewia and corallistids: they are smooth with
terminal zygosis in Macandrewia, while strong tubercles and non-
Definition terminal zygosis are found in Carallistes and allied genera. Both
families differ also in their ectosomal spicules: dichotriaenes in
'Lithistid' Demospongiae with ectosomal phyllotriaenesl corallistids and phyllotriaenes in Macandrewia; corallistids have
discotriaenes and complex desmas with triaenose (rarely monaxial) also completely different microscleres, usually streptasters (spi-
crepis; microscleres as microxeas. rasters) which do not occur in Macandrewia. These differences
provide empirical support to allocate Macandrewia to a different
Diagnosis family, as proposed by Schrammen (l924a).

Polymorphic; ectosomal spicules as strongly incised and den-

tate phyllotriaenes/discotriaenes; desmas with triaenose (rarely MACANDREWIA GRAY, 1859
monaxial) crepis, zygosis terminal; microscleres as microxeas
occurring in tbe ectosome. Synonymy

Scope Mac Andrewia Gray, 1859: 438; [Mac Andrewsia]

Bowerbank, 1868: 758, 823 (lapsus); Mac Andrewia Zittel, 1878a:
Monogeneric. 34, 57; Macandrewia Sollas, 1888: 340; Topsent, 1892a: 51;
Lendenfeld, 1903: 138 (partim); Lendenfeld, 1907: 357; Gruber,
Remarks 1993: 34; Dactylocalyx (partim) Bowerbank, 1869b: 76.

Schrammen (l924a) defined this family as 'Dicranocladina Type species

[=Corallistidae] with monaxial phyllotriaene-like dermalia'. This
was a mistake, as phyllotriaenes of Macandrewia azorica are Macandrewia azorica Gray, 1859: 438, pI. 15 (by monotypy).
clearly triaenose. So far Macandrewia has been considered a coral-
listid, based on apparently monaxial desmas; this was probably the Definition
reason for Kelly (2000) in her review of Neopeltidae to include
Macandrewia in Neopeltidae (considered as Dicranocladina). The 'Litbistids' with ectosomal dentate phyllotriaenes/discotri-
same author considered Macandrewia as belonging to one group aenes, complex choanosomal desmas witb triaenose crepis or
with Homophymia and Callipelta, which have monaxial ectosomal monocrepidial in nature, and microxeas as the only microscleres.
pseudophyllotriaenes. We reject this opinion, which Kelly (2000)
also based on supposedly non-triaenose crepis of ectosomal phyl- Diagnosis
lotriaenes in Macandrewia, while in fact crepis in ectosomal phyl-
lotriaenes of Macandrewia is very similar to that in phyllotriaenes Polymorphic; ectosomal spicules are dentate phyllotriaenes
occurring in Theonellidae, generally accepted as of triaenose type. and/or discotriaenes; choanosomal spicules are smooth desmas
Closer examination of the type material of Macandrewia azorica with chiefly terminal zygosis which have mostly a triaenose (rarely
Gray, as well as some other species, revealed that tbe desmas have monaxial) crepis; the only microscleres are ectosomal microxeas
in fact triaenose crepis in most cases (however, with one canal very forming a dense tangential ectosomal crust.

377
378 Porifera' Demospongiae· 'Lithistids" Macandrewiidae

Remarks

Lendenfeld (1903) synonymized Callipelta Sollas with

Macandrewia Gray, whereas the former is allocated here to the
Neopeltidae Sollas because of the monaxial nature of its ectosomal
spicules (pseudophyllotriaenes) and desmas. Currently eight
species are included in this genus, and probably all belong here.

Description of type species

Macandrewia azorica Gray (Figs 1-3).

Synonymy. Macandrewia azorica Gray, 1859: 483, pI. 15;
Sollas, 1888: 340; Carter, 1873c: 438,441; Topsent, 1904b: 60,
pI. 3: 2, pI. 8: 1; Gruber, 1993: 35; Dactylocalyx macandrewii;
Bowerbank, 1869b: 86, pI. 4: 5, pI. 5: 1-5. Topsent, 1892a: 51.
Materilzl examined. Holotype: BMNH 1851.7.28.16 - Fig. 1. Macandrewia azorica Gray, BMNH 1851.7.28.16, holotype
Azores, St. Michael's (dry specimen), 2 slides BMNH (scale 2 em).
1877.5.21.1464 (from type).

Fig. 2. Macandrewia azorica Gray, BMNH 1851.7.28.16, holotype. A, outer surface of the sponge with pores and dense crust of ectosomal microxeas
(scale SOO fLm); B, details of outer surface showing dense crust of microxeas (scale 100 fLm); C, outer (lower) surface of the choanosomal skeleton (scale
2 mm); D, inner (upper) surface of the choanosomal skeleton (scale 2 mm); E, details of choanosomal desmas (scale 200 fLm); F, articulation of the
choanosomal desmas (scale 100 fLm).
Porifera· Demospongiae • 'Lithistids' • Macandrewiidae 379

Fig. 3. Macandrewia azorica Gray, BMNH 1851.7.28.16, holotype A-D, ectosomal phyllotriaenes (scale 100 fLm); B-1, ectosomal microxeas (scale 20 fLm) ;
J, choanosomal oxea (scale 100 fLm).

Description. Cyathiform to flabellate, about 12 cm high, them very long); there is little, if any, relationship between crepis
l2cm wide, with short stem about 3cm long; walls 6-9mm thick, orientation and desma shape. The des rna clones are strongly
with undulating rounded margins. Outer surface smooth with branched at the end into long narrow spine-like processes which
small, irregularly but evenly distributed pores measuring grasp each other to form a very complex but rather loose terminal
37-58/Lm, leading to larger openings, about 200-300/Lm diame- articulation. The resulting skeleton is rather fragile and loosely
ter, in the choanosomal skeleton that open into choanosomal spaced; in longitudinal section the centres of articulation are
canals. Ectosome packed densely but irregularly with tangential aligned along vertical radial lines, producing a fan-shaped struc-
smooth microxeas. Inner surface smooth except for oscules, ture. At the growing edge desmas are strongly modified and elon-
0.3-2 mm in diameter, with slightly raised margins distributed gated into finger-like processes. Other choanosomal megascleres
evenly over the entire surface. The upper surface of the choanoso- are small, 585-690/Lm long and 10-13 /Lm thick, fusiform oxeas.
mal skeleton, apart from oscular openings, displays numerous Microscleres which are very common on the sponge surface are
choanosomal canal openings and subectosomal canals on the sur- smooth fusiform oxeas with blunt tips, measuring 38-63 /Lm long,
face which are both about 300-500 /Lm wide. Ectosomal megascle- 2-4/Lm thick. This species has been recorded from depths of
res are phyllotriaenes/discotriaenes with strongly incised clades; 454-l550m from the Azores, northern coast of Scotland and
short conical rhabd is l70-240/Lm long, c1adome 300-460/Lm in southern coast of Portugal.
diameter. Choanosomal skeleton consists of smooth complex des-
mas of variable morphology, either resembling tetraclones with Distribution
four rays emanating from the center, or rhizoclones with four or
more rays emanating from a short or long central beam; most of Caribbean, Azores, Portugal, Scotland, New Caledonia,
these desmas have clearly triaenose crepis with four rays (one of Philippines.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John NA. Hooper and Rob WM. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Phymaraphiniidae Schrammen, 1910

Andrzej Pisera1 & Claude Levi2

1 Instytut Paleobiologii, PolskaAkademia Nauk, ul. Twarda 51/55,00-818 Warszawa, Poland. (apis@twarda.pan.pl)
2Museum National d'Histoire Naturelle, Laboratoire de Biologie des Invertebres Marins et Malacologie, 57, Rue Cuvier, 75231 Paris
Cedex OS, France. (levi@mnhn.fr)

Phymaraphiniidae Schrammen (Demospongiae, 'lithistids') contains only two extant genera: Kaliapsis Bowerbank and Rimella Schmidt.
There are, however, several fossil (Mesozoic and Tertiary) genera (see chapter on fossillithistids). The genus Kaliapsis is known from
Bowerbank's original description in 1869 (1869b), from the 'South Seas' (no other data known), Madagascar, and recently has been found
in the Caribbean, while Rimella occurs in the Caribbean and Azores, NW Atlantic. Representatives of the family occur both in shallow
(coral reefs) and deep waters. The family is differentiated from others by the presence of trider-type desmas, discotriaenes to phyllotri-
aenes as ectosomal spicules, and microscleres are acanthoxeas, acanthorhabds to acanthostyles and streptasters with long rays. Genera are
differentiated by their habit and type of ectosomal spicules.
Keywords: Porifera; Demospongiae; 'lithistids'; Phymaraphiniidae; Kaliapsis; Rimella.

DEFINITION, DIAGNOSIS, SCOPE Scope

Synonymy There are two extant genera: Kaliapsis Bowerbank, 1869b,

Rimella Schmidt, 1879.
Phymaraphininae Schrarnmen, 1910: 104; Phymaraphiniidae
Schrammen, 1924a: 38; Lagneau-Herenger, 1962: 147. Phymara-
phinidae Levi, 1991: 80. Kaliapsidae de Laubenfels, 1936a: 175 Remarks
(partim).
This family was established by Schrammen (1910, 1924a)
Definition mostly for fossil genera, based on a very characteristic desma mor-
phology and phyllotriaene dermalia, and founded on the Upper
'Lithistid' Demospongiae with trider-type desmas and disco- Cretaceous genus Phymaraphinia Schrarnmen, 1910. Schrarnmen
triaenes to phyllotriaenes as ectosomal spicules; microscleres (1910) himself pointed out a close relationship between the Recent
are acanthorhabds to acanthostyles, and/or acanthoxeas, and genus Kaliapsis Bowerbank and his family. Our studies support
streptasters with long rays. his opinion. De Laubenfels (1936a) established a new family
Kaliapsidae, without diagnosis, embracing a wide array of various
Diagnosis 'lithistid' genera (apart from Kaliapsis also corallistids, pleromids
and theonellids), now considered clearly polyphyletic with genera
Polymorphic; endosomal spicules articulated very regular belonging to different families; in a later pUblication (de Laubenfels,
modified tetraclones (triders) which have one ray reduced to 1955b) he defined this family as 'typical rhizomorine sponges',
a knob or spine and without zygome, and strongly tuberculate; and added to it several fossil rhizomorine genera. As a result the
ectosomal spicules are phyllotriaenes and/or discotriaenes; micro- family Kaliapsidae de Laubenfels should be abandoned and the
scleres are acanthorhabds to acanthostyles and/or acanthoxeas, and inclusion of Kaliapsis as type genus relegates it into synonymy
streptasters with long spinose rays. with Phymaraphiniidae.

KEY TO EXTANT GENERA

(1) Encrusting in habit, ectosomal discotriaenes ......................................................................................................................... Kaliapsis

Clavate in habit, ectosomal phyllotriaenes ............................................................................................................................... Rimella

KAUAPSIS BOWERBANK, 1869 Definition

Synonymy Encrusting Phymaraphiniidae with tuberculated discotriaenes

(usually incised) as ectosomal spicules; microscleres are spinose
Kaliapsis Bowerbank, 1869b: 76, 337; Zittel, 1878a: 101, rhabds, and streptasters/amphiasters (not known from the type
152; Sollas, 1888: 333; Lendenfeld, 1903: 133; de Laubenfels, species).
1955b: E45. Gruber, 1993: 43.
Diagnosis
Type species
Thinly encrusting in habit; choanosomal spicules are strongly
Kaliapsis cidaris Bowerbank 1869b: 338 (by monotypy). tuberculated triders; ectosomal spicules are tuberculated or spinose

380
Porifera· Demospongiae· 'Lithistids'· Phymaraphiniidae 381

discotriaenes with incised margins; microscleres are spinose 338, pI. 25: 2-5; Carter, 1873: 441; Zittel, 1878a: 152, pI. 12: 12,
rhabds and streptasters/amphiasters (the last not reported from the 12a; Sollas, 1888: 334; Lendenfeld, 1903: 134.
type species). Material examined. Holotype: BMNH 1877.5.21 (slide) -
encrusting the base of Oculina rosea from the 'South Seas', no
Remarks other data given.
Description. Thinly encrusting plates without visible pores
In the holotype of K. cidaris (only slides are extant) no or oscules; holotype only 6 mm in diameter and about 0.6 mm
rnicroscleres other than rhabds were observed, while in the undoubted thick. Ectosomal spicules are strongly tuberculated or rnicrospined
Kaliapsis incrustans from Madagascar (see Vacelet & Vasseur, 1971; discotriaenes with a very short rhabd, usually strongly incised mar-
Vacelet et al., 1976) there are also amphiasters in addition to rhabd gins, and a tuberculated upper surface, measuring about 260-290
rnicroscleres. In our opinion this component was most probably over- fLm in diameter. Desmas are regular triders (modified tetraclones)
looked in Bowerbank's (1869b) very small sample. The genus with one clone forming brachyome in the form of polar button, or
Kaliapsis was established by Bowerbank (1869b), who in his earlier spine-bearing button; other clones have regular sculpturing of large
illustrations (Bowerbank, 1858, 1862a, 1864) refers to but does not tubercles (which bear in tum smaller low tubercles as sculpture)
formally name these spicules. Thus both the genus and the species placed in a ring-like fashion. Triders measure about 180 fLm in size.
Kaliapsis cidaris was formally established in 1869 (and not in 1858 Microscleres are fusiform microstrongyles (most probably
as often supposed by authors). Three species (K. cidaris Bowerbank, rnicrospined) and are about 28-48 fLm long and about 4-5 fLm thick.
1869b, K. aspera (Carter, 1880b) and K. permollis Topsent, 1890) Remarks. This is a poorly known species with Bowerbank's
were included into this genus. Examination of the original material (1869b) report the most recent, and furthermore only spicule
of K. permollis Topsent, 1890, revealed that in fact it is a preparations are extant - the original specimen was so small that it
Siliquariaspongia Hoshino, 1981b. The original placement of was used entirely for preparations.
K. aspera in Discodermia by Carter (1880b) seems more appropriate.
Its desmas are typical tetraclones, not resembling triders, and the sub- Distribution
sequent referral of this species to Kaliapsis was based on alleged sim-
ilarities of its spinose discotriaenes with those of K. cidaris. The type species was reported from an unknown locality in
Lendenfeld (1903) also considered this species to belong to the 'South Seas', with records also apparently from Madagascar
Theonellidae but this view is not supported here. and the Caribbean.

Description of type species

RIMELLA SCHMIDT, 1879
Kaliapsis cidaris Bowerbank, 1869b (Fig. 1).
Synonymy. Various unassigned spicules in Bowerbank, Synonymy
1858: 279, pI. 24: 32-38; 1862a: 759, pI. 28: 12; 1864: pI. 4:
102-103, pI. 5: 104-108. Kaliapsis cidaris Bowerbank, 1869b: Rimella Schmidt, 1879: 21.

Fig. 1. Kaliapsis cidaris Bowerbank, holotype BMNH 1877.5.21. A-B, triders (desmas) (scale 1 mm). C-E, ectosoma1 discotriaenes (scale 1 mm).
F, microsc1eres (scale 500 fLm).
382 Porifera· Demospongiae • 'Lithistids' • Phymaraphiniidae

Type species Description of type species

Rimella clava Schmidt, 1879: 21 (by monotypy). Rimella clava Schmidt (Figs 2-3).
Synonymy. Rimella clava Schmidt, 1879: 21, pI. 1: 2, pI. 2:
Definition 5,7,11; Sollas, 1888: 335. Not Racodiscula clava Topsent, 1892a:
49, pI. 1: 5-{i, pI. 8: 3; Topsent, 1904b: 58.
Clavate Phymaraphiniidae with as phyllotriaenes ectosomal
spicules.

Diagnosis

Clavate in habitus; ectosomal spicules are smooth phyllotri-

aenes; desmas are strongly tuberculated triders; microscleres
are acanthorhabds to acanthostyles and/or acanthoxeas, and
streptasters with long slender rays.

Remarks

This genus was described by Schmidt (1879) based on decid-

uous specimens from the Caribbean; Topsent described specimens
of Rimella (synonymized by him with Racodiscula) from the
Azores, which were very similar in habitus and attributed to R.
clava Schmidt, having all spicules present including microscleres.
However, we found it to be a different, closely related undescribed
new species. Rimella Schmidt was considered to be a junior syn-
onym of Racodiscula by Topsent (1892a, 1904b) and Lendenfeld
(1903). Although both genera share very similar ectosomal phyllo-
triaenes as well as microscleres they have very different desmas:
typical tetraclones in Racodiscula, and riders in Rimella. For this
reason we treat them as belonging to different families Fig. 2. Rimella clava Schmidt, syntype MeZ 6436 (orig. no. 275)
(Racodiscula in Theonellidae). (Schmidt, 1879, pI. 1: 2).

Fig. 3. Rimella clava Schmidt, MNHN DeL 1020 (from Schmidt's type MZUS). A, surface of choanosomal skeleton consisting of triders (scale 200 /-Lm).
B, trider from the choanosomal skeleton (scale 100 /-Lm). e, details of trider sculpture (scale 50 /-Lm). D, details of trider articulation (scale 50/-Lm).
Porifera' Demospongiae • 'Lithistids' • Phymaraphiniidae 383

Material examined. Syntypes: MCZ 6436 (orig. 275) - size, and with articulation general; tubercles are very high and
four different deciduous specimens, Gulf of Mexico; MZUS mushroom-shaped such that desmas resemble some dicran-
P0146 - Havana (fragment and slides DCL 1020 and MNHN DT oclones. These tubercles bear a sculpture of low tubercles. In
2202). Other material. MNHN DT 782 - Azores (Not Rimella an undescribed species from the Azores [Racodiscula (=Rimella)
clava, but an undescribed species). "clava" Schmidt, in Topsent, 1892a, 1904b] other megascleres
Description. Small cylindrical to clavate sponges, up to are strongyles; microscleres are acanthose fusiform microxeas,
3 cm long, lcm thick, sometimes displaying short lateral protuber- sometimes centrotylote, and rhabds, and streptasters with slender
ances or branches. Choanosomal skeleton bears numerous more- rays.
or-less longitudinal, but irregularly running subdermal canals Remarks. All original Schmidt specimens from the
visible through the ectosome. The type species is represented by Caribbean are deciduous, and as such a more precise characteriza-
deciduous specimens only but in closely related species from tion of the type species must await discovery of a new live material.
Azores [Racodiscula (=Rimella) "clava" Schmidt, in Topsent,
1892a, 1904b] ectosomal spicules are phyllotriaenes with wide Distribution
clads. Desmas of the type material are strongly tuberculated triders
(tetraclones with one ray reduced), measuring 230-320 f.Lm in Caribbean, Azores, Madagascar.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademicIPlenum Publishers, New York, 2002

'Lithistids'Incertae Sedis
Andrzej Pisera1 & Claude Levi2

1 Instytut Paleobiologii, Polska Akademia Nauk, ui. Twarda 51/55, 00-818 Warszawa, Poland. (apis@twarda.pan.pl)
2Museum National d'Histoire Naturelle, Laboratoire de Biologie des Invertebres Marins et Malacologie, 57, Rue Cuvier, 75231 Paris
Cedex 05, France. (levi@mnhnJr)

Five poorly known genera of lithistid sponges (Demospongiae, 'lithistids'), based on deciduous material and thus devoid of loose
spicules, remain of uncertain status, but most probably they are synonyms of other well-known genera. These include lithistids with tetr-
aclone desmas (Collectella Schmidt, 1880b; Collinella Schmidt, 1879), or with rhizoclone desmas (Arabescula Carter, 1873c; Plakidium
Lendenfeld, 1907; and Poritella Schmidt, 1879). The status of these genera might be decided when well-preserved material with all
spicules in place is found.
Keywords: Porifera; Demospongiae; 'lithistids'; incertae sedis; Collectella; Collinella; Arabescula; Plakidium; Poritella.
----------~-------------------------------------------

'LITIDSTIDS'WITHTETRACLONEDESMAS Synonymy. Collectella avita Schmidt, 1880b: 86, pI. 5: 1;

Sollas, 1888: 410; Theonella avita Lendenfeld, 1903: 128.
COLLECTELLA SCHMIDT, 1880 Material examined. Holotype: MZUS P0167 - Gulf of
Mexico, depth unknown, Schmidt original material. MNHN DT
Synonymy 2189 (slide); MNHN DCL 1161 - fragment and slide from MZUS
holotype.
Collectella Schmidt, 1880b: 86. Description. 'Knob-like, with somewhat flattened summit
on which vertical canals open' (after Sollas, 1888: 410). Desmas
Type species are massive and tuberculated on the whole length. Ectosomal
spicules (as illustrated by Schmidt, 1880b) are phyllotriaenes. No
Collectella avita Schmidt, 1880b: 86, pI. 5: 1 (by monotypy). microscleres are preserved.
Remarks. Ectosomal spicules illustrated by Schmidt
Definition (1880b) are clearly phyllotriaenes, not dichotriaenes as reported by
Sollas (1888). Schmidt (1880b) reported also the presence of
As for type species. 'Pachastrella-like spicules' among the whole sponge body: these
are probably either young desmas or foreign spicules. Lendenfeld
Diagnosis (1903) synonymized Collectella with Theonella. This may be true,
but phyllotriaenes also occur in Racodiscula, for example, and thus
As for type species. the final decision must be undertaken based on well-preserved
specimen with microscleres.
Remarks

Monotypic. [COLLINELLA] SCHMIDT, 1879

Description of type species Synonymy

Collectella avita Schmidt, 1880b (Fig. 1). [Collinella] Schmidt, 1879: 21; Sollas, 1888: 336 (preocc. for
Collinella Martens, 1871, Mollusca).

Type species

Collinella inscripta Schmidt, 1879: 21, pI. 1: 3, pI. 2: 12

(by monotypy).

Definition

As for type species.

Diagnosis

Fig. 1. Collectella avita Schmidt. Holotype MZUS P0167 (scale 200 f1m). As for type species.

384
Porifera· Demospongiae • 'Lithistids' • incerlae sedis 385

Remarks (fragments and slides from the original Schmidt material from
MZUS), MZUS P0174.
No formal diagnosis of the genus was ever given, as Schmidt Description. There are two specimens in the collection of
(1879) simply described a new species with this generic name, MCZ, which differ in morphology, both deciduous. The specimen
including it in tetracladine lithistids. Most probably this genus is a MCZ 6070 (illustrated by Schmidt, 1879, pI. 1: 3) is a clavate
junior synonym of Discodennia, Theonella or Racodiscula, but the sponge about 38 mm high, with expanded attachment. At the
absence of ectosomal spicules and/or microscleres precludes its rounded top there is an osculum leading to a deep atrial cavity.
formal synonymy. Lendenfeld (1907) and Van Soest & Stentofi Lateral surface bears several low elevated areas, with small open-
(1988) considered it a synonym of Discodennia. As [Collinella] ings at the center, and converging irregularly radial subdermal
Schmidt, 1879 is preoccupied by Collinella Martens, 1871 canals visible on the surface of the choanosomal skeleton. The
(Mollusca), a new name is theoretically required, but as we are other specimen is small, only 16mm high and 13mm wide, sub-
convinced that it is a junior synonym of another sponge genus, we spherical with short pedicel. At the summit there is a single oscu-
defer this action until new material becomes available to lum, about 1.6mm wide. The surface of the choanosomal skeleton
corroborate this opinion. displays numerous irregularly sinuous and branching subdermal
canals. No ectosomal spicules or microscleres were observed.
Description of type species Choanosomal desmas are small, massive tetraclones with rare
tubercles on clones, and strongly tuberculated articulating parts.
Collinella inscripta Schmidt (Figs 2-3). Remarks. Monotypic.
Synonymy. Collinella inscripta Schmidt, 1879: 21, pI. 1: 3,
pI. 2: 12; Sollas, 1888: 336; Discodermia inscripta Lendenfeld,
1903: 131. 'LITHISTIDS' WITH RHIZOCLONE DESMAS
Material examined. Cotypes: MCZ 6071, MCZ 6070 -
Cuba, depth 534m. MNHN DT 2266, MNHN DCL 1154 ARABESCUIA CARTER, 1873

Synonymy

Arabescula Carter, 1873c: 464; Zittel, 1878a: 103, 120.

Type species

Arabescula parasitica Carter, 1873c (by monotypy).

Definition

As for type species.

Diagnosis

As for type species.

Fig. 2. Collinella inscripta Schmidt. A, MCZ 6070 (scale IOmm). B-C, Remarks
MCZ 242 in lateral (B) and top view (C) (scale 5 mm).

This genus should probably be abandoned as unrecognizable,

as no determination can be made based on one layer of (probably
specialized) desmas, without any other features.

Description of type species

Arabescula parasitica Carter (Fig. 4).

Fig. 4. Arabescula parasitica Carter, 1873c. A, desmas, after Carter 1873c

Fig. 3. Collinella inscripta Schmidt. Choanosomal skelton of tetrac1ones, (no exact scale given by Carter, 1873c). B, desmas, after Zittel, 1878a
MZUS P0174 (scale 500 j.1m). (scale 500 j.1m).
386 Porifera' Demospongiae • 'Lithistids' • incertae sedis

Fig. 5. Plakidium acutum Lendenfeld, 1907. A, choanosomal skeleton as seen on the inner side (scale 500 fLm). B, details of the desmas from the surface of
the inner side (scale 100 fLm). C, view of the holotype (scale 1 em). D, choanosomal skeleton as seen on the outer side (scale 500 fLm). E, details of desmas
from the surface of the outer side (scale 100 fLm). P, desmas from the interior of the choanosomal skeleton (scale 100 fLm). All after Lendenfeld (1907).

Synonymy. Arabescula parasitica Carter, 1873c: 464, Siphonidium. As this character is not diagnostic for Siphonidiidae,
pI. 17: 7-9; Zittel, 1878a: 120, pI. 1: 11; Sollas, 1888: 409; and exotylostyles are absent, this allocation is questionable.
Corallistes parasitica Ridley, 1884b: 183; Coscinospongia
parasitica Lendenfeld, 1903: 138. Description of type species
Material examined. None. Recorded distribution: English
Channel and Indian Ocean (Seychelles), depth unknown. Plakidium acutum Lendenfeld (Fig. 5).
Description. Thinly encrusting sponge; neither ectosomal Synonymy. Plakidium acutum Lendenfeld 1907: 360, pI. 45:
spicules nor microscleres were reported. Desmas are rhizoclones. 8-13.
Remarks. As already pointed by Sollas (1888: 409) this is Material examined. None. Cape Verde Islands, Atlantic,
clearly a rhizomorine sponge, not Corallistes as suggested by depth of 1694 m.
Ridley (1884b) and Lendenfeld (1903). Description (based on Lenden/eld, 1907). Slightly folded
irregular plate, which is 77 X 40 mm long, and 4 mm thick, without
any visible attachment. The sponge is mostly deciduous. On the
PLAK/DIUM LENDENFELD, 1907 outer side there are numerous, densely spaced, 200/.Lm wide longi-
tudinal canals visible on the surface of the choanosomal skeleton.
Synonymy On the inner side occur rounded oscula (visible in partly preserved
dennal membrane), measuring 500 /.Lm in diameter and lying 1.5 mm
Plakidium Lendenfeld, 1907: 360. apart. No ectosomal spicules or microscleres have been observed.
Choanosomal desmas are thorny rhizoclones 200-350/.Lm in size.
Type species Remarks. The only known specimen of this species is mostly
deciduous, thus the final decision concerning the status of Plakidium
Plakidium acutum Lendenfeld, 1907: 360 (by monotypy). can only be made if well-preserved material is found. Probably, how-
ever, it will be synonymized with Leiodermatium. Lendenfeld
Definition (1907) described two types of desmas: those from the interior of the
choanosome, and those forming the outennost layer of the
As for type species. choanosomal skeleton, which are slightly modified rhizoclones.
Such differentiation of rhizoclones (and often tetraclones as well) is
Diagnosis (after Lendenfeld, 1907) a common feature in rhizomorine lithistids and cannot be given any
special taxonomic importance.
Plate-shaped Siphonidiidae without tylostyle rhabds.

Remarks PORITELLA SCHMIDT, 1879

Attribution of this genus to Siphonidiidae by Lendenfeld Synonymy

(1907) was based on the supposed presence of a superficial layer of
modified desmas, which were considered as characteristic of Poritella Schmidt, 1879: 27.
Porifera • Demospongiae • 'Lithistids' • incertae sedis 387

Fig. 6. Poritella decidua Schmidt, 1879, cotype MCZ 64116a (MCZ 237)
(scale 10mm). A, upper surface view. B, lower surface view.

Type species

Poritella decidua Schmidt, 1879: 27.

Definition
Fig. 7. Poritella decidua Schmidt, cotype MZUS P0167; choanosomal
As for type species. skeleton of rhizoclone desmas (scales: A, 400 f.Lm; B, 200 f.Lm).

Diagnosis

As for type species. DT2203 (slide), MNHN DCLI040 (slide) (both from original
MZUS material).
Remarks Description. Ear shaped, or irregular vase shaped sponge,
about 35 mm high and 32mm wide, with walls 10mm thick. Upper
This genus was synonimized with Leiodermatium Schmidt by side of the choanosomal skeleton with numerous densely distrib-
Lendenfeld (1903), but because all known specimens are clearly uted oscula, but without any order, which are 500 J.!.m in diameter.
deciduous, this allocation is questionable. Lower side of the choanosomal skeleton also with numerous
densely distributed pores, also distributed without order, which are
Description of type species about 200--250 J.!.m in diameter. Choanosomal skeleton very dense,
consisting of rhizoclone desmas with strong and numerous tuber-
Poritella decidua Schmidt, 1879 (Figs 6--7). cles. Articulation of desmas general. No ectosomal spicules or
Synonymy. Poritella decidua Schmidt, 1879: 27; Sollas, microscleres were observed.
1888: 351; Leiodermatium deciduum Lendenfeld, 1903: 147. Remark. Most probably this genus is synonymous with
Material examined. Holotype: MZUS P0167 - Mexican Leiodermatium Schmidt, but because all material is deciduous any
Gulf, depth 183-1472m. MCZ 6416a (MCZ 237, Cuba), MNHN such decision must be based on better-preserved material.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcadernicIPlenum Publishers, New York, 2002

tFossil 'Lithistids': An Overview

Andrzej Pisera

Instytut Paleobiologii, Polska Akademia Nauk, ul. Twarda 51/55,00-818 Warszawa, Poland. (apis@twarda.pan.pl)

The oldest fossillithistid sponges (Demospongiae, 'lithistids') are known from the Cambrian. Lithistids are very common in the fossil
record since the Ordovician, but their distribution is very punctuated: large faunas of lithistid demosponges are known from the
Ordovician, Silurian, Devonian, Pennian, Upper Jurassic, Upper Cretaceous and the Eocene. The Paleozoic faunas are dominated by
Orchocladina (extinct in the Permian) and Sphaerocladina (continuing up to the Recent); Mesozoic faunas are dominated by Tetracladina
and Rhizomorina, which are also common in the Tertiary and Recent lithistid faunas. Phylogeny of fossillithistids, especially relation-
ships between the Paleozoic and later fonns are still poorly understood, partly because only desmas are usually preserved in the fossil
state, thus no microscleres or rarely ectosomal spicules are known. These megascleres are crucial diagnostic characters in Recent lithistid
taxonomy. Given their good fossilization potential this fauna is diverse, with 13 suborders, 34 families and over 200 genera discussed in
the present work, although many more nominal genera of fossillithistids are known.
Keywords: Porifera; Demospongiae; lithistids; Dicranocladina; Didymmorina; Megamorina; Orchocladina; Rhizomorina; Sphaerocladina;
Tetracladina; Tricranocladina; Paleozoic; Mesozoic; Tertiary; fossil taxa.

INTRODUCTION (edited by J. Keith Rigby, Provo, Utah). This current work dis-
cusses 13 suborders, 34 families and over 200 genera of fossil
The classification of fossil 'lithistid' demosponges (fonnerly lithistids, although many more nominal genera are known.
order Lithistida), similar to most Recent taxonomic studies of
sponges, is based on spicule morphology and complement.
However, at the family level, especially among those with rhizo- PREVIOUS REVIEWS
clone desmas, the habit of the sponges is given a special impor-
tance (see especially Schrarnmen, 1924a). As shown by studies of The main authors associated with fossil lithistids are:
Recent lithistids, the same family (characterized by the same Zittel (1878a); Rauff (1893); Schrammen (1910, I 924a, 1937);
spicule components) encompasses sponges of very different mor- Moret (1926); de Laubenfels (1955b); Lagneau-Herenger (1962);
phologies, thus proving that paleontological classification in this Reid (1968d, 1970); Rigby (1983a, 1986, 1991b); Rigby & Dixon
respect may produce artificial groups of sponges and/or cause split- (1979); Rigby & Webby (1988); Rigby & Chatterton (1989);
ting. One of the reasons for such difficulties is the nearly total Levi (1991); Gruber (1993); Wiedenmayer (1994); Mehl-Janussen
absence of microscleres and frequent absence of ectosomal (1999); and Pisera (1999a).
spicules in the fossil record, which are important in Recent lithistid
taxonomy and phylogenetic reconstructions. The present overview
of fossillithistids does not constitute a revision of the group, which PALEOZOIC LITHISTIDS
is sorely needed but is clearly beyond the scope of the Systema
Porifera project, it presents rather the classification as it is cur- SUBORDER ORCHOCLADINA RAUFF, 1895
rently used. Lists of fossil genera are provided to ensure they are
included in any future revision of this group, even though no diag- Diagnosis
noses or illustrations of the genera are provided here. It would not
be possible to justify some taxonomic decisions in a short review Lithistid demosponges with dendroclones or chiastoclones as
such as this, as well as the enonnous amount of work that has yet choanosomal desmas. No ectosomal spicules known.
to be done prior to undertaking such a revision. These studies are
currently in progress (Pisera, unpublished). Stratigraphical range
As is the case with many other groups of fossil animals there is
profound difference between Paleozoic (for stratigraphic names and Middle Cambrian-Permian.
ages see Stratigraphical Chart; Fig. 22) and later Mesozoic to
Tertiary sponges, and the relationship between particular families or
groups often remains uncertain. These suggested phylogenies can be FAMILY ANTHASPIDELLIDAE ULRICH
found in Reid (1968d, 1970), Gruber (1993) and Mehl-Janussen IN MILLER, 1889
(1999), but some of their opinions must be treated with caution in the
light of more contemporary studies of Recent and fossillithistids. Synonymy
A more exhaustive review of all nominal genera and their
extensive synonymies and family allocations is in preparation for Anthaspidellidae Ulrich in Miller, 1889. Archaeoscyphiidae
the new edition of the Treatise on Invertebrate Palaeontology Rauff, 1894. Aulocopidae de Laubenfels, 1955b.

388
Porifera· Demospongiae· 'Lithistids'· fossil group 389

Fig. 1. Anthaspidellid choanosomal skelton composed of dendroclones Fig. 2. Dendroc1one desmas of anathaspidellid sponges (scale 100 fLm)
(scale 500 fLm) (after Eggink et al., 2001, fig. 13, photo V . van Hacht). (modified after Eggink et al. , 2001 , fig. 46, photo V.van Hacht).

Diagnosis Rigby & Webby, 1988 (Ordovician); Rankenella Kruse, 1983

(Cambrian); Rhopalocoelia Raymond & Okulitch, 1940
Orchocladina with dendroclones as choanosomal desmas (Ordovician); Schismospongia Rhebergen & Hacht, 2000
organized in ladder-like pattern (Figs 1-2); often vertical bundles (Silurian); Sommersetella Rigby & Dixon, 1979 (Silurian);
of oxeas accompany desmas; no ectosomal spicules known. Striataspongia Howell, 1952 (Late Devonian); Talacastonia
Beresi & Rigby, 1993 (Ordovician); Vandonia Rigby & Webby,
Scope (selected genera) 1988 (Ordovician); Vankempenia Von Hacht, 1994 (?Silurian);
Yarrrowigahia Rigby & Webby, 1988 (Ordovician).
Actinocoelia Finks, 1960 (Permian); Amplaspongia Rigby &
Webby, 1988 (Ordovician); Anthaspidella Ulrich & Everett, 1890 Stratigraphical range
(Ordovician) ; Antrospongia Rigby & Chatterton, 1989 (Silurian);
Archaeoscyphia Hinde, 1889 (Ordovician-Silurian); Aulocopella Middle Cambrian-Permian, maximum development in the
Rauff, 1894 (Ordovician); Aulocopoides Howell, 1952 (Late Ordovician.
Devonian); Aulocopium Oswald, 1847 (Ordovician-Silurian)
(fig. 3A-C); Australospongia Howel, 1952 (Late Devonian);
Calycocoelia Bassler, 1941 (Ordovician-Silurian); Canningella FAMILY CHIASTOCLONELLIDAE RAUFF, 1894
Rigby, 1986a (Late Devonian); Cauliculospongia Rigby & (NON DE LAUBENFELS, 1955)
Chatterton, 1989 (Silurian); Cockbainia Rigby, 1986a (Late
Devonian); Defordia King, 1983 (Permian); Diotricheum Van Diagnosis
Kempen, 1989 (Ordovician); Dunhillia Rigby & Webby, 1988
(Ordovician-Silurian); Fibrocoelia Van Kempen, 1978; Orchocladina with chiastoclones as choanosomal desmas; no
Fistulospongia Rigby, 1986a (Late Devonian); Gleesonia Rigby & ectosomal spicules known.
Webby, 1988 (Ordovician); Hudsonospongia Raymond &
Okulitch, 1940 (Ordovician); Lissocoelia Bassler, 1927
(Ordovician); Malongullospongia Rigby & Webby, 1988 Scope (selected genera)
(Ordovician); Multistella Finks, 1960 (Permian); Nevadocoelia
Bassler, 1927 (Ordovician); Patellispongia Bassler, 1941 Allassospongia Rigby, 1986a (Late Devonian);
(Ordovician-Silurian); Phacellopegma Gerth, 1927 (Carboniferous- Anthracosycon Girty, 1909 (Permian); Chiastoclonella Rauff, 1894
Permian); Perissocoelia Rigby & Webby, 1988 (Ordovician- (Ordovician-Silurian) (Fig. 3D); Collatipora Finks, 1960
Silurian) (Fig. 4); Playfordiella Rigby, 1986a (Late Devonian); (Permian); Dactylites Finks, 1960 (Permian); Insulipora Finks,
Protachilleum Zittel, 1877 (Ordovician); Psarodictyum Raymond & 1960 (Permian); Jereina Finks, 1960 (Devonian-Permian);
Okulitch, 1940 (Ordovician-Silurian); Pseudopalmatohindia Syltispongia Van Kempen, 1990b (Silurian).
390 Porifera· Demospongiae· 'Lithistids'· fossil group

Fig.3. A-C, Aulocopium aurantium Oswald, 1879, Silurian; lateral, lower side, and vertical cross-section views, respectively (scale 2 em) (modified after
Roemer, 1861). D, Chiastoclonella headi Rauff, 1894, Silurian, choanosomal skeleton of chiastoclones (scale 200 /Lm) (modified after Rauff, 1894).

SUBORDER HELOMORINA SCHRAMMEN, 1937

Diagnosis

Lithistid demosponges with heloclones as choanosomal

desmas; ectosomal spicules when present are dichotriaenes.

Stratigraphical range

Ordovician-Silurian, ?Permian (see also Mesozoic Helomorina).

FAMILY SACCOSPONGIIDAE RIGBY & DIXON, 1979

Diagnosis

Fig. 4. Perissocoelia sp. Silurian, coil. Natuurmuseum Gotland, Visby Paleozoic lithistid demosponges with heloclones as choano-
(scale 2 em) (after Eggink et al., 2001, fig. I, photo G. Strom, Visby). somal desmas.

Scope (selected genera)

Cliefdenospongia Rigby & Webby, 1988 (Ordovician);

Remarks Eochaunactis Rigby & Dixon, 1979 (Silurian); Finksella Rigby &
Dixon, 1979 (Silurian); Haplistionella Rigby & Dixon, 1979
Sometimes genera here included are regarded as real (Silurian); Saccospongia Bassler, 1889 (Ordovician).
Tetracladina, but this still awaits corroboration.
Remarks
Stratigraphical range
Lithistids with heloclones are generally considered to
Ordovician-Permian. belong to Megamorina (see Reid, 1968b; Rigby, 1983a). However,
Porifera· Demospongiae· 'Lithistids'· fossil group 391

investigations of Recent species with heloclones indicate that helo- 1988 (Ordovician); Parodospongia Rigby & Chatterton, 1989
clones and megaclones are completely different spicules, and thus (Silurian); Warrigalia Rigby & Webby, 1988 (Ordovician); Taplowia
these taxa should be separated. Rigby & Webby, 1988 (Ordovician).

Stratigraphical range Remarks

Upper Ordovician-Silurian, ?Upper Permian. The relationship between Paleozoic rhizomorines with
Mesozoic and younger rhizomorine lithistids remains very unclear
and requires further study.
SUBORDER RHIZOMORINA ZITTEL, 1878
Stratigraphical range
Diagnosis
Ordovician-Permian.
Lithistid demosponges with rhizoclones as choanosomal
desmas.
SUBORDER SPHAEROCLADINA SCHRAMMEN, 1910
Remarks
Diagnosis
The relationship between Paleozoic and Mesozoic
rhizomorine sponges remains unclear and needs further study. Lithistid demosponges with sphaeroclones or astroclones as
choanosomal desmas.
Stratigraphical range
Remarks
Ordovician-Permian (but see Mesozoic rhizomorines).
See remarks on Mesozoic Sphaerocladina below.

FAMILY HAPLISTIIDAE DE LAUBENFELS, 1955

Stratigraphical range
Diagnosis
See Astylospongiida, and Mesozoic sphaerocladine lithistids.
Paleozoic lithistid demosponges with rhizoclones as
choanosomal desmas.
FAMILY ASTYLOSPONGIIDAE ZITTEL, 1878
Scope
Diagnosis
Boonderooia Rigby & Webby, 1988 (Ordovician); Haplistion
Young & Young, 1877 (Silurian-Permian); Lewinia Rigby & Webby, Paleozoic lithistid demosponges with sphaeroclone desmas.

Fig. 5. A-C, Astylospongia praemorsa (Goldfuss, 1826), Silurian, lateral top, and vertical cross-section views respectively (scale 1 cm) (modified after
Rauff, 1894). D, Carpospongia globosa (Eichwald, 1830), choanosomal skeleton of sphaeroclones (scale 200/-Lm) (modified after Rauff, 1894).
E, Astylospongia sp. sphaeroclone (scale 100/-Lm) (modified after Rauff, 1894).
392 Porifera· Demospongiae· 'Lithistids'· fossil group

Scope (restricted) Scope

Astylostroma Rigby & Webby, 1988 (Ordovician); Arborohindia Rigby & Webby, 1988 (Ordovician);
Astylospongia Roemer, 1860 (Ordovician-Silurian) (Fig. 5A-C, E); Belubulaspongia Rigby & Webby, 1988 (Ordovician);
Astyloscyphia Rigby & Pisera, 2001 (Devonian); Astylospongiella Cotylahindia Rigby & Bayer, 1971 (Ordovician). Fenestrospongia
Rigby & Lenz, 1978 (Silurian); Astylotuba Rigby & Pisera, 2001 Rigby & Webby, 1988 (Ordovician); Hindia Duncan, 1879
(Devonian); Attungaia Pickett, 1969 (Devonian); Carpospongia (Ordovician-Permian) (Fig. 6); Mamelohindia Rigby & Webby,
Rauff, 1894 (Ordovician-Silurian) (Fig. 5D); Devonoscyphia 1988 (Ordovician); Palmatohindia Rigby & Webby, 1988
Rietschel, 1968 (Devonian); Garraspongia Picket & Rigby, 1983 (Ordovician); SadIe ria Rigby, 1986a (Late Devonian); Scheiia
(Devonian); lazwicella Rigby & Pisera, 2001 (Devonian); Tschemyschev & Stepanov, 1916 (?Carboniferous-Pennian);
Syltrochos Von Hacht, 1981 (?Silurian). Scheiella Finks, 1971 (Pennian); Scheielloides Rigby, 1986a (Late
Devonian).
Remarks
Stratigraphical range
A questionable astylospongiid Ellesmerospongia Rigby, 1970
was described from the Permian, which would make it the Ordovician-Permian.
youngest representative of the family, but its true affinities remain
uncertain. Relationships between Paleozoic and Mesozoic genera
with sphaeroclones has yet to be verified given the large time gap
between them. Preliminary comparisons of their desmas reveal, MESOZOIC-TERTIARY LITIDSTIDS
however, that apart from difference in their size (Palaeozoic sphae-
roclones are much larger), they are identical. Mostler & Balogh SUBORDER DICRANOCLADINA SCHRAMMEN, 1924
(1994) suggested Paleozoic and Mesozoic sphaerocladine lithistids
should be separated, previously considered to be a single group by Diagnosis
Reid (1968d, 1970), but this approach is not followed here.
Lithistid demosponges with dicranoclones as choanosomal
Stratigraphical range desmas and dichotriaenes as ectosomal spicules.

Ordovician- Late Devonian (?Permian). Remarks

SUBORDER TRICRANOCLADINA REID, 1968 Only one family is included.

Diagnosis Stratigraphical range

Lithistids with tricranoclones as choanosomal desmas. See Corallistidae.

Scope

Only one family Hindiidae Rauff, 1893 is included. FAMILY CORALLISTIDAE SOLLAS, 1888

Stratigraphical range Synonymy (restricted)

See Hindiidae. Pachinionidae Schrammen, 1924a. Phrissospongiidae

Lagneau-Heranger, 1962. Spinocladiidae Lagneau-Herenger,
FAMILY IDNDIIDAE RAUFF, 1893 1962. See also chapter on Recent Corallistidae.

Diagnosis Diagnosis

As for suborder. See Recent lithistids.

Fig. 6. Hindia sphaeroidalis Duncan, 1879, Silurian (modified after Rauff, 1894). A, habitus of the sponge (scale 1 em). B, cross section (scale as for A).
C, choanosomal tricranoclon (scale 100 /-Lm).
Porifera· Demospongiae • 'Lithistids'· fossil group 393

Fig. 7. Leiocarenus papillosus Schrammen, 1937, Upper Jurassic. A, cotype, Schrammen, 1937: pI. 20: 9 (scale 1 cm). B, J, choanosomal dicranoclones
in various orientations (scale 200 fLm).

Scope (selected genera)

Dicranoclonella Schrammen, 1937 (Late Jurassic);

Gignouxia Moret, 1926 (Late Cretaceous); Gilletia Lagneau-
Herenger, 1962 (Early Cretaceous); Iouea Laubenfels, 1955b
(=Phalangium Schrammen) (Late Cretaceous); Kyphoclonella
Kolb, 1910 (Late Jurassic); Leiocarenus Schrammen, 1937 (Late
Jurasssic) (Fig. 7); Pachinion Zittel, 1878a (Cretaceous) (Fig. 8A);
Phrissospongia Moret, 1926 (Late Cretaceous); Procorallistes
Schrammen 1901 (Late Cretaceous-Tertiary) (Fig. 8B);
?Pseudoverruclina Moret, 1926 (Cretaceous); Pycnoclonella
Lagneau-Herenger, 1962 (Early Cretaceous); Spinocladia
Lagneau-Herenger, 1962 (Early Cretaceous).

Remarks
Fig. 8. Corallistid morphology. A, Pachinion scriptum (Roemer, 1864),
Several fossil families of Dicranocladina have been proposed, Cretaceous (scale 1 cm). B, Procorallistes polymorphus Schrammen, 1901,
based mostly on their morphology, but these taxa are rejected here. Creatceous (scale 2cm).
394 Porifera· Demospongiae • 'Lithistids' • fossil group

Nevertheless, future revisions based on skeletal characters will Diagnosis

probably recognise distinctive characters that can define separate
families, but these characters are presently elusive. As for suborder.

Stratigraphical range Scope

Late Jurassic-Recent, common in the Late Jurassic and Late Coscinodiscus Schrarnmen, 1937 (Late Jurassic);
Cretaceous. Cylindrophyma Zittel, 1878a (Late Jurassic) (Fig. 9); Linochone
Schrammen, 1937 (Late Jurassic); Melonella Zittel, 1878a
(Jurassic).
SUBORDER DIDYMMORINA RAUFF, 1893
Remarks
Diagnosis
This family has undoubted representatives only from the Late
Lithistid demosponges with didymmoclones as choanosomal Jurassic. There are, however, dubious reports (Le., taxa with very
spicules; ectosomal spicules are dichotriaenes. poor preservation) of Cylindrophyma from the Early Jurassic
(Nutsubidze, 1965). Mehl & Fiirsich (1997) also placed their lithis-
Remarks tid lumarella (from the middle Jurassic) into Cylindrophymatidae,
but their sponge has sphaeroclone desmas and thus belongs to
Only one family is included. Vetulinidae.

Stratigraphical range Stratigraphical distribution

See Cylindorphymidae. Jurassic.

FAMILY CYLINDROPHYMIDAE SCHRAMMEN, 1937 SUBORDER HELOMORINA SCHRAMMEN, 1924

Synonymy Diagnosis

Cylindrophymidae Schrarnmen 1937. Cylindrophymatidae de Lithistid demosponges with heloclones as choanosomal

Laubenfels, 1955b. Coscinodiscidae Schrarnmen, 1937. desmas; ectosomal spicules when present are dichotriaenes.

Fig.9. Cylindrophyma milleporata Goldfuss. A, holotype of Goldfuss, 1833 (scale 2cm). B, choanosomal didymmoclones in various orienation (scale 200 j-Lm).
Porifera· Demospongiae· 'Lithistids'· fossil group 395

FAMILY ISORAPHINIIDAE SCHRAMMEN, 1924 Scope

(EMENDED)
Anomorphites Ko1b, 1910 (Late Jurassic); Amphilectella
Synonymy Schrammen, 1901 (Late Cretaceous); Brochodora (Late
Cretaceous); Doryderma Zittel, 1878a (Late Cretaceous);
Helomorinidae Schrammen, 1910. Helomorinides Moret, Heterostinia Zittel, 1878a (Cretaceous); Homalodora (Late
1926. Isoraphiniidae Schrammen, 1924a; de Laubenfels, 1955b; Cretaceous); lnodia Moret, 1926 (Late Cretaceous); Megalithista
Lagneau-Herenger, 1962. Isoraphiniadae Schrammen, 1937. Zittel, 1878a (Late Jurassic) (Fig. 10); Pachypoterion Hinde, 1883
(Cretaceous) (Fig. 11); Propleroma Moret, 1926 (Cretaceous-
Diagnosis Tertiary).

See chapter on Recent Isoraphiniidae lithistids. Remarks

Scope Fossil pleromids were split into several families by

Schrammen (1924a), based on vague morphological criteria which
lsoraphinia Zittel, 1878a (Late Cretaceous); Heloraphinia have little practical value unless we are dealing with known
Schrammen, 1937 (Jurassic-Cretaceous); Pachycothon species, and for this reason a conservative approach is here adopted
Schrammen, 1901 (Jurassic-Cretaceous). by synonymising these families. Nevertheless, it is possible that in
the future Pleromidae could be divided into different families
Remarks based on spicule characteristics of the included species, but any
proposed scheme requires substantial further study of these taxa.
Isoraphiniids are often considered to belong to Megamorina, Fossil pleromids display considerable morphological diversifica-
based on apparent continuity of megaclones and heloclones (Reid tion and may be plate like, massive irregular, cylindrical with atrial
1968b; Rigby 1983a). However, investigations by Levi (1991) and cavity, funnel-shaped or branched.
the present author (Pisera, unpublished) of megaclones and
heloclones in Recent lithistids have shown that these are very Stratigraphical distribution
different spicules, and thus Isoraphiniidae should be excluded from
Megamorina, as suggested previously by Schrammen (1937). Late Jurassic-Recent.

Stratigraphical distribution

Jurassic-Recent. SUBORDER SPHAEROCLADINA SCHRAMMEN, 1910

Diagnosis

SUBORDER MEGAMORINA ZITTEL, 1878 Lithistid demosponges with sphaeroclones or astroclones as

desmas; ectosomal spicules are known only in forms with astro-
Diagnosis clones, consisting of dichotriaenes.

Lithistid demosponges with megaclones as choanosomal Remarks

desmas; ectosomal spicules, where known, are dichotriaenes.
Mostler & Balogh (1994) suggested that Mesozoic sponges
Stratigraphical distribution with sphaeroclones belong to a separate group, which they called
Neosphaerocladina. Because there is no difference, except perhaps
Jurassic-Recent, loose megaclones appear in the Triassic. size, between the Paleozoic and Mesozoic sphaeroclones, the orig-
inal classification is here accepted, including both Paleozoic and
Mesozoic forms, in one suborder.

FAMILY PLEROMIDAE SOLLAS, 1888 Stratigraphical range

Synonymy (restricted) Silurian-Devonian, and Jurassic-Recent (loose spicules are

already known from the Ordovician).
Pleromidae Sollas, 1888. Doryderrnidae Schrammen, 1924a.
Heterostinidae Schrammen, 1924a. Megalithistidae Schrammen,
1937.
FAMILY LECANELLIDAE SCHRAMMEN, 1937
Diagnosis
Diagnosis
Polymorphic; with megaclones as desmas; ectosomal spicules
are dichotriaenes; other triaenes may be also present. See also Lithistid demosponges with astroclones as choanosomal
chapter on Recent Pleromidae. desmas; ectosomal spicules are dichotriaenes.
396 Porifera· Demospongiae • 'Lithistids' • fossil group

Fig. 10. Megalithistiajoraminosa Zittel, 1878, Upper Jurassic. A, habitus (scale 2cm). B-J, choanosomal megaclones (scale 200 j.Lm).

Remarks

The first ectosomal dichotriaenes were recently discovered

in situ in Lecanella (Pisera, 1997).

Stratigraphical range

Late Jurassic.

FAMILY VETULINIDAE LENDENFELD, 1903

Synonymy

Vetulinidae Lendenfeld, 1903. Sphaerocladiniidae

Schrammen, 1910. Mastosiidae de Laubenfels, 1955b.

Diagnosis

See chapter on Recent Vetulinidae.

Fig. 11. Pachypoterion robustum Hinde, 1883; Cretaceous. A, habitus Scope

(scale 3cm). B-C, details of choanosomal skeleton of megaclones (scales
1 mm) (modified after Hinde, 1883).
Cladodia Moret, 1926 (Late Cretaceous); Exodictya Moret,
1926 (Late Cretaceous); lumarella Mehl & Fiirsich, 1997 (Middle
Scope Jurassic); Macrobrochus Schramrnen, 1910 (Late Cretaceous);
Mastosia Zittel, 1878a (Late Jurassic) (Fig. 12); Ozotrachelus de
Lecanella Zittel, 1878a (Late Jurassic). Laubenfels, (=Pachytrachelus Schramrnen, 1910) (Late
Porifera· Demospongiae· 'Lithistids'· fossil group 397

Fig. 12. Mastosia wetzleri Zittel, 1878, Upper Jurassic. A, holotype habitus (scale 4 em). B-L, choanosomal sphaerocJones (scale 100 fLm).

Cretaceous); Regnardia Moret, 1926 (Late Cretaceous); Remarks

Rhytidodenna Schrammen, 1937 (Late Jurassic).
Lithistid sponges with rhizoclone desmas are traditionally
Remarks classed together into one subordinal group, Rhizomorina. No ecto-
somal spicules or microscleres are known in fossil forms, and thus
The genus lumarella (from the middle Jurassic) was the subdivision of fossil rhizomorine lithistids into families causes
originally attributed to Didymorina (Mehl & Fiirsich, 1997), serious problems. Schrammen (1924a), in an extreme action of
but as it displays sphaeroclones this allocation was clearly a splitting, established numerous rhizomorine families mostly based
mistake. on very vague morphological criteria (sometimes wall-thickness),
which is very difficult to apply especially when dealing with new
Stratigraphical range forms. In my opinion Schrammen's proposition is impossible to
accept and must be substituted for something more operational. For
Middle Jurassic to Late Cretaceous, and Recent; loose sphae- these reasons nominal families are listed here separately, and then
roclones are also known from the Tertiary. all genera, without attributing them to higher taxonomic categories.

Scope (families)
SUBORDER RHIZOMORINA ZITTEL, 1878
Amphichondridae Schrammen, 1924a; Amphitelionidae
Diagnosis Schrammen, 1924a; Aulosomidae Schrammen, 1924a;
Chonellidae Schrammen, 1924a; Cnemidiastridae Schrammen,
Lithistid demosponges with rhizoclones as choanosomal des- 1937; Cytoraceidae Schrammen, 1924a; Heterothelionidae
mas; no ectosomal spicules are known. Schrammen, 1924a; Hyalotragosidae Schrammen, 1937; Jereicidae
398 Porifera· Demospongiae· 'Lithistids'· fossil group

Schrammen, 1924a; Leiochonidae Schrammen, 1924a; Brimaud & Vachard, 1986 (Tertiary); Cytoracea Pomel, 1872
Leiodorellidae Schrammen, 1937; Lophiophoridae Schrammen, (Jurassic-Cretaceous). Epistomella Zittel, 1878a (Late Jurassic)
1924a; Oncophoridae Schrammen, 1924a; Platychoniadae (Fig. 14); Jereica Zittel, 1878a (Cretaceous-Tertiary); Jereopsis
Schrammen, 1937; Pyrgochoniadae Schrammen, 1937; Scytalidae Pomel, 1872 (Tertiary); Leiochonia Schrammen (Cretaceous);
de Laubenfels, 1955b; Seliscothonidae Schrammen, 1924a; Leiodorella Zittel, 1878a (Jurassic) (Fig. 13A); Lithostrobilus
Trachynotidae Schrammen, 1924a; Verrculinidae Schrammen, Schrammen, 1937 (Cretaceous-Jurassic); Lophiophora Schrammen,
1924a. 1924a (Late Cretaceous); Moretispongia Breistroffer, 1947
(=Marisca Pomel 1872, non-Gray 1840) (Cretaceous - Tertiary);
Scope (selected genera) Pachysalax Schrammen, 1910 (Late Cretaceous); Pachyselis
Schrammen, 1924a (Late Cretaceous); Phlyctia Pomel, 1872
Aulosoma Schrammen, 1924a (Cretaceous); Chonella Zittel, (Tertiary); Platychonia Zittel, 1878a (Jurassic); Plinthodermatium
1878a (Cretaceous-Tertiary); Chonellopsis Schrammen, 1937 Schrammen, 1910 (Cretaceous); Pseudocytoracea Lagneau-
(Jurassic); Cnemidiastrum Zittel, 1878a (Jurassic, Tertiary) Herenger, 1962 (Early Cretaceous); Seliscothon Zittel, 1878a
(Fig. 13B); Coscinostoma Schrammen, 1910 (Cretaceous- (Cretaceous-Tertiary) (Fig. 15); Pseudoseliscothon Moret, 1926
Tertiary); Coelosphaeroma Schrammen, 1910 (Cretaceous); (Cretaceous); Reiswigia Trammer, 1979 (Late Jurassic);
Coelocorypha Zittel, 1878a (Jurassic-Cretaceous); Cucumaltina Rhabdotum Schrammen, 1924a (Upper Cretaceous); Scytalia
Zittel, 1878a (Cretaceous); Stachyspongia Zittel, 1878a
(Cretaceous-Tertiary); Stichophyma Zittel, 1878a (Cretaceous-
Tertiary); Verruculina Zittel, 1878a (=Amphitelion Zittel, 1878a)
(Middle Jurassic-Tertiary).

B A c
Fig. 13. Mesozoic rhizomorine lithistids. A, Leiodorella expansa Zittel,
1878, Upper Jurassic (scale I cm) (after Zittel, 1878a). B, Cnemidiastrum Fig. 15. Seliscothon planum (Phillips, 1835), Cretaceous (scales I cm,
stella tum (Goldfuss, 1833) (scale 1 cm) (after Zittel, 1878a). 200 /-Lm) (modified after Hinde, 1883).

Fig. 14. Epistomelia clivosa (Quenstedt, 1878), Upper Jurassic. A, holotype of Quenstedt, 1878, pI. 131: 4 (scale 2cm). B-J, rhizoclone desmas (scale 100 /-Lm).
Porifera· Demospongiae • 'Lithistids' • fossil group 399

Remarks Stratigraphical distribution

Mesozoic to Tertiary rhizomorine lithistids are very diversi- Cretaceous-Recent; loose desmas also occur in the Late
fied morphologically and range from massive to cylindrical or Jurassic (Wiedenmayer, 1994).
nearly spherical, with or without an atrial cavity, or often with a
bundle of exhalant canals at the center, to leaf or vase shaped, with
or without canals visible on the surface or in the interior of the FAMILY PHYMATELLIDAE SCHRAMMEN, 1910
wall. Also, choanosomal rhizoclone desmas differ considerably.
These range from massive to rather slender, usually arched, and Synonymy (restricted)
supplied with various amounts of pointed thorns. Rhizoclones may
be organized into a fibrous-reticulate skeleton, or a dense disorgan- Phymatellinae Schrammen, 1910. Phymatellidae Schrammen,
ized skeleton. Often one or both surfaces display circular canal 1924a. Acrochordonidae Schrammen, 1924a.
openings raised above the surface, or the surface(s) may bear only
small pores. These characters, as used informally by Moret (1926), Diagnosis
together with desmas, will probably serve as the basis for a more
natural subdivision of rhizomorine lithistids into suprageneric units. See chapter on Recent Phymatellidae lithistids.

Stratigraphical range Scope (restricted)

Early Jurassic-Recent (loose rhizoclones already recorded Acrochordonia Schrammen, 1910 (Cretaceous);
from the Triassic), very common in the Late Jurassic and Late Achrochordiella Rigby, 1981 (Tertiary); Asterocalyx Moret, 1926
Cretaceous faunas. (Late Cretaceous); Astrolemma Schrammen, 1924a (Late
Cretaceous); Aulaxinia Zittel, 1878a (Cretaceous) (Figs 17A, 18);
SUBORDER TETRACLADINA ZITTEL, 1878 Callopegma Zittel, 1878a (Late Jurassic-Cretaceous) (Fig. 17D-E);
Callymatina Zittel, 1878a (Cretaceous); Craterella Schrammen,
Diagnosis 1901 (Cretaceous); Hallirhoa Lamouroux, 1821 (Cretaceous);
Jerea Lamouroux, 1821, emended Zittel, 1878a (Cretaceous)
Lithistid demo sponges with tetraclones or their modifications (Fig. 17C); Kalpinella Hinde, 1883 (Jurassic-Cretaceous);
as choanosomal desmas; ectosomal spicules where known are Myrmeciophytum Schrammen, 1910 (Cretaceous); ParaspeZaeum
dichotriaenes, discotriaenes, phyllotriaenes or anaxial plates. Schrammen, 1924a (Late Cretaceous); Phymatella Zittel,
1878a (Cretaceous); Pliegatella Brimaud & Vachard, 1986
Stratigraphical distribution

Triassic-Recent.

FAMILY PHYMARAPHINIIDAE SCHRAMMEN, 1910

Synonymy

Phymaraphininae Schrammen, 1910: 104. Phymaraphiniidae

Schrammen, 1924a; Lagneau-Herenger, 1962. Phymaraphinidae
Levi, 1991. Kaliapsidae de Laubenfels, 1936 (partim).

Diagnosis

See chapter on Recent lithistids.

Scope

Phymaraphinia Schrammen, 1910 (Cretaceous) (Fig. 16);

Cyclocnema Schrammen, 1910 (Late Cretaceous); Pholidocladia
Hinde, 1883 (Cretaceous); Prokaliapsis Schrammen, 1901
(Cretaceous); Lopadophorus Schrammen, 1910 (Cretaceous);
Polyrhypidium Schrammen, 1924a (Cretaceous); Stelidium
Schrammen, 1924a (Late Cretaceous).

Remarks

Phymaraphiniidae are common in the Late Cretaceous sponge

fauna of Europe; there is also an undescribed representative of this Fig. 16. Phymaraphinia infundibuliformis Schrammen, Upper Cretaceous
family from the Eocene of SW Australia (Pisera, unpublished). (scales 500 fLm). A, choanosomal triders. B, ectosomal phyllotriaenes in situ.
400 Porifera· Demospongiae • 'Lithistids' • fossil group

Fig. 17. Phymatellid morphology. A. Aulaxinia sulcifera (Roemer. 1864), Upper Cretaceous (scale 2cm). B, Turonia variabilis Michelin, 1846, cotype
(scale 3 cm). C, Jerea quenstedti Zittel, 1878, Upper Cretaceous (scale 2 cm). D, E, Cal/opegma acaulis Zitte, 1878, Upper Cretaceous (scale I cm).

(Cretaceous-Tertiary); Polyjerea Fromental, emended Zittel,

1878a (Late Cretaceous); Siphonia Parkinson, emended Zittel,
1878a (Cretaceous); Trachysycon Zittel, 1878a (Cretaceous);
Thecosiphonia Zittel, 1878a (Cretaceous); Turonia Michelin, 1846,
(Cretaceous) (Fig. 17B).

Remarks

Many of the genera included here are poorly known,

described only from single records, and probably a number of these
should be greatly reduced as they are usually based on only slight
morphological differences.

Stratigraphical distribution

Fig. 18. Aulaxinia sulcifera (Roemer, 1864), Cretaceous, choanosomal Late Jurassic-Recent, although the first phyllotriaenes are
skeleton composed of tetraclones (scale 500 fLm). known from the Triassic (Wiedenmayer, 1994).
Porifera· Demospongiae· 'Lithistids'· fossil group 401

FAMILY PLINTHOSELLIDAE SCHRAMMEN, 1910 (Fig. 19); Pycnodesma Schrammen, 1910 (Late Cretaceous);
Phymaplectia Hinde, 1883 (Cretaceous).
Synonymy
Remarks
Plinthosellinae Schrammen, 1910. Plinthosellidae Schrammen,
1924; Pisera, 2000: 293. Dactylotidae Schrammen, 1924a. This family is sometimes considered synonymous with the
Recent Neopeltidae Sollas, 1888, but there are strong differences in
Diagnosis the morphology of desmas, and probably also ectosomal spicules,
which are clearly monaxial in Neopeltidae and probably anaxial in
Lithistids with trider-like desmas and anaxial (?) siliceous Plinthosellidae. These data suggest their similarities are only
plates as ectosomal spicules. superficial.

Scope Stratigraphical range

Ingentilotus de Laubenfels, 1955b (=Dactylotus Schrammen, Late Cretaceous-Tertiary.

1910) (Cretaceous); Plinthosella Zittel, 1878a (Cretaceous-Tertiary)
FAMILY THEONELLIDAE LENDENFELD, 1903

Restricted synonymy

Theonellidae Lendenfeld, 1903 (partim). Astrocladiidae

Schrammen, 1924a. Astrocladinae Schrammen, 1910. Discoder-
minae Schrammen, 1910. Discodermiidae Schrammen, 1924a; de
Laubenfels, 1955b (partim). Lerouxidae Moret, 1926. Chenen-
doporidae Schrammen, 1924a.

Diagnosis

See chapter on Recent Theonellidae lithistids.

Scope (selected genera)

Fig. 19. Plinthosella squamosa Zittel, 1878; choanosomal skeleton Astrocladia Zittel, 1878a (Late Cretaceous) (Fig. 20C);
composed of trider-like desmas, Upper Cretaceous (scale I mm). Cladodermia Schrammen, 1924a (Late Cretaceous); Chenendopora

Fig. 20. Fossil theonellid morphology. A, Rhagadinia rimosa (Roemer, 1864), Cretaceous (scale 2cm). B, Phyllodermia antiqua (Schrammen, 1901),
Cretaceous (scale 2cm). C, Astrocladia subramosa (Roemer, 1864), Cretaceous (scale 3cm).
402 Porifera' Demospongiae· 'Lithistids" fossil group

C
I
gJ: (J)
i
0

i~
UJ.
~ c::UJ (!)ell
UJ c:t::2:
I
a..
HOLOCENE
I () QUATERNARY PLEISTOCENE
0 >- 1.75

t5
PLIOCENE
c::: NEOGENE 5.30
c:t: MIOCENE
OLIGOCENE
23.5
Z
~ PALEOGENE 33.7
I~ i~
EOCENE
PALEOCENE
53.0
65.0
LATE
() CRETACEOUS -~-----

EARLY
135.0
0
N
LATE
JURASSIC MIDDLE
0 EARLY
CJ)
LATE
203.0
ill
:2: TRIASSIC MIDDLE
EARLY
250.0
i LOPINGIAN
PERMIAN GUADALUPIAN
CISURALIAN
PENNSYLVANIAN
295.0
iCARBONIFEROUS MISSISSIPPIAN
355.0
gl LATE
O· DEVONIAN MIDDLE

~I EARLY
PRIDOLI
410.0

~I
LUDLOW
SILURIAN WENLOCK
LLANDOVERY
435.0
i LATE
ORDOVICIAN MIDDLE
EARLY
500.0
Fig. 21. Phyllodermia antiqua (Schrammen, 1901), Upper Cretaceous. LATE
A, choanosomal skeleton composed of tetraclones (scale 200 !Lm). B, ecto- CAMBRIAN MIDDLE
somal phyllotriaenes in situ (scale 500 !Lm). ........ I............ .L EARLY . ..............
545.0
*After International Stratigraphical Chart 2000
International Union of Geological Sciences

Lamouroux, 1821 (Cretaceous-Tertiary); Colossolacis Schrammen, Fig. 22. Geological time scale.
1910 (Late Creataceous); Eustrobilus Schrammen, 1910
(Cretaceous); Jereomorpha Moret, 1926 (Late Cretaceous);
Leiophyllum Schrammen, 1924a (Late Cretaceous); Lerouxia LITmSTIDS INCERTAE SEDIS
Moret, 1926 (Cretaceous-Tertiary); Mastophorus Schrammen,
1924a (Late Cretaceous); ?Microcladina Lagneau-Herenger, 1962 FAMILY MEGARHIZIDAE SCHRAMMEN, 1903
(Early Cretaceous); Microdendron Schrammen, 1910 (Late
Cretaceous); Phyllodermia Schrammen, 1924a (Cretaceous- Diagnosis
Tertiary) (Figs 20B, 21); PZacoscytus Schrammen, 1901 (Late
Cretaceous); Pseudojerea Moret, 1926 (Late Creraceous); Lithistid demo sponges with large choanosomal desmas
Rhagadinia Zittel, 1878a (Cretaceous) (Fig. 20A); Rhoptrum branched in numerous planes, slightly resembling astroclones.
Schrammen, 1910 (Cretaceous); Thamnospongia Hinde, 1883
(Cretaceous); Theonella (Tertiary-Recent); Verrucodesma Lagneau- Scope
Herenger, 1962 (Early Cretaceous).
Megarhiza Schrammen, 1901 (Cretaceous); ChaZaropegma
Remarks Schrammen, 1910 (Cretaceous).

Schrammen (1924a) established families Astrocladiidae and Remarks

Chendoporidae based on small differences in desma size and mor-
phology, in which the presence of ectosomal phyllotriaenes was The des mas of megarhizids are often considered to be similar
also reported (Pis era 2000), and which are here considered synony- to rhizoclones, but in fact these differ significantly from typical rhi-
mous with Theonellidae. Lerouxidae Moret, 1926 also differs zoclones. There is some resemblance to astroclones, and also to
vaguely only in desma shape, and as such are synonimized with desmas, for example, of Recent Petromica. For this reason the
Theonellidae (see Pisera, 2000). position of megarhizids remains uncertain.

Stratigraphical range Stratigraphical range

Late Jurassic-Recent. Late Cretaceous.

Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A Hooper and Rob w.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Order Poecilosclerida Topsent, 1928

John N.A. Hooper! & Rob W.M. Van Soest2

Poecilosclerida Topsent (Demospongiae) contains the highest diversity of species and possibly the greatest range of morphometric char-
acters of all Porifera, with 25 families, 129 genera and 50 subgenera recognised as valid, and several thousands of described species
worldwide, distributed from intertidal to abyssal depths. Four suborders are delineated based primarily on: chelae geometry (palmate-
arcuate-anchorate forms, and diancistra derivatives); presence/absence of discate microrhabds; presence/absence and geometry of other
microscleres; ectosomal megasclere geometry; and choanosomal megasclere diversity, geometry and regionalisation of megascleres
within the skeleton. The order is defined as containing demosponges with discrete organic and inorganic skeletons, with discrete siliceous
spicules, a skeletal organisation commonly localized to particular regions (at a minimum ectosomal and choanosomal skeletons but poten-
tially containing up to five regionally differentiated megasclere morphologies), microscleres consisting of meniscoid spicules (sigmas and
derivatives such as chelae, sigmancistras, diancistras, etc., with chelae synapomorphic for the order, although not all taxa contain them),
in addition to diverse other forms such as toxas, forceps, onychaetes, raphides (in trichodragmata), microrhabds, acanthomicrorhabds,
anisodiscorhabds, isochiadiscorhabds, spinorhabds, thraustoxeas, spirosigmata, thraustosigmata and microstyles.
Keywords: Porifera; Demospongiae; Poecilosclerida; Microcionina; Myxillina; Mycalina; Latrunculina.

DEFINITION, DIAGNOSIS, SCOPE types of megascleres (with or without distinctive geometries): the
larger usually choanosomal megascleres embedded in spongin
Synonymy fibres (=principal megascleres), and often smaller ones which are
free in the mesohyl (=subectosomal auxiliary megascleres), or
Poeciloscleridae Topsent, 1894c: 5. Poecilosclerina Topsent, localized at the surface (=ectosomal megascleres), or protruding
1928c: 41, 43. Poecilosclerida Topsent (nomen correctum de from spongin fibres in which they are embedded (=echinating
Laubenfels, 1955b: E2l). megascleres) - sometimes all the latter categories are present.
Conversely, various mycalid poecilosclerids have only a single
Definition category of megasclere ('mycalostyles'), with the presumption that
the differentiated ectosomal and echinating megascleres have been
Demospongiae with skeleton composed of discrete siliceous secondarily lost (Van Soest, 1984b), or alternatively, that their pres-
spicules; main skeleton composed of megascleres (monactinal, ence in most other poecilosclerids is an evolutionary achievement
diactinal or both) and spongin fibres in various stages of develop- (Hajdu et al., 1994a). Poecilosclerids usually have an abundantly
ment; both fibre and mineral skeletons always show regional differ- collagenous mesohyl, and microscleres include chelae (a synapo-
entiation such that megascleres are often differentiated into distinct morphy for the order), although not all taxa have them. Sexual
ectosomal and choanosomal components; microscleres include reproduction is predominantly viviparous, oviparous in two fami-
meniscoid forms such as chelae (unique to the order), sigmas and lies (Raspailiidae, and possibly Rhabderemiidae), and in those
sigmancistra-derivatives, and other diverse forms such as toxas, species incubating larvae they are parenchymella with uniform
raphides, microxeas and discate microrhabds; the order is predomi- flagellum size and bare posterior poles.
nantly viviparous with incompletely ciliated parenchymella larvae
(one oviparous family (Raspailiidae), and another suspected Remarks
oviparous family (Rhabderemiidae), is also included).
Poecilosclerida is the most diverse of all orders of Porifera
Diagnosis (Bergquist, 1978; Hartman, 1982), both in terms of numbers of
species and the range and diversity of morphological characters
Sponges characterized by a skeleton of both spicule and spon- (Bergquist & Fromont, 1988; Hajdu et al., 1994a). This diverse
gin elements, usually well-developed, either sometimes vestigial, in morphology is certainly also reflected in the many different opin-
which megascleres are monactinal, diactinal or both, and spongin ions concerning interpretation of character importance, character
development varies from well-developed horny fibres enclosing polarity, and indeed character homology itself, producing a prolif-
spicules to an interspicular collagen cement (Bergquist, 1978; eration of systematic schemes for the group (discussed in detail by
Hartman, 1982). A few groups have a fused (hypercalcified) basal Bergquist & Fromont, 1988).
calcitic skeleton or an articulated siliceous skeleton composed of Our present understanding of the concept of Poecilosclerida
desmas ('lithistid' or 'sublithistid'), along with free siliceous skele- stems from Ridley & Dendy's (1887) order Monaxonida, in
tons, with clues as to their identities provided by these free spicules, which poecilosclerid genera were placed in two families of the
and these desmas are interpreted as ancestral features, more com- suborder Halichondrina (Heterorrhapididae, lacking chelae; and
monly observed in the Mesozoic faunas. Simpson (1984) suggested Desmacidonidae with chelae). Topsent (1894c) refined this
that the order was characterized by at least two distinctly localized scheme, although splitting the non-cheliferous and cheliferous

403
404 Porifera· Demospongiae • Poecilosclerida

poecilosclerids into separate families Haploscleridae and as 'convenient' clades to manage large suites of taxa, whereas
Poeciloscleridae, respectively. Hentschel (1923) also provided a higher taxa are based on more sound phylogenetic principles,
major contribution to the present concept of Poecilosclerida in including published character analyses.
establishing a number of families, many of which are still recog-
nised today, but he persisted in retaining a subordinal distinction Scope
between taxa with or without chelae, and placed primary emphasis
on the presence or absence of spiny megascleres ('acanthostyles') The number of families recognised in the order varies
in his subordinal classification. Topsent (1928c) established according to different authors (e.g., Levi, 1973; Wiedenmayer,
Poecilosclerida to emphasize the unique possession of chelae 1977b; Bergquist, 1978; Hartman, 1982; Van Soest, 1984b;
microscleres, and his definition and families included have been Bergquist & Fromont, 1988). Recently, Hooper & Wiedenmayer
maintained more-or-Iess intact up to the present. De Laubenfels (1994) included 16 families in the order (although these
(1936a) purposefully, or otherwise, destabilised this phylogenetic were based solely on the published Australian fauna, with several
system in an attempt to simplify the poriferan nomenclature and families not yet 'officially' recorded for this fauna): twelve
classification, while explicitly acknowledging little heed for phylo- with chelae microscleres, three without chelae, and one of
genetic relationships among family-level taxa. His classification has uncertain placement. Hajdu et al. (1994a) presented a more
been adopted by a few authors, at family and sometimes higher tax- global view, recognising 17 families: Anchinoidae, Cladorhizidae,
onomic levels, up until the 1980s (e.g., Hoshino, 1981a, b; Pulitzer- Coelosphaeridae, Crambidae (=Crambeidae), Crellidae, Desma-
Finali, 1983), but this is now mostly rejected by contemporary cellidae, Guitarridae, Hamacanthidae, Hymedesmiidae, Iophonidae
authors who have more-or-Iess returned to Topsent's (1928c) (=Acarnidae), Microcionidae, Mycalidae, Myxillidae, Phorio-
scheme, following the lead of Levi (1973) and Bergquist (1978) in spongiidae, Raspailiidae, Rhabderemiidae, Tedaniidae.
particular. Bergquist & Fromont (1988) provide a comprehensive Latrunculiidae, included in Poecilosclerida by Levi (1973)
historical analysis of this systematics and the characters emphasized and Van Soest (1984b), has also commonly been assigned to
by the various authors in their respective classifications. Hadromerida (Reid, 1968d; Bergquist, 1978; Hartman, 1982).
Subsequent development of the Poecilosclerida systematics There is now more substantial evidence to support its inclusion in
has mostly concerned the allocation of genera to various families, Poecilosclerida (e.g., Kelly-Borges & Vacelet, 1995), but as
with these appearing and disappearing between the various demonstrated in this work, it is now considered to be polyphyletic
schemes. Hajdu et ai. (1994a) provide a brief outline of these based on ontogenetic differences in microrhabd morphology
family-level changes, and also proposed a subdivision of the order (,chessman spicules') between groups of taxa within the 'latrun-
based on recognizing monophyletic taxa within the vast and culiids'. Diacarnus and related genera (family Podospongiidae),
diverse assemblage of Poecilosclerida (see summary below). bearing spinose microrhabd (spinorhabd) microscleres, have sug-
There are now many contemporary monographic publications gested affinities with the Mycalidae based on possession of
on regional poecilosclerid faunas, each differing slightly in the a sigmoid protorhabd with presumed homology to diancistra
families and genera contained within, reflecting differences derivatives. Conversely, Latrunculia and related genera have
in opinions between authors as to the importance of certain diag- discate microrhabds (acanthomicrorhabds or discorhabds) (family
nostic characters over others: e.g., Levi (1963) (South Africa); Latrunculiidae), with a straight protorhabd, and remain uncertain
Wiedenmayer (1977b) and Van Soest (1984b) (Caribbean); as to their affinities within Poecilosclerida, and consequently are
Bergquist & Fromont (1988) (New Zealand); Wiedenmayer (1989) allocated here to a new monophyletic suborder Latrunculina.
(Bass Strait, Victoria), Hooper & Levi (1993a) and Hooper & Although this proposal is a preliminary phylogenetic hypothesis, as
Battershill (1998) (New Caledonia). There are also many contem- yet with little (published) empirical support, it does have merit in
porary publications that have reviewed or revised various families recognizing distinct ontogenetic differences in microrhabd mor-
of poecilosclerids, together reflecting the ecological importance, phologies between Podospongiidae and Latrunculiidae (Samaai,
the phylogenetic significance and the huge challenge of defining unpublished data).
this order within a phylogenetic framework: e.g., Raspailiidae In this present volume we recognise 25 families, 129 genera,
(Hooper, 1991), Microcionidae (Hooper, 1996a), Rhabderemiidae 50 subgenera and several thousands of described species, allocated
(Hooper, 1990a; Van Soest & Hooper, 1993), Acamidae, among four suborders: Microcionina (four families), Myxillina
Desmacididae and Myxillidae (Desqueyroux-Faundez & Van (eleven families), Mycalina (nine families) and Latrunculina incertae
Soest, 1996, 1997; Van Soest, 1998), Latrunculiidae (Kelly-Borges & sedis (one family) (see below). Poecilosclerids have been recorded
Vacelet, 1995), Crambeidae (Maldonado & Uriz, 1996b), from all marine environments, from the shallow intertidal to the
Desmacellidae (Van Soest, 1984b; Hooper et ai., 1991; Hooper, abyssal depths (Hartman, 1982).
1984b, 1996b), Mycalidae (Hajdu, 1999). Despite this growing lit-
erature, however, it is true that poecilosclerid systematics is still far Recent reviews
from resolved. Building on Hajdu et al. (1994a) proposal to
develop a systematics for Poecilosclerida based on monophyletic Levi, 1973; Wiedenmayer, 1977b, 1989; Bergquist, 1978;
clades the following chapters constitute major revisions of the Hartman, 1982; Van Soest, 1984b; Bergquist & Fromont, 1988;
order, focusing on subordinal, family and generic and subgeneric Hajdu et ai., 1994a; Hooper & Wiedenmayer, 1994; Kelly Borges &
levels of classification. In many cases subgeneric taxa are proposed Vacelet, 1995.

KEY TO SUBORDERS

(1) Chelae microscleres are absent ............................................................................................................................................................ 2

Chelae microscleres are present .......................................................................................................................................................... 6
Porifera· Demospongiae • Poecilosclerida 405

(2) Microscleres are absent (apart from raphides (trichodragmata) in a few genera) ................................... Microcionina (Raspailiidae)
Microscleres are present, although not chelae .. ...... ... ............................. ................ ................................ .............. ....... .................. ... ... 3
(3) Microscleres include sigmoid morphologies ....................................................................................................................................... 4
Microscleres include acanthose microrhabds .... ............... ................ ............ .... ............................ ....... .............. .......... ........... .... ......... 5
(4) Microscleres may include sigmas, microxeas, commata and raphides ...................................................... Mycalina (Desmacellidae)
Microscleres are peculiar sigmoid-like (spirosigmata, thraustosigmata), toxiform (thraustoxea) and rugose
microstyles ................................................................................................................ Microcionina (Rhabderemiidae, incertae sedis)
(5) Microscleres are distinctive acanthomicrorhabds with discate modifications (acanthose discorhabds), with single perpendicular
spicules typically arranged in a compact palisade, their bases buried in the ectosomal membrane and projecting
outside of the ectosome .......... ................. .......... ..................... ....................... ... .... ........ Latrunculina (Latrunculiidae, incertae sedis)
Microscleres are distinctive spinorhabds, often in two size categories, disposed in a dense crust or scattered in a band within the
ectosome ...... ................ ...... .......... ................ ...... ................ .......... ...... ...... .... ... ... ........... ..................... ....... Mycalina (Podospongiidae)
(6) Chelae microscleres are exclusively palmate isochelae, toxas often present ...................................................................................... 7
Chelae microscleres are tridentate-derived (arcuate, anchorate forms); toxas never present ............................................... Myxillina
(7) Ectosomal megascleres monactinal with small terminal spines; choanosomal megascleres include at least 2 morphologies localized
to distinct regions within the body, often spined .............................................................................................................. Microcionina
In addition to palmate chelae (iso- and aniso-morphologies), microscleres include sigmas or sigmancistra-derivatives; megascleres
consist only of a single smooth style with faintly constricted neck and/or faintly swollen tyle ('mycalostyles') ................ Mycalina

SUBORDINAL CLASSIFICATION OF cleistochelae (Fig. 4K), tetrapocilli (Fig. 4L), coelodisc (Fig. 4G),
POECILOSCLERIDA clavidisc (Fig. 4J), etc., some of which are diagnostic for particular
taxa.
Key characters Other microscleres. Meniscoid spicules, sigmas and deriva-
tives (including chelae, sigmancistras, diancistras), are symple-
Differentiation of suborders within Poecilosclerida funda- siomorphic for the clade Haplosclerida + Poecilosclerida (Van
mentally rests on the presence/ absence and morphology of chelae Soest, 1987a). Conversely, sigma-derivatives are only found in
microscleres (palmate versus tridentate-derived forms (arcuate and Poecilosclerida and many of these morphologies are apomorphic
anchorate chelae». The classification also relies on other charac- for particular taxa (e.g., placochelae for taxa such as Guitarridae;
ters, such as the presence or absence of toxa microscleres, the pres- Fig. 4A). Unmodified sigmas are also present in some poeciloscle-
ence/ absence, geometry and spination of ectosomal megascleres, rids (e.g., Desmacellidae; Fig. 4E). Similarly, sigmancistra-
and the number and localization of choanosomal megascleres derivatives such as diancistras (Hamacantha; Fig. 41) and clavidiscs
within the skeleton (Hajdu et al., 1994a). (Merlia; Fig. 4J) are apomorphic for Mycalina (Hamacanthidae
Chelae. Chelae morphotypes have been discussed in detail
by Hajdu et al. (1994a) based on a detailed SEM examination of all
reported forms in most nominal genera. These authors suggest that
there are three fundamental morphologies, differentiated by the
degree of fusion of alae to the shaft and the torsion of the shaft itself.
Unfortunately, this distinction is not always clear-cut, sometimes
being one of 'grade', with the consequence that intermediary forms
may make allocation to palmate, anchorate or arcuate categories
difficult (e.g., Hooper, 1996a). Nevertheless, in most cases these
categories hold up and their diagnostic utility as phylogenetic mark-
ers remains highly informative. Palmate isochelae (Fig. I) are
defined as having two incomplete lateral alae extensively fused with
the shaft, and a frontal ala of varying width. Arcuate chelae (Fig. 2)
have a well-formed frontal ala and more completely formed lateral
alae which in most cases are separated from the shaft. Intermediate
forms may have their lateral alae partially separated from the shaft
(and hence they may be potentially confused between either cate-
gory), but arcuate chelae are usually recognizable as having 'teeth-
like' alae, irrespective of the degree to which the lateral alae are
attached. Anchorate chelae (Fig. 3) have at least three completely
formed alae, with the lateral ones (i.e., those adjacent to the frontal
ala) completely detached from the shaft, in addition to which there
are two other lateral alae totally fused to the shaft and resembling
ridges. Modifications to these basic plans (such as unguiferous
forms; Figs 2D, 3B) are generally recognizable as belonging to Fig. 1. Palmate-derived isochelae (abbreviations: f, frontal ala fully-
one of these three categories based on the presence or absence of formed; I, lateral ala, incompletely formed , fused with shaft by its longest
axis; ff, lateral and frontal alae nearly completely detached from each
lateral ridges (anchorate vs. palmate forms, respectively). Other other). A--C, palmate isochelae with alae of variable length and thickness.
modifications to chelae are common amongst Poecilosclerida: e.g., D, fluted alae. E, winged shaft. F, anisochela. (A, C, Clathria australiensis;
birotules (Fig. 3D), anisochelae (Fig. IF), placochelae (Fig. 4A), B, C. oxyphila; D, C. I1Ulcropora; E, C. toxipraedita; F, Mycale sp.).
406 Porifera' Demospongiae • Poecilosclerida

and Merliidae). Birotules (Fig. 3D) are many-toothed chelae with

alae encircling the shaft, and thought to have originated from
anchorate chelae (Van Soest, 1987a). These are characteristic of the
myxillinid family Iotrochotidae. Toxas (Fig. 4B) are also symple-
siomorphic for Poecilosclerida + Haplosclerida. Their absence
from Myxillina is considered to be a derived character. Forceps,
recurved pincer-like rods (Fig. 4C-D), occur in both Mycalina and
Myxillina and their possession has been interpreted as independ-
ently derived in both, assuming they are indeed homologous
(Hajdu et at., 1994a). Onychaetes (Fig. 4F) are synapomorphic for
Tedaniidae, although their origin, and the affinities of this family,
are still equivocal (Hajdu et at., 1994a). Likewise, raphides (often
in trichodragmata; Fig. 4M) are homoplastic with a broad occur-
rence in most poecilosclerid assemblages and also found in several
taxa outside the Poecilosclerida (e.g., Halichondrida).
Two families have acanthomicrorhabds ('chessman spicules')
occurring in a surface crust. Previously these taxa were amalga-
mated in a single family (Latrunculiidae), but Kelly & Samaai and
Samaai & Kelly (this volume) demonstrate fundamental ontoge-
netic differences between two morphotypes of 'chessman'
spicules, indicating their homoplastic ontogenetic development
and negating any homology as previously thought. Podospongiidae
Fig. 2. Arcuate-derived isochelae (abbreviations: c. curved shaft; f, frontal have spinorhabds (Fig. 4N), bearing two whorls of spines arranged
alae well-formed; pf, partially fused frontal and lateral alae; I, lateral alae serially around the spicule, between a proximal and distal whorl of
partially separated from shaft; a, alae teeth like). A-C, arcuate isochelae
with alae of varying length. D, unguiferous arcuate isochelae.E, highly
modified displaster-like arcuate-derived chela. F, strongly deformed bluntly
spined arcuate-derived chela. (A, 'Clathria' grisea; B, Ectyodoryx sp.;
C, Hamigera dendyi; D, 'Clathria' scabida; E, Acanthancora clavi/obata;
F, Acanthancora cyanocrypta.)

Fig. 3. Anchorate-derived chelae (abbreviations: 3, three completely Fig. 4. Examples of other Poecilosclerida microscleres. A, spined chela,
formed alae, with the lateral ones completely detached from the shaft; including view of ala (Guitarra). B, wing-shaped toxa (Acarnus). C, forceps
2, two rudimentary lateral alae completely fused to the shaft resembling (Forcepia). D, small incurled forceps (Forcepia (Leptolabis)). E, sigma
ridges; u, unguiferous teeth-like alae). A, fully formed lateral and frontal (Mycale). F, onychaete (Tedania). G, coelodisc (Coelodischela). H, sigman-
alae. B, unguiferous arcuate isochelae with teeth-like alae (and enlarged cistra (Euchelipluma). I, diancistra (Hamacantha). J, clavidisc (Merlia).
view). c, reduced frontal teeth-like alae, but with persistent ridge on shaft. K, cleistochela (Clathria). L, tetrapocilla (Tetrapocillon). M, trichodrag-
D, birotule. E, polydentate anchorate spatulate chela. F, spherancora. (A-B, mata (bundle of raphides; Raspai/ia). N, spinorhabd (Diacarnus). 0, dis-
Monachora unguiculata; C, Crella incrustans; D, Iotrochota ace rata; E, corhabd (Latrunculia). P, thraustosigma (Rhabderemia). 0 thraustoxea
Ectyonopsis hartmani; F, Melonancora elliptica). (Rhabderemia). R, microstyle (Rhabderemia).
Porifera· Demospongiae • Poecilosclerida 407

spines, one of which may be longer than the other. This family has interstitially, in the ectosome, and/or echinating skeletal tracts.
suspected Mycalina affinities based on the fact that the protorhabd Microcionina can have up to five categories (with the exception of
of spinorhabds is sigmoid, thus with presumed homology to sig- Rhabderemiidae), whereas at the opposite end of the continuum,
mancistra derivatives, and with corroboratory support for this Mycalina and Latrunculina have only one category.
assertion from other skeletal and non-morphological characters. Skeletal architecture. The gross organic and inorganic
Latrunculiidae, of uncertain affinity within Poecilosclerida, have skeletal architecture, the structural differentiation of the inorganic
discate acanthorhabds (discorhabds) (Fig. 40), bearing apical and skeleton, and the distribution of mineral components in that struc-
basal whorls of spines and several crenulate discs in between these, ture are primary diagnostic features for many taxa (Levi, 1960b,
with the protorhabd lacking any sigmoid modification. Further, 1973; Bergquist, 1978; Hartman, 1982). However, when used
within Latrunculiidae there are several different morphologies of alone skeletal architecture is not necessarily a reliable indicator of
acanthodiscorhabds, each apomorphic for various genera: anisodis- phylogenetic affinities. Choanosomal skeletal structures such as
corhabds or 'typical chessman spicules' (in Latrunculia), isoconi- hymedesmioid, plumose, plumo-reticulate, regularly reticulate,
codiscorhabds or 'sceptres' (in Sceptrella), and isochiadiscorhabds isodictyal reticulate, renierioid reticulate, axially compressed and
or truncate discorhabds (in Tsitsikamma gen. nov.; Samaai & halichondroid arrangements may be characteristic of various gen-
Kelly) (refer to respective chapters in this volume). era, or sometimes of certain families, but higher taxa can rarely be
Similarly, the affinities of Rhabderemiidae based on their defined by these features alone, with the presumption that they are
microsclere morphologies remains enigmatic, with oxea- or toxa- more susceptible to adaptive pressures than are characters such as
like microscleres (thraustoxeas; Fig. 4Q), sigma-like spicules microsclere morphology (Ridley & Dendy, 1887; Dendy, 1905; see
(spirosigmata, thraustosigmata; Fig. 4P) and microstyles (Fig. 4R)
possibly analogous to toxas, sigmas and rhabds of other poe-
cilosclerids (refer to chapter Rhabderemiidae, this volume).
Rhabderemiidae remains incertae sedis within Poecilosclerida.
They may have affinities with Raspailiidae, based on characters
other than microscleres (skeletal structure similar to Auiospongus,
for example), although the possession of true toxas in some species
(Van Soest & Hooper, 1993) possibly supports a closer relationship
with Acamidae. Alternatively, Rhabderemiidae may be a com-
pletely independent line within Poecilosclerida, but there is so far
no molecular support to corroborate any relationship within
Poecilosclerida in general, or affinities with any family in particular.
Ectosomal megascleres. Ectosomal spicules are fundamen-
tal to the subclass classification, with Microcionina having predom-
inantly monactinal ectosomal megascleres. (1) Raspailiidae have
special bundles of styles or oxeas surrounding larger protruding
spicules; Microcionidae have basally spined ectosomal styles tan-
gential or paratangential; Acamidae have tangentially arranged
tylotes or anisostyles with spined bases tangential or paratangential;
Rhabderemiidae lack special ectosomal spicules, as do certain
genera of the other families (refer to chapters by Hooper in this
volume). (2) Myxillina have predominantly diactinal ectosomal
megascleres without basal spination, or with rare, irregular or coarse
spines (ectosomal spicules absent in several families) (refer to chap-
ters by Van Soest in this volume). (3-4) Mycalina and Latrunculina
lack ectosomal spicules altogether (presumed secondary losses; or
alternatively, independent evolutionary acquisitions on the part of
the other suborders) (refer to chapters by Hadju et ai., Van Soest
et aI., Kelly & Samaai and Samaai & Kelly in this volume).
Acanthostyles. Acanthostyles are widespread but not univer-
sal amongst Poecilosclerida, with their absence from various taxa
(e.g., Mycalina, Latrunculina) interpreted as either derived or prim- Fig. 5. Examples of poecilosc1erid acanthostyle geometries. A-D, G, I-L,
itive features (Hooper, 1991, 1996a; Hajdu et al., 1994a, respec- R, echinating acanthostyles. E, F. H, M, P, 'plocamiform' acan-
tively). These spicules echinate spongin fibres (e.g., Raspailiidae, thostrongyles. N, echinating style. 0, acanthorhabd. Q, cladotylote.
Microcionidae), core fibres (e.g., Myxillidae) or both (e.g., A, Chaetodoryx richardi (Coelosphaeridae). B, Hymedesmia zetlandica
(Hymedesmiidae). C, Phorbas perarmatus (Hymedesmiidae). D, Ploc-
Crellidae). Their diverse geometries (including patterns of spina-
amionida ambigua (Hymedesmiidae). E, Ectyonopsis ramosus
tion) (Figs 5--6) may be highly diagnostic for particular lower-level (Myxillidae). F, Ectyonopsis hartmani (Myxillidae). G, Crellomyxilla inter-
taxa (e.g., spatulate versus sharply pointed spines in the genus media (Myxillidae). H, Plocamiancora igzo (Myxillidae). I, Clathria
Echinodictyum (Raspailiidae», but these phylogenetic patterns are (Clathria) crassa (Microcionidae). J, Clathria (Thalysias) rubra
unclear at higher taxonomic levels (Refer to Hooper, 1991, 1996a (Microcionidae). K, Clathria (Thalysias) darwinensis (Microcionidae).
L, Clathria (Thalysias) phorbasiformis (Microcionidae). M, Antho (Acamia)
for additional illustrations). ridleyi (Microcionidae). N, Holopsamma arborea (Microcionidae). 0,
Diversity of megascleres. By definition PoeciloscIerida has Acanthorhabdusfragilis (Acarnidae). P, Wigginsia wigginsi (Acarnidae). Q,
at least two categories of megascleres, coring fibres and occurring Acamus innominatus (Acarnidae). R, Dolichacantha macrodon (Acamidae).
408 Porifera' Demospongiae • Poecilosclerida

although there are still no substantial molecular data to corroborate

this hypothesis.
Incorporation of detritus into skeleton. This feature is
widespread over a variety of otherwise unrelated taxa, including
various Poecilosclerida (Microcionina, Myxillina), Haplosclerida,
Dictyoceratida, Dendroceratida, Verongida etc., and is not attrib-
uted much significance above the species-level, or occasionally
genus-level of classification.

Microcionina Hajdu, Van Soest & Hooper, 1994a

Four families, with terminally spined ectosomal monactinal

megascleres (occasionally modified to quasidiactinal forms,
always asymmetrical); choanosomal megascleres diverse, consist-
ing of at least two categories localized to distinct regions within the
skeleton, or sometimes up to five categories including spicules
echinating fibres in many taxa; isochelae exclusively of palmate
origin, with diverse forms of toxas but lacking sigmas. Families:
Acarnidae, Microcionidae, Raspailiidae, Rhabderemiidae.

Myxillina Hajdu, Van Soest & Hooper, 1994a

Eleven families, with ectosomal megascleres basically diacti-

nal, although aniso-terminations are widespread, and terminal
spination of ectosomal megascleres rare or if present usually
coarse or irregular, with tridentate-derived chelae (iso- or aniso-,
arcuate, anchorate or birotulate), but lacking toxas. Families:
Chondropsidae, Coelosphaeridae, Crambeidae, Crellidae,
Dendoricellidae, Desmacididae, Hymedesmiidae, Iotrochotidae,
Fig. 6. Examples of poecilosclerid acanthostyle geometries (cont.). A-B,
rhabdostyles. C-D, ciavulate acanthostyles. E, G, bulbous acanthostyles.
Myxillidae, Phellodermidae fam. nov., Tedaniidae.
F, spatulate acanthostyle. H, 'plocamiform' acanthostrongyle. I-J, sagittal
triacts/tetracts (acanthoplagiotriaenes). A, Rhabderemia sorokinae (Rhab- Mycalina Hajdu, Van Soest & Hooper, 1994a
deremiidae). B, Rhabderemia pusilla (Rhabderemiidae). C, Ectyoplasia
tabula (Raspailiidae). D, Endectyon fruiticosa aruensis (Raspailiidae).
Nine families, with megascleres almost always a single
E, Raspailia (Hymeraphiopsis) irregularis (Raspailiidae). F, Echinodictyum
mesenterinum (Raspailiidae). G, Lithoplocamia lithistoides (Raspailiidae). smooth category, never echinating the skeletal tracts, consisting of
H, Plocamione pachysclera (Raspailiidae). I, Cyamon vickersi (Raspailiidae). (sub-)(tylo-)styles with faintly constricted neck and/or faintly
J, Trikentrionflabelliforme (Raspailiidae). swollen tyle ('mycalostyles'), with microscleres sigmancistra
derivatives and chelae which are mostly palmate; Podospongiidae,
with spinorhabds ('chessman spicules'), is also allocated to
discussion in Hajdu et al., 1994a, and Hooper, 1996a) (refer to illus- this suborder based on skeletal and other characters (see Kelly &
trations contained within the various poecilosclerid families in this Samaai, below). Families: Cladorhizidae, Desmacellidae,
volume). A contrary approach was taken by Berquist & Fromont Esperiopsidae, Guitarridae, Hamacanthidae, Isodictyidae,
(1988) who recognised the validity of different skeletal structures as Merliidae, Mycalidae, Podospongiidae.
primary diagnostic characters, in addition to, and in some cases
equal with other features such as microsclere morphology, resulting Latrunculina Kelly & Samaai, subord. nov.
in a proliferation of genera that confused their phylogenetic rela-
tionships. Chelae microsclere geometry, as a primary apomorphy, Monotypic (Latrunculiidae), with wispy wide-meshed reticu-
provides a partial resolution of these discrepancies. Choanosomal late to reinforced reticulate choanosomal skeleton composed of
skeletal architectures (e.g., hymedesmioid, microcionid, renieroid, anisostyles or strongyles, ectosomal skeleton compact tangential,
isodictyal, axially compressed etc.) are suspected to be homoplastic microscleres are disc ate acanthorhabds typically arranged verti-
developments largely incongruent with phylogenies based on cally in a palisade on the outside of ectosome; without chelae,
microsclere geometries (palmate - arcuate - anchorate origins), sigmas or toxa microscleres.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Suborder Microcionina Hajdu, Van Soest & Hooper, 1994

John N.A. Hooper

Queensland Museum, P.O. Box 3300, South Brisbane, Qld, 4101, Australia. (JohnH@qm.qld.gov.au)

Microcionina Hajdu et al. (Demospongiae, Poecilosclerida) contains 42 valid genera, 19 subgenera and over 1,000 described species of
sponges worldwide, with many more species awaiting description. The suborder is characterised in having terminally microspined ecto-
somal megascleres and up to five categories of structural megascleres, most frequently monactinal or derivatives from these, and
microscleres that include palmate chelae and diverse toxas, but never sigmas. Four families are included: (1) Microcionidae (viviparous)
with principal (subtylo)styles coring spongin fibres, echinating (acantho)styles perpendicular to fibres, and one or more category of acces-
sory (subtylo)styles forming a peripheral or extra-fibre skeleton, with microscleres palmate isochelae and diverse forms of toxas.
(2) Acarnidae (viviparous) with an ectosomal skeleton of basally spined tylotes arranged tangentially, with monactinal choanosomal
megascleres (styles) coring fibres, with or without echinating spicules, and microscleres palmate isochelae and toxas sometimes with
others such as bipocillae, modified anisochelae, microrhabds and raphides. (3) Raspailiidae (oviparous) often with some degree of axial
compression of the skeleton, typically with a hispid surface produced by brushes of small ectosomal megascleres surrounding protruding
choanosomal megascleres, diverse geometric modifications to echinating megascleres, and lacking microscleres other than raphides
(trichodragmata) in a few genera. (4) Rhabderemiidae (presumed oviparous) with rhabdostyles bearing a basal spiral twist, and peculiar
microscleres (perhaps superficially) resembling toxas (thraustoxeas), sigmas (spirosigmata, thraustosigmata) and rugose microstylote
spicules, and lacking chelae.
Keywords: Porifera; Demospongiae; Poecilosclerida; Microcionina; Microcionidae; Acarnidae; Rhabdermiidae; Raspailiidae.

DEFINITION, DIAGNOSIS, SCOPE axially compressed, to nearly halichondroid architectures, and pre-
dominantly with three skeletal regions defined by the presence of
Synonymy different forms of structural styles: (1) an axial (or choanosomal)
skeleton with spongin fibres enveloping principal styles echinated
Microcionina Hajdu, Van Soest & Hooper, 1994a. by acanthose or smooth styles; (2) an extra-axial (or subectosomal)
skeleton with tracts of auxiliary styles ascending to the surface;
Defmition (3) and an ectosomal skeleton with smaller auxiliary styles forming
a surface crust tangential, paratangential or perpendicular to the
Poecilosclerida with terminally spined ectosomal monactinal surface. One or more skeletal regions may be lost or modified
megascleres (occasionally modified to quasidiactinal forms); (Rhabderemiidae lacks any differentiation between choanosomal
choanosomal megascleres diverse, consisting of at least two cate- and ectosomal regions based on megasclere distribution, but has an
gories localized to distinct regions within the skeleton, or some- ectosomal layer of microscleres, although not forming a distinct
times up to five categories including spicules echinating fibres in crust). Megascleres are predominantly smooth (subtylo-)styles but
many taxa; isochelae of palmate origin, with diverse forms of toxas may be replaced by and/or supplemented with quasidiactinal or true
but lacking sigmas. diactinal forms, lost partially or completely and replaced with detri-
tus, or consisting only of a single category of rhabdostyles bearing
Diagnosis an extra spiral twist at the rounded extremity (Rhabderemiidae).
Microscleres include: palmate isochelae and diverse forms of
Sponges with diverse growth forms ranging from encrusting toxas (Microcionidae); supplemented with bipocillae, modified
to massive, arborescent to whip-like species, with ectosomal anisochelae, microrhabds and raphides (Acarnidae); absent com-
megascleres usually terminally microspined and up to five cate- pletely except for raphides (in trichodragmata) (Raspailiidae); or
gories of structural choanosomal megascleres, most frequently consist of peculiar toxifonn (thraustoxeas), sigmoid-like (spirosig-
monactinal or derivatives, and microscleres that include palmate mata, thraustosigmata) and rugose microstyles (Rhabderemiidae).
chelae and diverse toxas, never sigmas. Ectosomal spicules range
from apically spined tylotes (or anisostyles) forming a tangential Scope
ectosomal skeleton (Acarnidae), to terminally spined styles, rarely
modified to quasidiactinal or diactinal forms, forming a tangential Four families are included: Acarnidae, Microcionidae,
or paratangential skeleton (Microcionidae), to an erect skeleton Raspailiidae and Rhabderemiidae (the latter enigmatic and still
consisting of bouquets of smaller ectosomal styles, oxeas or anisox- incertae sedis). Together these families contain some of the most
eas surrounding protruding larger styles or oxeas (Raspailiidae), or diverse shallow-water faunas, and a few deeper-water representa-
absent altogether (Rhabderemiidae). Choanosomal skeletal struc- tives, with 42 genera and 19 subgenera recognised as valid, and
tures range from hymedesmioid, microcionid, plumo-reticulate, over 1,000 described species worldwide and many more waiting to
regularly reticulate, occasionally isodictyal or renieroid reticulate, be described.

409
410 Porifera· Demospongiae • Poecilosclerida • Microcionina

Remarks resembling the Microcionidae more than any other poecilosclerid

family, and certainly more than any other demosponge family.
Microcionina are not homogeneous in tenns of all taxa having There is also some early biochemical support for this relationship
the primary apomorphies (tenninally spined ectosomal megascle- (e.g., carotenoid proteins, general protein electrophoresis, free
res, multiple categories of structural megascleres, only palmate amino acid patterns; Lee & Gilchrist, 1985; Hooper, 1990b;
chelae, not arcuate or anchorate chelae or sigmas), and in this Hooper et al., 1992), but this evidence is equivocal because it is
regard the suborder may be artificial. Microcionidae and Acarnidae now outdated and has not yet been corroborated by more substan-
are the most closely related families in sharing these characters, tive genetic data. Rhabderemiidae has even less support for its
and are both viviparous (where known). This close relationship inclusion in Microcionina (or indeed within Poecilosclerida). Its
has some empirical support from biochemical analyses (e.g., reproductive strategy is unknown, presumed by some authors (e.g.,
carotenoid proteins, C28 fatty acids; Lee & Gilchrist, 1985; Levi, 1973) to be oviparous on the basis that incubated larvae have
Lawson et al., 1984). On the one hand Raspailiidae appears to be never been recorded, and both its megascleres and microscleres are
very different. It is oviparous (where known), lacks the poe- highly unusual for poecilosclerids. Its inclusion in both the order
cilosclerid chelae apomorphy (with only a few taxa having and suborder is based on some similarities in skeletal structure and
raphides), and has a unique ectosomal apomorphy consisting of spiculation to the Raspailiidae, particularly Hemectyonilla Burton
bouquets of oxeas or styles surrounding longer choanosomal oxeas and Aulospongus Nonnan (Hooper, 1991; Van Soest & Hooper,
or styles at the surface. On the other hand it shares some strong 1993; Hooper et al., 1999), but this is a working hypothesis and
skeletal features and megasclere geometries with Microcionidae requires corroboration from other data sets.
(reticulate or plumo-reticulate skeletons; both families having rep-
resentatives with axial compression of the skeleton; acanthostyles Recent revision
echinating fibres; large numbers of megasclere categories (up to
4-5) mostly localised to particular regions within the skeleton), Hajdu et al., 1994a.

KEY TO FAMILIES

(I) Ectosomal skeleton present ................................................................................................................................................................. 3

Ectosomal skeleton absent ................................................................................................................................................................... 2
(2) No special ectosomal megascleres although choanosomal rhabdostyles may protrude through the surface singly or in sparse bundles;
microscleres include peculiar thraustoxea, spirosigmata, thraustosigmata and rugose microstylotes ........................ Rhaderemiidae
Ectosomal spicules lost, with only choanosomal spicules (oxeas and/or styles) protruding through surface; microscleres absent apart
from occasional raphides ..... RaspaiJiidae (all of Cyamon, Amphinomia, most of Echinodictyum, some of Aulospongus, Raspailia
(Clathriodendron), Eurypon, Ceratopsion)
(3) Ectosomal skeleton fonned by tangentially arranged tylotes with spined bases; microscleres may include palmate isochelae and toxas,
sometimes with others (bipocillae, modified anisochelae, microrhabds and raphides) ....................................................... Acarnidae
Ectosomal skeleton composed of accessory (subtylo )styles, often with microspined bases, fonning a paratangential skeleton on or just
below the surface; microscleres include only palmate isochelae and toxas .................................................................. Microcionidae
Ectosomal skeleton with brushes of small completely smooth ectosomal styles or oxeas (or modified fonns of these) surrounding
long protruding single choanosomal styles or oxeas; microscleres absent except for raphides in some
taxa ......................................................................................................................................................... Raspailiidae (typical genera)

FAMILIES OF MICROCIONINA Acarnidae Dendy, 1922b

Microcionidae Carter, 1875c Acarnidae (including Iophonidae Burton, 1929a and Cornulidae
Levi & Levi, 1983b), contains thirteen valid genera and probably
Microcionidae (including Clathriidae Lendenfeld, 1884, several hundreds of species worldwide, although several genera are
Ophlitaspongiidae de Laubenfels, 1936a), contains nine genera, restricted to colder temperate waters or to very narrow geographic
twelve subgenera and approximately 460 described species world- distributions (regional endemics). Ectosomal skeleton typically with
wide (with many more species awaiting description), predominantly tangentially arranged tylotes with spined bases, and choanosomal
in shallow waters with a few recorded from deeper seas. Species have skeleton is reticulate in massive growth fonns or plumose in encrust-
monactinal principal structural (subtylo)styles coring spongin fibres, ing growth fonns, with or without echinating spiCUles. Megascleres
echinating (auxiliary) (acantho )styles perpendicular to fibres, and one are styles and microscleres are palmate isochelae and toxas, some-
or more categories of accessory (subtylo) styles fonning a peripheral times lost or sometimes with other microscleres such as bipocillae,
or extra-fibre skeleton. Only palmate isochelae microscleres are pres- modified anisochelae, microrhabds and raphides.
ent and these are only exceptionally modified to arcuate-like or
anchorate-like fonns as a result of torsion of the shaft and detachment RaspaiJiidae Hentschel, 1923
of alae. Toxa morphologies are diverse, including microxea-like toxas
in a few species. Skeletal structure ranges from hymedesmioid and Raspailiidae (including Euryponidae Topsent, 1928c) con-
microcionid in encrusting taxa, to plumo-reticulate and occasionally tains nineteen valid genera (one incertae sedis), seven subgenera,
axially compressed in some species, but typically reticulate in most and approximately 250 named species worldwide, mainly from
taxa, occasionally replaced completely by detritus. shallow waters and a few from abyssal depths. Species have a
Porifera' Demospongiae • Poecilosclerida • Microcionina 411

typically hispid surface, with genera differentiated mainly on the Rhabderemiidae Topsent, 1928c
basis of three morphological characters: (a) skeletal architecture
ranging from axial compression to reticulate, plumo-reticulate or Monogeneric, with 26 described species found predominantly
plumose skeletons; (b) the presence or absence of a specialized ecto- in shallow waters of all tropical and warm temperate seas, and one
somal skeleton (apomorphic for the family, whereby brushes of abyssal species. The family has unique apomorphies consisting of
small ectosomal megascleres surround long protruding single rhabdostyles bearing a basal spiral twist, and peculiar microscleres
choanosomal megascleres); and (c) geometric modifications to echi- resembling toxas (thraustoxeas), sigmas (spirosigmata, thraustosig-
nating megascleres, with about 15 distinct morphologies recognised. mata) and rugose rnicrostylote spicules.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Acarnidae Dendy, 1922

John N.A. Hooper

Queensland Museum, P.O. Box 3300, South Brisbane, Qld, 4101, Australia. (JohnH@qm.qld.gov.au)

Acarnidae Dendy (Demospongiae, Poecilosclerida, Microcionina) contains 37 nominal genera of which 13 are considered here to be
valid, and probably several hundreds of species. Acarnids are distributed worldwide, although several genera are predominant in colder
temperate waters, and several have very restricted (apparently endemic) geographic ranges. Major morphological characters diagnostic
for the family include massive, burrowing, excavating or encrusting growth forms with fistules, as well as non-fistulose genera with digi-
tate, flabellate, massive or encrusting growth forms; ectosomal skeleton with tangentially arranged tylotes with spined bases; choanoso-
mal skeleton reticulate (in massive) or plumose (in encrusting growth forms), with or without echinating spicules; megascleres include
styles; and microscleres are palmate isochelae and toxas, sometimes lost or sometimes with other microscleres such as bipocillae, modi-
fied anisochelae, microrhabds and raphides.
Keywords: Porifera; Demospongiae; Poecilosclerida; Microcionina; Acarnidae; Acanthorhabdus; Acarnus; Acheliderma; Cornulella;
Cornulurn; Darniria; Dolichacantha; Iophon; MegacieUa; Paracornulurn; Tedaniophorbas; Wigginsia; Zyzzya.
--------~---~~.--~ .---.~-~-~-----~ ---~~~

DEFINITION, DIAGNOSIS, SCOPE traditionally included in the family Coelosphaeridae Topsent. Levi &
Levi (1983b) subsequently split Coelosphaeridae into two groups
Synonymy for genera with palmate isochelae (Comulidae Levi & Levi) or
arcuate isochelae (Coelosphaeridae). Several non-fistular genera
Acarneae Dendy, 1922b. Iophoneae Burton, 1929a. Acarnidae had similar skeletal structure and spiculation, such as Acarnus
Boury-Esnault, 1973 (Not [Acarniidae] de Laubenfels, 1936a). and Iophon, possessing spined ectosomal tylotes and palmate
Comulidae Levi & Levi, 1983b. isochelae, but these were traditionally included in Myxillidae
(e.g., Topsent, 1928c; de Laubenfels, 1936a), which predominantly
Definition contain taxa with 'tridentate-derived' chelae (arcuate and
anchorate) and sigmas, but lack toxas. Hajdu et al. (1994a) proposed
Microcionina with apically spined tylotes forming a tangential to merge these two groups of genera by extending the scope
ectosomal skeleton. of Comulidae to include both fistulose and non-fistulose genera
with similar skeletal arrangements (ectosomal tylotes with
Diagnosis spined bases) and spiculation (palmate isochelae, toxas, lack of
sigmas).
Encrusting, massive, flabelJate or digitate growth forms, some- The correct name for the family is still contentious. Hajdu
times burrowing, many genera producing fistules. Ectosomal skele- et al. (1994a) noted that the inclusion of Iophon in this group
ton composed of tylotes, strongyles or modified tylote spicules necessitated recognition of Burton's (1929a) higher taxon
bearing microspined bases forming tangential and/or paratangen- Section Iophoneae, which they elevated to full family status.
tial tracts, often irregular or halichondroid in arrangement. Iophoneae was initially created only for Iophon and Iophonopsis
Choanosomal megascleres are styles or modified styles (anisoxeas) within the family Myxillidae. However, the same argument applies
forming reticulate structures in massive forms (isodictyal, isotropic, with the inclusion of Acarnus in this group. Dendy (1922b) had
anisotropic or more irregular skeletons) or plumose or hymedesmioid earlier created Section Acarneae for this genus, within the old fam-
skeletons in encrusting growth forms. Echinating spicules present ily Desmacidonidae, subfamily Ectyoninae, which therefore theo-
or absent, consisting of acanthostyles and/or unique cladotylotes. retically takes priority over Iophoneae. It could be argued that
Microscleres include palmate isochelae and toxas of various mor- Hajdu et al. (1994a), followed by Van Soest et al. (1994a) and
phologies (both sometimes lost), with some genera also having Desqueyroux-Faundez & Van Soest (1995) had thus established
other microscleres such as bipocillae, modified anisochelae, Iophonidae 'in prevailing usage' (Article 35.5 ICZN; Anon., 1999),
microrhabd-like spicules (modified microxeas or microstrongyles) in which case 'the older name is not to displace the younger name'.
and diamond-shaped microxeas (modified toxas). However, these authors overlooked the prior usage of Acarnidae as
a family level taxon by Topsent (1928c) and subsequently adopted
Scope by several authors (e.g., de Laubenfels, 1932; see discussion in
Hooper, 1987), negating the applicability of Article 35.5 of the
Acarnidae contains 37 nominal genera of which only 13 are ICZN.
presently considered to be valid. Under Hajdu et al.'s (1994a) concept of this group 36 nominal
genera were potentially allocated here. There are several recent
History and biology reviews outlining the scope and features of this group, including
comprehensive revisions of particular genera (Van Soest et al.,
Fistular poecilosclerid genera like Acheliderma, Cornulurn, 1991; Van Soest, Zea & Kielman, 1994; Desqueyroux-Faundez &
Darniria, Zyzzya with tylote ectosomal megascleres have been Van Soest, 1995).

412
Porifera' Demospongiae • PoeciioscIerida • Microcionina • Acarnidae 413

Differences with similar families (Hooper, 1996a». Some of the iophonid genera are clearly similar
to the Microcionidae (e.g., Acamus, Megaciella) in characters such
Acarnidae differs significantly from other Microcionina in as the geometry of megaScleres and microscleres, whereas others
possessing apically-spined tylotes or modified strongylote or (e.g., Acanthorhabdus, Dolichacantha) have been modified so
oxeote ectosomal spicules that typically form a tangential ectoso- extensively that their similarities are barely perceptible.
mal skeleton. By comparison the other families (Microcionidae,
Raspailiidae, Rhabderemiidae) have monactinal ectosomal Previous reviews
spicules which do not generally form a tangential skeleton
(although some species of Clathria are known to have modified Berquist & Fromont (1988), Van Soest, Zea & Kielman, 1994;
anisostyles with spines on both the base and apex of the spicule Desqueyroux-Faundez & Van Soest (1995).

KEY TO GENERA

(1) Ectosomal spicules are tylotes, mostly smooth, with apical spines .................................................................................................... 3
Ectosomal spicules modified .. .......... ........... ................. .......... ....... .... ........... .......... ........... ........... .......... ...... .......... ........... ........... ....... 2
(2) Ectosomal tylotes modified to acanthoxeas with apical spines ........................................................... Acanthorhahdus (non-fistular)
Ectosomal tylotes entirely smooth (secondary loss of spines) ............................................................. Tedaniophorbas (non-fistular)
(3) Choanosomal skeletal structure well-formed ...................................................................................................................................... 4
Choanosomal skeletal structure vestigial, consisting of single tylotes and scattered accessory spicules
(acanthoxeas) ........................................................................................................................................................ Cornulella (fistular)
(4) Choanosomal and ectosomal megascleres clearly differentiated in geometry and distribution .......................................................... 6
Ectosomal tylotes form both ectosomal and choanosomal skeletal structures .................................................................................... 5
(5) Apically-spined tylotes form both tangential ectosomal and plumo-reticulate choanosomal skeletons
(with or without a second category of choanosomal styles) .................................................................................. Cornulum (fistular)
Apically-spined tylotes form both tangential ectosomal tracts and plumose ascending choanosomal tracts
(with acanthostyles echinating hymedesmioid basal skeleton) ...................................................................... Paracornulum (fistular)
Apically-spined tylotes are the only megascleres ..................................................................................................... Damiria (fistular)
(6) Choanosomal megascleres consist only of styles, without accessory spicules ................................................................................... 8
More than one category of choanosomal megascleres, one greatly modified ..................................................................................... 7
(7) Accessory spicules consist of verticillately-spined strongyles (or strongyloxeas) ...................................................... Zyzzya (fistular)
Accessory spicules consist of one or more classes of cladotylotes ................................................................... Acarnus (non-fistular)
Accessory spicules consist of peculiar acanthostrongyles with numerous spined 'cladi' ...................... Dolichacantha (non-fistular)
Accessory spicules consist of 'plocarniform' acanthostrongyles echinating ascending tracts of styles ......... Wigginsia (non-fistular)
(8) With a regular, uni- or pauci-spicular isodictyal choanosomal reticulation of smooth or spined styles,
with or without echinating acanthostyles ............................................................................................................ Iophon (non-fistular)
With confused choanosomal skeleton divided into primary ascending multi spicular tracts and secondary interconnecting uni- or
bispicular tracts, both cored by basally spined styles, with or without echinating acanthostyles ................. Megaciella (non-fistular)
Choanosomal skeleton appears predominantly plumose due to the dominance of subectosomal tracts of ectosomal tylotes, whereas
the basal choanosomal skeleton is irregularly renieroid-reticulate composed of basally spined styles forming triangular or square
meshes; microscleres include elongated diamond-shaped microxeas ............................................................... Acheliderma (fistular)

ACANTHORHABDUS BURTON, 1929 (acanthorhabds) with heavy spines and microspined tylote apices;
choanosomal skeleton with smooth modified styles (anisoxeas)
Synonymy often provided with a terminal mucron, forming massive multispic-
ular plumose and reticulate columns, with acanthoxeas dispersed
Acanthorhabdus Burton, 1929a: 432. between (but not echinating) fibres; microscleres palmate
anisochelae with spurs.
Type species
Description of type species
Acanthorhabdus fragilis Burton, 1929a (by monotypy).
Acanthorhabdus fragilis Burton, 1929a (Fig. 1).
Definition Synonymy. Acanthorhabdus fragilis Burton, 1929a: 432,
text-fig. 5, pI. 4, fig. 2.
Non-fistular Acarnidae with apically-spined ectosomal acan- Material examined. Holotype: BMNH 1926.10.26.189 -
thorhabds, modified choanosomal mucronate anisoxeas and lack- Antarctica.
ing true echinating spicules. Description. Subspherical, massive growth form; surface
minutely conulose; texture friable; oscules raised on small papillae
Diagnosis or plush with surface; ectosomal skeleton an irregular surface crust
of acanthorhabds lying mostly tangential to surface in an irregular
Massive non-fistulose growth form; ectosomal skeleton nearly halichondroid criss-cross, but also protruding paratangen-
composed of a loose halichondroid tangential layer of acanthoxeas tially from surface; abundant tracts of anisochelae also scattered
414 Porifera· Demospongiae • Poecilosclerida • Microcionina • Acarnidae

I
A B c D
G

Fig. 1. Acanthorhabdus fragilis Burton (holotype). A, acanthoxea (acanthorhabd) (scale 300 /-Lm). B-C, choanosomal anisoxeas (scales 300/-Lm).
D, spurred palmate anisochelae (scale 25/-Lm). E, choanosomal skeleton (scale I mm). F, ectosomal skeleton (scale I mm). G, idealised reconstruction of
skeleton. H, holotype (approximately 25 cm) (H, from Burton, 1929a, pI. 4, fig. 2).

over surface; choanosomal skeleton composed of massive plumose Poecilosclerida is without doubt given its possession of chelae
and plumoreticulate columns of smooth anisoxeas with up to 15 microscleres.
spicules per tract, interdispersed with acanthorhabds in nearly hali-
chondroid arrangement and also sometimes interconnecting major Distribution
anisoxeote spicule tracts; megascleres include fusiform, heavily
spined ectosomal acanthoxeas (acanthorhabds) with spines curved Monotypic, endemic to Antarctica.
towards the middle of spicules and microspined apices (320-365 X
20-35 f.Lm), smooth choanosomal anisoxeotes often with distinct
mucron on the basal end and occasionally a single spine on the ACARNUS GRAY, 1867
point (450-590 X 15-42 f.Lm); microscleres spurred palmate
anisochelae (22-28 f.Lm). Synonymy
Remarks. Burton (1929a) initially included this monotypic
genus with the mycalid genera (his taxon Mycaleae) based on the Acamus Gray, 1867a. ? Fonteia Gray, 1867a. ? Trefortia
possession of anisochelae whereas the presence of acanthorhabds Deszo, 1880. Microtylotella Dendy, 1896. Acanthacamus Levi,
with spined tylote apices is not found in the mycalids but is more 1952. Taxonomic decision for synonymy: Hooper (1987), Van
consistent for the Acamidae (Van Soest, Zea & Kielman, 1994; Soest et al. (1991).
Desqueyroux-Faundez & Van Soest, 1995). This placement is
further supported by the shared possession of spurred anisochelae Type species
in Iophon. However, this genus is very atypical of Acamidae in
the geometry of ectosomal acanthorhabds, which resemble the Acamus innominatus Gray, 1867a (by monotypy).
acanthoxeas of Desmoxyidae more so than poecilosclerid
ectosomal spicules, and in the halichondroid reticulate nature of Definition
these spicules dispersed within the ectosomal and choanosomal
skeletons. Nevertheless, the inclusion of the genus in Non-fistular Acamidae with cladotylote echinating spicules.
Porifera' Demospongiae • Poecilosclerida • Microcionina • Acarnidae 415

Fig. 2. A-J, Acamus innominatus Gray (holotype). A, choanosomal style (scale 300 f.Lm). B-C, larger and smaller echinating cladotylotes (scales 200 f.Lm
and 50 f.Lm, respectively). D, ectosomal tylote (scale 200 f.Lm). E, accolada toxa (scale 200 JLm). F, oxhorn toxa (scale 50 f.Lm). G, deeply curved toxa (scale
50 f.Lm). H, palmate isochela (scale 25 f.Lm). I, peripheral skeleton (scale I mm). J, idealised reconstruction of skeleton. K, Acamus bergquistae Van Soest
et ai., 1991, specimen NTM Z855 (from Hooper, 1987, fig. 40; scale 50mm).

Diagnosis swollen microspined bases (180--280 X -4 /-Lm); choanosomal styles

robust, smooth, slightly curved at centre (280-460 X 11-22/-Lm);
Non-fistulose, encrusting to massive growth fonns; ectosomal echinating cladotylotes in two size classes: larger with entirely
tylotes with microspined ends, forming tangential or paratangential smooth shaft, smooth, rounded basal tyle and four apical clads
tracts (not brushes); choanosomal skeleton isodictyal, isotropic or (230--300 X 6-12 /-Lm); smaller with sparsely spined shaft, smooth,
anisotropic reticulate, occasionally plumo-reticulate or plumose or rounded basal tyle and four clads on apical end (80--115 X 2-3 /-Lm);
hymedesmioid in encrusting species; skeletal tracts cored by microscleres with three categories of toxas: oxhom toxas (50--160 X
smooth styles, echinated by 1 or 2 (exceptionally 3) categories of 2-4/-Lm); thin deeply curved toxas (35-150 X 1.5-2/-Lm); accolada
cladotylotes, echinating acanthostyles present or absent; microscle- toxas (220-450 X 0.5-1.5 /-Lm); palmate isochelae (9-24/-Lm).
res are palmate isochelae and toxas of several morphologies. Remarks. Acamus has been revised extensively and many new
species described over recent years (Levi, 1952; Van Soest, 1984b;
Description of type species Hooper, 1987; Van Soest, Hooper & Hiemstra, 1991; Hiemstra &
Hooper, 1991; Hooper & Levi, 1993a; Van Soest, Zea & Kielman,
Acamus innominatus Gray, 1867a (Fig. 2). 1994). The generic synonymy is expanded here to now include:
Synonymy. Acamus innominatus Gray, 1867a: 544 (based Fonteia Gray, 1867a (with type species Fonteia anomala Gray, 1867a
on unnamed spicule drawings of Bowerbank, 1864: pI. 3, figs (by monotypy) holotype unknown; Trefortia Deszo, 1880 (which is
73-76, pI. 18, fig. 292). Acamus carteri Ridley, 1884a: 354. probably a nomen nudum as neither the type species nor type material
Material examined. Holotype (schizotype): BMNH are known (de Laubenfels, 1936a:92»; Acanthncamus Levi, 1952
Bowerbank collection 309 - West Indies. (type species Acanthncamus souriei Levi, 1952 (by monotypy) holo-
Description. Thin or massively encrusting; surface optically type MNHN LBIMDCL1259); and Microtylotella Dendy, 1896 (type
smooth; oscules not apparent; texture soft; ectosomal skeleton species Microtylotella guentheri Dendy, 1896 (by monotypy) holo-
tangential layer of tylotes; choanosomal skeleton plumose major type NMV G2366). The first two genera are included with question
choanosomal tracts ascending to surface interconnected by given the dubious (or non-existent) status of type material. The latter
renieroid isotropic tracts, both cored by choanosomal styles in genus is here included into synonymy based on the discovery of
5-15 spicules per tract; renieroid component of skeleton dominates cladotylotes in the holotype. It is nevertheless unusual to the genus in
skeletal structure; choanosomal tracts echinated by cladotylotes, having ectosomal and choanosomal skeletal tracts penneated with
particularly at nodes; megascleres ectosomal tylotes with slightly sand and with a reduced spicule skeleton.
416 Porifera· Demospongiae • Poecilosclerida • Microcionina • Acarnidae

I
F
I
AB

c
Fig. 3. Acheliderma lemmiscata Topsent (holotype). A-B. acanthostyles (scale 50!Lm and 20 !Lm, respectively). C, choanosomal style (scale 100 !Lm).
D, ectosomal tylote (scale 50 !Lm). E, base of tylote (scale 2!Lm). F, diamond-shaped microxea (scale 50 !Lm). G, toxa (scale 20 !Lm). H, peripheral skeleton
(scale 250 !Lm) . I, idealised reconstruction of skeleton.

Distribution one species); choanosomal skeleton an irregular reticulation of

microspined styles and echinating acanthostyles; microscleres
Cosmopolitan in temperate and tropical seas, predominantly characteristically elongated diamond-shaped microxeas and toxas;
shallow water. chelae absent.

Description of type species

ACHEUDERMA TOPSENT, 1892
Acheliderma lemmiscata Topsent, 1892a (Fig. 3).
Synonymy Synonymy. Acheliderma lemmiscata Topsent, 1892a: 24.
Material examined. Holotype (schizotypes): MNHN
Acheliderma Topsent, 1892a. Astylinifer Topsent, 1927b. DTl70, BMNH 1953 .11.9.19 - Banyuls, Mediterranean.
Fusifer Dendy, 1896. Taxonomic decision for synonymy: Van Description. Encrusting growth form with small erect fis-
Soest et al. (1994). tules up to 10 mm high protruding through substrate; ectosomal
skeleton a tangential layer of tylotes with microspined bases over-
Type species laying a plumose subectosomal skeleton composed of the same
spicules extending to the surface from the basal choanosomal
Acheliderma lemmiscata Topsent, 1892a (by monotypy). skeleton; choanosomal skeletal structure predominantly plumose
due to the dominance of the subectosomal tracts of tylotes; basal
Definition choanosomal skeleton irregularly renieroid-reticulate composed of
choanosomal styles with microspined bases forming tracts of 1-3
Fistular Acarnidae with elongated diamond-shaped microxea spicules and triangular or square meshes; skeletal tracts echinated
microscleres. by acanthostyles; megascleres ectosomal tylotes with microspined
bases (250-310 X 3-4/-Lm), basally spined choanosomal styles
Diagnosis and subtylostyles (420-450 X 4-9 /-Lm), short acanthostyles evenly
spined (80-175 X 3-5 /-Lm); microscleres wing-shaped toxas
Fistulose growth forms; ectosomal spicules are tylotes with (90-130 X 1-2 /-Lm) and diamond-shaped microxeas (60-70 X
microspined bases lying tangential to surface (replaced by sand in 1 /-Lm); chelae absent.
Porifera' Demospongiae • PoeciloscIerida • Microcionina • Acarnidae 417

Remarks. Van Soest et al. (1994) suggest that the unusual Description of type species
diamond-shaped microxeas ('raphides fusiformes' of Topsent,
1892a), characteristic of this genus, are derived from toxas, and is Cornulella lundbecki Dendy, 1922b (Fig. 4).
most obvious in the transition series of microxeas-toxas seen in A. Synonymy. Cornulella lundbecki Dendy, 1922b:l03.
planum Topsent (1928c: 228). In the type species, however, this Material examined. Holotype: BMNH 1923.4.1.113 -
relationship is less obvious with spicules resembling 'true' microx- Seychelles, W Indian Ocean.
eas and toxas at each end of the continuum. All three nominal gen- Description. Delicate thin-walled fistules up to 13 mm high,
era (Acheliderma, Astylinifer (type species A. planum Topsent, 3 mm diameter, arising from thinly encrusting base; ectosomal
1928c (by monotypy) schizotype MNHN DT1288), and Fusifer skeleton with unispicular tangential layer of basally spined tylotes
(type species F. jistulatus Dendy, 1896 (by monotypy) syntype forming a criss-cross on the surface with vaguely differentiated
NMV G2356, schizotype BMNH 1902.10.18.61) have these ascending and transverse mu1tispicular tracts supporting the fistule
spicules and have similar growth form, skeletal structure and spic- wall; choanosomal skeleton vestigial, hymedesmioid in thinner
ulation and are now considered to be synonyms. Microspined parts of skeleton with individual tylotes and microrhabd-like
tylote ectosomal spicules support the inclusion of Acheliderma in (acanth)oxeas paratangentia1 to (but not echinating) basal layer of
Acarnidae, although in one South Australian species (A. jistulatus spongin, and tylotes forming multi spicular longitudinal tracts sup-
(Dendy, 1896» these have been secondarily lost and replaced by porting fistules and (acanth)oxeas dispersed singly amongst
sand, and in two other species (A. planum and A. lisannae) some or ty10tes; no fibres and poor collagen in mesohyl; megascleres tylotes
all ectosomal spicules are mucronated anisotylotes. with microspined bases (310-420 X 4-8/-Lm), small microrhabd-
The genus is intermediate between Paracornulum and like (acanth)oxeas with a roughened surface and sometimes with
Cornulum in having both echinating acanthostyles and structural a central or sub-central swelling (16-24 X 1-2/-Lm); microscleres
styles. The type species is somewhat unusual in having an exten- large palmate isochelae with thickened alae (42-58/-Lm); two
sive plumose subectosomal region, extending from the irregular sizes of toxas: larger slightly curved at centre with recurved arms
renieroid basal skeleton, occupying most of the sponge diameter. (470-640 X 5-8/-Lm), smaller sharply curved at centre, with
This is most evident in the fistular region (section BMNH recurved arms, rare (110-135 X 2-5/-Lm).
1953.11.9.19), whereas in the thinner encrusting section it is not so Remarks. Cornulella differs from Paracornulum in lacking
extensive (section MNHN DT170). In the BMNH material many accessory echinating acanthostyles although both genera have
arcuate and unguiferous arcuate isochelae were also seen, but these microrhabd-like acanthoxeas and lack structural styles in the
are presumably contaminants (possibly artifacts from sectioning), choanosomal skeleton (Van Soest et al., 1994). Van Soest et al.
not seen in other material or described in the literature. (1994) suggest that these microrhabds are thin, spined oxeas and
not true microrhabds, contrary to Dendy's (1922b) interpretation,
Distribution although they are extremely small and could be validly included as
a class of microscleres, particularly in the type species where their
Four valid species are currently recognised (Van Soest et al., ornamentation is poorly developed. The possession of basally
1994). Additional reviews are provided by Topsent (1892a: 24; spined ectosomal tylotes and palmate isochelae indicates affinities
1928c: 228-9), Dendy (1896:49) and Desqueyroux-Faundez & Van with the Acarnidae. The type species of Cornulella has quite
Soest (1996). Caribbean, Mediterranean, Azores, NE Atlantic and different choanosomal skeletal architecture from other Acarnidae,
South Australia, shallow water. lacking any trace of an organised reticulate skeleton and
having instead longitudinal and transverse multi spicular tracts
typical of fistular growth forms. This skeletal structure is more-or-
CORNULELLA DENDY, 1922 less typical for the genus, which varies from simply a loose collec-
tion of spicules to more organised tracts supporting the
Synonymy choanosome.
Van Soest et al. (1994) discuss relationships and provide com-
Cornulella Dendy, 1922b: 103. parative spicule dimensions for the six recognised species currently
included in Cornulella. All are similar in habit (thinly encrusting,
Type species possibly excavating, fistulose) and are readily differentiated in their
toxa morphology and spicule sizes.
Cornulella lundbecki Dendy, 1922b (by monotypy).
Distribution
Definition
Western Indian Ocean (Madagascar, Amirante, Seychelles),
Fistular Acarnidae with vestigial choanosomal skeleton of Caribbean, tropical Western Atlantic.
single tylotes and scattered acanthoxeas.

Diagnosis CORNULUM CARTER, 1876

Thinly encrusting, fistular growth forms; ectosomal skeleton Synonymy

a unispicular network of intercrossing tangential tylotes with spined
heads; choanosomal skeleton vestigial, made up of individual Cornulum Carter, 1876:309. Cornulotrocha Topsent, 1927b:9.
tylotes identical to those in the ectosomal skeleton, with accessory Coelosphaerella de Laubenfels, 1934. Melonchela Koltun, 1955b.
acanthoxeas distributed singly among the tylotes; microscleres Heterocornulum Levi & Levi, 1983b. Taxonomic decision for
palmate isochelae and frequently toxas. synonymy: Van Soest et al. (1994: 185) and Hooper (this work).
418 Porifera· Demospongiae • Poecilosclerida • Microcionina • Acarnidae

Fig. 4. A-F, Cornu leila lundbecki Dendy (holotype). A, portion of tylote (scale 20l1m). B, toxa (scale 100 11m). C, palmate isochela (scale 20 11m).
D, roughened microrhabd-like (acanth)oxea (scale 20 11m). E, section through peripheral skeleton (scale I mm). F, idealised reconstruction of the skeleton.
G, Cornulella tyro Van Soest et al., 1994 (from their fig. 3; scale I mm).

Type species Description of type species

Cornulum textile Carter, 1876 (by monotypy). Cornulum textile Carter, 1876 (Fig. 5).
Synonymy. Cornulum textile Carter, 1876: 309.
Definition Material examined. Holotype: BMNH 1882.7.28.76
(schizotype BMNH 1910.1.1.1363) - NEAtlantic.
Fistular Acamidae with apically-spined tylotes or strongyles Description. Growth form lobate-digitate, fistulose; surface
forming both tangential ectosomal and plumo-reticulate choano- optically smooth, concentrically striated; ectosomal skeleton mod-
somal skeletons, sometimes together with choanosomal styles or erately thick tangential layer of tylotes running longitudinally
aniso-strongyles, echinating spicules present in one species only. along surface in uni- or bispicular tracts, with occasional transverse
unispicular tracts overlaying ectosome; choanosomal skeleton
plumo-reticulate with multi spicular tracts of both choanosomal
Diagnosis styles and ectosomal tylotes forming diverging tract ascending
towards surface, cored by 10-15 spicules per tract, interconnected
Massive or encrusting growth forms, and with erect fistules or by smaller transverse tracts cored by the same spicules 5-8
pronounced pore areas; ectosomal skeleton consists of a compact spicules per tract, together forming triangular or occasionally
crust of spicule layers orientated tangential to the surface, square meshes; megascleres include ectosomal tylotes, often asym-
composed of smooth tylotes/strongyles with microspined apices, metrical, with slightly swollen bases usually microspined, occa-
or reduced to mucronate apices in one species; choanosomal sionally smooth (290-370 X 8-14 f,Lm); choanosomal styles vary
skeleton consists of tracts and scattered tylotes/strongyles as from true styles to aniso-strongyles with asymmetrical slightly
found in the ectosome, with monactinal spicules (styles) bearing swollen, micros pined or smooth bases and rounded or pointed tips
microspined bases in some species; echinating acanthostyles (390-480 X 15-22 f,Lm); toxas very abundant forming tangled
present in one species only, erect on a hymedesmioid basal skele- masses in between spicule tracts, usually greatly curved at centre
ton; microscleres palmate isochelae, toxas, and microstrongyles in and with slightly reflexed arms, occasionally modified (straight, sin-
one species. uous) (190-310 X 1-3 fLm); small palmate isochelae unmodified,
Porifera' Demospongiae • Poecilosclerida • Microcionina • Acarnidae 419

I
CD
A B

Fig. S. Cornulurn textile Carter (holotype). A, choanosomal style (scale 100 ....m). B, base of style (scale 30 .... m). C, ectosomal tylote (scale 100 ....m).
D, toxa (scale 100 .... m). E, palmate isochela (scale 5 .... m). F, choanosomal skeleton (scale 1 mm). G, ectosomal skeleton (TS) (scale I mm). H, bundles of
toxas (scale 50 .... m). I, idealised reconstruction of skeleton. J, specimens of Lundbeck (1910; from his pI. 2, figs 13-14; size 40mm).

very abundant, often completely lining the choanocyte chambers synonymy. It is remarkable in several features: its erect planar retic-
(11-15 f.Lm). ulate growth form (superficially resembling the microcionid
Remarks. Van Soest et al. (1994) suggested that the Clathria coppingeri and the raspailiid Echinodictyum cancellatum);
nominal genera Coelosphaerella (without styles) (type species extremely large size range of principal spicules, the largest protrud-
Cornulum johnsoni de Laubenfels, 1934 (by original designation) ing a long way through fibre bundles, reminiscent of Raspailiidae;
holotype USNM 22364), Xytopsene (without styles or toxas), the apparent lack of connecting fibres between the ascending
Heterocornulum (with microstrongyles), and Melonchela (with plumose spiCUle tracts; a ridge-like subterminal ornamentation on
peculiar anisochelae described as 'melonchelas' ) are all members toxas; and extremely abundant tracts of chelae microscleres
of Cornulum sharing the microspined condition of megascleres throughout the mesohyl. It is described as having peculiar
(occasionally verging towards entirely smooth), as opposed to the anisochelae ('melonchelas') but in fact these are cleistochelae and
more heavily and extensively basally spined condition of megas- belong to the palmate group of chelae. The presence of true diacti-
cleres seen in other acarnids. This synonymy is supported here with nal ectosomal spicules (tylotes, strongyles and intermediates, vary-
the exception of Xytopsene which is included in Damiria. Reviews ing in the degree of swelling of their bases), palmate chelae and
of species are found in Carter (1876:309), Lundbeck (1910: toxas indicate that Melonchela has affinities to genera such as
22), Koltun (1955b: 17; 1959: 187), Levi & Levi (1983b: 966), Acarnus and referred to the Acarnidae as a synonym of Cornulum.
Bergquist & Fromont (1988: 53) and Hooper (1996a). Heterocornulum (type species Heterocornulum virguliferum
Melonchela (redescribed in detail by Hooper, 1996a:85; type Levi & Levi, 1983b (by original designation) holotype MNHN
species Melonchela clathrata Koltun, 1955b (by monotypy) DCL2925) is burrowing/excavating with fistules; ectosomal skeleton
paratype BMNH 1963.7.29.7) is a doubtful inclusion in this is a dense criss-cross of tylotes or tylostyles with asymmetrical
420 Porifera· Demospongiae • Poecilosclerida • Microcionina • Acarnidae

ends, microspined on both ends, and abundant microstrongyles. tylotes bearing microspined bases; choanosomal skeleton slightly
The choanosomal skeleton is pulpy, occupying excavations in the irregularly halichondroid with loose reticulated multispicular tracts
substrate, with tylotes the same as on the sUlface, smooth small composed of thicker tylotes in bundles of up to 10 spicules per
choanosomal styles and smooth microstrongyles. Microscleres tract, vaguely subisodictyal in arrangement; megascleres are thin-
are thick accolada toxas, and palmate isochelae (Levi & Levi, ner ectosomal tylotes with microspined bases, frequently with
1983b: 966). asymmetrical bases (255-312 X 6-10 /-Lm), and thicker choanoso-
Cornulotrocha (type species Cornulotrocha cheliradians mal tylotes also with microspined bases (200-250 X 11-15 /-Lm);
Topsent, 1927b (by original designation) type material not exam- microscleres absent.
ined) also clearly belongs to this group, having a fistulose growth Remarks. Van Soest et al. (1994) broadened the definition of
form, lacking any differentiation between ectosomal and choanoso- Damiria to allow for the inclusion of species with toxas (D. toxifera
mal megascleres, which consist of subtylostyles modified to Van Soest et aI., 1994) although no other species have yet been
strongyles, often with mucronate apices; these spicules form weakly described with microscleres. Damiria is a sister genus of Zyzzya and
reticulate unispicular tracts in the choanosomal skeleton, with poor CornuZella based on similarities in the spination of tylotes although
fibres, and an irregular felt (tangential) ectosomal skeleton; echinat- Van Soest et al. (1994) concede that it might be a reduced
ing acanthostyles are erect on a hymedesmioid basal layer of spon- Cornulella or Paracornulum-like group having secondarily lost
gin; microscleres are palmate isochelae in rosettes lying between their chelae and acanthose oxeas/styles. Six species are currently
the choanosomal spicule tracts, and toxas (Topsent, 1928c: 226). included: D. simplex Keller, 1891, D. toxifera Van Soest etaZ., 1994,
The types species C. cheliradians Topsent differs only substantially D. leonorae Van Soest et al., 1994, D. testis Topsent, 1928c,
from Cornulum s.s. in having stylote to strongylote megascleres D. curvata (Vacelet, 1969) and D. jistuZatus (Carter, 1880b),
with modified mucronate apices, replacing apical spines, and whereas D. australiensis Dendy, 1896 was referred to Lissodendoryx
possessing echinating acanthostyles (unlike other species). given that it has smooth tylotes, oxeas and arcuate isochelae.
The type species of Xytopsene, Suberites jistulatus Carter from
Distribution the Gulf of Manaar (and allegedly also from SW Australia) (by orig-
inal designation, holotype lost), is fistulose, globular growth form,
Indo-west Pacific, Caribbean, NE Atlantic. cork-like consistency, has a cavernous choanosomal skeleton com-
posed of loose tracts of basally spined tylotes with microspined
bases, and has palmate isochelae microscleres. Unfortunately the
DAMIRIA KELLER, 1891 type specimen is no longer extant in the LFM or BMNH collections
(Hooper & Wiedenmayer, 1994: 139), and Carter's (1880b: 53)
Synonymy description and illustration is so brief that it cannot be ascertained
whether there are one or two size categories of megascleres.
Damiria Keller, 1891. Anisotylacanthaea Vacelet, 1969: 200. Nevertheless, from the information available the proposed synonymy
Xytopsene de Laubenfels, 1936a: 54. Taxonomic decision for between Xytopsene and Damiria is reasonable.
synonymy: Van Soest et al. (1994: 187). More uncertain is the placement of AnisotyZacanthaea.
The type species, A. curvata Vacelet from the Mediterranean
Type species (by original designation, holotype MNHN DJV689), is fistulose,
encrusting or massive, with a detachable ectosomal skeleton of tan-
Damiria simplex Keller, 1891 (by monotypy). gential tylotes ('acanthotylostrongyles'), with one swollen base
prominently microspined and the other completely smooth, strewn
Definition loosely on the surface. The choanosomal skeleton is a fragile,
cavernous reticulation of poorly developed tracts, without spongin
Fistular Acarnidae with apically microspined tylotes as the fibres, composed of both tylotes and styles with smooth bases and
only megascleres. acanthose shafts (although there is no clear evidence of spicules
being restricted to ectosomal or choanosomal regions of the skele-
Diagnosis ton). Microscleres are absent. Vacelet (1969) initially included the
genus in the Myxillidae based on similarities to the genera lanulum
Fistular growth form; ectosomal skeleton with tangential crust and Damiria, whereas Van Soest et al. (1994) suggest that it may
of tylotes with microspined apices; choanosomal skeleton irregu- be a synonym of Damiria although not conforming closely to it or
larly reticulate with tracts of larger tylotes of similar morphology the allied genera Zyzzya, Paracornulum and Cornulella.
as ectosomal spicules; no other megascleres; microscleres may
include toxas. Distribution

Description of type species Indo-Pacific (Madagascar, Red Sea, Amirante, Seychelles,

Indonesia), Caribbean, Azores, NE Atlantic.
Damiria simplex Keller, 1891 (Fig. 6).
Synonymy. Damiria simplex Keller, 1891: 309. Damiria
simplex variety jistulata Hentschel, 1912. DOLICHACANTHA HENTSCHEL, 1914
Material examined. Holotype: 2MB310b (schizotype
BMNH presently missing) - Red Sea. Synonymy
Description. Encrusting-massive, with robust fistules; sur-
face optically smooth; ectosomal skeleton with tangential crust of Dolichacantha Hentschel, 1914: 118.
Porifera· Demospongiae • Poecilosclerida • Microcionina • Acarnidae 421

Fig. 6. A-F, Damiria simplex Keller (holotype). A, ectosomal tylote (scale 50/-Lm). B, base of ectosomal tylote (scale 10 /-Lm). C, choanosomal tylote
(scale 50 /-Lm). D, base of choanosomal tylote (scale 10 /-Lm). E, peripheral skeleton (scale 100 /-Lm). F, idealised reconstruction of skeleton. G, Damiria tox-
ifera Van Soest et aI., 1994 (from their fig. 6; scale 2.5 mm).

Type species Synonymy. Dolichacantha macrodon Hentschel, 1914: 118.

Material examined. Holotype: ZMH S2337 - Gauss-
Dolichacantha macrodon Hentschel, 1914 (by monotypy). Station, Antarctica.
Description. Encrusting to massive growth encrusting forms
Definition up to 2 mm thick; ectosomal skeleton with peculiar long acantho-
strongyles (with spined bases) and tylotomotes (with smooth bases)
Non-fistulose Acarnidae with long ectosomal acan- arranged in bundles on the surface; choanosomal skeleton hyme-
thostrongyles bearing numerous spined 'cladi' along the apical desmioid with a basal skeleton with embedded small acanthostyles
third of shaft. and larger partially spined choanosomal subtylostyles standing
perpendicular to the substrate; bundles of tylotomotes and acan-
Diagnosis thostrongyles form tracts more-or-Iess ascending through the surface,
making the sponge very hispid; megascleres include very long smooth
Encrusting to massive growth forms; ectosomal skeleton with ectosomal tylotornotes with one tylote end and an asymmetrical non-
long acanthostrongyles intermingled with tylotornotes arranged in tylote end, both without apparent basal spination (776-1472 X 9-12
bundles on the surface; choanosomal skeleton hymedesmioid with f,Lm), peculiar acanthostrongyles with numerous spined 'cladi' on the
entirely spined acanthostyles and basally- or vestigially entirely apical third of the spicule and granular basal spination on the swollen
spined subtylostyles embedded in the basal skeleton, standing per- base (1200--1456 X 10-11 f,Lm), entirely spined echinating acan-
pendicular to the substrate, and bundles of tylotornotes and/or acanthostyles (176-360X9-12f,Lm), larger subtylostyles with vestigial
thostrongyles forming tracts more-or-less ascending through the spination on shaft or with spines restricted to the base (1120-1680 X
surface; microscleres include only uncommon palmate isochelae. 20-23 f,Lm), and palmate isochelae (15-23 f,Lm) (spicule dimensions
taken from both type material and Hentschel, 1914).
Description of type species Remarks. Re-examination of a slide preparation of the
holotype (kindly loaned by Mr Peter Stiewe, Zoologisches Institut
Dolichacantha macrodon Hentschel, 1914 (Fig. 7). und Zoologisches Museum, Hamburg, with a slide prepared by
422 Porifera· Demospongiae • Poecilosclerida • Microcionina • Acarnidae

D
I
I
E

Fig. 7. A-F, Dolichacantha macrodon Hentschel (holotype). A, cladote and basal ends of acanthostrongyle (scale 200 /-Lm). B, enlarged c1adote end of acan-
thostrongyle (scale 50 /-Lm). C, echinating acanthostyle (scale 100 /-Lm). D, apical and basal ends of ectosomal tylotomote (scale 100/-Lm). E, palmate isochelae
(scale 10 /-Lm). F, idealised reconstruction of skeleton. G, Dolichacantha shikotani Koltun, 1970 (from his pI. 3, fig. 4; specimen about 50mm long).

Rob Van Soest) showed that sigmas present in the tissue tease were and they may be derived from the common ectosomal tylotes/
loosely scattered between megascleres, whereas isochelae were strongyles. For the present the genus is included in Acarnidae given
firmly embedded within collagen throughout the mesohyl. Whereas its possession of ectosomal tylotornotes and palmate isochelae.
the former were abundant and the latter were relatively scarse, a fact This assignment is supported by the presence of basally-spined
also recorded by Hentschel (1914), it was decided that sigmas were subectosomal styles and echinating acanthostyles fonning a per-
contaminants whereas isochelae were native to the sponge. This pendicular hymedesmioid choanosomal skeleton (with the latter
conclusion is supported by Koltun's (1970: 205) description of a two spicule types common to both Acarnidae and Microcionidae,
second species, D. shikotani Koltun from the NW Pacific, in which as well as Hymedesmiidae). The alternative system, that it is
palmate isochelae were also reported but no sigmas. a member of Hymedesmiidae, is rejected given the possession of
The skeletal structure of the type species is difficult to deter- palmate chelae and tangential tornotes in both known species of
mine precisely, as the surviving portion of the holotype is Dolichacantha. Although this current assignment is equivocal
extremely small and consists of little more than a tease of tissue given that tylotornotes in D. macrodon lack apical spination,
containing bundles of spicules and some collagen. Hentschel's a characteristic of Acarnidae, tornotes in Koltun's (1970) species
(1914) description states that the architecture is hymedesmioid, D. shikotani do bear basal spines and it may be these have been
with acanthostyles (Le., both smaller echinating acanthostyles and secondarily lost in the type species.
larger subtylostyles, which are interpreted here to represent princi-
pal choanosomal styles) embedded in basal spongin and standing Distribution
perpendicular to the substrate. He also states that the ectosomal
skeleton has bundles of both tylotornotes and acanthostrongyles Cold water; Antarctica and vicinity of Kurile Islands, NW
lying on the surface, and also projecting a long way through it, and Pacific.
the acanthostyles are said to be mixed in with the tylotornotes.
The peculiar inventory of spicules in this genus make its fam-
ily assignment difficult. Tylotornotes may perhaps be interpreted IOPHON GRAY, 1867
as related to microspined tylotes, and the 'cladi' on the acan-
thostrongyles may indicate Acamus-affinities. Alternatively, the Synonymy
hooked acanthostrongyles may also be modified exotyles, in which
case they may have little phylogenetic significance (given that Iophon Gray, 1867a: 534. [Alebion] Gray, 1867a: 534 (preoc-
they occur in one form or another in Hadromerida and Mycalidae), cupied). Menyllus Gray, 1867a: 533. Ingallia Gray, I 867a: 537.
Porifera· Demospongiae • Poecilosclerida • Microcionina • Acarnidae 423

A
I I I B C

I
D

Fig. 8. lophon scandens (Bowerbank) (holotype). A, choanosomal styles (scale 50 fJ-m). B,juvenile choanosomal styles (scale 50 fJ-m). C, ectosomal tylote
(scale 50 .... m). D, spurred palmate anisochelae (scale 10 ....m). E, bipocillae (scale to ....m). F, peripheral skeleton (100 .... m). G, idealised reconstruction of
skeleton. H, ho)otype (from Bowerbank, 1874, pI. XLV, fig. 14; size about 9 em).

Pocillon Topsent, 1893c. lophonopsis Dendy, 1924. Burtonella Diagnosis

de Laubenfels, 1928. lophonota de Laubenfels, 1936a.
Hymedesanisochela Bakus, 1966b. Taxonomic decision for syn- Non-fistulose, massive, branching or encrusting growth
onymy: de Laubenfels (1936a: 85), Van Soest et al. (1994: 185), forms; ectosomal skeleton composed of tylotes with microspined
Desqueyroux-Faundez & Van Soest (1996: 7), Hooper (this work). bases, lying tangential to surface, intercrossing and in erect
brushes; choanosomal skeleton an isodictyal rounded, triangular or
Type species square-meshed reticulation of smooth or spined choanosomal
styles, arranged singly or in 2-3 per row, with (s.\.) or without (s.s.)
Halichondria scandens Bowerbank, 1866 (by original desig- echinating acanthostyles; rnicroscleres inequiended bipocilla and
nation; reinstated by Ridley & Dendy, 1886). palmate anisochelae with spurs, toxas absent.

Definition Description of type species

Non-fistular Acarnidae with a regular, uni- or pauci-spicular lophon scandens (Bowerbank, 1866) (Fig. 8).
isodictyal choanosomal reticulation of smooth or spined styles, Synonymy. Halichondria scandens Bowerbank, 1866: 259;
with or without echinating acanthostyles. 1874: p\. XLV. Iophon scandens; Dendy, 1924: 347.
424 Porifera· Demospongiae • Poecilosclerida • Microcionina • Acarnidae

Material examined. Holotype (schizotypes): BMNH (Dendy, 1924), and consequently it also falls into synonymy with
1910.1.1.2255,2256,2257,2258 - Shetland. Iophon.
Description. Growth form ranges from encrusting to Hymedesanisoehela (type species H. rayae Bakus, 1966b (by
cushion-shaped, often epizootic; surface optically smooth; oscules original designation) holotype not examined) differs from Iophon
small, scattered; ectosome with irregularly scattered tangential and s.s. only in having an encrusting habit and two size classes of
paratangential bundles of tylotes and many chelae also scattered on (acantho)styles forming hymedesmioid skeletal structure, stylote
surface; choanosomal skeleton with fairly regular isodictyal reticu- modifications to tylotes, and lacking bipocillae, whereas in other
lation of choanosomal styles, arranged 2-3 per row, forming rela- respects it is typical of the genus. Neither of these characters pro-
tively wide, mostly triangular or square meshes, occasionally vides any reasonable synapomorphy for Bakus's genus and hence it
rounded, and ascending spicule tracts slightly thicker than trans- is synonymised here. Other nominal genera included into syn-
verse connecting tracts producing a slightly plumo-reticulate onymy with Iophon are: [Alebion] Gray (preoccupied, with type
appearance; echinating acanthostyles absent; mesohyl moderately species A. proximum Ridley, 1881 (by subsequent designation;
light with numerous chelae and juvenile (wispy) styles scattered Ridley, 1881 (Vosmaer, 1882»; Menyllus Gray (type species
between skeletal meshes; megascleres ectosomal tylotes with faint Haliehondria ingalli Bowerbank, 1866 (by original designation»,
to prominent microspination (145-210 X 3-9IJ-m), choanosomal Ingallia Gray (type species I. eupulifera Gray, 1867a (by mono-
styles with small spines on the basal end, or sometimes over typy», Burtonella de Laubenfels (type species B. melanokhemia
the entire spicule or completely smooth (195-250 X 8-13IJ-m); de Laubenfels, 1936a (by original designation» and Iophonota
microscleres are rare inequiended bipocilla with saucer-shaped de Laubenfels (type species Iophon aeeratus Hentschel, 1914
alae (7-15IJ-m long) and abundant palmate anisochelae with spurs (by original designation». Although none of the holotypes of the
(15-32IJ-m long); toxas absent. type species have been seen by the author this synonymy has been
Remarks. Van Soest et al. (1994) and Desqueyroux- verified by Bakus (1966b: 479), Bergquist & Fromont (1988: 90), Van
Faundez & Van Soest (1996) provide a detailed discussion of the Soest et al. (1994), and Desqueyroux-Faundez & Van Soest (1996).
genus and its synonymy. Iophon lacks toxas, common to many
species of this family, but shares palmate anisochelae (cleis- Distribution
tochelae) with Melonehela (=Comulum) and Aeanthorhabdus.
Mycalidae also share this latter character but this is interpreted as a Cold and temperate waters of most oceans and seas, not yet
convergence by Van Soest et al. (1994). In skeletal structure Iophon known from the shallow tropics.
resembles, and has been traditionally assigned to Myxillidae, but
in the revised sense of Hajdu et al. (1994a) the latter group is
restricted to taxa with anchorate chelae and sigmata. This decision MEGACIELLA HALLMANN, 1920
de-emphasises the primary importance of the 'classic myxillid'
isotropic skeletal reticulation but this certainly occurs in distantly Synonymy
related taxa (e.g., Antho of Microcionidae, Lissodendoryx of
Myxillidae, Eetyoplasia and Amphinomia of Raspailiidae). Megaeiella Hallmann, 1920. Myxiehela de Laubenfels,
Re-examination of relevant type material of both Iophon 1936a. Taxonomic decision for synonymy: Van Soest et al.
seandens (redescribed above) and I. hyndmani (Bowerbank, 1858) (1994: 187).
(schizotypes BMNH Bowerbank collection 147,935) showed that
they are not conspecific as suggested by Desqueyroux-Faundez & Type species
Van Soest (1996). The two species differ quite substantially in the
geometry, size and spination of their megascleres, possession of Amphileetus pilosus Ridley and Dendy, 1886 (by original
echinating acanthostyles in the latter, relative abundance of designation).
bipocillae chela in the latter (scarse in the former), and differences
in their gross morphologies. Iophon seandens has also been misin- Definition
terpreted by Dendy (1924) and subsequent authors as having echi-
nating acanthostyles, whereas type material clearly shows that it Non-fistular Acarnidae with confused choanosomal skeleton
does not. On this basis it falls into Dendy's (1924) concept of divided into primary ascending multispicular tracts and secondary
Iophonopsis (type species Haliehondria nigrieans Bowerbank, interconnecting uni- or bispicular tracts cored by basally spined
1866 (by original designation) holotype not examined), created styles, with or without echinating acanthostyles.
specifically for 'Iophon-like' species lacking echinating acan-
thostyles. This makes Iophonopsis redundant given that the type Diagnosis
species of both nominal genera, Iophon and Iophonopsis, both lack
echinating megascleres. In any case, the generic distinction based Non-fistulose, branching, lobate, flabellate or encrusting
on the absence or presence of echinating spicules is not considered growth forms; ectosomal skeleton composed of tangential or
of great phylogenetic value at the generic level of classification, paratangential layer of ectosomal tylotes with microspined bases,
with precedent seen in other genera of Poecilosclerida (e.g., often sinuous or curved; choanosomal skeleton reticulate or con-
Aeamus versus Aeanthaeamus (Van Soest et al., 1991), Clathria fused (hymedesmioid in encrusting species), with ascending multi-
versus Isociella and Axociella (Hooper, 1996a». Poeillon (type and paucispicular tracts of smooth choanosomal styles with spined
species Isodietya implicita Bowerbank (by original designation) bases, interconnected by secondary uni- or bispicular tracts bound
junior synonym of Haliehondria hyndmani Bowerbank (Dendy, together with collagen producing irregular wide meshes; echinat-
1924: 347), holotype not examined) is an objective synonym of ing acanthostyles present or absent; microscleres are palmate
Iophonopsis through synonymy of their respective type species isochelae and two morphologies of toxas.
Porifera· Demospongiae • Poecilosclerida • Microcionina • Acarnidae 425

I
B

A
I

Fig. 9. Megaciella pilosus (Ridley and Dendy) (lectotype). A, choanosomal style (scale 100 j.Lm). B, base of choanosomal style (scale 50 j.Lm). C, ectoso-
mal tylote (scale 50 j.Lm). D, base of ectosomal tylote (scale 20 j.Lm). E, accolada toxa (scale 250 j.Lm). F, wing-shaped toxa (scale 50 j.Lm). G, palmate
isochela (scale 2 j.Lm). H, ectosomal skeleton (scale 200 j.Lm). I, peripheral choanosomal skeleton (scale 200 j.Lm). J, idealised reconstruction of skeleton.
K, lectotype (from Ridley & Dendy, 1887: pI. XXV, fig. 3; length about 8 em).

Description of type species are not associated with the ectosomal skeleton but scattered
throughout the mesohyl. Conversely, 'bundles of oxeas' described by
Megaciella pilosus (Ridley and Dendy, 1886) (Fig. 9). Ridley & Dendy (1887) are actually bundles of ectosomal tylotes.
Synonymy. Amphilectus pilosus Ridley and Dendy, 1886: Hallmann (1920) tacitly included MegacieUa within the
350; Megaciella pilosus; Hallmann, 1920: 772. Microcionidae, with closest affinities to Artemisina (e.g., lax skele-
Material examined. Lectotype: BMNH 1887.5.2.125 - tal construction and supposedly comparable spiculation), differing
Marion Is, Kerguelen. in lacking ornamentation on toxas. However, both genera have
Description. Lobate flabellate growth form; surface shaggy, quite different ectosomal skeletons (basally spined diactinal tylotes
ridged, hispid; ectosomal skeleton with tangential or paratangential in Megaciella, monactinal ectosomal skeleton in Artemisina), and
layer of tylotes, often in bundles; subectosomal skeleton radial or consequently Van Soest (1984b) referred it to Myxillidae, whereas
plumose, with erect choanosomal principal styles protruding from Van Soest et al. (1994) are not completely certain it belongs to
peripheral fibres through surface; choanosomal skeleton reticulate, Acamidae or Microcionidae (with the independent acquisition of
with ascending multi- or paucispicular tracts of choanosomal prin- tylotes assumed in the former case). However, the type species
cipal styles, interconnected by secondary uni- or bispicular tracts does appear to have true tylotes with microspined bases, which
within light spongin, producing irregular wide meshes; echinating under the revised scheme of Hajdu et al. (1994a) Megaciella is
spicules absent; megascleres are ectosomal tylotes, often curved more appropriately included in Acamidae. In fact, the possession
or sinuous, with slightly swollen, microspined bases (230-295 X of palmate isochelae and toxas being the only real synapomorphy
3-5 f1m), very large smooth choanosomal principal styles between MegacieUa and the Microcionidae (i.e., Microcionina).
(565-1215 X 5-8 f1m); microscleres minute palmate isochelae Another Iophonid, Acamus, is also similar in this respect.
(4-6 f1m), two sizes of toxas, very large accolada toxas (490- The type species of Megaciella lacks echinating acan-
1200 X 2-4 f1m) and wing-shaped toxas (70-195 X 1-3 f1m). thostyles, whereas Van Soest et al. (1994) and Desqueyroux-
Remarks. Ridley & Dendy (1886, 1887) mistakenly consid- Faundez & Van Soest (1996) remark that 'similar species with
ered the very large accolada toxas to be 'long thin centrally curved acanthostyles exist', but mention only one: Myxichela microtoxa
oxeas' in the type species Amphilectus pilosus, and moreover, they de Laubenfels, 1935.
426 Porifera· Demospongiae • Poecilosclerida • Microcionina • Acarnidae

Van Soest et al. (1994) include Myxichela de Laubenfels in standing perpendicular to surface. Subectosomal skeleton radial or
synonymy with Megaciella. The type species Lissodendoryx taw- plumose, with tracts of smaller tylotes ascending to surface, and
iensis Wilson (by original designation, holotype USNM21272) has acanthostyles also scattered throughout mesohyl. Ectosome with
a lamellate growth fonn, an ectosomal skeleton composed of erect tangential tracts of larger tylotes forming a nearly halichondroid
brushes of spined choanosomal styles protruding through the sur- reticulate criss-cross on the surface. Megascleres are tylotes with
face, and a tangential or paratangential layer of ectosomal granular-microspined bases, of 2 size categories: mainly ectosomal
tylostrongyles (or subtylotes or subtylostyles). The choanosomal (380-440 X 4-11 f.Lm) and mainly choanosomal (200-265 X
skeleton is a multispicular reticulation of differentiated primary 4-8 f,LID), and acanthostyles (with some oxeote or strongylote modi-
ascending and smaller connecting fibres composed of poor spon- fications) (85-150 X 3-5 f.Lm). Microscleres palmate isochelae
gin, found mainly at nodes of spicule tracts, cored by spinous (12-17 f.Lm) and oxhorn toxas, slightly curved or straight (30-79 X
styles, and tracts becoming plumose in the periphery; microscleres 1.5-3.5 f.Lm).
are toxas and palmate isochelae. Further references: Ridley & Remarks. Paracornulum does not appear to be as closely
Dendy (1886: 350); Hallmann (1920: 772); Desqueyroux- related to microcionids as inferred by Hallmann (1920), showing
Faundez & Van Soest (1996); Hooper (1996a: 84). similarities mainly in the possession of echinating acanthostyles,
hymedesmioid basal spongin fibres and palmate isochelae and
Distribution toxas. Van Soest et al. (1994) referred other species previously
assigned to Paracornulum to Cornulella, Cornulum and Zyzzya,
Disjunct, relatively deeper water species-Kerguelen, Sulu Sea including those included there by Hallmann (1920) and Bergquist &
(Indonesia), California. Fromont (1988). Another species assigned to Paracornulum at one
time or another (Cliona purpurea Hancock) remains poorly known
and currently uncertain in its placement. Based on its ectosomal
PARACORNULUM HALLMANN, 1920 characteristics and megasclere spiculation Paracornulum was
referred to Cornulidae Levi & Levi (1983b: 966), whereas
Synonymy under the scheme of Hajdu et al. (1994) it belongs to Acarnidae,
similar to the fistular genera Cornulum Carter, Iophon Gray and
Paracornulum Hallmann, 1920: 772. [Cornulacantha] Levi & Zyzzya de Laubenfels. Unlike these genera Paracornulum has
Levi, 1983b: 966; Desqueyroux-Faundez & Van Soest, 1996: 26 true acanthostyles echinating the basal skeleton, similar to
(nomen nudum). Taxonomic decision for synonymy: Levi & Levi Acheliderma.
(1983b: 966), Desqueyroux-Faundez & Van Soest (1996: 26). Levi & Levi (1983b) and Desqueyroux-Faundez & Van Soest
(1996) mention a genus name 'Cornulacantha Levi' as being a syn-
Type species onym of Paracornulum, but the name was not found in the literature
or Zoological Record. Levi (pers.comm.) stated that he intended to
Cornulurn dubium Hentschel, 1912 (by original designation). create the name for the South African species, Paracornulum
coherens Levi, 1963, but during final editing of his (1963) manu-
Definition script he chose instead to use Hallmann's genus Paracornulum. Van
Soest et al. (1994: 184) retained P. coherens in Paracornulum only
Fistular Acamidae with acanthostyles echinating the on a tentative basis (given that its robust, smooth acanthostyles and
hymedesmioid basal skeleton overlain by apically-spined tylotes ectosomal strongyles are notably different from those of the type
forming both plumose ascending choanosomal tracts and tangen- species), perhaps justifying the recognition of a separate genus,
tial ectosomal tracts. however 'Cornulacantha' remains a nomina nudum.

Diagnosis Distribution

Encrusting-fistular growth fonn, with ectosomal skeleton Aru I., Indonesia, and South Africa.
composed of intercrossing halichondroid tracts of larger tylotes
with granular or microspined apices, and choanosomal skeleton
composed of similar but slightly smaller spicules forming radial or TEDANIOPHORBAS DE LAUBENFELS, 1936
plumose tracts arising from a hymedesmioid layer of basal spongin
echinated by acanthostyles perpendicular to the substrate; Synonymy
microscleres are palmate isochelae and toxas.
Tedaniophorbas de Laubenfels, 1936a.
Description of type species
Type species
Paracornulum dubium (Hentschel, 1912) (Fig. 10).
Synonymy. Cornulum dubium Hentschel, 1912: 346. Amphilectus ceratosus Ridley & Dendy, 1886 (by original
Paracornulum dubium; Hallmann, 1920: 772. designation).
Material examined. Holotype: SMF964 (schizotype
BMNH 1930.8. 13.83a) -Aru I., Indonesia. Definition
Description. Encrusting to fistular growth fonn. Surface
smooth, even. Choanosomal skeleton hymedesmioid, with spongin Non-fistular Acamidae with well-developed reticulate
fibres lying on substrate, bases of acanthostyles embedded and spongin fibre skeleton but lacking any coring or echinating
Porifera' Demospongiae • Poecilosclerida • Microcionina • Acarnidae 427

A
I
I
BCD
II

Fig. 10. A-I, Paracornulum dubium (Hentschel) (holotype). A, ectosomal tylote (scale 50 fl-m). B, base of ectosomal tylote (scale 10 fl-m) . C, choano-
somal tylote (scale 50 fl-m). D, echinating acanthostyles (scale 25 fl-m). E, oxhom toxa (scale 20 fl-m) . F, palmate isochela (scale 5 fl-m). G, ectosomal
skeleton (scale 250 fl-m). H, peripheral skeleton (scale 250 fl-m). I, idealised reconstruction of skeleton. J, Paracornulum coherens Levi. 1963 (from his
pI. 4G; about 5 cm diameter).

spicules, having instead apically-smooth ectosomal tylotes Material examined. Holotype: BMNH 1887.l.24.7 - Port
scattered between fibres and forming a tangential ectosomal Jackson, NSW, Australia.
skeleton. Description. Massive, lobate growth form, spongy, elastic tex-
ture, surface uneven, microconulose; ectosome with distinct, darker,
Diagnosis detachable cortex, with a crust of isochelae and irregularly scattered
ectosomal tylotes, in bundles or individually; choanosomal skeleton
Massive, lobate growth form; ectosomal skeleton detachable, more-or-less evenly reticulate, well developed horny spongin
composed of scattered smooth tylotes, singly or in bundles, form- fibres with large, even meshes; fibres completely aspicular although
ing a tangential skeleton; choanosomal skeleton well developed, ectosomal tylotes scattered between fibres, individually or forming
reticulate, composed of horny spongin fibres lacking coring or plumose tracts particularly near the periphery, and abundant chelae
echinating spicules, but with ectosomal tylotes scattered between throughout the skeleton; ectosomal tylotes with prominently swollen,
fibres and aggregated into loose wispy tracts near the periphery; smooth bases (175-235 X 3-7 /-Lm); palmate isochelae large, heavily
microscleres are palmate isochelae; toxas absent. silicified, with prominently thickened lateral alae (18-25 /-Lm).
Remarks. Tedaniophorbas is currently a monotypic, orphan
Description of type species genus, which Van Soest et al. (1994) suggested might belong to
Coe1osphaeridae but also closely resembled the Acamidae. It was
Tedaniophorbas ceratosus (Ridley & Dendy, 1886) (Fig. 11). also suspected from their respective published descriptions
Synonymy. Amphilectus ceratosus Ridley & Dendy, that Tedaniophorbas might be a reduced Megaciella, however,
1886: 350. Tedaniophorbas ceratosus; de Laubenfels, 1936a: 95. subsequent comparisons of their respective type species showed
428 Porifera· Demospongiae • Poecilosclerida • Microcionina • Acarnidae

II
AB
I
c

Fig. 11. Tedaniophorbas ceratosus (Ridley & Dendy) (holotype). A, tylote (scale 50 !Lm). B, base of tylote (scale 10 !Lm). C, palmate isochelae
(scale 10 fJ-m). D, ectosomal skeleton (scale 250 !Lm). E, peripheral skeleton (scale 250 !Lm). F, idealised reconstruction of skeleton. G, holotype
(from Ridley & Dendy, 1887: pI. XXV, fig. 2; diameter about 3cm).

marked contrasts in skeletal architecture (horny reticulate, aspicular Type species

versus reticulate, heavily spiculose with massive spicules but poor
fibre development), composition of choanosomal skeletons (com- Wigginsia wigginsi de Laubenfels, 1953 (by original
pletely aspicular versus completely spicular), ornamentation of designation).
ectosomal tylotes (smooth heads versus spined), toxa morphology
(absent versus two forms, respectively). The type species differs
from other acamids in lacking spined bases on ectosomal tylotes, Definition
and it is assumed that these spines have become secondarily lost.
Non-fistulose Acamidae with 'plocamiform' acan-
Distribution thostrongyles echinating tracts of choanosomal styles forming
isodicyal tracts.
Known only from Port Jackson, NSW (Australia).
Diagnosis

WIGGINSIA DE LAUBENFELS, 1953 Encrusting lamellate non-fistular sponge with tangential ecto-
somal tylotes bearing faint basal spination, thin, vaguely ascending
Synonymy tracts of smooth or faintly basally spined styles and a secondary
skeleton of 'plocamiform' acanthostrongyles forming a tight
Wigginsia de Laubenfels, 1953: 7. meshed isodictyal skeleton. Microscleres palmate isochelae only.
Porifera· Demospongiae • Poecilosclerida • Microcionina • Acarnidae 429

Fig. 12. Wigginsia wigginsi de Laubenfels (holotype). A, choanosomal (acantho-)styles, including juvenile form (scale 100 J.Lm). B, 'plocamiform'
acanthostrongyles (scale 50 J.Lm). C, ectosomal tylotes (scale 50 J.Lm). D, palmate isochelae (scale 10 J.Lm). E, ectosomal skeleton (scale 100 J.Lm).
F, choanosomal skeleton (scale 100 J.Lm). G, isodictyal skeleton formed by 'plocamiform' acanthostrongyles (scale 100 J.Lm).

Description of type species tyles, often asymmetric between ends, and smooth or only very
faintly microspined bases (190-245 X 5-9 fLm); choanosomal
Wigginsia wigginsi de Laubenfels (Fig. 12). (acantho-)styles, long curved shaft, tapering and sharply pointed,
Synonymy. Wigginsia wigginsi de Laubenfels, 1953: 7-9, the larger entirely smooth or with few basal spines, the smaller
fig. 4. more heavily spined at the base and distal portion of shaft
Material examined. Holotype: USNM 23222 - 24 km from (175-484 X 11-19 fLm), with raphidiformjuvenile styles also com-
Point Barrow, Alaska, 153 m depth. mon (about 120 X 2 fLm); 'plocamiform' acanthostrongyles with
Description. Small lamellate growth form encrusting rock, slightly curved shaft, spines dispersed over whole spicule but
3 mm high X 50 mm wide X 3 mm lamellae thickness, with smaller thicker on basal and central portions of the spicule, with evenly
lamellae abutting larger ones; surface with small elevations 1 mm rounded or asymmetrical ends (138-174 X 8-13 fLm). Microscleres
high each covered with smaller microconules; red alive; oscules palmate isochelae, with shaft contorted and reduced alae to resem-
not observed; consistency firm, harsh. Ectosomal skeleton dense, ble pseudo-arcuate forms (25-36 fLm).
with tangential layer of tylotes, 2-3 mm deep, through which protrude Remarks. This monotypic genus is referred to Acarnidae
long choanosomal (acantho-)styles in bundles erect on the surface, from Myxillidae, to which it was originally assigned, based on the
their bases embedded in terminal ascending spongin fibres. possession of faintly spined bases of ectosomal tylotes, some
Choanosomal skeleton with ascending multispicular tracts of large choanosomal styles also have faintly microspined bases, isochelae
styles, heavily 'echinated' by 'plocamiform' acanthostrongyles, the are palmate, and it lacks sigmas. It is unique in Acarnidae in having
latter forming close-set parallel tracts and producing a regular iso- 'plocamiform' acanthostrongyles that form a secondary isodictyal
dicyal reticulation ('log cabin architecture' of de Laubenfels). reticulate skeleton, reminiscent of other 'plocamiform' Microcionina
Megascleres short thick ectosomal tylotes with slightly swollen such as Antho (Microcionidae) and Plocamione (Raspailiidae).
430 Porifera· Demospongiae • PoeciloscIerida • Microcionina • Acarnidae

I
B

A
I cI I I D E

Fig. 13. Zyzzyafuliginosa (Carter) (holotype). A, ectosomal tylote (scale 50ILm). B, base oftylote (scale IOlLm). C, acanthostrongyle (scale 50lLm). D, acan-
thotylote (scale 50 ILm). E, tylostyle (scale 50 ILm). F, palmate isochela (from QM specimen, scale 5 ILm). G, ectosomal skeleton (scale 250 ILm). H, choanoso-
mal skeleton (scale 250 ILm). I, idealised reconstruction of skeleton. J, specimen NTM Z2891 (from Hooper & Krasochin, 1989: fig. 1; specimen approx. 20 em).

The species has not been recorded since it was first described from Type species
the Arctic, and its skeleton and spicules have not been previously
illustrated. Measurements provided by de Laubenfels (1953) for Piocamia massalis Dendy, 1922b (by original designation)
spicules were also in error. (junior synonym of Suberitesfuliginosus Carter, 1879b; taxonomic
decision for synonymy Van Soest et ai., 1994).
Distribution
Definition
Known only from the Alaskan Arctic.
Fistulose Acarnidae with verticillately-spined strongyles
(or strongyloxeas).
ZYZZYA DE LAUBENFELS, 1936
Diagnosis
Synonymy
Massive, burrowing, fistulose or cryptic encrusting growth
Zyzzya de Laubenfels, 1936a. Damirina Burton, 1959a. form with solid apical fistules or blind papillae; ectosomal skeleton
Histodermopsis de Laubenfels, 1936a. Taxonomic decision for thick detachable crust of tangentially orientated tylotes with
synonymy: Van Soest et ai. (1994: 165). microspined bases; choanosomal skeleton with distinctive irregular
Porifera· Demospongiae • PoeciloscIerida • Microcionina • Acarnidae 431

or plumose multi spicular, widely spaced tracts ascending to the Remarks. Damirina Burton (with type species D. verticilata
surface cored by the same tylotes, interspersed with a unispicular, Burton (by original designation, holotype BMNH 1936.3.4.510) is
irregular isodictyal reticulation of verticillately-spined strongyles an objective synonym of Zyzzya due to synonymy of their respec-
(or strongyloxeas); microscleres if present are palmate isochelae tive type species. Histodermopsis de Laubenfels is also included
and toxas. here in synonymy with Zyzzya based on Lundbeck's (1910)
comprehensive description and illustrations of the type species.
Description of type species The holotype of H. coriacea Lundbeck from the North Sea
(by original designation) has yet to be found (Zool. Mus.
Zyzzyajuliginosa (Carter, 1879b) (Fig. 13). Copenhagen - missing) but is described as being elongated,
Synonymy. Suberites juliginosus Carter, 1879b: 347. Zyzzya bulbous, fistulose growth form, with a tangential ectosomal skele-
fuliginosa; Van Soest et aI., 1994: 168. Plocamia massalis Dendy, ton composed oftylotes (and strongylotes) with microspined bases
1922b: 78. Dendoricella massalis; Topsent, 1928c: 64. and swelling of variable development, an irregularly reticulate
Lissodendoryx massalis; Burton, 1935a: 400; Thomas, 1973: 32. choanosomal skeleton composed of scattered acanthostrongyles
Zyzzya massalis; de Laubenfels, 1936a: 64. Damirina verticillata and acanthoxeas without definite arrangement, no echinating
Burton, 1959a: 240. Paracornulum atoxa Vacelet et al., 1976: 59. spicules, and no microscleres. De Laubenfels (1936a:72) included
Material examined. Holotype of S. fuliginosus: BMNH other species in Histodermopsis, some with arcuate chelae and
1846.8.5.8 - Torres Strait, far north Queensland. Holotype of sigmas, but these were misplaced, and only the type species is here
P. massalis: BMNH 1921.11.7.67 - Mauritius. included.
Description. Excavating, thinly encrusting, fistular, massive This genus was comprehensively revised by Van Soest et al.
to pear-shaped; endolithic parts insubstantial, invasive of the sub- (1994), including a review of all published Indo-west Pacific popu-
strate, but without producing distinct galleries or chambers; surface lations. From this revision only three species are now recognised:
smooth, consistency generally fragile, live colouration brown to the type species (with several junior synonyms listed above),
black; ectosomal skeleton multilayered crust of intercrossing tan- Z. papillata (Thomas, 1968) from India and the Seychelles, Z. invemar
gentially arranged tylotes; choanosomal skeleton unispicular iso- Van Soest et al., 1994 from Colombia, and tentatively Z. coriacea
dictyal reticulation of acanthostrongyles plus many ectosomal (Lundbeck) from the North Sea (pending re-examination of its
tylotes interspersed, as well as multi spicular tracts traversing holotype). The unique feature of this genus is the possession of
basal part; megascleres ectosomal tylotes with well developed, verticillately-spined strongyles, but otherwise it conforms quite
often asymmetrical, microspined bases (173-590 X 1.5-20 fJ..m), closely to other Acamidae in ectosomal and microsclere spiculation.
occasionally modified to stylote spicules; acanthostrongyles
verticillately-spined, or simply regularly or irregularly spined, or Distribution
occasionally smooth on shaft, rarely stylote (109-300 X 3-28 fJ..m);
microscleres palmate isochelae, but often absent from some Widespread Indo-Pacific, from Zanzibar to Fiji, and
specimens (including holotype) (14-22 fJ..m). Colombian Carribean, one species from sub-Arctic deeper waters.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Microcionidae Carter, 1875

John N.A. Hooper

Queensland Museum, P.O. Box 3300, South Brisbane, Qld, 4101, Australia. (lohnH@qm.qld.gov.au)

Microcionidae Carter (Demospongiae, Poecilosclerida), including Clathriidae, Ophlitaspongiidae, contains 77 nominal genera of which
only 9 genera and 12 subgenera are considered to be valid (one incertae sedis). There are approximately 470 described (valid) species
worldwide, living predominantly in shallow waters with a few recorded from deeper seas, and with many other species still collected but
remaining undescribed. Microcionids typically have three skeletal regions delineated by the distribution of different structural megascle-
res: (1) the choanosomal skeleton (with principal monactinal spicules enclosed within spongin fibres and spined monactinal spicules typ-
ically echinating fibres; in two groups this is replaced by a basal or axial renieroid skeleton of smooth or acanthose styles or strongyles,
with or without echinating spicules); (2) an extra-fibre subectosomal skeleton (with tracts of larger auxilIary monactinal spicules ascend-
ing to the surface); (3) and a non-tangential ectosomal skeleton (with smaller auxillary styles forming a surface crust perpendicular to the
surface). Megascleres are predominantly smooth ectosomal and choanosomal styles, with some diactinal and acanthose modifications.
Microscleres are palmate isochelae, only exceptionally modified to superficial arcuate-like or anchorate-like forms (produced by torsion
of the shaft and detachment of alae), and toxas with diverse morphologies including microxea-like and raphidiform toxas in few species.
Skeletal structures range from 'hymedesmioid' and 'microcionid' in encrusting taxa, to plumo-reticulate and occasionally axially com-
pressed in some species, but usually irregularly reticulate in most taxa. Occasionally spicules are partially or completely replaced by detri-
tus. Two subfamilies are recognised: Ophlitaspongiinae de Laubenfels (with a secondary renieroid spongin fibre and/or spiculose skeleton
overlaying a primary reticulate, plumo-reticulate, plumose or hymedesmioid spiculo-spongin skeleton) and Microcioninae Carter (lacking
a secondary renieroid reticulate skeleton, having only a reticulate, plumo-reticulate, plumose hymedesmoid, microcionid or axially com-
pressed primary skeleton).
Keywords: Porifera; Demospongiae; Poecilosclerida; Microcionina; Microcionidae; Microcioninae; Clathria (Clathria); C. (Wilsonella);
C. (Microciona); C. (Dendrocia); C. (Axosuberites); C. (Isociella); C. (Thalysias); Holopsamma; Echinochalina (Echinochalina);
E. (Protophlitaspongia); Pandaros; Ophlitaspongiinae; Antho (Antho); A. (Acarnia); A. (Isopenectya); Artemisina; Echinoclathria;
Ophlitaspongia; Sigmeurypon (incertae sedis).
- -...- - - - - - - - - - - - -

DEFINITION, DIAGNOSIS, SCOPE Diagnosis

Synonymy Microcionina with encrusting, lobate, arborescent or flabel-

late growth forms. Skeleton differentiated into choanosomal
Microcionina Carter, l875c. Microcionidae Hentschel, 1923. (axial), subectosomal (extra-axial) and ectosomal regions. Axial
Clathriidae Lendenfeld, 1884. Ophlitaspongiidae de Laubenfels, skeleton formed by unispicular or multispicular tracts of choanoso-
1936a. mal (principal) megascleres, typically coring spongin fibres or
sometimes simply bound together by collagen. Fibres echinated by
(acantho-) styles (accessory spicules). Skeletal structures include
Definition isodictyal, renieroid, reticulate, plumo-reticulate, plumose or
hymedesmioid, but never radial. Subectosomal (extra-axial) skele-
Microcionina with terminally spined ectosomal styles, rarely ton formed by tracts of subectosomal (auxiliary) spicules, usually
modified to quasidiactinal or diactinal forms; three skeletal regions dispersed outside of fibres, rarely well organised but usually with
defined by the presence of different forms of structural styles: some degree of difference between axial and extra-axial regions.
(1) choanosomal (axial) skeleton (with spongin fibres enveloping Ectosomal skeleton ranges from membraneous, or with protruding
principal styles echinated by acanthose or smooth styles; in two larger (subectosomal auxiliary) spicules, or with a special category
groups this is replaced by a basal or axial renieroid skeleton of of smaller (ectosomal auxiliary) spicules. Principal megascleres
smooth or acanthose styles or strongyles, with or without echinat- monactinal, predominantly smooth or only partially spined, occa-
ing spicules); (2) subectosomal (extra-axial or extra-fibre) skeleton sionally vestigial or absent completely, supplemented by another
(with individual or tracts of auxiliary styles ascending to the sur- category of acanthose diactinal spicules, or sometimes replaced by
face); and (3) ectosomal skeleton (with smaller auxiliary styles detritus. Auxiliary megascleres usually monactinal, rarely quasi-
forming a surface crust tangential, paratangential or perpendicular diactinal, smooth shaft and basal spines, more slender than
to the surface). One or more skeletal regions may be lost or modi- choanosomal spicules. Echinating styles or subtylostyles smooth,
fied. Megascleres predominantly smooth styles but may be modi- partially or completely spined. Renieroid skeleton composed of
fied and/or supplemented by quasidiactinal or diactinal forms, or acanthose or occasionally smooth styles or 'dumbell' spicules.
lost completely and replaced with detritus. Microscleres include Microscleres include toxas of several morphologies (including
palmate isochelae and diverse forms of toxas. raphidiform and rarely microxeote toxas), and palmate isochelae.

432
Porifera· Demospongiae • Poecilosclerida • Microcionina • Microcionidae 433

Some species have modified chelae, arcuate-like or anchorate-like, Differences with similar families
a result of torsion of the shaft and detachment of alae, but these are
exceptional. Larvae viviparous. Microcionidae is very difficult to define in terms of one
or more apomorphies. It differs from other families of
Scope Microcionina in lacking certain features: such as apically spined
diactinal ectosomal tylotes or tornotes characteristic of Acarnidae;
Microcionidae is a highly specious family, containing about small oxeote or stylote ectosomal spicules forming bouquets,
580 described species (of which about 470 are 'valid'), with many unique to Raspailiidae; or having rhabdostyles and several peculiar
other undescribed species already known from various collections microscleres seen in Rhabderemiidae. Similarly, the four families
worldwide (Hooper, unpublished data). 82 nominal genera have of Microcionina differ from the Myxillina and Mycalina in lacking
been included in the family at one time or another, of which 77 particular characters, as opposed to possessing unique ones
were recognized as (potentially) residing here (Hooper, 1996a, and (Myxillina with tridentate-derived chelae; Mycalina with sigman-
more recent additions). Most of these genera were merged into cistra derivatives), yet both these latter suborders may also have
others (e.g., Levi, 1960b; Simpson, 1968; Van Soest, 1984b; palmate chelae in addition to their derived microscleres.
Bergquist & Fromont, 1988; Hooper, 1990b; Hooper & Following a recent comprehensive revision (Hooper, 1996a)
Wiedenmayer, 1994; Hooper, 1996a; Howson & Chambers, 1999), Microcionidae is now restricted to genera having predominantly
leaving only 9 valid genera and 12 subgenera: Clathria (with sub- smooth monactinal ectosomal and choanosomal spicules.
genera Clathria, Wilsonella, Microciona, Dendrocia, Axosuberites, It excludes certain microcionid-like genera with true tylotes or
Isociella, Thalysias), Antho (with subgenera Antho, Acarnia, strongylotes as their ectosomal spicules (e.g., Acarnus,
Isopenectya), Echinoclathria, Holopsamnw, Echinochalina (with Megaciella). These taxa are now included in Acarnidae, as defined
subgenera Echinochalina, Protophlitaspongia), Artemisina and by their ectosomal features (Hajdu et al., 1994a). However, the
Ophlitaspongia, plus Pandaros and Sigmeurypon as incertae sedis. definition barely differentiates species with modified or reduced
Microcionidae are found in all seas, are sometimes highly diverse quasidiactinal (styloid) auxiliary megascleres (e.g., several
and prevalent in shallow coastal waters, comprising up to 16% of Echinoclathria, Holopsamnw and Echinochalina species), or
species in some regional faunas (Hooper & Levi, 1993a; Hooper, quasimonactinal (amphistrongylote or tornote-like) auxiliary
1996a). Most species have been recorded from shallow-waters but megascleres (e.g., Protophlitaspongia). These modified auxiliary
some range down to 2500m depth (Hartman, 1982), and it is likely spicules are usually asymmetrical and are interpreted here as
that the family is substantially more diverse than presently known. convergent upon true diactinal spicules. These anomalous micro-
cionids share certain characteristics of both Microcionidae and
History and biology Acarnidae, and the importance of these characters at higher levels
of systematics must therefore be questioned, or a certain level of
The disagreement as to which of Microcionidae Carter and homoplasy must be accomodated in the phylogeny of the suborder.
Clathriidae Hentschel had priority was settled by Hooper (1996a) Similarly, the definition given above cannot always clearly
based on Article 40 of the International Code of Zoological distinguish some Microcionidae from some Raspailiidae, but this is a
Nomenclature (Anon., 1999). Hooper (1996a) also provided
a comprehensive revision of the family based on extensive
re-examination of type material of all nominal type species of gen-
era and over 400 nominal species, primarily from the Indo-west
Pacific region.
Several substantial taxonomic publications have appeared for
this family, including descriptions of regional faunas, analyses of
morphological characters, chemotaxonomy and/or detailed taxo-
nomic revisions (Hallmann, 1920; Levi, 1960b; Van Soest, 1984b;
Bergquist & Fromont, 1988; Hooper, I 990b, 1996a; Hooper &
Levi, 1993a; Hooper et al., 1992). These studies have produced
several schemes based on the emphasis of certain characters over
others, such as patterns of skeletal architecture (Levi, 1960b),
megasclere form and distribution within the skeleton (Hallmann,
1920), ectosomal structure and composition (Van Soest, 1984b),
microsclere geometry and diversity (e.g., de Laubenfels, 1936a).
More recently all these features, based on a consensus of opinions,
were combined into a contemporary classification (Bergquist &
Fromont, 1988), with the net result being a proliferation of genera
not all of which can be justifiably supported on phylogenetic
grounds. This 'consensus' classification is rejected here, although Fig. 1. Idealised microcionid skeletal structure. Key: I, ectosomal skeleton.
ultimately molecular data may confirm or refute this opinion. 2, subectosomal skeleton. 3, choanosomal skeleton. 4, basal spongin fibre.
The most recent phylogenetic analysis (Hooper, 1996a) was 5, echinating acanthostyle. 6, reticulate fibre skeleton. 7, isotropic extra-
based primarily on structural features of the skeleton, particularly fibre skeleton. 8, detrital entrapping fibre. 9, renieroid reticulate secondary
fibre skeleton. 10, microcionid radial fibre skeleton. II, hymedesmioid
skeletal architecture and skeletal differentiation. Lesser support spicule skeleton. 12, plumose/dendritic fibre skeleton. 13, coring principal
was given to megasclere geometry and virtually no support was spicules. 14, subectosomal auxuliary spicules. 15, spicate spicule skeleton.
accorded microsclere modification (cf. Hajdu et al., 1994a). 16, ectosomal auxiliary spicules. Modified from Hooper (1996a).
434 Porifera· Demospongiae • PoeciloscIerida • Microcionina • Microcionidae

Fig. 2. Range of skeletal structures in Microcionidae. Arrows indicate diagnostic features (as mentioned in the Key to genera and subgenera).
A-I, Microcioninae. A, Clathria (Clathria) (C. (C.) lipochela Burton). B, Clathria (Microciona) (C. (M.) antarctica (Topsent)). C, Clathria (Dendrocia)
(C. (D.) pyramida Lendenfeld). D, Clathria (Wilsonella) (C. (w.) australiensis Carter). E, Clathria (Axosuberites) (C. (A.) canaliculatus (Whitelegge)).
F, Clathria (lsociella) (C. (I.) eccentrica (Burton)). G, Clathria (Thalysias) (C. (7:) rubra (Lendenfeld)). H, Holopsamma (H. crassa Carter). I,
Echinochalina (Echinochalina) (E. (E.) australiensis (Ridley)). A-I, modified from Hooper (1996a).

problem of semantics rather than a biological one. In general, most Theoretically, this is a viable system for the suprafamily classifica-
species of Raspailiidae have well-compressed axial skeletons, tion of Microcionidae, but in practical terms it is not always possi-
well-differentiated axial and extra-axial skeletons, and special ble to distinguish between true tridentate-derived chelae and
bouquets of ectosomal spicules surrounding the larger choanoso- palmate chelae with 'arcuate' or 'anchorate' modifications (i.e., par-
mal spicules protruding through the surface. In contrast, most tial torsion of the shaft and partial or complete detachment of alae
Microcionidae lack these two former features, or they are only from the shaft). These few anomalies have yet to be fully resolved.
poorly developed and probably convergent, perhaps related to habit
(e.g., flexuous whip-like growth forms). Nevertheless, there are Previous reviews
examples in both families where the boundaries between taxa
blur, such as the microcionid-like Raspailia (Clathriodendron) De Laubenfels (1936a: 112), Levi (1960b: 50), Simpson
arbuscula (see Hooper, 1991: figs 19-20), and the raspailiid- (1968: 102), Van Soest (1984b: 90), Wiedenmayer (1989: 56),
like Clathria (Axosuberites) canaliculata (see Hooper, 1996a: Bergquist & Fromont (1988: 106), Hooper & Wiedenmayer (1994:
figs 118-119). Conversely, the two families are consistently differ- 252), Hooper (1996a: 1).
entiated by this latter ectosomal feature, and possession of chelae
microscleres in Microcionidae (absent in Raspailiidae), which
appear to be more important characters than skeletal structure. PROPOSAL FOR A DIVISION OF M1CROCIONIDAE
Hajdu et al. (1994a) restricted Microcionina (and hence
Microcionidae) to taxa with only palmate isochelae, tacitly Hooper's (1996a) analysis of Microcionidae showed that
excluding several microcionid-1ike genera specifically created there were two fundamental groups of genera within the family,
for species with tridentate-derived (arcuate or anchorate) chelae. differentiated primarily by their skeletal structure, the complexity
Porifera· Demospongiae • Poecilosclerida • Microcionina • Microcionidae 435

Fig. 3. Range of skeletal structures in Microcionidae (cont.). A-B , Microcioninae. A, Echinochalina (Protophlitaspongia) (E. (P.) tuberosa Hooper).
B, Pandaros (P. acanthifolium D&M). C-H, Ophlitaspongiinae. C, Antho (Antho) (A. (A.) tuberosa (Hentschel» . D, Antho (Acamia) (A. (A.) ridleyi
(Hentschel». E, Antho (Isopenectya) (A. (I.) chartacea (Whitelegge» . F, Artemisina (A. jovis Dendy). G, Echinoclathria (E. parkeri Hooper).
H, Ophlitaspongia (0. papilla Bowerbank). A.C-G. modified from Hooper (l996a). B. modified from Van Soest (l984b). H. modified from Howson &
Chambers (1999).

or differentiation of the skeleton, and the presence or absence, The possibility that designated subgenera should be elevated to
respectively, of a secondary renieroid reticulate skeleton overlaying full generic status was rejected by Hooper (1996a), on the basis that
the primary (reticulate, plumo-reticulate, plumose or hymedesmioid) the characters inferring polyphyly (e.g., partial substitution of cor-
skeleton. (1) One group, contammg Artemisina, Antho, ing megascleres by detritus in Wilsonella, and loss of principal
Echinoclathria and Ophlitaspongia, has suggested myxillid-like spicules completely in Dendrocia) are homoplasious, representing
features, most possessing a secondary renieroid skeleton overlaying convergences through functional acquisition or secondary loss of
a primary reticulate, plumo-reticulate, plumose or hymedesmioid particular features, rather than real apomorphies. Thus, some of
spiculo-spongin skeleton. (2) The other group, containing Clathria, these subgeneric taxa may be artificial given the existence of these
Echinochalina, Holopsamma and Pandaros, has suggested homoplasies, but the use of this subfamily classification allows us to
raspailiid-like features including similarities to typical raspailiid construct both a working phylogenetic hypothesis as well as the
genera (e.g., Clathria (Axosuberites) and Raspai/ia) and atypical production of a useful, working classification. Of dubious phyloge-
raspailiid genera (e.g., Echinochalina (Echinochalina) and netic value are the possession of acanthose strongyles in place of
Echinodictyum, respectively) (Figs 1-3). Within the second group acanthose styles in Antho and Plocamia, presence of detritus within
there are also several subgenera of Clathria indicated as being possi- fibres in Clathria (Wilsonella), and encrusting habit in Clathria
bly polyphyletic (Hooper, 1996a). This second group has only a pri- (Microciona). Conversely, other subgenera have a more substantial
mary skeletal structure, with reticulate, plumo-reticulate, plumose phylogenetic basis and are more easily justified within the classifi-
hymedesmoid, microcionid or axially compressed skeletal architec- cation presented here (e.g., stylote versus oxeote structural mega-
tures. These groups are elevated here to subfamily status: scleres in Echinochalina and Protophlitaspongia; presence or
Ophlitaspongiinae and Microcioninae, respectively. absence of ectosomal specialisation in Thalysias and Clathria).
436 Porifera· Demospongiae • Poecilosclerida • Microcionina • Microcionidae

KEYS TO MICROCIONIDAE

Key to subfamilies

(1) With a single primary skeletal structure: reticulate, plumo-reticulate, plumose hymedesmioid, microcionid or axially compressed
(Figs 2A-I, 3A-B) ........................................................................................................................................................ Microcioninae
With a secondary renieroid spongin fibre and/or spiculose skeleton overlaying a primary reticulate, plumo-reticulate, plumose or
hymedesmioid spiculo-spongin skeleton (Fig. 3C-H) ........................................................................................... Ophlitaspongiinae

Key to genera and subgenera (Refer to Figs 1-3 for skeletal structures)

(1) Choanosomal skeleton more-or-less undifferentiated, unstructured ................................................................................... Artemisina

Choanosomal skeleton well structured, ranging from hymedesmioid to reticulate, but lacking any differentiated components ....... 2
Choanosomal skeleton well structured, predominantly reticulate, differentiated into two distinct structural components (e.g., primary
renieroid skeleton overlaid by secondary plumose or radial skeletons) .............................................................................................. 7
(2) Choanosomal fibres cored by one or more category of principal spicules ................ ................................. ........................ ................. 3
Choanosomal fibres cored by auxiliary spicules partially (or sometimes wholly) replaced by detritus ............................................. 5
Choanosomal fibres cored by auxiliary spicules identical to those in ectosomal and subectosomal skeletons .................................. 6
Choanosomal fibres form a ladder-like isodictyal reticulation, but only primary fibres cored by plumose columns of megascleres and
transverse fibres virtually clear ..................................................................................................................................... Ophlitaspongia
Choanosomal fibres or skeletal tracts cored by auxiliary spicules different from those in peripheral skeleton .................... Pandaros
(3) Choanosomal skeleton without any marked axial compression or differentiated axial and extra-axial regions ................................. 4
Choanosomal skeleton with noticeably compressed axis and well differentiated axial and extra-axial (radial, plumose or plumo-
reticulate) regions; lacking echinating megascleres ........................................................................................ Clathria (Axosuberites)
Choanosomal skeleton evenly renieroid reticulate throughout, with well developed spongin fibres cored by smooth principal styles;
lacking echinating megascleres .............................................................................................................................. Clathria (Isociella)
Choanosomal skeleton hymedesmioid or microcionid, with basal layer of spongin lying on substrate without ascending fibre nodes
ranging to ascending non-anastomosing columns of spongin fibres; bases of principal styles perpendicular to the substrate or forming
plumose tracts in ascending fibres, and with acanthostyles or smooth styles echinating megascleres, differentiated from principal
styles coring fibres ............................................................................................................................................ Clathria (Microciona)
(4) With a single category of subectosomal auxiliary spicule forming the ectosomal skeleton, producing tangential, paratangential or
plumose tracts; choanosomal skeleton predominantly reticulate or plumo-reticulate ........................................... Clathria (Clathria)
With two categories of auxiliary spicules, smaller ectosomal spicules generally overlaying larger subectosomal spicules forming
discrete bundles or continuous palisade on surface .............................................................................................. Clathria (Thalysias)
(5) Echinating acanthostyles differentiated from principal spicules coring fibres ................................................... Clathria (Wilsonella)
Echinating styles or acanthostyles identical to principal styles coring spongin fibres .................................................... Holopsamma
(6) Echinating acanthostyles differentiated from auxillary spicules coring the fibres ............................................. Clathria (Dendrocia)
Fibres cored by a single category of auxillary styles identical to those forming subectosomal and ectosomal skeletons, and echinated
by smooth styles derived from principal spicules ............................................................................. Echinochalina (Echinochalina)
Fibres cored by a single category of auxiliary oxeas or styles different from principal oxeas, anisoxeas or styles that echinate
fibres ........................................................................................................................................... Echinochalina (Protophlitaspongia)
(7) Primary skeleton renieroid, cored by axially or basally compressed tracts of acanthostyles; secondary skeleton subisodictyal or
plumo-reticulate, cored by smooth principal styles in plumose tracts .......................................................................... Antho (Antho)
Primary skeleton renieroid, cored by axially or basally compressed tracts of acanthostrongyles; secondary skeleton subisodictyal or
plumo-reticulate, cored by smooth principal styles in plumose tracts ....................................................................... Antho (Acarnia)
Primary skeleton composed of axially compressed spongin fibres cored by renieroid tracts of sparsely spined principal styles inter-
mingled with plumose or plumo-reticulate tracts of smooth principal styles, overlaid by secondary extra-axial plumose skeleton cored
by larger smooth principal styles .......................................................................................................................... Antho (Isopenectya)
Primary renieroid reticulate skeleton cored by smooth principal styles and echinated by identical spicules, with secondary radial
extra-axial skeleton on exterior edge of skeleton cored only by larger smooth principal styles .................................. Echinoclathria

SUBFAMILY MICROCIONINAE CARTER, 1875 Definition

Synonymy Microcionidae lacking a secondary renieroid skeleton, with

only a reticulate, plumo-reticulate, plumose hymedesmioid, micro-
Microcionina Carter, 1875c. Microcionidae Hentschel, cionid or axially compressed primary skeleton.
1923.
Scope
Type genus
Four genera and seven subgenera: Clathria (Clathria
Clathria Schmidt, 1862. (Clathria), C. (Wilsonella), C. (Microciona), C. (Dendrocia),
Porifera· Demospongiae • Poecilosclerida • Microcionina • Microcionidae 437

C. (Axosuberites), C. (lsociella), C. (Thalysias)), Echinochalina, SUBGENUS CLATHRIA SCHMIDT, 1862

Holopsamma and Pandaros.
Synonymy

CLATHRIA SCHMIDT, 1862 Clathria Schmidt, 1862: 57. [Clatharia] Kumar, 1925: 221
(lapsus). Allocia Hallmann, 1920: 768 (type species Spanioplon
Synonymy cheliferum Hentschel, 1911: 362 (by original designation and
monotypy), holotype 2MB 4440). Antherochalina Lendenfeld,
Clathria Schmidt, 1862: 57 (see subgenera for full synonymy). 1887c: 741, 786 (type species Antherochalina crassa Lendenfeld,
1887c: 787 (by subsequent designation; Burton, 1934a: 558), holo-
Type species type BMNH 1886.8.27.450). Bipocillopsis Koltun, 1964b: 79 (type
species Bipocillopsis nexus Koltun, 1964b: 80 (by monotypy) holo-
Clathria compressa Schmidt, 1862: 58 (by subsequent type ZIL 10644). Dictyociona Topsent, 1913b: 579, 618 (type
designation; Schmidt, 1864: 35). species Microciona discreta Thiele, 1905: 447 (by monotypy),
holotype 2MB 3302). Labacea de Laubenfels, 1936a: 125 (type
Definition species Clathriajuncea sensu Burton, 1931a: 343 (by original des-
ignation), schizotype BMNH 1926.2.19.2 (unconfinned)). Ligrota
Microcioninae with auxiliary styles in one or two categories de Laubenfels, 1936a: 125 (type species Clathria lobata Vosmaer,
fonning various ectosomal structures ranging from membraneous 1880: 151 (by original designation) holotype RMNH 276).
paratangential to a dense erect palisade of brushes; choanosomal Litaspongia de Laubenfels, 1954: 162 (type species
skeleton well-structured, hymedesmioid to reticulate, with mostly Ophlitaspongia arbuscula Row, 1911: 347 (by original designa-
smooth styles enclosed within spongin fibres and fibres echinated tion) holotype BMNH 1912.2.1.63). Paresperia Burton, 1930c:
by mostly acanthose smaller styles. 501 (type species Paresperia intermedia Burton, 1930c: 501 (by
monotypy) holotype BMNH 1910.1.1.912). Marleyia Burton,
Diagnosis 1931a: 346 (type species Marleyia irregularis Burton, 1931a: 346
(by original designation) holotype NM 1279). Pitalia Gray, 1867a:
Thinly encrusting, massive, digitate, lamellate, flabellate, 524 (type species Reniera jrondiculata Schmidt, 1864: 39 (by
arborescent or occasionally fistulose growth fonns. Ectosomal monotypy) fragment of holotype BMNH 1910.1.1.542). Ramoses
skeleton composed of monactinal auxiliary spicules in one or two de Laubenfels, 1936a: 109 (type species Clathria pauper
categories forming structures ranging from sparse, mostly mem- Brondsted, 1927: 3 (by original designation and monotypy) schizo-
braneous (subgenus Microciona), sparse, paratangential (subgenus type BMNH 1930.11.5.2). Thalyseurypon de Laubenfels, 1936a:
Clathria) to a dense erect palisade of brushes on the surface (sub- 107 (type species Spongia raphanus Lamarck, 1814: 444 (by orig-
genus Thalysias). Choanosomal skeletal tracts usually enclosed inal designation) holotype MNHN DT572). Taxonomic decision
within spongin fibres, sometimes simply with nodal spongin; fibres for synonymy: Hooper (1996a), and this work.
cored by smooth, basally spined or partially spined principal mon-
actinal megascleres, usually geometrically different from auxiliary Type species
megascleres, sometimes secondarily lost and cored by single cate-
gory of auxiliary subtylostyle (subgenus Dendrocia), or sometimes Clathria compressa Schmidt, 1862: 58 (by subsequent desig-
replaced partially or fully by detritus in fibres (subgenus nation; Schmidt, 1864: 35).
WilsoneUa). Echinating megascleres partially or entirely acan-
those, occasionally smooth or vestigial spination, sometimes sec- Definition
ondarily lost (subgenera Axosuberites, [sociella). Choanosomal
structure ranges from hymedesmioid, leptoclathrid to microcionid Clathria with only a single category of auxiliary style
plumose (subgenus Microciona), renieroid (subgenus [sociella), fonning a sparse paratangential ectosomal skeleton; choanosome
plumo-reticulate or reticulate, with (subgenus Axosuberites) or without marked difference between axial and extra-axial
without compressed axis and radial extra-axial regions. regions.
Microscleres include palmate isochelae and modified fonns, and
toxas with smooth or spined points, occasionally absent. Diagnosis

Remarks Predominantly massive, lamellate, digitate, flabellate or

arborescent growth fonns; ectosomal skeleton composed of a sin-
The above definition is necessarily broad to encompass the gle undifferentiated category of auxiliary megasclere; choanoso-
seven subgenera included in Clathria. These characters span a mal skeletal structure plumo-reticulate or reticulate, usually without
wide spectrum of states, most of which are interpreted as second- marked difference between axial and extra-axial regions; spongin
ary losses rather than unique apomorphies, and many characters fibres cored by completely smooth, basally spined or partially
show intennediate states making it virtually impossible to maintain spined principal megascleres, geometrically differentiated from
strict generic boundaries recognised by earlier authors. auxiliary megascleres, but sometimes secondarily lost; echinating
megascleres entirely or partially acanthose, occasionally smooth,
Distribution sometimes secondarily lost. Microscleres include palmate
isochelae and modified fonns, and toxas with smooth or spined
Worldwide, predominantly shallow waters. points.
438 Porifera· Demospongiae • PoeciloscIerida • Microcionina • Microcionidae

c
B

A
Fig. 4. Clathria (Clathria). A-I, C. (C.) compressa Schmidt. A, choanosomal principal style. B, subectosomal auxiliary subtylostyle. C, echinating
acanthostyle (scale A-C, 50 f.lm). D, palmate isochelae. E, spined accolada toxa (scale D-E, 10 f.lm). F, fragment of holotype BMNH 1867.7.2.6.78 (scale
30mm). G, ectosomal skeleton (scale 150 f.lm). H, choanosomal skeleton (scale 150 f.lm). I, fibre structure (scale 50 f.lm) .

Description of type species (75-116 X 6-9/-Lm). Microscleres small palmate isochelae

(6-11 /-Lm) and forceps-shaped or accolada toxas with spinose
Clathria compressa Schmidt, 1862 (Fig. 4). extremities (32-148 X 1.5-4/-Lm).
Synonymy. Clathria compressa Schmidt, 1862: 8; Thalysias Remarks. Of the 160 named species originally described
compressa; de Laubenfels, 1936a: 105. in, or subsequently referred to Clathria (Clathria), or one of its
Material examined. Holotype: LMJG 15509 (schizotypes synonyms listed above, 115 are thought to be valid and most appro-
BMNH 1867.7.26.78, BMNH 1910.1.1.2362, 2363) - Adriatic. priately placed in this subgenus (the others being junior synonyms
Refer to Hooper (1996a) for full list of other type material examined. of established species or referred to other families).
Description. Erect, arborescent, thinly lamellate, branching Pitalia Gray is a new synonym included here in Clathria
growth form. Surface even, not hispid. Choanosomal skeleton (Clathria), overlooked by Hooper (1996a). Paresperia Burton also
regularly reticulate, with well developed spongin fibres forming belongs to Clathria, as originally suspected by Burton (1930c).
regular or irregular anastomoses of differentiated primary and Van Soest & Stone (1986: 45) suggested it was closer to
secondary spongin fibres. Fibres cored by choanosomal principal Esperiopsis (? Mycalidae), and was followed by Hooper (1996a:
subtylostyles in muItispicular ascending tracts and uni- or bispicu- 86) in this determination. Van Soest et at. (2000) returned it to
lar transverse connecting tracts, and echinated by acanthostyles Clathria, although again with question, and re-examination of type
perpendicular to or at acute angles to spongin fibres. Ectosomal material in this work supports its allocation to Microcionidae rather
skeleton with tangential layer of subectosomal auxiliary subty- than to Mycalidae. The species is poorly known only from a single
lostyles, of a single size category. Megascleres basally spined record from Norway, and type material is imperfect (holotype
choanosomal principal subtylostyles (220-345 X 8-14/-Lm), slides BMNH 1910.1.1.912, 2606-7). Re-examination
entirely smooth subectosomal auxiliary subtylostyles (164-235 X of these slides by Hooper (1996a: 86) failed to adequately fix
3-6/-Lm), and echinating acanthostyles with even spination the affinities of this species. The choanosomal skeleton is loosely
Porifera· Demospongiae • Poecilosclerida • Microcionina • Microcionidae 439

reticulate, formed by smaller acanthose styles (principal spicules), Synonymy. Wilsonella australiensis Carter, 1885d: 366;
bearing an even covering of minute spines (330 X 9/-Lm), and Clathria australiensis; Dendy, 1896: 33; Clathria (Wilsonella)
larger subtylostyles with microspined bases (auxiliary spicules) australiensis; Hooper, 1996a: 183, figs 83-84. Clathria australien-
(510 X 8 /-Lm) also scattered throughout the choanosomal skeleton, sis var. spinulata Hentschel, 1911: 374; Clathriopsamma lobosa
although this skeleton is not obviously divisible into primary or Lendenfeld, 1888: 149; Thorecta ramsayii Lendenfeld, 1888: 149;
secondary tracts as seen in many other Clathria. Echinating acan- Sigmatella corticata var. elegans Lendenfeld, 1888: 199 (Not
thostyles are absent. Both principal and auxiliary megascleres form Clathria australiensis; Levi, 1967a: 22. Not Ophlitaspongia aus-
a loose tangential ectosomal reticulation, and microscleres consist traliensis Ridley, 1884a: 442. Not Echinochalina australiensis;
only of palmate isochelae (24/-Lm long). Assigning this species Thiele, 1903a: 961).
accurately to a subgenus is currently impossible given the poor Material examined. Lectotype: BMNH 1886.12.15.43 -
quality of type material. Its lack of echinating acanthostyles indi- Port Phillip, Victoria, Australia. Other material. Refer to Hooper
cates possible affinities to Clathria (Isociella) and C. (Axociella), (1996a: 183).
but both of these subgenera have distinctive skeletal structure Description. Shape lobate, lobate-digitate, club-shaped,
(isodictyallrenieroid reticulate, and axially compressed, extra-axial thickly lamellate, thickly encrusting-bulbous or rarely fistulose
radial to plumo-reticulate, respectively), and neither has spinose growth forms; large oscules on apical or lateral margins of surface
principal spicules. Isociella is also exclusively Gondwanan in dis- lobes; surface arenaceous; ectosome membraneous, without spe-
tribution. Paresperia is, by default, left in Clathria (Clathria), and cialized dermal megascleres; choanosomal skeleton irregularly
for the time being is considered a greatly reduced form of this reticulate, with clearly differentiated primary and secondary spon-
genus (lacking acanthostyles and any definite skeletal structure). gin fibres, forming a vaguely longitudinal reticulation with cav-
ernous meshes; primary ascending fibres producing ascending
lines abundantly cored by detritus, lightly cored by auxiliary styles,
SUBGENUS WILSONELLA CARTER, 1885 heavily echinated by acanthostyles, particularly at fibre nodes;
smaller secondary spongin fibres mainly transverse, connecting
Synonymy with primary elements, with no or little detritus, paucispicular
tracts of auxiliary styles and lightly echinated by acanthostyles;
Wilsonella Carter, 1885d: 366 (Not Hallmann, 1912: 242). auxiliary megascleres coring fibres occupy only a small proportion
Clathriopsamma Lendenfeld, 1888: 227 (type species of fibre diameter; megascleres are choanosomal auxiliary styles
Clathriopsamma reticulata Lendenfeld, 1888:227 (by subsequent coring fibres differ from subectosomal auxiliary styles; only in
designation; Hallmann, 1920: 771), lectotype AM G9135). Aulenella being slightly thicker and lacking characteristic apical microspines
Burton & Rao, 1932: 345 (type species Aulenella foraminifera of the latter; coring spicules relatively thin, straight, smooth,
Burton & Rao, 1932: 345 (by original designation) holotype 1M mostly hastate, with slightly subtylote and occasionally micro-
P1167/1). Psammotoxa de Laubenfels, 1936a: 99 (type species spined bases (62-152 X 2.5-4 /-Lm); subectosomal auxiliary styles,
Phoriospongia guettardi Topsent, 1933: 19 (by original designa- dispersed between fibres and in dermal skeleton, straight, usually
tion), lectotype and paralectotype MNHN DT532, 3398, respectively hastate, subtylote bases with microspines on both points and bases
(here designated». Taxonomic decision for synonymy: Hooper (92-152 X 2.5-4 /-Lm); echinating acanthostyles small, evenly spin-
(1996a), and this work. ose or with granular, vestigial spines, slightly subtylote bases,
fusiform points (49-68 X 2-4.5 /-Lm); microscleres are palmate
Type species isochelae relatively large, unmodified (11-18 /-Lm); toxas oxhorn,
uncommon, rare in some specimens, often forming trichodrag-
Wilsonella australiensis Carter, 1885d: 366 (by monotypy). mata, usually with wide, angular, central curves and slightly
reflexed points (35-89 X 0.5-1 /-Lm).
Definition Remarks. This group of arenaceous microcionids has
already become well known under the name of Clathriopsamma
Clathria with sand grains and foreign spicules partially or (e.g., Hooper, 1990b; Hooper & Levi, 1993a), whereas Hooper
completely replacing coring spicules inside fibres; coring spicules (1996a) noted that the inclusion of C. (Wilsonella) australiensis in
same or very similar geometry to auxiliary spicules located outside this group unfortunately means that the senior name Wilsonella
fibres; skeletal architecture reticulate. (1885) must take precedence over Clathriopsamma (1888).
Unlike Clathria (Dendrocia), in which there is only a single
Diagnosis category of coring and extra-fibre megasclere, most Clathria
(Wilsonella) have more than one form of auxiliary style, one coring
Lobate, clavulate, lamellate or bulbous growth forms; surface the fibres (choanosomal megascleres) and one outside of fibres
arenaceous, and ectosome either lacking any megascleres or with (subectosomal megascleres). In some cases (e.g., c. (W) aus-
sparsely scattered auxiliary styles; choanosomal skeleton reticulate, traliensis, C. (W) ensiae), these spicules are only slightly different
cavernous, with fibres cored by sand grains and detritus and varying in their geometry, although showing clear differences in their
quantities of principal spicules, and echinated by acanthostyles; spination; in others (e.g., c. (W) reticulata, C. (W) mixta) these
principal and auxiliary styles usually poorly differentiated in their spicules are quite different; whereas in one (c. (W) abrolhosensis)
geometry; microscleres include palmate isochelae and toxas. there is no apparent difference, and this is interpreted as a conver-
gence or subsequent loss of a spicule category. Clathria
Description of type species (Dendrocia) and Clathria (Wilsonella) can also be distinguished by
their skeletal architecture - being predominantly plumose in the
Clathria (Wilsonella) australiensis (Carter, 1885d) (Figs 2D, 5). former and reticulate in the latter.
440 Porifera' Demospongiae • Poecilosclerida • Microcionina • Microcionidae

t o .-

C--.,

'.
r

B
Fig. 5. Clathria (Wilsonella). A-L, C. (w.) australiensis (Carter). A, subectosomal auxiliary subtylosty1e. B, choanosomal auxiliary subtylostyle. C, echi·
nating acanthostyle. D, Oxhom toxa. E, palmate isochela (scale A-E, 25 /-Lm). F, holotype BMNH 1886.12.15.43 (scale 30 mm). G, choanosomal skeleton
(scale I mm). H, peripheral skeleton (scale 500 /-Lm). I, continuum in basal and apical spination on auxiliary subtylostyles (scale 5 /-Lm). J, echinating acan-
thostyle (scale 20 /-Lm). K, oxhom toxas (scale 20 /-Lm). L, palmate isochelae (scale 5 /-Lm). A-L, modified from Hooper (l996a).

Psammotoxa de Laubenfels, 1936a is also a synonym of SUBGENUS MICROCIONA BOWERBANK, 1862

Clathria (Wilsonella), previously overlooked by Hooper (1996a).
Its type species, Phoriospongia guettardi Topsent was a new name Synonymy
proposed for two specimens erroneously identified as Spongia
carduus by Lamarck, 1814. The lectotype re-examined is of massive Microciona Bowerbank, 1862b: 1109. [Abila] Gray, 1867a:
flabellate growth form, with well developed spongin fibres form- 539 (preocc.) (Not Abila Gray, 1867a: 522) (type species
ing a nearly regular reticulation of primary (ascending) and sec- Microciona laevis Bowerbank, 1866:124 (by monotypy) holotype
ondary (connecting) tracts, partially cored by foreign detritus and BMNH 1877.5.21.1543). [Aaata] de Laubenfels, 1930: 27 (pre-
multispicular tracts of smooth styles, the latter confined mainly to occ.). Anaata de Laubenfels, 1932: 89 (replacement name for
the axis of the fibre, and echinated sparsely by acanthostyles. The [Aaata] de Laubenfels) (type species Aaata spongigartina de
ectosomal skeleton was not observed (dry material, from which the Laubenfels, 1930: 27 (by original designation) holotype USNM
surface is abraded). Microscleres consist of two forms of toxas, 21428). Axocielita de Laubenfels, 1936a: 118 (type species
long and sinuous ones that form dragmata, and smaller wing- Microciona similis Stephens, 1915b: 441 (by original designation)
shaped forms. holotype RSME 1921.143.1447). Fisherispongia de Laubenfels,
Porifera· Demospongiae • Poecilosclerida • Microcionina • Microcionidae 441

1936b: 460 (type species Fisherispongia ferrea de Laubenfels, 1866b: 149; Schmidt, 1866a: 15; cf. Microciona prolifera;
1936b: 460 (by original designation) holotype USNM 22239). Vosmaer, 1935: 604.
Holorodesmia Topsent, 1928c (type species Hymedesmiaflaccida Material examined. Holotype: BMNH 1930.7.3.225 - NE
Topsent, 1927b (by monotypy». Hymantho Burton, 1930c: 503 Atlantic. Refer to Hooper (1996a) for full list of other type material
(type species Hymantho normani Burton, 1930c: 503 (by original examined.
designation) holotype BMNH 1910.1.1.791). Hymeraphia, in part, Description. Encrusting growth form. Surface hispid,
Hentschel, 1912: 377 (not Hymeraphia Bowerbank, 1864: 189). uneven. Choanosomal skeleton hymedesmioid, with spongin fibres
Leptoclathria Topsent, 1928c: 298 (type species Leptoclathria reduced to basal layer lying on substrate, bearing erect, non-
haplotoxa Topsent, 1928c: 298 (by monotypy) holotype MNHN anastomosing, rarely branching, scattered fibre nodes perpendicular
DT1101). Ophistospongia Gray, 1867a: 514 (type species to substrate ('microcionid' fibres), each cored by plumose ascend-
Ophistospongia australis Gray, 1867a (by monotypy), from ing columns of choanosoma1 principal subtylostyles, wholly or
Bowerbank, 1864: pI. 17, fig. 288; holotype unknown). partly embedded in fibres, with points of spicules usually project-
Paratenaciella Vacelet & Vasseur, 1971: 103 (type species ing through ectosome. Echinating acanthostyles also erect on fibre
Paratenaciella microxea Vacelet & Vasseur, 1971: 103 (by original nodes. Subectomal skeleton with tangential layer of subectosomal
designation) holotype MNHN DN27). Pseudanchinoe Burton, auxiliary subtylostyles, singly or in bundles on surface. Ectosomal
1929a: 433 (type species Stylostichon toxiferum Topsent, 1913b: skeleton without specialized spiculation, but choanosomal and
621 (by original designation and monotypy) holotype MNHN subectosomal spicules protude through surface. Megascleres
DT1612). Sophax Gray, 1867a: 521 (type species Microciona choanosomal principal subtylostyles with smooth or microspined
fallax Bowerbank, 1866: 128 (by monotypy) lectotype BMNH bases (102-385 X 11-22/-Lm), evenly spined echinating acan-
1910.1.1.71). Wetmoreus de Laubenfels, 1936a: 112 (type species thostyles (62-87 X 4-9/-Lm), and smooth or basally spined subec-
Microciona novaezealandica Brondsted, 1924b: 463 (by original tosomal auxiliary subtylostyles (122-295 X 2-8 /-Lm). Microscleres
designation) holotype UZM not found). Taxonomic decision for palmate isochelae (uncommon) (12-17 /-Lm) and oxhom to wing-
synonymy: Hooper (1996a), and this work. shaped toxas (42-110 X 1.5-4/-Lm).
Remarks. Of 118 named species originally described in, or
Type species subsequently referred to Microciona or one of its synonyms listed
above, 103 appear to be valid taxa (the others being junior syn-
Microciona atrasanguinea Bowerbank, 1862b (by subsequent onyms of established species) (Hooper, 1996a). Microciona (s.s.)
designation; Bowerbank, 1864: 188). differs from Clathria (s.s.) in having an encrusting growth form, a
reduced hymedesmioid skeleton with erect fibre nodes cored by
Definition plumose tracts of principal and echinating spicules standing erect
on the substrate (='microcionid' architecture of Levi, 1960b), and
Clathria with persistently encrusting growth form, with smooth toxas. The critical difference between these genera,
hymedesmioid skeletal architecture consisting of a basal layer of therefore, is the possession of the plumose, non-anastomosing
spongin, typically with ascending, plumose, non-anastomosing fibre nodes, whereas other encrusting genera in the family have
spongin fibre nodes, and megascleres embedded and erect on basal simply hymedesmioid skeletal construction (e.g., Leptoclathria,
layer. Anaata, Hymantho), a basal renieroid reticulation (e.g., some
Antho, Plocamilla), or an isodictyal fibre skeleton arising from
Diagnosis a hymedesmioid base (Ophlitaspongia). By comparison, erect or
massive genera of Microcionidae commonly have reticulate or
Thinly or thickly encrusting growth forms; ectosomal plumo-reticulate skeletons (e.g., Clathria, Holopsamma), renieroid
skeleton with a single undifferentiated category of auxiliary or subisodictyal skeletons (e.g., Isociella, Isopenectya, Pandaros),
styles sparsely dispersed on or near surface, lying paratangential or virtually halichondroid skeletal architecture (e.g., Artemisina).
or tangential to surface; choanosomal skeleton essentially There are many contemporary authors who maintain this distinc-
hymedesmioid, with a basal layer of spongin bearing echinating tion between Microciona and erect non-plumose genera, such as
and principal styles embedded in and perpendicular to it, forming Clathria at the generic level (e.g., Levi, 1969; Wiedenmayer,
single or plumose ascending columns, and in the latter case having 1977b; Pulitzer-Finali, 1983; Uriz, 1984a,b; Wintermann-Kilian &
spicule bundles partially enclosed by ascending, plumose, non- Kilian, 1984; Boury-Esnault & Lopes, 1985; Bergquist & Fromont,
anastomosing spongin fibres (fibre nodes); principal styles fre- 1988), whereas Van Soest (1984b) and Hooper (1996a) recognise
quently with prominent basal tyle and/or with basal spination; the taxon only at the subgenus level because this particular skeletal
microscleres palmate isoche1ae and various toxa morphologies. architecture is clearly linked to some extent by the ontogeny of
the sponge individual (Simpson, 1968), and probably largely a
Description of type species phenotypic response to prevailing ecological conditions. Hooper
(1996a) included Seriatula Gray (1867a: 515) (type species
Clathria (Microciona) atrasanguinea (Bowerbank, 1862b) Spongia seriata Grant, 1826c: 116 (by monotypy) holotype
(Fig. 6). BMNH 1847.9.7.14) into synonymy with Ophlitaspongia
Synonymy. Microciona atrasanguinea; Bowerbank, Bowerbank, and hence into synonymy with Microciona (follow-
1862a: 824; Microciona atrosanguinea; Gray, 1867a: 535; ing Simpson, 1968), but this was shown to be an incorrect assump-
Microciona atrasanguineum; Cuenot, 1903: 4; Clathria tion whereby S. seriata is identical to Halichondria panicea
(Microciona) atrasanguinea; Van Soest, 1993b: 103; Amphilectus Pallas (Howson & Chambers, 1999). Differences between
atrasanguineus; Vosmaer, 1880: 115; Plumohalichondria atrasan- Microciona and other genera are discussed below (see
guinea; Hanitsch, 1890: 207; Scopalina atrosanguinea; Schmidt, Ophlitaspongia).
442 Porifera' Demospongiae • Poecilosclerida • Microcionina • Microcionidae

Fig. 6. Clathria (Microciona). A-G, C. (M.) atrasanguinea (Bowerbank). A, choanosomal principal subtylostyles (scale 100 !Lm). B, echinating acan-
thostyles (scale 25!Lm). C, subectosomal auxiliary subtylostyles (scale 100 !Lm). D, oxhorn and wing-shaped toxas (scale 50 !Lm). E, palmate isochela
(scale 5!Lm). F, skeletal structure (scale 200 !Lm). G, reconstruction of skeleton.

SUBGENUS DENDROCIA HALLMANN, 1920 Diagnosis

Synonymy Massive, lobate, globular, digitate and flabellate growth

forms; with a single undifferentiated category of smooth auxiliary
Dendrocia Hallmann, 1920: 767. Paradoryx Hallmann, 1920: spicule (style, subtylostyle or modified style) forming plumose
767 (type species Clathria dura Whitelegge, 1901: 83 (by original or plumo-reticulate choanosomal tracts, ectosomal brushes and
designation) holotype AM G3046). Wilsonella in part; sensu also dispersed between skeletal tracts. Echinating acanthostyles
Hallmann, 1912: 242 (not Carter, 1885d: 366). Taxonomic decision usually heavily spined and distributed evenly over skeletal tracts.
for synonymy: Hooper (1996a). Microscleres include isochelae ranging from typical palmate form
(straight shaft, lateral alae fused to shaft), modified palmate
Type species forms (thickened, curved shaft, partially detached lateral alae) to
'anchorate-like' forms (alae detached from shaft, shaft with lateral
Clathria pyramida Lendenfeld, 1888: 222 (by original ridge); toxas if present include oxhorns.
designation).
Description of type species
Definition
Clathria (Dendrocia) pyramida Lendenfeld, 1888 (Figs 2C, 7).
Clathria with only a single undifferentiated category of Synonymy. Clathria pyramida Lendenfeld, 1888: 222;
structural megasclere (auxiliary styles) forming choanosomal and Wilsonella pyramida; Hallmann, 1912: 240; Dendrocia pyramida;
ectosomal skeletons. Hallmann, 1920: 767; Clathria (Wilsonella) pyramida; Hooper,
Porifera· Demospongiae • Poecilosclerida • Microcionina • Microcionidae 443

Fig. 7. Clathria (Dendrocia). A-I, C. (D.) pyramida Lendenfeld. A, echinating acanthostyle. B, subectosomal auxiliary subtylostyle. C, palmate isochelae
(scale A-C, 50 jJom). D, paraIectotype of C. alata NMVG2283 (scale 30 mm). E, section through choanosomal skeleton (scale 500 jJom). F, peripheral skeleton
(scale 1 mm). G, fibre characteristic (scale 100 jJom). H, echinating acanthostyles and pattern of spination (scales L-R, 20, 50, 10 jJom respectively).
I, palmate isochela (scale 10 jJom). A-I, modified from Hooper (l996a).

1996a: 242; Clathria alata Dendy, 1896: 34; Wilsonella alata auxiliary spicules; subectosomal auxiliary styles thin, hastate,
Hallmann, 1912: 241. Dendrocia alata Hallmann, 1920: 767. straight or slightly curved towards basal end, usually with smooth,
Material examined. Lectotype: AMG9047 - Port Jackson, slightly subtylote bases that taper towards end into a small point
NSW, Australia. Other material. Refer to Hooper (l996a: 242). (partially mucronate), points hastate (208-321 X 4-8 /-Lm); acan-
Description. Shape massive, lobate, lobo-digitate, irregular tho styles club-shaped, subtylote, with large spines mostly confined
growth form with small lobate surface projections; large oscules on on base and more-or-less aspinose towards point; spines usually
apex of lobes, each with slightly raised membraneous lip; smooth, robust (87-135 X 8-12 /-Lm); microscleres isochelae large, very
relatively even, bulbous, fleshy surface; ectosome microscopically abundant, primarily palmate, heavily silicified, usually with thick-
hispid with dense crust of subectosomal auxiliary styles forming ened and slightly curved shaft, large lateral alae completely fused
continuous, erect, regular (straight) or irregular (stellate, paratan- to shaft resembling 'wings', with front ala free, but sometimes with
gential), plumose palisade; choanosomal skeleton irregularly reduced alae and sigmoid curvature (21-27 /-Lm); toxas absent.
plumo-reticulate, with sinuous spongin fibres ascending to surface Remarks. Seven species are included in Clathria
in meandering tracts; fibres anastomose more frequently at axis (Dendrocia), all of which are endemic to temperate Australian
than in peripheral skeleton; peripheral fibres and skeletal tracts waters, with hypothesised Gondwanan origins.
often diverge becoming plumose in subectosomal region, or form a Dendrocia, like Wilsonella, differs from other microcionids in
paratangential layer immediately below erect ectosomal skeleton; having auxiliary styles both coring fibres and forming the ectoso-
fibres cored by muitispicular, sinuous tracts of subectosomal auxil- mal skeleton. However, whereas Wilsonella has two categories of
iary styles; acanthostyles heavily echinate most fibres; choanoso- auxiliary spicules and detritus is incorporated into the skeleton,
mal principal megascleres absent, or at least undifferentiated from Dendrocia has only one category of structural spicule throughout
444 Porifera· Demospongiae • Poecilosclerida • Microcionina • Microcionidae

Fig.8. Clathria (Axosuberites). A-K, C. (A.) cylindrica (Ridley & Dendy). A, choanosomal principal subtylostyle (coring fibres). B, subectosomal auxil-
iary style (coring extra-axial skeleton). C, ectosomal auxiliary subtylostyles (scale A-C, 100 J.Lm). D, oxhom toxa. E, palmate isochelae (scale D-E,
25 J.Lm). F, holotype BMNH 1887.5.2.96 (scale 30 mm). G, choanosomal skeleton (scale 1 mm). H, fibre characteristic (scale 100 J.Lm). I, cross-section
through branch (scale 1 mm). J, palmate isochelae (scale 10 J.Lm). K , oxhom toxas (scale 20 J.Lm). A-K, modified from Hooper (l996a).

the skeleton. Dendrocia also has a characteristic dendritic or Tenaciella Hallmann, 1920: 772 (type species Esperiopsis canalic-
plumo-reticulate skeletal architecture, whereas Wilsonella is invari- ulata Whitelegge, 1906: 471 (by monotypy) lectotype AM G4325).
ably reticulate. In having only one geometric form of auxiliary Taxonomic decision for synonymy: Hooper (1996a), and this work.
spicule producing the extra-fibre skeleton (i.e., subectosome and
ectosomal regions) the genus has closest affinities with Clathria, Type species
whereas in skeletal structure (with a continuous ectosomal palisade
of spicules) the genus resembles the Thalysias condition. For this Axosuberites fauroti Topsent, 1893a: 179 (by monotypy).
reason Dendrocia is enigmatic, and Hooper (1990b) maintained it
as a separate taxon, whereas earlier Van Soest (1984b) had indi- Definition
cated that it was probably a synonym of Clathria.
Clathria with axial skeleton compressed and well differentiated
from plumose(-reticulate) extra-axial skeleton; echinating spicules
SUBGENUS AXOSUBERITES TOPSENT, 1893 absent; ectosomal skeleton with special category of (smaller)
auxiliary style forming surface brushes.
Synonymy
Diagnosis
Axosuberites Topsent. 1893a: 179. Axociella Hallmann. 1920:
779 (type species Esperiopsis cylindrica Ridley & Dendy, 1886: Digitate. whip-like. arborescent and flabellate growth forms;
340 (by original designation) holotype BMNH 1887.5.2.96). with well differentiated axial and extra-axial skeletal architecture,
Porifera· Demospongiae • Poecilosclerida • Microcionina • Microcionidae 445

reminiscent of Raspailiidae; axial skeleton markedly compressed, fibres but very few transverse uni- or aspicular connecting fibres;
reticulate; extra-axial skeleton radial, plumose or plumo-reticulate megascleres include choanosomal principal subtylostyles coring
composed of large subectosomal auxiliary styles-subtylostyles; axial fibres relatively short, slightly curved at centre, with smooth
specialised ectosomal skeleton present composed of smaller auxil- subtylote or sometimes evenly rounded bases and fusiform points
iary spicules; echinating megascleres absent although principal (215-395 X 5-15 /-Lm); subectosomal auxiliary styles forming
spicules may protrude through fibres at acute angles; microscleres extra-axial bundles much longer and thicker than principal
absent or may include palmate isochelae and toxas. spicules, with smooth rounded or tapering (hastate) bases, and
fusiform points (424--725 X 15-29 /-Lm); ectosomal auxiliary sub-
Description of type species tylostyles straight, variable length, with smooth rounded or slightly
subtylote bases and fusiform points (208-575 X 6-10 /-Lm);
Axosuberitesfauroti Topsent, 1893a: 179 (by monotypy) (Not microscleres palmate isochelae unmodified, differentiated into two
figured; referto Hooper, 1996a: fig. l1c-d). size classes, with long lateral alae entirely fused to shaft and com-
Synonymy. Axosuberites fauroti Topsent, 1893a: 179-181, pletely fused front ala (6-13,19-25 /-Lm); toxas oxhom, thick, with
fig. 3. ? Rhaphidophlus fauroti; Van Soest, 1984b: 130. Clathria wide central curvature and slightly reflexed points (45-130 X
(Axociella)fauroti; Hooper, 1996a: 247. 2.5-6/-Lm).
Material examined. Holotype: MNHN DTl859 (fragment)- Remarks. The inclusion of Axosuberites into synonymy
Tadjoura, Gulf of Aden, Arabian Gulf. with Axociella (Van Soest, 1984b; Hooper, 1996a) unfortunately
Description. Flabellate, flattened digitate growth form. requires that the former name takes precedence, based on seniority,
Surface hispid, conulose. Choanosome with compressed reticulate a fact overlooked by Hooper (1996a). The type species of both
axis and plumose extra-axial skeleton, with only light spongin nominal genera are redescribed here whereas the latter is better
fibres. Axial fibres produce close-meshed reticulation of multi- known from the recent literature.
spicular tracts cored by choanosomal principal subtylostyles form- The existence of microcionids with compressed axial skeletons
ing criss-cross reticulation, tracts plumose near periphery. and differentiated axial and extra-axial skeletons supports the pro-
Echinating megascleres absent. Subectosomal extra-axial skeleton posal for a closer relationship between Microcionidae and
well differentiated from axial region, with ascending plumose Raspailiidae, as proposed by Hooper (1990b, 1991), and formalised
columns of larger subectosomal auxiliary subtylostyles arising further by Hajdu et al. (1994a) in the subordinal classification of
from peripheral choanosomal skeleton. Ectosome with brushes of Poecilosclerida as suborder Microcionina. In particular, Axociella is
smaller auxiliary subtylostyles overlaying larger subectosomal most reminiscent of Raspailia (Syringella) and Ectyoplasia. Crucial
spicules. Megascleres entirely smooth choanosomal prin- characters differentiating the Axosuberites group from typical
cipal subtylostyles-tylostyles, and two size classes of auxiliary Raspailiidae are the possession of chelae and absence any true echi-
subtylostyles-tylostyles, both with smooth bases. Microscleres nating spicules in the former, versus possession of a specialized
absent. ectosomal skeleton (composed of small styles or oxeas in brushes
Description of type species ofAxociella. surrounding larger protruding spicules) in the latter (see Hooper,
Clathria (Axociella) cylindrica (Ridley & Dendy, 1886) (Fig. 8). 1991, 1996a). Despite contrary arguments by Van Soest (1984b),
Synonymy. Esperiopsis cylindrica Ridley & Dendy, 1886: Axosuberites is considered here to be sufficiently different from
340; Axociella cylindrica Hallmann, 1920: 780 (Not Axociella Thalysias in its skeletal construction to be differentiated at the
cylindrica; Sim & Byeon, 1989: 39); Clathria cylindrica; Hooper supraspecific level (although not for the reason that it lacks echinat-
& Wiedenmayer, 1994: 262. Clathria (Axociella) cylindrica; ing acanthostyles, as suggested by Simpson, 1968).
Hooper, 1996a: 251, figs 120--121. (Not Rhaphidophlus cylindri-
cus Kieschnick, 1900: 53).
Material examined. Holotype: BMNH 1887.5.2.96 - off
Port Jackson, NSW, Australia. Other material. Refer to Hooper SUBGENUS ISOCIELLA HALLMANN, 1920
(1996a: 251).
Description. Shape thin, cylindrical digitate, branching, Synonymy
whip-like, with thin, dichotomously branched, cylindrical or
slightly flattened, distally tapering branches; firm, flexible, tough Isociella Hallmann, 1920: 784.
consistency; surface even, felt-like, unomamented, prominently
hispid; ectosome with sparse, plumose brushes of small ectosomal Type species
auxiliary subtylostyles forming discrete, discontinuous bundles on
surface, arising from ends of radial skeletal columns, scattered Phakellia jlabellata, in part (sensu Ridley & Dendy, 1886:
around larger, protruding subectosomal auxiliary styles which 478) (by monotypy) (Not P. jlabellata Carter, 1885d: 363).
project some distance through surface; choanosomal skeletal struc- Synonymous with Phakellia jacksoniana (Dendy, 1897: 236)
ture with clearly differentiated axial and extra-axial components, (holotype BMNH 1887.5.2.9), both junior synonyms of Clathria
markedly compressed in axis and radial in extra-axis; compressed macropora, in part, Lendenfeld, 1888: 221 (holotype AMZ466).
axial skeleton with longitudinal fibres cored by short choanosomal Taxonomic decision for synonymy: Hooper (1996a).
principal subtylostyles, interconnected by thinner pauci- or aspicu-
lar fibres; axial fibres heavily collagenous, and fibre anastomoses Definition
very close-meshed forming elongate reticulation; echinating
megasc1eres absent; radial extra-axial skeleton with large auxiliary Clathria with evenly renieroid reticulate choanosomal skeleton,
styles perpendicular to axis, forming pauci- or multispicular radial with spongin fibres cored by entirely smooth principal styles;
(non-plumose) tracts associated with very light, ascending spongin echinating megascleres absent.
446 Porifera' Demospongiae • Poecilosclerida • Microcionina • Microcionidae

I
'H

Fig. 9. Clathria (lsociella). A-H, C. (I.) macropora Lendenfeld. A, choanosomal principal styles. B, subectosomal auxiliary subtylostyles (scale A-B ,
100 fLm). C, modified palmate isochelae (scale 25!Lm). D, paralectotype of Phakellia jacksoniana BMNH 1887.5.2.8 (scale 30mm). E, choanosomal
skeleton (scale 1 mm). F, semi-renieroid fibres (scale 500 !Lm). G, bases of subectosomal auxiliary styles (scale 10 fLm). H, modified palmate isochelae
(scale 5!Lm). A-H, modified from Hooper (l996a).

Diagnosis Synonymy. Phakellia jlabellata Ridley & Dendy, 1886: 478

(Not Phakelliaflabellata Carter, 1885d: 363); Clathria macropora,
Massive, bulbous, lobate, digitate and flabellate growth in part, Lendenfeld, 1888: 221. Clathria (lsociella) macropora;
forms; skeletal structure with relatively homogeneous renieroid or Hooper, 1996a: 271. (Not Plectispa macropora Lendenfeld, 1888:
subrenieroid, wide-meshed reticulation; main skeleton with primary 226. Not Clathria macropora Whitelegge, 1901: 91. Not Wilsonella
plumose ascending, multi spicular tracts cored by smooth macropora; Hallmann, 1912: 203. Not Plumohalichondria australis
choanosomal styles, interconnected by secondary, uni- or pau- Whitelegge, 1901: 90); lsociellajlabellata Hallmann, 1920: 784;
cispicular tracts cored by same spicules, and choanosomal spicules Phakellia jacksoniana Dendy, 1897: 236; lsociella jacksoniana;
sometimes diverging and forming plumose brushes at surface; Bergquist & Tizard, 1967: 187.
echinating megascleres absent; ectosomal skeleton with single cat- Material examined. Holotype: AM Z466 - Port Stephens,
egory of auxiliary spicule, tangential, paratangential. Microscleres NSW, Australia. Other material. Refer to Hooper (1996a: 271).
palmate-derived isochelae and toxas. Description. Shape irregularly flabellate-digitate or flabellate,
planar, with short cylindrical stalk, one or more thinly lobate, bifur-
Description of type species cated branches, with rounded, digitate, uneven or shaggy margins;
oscules small, in special areas (sieve-plates) scattered over surface
Clathria (lsociella) macropora Lendenfeld, 1888 (Fig. 9). of branches, with series of stellate subdermal drainage canals
Porifera· Demospongiae • Poecilosclerida • Microcionina • Microcionidae 447

surrounding each osculum; branches with separate inhalant and Esper, 1797: 195 (by original designation; Ehlers, 1870: 18)
exhalant faces, one porous, rugose, with irregular longitudinal schizotypes 2MB 4577). Tenaeia Schmidt, 1870: 56 (type species
ridges, microconules or irregular striations, other smooth, membra- Tenaeia clathrata Schmidt, 1870: 56 (by monotypy) (schizotype
neous; ectosome membraneous, hispid, with points of choanoso- BMNH 1870.5.3.156); junior synonym of Spongia virgultosa
mal principal styles protruding through surface, individually or in Lamarck, 1814: 444 (fragments of holotype MNHN DNBE1344,
multispicular plumose bundles; surface skeleton with relatively 1338». [Echinonema] Carter, 1875c: 194 (nomen nudum); Carter,
sparse tangential, paratangential or sometimes plumose erect skele- 1881b: 378 (type species Eehinonema typieum Carter, 1881b: 377
ton of small subectosomal auxiliary styles projecting between prin- (by typonymy) (lectotype BMNH 1877.5.21.149); junior synonym
cipal spicules, sometimes surrounding (in proximity of) principal of Spongia eaetiformis Lamarck, 1814: 440 (lectotype MNHN
spicules reminiscent of Raspailiidae; choanosomal skeleton with DT580». Thalassodendron Lendenfeld, 1888: 222 (type species
very slightly compressed axis and plumose sub-isodictyal extra- Thalassodendron typiea Lendenfeld, 1888: 223 (by indication)
axis; axial region with moderately heavy spongin fibres, forming (holotype unknown». Stylotellopsis Thiele, 1905: 456 (type
tight irregularly reticulate meshes cored by paucispicular tracts of species Stylotellopsis amabilis Thiele, 1905: 456 (by monotypy)
choanosomal principal styles; axis (corresponding to centrallamel- holotype 2MB 3309). Colloclathria Dendy, 1922b: 74 (type
lae and basal stalk) has few ascending, primary tracts, forming species Colloclathria ramosa Dendy, 1922b: 74 (by monotypy)
multispicular, halichondroid structures, producing few multispicu- holotype BMNH 1921.11.7.64). Damoseni de Laubenfels, 1936a:
lar, dendritic tracts running from basal stalk to periphery; extra- 110 (type species Hymeraphia miehaelseni Hentschel, 1911: 351
axial skeleton with plumose spicule tracts bound by collagen (by original designation) fragment of holotype SMF969T).
(without fibre component), ascending to surface, cored by uni-, Taxonomic decision for synonymy: Hooper (1996a).
pauci- or less frequently multi spicular tracts of choanosomal
principal styles; primary ascending extra-axial spicule tracts per- Type species
pendicular to axis, interconnected by more-or-Iess transversely ori-
entated, smaller secondary uni- or paucispicular tracts producing Spongia virgultosa Lamarck, 1814 (by subsequent designation;
the predominantly subisodictyal structure; peripheral spicule tracts de Laubenfels, 1936a: 104) (holotype MNHN missing, schizotype
more plumose than deeper choanosomal tracts; meshes produced BMNH 1954.2.20.67); junior synonym of Spongia juniperina
by spicule-fibre anastomoses in extra-axial region rectangular or Lamarck, 1814 (lectotype MNHN DT570) (de Laubenfels,
triangular in shape; echinating acanthostyles absent; megascleres 1936a: 104).
choanosomal principal styles long or short, thick, slightly curved at
centre, less often straight, with rounded or slightly tapering, Definition
smooth bases, varying from fusiform to hastate points (369-552 X
21-35/-Lm); subectosomal auxiliary styles variable in length, thin, Clathria with two categories of auxiliary styles forming a
straight or very slightly curved, with basal terminations varying specialized ectosomal skeleton, the smaller usually forming dis-
from evenly rounded, tapering hastate, quasi-diactinal mucronate crete bundles or a continuous palisade perpendicular to the surface;
or slightly subtylote, and with hastate points (191-424 X 4-9/-Lm); echinating megascleres present.
microscleres palmate isochelae with highly modified, relatively
small alae bearing wing-shaped fluted processes; lateral alae Diagnosis
entirely fused to shaft; front ala complete or bifurcated with medial
tooth; chelae frequently twisted or occasionally anisochelate Thickly encrusting, massive, globular, digitate, flabellate and
(8-16 /-Lm); toxas absent. arborescent growth forms; with specialized ectosomal skeleton of
Remarks. Four species of Isociella are known for the differentiated size classes of auxiliary (subtylo)styles, with smaller
Australian fauna, three from the tropical coasts of WA, NT and Qld, ectosomal spicules usually overlaying larger subectosomal ones
and one temperate species from NSW. The type species is also known forming erect brushes in a continuous palisade, or discrete bundles,
from SE Indonesia, and only one other species is known from the New paratangential or rarely tangential to surface. Choanosomal skele-
Zealand fauna. Isociella has a slightly compressed axial skeleton par- ton without any marked differentiation between axial and extra-
tially offset from a diverging, plumose, subisodictyal reticulate extra- axial regions. Echinating acanthostyles usually present.
axial skeleton, showing vague structural similarities to Ceratopsion Microscleres are palmate isochelae and diverse forms of toxas.
and Raspailia (Syringella) (family Raspailiidae). This skeletal struc-
ture could also justify its inclusion in the subgenus Axosuberites, but it Description of type species
is considered here that the subisodictyal reticulation dominates the
skeleton and is more characteristic of Isociella than Axosuberites, Clathria (Thalysias)juniperina (Lamarck, 1814) (Fig. 10).
whereas in the latter subgenus the axial compression and differentia- Synonymy. Spongia juniperina Lamarck, 1814: 444;
tion between axis and extra-axis are predominant features. Clathriajuniperina; Hooper & Wiedenmayer, 1994: 270. Clathria
(Thalysias) juniperina; Hooper, 1996a: 346, figs 176-177 (Not
Pandaros juniperina; Duchassaing & Michelotti, 1864: 90. Not
SUBGENUS THALYSIAS DUCHASSAING & Thalysias juniperina de Laubenfels, 1936a: 105); Rhaphidophlus
MICHELOTTI, 1864 clathratus Hallmann, 1912: 209 (Not Tenacia clathrata Schmidt,
1870: 56).
Synonymy Material examined. Lectotype: MNHN DT570 - SW coast
of Australia. Other material. Refer to Hooper (1996a: 346).
Thalysias Duchassaing & Michelotti, 1864: 82. Description. Shape ranges from thickly encrusting to fron-
Rhaphidophlus Ehlers, 1870: 19 (type species Spongia eratitia dose, lamellate, clathrous, with or without free or anastomosing
448 Porifera· Demospongiae • Poecilosclerida • Microcionina • Microcionidae

;:
C~
B'
A
Fig. 10. Clathria (Thalysias). A-F, I-L, C. (T.) juniperiTUl (Lamarck). G-H, C. (T.) reinwardti Vosmaer. A, larger subectosomal auxiliary style. B, smaller ecto-
somal auxiliary style. C, choanosoma! principal style. D, echinating acanthostyle (scale A-D, 100 ILm). E, sinuous, accolada and u-shaped toxas. F, palmate
isochelae (scale E-F, 25 ILm). G, paralectotype MNHN DT3354. H, choansomal skeleton (scale 1 mm). I, echinated fibre (scale 200 ILm). J, echinating acan-
thostyle (scales L-R, 20 and 10 ILm). K, palmate isochelae (scale 5ILm). L, toxas (scales L-R, 10,2 and 20 ILm). A-L, modified from Hooper (1996a).

branches; surface relatively smooth, even, with white subdermal with multiple curves (sinuous), with smooth subtylote bases
canals in encrusting forms, or irregularly microconulose or (169-310 X 4-6.5 f.Lm); ectosomal auxiliary subtylostyles with
clathrous in more massive forms; ectosomal skeleton crust-like, prominent subtylote, smooth bases (93-110 X 2-4.5 f.Lm); acan-
easily detachable, relatively thin but dense palisade of erect or thostyles small, stubby, with rounded or only slightly subtylote
paratangential brushes supported by paratangential tracts of larger bases, with few spines and extensive aspinose regions on necks and
subectosomal auxiliary megascleres immediately below surface; points; spines large, bulbous, erect (45--65 X 5-8 f.Lm); microscle-
peripheral fibres immediately subectosomal with vaguely ascend- res palmate isochelae unmodified, incompletely differentiated into
ing multi spicular subectosomal tracts arising to surface; choanoso- two size classes; lateral alae entirely fused to shaft, approximately
mal skeleton irregularly reticulate, with very heavy spongin fibres same length as front ala and completely detached from front ala
forming oval meshes; fibres usually with paucispicular core of (6-9, 12-16.5 f.Lm); toxas very variable in length, mostly thin,
subectosomal auxiliary styles occupying only a small proportion of ranging from accolada forms with large central curvature and
fibre diameter, and fewer choanosomal principal styles which are slightly reflexed points, large curvature and simply u-shaped, to
entirely enclosed in, or project from fibres; in some cases fibres asymmetrical sinuous forms (55-180 X 0.8-1.5 f.Lm).
completely uncored, whereas others contain abundant, disorgan- ReTlUlrks. Of 137 named species described in, or referred to
ized auxiliary megascleres; fibres typically heavily echinated, Thalysias or one of its synonyms, 93 are thought to be valid.
some enveloping echinating megascleres entirely, some fibres Hooper (1996a) notes that the taxon Clathria juniperina as
without echinating megascleres; megascleres choanosomal princi- presently understood (e.g., Van Soest, 1984b) is probably com-
pal styles straight or slightly curved near basal end, with smooth, posite, incorporating two or more sibling species, with disjunct
rounded or very slightly subtylote bases (170-280 X 9-12 f.Lm); populations from the Caribbean (e.g., Hartman, 1955; Simpson,
subectosomal auxiliary subtylostyles straight or curved, sometimes 1968; Wiedenmayer, 1977b; Van Soest, 1984b), and Indian Ocean
Porifera· Demospongiae • Poecilosclerida • Microcionina • Microcionidae 449

(Lamarck, 1814, 1816) (i.e., of Spongia juniperina). The accepted remainder being either junior synonyms or more appro-
interpretation is that the nominotypical population from the Indo-west priately included in other genera), of which all live in the Indo-west
Pacific be known as C. (T.) juniperina whereas the West Indies popu- Pacific.
lation accept the most senior available name for several Caribbean
species presently included in synonymy with Spongia virgultosa.
The primary and only consistent morphological feature that SUBGENUS ECHINOCHAUNA TmELE, 1903
distinguishes Thalysias from other Clathria-like taxa is the
presence of a specialized ectosomal skeleton, consisting of two dif- Synonymy
ferentiated categories of auxiliary subtylostyles which form
brushes on the surface (either as discrete brushes (s.s.) or a con- Echinochalina Thiele, 1903a: 961. Tablis de Laubenfels,
tinuous palisade). It differs from Axociella in having echinating 1936a: 76 (type species Echinochalina anomala Hallmann, 1912:
acanthostyles. 292 (by original designation) holotype AM G 10548).
Echinoclathria sensu Uriz, 1988: 89 (Not Echinoclathria Carter,
1885d). Taxonomic decision for synonymy: Hooper (1996a).
ECHINOCHAUNA TIDELE, 1903
Type species
Synonymy
Ophlitaspongia australiensis Ridley, 1884a: 442 (by sub-
Echinochalina Thiele, 1903a: 961 (see subgenera for full sequent designation; Hallmann, 1912: 288).
synonymy).
Definition
Type species
Echinochalina with smooth monactinal, quasi-monactinal or
Ophlitaspongia australiensis Ridley, 1884a: 442 (by subse- secondarily derived true diactinal auxiliary megascleres coring
quent designation; Hallmann, 1912: 288). fibres, and smooth or spined monactinal principal spicules echinat-
ing fibres.
Definition
Diagnosis
Microcioninae with monactinal, quasi-monactinal or thin
diactinal auxiliary megascleres both coring spongin fibres and Thickly enrusting, massive, lobate, bulbous, clavulate,
forming extra-fibre and ectosomal tracts, and principal spicules branching, clathrous, honeycomb-reticulate, and digitate growth
(styles or oxeas) echinating fibres. forms; ectosomal skeleton, interstitial skeleton and spongin fibres
cored by smooth auxiliary megascleres (subtylostyles, tornostyles
Diagnosis or quasi-stongyles); fibres echinated by smooth or spined principal
styles; ectosomal skeleton with single or tracts of auxiliary megas-
Thickly encrusting, massive, lobate, bulbous, clavulate, cleres tangential or paratangential to the surface; choanosomal
clathrous, honeycomb-reticulate, tubulo-digitate, digitate and architecture reticulate with differentiated multi spicular primary
arborescent growth forms; with monactinal, quasi-monactinal or and paucispicular secondary tracts composed of auxiliary megas-
thin diactinal auxiliary megascleres tangential or erect on ecto- cleres; microscleres if present palmate isochelae or toxas.
some; choanosomal skeleton irregularly reticulate; fibres cored by
tracts of auxiliary megascleres, identical to those in ectosomal Description of type species
skeleton, and echinated by principal megascleres varying from true
monactinal, quasi-diactinal to true diactinal forms, smooth or acan- Echinochalina (Echinochalina) australiensis (Ridley, 1884a)
those; microscleres may include palmate isochelae and toxas. (Figs 21, 11).
Synonymy. Ophlitaspongia australiensis Ridley, 1884a:
Remarks 442. Echinochalina australiensis; Thiele, 1903a: 961.
Echinochalina (Echinochalina) australiensis; Hooper, 1996a: 518,
The genus contains two groups of species: one (Echinochalina figs 278-279.
(Echinochalina» with true monoactinal spicules, showing superfi- Material examined. Holotype: BMNH 1881.10.21.299 -
cial affinities with Holopsamma (having honeycomb reticulate Port Molle (Airlee), Qld, Australia. Other material. Refer to
growth forms) and the Raspailiidae (fibre characteristics), and the Hooper (1996a: 518).
other with quasi-diactinal or secondarily modified true diactinal Description. Shape massive, branching, clathrous,
megascleres (Echinochalina (Protophlitaspongia» superficially honeycomb-reticulate growth form, fibre-branches flattened, with
resembling Niphatidae (Haplosclerida). Both groups are linked by cavernous, angular meshes; oscules small situated between adja-
the common possession of auxiliary megascleres coring fibres and cent fibre bundles; conulose surface produced by anastomosing
principal megascleres echinating fibres. fibre bundles ('lacunae'), interconnected by translucent ectosomal
membrane stretched between adjacent conules; ectosome mem-
Distribution braneous, with thinner choanosomal, undulating fibres lying
immediately below ectosome cored by paucispicular tracts of aux-
Twenty six species have been included in Echinochalina iliary megascleres, and with irregular layer of auxiliary megascle-
at one time or another, of which only twenty are valid (with the res scattered tangential to surface; choanosomal skeleton vaguely
450 Porifera· Demospongiae • Poecilosclerida • Microcionina • Microcionidae

II
I J
Fig. 11. Echinochalina (Echinochalina). A-J, E. (E.) australiensis (Ridley). A, auxiliary subtylostyles/tomostyles (interstitial and coring fibres). B, prin-
cipal subtylostyles (echinating fibres) (scale A-B, 50 /-Lm). C, holotype BMNH 1881.10.21.299 (scale 30mm). D, choanosomal skeleton (scale 500 /-Lm).
E, fibre characteristics (scale 100 /-Lm). F, skeletal reconstruction. G, principal subtylostyle (scale 50 /-Lm). H, principal spicule terminations (scale 10 /-Lm).
I, auxiliary tornostyle (scale 100 /-Lm). J, auxiliary spicule terminations (5 /-Lm).

regularly reticulate composed of primary, distinctly ascending, Type species

multispicular fibres interconnected at irregular intervals by numer-
ous transverse, paucispicular, secondary fibres; fibre anastomoses Siphonochalina bispiculata Dendy, 1895: 246 (by original
form rounded or elongate, narrow meshes; fibres cored by auxiliary designation).
megascleres, sparsely echinated by principal subtylostyles; megas-
cleres principal subtylostyles (echinating fibres) straight, smooth, Definition
tapering, rounded or slightly subtylote bases, fusiform or slightly
telescoped points (83-131 X 6-9.5 jl.m); auxiliary megascleres Echinochalina with diactinal or quasi-monactinal auxiliary
(coring fibres and interstitial) straight, thin, subtylostyles, tomo- megascleres coring fibres, and diactinal or quasi-monactinal prin-
styles or quasi-strongyles, with smooth, rounded, or very slightly cipal spicules echinating fibres.
subtylote bases, tapering-rounded points (147-192 X 1.5-4.5 jl.m);
microscleres absent. Diagnosis
Remarks. Refer to Protophlitaspongia.
Thickly encrusting, massive, lobate, bulbous-digitate, digitate
and arborescent growth forms; ectosomal skeleton, interstitial
SUBGENUS PROTOPHUTASPONGIA BURTON, 1934 skeleton and spongin fibres cored by auxiliary diactinal or quasi-
monactinal spiCUles; fibres echinated by principal diactinal or quasi-
Synonymy monactinal spicules; ectosomal skeleton with erect plumose brushes
of auxilIary spicules arising from peripheral spongin fibres;
Protophlitaspongia Burton, 1934a: 562 (Not Protophlita- choanosomal skeleton reticulate to sub-isodictyal with differentiated
spongia; de Laubenfels, 1954: 96; Pulitzer-Finali, 1986: 138). primary multispicular and secondary paucispicular tracts of auxiliary
Echinochalina (Protophlitaspongia); Hooper 1996a: 543. megascleres; microscleres if present palmate isochelae or toxas.
Porifera· Demospongiae • Poecilosclerida • Microcionina • Microcionidae 451

H
Fig. 12. Echinochalina (Protophlitaspongia). A, E. (E.) bispiculata (Dendy). A, auxiliary oxeas (interstitial and coring fibres). B, principal oxea (echinating
fibres) (scale A-B, 100 fJ-m). C, lectotype NMV G2319 (scale 30 mm). D, choanosomal skeleton (scale 500 fLm). E, fibre characters (100 fJ-m) . F, spiculo-
spongin fibre characteristics (scale 100 fJ-m). G, auxiliary oxea and termination (scales 50, 10 fJ-m). H, principal oxea and terminations (scales L-R,
°
50, 1 fJ-m). I, skeletal reconstruction.

Description of type species reticulate; ectosome microscopically hispid with scattered plumose
brushes of long, thin auxiliary oxeas, arising from the points of
Echinochalina (Protophlitaspongia) bispiculata (Dendy, peripheral fibres and paratangential to surface; choanosomal fibres
1895) (Figs 3A, 12). immediately below surface although spicule tracts more plumose
Synonymy. Siphonochalina bispiculata Dendy, 1895: 246; in peripheral skeleton than at core; choanosomal skeleton regularly
Diplodermia bispiculata; Hallmann 1912: 255; Protophlitaspongia reticulate to subisodictyal, with well developed fibres divided
bispiculata; Burton, 1934a: 562; Echinochalina bispiculata; Hooper into primary, ascending, multi spicular fibres forming distinctly
& Levi, 1993a: 1279. Echinochalina (Protophlitaspongia) bispicu- plumose tracts within core of fibre, interconnected at more-or-less
lata; Hooper, 1996a: 543, figs 296-297. regular intervals by slightly thinner secondary, transverse, uni-,
Material examined. Lectotype: NMV G2319 - Port Phillip pauci- or aspicular fibres; fibres cored by long, thin auxiliary oxeas
Heads, Vic., Australia. Other material. Refer; to Hooper (1996a: and echinated by short, stout principal oxeas; fibre anastomoses
543). form cavernous ovoid meshes; principal megascleres echinating
Description. Shape massive, subspherical, irregularly fibres straight, short, thick, invariably smooth, hastate oxeas, with
lobate, with short bulbous surface lobes, or subcylindrical, flat- abruptly pointed or telescoped ends (31-45 X 2-5.5 j.Lm); auxiliary
tened digitate sponge; oscules small, scattered over surface, partic- megascleres coring fibres long, thin, straight, hastate or rarely
ularly on apex of surface lobes; surface optically even, minutely fusiform oxeas (105-266 X 2-5 j.Lm); microscleres absent.
452 Porifera· Demospongiae • Poecilosclerida • Microcionina • Microcionidae

Remarks. In species of Protophlitaspongia structural completely replaced by detritus although those echinating fibres usu-
megascleres (coring and echinating fibres) are diactinal and do not ally always present. Microscleres palmate isochelae and rarely toxas.
appear to be modified from monactinal forms, unlike all other
microcionids. Consequently, the inclusion of this taxon in the Description of type species
Microcionidae is equivocal but supported by the possession of
more-or-less plumose ascending, primary spicule tracts, true echi- Holopsarnma crassa Carter, 1885b (Figs 2H, 13).
nating megascleres, isochelae and toxa microscleres in several Synonymy. Holopsarnma crassa Carter, 1885b: 211; Aulena
species, and the possession of true monactinal ectosomal spicules in crassa Lendenfeld, 1889a: 101; Echinoclathria crassa; Hallmann,
several species. The coring and echinating megascleres are equated 1912: 287 (Not Antherochalina crassa Lendenfeld, 1887c: 787);
here with auxiliary and principal spicules, respectively, of typical Halrne rnicropora Lendenfeld, 1886a: 303; Halrne globosa
Echinochalina. Dendy (1896) originally included the type species Lendenfeld, 1886a: 303; Echinoclathria globosa; Wiedenmayer,
in the Haplosclerida, but remarked on its unique spicule arrange- 1989: 63.
ment, particularly the ectosomal structure and fibre echination. Ma1erialexamined. Lectotype: BMNH 1886.12.15.313-Port
Burton (1934a) subsequently assigned the type species to Phillip, Vic., Australia. Other material. Refer to Hooper (1996a: 487).
Microcionidae, for similar reasons as those outlined above, whereas Description. Shape massive, subcylindrical, or lobate-
de Laubenfels (1936a) referred it to the Desmacididae because, he digitate honey-combed reticulate sponge, large oscules slightly
suggested, the hastate diactinal megascleres closely resembled recessed within reticulate meshes, surrounded by thin membrane-
those of Guitarra and Liosina, although it lacked poecilosclerid ous lip in life; surface reticulate, lacunose, consisting of irregularly
microscleres. Nevertheless, E. (P.) bispiculata is included with meandering ridges forming large meshes producing characteristic
Echinochalina since it has a paratangential ectosomal skeleton com- honey-comb growth form, in life covered by moderately thin,
posed of auxiliary megascleres, reticulate spongin fibres and echi- translucent dermal membrane stretched between adjacent ridges;
nating principal spicules, whereas other species described by de ectosome heavily arenaceous, with a thick sandy external cortex
Laubenfels (1936a) (P. aga, P. ada and P. antillana) were referred to covered by a fine membraneous ectosomal skeleton usually con-
the Desmacididae and Haplosclerida (Hooper, 1996a). taining a tangential layer of auxiliary spicules, in tracts or scattered
singly across surface, but easily detached; subectosomal region
undifferentiated from choanosome, fibres immediately subectoso-
HOLOPSAMMA CARTER, 1885 mal; choanosomal skeleton irregularly reticulate, with heavy, rela-
tively homogeneous, lamellated spongin fibres fully cored by sand
Synonymy grains, with or without a core of choanosomal principal styles;
fibres usually always echinated by principal styles but sometimes
[Halrne] Lendenfeld, 1886a: 285 (preocc.) (Not Halrne echinating spicules reduced in heavily arenaceous specimens;
Pascoe, 1869). (type species Holopsarnma larninaefavosa Carter, choanosomal principal styles coring and echinating fibres short,
1885b: 212 (by subsequent designation; de Laubenfels, 1936a: 17) thin, straight, with rounded or tapering smooth bases, fusiform,
holotype BMNH 1886.12.15.312). Holopsarnrna Carter, 1885b: sharply pointed (53-81 X 1.2-6/-Lm); subectosomal auxiliary
211. Plectispa Lendenfeld, 1888: 225 (type species Plectispa spicules long, slender, strongylote styles, straight or slightly curved
rnacropora, in part, Lendenfeld, 1888: 226 (by subsequent designa- or sinuous, with slightly subtylote bases, evenly rounded or bifid
tion; Hallmann, 1912: 204) holotype AM G9159). Aulena; in part, points (188-261 X 1.5-6/-Lm); microscleres palmate isochelae
Lendenfeld, 1888: 228 (Not Lendenfeld, 1886a: 309). Taxonomic small, straight shaft, long, thick lateral alae fully fused to shaft,
decision for synonymy: Hooper (1996a). nearly completely detached from front ala, front ala shorter than
lateral alae (8-15 /-Lm); toxas absent.
Type species Remarks. The genera Holopsarnrna and Echinoclathria,
as defined here, have been confused throughout the literature and
Holopsarnma crassa Carter, 1885b: 211 (by subsequent desig- several names (Echinoclathria, Ophlitaspongia, Plectispa, Halrne)
nation, de Laubenfels, 1936a: 98) (lectotype BMNH have been used interchangeably for all species. However,
1886.12.15.313; Hooper & Wiedenmayer, 1994), a senior synonym Holopsarnrna differs substantially from Echinoclathria in having a
of Halrne globosa Lendenfeld, 1886a: 303 (lectotype BMNH distinctive 'honeycombed reticulate' growth form, homogeneous
1886.8.27.71) (cf. Wiedenmayer, 1989: 63). fibre reticulation, a single size class of choanosomal spicule found
both inside fibres and echinating fibres (lacking the larger,
Definition extra-axial styles found in the peripheral skeleton of most
Echinoclathria). When respective type species are compared it is
Microcioninae of 'honeycomb reticulate' growth form with surprising why these genera have ever been confused at all.
coring and echinating spicules identical in geometry, or coring Distribution. Twenty four named species have been
spicules replaced partially or completely by detritus. referred to this genus at one time or another, of which only ten are
well established, all of which are endemic to the Gondwanan fauna.
Diagnosis

'Honeycombed reticulate' growth form consisting of tightly PANDAROS DUCHASSAING & MICHELOTTI, 1864
anastomosing flattened fibre-branches (lacunae); choanosomal
skeleton simply reticulate, without any axial compression or differ- Synonymy
entiation between axial and extra-axial skeletons; principal spicules
core and echinate fibres, those inside fibres sometimes partially or Pandaros Duchassaing & Michelotti, 1864: 88.
Porifera· Demospongiae • PoeciioscIerida • Microcionina • Microcionidae 453

I.H
Fig. 13. Holopsamma. A-J, H. crassa Carter. A, subectosomal auxiliary subtylostyles/strongyles. B, principal style (coring and echinating fibres) . C,
palmate isochela (scale A-C, 50!-Lm). D, specimen SAM TS4085 . E, lectotype BMNH 1886.12.15.313 (scale D-E, 30mm). F, choanosomal skeleton
(scale 1 mm). G, fibre characteristics (scale 200 !-Lm). H, palmate isochelae (scale 5 !-Lm). I, base and bifid point of auxiliary spicule (scale 5 !-Lm). J, base
and point of principal spicule (scale 10 !-Lm).

Type species Description of type species

Pandaros acanthifolium Duchassaing & Michelotti, 1864: 90 Pandaros acanthifolium Duchassaing & Michelotti, 1864
(by subsequent designation; de Laubenfels, 1936a: 123). (Figs 3B, 14).
Synonymy. Pandaros acanthifolium Duchassaing &
Definition Michelotti, 1864: 90; Pandaros arbusculum, in part, Duchassaing &
Michelotti, 1864: 88; Thalyseurypon conulosa Hechtel, 1965: 44.
Microcioninae with flattened spongin fibres cored by smooth Materio.l examined. Lectotype: TM POR57 (fragments
slightly rhabdose auxiliary styles and echinated by rare acanthose USNM31020, MNHN DNBE1309, BMNH 1928.11.12.15a,16a)-
and smooth styles. St. Thomas, Virgin Is., Caribbean.
Description. Bushy arborescent growth form. Surface
Diagnosis highly conulose, with flattened or lobate lamellae. Choanosomal
skeleton reticulate, with well developed flattened spongin fibres
Bushy arborescent growth form; with well developed (trabeculae) cored by choanosomal principal subtylostyles lying in
choanosomal skeletal reticulation of spongin fibres woven into all directions within fibres (from isodictyal reticulate to echinating)
flattened anastomosing branches, or forming continuous sheets and with sparse acanthostyles echinating or also incorporated
without any regular architecture; fibres cored by auxiliary styles, into fibres. Subectosomal skeleton radial, reduced to single long
subtylostyles to tylostyles, predominantly smooth, distributed subectosomal auxiliary subtylostyles protruding through surface
within fibres in criss-cross fashion; fibres echinated by rare acan- and also scattered throughout mesohyl. Ectosome without special
thostyles, rarely smooth styles; few anisotomotes, verging on spicules. Megasc1eres smooth choanosomal principal subty-
oxeas, scattered throughout mesohyl; sparsely dispersed tangential lostyles-tylostyles, often with slightly rhabdose bases and terminal
subectosomal auxiliary spicules dispersed throughout peripheral or subterminal basal swellings, and often polytylote or irregular
skeleton; microsc1eres absent. terminations, with large size range (84-435 X 3-11 /-Lm), long
454 Porifera· Demospongiae • Poecilosclerida • Microcionina • Microcionidae

Fig. 14. Pandaros. A-G, P. acanthifolium Duch. & Mich. A, subectosomal auxiliary subtylostyle (scale 100 fLm). B, sinuous raphidifonn anisotomotes
(scale 50 fLm). C, principal subtylostyles (scale 100 fLm). D, 'echinating' acanthostyle (scale 50 fLm). E, section through peripheral skeleton (scale
500 fLm). F, part of lectotype BMNH 1928.l1.12.15a (scale 30 mm). G, skeletal reconstruction (after Van Soest, 1984b).

curved or straight subectosomal auxiliary subtylostyles spicules than they do echinating (accessory) spicules typical of
(340--665 X 5-13 /Lm), lightly acanthose or rarely smooth styles other Microcionidae; furthermore, these spicules are only rarely
'echinating' fibres (160-195 X 5-8 /Lm), and long curved or sinu- seen echinating fibres, but more commonly they are incorporated
ous, raphidiform spicules verging on anisotomotes (303-455 X into them together with the principal megascleres; (3) the
2-4/Lm). Microscleres absent. long subectosomal auxiliary subtylostyles that protrude through
Remarks. The above diagnosis is compiled from the surface (more similar to a reduced Raspailiidae, such as
re-examination of type and recent material of Pandaros acanthi- Echinodictyum or Ceratopsion, than to typical Microcionidae); and
folium (schizotype MNHN DNBE1309, specimen BMNH (4) the possession oflong raphidiform, sinuous or curved spicules
1884.7.11.2) and the redescription of live populations by Van Soest (classed as 'microscleres' with question by Van Soest, 1984b) that
(1984b). The important features of this species and genus Pandaros have irregular terminations - sometimes stylote and sometimes
are: (1) the prominently flattened fibres cored by smooth slightly strongylote - which he calls anisotomotes.
rhabdose principal subtylostyles-tylostyles (more reminiscent of Pandaros is a borderline taxon that could be legitimately
Rhabderemiidae than Microcionidae); (2) the sparse, lightly spined included in either Raspailiidae or Microcionidae. In fact Van
styles which more closely resemble a second category of principal Soest (1984b) noted that only the presence of rare echinating
Porifera· Demospongiae • Poecllosclerida • Microcionina • Microcionidae 455

acanthostyles in P. acanthifolium gives any cause to link it to the Definition

Microcionidae at all. He speculated that it might be necessary to
erect a separate family for the species, or even remove it from the Ophlitaspongiinae with choanosomal skeleton modified to a
Poecilosclerida altogether, as it also shows affinities with axinellid basal or axial renieroid reticulation of acanthose or occasionally
genera such as Ptilocaulis. There are no microscleres to give any smooth styles and!or strongyles, overlaying a plumose(-reticulate)
further clues as to its affinities. Pandaros is maintained as a sepa- subectosomal skeleton of smooth principal styles, with or without
rate genus and included in Microcionidae on a tentative basis until echinating spicules.
further evidence is forthcoming. Of the twelve species previously
referred to Pandaros, only the type species clearly belongs here. Diagnosis
Raspailia kasumiensis Tanita (MMBS SIS-052), which Hoshino
(1981a) referred to Thalyseurypon, should also be allocated to Encrusting, massive, bulbous, lobate, digitate and flabellate
Pandaros in its skeletal architecture and spiculation (as suggested growth forms; skeleton with 2 distinct components: (1) primary
earlier by Hooper, 1990b). (basal or axial) renieroid choanosomal skeleton replacing usual
Wiedenmayer (1977b) merged Thalyseurypon with Pandaros, microcionid choanosomal skeleton, composed of acanthostyles and!
because he considered that the type species of the former (viz., or acanthostrongyles; (2) secondary (extra-axial, subectosomal)
Spongia raphanus Lamarck) had architecture closely comparable skeleton composed of smooth choanosomal styles forming
to P. acanthifolium (Hechtel, 1965), but this is not upheld here. The dendritic, plumose, subisodictyal or plumo-reticulate tracts, or
only features these genera have in common is in lacking microscle- simply echinating main spiCUle tracts; secondary skeleton usually
res. Wiedenmayer (1977b) also speculated that the genus had a arising from nodes of renieroid skeleton, or ascending upwards
close relationship with Echinoclathria (=Holopsamma as defined from basal spongin fibres, with or without axial compression;
here), based on alleged similarities in skeletal architecture, and he spongin fibres relatively poorly developed; additional category
suggested that the two genera probably intergrade in habit and of echinating acanthostyles present or absent; ectosomal skeleton
spiculation, but these suggested affinities were not noted from tangential, paratangential or plumose tracts of 1 or 2 categories
re-examination of relevant specimens. of auxiliary styles; microscleres diverse forms of isochelae and
toxas.
Distribution
Remarks
Known only from the Caribbean.
Under Van Soest & Stone's (1986) system all microcionids
having a renieroid basal (or axial) skeleton composed of acanthose
megascleres are grouped in a single genus (Antho s.l.). This system
SUBFAMILY OPHLITASPONGllNAE was supported by Hooper (1996a) with the distinction that three
DE LAUBENFELS, 1936 subgenera are recognised within Antho based on structure and
composition of the renieroid skeleton: the nominotypical subgenus
Synonymy Antho (with predominantly (acantho)styles forming the renieroid
skeleton, less often acanthostrongyles, without echinating
Ophlitaspongiidae de Laubenfels, 1936a. acanthostyles), Acamia (with predominantly (acantho)strongyles
forming the renieroid skeleton, less often acanthostyles, and
Defmition a special category of echinating acanthostyles overlap the main
skeleton), and !sopenectya (with an axially compressed and extra-
Microcionidae with a secondary renieroid spiculo- axially renieroid reticulate skeleton composed of two forms of
spongin skeleton (see Fig. 1, symbol 9) overlaying a primary retic- choanosomal spicules inside spongin fibres, overlaid by a second
ulate, plumo-reticulate, plumose or hymedesmioid skeletal extra-axial plumose skeleton). It could be argued that !sopenectya
architecture. (s.s.) could also be included in Echinoclathria, given the close
resemblance in growth form and renieroid skeletal structure with
Scope the type species E. leporina, but in !sopenectya the renieroid skele-
ton is composed of sparsely spined principal styles (clearly differ-
Four genera and three subgenera: Antho (Antho (Antho), entiated from the larger smooth styles of the extra-axial skeleton),
A. (Acamia), A. (!sopenectya», Artemisina, Echinoclathria and with differentiated axial (compressed) and extra-axial (renieroid)
Ophlitaspongia. regions, overlaid by a second extra-axial (plumose) skeleton com-
posed oflarger, smooth principal styles. In Echinoclathria, as rede-
fined by Hooper (1996a) based on its type species, megascleres of
ANTHO GRAY, 1867 the renieroid skeleton are exclusively smooth, and the larger,
smooth principal styles which form a radial skeleton are only found
Synonymy on the surface, embedded in peripheral fibres. Nevertheless, both
genera, Antho and Echinoclathria, differ from other microcionids
Antho Gray, 1867a: 524 (see subgenera for full synonymy). in having a basically renieroid skeletal construction and it is possi-
ble that Echinoclathria are highly derived forms of Antho (loss of
Type species spinated principal spicules, loss of extra-fibre skeleton, loss of
spined acanthostyles (geometrically different from principal
Myxilla involvens Schmidt, 1864: 37 (by monotypy). spicules».
456 Porifera· Demospongiae • Poecilosclerida • Microcionina • Microcionidae

Distribution in part; Topsent, 1892b: 23; Clathria morisca Schmidt, 1864: 37,
45; Dictyoclathria morisca; Topsent, 1920a: 18; Plocamia incon-
Indo-Pacific, W Indian Ocean, NE and NW Atlantic, stans; Topsent, 1925c: 661; Holoplocamia inconstans; de
Mediterranean. Laubenfels, 1936a: 75. Antho inconstans; Ackers, Moss & Picton,
1992: 140; Isodictya beani Bowerbank, 1866: 274; Dictyoclathria
beanii; Arndt, 1935: 81; Amphilectus beanii; Vosmaer, 1880: 115;
SUBGENUS ANTHO GRAY, 1867 Clathria beanii; Ridley, 1881: 485; Myxilla beanii; Topsent,
1892b: 23; Artemisina mediterranea Babic, 1921: 87; Microciona
Synonymy virgula Sara & Siribelli, 1960: 77; ? Artemisina paradoxa Babic,
1921: 87; Clathria paradoxa; Burton, 1930c: 528; Antho paradoxa;
Antho Gray, 1867a: 524. Anomoclathria Topsent, 1929a: 26 Pulitzer-Finali, 1983: 610. ? Raspailia incrustans Swarczewsky,
(Not Topsent, 1932a: 103) (type species Alcyonium opuntioides 1906: 52.
Lamarck, 1815: 164 (by original designation) lectotype MNHN Material examined. Holotype: LMJG (fragment BMNH
DT654). Anthoarcuata Bakus, 1966b: 431 (type species 1867.3.11.92) - Adriatic.
Anthoarcuata graceae Bakus, 1966b: 431 (by original designation Description. Thinly encrusting growth forms; surface
and monotypy) holotype USNM 36284 (161848)). Dictyoclathria rugose, hispid; choanosomal skeleton renieroid reticulate with
Topsent, 1920a: 18 (type species Clathria morisca Schmidt, 1868: acanthostyles-strongyles coring spongin fibres, or simply united
9 (by original designation) (schizotype BMNH 1868.3.2.21); jun- at nodes by variable quantities of spongin, producing triangular
ior synonym of Antho involvens (Schmidt, 1864) (Levi, 1960b: 57) and rectangular skeletal meshes; junctions of skeletal meshes with
schizotype BMNH 1867.3.11.92). [Dyctioclathria] Ferrer- principal choanosomal styles echinating fibre nodes, standing erect
Hernandez, 1921: 172 (lapsus). Isociona Hallmann, 1920: 768 or at oblique angles, in tufts or singly; true echinating megascleres
(type species Lissodendoryx tuberosa Hentschel, 1911: 326 (by absent (i.e., undifferentiated from choanosomal principal styles);
monotypy) holotype 2MB 4417). Jia de Laubenfels, 1930: 28 ectosome contains tangential or paratangential multi spicular
(type species Jiajia de Laubenfels, 1930: 28 (by original designa- brushes of subectosomal auxiliary styles protruding through
tion) holotype USNM 21510). Plocamilla, in part, Burton, 1935a: surface; megascleres acanthose principal styles/strongyles of
402 (Not Plocamilla Topsent, 1928c: 63). ? Quintoxilla de renieroid basal skeleton (124-187 X 2-12 /-Lm), smooth or slightly
Laubenfels, 1950a: 18 (type species Myxilla arcitenens Topsent, acanthose principal styles/subtylostyles of the choanosomal skele-
1892b: 24, by original designation; no extant type material). ton (164-335 X 9-14 /-Lm), and smooth subectosomal auxiliary
Quizciona de Laubenfels, 1936a: 111 (type species Microciona styles, often with basal spines (175-386 X 4-7 /-Lm); microscleres
heterospiculata Brondsted, 1924b: 465 (by original designation) palmate isochelae (13-20/-Lm), wing-shaped and accolada toxas
schizotype BMNH 1901.12.26.13). Taxonomic decision for syn- (22-175 X 1-4/-Lm).
onymy: Hooper (1996a), and this work. Remarks. Twenty two named species have been included in,
or referred to, Antho (Antho), although only twelve of these are
Type species currently widely accepted as being valid. The subgenus was
recently revised (Hooper, 1996a) and needs no further discussion
Myxilla involvens Schmidt, 1864: 37 (by monotypy). here apart from some comment on the inclusion of Quintoxilla de
Laubenfels in the above synonymy, with question. The type species
Definition is described as thinly encrusting, having large acanthostyles 330 X
4-5/-Lm, echinating acanthostyles 190/-Lm long, and ectosomal
Antho with predominantly (acantho)styles forming the smooth tylostyles or tylostrongyles (no sizes given). Microscleres
renieroid skeleton, less often acanthostrongyles, without echinat- are toxas 90-330 /-Lm long, 'palmate tridentate' chelae 20/-Lm long,
ing acanthostyles. and flexuous raphides which could be young toxas 100/-Lm long,
often forming bundles (toxodragmata). No skeletal structure was
Diagnosis described, but Topsent's (1892b) inclusion of the taxon in Myxilla
was presumbably on account of an observed reticulate arrangement
Encrusting, lobate and digitate growth forms; with a primary of the skeleton, and this, together with megasclere and microsclere
basal renieroid choanosomal skeleton composed of acanthostyles geometries, suggests affinities with Antho. De Laubenfels' (1936a)
and/or acanthostrongyles; secondary extra-axial (subectosomal) assumption that chelae are anchorate is unfounded. Unfortunately
skeleton plumose, plumo-reticulate, or simply composed of there appears to be no surviving type material (or slides) of the type
choanosomal styles echinating (project from) basal renieroid skele- species (Hooper, Van Soest, personal observations of the MNHN
ton; spongin fibres poorly developed; special category of echinating collections), so these details and the true allocation of this taxon
acanthostyles absent; ectosomal skeleton with tangential, paratan- cannot be verified. It remains Antho (Antho) incertae sedis.
gential, or plumose tracts of one size of auxiliary styles or subty-
lostyles; microscleres include diverse forms of isochelae and toxas.
SUBGENUS ACARNIA GRAY, 1867
Description of type species
Synonymy
Antho (Antho) involvens Schmidt, 1864 (Figs 3C, 15).
Synonymy. Myxilla involvens Schmidt, 1864; Hymedesmia Acarnia Gray, 1867a: 515. Plocamia Schmidt, 1870: 62 (type
involvens; Schmidt, 1866a: 16; Antho involvens; Gray, 1867a: 524; species Plocamia gymnazusa Schmidt, 1870: 62 (by subsequent des-
Desmacodes involvens; Vosmaer, 1880: 108; Myxilla banyulensis, ignation; Burton, 1935a: 401 (holotype possibly LMJG, schizotype
Porifera· Demospongiae • Poecilosclerida • Microcionina • Microcionidae 457

Fig. 15. Antho (Antho) . A-G, A. (A.) involvens (Schmidt). H-I. A. (A.) tuberosa (Hentschel). A, subectosomal auxiliary styles. B , smooth principal styles
of the choanosomal skeleton (scale A-B , 50 fLm) . C,acanthostyles/strongyles of the renieroid basal skeleton (scale 100 fLm). D, wind-shaped and accolada
toxas (scale 25 fLm). E, palmate isochelae (scale 10 fLm) . F, ectosomal skeleton. G. choanosomal skeleton (scale F-G, 250 fLm). H, choanosomal skeletal
structure (scale 250 fLm) . I,renieroid skeleton (scale 50 fLm) .

MNHN DCL1105L» (Fig. 17). [Dirrhopaluml Ridley, in Ridley & Burton, 1932b: 320 (type species Protoclathria simplicissima
Duncan, 1881: 477 (unjustified replacement name for Plocamia) . Burton, by monotypy, type material not examined). Holoplocamia
Plocamiopsis Topsent, 1904b: 155 (type species Plocamiopsis de Laubenfels, 1936a: 75 (type species Holoplocamia penneyi de
signata Topsent, 1904b: 155 (by monotypy) ho1otype MNHN Laubenfels, 1936a: 75 (by original designation) holotype USNM
DT947). Heteroclathria Topsent, 1904c: 93 (type species 22460). Echinoplocamia Burton, 1959a: 252 (type species
Heteroclathria hallezi Topsent, 1904c: 94 (by original designation Echinoplocamia arbuscula Burton, 1959a: 252 (by original desig-
and monotypy) schizotype MNHN DT1884). Lissoplocamia nation) holotype BMNH 1936.3.4.413). Taxonomic decision for
Br~ndsted, 1924b: 470 (type species Lissoplocamia prima synonymy: Hooper (1996a), and this work.
Br~ndsted, 1924b: 470 (by original designation) holotype not seen;
specimen MNHN DCL637 ex. South Africa). Plocamilla Topsent, Type species
1928c: 63 (Not Burton, 1935a: 402) (type species Isodictya cori-
acea Bowerbank, 1874b: 136 (by original designation; Topsent, Hymeniacidon cliftoni Bowerbank, 1862a (by monotypy)
1928c: 63) holotype BMNH 1910.1.1.251). Anomoclathria; in (junior synonym of Spongia frondifera Lamarck, 1814; Hooper,
part, Topsent, 1932: 103 (Not Topsent, 1929a: 26). Protoclathria 1996a: 422).
458 Porifera· Demospongiae • Poecllosclerida • Microcionina • Microcionidae

Fig. 16. Antho (Acarnia). A-L, A. (A.) frondifera (Lamarck). A, subectosomal auxiliary subtylostyles. B, choanosomal principal subtylostyles. C, acan-
thotylostrongyles of the renieroid skeleton. D, wing-shaped toxa. E, palmate isochelae (scales A-E, 50/-Lm). F, lectotype MNHN DT565 scale 30 mm.
G, choanosomal skeleton (scale 500 /-Lm). H, renieroid skeleton (scale 100 /-Lm). I, toxa (scale 50 /-Lm). J, acanthotylostrongyles (scale 50 /-Lm). K, palmate
isochelae (scale 5 /-Lm). L, bases of choanosomal principal style and auxiliary subtylostyle (scale 5/-Lm).

Definition standing perpendicular to base or axis, joining with echinating

megascleres to produce ascending plumose skeletal tracts. Extra-
Antho with predominantly (acantho)tylostrongyles forming axial (subectosomal) skeleton plumose, dendritic, or subisodictyal,
the renieroid skeleton, less often acanthostyles, and a special cate- composed of choanosomal styles, originating from substrate or
gory of echinating acanthostyles overlap the main skeleton. simply confined to periphery, forming tangential, paratangential
or plumose extra-axial tracts. Ectosomal skeleton with or without
Diagnosis specialized spiculation (one or two categories of auxiliary styles).
Microscleres include diverse forms of isochelae and toxas.
Encrusting, lobate, digitate and flabellate growth forms; with
regular basal or axial renieroid skeleton of acanthostrongyles (less Description of type species
frequently acanthostyles or smooth tylostrongyles), with or with-
out spongin fibres. Renieroid tracts may be echinated by acan- Antho (Acamia)frondifera (Lamarck, 1814) (Fig. 16).
thostyles at spongin fibre nodes. Basal renieroid skeleton overlays Synonymy. Spongia frondifera Lamarck, 1814: 445;
hymedesmioid, leptoclathriid or microcionid main skeleton com- Lamarck, 1816: 374. Anomoclathria frondifera; de Laubenfels,
posed of echinating (acantho-)styles and/or choanosomal styles, 1936a: 108. Antho frondifera; Hooper & Wiedenmayer, 1994: 256.
Porifera· Demospongiae • Poecilosclerida • Microcionina • Microcionidae 459

Antho (Plocamia) frondifera; Hooper, 1996a: 422, figs 216-217. current authors. However, one of those species (A. (Acarnia)
Anomoclathria opuntioides var. frondifera Topsent, 1929a: 26-29, erecta) is very poorly known, and it is also possible that a number
figs 10-14; Topsent, 1932a: 103, pI. 1, figs 6-7. Hymeniacidon of other species will be eventually merged. The nominal genus
cliftoni Bowerbank, 1862a: 773, pI. 30, fig. 9; Bowerbank, 1864: Echinoplocamia was also overlooked by Hooper (1996a) and is
276, figs 70, 291. Acamia cliftoni; Gray, 1867a: 515. (Not included here as a new synonym within the subgenus.
Alcyonium opuntioides Lamarck, 1815: 164.) It is possible that species with smooth tylostrongyles forming
Material examined. Lectotype: MNHN DT565 - ? Sw. the renieroid skeleton ('sausage-shaped spicules' of Ridley, in
Australia (Turgot collection). Paralectotype: MNHN DT3356 - Ridley & Duncan, 1881) should be separated into a separate taxon
same details. Holotype of H. cliftoni: BMNH 1877.5.21.608 (frag- (e.g., subgenus Plocamia). These spicules superficially appear
ments BMNH 1877.5.21.616, 1185,218) - SW Australia. to be geometrically distinct from (and therefore possibly not
Description. Lobate, thickly flabellate, digitate fans with homologous with) acanthotylostrongyles in typical Acarnia
uneven, digitate margins and irregular lobate surface. Large oscules (Fig. 17). These smooth smooth 'sausage-shaped spicules' are found
scattered evenly over surface and lateral margins of digits, with in the nominal genera Plocamia (including Dirrhopalum),
remnants of stellate drainage canals converging on each oscule. Heteroclathria and Lissoplocamia. However, skeletal structures are
Ectosomal skeleton membraneous, heavily collagenous, with some identical between both groups without any other obvious corrobo-
embedded detritus and sparse tangential and paratangential tracts or ratory characters to support their differentiation. Another nominal
single auxiliary subtylostyles scattered near periphery, sometimes
forming bundles protruding through ectosome; points of (smooth)
choanosomal principal styles from ascending plumose tracts pro-
trude only slightly through collagenous surface membrane; subecto-
somal skeleton virtually undifferentiated from choanosome,
although peripheral choanosomal styles of plumose skeleton
slightly more dense, diverging, than tracts in skeletal core.
Choanosomal skeleton with two distinct components: (1) ascending
plumose and (2) basaUaxial renieroid (in some places isodictyal).
Plumose skeleton with pauci- or multispicular tracts of smooth
choanosomal principal styles ascending to surface, rarely branching f
or anastomosing; tracts associated with, but not necessarily coring,
heavy, dark brown, spongin-coated algal filaments (ostensibly
Ficus; Topsent, 1932a), which dominate skeleton; filaments up to
250 f.Lm diameter, 300-400 f.Lm apart, branching, diverging from
base of sponge through sponge surface. Renieroid skeleton com-
posed of 1 or 2 acanthotylostrongyles abreast forming square or tri-
angular meshes up to 120 f.Lm diameter, even mesh size throughout
skeleton, overlaying plumose skeleton; some detritus scattered
between renieroid skeletal meshes, usually coated with spongin;
mesohyl not intact although some granular collagen containing
microscleres scattered between spicule meshes. Megascleres:
choanosomal principal styles entirely smooth, short, robust,
slightly curved at centre, with rounded or slightly subtylote bases,
fusiform points (88-118 X 4--13 f.Lm). Acanthotylostrongyles of the
renieroid skeleton thick or thin, rounded or slightly subtylote at both
ends, heavily spined particularly at points, spines large, conical or
slightly recurved, sharply pointed (85-103 X 3-14 f.Lm). Subecto-
somal auxiliary subtylostyles long, very slender, curved at centre or
B
sinuous, subtylote usually microspined bases, fusiform or occasion-
ally telescoped points (120-184 X 1-2.5 f.Lm). Microscleres:
palmate isochelae large, unmodified, with front and lateral alae
approximately same length, lateral alae entirely fused to shaft, front
ala detached along lateral margin (15-20 f.Lm long). Toxas wing-
shaped, short, moderately thick, with large central curvature and
slightly reflexed points (40-116 X 1-2 f.Lm).
Remarks. The inclusion of Hymeniacidon cliftoni
Bowerbank into synonymy with the type species of Plocamia
(Spongia frondifera Lamarck) unfortunately requires that the sub-
A
genus name be emended to Acarnia. Previously, Hooper (1996a: 422)
suspected that the two species were synonymous but the available
type material of Acarnia was so poor to be uncertain. Subsequent
C E
Fig. 17. Antho (Acamia), cont. A-E, A. (A.) gymnazusa (Schmidt, 1870)
examination of a better slide preparation confirms this synonymy.
(type species of Plocamia Schmidt). A, echinating acanthostyle (scale 50 j..l.m).
Twenty two nominal species have been referred to this sub- B, subectosomal auxiliary subtylostyle. C, choanosomal principal styles (scale
genus, all of which are presently recognized as valid species by B-C, 100 j..l.m). D, tylostrongyles (scale 50 j..l.m). E, toxas (scale 10 j..l.m).
460 Porifera· Demospongiae • Poecilosclerida • Microcionina • Microcionidae

genus, Protoclathria Burton, 1932b, is also included here as a Description. Shape thinly flabellate with long, thickly
synonym of Antho (Acamia), overlooked by Hooper (1996a). The cylindrical stalk, very thin lamellae, slightly digitate or evenly
type species, P. simplicissima Burton from Tristan da Cunha, is a rounded margins; firm, barely compressible, flexible, slightly
massive pyriform sponge with a hispid, even surface. The skeleton spiky; optically smooth, even surface; ectosome prominently
is renieroid, composed of dense reticulation of triangular, mostly hispid, with pauci- or multispicular plumose brushes of larger,
multispicular meshes cored by a single category of smooth smooth choanosomal principal styles protruding through surface,
subtylostyle (260 X 15 /-Lm), occasionally with oxeote modifica- forming a vestigial radial extra-axial skeleton, arising from pauci-
tion, intermingled with modified acanthostrongyles (180 X 12/-Lm) or multi spicular tracts of (marginally) smaller smooth principal
coring fibres, apparently not echinating them. No ectosomal styles in subectosomal region; subectosomal auxiliary subty-
spicules were described and microscleres are also apparently lostyles tangential, paratangential, or rarely plumose, at base of
absent. It is likely that this species was imperfectly described protruding choanosomal spicule brushes; peripheral skeleton
by Burton (1932b) as no mention was made of a secondary relatively cavernous in comparison to the central choanosomal
plumose or plumo-reticulate skeleton common to all members skeleton, moderately heavily collagenous; choanosomal skeletal
of Antho. structure with 3 distinct components: (1) slightly compressed spon-
gin fibres forming close-meshed anastomoses at core of skeleton,
more cavernous towards surface, (2) renieroid skeleton composed
SUBGENUS ISOPENECTYA DALLMANN, 1920 of acanthose styles, overlaying other structures, (3) longitudinal,
ascending tracts of smooth principal styles, marginally smaller
Synonymy than those protruding through surface, forming subisodictyal tracts
at core, more plumose in periphery, and usually (but not invariably)
[sopenectya Hallmann, 1920: 789. Clathriella Burton, 1935c: associated with larger spongin fibres; spongin fibres in axial skele-
73 (type species Clathriella primitiva Burton, 1935c: 73 (byorigi- ton heavy, producing irregularly oval or elongate meshes, cored by
nal designation) holotype BMNH 1938.7.4.93). Taxonomic deci- uni- or bispicular tracts of smaller, smooth choanosomal principal
sion for synonymy: Hooper (1996a). styles; fibres closer to surface regularly anastomosing, wide-
meshed, forming regularly renieroid (triangular) spicule meshes
Type species and oval or elongate fibre meshes, cored by uni- or bispicular tracts
of smaller acanthose styles; plumose extra-fibre skeleton com-
Clathria (?) chartacea Whitelegge, 1907: 497 (by monotypy). posed of uni-, pauci- or multi spicular ascending tracts of smooth
choanosomal styles standing perpendicular to axis, becoming
Definition increasingly plumose, larger, and typically multispicular towards
periphery; echinating megascleres absent; megascleres smooth
Antho with an axially compressed and extra-axially renieroid choanosomal principal styles long, thick, slightly curved or
reticulate skeleton composed of two forms of choanosomal straight, with rounded or very slightly subtylote bases, rarely with
spicules inside spongin fibres, overlaid by a second extra-axial basal microspination, fusiform points (168-274 X 13-17 /-Lm);
plumose skeleton. acanthose choanosomal styles of renieroid skeleton short, thick,
fusiform, slightly curved or straight, with rounded or slightly sub-
Diagnosis tylote bases, lightly microspined bases and points, with fewer
spines scattered on shaft, occasionally completely smooth shaft
Bulbous, arborescent and flabellate growth forms; with three (92-127 X 9-12.5 /-Lm); subectosomal auxiliary subtylostyles
skeleton components: (1) renieroid reticulation of acanthose styles, short, thin, usually straight, with prominent subtylote, typically
(2) overlayed by isodictyal or subisodictyal reticulation of smooth microspined bases, hastate points, abrupt points, or sometimes tele-
styles coring spongin fibres, (3) surmounted by plumose or radial scoped or bifid points (163-243 X 2-4.5 /-Lm); microscleres absent.
extra-axial skeleton of larger smooth styles, perpendicular to axis, Remarks. Incuding the present work [sopenectya contains
in peripheral region; skeleton may be slightly compressed at core, only four species, three from the SW. Pacific and one from the NW
spongin fibres only moderately developed; echinating megascleres Pacific Ocean. All species lack microscleres but this is interpreted
absent; ectosomal skeleton with single category of auxiliary subty- as a secondary loss and consequently not given primary diagnostic
lostyle forming tangential or paratangential tracts; microscleres importance. Hallmann (1920) created [sopenectya for the type
absent. species primarily on the basis of having a renieroid skeleton, with
two categories of choanosomal styles, without echinating acan-
Description of type species thostyles, and without microscleres. The type species has obvious
affinities with Antho. It differs from other microcionids with
Antho (lsopenectya) chartacea (Whitelegge, 1907) myxillid-like renieroid skeletons (viz., Antho (Antho), Antho
(Figs 3E, 18). (Acamia), Clathria (Clathria) and Clathria ([sociella» in having a
Synonymy. Clathria (?) chartacea Whitelegge, 1907: 497. condensed axis and more-or-less plumose extra-axial skeletons
[sopenectya chartacea; Hallmann, 1920: 789. Antho chartacea; cored by smooth choanosomal (principal) styles, in one or more
Rudman & Avern, 1989: 335. Antho (lsopenectya) chartacea; size categories, together with the usual renieroid structure overlay-
Hooper, 1996a: 433, figs 221-222. Antherochalina perforata ing the remainder of the skeleton composed of acanthose (or some-
Lendenfeld, 1887c: pl.22, fig.44 (Not Antherochalina perforata times smooth) styles different from principal spicules.
Lendenfeld, 1887c: 788; Lendenfeld, 1888: 89-90). The subgenus superficially resembles Echinoclathria lepo-
Material examined. Holotype: AMZ436 - Off Coogee, rina, mainly in the emphasis of the compressed central skeleton and
NSW, Australia. Other material. Refer to Hooper (1996a: 433). subrenieroid skeletal structure in both species, whereas megascleres
Porifera· Demospongiae • Poecilosclerida • Microcionina • Microcionidae 461

A
Fig. 18. Antho (lsopenectya). A-K, A. (I.) chartacea (Whitelegge). A, choanosomal principal style. B, subectosomal auxiliary style. C, acanthostyle of the
renieroid skeleton (scale A-C, 25 fLm). D, holotype AMZ436 (scale 30 mm). E, choanosomal skeleton (scale 500 fLm). F, fibre characteristics (scale 100 fLm).
G, acanthostyle (scale 50 fLm). H, acanthostyle shaft spines (scale 5 fLm). I, base of principal style (scale 10 fLm). J, base of auxiliary style (scale 5 fLm).
K, tips of auxiliary styles (scale 5 fLm).

forming these skeletons are quite different. Choanosomal megas- ECHINOCLATHRIA CARTER, 1885
cleres in A. (I.) chartacea are clearly differentiated: small acan-
those styles forming the renieroid skeleton (not echinating fibres), Synonymy
small smooth styles forming a secondary radial ascending skeleton,
and larger smooth styles forming the peripheral perpendicular [Echinoclathria] Carter, 1884b: 204 (nomen nudum). Echino-
skeleton. By comparison, in E. Zeporina there is a smaller size class clathria Carter, 1885d: 355 (Not Uriz, 1988: 89).
of smooth principal styles both coring and echinating heavy spon-
gin fibres, forming a renieroid skeletal structure, and a second, Type species
larger class of smooth principal styles forming a sparse radial or
plumose peripheral skeleton (embedded in peripheral fibres). This Echinoclathria tenuis Carter, 1885d: 355 (by subsequent des-
latter structure links the two groups. Antho chartacea should also ignation; Burton, 1934a: 562) (holotype BMNH 1886.12.15.147);
be contrasted with the renieroid raspailiid genus Amphinomia, junior synonym of Spongia Zeporina Lamarck, 1814: 444 (Topsent,
which also has acanthose structural spicules (Hooper, 1991). 1932a: 101) (holotype MNHN LBIM DT567).
462 Porifera· Demospongiae • Poecilosclerida • Microcionina • Microcionidae

F
Fig. 19. Echinoclathria. A-H, E. Jeporina (Lamarck). A, subectosomal auxiliary subtylostyles. B, principal styles/subtylostyles (coring and echinating
fibres) (scale A-B, 25 f.Lm). C, holotype of junior synonym E. tenuis (scale 30 mm). D, choanosomal skeleton (scale 500 f.Lm). E, fibre characteristics (scale
100 f.Lm). F, base of larger principal style (protruding through surface) (scale 10 f.Lm). G, base of smaller principal style (in renieroid skeleton; scale 5 f.Lm).
H, reconstruction of choanosomal skeleton.

Definition skeleton cored by smaller, smooth principal styles, echinated by

identical spicules (occasionally absent), typically very well devel-
Ophlitaspongiinae with the choanosomal skeleton consisting oped spongin fibres sometimes slightly compressed at axis, more
of a relatively homogeneous renieroid spongin fibre reticulation openly reticulate towards periphery; and (2) a vestigial radial extra-
cored by smaller, smooth principal spicules and echinated by the axial skeleton perched on the external surface, barely extending into
same spicules, and a vestigial radial extra-axial skeleton composed choanosome, consisting of larger, smooth principal spicules, with
of larger principal spicules forming plumose brushes on the exter- identical geometry to those at core, forming radial or plumose
nal surface. brushes on surface. Ectosomal skeleton with single size class of
auxiliary subtylostyle lying paratangentially or embedded perpendi-
Diagnosis cular to surface. Microscleres include toxas and palmate isochelae.

Thickly encrusting, bulbous, claviform, bushy, digitate, Description of type species

lamellate, arborescent and flabellate growth forms; with two distinct
skeletal components: (1) predominantly renieroid reticulate main Echinoclathria Zeporina (Lamarck, 1814) (Fig. 19).
Porifera· Demospongiae • Poecilosclerida • Microcionina • Microcionidae 463

Synonymy. Spongia leporina Lamarck, 1814: 444. Echino- (e.g., E. leporina, E. confragosa (Hallmann» there are obvious size
clathria leporina; Topsent, 1932a: 101. Ophlitaspongia leporina; differences between principal styles coring fibres in the choano-
Burton, 1934a: 558. Echinoclathria tenuis Carter, 1885d: 355. some and those protruding through the surface, whereas in others
Ophlitaspongia tenuis; Dendy, 1896: 37 (Not Clathria tenuis (e.g., E. nodosa Carter) there is no obvious size differences
Hentschel, 1911: 377; Parish etal., 1991: 56). Phakelliapapyracea between principal styles at the core and those at the periphery,
Carter, 1886c: 379. Antherochalina tenuispina Lendenfeld, although structurally these are similar to the first condition. In oth-
l887c: 789. ers (e.g., E. egena Wiedenmayer, E. waldoschmitti de Laubenfels)
Material examined. Holotype: MNHN DT567 there is further reduction whereby the extra-fibre skeleton is virtu-
'Australian Seas'. Other material. Refer to Hooper (1996a: 462). ally absent and all spicules are vestigial, poorly silicified. Further
Description. Shape persistently very thin, flabellate digits discussion of the genus, particularly in relation to its alleged simi-
ranging from single elongate planar fans with evenly rounded mar- larities to Ophlitaspongia (sensu Howson & Chambers, 1999), are
gins, to bifurcate palmate digits growing in more than one plane, with presented below (see Ophlitaspongia).
uneven margins, usually with cylindrical basal stalk; oscules small
dispersed over margins of digits; surface even; ectosome micro- Distribution
scopically hispid, with larger sizes of principal style/subtylostyle
protruding through surface, singly or in brushes, forming vestigial Eighteen species are confined to the Indo-west Pacific region,
plumose or radial extra-axial skeleton in peripheral region, with a and another eleven have been recorded from the Caribbean,
distinct tangential layer of subectosomal auxiliary subtylostyles, in SW Atlantic, Arctic, Mediterranean and W Indian Ocean.
pauci- or multispicular tracts, underlying erect principal spicule
brushes; choanosomal skeleton with three components: (1) irregu-
larly isodictyal, slightly compressed axis with single, thickened ARTEMISINA VOSMAER, 1885
central core of heavy fibres, vaguely separated into primary,
plumose, arborescent, ascending pauci- or multi spicular fibres pro- Synonymy
ducing radial tracts, and secondary, mostly transverse uni- or pau-
cispicular, regularly renieroid fibres; (2) more open-reticulate Artemisina Vosmaer, 1885a: 25. [Artenisinal Burton, 1934b:
renieroid or subrenieroid extra-axial region; and (3) (vestigial) 54 (lapsus). Qasimella Thomas, 1974: 311 (type species Qasimella
plumose or radial skeleton in peripheral region with spicule tracts indica Thomas, 1974: 311 (by original designation) holotype
increasingly plumose, protruding through fibres, in peripheral CMFRI T841l not seen). Taxonomic decision for synonymy:
skeleton; fibres cored and sparsely echinated by smaller choanoso- Hooper (l996a).
mal principal styles/subtylostyles in choanosomal skeleton (larger
in peripheral skeleton); echinating principal spicules located pre- Type species
dominantly on primary fibres; fibre anastomoses form rectangular,
triangular, or less often oval meshes more compressed in axis than Artemisina suberitoides Vosmaer, 1885a: 25 (by monotypy)
at periphery; megascleres choanosomal principal styles larger in (holotype ZMA POR443); junior synonym of Suberites arciger
peripheral region than in axis, thick, straight or slightly curved, Schmidt, 1870: 47 (Burton, 1930c: 528) (schizotype BMNH
with smooth, evenly rounded, or slightly tapering subtylote bases, 1870.5.3.90).
sometimes quasi-oxeote, rarely microspined bases, usually with
fusiform points (62-305 X 4-14 !-Lm); subectosomal auxiliary sub- Definition
tylostyles long, thin, straight, slightly curved, or frequently sinu-
ous, with predominantly smooth, occasionally microspined, Ophlitaspongiinae lacking a distinctive choanosomal skeleton
subtylote bases, hastate points (148-321 X 1.5-4.2 !-Lm); or definite spongin fibres, lacking echinating spicules, and having
microscleres absent. a nearly radial ectosomal skeleton.
Remarks. Sixty nine species names have been referred to or
included in Echinoclathria (or one of its synonyms) at one time or Diagnosis
another, but of these only 29 are appropriately referred here (most
of the remainder belonging either to Holopsamma or Clathria, or Massive, cushion-shaped, bulbous, clavulate, tubular, digitate
are junior synonyms or homonyms of other taxa). and flabellate growth forms; without choanosomal fibres or indefi-
Echinoclathria is similar to Antho (/sopenectya), as noted nite fibres, whereas skeletal architecture consists of vaguely ascend-
above, differing in having only two skeletal components: a rela- ing longitudinal tracts of spicules bound by abundant collagen,
tively homogeneous renieroid choanosomal skeleton composed of cored by smooth choanosomal principal subtylostyles in a more-or-
smaller, smooth principal spicules; a vestigial radial extra-axial less confused halichondroid reticulation of vaguely multispicular
skeleton on the external surface. /sopenectya has in addition a ascending and scattered transverse megascleres; echinating megas-
renieroid skeleton of acanthose spicules, and the smooth principal cleres absent; subectosomal peripheral skeleton more radially
styles form longitudinal tracts extending all the way from the axis arranged; ectosome membraneous, skin-like, with smooth styles of
to the surface and beyond. The difficulty lies in trying to determine a single size category protruding through surface, forming paratan-
affinities of species in either genus with reduced structural charac- gential or erect, discrete spicule bundles; microscleres palmate or
ters. For example, A. (/.) punicea Hooper has spined renieroid arcuate isochelae and toxas with smooth or spined points.
spicules whereas E. riddlei Hooper has smooth renieroid spicules,
and both species have a reduced extra-fibre skeleton. Description of type species
Within Echinoclathria most of the variability concerns the
development of the extra-fibre skeleton. In some species Artemisina arcigera (Schmidt, 1870) (Fig. 20).
464 Porifera· Demospongiae • Poecilosclerida • Microcionina • Microcionidae

I
F
A C
Fig.20. Artemisina. A-I, A. arcigera. J, A. jovis Dendy. A, choanosomal principal subtylostyle (scale I ()() !Lm). B, base of principal subtylostyle (scale 20 !Lm).
C, subectosomal auxiliary style (scale 50 !Lm). D, base of auxiliary style (scale 10 !Lm). E, wing-shaped toxas (scale 50 !Lm). F, spined tip of toxa (scale
2 !Lm). G, palmate isochelae (scale 5 !Lm). H, fragment of holotype BMNH 1870.5.3.90 (scale 30mm). I, section through peripheral skeleton (scale 250 !Lm).
J, reconstruction of peripheral skeleton.

Synonymy. Suberites arciger Schmidt, 1870: 47; Artemisina (8-14/Lm) , and wing-shaped toxas with spinous extremities
suberitoides Vosmaer, 1885a: 25; Artemisina arcigera; Lundbeck, (55-280 X 2-4/Lm).
1905: 110; Artemisina arcigera; Burton, 1959b: 42. Remarks. Defining Artemisina in phylogenetic terms is
Material examined. Holotype: ZMUC (not seen) (fragment most difficult. The taxon has no real distinctive features, although
BMNH 1870.5.3.90) -Arctic. it differs from other Microcionidae in lacking a distinctive
Description. Massive, subspherical growth form. Surface choanosomal skeleton or definite spongin fibres (also found in
skin-like, microscopically hispid, with few raised oscules; texture the nominal genus Qasimella Thomas which Hooper, 1996a
distinctly stringy. Choanosomal fibres indefinite or absent, overall synonymised with Artemisina), lacking echinating spicules, and
architecture plumo-reticulate, nearly halichondroid in places, com- having a nearly radial ectosomal skeleton reminiscent of some
posed of multi spicular ascending and paucispicular transverse Ceratopsian (Raspailiidae; Hooper, 1991). These characteristics,
tracts of choanosomal principal styles, bound together with colla- two of which might be interpreted as reductions or secondary
gen. Echinating megasc1eres absent. Ectosomal skeleton plumose, losses and the third as a convergence, are the only definable mor-
composed of single category of subectosomal auxiliary styles phological apomorphies. Moreover, ectosomal structure varies
forming discontinuous palisade of discrete brushes. Megasc1eres between several species, ranging from the typical condition com-
smooth choanosomal principal subtylostyles (416-624 X 9-12 posed of erect brushes (e.g., A. arcigera) to a tangential layer of
/Lm) and smooth fusiform subectosomal auxiliary styles or subty- spicules in criss-cross fashion (e.g., A. meZana Van Soest). This
lostyles (275-390 X 8-16 /Lm). Microsc\eres palmate isochelae variability is equivalent to the MycaZe subgenera Carmia and
Porifera· Demospongiae • Poecilosclerida • Microcionina • Microcionidae 465

Aegagropila, respectively (e.g., Topsent, 1924). At least one Definition

species lacks a specialized ectosomal skeleton completely (e.g.,
A. transiens Topsent). Ophlitaspongiinae with isodictyal reticulate spongin fibre
Some species of Artemisina have honeycombed reticulate skeleton arising from a hymedesmioid basal fibre skeleton; mineral
growth forms, approaching the characteristic Holopsamma mor- skeleton is predominantly or exclusively plumose, with no or very
phology, but there is no consistency or pattern of gross morpholo- few spicules coring transverse fibres; echinating megascleres
gies amongst Artemisina, and in any case it is unlikely that the absent although principal subtylostyles protrude through fibres in
genus can be defined on that basis. De Laubenfels (1936a) and plumose arrangement; and megascleres are exclusively smooth
Ristau (1978b) suggested that Artemisina could be characterized by subtylostyles.
the absence of echinating acanthostyles and by the presence of spin-
ous extremities on toxas. Neither character has much systematic Diagnosis
value at the generic level. In the present interpretation echinating
acanthostyles represent the retention of an ancestral character, in Encrusting growth form; with an isodictyal reticulate, ladder-
which case their presence or absence does not constitute a valid rea- like, spongin fibre skeleton composed of parallel ascending fibres
son to define a phylogenetic grouping, and in any event they occur with regular cross-connecting fibres arising from a hymedesmioid
and disappear throughout numerous microcionid and raspailiid taxa. basal layer of spongin fibre. Mineral skeleton exclusively plumose,
Similarly, toxas with spinous extremities are also known to occur in with only ascending fibres cored by plumose columns of entirely
many Microcionidae, including the type species Clathria compressa smooth principal subtylostyles, with very few or no spicules coring
Schmidt (and other species Microciona spinarchus Carter & Hope, the transverse connecting fibres. Tracts of principal subtylostyles
M. coccinea Bergquist, M. rubens Bergquist, Eurypon acanthotoxa protrude through ascending fibres, diverging towards the surface.
Stephens, M. spinatoxa Hoshino, Stylostichon toxiferum Topsent, True echinating spicules absent but principal styles project from
Labacea juncea Burton, Plocamia ridleyi Hentschel, and fibres as echinating spicules and/or in spicate arrangement.
Ophlitaspongia thielei Burton). They also occur in genera which Microscleres include toxas and palmate isochelae (the latter absent
have an ectosomal structure consistent with the Myxillidae (e.g., in the type species).
Melonchela clathrata Koltun), and consequently it is not parsimo-
nious to base a definition of Artemisina on this feature. Similarly, Description of type species
several other species of Artemisina have smooth toxas (e.g., A.
melana), and one (A. archegona Ristau) has oxeote toxas similar to Ophlitaspongia papilla Bowerbank, 1866 (Figs 3H, 21).
Paratenaciella microxea Vacelet and Vasseur. Synonymy. Ophlitaspongia papilla Bowerbank, 1866: 14; ?
Thus, in the broad sense Artemisina contains a heterogenous Clathria papilla; Schmidt, 1870: 77; Echinoclathria papilla;
assemblage of species, which prompted Burton (1930c) to divide Hanitsch, 1894a; Ophistospongia papilla; Gray, 1867a: 515.
the group into three sections based on the number of megasclere Material examined. Holotype: BMNH 1910.1.1.395 -
categories present. The simplest forms have only one category Vazon Bay, Guernsey.
of spicule (choanosomal principal megascleres; e.g., A. transiens Description (partly from Howson & Chambers, 1999, and
Topsent); the typical form has two categories of megascleres re-examination of type material). Thinly encrusting, with firm
(larger choanosomal principal styles and smaller subectosomal elastic texture, smooth and finely hispid surface with regularly
auxiliary styles; e.g., A. arcigera); and the third form has an incom- arranged oscules flush with surface, lacking subdermal drainage
pletely differentiated series of three megasclere types (two canals. Ectosomal skeleton consists of sparse paratangential bun-
choanosomal spicules and one subectosomal spicule; A. plumosa dles of auxiliary sUbtylostyles usually at the ends of ascending
Hentschel). In all these forms species are only really united in their choanosomal tracts, and also scattered tangential to the surface
lax choanosomal skeletal structure. together with toxas. Choanosomal skeleton consists of parallel
columns of well developed spongin fibres arising from a
Distribution hymedesmioid basal fibre skeleton, with ascending fibres regularly
interconnected by transverse ones, forming a ladder-like isodictyal
Predominant in colder waters including Antarctica, Arctic, reticulation. Ascending fibres cored by plumose tracts of choanoso-
Greenland, Iceland, New Zealand, NE and SW Atlantic, NW, NE mal principal subtylostyles whereas transverse fibres are rarely
and SW Pacific, but also recorded from Mediterranean, Caribbean, cored. True echinating megasc1eres absent. Choanosomal spicule
central Indian Ocean and SW Pacific. tracts protrude through the ectosome in sparse bundles, and in some
places they can be said to be 'echinating' whereas in others merely
'spicate' (i.e., protruding from fibres in plumose fashion). Mesohyl
OPHLITASPONGIA BOWERBANK, 1866 is highly collagenous and contains scattered auxiliary subtylostyles
and toxas. Megascleres consist choanosomal principal subty-
Synonymy lostyles, completely smooth, often curved, widest at the centre and
tapering towards both terminations, with a well developed subtylote
Ophlitaspongia Bowerbank, 1866: 14, 378. (Not swelling and constricted 'neck' (64-143 X 5-13/-lm). Subectosomal
Ophlitaspongia of most authors). auxiliary subtylostyles short, thin, straight, entirely smooth, with
well developed subtylote basal swelling (71-167 X 2-3/-lm).
Type species Microscleres consist of oxhorn to wing-shaped toxas in a single but
wide-ranging size class (64-127 X 2-4/-lm); chelae absent.
Ophlitaspongia papilla Bowerbank, 1866: 378 (by original Remarks. The history of Ophlitaspongia is complex and
designation). tortuous, based largely on misidentifications, misdiagnoses and
466 Porifera· Demospongiae • PoecUoscierida • Microcionina • Microcionidae

Fig. 21. Ophlitaspongia. A-F, O. papilla Bowerbank. A, choanosomal principal subtylostyles. B, subectosomal auxiliary subtylostyle (scale A-B,
50 f.lm). C, oxhom and wing-shaped toxas (scale 50 f.lm). D, section through peripheral skeleton (scale 200 f.lm). E, fibre characteristic (scale 100 f.lm).
F, skeletal reconstruction.

incorrect assumptions concerning type material. The genus was Howson & Chambers (1999) without any full or appropriate dis-
erected by Bowerbank (1866) with O. papilla nominated as the cussion of the significance or implications of its important mor-
type species (with holotype BMNH 1910.1.1.395). Simpson phological characters within the Microcionidae, and consequently
(1968) subsequently proposed that O. papilla was a synonym of a detailed resume of these justifications is provided here.
O. seriata (Grant, 1826c; with alleged holotype being BMNH Howson & Chambers (1999) discovered that the concept of
1847.9.7.14; but see below), which therefore became the senior "0. seriata" of authors was based on a false premise. They redis-
name for the type species, and which was also reportedly cytologi- covered Grant's original slide preparation of Spongia seriata
cally identical to Microciona atrasanguinea Bowerbank, 1862b (UCLZ B73; 'Spongia seriata Gr "variety of S. papillaris ?"') which
(the type species of Microciona Bowerbank, 1863; holotype they found was identical to Halichondria panicea Pallas (i.e.,
BMNH 1930.7.3.225). Thus, under this interpretation, adopted Grant's, 1826c diagnosis of S. seriata was incorrect). They negated
by many contemporary authors (including Hooper, 1996a), the long-held assumption that Johnson's (1842) material (BMNH
Microciona became the senior-most available name for a group of 1847.9.7.14) was the holotype of S. seriata (e.g., Simpson, 1968;
largely encrusting taxa characterised in having smooth echinat- see Hooper, 1996a: 62); and that Johnston (1842) had in his posses-
ing megascleres and hymedesmioid or microcionid skeletal sion material of a new species (BMNH 1847.9.7.14), which
architecture. However, the genus was recently resurrected by he misidentified as Grant's taxon. Johnson's (1842) material is
Porifera· Demospongiae • Poecilosclerida • Microcionina • Microcionidae 467

identical to Bowerbank's (1866) material of O. papilla (BMNH the skeleton reportedly consists of a reticulate network of horny
1910.1.1.395), and therefore, O. papilla Bowerbank becomes the fibres (reminiscent of Ophlitaspongia s.s.). Levi states that the
next available species name for the concept of "0. seriata" sensu South African and North Atlantic populations are 'in all points
Johnston and authors, and the type species of Ophlitaspongia. identical', but this has yet to be corroborated through re-examination
Similarly, comparisons between the holotype of Microciona of relevant type material (although it is highly probable that Levi's
atrasanguinea (BMNH 1930.7.3.225) and Ophlitaspongia papilla voucher material does not exist).
show that Simpson (1968) was incorrect in merging the two taxa, Many other records of Ophlitaspongia species in the literature
and in his conclusion that Ophlitaspongia was a synonym of can also be referred to Echinoclathria. A comparison between the
Microciona. The two species are not at all closely related in their emended definition of Echinoclathria (Hooper, 1996a, differing from
skeletal details, virtually having only their encrusting growth forms previous concepts ofthe genus; e.g., Hallmann, 1911, 1912), Clathria
in common. Not only was spicule morphology very incongruent, and Microciona (as revised by Hooper, 1996a), and Ophlitaspongia
but also skeletal structure, the number and distribution of spicule (as revised by Howson & Chambers, 1999, and summarised here)
categories present, and spicule sizes differed significantly between shows each to have substantially different fibre structure, skeletal
the two species. This finding therefore raises questions concerning structure and distribution of spicule types within fibres. There is
the systematic value of Simpson's (1968) unique cytological data no doubt that each serves its purpose to group like-taxa, but it is
used at the supraspecific level given its flawed taxonomic basis. still equivocal at what level they diverge (genus or subgenus).
Differences between Ophlitaspongia and Microciona are clear Echinoclathria has two distinct skeletal components: (1) an evenly
from their respective type material, and these descriptions (see renieroid reticulate skeleton, including a slightly compressed axis,
above) form the bases of the respective diagnoses. Ophlitaspongia with all axial fibres cored by smaller principal choanosomal styles
has a very prominent spongin skeleton that forms an isodictyal retic- and echinated by the same principal choanosomal styles; (2) the
ulation, ranging from a hymedesmioid basal fibre skeleton to near extra-axial or peripheral skeleton is radial, vestigial, with the largest
the surface, whereas the mineral skeleton is exclusively plumose, principal spicules perched on the outer radial fibres and projecting
principal spicules coring only ascending fibres and no or only very through the surface individually or in bundles; in addition to this there
few spicules coring the transverse fibres; there are no true echinating are also auxiliary styles or subtylostyles tangential or erect on the sur-
spicules (although principal subtylostyles protrude through fibres in face. By comparison, Ophlitaspongia has a homogeneous isodictyal
plumose arrangement); and megascleres are exclusively smooth sub- reticulate fibre skeleton in both the basal and peripheral regions, with
tylostyles. By comparison, Microciona has a plumose, ascending a compressed layer of spongin fibres lying on the substrate and regu-
(i.e., non-reticulate) spongin fibre skeleton with fibre nodes arising larly reticulate spongin fibres arising from this base, producing a
from a hymedesmioid spongin base ('microcionid' architecture), and regular isodictyal fibre network divided into ascending primary and
adjacent fibre nodes do not form a reticulation (i.e., they are exclu- secondary transverse elements. Mostly only the primary ascending
sively plumose). The spicule skeleton consists of choanosomal prin- fibres are cored by principal choanosomal spicules producing a
cipal subtylostyles (larger, smooth or basally spined styles) forming strictly plumose mineral skeleton; there is no differentiated radial
ascending plumose tracts usually perched on the ends of spongin skeletal component in the periphery (as in Echinoclathria); there are
fibre nodes, well differentiated from smaller echinating (acantho-) no true echinating spicules (as in Echinoclathria); and whereas the
styles which are dispersed over the basal layer of spongin and fibre skeleton is reticulate the mineral skeleton is plumose.
ascending fibre nodes, individually or in bundles. Both genera have In resurrecting Ophlitaspongia, and recognising these differ-
smooth wing-shape toxas, and smooth auxilliary subtylostyles ences between it and Echinoclathria, it could be argued that some
forming an extra-fibre and/or dermal skeleton. of the species described by Hooper (1996a) may be conceivably
At present Ophlitaspongia is restricted to two species in the included in it. However, these arguments are rejected as follows.
NE Atlantic (Howson & Chambers, 1999). Although other authors (1) Echinoclathria egena Wiedenmayer, 1989 lacks true echinating
have reported alleged populations of O. papilla (or O. seriata sensu spicules, and its skeletal strucutre is dendro-reticulate. However,
Johnson) from New Zealand (Bergquist et al., 1968, 1969, 1973) although spicules coring the transverse connecting fibres are
and South Africa (Levi, 1963), it is highly unlikely these are con- reduced, as compared to those in ascending fibres at least, many
specific with the NE Atlantic species given the reported differ- reticulate tracts are definitely present such that overall mineral
ences between the highly disjunct populations. However, this skeletal structure remains reneiroid reticulate (certainly not
assumption is difficult to test given that no voucher material has plumose as in Ophlitaspongia). Moreover, ascending tracts
been located (NMNZ, AMS or MNHN). The NZ species was never of spicules are confined entirely to within the axes of fibres, not
formally described, although Bergquist & Sinclair (1968) stated diverging, protruding through fibres or as plumose columns as
that 'there is ... no doubt that this sponge is conspecific with in Ophlitaspongia. This species should, therefore, be left
O. seriata Bowerbank from Britain'. However, from the brief in Echinoclathria and the alleged similarities between it and NE
description provided by Hogg (1967) it appears that the NZ species Atlantic Ophlitaspongia species should be attributed to (i.e.,
is not related to O. papilla, but closer to Microciona aceratoobtusa hypothesised as) a secondary loss of connecting transverse spicule
(i.e., having smaller smooth echinating spicules clearly differenti- tracts, with the consequent reduction in renieroid structure, and
ated from the larger principal spicules in fibres; the mineral skele- secondary loss of echinating spicules. (2) Echinoclathria subhisp-
ton is plumose as is the fibre skeleton, with plumose bulbous fibre ida Carter, 1885d also has few transverse connecting spicule tracts,
nodes but no reticulations between fibres). It is clear that Hogg producing a nearly radial mineral skeleton. But true echinating
(1967) did not re-examine any type material, and this New Zealand spicules are present; spongin fibres are well compressed in the
population is more appropriately referred to Microciona. Levi's axial skeleton and clearly diverging in the peripheral region (not
(1963) material of '0. seriata' from South Africa probably also homogeneously isodictyal renieroid as in Ophlitaspongia);
represents a separate species of Microciona, with differentiated spicules are confined to straight tracts in the axes of fibres and not
coring and echinating subtylostyles (albiet both smooth), although diverging-plumose as in Ophlitaspongia. This species should also
468 Porifera· Demospongiae • Poecilosclerida • Microcionina • Microcionidae

be retained in Echinoclathria, with the assumption that transverse skeleton composed of oxeas (? anisoxeas) and scattered raphides (?
spicule tracts have again been secondarily lost. (3) Some of the raphidiform toxas) and sigmas (? chelae).
N Atlantic and Mediterranean species included in Echinoclathria
by Hooper (1996a: 480) are speculative, such as E. beringensis Diagnosis
(Hentschel, 1929), E. hjorti Arnesen, 1920, E. translata (Pulitzer-
Finali, 1977), but these do not clearly fit into the revised concept of Thinly encrusting growth form. Surface even and hispid.
Ophlitaspongia either (Howson & Chambers, 1999), and their pre- Choanosomal skeleton basally compressed uncored spongin fibres
cise assignment requires re-eva1uation. Earlier interpretations of lying on the substrate and a thin mesohyl containing raphides and
Echinoclathria, conceived (and grossly misinterpreted) mainly by sigmas. Subectosomal skeleton with long smooth styles embedded
Hallmann (1912) are invalid, as unfortunately was Wiedenmayer's in basal spongin, standing perpendicular to substrate and protrud-
(1989) assessment of the genus. Echinoclathria as redefined by ing through ectosome. Echinating acanthostyles present and erect
Hooper (1996a) includes only those species agreeing with the type on substrate, interspersed with choanosomal megascleres.
species, E. leporina, with most others redistributed to Holopsamma, Ectosomal skeleton with protruding styles from choanosome and
Echinochalina or Clathria (Wilsonella). In my opinion none of the bundles of raphides lying tangential to surface. Structural mega-
Echinoclathria described by Hooper (1996a) should be transferred scleres styles and acanthostyles. Microscleres raphides occurring
or returned to Ophlitaspongia (with the possible exception of these singly or in bundles (trichodragmata).
three N Atlantic and Mediterranean species).
Wiedenmayer (1989: 59) provided a succinct discussion Description of type species
of Ophlitaspongia (sensu stricto), Ophlitaspongia (of authors)
and Echinoclathria. He noted that Simpson (1968) merged Sigmeuryponjascispiculiferum (Carter, 1880b) (not illustrated)
Ophlitaspongia in Microciona based on re-examination of 'topo- Synonymy. Microcionajascispiculiferum Carter, 1880b: 44.
types' of the type species O. papilla. Wiedenmayer (1989) upheld Sigmeuryponjascispiculiferum; Hooper, 1991: 1394.
Simpson's decision (for no apparent reason, apart from the fact Material examined. None. Holotype: LFM (destroyed).
that Simpson had apparently already done the ground-work of Description (from Carter, 1880b). Thinly encrusting plate-
re-examining specimens collected from the vicinity of the type like growth form. Surface even and hispid. Ectosomal skeleton
locality, and therefore there was no further reason to question his with bundles of raphides (trichodragmata) (280/Lm long) lying
conclusions). Wiedenmayer was followed by several other authors, tangential to the substrate and also forming erect brushes.
including Hooper (1996a), correctly noting that most species Choanosomal skeleton basally compressed hymedesmioid fibres
of Ophlitaspongia were more closely related to species of with rhabdose bases of long styles (980 X 25 /Lm) and echinating
Echinoclathria or Echinochalina (i.e., renieroid reticulation, echi- acanthostyles (70/Lm long) erect on the substrate, with the former
nating spicules, etc., as defined above), whereas the type species projecting a long way through the surface; microscleres raphides
O. papilla was clearly different. Of contemporary authors Levi and (?) sigmas (8/Lm chord length) dispersed throughout mesohyl.
(1960b) and Van Soest (1984b) provided virtually monospecific Structural megascleres subectosomal styles, ectosomal oxeas and
definitions for Ophlitaspongia, which in retrospect they were cor- acanthostyles. Microscleres ? sigmas and raphides occurring singly
rect and it is this concept which is maintained here (i.e., isodictyal or in bundles (trichodragmata).
reticulate fibre skeleton, plumose mineral skeleton, homogeneous Remarks. There is no surviving material of M. jascispi-
coring spicules, no echinating spicules) and not one which includes culiferum. The holotype was destroyed in the LFM during World
Microciona or Echinoclathria as potential synonyms. War II (Ms Shirley Stone, pers. comm.), and no spicule prepara-
tions were discovered in the BMNH. Therefore, the type species of
Distribution Sigmeurypon is only known from Carter's (1880b) brief original
description. It is possible that raphides recorded by him are raphid-
Restricted to the NE Atlantic. iform toxas, which are known to occur in several microcionids
(e.g., Hooper et al., 2000). Furthermore, it is also possible that
'sigmas' recorded by Carter (1880b) in this species are actually
SIGMEURYPON TOPSENT, 1928 (INCERTAE SEDIS) examples of sigmoid isochelae. These possibilities prompted
de Laubenfels (1936a: 110) to refer this species to his genus
Synonymy Damoseni in the family Microcionidae, but it is not possible to con-
firm or refute those ideas without checking type material. If these
Sigmeurypon Topsent, 1928c: 59. assumptions are correct then it is probable that it is no more than
a thinly encrusting Clathria (Microciona). However, the species
Type species and genus are incertae sedis with no resolution possible without
new material.
Microciona jascispiculiferum Carter, 1880b (by original
designation). Distribution

Definition Monotypic, from Sri Lanka.

? Microcionidae with hymedesmioid spongin fibres cored by

erect long styles and acanthostyles, with a tangential ectosomal
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Raspailiidae Hentschel, 1923

John N.A. Hooper

Queensland Museum, P.O. Box 3300, South Brisbane, Qld, 4101, Australia. (JohnH@qm.qld.gov.au)

Raspailiidae Hentschel (Demospongiae, Poecilosclerida), including Euryponidae Topsent, contains 60 nominal genera of which 19 are
valid (one incertae sedis), with 7 subgenera, and approximately 270 named species worldwide, mainly from shallow waters and a few
from abyssal depths. Species have a typically hispid surface, with genera differentiated mainly on the basis of three morphological char-
acters: skeletal architecture ranging from axial compression to reticulate, plumo-reticulate or plumose skeletons; the presence or absence
of a specialized ectosomal skeleton (apomorphic for the family, whereby small ectosomal megascleres form bouquets surrounding the
longer choanosomal or subectosomal megascleres that penetrate the surface); and geometric modifications to echinating megascleres
(with about 15 distinct morphologies recognised). Five new subfamilies are established, based on geometry of echinating megascleres and
skeletal structure.
Keywords: Porifera; Demospongiae; Microcionina; Raspailiidae; Amphinomia; Aulospongus; Axechina; Cantabrina; Ceratopsion;
Cyamon; Cyamoninae subfam. nov.; Echinodictyinae subfam. nov.; Echinodictyum; Ectyoplasia; Endectyon; Eurypon; Hymeraphia;
Lithoplocamia; Plocamioninae subfam. nov.; Plocamione; Raspaciona; Raspailia; Raspailiinae; Rhabdeurypon; Thrinacophora;
Thrinacophorinae subfam. nov.; Trikentrion; Waltheramdtia.
- - - - ..- - . - - . - - - - - . - - . - - - -

DEFINITION, DIAGNOSIS, SCOPE referred to Dictyonellidae (Halichondrida) and Microcionidae,

respectively (see chapters by Van Soest et al., and Hooper, this
Synonymy volume).
Approximately 270 species have been published worldwide,
Raspailiidae Hentschel, 1923. Euryponidae Topsent, 1928c. 56 from Australia alone, with a potential worldwide fauna of about
500 species given that many undescribed species have been discov-
Definition ered and many more probably await discovery, particularly thinly
encrusting taxa (Hooper, unpublished collections). For example,
Microcionina with a special category of smaller ectosomal Hymeraphia is prevalent on soft sediments of deeper coastal waters
styles, oxeas or anisoxeas forming discrete bouquets around the in the North Atlantic with few records from elsewhere, so it is pos-
protruding larger styles or oxeas. sible that there are many similar species throughout the other ocean
systems.
Diagnosis
History and biology
Encrusting, massive, lobate, fan-shaped or branching growth
forms, usually with a very hispid surface. Specialised ectosomal Hentschel's (1923) initial concept of Raspailiidae allowed for
skeleton typically present, consisting of small thin styles, oxeas or the inclusion of species with meniscoid microscleres (chelae,
anisoxeas forming bouquets around long styles or oxeas that pene- sigmas), whereas Topsent (1928c) restricted the family to include
trate the surface. Choanosomal skeletal structure is typically reticu- only species lacking microscleres or having only raphides. The
late with a compressed axial region and plumo-reticulate extra-axial synonymy of Euryponidae and Raspailiidae was proposed by
(peripheral) skeleton, but may range from hymedesmioid and Bergquist (1970: 31), although Levi (1973: 608) subsequently
microcionid in encrusting taxa, to plumose, radial or evenly reticu- recognised both as distinct families. Both families have a 'ras-
late (non-compressed) skeletons in other taxa. Spongin fibres usu- pailiid' ectosomal skeleton, considered here to be their primary
ally completely enclose coring spicules (choana somal styles, oxeas apomorphy, whereas differences in their respective skeletal struc-
or both). Echinating acanthostyles (or modified styles) echinate tures (hymedesmioid versus axially compressed, reticulate skele-
fibres, standing perpendicular to them. Microscleres are usually tons) do not warrant their recognition at the family level.
absent, although a few genera have raphides in bundles (trichodrag- Definitions of the family were progressively developed by
mata). Where known, reproduction is oviparous and probably Bergquist (1978: 167), Hartman (1982: 647) and Hooper (1991:
typical for this family. 1185), and the major characters used to differentiate genera were
revised by Hooper (1991) and Hooper et al. (1999).
Scope The higher systematics of the Raspailiidae has been debated
by several authors (e.g., Ridley & Dendy, 1887; Topsent, 1894c,
Sixty generic names have been included in the family at one 1928c; Dendy, 1905; Vosmaer, 1912; Wilson 1921; Bergquist,
time or another, but only nineteen ofthese are valid (Hooper, 1991, 1970; Hooper, 1991). It was initially included in Poecilosclerida
and this work), one of which is incertae sedis, and with seven sub- (Hentschel, 1923; Topsent, 1928c) but subsequently referred to
genera recognised. Hooper (1991) also included Tethyspira and Axinellida on the basis that many taxa had compressed axial skele-
Sigmeuryon as incertae sedis in Raspailiidae whereas these are tons and clearly differentiated radial, plumose or plumo-reticulate

469
470 Porifera· Demospongiae • Poecilosclerida • Microcionina • Raspailiidae

extra-axial skeletons, and where known, species were thought surveyed in the North Atlantic and Mediterranean (e.g., Topsent,
to be oviparous (e.g., Bergquist, 1970, 1978; Hartman, 1982; 1928c; Vosmaer, 1935; Boury-Esnault & Lopes, 1985), Caribbean
Wiedenmayer, 1989). Hooper (1991) and Hooper et al. (1992) sub- (see review in Hooper et al., 1999), southeast Indonesian (Hentschel,
sequently returned the family to Poecilosclerida based on morpho- 1912), New Zealand (Bergquist, 1970), Australian (Hooper, 1991)
logical and biochemical similarities to the Microcionidae (Hooper, and New Caledonian regions (Hooper & Levi, 1993), although many
1991), and it now forms one of the four families of Microcionina other publications record one or few species from diverse localities.
(Hajdu et al., 1994a). The presence of homologous echinating Deeper-water species have been infrequently recorded by Ridley &
acanthostyles in both Raspailiidae and Microcionidae is an obvious Dendy (1887) and Levi & Levi (1983b).
morphological clue to their common ancestry, but this feature is
interpreted as representing the retention of an ancestral character Differences with similar families
state, and as such the group cannot be defined solely on this basis.
The alternative classification, in which Raspailiidae is Raspailiidae shares the symplesiomorphy of echinating
assigned to a now more-or-Iess defunct concept of 'Axinellida', acanthostyles with other families of Microcionina. Furthermore,
rests solely on supposed homologies in skeletal architecture some taxa (e.g., Raspailia s.s.) show identical construction to
between representative taxa (i.e., the possession of a differentiated some microcionids (e.g., Axociella s.s.), and it has been suggested
axial and extra-axial skeleton and a compressed axial skeleton - (Hooper, 1991) that a compressed axial skeleton, and differentiated
termed here an 'axinellid' skeleton). However, it is obvious from axial and extra-axial skeletons (used to define 'axinellids') has
examination of all Raspailiidae genera that the family demonstrates been independently acquired several times throughout the Porifera.
a wide range of architectural types, and many of these structures This interpretation de-emphasises the primary importance of skele-
show only vague or no similarities with 'axinellid' forms at all. In tal structure in sponge systematics. This, together with chemotaxo-
fact the reverse is true: many skeletal types in the family clearly nomic evidence (Hooper et al., 1992) was used to justify the return
show structural relationships with other poecilosclerids. For exam- of Raspailiidae to the Poecilosclerida (Hooper, 1991), although
ple, the raspailiid genus Aulospongus is peculiar in having tubular, these data require corroboratory support from more contemporary
plumose, skeletal fibre bundles (Hooper, 1991; Hooper et al., 1999), molecular studies.
whereas identical fibres are known for the Clathria 'parthena' A generic classification of the Raspailiidae centres around
species group of Microcionidae (Hooper, 1996a). Similarly, ras- three principal diagnostic features: skeletal architecture, presence
pailiids with typical 'axinellid' skeletal structure (e.g., Raspailia or absence of a specialized ectosomal skeleton, and modifications
(Raspailia) vestigifera) have structural counterparts in the to echinating megascleres (with about 15 distinct morphologies,
Microcionidae (e.g., Clathria (Axosuberites) canaliculata which several apomorphic for various genera). These many examples,
has a distinctive compressed axis, plumose-radial extra-axis, in comparing microcionid and raspailiid species with similar features,
addition to isochelae microscleres). Several other homologies support the hypothesis that these two families are most closely
between Raspailiidae and Microcionidae are also apparent. related amongst the Microcionina.
Incorporation of detritus into spicule-bearing fibres is a common
trait amongst the Poecilosclerida (e.g., Clathria (Wilsonella», and it is Previous reviews
also known in several Raspailiidae (e.g., Raspailia (Clathriodendron),
Echinodictyum arenosum). Several raspailiids have completely lost Bergquist (1970), Levi (1973), Hooper (1991), Hooper et al.
their echinating megascleres (e.g., Raspailia (Parasyringella), (1999).
Ceratopsion, Thrinacophora), and this is also a feature of some
microcionids (e.g., Clathria (Axosuberites), C. (Isociella». One ras-
pailiid (Aulospongiella monticularis) incorporates echinating acan- PROPOSAL FOR SUBDIVISION OF RASPAILIIDAE
thostyles secondarily into fibres (Hooper, 1991; Hooper et al.,
1999), which is also a feature peculiar to the microcionid Clathria Phylogenetic relationships between genera of Raspailiidae
'phorbasiformis' species group (Hooper, 1996a). Some raspailiids were postulated by Hooper (1991), recognising three major groups
lack the specialized ectosomal structure characteristic of the family (based on skeletal structure) and six subgroups based on differ-
(e.g., Raspailia australiensis), and have instead a continuous crust ences in spicule geometries forming their choanosomal (axial),
of spicules also seen in a particular group of Microcionidae subectosomal (extra-axial) and ectosornal skeletons, in addition to
(Clathria (Thalysias». Thus, morphological comparisons alone echinating spicule morphologies. Synapomorphy for all these
suggest that raspailiids are more closely related to Poecilosclerida, groups is the possession of a specialized (raspailiid) ectosomal
such as Microcionidae, than they are to axinellids, such as skeleton, and symplesiomorphy is the possession of echinating
Hemiasterellidae and Axinellidae, and there is also some biochemi- acanthostyles: most but not all species have these features, and
cal support for this idea (e.g., Hooper et al., 1992). where absent their obvious affinities to Raspailiidae are indicated
Revisions of New Zealand, Australian and New Caledonian by one or more other features shared with other raspailiids.
species were undertaken by Bergquist (1970), Hooper (1991), and Five of these six subgroups are elevated here to subfamilies
Hooper & Levi (1993), respectively. Several biochemical charac- (based predominantly on acanthostyle geometry and skeletal struc-
teristics of the family have been described by Hooper et al. (1992), ture), whereas one subgroup is poorly resolved and incorporated in
who noted a disproportionately high level of antibiotic activity the nominotypical subfamily. Subgroups delineated by Hooper
against several human pathogens, as compared to other families of (1991) are as follows.
sponges sampled. 'Raspailoids', including Raspailia, Ectyoplasia and
Species are widely distributed, from shallow-waters, where Endectyon, with a noticeably compressed axial skeleton composed
they are more diverse (Hooper, 1991), to at least 2460m depth of criss-cross reticulation fibres and/or spicules. Echinating acan-
(Hartman, 1982). Shallow water species have been reasonably well thostyles are microcionid-like club-shaped with small granular
Porifera· Demospongiae • Poecilosclerida • Microcionina • Raspailiidae 471

or erect spines, ranging to club-shaped with strongly recurved or lack any marked differentiation between axial and extra-axial
clavulate spines on the basal and distal ends of spicules, and two skeletons. There are no unique echinating spicule geometries
subgenera have echinating acanthorhabdostyles. One species has within genera, showing a similar range of geometries as the 'ras-
raphides in trichodragmata. This subgroup is elevated to Raspailiinae pailoids'. For example, Aulospongus has echinating rhabdostyles
Hentschel, 1923. similar to Raspailia (Raspaxilla) and Endectyon (Hemectyon);
'Axinelloids', including Ceratopsion, Axechina and Eurypon is essentially an encrusting Raspailia with hymedesmioid
Thrinacophora, lacking echinating megascleres altogether. Two skeletal arrangement (as opposed to a compressed axial reticulate
genera have raphides in trichodragmata. Differentiation between skeleton in the latter), and its echinating acanthostyles are identical
axial and extra-axial skeletons is more prominent. This subgroup is to those seen in many Raspailia. Several species of Eurypon and
elevated to Thrinacophorinae subfam. nov. one of Aulospongus have raphides in trichodragmata. These genera,
'Triaenoids', including Trikentrion and Cyamon, having therefore, are justifiably included in the subfamily Raspailiinae.
echinating spicules modified to sagittal mono-, di-, tri-, tetr- or 'Myxilloids', including only Amphinomia and Echinodictyum,
pentactinal acanthose megascleres (acanthoplagiotriaenes). These having regularly reticulate choanosomal skeletal structure, with
taxa share other common features including the possession of only extra-axial skeleton vesigial or virtually absent, and all but one
a slightly compressed axial skeleton composed of plumo-reticulate species lack ectosomal specialisation. Echinating megascleres are
spicule tracts forming more-or-Iess rectangular, renieroid meshes, microcionid-like club-shaped acanthostyles. This subgroup is ele-
or one that is reduced to a basal spongin layer in encrusting species vated to Echinodictyinae subfam. nov.
with erect spicules in hymedesmioid arrangement. One species has 'Plocamoids', includes Lithoplocamia and Plocamione,
raphides in trichodragmata. This subgroup is elevated to Cyamoninae having acanthostrongyles or acanthotylostrongyles forming the
subfam. nov., and to it is added the monotypic Waltherarndtia that choanosomal skeleton. This subgroup is elevated to Plocamioninae
has presumably lost these triaenes but retains other features com- subfam. nov. Cantabrina is excluded from this hierarchy. It is
mon to these taxa incertae sedis, linked to the Raspailiidae only tenuously by its pos-
'Hymedesmioid-microcionoids', including Aulospongus, session of smooth rhabdostyles (which are rare and possibly con-
Raspaciona, Eurypon, Rhabdeurypon and Hymeraphia, have loosely taminants), and may not belong in Raspailiidae at all. If it is
aggregated or plumose skeletal structure, or hymedesmioid structure eventually excluded from Raspailiidae it most closely resembles
in encrusting taxa (i.e., megascleres erect on basal spongin), and the Halichondriidae in skeletal structure.

KEYS TO RASPAILIIDAE

Key to Subfamilies

(I) Echinating megascleres present .......................................................................................................................................................... 2

Echinating megascleres absent ............................................................................................................................... Thrinacophorinae
Echinating megascleres absent .......................................................................................... Raspailiinae (Raspailia (Parasyringella»
(2) Echinating megascleres club-shaped or rhabdose ............................................................................................................................... 3
Echinating megascleres modified to sagittal acanthoplagiotriaenes ................................................................................ Cyamoninae
(3) Choanosomal skeleton cored by oxeas or styles ................................................................................................................................. 4
Choanosomal skeleton cored by acanthostrongyles or acanthotylostrongyles ('peg-top' spicules) ............................ Plocamioninae
(4) Choanosomal skeleton with greater or lesser differentiation between axial and extra-axial regions, ectosome with specialised
raspailiid skeleton .... ............... ................. ... ........ ............... ........... ...... ........ ... .......... ................. .......... ...... .... ...... .............. Raspailiinae
Choanosomal skeleton regularly reticulate, extra-axial skeleton vesigial or virtually absent, and all but one species lack ectosomal
specialization ................................................................................................................................... '" ................ ........ Echinodictyinae

Key to Genera

(1) Echinating spicules are microcionid-like, club-shaped, with rounded or sharp points, subtylote bases, and with evenly or unevenly
distributed spines .. ........ ....... ... ........ ... ...... ....... ... ........... .... .................. ....... .... ....... ........ ........ .................................... .......... .......... ....... 2
Echinating spicules are club-shaped with clavulate points; axial and extra-axial skeletons are composed of a single category of undif-
ferentiated choanosomal megascleres ................................................................................................................................ Ectyoplasia
Echinating spiCUles are acanthose, club-shaped or strongylote, with strongly curved hooks on the base and shaft (cladotylote), and
these spicules are usually confined to a particular region within the skeleton .................................................................... Endectyon
Echinating spicules are acanthostyles with smooth rhabdose bases, and large recurved spines are distributed over
the shaft ......................................................................................................................................................................... '" Aulospongus
Echinating spicules are smooth rhabdostyles (rare and possibly contaminants) ...................................... Cantabrina (incertae sedis)
Echinating spicules are acanthostyles with bulbous tylote bases, with or without spines on the points and other modifications to the
distal portion .... ....... ........... .......... ................. .......... ................. ....... ....... ....... .... ............ ................ ........... ...... .......... ......... Hymeraphia
Echinating spicules are absent, but diactinal acanthorhabds form a compact basal mass throughout
the skeleton ... ... ......... ... ... ... ........... ... ............. .......... ..... ............. .......... .... .......... .............................. ... ................ ........... Rhabdeurypon
Echinating spicules are sagittal monact-, diact- or tetractinal (acanthoplagiotriaenes) with only one
spined ray ...................... '" ... ..................................... .... ....... .......... ...... ........... .......... ...... .......... ...... ..... ...... .......... ................ Trikentrion
472 Porifera· Demospongiae • Poecilosclerida • Microcionina • Raspaillidae

Echinating spicules are sagittal tetract- or pentactinal (rarely with fewer rays) (acanthoplagiotriaenes) with all or most
rays spined ............. ...... ...... .......... ...... ....... ... .......................... ...... .............. ................... .......... ..... ................. ... ....... ... ... ........... Cyamon
Echinating spicules are absent ......................................................................................................................................................... 3, 8
(2) Choanosomal skeleton consists of slightly axially compressed reticulation of spongin fibres and/or spicules .......... Raspailia ... 3
Choanosomal skeleton microcionid, basally compressed fibres lying on the substrate, with or without ascending fibre nodes ....... 6
Choanosomal skeleton reticulate without any trace of axial compression .......................................................................................... 7
Choanosomal skeleton consists of loosely aggregated or plumose axial fibres ................................................................. Raspacwna
(3) With echinating acanthostyles ............................................................................................................................................................. 4
Without echinating acanthostyles .............................................................................................................. Raspailia (Parasyringella)
(4) Choanosomal skeleton consists of an axially compressed reticulation of spongin fibres and/or spicule tracts, with at least some degree
of differentiation between axial and extra-axial skeletons .... .... ..... .... ........ ... .................. ... ... ... ....... ......... ....... ... ... .......... ........... ....... 5
Choanosomal skeleton consists of a reticulation of spongin fibres and/or spicule tracts, without any trace of axial compression, and
with reduced differentiation of the axial and extra-axial skeletons ........................................................ Raspailia (Clathriodendron)
(5) Echinating spicules are club-shaped acanthostyles, with subtylote bases and straight shafts ............................ Raspailia (Raspailia)
Echinating spicules are acanthose rhabdostyles ............................................................................................... Raspailia (Raspaxilla)
Echinating spicules have smooth, swollen, tylote bases ......................................................................... Raspailia (Hymeraphiopsis)
(6) Spicules in the axial or basal skeleton are acanthostrongyles or acanthotylostrongyles ................................................... Plocamwne
No true choanosomal megascleres occur in the axial skeleton, although large extra-axial spicules and echinating acanthostyles may
be embedded in spongin fibres ............................................................................................................................................... Eurypon
(7) Choanosomal spicules are styles or subtylostyles with spines on both basal and distal ends, forming a renieroid
reticulate skeleton ........................ ... ............... ... ...... .......... ...... ...... .... .......... .................. ............... ........... ................ ......... Amphinomia
Choanosomal spicules are exclusively oxeas ...... ... ...... ....... ... ...... ....... ... ................ ...... ............ .......... ... ....... ............. ... Echinodictyum
Choanosomal spicules are acanthostrongyles or acanthotylostrongyles forming a secondary (sub)
isodictyal reticulation .................................................................................................................................................... Lithoplocamia
(8) Choanosomal skeleton consists of dense axial compression of criss-crossed spiCUles, without
axial fibres .. ....... ..................... ...... .......... .......... .... ... ....... ... ...... ... ....... ... ... .... ....... ..... ... ............................. ... ...... ...... ...... Thrinacophora
Choanosomal skeleton with a renieroid reticulate skeleton of multi spicular primary fibres interconnected by uni- or paucispicular
secondary fibres, forming elongate meshes, slightly more compressed at the axis than in the periphery ................... Waltherarndtia
Choanosomal skeleton consists of axially compressed reticulation of spongin fibres and/or spicule tracts ...................................... 9
(9) Spicules in the axial skeleton are sinuous styles or anisoxeas .......................................................................................... Ceratopswn
Spicules in the axial skeleton include both spined oxeas and smooth anisoxeas ... ........... ... ... .......................................... .... Axechina

SUBFAMILY RASPAILIINIAE NARDO, 1833 Definition

Definition Raspailiidae with a more-or-less compressed axial skeleton

and a radial, plumose or simply reticulate extra-axial skeleton,
Raspailiidae with echinating megasclere geometry ranging from with choanosomal spicules consisting of 2-3 or more different
microcionid-like club-shaped acanthostyles with small granular or size classes (styles and/or oxeas), and echinating acanthostyles
erect spines, to club-shaped with strongly recurved or clavulate spines microcionid-like or secondarily modified.
on the basal and distal ends of spicules, to acanthose rhabdostyles.
Diagnosis
Scope
Arborescent, lobo-digitate to massive growth form, typically
Eight genera and seven subgenera. Raspailia (R. (Raspailia), with cylindrical branches and basal holdfast. Surface even or
R. (Clathriodendron), R. (Raspaxilla), R. (ParasyringeUa), rugose, often optically hispid. Choanosomal skeleton always with
R. (Hymeraphiopsis», (Aulospongus, Ectyoplasia, Endectyon fibres and spiCUles in distinct tracts: axial skeleton typically com-
(E. (Endectyon), E. (Hemectyon», Eurypon, Hymeraphia, pressed with widely spaced reticulate fibres cored by styles, but
Raspaciona and Rhabdeurypon. degree of compression varies considerably between species. Extra-
axial skeleton typically plumo-reticulate, with extra-axial spicule
tracts standing perpendicular to axial skeleton, cored by large
RASPAILIA NARDO, 1833 styles or oxeas which ascend to and poke through surface in uni- or
paucispicular brushes, although this may be reduced to simple
Synonymy plumose structure. Peripheral spicule tracts may be multispicular
or reduced to brushes of subectosomal spicules embedded in the
RaspeUa Nardo, 1833: 522. Raspailia Nardo, 1847a: 3 (nom. subdermal region. Ectosome typically with specialized skeleton
emend.). See subgenera for full synonymy. of small styles grouped into brushes standing perpendicular to
surface, surrounding bases of protruding extra-axial megascleres,
Type species but these may be secondarily lost. Fibres usually echinated by
acanthostyles or modified forms, sometimes secondarily lost.
RaspaiUa typica Nardo, 1833 (by monotypy). Structural megascleres styles, oxeas or anisoxeas, typically 3,
Porifera· Demospongiae • Poecilosclerida • Microcionina • Raspailiidae 473

sometimes 2, distinct size categories; echinating acanthostyles undescribed (Hooper, unpublished data). Species are distributed
morphologically similar to microcionid sponges. Microscleres worldwide, predominantly in shallow waters.
absent.

Remarks SUBGENUS RASPAILIA NARDO, 1833

The genus Raspailia was originally reviewed by Vosmaer Synonymy

(1912, 1935) and subsequently by Hooper (1991), with both revi-
sions based on restricted faunas (European and Australian species, Raspellia Nardo, 1833: 522; Burton 1937: 33. Raspailia
respectively). Vosmaer's (1935) revision must be accepted with Nardo, 1847a: 3 (nom. emend.); Schmidt, 1862: 59; 1866a: 14;
caution as he synonymized about 20 nominal species, including Ridley & Dendy, 1887: 188; Hanitsch, 1889: 161; Lendenfe1d,
Nardo's type species, under Raspailia hispida (Montagu), an act 1890b: 401; Topsent, 1894c:16; Pick, 1905: 19; Hentschel, 1912:
which very few authors subsequently accepted. Wilson (1921) 4l3; Vosmaer, 1912: 313; Wilson, 1921: 54-60; Vosmaer, 1935:
made a better attempt in redefining the genus. From an examina- 766; Bergquist, 1970: 26; Hooper, 1991: 1195. [Rasalia] Gray,
tion of a number of species he arrived at a broad definition for I 867a: 522 (lapsus). [Raspalia] Gray, 1867a: 523 (lapsus). [Abila]
Raspailia s.l., and simultaneously succeeded in conveying the Gray, 1867a: 522 (preocc.) (Not Abila Gray, 1867a: 539). Abilana
enormous difficulty involved in defining this large and diverse Strand, 1928: 33 (replacement name for [Abila] Gray;
group. There is obviously a high diversity of character states found de Laubenfels, 1936a: 102). Valedictyum de Laubenfels, 1936a:
in the genus, encompassing most characters: growth forms of 102. Bethia de Laubenfels, 1936a: 108. Taxonomic decision for
Raspailia range from massive, branching to lobate; skeletal struc- synonymy: Hooper (1991), Hooper & Levi (1993), and this work.
tures vary from compressed axis/radial extra-axial skeleton, to
reduced extra-axial, plumo-reticulate extra-axial, reduced plumose Type species
extra-axial, or simply reticulate skeletons (nominal genus
Clathriodendron); structural megascleres may be exclusively sty- Raspailia typica Nardo, 1833 (by monotypy).
lote, rhabdostyles (nominal genus Axinectya), stylote and oxeote
together, or exclusively oxeote; ectosomal structure ranges from Definition
the 'raspailiid' condition (protruding extra-axial (subectosomal»
megascleres surrounded by special ectosomal brushes, to extra- Raspailia with microcionid-like acanthostyles, myxillid-like
axial megascleres protruding through the surface but without acanthostyles or thin vestigial acanthostyles.
any specialized ectosomal megascleres, or with only special der-
mal spicules scattered over the surface, or lacking ectosomal spe- Diagnosis
cialisation at all (nominal genus Echinaxia); and geometry of
echinating spicules ranging from cylindrical microcionid-like Digitate and branching Raspailia, usually with a highly hispid
spicules, acanthose rhabdostyles (nominal genus Raspaxilla), or surface; choanosomal skeleton composed of distinctive axial and
secondarily lost (nominal genera Raspailopsis, SyringeZZa and extra-axial components; axial skeleton compressed with reticulate
Parasyringella). spongin fibres cored by multispicular tracts of principal styles or
Hooper (1991) used subgeneric taxa to overcome this prob- sometimes oxeas, and echinated by acanthostyles; extra-axial
lem, recognising five subgenera differentiated by the geometry and skeleton primarily plumose or radial, with longer principal spicules
spination of echinating acanthostyles. forming mu1tispicular tracts and interconnected by sparse uni- or
Another nominal genus referred here to Raspailia was paucispicular fibres; extra-axial tracts eventually protruding
overlooked by Hooper (1991). Bethia de Laubenfels, 1936a: 108 through the surface, with the larger spicules surrounded by bou-
(type species Dictyocylindrus laciniatus Carter, 1979b: 296, by quets of ectosomal styles or anisoxeas at the point of their insertion
original designation), is clearly as raspailiid, and from its descrip- to the surface. Megascleres include styles (and/or or less com-
tion belongs to Raspailia (Raspailia). The type species, monly oxeas or oxeote modifications to styles) of at least 2-3 size
from Mauritius, is described as a hemispherical, long radiated classes, varying from robust and long to small, sinuous and vestig-
laciniate sponge with extremely hispid surface, a compressible tex- ial, and echinating acanthostyles that are microcionid-like with
ture, has subradiate plumo-reticulate architecture (reminiscent of a subtylote base, usually a straight shaft, and evenly spined or with
Suberitidae according to de Laubenfels, 1936a) with indistinct an aspinose region proximal to the basal swelling.
spongin fibres/spicule tracts becoming more plumose towards
the periphery, cored by very long choanosomal subtylostyles
(over 4mm long, about 35 fLm wide), protruding through the sur- Description of type species
face for long distances, small ectosomal oxeas (about 550 fLm long
and 17 fLm wide), and short echinating acanthostyles with well Raspailia typica Nardo, 1833 (Fig. 1).
developed slightly swollen base, slightly rounded point, and spines Synonymy. Raspailia typica Nardo, 1833: 522; Raspalia
concentrated mainly at extremities (about 95 fLm long, 17 fLm viminalis Schmidt, 1862: 59.
wide), sparsely echinating fibres. Microscleres are absent. Material examined. Ho10type: Unknown. Fragments of
Schmidt's specimen MNHN DCL 1237L, BMNH 1867.3.11.8 -
Distribution Adriatic Sea. Schizotype of R. viminalis. MNHN DCL 1238L -
Sebenico, Adriatic Sea.
There are more than one hundred described species of Description. Erect, digitate, branching growth form;
Raspailia, and at least the same number known but which remain choanosomal skeleton with differentiated axial and extra-axial
474 Porifera· Demospongiae • Poecilosclerida • Microcionina • Raspailiidae

B c
A I
D E
I

Fig. 1. Raspailia (Raspailia). A-E, R. (Raspailia) typica (schizotype BMNH 1867.3.11.8). A-B, choanosomal principal style and subtylostyle (scale
400 !Lm). C, base of principal subtylostyle (scale 25 !Lm). D, ectosomal style and anisoxea (scale 150 !Lm). E, echinating acanthostyle (scale 30 !Lm).
F-H, R. (Raspailia) vestigifera Dendy. F, specimen NTMZ910 (scale 5 em). G, choanosomal skeleton (scale 500 !Lm). H, ectosomal skeleton (scale 300 !Lm).

skeletons; compressed axis composed of several, thin, parallel, (1140-1950 X 11-23 ,.,..m), ectosomal styles or anisoxeas slightly
multispicular tracts running longitudinally through branches sepa- curved, tapering to hair-like point at one end (470-595 X 3-9,.,..m),
rated by relatively wide meshes; extra-axial skeleton more-or-less echinating acanthostyles with subtylote base and evenly spined
at right angles to axis, running horizontally through branches, (88-112 X 4-7 ,.,..m); microsc1eres absent.
composed of slender, slightly plumose tracts 1-5 principal spicules Remarks. See Remarks for the genus. Re-examination of
abreast bound by light collagen, interconnected by unispicular type material confirms the synonymy between Raspailia and Abila
tracts of principal spicules together forming a loose, wide-meshed (type species RaspailiaJreyerii Schmidt, 1862: 60, by monotypy)
reticulation; extra-axial tracts ascending to surface and terminating and Valedictyum (type species RaspaiUa vestigifera Dendy, 1896,
in single principal spicule protruding through surface; spicule by monotypy).
tracts echinated by acanthostyles, particularly in axis; specialised
ectosomal skeleton consists of small ectosomal styles or oxeas
forming brushes around single protruding principal style at point of Distribution
insertion in surface; principal styles long slender curved or some-
times sinuous, with subtylostylote or more well-developed tyles Worldwide, shallow seas.
Porifera· Demospongiae • PoeciioscIerida • Microcionina • Raspailiidae 475

SUBGENUS RASPAXILIA TOPSENT, 1913 persist all the way to the surface; echinating rhabdostyles pre-
dominant in extra-axial skeleton; megascleres include long cho-
Synonymy anosomal styles, slightly curved centrally or straight, with evenly
rounded, smooth, non-rhabdose bases (550-900 X 10-16 f.Lm);
Raspaxilla Topsent, 1913b: 616; Bergquist, 1970: 28-30; extra-axial styles long, thick, straight or slightly curved, entirely
Hooper, 1991: 1195, 1245; Hooper et al., 1999: 673. Echinaxia smooth (1100-1450 X 12-18 f.Lm); ectosomal styles wispy, rhaphid-
Hallmann, 1916b: 543; Hallmann, 1917b: 391; de Laubenfels, iform, straight, centrally curved or sinuous (450-650 X 2-3 f.Lm);
1936a: 102; Bergquist, 1970: 30; Hooper, 1991: 1195; Hooper echinating rhabdostyles moderately long, with slightly rhabdose
et al., 1999: 681. Axinectya Hallmann, 1917b: 393; Hooper, 1991: and subtylote bases, entire spicule smooth or small, erect spines on
1195; Hooper et al., 1999: 688. Taxonomic decision for synonymy: apical two-thirds of spicule only (140-370 X 8-18 f.Lm); micro-
Hooper (1991), Hooper et al. (1999). scleres absent.
Remarks. Raspai/ia (Raspaxilla) differs from typical
Type species Raspailia in having rhabdose echinating acanthostyles. It differs
from other rhabdose raspailiids, notably Aulospongus, in having
Raspaxilla phakellina Topsent, 1913b (by monotypy). echinating rhabdostyles geometrically very different from the usu-
ally longer choanosomal styles (the latter without any basal rhabd);
Definition extra-axial styles forming a radial skeleton perpendicular to the
axis; and well differentiated axial and extra-axial skeletons (the
Raspailia with echinating rhabdostyles geometrically very former compressed, the latter plumo-reticulate and/or radial).
different from the usually longer choanosomal styles (the latter Placement of all species, however, is not always straightforward
without any basal rhabd); extra-axial styles forming a radial skele- given that some taxa may lose certain characters (e.g., extra-axial
ton perpendicular to the axis; and well differentiated axial and skeleton becomes reduced to single long extra-axial spicules
extra-axial skeletons (the former compressed, the latter plumo- embedded in axis and forming a radial skeleton; or the extra-axial
reticulate and/or radial). spicules are lost completely). There is also a correlation between
the localisation of echinating rhabdostyles in the peripheral skele-
Diagnosis ton and the degree of axial compression. In species with very com-
pressed skeletons the extra-axial skeleton is reduced to single long
Digitate, bushy, lobate, lamellate, arborescent, flabellate and extra-axial spicules (without reticulate connections) and the echi-
vasiform Raspai/ia; with echinating acanthose rhabdostyles; larger nating rhabdostyles are 'pushed' into the ectosomal skeleton where
choanosomal styles completely smooth, without any basal rhabd, they form brushes or a continuous palisade of rhabdostyles
geometrically different from smaller acanthose echinating (Hooper et al., 1999). Echinaxia (type species Axinella frondula
spicules; axial skeleton well differentiated from extra-axial skele- Whitelegge, 1907, by original designation) and Axinectya (type
ton; axial skeleton compressed, composed of reticulate tracts cored species Axinella mariana Ridley & Dendy, 1886, by original
by choanosomal styles; extra-axial skeleton plumo-reticulate, with designation) are clearly synonymous with Raspaxilla.
plumose ascending tracts interconnected by transverse tracts both
cored by choanosomal styles (forming a reticulation), or reduced Distribution
to radial tracts of single long extra-axial styles embedded in and
perpendicular to axis, protruding through the surface; echinating Sixteen species are currently assigned to Raspai/ia (Raspaxi/la),
rhabdostyles more abundant in peripheral skeleton. with a wide geographic distribution, ranging from the Indo-west
Pacific (north and south New Zealand, northwest Australia,
Description of type species northern Great Barrier Reef, central NSW, New Caledonia, Japan,
Micronesia), Caribbean (North Carolina), central east Pacific and the
Raspailia (Raspaxil/a) phakellina Topsent, 1913b (Fig. 2). antarctic-subantarctic region (Hooper et al., 1999).
Synonymy. Raspaxilla phakellina Topsent, 1913b: 617;
RaspaiUa (Raspaxilla)phakellina; Hooper, 1991: 1196.
Material examined. Holotype: MOM (fragment MNHN SUBGENUSCIATHRIODENDRON
DT1614) - Antarctica. Other material. BMNH 1928.2.15.781a, LENDENFELD, 1888
846a - Falkland Islands.
Description. Erect, digitate, arborescent, with enlarged Synonymy
basal holdfast attachment and branching in one plane; surface
slightly conulose, hispid; ectosomal skeleton with long extra-axial Clathriodendron Lendenfeld, 1888: 215; Kirk, 1911: 579;
styles protruding through the surface, surrounded at their bases by Hentschel, 1911: 383; Hallmann, 1912: 295; Topsent, 1894c: 19; de
wispy bundles of ectosomal anisoxeas forming stellate bundles Laubenfels, 1936a: 102; Bergquist, 1970: 30. Dictyocy/indrus
nearly parallel to the surface; axial and extra-axial skeletons well Bowerbank, 1862b: 1108 (in part); Gray, 1867a: 519; Topsent, 189Oc:
differentiated; choanosomal skeleton with a compressed, strongly 289; de Laubenfels, 1936a: 102. (Not Dictyocylindrus; Carter, 1879b:
reticulate axis composed of multispicular fibres cored by 297). Taxonomic decision for synonymy: Hooper (1991).
choanosomal styles and echinated sparsely by echinating rhab-
dostyles; extra-axial skeleton plumo-reticulate, with ascending Type species
multispicular fibre bundles cored by choanosomal styles and pro-
fusely echinated by rhabdostyles, diverging towards the periphery, Clathriodendron arbuscula Lendenfeld, 1888 (by subsequent
interconnected by transverse pauci- or multi spicular fibres which designation; Hallmann, 1912: 295).
476 Porifera· Demospongiae • PoeciloscIerida • Microcionina • Raspailiidae

Fig. 2. Raspai/ia (Raspaxilla). A-F, R. (RaspaxiUa) phakellina Topsent (schizotype). A, choanosomal style (scale 100 fLm). B, echinating rhabdostyle
(scale 50 fLm). C, extra-axial style (scale 200 fLm). D, ectosomal styles (scale 50 fLm). E, ectosomal skeleton (scale 250 IJ.m). F, choanosomal fibre bundle
(scale lOOfLm). G-I, R. (Raspaxi/la) wardi Hooper (holotype NTMZ13l9). G, holotype (scale 3cm). H, axial skeleton (scale 40fLm). I, echinating
rhabdostyle (scale 20 fLm).

Definition Description of type species

Reduced Raspailia lacking any axial compression or any dif- Raspailia (Clathriodendron) arbuscula Lendenfeld, 1888
ferentiation between axial and extra-axial skeletons, but retaining (Fig. 3).
two or more forms of choanosomal structural megascleres. Synonymy. Clathriodendron arbuscula Lendenfeld, 1888:
215; Echinonema anchoratum var. ramosa Lendenfeld, in part (as
Diagnosis Echinonema ramosa); Whitelegge 1901: 81; Raspailia agminata
Hallmann, 1914b: 438; Halichondria rubra var. digitata Lendenfeld,
Digitate, lobate or massive Raspailia with microconulose or 1888: 81; Clathriodenderon nigra Lendenfeld, 1888: 216 (in part).
less commonly hispid surface; choanosomal skeleton exclusively Material examined. Lectotype: AM G9045 - Port Jackson,
reticulate, with little differentiation between axial and extra-axial NSW. Holotype of C. nigra var.jacksoniana: BMNH 1887.1.24.64-
regions although peripheral skeleton may still contain the vestiges of Port Jackson, NSW.
radial or plumose tracts; spongin fibres generally well developed, Description. Shape elongate, laterally flattened, digitate
cored by multi spicular tracts of principal styles often together with fronds connected to common base; surface prominently hispid,
sand grains or other detritus, and echinated by microcionid-like acan- with arenaceous particles embedded in ectosome producing uneven
thostyles; longer styles may be present in the peripheral skeleton, conules dispersed over surface; texture harsh, branches easily flex-
protruding through the surface, sometimes surrounded by bouquets ible, relatively fragile (easily tom); ectosomal skeleton moderately
of ectosomal styles/anisoxeas, but often lacking a specialised ectoso- arenaceous; long subectosomal megascleres protrude through sur-
mal skeleton; ectosome often highly collagenous and arenaceous. face, sometimes surrounded by sparse brushes of thin ectosomal
Porifera· Demospongiae • Poecilosclerida • Microcionina • Raspailiidae 477

A B

Fig. 3. Raspailia (Clathriodendron). A-Eo R. (Clathriodendron) arbuscula (lectotype AMG9045). A, subectosomal extra-axial style and anisoxea (scale
300 fLm). Bo ectosomal auxiliary anisoxea (scale 100 fLm). Co echinating acanthostyles (scale 25 fLm). Do skeleton (scale 500 fLm). Eo lectotype (scale 3 em).

styles or anisoxeas; bases of subectosomal megascleres embedded aspinose region near basal region (85-101 X 9-11 f,Lm); microscle-
in axial skeleton, enveloped by heavy fibrous spongin, protruding res absent
through surface as single spicules or paucispicular brushes, form- Remarks. Clathriodendron (including junior synonym
ing extra-axial skeleton; subectosomal megascleres near centre of Dictyocylindrus, with type species Spongia hispida Montagu,
axial skeleton smaller than spicules occurring towards periphery; 1814, by original designation) is superficially quite different from
no compression of choanosomal skeleton nor any well differenti- Raspailia lacking axial condensation, any marked axial and extra-
ated extra-axial skeleton; choanosomal skeleton with very heavy axial skeletal differentiation, and often lacking the specialized
spongin fibres forming irregular reticulation of longitudinal tracts, ectosomal skeleton typical of other raspailiids. In its aberrant
without special category of choanosomal megascleres, but with skeletal structure it shows closest similarities to Echinodictyum.
aspicular fibres or echinating spicules secondarily incorporated Some species are also known to incorporate detritus into their
into fibres; skeletal tracts heavily echinated by acanthostyles; prin- skeletons, and in that respect they are homologous to the relation-
cipal megascleres absent; subectosomal extra-axial megascleres ship demonstrated in the Microcionidae between Clathria s.s. and
long, setaceous, straight or slightly curved towards basal end, with Wilsonella, the latter used as a 'convenient' subgenus for arena-
rounded stylote, sometimes subtylote, less commonly asymmetri- ceous species of Clathria (Hooper, 1996a). Hallmann (1912), fol-
cal oxeote ends (820-1545 X 11-26f,Lm); ectosomal auxiliary lowed by Wiedenmayer (1989), considered that C. arbuscula was
styles or anisoxeas relatively small, thin, slightly curved at centre sufficiently different from Raspailia to maintain the two taxa.
or base, with fusiform points and rounded stylote or fusi- Conversely, Hentschel (1911), followed by Hooper (1991) sug-
form oxeote bases (253-412 X 4-6 f,Lm); echinating acanthostyles gested that Clathriodendron was synonymous with Raspailia due
relatively smail, subtylote, with large granular spination, an to the existence of species that otherwise clearly belonged to
478 Porifera' Demospongiae • Poecilosclerida • Microcionina • RaspaiJiidae

Raspailia, but with exclusively or nearly exclusive reticulate branch diameter, but nevertheless forming thick bundles, arising tan-
choanosomal skeletons and no differences in their echinating gentially or obliquely from axis, and composed of plumose brushes of
acanthostyle geometries. longer styles or anisoxeas embedded in the axial skeleton and pro-
truding only a relatively short distance through the surface; axial core
Distribution compressed, comprising 60-80% of branch diameter, composed of
discrete, closely set, fasciculated bundles of shorter choanosomal
Common in Indo-west Pacific region, present in the principal styles usually running longitudinally through branches but
Caribbean, and possibly worldwide in distribution. also at more oblique angles, producing solid but flexible skeleton.
Megascleres: both categories of structural megasclere are very similar
in their geometry, differing mainly in size and location within skele-
SUBGENUS PARASYRINGELLA TOPSENT, 1928 ton. Shorter choanosomal principal (axial) styles straight or slightly
curved, with evenly rounded or tapering fusifonn bases and tapering
Synonymy to sharp points (232-365 X 6-9,J.m); longer subectosomal (extra-
axial) styles, thick, relatively straight, with fusifonn tapering bases
[Syringella] of authors; Ridley, 1884a: 460; Pick, 1905: 18; and sharply pointed tips (543-964 X 9-25 fLm); ectosomal (auxiliary)
Topsent, 1892a: 123, 1904b: 138, 1928c: 42; Burton, 1934b: 42; styles or anisoxeas range from thin wispy flexuous anisoxeas to
de Laubenfels, 1936a: 121 (Not Syringella Schmidt, 1868: 10). thicker straight or slightly curved styles (145-237 X 0.5-5 fLm);
Parasyringella Topsent, 1928c: 287; de Laubenfels, 1936a: 102. echinating megascleres absent; microscleres absent.
Raspailopsis Burton, 1959a. Taxonomic decision for synonymy: Remarks. The taxon Parasyringella is used here as a con-
Hooper (1991), and this work. venient subgenus, to delineate a group of Raspailia species lacking
echinating megascleres, and includes the junior synonym
Type species Raspailopsis and [Syringella] of authors, Ridley (1884a) being the
first (not of Schmidt, 1868). Burton (1959a) created Raspailopsis
Raspailia (Parasyringella)falcifera Topsent, 1892a (by origi- (type species Raspailopsis cervicornis Burton, 1959a, by original
nal designation; schizotype MNHN DT901). designation; holotype BMNH 1936.3.4.604) for raspailiids that
lacked echinating acanthostyles whereas [Syringella] (type species
Detinition Raspailia syringella Schmidt, 1868: 10, by monotypy, holotype
unknown) was already in use by earlier authors for much the same
Raspailia which have secondarily lost their echinating purpose. In any case, some specimens of the type species of
megascleres. Raspailopsis, R. cervicornis (BMNH 1936.3.4.521 and BMNH
1936.3.4.522) from the South Arabian coast did have echinating
Diagnosis acanthostyles present on the peripheral fibres of the axial skeleton,
and in this regard Raspailopsis is undoubtedly a synonym of
Unbranched cylindrical, digitate or whip-like Raspailia usu- Raspailia. Hooper (1991) followed the prior use of the subgeneric
ally with a hispid surface; skeleton divided into distinct axial and name [Syringella] for this group, but also noted that Schmidt's type
extra-axial regions; axial skeleton compressed- highly compacted species, R. syringella, did not belong. Consequently, the name
reticulation of fibres and multispicular tracts running mainly longi- 'Syringella' cannot be used as a valid taxon, with the next available
tudinally through branches to produce a flexible skeleton, cored by name being Parasyringella Topsent. Only a poor microscope slide
shorter principal styles; extra-axial skeleton composed of multispic- preparation of R. falcifera has been examined from the MNHN col-
ular tracts cored by larger extra-axial styles together forming dis- lections, so the description of Ridley's species is provided instead
crete bundles embedded in the axial skeleton and extending radially as 'representative' of the subgenus.
to the surface, with single or bundles of extra-axial styles protruding
a short way through the surface, and surrounded by bouquets of Distribution
ectosomal styles; megascleres include at least 2-3 size classes of
styles, with the smaller ones sometimes modified to anisoxeas. Uncommon, Indo-Pacific.

Description of 'representative' species

SUBGENUS HYMERAPHIOPSIS HOOPER, 1991
Raspailia (Parasyringella) australiensis Ridley, 1884a (Fig. 4).
Synonymy. Raspailia (Syringella) australiensis Ridley, Synonymy
1884a: 460.
Material examined. Lectotype: BMNH 1882.2.23.253 - Hymeraphiopsis Hooper, 1991: 1270.
Port Darwin, NT.
Description. Shape unbranched cylindrical digits, bifurcate, Type species
with short holdfast, long slender woody stalks; texture firm, flexible,
with woody central stem (=axial skeleton) and fleshy branches (pro- Raspailia irregularis Hentschel, 1914 (by original designation).
duced by extra-axial region); surface optically even, microscopically
rugose and hispid, with minute conules and ridges; ectosomal skele- Definition
ton with sparse brushes of thin ectosomal styles surrounding multi-
spicular brushes of larger extra-axial styles protruding through the Raspailia having acanthostyles with smooth and very swollen
surface; these extra-axial spicule tracts comprise only 20-40% of the tylote bases.
Porifera· Demospongiae • Poecilosclerida • Microcionina • Raspailiidae 479

c D
I I
B

A I

Fig. 4. Raspailia (Parasyringella). A-E, R. (Parasyringella) australiensis (lectotype BMNH1882.2.23.253). A, subectosomal (extra-axial) style (scale
200 !Lm). B, choanosomal principal (axial) styles (scale 60 !Lm). C, ectosomal (auxiliary) style/anisoxea (scale 50 !Lm). D, choanosomal skeletal structure
(scale 500 !Lm). E, ectosomal and extra-axial skeleton (scale 100 .... m) . F, specimen (QM GL850) (scale 3 cm).

Diagnosis branches aculeate, irregularly conulose and prominently hispid;

texture compressible, flexible; ectosomal skeleton with paucispicu-
Monotypic. Refer to type species description below. lar brushes of ectosomal styles or anisoxeas, dispersed over surface
and intermingled with erect brushes of acanthostyles just below
Description of type species ectosome; brushes not associated with protruding subectosomal
extra-axial spicules; subectosomal extra-axial megascleres embed-
Raspailia (Hymeraphiopsis) irregularis Hentschel, 1914 ded in axial skeleton, protruding up to 2 mm from surface, domi-
(Fig. 5). nating skeletal architecture so much that axial skeleton resembles a
Synonymy. Raspailia irregularis Hentschel, 1914: 121; cluster of spiCUles surrounding bases of subectosomal megascleres;
Eurypon miniaceum; Burton, 1932b:325; (Not Eurypon miniaceum choanosomal axial skeleton a series of compressed axial clusters
Thiele, 1905: 446). of echinating megascleres, without fibres or any specialized
Material examined. Holotype: ZMH (not seen). Specimen choanosomal megascleres, united by granular spongin and forming
of Burton (1932b): BMNH1928.2.15.244 - South Georgia, plumose brushes; echinating spicule brushes near periphery
S Atlantic. directed outwards, and points of acanthostyles protrude up to
Description. Shape small, bushy, branching, with multiple but not through ectosome; bases of subectosomal extra-axial
points of attachment, several basal holdfasts and short stalks, and spicules embedded in core of axial skeleton; principal megascleres
flattened, irregularly bifurcate and bushy branches; surface of absent; subectosomal extra-axial spicules very long and thick
480 Porifera' Demospongiae • Poecilosclerida • Microcionina • RaspaiIiidae

I
Fig. 5. Raspai/ia (Hymeraphiopsis). A-F, R. (Hymeraphiopsis) irregularis (specimen BMNH 1928.2.15.244). A, subectosomal styles (scale 450/-Lm).
B, echinating acanthotylostyles (scale 100 /-Lm). C, ectosomal style and anisoxea (scale 250/-Lm). D, section tbrough peripheral skeleton (scale 250/-Lm).
E, choanosomal skeleton (scale 250/-Lm). F, specimen (scale 3 cm).

styles, curved near basal end, with enlarged but not necessarily Distribution
tylote bases, tapering to sharp points (1969-3016 X 35-54 /-Lm);
ectosomal auxiliary megascleres relatively long thin styles or Monotypic, Antarctica.
anisoxeas, sometimes straight but mostly with prominent curvature
at centre, with rounded tapering hastate bases, sometimes oxeote,
tapering to sharp points (435-712 X 2-11 /-Lm); echinating acan- AULOSPONGUS NORMAN, 1878
thosty1es straight, thin or thick, with grossly enlarged tylote base,
evenly cylindrical shaft and sharply pointed at apex, with vestigial Synonymy
spination on shaft or sometimes confined to points (181-301 X
12-24 /-Lm); microscleres absent. Aulospongus Norman, 1878: 267; Dendy, 1889: 89; Dendy,
Remarks. Hymeraphiopsis is currently monotypic, known 1922b: 61; Burton, 1937: 38; Hooper, 1991: 1307; Hooper & Levi,
only from several records of the Antarctic type species including a 1993: 1294: Hooper et al., 1999: 651 (Not Aulospongus; de
comprehensive redescription of type and new material by Hooper Laubenfels, 1936a: 100). Aulospongiella Burton, 1956: 141.
(1991). It differs from other subgenera of Raspailia essentially in Heterectya Hallmann, 1917b: 393. Rhaphidectyon Topsent, 1927b:
its possession of acanthostyles with swollen bulbous tyles and 15. Hemectyonilla Burton, 1959a: 254. Trachostylea Topsent,
spines confined mainly to the apical points of spicules. The type 1928c: 166. Taxonomic decision for synonymy: Hooper (1991),
species also differs from typical Raspailia in having echinating Hooper et al. (1999), this work.
acanthotylostyles clustered around the bases of the long extra-axial
styles, deep within the choanosomal skeleton, and with ectosomal Type species
styles/anisoxeas scattered mainly near the peripheral skeleton,
below the surface, and rarely forming bouquets around these Haliphysema tubulatus Bowerbank, 1873c (by original
protruding spicules. designation).
Porifera· Demospongiae • Poecilosclerida • Microcionina • Raspailiidae 481

Definition Description. Growth form subspherical, massive, tubular,

composed of amalgamated fibre-bundles that extend to the surface
Raspailiidae with at least two size classes of rhabdostyles of and produce a compartmentalised surface of discrete conules; ecto-
similar geometry, the larger (smooth or partially spined) core spon- some with vestigial 'raspailiid skeleton' composed of sinuous or
gin fibres, and the smaller (partially spined) echinate fibres rhaphidiform ectosomal auxiliary styles scattered sparsely and
although neither are localised to any region of the skeleton; indiscriminantly over the surface, and with plumose bundles of
choanosomal skeletal structure is predominantly plumose, with both larger and smaller rhabdostyles protruding through the surface
spicules and fibres amalgamated into bulbous tracts ('fibre- mainly at the ends of conules; no subectosomal extra-axial spicules;
bundles'), more-or-Iess compacted in the axial skeleton, becoming adjacent surface conules interconnected by aspicular (membra-
increasingly plumose as they ascend to the periphery, eventually nous) collagen; choanosomal skeleton exclusively plumose with
producing a shaggy, compartmentalised or conulose surface; axial fibre-bundles fused relatively closely towards the base of the
and extra-axial skeletons undifferentiated apart from greater sponge and axis of the skeleton, becoming increasingly plumose
amalgamation of fibre-bundles in the axis. towards the periphery, and eventually completely discrete/ com-
partmentalised at the surface; fibre-bundles composed of rhabdo-
Diagnosis styles, both coring and echinating fibres, forming ascending
multispicular columns, branching or unbranched towards their
Growth forms tubular, cup-shaped, lobate, lamellate or erect apex, bonded together with light granular collagen; larger smooth
cylindrical-digitate; individual lobes or branches are composed of choanosomal principal rhabdostyles extend out from fibres in
large fibre-bundles amalgamated at the core or base of the sponge, plumose bundles, particularly near periphery of skeleton; smaller
diverging and becoming increasingly plumose towards the periph- spined rhabdostyles heavily echinate fibres and fibre nodes; mega-
ery, eventually producing a compartmentalised surface of discrete scleres consist of larger coring choanosomal principal rhabdostyles
lobes or shaggy surface processes. Ectosomal skeleton ranges with slightly subtylote or rounded bases, slight basal rhabd,
from: well developed, 'specialised raspailiid' (consisting of long fusiform points, completely smooth (304-462 X 16--24 J.Lm);
subectosomal extra-axial styles protruding through the surface, smaller echinating rhabdostyles with entirely smooth, slightly
surrounded by sparse brushes of ectosomal auxiliary spicules); rhabdose, slightly swollen bases, and small spines covering
vestigial (with wispy raphidiform or sinuous ectosomal auxiliary apical half of spicule (109-126 X 5-10 J.Lm); long, thin, curved,
spicules scattered sparsely and indiscriminately over the surface); sinuous or rhaphidiform ectosomal auxiliary styles (212-250 X
or absent completely (with only choanosomal principal spicules 2-3 J.Lm); subectosomal extra-axial spiCUles absent. Microscleres
protruding through the surface, forming discrete surface bundles). absent.
Long subectosomal extra-axial spicules produce a hispid surface in Remarks. Re-examination of type material by Hooper et al.
some species. Choanosomal skeletal structure predominantly (1999) confirmed the synonymy proposed above between
plumose (with very few reticulate connecting fibres, and these Aulospongus and Aulospongiella (type species Axinella monticularis
mainly towards the axis), with virtually no differentiation between Ridley & Dendy, 1886, by original designation and mono-
axial and peripheral skeletons. Ascending fibres nearly fully cored typy; holotype BMNH 1887.5.2.20), Heterectya (type species
by larger choanosomal principal rhabdostyles, forming dense Raspailia (?) villosa Thiele, 1898, by original designation; holo-
plumose bundles particularly on fibre nodes, and echinated by type 2MB 2204), Rhaphidectyon (type species Rhaphidectyon
smaller rhabdostyles, together producing bulbous spiculo-spongin spinosum Topsent, 1927b, by original designation and mono-
tracts (termed here 'fibre-bundles'); smaller echinating rhab- typy; schizotypes MNHN DT 1139, BMNH 1930.7.1.39) and
dostyles more-or-Iess evenly dispersed throughout the skeleton; Hemectyonilla (type species Stylostichon involutum Kirkpatrick,
interconnecting fibres, if present, are aspicular or paucispicular, 1903b, by original designation and monotypy; holotype BMNH
and generally confined to the axial region. Megascleres consist of 1902.11.16.33). Trachostylea Topsent (type species T. semota
larger choanosomal principal rhabdostyles usually with a relatively Topsent, 1928c, by monotypy, holotype not seen, possibly MOM)
slight basal rhabd, entirely smooth or with recurved spines only on should also be referred here (Van Soest, pers.comm.). It has two
apical part of spicules. Smaller echinating rhabdostyles in one or size classes of acanthose styles, the smaller more rhabdose and
two categories have more pronounced basal rhabd, often promi- subtylote than the larger, the larger only slightly curved near the
nently subtylote, entirely smooth or with spines only at the apex base. Topsent (1928c) described the skeleton as a relatively lax
of spicules, or covering most of the spicule except for the base, or unispicular reticulate network, not very regular, without distinct
rarely completely spined. Subectosomal extra-axial styles or tracts and lacking apparent spongin, and the larger spicules only
anisoxeas, if present are long and protrude through the surface. slightly projecting through as a surface hispidation. The species
Ectosomal auxiliary styles or anisoxeas, if present are wispy, sinu- remains poorly known only from its original description and its
ous or raphidiform, often vestigial. Raphide microscleres are present precise assignment requires corroboration from re-examination of
in only one species. type material.
Rhabdostyles have traditionally been accorded primary diag-
Description of type species nostic significance for Aulospongus yet these also occur in two
other raspailiid genera (Raspailia (Raspaxilla) including synonyms
Aulospongus tubulatus (Bowerbank, 1873c) (Fig. 6). Echinaxia and Axinectya, and Endectyon (Hemectyon», and osten-
Synonymy. Haliphysema tubulatus Bowerbank, 1873c: 29; sibly also in Cantabrina (included here as incertae sedis in
Aulospongus tubulatus; Norman, 1878: 267; Axinella tubulata; Raspailiidae). They are also found in other Poecilosclerida
Dendy, 1889: 89. (Rhabderemia, Rhabderemiidae; Biemna, Desmacellidae), and the
Material examined. Holotype: BMNH 1873.7.21.9 - Halichondrida families Desmoxyidae (Halicnemia Bowerbank,
Ceylon (Sri Lanka). Higginsia Riggin), and Bubaridae (Rhabdoploca Topsent,
482 Porifera· Demospongiae • Poecilosclerida • Microcionina • Raspailiidae

A I
Ic
B

Fig. 6. Aulospongus. A-G, A. tubulatus (Bowerbank) (holotype). A, choanosomal principal rhabdostyles (scale 100 /Lm). B, echinating rhabdostyles (scale
10 /Lm). C, ectosomal auxiliary styles (scale 50 /Lm) . D, holotype (scale 3 cm). E, Dendy (l922b) 'Investigator' specimen (scale 3 cm). F, ectosomal skele-
ton (scale 250 /Lm). G, choanosomal skeleton (scale 250 /Lm).

Hymerhabdia Topsent, and Monocrepidium Topsent), with the Hooper et al. (1999) found the remnants of a specialised
implication that they have been derived independently within each raspailiid ectosomal skeleton to be present in both the type species
group (homoplasic developments). Amongst the rhabdose ras- of Aulospongus (A. tubulatus) and Hemectyonilla (Stylostichon
pailiid genera, however, Aulospongus differs in having two size involutum), a character overlooked by previous authors. This dis-
categories of rhabdostyles (the larger, smooth or partially spined covery (a) confirmed the synonymy between Aulospongus and
rhabdostyles coring fibres, and the smaller, usually spined rhab- Hemectyonilla, proposed tentatively by Hooper (1991), (b) provided
dostyles echinating fibres); a plumose skeletal structure composed more firm evidence in support of the inclusion of Aulospongus in
of ascending compressed fibre-bundles (with few or no reticulate the Raspailiidae, previously considered to be relatively atypical of
elements, and in cases where reticulate interconnecting fibres are the family, and (c) illustrated that ectosomal specialisation is easily
present these are usually aspicular and disappear completely in the lost within a raspailiid taxon, ranging from well developed, typical
peripheral skeleton); lacking any differentiation between axial and 'raspailiid condition' (e.g., A. gardineri (Dendy», vestigial with few
extra-axial regions of the skeleton (although plumose fibres are sinuous ectosomal spicules (e.g., the type species), to absent com-
slightly more compressed in the axis than in the periphery). In pletely (e.g., A. monticularis (Ridley & Dendy».
Aulospongus, unlike other rhabdose raspailiid genera, the larger A detailed discussion of the genus in relation to other rhab-
choanosomal principal rhabdostyles bear a striking resemblance to, dose raspailiids, and a phylogenetic and biogeographic analysis of
and are probably commonly derived from, the smaller echinating species is provided by Hooper et al. (1999).
rhabdostyles. These smaller rhabdostyles are found in all three
rhabdose raspailiid genera, and are probably homologous and Distribution
derived from typical raspailiid echinating acanthostyles. In contrast
Raspaxilla and Hemectyon, as in most raspailiids, have fibres cored Aulospongus contains 11 species and has a disjunct geo-
by non-rhabdose, smooth styles of distinctly different geometry graphic distribution, from the N Atlantic (Sao Vicente and Cape
and origin from the rhabdose echinating spicules. Verde Islands), SW Indian Ocean (Natal), W and central Indian
Porifera· Demospongiae • Poecilosclerida • Microcionina • Raspailiidae 483

Ocean (Zanzibar, Kenya, Gulf of Aden, Arabian Gulf, S Arabian distance; choanosomal and subectosomal megascleres undifferen-
coast, Amirante, India, Sri Lanka), NW Pacific (Japan) and SW tiated; choanosomal skeleton with distinct, irregularly reticulate
Pacific Ocean (New Caledonia), E Pacific (Gulf of California), and axis, radial non-reticulate extra-axis, and plumose ectosomal
Caribbean (Colombia and Jamaica). Biogeographic analysis spicule brushes; well developed spongin fibres cored by pauci-
showed it to have a pan-equatorial, predominantly tropical- spicular tracts of choanosomal styles (438-843 X 17-29 f.Lm)
sUbtropical distribution, with rare incursions into cooler temperate becoming plumose towards periphery; fibres moderately heavily
waters, in contrast to the other rhabdose raspailiids which are either echinated by clavulate acanthostyles (111-146 X 6--11 f.Lm);
centred on the Pacific rim, distributed in tropical, temperate and sub ectosomal megascleres completely undifferentiated from
antiboreal waters, extending into the Antarctic (Raspaxilla) or choanosomal spicules.
known only from the Caribbean (Hemectyon). Remarks. The type species of Ectyoplasia shows classical
raspailiid ectosomal specialization, with brushes of ectosomal aux-
iliary spicules surrounding the bases of protruding spicules. By
ECTYOPIASIA TOPSENT, 1930 comparison, E. frondosa (Lendenfeld) and E. ferox (Duchassaing &
Michelotti) have an atrophied ectosomal skeleton containing only
Synonymy sparsely dispersed tangential ectosomal spicules. Topsent (1930)
suggested that the genus differed from Raspailia (s.s.) in having
Ectyoplasia Topsent, 1930: 23--4; de Laubenfels, 1936a: 102; less compressed skeletal architecture, clavulate tips on acan-
Wiedenmayer, 1977b: 158. thostyles, and 'exceptional differentiation' of the exhalant pores
(oscula) along the lateral margins of branches. The latter feature
Type species has little systematic importance at the generic level, and in any case
it does not occur in either E. frondosa or E. vannus Hooper.
Spongia tabula Lamarck, 1814 (by original designation). Similarly, supposed differences in skeletal architecture between
the two genera are not upheld upon comparison with other
Defmition species (e.g., compare Raspailia phakellopsis Hooper, 1991
with E. frondosa, and R. darwinensis Hooper, 1991 with E. ferox).
Raspailiidae with a single category of undifferentiated Thus in Topsent's (1930) conception of Ectyoplasia this leaves
choanosomal styles, and club-shaped acanthostyles bearing large only acanthostyle morphology, or modifications to acanthostyles,
clavulate spines on the apex and a smooth bulbous base. as a primary distinguishing feature although Hooper (1991) also
noted that Ectyoplasia has well differentiated axial and extra-axial
Diagnosis skeletons like Raspailia s.s. but in which spicules forming both
structures are identical; i.e., the genus lacks specialized subectoso-
Flabellate, flattened arborescent or tubular growth form, with mal megascleres.
even or slightly corrugated hispid surface. Choanosomal skeleton
slightly compressed axial reticulation of spongin fibres cored by Distribution
styles or anixoxeas, echinated by clavulate acanthostyles; subecto-
somal extra-axial skeleton of radially arranged plumose ascending Four species are described from tropical and subtropical
tracts, composed of undifferentiated choanosomal styles protruding Australia, Indonesia, Micronesia (Hooper, 1991; Hooper et al.,
through ectosome, interconnected by uni- or paucispicular trans- 1999), and Caribbean (e.g., Wiedenmayer, 1977b).
verse spicule tracts producing regular reticulation; ectosomal
region with a specialized skeleton of small styles or anisoxeas typi-
cally forming brushes around choanosomal styles at surface, but ENDECTYON TOPSENT, 1920
sometimes lying tangential to surface. Structural megascleres styles
or rhabdostyles of two sizes, sometimes with anisoxeote or strongy- Synonymy
lote modifications; echinating spicules always acanthostyles with
clavulate points; microscleres absent. Endectyon Topsent, 1920a: 25 (see subgenera for full
synonymy).
Description of type species
Type species
Ectyoplasia tabula (Lamarck, 1814) (Fig. 7).
Synonymy. Spongia tabula Lamarck, 1814: 374. Phakellia tenax Schmidt, 1870 (by original designation).
Ectyoplasia tabula; Topsent, 1930: 23.
Material examined. Holotype: MNHN DT553 - East of Definition
Cape Leeuwin, Western Australia.
Description. Flattened bifurcate digitate, stalked, with Raspailiidae with clavulate modifications to acanthostyle
more-or-less regular lateral surface ridges, sandpaper-like stiff tex- geometry, and acanthostyles confined to a particular region outside
ture; ectosomal skeleton discrete bundles of ectosomal auxiliary of the skeletal axis.
styles or anisoxeas (244-331 X 4-7 f.Lm) surrounding bases of pro-
truding choanosomal styles extending from extra-axial skeleton; Diagnosis
extra-axial skeleton more-or-less radial non-plumose bundles of
choanosomal axial spicules, in uni- or paucispicular tracts, Prominently hispid, conulose surface, and typically arborescent
extending from axis and protruding through surface for only short growth forms. Skeleton always with marked axial and extra-axial
484 Porifera' Demospongiae • Poecilosclerida • Microcionina • Raspailiidae

E
AB

Fig. 7. Ectyoplasia. A-F, E. tabula (holotype). A, choanosomal styles (scale 200 f,l-m). B, ectosomal auxiliary style (scale 100 f,l-m). C, clavulate acan-
thostyles (scale 50 f,l-m). D, specimen NTM Z2677) (scale 3 em). E, ectosomal skeleton (scale 150 f,l-m). F, choanosomal skeleton (scale 150 f,l-m).

differentiation of skeleton; axial skeleton with well developed extra-axial and axial regions, or forming plumose brushes along
spongin fibres forming compressed reticulation, cored by stout length of extra-axial tracts, or localized exclusively to base of
choanosomal styles; extra-axial subectosomal skeleton radial or sponge (nominal genus Basiectyon). Structural megascleres
plumose, with multi- or paucispicular tracts of long subectosomal smooth styles of 2-3 size categories; echinating megascleres mod-
styles (subgenus Endectyon) or choanosomal styles (subgenus ified acanthostyles and/or acanthostrongyles with peculiar strongly
Hemectyon), sometimes connected by unispicular tracts forming curved (clavulate) hooks on shaft, base, and/or apex. Microscleres
hexagonal meshes, usually protruding through surface. Ectosomal absent.
skeleton varies from typical raspailiid condition, with thin ectoso-
mal styles grouped in brushes around protruding subectosomal Remarks
styles (subgenus Endectyon), to surface brushes composed of
subectosomal styles only (nominal genus Basiectyon), to brushes Endectyon differs from typical raspailiids (e.g., Raspailia)
of acanthostyles surrounding choanosomal styles (subgenus in having clavulate modifications to acanthostyle geometry.
Hemectyon). Erect brushes of echinating acanthostyles located on Moreover, unlike species of Raspailia in which echinating spicules
outer margin of axial skeleton, surrounding boundary between are relatively evenly dispersed throughout the skeleton, Endectyon
Porifera· Demospongiae • Poecilosclerida • Microcionina • Raspailiidae 485

has these confined to a particular region of the skeleton (i.e., out- outside the axial skeleton, and have recurved (clavulate) hooks on
side the axis). Some species of Endectyon show reduced character- both ends but lack any basal rhabd.
istics from the typical condition. The affinities of these species are
not completely obvious, but the most reasonable interpretation is Description of type species
that they are forms of Endectyon (Hooper, 1991). Hooper et at.
(1999) proposed the recognition of two subgenera. Endectyon tenax (Schmidt, 1870) (Fig. 8).
Species of Endectyon (Endectyon) (including Basiectyon, Synonymy. Phakellia tenax Schmidt, 1870: 62. Endectyon
type species Basiectyon pilosus Vacelet, 1961a, by monotypy; lec- tenax; Topsent, 1920a: 23. Endectyon (Endectyon) tenax; Hooper
totype SMEM 1705(1), schizotype MNHN DNBE 718L) have et at., 1999: 695.
acanthostyles with recurved (clavulate) hooks on both ends and Material examined. Holotype: LMJG unregistered, schizo-
lack any basal rhabd, and with a specialised ectosomal skeleton. types MNHN DCL2163, 1194, BMNH 1870.5.3.170 - Florida.
Endectyon (Hemectyon) has a more openly reticulate axial skeletal, Description. Erect, arborescent growth form; choanosomal
lacking differentiated subectosomal megascleres in the extra-axial skeleton with compressed axial skeleton composed of stout
skeleton, lacking a specialized ectosomal skeleton, and the bases of choanosomal styles (330--400 X 15-22 /-Lm) enclosed in well
acanthostyles are predominantly smooth, subtylote, and some are developed spongin fibres; extra-axial skeleton plumose, composed
slightly rhabdose. Basiectyon also has a loosely reticulate axial oflong subectosomal styles (1000-1300 X 20-22 /-Lm) with little or
skeleton, lacks ectosomal specialization, and its acanthostyles are no spongin, protruding a long way and hispidating surface, also
localized at the base ofthe sponge, and based on acanthostyle mor- with irregular reticulate spicule skeleton composed of choanoso-
phology it appears to be most closely related to Endectyon mal styles extending up to surface; protruding auxiliary spicules
(Endectyon). surrounded by bundles of ectosomal styles (220-230 X 2-3 /-Lm) on
Based on acanthostyle geometry, acanthostyle distribution, surface; echinating acanthostrongyles with clavulate spines on
and axial and extra-axial skeletal structure all three genera share both ends and partially on shaft (122-133 X 15-18 /-Lm), implanted
features which argue for their inclusion into a single taxon (see at diverse angles, usually in heavy concentrations, only in extra-
Topsent 1920a). axial tracts, producing a unispicular network between tracts in
deeper parts of subectosome, standing erect on ectosome closer to
Distribution surface.
Remarks. See remarks for subgenus Endectyon (Hemectyon).
Twelve species have been described from the Mediterranean
(Schmidt, 1870; Vacelet, 1961a), North Atlantic (Schmidt, 1870; Distribution
Topsent, 1892c; Burton, 1930c; Cabioch, 1968a), Indian Ocean
(Ridley & Dendy, 1886; Dendy, 1887; Thomas, 1976), and tropical Caribbean, Mediterranean, North Atlantic, Indo-Pacific
Australian seas (Lamarck, 1814; Hooper, 1991) (Hooper et at., 1999). (Hooper, 1991; Hooper et at., 1999).

SUBGENUS ENDECTYON TOPSENT, 1920 SUBGENUS HEMECTYON TOPSENT, 1920

Synonymy Synonymy

Endectyon Topsent, 1920a: 25; Burton, 1937: 34; Cabioch, Hemectyon Topsent, 1920a: 27; Hooper, 1991: 1284; Hooper
1968b: 224; Thomas, 1976: 169; Hooper, 1991: 1284. Basiectyon et at., 1999: 695.
Vacelet, 1961a: 37. Taxonomic decision for synonymy: Hooper
(1991), Hooper et al. (1999). Type species

Type species Raspailia (?) hamata Schmidt, 1870 (by original designation).

Phakellia tenax Schmidt, 1870 (by original designation). Definition

Definition Endectyon with subtylote echinating rhabdostyles bearing

clavulate spines only on apex, grouped around protruding
Endectyon with recurved (clavulate) hooks on both ends and choanosomal styles in the ectosomal skeleton.
lacking any basal rhabd on echinating megascleres, and lacking
a special ectosomal skeleton composed of ectosomal styles/ Diagnosis
anisoxeas grouped around long extra-axial styles.
Arborescent growth form; echinating acanthostyles bearing
Diagnosis clavulate spines only on apex of spicule (not on base), and base is
smooth and sometimes slightly rhabdose; echinating spicules
Arborescent growth form; with a compressed axial skeleton localised outside the axial skeleton, usually at the junction of axial
cored by stout choanosomal principal styles, and plumose extra- and extra-axial skeletons, and/or forming plumose brushes along
axial skeleton cored by long subectosomal auxiliary styles produc- extra-axial skeleton, and often also produce spicule brushes at the
ing a hispid surface, surrounded by bundles of ectosomal auxiliary surface; axial skeleton compressed reticulate; extra-axial skeleton
styles at the surface. Echinating acanthostyles are confined to plumose(-reticulate) cored by choanosomal styles.
486 Porifera· Demospongiae • Poecllosclerida • Microcionina • Raspalliidae

Fig. 8. Erulectyon (Erulectyon). A-F, E. (Endectyon) tenax (fragment ofholotype BMNH 1870.5.3.170). A, subectosomal style (scale 200!Lm). B, choanoso-
mal styles (scale 100 !Lm). C, ectosomal styles (scale 50 !Lm). D, acanthostrongyles (scale 50 !Lm). E, choanosomal skeleton (scale 300 !Lm). F, peripheral
skeleton (scale 300 !Lm). G-H, E. (Endectyon) elyakovi Hooper, 1991 (paratype NTMZ2738). G, paratype (scale 3 em). H, skeletal structure (scale 500 !Lm).

Description of type species well differentiated. Axial skeleton strongly reticulate, compressed.
with heavy fibres cored by small choanosomal principal styles in
Endectyon (Hemectyon) hamata Schmidt, 1870 (Fig. 9). multi spicular tracts mostly running longitudinally through branches,
Synonymy. Raspailia (?) hamata Schmidt, 1870: 62; and with few echinating rhabdostyles. Extra-axial skeleton radial-
Hemectyon hamatum; Topsent, 1920a: 26; Endectyon hamata; reticulate, without fibre-bundles, with ascending paucispicu1ar tracts
Hooper, 1991: 1284; Endectyon (Hemectyon) hamata; Hooper interconnected by unispicular transverse tracts of choanosomal prin-
et al .• 1999: 695. cipal styles. Echinating rhabdostyles predominantly on exterior sur-
Material examined. Holotype: MZUS P0151 (not seen). face of primary (ascending) extra-axial fibres. with greatest numbers
fragment MNHN DT2161 - 'West Indies'. concentrated at the surface in brushes. MegascIeres consist of
Description. Growth form arborescent, cylindrical branches. choanosomal principal styles slightly curved centrally, without basal
Surface slightly corrugated. Colour pale brown in dry state. rhabd, entirely smooth (270--615 X 8-18 fLm). Echinating rhab-
Specialised 'raspailiid ectosomal skeleton' absent. with only pro- dosty1es with very slight basal rhabd, smooth slightly swollen base.
truding bundles of a few choanosomal principal styles surrounded at and large cIavulate spines only on apical extremity or apical 1I3rd of
their base by multispicular plumose bundles of rhabdostyles, spicule at most (120-150 X 5-10 fLm). Subectosomal extra-axial
although vestigial ectosomal auxiliary styles are scattered within the styles absent. Ectosomal styles vestigial and scattered within the
choanosomal skeleton. Axial and extra-axial skeleton moderately choanosome (220-275 X 1-2fLm). MicroscIeres absent.
Porifera· Demospongiae • Poecilosclerida • Microcionina • Raspailiidae 487

D
c
Fig. 9. Endectyon (Hemectyon) . A-D, E. (Hemectyon) hamata (fragment of holotype MNHNLBIMDT2161). A, choanosomal skeleton (scale I mm).
B, peripheral skeleton (scale 200 /Lm), C, echinating rhabdostyles in fibres (scale 50/Lm). D, echinating rhabdostyIe (scale 50/Lm).

Remarks. Hemectyon was merged into synonymy with localised outside the axial skeleton (usually at the junctions of axial
Endectyon by Hooper (1991), on the basis that the two differed in and extra-axial skeletons).
only a few characters: Endectyon (s.s.) with a special ectosomal Comparisons between other raspailiids with rhabdose acan-
skeleton composed of ectosomal styles/anisoxeas grouped around thostyles (Autospongus and Raspailia (Raspaxilla» are slightly mis-
long extra-axial styles, whereas Hemectyon lacked both true leading. Echinating acanthostyles in Endectyon (Hemectyon) are not
ectosomal and extra-axial spicules (having instead acanthostyles truly rhabdose, like the other two genera, with the slight basal curva-
grouped around protruding choanosomal styles), and the latter also ture often overemphasised by the basal swelling on these spicules.
had more openly reticulate axial skeleton than Endectyon (s.s.). Nevertheless, rhabdostyles in these three genera may be potentially
More importantly though, in Hemectyon the bases of acanthostyles confused. The subgenus contains only the type species, although
are predominantly smooth, subtylote, and some are slightly rhab- arguably E. fruticosa (Dendy), E. fruticosa aruensis (Hentschel), and
dose, whereas those of Endectyon have recurved (c1avulate) hooks E. xerampelina (Lamarck) could also be included given that some
on both ends and lack any basal rhabd. On this basis Hooper et at. (but not all) of their echinating acanthostyles have smooth, swollen,
(1999) used Hemectyon as a convenient subgenus within Endectyon, slightly rhabdose bases with c1avulate spines mainly on the apex of
both having in common c1avulate spines on acanthostyles, the latter the spicule. They are not included, however, because other spicules
488 Porifera· Demospongiae • Poecilosclerida • Microcionina • Raspailiidae

also have clavulate spines on their bases and more closely resemble Description. Encrusting growth form; choanosomal skeleton
those of Endectyon species (see Hooper, 1991). with microcionid structure consisting of a basal layer of spongin on
the substrate giving rise to individual fibre nodes, each node cored
Distribution by thick bundles of long, completely smooth subectosomal
(sub)tylostyles (685-2310 X 11-28 f.lm) forming plumose columns
Caribbean. that penetrate and extend a long way through the surface, and echi-
nated by thick bundles of acanthostyles (64-472 X 5-19 f.lm); acan-
thostyles most abundant on basal layer of spongin; acallthostyies
EURYPON GRAY, 1867 completely spined, with bulbous tylote bases, and showing a large
size range; ectosomal skeleton with ectosomal styles (418-695 X
Synonymy 3-5 f.lm) forming plumose bouquets surrounding the base of where
subectosomal (sub)tylostyles pierce the surface, and also lying
Eurypon Gray, 1867a: 521; de Laubenfels, 1936a: 107; paratangential to surface. Microscleres absent.
Bergquist, 1970: 31. Epicles Gray, 1867a: 521; de Laubenfels, Remarks. The use of the name Eurypon over Epicles (type
1936a: 110; Bergquist, 1970: 31. Dragmatyle Topsent, 1904b: species Hymeraphia radiata Bowerbank, 1866, by monotypy),
195. Acantheurypon Topsent, 1927b: 15; Topsent, 1928c: 291. was discussed by Hooper (1991), where the latter has page-line
Tricheurypon Topsent, 1928c: 295; de Laubenfels, 1950a: 80; priority over the former. Similarly, Hooper (1991) broadened the
Wiedenmayer, 1977b: 159. Protoraspailia Burton & Rao, 1932: definition of Eurypon to include species with spined as well as
342. Leptochelastra de Laubenfels, 1936a: 111. Quixilla de smooth choanosomal megascleres (to accommodate species of
Laubenfels, 1936a: 88. Proraspailia Levi, 1958: 27. Taxonomic Acantheurypon (type species Hymeraphia pilosella Topsent,
decision for synonymy: Hooper (1991), and this work. 1904b, by original designation), with or without raphidiform
microscleres to allow the inclusion of species referred to
Type species Tricheurypon (type species Hymeraphia viridis Topsent, 1889,
by monotypy) and Protoraspailia (type species Proraspailia
Hymeraphia clavata Bowerbank, 1866 (by monotypy). polyplumosa Levi, 1958, by monotypy), and with the addition of
long raphidiform oxeote spicules scattered throughout the mesohyl
Definition (Protoraspailia, with type species ProtoraspaiZia cactoides Burton &
Rao, 1932: 343, by original designation). To this list should be
Typically encrusting Raspailiidae with microcionid skeletal added three other nominal genera, overlooked by the Hooper
structure in which fibre nodes ascend from the basal layer of spongin. (1991) revision. Dragmatyle (type species Dragmatyle Zictor
Topsent, 1904b, by original designation), is very similar to
Diagnosis Tricheurypon, and is included in synonymy here following the sug-
gestion of Hallmann (1917c). Dragmatyle has raphides in tri-
Encrusting, massive or digitate growth forms. Surface hispid, chodragmata, ectosomal oxeas (described as tomotes but usually
even, granular or conulose. Encrusting species have a microcionid sharply pointed and flexuous) forming tangential and paratangen-
choanosomal skeletal structure with a basally compressed layer of tial tracts (but not a specialised raspailiid skeleton), and also lacks
spongin fibres lying on the substrate producing small spongin-fibre echinating acanthostyles - thus representing a greatly reduced
nodes echinated by acanthostyles, and radially disposed extra-axial Eurypon. Leptochelastra de Laubenfels, 1936a (type species
skeleton composed of subectosomal styles standing perpendicular to Hymeraphia toureti Topsent, 1894a: 30, by original designation;
and embedded in basal fibres. Massive species have slightly axially slide of holotype MNHN DT1840, from the Banc de Campeche,
compressed plumose tracts of extra-axial styles, often forming fan- viewed by R.W.M. Van Soest, pers. comm.). It has tylostyles with
like bundles, and tracts lightly echinated by acanthostyles. Extra- grossly swollen heads, typical of Eurypon, in combination with
axial styles may be partially or entirely spined (Acantheurypon). short spined acanthostyles and bundles of thin oxeas gathered
Ectosomal specialization present (s.s.) or absent; if present ectosomal around the heads of the large tylostyles. Microscleres were not
skeleton consists of fine monactinal (or diactinal) spicule brushes observed. Topsent's (1894a) description mentions dubious chelae
surrounding single protruding extra-axial styles. Structural megas- and asters, but these are indeed foreign as he suspected. Quixilla de
cleres consist of 1-2 categories of styles or subtylostyles (rarely Laubenfels, 1936a: 88 (type species Hymeraphia lacazei Topsent,
modified to oxeas); echinating acanthostyles microcionid-like, typi- 1891a: 541, by original designation, type material not examined),
cally long and with subtylote bases. Several species (Tricheurypon, is also clearly a synonym of Eurypon. The type species, from
Dragmatyle = Eurypon) have raphides in trichodragmata. Roscoff, is thinly encrusting, with a highly hispid surface. Skeletal
structure is hymedesmioid, consisting of long subtylostyles (up to
Description of type species 2 mm long, 20 f.lm wide), with their bases embedded in basal spon-
gin and points protruding a long way through the surface, club-
Eurypon clavata (Bowerbank, 1866) (Fig. 10). shaped echinating acanthostyles with heavy spines distributed all
Synonymy. Hymeraphia clavata Bowerbank, 1866: 143. over the spicule (75-80 f.lm long, 10 f.lm wide), forming a dense
Eurypon clavatum; Vacelet, 1969: 186. Eurypon clavata; Hooper, erect palisade on the basal spongin, and ectosomal oxeas (230 f.lm
1991: 1313. long, 7 f.lm wide), forming a specialized raspailiid skeleton consist-
Material examined. Holotype: BMNH 1877.5.21.1556 ing of bundles surrounding the protruding long subtylostyles. No
(including 2 unregistered BMNH slides made from type, microscleres.
and MNHN DT942) - Shetland Is. Other material. BMNH This broad definition of Eurypon is consistent with the
1953.11.11.118. treatment of the diverse and similar (homologous) features
Porifera· Demospongiae • Poecilosclerida • Microcionina • Raspailiidae 489

A c

F
Fig. 10. Eurypon. A-F, E. clavatum, holotype. A, subectosomal subtylostyles (scale 200 f.Lm). B, ectosomal styles (scale 100 f.Lm). C, echinating acanthostyles
(scale 100 f.Lm). D, section through peripheral skeleton (scale 300 ILm). E, ectosomal skeleton (scale 100 f.Lm). F, microcionid fibre clump (scale 100 f.Lm).

in Microcionidae, in which genera containing smooth or spined 1970: 32), but given the diversity of those megascleres within obvi-
megascleres (e.g., Clathria Schmidt, and Dictyociona Topsent or ous microcionids (i.e., chelae-bearing taxa), in some instances this
Anaata de Laubenfels), and with or without microscleres (e.g., difference must be questioned also. It is possible that Eurypon
Clathria and Abila Gray) have been merged (van Soest, 1984b; will be eventually divided amongst Raspai/ia (species of Eurypon
Hooper, 1996a). This comparison with Microcionidae is most with ectosomal oxeas), Clathria (Clathria) (without ectosomal
appropriate for Eurypon, and many species have been removed megascleres), and Clathria (Thalysias) (with ectosomal styles or
from Raspailiidae (or Euryponidae) and synonymized with various subtylostyles).
microcionid genera (e.g., Eurypon asodes de Laubenfels, 1930: 27; Similarly, it could be argued that Eurypon and Hymeraphia
and E. microchela Stephens, 1916: 240 were transferred to are also potential synonyms, differing only substantially in having
Dictyociona (de Laubenfels, 1936a); E. rhopalophora (Hentschel, microcionid versus hymedesmioid skeletal structure, respectively,
1912: 380), and E. tenuissima Stephens, 1916: 240 were trans- and the possession of peculiar spination on acanthostyles in the lat-
ferred to Microciona (Burton, 1959a; Levi, 1960b». Those species ter genus. This may be a case for recognising these two taxa as sub-
are now recognized as merely being encrusting Clathria-like genera within a single genus (in which case Hymeraphia would
sponges (Hooper, 1996a). Another aspect which has been used to have priority), although this action is not formally proposed here
differentiate the raspailiid-like taxa (Eurypon) from microcionid without a larger study of the many encrusting raspailiid species and
species (Clathria s.l.) is acanthostyle morphology (e.g., Berguist, the diversity of their skeletal structures and spicule ornamentation.
490 Porifera' Demospongiae • Poecilosclerida • Microcionina • Raspailiidae

Eurypon differs from Raspailia in having only up to three cate- Description. Thinly encrusting growth form; choanosomal
gories of megascleres (echinating acanthostyles, ectosomal styles/ skeleton hymedesmioid with compressed basal layer of spongin in
oxeas, and a single category of long spicule which is equated with which the tylote bases of both long subectosomal (sub)tylostyles
the long subectosomal extra-axial megascleres found in typical (295-1875 X 6-19 fLm) and echinating acanthostyles (64-147 X
Raspailia); basal rather than axial compression of the fibre skeleton, 6-12 fLm) are embedded, standing erect on the substrate; acan-
lacking true choanosomal megascleres, and with exclusively plumose thostyles have slightly rhabdose, smooth, swollen tubular bases
architecture (or microcionid fibre structure in thinly encrusting and stellate-spined points; slender straight ectosomal oxeas
fonns). By comparison, typical Raspailia have four categories of (220-735 X 2-4 fLm) fonn bouquets around the point of penetra-
megascleres. This is, however, a tentative separation based on skeletal tion ofthe longer subectosomal (sub)tylostyles through the surface.
architecture (which is not always a reliable diagnostic character). Microscleres are absent.
Remarks. The type species was redescribed by Ackers et al.
Distribution (1992) from a living population. In their description they mention
the presence of a specialised raspailiid ectosomal skeleton of bou-
Worldwide. quets of oxeas surrounding protruding long styles, whereas in the
type specimen (and associated slide preparations) these spicules
were uncommon (and hence previously overlooked by Hooper,
HYMERAPHIA BOWERBANK, 1864 1991), and the specialised ectosomal skeleton was not observed in
type material preparations.
Synonymy Hymeraphia is unusual amongst Raspailiidae in its
hymedesmioid skeletal structure, without any observable fibre
Hymeraphia Bowerbank, 1864: 189 (Not Hymeraphia; nodes or ascending fibre tracts, superficially resembling the micro-
Hentschel 1912: 377). Mesapos Gray, 1867a: 543. Taxonomic cionid Leptoclathria (= Clathria) condition. Unlike the latter genus
decision for synonymy: Hooper (1991). Hymeraphia lacks any differentiated choanosomal or subectosomal
spicules, has modified acanthostyles, lacks microscleres, and has a
Type species specialised raspailiid ectosomal skeleton unique to the family. By
comparison, Hymeraphia sensu Hentschel (1912) is a microcionid,
Hymeraphia stellifera Bowerbank, 1864 (by original with chelae, toxas, and ectosomal megascleres that do not fonn
designation). surface bouquets. -ether thinly encrusting raspailiids, such as
Eurypon, have plumose skeletal columns whereas Hymeraphia is
Definition strictly hymedesmioid. Mesapos Gray is an objective synonym
(with type species Hymeraphia stellifera Bowerbank, 1864, by
Persistently encrusting Raspailiidae with hymedesmioid monotypy).
skeletal structure, echinating acanthostyles having bulbous tylote
bases, with or without spines on the points, and often with other Distribution
modifications to the distal portion of the spicule.
Most species records are from the North Atlantic and
Diagnosis Antarctica, whereas the genus is probably more widely distributed.

Thinly encrusting growth form. Choanosomal skeleton

reduced to basal membrane lying on substrate; bases of echinating RASPACIONA TOPSENT, 1936
acanthostyles and extra-axial styles embedded in basally com-
pressed fibres, in radial or hymedesmioid arrangement, standing
perpendicular to substrate, not grouped into brushes or other struc- Synonymy
tures. Choanosomal megascleres absent. Extra-axial styles usually
protrude a long way through the ectosome, surrounded by bouquets Raspaciona Topsent, 1936: 49; Sara, 1958: 254; Vacelet,
of ectosomal oxeas or styles at their point of penetration through 1961a: 36; Pulitzer-Finali, 1977: 41.
the surface, or the ectosomal skeleton may be reduced to a tangen-
tial layer of oxeas/styles scattered over the surface. Megascleres Type species
include longer styles or tylostyles, thinner ectosomal styles/ oxeas,
and smaller tylostyles with prominently swollen tylote bases echi- Halichondria aculeata Johnston, 1842 (by original designation).
nating the basal layer of spongin (those of type species also have
modified stellate-acanthose points and slightly rhabdose bases). Definition
Microscleres absent.
Rapailiidae without axial compression of the skeleton but
Description of type species with loosely aggregated or plumose tracts of a single category of
long styles.
Hymeraphia stellifera Bowerbank, 1864 (Fig. 11).
Synonymy. Hymeraphia stellifera Bowerbank, 1864: 189. Diagnosis
Material examined. Holotype: BMNH 1877.5.21.460
(fragments BMNH 1910.1.1.87, 2354, 2355, MNHN DT2501) - Lobo-digitate, or ramose-bushy growth forms; surface hispid
Shetland, English Channel. and prominently microconulose. Choanosomal skeleton plumose,
Porifera· Demospongiae • Poecilosclerida • Microcionina • Raspailiidae 491

I
Fig. 11. Hymeraphia. A-E, H. stellifera, holotype. A, subectosomal subtylostyles (scale 200 j.l-m). B, ectosomal oxeas (scale 100 j.l-m). C, echinating acan-
tho styles (scale 50 j.l-m). D, section through peripheral skeleton (scale 500 j.l-m). E, bundles of erect spicules (scale 50 j.l-m).

consisting of basally compressed spongin fibres ascending through Description of type species
branches and diverging, but without any special category of
choanosomal megasclere. Extra-axial skeleton with weakly devel- Raspaciona aculeata (Johnston, 1842) (Fig. 12).
oped, ascending, plumose columns of long flexuous subectosomal Synonymy. Halichondria aculeata Johnston, 1842: l3l.
styles protruding through the ectosome, branching or forming Raspaciona aculeata; Vacelet, 1961a:36.
occasional anastomoses, and forming shaggy surface micro- Material examined. Holotype: BMNH 1877.5.2l.956,
conules. Ascending extra-axial tracts echinated by acanthostyles, schizotype MNHN DNBE 320L - Mediterranean.
concentrated around the base of the plumose skeletal columns. Description. Lobo-digitate growth form; choanosomal
Spongin predominant in the basal region, at the point of contact skeleton microcionid with compressed basal layer of spongin with
between plumose (extra-axial) tracts and compressed (axial) fibres; erect fibre nodes cored by erect plumose columns of long thin and
very little spongin occurs elsewhere in skeleton. Ectosomal skele- flexuous subectosomal styles (1150--1610 X 9-18 f.1m), surrounded
ton with specialized spiculation of small styles arranged in bou- at their bases by bundles of small and evenly spined echinating
quets around the protruding extra-axial styles. Megascleres include acanthostyles (104-126 X 7-12 f.1m); extra-axial plumose columns
styles or subtylostyles of 2 sizes (ectosomal and subectosomal), of long styles protruding through surface, occasionally branching
and 1-2 size classes of acanthostyles ranging from basally spined and anastomosing; ectosomal skeleton with sparse bundles of flex-
to entirely spined. Microscleres absent. uous wispy ectosomal styles (334-638 X 3-6 f.1m) surrounding
492 Porifera· Demospongiae • Poecilosclerida • Microcionina • Raspailiidae

points of protruding extra-axial styles on the surface. Microscleres Distribution

absent.
Remarks. Raspaciona has a spiculation and ectosomal struc- Three species, all from the Mediterranean (Johnston, 1842;
ture typical of Raspailiidae, but a plumose choanosomal and subecto- Sara, 1958; Vacelet, 1961 a; Pulitzer-Finali, 1977).
somal architecture reminiscent of several genera of Halichondriidae.
Variability in skeletal morphology between specimens of different
growth forms has been well illustrated by Topsent (l925c) and RHABDEURYPONVACELET,1969
Pulitzer-Finali (1977). Those authors showed that megasclere size
and ornamentation varied quite considerably, and it has been sug- Synonymy
gested that Sara's (1958) R. robusta and R. calva are merely different
morphs of R. aculeata (e.g., Vacelet, 1961a). However, the acan- Rhabdeurypon Vacelet, 1969: 188.
thostyles of R. calva are illustrated with swollen tylote bases, much
the same as those found in Hymeraphia species (Sara, 1958: fig. 21). Type species
Although Raspaciona lacks any remarkable characters which can dis-
tinguish it readily from other genera, it can be differentiated from Rhabdeurypon spinosum Vacelet, 1969 (by original designation).
other species with plumose skeletal architecture (e.g., Aulospongus)
by its Raspailia-like spiculation, and from the typical Raspailia Definition
condition by its strictly plumose non-reticulate architecture.
Raspailiidae lacking echinating spicules but diactinal acan-
thorhabds are dispersed throughout the skeleton forming a com-

A pact mass.

Diagnosis

Thinly encrusting growth form. Surface even and hispid.

Choanosomal skeleton hymedesmioid with basally compressed
spongin lying on substrate. Choanosomal megascleres and true
echinating spicules absent, but mesohyl contains acanthorhabds
and spined microrhabds scattered throughout basal skeleton, with-
out apparent order. Extra-axial skeleton with long smooth subecto-
somal subtylostyles embedded in and perpendicular to substrate,
protruding through surface. Ectosome with specialized skeleton of
oxeas in bouquets surrounding extra-axial styles. Structural mega-
scleres smooth choanosomal styles or subtylostyles of one category,
ectosomal auxiliary oxeas, and diactinal acanthorhabds possibly
highly modified acanthostyles. Microscleres spined microrhabds.

D Description of type species

Rhabdeurypon spinosum Vacelet, 1969 (Fig. 13).

E Synonymy. Rhabdeurypon spinosum Vacelet, 1969: 188.
Material examined. Holotype: MNHN DN4 - Cassidaigne,
Mediterranean.
Description. Encrusting growth form; choanosomal skele-
ton hymedesmioid with basal layer of spongin lying on substrate
and bases of long subectosomal (sub)tylostyles (1120-3250 X
15-22/-Lm) embedded and their points penetrating the surface, sur-

c
rounded by bouquets of ectosomal oxeas (450-2000 X 2.5-6/-Lm)
at their point of insertion through the surface; basal skeleton is
a compact mass of acanthorhabds (40-110 X 3-8 /-Lm) orientated
in all directions, but not truly echinating. Spined microrhabds
(50-60 X 0.5--0.7 /-Lm) present, scattered throughout the mesohyl,
possibly juvenile forms of acanthorhabds.
Remarks. The diactinal acanthorhabds of Rhabdeurypon
are unusual, superficially resembling discorhabds of Negombo
(Halichondrida, Desmoxyidae), acanthoxeas of Histodermella
(Poecilosclerida, Coelosphaeridae), and also similar to diactinal
forms of acanthose spicules of Tethyspira (Halichondrida,
Fig. 12. Raspaciona. A-E, R. aculeata, holotype. A, subectosomal style
(scale 200 11m). B, ectosomal styles (scale 200 11m). C, echinating Dictyonellidae) and perhaps even Didiscus (Halichondrida,
acantbostyles (scale 30 11m). D, fibre (scale 200 11m). E, section through Desmoxyidae). If it were not for the possession of a specialized
peripheral skeleton (scale 500 11m). raspailiid skeleton (consisting of bouquets of ectosomal oxeas
Porifera • Demospongiae • Poecilosclerida • Microcionina • Raspailiidae 493

E
Fig. 13. Rhabdeurypon. A-F, R. spinosum, holotype. A,subectosomal subtylostyle (scale 500 j..l.m). B, ectosomal oxeas (scale 400 j..l.m). C, acanthorhabds
(scale 50 j..l.m). D ,spined microrhabds (scale 30 j..l.m). E, section through peripheral skeleton (scale I mm). F, holotype (scale 2 cm).

surrounding the protruding larger styles) and a plumose extra-axial THRINACOPHORA RIDLEY, 1885
skeleton (composed of larger (sub)tylostyles) the allocation of
Rhabdeurypon to any particular genus or family would be very dif- Synonymy
ficult. Vacelet (1969) noted that acanthorhabds are not echinating
but merely dispersed throughout the mesohyl - albiet forming a Thrinacophora Ridley, 1885: 572; Ridley & Dendy, 1886:
compact basal mass - and therefore perhaps they should not be 483, 1887: 193; Dendy, 1905: 186 (in part); Hallmann, 1917c:
regarded simply as modified acanthostyles. Hooper (1991) sug- 634-7; Hooper, 1991: 1339.
gested that the microscleres were smooth microxeas whereas more
detailed study of the type material confirms that these are spined Type species
and possibly represent juvenile forms of acanthorhabds.
Thrinacophora funiformis Ridley & Dendy, 1886 (by
Distribution monotypy).

Monotypic, so far known only from the Mediterranean Definition

(Vacelet, 1969).
Raspailiidae lacking echinating spicules, having a densely
compressed axial skeleton, without axial fibres, cored by short
SUBFAMILY THRINACOPHORINAE SUBFAM. NOV. oxeas or styles in a criss-crossed reticulation, and a plumose extra-
axial skeleton composed of longer styles or anisoxeas. Microsclere
Definition are raphides in trichodragmata.

Raspailiidae lacking echinating megascIeres, with well Diagnosis

differentiated axial and extra-axial skeletons.
Cylindrical, arborescent or encrusting and massive growth
Scope forms. Surface not prominently hispid but evenly conulose. Axial
and extra-axial skeletons well differentiated. Choanosomal skeleton
Three genera. Thrinacophora, Ceratopsion and Axechina. markedly axially compressed, with a dense reticulate core occupying
494 Porifera· Demospongiae • Poecilosclerida • Microcionina • Raspailiidae

Fig. 14. Thrinacophora. A-F, T. funiformis (holotype). A, holotype (scale 5 cm). B, choanosomal oxeas (scale 50 !Lm). C, subectosomal anisoxea (scale
200 !Lm). D, ectosomaI styles (scale 100 !Lm). E, trichodragmata (scale 50 !Lm). F, section through peripheral skeleton (scale 1 mm).

a large proportion of branch diameter composed of short, stout Description. Cylindrical digitate, branching growth form;
oxeas, anisoxeas or occasionally styles. Echinating megascleres choanosomal skeleton axially compressed with a dense reticulate
absent. Extra-axial skeleton plumose, with uni- or paucispicular core composed of short curved choanosomal oxeas (220-345 X
tracts radiating towards surface, cored by long, thick subectosomal 6-13 fLm) without apparent associated spongin fibres; extra-axial
styles or anisoxeas. Ectosomal skeleton with a special category of skeleton plumose to radial composed of long paucispicular tracts
styles (sinuous and apically pronged in the type species), forming containing long straight subectosomal styles and anisoxeas
erect bouquets or paratangential brushes around extra-axial (722-1640 X 16-24 fLm) radiating from the axis and protruding
spicules, and also scattered within the mesohyl. Structural mega- through surface; ectosomal styles, usually sinuous and apically
scleres short oxeas and long styles, some with strongylote or oxeote prolonged (345-462 X 4-6 fLm) form erect or paratangential bou-
modifications, and ectosomal styles. Microscleres raphides quets around protruding subectosomal tracts; microscleres
occurring singly or in bundles (trichodragmata). raphides occurring in trichodragmata (85-110 X 10-15 fLm).
Remarks. Hallmann (l917c) restricted Thrinacophora to
Description of type species include only species containing a specialized ectosomal skeleton
composed of styles. He redistributed other nominal species (i.e.,
Thrinacophorafuniformis Ridley & Dendy, 1886 (Fig. 14). Thrinacophora, sensu Dendy, 1905) amongst his new genera
Synonymy. Thrinacophora funiformis Ridley & Dendy, Dragmaxia, Dragmacidon and Axidragma (=Tragosia Gray) in
1886: 484. the Axinellidae. However, these Thrinacophora clearly belong to
Material examined. Holotype: BMNH 1887.5.2.53 - off the Raspailiidae as indicated by their special raspailiid ectosomal
Bahia, Brazil. skeleton.
Porifera' Demospongiae • Poecilosclerida • Microcionina • Raspailiidae 495

Thrinacophora is most similar to Ceratopsion, both possess- from a continuous palisade on the surface to discrete bouquets
ing a specialized ectosomal skeleton, large extra-axial megascleres of spicules grouped around the projecting extra-axial megas-
and a more-or-less radial arrangement of the extra-axial skeleton cleres where they penetrate the surface. Structural megascleres
which protrudes a long way through the surface, well developed include at least two categories of styles, anisoxeas or strongyles
axial and extra-axial differentiation, and lacking echinating mega- (one often sinuous), oxeas or anisoxeas, and thin ectosomal oxeas
scleres. Thrinacophora has a dense axial reticulation of short, stout or styles. Microscleres usually absent or may include raphides in
choanosomal spicules, usually oxeas but occasionally anisoxeas or trichodragmata.
rarely styles, and this axial region lacks any evidence of spongin
fibres. The criss-cross of axial spicules, which occupies most of the Description of type species
diameter of branches, differentiates Thrinacophora from Raspailia
(Parasyringella). Earlier authors (e.g., Ridley & Dendy, 1887; Ceratopsion expansa (Thiele, 1898) (Fig. 15).
Hallmann, 1916b) considered that the possession of raphides was Synonymy. Ceratopsis expansa Thiele, 1898: 57.
an important generic diagnostic character, and placed secondary Ceratopsion expansa; Hooper, 1991: 1327.
emphasis on ectosomal specialization (used here to define the Material examined. Holotype (fragment): MNHN
Raspailiidae), megasclere geometry and skeletal architecture. In DCL981L - Sagami Bay, Japan. Comparative material. Holotype
Thrinacophora, at least, it is unlikely that the possession of of Ceratopsion dichotoma (Whitelegge): AM G4353 (see Hooper,
raphides can define the genus (as supposed by Ridley & Dendy, 1991).
1887: 193), and more recently this was confirmed by the descrip- Description. Erect, palmate-flabellate, digitate growth
tion of a species of Ceratopsion bearing raphides (Alvarez & Van form; choanosomal skeleton a compressed axial reticulation of
Soest, 1993). fibres cored by robust choanosomal oxeas (230-270 X 4-11 J.Lm)
forming longitudinal tracts through branches, with a well differen-
Distribution tiated radial extra-axial skeleton composed of curved subectosomal
styles (750-1200 X 20-30 J.Lm) embedded in and perpendicular to
Four species are known so far, from Brazil (Ridley & Dendy, axis and protruding through surface; ectosomal skeleton composed
1886, 1887), Puerto Rico (Wilson, 1902), Philippines (Ridley & of small ectosomal oxeas (80-110 X 2-3 J.Lm) forming a nearly
Dendy, 1887), Arafura Sea and Moluccas, Indonesia (Kieschnick, continuous palisade on surface mainly clustered around the pro-
1896; Hentschel, 1912) and Northwest Australia (Hooper, 1991). jecting bundles of subectosomal styles. Microscleres absent.
Remarks. Ceratopsion includes species which have either
monactinal or diactinal axial megascleres and extra-axial mega-
CERATOPSION STRAND, 1928 scleres (Hooper, 1991). Hallmann (1916b) used Ceratopsion as
a catch-all for various 'Axinellidae with microxeas' (excluding
Synonymy and type species sigma-bearing and desmoxyid genera). However, Bergquist (1970)
showed that these 'microxeas' were in fact ectosomal megascleres,
[Ceratopsis] Thiele, 1898: 57 (preocc.); Hallmann 1916b: 541. raphidiform in some species, and on this basis the genus was
Ceratopsion Strand, 1928: 33; de Laubenfels 1936a: 132; Bergquist referred to Raspailiidae (Hooper, 1991).
1970: 18; Hooper, 1991: 1327; Alvarez & Van Soest, 1993: 629; Ceratopsion lacks echinating megascleres, but has other fea-
Hooper & Levi, 1993: 1287. Ommatosa de Laubenfels, 1936a: 132. tures found in most raspailiids. On paper the genus is difficult to
distinguish from Raspailia (Parasyringella), but the presence of
Type species sinuous axial styles or anisoxeas in some species, and the posses-
sion of distictly radial extra-axial skeleton of styles, anisoxeas or
Ceratopsis expansa Thiele, 1898 (by original designation). strongyles may serve to differentiate these two genera. The genus
was previously defined as lacking raphides whereas Alvarez & Van
Definition Soest (1993) described a species from the Florida Keys with tri-
chodragmata, and they remark further on the affinities between
Raspailiidae lacking echinating spicules, having an axially Thrinacophora and Ceratopsion.
compressed choanosomal skeleton with reticulate fibres cored by Only two species of Ceratopsion have a 'classic' ectosomal
sinuous styles or anisoxeas and a well differentiated radial extra- skeleton consisting of bouquets of ectosomal styles or oxeas sur-
axial skeleton cored by longer megascleres. rounding the projecting extra-axial megascleres (c. cuneiformis
Bergquist, C. montebelloensis Hooper), whereas all other species
Diagnosis have a more dense palisade of ectosomal brushes erect on the sur-
face. Ceratopsion davata, C. microxephora and C. minor form a
Erect, lamellate or arborescent growth forms. Surface granu- group possessing sinuous strongyles in the axial skeleton, reminis-
lar and hispid. Axial and extra-axial skeletons well differentiated. cent of the Bubaridae. Axinella aurantiaca Lendenfeld, 1888,
Choanosomal skeleton a compressed axial reticulation of spongin should also be transferred to Ceratopsion.
fibres, deficient in collagen, cored by sinuous or straight styles, Ommatosa de Laubenfels was previously considered a junior
anisoxeas or strongyles. Echinating megascleres absent. Extra- synonym of Bubaris by Carballo (2000: 585), as also suggested by
axial skeleton of radially arranged subectosomal styles, anisoxeas Hooper & Levi (1993) for Bubaridae, whereas re-examination
or strongyles, including sinuous andlor straight forms, embedded of the holotype of the type species, Axinella rugosa (Schmidt,
in and perpendicular to axial core, projecting through surface. 1870: 61), by Belinda Alvarez (pers. comm.) confirmed that it
Specialized ectosomal skeleton present (s.s.) or absent, and where belongs to Ceratopsion, and furthermore the type species was the
present consisting of oxeas or styles forming structures ranging senior synonym of C. crustosum Alvarez & Van Soest, 1993.
496 Porifera· Demospongiae • Poecilosclerida • Microcionina • Raspailiidae

c D

Fig. 15. Ceratopsion. A, C. expansa (fragment of holotype). A, spicules (scale 500 /Lm). B-G, C. dichotoma (holotype). B, holotype (scale 3 em).
C, subectosomal style (scale lOO/Lm). D, choanosomal styles (scale loo/Lm). E, ectosomal style (scale loo/Lm). F, ectosomal skeleton (scale 2oo/Lm).
G, section through peripheral skeleton (scale 500 /Lm).

Distribution Type species

Eleven species are known, from Florida (Schmidt, 1870; de Axechina raspailioides Hentschel, 1912 (by original
Laubenfels, 1936a; Alvarez & Van Soest, 1993), Mediterranean designation).
(Pulitzer-Finali, 1983: 520), Japan (Thiele, 1898), New Zealand
(Bergquist, 1970: 18), north, east and west coasts of Australia Definition
(Whitelegge, 1907; Hooper, 1991), New Caledonia (Hooper &
Levi, 1993), South Africa (Kirkpatrick, 1903b: 242), and Indonesia Raspailiidae lacking echinating acanthostyles, having an axi-
(Hentschel, 1912). ally compressed choanosomal skeleton divided into two compo-
nents: a central tightly reticulate region cored by oxeas with spined
points, and a peripheral radial region formed by smooth anisoxeas
AXECHINA HENTSCHEL, 1912 projecting outwards.

Synonymy Diagnosis

Axechina Hentschel, 1912: 417; de Laubenfels, 1936a:102; Bushy flabellate growth form. Surface rugose, conulose, and
Hooper, 1991: 1344. hispid. Choanosomal skeleton with two components: compressed,
Porifera· Demospongiae • Poecilosclerida • Microcionina • Raspailiidae 497

B
Fig. 16. Axechina. A-F, A. raspailoides. A, specimen NTM Z2385 (scale 3 em). B-F, holotype. B, subectosomal style (scale 500 j.Lm). C, choanosomal
anisoxeas (scale 100 j.Lm). D, choanosomal spined oxeas (scale 50 j.Lm). E, ectosomal styles (scale 300 j.Lm). F, peripheral skeletal structure (scale 150 j.Lm).

tightly reticulate axis cored by oxeas with spined points; radial Description. Bushy arborescent, stalked, cylindrical
peripheral region formed by columns of smooth anisoxeote branches, prominently conulose or clathrous; surface prominently
spicules projecting outwards from axis. Echinating megascleres hispid; choanosomal skeleton with well differentiated axial and
absent. Extra-axial skeleton radial, with tracts of long, thick, extra-axial regions; axial skeleton with two components:
subectosomal styles embedded in the outer (radial) portion of the a compressed axial formed by tightly reticulate tracts of choanoso-
axis, ascending to surface and projecting through it. Ectosomal mal oxeas with spined points (195-264 X 8-16 fLm) forming a
skeleton composed of peculiarly curved or sinuous toxiform styles dense central criss-cross running longitudinally through branches,
with spined terminations, grouped around a central extra-axial and a plumose or radial region at the periphery of the axial skeleton
style in a bouquet. Structural megascleres include oxeas with formed by columns of smooth anisoxeote spicules (305-425 X
spined terminations, smooth anisoxeas, large styles or anisoxeas 16--24 fLm) embedded in the axis and directed outwards singly or in
with rounded bases, and dermal flexuous toxiform styles with paucispicular bundles, often also penetrating the surface; extra-
spined terminations. Microscleres absent. axial skeleton radial, with long subectosomal styles (1044-2760 X
15-20 fLm) embedded in the radial portion of the axial skeleton,
Description of type species projecting a long way through the surface; ectosomal skeleton with
bouquets of sinuous ectosomal styles (231-305 X 2-6 fLm)
Axechina raspailioides Hentschel, 1912 (Fig. 16). surrounding the longer extra-axial styles; echinating spicules
Synonymy. Axechina raspailioides Hentschel, 1912: 417. absent; microscleres absent.
Material examined. Holotype: SMF991 (schizotype MNHN Remarks. The type and only species of Axechina has a typical
DCL2261) - Arafura Sea. raspailiid ectosome formed by bouquets of special styles, clearly
498 Porifera· Demospongiae • Poecilosclerida • Microcionina • Raspailiidae

indicating its affinities with this family although previously the no or few anastomoses. Extra-axial skeleton with few very long
genus was included in Axinellidae (Hentschel, 1912). In fact the thin extra-axial styles, embedded in choanosomal fibre nodes and
axial skeletal architecture and possession of megascleres with protruding through surface. Extra-axial spicules also scattered
spined terminations is reminiscent of Reniochalina (AxineUidae) throughout mesohyl. Fibres very heavily echinated by sagittal tri-
but this comparison is superficial and they have little else in com- acts, tetracts, or forms even further modified, producing almost
mon. Axechina could possibly be reasonably included with rigid interlocking secondary skeleton. Ectosomal specialization
Ceratopsion on the basis of its spiculation, or Thrinacophora in absent. Structural megascleres styles or subtylostyles of 2-3 sizes;
having at least part of its axial skeleton consisting of a dense criss- thinner may have few spines and subtylote swellings on distal and
cross reticulation of spicules without obvious fibres, but it fits into basal ends, respectively; echinating spicules sagittal tetracts or
neither genus easily. The genus is retained on a tentative basis, dif- pentactinal megascleres (acanthoplagiotriaenes), less commonly
ferentiated from others by the presence of the following features: diacts or triacts, with all (s.s.) or at least more than one acanthose
an axial reticulate core, a radial (or plumose) region at the periph- ray. Microscleres absent.
ery of the axis, a radial extra-axial skeleton, and a plumose ectoso-
mal skeleton, the absence of choanosomal fibres or definite axial Description of type species
spicule tracts (cf. Thrinacophora), the presence of basal and apical
spination on both choanosomal and ectosomal megascleres, and Cyamon vickersii (Bowerbank, 1866) (Fig. 17).
the absence of echinating megascleres. Synonymy. Dictyocylindrus vickersii Bowerbank, 1866: 267.
Cyamon vickersii; Dendy, 1922b: 108.
Distribution Material examined. Holotype: BMNH 1877.5.21.1887 -
West Indies.
Monotypic, endemic to southeast Indonesia and northwest Description. Lamellate, columnar, cushion-shaped or
Australia. massive-encrusting growth forms; choanosomal skeleton microcionid
with compressed basal layer of spongin with individual fibre nodes
erect on basal spongin, cored by smaller stout choanosomal styles
(564-745 X 19-30 /-Lm) in columns of 2-4 spicules, also projecting
SUBFAMILY CYAMONINAE SUBFAM. NOV. from fibres in plumose tufts; extra-axial skeleton with diverging
plumose columns of both choanosomal styles and longer thin
Definition subectosomal styles (935-1368 X 9-14 /-Lm) embedded in micro-
cionid fibres and protruding through surface; peripheral
Raspailiidae with echinating spicules modified to sagittal fibres echinated by sagittal triactinal, tetractinal and pentactinal
acanthoplagiotriaenes, with one species having trichodragmata. acanthoplagiotriaenes with all rays bearing spines (98-132 X
10-16 /-Lm), and including juvenile forms (superficially resembling
Scope orthotriaenes of the Astrophorida) bearing vestigial spination
on all rays (62-125 X 3-8 /-Lm); acanthoplagiotriaenes form an
Three genera. Cyamon Trikentrion and Waltheramdtia. almost interlocking secondary skeleton within the choanosome;
ectosomal skeleton without specialised brushes typical of most
raspailiids but with ectosomal auxiliary styles (212-415 X
CYAMON GRAY, 1867 2-6 fLm) scattered between skeletal columns and forming bundles
within choanosomal and subectosomal regions. Microscleres
Synonymy absent.
Remarks. Bowerbank's (1866) original description of
Cyamon Gray, 1867a: 546; Dendy, 1922b: 107; de Laubenfels, C. vickersii is inadequate and barely characterises the genus,
1936a: 80; Levi, 1973: 609; Sim & Bakus, 1986: 17. whereas Carter's (1879b: 292) redescription of the type specimen
is much more detailed and useful. The BMNH holotype is dry but
Type species still in good condition, and from that material (and accompanying
slide preparations) it is confirmed that skeletal structure consists of
Dictyocylindrus vickersii Bowerbank, 1866 (by monotypy). plumose columns of choanosomal and subectosomal styles, there is
no special ectosomal skeleton but a thinner (third, ectosomal) cate-
Definition gory of styles is present within the choanosomal skeleton, and
there is also an almost interlocking secondary skeleton composed
Raspailiidae with acanthostyles modified to sagittal triactinal of sagittal acanthoplagiotriaenes. This structure is quite different
or tetractinal acanthoplagiotriaenes with all rays spined. from Trikentrion, but it is probably one related to Cyamon's growth
form. Nevertheless, there are also significant differences in the
Diagnosis geometry and spination of sagittal acanthoplagiotriaenes between
the two genera: Trikentrion has monact-, diact-, triact- or rarely
Encrusting to massive growth form. Surface hispid and usu- tetractinal sagittal spicules, with only one spined ray; Cyamon
ally conulose. Choanosomal skeleton basally compressed layer of (s.s.) has rarely triactinal, mostly tetractinal and sometimes pen-
spongin fibres lying on substrate, with microcionid-like plumose tactinal sagittal spicules, and all rays are spined, at least at their
spongin-fibre nodes ascending to surface. Basal fibres cored by apex. Dendy (l922b: 108) and de Laubenfels (1936a: 80) also
pauci- or multispicular plumose tracts of choanosomal styles, with redescribed additional material of C. vickersii from Amirante
Porifera· Demospongiae • Poecilosclerida • Microcionina • Raspailiidae 499

CD EA
I

Fig. 17. Cyamon. A-G, C. vickersii. A, holotype (scale 3cm). B, specimen BMNH 1931.1.1.19 (scale 5cm). C, subectosomal style (scale 250IJ.m).
D, choanosomal styles (scale 200 IJ.m). E, ectosomal styles (scale 100 IJ.m). F, acanthoplagiotriaenes including juvenile form (scale 100 IJ.m). G, section
through peripheral skeleton (scale 500 IJ.m).

(BMNH 1931.1.1.19; Fig. 17B) and the West Indies, respectively, TRlKENTRION EHLERS, 1870
although de Laubenfels (1936a) questioned the conspecificity of
Dendy's material. Nevertheless, from those accounts it appears that Synonymy
Cyamon differs from Trikentrion in several significant respects and
is maintained here as a separate genus. Trikentrion Ehlers, 1870: 6, 31; Carter, 1879b: 291; Hentschel,
1912: 373; Topsent, 1928c: 58; de Laubenfels, 1936a: 80; Levi,
Distribution 1973: 609. [Tricentrium] Ridley, 1879 (in Zoological Record 16: 5)
(lapsus). Plectronella Sollas, 1879b: 17; de Laubenfels, 1936a: 81.
Seven species of Cyamon are presently known: West Indies Taxonomic decision for synonymy: Hooper (1991).
(Bowerbank, 1866; de Laubenfels, 1936a), southern California (de
Laubenfels, 1930; Dickinson, 1945; Sim & Bakus, 1986), Gulf of Type species
Manaar, Indian Ocean (Carter, 1880b), and Indonesia (Hentschel,
1912; Hooper, 1991). Spongia muricata (Pallas, 1766) Esper, 1794 (by monotypy).
500 Porifera· Demospongiae • Poecilosclerida • Microcionina • Raspailiidae

c
I,

Fig. 18. Trikentrion. A-G, T. muricata. A, specimen BMNH 1872.10.19.1 (scale 3 em). B-G, schizotype 2MB7160. B, section through peripheral skele-
ton (scale 300 fLm). C, choanosomal oxeas (scale 50 fLm). D, subectosomal style (scale 100 fLm) . E, ectosomal styles (scale 50 fLm). P, aeanthoplagiotriaenes
(scale 50 fLm) . G, trichodragmata (scale 20 fLm). H-J, T. flabelliforme Carter. H, specimen (NTM Z2383) (scale 5 em). I, section through choanosomal
skeleton (scale 500 fLm). J, aeanthoplagiotriaenes in situ (scale 100 fLm).

Definition with only slightly compressed axial fibres running longitudinally

through axial core. Fibres cored by pauci- or multispicular tracts
Raspailiidae with acanthostyles modified to sagittal monacti- of oxeas. Axial and extra-axial skeletons not markedly differenti-
nal, diactinal or tetractinal acanthoplagiotriaenes with only one ated; extra-axial skeleton consists of ascending multi- or
spined ray. paucispicular primary fibres cored by choanosomal oxeas, intercon-
nected by uni- or paucispicular secondary fibres, together producing
Diagnosis a more-or-Iess regular reticulation. Single or brushes of subectoso-
mal styles embedded at ends of ascending primary fibres, poking
Arborescent, digitate and flabellate growth forms. Surface through peripheral skeleton. Peripheral fibres moderately heavily
usually hispid, and even or rough and microconulose. Choanosomal echinated by sagittal triacts. Ectosome with specialized skeleton of
skeleton with poorly developed wide meshed reticulation, ectosomal styles in brushes surrounding bases of extra-axial styles.
Porifera' Demospongiae • Poecilosclerida • Microcionina • Raspaillidae 501

Megascleres styles of two sizes, choanosomal oxeas, echinating Definition

megascleres sagittal triacts (or acanthoplagiotriaenes) with only
one spined ray. Microscleres may include raphides, occurring Cyamoninae which have lost sagittal triaenes but retain
singly or in bundles (= trichodragmata). a renieroid reticulate skeleton slightly compressed in the axial
region and more open with elongate meshes in the periphery.
Description of type species
Diagnosis
Trikentrion muricata (Pallas, 1766) (Fig. 18).
Synonymy. Spongia muricata (Pallas, 1766) Esper, 1794: Raspailiidae, monotypic, flabellate, stalked, with regularly
185. Trikentrion muricata; Carter, 1879b: 293. spaced depressed oscules scattered over an optically smooth sur-
Material examined. Holotype: Unknown, schizotype face; specialized ectosomal skeleton present consisting of bundles
2MB7160 - West Africa. of microxeas with spined tips surrounding protruding choanosomal
Description. Arborescent, digitate growth form; choanoso- styles; choanosomal skeleton renieroid reticulate with differenti-
mal skeleton with wide meshed reticulation, with little or no axial ated primary multi spicular tracts interconnected by uni- or pau-
compression, composed of relatively close-meshed paucispicular cispicular tracts of styles forming regular meshes, slightly
tracts cored by choanosomal oxeas (235-320 X 8-18 /-Lm); extra- compressed in the axis and more elongate and regular in the
axial skeleton plumoreticulate with ascending primary multispicu- periphery forming longitudinal serial meshes, with choanosomal
lar tracts interconnected by transverse tracts of choanosomal oxeas; styles protruding at the surface. Megascleres consist of styles in
sparse subectosomal styles (555-964 X 6-15 /-Lm) embedded at two size classes, the smaller with spined bases, and ectosomal
ends of primary ascending spicule tracts protruding a short dis- microxeas with spined tips. Echinating sagittal spicules absent.
tance through the surface, surrounded by sparse brushes of ectoso-
mal anisoxeas, occasionally styles (195-315 X 2-4/-Lm) forming Description of type species
bouquets around protruding subectosomal styles; peripheral fibres
echinated by nearly symmetrical sagittal diacts with only the Waltheramdtia caliculatum (Kirkpatrick) (Fig. 19).
longest ray spined at its apex (132-165 X 12-18 /-Lm); microscleres Synonymy. Hymeniacidon caliculatum Kirkpatrick, 1903b:
are raphides in trichodragmata (65-85 X 10-15 /-Lm). 241-242, pI. 5, fig. 8, pI. 6, fig. 12. Hymeniacidon caliculatum var.
Remarks. Trikentrion is similar to Cyamon in having highly osculatum Kirkpatrick, 1903b: 242, pI. 6, fig. 13.
modified echinating spicules called sagittal triacts (or acanthopla- Material examined. Holotype: BMNH 1902.11.16.20
giotriaenes), usually triradiate or further modified to quadriradiate (slide) - South Africa, E London, 170 m, Gilchrist collection. Other
and other forms. On this basis alone these genera could be eventu- material. Holotype of H. caliculatum var. osculatum: BMNH
ally synonymised but they differ in other significant details: 1902.11.16.21 - South Africa, E London coast, 170m, Gilchrist
Trikentrion has only one spined rayon echinating spicules, it collection.
has diactinal choanosomal spicules, a non-plumose skeleton, and Description (partly from Kirkpatrick, 1903b and re-examination
a specialized raspailiid ectosomal skeleton. For those reasons of type material). Flabellate, with a thick flat lamella, 0.7-1.5 cm
they are maintained separately here (see also remarks for thick, originally 19 cm high, 17 cm wide, on a broad stalk. Surface
Cyamon). optically smooth but rough to touch due to projecting tufts of
Plectronella (type species Plectronella papillosa Sollas, spicules, with a subsurface lace-like reticulation visible below the
1879b, by monotypy) was merged with Trikentrion in 1879 (by dermal membrane. Oscules small 2 mm in diameter on one face of
Ridley in the Zoological Record for that year), and again by de the lamella, in oscular depressions. Colour brick red alive, orange
Laubenfels (1 936a), when it was found that its type species was brown in alcohol. Ectosomal skeleton with microxeas in brushes
synonymous with T. muricata from West Africa. surrounding the protruding choanosomal megascleres or lying
paratangential to the ectosome. Choanosomal skeleton renieroid
Distribution reticulate with multispicular primary tracts (50-100/-Lm diameter)
connected by uni- to paucispicular secondary ones, at relatively
Four species are presently known: West Africa (Esper, 1874; regular intervals, forming rectangular meshes approximately 200 X
Ehlers, 1870; Carter, 1879b), Gulf of California (Dickinson, 1945), 50 /-Lm in dimension. Meshes arranged serially and longitudinally in
tropical Australia and Indonesia (Carter, 1882b; Hallmann, 1914b; a central lamella, more compressed near the axis of lamella than at
Hooper, 1991). the surface where they become elongated with respect to the sur-
face. Megascleres larger styles thick and slightly curved, with
rounded bases, 210-(233.8}-290 J.Lm long, 5-(l5.9}-20 J.Lm wide;
WALTHERARNDTIA DE LAUBENFELS, 1936 smaller styles with 1-3 microspines on the base, 115-{141.3}-225 J.Lm
long, 5-(5.3}-8 J.Lm wide; microxeas with a row of 3 or more (up to
Synonymy 8) spines near their tips, 40-(63.8}-75 J.Lm long, < 1 J.Lm wide.
Remarks. The genus is monotypic and originally assigned
Waltheramdtia de Laubenfels, 1936a: 131. to Axinellidae by de Laubenfels (1936a). However, both the
spicule complement and the organisation of the skeleton do not
Type species agree with the concept ofAxinellidae adopted in this work (see
chapter by Alvarez de Glasby), and the possession of a specialized
Hymeniacidon caliculatum Kirkpatrick, 1903b: 241 (by 'raspailiid' ectosomal skeleton indicates closer affinities with
original designation). Raspailiidae, but its inclusion in this family remains a tentative
502 Porifera· Demospongiae • Poecilosclerida • Microcionina • Raspailiidae

Fig. 19. Waltherarndtia de Laubenfels. A-F, holotype Hymeniacidon caliculatum var. osculatum Kirkpatrick, BMNH 1902.11.16.21 . A-C, spicules (scale
50 /Lm). D, SEM of tips of microxeas. E, holotype. F, SEM of choanosomal skeleton (scale 100 /Lm).

hypothesis. Kirkpatrick (1903b: 242) noted that the choanosomal Distribution

skeleton was transitional between renierioid and axinellid architec-
ture, showing both a slight axial compression at the core of the Known only from the East London coast, South Africa.
lamella and regular reticulate meshes near the surface. While its
affinities to Raspailiidae are obvious from the possession of a special-
ized 'raspailiid' ectosomal skeleton, its relationships within SUBFAMILY PLOCAMIONINAE SUBFAM. NOV.
Raspailiidae remain speculative. It is tentatively assigned to the
Cyamoninae based on the possession of styles with spines at their Definition
base which occur in other Raspailiidae, and that the ectosomal skele-
ton composed of microxeas bearing basal spines is reminiscent of Raspailiidae with acanthostrongyles or acanthotylostrongyles
Trikentrion flabelliformis (Alvarez de Glasby and Van Soest, pers. ('peg-top' spicules) forming choanosomal skeletal tracts.
comm.; who have both personally examined the type material), in
addition to the renieroid and slightly compressed choanosomal skele- Scope
tal architecture reminiscent of other Cyamoninae. In this placement it
is hypothesized that the genus has lost its sagittal spicules. Two genera. Plocamione and Lithoplocamia.
Porifera· Demospongiae • Poecllosclerida • Microcionina • Raspailiidae 503

PLOCAMIONE TOPSENT, 1927 order Poecilosclerida related to the Myxillidae, Crellidae or

Microcionidae (Plocamia Schmidt, 1870, [Dirrhopalum] Ridley
Synonymy in Ridley & Duncan, 1881, Lissoplocamia Brondsted, 1924b,
Heteroclathria Topsent, 1904c, Plocamiopsis Topsent, 1928c,
Plocamione Topsent, 1927b: 16; 1928c: 63. Raspeloplocamia Echinoplocamia Burton, 1959a, Plocamionida Topsent, I 927b,
Burton, 1935a: 402. Axoplocamia Burton, 1935a: 402; Bergquist & Plocamissa Burton, 1935a, Plocamiancora Topsent, 1927b,
Fromont, 1988: 122. Taxonomic decision for synonymy: Hooper Damiria Keller, 1891, Damiriella Burton, 1935a, and Damiriopsis
(1991), and this work. Burton, 1928a). Two of these belong to the Raspailiidae:
Plocamione (including Raspeloplocamia, with type species
Type species Plocamia clopetaria Schmidt, 1870, by original designation; and
Axoplocamia with type species Bubaris ornata Dendy, 1924, by
Plocamione dirrhopalina Topsent, 1927b (by monotypy). original designation), and Lithoplocamia (see below). Both have
modified acanthostyle 'peg top spicules' but differ essentially in
Definition their skeletal structure and disposition of these spicules.
Five of the six known species are typical of Raspailiidae in
Raspailiidae with an axially or basally compressed micro- having a specialized raspailiid ectosomal skeleton and curved or
cionid skeleton of which the axial skeleton is formed by vermiform 'sausage shaped' spicules (acanthostrongyles) which
acanthostrongyles or acanthotylostrongyles. are characteristic of 'plocarniform' sponges (sensu Burton, 1935a).
Specialised ectosomal bouquets and 'sausage shaped' spicules are
Diagnosis absent in P. pachysclera (Levi & Levi), but that species has other
features that suggest a close relationship with the group. Several
Encrusting or digitate growth forms. Surface even and other species have also been included in this genus or one of its
hispid. Choanosome basally or axially compressed reticulation of synonyms by authors at one time or another but these have been
choanosomal acanthostrongyles, forming thick longitudinal core; demonstrated to have affinities with Microcionidae (Hooper, 1991,
echinating acanthostyles embedded in core, standing perpendicular 1996a). This includes Plocamia gymnazusa Schmidt, the type
to axis. Extra-axial skeleton radial or plumose columns of long species of Plocamia Schmidt (not Plocamia Lamouroux) and
subectosomal styles. Ectosomal skeleton with or without special- [Dirrhopalum] Ridley in Ridley & Duncan, 1881, which Hooper
ized skeleton of ectosomal styles or anisoxeas forming brushes (1996a) showed was a microcionid related to Antho.
around extra-axial spicules. Structural megascleres include two
categories of styles or subtylostyles; echinating spicules evenly Distribution
spined acanthostyles with subtylote bases, sometimes entirely
smooth, partially acanthose, or with smooth, spined or tuberculate Six species are known from the North Atlantic (Duncan in
bases. Choanosomal axial or basal acanthostrongyles or acanthoty- Ridley & Duncan, 1881; Topsent, 1927b) and southwest Pacific
lostrongyles, straight or slightly curved, distinctly acanthose, (Dendy, 1924; Levi & Levi, 1983b).
merely tuberculate, or rarely entirely smooth (= 'peg-top' spicules
of Ridley in Ridley & Duncan, 1881). Microscleres absent.
LITHOPLOCAMIA DENDY, 1922
Description of type species
Synonymy
Plocamione dirrhopalina Topsent, 1927b (Fig. 20).
Synonymy. Plocamione dirrhopalina Topsent, I 927b: 16. Lithoplocamia Dendy, 1922b: 79. ? lanulum de Laubenfels,
Material examined. Holotype: MOM (not seen), schizotype 1936a: 79. Monectyon Levi & Vacelet, 1958: 236. Taxonomic deci-
MNHN DTl245 - Azores. Comparitive material. P. clopetaria sion for synonymy: Hooper (1991), and this work.
(Schmidt, 1870): fragments of holotype MNHN DCLlI06L,
BMNH 1870.5.3.78. P. ornata (Dendy, 1924): holotype BMNH Type species
1923.10.1.126. P. pachysclera (Levi & Levi, 1983b): holotype
MNHN DCL2448. Lithoplocamia lithistoides Dendy, 1922b (by monotypy).
Description. Encrusting growth form; choanosomal skele-
ton hymedesmioid with basal layer of spongin on substrate cored Definition
by longitudinal tracts of choanosomal acanthostrongyles (70-95 X
7-10 /-Lm) forming thick basal crust or loosely reticulate basal Raspailiidae with a secondary basal reticulate skeleton lack-
skeleton, with bases of echinating acanthostyles (175-280 X ing any trace of axial compression, formed by (sub )isodictyal tracts
11-14/-Lm) and long subectosomal styles (800-3500 X 18-35 /-Lm) of acanthostrongyles or acanthotylostrongyles.
embedded in spicule reticulation and standing perpendicular to
substrate; ectosomal skeleton with bouquets of ectosomal styles Diagnosis
(455-700 X 3.5-5 /-Lm) surrounding protruding subectosomal
spicules. Massive or semi-encrusting growth forms. Surface often
Remarks. There is a large number of species and genera sculptured by drainage canals. Choanosomal skeleton lacks axial
of 'plocarniform' sponges which have a basal or axial skeleton compression, consisting of a dense regular isodictyal (s.s.) or irreg-
of acanthose diactinal megascleres, but most of those taxa are ular subisodictyal secondary reticulation of acanthostrongyles.
relatively poorly known. Most are obvious members of the Primary skeleton consists of radial or plumose tracts of styles
504 Porifera· Demospongiae • PoeciloscIerida • Microcionina • Raspailiidae

B
I

Fig. 20. Plocamione. A-F, P. dirrhopalina (holotype). A, subectosomal style (scale 500 !Lm). B, ectosomal style (scale 200 !Lm). C, echinating acan-
thostyle (scale 100 !Lm). D, choanosomal acanthostrongyles (scale 50 !Lm). E, section through peripheral skeleton (scale 500 !Lm). F, choanosomal spicule
bundle (scale 300!Lm). G-J, P. pachysclera (holotype). G, holotype (scale 2cm). H, basal end of large extra-axial style (scale 50!Lm). I, choanosomal
styles (scale 100 !Lm). J, acanthostyles (scale 50 !Lm). K, P. ornata (holotype), section through peripheral skeleton (scale 200 !Lm).

ascending to the surface and protruding slightly through it, only Synonymy. Lithoplocamia lithistoides Dendy, 1922b: 79.
visible near the surface and usually obscured by the dense second- Material examined. Holotype: BMNH 1921.11.7.68
ary isodictyal reticulation. Ectosome with specialized skeleton of Mauritius.
long, slender oxeas forming sparse bouquets at the ends of primary Description. Massive growth form; choanosomal skeleton
skeletal tracts. Structural megasc\eres are smooth styles or subty- composed of a subisodictyal (secondary) reticulation of acan-
lostyles, ectosomal oxeas, diactinal or pseudodiactinal acanthostrongyles (122-226 X 12-24 /-Lm) occupying majority of
thostrongyles, with or without (s.s.) echinating acanthostyles. choanosome, with radial or plumose tracts of choanosomal styles
Microsc\eres absent. (252-427 X 9-14 /-Lm) running towards surface; no axial compres-
sion or differentiation between axial and extra-axial regions;
no true echinating spicules present; ectosomal skeleton present
Description of type species consisting of bouquets of long thin rhaphidiform oxeas (322-
435 X 3-5 /-Lm) surrounding the protruding choanosomal spicules.
Lithoplocamia lithistoides Dendy, 1922b (Fig. 21). Subectosomal spicules and rnicrosc\eres absent.
Porifera· Demospongiae • Poecilosclerida • Microcionina • Raspailiidae 505

·B

Fig. 21. Lithoplocamia. A-F, L. lithistoides (holotype). A, choanosomal styles (scale 100 fLm). B, ectosomal oxeas (scale 100 fLm). C, acanthostrongyles
(scale 100 !Lm). D, holotype (scale 2cm). E, ectosomal skeleton (scale 200 fLm) . F, choanosomal subisodictyal reticulation (scale 200 fLm).

Remarks. The diagnosis of this genus has been emended curved acanthostyles characteristic of Lithoplocamia (and other
above from Hooper (1991) due to the discovery of some additional 'plocamiid' sponges), but if any relationship exists between these
features overlooked in the type material. The genus shows some two forms of megascleres it is probably more likely that the reverse
superficial similarities to Antho (Microcionidae) and Damiria is true. Levi & Levi (1983b) also compare Lithoplocamia with
(Acarnidae), and Dendy (1922b) suggested that the type species Endectyon, the former having a massive and non-branching growth
was most closely related to Plocamia massalis (Microcionidae). form and a compact isodictyal or subisodictyal reticulation of
However, Lithoplocamia is clearly a raspailiid given the presence acanthostrongyles, whereas Endectyon has acanthose spicules
of a specialised raspailiid skeleton, and similarities with poe- echinating plumose columns of choanosomal axial styles. In this
cilosclerid 'plocamiform' sponges are based on their common pos- regard Lithoplocamia must also be compared with the renieroid
session of a secondary isodictyal skeleton of acanthose spicules. reticulate raspailiid genus Amphinomia Hooper.
Dendy (1922b) and Levi & Levi (l983b) remark on the close The nominal genus Monectyon (type species Monectyon
resemblance in the basal skeletal construction between this genus atlanticus Levi & Vacelet, 1958, by monotypy) was described as
and 'Lithistida' with monocrepidial desmas (e.g., Lithobubaris having a skeleton composed only of acanthostrongyles, without
Vacelet =Sulcastrella Schmidt). Intermediate forms also exist (e.g., other megascleres present at all, but Levi & Levi (1983b) suggest
Cerbaris Topsent; family Bubaridae). Dendy speculates further that additional material of M. atlanticus may eventually show
that monocrepidial des mas may have developed from the that the species does possess true ectosomal auxiliary and
506 Porifera· Demospongiae • Poecilosclerida • Microcionina • Raspailiidae

subectosomal extra-axial spicules, and consequently Hooper Definition

(1991) synonymised Lithoplocamia and Monectyon.
lanulum de Laubenfels, 1936a: 79 (type species Isodictya Raspailiidae with an exclusively reticulate choanosomal
spinispiculum Carter, 1876: 310, by original designation), also skeleton, without any trace of axial compression, cored exclusively
appears to belong to this genus. The type species, from 'Cape by smooth oxeas, and vestigial radial extra-axial and ectosomal
St. Vincent', is described as laminiform, thickly encrusting, with skeletons.
porous, even surface and friable consistency. Fibre skeleton forms
areolate wide-meshed isodictyal reticulation, with fibres cored by Diagnosis
regular lines of acanthostrongyles arranged end-to-end (plocami-
form arrangement). Acanthostrongyles are peculiar in that they Erect, vasiform, flabellate, ramose or massive growth forms.
are bent at each end, in the same direction, and heavily spined at Surface typically rugose, with ridges, conules and other processes.
the middle of the spicule but not at the ends. It is possible that there Texture characteristically harsh, brittle and flexible, reflecting high
are other types of spicules present, overlooked by Carter (1876), ratio of silica to spongin in skeleton. Axial and extra-axial skele-
but the type specimen, if it still exists, has not been located and tons virtually undifferentiated. Choanosomal skeleton not com-
re-examined. This species seems close to the plocamiform group pressed, irregularly reticulate. Spongin fibres usually massive,
of raspailiids, in particular Lithoplocamia, to which it is assigned fully cored by oxeas, and echinated by acanthostyles. Extra-axial
with question (pending rediscovery of the type material). skeleton vestigial, consisting of individual subectosomal styles
embedded in peripheral fibres, and projecting through ectosome or
Distribution merely dispersed between fibres. Ectosomal skeleton usually mem-
braneous, sometimes skin-like, typically without specialized spicu-
Known only from Mauritius and Seychelles (Dendy, 1922b), lation (except in one species). Ectosomal region contains heavy
New Caledonia (Levi & Levi, 1983b), and eastern Atlantic (Levi & deposits of type B spongin, frequently with dense deposits of pig-
Vacelet, 1958), all relatively deep water species. ment granules. Structural megascleres are oxeas of 1-2 sizes,
together with acanthostyles, long or short subectosomal styles, and
short slender ectosomal styles in one species. Microscleres absent.

SUBFAMILY ECHINODICTYINAE SUBFAM. NOV. Description of type species

Echinodictyum mesenterinum (Lamarck, 1814) (Fig. 22).

Definition Synonymy. Spongia mesenterina Lamarck, 1814: 444.
Echinodictyum mesenterinum; Carter, 1882b: 114. Spongia bilamel-
Raspailiidae with regularly reticulate choanosomal skeletal lata Lamarck, 1816: 436 (in part, var. B). Echinodictyum bilamella-
structure, extra-axial skeleton vestigial or virtually absent, and all tum; Ridley, in Ridley & Duncan, 1881: 493. Kalykenteron elegans
but one species lacks ectosomal specialisation. Echinating mega- Lendenfeld, 1888: 216. Echinodictyum elegans; Hallmann, 1912:
scleres are microcionid-like club-shaped acanthostyles. 171. Kalykenteron silex Lendenfeld, 1888: 217. Echinodictyum
topsenti de Laubenfels, 1936a: 63. Thalassodendron typica;
Scope Whitelegge, 1901: 86 (in part) (Not Thalassodendron typica
Lendenfeld, 1888: 233). Echinonema vasiplicata Carter, 1882b: 114.
Two genera. Echinodictyum and Amphinomia. Material examined. Lectotype: MNHN DT568 - 'Australian
Seas'. Comparative material. Refer to extensive list in Hooper
(1991).
Description. Erect, vasiform or cup-shaped, on short stalk;
ECHINODICTYUM RIDLEY, 1881 harsh texture, only slightly compressible or flexible, difficult to
tear; exterior surface conulose, prominently fibrous, greatly undu-
lating, interior surface more compressed; ectosome membranous,
Synonymy without specialized spiculation, with tympanic skin-like membrane
stretched between adjoining surface conules and subectosomal
Echinodictyum Ridley, in Ridley & Duncan, 1881: 493; Ridley, styles (112-318 X 4-7f.Lm) piercing surface skeleton mainly on
1884a: 454; Ridley & Dendy, 1887: 164; Topsent, 1894c: 19; Dendy, conules; choanosomal skeleton irregularly reticulate, without axial
1896:44,1905: 175, 1916a: 129; Thiele, 1899: 15;EJeschnick, 1900: compression or any axial and extra-axial differentiation, meshes
570; Hentschel, 1911: 385, 1912: 369; Hallmann, 1912: 171-5; cavernous, ovoid; spongin fibres heavy, multi spicular, cored by
Burton, 1931a: 348; Burton & Rao, 1932: 347; Levi, 1965: 19, 1969: choanosomal oxeas (112-276 X 3-15 /-Lm); echinating acan-
966; Hooper, 1991: 1347. Kalykenteron Lendenfeld, 1888: 216; thostyles (71-105 X 6-11 /-Lm) with prominent swollen, spined
Hallmann, 1912: 171. Kieplitela de Laubenfels, 1954: 116. bulb at apex, evenly dispersed over fibres.
Taxonomic decision for synonymy: Hooper (1991). Remarks. Echinodictyum, including synonyms Kaly-
kenteron (type species K. elegans Lendenfeld, 1888, by subsequent
Type species designation; Hooper, 1991), and Kieplitela (type species
K. antrodes de Laubenfels, 1954, by monotypy), is atypical of
Spongia bilamellata, var. B, Lamarck, 1816 (in part) (by orig- Raspailiidae. Its affinities with the family lie mainly in the
inal designation), junior synonym of Spongia mesenterina presence of echinating acanthostyles and subectosomal styles,
Lamarck, 1814 (Topsent, 1932a: 101). which appear to be remnants of an extra-axial skeleton. This weak
Porifera· Demospongiae • Poecilosc1erida • Microcionina • Raspailiidae 507

D F

Fig. 22. Echinodictyum . A-F, E. mesenterinum. A, specimen NTM ZI171 (scale Scm). B-C, specimen NTM Z264S. B, SEM section through choanoso-
mal skeleton (scale SOO !Lm). C, SEM section of echinated fibre (scale 200 !Lm). D-F, holotype. D, subectosomal style (scale 100 !Lm). E, echinating
acanthostyles (scale SO !Lm). F, choanosomal oxeas (scale 100 !Lm). G, E. nidulus (specimen NTM Z2882), ectosomal specialisation (scale 100 ILm).

evidence, however, was strengthened by the discovery of a special between Echinodictyum and the microcionid genus Echinochalina
raspailiid ectosomal skeleton in one species (E. nidulus) (Hooper, Thiele. Echinochalina is reticulate, with minimal spongin, with
1991). Echinodictyum is most similar to the nominal raspailiid either oxeas, quasi-monactinal or quasi-diactinal megascleres cor-
genus Clathriodendron, the type species of which also has a pre- ing fibres, and it has smooth (s.s.) echinating styles (Hooper,
dominantly reticulate skeleton, lacks any specialized ectosomal 1996a), whereas Echinodictyum invariably has acanthostyles.
skeleton, and in which the fibres are echinated by microcionid-like
acanthostyles (i.e., with evenly distributed recurved spines, and
with or without an aspinose 'neck'). Echinodictyum is exclusively Distribution
reticulate, with only vestigial development of the extra-axial skele-
ton (occurring as individual extra-axial styles), and choanosomal Thirty six species are known so far from the Indo-west Pacific
megascleres are always short, stout oxeas. By comparison, region, and probably at least double this number worldwide.
Clathriodendron has more obvious affinities with Raspailia, with Species have been described from circum-Australia (see Hooper,
at least some differentiation of the axial and extra-axial skeleton 1991), central Indian Ocean (Carter, 1886d; Dendy, 1905; Thomas,
and with choanosomal styles or modified styles coring well 1968b), western Indian Ocean (Carter, 1879b), South Africa
developed spongin fibres. The comparison should also be made (Burton, 1931a), Arabian Gulf (Ridley, 1881), Indo-Malay
508 Porifera· Demospongiae • PoeciloscIerida • Microcionina • Raspaillidae

Fig. 23. Amphinomia. A-G, A. sulphurea (holotype). A, extra-axial styles (scale 50 !Lm). B, choanosomal styles (scale 100 !Lm) . C, echinating acan-
thostyles (scale 50 !Lm). D, holotype (scale 3 cm). E, echinated fibre (scale 50 !Lm). F, choanosomal renieroid reticulation (scale 50 !Lm). G, choanosomal
skeleton (scale 500 !Lm).

archipelago (Thiele, 1889; Brondsted, 1929), central west Atlantic Definition

(Hechtel, 1983), North Atlantic (Levi, 1969), west Africa (Topsent,
1906a), Red Sea (Keller, 1889; Topsent, 1892b; Levi, 1965), Raspailiidae having a renieroid reticulate choanosomal skele-
Mediterranean (Topsent, 1892b), and West Indies (Duchassaing & ton, lacking any axial compression, cored by choanosomal styles
Michelotti,1864). with spines on both basal and distal ends.

Diagnosis

AMPHINOMIA HOOPER, 1991 Massive flabellate-Iobate growth form. Surface fleshy, uneven
and not hispid. Choanosomal skeletal architecture regularly
Synonymy renieroid-reticulate, without any axial compression or marked dif-
ferentiation between axial and extra-axial skeletal tracts. Spicules
Amphinomia Hooper, 1991 : 1321. and fibres form ascending multispicular tracts, interconnected by
uni- or paucispicular transverse tracts, and fibres are lightly echi-
Type species nated by acanthostyles. Extra-axial skeleton vestigial, with subec-
tosomal styles scattered throughout the skeleton especially near the
Amphinomia sulphurea Hooper, 1991 (by original designation). periphery. Ectosomal skeleton thickly membraneous, without
Porifera· Demospongiae • Poecilosclerida • Microcionina • Raspailiidae 509

Fig. 24. Cantabrina. A-E, C. erecta. A, paratype (scale I em). B, style (scale 200 j-Lm). C, section through peripheral skeleton (scale 300 j-Lm).
D-E, MNHN fragment of paratype. D, rhabdostyle (scale 100 iJ,m). E, section through peripheral skeleton (scale 400 j-Lm).

specialized spiculation. Megascleres include smooth subectosomal renieroid reticulate, with ascending multispicular tracts intercon-
styles, choanosomal styles with large spines on both basal and dis- nected by transverse uni- or paucispicular tracts near centre of
tal ends, and evenly spined echinating acanthostyles. Microscleres skeleton but more obviously renieroid near periphery, with triangu-
absent. lar, hexagonal or square meshes; light spongin fibres cored by
choanosomal styles with spined points and bases (152-274 X
Description of type species 18-26 J.1m), some uncored; echinating acanthostyles dispersed over
fibres (102-138 X 4-11 J.1m); subectosomal extra-axial styles scat-
Amphinomia sulphurea Hooper, 1991 (Fig. 23). tered between fibres throughout skeleton (178-248 X 4-12 J.1m);
Synonymy. Amphinomia sulphurea Hooper, 1991: 1322. microscleres absent.
Material examined. Holotype: NTMZ1787 - Amphinome Remarks. Amphinomia differs from other raspailiids in its
Shoals, Northwest Shelf, Western Australia. regularly renieroid reticulate skeleton, without any trace of axial
Description. Massive, thickly lobate-lamellate growth form, compression, cored by choanosomal styles bearing both basal and
with flabellate lobes, and rounded or slightly undulating margins; distal spination. These megascleres are unusual but not unique
large oscules on edges of lobes; surface thickly glabrous, amongst Porifera. Hooper (1991) documents several other species
fleshy, prominently crinkled and uneven, produced by irregular, with similar spicules, many from the older literature (e.g., Bowerbank,
interconnected low ridges, depressions and microconules; ectoso- 1864), although none of them are raspailiids. Amphinomia
mal skeleton membraneous, heavily collagenous, with few sulphurea is superficially similar to the massive suberitids
choanosomal styles protruding; choanosomal skeleton regularly (Hadromerida) in growth form and texture; it also shows superficial
510 Porifera· Demospongiae • Poecilosclerida • Microcionina • Raspailiidae

similarities in skeletal construction to members of the order Diagnosis

Agelasida, Poecilosclerida families Microcionidae (such as Antho
(]sopenectya», Acarnidae (Acamus) and Myxillidae (Myxilla). Erect lobate growth form. Conulose and hispid surface.
Based on its affinities with Lithoplocamia, and in possessing the Choanosomal skeleton halichondroid in the axis composed of a
remnants of an extra-axial skeleton and echinating acanthostyles, disorganized criss-cross of long thick styles. No fibres present.
the genus was referred to Raspailiidae although it has potential Subectosomal extra-axial skeleton increasingly plumose towards
affinities to the Microcionidae and Acamidae. It differs from the periphery, with long thick styles protruding through surface
Lithoplocamia by the different geometries of their basal choanoso- only at the tips of conules. Echinating megascleres rare. Ectosomal
mal spicules (monactinal versus diactinal, respectively), and the lat- skeleton with long thin styles, sinuous or straight, in sparse bundles
ter having a secondary (sub)isodictyal reticulate skeleton composed associated with plumose tracts near the surface. Structural mega-
of spined acanthostrongyles, an extra-axial plumose or radial skele- scleres are long styles of two categories and rare smooth echinating
ton of smooth choanosomal styles, lacking echinating acanthostyles styles with slightly rhabdose bases. Microscleres absent.
and having a specialised raspailiid ectosomal skeleton.
Acanthostylotella Burton & Rao, 1932 (type species Stylotella Description of type species
comuta Topsent, 1897a: 464, from Ambon, Indonesia and the
Maldives) is described as having a skeleton composed of uni- to Cantabrina erecta Ferrer-Hernandez, 1914 (Fig. 24).
paucispicular tracts of styles, smooth or with terminal rudimentary Synonymy. Cantabrina erecta Ferrer-Hernandez, 1914a: 453.
spines, forming an irregular anisodictyal reticulation with weak Material examined. Paratype. BMNH 1930.1.21.9, schizo-
spongin fibres. The subsequent descriptions of Topsent (1897a), type MNHN LBIM DCL174L-Spain.
Burton & Rao (1932) and Desqueyroux-Faundez (1981) are still Description. Erect lobate bulbous growth form. Surface
very inconclusive as to its true identity, with the possibility that it conulose and hispid. Ectosomal skeleton membraneous, with pau-
belongs amongst the Raspailiidae, and with terminal spination cispicular bundles of thinner (? ectosomal) styles (long thin,
on choanosomal styles (perhaps superficially) reminiscent of straight or sinuous; 640-920 X 4-6/Lm) clustered around the
Amphinomia. However, no true echinating acanthostyles or extra- plumose choanosomal tracts in the peripheral skeleton, although
axial styles were reported and it allocation remains incertae sedis these do not appear to penetrate the surface; subectosomal skeleton
until original material is re-examined. plumose, with multispicular ascending tracts of thicker
(? choanosomal ) styles (very long, thick, with evenly rounded
Distribution base; 950-1475 X 12-22/Lm) diverging towards and penetrating
the surface only on the tips of conules; choanosomal skeleton
Monotypic, endemic to northwest Australia. halichondroid reticulate, with disorganized criss-cross of thicker
(? choanosomal) styles. No fibres present. Mesohyl only lightly
invested with spongin towards the axis becoming more dense
CANTABRINA FERRER-HERNANDEZ, 1914 near the periphery. Echinating styles entirely smooth with slightly
(INCERTAE SEDIS) rhabdose base (220 X 6/Lm), very rare or possibly contaminants.
Structural megascleres consist of two categories of long styles, the
Synonymy larger often with telescoped points and rounded bases, and smooth
echinating styles with slightly rhabdose bases. Microscleres
Cantabrina Ferrer-Hernandez, 1914a: 453; Ferrer-Hernandez, absent.
1914b: 22; Hooper, 1991: 1394. Remarks. The affinities of this monotypic genus are still
uncertain. Echinating megascleres supposedly native to this
Type species species (Ferrer-Hernandez, 1914a,b) are extremely rare in the
paratype (including several BMNH slide preparations) a spicule
Cantabrina erecta Ferrer-Hernandez, 1914 (by monotypy). slide also made from this material in MNHN. It is possible that
these rhabdostyles are contaminants in which case it may belong to
Definition Halichondriidae given its classic halichondrid skeletal structure.

Raspailiidae with rare echinating rhabdostyles completely Distribution

smooth, a halichondroid reticulate choanosomal skeleton becom-
ing plumose near the periphery. Monotypic, Spain.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Rhabderemiidae Topsent, 1928

John N.A. Hooper

Queensland Museum, P.O. Box 3300, South Brisbane, Qld, 4101, Australia. (JohnH@qm.qld.gov.au)

Rhabderemiidae Topsent (Demospongiae, Poecilosclerida) contains four nominal genera of which only one is now valid, currently with
26 described species predominantly from shallow waters of all tropical and warm temperate seas (with one deep water species known
from 1360 m depth). The family is still of uncertain relationship, having a unique apomorphy in the form of smooth or spined rhabdostyles
bearing a basal spiral twist, and peculiar usually rugose microscleres resembling oxeas or toxas (thraustoxeas), sigmas (spirosigmata,
thraustosigmata) and microstyles, although these are probably analogous to toxas and sigmas found in other poecilosclerids. Species are
usually thinly encrusting but also include branching, club-shaped, digitate or massive forms, usually fleshy, with species differing mainly
in their microsclere composition and geometry and spicule dimensions.
Keywords: Porifera; Demospongiae; Poecilosclerida; Microcionina; Rhabderemiidae; Rhabderemia.

DEFINITION, DIAGNOSIS, SCOPE 1936a; Bergquist, 1968). Hooper (1990a) returned the family to
Poecilosclerida on the basis that its monactinal choanosomal
Synonymy megascleres, the diversity and geometry of microscleres are
homologous to those of poecilosclerids. However, Van Soest &
Rhabderemiidae Topsent, 1928c: 64. Hooper (1993) suggest that there is doubt over the homology of the
sigmoid toxiform microscleres between Rhabderemia and other
Definition poecilosclerids (hence the use of the terminology of 'spirosigmata'
and 'thraustosigmata' as opposed to 'normal sigmata' and 'thraust-
Microcionina of uncertain relationship, with smooth or oxea' as opposed to 'normal oxeas'), given their persistently
spined rhabdostyles bearing an extra spiral twist at the rounded spined condition, but they retained the family taxon for 'practical
extremity and microscleres include peculiar toxiform (thraus- purposes'.
toxeas), sigmoid-like (spirosigmata, thraustosigmata) and rugose This definition of Rhabderemiidae in the Poecilosclerida
microstyles. was developed from those of contemporary authors (Levi, 1973;
Bergquist, 1978; Hartman, 1982; Hooper, 1990a; Van Soest &
Diagnosis Hooper, 1993), whereas earlier definitions (Topsent, 1928c; de
Laubenfels, 1936a) were non-committal in the higher taxonomic
Thinly and thickly encrusting, massive, lobate, anastomosing, placement of the family. The 26 described species in this mono-
digitate, arborescent and lobate-flabellate growth forms; megascle- generic family are nearly cosmopolitan (excluding arctic and
res consist of smooth or distally spined monactinal choanosomal antarctic seas), and confined largely to cryptic habitats (particu-
rhabdostyles bearing a basal spiral twist forming hymedesmioid larly in coral reefs). Some morphological trends are apparent
(Fig. lA-B), plumose, plumo-reticulate (Fig. 2B) or reticulate amongst species: rhabdostyles (an autapomorphy for the family)
skeletal structures, usually with poorly developed spongin fibres, may lose their spiral twist in the rhabdose end and lose spination on
with rhabdostyles usually forming diverging plumose tracts within their distal ends; and microstyles, thraustoxeas, contorted sigmata
fibres; microscleres if present include rugose oxeote or toxa-like and thraustosigmata may be lost in some species (Van Soest &
spicules (thraustoxeas), rugose sigma-like spicules (spirosigmata, Hooper, 1993).
thraustosigmata) and rugose microstyles.
Differences with similar families
Scope
Rhabderemiidae shows some similarities in skeletal structure
Monogeneric, with 26 described species predominantly tropical- and spiculation to the Raspailiidae, particularly Hemectyonilla
subtropical. Burton and Aulospongus Norman (Hooper, 1991; Hooper et al.,
1999), but was excluded from it in having unique rugose sigmoid,
History and biology toxiform and microstylote microscleres (whereas most raspailiids
lack microscleres, or when present they consist only of smooth
Rhabderemiidae is a monogeneric family since the included raphides). Rhabderemiids also lack any evidence of an extra-axial
genera (Rhabderemia, Rhabdosigma, Nisibaris and Stylospira) skeleton, but this feature is not always obvious in many raspailiids
were synonymised (Hooper, 1990a; Van Soest & Hooper, 1993). either (Hooper, 1991). Although typical raspailiids such as
Until recently the family was included in the polyphyletic order Raspailia Nardo have well differentiated axial and extra-axial
Axinellida (Levi, 1973; Hartman, 1982), although it has also skeletons, others such as Echinodictyum Ridley have simply reticu-
been assigned to the Poecilosclerida (e.g., Topsent, 1928c) late skeletons, but generally in the Raspailiidae there is always at
and Hadromerida (in the family Spirastrellidae) (de Laubenfels, least some remnants of an extra-axial skeleton. Although some

511
512 Porifera· Demospongiae • Poecilosclerida • Microcionina • Rhabderemiidae

A
I
I
c

Fig. 1. A-F, Rhabderemia minutula (Carter) (holotype). A, rhabdostyles (scale 50 !Lm). B, base of rhabdostyle (scale 10 !Lm). C, spirosigmata (scale 10 !Lm).
D, microstyles (scale 10 !Lm). E, ectosomal skeleton (scale 250 !Lm). F, choanosomal skeleton (scale 100 !Lm). G, Rhabderemia sorokinae Hooper,
in situ, Papua New Guinea (photo author) .

Fig. 2. Idealised rhabderemiid skeletal structures. A, hymedesmioid species. B, plumo-reticulate species.

Porifera· Demospongiae • Poecilosclerida • Microcionina • Rhabderemiidae 513

degree of relationship is inferred between these two families, with Description of type species
vague similarities appearing in skeletal architecture and the presence
of rhabdostyles in genera of both (e.g., Rhabderemia and Rhabderemia minutula (Carter, 1876) (Figs lA-F).
Aulospongus, respectively), this relationship is still not completely Synonymy. Microciona minutula Carter, 1876: 479;
clear. Although the possession of rhabdostyles partially defines Rhabderemia minutula; Van Soest & Hooper, 1993: 323 (Not
Rhabderemiidae these also occur in other demosponges: the Rhabderemia minutula; Topsent, 1904b: 152; Topsent, 1918: 541;
Poecilosclerida families Raspailiidae~Aulospongus Norman, Levi, 1956b: 393; Boury-Esnault, 1971b: 306; Biblioni & Gili,
Raspaxilla Topsent, Echinaxia Hallmann, Axinectya Hallmann, 1982: 231; Biblioni et al., 1990: 327; Pulitzer-Finali, 1983:
Hemectyon Topsent) and Desmacellidae (Biemna Gray); and the 533; Hooper, 1990a: 77). Microciona pusilla Carter, 1876: 239
Halichondrida families Desmoxyidae (Halicnemia Bowerbank, (lapsus); Microciona pusilla Carter, 1880a: 437; Rhabderemia
Higginsia Higgin), and Bubaridae (Rhabdoploca Topsent, pusilla; Topsent, 1892a: 116 (Not Rhabderemia pusilla; Dendy,
Hymerhabdia Topsent, Monocrepidium Topsent). These spicules are 1922b: 85).
likely homoplastic developments, derived independently within each Material examined. Holotype: BMNH 1902.11.16.32 -
group, and there are some significant but subtle differences in rhab- uncertain locality, possibly Caribbean. Other material. See below
dostyle morphology between these groups to support this contention (also refer to Van Soest & Hooper, 1993: 324).
(Hooper et al., 1999). For example, rhabdostyles of Rhabderemia Description. Encrusting and partially insinuating (excavat-
usually have an extra spiral twist at the rounded extremity (i.e., in a ing) growth form; choanosomal skeleton hymedesmioid with basal
third plane), whereas amongst the rhabdose raspailiids there is either layer of spongin charged with microstyles and rhabdostyles erect on
only a slight basal rhabd, represented by a small curvature at the base basal spongin, singly or in sparse plumose bundles; ectosomal
of the spicule, or in species with a well developed basal rhabd this skeleton charged with microscleres predominantly microstyles and
occurs in only two planes. Nevertheless, the status of this mono- sigmoid spicules; megascleres rhabdostyles bearing spirally twisted
generic family is still uncertain, with the suggestion that the family base and slightly rugose or more prominently spined distal end,
taxon may be superfluous (Van Soest & Hooper, 1993). possibly 2 size categories (81-352X4-9flom); microscleres rugose
microstyles with subtylote subterminal swellings (90-135 X 1 flom),
Previous reviews and rugose sigmata-like spicules either irregular or contort
spirosigmata (9-18 flom long).
Hooper (1990a), Van Soest & Hooper (1993). Remarks. Carter's (1876) taxon M. pusilla is invalid
because he emended the name himself to 'minutula' in a subse-
RHABDEREMIA TOPSENT, 1890 quent publication (Carter, 1880a), and therefore the earliest avail-
able name for the type species is M. minutula. Van Soest & Hooper
Synonymy (1993) redescribed the type species based on recent material from
the Caribbean, restricting the species to shallow-water West Indies
Rhabderemia Topsent, 1890b: 28; Topsent, 1892a: 115; populations, hence the number of invalid published records of the
Dendy, 1905: 180; Topsent, 1928c: 309; de Laubenfels, 1936a: 144; species from other localities (many of which represented new
Levi, 1973: 606; Hooper, 1990a: 66; Van Soest & Hooper, 1993: sibling species).
323. Rhabdosigma Hallmann, 1916b: 520; Hallmann, 1917b: 398; The synonymy of Rhabderemia proposed by Hooper (1990a)
Topsent, 1928c: 312; de Laubenfels, 1936a: 144; Hooper, 1990a: and Van Soest & Hooper (1993) is confirmed through examination
66; Van Soest & Hooper, 1993: 323. Stylospira de Laubenfels, of relevant type material: Rhabdosigma Hallmann (type species
1934: 10. Nisibaris de Laubenfels, 1936a: 144; Hooper, 1990a: 66; Sigmaxinella mammillata Whitelegge, 1907 (by original desig-
Van Soest & Hooper, 1993: 323. [Hallmannia] Burton, 1931a: 352 nation) holotype AMG4356); Stylospira de Laubenfels (type
(preocc.); Hooper, 1990a: 66 (not Hallmannia Burton, 193Oc: 519). species Stylospira mona de Laubenfels, 1934 (by original designa-
Taxonomic decision for synonymy: Van Soest & Hooper (1993). tion) holotype USNM22324); and Nisibaris de Laubenfels (type
species Hallmannia spirophora Burton, 1931a: 352 (by original
Type species designation) holotype NM1254). Hallmannia sensu Burton
(1931a: 352) is not con specific with Hallmannia Burton (1930c:
Microciona pusilla Carter, 1876 (by subsequent designation; 519), with the former referred to R. spirophora (Burton) by Van
Dendy, 1905: 180) (junior synonym of Microciona minutula Carter, Soest & Hooper (1993), and the latter (having seniority) a junior
1880b; Van Soest & Hooper, 1993: 323). synonym of Biemna (with B. aruensis Hentschel as its type
species).
Definition
Distribution
As for family.
Found predominantly in shallow waters of all tropical and
Diagnosis warm temperate seas, although one species has been recorded
from 1360 m (Boury-Esnault et al., 1995), none from colder waters
As for family. (Van Soest & Hooper, 1993).
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Suborder Myxillina Hajdu, Van Soest & Hooper, 1994

Rob W.M. Van Soest

Zoological Museum, University of Amsterdam, P.O. Box 94766,1090 GT Amsterdam, The Netherlands. (soest@science.uva.nl)

Suborder Myxillina Hajdu et ai. (Demospongiae, Poecilosclerida) are characterized by possession of tridentate chelae and absence of
toxas. Most Myxillina have differentiated choanosomal and ectosomal megascleres, but this may be lost secondarily. Eleven families are
distinguished based on microsclere morphology, megasclere types, and skeletal architecture. A key to the families is provided.
Keywords: Porifera; Demospongiae; Poecilosclerida; Myxillina; Chondropsidae; Coelosphaeridae; Crambeidae; Crellidae;
Dendoricellidae; Desmacididae; Hymedesmiidae; Iotrochotidae; Myxillidae; Phellodermidae fam. nov.; Tedaniidae.

DEFINITION, DIAGNOSIS, SCOPE skeletal plan or - in thinly encrusting forms - perpendicular to and
penetrating the surface, and macroscopically hispid. The auxiliary
Definition megascleres tend to be smaller, usually entirely spined, echinating the
skeletal tracts, the nodes of the skeletal reticulum or - in thinly
Poecilosclerida with tridentate or polydentate chelae micro- encrusting forms - arranged in groups ('bouquets') around a single
scleres; palmate chelae absent; toxas absent; sigmas usually present. main megasclere. In one family they simulate ectosomal spicules to
Differentiated ectosomal and choanosomal megascleres, although form a surface crust. Auxiliary megascleres are frequently lost or in
either or all may be lost secondarily. Ectosomal megascleres typi- some cases undifferentiated in shape from the main megascleres.
cally diactinal, commonly with aniso-terminations. Choanosomal Several groups have their choanosomal skeleton partly or wholly
megascleres usually styles, rarely oxeas or strongyles. replaced by a reticulation of sand grains and other foreign material.
Chelae. Chelae deviate from the typical poecilosclerid
Diagnosis palmate chelae in having at least three clearly developed alae:
a median fluke and two flanking alae. These 'tridentate' chelae may
Myxillina basically have encrusting shape and soft consis- be further differentiated into 'arcuate' chelae (which have their
tency; some are very thinly encrusting; occasionally branching flanking alae still partly attached to the shaft without visible devel-
or of firm consistency, or bladder-like and fistular. They share a opment of further alae on the shaft) and 'anchorate' chelae (which
distinction with Microcionina in having separate ectosomal have incipient extra alae, called 'fimbriae', which also may extend
and choanosomal megascleres. a long way along the shaft). The alae of both arcuate and anchorate
chelae are normally rounded blades (called 'spatulate') but may
Description of characters be occasionally pointed, looking like predator's teeth (called
'unguiferate'). Both spatulate and unguiferate chelae often develop
Ectosomal megascieres. Ectosomal megascleres, called extra alae (called 'polydentate'). In one family species possess
'tomotes' in this suborder, are most frequently arranged as palisades probable derivations of polydentate anchorate chelae in the shape
or bouquets; their shape is basically diactinal (oxea-, strongyle- or of 'double-umbrella' microscleres (called 'birotulas'). Arcuate
tylote-like), but very frequently shape and ornamentation of both chelae may become deformed to shapes dissimilar to the original
ends are slightly different (then called 'anisotomotes'). In one fam- type (e.g., 'abyssochelae'). Chelae also often occur in two size cat-
ily their shape is so nearly a subtylostyle that these are likely to be egories, which frequently differ slightly in shape. Occasionally,
truly monactinal and their homology with other tomotes is doubtful; they may be asymmetrical, twisted, or otherwise deformed. In a
in this case the name 'tomote' is avoided. The tomote shafts are few genera chelae occur with spines or hooks on their shaft. Chelae
smooth almost without exception; their endings may be variously may occasionally be lost in species which otherwise share convinc-
sharply pointed, mucronate, blunt, swollen, microspined or bearing ing similarities with various Myxillina. One family lacks chelae
one or several larger spines. The extent to which the tomotes pene- entirely and its membership of Myxillina is tentative based on
trate into the choanosomal skeleton varies considerably and in some similarities of its tomotes with those of other Myxillina.
families they replace partly or wholly the choanosomal megascle- Other microscleres. Sigmas (shared with Mycalina, but
res. In some families the tomotes are grouped palisade-like around lacking in Microcionina) are frequent, often in two size categories,
slightly raised rounded pore-fields called areolated porefields or but lacking entirely in one family. Toxas are absent. Trichodragmas
'areolae' (in French: 'cribles'). and single raphides are occasionally found. One family has special
Choanosomal megascieres. Choanosomal megascleres are raphide-like microspined microscleres (called 'onychaetes').
basically styles (occasionally oxeas or strongyles), which may be
smooth, lightly or more heavily spined on and around the head, or Scope
entirely spined. Like in many Microcionina these are often of two cat-
egories: main and auxiliary, usually differentiated in size, ornamenta- Eleven families are presently assigned to Myxillina:
tion and location within the skeleton. Main megascleres tend to be Chondropsidae Carter, 1886: 122; Coelosphaeridae Dendy, 1922:
smooth or lightly spined, usually longer and thicker forming the basic 95; Crambeidae Levi, 1963: 16; Crellidae Dendy, 1922: 92;

515
516 Porifera· Demospongiae • Poecilosclerida • MyxiUina

A B ZONE I

ZONE
,

ZONE II
ZONE

Ul:J
II

~
ZONE III

ZONE
III

Fig. 1. A, Idealized skeletal 'bauplan' of Myxillina, consisting of three distinct zones. I, ectosomal skeleton. II, subectosomal skeleton. III, choanosomal
(or basal) skeleton. B, megasclere types characteristic for the three zones.

Dendoricellidae Hentschel, 1923: 406; Desmacididae Schmidt, taxa, concluding that there was no concordance between broad sets
1870: 52; Hymedesmiidae Topsent, 1928c: 250; Iotrochotidae of characters such as habit, skeletal architecture, surface characters
Dendy, 1922: 96; Myxillidae Dendy, 1922: 85; Phellodermidae and spicule complement. This lack of consistency has led to a pro-
fam. nov.; Tedaniidae Ridley & Dendy, 1886: 335. liferation of proposed families and genera: currently 189 nominal
genera belong to the suborder Myxillina as recognized here.
Debate continues over the validity of such characters as the precise
TAXONOMIC HISTORY nature of chelae morphology, ornamentation of the megascleres,
absence or presence of categories of megascleres and microscleres,
The suborder was only recently erected from are-evaluation plumose versus reticulate architecture, etc. It is stressed here that
of all poecilosclerid characters (Hajdu et al., 1994a). The posses- there is no single classification that has gained wide acceptance.
sion of 'tridentate' chelae (Tedaniidae excepted) and lack of toxas The classification presented here introduces changes and novelties
are pivotal independent characters that distinguish the taxonomy, to the established classifications (including the most recent sum-
forming a strong set of synapomorphies that complement the pos- maries) based on re-examination of type and other specimens, and
session of other shared non-exclusive characters (such as diactinal from a thorough survey of literature. However, this proposed
tomotes, frequent occurrence of sigmas in more than one size cate- scheme is not static, and serves as a sound objective basis for future
gory, and spined auxiliary styles). Previous attempts to classify proposals investigating non-skeletal characters such as histology,
the large numbers of Poecilosclerid genera (Topsent, 1928c; de reproduction, nucleic acid sequences and secondary metabolite
Laubenfels, 1936a; Van Soest, 1984b; Bergquist & Fromont, 1988), content.
although arriving at widely diverging schemes, emphasized skele-
tal architectural features, but disregarded, to a large extent, Basic skeleton
microsclere morphology. Only de Laubenfels (1936a) formalized
his ideas at the suprafamilial level and erected several suborders, An idealized view of basic Myxillina skeletal types and their
including the Myxilliformes. This name has not been adopted here likely development from an ancestral skeleton is presented in
because of extensive differences in content between that group and Figure 1A. This ancestral skeleton 'bauplan' is based on a combi-
the Myxillina (allowed by the ICZN Article 1; Anon., 1999). nation of skeletal structures within the order and derived from the
generalized skeleton of typical Microcionina (which is likely the
most closely related sister group on account of shared possession
REMARKS of spined auxiliary styles and differentiation of megascleres into
ectosomal and choanosomal spicules). The Myxillina skeletal
Spicule morphology versus skeletal architecture 'bauplan' is assumed to have had at the surface bouquets or
palisades of smooth diactinal tomotes (zone I in Fig. lA), grouped
Hajdu et al. (1994a) discussed the distribution of the various around the peripheral ends of one or a few long smooth styles
characters amongst poecilosclerid sponges over the established (possibly with spined heads) (zone II in Fig. 1A). These are erect
Porifera· Demospoogiae • Poecilosclerida • Myxillioa 517

\\V! \\III

E F G H I

L M o p
~l
l' W ~
u_~ li_

...... .......
WI .\\/1
] 'ttl

'. :

R T u v w x y
Fig.2. Schematic drawings of Myxillina skeletal types. A-B, ancestral 'bauplan'. C-N, skeletal types with suppressed zones II-III. O-Q, with suppressed
zones III and deemphasized zone I. R-Y, with emphasized zones III and suppressed zone II.

on the nodes of an isotropic choanosomal skeleton made up of The skeleton of the Dendoricellidae and Phellodermidae
acanthose or smooth styles (zone III in Fig. IA). The nodes of this (Fig. 2L) may not be easily derived from either the
reticulation have echinating styles of approximately the same Hymedesmiidae/Crellidae basal type, nor from the Myxillidae/
shape and size as those of the meshes. This ancestral skeleton, Coelosphaeridae/ Tedaniidae type, and we assume it underwent
more-or-less preserved in Plocamionida, is fundamentally similar a separate development. In these family there are no echinating
to skeletons found in many Microcionina (e.g., Clathria, Antho, spicules and a regularly reticulated skeleton is rare. Instead, skele-
and Raspailia). From this ancestral 'bauplan', the following devel- tons are hymedesmioid or plumose/irregularly plumoreticulate
opments led to the skeletal types observed in the various families (pointing in the direction of the former line), and chelae are similar
and genera (summarized in Fig. 2). to those of the latter line. It is necessary to stress here that skeletal
Loss of the basal isotropic reticulation led to hymedesmioid architecture is probably highly adaptive, related to form (habit) and
and plumose skeletons depicted in Figure 2C-Q; the variously size of sponges. Considerable convergence may have taken place and
stronger or suppressed development of tomote bundles gave rise to similar architecture may be found in sponges with very different
generic types predominant in the family group Hymedesmiidae spiculation. Broadly descriptive terms as 'isotropic', 'plumo-
(Fig. 2J-K) and Crellidae (Fig. 2F-I). Crellidae emphasized the reticulate' or 'hymedesmioid' do not appear to have phylogenetic
small acanthose megascleres and built them into unique ectosomal significance at higher level (e.g., families), but may serve to distin-
crusts. guish between related genera and species. Likewise, fistular or blad-
Loss of long choanosomal styles led to the proliferation of der-like growth form is considered to be an adaptive character with
reticulate skeletons found in Myxillidae and Coelosphaeridae and little significance for higher taxonomic levels, but it may serve to
Tedaniidae (Fig. 2R-Y). Again, the variously stronger or weaker distinguish between related genera and species within a family-level
development of ectosomal tomotes may be regarded to account for taxon.
loss of megasclere diversity observed in all three families.
Loss of both zone II and zone III led to skeletal types found in Megascleres
Desmacididae (Fig. 2D), Iotrochotidae and Crambeidae (Fig. 2E),
several fistular Coelosphaeridae (Fig. 2C) and many Chondropsidae Megasclere types may be related to the zones depicted in
(Fig. 2M-N). It is hypothesized that Chondropsidae lost their Figure IA. Zone I tomotes (cf. Fig. IB) vary from smooth
choanosomal spicules through replacement by foreign material as strongyles, spined or smooth tylotes to subtylostyles. Zone II
the skeletal support. megascleres are normally styles or tylostyles with smooth or more
518 Porifera· Demospongiae • Poecilosclerida • Myxillina

B c

Fig.3. Arcuate chelae and their derivatives.

frequently spined heads. Zone III megascleres are short, often those of the suborder Microcionina), and thus is considered to be
entirely spined styles, occasionally strongyles. Zone I tornotes a 'switch-onlswitch-off' character. Their significance as phyloge-
are normally easily recognized, but zone II and III megascleres netic markers at the supraspecific level is dubious but in view of the
may intergrade and become indistinguishable. Zone I megasclere often great numbers of species in the various families and genera of
types appear to be unrelated to the skeletal architecture, nor to the Myxillina it may be convenient to have a taxonomic rank to unite
Zone II and III megascleres. There also appears to be considerable similar sponges differing only in the lack or possession of these
intra-specific and supra-specific variation of tornote morphology spicule types. It is proposed to retain these groupings at the sub-
and ornamentation. Zones II and III spicules show only limited genus level, similar to solutions proposed for the Microcionina
intra-specific and supra-specific variation and have an overall genera. Similar pragmatic solutions are suggested for presence or
less diverse morphology and ornamentation. Where they are absence of other spicule types, such as chelae or sigmas.
present, they appear to be unrelated to skeletal architecture.
Sand incorporation
Microscleres
It is a relatively common phenomenon for Myxillina to incor-
Microscleres provide a most informative set of characters by porate sand grains and other detritus, partly or wholly replacing
their diversity and intricate and complicated micromorphology. native megascleres. Nevertheless, this is not exclusive to
Within Myxillina chelae may be derived from a basic arcuate type Myxillina, as it is also commonly observed in Microcionina, and
and are variously modified from this arcuate type (Fig. 3) or, outside the Poecilosclerida, such as in the haplosclerid family
by addition of alae, from an anchorate type (Fig. 4). Because of Callyspongiidae, and many dictyoceratids such as Thorectidae. It
the complicated morphology, the chelae are considered to is striking to note that by far the highest proportion of poeciloscle-
reflect phylogenetic relationships at both the family and the genus rid species with sand skeletons occur in South Australia
levels. Further microscleres encountered (Fig. 5) are less infor- (Wiedenmayer, 1989). The ecological and evolutionary implica-
mative of phylogenetic relationships, with the exception of ony- tions for this are still a major sponge conundrum. Sand skeletons
chaetes (Fig. 5J), which are a synapomorphy for Tedaniidae, and cannot be used to unite genera into families within Myxillina with-
forcipes (Fig. 5C-D), which are a synapomorphy of a genus out violating otherwise highly informative microsclere and megas-
Forcepia s.l. clere distribution patterns. Consequently, there is no clear
indication to assign sand sponges to taxa above the genus level, and
Echinating acanthostyles even at the genus level this feature is a doubtful synapomorphy.
The family Chondropsidae contains the majority of the sand gen-
Presence or absence of echinating acanthostyles occurs in era, but arenaceous species and genera also occur in most of the
many otherwise not-closely related families and genera (including other families (Crellidae excepted).
Porifera. Demospongiae • Poecilosclerida • Myxillina 519

K
Fig. 4. Anchorate chelae and their derivatives.

A c D E G

J K L
H
Fig.S. Micro c1eres (other than chelae) occurring in various Myxillina group.
520 Porifera· Demospongiae • Poecilosclerida • Myxillina

MYXILLINA FAMILY-LEVEL CHARACTERISTICS hymedesmioid growth form and architecture is thought to grade
into anchinoid architecture; spiculation is largely similar in all
Eleven families are tentatively recognized in Myxillina, but they members of these two nominal families. In support of this hypoth-
still need further corroboration. Most Myxillina have arcuate chelae, esis is the existence of a similar continuum from microcionid to
but three families (MyxiUidae, Desmacididae and Crambeidae) have anchinoid-like architecture in some Microcionidae (e.g., Clathria
exclusively anchorate chelae. Such restriction has resulted in the - (Thalysias) phorbasiformis Hooper, 1996a), suggesting that these
possibly artificial - separation of genera previously united within skeletal structures pertain to grades of construction rather than
a single family Myxillidae (e.g., Lissodendoryx and Myxilla). So far phylogenetic clades.
only a single exception to this chela-type distribution has been Crellidae is closely related, sharing the areolate porefields
observed: the genus Crellomima, which is a typical representative of with many Hymedesmiidae, but having the tangential crust of
Crellidae in most respects, but has (polydentate) anchorate chelae. acanthose megascleres as a clearly distinctive trait.
This casts some doubt on the validity of using the anchorate condition Chondropsidae remain problematic because of their reduced
of the chelae as a synapomorphy for genera and families (as was also nature; they are united in having a single megasclere category in
noted by Hooper (1996a) for a few Microcionidae). the form of thin smooth strongyles or styles.
Birotulas are considered to be homologous for all genera that Dendoricellidae has been revived based on the combination of
possess them, and they are probably derived from anchorate chelae; arcuate chelae as microscleres (often with sigmas) and oxeas
genera bearing them are consequently assigned to a single family (tomote derivatives) as the only megascleres present; architecture
lotrochotidae, a sister family to the Myxillidae and Desmacididae. is plumose.
Crambeidae is exceptional in having subtylostyle-like ectoso- Desmacididae has been restricted to encompass sponges with
mal megascleres; they are included in Myxillina on the basis of a similar spicule complement to that of Dendoricellidae but in
their anchorate chelae. these chelae are anchorate and skeletal architecture is reticulate.
Fistular or bladder-like growth forms are considered to be an A new family Phelloderrnidae is proposed to accommodate
adaptive, non-phylogenetic character and consequently Coelos- species possessing arcuate chelae and exclusively styles for
phaeridae sensu Topsent (1928c) is united with non-fistular genera megascleres.
with reticulate skeletons into Coelosphaeridae s.l. Tedaniidae is exceptional in lacking microscleres other than
Anchinoidae Topsent (1928c: 284) is here merged with onychaetes, and lacking auxiliary acanthostyles; its assignment to
Hymedesmiidae as a consequence of discussions in the literature: Myxillina is based on similarities in tomote shape.

KEY TO FAMILIES

(1) Microscleres exclusively onychaetes (very thin spined or rugose unequal-ended spicules) (Fig. 5J) ................................ Tedaniidae
Microscleres include birotulas (double-umbrella microscleres) (Fig. 41) ...................................................................... Iotrochotidae
No onychaetes or birotulas .................................................................................................................................................................. 2
(2) Megascleres exclusively of one type, either oxeas, styles or strongyles ............................................................................................. 3
Megascleres of diverse types (may include oxeas in combination with other types) or entirely absent ............................................. 5
(3) Megascleres oxeas only ....................................................................................................................................................................... 4
Megascleres thin strongyles only (occasionally thin styles only); chelae arcuate or absent (Fig. 3) .......................... Chondropsidae
Megascleres robust styles only; chelae arcuate or derivates thereof (Fig. 3) .............................................................. PheUodermidae
(4) Skeleton reticulate, microscleres anchorate chelae (Fig. 4) ........................................................................................... Desmacididae
Skeleton plumose or irregular, microscleres arcuate chelae (Fig. 3) .......................................................................... Dendoricellidae
(5) Megascleres consisting of peripheral thinner subtylostyles and choanosomal thicker styles; chelae exclusively
anchorate (Fig. 4A, B, E, G) ........................................................................................................................................... Crambeidae*
Megascleres a combination of diactinal and monactinal spicules, or entirely absent ......................................................................... 6
(6) Ectosomal skeleton consisting of vertical brushes of megascleres fanning out and becoming tangential or
megascleres entirely absent ................................................................................................................................................................. 7
Ectosomal skeleton a crust of entirely spined oxeas or styles ................................................................................................ Crellidae
(7) Chelae anchorate or polydentate (Fig. 4) ............................................................................................................................ Myxillidae
Chelae arcuate or absent (Fig. 3) ......................................................................................................................................................... 8
(8) Choanosomal skeleton plumose or hymedesmioid; ectosome with areolated porefields .......................................... Hymedesmiidae
Choanosomal skeleton reticulate or absent (in fistular forms); no areolated pore fields .......................................... Coelosphaeridae
Choanosomal skeleton a reticulation of sand grains, neither megascleres nor microscleres ......................................... Chondropsidae
* Several species in this family show considerable infraspecific variability in spicule presence and shape. Individuals showing reduced and modified
spiculation may key out in other families.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Chondropsidae Carter, 1886

Rob W.M. Van Soest

Zoological Museum, University of Amsterdam, P.O. Box 94766,1090 GT Amsterdam, Netherlands. (soest@science.uva.nl)

Chondropsidae Carter (Demospongiae, Poecilosclerida) is revived to encompass Myxillina genera, which possess only a single thin
megasclere type, usually strongyles or strongylote spicules, in plumose arrangement and arcuate chelae as microscleres. The latter may
sometimes be secondarily lost or replaced by sigmas, which occur in some species. Sand may replace partly or wholly the bundles of
megascleres in several genera and species. Surface usually reticulate or provided with poresieves. Five genera are considered valid, among
which Phoriospongia is revived to accomodate sand sponges with a reduced skeleton of styles instead of the usual strongyles.
Keywords: Porifera; Demospongiae; Poecilosclerida; Myxillina; Chondropsidae; Batzella; Chondropsis; Phoriospongia; Psammoclema;
Strongylacidon.
- - - - - - - - - - - - - - - - -

DEFINITION, DIAGNOSIS, SCOPE Taxonomic remarks

Synonymy Tomotes. The strongyles are presumably of tomote deriva-

tion; many are thin and have often slightly asymmetrical ends
Chondropsina Carter, 1886a: 122; Phoriospongiinae and may occasionally be stylote. The styles of one genus
Lendenfeld, 1889a; Psammascidae de Laubenfels, 1936a; (Phoriospongia) show slightly swollen tyles, indicating possible
Stylotellinae sensu Wiedenmayer, 1989 (not Lendenfeld, 1888). homology with choanosomal principal megascleres.
Sand. Many members of this family have their skeleton
Definition partly or wholly replaced by sand grains.
Microscleres. Based on the presence or absence of
Myxillina with only thin smooth strongyles or occasionally microscleres, there is a series of intergrading 'genera': Batzella (no
thin smooth styles as megascleres, in an irregular plumose arrange- microscleres), Phoriospongia (sigmas), Strongylacidon (unguifer-
ment; no acanthostyles; microscleres arcuate or unguiferate chelae ate chelae + sigmas), Burtonispongia (arcuate chelae + sigmas).
and/or sigmas. It is here proposed to ignore the presence or absence of sigmas
and/or chelae at the genus level. The entire absence of microscleres
Diagnosis (Psammoclema, Batzella) makes allocation to family arguable, and
consequently these genera are retained for possible allocation to
Encrusting, massive, flabellate or digitate growth forms. other groups, even though the distinction among them is probably
Ectosomal skeleton frequently absent, replaced by arenaceous artificial. Batzella and Chondropsis differ in the presence or
or spicular detritus, but often with areolate porefields on sur- absence of a sand reticulation, which may also be an artificial
face. Ectosomal smooth strongyles or occasionally styles, often rare distinction.
or secondarily lost. Choanosomal spicules, if present, are Affiliations. This is a small somewhat suspect, dustbin-
auxiliary megascleres of ectosomal origin, strongyles or occasionally type group. Possibly various genera could be reduced members
styles, whereas principal spicules appear to be absent. Microscleres of Hymedesmiidae.
are arcuate isochelae and sigmas, but these may be often absent.

Scope Previous reviews

Sixteen genera are assigned to the family, of which five are con- Topsent (1928c), Wiedenmayer (1989 as Stylotellinae).
sidered valid: Batzella, Chondropsis, Phoriospongia, Psammoclema,
Strongylacidon.

KEY TO GENERA

(1) Megascleres present ............................................................................................................................................................................ 2

No proper megascleres, the skeleton is made up of foreign material ............................................................................ Psammoclema
(2) Megascleres styles ........................................................................................................................................................ Phoriospongia
Megascleres strongyles ........................................................................................................................................................................ 3
(3) Chelae and/or other microscleres present ........................................................................................................................................... 4
No microscleres ................................................................................................................................................................................... 5
(4) Sand grains and foreign debris dominating the fibres ...................................................................................................... Chondropsis
No sand or foreign debris ............................................................................................................................................. Strongylacidon

521
522 Porifera· Demospongiae • Poecilosclerida • Myxillina • Chondropsidae

(5) Sand grains and foreign debris dominating the fibres ...................................................................................................... Chondropsis
No sand or foreign debris ......................................................................................................................................................... Batzella

BATZELIA TOPSENT, 1893 CHONDROPSIS CARTER, 1886

Synonymy Synonymy

Batzella Topsent, 1893c: 34. Chondropsis Carter, 1886a: 122. Sigmatella Lendenfe1d,
1888: 198; Collosclerophora Dendy, 1917b: 320; Burtonispongia
Type species de Laubenfels, 1936a: 52.

Halichondria inops Topsent, 1891a: 533 (by original Type species

designation).
Chondropsis arenifera Carter, 1886a (monotypy).
Definition
Definition
Chondropsidae with strongyles only, no microscleres, no sand.
Chondropsidae with skeletal columns cored by strongyles and
Diagnosis foreign material (sand, broken spicules); sigmas and/or arcuate or
unguiferate chelae may be present.
Thinly encrusting to massive, with simple choanosomal skeleton
of columns of strongyles, without a proper ectosomal skeleton. No Diagnosis
further spicules. Half a dozen species from different parts of the world.
Sand sponges with feebly developed spicular skeletons con-
Previous review sisting of strongyles. Very small sigmas may be found in a few
species, and one species also has reduced arcuate chelae.
Van Soest et al., 1996: 91.
Previous review
Description of type species
Wiedenmayer (1989: 71, as Phoriospongia in part).
Batzella inops (Topsent, 1891a) (Fig. 1A).
Synonymy. Halichondria inops Topsent, 1891a: 533, Remarks
pI. XXII fig. 1; Topsent, 1893c: xxxiv; Batzella inops; Topsent,
1894b: 38. The genus is revived despite Wiedenmayer's recent syn-
Material examined. Holotype (slide): MNHN-LBIM DT onymization of it with Phoriospongia, because the type of the
2109 - fie Verte, Roscoff, France, microscopic slide labeled 'type'. latter genus has exclusively styles, many of which are clearly mon-
Description. According to the original description, it is actinal, clearly different from the strongyles of the present genus.
thinly encrusting, smooth and soft. Colour yellow. No other particu- The alleged variability of megasclere endings is not present in the
lars are known. Skeleton. The spicule bundles in the microscopic type species of Chondropsis.
slide of the type are vague, not visibly bound by spongin, and they
follow a wispy course. There are no obvious bouquets at the surface Description of type species
(but these were reported by Topsent), and an ectosomal cover of
spicules is not apparent. Spicules strongyles only, 170-200 X Chondropsis arenifera Carter, 1886a (Fig. lB-C).
2-3 jJ.m, they are isodiametric, in general equally shaped at both Synonymy. Chondropsis arenifera Carter, 1886a: 122;
apices, but a few may be found with one end somewhat mucronate. Phoriospongia arenifera; Wiedenmayer, 1989: 71.
There is very little variation in length and thickness and all belong Material examined. Lectotype (here designated): BMNH
obviously to a single category. Most have a wide axial canal. 1886.12.15.149 - Port Philip Heads, SE Australia (one of Carter's
Reproduction. August-September. Its larvae have been described as syntypes).
big, yellow and having a bare posterior pole (Topsent, 1894b), Description. This is a massively rounded sponge, with
which is characteristic of Poecilosclerida. Distribution and ecology. smooth or slightly conulose surface with a fine reticulate pattern.
Known with certainty only from Roscoff, but recorded from various Numerous irregular oscules. Sandy and soft. Colour buff or reddish
localities in the Mediterranean, under stones in the mid-littoral. grey. Skeleton (Fig. lB). Predominantly a reticulation of rather
Remarks. Several of the later records of this species are sus- coarse sand-tracts glued together by spongin, at the periphery
pect, because they quote different colours and more than one type of mixed with wispy tracts of thin strongyles. Size of meshes
spicule (including styles). Reduced specimens of Crambe crambe ca. 400-800 jJ.m, thickness of tracts ca 80-250 jJ.ID. At the surface the
may be easily mistaken for Batzella. Other records of the genus need skeletal meshes are further subdivided into smaller meshes by bun-
critical re-examination. Reliably assigned species appear Caribbean dles of strongyles. Spicules (Fig. 1C). Thin flexuous strongyles
B. rosea Van Soest (1984b) and Australian B. jrutex Pulitzer-Finali 160-190 X 1-2 jJ.ID (Carter gives size 280 X 3 jJ.m, but no matching
(1981). spicules were found in the examined type; Carter also mentions thin
Porifera • Demospongiae • Poecilosclerida • Myxillina • Chondropsidae 523

..-

1-----l1250

A
I
50
B

Fig. 1. A, Batzella inops Topsent, 1891a (as Halichondria), drawing of skeleton and spicules made from a slide of the holotype. B-C, Chondropsis arenifera Carter
(1886a). B, drawing of skeleton and spicule made from a slide of the holotype. C, SEM photo of spicule made from the holotype (scale I /Lm). D, Chondropsis
arenacea (Dendy, 1917b, as Collosclerophora) , drawing of spicules and 'colloscleres' reproduced from Dendy pI. II fig. 8 (size see text). E, Chondropsis australis
(Dendy, 1896 as Desmacidon), type of Bunonispongia de Laubenfels (1936a), drawing of skeleton and spicules made from a slide of the holotype.

styles, but these are likely to be the same spicule type as the dominant sausage-shaped inclusions called 'colloscleres', the nature of
strongyles). Distribution and ecology. South Australia, 36m depth. which is uncertain. No phylogenetic significance of these inclu-
Remarks. According to Wiedenmayer (1989) C. arenifera is sions is apparent. Hooper & Wiedenmayer (1994) considered this
a junior synonym of Phoriospongia argentea (Marshall, 1880, as to be a junior synonym of Batzella, but in view of the possession of
Dysidea). No confirmation of this can be offered; Marshall's sand tracts it is here referred to Phoriospongia.
description makes no reference to any proper spicules, and The genus Sigmatella Lendenfeld, 1888 was erected for type
Wiedenmayer did not provide subsequent evidence to support his species Sigmatella macropsamma Lendenfeld (1888: 198) (by sub-
assignment. sequent designation, de Laubenfels, 1936a: 99), type 2MB 1206
The genus Collosclerophora Dendy (1917b: 320, pI. II (not examined). The type species designation is remarkable, as
figs 1-16) was erected (by monotypy) for South Australian de Laubenfels chose the one species as type which: (1) did not have
Collosclerophora arenacea Dendy (1917b), type not examined, not the sigmas mentioned in Lendenfeld's definition; (2) was described
present in BMNH. This is a typically massive sand sponge with only very cursorily; and (3) was not treated in Lendenfeld's
reduced spiculation of strongyles only (Fig. 10),220 x 2.5/Lm, in monograph (1 889a), in which Sigmatella was newly erected (prob-
combination with a system of columns of sand-filled tracts. ably because the 1888 publication was not yet published when
The genus was distinguished for its possession of gelatinous he wrote the manuscript). In both publications, Sigmatella australis
524 Porifera· Demospongiae • PoecUosclerida • MyxUlina • Chondropsidae

was described as the first species, which moreover possesses the sig- Synonymy. Phoriospongia solida Marshall, 1880: 122,
mas, and would have been the logical choice as type. Nevertheless, de pI. vm figs 12-17.
Laubenfels' type species designation must be accepted. Lendenfeld's Material examined. None. Whereabouts of the two type
description mentions a flabelliform sponge, size 8 X 7 X 1.7 cm, specimens from Tasmania, collected by Haeckel, is unknown
with radial longitudinal sharp ridges on the surface, scattered (according to Hooper & Wiedenmayer, 1994: 174).
oscules. Fibres of the skeleton are composed of sand grains and very Description (from MarshaU, 1880). Conical sponges
abundant strongyles, up to 260 X 3 fJ.m (called 'microsclera'). (Fig. 2A), covered by a slimy skin, through which larger and
The genus Burtonispongia de Laubenfels, 1936a: 52 was smaller foreign debris is visible. A few oscules of 0.5-2 mm diam-
erected (by original designation) for type species Desmacidon eter. Skeleton. In cross section there is only a mass of sand and
australis Dendy, 1896 (slides of type specimens BMNH shell debris subdivided by an irregular system of canals. No fibres
1902.10.18.268,270 were examined). This is a grey massive irregu- or surface reticulation present. The megascleres do not appear to be
lar sponge with numerous vents and sandy texture. Ectosome arranged in tracts. Spicules. Thin smooth stylote and subtylostylote
(Fig. IE) with characteristic pattern made by choanosomal sand megascleres, some of which have a distinct terminal or subterminal
columns reaching the surface between the end of which is a reticu- tyle, 200-300 fJ.m; sigmas of c- and s-shape, no size quoted.
lation of spicule bundles. The choanosomal skeleton (Fig. IE) con- Remarks. The drawing of the sty10te spicules in Marshall's
sists largely of stout sand columns, up to 900 fJ.m in diameter, lying pI. vm fig. 16 (here reproduced) is such, that these are unlikely to rep-
at distances of 200-1500 fJ.m, but at the surface there are distinct resent modi-fied strongyles. For that reason, Wiedenmayer's proposal
bundles of spicules 20-40 fJ.m in diameter containing up to 20 for synonymy of Phoriospongia and Chondropsis is here refuted, and
spicules in cross section, ending in brushes of spicules forming a both are considered valid genera. A second species of Phoriospongia,
subdermal reticulation with meshes of 50 fJ.m diameter. Microscleres p. reticulum, was described by Marshall (1880) likewise from
scattered in the ectosome. Megascleres thin strongyles (Fig. IE) of Tasmania. It differs primarily in the polygonal ornamentation of the
160-193 X 2-3 fJ.m. Microscleres thin, reduced isochelae (Fig. IE) surface.
of ca. 10-17 ILm and thin sigmas (Fig. IE) of 20-30 fJ.m. The defini- The genus Stylotrichophora Dendy, 1895 was erected
tion of Chondropsis is extended to include species with chelae (by monotypy) for type species Stylotrichophora rubra Dendy
because of the similarities of this species to other Chondropsis (1895: 259). Three slides of the holotype, BMNH 1902.10.18.324,
species. were studied (Fig. 2B). It has stylote spicules in a large size, range
up to 270 X 4 fJ.m, the shorter of which tend to be concentrated in
the peripheral region. There are also smooth sinuous thin raphide-
PHORJOSPONGIA MARSHALL, 1880 like spicules. The skeletal columns of 250 fJ.m in diameter con-
sist predominantly of sand. This is a clear junior synonym of
Synonymy Hircinia jlabellopalmata Carter (1885c: 313), type species of
Tedaniopsamma Burton, 1934b: 599 (by original designation).
Phoriospongia Marshall, 1880: 122. Stylotrichophora Dendy, The type specimen of Hircinia jlabellopalmata, BMNH
1895: 259; Tedaniopsamma Burton, 1934b: 199. ? Kaneohea de 1886.12.15.240, from Port Philip Heads, was re-examined. It has
Laubenfels, 1950b: 23. a flabellate digitate growth form; ectosomal skeleton arenaceous;
choanosomal skeleton composed of a reticulate skeleton with well
Type species developed fibres cored by and axial column of sand and smooth
styles, 165-240 X 2--4 fLm; auxiliary spicules are smooth long
Phoriospongia solida Marshall, 1880: 122 (by subsequent raphides, 100-200 X 0.5 fJ.m. Large numbers of fine toxas noticed
designation; de Laubenfels, 1936a: 99). in the sections are considered foreign.
The genus Kaneohea de Laubenfels, 1950b: 23 was erected for
Definition type species (by monotypy) Kaneohea poni de Laubenfels, 1950b:
23, fig. 14 from Hawaii. Holotype (not examined) USNM 22751.
Chondropsidae with skeletal columns cored by styles and This is a thin purple encrustation with a skeleton of foreign material
foreign material (sand, broken spicules); sigmas usually present. making an isodictyal reticulation. No microscleres are reported, but
proper megascleres are given as strongyles of 90 X 5 fJ.m, styles of
Diagnosis 120 X 4-5 fJ.m, and oxeas 120 X 5 fJ.m; the described raphides of
90 X 0.5 fJ.m might very easily be juvenile megascleres. The assign-
Sandy, variable in amount and distribution of foreign detritus. ment of this sponge to Phoriospongia as a junior synonym is tenta-
Megascleres mostly frail subtylostyles or styles, with frequent tive, based on the assumption that the oxeas are not proper to the
transitions. Spicules scattered or in vaguely radial to plumose sponge. There is also a possibility that this is Desmapsamma with
arrangement. Microscleres in the form of small sigmas occur in rare or absent microscleres, but that is mere speculation.
a few species. About a dozen species.

Previous review PSAMMOCLEMA MARSHALL, 1880

Wiedenmayer (1989: 71). Synonymy

Description of type species Psammoclema Marshall, 1880: 109. Psammascus Marshall,

1880: 93. Psammopemma Marshall, 1880: 113. Sarcocornea
Phoriospongia solida Marshall, 1880 (Fig. 2A). Carter, 1885b: 214.
Porifera· Demospongiae • Poecilosclerida • Myxillina • Chondropsidae 525

A B I
00

Fig. 2. A, Phoriospongia solida Marshall, 1880, drawing of shape and spicules reproduced from Marshall's pI. VIII figs 12, 16-17 (sizes see text),
B, Phoriospongiajlabellopalmata (Carter, 1885a as Hircinia), senior synonym of Stylotrichophora rubra Dendy (1896) and type of Tedaniopsamma
Burton (1934b), drawing of skeleton and spicules made from a slide of the holotype of Stylotrichophora rubra Dendy, C, Psammoclema ramosum
Marshall (1880), photos of shape and skeleton of lectotype reproduced from Wiedenmayer (1989, pI. 9 fig. 4, pI. 28 figs 3-4) (scale, see text).
D, Psammoclema decipiens (Marshall, 1880 as Psammascus), drawing of shape and skeleton reproduced from Marshall's pI. VI figs 1,3,5 (scale, see text).
E, Psammoclema nodosum (Carter, l885a as Sarcocomea), photo ofBMNH holotype, scale 19 X 8 X 13 em, copied from Wiedenmayer (1989, pI. 8 fig. 11).

Type species Previous review

Psammoclema ramosum Marshall, 1880: 109 (by monotypy), Wiedenmayer (1989).

Definition Remarks

Chondropsidae without proper spicules, just sand. Through its lack of spicules members of this genus have been
mistaken for keratose sponges associated with Dysidea or Hyrtios,
Diagnosis The firm sandy consistency, the lack of conules and the irregular
sand columns of Psammoclema readily distinguish it from Dysidea
Ramose and massive growth forms; ectosomal skeleton arena- and Hyrtios. De Laubenfels (1936a: 59) suggested the genus is ajun-
ceous, detachable; choanosomal skeleton dendritic tracts composed ior synonym of Dactylia Carter (1885), but this is clearly incorrect as
only of sand grains united by spongin, or amorphously packed with that is a member of Cal1yspongiidae. Also, even if both would have
sand; no megascleres or microscleres. Half a dozen species. been synonymous, Psammoclema would have been the senior name.
526 Porifera· Demospongiae • Poecilosclerida • Myxillina • Chondropsidae

c
')

E F
Fig. 3. A-C, Strongylacidon zanzibarense Lendenfeld, 1897c. A, drawing of choanosome and spicules reproduced from Lendenfeld's pI. X figs 106-116
(sizes, see text). B-C, SEM photos of spicules from the holotype. B, strongyle. C, chela (scale I !Lm). D-E, Strongylacidon osburnensis (George & Wilson,
1919 as Phoriospongia), type species of Xytopsues de Laubenfels (l936a). D, photo of type specimen, reproduced from George & Wilson's pI. LXI fig. 24
(size, see text). E, drawing of spicUles reproduced from George & Wilson's pI. LXVI fig. 60 (sizes, see text). F, Strongylacidon kaneohe (de Laubenfels,
1950b as Xytopsiphum), drawing of spicUles reproduced from de Laubenfels' fig. 5 (sizes, see text).

Description of type species surface. Oscules distributed over one side of the branches, flush,
1-2 mm in diameter. Consistency firm, compressible. Colour dark
Psammoclema ramosum Marshall, 1880 (Fig. 2C). purple or reddish grey. Skeleton of the periphery is a dense palisade
Synonymy. Psammoclema ramosum Marshall, 1880: 109, of broken spicules and sand grains forming a tight-meshed surface
pI. VII figs 12-15, pI. VIII figs 1-5; Polt!jaeff, 1884: 43, pI. III fig. 8, skeleton, with large subdermal lacunae underneath. The choanoso-
pI. IV fig. I; Wiedenmayer, 1989: 80, pI. 9 figs 3--6, pI. 28 figs 3-4. mal skeleton is an irregularly anastomosing system of sand
Chalinopsilla arborea var. ramosum; Lendenfeld, 1889a: 152. columns, which may be up to 1000 /-Lm or more in diameter, run-
Materilll examined. Lectotype (not examined): PMJ 134 ning the length of the branches in the centre and forming an axial
(designated by Wiedenmayer, 1989: 80). Paralectotypes (not core, with dichotomously branching columns directed towards the
examined): 2MB 2632 (see Hooper & Wiedenmayer, 1994: 175). periphery. Spicules absent. Distribution. Bass Strait, S Australia.
Specimens of Polejaeff: BMNH 1885.8.8.12-14 - Bass Strait, Remarks. Although spicules are absent, the general struc-
'Challenger' Expedition. stat. 162,38 fms. ture of the skeleton resembles that of Chondropsis and
Description. The lectotype (stated to be in the Jena Museum, Phoriospongia, and family membership of Psammoclema appears
see Fig. 2C) as well as the specimen collected by the 'Challenger' certain.
expedition from the same area as the type, are irregularly branching The genus Psammascus Marshall, 1880 was erected (by
sponges, branches 8-10 mm in diameter, with smooth velvety monotypy) for type species Psammascus decipiens Marshall
Porifera' Demospongiae • Poecilosclerida • Myxillina • Chondropsidae 527

(1880: 93, pI. VI figs 1-5, here reproduced as Fig. 2D), and if Diagnosis
indeed congeneric with Psammoclema would have page priority.
However, Wiedenmayer (1989: 75) was unable to find the type Encrusting to massive; ectosomal skeleton with protruding
and proposed to use Psammoclema as the senior name for these brushes of stongyles from the ascending choanosomal fibres;
sponges, because the holotype of its type species is still extant. The choanosomal skeleton plumose in encrusting specimens becoming
type specimen of Psammascus decipiens is described to have plumoreticulate in more massive specimens; fibres cored by
a tubular shape, soft consistency and a discrete fibre network. strongyles and occasionally some detritus; microscleres are
The genus Psammopemma Marshall, 1880: 113 was erected unguiferous-arcuate isochelae and/or sigmas. About 15 species.
for Psammopemma densum Marshall, 1880: 113, pI. VIII figs 6-11
from Tasmania (by monotypy). The holotype (not examined) is in Previous reviews
2MB according to Wiedenmayer (1989: 77). This is now a flat-
tened mass of about 1.5 x 0.8 cm, probably a fragment of the orig- Van Soest (l984b), Wiedenmayer (1989).
inal specimen of 3 X 3.5 cm. Subsequently identified specimens
may be repent-ramose and up to 9 X 5 X 4 cm (Wiedenmayer, Description of type species
1989). Marshall depicts the specimen with a group of oscules on
the upper surface. The specimen consists mostly of sand and shell Strongylacidon zanzibarense Lendenfeld, 1897c (Figs 3A-B).
fragments and/or spicule debris, with some cementing spongin. Synonymy. Strongylacidon zanzibarense Lendenfeld,
Marshall claims generic status for his species because of the appar- 1897c: 110, pI. X figs 106-116.
ent lack of foreign material in the surface membrane. In all other Materiol examined. Holotype: SMF 1570 - Zanzibar.
aspects the type conforms to Psammoclema, and the variability of Fragment from type: BMNH 1908.9.24.150; slide BMNH
the external sand coat in these and similar sponges is such that 1897.3.25.73.
generic distinctness is unwarranted. Wiedenmayer's (1989) deci- Description. Encrusting on the carapace of a crab, 5-8 mrn
sion to consider Psammopemma a synonym of Psammoclema thick, lateral expansion 2 cm. Surface irregular, possibly with small
is here endorsed. Psammopemma has been associated with pore-sieves ('Hockern'). Grey-brown in alcohol. There is algal
Dysideidae by some authors (e.g., Hentschel, 1929: 912), but there thallus in most of the sponge body. Skeleton, a reticulation of
is little reason for this. spongin-enforced tracts of megascleres, about 50 fLm in diameter,
The genus Sarcocomea Carter, 1885b was erected (by mono- and many loose spicules. The tracts rise up from a spongin plate
typy) for type species South Australian Sarcocomea nodosa Carter overlying the algal tissue. Spicules, strongyles, straight, cylindri-
(1885b: 214). The type specimen, BMNH 1886.12.15.295, was cal, 160-260 X 3-4 fLm; unguiferate chelae, with three teeth some-
reexamined. It is a large crest-like mass, 19 X 8 X 13 cm, lobate, what dissimilar in length on opposite sides ('anisochelae').
with oscules on the rim, hard in dry condition (Fig. 2E). Surface Distribution and ecology. Zanzibar, shallow-water.
smooth. The interior structure conforms closely to P. ramosum but Remarks. The genus Xytopsues de Laubenfels (l936a: 55)
is coarser and has more spongin in the choanosomal columns, was erected by original designation for type species Phoriospongia
which may anastomose into complex fenestrated sheets. Bergquist osbumensis George & Wilson (1919: 154, pI. LXI fig. 24,
(1980b) assigned this genus to Dysidea, but that appears incorrect pI. LXVI fig. 60, here reproduced as Fig. 3C-D) (=junior syn-
in view of the similarity to Psammoclema ramosum. onym of Desmacidon grise us Schmidt, 1870). A slide of the type of
Schmidt, BMNH 1870.5.3.37, 'Florida 80', as well as de
Laubenfels' 1950 specimen, USNM 23424, Bermuda, labeled as
STRONGYLACIDON LENDENFELD, 1897 Xytopsues griseus, were re-examined. These are fist- or head sized,
or thin crusts, soft, slimy, with tuberculate surface; USNM 23424
Synonymy is filled with the strands of an alga of the genus Jania, whereas
George & Wilson's material contained sand grains in the fibres.
Strongylacidon Lendenfeld, 1897c: 110. Xytopsues de Spicules strongyles, 150-180 X 1-3 fLm, unguiferous tridentate
Laubenfels, 1936a: 55. Xytopsiphum de Laubenfels, 1950b: 12. chelae, 12-18 fLm, and sigmas 10-30 fLm. The extent to which sand
is incorporated determines whether this should be assigned to
Type species Strongylacidon or to Chondropsis.
The genus Xytopsiphum de Laubenfels, 1950b was erected
Strongylacidon zanzibarense Lendenfeld, 1897c: 110 (by (monotypy) for Hawaiian type species Xytopsiphum kaneohe de
monotypy). Laubenfels (1950b: 12, fig. 5, here reproduced as Fig. 3E). The
holotype USNM 22742 was re-examined. This is a thinly encrust-
Definition ing, dark-coloured, smooth sponge. The skeleton shows spicule
tracts with up to 150 spicules in cross section, and scattered
Chondropsidae with strongyles and arcuate, often unguiferate tangential spicule at the surface. Spicules are thin strongyles,
chelae (may be absent), occasionally sigmas (may be absent); sand 200-210 X 3-4 fLm and reduced unguiferate chelae, 10-15 fLm.
may be present in low quantities but is not dominating the fibres. It is very similar to the type species of Strongylacidon.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademicIPlenum Publishers, New York, 2002

Family CoeJospbaeridae Dendy, 1922

Rob W.M. Van Soest

Zoological Museum, University of Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, Netherlands. (soest@science.uva.nl)

Coelosphaeridae Dendy (Demospongiae, Poecilosclerida), originally intended to receive all Poecilosclerida with fistules, parchment-like
surface membrane and hollow bladder-like shape, is here rearranged to unite Myxillina possessing arcuate chelae, ectosomal smooth
tomotes and if present a reticulate choanosomal skeleton. Fistular shape is considered only significant at the genus level. Genera and
species with fistular shape but possessing birotulas palmate chelae andlor toxas previously assigned to Coelosphaeridae are here excluded
from the family and reassigned to Iotrochotidae (genus Amphiasterella), Microcionina (fistular Acamidae) and Mycalina (isodictyid genus
Coelocarteria). In the newly revised family seven genera are considered valid, but several are further subdivided into convenient subgenera.
Keywords: Porifera; Demospongiae; Poecilosclerida; Myxillina; Coelosphaeridae; Chaetodoryx; Coelosphaera; Forcepia;
Histodermella; Inflatella; Lepidosphaera; Lissodendoryx.

DEFINITION, DIAGNOSIS, SCOPE the other hand, several genera traditionally assigned to
Coelosphaeridae because they possess fistular habit, but otherwise
Synonymy dissimilar in spiculation, are reassigned to disparate families:
Amphiasterella to Iotrochotidae because of its possession of birotu-
Coelosphaereae Dendy, 1922b: 95. Coelosphaeridae las, Coelocarteria to Isodictyidae because of its possession of
Hentschel, 1923: 406. palmate chelae in combination with strongyloxeas, genera possess-
ing microspined tylotes in combination with palmate isochelae
Definition andlor toxas (Cornulum s.s., see Van Soest et al., 1994) to the
Microcionine family Acamidae. Several of the genera of the newly
Myxillina with reticulate choanosomal skeleton and arcuate defined Coelosphaeridae, e.g., Lissodendoryx and Forcepia have
isochelae. large numbers of species and wide geographic and ecological
amplitude. Others are rare deep-water forms (Histodermella). Two
Diagnosis genera (Inflatella, Lepidosphaera) are assigned to Coelosphaeridae
provisionally, because affinities cannot be determined for their
Fistular-hollow, branching, massive or encrusting sponges. lack of arcuate chelae.
Surface smooth in fistular forms. Non-fistular representatives have
their surface often irregularly pitted and punctate, but areolated Taxonomic remarks
pore-fields are absent. Skeleton reticulate, in fistular forms fre-
quently vestigial. Ectosomal tomotes diactinal (often tylote). Skeletal architecture. Coelosphaeridae are similar to myxil-
Choanosomal megascleres smooth or acanthose styles, occasion- lid sponges in the possession of a skeletal architecture of reticulate
ally oxeas or strongyles. In fistular forms choanosomal megascle- tracts forming an isodictyal skeleton, with the usual brushes of
res may be lost. Next to arcuate isochelae, microscleres include tomotes assuming a partially tangential position. Skeletal zone II
sigmas and raphides. (see chapter on suborder Myxillina) is usually not developed. In the
bladder-like fistular forms the choanosomal skeleton is absent or
Scope reduced to a few wispy bundles traversing the interior, and the ecto-
somal skeleton becomes strictly tangential. In one case (Forcepia
39 nominal genera of which seven are valid: Chaetodoryx, subgenus Leptolabis), the choanosomal skeleton is hymedesmioid.
Coelosphaera, Forcepia, Histodermella, Inflatella, Lepidosphaera, Both types of reduced skeletons may be derived from the basic
Lissodendoryx; and in three of which (Coelosphaera, Forcepia and reticulate structure by suppression and reduction, of zone II and ill
Lissodendoryx) subgeneric units are recognized. in bladder-shapes, and of zone ill in hymedesmioid sponges.
Echinating acanthostyles. Like in other families of
History and biology Poecilosclerida sponges with very similar skeletons and spicules
may differ only in the presence or absence of echinating acan-
Originally (Dendy, 1922b; Hentschel, 1923; Topsent, 1928c) thostyles. Although the repetitive occurrence indicates low phy-
this family was restricted to hollow, bladder-like, spherical, club- logenetic significance of presence or absence of acanthostyles,
shaped, tubular, and cushion-shaped growth forms. However, such it is considered practical to distinguish these variants at the
growth forms are not exclusively occurring in the Myxillina and do subgenus level in the larger genera (Lissodendoryx, Forcepia,
not constitute reliable characters for family recognition. Non- Coelosphaera).
fistular sponges with spiculation very similar to those of fistular Previous reviews. Dendy (1922b), Hentschel (1923).
sponges are indicative of a larger group comprising this family Topsent (1928c), Levi (1973), Van Soest (1984b), Bergquist &
including genera as Lissodendoryx s.l. and Forcepia s.l. On Fromont (1988); Hajdu et al. (1994a).

528
Porifera· Demospongiae • Poecilosclerida • Myxillina • Coelospbaeridae 529

KEY TO GENERA

(1) No microscleres, just one type of megascleres ...................................................................................................................... Inflatelfa

Microscleres (chelae, sigmas, microxeas, trichodragmas, forceps, or ecailles) present ...................................................................... 2
(2) Microscleres are peculiarly disc-shaped ('ecailles') situated at the surface ................................................................. Lepidosphaera
No disc-shaped ectosomal spicules ..................................................................................................................................................... 3
(3) Microscleres include forceps (labis) ....................................................................................................................................... Forcepia
No forceps .................................. ......................................................................................................................................................... 4
(4) Spiculation includes entirely spined oxeas .......................................................................................................................................... 5
No entirely spined oxeas ...................................................................................................................................................................... 6
(5) Spined oxeas are oxychaetes (microxeas finely spined all over) ..................................................................................... Chaetodoryx
Spined oxeas are megasclere-sized with coarse hook-like spines .................................................................................. Histodermelfa
(6) Sponge hollow, bladder-like with leathery or parchment-like skin; normally with longer or shorter
tubular fistules .................................................................................................................................................................. Coelosphaera
Sponge massive, lobate or branching, but not hollow, bladder-like .............................................................................. Lissodendoryx

CHAETODORYX TOPSENT, 1927 Colour dark purple in life, white in alcohol. Ectosomal megascleres
irregularly arranged in the surface membrane, but erected in bundles
Synonymy around the oscules. Choanosomal skeleton (Fig. lB) a reticulation
of acanthostyles, with smaller echinating acanthostyles at the nodes.
Chaetodoryx Topsent, 1927b: 11. Coelectys Topsent, 1936: 12. Oxychaetes and chelae distributed throughout the interior. Spicules
(Fig. lB), ectosomal anisotylotes, smooth, straight, with elongated
Type species but slightly different swollen ends, often polytylote, 155-175 X
3 jJ.m; larger acanthostyles, slightly curved, lightly spined all-over,
Chaetodoryx richardi Topsent, 1927b: II (by monotypy). 175-195 X 10-12jJ.m; smaller acanthostyles, short and fat, spined
all-over, 6-8jJ.m. Microscleres, arcuate isochelae, curved, relatively
Definition elongated, 55-59jJ.m; long, thin, uniformly spined oxeas (oxy-
chaetes), 225-245 X 2-3jJ.m. Distribution and ecology. Boavista,
Coelosphaeridae with smooth ectosomal tylotes, choanoso- Cape Verde Islands, on shells at 91 m.
mal reticulation of styles, echinated by acanthostyles; microscleres Remarks. Although dissimilar in body shape and surface
include oxychaetes. characters, the type species (by monotypy) of Coelectys Topsent,
1936, C. insinuans Topsent (1936: 12, figs 2-3) from Monaco
Diagnosis (30-60 m) has the same spiculation as C. richardi. The holotype,
MNHN D.T. 58, was reexamined. It is a fistulose specimen
Encrusting, massive or fistular-insinuating sponges. Surface (Fig. 1C) overgrowing a Hyrtios. Colour white or pale greenish
smooth or furrowed. Ectosomal skeleton of tylotes or anisotylotes. yellow. Spicules include smooth tylotes with elongate heads
Choanosomal skeleton reticulate, made up of spined styles, with (Fig. 1D), 225--400 X 3-5.5jJ.m, acanthostyles (Fig. lE), 75-160 X
echinating acanthostyles. Microscleres include arcuate isochelae 3jJ.m; microscleres include arcuate isochelae (Fig. IH), differenti-
and rugose oxychaetes. ated into smaller normal isochelae, about 20 jJ.m, and elongate ones
with pointed teeth of 40 jJ.m, and finely spined curved microxeas
Scope (Fig. IF), 100-110 X ljJ.m. Large thin sigmas (Fig. IG) were
observed in the type specimen, but not recorded by Topsent. The
Two species, both from the Mediterranean-Atlantic area. genus Histodermella (cf. below) also has spined oxeas, but these
are unlike the oxychaetes of Chaetodoryx. By their size and posi-
Previous review tion these are considered as choanosomal megascleres. The thin
acanthose oxychaetes of Chaetodoryx mimic tedaniid onychaetes,
Topsent (1928c). but these are stylote, whereas the present microscleres are clearly
oxeote.
Description of type species

Chaetodoryx richardi Topsent, 1927b (Fig. lA-B). COELOSPHAERA THOMSON, 1873

Synonymy. Chaetodoryx richardi Topsent, 1927b: 11;
Topsent, 1928c: 242, pI. IT fig. 8, pI. VITI figs 8-9. Synonymy
Material examined. Ho1otype (slide): MNHN D.T. 1118 -
labeled 'Tedanectyon richardi n.g. n.sp., St. 1203, 1901'. Coelosphaera Thomson, 1873: 484. [Histodermal Carter,
Description (largely from Topsent, 1928c). Massively I 874a: 220 (preocc.). Histioderma Carter, 1886c: 452. [Sideroderma]
encrusting (Fig. IA), 2.7 cm long, 0.3 cm high, convex at one side Ridley & Dendy, 1886: 348 (preocc.). Siderodermella Dendy,
but with a deep furrow on the other; it is attached to a shell. The sur- 1922b: 105. Histodermion Topsent, 1927b: 9. Xytopsoocha de
face of the convex side bears numerous aquiferous apertures impos- Laubenfels, 1936a: 55. Naauna de Laubenfels, 1950a: 78.
sible to differentiate into oscules and pores. The consistency is firm. Coelosphaericon Bakus, 1966a: 33.
530 Porifera· Demospongiae • Poecilosclerida • Myxillina • Coelospbaeridae

Fig. 1. Chaetodoryx. A-B, C. richardi Topsent (1927b), habitus of holotype and spicules (scales: A, I cm, B, sizes see text) reproduced from Topsent
(1928e pi. II fig. 8 and pi. VIII figs 8). C-H, C. ;ns;nuans Topsen! (1936 as Coelectys). C, holotype fragments (scale I em). D-H, SEM photos of spicules
made from the holotype (scale 10 J.Lm, except F, I J.Lm).

Type species tangential crust; choanosomal skeleton consists of poorly devel-

oped tracts and scattered smooth spicules; megascleres are smooth
Coelosphaera tubifexThomson, 1873: 484 (by monotypy). tylotes or strongyles, of one size only, to which acanthostyles may
be added; microscleres are arcuate isochelae, sigmas and raphides
Definition (often in trichodragmata), some of which may be absent. The
genus Histiodennion is here included as a subgeneric unit
Coelosphaeridae with hollow bodies provided with fistules; of Coelosphaera since it differs only from species of the latter in
megascleres tylotes or strongyles, to which echinating acan- the possession of echinating acanthostyles. Coelosphaera s.l. has
thostyles may be added; microscleres arcuate isochelae, sigmas a cosmopolitan distribution. About 30-35 species have so far
and raphides (may be absent). been described.

Diagnosis
Previous reviews
Massive, bladder-like, or encrusting, burrowing growth forms
with erect fistules; smooth ectosomal tylotes form a compact Van Soest (1984b), Bergquist & Fromont (1988).

Key to subgenera of Coelosphaera

(I) Spiculation includes acanthostyles ................................................................................................................................. Histiodermion

No acanthosty1es ............................................................................................................................................................. Coelosphaera
Porifera' Demospongiae • Poecilosclerida • Myxillina • CoeJospbaeridae 531

Fig. 2. Coelosphaera. A, C. (Coelosphaera) tubifex Thomson (1873), drawing of type specimen reproduced from Thomson (1873).
B-F, C. (Coelosphaera) navicelligera (Ridley, 1884a as Sideroderma), SEM photos of spicules from the holotype (scales: B, lOG !Lm, C and D left,
10 !Lm, D right and E-F, l!Lm). G, C. (Coelosphaera) peltata (Topsent, 1904b as Desmacidon), type of Naauna de Laubenfels (1950a), chelae reproduced
from Topsent (l904b) (size 50 !Lm). H-K, C. (Coelosphaera) hatschi (Bakus, 1966a as Coelosphaericon). H, habitus of BMNH paratype (scale I cm).
I-K, SEM photos of spicules from BMNH paratype (scales: B, 100 !Lm, C and D left, 10 !Lm, D right and E-F, l!Lm). L, C. (Histodermion) dividuum
(Topsent, 1927b), drawing of spicules reproduced from Topsent (l928c: pI. VITI fig . I,sizes see text).

SUBGENUS COELOSPHAERA THOMSON, 1873 Porcupine stat. 24, North Atlantic, one of Carter's syntypes' .
Paratype: BMNH 1882: 7:28:38 - North Atlantic, 'Porcupine'stn.
Type species 2 (syntype of Histodenna appendiculata).
DescriptWn. Subglobular (Fig. 2A) with several long nar-
Coelosphaera tubifex Thomson, 1873: 484 (by monotypy). row tubular fistules of different lengths, some of which are open
and presumably function as oscules. Inside virtually hollow. Size
Definition not recorded. Surface smooth. Consistency tough, parchment-like.
Colour light grey. Skeleton a dense tangential crust of intercrossing
Coelosphaera without echinating acanthostyles. megascleres at the surface; choanosome without spicule tracts
or fibres, just a pulpy mass of organic material, containing the
Description of type species canal system and loose spicules. Spicules smooth tylotes of
520 X 6/Lm, long smooth stylotes resembling the tylotes but
Coelosphaera (Coelosphaera) tubifexThomson, 1873 (Fig. 2A). with one end tylote and the other bluntly stylote, up to 900 X
Synonymy. Coelosphaera tubifex Thomson, 1873: 484; ? 14/Lm; arcuate isochelae of about 25-30/Lm. Distribution
Carter, 1876: 472. Histodenna appendiculatum Carter, 1874a: 220, and ecology. North Atlantic and Arctic, deep sea, 200--1500 m
pI. XIV figs 23-25, pI. XV fig. 39; Carter, 1876: 472; depth.
Coelosphaera appendiculata; Dendy, 1922b: 102. Remarks. This species has been ignored in the literature,
Material examined. Neotype (proposed herein): BMNH whereas a very similar species, C. appendiculata (Carter, 1874a
1882.7.28.27 - labeled 'Histodenna appendiculata Carter, as Histodenna) is recorded several times. It is likely that both are
532 Porifera' Demospongiae • Poecilosclerida • Myxillina • CoeJospbaeridae

synonyms. Dendy (1922b) suggested that the type specimens SUBGENUS HISTODERMION TOPSENT, 1927
of Histoderma appendiculatum (BMNH 1882.7.28.27, 38) from
H.M.S. 'Porcupine' Expedition stat. 24, are the same material as Type species
that described as Coelosphaera tubi/ex by Thomson, but the proof
of that is now wanting. Carter himself (1876) hesitatingly admitted Histodermion dividuum Topsent, 1927b: 9 (by monotypy).
that the two could be synonymous. In the absence of Thomson's
material, the type of Carter registered as BMNH 1882.7.28.27 is Definition
here proposed as a neotype for Thomson's species. A second North
Atlantic species is C. physa (Schmidt, 1875 as Desmacidon) differ- Coelosphaera with echinating acanthostyles.
ing from C. tubi/ex in lacking sigmas and possessing trichodrag-
mas instead. Carter (1886c: 452) realized his name [Histoderma] Description of type species
was preoccupied by a fossil annelid, and replaced it by
Histioderma, with the same type species, and thus it becomes a Coelosphaera (Histodermion) dividuum Topsent, 1927b
junior synonym of Coelosphaera. (Fig.2L).
[Sideroderma] Ridley & Dendy (1886: 348) was erected (by Synonymy. Histodermion dividuum Topsent, 1927b: 9;
original designation) for type species Crella navicelligera Ridley Topsent, 1928c: 225, pI. VIII fig. 1.
(1884a: 571). Similar to Histoderma, the name [Sideroderma] Material examined. Holotype: MOM (not seen). Slide
Carter was found to be preoccupied by Sideroderma Lendenfeld of holotype: MNHN D.T. 1170 - labeled 'n.g.n.sp. st. 1420,
(1883) for a different sponge now considered a Polymastia, 1902'.
and Dendy (1 922b: 105) proposed the replacement name Description (largely from Topsent, 1928c). Encrusting on
Siderodermella. The New Guinean sponge S. navicelligerum blackish gorgonian branches, surface smooth, glistening, with
(Fig. 2B-F) is a pulpy mass with long hollow fistules. Megascleres cylindrical fistules of several mm high, rarely perforated at the end
are exclusively tylotes (Fig. 2B), 280--595 X 6-12 fJ-m. Micro- (presumably oscular). Skin tears off easily. Inside mostly hollow,
scleres are two sizes of sigmas (Fig. 2D), 60 fJ-m and 19 fJ-m, but some organic material is gathered at the base. Consistency
raphides (Fig. 2C) in wispy trichodragmas of 450 fJ-m long, arcuate fragile and soft. Colour yellowish or white (alcohol). Ectosomal
isochelae (Fig. 2E) of about 20 fJ-m and tiny 'navicelliform' thick- parchment of tangential tylotes. Thin, scarce, choanosomal
shafted chelae (Fig. 2F) of about 10 fJ-m. tracts of tylotes, 3-4 spicules in diameter, traverse the vast hollow
The genus Xytopsoocha de Laubenfels (1936a: 55) was pro- spaces. At the base, single, long acanthostyles are erect on the
posed for the curious North Atlantic deep water species Gellius substrate or are lying loose. Microscleres concentrated at the
macrosigma Topsent, 1890b: 65 (by original designation, a slide of surface. Spicules (Fig. 2L), tylotes, occasionally strongylote,
the type was reexamined, but not illustrated here). This is a thin swollen in the middle part, quite variable in size, 425-740 X
yellow encrustation with tylotes of 700 X 10 fJ-m as megascleres, 8-15/-lm; acanthostyles, entirely spined, long, 450-470 X 13-
and two size categories of sigmas, the largest of which is enor- 16 fJ-m, arcuate isochelae, comparatively elongated, with well-
mous, 415 X 8 fJ-m, the smaller 30 fJ-m. Trichodragmas of 15-65 fJ-m developed teeth, quite variable in size but not clearly divisible,
are present. The giant sigmas can hardly be considered of generic 25-50 fJ-m. Distribution and ecology. Azores, deep water, 2460m
importance. Membership of Coelosphaeridae and synonymy with depth.
Coelosphaera is tentative, based entirely on the assumption that the Remarks. Coelosphaera (Histodermion) cryosi (Boury-
chelae have become lost secondarily. Esnault et al., 1994b) is a closely related species differing in
The genus Naauna was erected (by original designation) by having much smaller acanthostyles and possessing sigmas; the
de Laubenfels (1950a: 78) for type species Desmacidon peltatus chelae occur in two size categories.
Topsent (1904b: 204), from deep water (600m depth) off the
Azores. A slide of the type, MNHN DT. 1014, was examined. The
type specimen, kept in the Monaco Museum, was apparently dam-
aged when collected. Its skeleton is described as having a fragile FORCEPIA CARTER, 1874
skin and pulpy interior. The only megascleres are strongyles of
500--530 X 11-13 /-lm; microscleres are peculiar palmate-like Synonymy
isochelae (Fig. 2G) of 50 fJ-m here interpreted as arcuate, and
trichodragmas of 130 /-lm. Forcepia Carter, 1874a: 248. Forcipina Vosmaer, 1885a: 26;
The genus Coelosphaericon Bakus (1966a) was erected for Forcepina Vosmaer, 1887. Leptolabis Topsent, 1901b: 353.
the North Pacific species C. hatschi Bakus (1966a: 33). A paratype Trachyforcepia Topsent, 1904b: 181. Clinolabis Topsent, 1927b.
specimen, BMNH 1965.6.171.1 (Fig. 2H-K) was re-examined. Wilsa de Laubenfels, 1930: 27. Labisophlita de Laubenfels, 1936a:
This has the shape (Fig. 2H) and structure of Coelosphaera, but 120. Marcusoldia de Laubenfels, 1936a: 87. Tedandoryx de
in addition to the usual spicule complement of tylotes (Fig. 21), Laubenfels, 1954: 130. Ectoforcepia Cabioch, 1968b: 232.
525-1010 X 12-22/-lm, and arcuate chelae (Fig. 2K), 30--42 /-lm, it
has toxiform microxeas (Fig. 2J). Since proper toxas are not found Type species
in any other myxilline sponge it is assumed to be a unique feature
of a single species. This is supported by the undulating irregular Forcepia colonensis Carter, 1874a: 248 (by monotypy).
outline of these spicules (Fig. 2J, at right). In other respects it
conforms closely to Coelosphaera. If more species with similar Definition
spiculation will be found, Coelosphaericon may be revived as
subgenus of Coelosphaera. Coelosphaeridae with forceps microscleres.
Porifera· Demospongiae • Poecilosclerida • Myxillina • Coelospbaeridae 533

Diagnosis (Carter, 1886c) for this South Australian species. This is also in
accordance with Wiedenmayer's (1989) proposals concerning this
Encrusting to massive growth forms; ectosomal skeleton com- species. As a neotype needs to be from the same general area to
posed of tangential smooth ectosomal tylotes; choanosomal skele- avoid possible confusion with species from other areas, it is neces-
ton composed of choanosomal styles or ectosomal tylotes forming sary to find a neotype among material described from the
a hymesmioid structure in encrusting species or reticulate architec- Caribbean. The Caribbean currently is known to have four species
ture in massive forms; microscleres are smooth or spined forceps of Forcepia: viz., F. trilabis (Boury-Esnault, 1973), F. grandisigmata
(labis), arcuate isochelae, sigmas. Occurrence cosmopolitan. About Van Soest (1984b), F. vermicola Lehnert & Van Soest, 1996 and
40 species. an unnamed Forcepia from the Caribbean (e.g., Colombia and
Cura.. ao; Zea, Van Soest unpublished data). Of these, F. trilabis
Previous reviews sensu Van Soest (1984b) shows considerable similarity with
F. colonensis in possessing unusually large forceps (up to 258 /-Lm
Carter (1874a), Dendy (1895), Van Soest (1984b). versus 260/-Lm of Carter's description), whereas the other three
species have forceps less than 160/-Lm long. Brazilian F. trilabis
Taxonomic remarks sensu Boury-Esnault, 1973, shows a minor difference, viz., an
extra category of smooth forceps, and possibly concerns a species
The original material of Forcepia colonensis came from 'an different from the Caribbean material. It is judged likely that
arenaceous deposit in the neighbourhood of Colon or Aspinwall, Caribbean F. trilabis, and more in particular, the specimen
Panama' collected by Mr F. Kitton of Norwich. No traces of it have described from deep water off Barbados by Van Soest (1984b) con-
been found in the collections of the Natural History Museum, forms to F. colonensis. It is proposed here to adopt that specimen as
London, during a search by Ms Clare Valentine. The description of the neotype of F. colonensis. This action also settles the status of
the type was based on characteristic spicules (large forceps) in the the genus EctoJorcepia Cabioch (1968b), erected for Forcepia
fossil sample. In the declared absence of type material, it is desir- species lacking choanosomal styles. Since the neotype does not
able to assign a neotype if stability of a species or genus is threat- possess the choanosomal styles (nor were these mentioned in the
ened. In the light of the fact that several rivalling name synonyms description of Carter), F. colonensis would belong to both Forcepia
(cf. above) are in existence, it is urgent to establish the characters and EctoJorcepia, thus making the latter a junior synonym (cf. also
of the type of Forcepia (see Article 75 of the ICZN code; Anon., below). Previously, sponges possessing forceps microscleres in
1999). However, due to the paucity of characters in the original combination with ectosomal smooth tylotes, were scattered over
description this is not a straightforward matter. All material labeled several families (Myxillidae, Hymedesmiidae and Tedaniidae)
as 'Forcepia colonensis' in the BMNH collection is from the South because of disparate skeletal structure and variability of
Australian Bracebridge Wilson collection, described subsequently forceps shape. However, the myxilline forceps with its rugose or
by Carter (1885a) and Dendy (1896) under this name (Ms Clare spined surface is a unique spicule type, here interpreted as a
Valentine in litteris). For obvious geographic reasons alone, it is synapomorphy for a single genus of sponges. In view of the fact
highly unlikely that this material conforms to the type from that hymedesmioid structure coincides with the possession of
Panama. Moreover, the size of the forceps of the South Australian echinating acanthostyles in several species bearing forceps, it is
material is less half that of the Caribbean specimen. Consequently proposed to recognize a separate subgenus Forcepia (Leptolabis)
this material does not satisfy the qualifying conditions for assign- for these, differentiated from the nominal subgenus Forcepia
ing a neotype. The specimens Dendy classed as F. colonensis (Forcepia) which are reticulate. Future subdivisions may be possi-
included Carter's (1886c) Suberites biceps and accordingly ble on the diverse shapes and sizes of the forceps, but for the time
it is proposed to use the combination Forcepia biceps being this is not pursued.

Key to subgenera of Forcepia

(1) Spiculation includes basal acanthostyles in a hymedesmioid arrangement ......................................................................... Leptolabis

No basal acanthostyles; if there are spined styles, then they are the structural megascleres making the choanosomal
reticulation ............................................................................................................................................................................... Forcepia

SUBGENUS FORCEPIA CARTER, 1874 Description of type species

Synonymy Forcepia colonensis Carter, 1874a (Fig. 3A-C).

Synonymy. Forcepia colonensis Carter, 1874a: 248, pI. XV
Forcepia Carter, 1874a: 248. Forcipina Vosmaer, 1885a: 26 fig. 47. Forcepia trilabis; Van Soest, 1984b: 66, pI. VI figs 1-2,
(Forcepina Vosmaer, 1887= ? lapsus). TrachyJorcepia Topsent, text-fig. 24 (not Boury-Esnault, 1973: 280, fig. 32).
1904b: 181. Wilsa de Laubenfels, 1930. Labisophlita de Material examined. Neotype: ZMA POR. 4564 - Barbados,
Laubenfels, 1936a: 120. Tedandoryx de Laubenfels, 1954: 130. 0.5 mile off Holetown, 100m, coIl. P. Wagenaar Hummelinck,
EctoJorcepia Cabioch, 1968b: 232. #1442 (originally described as F. trilabis), including four slides.
Description (from Van Soest, 1984b). A thin crust on a con-
Definition glomerate of coral rubble and worm tubes. Size about 1-2cm2 .
Surface smooth, no apparent oscules. Consistency soft. Colour. grey-
Forcepia with reticulate skeleton and lacking echinating ish brown in alcohol. Skeleton. Ectosomal skeleton a crust of inter-
acanthostyles. crossing megascleres; choanosomal skeleton mostly an irregular
534 Porifera· Demospongiae • Poecilosclerida • Myxillina • Coelospbaeridae

Fig. 3. Forcepia. A-C, F. (Forcepia) colonensis Carter (1874a), SEM photos of spicules from the neotype ZMA POR. 4564 (scales: 10 /Lm, except detail
of B right, I /Lm). D-H, F. (Forcepia) jorcipis (Bowerbank, 1866), type of Forcipina Vosmaer (1885a), SEM photos of spicules of one of Bowerbank's
paratypes, BMNH 1882:7:28:30 (scales: 10 /LID, except E, 100 /LID).

arangement of single spicules. Spicules. Tylotes (Fig. 3A) smooth, Carter's description of Forcepia as valid. Later (1887) he expressed
with well-developed tyles, rather uniform in length, 330-360 X doubts (and changed the name into Forcepina for no apparent rea-
4-7 fLm; large arcuate isochelae (Fig. 3C) with reduced alae, son; this name is either a lapsus or a nomen nudum). One of
20-38 fLm; small normal-shaped arcuate isochelae (Fig. 3C), Bowerbank's types, BMNH 1882.7.28.30 from Shetland, was reex-
15-20 fLm; large forceps (Fig. 3B), spined all over, 200-260 X amined (Fig. 3D-H). It is a massive sponge with irregular surface.
3.5-4.5 fLm (Carter gives 260 fLm). Distribution and ecology. The skeleton is an irregular reticulation of vague tracts (2-6
Panama, Barbados, on coral rubble bottom, at 100 m depth. spicules), forming an aniostropic skeleton. Spicules are smooth
Remarks. Van Soest recorded also a single smaller acan- tylotes (Fig. 3D), 231-302 X 4-6 fLm; smooth styles (Fig. 3E),
those forceps, but subsequent examination of the slides and SEM 495-572 X 9-21 fLm; arcuate chelae (Fig. 3H) in two size categories,
preparation failed to confirm its presence. Presumably Van Soest 38-65 fLm and 22-30 fLm; forcepses (Fig. 3F-G) occur in three cate-
(1984b) mistook a broken thin growth stage of the large forceps for gories, all acanthose, 351-432 X 3-4 fLm, 75-91 X 1 fLm and 20-33 X
it. Forcepia trilabis Boury-Esnault (1973: 280) appears quite close 0.5 fLm, the smaller ones (Fig. 3G) have one longer, curved leg.
to F. colonensis in its present interpretation, but differs in having a The genus Trachyforcepia Topsent (1904b: 181) was erected
smooth category of large forceps, as well as a definite smaller cate- for type species Forcepia groenlandica Fristedt, 1887: 453 (by
gory of acanthose forceps. It is likely to be a closely related but original designation), which has spined choanosomal styles instead
separate species. of the usual smooth ones. A slide of the holotype was found in the
The genus Forcipina Vosmaer (1885a: 26, pI. V figs 60--66), Natural History Museum, London, BMNH 1910.1.1.1322, labeled
with type Halichondriaforcipis Bowerbank, 1866: 244 (by original 'from Herr Fristedt, Gronland, Forcepia groenlandica, Norman
designation), was erected by Vosmaer because he could not accept colI.' The skeleton at the surface is a tangential crust of tylotes.
Porifera· Demospongiae • Poecilosclerida • Myxillina • Coelospbaeridae 535

A
n

Fig. 4. Forcepia. A, Forcepia (Forcepia) groenlandica Fristedt (1887), type of Trachyforcepia Topsent (l904b), drawing of skeleton and spicules made
from a holotype slide in BMNH. B, Forcepia (Forcepia)forcipula Lundbeck (1905 as Esperiopsis), type of Labisophlita, drawing of shape and spicules
reproduced from Lundbeck's pI. I fig. 5 and pI. VIII fig. 5 (for sizes see text). E-G, Forcepia (Forcepia) lissa (de Laubenfels, 1954 as Tedandoryx), SEM
photos of spicules made from the holotype (scale 10 /-Lm). H, Forcepia (Forcepia) carteri Dendy (1896), drawing of skeleton and spicules made from a
BMNH type slide.

Choanosomal skeleton (Fig. 4A) an isotropic reticulation of 1-3 lacking (no tylotes are mentioned and the chelae are described as
spicules per side. No spongin, no echinating acanthostyles. palmate). Recently, (Lee, 2002) this species was redescribed from
Spicules (Fig. 4A). Ectosomal smooth tylotes, slightly anisotylote original material and an additional fresh specimen, and assigned to
but with tyles well-developed, about 400 f.1m; choanosomal acan- Forcepia (Forcepia) on account of its possession of forceps and
thostyles, lightly spined all over, robust, straight, 500-550 f.1m; arcuate chelae.
small arcuate isochelae (Fristedt says they are anchorate, but in the The genus Labisophlita de Laubenfels (1936a: 120) was
type slide they are clearly arcuate) about 35 f.1m; large, somewhat erected (original designation) for type species Esperiopsis forcipula
flagelliform sigmas, 100-150 f.1m; small finely acanthose forceps, Lundbeck (1905: 17, pI. I fig. 5, pI. VIII fig. 5). The BMNH collec-
with one leg longer and flagelliform, about 50 f.1m. Lehnert & Van tion holds a topotypical specimen, BMNH 1910.1.1.644 from
Soest (1996) employed the name Trachyforcepia as a sub-genus Davis Strait, 160-180m, Norman collection, which was examined.
for Forcepia species with echinating acanthostyles, but these The label suggests this material was sent/donated by Lundbeck.
are absent in F. groenlandica. For Forcepia species with It shows a tangential ectosomal skeleton of smooth styles, up
hymedesmioid architecture, like the species described by Lehnert & to 680 f.1m and rare thin smooth tylotomotes up to 470 f.1m, carried
Van Soest, the subgenus Leptolabis is available (cf. below). by choanosomal spicule tracts (Fig. 4B). The tylotomotes were
The genus Wilsa de Laubenfels, 1930: 27 was erected for type not mentioned by Lundbeck and that was enough reason for
species Wilsa hymena de Laubenfels, 1930: 27, more extensively de Laubenfels (1936a: 120) to erect a new genus. Microscleres
described in de Laubenfels (1932: 72, fig. 39). The description of include two size categories of arcuate isochelae, about 45 and
this sponge is ambiguous, with major Forcepia features apparently 10 f.1m (Lundbeck says the smaller are palmate, but under high
536 Porifera· Demospongiae • Poecilosc1erida • Myxillina • Coelospbaeridae

magnification they appear to be arcuate), two size categories of sig- Description of type species
mas, 30 and 85 /-Lm, the smaller of which is sharply pointed, and
two size categories of small forceps, up to 20/-Lm (Lundbeck Forcepia (Leptolabis) luciensis (Topsent, 1888) (Fig. 5A-E).
mentions only one). Synonymy. Dendoryx luciensis Topsent, 1888: xxxvii.
The genus Tedandoryx de Laubenfels, 1954 was erected for Leptosia exilis Topsent, 1892b. xxii. Leptoiabis luciensis; Topsent,
type species Tedandoryx lissa de Laubenfels (1954: 130). The 1901b.353;Topsent, 1928c.56
type specimen, USNM 22906, was re-examined. Its ectosomal Material examined. Holotype (slides): MNHN D.T. 2560,
skeleton is a tangential crust of tylotes and forcipes, carried by ill- 2561 - Normandy, W France, labeled 'Luc s Mer, 12-IX-92'.
defined bundles of megascleres consisting of a mixture of tylotes Other material. ZMA POR. 6621: Mauritania Exped. stat. 27-9, off
and (subtylo-)styles. Single styles are erect at the base of the Banc d'Arguin, 16m depth.
sponge. Spicules (Fig. 4C-G). Ectosomal tylotes (Fig. 4C), with Description. Thinly encrusting with upright papillae.
prominent elongated heads, 243-(265.5)-285 X 2-(3.8)-5 /-Lm; Colour yellow. The skeleton shows a rather irregular arrangement
subtylostyles (Fig. 4D) with elongated swollen heads, 270- of bundles of megascleres with at the base of the sponge single
(290.6)-303 X 6-7/-Lm; chelae in two categories, large thick small acanthostyles. Spicules (Fig. 5A-E), tylotes (Fig. 5A), with
strongly curved arcuate chelae (Fig. 4F), with only two alae in well-developed heads, 230-310 X 2-5 /-Lm; acanthostyles (Fig.
stead of the usual three, 23-(25.1)-27/-Lm, and small arcuate 5B), 65-100 X 4.5 /-Lm; no other megascleres; microscleres include
chelae (Fig. 4G) of normal form: 10-(1O.8)-12/-Lm; the forcepses two size categories of well-developed arcuate isochelae (Fig. 5E),
(Fig. 4E) of 120 X 1-1.5/-Lm are straight and thin, and often bro- 12-18 and 33-35 /-Lm, normal small forceps (Fig. 5C) and peculiar
ken at the joint. De Laubenfels did not recognize the paired legs of small forceps of which one leg may be strongly incurled (Fig. 5D),
the forcepses and considered them as separate onychaete-like 5-10 /-Lm. Distribution. W coast of France, S coast of England,
spicules. This explains why he assigned this genus to Tedaniidae Mediterranean, Mauritania, Azores.
close to Tedania. Remarks. Topsent (1928c) described also sigmas of 35-90
The genus Ectoforcepia Cabioch (1968b: 232) was erected /-Lm in Mediterranean specimens, but these are absent in the type
for the group of Forcepia species without choanosomal styles, and in the specimens from the Atlantic (see Topsent, 1928c: 185).
apparently having replaced these by the ectosomal tylotes. The genus Clinolabis Topsent, 1927b was erected (by mono-
However, as demonstrated above, the type of Forcepia also does typy) for C. dentifera Topsent (1927b: 14) from deep water
not possess the styles, so it conforms to Ectoforcepia, making it a (2460 m) off the Azores (Fig. 5F). A spicule slide of the type was
junior synonym. Type species was named the South Australian examined, MNHN D.T. 1162, labeled "n.g.n.sp. st. 1420, 1902",
Forcepia carteri Dendy (1896: 25). This is a greyish-yellow or showing a complement of smooth polytylote tylotes, 300-455 X
ochre-yellOW massive, irregular sponge chiefly composed of 6-7/-Lm, heavily spined acanthostyles in two size categories,
sand. A slide of the type, BMNH 1902.10.18.286 - RN 607, was 320-370 X 20-24/-Lm and 140-160 X 14-15 /-Lm, unguiferate-
examined. The ectosome is a sand crust of up to 1 rom in thick- arcuate isochelae, 30-46/-Lm, large sigmas, 95-175 /-Lm, and acan-
ness. The skeleton (Fig. 4H) contains next to sand also a vague those forceps in two size categories, 65-75 and 23-28 /-Lm. Topsent
reticulation of spicule bundles which become clear plumose stressed the fact that the chelae were 'isancres' but they never have
tracts of up to 20 spicules in cross section, 50-150 /-Lm in diame- more than three teeth and show no similarity to anchorate chelae.
ter, near the surface. Megascleres (Fig. 4H) are anisotylotes and In all other aspects this species conforms to Leptolabis.
strongyles of 228-(257.4)-303 X 3-(3.7)-4.5 /-Lm. Micro- The genus Marcusoldia de Laubenfels (1936a: 87) was
sc1eres (Fig. 4H) consist of isochelae with reduced alae: erected (by original designation) for type Leptolabis tenuissima
12-(16.6)-22/-Lm (possibly in two size categories of ca. 12-15 Hentschel (1911: 362, fig. 41, here reproduced as Fig. 5G),
and 18-22/-Lm) and thin equal-legged rugose forceps many of because it would have palmate isochelae in combination with for-
which are broken at the sharp angled junction of the the two legs: ceps. Although no material was examined, de Laubenfels was
63-(79.9)-91 /-Lm. clearly in error. According to Hentschel's description, it is a thinly
encrusting sponge with basal acanthostyles, in at least two size
categories, 64-144/-Lm, ectosomal strongyles-tylotes, 120-168 X
2-3/-Lm, a single category of arcuate chelae, 12-15/-Lm (the draw-
SUBGENUS LEPTOIABIS TOPSENT, 1901 ing and the text both confirm the arcuate nature of the chelae), two
sizes of sigmas, 27--60/-Lm and 8/-Lm, and are small and relatively
Synonymy heavily spined forcepses, 30-38 X 1-2/-Lm.

Leptolabis Topsent, 1901b: 353; Clinolabis Topsent, 1927b:

14; Marcusoldia de Laubenfels, 1936a: 87. HISTODERMELIA LUNDBECK, 1910

Type species Synonymy

Dendoryx luciensis Topsent, 1888: xxxvii (by subsequent Histodermella Lundbeck, 1910: 14. Hiltonus de Laubenfels,
designation; Topsent, 1928c: 56). 1936a.

Definition Type species

Forcepia with hymedesmioid skeleton, with basal Histodermella ingolfi Lundbeck, 1910: 14 (by subsequent
acanthostyles erect on the substrate. designation; de Laubenfels, 1936a: 72).
Porifera· Demospongiae • Poecilosclerida • MyxilIina • Coelosphaeridae 537

Fig. 5. Forcepia (Leptolabis). A-E, Forcepia (Leptolabis) luciensis (Topsent, 1888 as Dendoryx), SEM photos of spicules made from ZMA POR 6621
(from Mauritania) (scales: 10 /-Lm, except C-D, l/-Lm). F, Forcepia (Leptolabis) dentifera (Topsent, 1927b as Clinolabis), drawing of spicules reproduced
from Topsent (I92Sc, pI. IX fig. 25) (sizes see text). G, Forcepia (Leptolabis) tenuissima (Hentschel, 1911), type of Marcusoldia de Laubenfels (1936a),
drawing of spicUles reproduced from Hentschel, 1911: fig. 41 (sizes see text).

Definition Description (from Lundbeck, 1910). Hollow, bladder-like

main body (Fig. 6A) with long thin fistules, which may be branched.
Coelosphaeridae with heavily spined acanthoxeas as Surface smooth. Oscules at the end of some of the fistules, others are
choanosomal megascleres. closed and are presumably inhalant. Consistency leathery. Skeleton
of the ectosome a feltwork of tangentially placed megascleres, of the
Diagnosis choanosome virtually absent, consisting of a few irregular tracts.
Spicules, tylotes (Fig. 6B), often somewhat swollen in the middle,
Spherical base with erect fistules; ectosomal skeleton consists 290--650 X 7-17 f.Lm; heavily spined acanthoxeas (Fig. 6C),
of smooth ectosomal tylotes (or strongyles) forming a thick tan- 170--210 X 4-14 f.Lm; sigmas (Fig. 6D) thin, 56--61 f.Lm; arcuate
gential layer; choanosomal skeleton consists of irregular tracts of chelae (Fig. 6E), occasionally somewhat deformed, 21-28 f.Lm.
scattered tylotes (or strongyles), and acanthoxeas or acan- Distribution and ecology. Arctic, deep water, 1438 m depth.
thostrongyles form a layer at right angles to the ectosomal tylotes Remarks. A second species of Histodermella is New
and are scattered throughout the choanosome; microscleres Zealand H. australis Dendy (1924: 373, pI. XV figs 24-27),
are arcuate isochelae and sigmas, sometimes also raphides in redescribed from additional material by Bergquist & Fromont
trichodragmata. (1988: 49, pI. 19 figs B-F).
De Laubenfels (1936a: 72) erected a genus Hiltonus for this
Previous reviews species on the alleged presence of acanthostyles instead of acanthox-
eas. This was based on the drawing of a blunt-ending acanthoxea next
Lundbeck (1910), Bergquist & Fromont (1988). to a normal acanthoxea in Dendy's illustration. The type specimen of
H. australis, BMNH 1923.10.1.155 was reexamined (Fig. 6F-H).1t is
Description of type species closely similar to H. ingolfi in shape and spicules (tylotes 700 X 20
f.Lm, acanthoxeas 170 X 34 f.Lm, arcuate isochelae 18-28 f.Lm, sigmas
Histodermella ingolfi Lundbeck, 1910 (Fig. 6A-E). 30--44 f.Lm) differing merely in the possession of sigmas and the
Synonymy. Histodermella ingolfi Lundbeck, 1910: 14, pI. II thicker acanthoxeas (Fig. 6G). Stylote acanthoxeas are rare and obvi-
figs 1-4, pI. N fig. 4. ously the same spicule type. Histodermella may be mistaken for the
Material examined. Syntypes: Not examined. There are acamid genera Zyzzya de Laubenfels (1936a) or Cornulella Dendy
several type specimens, assumed to be in the Copenhagen (1905), if studied superficially, as both have diactinal acanthose auxil-
Museum. Other material. BMNH 1953.11.11.91- a slide made of iary megascleres in combination with tylotes as principal megascleres.
the specimen described by Stephens (1920), no. SR 480, from the However, the tylotes of the latter two genera are distinctly microspined
W coast of Ireland. at both ends, they lack sigmas, and have palmate isochelae.
538 Porifera· Demospongiae • Poecilosclerida • Myxillina • Coelosphaeridae

D
c
Fig. 6. Histodennella. A-E, Histodennella ingolfi Lundbeck (1910), drawings of habit (A) and spicules (B-E) reproduced from Lundbeck, 1910 (sizes see
text). F-H, Histodennella australis Dendy (1924), type of Hiltonus de Laubenfels (1936a), SEM photos of spicules from the holotype (scales: F, 100/-Lm;
G, 10 /-Lm; H, l/-Lm).

INFLATELLA SCHMIDT, 1875 injlata Hansen, 1885: 7, pI. I fig. 4. Joyeuxia viridis; Topsent, 1904b:
205, pI. ill fig. 12 (not: Topsent, 1892a: 94 =Inflatella viridis).
Synonymy Material examined. None. Syntypes: 5 specimens, not
located. A slide is present in the Berlin Museum (ZMB 6792)
Inflatella Schmidt, 1875; 117. Joyeuxia Topsent, 1892a; 94. according to Desqueyroux-Faundez & Stone (1992). This is here
designated the lectotype.
Type species Description (mostly derived from Lundbeck, 1910).
Stalked bladders (Fig. 7A-B) with 1-4 short fistules on the smooth
Inflatella pellicula Schmidt, 1875: 117 (by monotypy). upper surface. Size up to 2 cm high, fistules up to 4 mm long.
Oscules at the end of a fistule. Interior virtually hollow, skin thin
Definition but firm. Consistency leathery. Skeleton of the ectosome is a felted
mass of tangential megascleres, intercrossing in all directions.
Coelosphaeridae with fistular shape, lacking choanosomal Choanosomal skeleton an irregular system of loose spicule tracts,
megascleres and microscleres; the only spicules present are little spongin. Spicules (Fig. 7C), smooth, strongylote, thickest in
strongylote spicules. the middle, tapering to somewhat unequal endings, 420 X 640 X
10--28 /-Lm. Distribution and ecology. Norway, North Atlantic, deep
Diagnosis water below 200 m depth.
Remarks. If the strongylote spicules were truly strongyles,
Massive, spherical with erect fistules; ectosomal skeleton this genus should probably be assigned to Chondropsidae.
a compact tangential crust of diactinal spicules; choanosomal However, in the absence of a choanosomal skeleton and microscle-
skeleton consists of thin tracts of scattered diactinal spicules, the res the assignment of this genus to Coelosphaeridae is based on
same as on the surface; diactinal megascleres are of a single sort, similarity to Coelosphaera and Histodermella in habit and mega-
being smooth ectosomal strongyles or anisotylotes; microscleres sclere shape, but this assignment is tentative. Vacelet &
absent. Half a dozen species. Vasseur (1971), followed by Desqueyroux-Faundez (1987)
used Inflatella as a genus of Phloeodictyidae related to Oceanapia
Previous reviews but with strongyles instead of oxeas. However, the type of
Inflatella shows little skeletal resemblance to Oceanapia and the
Lundbeck (1910: 18), Bergquist & Fromont (1988: 51). spicules are anisostrongyles with tapering ends, unlike those of
Oceanapia.
Description of type species The genus Joyeuxia Topsent, 1892a was erected (by mono-
typy) for green-coloured J. viridis Topsent (1892a: 94, pI. II fig. 8,
Inflatella pellicula Schmidt, 1875 (Fig. 7A-C). pI. X fig. 19) from deep water off the Azores. Two slides of one of
Synonymy. Inflatella pellicula Schmidt, 1875: 117, pI. I the type specimens were examined in MNHN, D.T. 64 and 65,
fig. 5; Lundbeck, 1910: 18, pI. IT figs 7-9, pI. N fig. 6. Reniera labeled 'Camp. Hirondelle, 1888, 100'. The species has a bulbous
Porifera' Demospongiae • Poecilosclerida • Myxillina • Coelosphaeridae 539

Fig. 7. A-D, Inflatella . A-C, Inflatella pellicula Schmidt (1875) (sizes see text). A, Schmidt's figure of the habit. B, various shapes reproduced from
Lundbeck (1910, pI. II figs 7-9, pI. IV fig. 6). C, spicules reproduced from Lundbeck (1910). D, Inflatella viridis (Top sent, 1892a as Joyeuxia), habit
and megasclere reproduced from Topsent (l892a, pI. II fig . 8, pI. X fig. 19) (sizes see text) . E-G, Lepidosphaera hindei Levi & Levi (1979). E, habit
(scale 1 cm). F, tylotes (scale 100 11m). G, 'ecaille' (scale 10 11m), all reproduced from Levi & Levi (1979, figs 1-5).

form and is largely hollow (Fig. 8D). There is no choanosomal Definition

skeleton, only a thick ectosome consisting of intercrossing
strongyles, which are slightly thicker in the middle, size 765 X 9 fJ-m. Coelosphaeridae with ectosomal crust of 'ecailles', semicircu-
The species has also been reported from the Caribbean by Burton lar disc-shaped spicules; megascleres tylotes only, no microscleres.
(1954). It is a clear Inflatella, but definitely a different species
(except for Topsent's 1904b record). The major differences with Diagnosis
I. pellicula are the distinctive colour and the somewhat longer
strongyles. The genus name has been used for a number of unre- Basal mass burrowing with erect fistules protruding through sub-
lated sponges (e.g., Injlatella bartschi de Laubenfels (1934: 21), strate; ectosomal skeleton consists of a rigid cortex composed of
I. dura Vacelet & Vasseur (1971: Ill), I. perlucida Desqueyroux- peripheral layer of platelet-like discs, 'ecailles'; choanosomal skeleton
Faundez (1987: 211), all probably belonging to the phloeodictyid pulpy with smooth tylotes dispersed throughout; microscleres absent.
genus Oceanapia), and it is likely that only a few species belong
to it. These appear to be confined to cold and deep waters in both Description of type species
the North Atlantic (I. pellicula, I. viridis), the North Pacific
(I. globosa Burton, 1955,fide Koitun, 1959: 105), and the Southern Lepidosphaera hindei Levi & Levi, 1979 (Fig. 7E-G).
Oceans (I. belli Kirkpatrick, 1907a: 203). Dendy, 1924 followed by Synonymy. Lepidosphaera hindei Levi & Levi, 1979: 443,
Bergquist & Fromont (1988: 51) assigned a species with exclu- figs 1-5.
sively oxeas to this genus, viz., Injlatella spherica Dendy (1924: Material examined. None. Holotype: MNHN DCI 2447 -
373). This is here transferred to Dendoricellidae on account of its could not be found in 1996 (Claude Levi, pers.comm.).
similarities in shape, and spicule sizes and form to Pyloderma Description (from Levi & Levi, 1979). Fistular sponge, con-
latrunculioides. sisting of a basal mass of about I rom 3 fixed on a mass of calcareous
debris, issuing a single tube of 3 mm long and 0.35 rom diameter
(Fig. 7E). The tube is open at the end. Skeleton. Of the surface,
a rigid cortex of semicircular disc-shaped 'ecailles', overlapping one
LEPIDOSPHAERA LEVI & LEVI, 1979 another partly, to form a continuous cover. Choanosome pulpy, with
dispersed tylotes. Spicules. Ectosomal 'ecailles' (Fig. 7G), smooth,
Synonymy ellipsoid, but with one straight and one curved side, 95-100 X 32-35
fJ-m; tylotes (Fig. 7F), with prominent heads, 130-150 X 5-6 fJ-m.
Lepidosphaera Levi & Levi, 1979: 443. Distribution and ecology. Off New Caledonia, dredged from 200m
offshore from the reefs; no further records.
Type species Remarks. The unique 'ecailles' are unprecedented in Recent
sponges, and their origin and relationships with other spicule types
Lepidosphaera hindei Levi & Levi, 1979: 443 (by monotypy). remains a matter of speculation. Regrettably, type material can not
540 Porifera' Demospongiae • PoeciloscIerida • Myxillina • Coelosphaeridae

be located anymore, but we rely on Prof. Levi's vast experience in occasionally absent or modified to strongyles or oxeas; arcuate
the assumption that these 'ecailles' are truly sponge spicules and not chelae.
foreign material from some unknown source used by the sponge to
enforce its surface. Prof. Levi pointed out that Hinde & Holmes
(1892) recorded very similar fossil spicules (semicircular platelets Diagnosis
of 92 X 27/-Lm) from the Oamaru deposits, New Zealand. The
membership of the family Coelosphaeridae is determined by Massive, lobate, or flabelliform sponges, with irregular or
elimination, based on the presence of tylotes in combination with clathrate surface. Ectosomal skeleton with smooth ectosomal
fistular shape. No firm evidence for this assignment can be offered tylotes or strongyles forming tangential tracts and surface brushes;
without further corroboratory support from re-examination of choanosomal skeleton composed of smooth or acanthose choanoso-
material. mal styles, sometimes oxeas or strongyles, forming an isodictyal
reticulate architecture of single spicules; echinating spicules may be
present; microscleres are arcuate isochelae, sigmas, and raphides in
USSODENDORYX TOPSENT, 1892 trichodragmata. Cosmopolitan distribution, many species.

Synonymy
Previous review
Lissodendoryx Topsent, 1892a: 97. Paramyxilla Dendy, 1905:
233. Ectyodoryx Lundbeck, 1909: 445. Anomodoryx Burton, 1934a: Hofman & Van Soest (1995: 78).
555. Damiriella Burton, 1935c: 404. Jones de Laubenfels, 1936a:
79. Waldoschmittia de Laubenfels, 1936a: 95. Zetekispongia de
Laubenfels, 19~6b: 446. Zetekopsis de Laubenfels, 1936a: 85. Taxonomic remarks
Zottea de Laubenfels, 1936a: 64. Damiriana de Laubenfels, 1950a:
14. Acanthodoryx Levi, 1961b: 516. The present concept of Lissodendoryx is widened compared
to the latest revision (see Hofman & Van Soest, 1995) with inclu-
Type species sion of species having other tomote morphologies than strictly
smooth tylotes, as well as with species lacking sigmas.
Tedania leptoderma Topsent, 1889: 49 (by original designa- The widened concept is accompanied by a proposal to recognize
tion) (this is generally considered a junior synonym of five subgenera: Lissodendoryx (full spicule complement but no
Halichondria isodictyalis Carter, 1882a). echinating spicules), Ectyodoryx (echinating acanthostyles),
Anomodoryx (single megasclere type consisting of strongyles or
Definition tylotes), Waldoschmittia (oxeas as choanosomal megascleres) and
Acanthodoryx (plumose skeleton). It is assumed that the gender of
Coelosphaeridae with ectosomal tornotes in the form Lissodendoryx is female, although no direct evidence for such an
of tylotes or strongyles; choanosomal smooth or spined styles, assumption is available.

Key to subgenera of Lissodendoryx

(1) Only a single megasclere type .......................................................................................................................................... Anomodoryx

At least two megasclere types (differentiated ectosomal and choanosomal megascleres) .................................................................. 2
(2) Choanosomal megascleres oxeas ................................................................................................................................. Waldoschmittia
Choanosomal megascleres smooth or spined styles ............................................................................................................................ 3
(3) Skeletal architecture plumose with thick tracts of (acantho-) styles .............................................................................. Acanthodoryx
Skeletal architecture reticulate, usually isotropic ........ ................ ...................... ........... ......... ....... .......... ...... .......... ... ... ........... ............ 4
(4) Spicules include a second category of smaller acanthostyles ............................................................................................. Ectyodoryx
No second category of smaller acanthostyles ................................................................................................................ Lissodendoryx

SUBGENUS USSODENDORYX TOPSENT, 1892 Definition

Synonymy Lissodendoryx with a full complement of megascleres includ-

ing ectosomal tylotornotes and choanosomal styles, which mayor
Lissodendoryx Topsent, 1892a: 97. ? Paramyxilla may not bear spines; no smaller category of echinating acanthostyles
Dendy, 1905: 233. Jones de Laubenfels, 1936a: 79. microsc1eres inc1ude arcuate isochelae and sigmas, which may be
Zetekopsis de Laubenfels, 1936a: 85. Zottea de Laubenfels, absent. Raphides may be present. Distribution. Circumglobal warm
1936a: 64. water, occasionally also in temperate and cold water.

Description of type species

Type species
Lissodendoryx (Lissodendoryx) isodictyalis (Carter, 1882a)
Tedania leptoderma Topsent, 1889: 49. (Fig.8A).
Porifera· Demospongiae • Poecilosclerida • MyxiUina • Coelosphaeridae 541

-I ,I ...

I
Fig. 8. Lissodendoryx (Lissodendoryx). A, Lissodendoryx (Lissodendoryx ) isodictyalis (Carter, 1882a as Halichondria), drawing of skeleton and spicules
reproduced from Van Soest (l984b, fig. 19). B, Lissodendoryx (Lissodendoryx) amaknakensis (Lambe, 1894 as Myxilla) , type of Jones de Laubenfels
(1936a), drawing of shape and spicules reproduced from Lambe (1894, pI. II fig . 10) (sizes see text) . C, Lissodendoryx (Lissodendoryx) indistincta
(Fristedt, 1887 as Hastatus), type of Zetekopsis de Laubenfels (1936a), drawing of skeleton and spicules made from BMNH type slide. D-H,
Lissodendoryx (Lissodendoryx) grata (Thiele, 1903a as Myxilla) , type of Zottea de Laubenfels (l936a), SEM photos of spicules from the SMF holotype,
reproduced from Hofman & Van Soest (1995: figs 1-8) (scales: D- E, lOj.Lm, F-G, I j.Lm). I,Lissodendoryx (Lissodendoryx) infrequens Carter (1881b:
369, as Halichondria), type of Paramyxilla Dendy (1905) , drawing of spicules reproduced from Carter, l881b: pI. VIII fig . 9(sizes see text).

Synonymy. Halichondria isodictyalis Carter, 1882a: 285, Description. Lobate masses, which may attain considerable
pI. XI fig. 2. Tedania leptoderma Topsent, 1889: 49; Lissodendoryx size, often more than 10 cm in diameter. Surface irregular, ridged,
leptoderma; Topsent, 1892a: 97; Lissodendoryx isodictyalis; undulating, but smooth inbetween. Oscules may be prominent, on
Topsent, 1897a: 456. Esperiopsis fragilis Verrill, 1907: 333, volcano-shaped lobes and deep. Consistency soft, easily torn.
pI. XXXV-C figs 1-3. Colour light greyish green, occasionally pale yellowish. Skeleton
Material examined. Syntypes (8 specimens, not examined): (Fig. 8A) of the ectosome loose or bundled tylotes, not coherent.
CLM Sp. 57-58, 62-66, 73 - Puerto Caballo, Nassau, Bahamas Choanosomal skeleton reticulate, isotropic, with single styles or
(cf. Wiedenmayer, 1977b: 258, table 51). Other material. paucispicular bundles. Spicules (fig. 8A) tylotes with well-
Specimens of L. isodictyalis: ZMA collection (see Van Soest, developed heads, occasionally polytylote, heads usually smooth, but
1984b). Holotype of Tedania leptoderma: MNHN, not examined. occasionally with a few spines, 150-230 X 2-4 fLm; styles, usually
542 Porifera· Demospongiae • PoeciloscIerida • Myxillina • Coelospbaeridae

smooth, but not infrequently with a few spines near the blunt end, SUBGENUS ACANTHODORYX LEVI, 1961
occasionally subtylostylote, 130-200 X 2-6 fLm; arcuate isoche1ae
in two distinct size categories, 19-34 and 8-16 fLm; sigmas in two Synonymy
distinct size categories, 23-45 and 11-20 fLm. Distribution and
ecology. West Indies, E coast of North America, also recorded from Acanthodoryx Levi, 1961b: 516.
the E Atlantic and E Pacific; mangroves and sea grass meadows.
Remarks. Closely similar species (e.g., L. similis Thiele, Type species
1899: 18, and L. ternatensis (Thiele, 1903a: 952 as Hamigera) are
found in a circumtropical belt, and may be considered disjunct Acanthodoryxfibrosa Levi, 1961b: 516 (by monotypy).
populations of a former widespread parent species.
The genus Jones de Laubenfels (1936a: 79) was erected (orig- Definition
inal designation) for type species Myxilla amaknakensis Lambe,
(1894: 122, pI. II fig. 10, here reproduced as Fig. 8B). This has Lissodendoryx with radiating choanosomal skeleton of
sparsely spined, blunt-ending tomotes, 137 X 8 fLm, entirely spined thick bundles of acanthostyles, showing little anastomosing, no
styles, 144 X 13 fLm, in a renieroid reticulation; microsc1eres sigmas.
appear to be arcuate isochelas, 22 fLm; no sigmas. It belongs to
a group of Lissodendoryx species with spined styles, characteristic Previous review
of cold water.
The genus Zetekopsis de Laubenfels (1936a: 85) was erected Levi (1961b: 516).
(by original designation) for type species Hastatus indistinctus
Fristedt (1887: 444) on account of alleged spined sigmas. A slide Description of type species
of the holotype was found in the Natural History Museum,
London, BMNH 1910.1.1.1424, labeled 'From Herr K. Fristedt, Acanthodoryxfibrosa Levi, 1961b (Fig. 9A-D).
Spitzbergen, Ar 1872'. The skeleton (Fig. 8C) is an isotropic Synonymy. Acanthodoryx fibrosa Levi, 1961b: 516,
reticulation of single styles or bundles of 2-3. Ectosomal tylotes fig. 8.
are mucronate at one end, normally tylote at the other, 200 X Material examined. Holotype: MNHN DCI. 700 -
5 fLm; styles are lightly spined at both apices but smooth inbe- Zamboanga, Philippines.
tween, 350 X 14 fLm. Chelae consist of larger normal arcuate Description. Massive sponge (Fig. 9A), size about 2.5 X 4cm,
isochelae, 25 fLm, and smaller unguiferate isochelae, about 7 fLm composed of a mass of radiating fibres covered by a thin membrane.
(considered sigmas by Fristedt). Sigmas are normal shaped and Colour white in alcohol, probably red in life. Skeleton plumose,
relatively large, 50 fLm. No sigmas with 'very long spines' attrib- strong choanosomal fibres form a system of radiating tracts, occa-
uted to this species by de Laubenfels were found. The species sionally anastomosing and with divergent spicules, but not echinated.
belongs to the group of cold water Lissodendoryx species with Fibres 50-160 fLm in diameter. The ectosomal skeleton is poorly
spined styles. developed. Spicules (Fig. 9B-D), ectosomal tylotes, elliptical heads
The genus Zottea de Laubenfels (1936a: 64) was erected (by slighty inequiended, 200-240 X 3-4 fLm; acanthostyles, lightly
original designation) for type species Myxilla grata Thiele (1903a: spined allover, in a large size variation, up to 150 X 170 X 7 fLm;
954, fig. 19) on account of its choanosomal spined strongyles arcuate isochelae in two size categories, 33-34 and 20-21 fLm.
(Fig. 8E). The holotype from the Senckenberg Museum, SMF 1783, Distribution and ecology. Philippines, shallow water coral reefs.
from Temate, Indonesia, was re-examined (see also Hoffman & Remarks. So far only a single species can be attributed to
Van Soest, 1995: 83). This is a massive lobate sponge with ectoso- this subgenus. The structure of the skeleton reminds of the genus
mal smooth tylotes (Fig. 8D), 190-235 X 2-6 fLm, and an isod- Phorbas in the family Hymedesmiidae, but the choanosomal tracts
ictyal skeleton of acanthose strongyle-like spicules (in fact they are in that genus are truly plumose with no anastomosing and they
stylote spicules with a blunt and a more pointed end), 130-180 X are usually echinated by acanthostyles; the surface characters
5-16 fLm. The microsc1eres are normal arcuate isochelae (Fig. 8G), of Phorbas with the areolae are lacking in the present species.
16-32 fLm, sigmas (Fig. 8F), 12-26 fLm, and trichodragmas
(Fig. 8H), 45-140 fLm. The species falls within the synonymy of
the present subgenus. SUBGENUS ANOMODORYX BURTON, 1934
The genus Paramyxilla Dendy (1905: 233) was erected for
type species Halichondria infrequens Carter (1881b: 369, pI. vrn Synonymy
fig. 9, here reproduced as Fig. 81). The type specimen was not
found during a recent search in the BMNH, so we cannot be sure of Anomodoryx Burton, 1934a: 555. Damiriella Burton,
its exact affinities. It possesses spined oxeas, about 130 X 4-5 fLm, 1935a: 404.
as choanosomal megasc1eres; ectosomal spicules are the usual
tylotes, about 260 X 3 fLm, and microscleres comprise arcuate Type species
isochelae, about 40 fLm, and sigmas about 60 fLm (sizes calculated
from Carter's drawings). On this basis it may be considered close Desmacidon dendyi Whitelegge, 1901: 78 (by original
to Lissodendoryx (Lissodendoryx) grata. However, it may also be a designation).
member of Lissodendoryx (Waldoschmittia), of which the type has
smooth oxeas (cf. below). The spined oxeas are unusual, so far Dermition
unique, and its importance as a phylogenetic character remains to
be studied. Lissodendoryx with a single megasclere type.
Porifera· Demospongiae • Poecilosclerida • Myxillina • Coelospbaeridae 543

Fig. 9. A-D, Lissodendoryx (Acanthodoryx)fibrosa (Levi, 1961b). A, habit of holotype (scale 1 cm). B-D, SEM photos of spicules made from the holo-
type (scale 10 !Lm). E-H, Lissodendoryx (Anomodoryx) dendyi (Whitelegge, 1901 as Desmacidon). E, habit of holotype reproduced from Whitelegge
(1901) (scale 1 cm). F-H, SEM photos of spicules from the type (scales: F, l!Lm; G-H, lQ!Lm). I, Lissodendoryx (Anomodoryx) cavernosa (Topsent,
1892a as Damiria), type of Damiriella Burton (1935c), drawing of spicules reproduced from Topsent (1936: fig. 4) (sizes see text).

Previous review Description. Massive, clathrate sponge (Fig. 9E).

Reticulated, grooved surface, space between ridges 2-3 mm.
Burton (1934a: 554). Oscules scattered, about 5 mm in diameter. Consistency spongy,
somewhat brittle. Colour yellowish grey (dry condition). The frag-
Description of type species ment examined in the Natural History Museum, London, was
1.5 X 1.5 X 1.5 cm, original specimen 15 X 10cm (Fig. 9E). At the
Lissodendoryx (Anomodoryx) dendyi (Whitelegge, 1901) surface individual megascleres are scattered in the dermal mem-
(Fig. 9E-H). brane; the choanosomal skeleton is an irregular system of wispy
Synonymy. Desmacidon dendyi Whitelegge, 1901: 78, pI. X fibres of spongin filled with spicules, of about 100 /Lm in diameter,
fig. 9; Homoeodictya dendyi; Hentschel, 1911: 319, fig. 17; interconnected irregularly with thinner bundles, 3-4 spicules thick.
Anomodoryx dendyi; Burton, 1934a: 555. Meshes 150-200/Lm. Spicules. Tylotes (Whitelegge calls them
Material examined. Holotype (fragment): BMNH strongyles, but the megascleres have distinct tyles (Fig. 9F) and are
1927.6.23.9 - Port Stephens, New South Wales, Australia (from truly equiended tylotes), 150-250 X 2.5-9 /Lm; arcuate chelae
AMG304Z). (Fig. 9H) in a variable but continuous size, 18-25 /Lm (Whitelegge
S44 Porifera' Demospongiae • Poecilosclerida • Myxillina • Coelosphaeridae

mentions two sizes, 25 and 18 /-Lm); sigmas (Fig. 9G), 35-40/-Lm. Gronland, Exp. af Sophia, ar 1883, from Herr K. Fristedt, Norman
Distribution and ecology. Central coast, New South Wales, CoIl. (with sections), and 'Hastatus foliatus Fristedt, Greenland,
Australia, shallow water. from K. Fristedt, Norman Coli.", prepared from slide 10.1.1.1435'
Remarks. The genus Damiriella was erected by Burton, (spicule mount).
1935a: 404 for type species Damiria cavernosa Topsent (1892b: Description (from Fristedt, 1887). Leaf-shaped. Height up
xxii) (by original designation), from Cap l' Abeille, Banyuls area. to 3.5 cm, thickness 3 mm. Surface even and smooth. Oscules
Topsent's original description mentions a cavernous, massive, numerous, small. Consistency fragile, soft. Colour grey in alcohol.
brown soft sponge, with ectosomal smooth tylotes, 300 X Skeleton (Fig. lOA, few ectosomal tomotes scattered near the sur-
6-10 /-Lm, a reticulation of smooth strongyles, 250 X 15 /-Lm, and face; choanosomal skeleton an irregular isotropical reticulation of
isochelae which are not further characterized. Later, Topsent bundles of 3-5 acanthostyles echinated by smaller acanthostyles.
(1936: 19, fig. 4, here reproduced in Fig. 91) redescribed the Spicules (Fig. lOA), ectosomal tomotes, tylote-like with unequal
species on the basis of new material collected off Monaco. mucronated heads, 200-280 X 4 /-Lm; large structural acan-
Hoffman & Van Soest (1995) on the basis of this description thostyles, up to 300 X 20 /-Lm; smaller acanthostyles, entirely but
assumed that ectosomal and choanosomal megascleres were differ- weakly spined, 130-300/-Lm; arcuate isochelae, strongly but evenly
entiated into tylotes and strongyles. For the present study, a MNHN curved, possibly in two size categories, 25-35/-Lm; sigmas small
slide of the later Topsent material was examined labeled and thin, 20/-Lm. Distribution and ecology. Norway; North Atlantic,
"Lissodendoryx cavernosa Monaco, 19.IV.27' registered as D.T. Arctic, also recorded from South Africa; deep water, 70-1948m
74. It has plumose bundles of tylote-like tomotes, 275-350 X 7-9 depth.
/-Lm, fanning out at the surface to form a tangential skeleton. The Remarks. Lundbeck (1909) proposed to subdivide the gen-
tylotes vary in shape to include strongylote and even stylote forms, era Myxilla and Lissodendoryx (which were stated to differ only in
but they are obviously the same structural type. The choanosomal possession of anchorate vs. arcuate chelae) further into those with
skeleton is a reticulation of smooth stylotes, rarely provided with a smaller echinating acanthostyles (Ectyomyxilla and Ectyodoryx)
few spines, and not infrequently strongylote or even tylote, and those without them (Myxilla and Lissodendoryx s.s.). This arti-
235-325 X 7-10 /-Lm. In fact, the ectosomal and choanosomal ficial distinction is here maintained only at the subgenus level. It
spicules are not at all clearly differentiated, and Topsent's (1936) should be noted that Ectyomyxilla was a nomen nudum since no
attempt to consider the choanosomal spicules as (acantho-)styles type species was mentioned (cf. chapter on Myxillidae). Lundbeck
was misleading. The microscleres are relatively elongated larger (1909) reexamined Fristedt's type specimen and ascertained it had
arcuate chelae 23-30/-Lm and smaller reduced arcuate chelae the echinating spicules (be it few in number). This is confirmed
10-13 /-Lm. Pulitzer-Finali (1978: 56) described further material, from the slide material examined. Other sympatric Lissodendoryx
confirming that the megascleres are mere variations of the same (Ectyodoryx) species are Ectyodoryx atlanticus Stephens
(probably ectosomal) spicule type. Thus, the only difference with (1920: 43) and E. multiformis Br~ndsted (1932: 10). These differ
Anomodoryx is the absence of sigmas, which is considered to be of substantially in shape (thinly encrusting in E. atlanticus) and
specific level. spicule sizes and categories (megascleres twice as long in
E. atlanticus; several categories of microscleres in E. multiformis).
Both are to be renamed Lissodendoryx (Ectyodoryx) atlantica and
SUBGENUS ECTYODORYX LUNDBECK, 1909 L. (E.) multiformis.

Synonymy
SUBGENUS WAWOSCHMI1TIA DE
Ectyodoryx Lundbeck, 1909: 445. LAUBENFELS, 1936

Type species Synonymy

Hastatus foliatus Fristedt, 1887: 443 (by original ? Paramyxilla Dendy, 1905: 233. Waldoschmittia de
designation). Laubenfels, 1936a: 95. Zetekispongia de Laubenfels, 1936b: 446.
Damiriana de Laubenfels, 1950a: 14.
Definition
Type species
Lissodendoryx with echinating acanthostyles.
Crella schmidti Ridley, 1884a (by original designation).
Previous reviews
Definition
Lundbeck (1909: 445), Hofman & Van Soest (1995: 81).
Lissodendoryx with oxeas as choanosomal megascleres.
Description of type species
Description of type species
Lissodendoryx (Ectyodoryx)foliata (Fristedt, 1887) (Fig. lOA).
Synonymy. Hastatus foliatus Fristedt, 1887: 443, pI. 25 Lissodendoryx (Waldoschmittia) schmidti (Ridley, 1884a)
figs 7-12; Ectyodoryxfoliata; Lundbeck, 1909: 445. (Fig. lOB-G).
Material examined. ? Holotype (slides): BMNH Synonymy. Crella schmidti Ridley, 1884a: 432, pI. XLI
1910.1.1.1435, 1435A - labeled "Hastatus foliatus Fristedt, fig. A; Damiria schmidti; Topsent, 1897a: 455. Damiria australiensis
Porifera' Demospongiae • Poecilosclerida • Myxillina • Coelosphaeridae 545

Fig. 10. A, Lissodendoryx (Ectyodoryx) Joliata (Fristedt, 1887 as Hastatus), drawing of skeleton and spicules made from a BMNH type slide. B-G,
Lissodendoryx (Waldoschmittia) schmidti (Ridley, 1884a as Crella). B, drawing of skeleton and spicules made from the BMNH holotype. C-F, SEM photos
of spicules from the BMNH holotype (scales: I /-Lm, except A left and B left, to /-Lm). G, drawing of spicules of Lissodendoryx (Waldoschmittia) zonea de
Laubenfels (1936b as Zetekispongia), junior synonym of L. (W) schmidti, reproduced from his fig. 40 (sizes see text). H, drawing of spicules of Lissodendoryx
(Waldoschmittia) hawaiiana de Laubenfels (1950a as Damiriana), junior synonym of L. (W) schmidti, reproduced from his fig. 7 (sizes see text).

Dendy, 1896: 28; Lindgren, 1898: 25, pI. 17 fig. 10, pI. 19 fig. 15; Hofman & Van Soest, 1995. Type slide of Damiria australiensis:
Myxilla schmidti; Thiele, 1903a: 954; Dendoricella schmidti; BMNH 1929.1.26.39. Holotype of Zetekispongia zonea: USNM
Hentschel, 1911: 328, fig. 22; Waldoschmittia schmidti; de 22215 (considered synonymous with the former). Holotype of
Laubenfels, 1936a: 95. Zetekispongia zonea de Laubenfels, 1936b: Damiriana hawaiiana: USNM 22737 (also considered
446, fig. 40. Damiriana hawaiiana de Laubenfels, 1950a: 14, synonymous).
fig. 7; Damiriana schmidti; Levi, 1958: 30, fig. 25. Lissodendoryx Description. Massive, lobate, riddled with channels of
aspera; Hofman & Van Soest, 1995: 87, figs 19-24 (not 1-4mm diameter. Surface covered with narrow longitudinal ridges,
Halichondria aspera Bowerbank, 1875b: 287). rough, but smooth in between. Oscules small and few, lying in
Material examined. Holotype: BMNH 1881.18.23.329 depressions. Consistency crumbly, colour dirty yellowish white (in
(dry), BMNH 1881.18.23.328 (slides with sections). alcohol), probably red in life. Skeleton (Fig. lOB). The ectosomal
Other material. Several specimens from Indonesia cited by skeleton consists of brushes of tylotes lying at distances of 250 Il-m
546 Porifera' Demospongiae • Poecilosclerida • Myxillina • Coelospbaeridae

enclosing distinct subdennal spaces. The tylotes fan out at the sur- makes up the bulk of the material of Halichondria aspera and was
face to fonn a tangential skeleton. The sections show an irregular obviously the material intended for it by Bowerbank, Hofman &
isotropic skeleton of single oxeas or 2-3 per side fonning meshes van Soest's (1995) use of the species name for Lissodendoryx
of 150 X 250 IJ,m. Scattered microscleres occur especially at the material with oxeas as the structural megascleres is thus incorrect
surface. Spicules. Ectosomal tylotes (Fig. 1OC), entirely smooth, (see also chapter Tedaniidae).
with distinct, elongate heads, slightly flattened at the end: The genus Zetekispongia de Laubenfels, 1936b was erected
210--(228.9)-264 X 4-(5.3)-7IJ,m; oxeas (fig. 1OD), smooth, with (by monotypy) for the type species Z. zonea de Laubenfels (1936b:
sharp occasionally mucronate ends: 183-(194.5)-210 X 6-(7.4)-8 446, fig. 40, here reproduced as Fig. lOG). The type specimen
IJ,m; arcuate chelae (Fig. lOE) in two size categories, chela I USNM 22215 from the Pacific coast of Panama was re-examined.
29--(33.3)-39IJ,m; chela II 18--(20.5)-24IJ,m; sigmas (Fig. lOF) in This is a massive sponge with reticulate skeleton of smooth oxeas,
two size categories, sigma I: 26-(28.9)-31IJ,m; sigma II: 210--2151J,m, with ectosomal tylotes, 170--200 IJ,m, and arcuate
16-(18.4)-21IJ,m. Distribution and ecology. Described originally isochelae, 36 and 16-24IJ,m, and sigmas, 16-24IJ,m. This is a
from Port Jackson, SE Australia, but subsequently recorded from all junior synonym of L. (Waldoschmittia) schmidti.
over the Indo-Pacific. If synonymy is correct, then the species also The genus Damiriana de Laubenfels, 1950a was erected (by
occurs in Hawaii and along the W coast of Panama; shallow-water. monotypy) for D. hawaiiana de Laubenfels (1950a: 14, fig. 7, here
Remarks. This species was considered a junior synonym reproduced in Fig. lOH). The type specimen USNM 22737 was
of Halichondria aspera Bowerbank, 1875b from the Straits of reexamined. This is a massive red sponge, smooth with some digi-
Malacca, type of the genus Xytopsihis de Laubenfels (1936a) by tations; oscules relatively large. Spicules include ectosomal
Hofman & Van Soest (1995: 87). Bowerbank's description is mis- smooth tylotes, 170 X 41J,m; smooth oxeas, 200--230 X 8-9 IJ,m;
leading in giving the spicule combination of tylotes, oxeas, styles, arcuate isochelas, 15-27 IJ,m; sigmas 131J,m. This is also a likely
two size categories of chelae, sigmas. The type specimen and two junior synonym of L. (Waldoschmittia) schmidti.
microscopic slides are kept in the BMNH London, unregistered. It The genus Paramyxilla Dendy (1905: 233) was erected for
is an irregularly lobate sponge of 8 X 6 X 3 cm, with an irregularly type species Halichondria infrequens Carter (1881b: 369, pI. VIII
pitted surface. At its undersurface there is a smaller cushion-shaped fig. 9, here reproduced as Fig. 81), see description and discussion
sponge attached, size 3 X 2 X 0.5 cm, with a smooth surface. The above under Lissodendoryx (Lissodendoryx). It possesses spined
larger sponge is a Tedania, probably matching the description of oxeas as choanosomal megascleres, which spined oxeas are
T. dirhaphis Hentschel (1912). No chelae or sigmas are found in unusual, so far unique. It may be close to either Lissodendoryx
this specimen. However, the smaller sponge is a Lissodendoryx, (Lissodendoryx) grata (cf. above) or to Lissodendoryx
keying out as L. tematensis (Thiele, 1903a). Since the Tedania (Waldoschmittia) schmidti.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Crambeidae Levi, 1963

Rob W.M. Van Soest

Zoological Museum, University of Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, Netherlands. (soest@science.uva.nl)

Crambeidae Levi (Demospongiae, Poecilosclerida) is characterized by the possession of anchorate chelae in combination with
monactinal ectosomal megascleres in addition to thicker choanosomal styles or tylostyles. Peculiar acanthose microxeas occur in three of
the four genera considered valid, whereas two of the four genera are sublithistid, having desmoid spicules. Several species of two genera
exhibit unusually large variability of spicule presence and morphology.
Keywords: Porifera; Demospongiae; Poecilosclerida; Myxillina; Crambeidae; Crambe; Discorhabdella; Lithochela; Monanchora.
-.----- --------- - , , _. . . __ __ . _
. .. _--- . __. -_._.. -_._- -----------

DEFINITION, DIAGNOSIS, SCOPE Later, Levi (1973) associated Crambe with desma-free genera
Monanchora and Echinostylinos, and assigned these to a large fam-
Synonymy ily Esperiopsidae (thus, in fact, returning it to the position given to
it by Topsent, 1928). Van Soest (1990) discussed the similarities in
[Crambidael Levi, 1963: 16 (preocc.). Crambeidae chela-shape between Crambe and Monanchora and Van Soest et al.
Maldonado et al., 2001: 1261. (1996a) added to this the shared chemistry of these genera. The lat-
ter authors made the suggestion that Crambe and Monanchora
Definition were likely to be synonyms, but in view of the fact that several
species exist with and without the pseudastrose desmoid spicules
Myxillina with subty1osty1es as ectosomal megascleres and it is practical to retain both as separate genera. The contents of
anchorate-unguiferate chelae as microscleres; occasionally also the family are here extended to include Discorhabdella and
spined microrhabds. sigmoid spicules, and astro- or sphaeroclone Lithochela, which share important similarities. Maldonado &
desma-like spicules and derivates thereof are present. Uriz (1996) demonstrated the relatedness of Crambe and
Discorhabdella. Lithochela is added because of the combination of
Diagnosis unguiferate chelae, spined microrhabds and ectosomal styles. Its
desmas, though differing in shape and skeletal position, are consid-
Encrusting, massive, ramose or tubular growth forms. ered homologous to those of Crambe. Crambeidae are encrusting
Ectosomal megascleres consist of smooth subtylosty1es, usually to irregularly ramose or tubular sponges occurring on rocks, corals
arranged in brushes or bundles perpendicular to surface. and mangroves, mostly in shallow water, but some genera occur
Choanosomal megascleres are smooth or acanthose styles- down to 180m depth. Reproduction of Crambe is viviparous and
tylostyles forming hymedesmioid, plumose or plumoreticu1ate larvae - exhibiting the usual bare posterior pole - measure 850 X
skeletal structures. Peculiar desma-like spicules, likened to astro- 450 fLm (Uriz et al., 1998). Chemistry known from Crambe and
clones and sphaeroclones, occur in two genera. Microscleres con- Monanchora includes polycyclic guanidine alkaloids (Berlinck et
sist of spatulate or unguiferate anchorate isochelae, occasionally at., 1990; Van Soest et aI., 1996a), which are used in chemical
modified to sigma-like reduced forms which in some species can- defense (Turon et al., 1996); the compounds are also found to be
not be distinguished from true sigmas; finely spined microxeas active in vitro against HIV.
occur in some species of three genera.
Taxonomic remarks
Scope
Synonymy. [Crambidael Levi, 1963. This is a junior homo-
Ten nominal genera are included, of which four are consid- nym of Crambidae Latreille, 1810 (Insecta: Lepidoptera).
ered valid: Crambe, Discorhabdella, Lithochela and Monanchora. However, the name originally given by Levi should be emended to
Crambeidae, since the stem of Crambe is not obviously Cramb-
History and biology (which is the case for the insect Crambus). Thus, the family
name should be constructed from stem Crambe- by adding -idae as
Crambe was originally assigned to the large group of suffix. In this way, Crambeidae becomes available as an emended
'Desmacidonidae' by Vosmaer (1880) and retained there by family name in Porifera: Poecilosclerida: Myxillina (see Maldonado
Topsent (1928). De Laubenfels (1936a) assigned the genus to et al., 2001).
a family Monanthidae based on shared possession of desmas of Contents. The contents of this family are here newly
Crambe and e.g., Monanthus. The latter genus is here assigned to assigned. The sponges assembled here share the combination of
the family Desmanthidae ('Lithistida'). Levi (1963) recognized the anchorate chelae with monactinal tomotes. Also the choanosomal
unique nature of the Crambe desmas (they are in fact polyaxonic megascleres are thick, often knobbed or tuberculate tylostyles or
pseudastrose spicules unlike proper tetracrepid or monocrepid des- derivations thereof. Echinating acanthostyles are rare, but may be
mas), and accordingly erected a new family Crambidae for it. represented in one genus. Several species of three of the four

547
548 Porifera· Demospoogiae • PoecUosclerida • Myxillina • Crambeidae

genera of this family show a remarkable intraspecific variation in upon which tylostyles are erected, heads lodged between the
spicule presence, size and shape making identification unusually desmas (Maldonado & Benito, 1991; Driz & Maldonado, 1995).
complicated. Diagnostic in such cases are the presence of two mor- The sphaeroclone desmas are similar to those described for the
phological megasclere types, relatively long and thin ectosomal recent 'lithistid' Vetulina (Uriz & Maldonado, 1995), but since that
spicules (variously subtylostyles, strongyles or anisostrongyles) genus - along with a series of fossils classed in the family
and relatively short and fat choanosomal spicules (variously Chiastoclonellidae (with no extant representatives; see chapter
tylostyles, styles or strongyles). on fossillithistids, this volume) - has no spicules other than the
Desmoids. Species of two genera, Crambe and Lithochela, desmas, inclusion in the family Crambeidae is postponed until fur-
are provided with desmoid spicules. Those of Crambe are polyax- ther evidence is provided. Lithochela desmas appear to be
one spicules in the fonn of astro- or sphaeroclones (Driz & monocrepid, but some similarity with Crambe crambe desmoids is
Maldonado, 1995; Maldonado et al., 2001). In several species present.
of Crambe these desmas are interlocked to fonn a basal layer Previous review. Maldonado et al. (2001).

KEY TO GENERA

(1) Megascleres include short, spined, club-shaped pseudastrose tylostyles ..................................................................... DiscorlUlbdella
No short club-shaped acanthostyles ...... ................................................ .......... ........................................ .......... ...... ........ .................. ... 2
(2) Spicules include desrnas or irregular desmoid polyaxones (the latter may be rare) ............................................................................ 3
No desmas or desmoid polyaxones ....... .......... ...... .......... ...... .................. ...... ...................................................... .... .... .... .............. ....... 4
(3) Desmas are elongate monocrepids forming a reticulated choanosomal skeleton ................................................................ Lithochela
Desmas are small polyaxone or lumpy-lobate spicules concentrated at the base of the sponge .............................................. Crambe
(4) Live appearance includes white-lined veinal channel patterns ........................................................................................ Monanchora
Live appearance with unlined veinal patterns ............................... .......... ........ ...................................................... .................... Crambe

CRAMBE VOSMAER, 1880 Previous reviews

Synonymy Vacelet & Boury-Esnault (1982), Maldonado & Benito (1991),

Driz & Maldonado (1995), Maldonado & Driz (1996: 371).
Crambe Vosmaer, 1880: 135. [Plicatella] Schmidt, 1870: 45
(preocc.). Tetranthella Lendenfeld, 1894: 101. Description of type species

Type species Crambe crambe (Schmidt, 1862) (Figs 1, 2A).

Synonymy: Suberites crambe Schmidt, 1862: 66, pI. VI
Suberites crambe Schmidt, 1862: 66 (by original designation). fig. 9. Suberites fruticosus Schmidt, 1862: 66, pI. VI fig. 10.
Reniera labyrinthica Schmidt, 1864: 39, pI. IV fig. 9. Crambe
Definition harpago Vosmaer, 1880: 135. Crambe crambe; Riitzler, 1965: 32;
Maldonado & Driz, 1996: 372, figs IF, 2F, 3G. Stylinos brevicuspis
Crambeidae with astroclone or sphaeroclone desmoid Topsent, 1892b: xx. Tetranthellajruticosa; Lendenfeld, 1894: 101.
spicules, spatulate or unguiferate anchorate isochelae, and spined Hemimycale ambigua Sara, 1960a: 452, fig. 4. Hemimycale
microxeas (may all be absent). Live appearance shows veinal brevicuspis; Sara & Siribelli, 1960: 44.
patterns without white or yellow lining. Material examined. Lectotype (not examined): LMJG
15265 - Lissa, Adriatic. Paralectotype of Suberites crambe: LMJG
Diagnosis 15672 - no locality data, Adriatic. Other material. Paralectotype of
Suberitesfruticosus: LMJG 15667 - no locality data, Adriatic.
Thin flat crusts or thicker tubercular masses, occasionally Description (based on re-examination of the paralectotype
more elaborately lobate; surface translucent, slightly hispid, occa- specimens of Suberites crambe and Suberites fruticosus). Thin
sionally conulose, detachable, with clear veinal channel pattern in flat crusts (0.5 cm) or thicker masses (up to 1.5 cm), smooth or
life. Ectosomal skeleton composed of brushes of thin styles- with upright tubercles or projections. Surface with distinctly
subtylostyles (occasionally strongyles), which may fan out into a swollen veins in life (Fig. IF), slightly hispid or conulose. Lateral
tangential layer; choanosomal skeleton composed of desmas form- size may be considerable, up to 0.5 m2 (Becerro et al., 1994).
ing a semi-rigid basal skeleton (often absent), and stylote spicules Colour orange-red. Skeleton (Fig. 2A), near the surface brushes of
forming a plumose skeleton. Megascleres are thicker styles ectosomal megascleres fan out and many single spicules occur tan-
and thinner subtylostyles (may be modified into strongyles); gentially. In the choanosome bundles of megascleres (up to 10 or
microscleres are pseudastrose desmoid spicules (often absent), more in cross section) rise up from a basal spongin plate, occasion-
spined microxeas (often absent) and anchorate isochelae (occa- ally single megascleres are in erect position. Binding spongin is
sionally absent or reduced to sigmoid spicules). So far 6-7 variable and bundles may be loose and irregular, 50-130 11m in
species including one fossil have been assigned to this genus diameter. At the base peculiar pseudastrose desmas may be pres-
(Maldonado & Driz, 1996). ent, but these are rare and often absent in specimens. Likewise
Porifera· Demospongiae • Poecilosclerida • Myxillina • Crambeidae 549

Fig. I. Crambe crambe (Schmidt, 1862 as Suberites). A-E, SEM photos of spicules made from paralectotype specimens of Suberites crambe (A-D) and
S. fruticosus (E) (scales 10 j.Lm). F. specimen photographed in situ (photo B.E. Picton).

chelae may be present, rare or absent. Sand and other material are and calcareous tubes, and coralline algae; shallow-water down to
frequently incorporated in the basal parts. Spicules (measurements 367m depth.
taken from the paralectotype), styles from the peripheral bundles Remarks. The type species C. crambe is atypical in the
(Figs lA, 2A), smooth, straight, faintly constricted underneath the genus for its poorly developed astroclone desmas. Compared to
head, occasionally clearly subtylostylote, 316-(351)-381 X 7 11m C. acuata (Levi, 1958 as Folitispa), C. talliezi Vacelet & Boury-
(Riitzler, 1965: 225-320 X 3.5-5.5 11m); styles from the base of the Esnault (1982), c. tube rosa Maldonado & Benito (1991) and
sponge (Figs lB, 2A), longer and thicker than those of the periph- C. erecta Pulitzer-Finali (1993), they are reduced in form, asym-
eral bundles, straight, smooth, mostly faintly swollen subtermi- metrical and mostly not interlocked. Maldonado & Uriz (1996)
nally, 423-(489.3)-539 X 11-(14.6)-17 11m (Riitzler, 1965: mention the occurrence of elaborately shaped sphaeroclones in
370--400 X 8-13 11m); unguiferate anchorate chelae, often with slides of Topsent from Banyuls, but failed to provide evidence that
reduced shortened teeth (Figs 1D, 2A), 28-(38.3)-43 11m; astrose this material belonged with certainty to Crambe. The variability
desmoid spicules (Figs lC, E, 2A) with variable number of rays in spicule presence and shape in this species is remarkable and
with rounded knobbed ends and of unequal length, cladome parallels that known for Monanchora arbuscula (cf. below). The
38-(68.5)-99, rays 10--(35.7)-47 X 7-(10.6)-14 fLm. Distribution paralectotype specimen of S. crambe here examined has only
and ecology. Mediterranean, Atlantic coast of Portugal, Canary a few chelae, some of which are reduced (Fig. 2D), and relatively
Islands; encrusting on vertical rocks and ceilings of caves, on shells few desmoid spicules (Fig. 2C). The lectotype of S. jruticosus,
550 Porifera· Demospongiae • Poecilosclerida • Myxillina • Crambeidae

----=..

1
100
.t 1
A B

- lOG
...1
fJ
1 D c
Fig. 2. A, Crambe crambe (Schmidt, 1862 as Suberites), drawing of skeleton and spicules made from a slide of a paralectotype. B, Discorhabdella
incrustans Dendy (1924), drawing of skeleton and spicules made from a slide of the holotype. C, Discorhabdella tuberoscapitata (Topsent, 1890b as
Hymeraphia), type of Cionanchora de Laubenfels (1936a), drawing of the spicUles reproduced from Topsent (1892a: pI. XI fig. 6) (sizes see text).
D, Lithochela conica Burton (1928), drawing of skeleton and spicUles made from a slide of the paralectotype.

LMJG 15105, has abundant desmoids (Fig. 2E). Fossil spicules reduced spiculation. Recent specimens in the ZMA collection, e.g.,
described from the Oligocene of New Zealand (Hinde & Holmes, ZMA POR. 10966, combine a shape reminiscent of Schmidt's
1892) may belong to this species (described as Crambe oamaruen- drawing of R. labyrinthica in combination with a spiculation of
sis). Many other specimens lack both microscleres and desmoids, exclusively thin strongyles.1t is tentatively assumed that Schmidt's
and moreover have their spicules modified into strongyles. It is material and ZMA POR. 10966 are Crambe crambe. The genus
suspected that some of the specimens and species assigned to gen- name [Plicatella] Schmidt is preoccupied by the mollusc genus
era Batzella and Hemimycale are in reality specimens of Crambe or Plicatella Swainson, 1840.
Monanchora (cf. below). The genus Tetranthella Lendenfeld, 1894: 101 was erected for
The synonymy of Reniera labyrinthica Schmidt, 1864, type of type species Suberites fruticosus Schmidt, 1862: 66 (by original des-
[Plicatella] Schmidt, 1870: 45 (by subsequent designation, de ignation). As demonstrated above this is a clear synonym of Crambe
Laubenfels, 1936a), with Crambe crambe likewise is based on crambe, and thus Tetranthella is a junior synonym of Crambe.
Porifera· Demospongiae • Poecilosclerida • Myxillina • Crambeidae 551

Fig. 3. Discorhabdella incrustans Dendy, 1924, SEM images of spicules of type. A, ectosomal subtylostyle. B, choanosomal tylostyle with warty head.
C, mace-shaped pseudastrose tylostyle. D, polydentate isochela (photos M. Maldonado).

DISCORHABDELLA DENDY, 1924 Description. Thinly encrusting, approx. 1.5 mm thick, with
spined surface. Skeletal architecture hymedesmioid (Fig. 2B).
Synonymy Main styles huge, singly erect on the substrate, with lumpy heads,
but otherwise smooth; distance between them ca. 100-150 fLm.
Discorhabdella Dendy, 1924: 376. Cionanchora de Base of huge styles surrounded by masses of pseudastrose
Laubenfels, 1936a: 108. tylostyles echinating the substrate. Smooth subtylostyles are
arranged in bundles with heads downwards around the shaft of the
Type species main styles. Microscleres zoned, chelae at the periphery, spined
microrhabds deeper in the sponge. Spicules (Figs 2B, 3).
Discorhabdella incrustans Dendy, 1924: 376 (by monotypy). Ectosomal subtylostyles (Fig. 3A), smooth, with clearly swollen
heads, 357-(496.8)-592 X 10-(12.6)-15 fLm; choanosomal (tylo-)
Definition styles (Fig. 3B) with irregular swollen lumpy heads, mostly broken
in the slides, so length is tentative, 900-1700 X 28-(43.1)-61 fLm;
Crambeidae with hymedesmioid skeleton of choanosomal club-shaped ('mace' -shaped) pseudastrose spicules (Fig. 3C), with
styles with swollen lumpy bases and tuberculate club-shaped a swollen crown covered with spines, a short smooth middle part
pseudoastrose tylostyles. and a swollen and spined end, length 36-(45.4)-53 fLm and width
of the crown 32-(37.3)-43 fLm; anchorate-unguiferate isochelae
Diagnosis (Fig. 3D) with up to 7 long well-developed alae, occasionally with
reduced alae, 33-(44.0)-51 fLm; spined microxeas (Fig. 2B) with
Smooth ectosomal subtylostyles, long choanosomal styles two swollen and spined lumps at a short distance from either end,
with swollen lumpy bases and tuberculate club-shaped pseudoas- 26-(31.4)-34 fLm. Distribution and ecology. Off Three Kings
trose or heavily spined tylostyles forming erect hymedesmioid Islands, N New Zealand, dredged from 180 m depth.
skeleton; microscleres anchorate unguiferous isochelae and may Remarks. Boury-Esnault et al. (1992a) retained this genus
include spined microxeas with two lumpy swellings or sigma-like in the family Hymedesmiidae. Maldonado & Uriz (1996) pointed
spicules. Four species from allover the world. out the similarities between Discorhabdella and Crambe, but did
not assign either genus to a definite family. Similarity with the
Previous reviews spicule complement of Crambe (shared morphologies of chelae,
spined microxeas and pseudastrose spicules) is so great that
Dendy (1924), Boury-Esnault et al. (1992a), Maldonado & the mere sharing of hymedesmioid skeletal architecture with
Uriz (1996), Maldonado et al. (2001). Hymedesmia is insufficient to follow Boury-Esnault et al. (1992a).
Discorhabdella incrustans falls naturally to the family Crambeidae
Description of type species on account of ectosomal subtylostyles, tylostylote choanosomal
spicules and anchorate chelae. The pseudastrose styles have been
Discorhabdella incrustans Dendy, 1924 (Figs 2B, 3A-D). interpreted as homologous to echinating acanthostyles of
Synonymy. Discorhabdella incrustans Dendy, 1924: 376, Hymedesmiidae and other Myxillina families by Boury-Esnault
pI. XV figs 34-38; Boury-Esnault et al., 1992a: 5; Uriz & et al. (1992a). However, they share similarities to pseudastrose
Maldonado, 1995: 3, figs Ie, 2d, 5a-b, 6b-c; Maldonado & Uriz, desma-like spicules in Crambe and they have peculiar morphology
1996: 373, figs 6E, 7D, 8C, 9C, lOF-G. with spines confined to both swollen ends. Maldonado & Uriz
Material examined. Holotype (slides): BMNH 1923. (1996) demonstrated the spines are unlike those of normal acan-
10.1.157 - 'Terra Nova' Expedition, RN.xLVII-3, Three Kings thostyles in having an axial canal (thus are 'polyaxial'). It is possi-
Island, New Zealand, 180m depth. ble that these spicules derive ultimately from choanosomal
552 Porifera· Demospongiae • Poecilosclerida • Myxillina • Crambeidae

Fig.4. LithocheZa conica Burton (1929b). A-B, photos of habit ofBMNH lectotype (A) and paraleetotype (B) (scale 1 em). C-H, SEM photos of spicules
of lectotype (scales: 10 j.Lm, except G left, 1 j.Lm).

tylostyles. Fossil spicules described from the Oligocene of New Type species
Zealand (Hinde & Holmes, 1892) may belong to this species.
The type species of the genus Cionanchora de Laubenfels, Lithochela conica Burton, 1929b: 9 (by monotypy).
1936a: 108, Hymeraphia tuberosocapitata Topsent, 1890b: 68 (by
original designation; full description and illustration in Topsent, Definition
1892a: 113, pI. XI fig. 6, here reproduced as Fig. 2C) is a clear
Discorhabdella. A microscopic slide of the holotype, which is Crambeidae with desmas forming the main skeleton,
lodged in MOM, was examined in the Paris Museum, MNHN D.T. and possessing robust ectosomal subtylostyles and unguiferate
938, labeled 'Camp. Hirondelle 1888, No. 65, 159', as well as a chelae.
slide of a topotypical specimen MNHN D.T. 939, labeled 'PA 60
1895" (recorded in Topsent, 1904b), both from deep water, 550--736 Diagnosis
m, off the Azores. The skeletal architecture of H. tuberosocapitata
is hymedesmioid with single large tylostyles, about 650 f.lm long Thin-walled, stalked hollow tubes with desma-reticulation
with tuberose heads, about 28 f.lm in diameter, erect on the forming the support of the tube wall. Ectosomal brushes of robust
substrate, surrounded by small tuberose acanthotylostyles of about styles. Scattered isochelae with long thin teeth. Spined microxeas
130 f.lm long (interpreted here as homologous to the pseudastrose present. Only a single species known so far.
spicules of D. incrustans). Ectosomal subtylostyles are robust, with
elliptical heads and short points, length averaging 330 f.lm. Description of type species
Anchorate isochelae with prominent fimbriae, 4-5 teeth, averaging
25 f.lm. No spined microxeas. For SEM images and light micro- Lithochela conica Burton, 1929b (Figs 2D, 4A-H).
graphs of this species see Maldonado & Uriz (1996) and Hooper Synonymy. Lithochela conica Burton, 1929b: 9, pI. I
(1996a), the latter author erroneously referring the taxon to figs 9-10, pI. II figs 11-19.
Microcionidae as a synonym of Clathria (Microciona). Material examined. Lectotype (designation herein): BMNH
Further species were described by Boury-Esnault et al. (1992a) 1926.4.14.103. Paralectotype: BMNH 1926.9.3.79 (3 slides RN
and Maldonado et al. (2001). Discorhabdella hindei Boury-Esnault 169 and RN 204).
et al. (1992a) from 600m in the Mediterranean; is similar to Description. Shaped like a carrot, fistular-tubular, stalked
D. incrustans but possesses sigma-like microscleres in addition to (Fig. 4A-B), up to 5 cm high, I cm diameter. Flattened at the apex
the usual unguiferate anchorate chelae; like D. tuberosocapitata, it with a single central papillate oscule, 1 mm in diameter. Inside the
lacks spined microxeas. The variability in presence of spined central core is hollow, surrounded by a thick wall. Consistency
microrhabds and sigma-like microscleres in this genus parallels that fragile, easily damaged. Colour brownish yellow. Skeleton (Fig. 2D)
found in the genus Monanchora (cf. Van Soest et al., 1996a). of the surface a tangential feltwork of styles. Inside a reticulation of
styles and larger and smaller desmas form the skeleton of the thick
wall. The reticulation is made up of longitudinal tracts of styles
UTHOCHELA BURTON, 1929 connected by single or bundles of elongate desmas. Microscleres
extremely abundant along the inner wall. Spicules, styles of the
Synonymy ectosome (Figs 2D, 4C) and the choanosomal tracts (Figs 2D, 4D),
curved, sometimes irregular, 360--450 X 22 f.lm; desmas, probably in
Lithochela Burton, 1929b: 9. two categories, the larger ones (Figs 2D, 4E) elongated, rhabd-like
Porifera· Demospongiae • Poecilosclerida • Myxillina • Crambeidae 553

Fig. 5. A-E, Monanchora clathrata Carter, 1883c. A, habitus of BMNH holotype (scale 1 cm). B-E, SEM photos of spicules of the holotype
(scales: 10 j.Lm, except D left, 1 j.Lm). F-G, Monanchora enigmatica (Burton & Rao, 1928 as Ectyobatzella). F, SEM photos of ectosomal spicules made
from the holotype. G, ditto of choanosomal spicule.

with simple flat extensions at the end which connect with the tracts Dermition
of styles, 270-360 X 36 /-Lm, and the smaller branching and
knobbed (Figs 2D, 4F), cladomes 180-200 /-Lm, thickness of rhabds Crambeidae without pseudoastrose spicules or desmas;
20/-Lm; microscleres are unguiferous isochelae (Figs 2D, 4H), microscleres spatulate or unguiferate anchorate isochelae (may be
23-40/-Lm, and spined microxeas (Figs 2D, 4G), often with a cen- absent), reduced sigmoid chelae (may be absent), and spined
tral 'tyle', 55-100/-Lm (not reported by Burton). Distribution and microxeas (may be absent). In life, the surface has characteristic
ecology. Natal coast, South Africa, collected from green mud bot- white- or yellow-lined veinal channel pattern.
tom at 180 m depth.
Remarks. This is a unique species with no close relatives. Diagnosis
The skeletal architecture is highly correlated with the unusual fistu-
lar shape of the body. In this aspect Lithochela is quite different Encrusting to lobate or ramose. Surface smooth or extended into
from Crambe. However, some similarities may be present in the corrugate or spined projections. In life, superficial canals characteris-
sphaeroclone-type desmas of Crambe and the smaller desmas of tically swollen and with a lighter-coloured lining. Skeletal arrange-
Lithochela. There is no differentiation in style types, but the larger ment is simple: irregular plumose bundles of megascleres, which
desmas may be considered derived from the tylostyles found in the tend to form loose brushes at the surface without forming a definite
remaining crambeids, because as Burton already demonstrated, ectosomal skeleton. Spongin variably developed but may be consid-
they seem to grade into style-like spicules. Also, the spined erable. Spicule complement rather variable, both among and within
microxeas, reported for the first time, are similar to those of several species. Basically, the two size categories of megascleres characteris-
Crambe, Monanchora, and Discorhabdella species. tic for the family are present, but these may be indistinguishable in
some species or modified to strongyle-like spicules. Microsc1eres
may include a full complement, then consisting of two categories of
MONANCHORA CARTER, 1883 anchorate-unguiferate chelae and spined microxeas; frequently one
or more of these may be absent or reduced. In extreme cases only
Synonymy reduced and modified megascleres may be found (Ectyobatzella
form) and then assignment to Monanchora remains problematic.
Monanchora Carter, 1883c: 369. Amorphoclada Topsent,
1930. Ectyobatzella Burton & Rao, 1932: 332. Fasubera de Previous reviews
Laubenfels, 1936a: 119. Folitispa de Laubenfels, 1936a: 119.
[Okadaial de Laubenfels, 1936a: 120 (preocc.). Neofolitispa Van Soest (1984b, 1990), Bergquist & Fromont (1988), Van
Bergquist, 1965: 172. Soest et al. (1996a).

Type species Description of type species

Monanchora clathrata Carter, 1883c: 369 (by monotypy). Monanchora clathrata Carter, 1883c (Figs 5A-E, 6A).
554 Porifera· Demospongiae • Poecilosclerida • Myxillina • Crambeidae

I
50
,I.

----...

A I
10
B

On
I
50

I
"
c uu
00

c D
Fig. 6. A, Monanchora clathrata Carter (1 883c), drawing of skeleton and spicules made from a slide of the type. B, Monanchora laevissima
(Dendy, 1922b as Hymedesmia), type of Folitispa de Laubenfels (l936a), drawing of skeleton and spicules made from a slide of the type. C, Monanchora
unguiculata (Dendy, 1922b as Amphilectus), type of [Okadaial de Laubenfels (l936a), drawing of skeleton and spicules made from a slide of the type.
D, Monanchora enigmatica (Burton & Rao, 1932 as Ectyobatzelia), drawing of skeleton and spicules made from a slide of the type.

Synonymy. Monanchora clathrata Carter, 1883c: 369, anchorate-unguiferate chelae with 3-5 teeth, 26-(30.8)--36 f.Lm,
pI. XV fig. 10. reduced unguiferate chelae with spiky teeth in the same size range;
Material examined. Holotype: BMNH 1973.12.5.1 - microspined microxeas (Figs 5E, 6A), 36-57 X <0.5 f.Lm.
Fre-mantle, Bowerbank collection; two slides of the type with Distribution. Western Australia.
same number. Remarks. The similarity with Crambe is strong, including
Description (based on a beach-worn specimen, partly after similar chemistry (e.g., Braekman et aI., 1996; cf. also Van Soest
Carter, 1883c). Massive, clathrous, crumb-of-bread texture et al., 1996a), but the two genera differ in the presence or absence
(Fig. SA). Size 6 X 4 X 4.5 cm. Colour 'tawny'. Skeleton (Fig. SA), of desma-like spicules. A dozen species have been described. Some
a reticulation of cored spongin fibres and interconnecting spicules. species of the genus are notorious for frequently lacking a full
Fibres up to 150 f.Lm in diameter, cored by up to 4-5 spicules; inter- complement of spicules. For example, in Monanchora arbuscula
connecting spicules 1-3. Spicules (Figs 5A-E, 6A). subtylostyles, (Duchassaing & Michelotti, 1864) the following combinations of
248-(268.6)--297 X 4-(5.2)--7 f.Lm; tylostyles, robust, smooth, with shape and spiculation have been found: ramose, with conulose sur-
faintly developed tyle, 330--(343.4)--362 X 10--(12.8)-14 f.Lm; face, with a full complement of megascleres and microscleres,
Porifera· Demospongiae • Poecilosclerida • Myxillina • Crambeidae 555

including spined microxeas; ramose, with conulose surface, with similar, 29-(31.3)-34f.Lm and 15-(18.4)-22f.Lm. Further species
subtylostyles, thicker styles, reduced sigma-like chelae; ramose, assigned to [Okadaia] by de Laubenfels were Esperiopsis
with conulose surface, with anisostrongyles and thicker strongyles; viridis Kieschnick (1898: 59), E. rigida Lambe (1892: 60),
thickly encrusting, with conulose surface, with full complement E. vancouveri Lambe (1892: 68), and E. quatsinoensis Lambe
of spicUles; thinly encrusting, with smooth surface, with subty- (1892: 69). [Okadaia] de Laubenfels was found to be preoccupied
lostyles, styles, reduced sigmoid chelae, and anchorate chelae; by Okadaia Baba (1930), and de Laubenfels (1949a: 15) replaced
thinly encrusting with smooth surface, with subtylostyles, styles it by Neoesperiopsis; however, in the same publication he named as
and reduced sigmoid chelae; and, thinly encrusting with smooth the type of Neoesperiopsis, N. deichmannae, a species unrelated
surface with subtylostyles and thicker styles, no microscleres. to Amphilectus unguiculatus and from descriptions and studies of
The genus Ectyobatzella Burton & Rao, 1932 was erected for the type material by Hartman (1958: 45) a probable synonym
Indian Ocean Ectyobatzella enigmatica Burton & Rao (1932: 332) of Isodictya palmata (suborder Mycalina, family Isodictyidae).
(by monotypy), a lobate sponge with irregularly conulose to digi- The genus Neofolitispa Bergquist (1965: 172), with type
tate surface. A microscopic slide of the holotype from the Indian species the Philippine sponge Monanchora dianchora de Laubenfels
Museum is kept in the Natural History Museum, London, BMNH (1935a) (by original designation), was erected as a 'new name' for
1934.11.24.100. This contains a cross section (cf. Fig. 6D) show- the preoccupied Okadaia (cf. above), but since the two do not have
ing a plumose-dendritic skeleton of bundles of thin strongyles the same type species, Neofolitispa effectively became a new genus,
enclosed partly in spongin (increasing towards the base of the not merely a new name. Bergquist was reluctant to assign
sponge). The bundles fan out towards the surface without forming A. unguiculatus to Neofolitispa without having studied its type
a special ectosomal skeleton. Thicker and shorter styles are scat- specimen. She distinguished Neofolitispa from Monanchora
tered singly in the interior and echinate the bundles of strongyles. and Folitispa by the lack of differentiation in megascleres and the
Spicules, strongyles (Figs 5F, 6D), straight, occasionally styles possession of two categories of chelae instead of one. In view of
(Figs SF, 6D) or with irregular ends, 162-(177.2)-189 X the above mentioned variability in spicule presence and form
1.5-(2.6)-3.5 f.Lm; (tylo-)styles (Figs 5G, 6D), smooth, with con- Neofolitispa is regarded as a synonym. The whereabouts of the type
stricted 'neck', 114-(128.9)-144 X 4-(5.6)-7 f.Lm; no microscle- specimen of M. dianchora is unknown, it is not in USNM (Ms K.
res. This species is assigned to Monanchora on the basis of the Smith, pers. comm.); it is a light orange-red, thickly encrusting,
variability observed in Caribbean Monanchora arbuscula, in which lamellate mass, surface irregular. Ectosomal spicules arranged tan-
the morphology of the megascleres and the presence of the gentially, choanosomal spicules in confusion. Megascleres
microscleres shows extremes from fully characteristic - including 'tylostyles', differentiated only in size, in the ectosomal region they
subtylostyles, tylostyles, anchorate chelae, reduced sigmatose measure approx. 283 X 4 f.Lm, in the choanosome 270-370 X
chelae and spined microrhabds - to largely reduced with thin 5-9 f.Lm; microscleres anchorate unguiferate chelae of 33 f.Lm and
strongyles as the only spicules (see Van Soest et al., 1996a). reduced unguiferate chelae with three minute teeth of 20 f.Lm. Folitispa
The genus Folitispa de Laubenfels (1936a: 119) was erected pingens de Laubenfels (1954: 159) is ajunior synonym of this species.
for Indian Ocean Hymedesmia laevissima Dendy (1922b: 81) (by The genus Fasubera de Laubenfels, 1936a: 119 was erected
original designation). Slides of the type were re-examined BMNH for type species Hymedesmia lipochela Dendy, 1922b: 82, pI. 6 fig. 3,
1931.1.7.69a ('Sealark', RN CXXV.6). This is a thinly encrusting pI. 15 fig. 2 (by original designation), from Cargados Carajos in
sponge with a transparent membrane showing irregular clusters of the Western Indian Ocean. Slides of the holotype (BMNH
pores. The choanosomal skeleton is hymedesmioid (Fig. 6B) with 1931.11.7.70) were reexamined. This is an irregularly lobate ('cau-
single large styles erect on the substrate (head down) and bundles liflower') sponge, with few small oscules. The skeleton consists of
of subtylostyles carrying the ectosome. Spicules are ectosomal lax spicule bundles and a lot of coarse sand. Dendy's contention
subtylostyles with pointed or telescoped ends, which are occasion- that the sand is organized as part of the skeleton cannot be con-
ally rugose or grooved, 240-(307.7)-364 X 4-(5.1)-7 f.Lm, robust firmed, but individual spicule tracts arise from these concentrations
choanosomal tylostyles, 288-(473.5)-615 X 15-(21.4)-27j.lm; of sand and they are 'echinated' by the spicules. At the surface
microscleres are spatulate achorate chelae and reduced chelae the spicule bundles fan out and form characteristic bouquets.
of similar length, not interpreted as a separate category: Megascleres subtylostyles 280-396 X 2-7 f.Lm, thick (tylo-)styles
26-(29.1)-31 f.Lm. It is a clear Monanchora lacking spined with swollen heads, 306-430 X 12-18 f.Lm, and microscleres (not
microrhabds; the hymedesmioid architecture is not sufficient described by Dendy) thin spined microxeas, up to 30-42 X 0.5 f.Lm.
grounds for retaining the genus as separate from Monanchora. Despite the lack of chelae, this species appears to be a clear
Hooper (1996a: 45) erroneously included this nominal genus into Monanchora, based on the characteristic spiculation, including
synonymy with Clathria (Microciona). the spined microxeas. Accordingly, Fasubera is assigned to
The genus [Okadaia] de Laubenfels (1936a: 120) was erected Monanchora as a junior synonym. Amorphoclada Topsent, 1930
for type species Amphilectus unguiculatus Dendy (1922b: 58) (by was erected (designation herein) for type species Chondrocladia
original designation). Examined material comprised two micro- alaskensis Lambe, 1894: 119, together with a second species
scopic slides, BMNH 1921.11.7.47a, Indian Ocean, '''Sealark' C. pulchra Lambe, 1894. This is a junior syndrome of Monanchora.
sponges, #RN CXID.l 0". This thinly encrusting species shows
the typical simple plumose skeleton of bundles of subtylostyles
(Fig. 6C), length 1-2mm, thickness ca. 15 spicules in cross section ACKNOWLEDGEMENTS
(50-250 f.Lm), at distances of 200-350 f.Lm. No ectosomal specializa-
tion. Megascleres not distinguishable into two distinct categories, Dr Manuel Maldonado (CEAB, Blanes) kindly provided SEM
subtylostyles with variable sharp or blunt ends grading into each images of the spicules of Discorhabdella incrustans (figs 3A-D).
other, 284-(316.4)-350 X 3-(4)-10 f.Lm. Microscleres reduced Also, his comments and remarks on this chapter are gratefully
unguiferate chelae in two size categories which are morphologically acknowledged.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Crellidae Dendy, 1922

Rob W.M. Van Soest

Zoological Museum, University of Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, Netherlands. (soest@science.uva.nl)

Crellidae Dendy (Demospongiae, Poecilosclerida) are characterized by a tangential ectosomal crust made up of acanthostyles or acan-
thoxeas. Prior to the present revision, Crellidae were a substantial group with 22 nominal genera. However, structural diversity appeared
to be low which is reflected in only five genera and four subgenera considered valid. The crellid choaonosomal skeleton is hymedesmioid
or plumose, with choanosomal megascleres consisting of peripheral tomotes and basal acanthostyles. The latter may be absent.
Echinating acanthostyles are present in several species. Microscleres are commonplace arcuate isochelae (one species excepted) and sig-
mas, either of which may be absent. The surface of most species features prominent areolated porefields, shared with the sisterfamily
Hymedesmiidae.
Keywords: Porifera; Demospongiae; Poecilosclerida; Myxillina; Crellidae; Anisocrella; Crella; Crella (Crella); Crella (Pytheas); Crella
(Yvesia); Crella (Grayella); Crellastrina; Crellomima; Spirorhabdia.

DEFINITION, DIAGNOSIS, SCOPE on account of the easily recognized surface skeleton and early
attempts (e.g., by Topsent, 1892a, 1904b, 1928c) to differentiate
Synonymy genera sharing this feature. Crellidae are concentrated in cold-
temperate and deep-water habitats, but several species, e.g., Crella
Crelleae Dendy, 1922b: 92. Crellidae Hentschel, 1923: 406. cyathophora, are also found in tropical reef habitats. Distribution is
worldwide.
Definition

Myxillina with a tangential crust of spined ectosomal spicules Taxonomic remarks

(oxeas, anisoxeas or styles), and a choanosomal plumose skeleton
of smooth tomotes. A basal skeleton of acanthostyles erect on the Acanthose megascleres. The major synapomorphy for the
substrate may be present. Surface provided with areolated pore family is the ectosomal tangential crust of spined megascleres.
fields the walls of which are built by spicules arranged in parallel. These are usually acanthoxeas, but they may have one or both ends
Microscleres arcuate chelae, and occasionally sigmas (the latter so blunt and then resemble acanthostyles or acanthostrongyles.
far recorded in only two species). Through the frequently observed intermediate spicule forms such as
tapering blunt endings, often somewhat inequiended acanthoxeas,
Diagnosis etc., it is a safe assumption that these are all a single spicule type.
Genus proliferation. This is a small family with probably
Encrusting, massive, club-shaped and branching growth forms. only a single well-established genus, the other four being peculiar
Choanosomal skeleton regularly reticulate or plumo-reticulate, single or pairs of species. In the past, all possible combinations of
composed of bundles of smooth oxeas. Ectosomal skeleton with presence and absence of spicule types and forms have been used to
a thick crust of tangentially placed acanthostyles and/or acanthox- erect genera. Many of these overlapped in characters and thus it is
eas. Spicules also fortify the walls of elevated surface areolae. not likely that they are natural groups. Accordingly, these are
Acanthose spicules may also be embedded erect on basal spongin merged and we are left with only a few genera with Crella as the
(=echinating basal acanthostyles), and dispersed within the major genus. In addition, it is proposed to recognize some sub-
choanosome between the tracts of smooth diactines. True echina- generic units in Crella to facilitate classification and recognition of
tion of the tomote bundles has not been recorded so far. the large number of species.
Microscleres consist of arcuate isochelae, occasionally anisochelate, Family status. The family shares the surface areolae with
and sigmas. Hymedesmiidae and it may be argued that the two ought to be
recognized at the level of subfamilies of a larger family uniting
Scope the two. However, in view of the distinct surface skeletons,
including differential skeletal structure of the areolae (supported
23 nominal genera of which five genera are considered valid, by acanthose spicules in Crellidae, smooth spicules in
and four subgenera: Anisocrella, Crella, Crella (Crella), Crella Hymedesmiidae) the two may be readily told apart and any pro-
(Pytheas), Crella (Yvesia), Crella (Grayella), Crellastrina, posal to merge them would be premature without corroboratory
Crellomima, Spirorhabdia. evidence to do so.

History and biology

Previous reviews
The family was recognized as a formal unit for the first time
by Dendy (1922b), and this was followed by most later authors Hentschel, 1923, Topsent, 1928c, Levi, 1973.

556
Porifera· Demospongiae • PoeciioscIerida • Myxillina • Crellidae 557

KEY TO GENERA

(1) Chelae include peculiar, reduced anisochelae next to normal isochelae ............................................................................ Anisocrella
Only normal isochelae or chelae absent ............................................................................................................................................... 2
(2) Surface skeleton made up of strongly spined aster- or spiraster-like spicules which are modified acanthoxeas ................................ 3
No astrose acanthoxeas ........................................................................................................................................................................ 4
(3) Basal acanthostyles absent, stylote tomotes ...................................................................................................................... Crellastrina
Basal acanthostyles present, oxeote tomotes ................................................................................................................... Spirorhabdia
(4) Chelae normally arcuate with rounded alae or chelae absent ...................................................................................................... Crella
Chelae polydentate-anchorate (sharp-teethed) ................................................................................................................... Crellomima

ANISOCRELLA TOPSENT, 1927 categories, both entirely spined, up to 200 X 27/-Lm and 50-70 /-Lm
(the latter are from Vacelet, 1969). Microscleres: arcuate isochelae,
Synonymy small but of the usual shape, 13-15/-Lm; reduced, often
inequiended chelae called 'anisancres arquees' , 15-17 /-Lm.
Anisocrella Topsent, 1927b: 11. Distribution and ecology. Azores, Western Mediterranean, deep
water, 500-2460m depth.
Type species Remarks. This is a mono specific genus, based primarily on
the peculiar chelae setting it apart from mainstream Crella. If no
Anisocrella hymedesmina Topsent, 1927b: 11 (by monotypy). further similar species will be found, it may be better to consider it
a subgenus in the future. The spicule sizes quoted by Vacelet
Definition (1969) and Boury-Esnault et al. (1994b) differ somewhat from
Topsent's specimen (acanthoxeas only up to 115 /-Lm, tomotes up to
Crel\idae with hymedesmioid architecture; next to normal 222 X 5 /-Lm; anisancres up to 22 /-Lm), but it is likely that this con-
isochelae there are peculiar reduced anisochelae. stitutes merely infraspecific variation. The drawings of the chelae
of Boury-Esnault et al., 1994b remind more of palmate chelae
Diagnosis than of arcuate chelae, but they call them 'arquees' nevertheless;
also, although no acanthoxeas are drawn, these are mentioned in
Encrusting growth form; ectosomal skeleton composed of tan- the description. The hymedesmioid condition of the skeleton in
gential layer of acanthoxeas; choanosomal skeleton composed of combination with the obvious Crellidae ectosome again demon-
smooth diactinal spicules in bundles and an erect basal layer of acan- strates the likely polyphyletic nature of the hymedesmioid skeletal
thostyles; microscIeres are arcuate isochelae and reduced anisochelae. structure.

Previous review

Boury-Esnault et al. (1994b). CRELLA GRAY, 1867

Description of type species Synonymy

Anisocrella hymedesmina Topsent, 1927b (Fig. lA). Crella Gray, 1867a. Grayella Carter, 1869b: 190. Pytheas
Synonymy. Anisocrella hymedesmina Topsent, 1927b: 11; Topsent, 1890b: 31. Yvesia Topsent, 1892a: 103. Pseudoclathria
Topsent, 1928c: 234, pI. VIII fig. 5; Vacelet, 1969: 198, fig. 35; Dendy, 1897: 258. Kowalevskyella Swartschewsky, 1905: 40.
Boury-Esnault et al., 1994b: 116, fig. 88. Crellina Hentschel, 1914: 110. Grella de Laubenfels, 1936a: 64.
Material examined. Holotype (not seen): MOM. Tisrone de Laubenfels, 1936a: 78. Yvesiella de Laubenfels, 1936a:
Schizotypes: MNHN DT 1172 - 3 slides (two sections, one spicule 78. Yvesiorbas de Laubenfels, 1936a: 65. Crellinspira de
mount) made from the holotype, labeled "St. 1349, 1902". Other Laubenfels, 1936a: 79. Pytheilla de Laubenfels, 1936a: 64.
material. MNHN DT 1173 - 1 slide from stn. 1420. Ramosichela de Laubenfels, 1950b: 18. Naniupi de Laubenfels,
Description (mostly based on Topsent, 1928c). Thin 1950b: 19. Crelloxea Hechtel, 1983: 70.
smooth crusts on deep water corals, 2-3 cm2 (Vacelet, 1969).
Boury-Esnault et al., 1994b report areolated porefields. Greyish Type species
in alcohol (Topsent, 1928c; Vacelet, 1969), maroon in alcohol
(Boury-Esnault et al., 1994b). Skeleton. Hymedesmioid, with Cribrella elegans Schmidt, 1862 (by original designation).
acanthostyles erect on the substrate; bundles of tomotes connect
the basal skeleton with the surface and carry the tangential skeleton Definition
of acanthoxeas. Spicules (Fig. lA). Ectosomal acanthoxeas, robust,
curved, heavily spined, 165-230 X 8-10 /-Lm (Vacelet, 1969 gives Crellidae with ectosomal acanthoxeas or acanthostyles,
only 80-115 /-Lm); tomotes smooth, faintly polytylote, inequiended, oxeote, strongylote or stylote tomotes; basal acanthostyles may be
mucronate or rounded, 130-145 X 2/-Lm; acanthostyles in two size present; microscleres arcuate isochelae (may be absent).
558 Porifera· Demospongiae • Poecilosclerida • Myxillina • Crellidae

D
B c
"

Fig.1. A, Anisocrella hymedesmina Topsent, 1927b, drawing of spicules reproduced from Topsent (1928c, pI. VIII fig. 5) (sizes see text). B-D, Crella
(Crella) elegans (Schmidt, 1862 as Cribrella). B, drawing of shape and acanthoxea reproduced from Schmidt (1862, pI. VII fig. 3) (scales see text).
e, drawing of spicules copied from Topsent (1925c: fig. 20) (sizes see text). D, drawing of surface areola and spicules reproduced from Bibiloni, 1981,
fig. 7. E, Crella (Crella) tubifex (Hentschel, 1914 as Crellina), drawing of shape and spicules reproduced from his pI. IV fig. 12, pI. VII fig. 14 (scales see
text). F, Crella (Crella) gracilis (Swartschewsky, 1905 as Kowalevskyella) (=Crella elegans), drawing of peripheral skeleton and spicules, reproduced
from Swartchewsky, 1905: pI. 7 fig. 2) (scales see text).

Diagnosis Taxonomic remarks

Encrusting, lobate or pedunculate; ectosomal skeleton com- The type species was originally described in the genus
posed of a tangential layer of acanthoxeas or acanthostyles; [Cribrellal Schmidt, 1862, but this is preoccupied by Cribrella
choanosomal skeleton composed of smooth tomotes or strongyles Agassiz, 1835. [Cribrellal sensu Schmidt, 1862 is a senior preoccu-
arranged in tracts which may be echinated by acanthostyles, or in pied synonym of Hamigera (subsequent type species designation by
encrusting forms there may be an erect basal layer of acan- Desqueyroux-Faundez & Stone, 1992). A slight complication with
thostyles; microscleres may include arcuate isochelae, sometimes the name Crella may arise from the fact that the only extant material
absent, and occasionally sigmas. More than 50 species. labeled as Cribrella elegans in the LMJG collections is not that
species (cf. below). Since there is no locality data with the LMJG
Previous review specimens, and Schmidt's plate and drawing are reasonably obvious,
it is almost certain that the LMJG material is not the original
Topsent (1928c). specimen from the Zara Canal. A neotype designation can solve this
Porifera· Demospongiae • Poecilosclerida • Myxillina • Crellidae 559

problem. As indicated above, there are 15 generic names available characters, which were not described by the original authors or were
covering the present concept of Crella. For reasons so far unclear this incorrectly described (genera Pytheilla and Crellinspira). Compared
proliferation of genera, distinguished only on minor features as the to other families, structural diversity of Crellidae is quite limited and
shape of the tomotes and ectosomal acanthose megascleres, presence the recognition of so many generic names for such trivial differences
or absence of echinating acanthostyles and the presence or absence between individual species is unprecedented. For convenience sake,
of chelae, exceeds the normal diversity of such artificial 'genera' dis- four of the generic names, Crella, Grayella, Pytheas, and Yvesia
tinguished in related Poecilosclerids. As usual, de Laubenfels takes are here regarded as subgenera to facilitate identification of the
care of more than half the number of synonyms. To add to the confu- large numbers of species. In a phylogenetic sense these subgenera
sion, two type species of de Laubenfels's genera were attributed are artificial, they do not comprise natural groups.

Key to subgenera of Crella

(1) Chelae present ..................................................................................................................................................................................... 2

No chelae ............................................................................................................................................................................................. 3
(2) Basal echinating acanthostyles ................................................................................................................................... Crella (Pytheas)
Acanthostyles either absent, or of a single kind concentrated at the surface; no basal echinating acanthostyles ......... Crella (Yvesia)
(3) Basal acanthostyles echinating the substrate present .................................................................................................... Crella (Crella)
No echinating acanthostyles ...................................................................................................................................... Crella (Grayella)

SUBGENUS CRELLA GRAY, 1867 the original material, since Schmidt's drawing of the original speci-
men clearly depicts an acanthoxea, which are absent in the Graz and
Synonymy Strasbourg Museum specimens. A neotype should be assigned to cre-
ate stability in this group with so many generic names. Prof. Levi (in
Crella Gray, 1867a. Kowalevskyella Swartschewsky, 1905: 40. litteris) proposed to designate as the neotype specimen the material
Crellina Hentschel, 1914: 110. described by Topsent (1925c) from the Bay of Naples, now housed in
the Strasbourg Museum, with slides and a fragment in MNHN regis-
Type species tered as MNHN DC11190. This collective material is here designated
as the neotype.
Cribrella elegans Schmidt, 1862 (by original designation). Swartschewsky (1905: 40) erected the genus Kowalevskyella
for type species (by monotypy) Kowalevskyella gracilis
Definition Swartschewsky, 1905: 40, pI. II fig. 7, pI. VII fig. 2 (the latter figure
is here reproduced as Fig. IF). This specimen has all the characters
No chelae; basal or echinating acanthostyles present. of Crella elegans, including red colour, surface areolae, lack of
microscleres, and possession of oxeote tomotes, acanthoxeas and
Description of type species acanthostyles in the appropriate size. It is proposed here to syn-
onymize gracilis with elegans, and thus Kowalevskyella becomes
Crella (Crella) elegans (Schmidt, 1862) (Fig. lB-D). an objective synonym of Crella (Crella).
Synonymy. Cribrella elegans Schmidt, 1862: 70, pI. VII fig. 3 Hentschel (1914: 110) expressly erected a genus Crellina for its
(Schmidt's text gives fig.1 but this represents Tethya). Kowalevskyella possession of echinating acanthostyles in combination with the lack
gracilis Swartschewsky, 1905: 40, 53, pI. II fig. 7, pI. VII fig. 2. of chelae, because Schmidt's original description did not mention the
Material examined. Holotype: No type material could be acanthostyles. Crellina with type species (by monotypy) c. tubifex
located. Neotype (designated herein): MNHN DCl1190 - Bay of Hentschel (1914: 110, pI. IV fig. 12, pI. VII fig. 14) clearly falls under
Naples (see Topsent, 1925c). Other material. ZMA POR 54, the definition of Crella (Crella), although its type species (Fig. IE)
Banyuls, 10m, colI. G. Kleeton, VIII-1962. appears only distantly related to C. elegans. The type (not re-
Description. (largely based on Topsent, 1925c) Encrusting, examined) is described as massive, with elongated tubes, which are
smooth, colour pink. Surface provided with numerous red- possibly homologous to the surface areolae of other species. The
coloured areolae (Fig. IB). Consistency firm. Skeleton. Ectosomal ectosomal crust is formed by acanthostrongyles of 440-560 X
crust of densely intercrossing acanthoxeas to form a firm leathery 12.5-15jLm, the choanosomal skeleton consists of ill-defined bundles
crust; choanosomal skeleton consisting of thick bundles of of tomotes of 472-650 X 7-8jLm and the acanthostyles are erect on
tomotes; basal acanthostyles echinate the substrate (not mentioned the substrate, 176-392 X 10011jLm. It should be noted that the mate-
by Schmidt). Spicules: ectosomal acanthoxeas, densely spined all rial identified as Crellina tubifex by Burton, BMNH 1928.11.15.225
over: 70-95 X 3-15jLm; smooth, faintly polytylote, oxeote was found to belong to Acanthorhabdus fragilis (family Acarnidae).
tomotes, sharply pointed, 210-250 X 3-5jLm; acanthostyles in
a single category: 110-150 X 5-10 jLm. Distribution and ecology: SUBGENUS GRAYELLA CARTER, 1869
Mediterranean, NW Spain; shallow-water.
Remarks. The alleged type material in the Graz Museum, Synonymy
LMJG 15526 (no locality data) was reexamined and discovered to
be a specimen of Phorbas ficititius. Prof. Levi (in litteris) confirms Grayella Carter, 1869b: 190. Pseudoclathria Dendy, 1897:
that the Schmidt material in the Strasbourg Museum labeled as 258. Grella de Laubenfels, 1936a: 64. Yvesiorbas de Laubenfels,
Cribrella elegans likewise concerns Phorbas fictitius. This cannot be 1936a: 65. Crelloxea Hechtel, 1989: 70.
560 Porifera· Demospongiae • Poecilosclerida • Myxillina • Crellidae

1
F ij ~
6U
G
H

/!: ":::I:~

Fig. 2. Crella (Grayella) species. A~, Crella (Grayella) cyathophora (Carter, 1881 as Grayella). A-B, SEM photos of spicules made from the holotype
(scale 1 !Lm). C, drawing of skeleton and spicules made from a slide of the holotype. D-E, Crella (Grayel/a) compressa (Carter, l886c as Halichondria),
type of Pseudoclathria, SEM photos of spicules made from the holotype (scale I fLm) . F, Crella (Grayel/a) pulvinar (Schmidt, 1868 as Myxilla), type of
Yvesiorbas, drawing of spicules copied from Topsent (1925c: fig. 22 as Crella mollior) (sizes see text). G, Crel/a (Grayel/a) camosa (Topsent, 1904b, as
Yvesia), drawing of spicules reproduced from his pI. XV fig. 19 (sizes see text). H, Crella (Grayella) spinosa (Hechtel, 1983 as Crelloxea), drawing of
spicules reproduced from his fig. 11 (sizes see text).

Type species diameter (containing 6--20 spicules). Spicules (Fig. 2A-B).

Ectosomal acanthoxeas, thin, finely spined, frequently faintly cen-
Grayella cyathophora Carter, 1869b: 190 (by monotypy). trotylote, 96--(104.5)-117 X 2-(2.6)-3 ""m; choanosomal tomotes
are strongyles, occasionally stylote, 279-(291.8)-306 X 2.5-(3.8)-
Definition 5 ""m. Distribution and ecology: Red Sea; common in the Indian
Ocean; shallow-water reefs.
No chelae, no basal or echinating acanthostyles. Remarks. The species differs from Crella elegans in lacking
basal acanthostyles, a common enough phenomenon, not worthy
Description of type species of generic distinction. Topsent (1892a: 103) declared Grayella
cyathophora a member of his genus Yvesia without acknowledging
Crella (Grayella) cyathophora (Carter, 1869b) (Fig. 2A-C). the fact that Grayella preceded Yvesia by 23 years. Later (l928c),
Synonymy. Grayella cyathophora Carter, 1869b: 190, pI. VII he recognized Grayella as separate from a restricted Yvesia.
figs 1-10; Yvesia cyathophora; Topsent, 1892a: 103; Crella The genus Pseudoclathria Dendy (1897: 258) was erected (by
cyathophora; Dendy, 1922b: 95 original designation) for type species Halichondria compressa
Material examined. Holotype (slide): BMNH 1869.6.25.48- Carter (1886c: 450), redescribed by Dendy (1897) on the basis of
Gulf of Suez. Carter's holotype (BMNH 1886.12.15.9) and additional speci-
Description. Massively encrusting on rocks; type specimen mens. A further specimen BMNH 1886.12.15.392 also labeled as
3.5 cm across, 1.25 cm thick. Surface smooth with numerous sub- "Halichondria compressa" belongs to Hymeniacidon. Carter's type
circular, elevated areolae. Oscules likewise elevated. The type slide is an elaborate flabelliform sponge, with a broad margin bearing
contained a surface peel with three areolae and cross sections. The oscules. The surface skeleton is a crust of intercrossing acan-
areolae are 1500--1800 /-Lm in diameter are slightly raised above the thostyles (Fig. 2B), 108-122 X 7-9 /-Lm, which are also scattered in
surface and their walls are supported by perpendicular bundles of the interior. The choanosomal skeleton is a system of long, infre-
4--5 tomotes. Skeleton (Fig. 2C). The surface between the areolae quently branching spongin fibres with a core of thin, straight, sub-
has a tangential crust of intercrossing acanthoxeas. The choanoso- tylostylote tomotes (Fig. 2D), 195-225 X 4--5 /-Lm, surrounded
mal skeleton consists of dendritic tracts of tomotes, 20--70/-Lm in by a bark of acanthostyles similar to the ectosomal acanthostyles
Porifera· Demospongiae • Poecilosclerida • Myxillina • Crellidae 561

(Fig. 2E). No echinating acanthostyles. By its elaborate shape, and Description (based on Topsent, 1892a, 1904b). Encrusting,
by the intrusion of acanthostyles into the choanosomal bundles of soft, smooth; easily detachable 'skin'. Colour black (Fig. 3A).
tomotes, the species is an unusual Crella (Grayella), but its charac- Skeleton. The main skeleton consists of thick columns of tomotes;
ters fall within the definition of the subgenus. these carry the tangential ectosome of acanthostyles; many acan-
The genus Grella was erected by de Laubenfels (l936a: 64) thostyles distributed singly in the interior. Spicules (Fig. 3A).
for the type species (original designation) Yvesia camosa Topsent Ectosomal acanthostyles, long, thin, finely spined, 375 JJ.m;
(l904b: 198). This is a thickly encrusting Azoran deep water tomotes tylote, with distinct tyles, smooth, 375 JJ.m; basal
sponge (100-1250m) with a smooth dark coloured surface and acanthostyles in the same size as the ectosomal ones, but with
a greyish interior. A spicule slide made of the Monaco Museum the lower half of the shaft smooth; microscleres arcuate isochelae,
holotype was examined in MNHN, labeled D.T. 1031, '68 P.A., 30 JJ.m. Distribution and ecology: Azores; deep water, 736 m depth,
1895' . The slide contains fusiform ectosomal acanthoxeas on coral.
(Fig. 2G): 160-180 X 5-6 JJ.m and smooth polytylote choanosomal Remarks. The slide of the holotype examined did not con-
anisotomotes (Fig. 2G) with mucronate apices: 600 X 10 JJ.m. The tain an obvious second category of acanthostyles, but Topsent
choanosomal skeleton is described by Topsent as consisting reports a second category of the same size as the ectosomal ones,
of straight parallel tracts of tomotes up to 100 JJ.m in diameter but with the lower half smooth. These are assumed to be a different
carrying a surface crusts of acanthoxeas; these are also scattered category from the ectosomal spicules, echinating the substrate.
singly between the choanosomal tracts. Later (l928c), Topsent The genus Crellinspira de Laubenfels (1936a: 79) was erected
assigned specimens with echinating acanthostyles to this species, for type species (original designation) Crella donsi Burton (l931c:
and this would cause it to be transferred to Crella. 136, fig. 1), a 1 mm thin encrustation collected under a stone at the
The genus Yvesiorbas de Laubenfels (1936a: 65) was erected low water mark at the Lofoten in northern Norway. The type is in
for type species (original designation) Myxilla pulvinar Schmidt the Trondheim Museum, but microscopical slides of the holotype
(1868: 14, pI. II fig. 14) from Algiers, redescribed by Topsent are in the Natural History Museum, London, BMNH
(1939). Type slide BMNH 1868.3.2.63, from Algiers, was exam- 1931.1O.28.30a and these were re-examined. The sponge has the
ined. It is a thinly encrusting crellid, with acanthoxeas of 50-80 X usual crust of acanthose megascleres leaving contracted areoles
1.5 JJ.m (Pulitzer-Finali, 1978), and strongylote tomotes of charged with microscleres; the choanosome has plumose bundles
290-410 X 2--4 JJ.m (isochelae recorded by Schmidt are foreign) of smooth tomotes and scattered single acanthostyles similar to
(Fig. 2F). This conforms to Grayella; however, Schmidt's type those of the ectosomal crust. At the base of the sponge there are
slide contained rare acanthostyles, so some uncertainty is apparent. long acanthostyles echinating the substrate (Fig. 3B). Spicules
The genus Crelloxea Hechtel, 1983 was erected for type species (Fig. 3B). Smooth oxeote tomotes, sometimes faintly polytylote,
(by monotypy) c. spinosa Hechtel (1983: 70) from Brazil. This is a with sharp or occasionally mucronate ends, 171-(204.0)-243 X
thinly encrusting sponge with surface crust of acanthoxeas (Fig. 2H) 2.5-(3.2)--4 JJ.m; ectosomal acanthostyles (acanthostyles II), thick-
and acanthostrongyles, 80-140 X 2-9 JJ.m, and choanosomal est some distance from the blunt end, fully spined,
oxea-shaped tomotes (Fig. 2E), 295--407 X 10-24 JJ.m, arranged in 69-(84.1)-98 X 3-(4.0)-5 JJ.m; long echinating acanthostyles,
uni- to paucispicular tracts. The unusual feature of the sponge is 153-(178.0)-210 X 5-(5.8)-7 JJ.m; arcuate isochelae, not reported
the fusiform oxea-shape of the tomotes. by Burton ('microscleres absent') but nevertheless definitely pres-
ent (dozens of similar chelae were observed in situ in the sections),
18-(20.1)-24 JJ.m. Thus, the type species of Crellinspira does not
SUBGENUS PYTHEAS TOPSENT, 1890 conform to the definition given by de Laubenfels. It clearly falls
into the synonymy of Pytheas. Some sigmas were also noted, but
Synonymy these were probably foreign.
The genus Ramosichela de Laubenfels (1950b: 18) was
Pytheas Topsent, 1890b: 31. Crellinspira de Laubenfels, erected for type species (by original designation) Grayella
1936a: 79. Ramosichela de Laubenfels, 1936a: 18. Naniupi de akraleitae Brl'lndsted (1932: 14, fig. 7). This is a thinly encrusting
Laubenfels, 1950b: 19. deep-water sponge (not re-examined) on hydroids. The ectosomal
skeleton is a crust of acanthostyles, around 100 X 4 JJ.m; the
Type species choanosomal skeleton consists of bundles of polytylote, stylote
tomotes (fig. 3C), 340-420 X fr-7 JJ.m; basal acanthostyles up to
Pytheas aterTopsent, 1890b (by monotypy). 280 X 8 JJ.m. The arcuate chelae are about 38 JJ.m. It conforms
closely to Pytheas.
Definition The genus Naniupi de Laubenfels, 1950b was erected for type
species (by monotypy) N. ula de Laubenfels (l950b: 19, fig. 11).
Crella with isochelae; basal or echinating acanthostyles present. The holotype USNM 22740 was re-examined. It is a thin encrusta-
tion. It has ectosomal acanthoxeas (Fig. 3D) of 110 X 4 JJ.m,
Description of type species choanosomal stylote tomotes of 190 X 4 JJ.m, echinating acan-
thostyles of 130 X 7 JJ.m and arcuate chelae of about 2 JJ.m. In addi-
Crella (Pytheas) atra(Topsent, 1890b) (Fig. 3A). tion it has some of the acanthoxeas curled in such a way that de
Synonymy. Pytheas ater Topsent, 1890b: 31; Topsent, Laubenfels deemed them sufficient reason for erecting a new
1892a: 110, pI. I fig. 1, pI. XI fig. 1. genus. They are uncommon and obviously derived from normal-
Material examined. Holotype: MOM (not seen). shaped acanthoxeas. The only difference with the type of Pytheas
Schizotype: MNHN DT. 996 (slide from the holotype) - labeled is the fact that it has ectosomal acanthoxeas instead of acan-
'Camp. Hirondelle, 1888 No. 65,1888:106'. thostyles.
562 Porifera' Demospongiae • Poecilosclerida • Myxillina • Crellidae

..
,

l(i
j B

) n y.

I I

I
~
I I ~
~

U C D

Fig. 3. Crella (Pytheas) species. A, Crella (Pytheas) atra (Topsent, 1890b as Pytheas), drawing of shape and spicules, reproduced from Topsent (1892a
pI. I fig. 1 and pI. XI fig. I) (scales see text). B, Crella (Pytheas) donsi Burton (1931), type of Crellinspira, drawing of skeleton and spicules made from a
slide of the holotype. C, Crella (Pytheas) akraleitae(BfPndsted, 1932 as Grayella), type of Ramosichela, drawing of spicules reproduced from his fig. 7
(sizes see text). D, Crella (Pytheas) ula (de Laubenfels, 1950b as Naniupi), drawing of spicules reproduced from his fig. 11 (sizes see text).

SUBGENUS YVESIA TOPSENT, 1892 Description of type species

Synonymy Crella (Yvesia) albula (Bowerbank, 1866) (Fig. 4A-D).

Synonymy. Halichondria albula Bowerbank, 1866: 268;
Yvesia Topsent, 1892a: 103. Pytheilla de Laubenfels, 1936a: Bowerbank, 1874b: 112, pI. XLV figs 21-24; Dendoryx albula;
64. Tisrone de Laubenfels, 1936a: 78. Yvesiella de Laubenfels, Gray, 1867a: 536; Myxilla albula; Vosmaer, 1880: 128. Myxilla
1936a: 78. exigua Hansen, 1885: 11, pI. II fig. 2; Grayella albula; Lundbeck,
1910: 37; Yvesia albula; Topsent, 1892a: 103; Crella albula;
Type species Burton, 1930a: 494, 506, fig. 5.
Material examined. Holotype: BMNH 1930.7.3.242 - from
Halichondria albula Bowerbank, 1866 (by subsequent desig- Shetland.
nation; Topsent, 1928c: 51). Description. Very thinly encrusting (on a shell), not exceed-
ing 1 cm in widest expansion; whitish sponge. Surface smooth.
Definition Oscules (or pore fields) scattered. Skeleton (Fig. 4D). Ectosomal
skeleton a tangential crust of acanthostyles; choanosomal skeleton
Crella with chelae; no basal echinating acanthostyles. irregular bundles of tomotes. Spicules (Fig. 4A-D). Ectosomal
Porifera· Demospongiae • Poecilosclerida • Myxillina • Crellidae 563

.,
"

E F

Fig.4. Crella (Yvesia) species. A-D, Crella (Yvesia) alhula (Bowerbank, 1866 as Halichondria). A-C, SEM photos of spicules from tbe holotype (scales:
A left and B, 10 fLm; A right and C, 1 fLm). D, drawing of skeleton and spicules made from a slide of the holotype. E, Crella (Yvesia) topsenti (Babi~, 1922
as Grayella), type of Pytheilla, drawing of spicules reproduced from his fig. U (sizes see text). F, Crella (Yvesia) spinulata (Hentschel, 1911 as Grayella),
type of Tisrone, drawing of spicules reproduced from his fig. 29 (sizes see text).

acanthostyles (Fig. 4B,D), some verging towards acanthoxeas, spicules. It is distinct from Pytheas in lacking basal echinating
curved, 100-120 X 2-3 /-Lm; tomotes (Fig. 4A, D) are style-like acanthostyles.
with a rounded and a pointed end, smooth, straight, 210--360 X The genus Pytheilla de Laubenfels, 1936a: 64 was erected for
2--6/-Lm; arcuate isochelae (Fig. 4C, D), 18-28/-Lm. Distribution type species (by original designation) Grayella topsenti Babi,<
and ecology. Shetland and Norway, mostly deep water, 7-180m (1922: 246, text-fig. U, here reproduced in Fig. 4E). This is
depth. encrusting other sponges or various substrates. Colour rose-violet
Remarks. The genus Yvesia Topsent, 1892a was originally or yellowish violet. Skeleton consisting of an ectosomal crust of
erected for Crella s.l., but later (1928c), Topsent designated acanthoxeas, 80--133 X 4-5 /-Lm, carried by choanosomal bundles
Halichondria alhula Bowerbank (1866) as type species, and of tomotes, 185-300 X 4-7 /-Lm. In contrast to what de Laubenfels
restricted its use for species with chelae, stylote tomotes and writes, Babi,< does not report the occurrence of echinating spicules
acanthostyles. Here the definition is widened again to include and thus the grounds for maintaining a (sub-)genus Pytheilla
diactinal tomotes and ectosomal spicules, in accordance with the separate from Yvesia are lacking. Microscleres: unguiferate arcuate
tendency in the type species to have variable endings of these isochelae, 12-19 /-Lm, and sigmas, 12-35 /-Lm.
564 Porifera· Demospongiae • Poecilosclerida • Myxillina • Crellidae

Fig. 5. A-D, Crella (Yvesia) pyrula (Carter, 1876 as Cometella), type of Yvesiella. A-C, SEM photos of spicules made from the holotype (scale 10 !Lm).
D, drawing of skeleton and spicules made from a slide of the holotype. E, Crellomima imparidens Rezvoi (1925), drawing of skeleton and spicules repro-
duced from his fig. 4 (scales see text). F-I, Crellomima incrustans Hentschel (1929), type of Damonilla, SEM photos of spicules made from a specimen in
BMNH (scales: 10 fLill, except I, I fLill).

The genus Tisrone de Laubenfels (1936a: 78) was erected for the stalk. Several embryos of 200-300/-Lm long are present in the
type species (by original designation) GrayeUa spinulata Hentschel slides at the base of the peripheral zone. Spicules (Fig. SA-D):
(1911: 340, fig. 29, here reproduced in Fig. 4F). The type (not smooth tomotes with blunt (Fig. SA) or sharp ends (Fig. SD), occa-
re-examined) is an encrusting-lobate sponge, with surface covered sionally faintly polytylote: 423-(S21.8)-S86 X 7-(8.7)-11 /-Lm;
in porefields; colour orange-red or grey-violet (alcohol). Skeleton ectosomal acanthostyles (Fig. SB, D), thickest some distance away
the usual dennal crust of acanthoxeas, 120-1S2 X 4--S /-Lm, and from the blunt end, fully spined: 123-(146.7)-166 X 8-(11.2)-
choanosomal tracts of oxeote tomotes, 264--328 X 3-S /-Lm, mixed 14/-Lm; arcuate isochelae (Fig. SC,D), middle part of the shaft
with acanthoxeas. Microscleres arcuate isochelae, 16-20/-Lm. This occasionally swollen: 21-(23.4)-26/-Lm. These characters confonn
combination of characters clearly confonns to Yvesia. to those of Yvesia. This species was recorded from several deep
The genus Yvesiella de Laubenfels, 1936a: 78 was erected for water localities along the coasts of Europe and North Africa (Boury-
the pedunculate type species (original designation) Cometella pyrula Esnault et al., 1994b).
Carter (1876: 388, pI. XIV fig. 20, pI. XV fig. 38) from deep water
(S12m) in the North Atlantic. The type BMNH 1882.7.2.8.11, was
re-examined. It has a thick (30-S0 /-Lm) multilayered crust of acan- GENUS CRELLOMIMA REZVOI, 1925
those megascleres (acanthostyles). The choanosomal skeleton (Fig.
SD) consists of a 800-1000 /-Lm thick peripheral zone in which thick Synonymy
bundles of tomotes (100 /-Lm at the periphery thickening to 200 /-Lm in
the centre of the sponge) carry the surface skeleton. These bundles Crellomima Rezvoi, 1925: 198. Damonilla de Laubenfels,
converge into the centre of the sponge where they fonn the axis of 1936a: 6S.
Porifera' Demospongiae • Poecilosclerida • Myxillina • Crellidae 565

Type species belonging to a species of [otroata, are found in cuts of the speci-
men. Some doubts about the true nature of this sponge remain, and
Crellomima imparidens Rezvoi, 1925: 198 (by monotypy). a reliable fresh record of the species is much needed. Nevertheless,
it is clear that Damonilla is a junior synonym of Crellomima.
Definition

Crellidae with anchorate chelae. CRELLASTRINA TOPSENT, 1927

Diagnosis Synonymy

Encrusting, with hymedesmioid skeleton of acanthostyles Crellastrina Topsent, 1927b. Aaaba de Laubenfels, 1936a: 65.
with heads embedded in a basal layer of spongin; choanosomal
skeleton of bundles of smooth tomotes; ectosomal skeleton the Type species
usual crust of acanthose megascleres; microscleres tridentate,
quadridentate or polydentate anchorate chelae. Two species. Yvesia alecto Topsent, 1898b: 248 (by original designation).

Description of type species Definition

Crellomima imparidens Rezvoi, 1925 (Fig. 5E). Crellidae with peculiar astrose derivations of spined oxeas
Synonymy. Crellomima imparidens Rezvoi, 1925: 198, forming a tangential crust; no chelae.
fig. 4; Koltun, 1959: 172, fig. 132.
Material examined. None. Whereabouts of the type are Diagnosis
unknown.
Description. Thinly encrusting, smooth, with leathery surface Encrusting, with an ectosomal crust of acanthoxeas
(Fig. 5E), covered with small (0.5 mm) papillae spaced 1-1.5 mm and astrose derivations; choanosoma1 skeleton of strongylote
apart. Colour bright rose. Skeleton. Ectosomal skeleton a crust of or stylote tomotes; no acanthostyles, no microscleres. One
acanthostyles tangentially arranged. Choanosomal skeleton of species.
obliquely arranged bundles of tomotes; at the base there are single
acanthostyles erect on the substrate. Spicules (Fig. 5E). Ectosomal Description of the type species
acanthostyles, 93-190 X 4--7/-Lm; oxeote tomotes, polytylote,
160-240 X 5-7/-Lm (Koltun gives 200-416 X 4--8/-Lm); basal acan- Crellastrina alecto (Topsent, 1898b) (Fig. 6A).
thostyles, 92-214 X 6-15/-Lm; spatulate tridentate or quadridentate Synonymy. Yvsia alecto Topsent, 1898b: 248; Topsent,
anchorate chelae, 16-22/-Lm. Distribution and ecology. Arctic, 1904b: 196, pI. XV fig. 16; Crellastrina alecto; Topsent, 1928c:
8-320m depth. 51; Aaaba alecto de Laubenfels, 1936a: 65.
Remarks. The type species is clearly described by Rezvoi, Material examined. Holotype: MOM (not seen).
and it was recorded again by Koltun, but cannot be considered well Schizotype: MNHN DT 1023 - slide of the holotype, labeled
known. The possession of anchorate chelae (the drawing of Rezvoi "Yvesia a1ecto n.sp. PA 1897,111".
cannot be interpreted otherwise) is an exception in the family Description (based on Topsent, 1904b). Small crust on a
Crellidae, and if this family is considered the sister family to deep water coral, thin, translucent, smooth, parchment-like. Colour
Hymedesmiidae, then the anchorate chelae are even more unusual. grey in alcohol (brown interior). Skeleton. The acanthoxeas and
This diminishes the value of chela micromorphology as a marker asters form a detachable tangential crust, carried by bundles of
for this group of sponges, unless the Crellomima chelae are tomotes. Spicules (Fig. 6A). Ectosomal acanthoxeas, curved, with
regarded as a separate independent conversion from ancestral arcu- the entire surface provided with long conical spines or tubercles,
ate chelae. Close relationship with Ectyomyxilla (Crellomyxilla) often partly finely spined or rugose, 120 X 6/-Lm (without spines),
(cf. family Myxillidae) is not likely because the skeletal architec- spines 15-18/-Lm long; asters of various shapes, looking similar
ture of that group is reticulate and the arrangement of the ectoso- to the spined oxeas and probably reduced derivations thereof:
mal acanthostyles is perpendicular, not tangential. 27-30 I-Lm in diameter; tomotes are strongyles with unequal end-
De Laubenfels (1936a: 65) erected Damonilla for type species ings, maybe to be called strongylostyles, with one end often with a
(by original designation) Crellomima incrustans Hentschel (1929: slight tyle: 540 X 7/-Lm. Distribution and ecology. Azores, deep
889, pI. XllI fig. 4). This thinly encrusting sponge has irregular sur- water, 600 m depth.
face, made up of a crust of acanthostrongyles and acanthostyles of Remarks. This is a monospecific genus based on very pecu-
113-156 X 6-7 f.1m; choanosomal oxeote tomotes are 210-280 X liar derivations of the ectosomal acanthoxeas. Some of the asters
4--7/-Lm, basal acanthostyles are 145-224 X 6-11 f.1m, and appear deceptively like true asters and they may take in extreme
microscleres are polydentate (5-7 teeth) chelae of 16-21 f.1m. forms a shape not unlike those of Jaspis or Hemiasterellidae, for
A specimen and a slide labeled Damonilla incrustans in The Natural example. Some of the acanthoxeas also resemble to some extent
History Museum London collection, BMNH 1936.11.20.14, from the peculiar spiraster-like spicules of the latrunculiid genus
'between Franz-Joseph Land and Lenin land, Zool. Inst. Acd. Sci. Sceptrintrus and their possible close relationship cannot be
Leningrad' appears to belong to this species. It is a hard yellow excluded. The assignment to Crellidae or even Poecilosclerida is
crust, with soft interior. It has the structure and contains the spicules tentative as no chelae or other microscleres, nor acanthostyles are
(Fig. 5F-I) recorded by Hentschel, including the polydentate found. Thus, this genus lacks any of the Poecilosclerida synapo-
anchorate chelae (Fig. 51), but some other spicules, presumably morphies. However, the crellid nature of the genus is strengthened
566 Porifera· Demospongiae • PoeciloscIerida • Myxillina • Crellidae

c
A B

Fig. 6. A, Crellastrina alecto (Topsent, 1898b as Yvesia), drawing of spicules reproduced from Topsent (1904b, pI. XV fig. 16). B-C, Spirorhabdia vidua
(Schmidt, 1875 as Spirastrella). B, drawing of spicules made from a slide of the holotype, reproduced from Thiele (1903b, fig. 27). C, drawing of spined
spicules reproduced from Topsent (1918, fig. xvi) (sizes see text).

by the occurrence of species like Spirorhabdia vidua (cf. below) Synonymy. Spirastrella vidua Schmidt, 1875: 120.
showing similarities to it, but possessing basal acanthostyles. Spirastrella aculeata Topsent, 1892a: 127, pI. VIII fig. 10;
The genus Aaaba de Laubenfels, 1936a: 65 is an objective Hymedesmia vidua; Thiele, 1903b: 393, fig. 27 Spirorhabdia
synonym because it has the same type species (by original designa- vidua; Topsent, 1918: 556, fig. xvi.
tion) as Crellastrina. Material examined. None.
Description (from Thiele, J903b). Thinly encrusting, size
about 1 cmz, smooth and even surface. Colour whitish. Skeleton.
Hymedesmioid with acanthostyles erect on the substrate, and bun-
SPIRORHABDIA TOPSENT, 1918 dles of tomotes. Ectosome a tight mass of spined rhabds forming
a coherent surface crust. Spicules (Fig. 6B). Ectosomal acan-
Spirorhabdia Topsent, 1918: 556. thorhabds, spines or thorns conical, with a terminal thickening,
45-50 X 5-8 f.Lm; tomotes, faintly polytylote, oxeote, 340-350 X
Type species 6 f.Lm; acanthostyles, spined allover, up to 350 X 15 f.Lm.
Distribution and ecology. Norway, Azores, deep water, below 190 m
Spirastrelia vidua Schmidt, 1875: 120 (by original designation). depth.
Remarks. This is a monospecific genus based on very pecu-
Definition liar derivations of the ectosomal acanthoxeas. It is here assigned
to Crellidae for the first time; previous asignment was to
Crellidae with polytylote ectosomal oxeote tomotes, basal Hymedesmiidae, but the surface crust of acanthose spicules of admit-
acanthostyles, and profusedly spined rhabds in an ectosomal crust. tedly peculiar shape, fits Crellidae much better than Hymedesmiidae.
Thiele (1903b) re-examined Schmidt's type material and expressly
Diagnosis stated that the spines of the acanthorhabds were not spirally arranged;
however, Topsent (1918) suggested that developmental forms demon-
Thinly encrusting, with dense ectosomal cover of acan- strated spiral arrangement, and he proposed to call these spicules
thorhabds, carried by bundles of choanosomal oxeote tomotes; basal 'spirorhabds', likening them to the spicules of Latrunculia and
acanthostyles echinate the substrate. No microscleres. One species. Sceptrintus. In view of the presence of acanthostyles, the spirorhabds
are unlikely to be homologous to spirasters of Spirastrella, but affin-
Description of type species ity with Latrunculia cannot be excluded, since that enigmatic genus
shows poecilosclerid affinities in its chemistry (similar to that of
Spirorhabdia vidua (Schmidt, 1875) (Fig. 6B-C). Zyzzya and Mycale).
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Dendoricellidae Hentschel, 1923

Rob W.M. Van Soest

Zoological Museum, University of Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, Netherlands. (soest@science.uva.nl)

Dendoricellidae Hentschel (Demospongiae, Poecilosclerida) is revived in an emended form wihin the suborder Myxillina for species
having only oxeas as megascleres and arcuate isochelae as microscleres, in addition to which sigmas may be present. The family resem-
bles the Myxillina family Desmacididae, but differs in possessing arcuate chelae and having spicules arranged in a plumose fashion, as
opposed to anchorate chelae and reticulate skeletons in Desmacididae. It is similar to the Myxillina family Phellodermidae, but that has
exclusively styles. There is also resemblance to the Mycalina family Isodictyidae, but these have palmate chelae. Three valid genera are
recognized in the family, viz., Dendoricella, Fibulia and Pyloderma.
Keywords: Porifera; Demospongiae; Poecilosclerida; Myxillina; Dendoricellidae; Dendoricella; Fibulia; Pyloderrna.

DEFINITION, DIAGNOSIS, SCOPE History and biology

Synonymy The name of the family was previously employed for a differ-
ent set of genera by Hentschel (1923). The former family
Dendoricellidae Hentschel, 1923. Desmacididae or Desmacidonidae was loosely employed for poe-
cilosclerid sponges having exclusively oxeas as megaScleres. As
Definition outlined in the chapter on the suborder Myxillina, and also again in
the remarks on the family Desmacididae, microsclere shape and
Myxillina with megascleres exclusively oxeas not differenti- distribution does not match the possession of exclusively oxeas as
ated morphologically into ectosomal and choanosomal forms. The megascleres and accordingly it is proposed to spread the former
skeletal architecture is plumose. Microscleres arcuate chelas. desmacidid genera over four families which are not otherwise con-
Sigmas may be present. sidered closely related (Desmacididae s.s., Dendoricellidae, and
Phellodermidae, all in the suborder Myxillina, and Isodictyidae in
Diagnosis the suborder Mycalina). The present family is emended from
Hentschel's concept (1923: 406) to accommodate Myxillina with
Shape usually elaborate, massive with fistules or long projec- the combined possession of exclusively oxeas as megascleres and
tions, branching or flabellate. Spicules sharp-pointed oxeas, some- arcuate chelae as microscleres. These properties, with the addition
times differentiated into distinct smaller ectosomal and larger of plumose instead of reticulate arrangement of the megascleres,
choanosomal sizes. Arrangement of megascleres plumose, occa- distinguish this small group of genera from the equally small
sionally almost plumoreticulate; in the case of fistular growth group of Desmacididae. Isodictyidae are reticulate and possess
fonns there is a parchment like tangential ectosomal skeleton and palmate isochelae. Members of the family Dendoricellidae occur
a scanty choanosomal skeleton. Microscleres scattered arcuate predominantly in cold or temperate waters, frequently at greater
isochelae, sometimes modified to resemble peculiarly round sig- depths.
moids, occasionally entirely absent. In the latter case membership
of this family is based on circumstantial similarities in megasclere Previous reviews
shape and arrangement.
Lundbeck (1905: 126), Hentschel (1923: 406).
Scope

Six nominal genera, three of which are considered valid:

Dendoricella, Fibulia and Pyloderrna.

KEY TO GENERA

(1) Oxeas differentiated in smaller ectosomal and larger choanosomal size categories ........................................................................... 2
Only a single size category of oxeas .......................................................................................................................................... Fibulia
(2) Shape bladder-like with hollow fistular projections ............................................................................................................ Pyloderma
Shape lobate or fan-shaped, not fistular ........................................................................................................................... Dendoncella

567
568 Porifera' Demospongiae • PoeciloscIerida • MyxilIina • DendoricelIidae

)1
B

( I
.

Fig. 1. A, Dendoricella rhopalum Lundbeck (1905), drawing of shape and spicules reproduced from his pI. IV figs 4-5, pI. XIV fig. 1 (scales see text).
B-E, Fibulia carnosa Carter (1886). B, drawing of skeleton and spicules made from a slide of the holotype. C, holotype (scale I cm). D-E, SEM photos of
spicules made from the type (scale 1 fLm).

DENDORICELLA LUNDBECK, 1905 dendritic tracts; microscleres are arcuate isochelae of normal
shape, and sometimes sigmas (absent in the type species).
Synonymy
Diagnosis
Dendoricella Lundbeck, 1905: 127.
Flabellate or club-shaped sponges with irregular surface.
Type species Smaller oxeas concentrated at the surface in bouquets.
Choanosomal skeleton dendritic-plumose, consisting of bundles of
Dendoricella rhopalum Lundbeck, 1905: 127 (by subsequent spicules larger than those of the surface. Not clearly reticulated
designation; de Laubenfels, 1936a: 53). although spicule tracts may be connected. Microscleres arcuate
isochelae and sigmas (may be absent).
Definition
Description of type species
Dendoricellidae with size differentiation in megascleres:
smaller ectosomal oxeas, and larger choanosomal oxeas forming Dendoricella rhopalum Lundbeck, 1905 (Fig. lA).
Porifera· Demospongiae • Poecilosclerida • Myxillina • Dendoricellidae 569

Synonymy. Dendoricella rhopalum Lundbeck, 1905: 127, Material examined. Holotype: BMNH 1886.12.15.71; slide
pI. IV figs 4--5, pI. XIV fig. 1. BMNH 1902.10.18.36 -labeled as "Desmacidon camosa", South
Material examined. None. Australia.
Description (from Lundbeck, 1905). This is a club-shaped Description. Cylindrical, thickly ramose or massively digi-
to stalked-flabellate species (Fig. lA). Live colouration unknown. tate sponges with slippery smooth surface (Fig. 1C). Size up
Skeleton a plumose-dendritic system of bundles of oxeas. These to 15 cm long. Scattered oscules on slightly raised papillae.
are apparently divisible in oxea-like tomotes of the surface smaller Consistency solid, fleshy. Colour dark red (beige in alcohol).
than the choanosomal oxeas, but morphologically they are similar. Skeleton (Fig. lB) at the surface consists of evenly distributed
Spicules, ectosomal oxeas, 340-429 X 7-11jLm, choanosomal brushes of oxeas surrounding the pores. Interiorly there is an axial,
oxeas, 680-980 X 14--22jLm. Microscleres normal-shaped arcuate massively developed system of spicule columns, up to 1 mm in
isochelae and thin growth forms resembling sigmas, 34--43 jLm. diameter, branching off and thinning out to spicule tracts of less
Distribution and ecology. Denmark Strait, between Iceland and than 20 jLm diameter towards the surface, where they fan out to
Greenland, deep water, 2000-3000m. form the ectosomal brushes. The fibre system is dendritic-plumose,
Remarks. The genus is close to Pyloderma, differing prima- with few anastomoses. Spicules (Fig. lB, D-E), oxeas of variable
rily in the non-fistular habit and punctate surface characters, size and endings, tending to be smaller at the surface but there
whereas Pyloderma has a parchment-like surface skeleton and are no clearly different size categories, 155-245 X 10-15jLm;
bears fistules. Several of the species assigned to Dendoricella by microscleres peculiarly reduced, sigmiform chelae (Fig. IE), occa-
Lundbeck (1905: 127) are no longer considered related (erella sionally with unequal endings, 8-15jLm. Distribution and ecology.
schmidti and Damiria cavemosa belong to Lissodendoryx), but South Australia, shallow water, 9-33 m.
other species of Lundbeck's list appear to be good members of Remarks. The nomenclatorial confusion over this genus was
Dendoricella, viz., Desmacidon abyssi Topsent (1904b: 204) and resolved by Hechtel (1965) and Hartman (1967). In summary, the
Dendoricella obesichela Lundbeck (1905: 130). The latter has sig- confusion stems from the fact that Burton (1929a: 424) assumed that
mas in addition to the arcuate chelae. Several other Dendoricella Fibulia was a misprint for Carter's (1882b) genus Fibularia, estab-
may hide under the genus name Isodictya, a genus with palmate lished for Fibularia massa. Later (1936: 142), Burton admitted that
chelae and reticulate skeleton. the evidence for that assumption was lacking and thus that Fibulia
should be reinstated with Plumocolumella as objective (or homo-
typic) synonym, both having the same species as the type.
Remarkably, he ignored this decision a few years later and used
FlBUUA CARTER, 1886 Plumocolumella again (Burton, 1938: 11). De Laubenfels (1936a: 51)
independently assumed that Fibulia was intended as a replacement
Synonymy name for Fibularia Carter (1882b), since that was preoccupied by
the echinoid genus Fibularia Lamarck (1815), a fact established
Fibulia Carter, 1886: 51. Plumocolumella Burton, 1929a: 424. by Vosmaer (1887). De Laubenfels (1936a) unfortunately assumed
Xytopsaga de Laubenfels, 1936a: 54. Fibulia to have massa as the type, despite the fact that Carter (1886)
did not give any indication that he was aware of the preoccupation or
Type species that he intended his name as a replacement. Since camosa is the
only species mentioned when Fibulia was used for the first time, it
Fibulia camosa Carter, 1886: 51 (by monotypy). is firmly established as the type species. Fibulia is similar to
Desmacidon in having the oxeote tomotes as structural megascleres
Definition arranged in a system of spicule tracts. The genera differ in the shape
of the isochelae (anchorate in Desmacidan) and skeletal architecture
Desmacididae with a plumose or confused skeleton of multi- (reticulate in Desmacidon). The two are here considered only
spicular tracts; microscleres arcuate or reduced isochelae. distantly related.
De Laubenfels (1936a: 54) erected the genus Xytopsaga for
Diagnosis type species Plumocolumella myxilloides Burton (1929a: 288) (by
original designation). The holotype BMNH 1928.2.15.321, and an
Massive; ectosomal skeleton regularly reticulate composed of additional specimen BMNH 1928.2.15.788 and one slide BMNH
the same spiCUles as in the choanosomal skeleton, and with brushes 1928.2.15.788a, from the Falkland Islands, 81-267m, were exam-
of spicules protruding through the surface; choanosomal skeleton ined. This is an erect arborescent sponge, with flattened anasto-
plumose or thoroughly confused, with curved or sinuous primary mosing branches, with smooth surface, but showing numerous
spongin fibres interconnected frequently by smaller secondary scattered circular depressions. The skeleton of this species is simi-
fibres, both cored by bundles of small thin oxeas or oxeotes lar to that of Fibulia camasa, with oxeas 248-333 X 5-8jLm,
ascending to surface and producing plumose dermal brushes, arranged in irregular wispy bundles ending in surface bmshes.
no echinating spicules; microscleres are arcuate or irregularly It differs from F. camasa in having nomlal arcuate isochelae,
unguiferate isochelae and sigmas. Half a dozen species. 26-33jLm, instead of reduced chelae. In view of the great similar-
ity in structure and further spiculation, the chelae of Fibulia are here
Description of type species considered highly deformed arcuate isochelae. A further species
to be allocated is Fibulia ancharata (Carter, 1881b: 382, pI. xvm
Fibulia camosa Carter, 1886 (Fig. lB-E). fig. 3, as Axos). The holotype and two paratype specimens together
Synonymy. Fibulia camosa Carter, 1886: 51; Plumacalumella labeled as BMNH 1871.5.12.34, the former from South Australia, the
camasa; Burton, 1929a: 424. latter from South Africa, were examined. This species has oxeas
570 Porifera· Demospongiae • PoeciioscIerida • MyxiUina • DendoriceUidae

I
,.

~ C [ J

Fig. 2. Pyloderma spp. A-B, Pyloderma latrunculioides (Ridley & Dendy, 1887 as Halichondria). A, photo of lectotype (scale I cm). B, drawings of habit,
detail of papilla, ectosomal skeleton and spicules reproduced from Ridley & Dendy (1887, pI. I fig.5, pI. II fig. I, pI. XLVI fig. 5) (scales see text).
C-E, Pyloderma demonstrans Dendy (1924). C, drawings of habit and spicules reproduced from Dendy (1924, pI. IX fig. 4, pI. XV figs 22-23) (scales see
text). D-E, SEM photos of spicules made from the type (scale D left, 100 f1m; D right and E, 10 f1m).

approximately 300 X 12 fLm and unguiferate chelae, 15-20 fLm. Definition

Hooper & Wiedenmayer (1994: 303) list an additional four species
from South Australia. Dendoricellidae with fistular shape.

Diagnosis
PYLODERMA KIRKPATRICK, 1907
Fistular, smaller and larger oxeas of same shape. Type
Synonymy species lacks chelae, but a second species, with similar shape,
identical oxeas, shape and size, has arcuate chelae. At least
Pylodemw Kirkpatrick, 1907a. Manawa Bergquist & Fromont, 2 species.
1988: 52.
Description of type species
Type species
Pylodemwlatrunculioides (Ridley & Dendy, 1886) (Fig. 2A-B).
Halichondria latrunculioides Ridley & Dendy, 1886: 326 Synonymy. Halichondria latrunculioides Ridley & Dendy,
(by subsequent designation; de Laubenfels, 1936a: 72). 1886: 326; Ridley & Dendy, 1887: 6, pI. I fig. 5, pI. II fig. 1,
Porifera' Demospongiae • Poecilosclerida • Myxillina • Dendoricellidae 571

pI. XLVI fig. 5; Van Soest et aI., 1990: 49, fig. 82; Pyloderma Remarks. Van Soest et al. (1990) assigned this genus to the
latrunculioides; Kirkpatrick, 1908c: 51; Dendy, 1924: 370; family Halichondriidae, but the close similarity with chela-bearing
Inflatella latrunculioides; Hentschel, 1914: 83. Pyloderma demonstrans Dendy (1924: 370) now renders this
Material examined. Lectotype (here figured in Fig. 2A), assignment untenable. The genus Manawa was erected by
and paralectotype: BMNH 1887.5.2.197 - "Challenger" collection, Bergquist & Fromont (1988: 52) for type species Pyloderma
Argentina; including two slides, and a further slide (BMNH demonstrans Dendy (1924: 370) (by original designation). The
1891.10.3.4). type specimen, BMNH 1923.10.1.152 from the 'Terra Nova'
Description. Erect-Iobose, pear-shaped (Fig. 2A), with Expedition, stat. 96, North Cape, New Zealand, was re-examined.
uneven, parchment-like surface (Fig. 2B). There are two specimens in This is a spherical sponge (Fig. 2C) with fistulose inhalant and
the BMNH jar and the specimen chosen as the lectotype is the one exhalant surface papillae; the ectosomal skeleton is a compact tan-
pictured by Ridley & Dendy. Oscule and pore areas elevated, rounded gential crust of oxeas; choanosomal skeleton consists of coarse
or elongated. Consistency soft, spongy. Colour light grey (alcohol). irregular, poorly defined tracts of oxeas, also dispersed throughout
Skeleton a dense ectosomal crust of felted, intercrossing oxeas, the choanosome; microscleres are arcuate isochelae. Oxeas
largely independent of the choanosome, which shows irregular tracts (Fig. 2C-D) measure about 500 X 12 f.Lm, chelae (Fig. 2E) 28 f.Lm.
and loose oxeas. Spicules, oxeas only, interpreted as tomotes, sharply We concur with Dendy (1924), that the presence of chelae is the
pointed, often inequiended, 700-1250 X 22-31 f.Lm. Distribution and only significant difference with P. latrunculioides and this is
ecology. Off the coast of Argentina, deep water, 1100 m. judged to be of specific value only.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademicIPlenum Publishers, New York, 2002

Family Desmacididae Schmidt, 1870

Rob W.M. Van Soest

Zoological Museum, University of Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, Netherlands. (soest@science.uva.nl)

Desmacididae Schmidt (Demospongiae, Poecilosclerida) are Myxillina characterized by the possession of anchorate chelae in combina-
tion with a reticulate skeleton of exclusively oxeas. By this definition the family contents are here newly restricted to the type genus
Desmacidon and its closest relative Desmapsamma. All other genera previously assigned by various authors to this family are referred to
other families: Myxillina with exclusively oxeas and arcuate isochelae are united in the family Dendoricellidae, whereas Myxillina with
exclusively strongyles and arcuate chelae are assigned to Chondropsidae, and Myxillina with exclusively styles and arcuate chelae are
assigned to Phellodermidae. Poecilosclerid sponges with exclusively diactinal spicules and palmate chelae are referred to the Mycalina
family Isodictyidae.
Keywords: Porifera; Demospongiae; Poecilosclerida; Myxillina; Desmacididae; Desmacidon; Desmapsamma.

DEFINITION, DIAGNOSIS, SCOPE History and biology

Synonymy The taxonomic history of Desmacididae is complicated and

full of erroneous assumptions and decisions, as was explained at
Desmacidinae Schmidt, 1870: 52 (in part) (emended to Des- length by Hajdu et ai. (1994a). This was the reason for these authors
macididae; Wiedenmayer, 1977b: 79). Desmacidonidae Gray, 1872a: to make a case for abandoning this name. However, since it is one
449 (in part). Desmacidontidae de Laubenfels, 1955b: E37 (in part). of the oldest names in sponge systematics, and since the priority
rule also applies to family names, it is not possible to maneuver
Definition it into synonymy. It is here newly proposed to restrict the use
of the family name to Desmacidon and its immediate relative
Myxillina with exclusively oxeote megascleres; no morpho- Desmapsamma, despite the similarity (microscleres) of
logical differentiation in ectosomal and choanosomal megascleres; Desmacidon to Myxilla in the family Myxillidae. Justification is
anchorate or unguiferate tridentate chelae and sigmas. found in the apparent suppression of choanosomal megascleres in
favour of the ectosomal tomotes, which are smooth oxeotes.
Diagnosis Several species are fouling organisms, and they may incorporate
considerable amounts of sand. Reproduction is viviparous where
Encrusting, massive, lobate, fan-shaped or branching sponges. known (Desmapsamma).
The ectosomal skeleton consists of perpendicular bundles or bou-
quets of megascleres and numerous microscleres. The bouquets of Taxonomic remarks
megascleres may also be at the end of longer subectosomal spicule
bundles traversing larger subdermal lacunae. Choanosomal skele- The possession of only tomote-derived spicules is shared with
ton reticulated, either an isodictyal network making small meshes genera such as Fibulia, Amphiastrella, several Coelosphaeridae
one spicule in length and width, or forming a coarser mesh in an and most Chondropsidae. From these, members of Desmacididae
anisotropic reticulation of thick spicule tracts. Desmacididae occur differ in having a reticulate skeleton and anchorate chelae. Sponges
in temperate and tropical shallow-water habitats. with only styles as megascleres and arcuate chelae are united in the
new family Phellodermidae.
Scope Previous reviews. Hajdu et ai., 1994a; Desqueyroux-
Faundez & Van Soest, 1996.
Two genera are included, both of which are considered valid:
Desmacidon, Desmapsamma.

KEY TO GENERA

(1) Skeletal architecture isotropic or irregular, making meshes one spicule in diameter, no distinct spicule tracts; sand is normally
incorporated in the ectosomal and choanosomal skeleton ........... ...... .... ...... ...... .......... .................... ....... ........... ....... ... Desmapsamma
Skeletal architecture plumoreticulate, consisting of thick spicule bundles making large meshes; no sand ...................... Desmacidon

572
Porifera • Demospongiae • Poecilosclerida • Myxillina • Desmacididae 573

~ c
I J

Fig.!. A-E, Desmacidonfruticosum (Montagu, 1818 as Spongia). A, habit oflectotype encrusting a barnacle (scale 1 cm). B-D, SEM photo ot spIcules
made from the lectotype (scale 10 !Lm). E, drawing of skeleton and spicules made from a slide of a specimen from Roscoff. F-J, Desmapsamma anchor-
ala (Carter, l882a as Fibularia). F, holotype fragments (scale I cm). G-I, SEM photos of spicules made from the holotype (scale 10 !Lm). J, drawing of
skeleton and spicules made from a slide of a specimen from Cura",ao.

DESMACIDON BOWERBANK, 1861 composed of smooth diactinal megascleres forming an isodictyal

reticulation; microscleres are spatuliferous anchorate isochelae and
Synonymy sigmas. Probably not more than half a dozen species, recorded
mostly from temperate and subtropical waters.
Desmacidon Bowerbank, 1861: 372.

Type species Previous reviews

SpongiafruticosaMontagu, 1818: 112 (by original designation). Ridley & Dendy (1887: 103); Bergquist & Fromont (1988:
37); Desqueyroux-Faundez & Van Soest (1996: 45).
Definition
Description of type species
Desmacididae with an anisotropic reticulation of spongin
enforced spicule tracts and anchorate isochelae. Desmacidon fruticosum (Montagu, 1818) (Fig. lA-E).
Synonymy. Spongia fruticosa Montagu, 1818: 112;
Diagnosis Halichondria fruticosa; Johnston, 1842: 103, pI. XN fig. 1;
Desmacidon fruticosum; Bowerbank, 1861: 372; Bowerbank,
Erect growth forms; ectosomal skeleton composedof smooth 1866: 345; Bowerbank, 1874b: 155, pI. LXI figs 1-7; Ackers
oxeote spicules arranged in bouquets; choanosomal skeleton et at., 1992: 119, fig. 42.
574 Porifera· Demospongiae • Poecilosclerida • Myxillina • Desmacididae

Material examined. Lectotype: BMNH 1930.7.3.410. Other forming a paucispicular isotropic reticulation of short spicule tracts
material. ZMA POR. 286 - Atlantic coast of France, Roscoff, composed of oxeas similar to those of the ectosome; megascleres
18-25 m, colI. G.J. Kleeton. ZMA POR. 2435 - Mediterranean, slender oxeas; microscleres are anchorate isochelae and sigmas.
Banyuls, colI. J.H. Stock. Two species from the tropical Atlantic and Pacific.
Description (mostly derived from Ackers et al., 1992).
Massive-lobose (Fig. lA); substantial growths often with massive Previous reviews
tubular processes. Up to 22 cm high. The processes are simple or
coalescent, often fusing at the bases and so resembling malformed, Burton (1934a), Desqueyroux-Faundez & Van Soest (1996),
band-like structures. Sometimes the thicker branches have a longi- Van Soest (1998).
tudinal groove in one side. Surface bristly. Oscules are usually
small, scattered and numerous. Consistency firm, compressible. Description of type species
Exudes large amounts of slime when removed from the water. No
smell. Colour yellow to orange. Skeleton (Fig. IE) of the ectosome Desmapsamma anchorata (Carter, 1882a) (Fig. IF-J).
consists of bouquets of tornotes, possibly slightly shorter than Synonymy. Fibularia anchorata Carter, 1882a: 283;
those of the choanosomal tracts. Choanosomal skeleton is an irreg- Desmapsamma anchorata: Burton, 1934a: 547 (only West Atlantic
ular reticulation of thick multi spicular tracts of tornotes, about specimens); Burton, 1956: 131; Levi, 1959: 134, pI. 5 fig. 3;
100 fLm in diameter, forming meshes of 800 X 500 fLm. Many Hechtel, 1965: 21, pI. II fig. 4; Hartman, 1967: 20, pI. 7 fig. 1;
loose individual spicules. Spicules (Fig. lB-E), tornotes, with Van Soest, 1984b: 35, text-fig. 10, pI. II figs 2-5; Van Soest, 1998:
oxeote or mucronate ends, 184-236 X 5-8 fLm; spatuliferous 433. Desmacidon reptans Ridley & Dendy, 1886: 345; Ridley &
anchorate chelae oftwo sizes, 15-20 and 32-49 fLm, and sigmas of Dendy, 1887: 105, pI. XXIII fig. 7. Desmacidon carterianum
two sizes, 15-20 and 50-63 fLm. Distribution and ecology. British Arndt, 1927: 147.
Isles; France; Spain, Mediterranean, on mud, sand, gravel, rock, or Material examined. Holotype: BMNH 1939.3.24.12 -
broken shell. Recent records are from 30 m or greater depths. Antigua, Carter collection. Other material. Many specimens in
Remarks. The alleged holotype is of uncertain status as ZMA from Cura~ao and Cape Verde Islands.
most of Montagu's material is no longer extant, and the label with Description. Upright, somewhat ramose masses (Fig. IF)
the specimen is not original; nevertheless it is here considered the with oscules on elevations, or sprawling clumps of volcanoe-
lectotype. The microscleres of this species are similar to those of shaped oscular tubes. Up to 15 cm long or more, up to 15 cm in
various Myxilla species, but lack of spined styles is a good micro- diameter (largest type fragment 3 cm long, 1 cm diameter). Oscules
scopic marker to tell this apart from species of that genus. Other 1-5 mm in diameter. Consistency compressible, rather soft; slimy.
species of Desmacidon: the genus name is widely used in the liter- Colour. Variably pale-purplish pink to dirty pink (salmon-colour),
ature, but few of those records seem to correspond to the characters or dark brown, depending of the quantity and colour of sand grains
of the type species. They are likely or certainly members of other available in the environment. Skeleton (Fig. lJ). Ectosomal pal-
genera, e.g., Fibulia, Pyloderma or Isodictya. isade of bundles of diverging oxeas covered and partly obscured by
variable quantities of sand grains; single oxeas may also form an
irregular surface reticulation. The cohesiveness of the ectosomal
DESMAPSAMMA BURTON, 1934 skeleton is influenced strongly by the quantity of foreign material.
Choanosomal skeleton. Basically a renieroid, irregularly isotropic
Synonymy skeleton of bundles of oxeas, 3-6 per side, but here and there
longer ascending primary tracts are visible. The bundles and tracts
Desmapsamma Burton, 1934a: 547. may be strengthened and partly replaced by variable quantities of
sand grains and broken spicules. Spicules (Fig. IG-J). Oxeas, slen-
Type species der, sharply pointed, 140-190 X 3-9 fLm; anchorate chelae in
2 size categories, the larger with fimbriae along the entire length of
Fibularia anchorata Carter, 1882a: 283 (by original the shaft, 15-20 and 8-10 fLm; sigmas in 2 size categories, 19-39
designation). and 11-16 fLm. Distribution and ecology. Tropical Atlantic, on both
African and American coasts, in shallow reef habitats, lagoons,
Definition often fouling.
Remarks. Desqueyroux-Fa6.ndez & Van Soest (1996: 45)
Desmacididae with paucispicular isotropic reticulation; sand suggested that the genus is very close to Desmacidon and should be
is normally incorporated in variable quantities. recognized only at the subgeneric level. The genus was recently dis-
cussed by Van Soest (1998), who erected D. vervoorti, with type from
Diagnosis Indonesia, for Indo-Pacific records of D. anchorata. This differs sub-
tly from the Caribbean D. anchorata in colour, surface characteristics,
Erect, and repent-ramose growth form; ectosomal skeleton and in a third size category of sigmas. In view of the fact, that at least
tough, arenaceous, with oxeas in bouquets and sand grains forming one other species similar to the type species exists, sharing the
a narrow-meshed reticulation sometimes completely replacing isotropic reticulation of the oxeas as well as a variable sand cover, it
oxeas; choanosomal skeleton arenaceous, also with smooth oxeas seems preferable to keep Desmapsamma as a separate genus.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Hymedesmiidae Topsent, 1928

Rob W.M. Van Soest

Zoological Museum, University of Amsterdam, P.O. Box 94766,1090 GT Amsterdam, Netherlands. (soest@science.uva.nl)

Hymedesmiidae Topsent (Demospongiae, Poecilosclerida) is revised substantially to form a new concept within Myxillina for sponges
with hymedesmioid or plumose skeletal arrangement in combination with the possession of arcuate chelae and smooth diactinal ectoso-
mal megascleres. The formerly employed family Anchinoidae is here merged with a former concept of Hymedesmiidae, because no clear
line can be drawn between the hymedesmioid and the anchinoid skeletal architecture. The family is close to Crellidae, sharing
hymedesmioid and/or plumose skeletal arrangement and the possession of areolated porefields, but differs in lacking the surface special-
ization of tangentially arranged acanthoxeas and -styles. Ten genera and two subgenera, of 34 nominal genera, are considered valid. The
species described as Lissopocillon dendyi Ferrer-Hernandez, 1916 is assigned to Phorbas and because the combination Phorbas dendyi is
already occupied by P. dendyi (Topsent, 1913) a new name Phorbasferrerhernandezi n. sp. is here proposed.
Keywords: Porifera; Demospongiae; Poecilosclerida; Myxillina; Hymedesmiidae; Acanthancora; Hamigera; Hemimycale; Hymedesmia;
Hymedesmia (Hymedesmia); Hymedesmia (Stylopus); Kirkpatrickia; Myxodoryx; Phorbas; Plocamionida; Pseudohalichondria; Spanioplon.

DEFINITION, DIAGNOSIS, SCOPE Scope

Synonymy Thirty four nominal genera of which ten genera and two
subgenera are considered valid. Acanthancora, Hemimycale,
Hymedesmiidae Topsent, 1928c: 54. Anchinoidae Topsent, Hamigera, Hymedesmia, Hymedesmia (Hymedesmia), Hymedesmia
1928c: 57. Phorbasidae de Laubenfels, 1936a: 62. (Stylopus), Kirkpatrickia, Myxodoryx, Phorbas, Plocamionida,
Pseudohalichondria, Spanioplon.
Definition
History and biology
Myxillina with smooth ectosomal tomotes (basically diacti-
nal, but may be anisotomotes or styles), a peripheral choanosomal This family unites the former families Anchinoidae and
plumose skeleton of bundles of smooth tomotes, a basal skeleton of Hymedesmiidae sensu Topsent, 1928c: 54 (excepting those genera
acanthostyles, singly or in plumose bundles (may be absent), echi- with anchorate or palmate chelae, or with forceps), because no clear
nating acanthostyles (may be absent). Additional non-echinating line can be drawn between the hymedesmioid and the anchinoid
acanthostyles or acanthoxeas may occasionally be present. skeletal architecture. Hymedesmioid architecture grades into
Microscleres include arcuate chelas and sigmas, but may be absent. plumose architecture by strengthening and elongation of the subder-
mal tomote bundles. In fact the hymedesmioid architecture occurs
Diagnosis in many, often unrelated, families (Raspailiidae, Microcionidae,
Crambeidae, Crellidae), and thus using it to unite an arbitrary group
Encrusting, massive and branching sponges; surface with of sponges with arcuate chelae appears not justified. It is proposed
characteristic groups of pores (areoles) (shared with Crellidae). to abandon hymedesmioid architecture as a family character (it may
Megascleres smooth diactinal spicules (oxeote, tylote or strongy- dubiously be retained as a genus character).
lote, occasionally stylote tomotes), grouped together or scattered in
the ectosome, but never forming a crust, lying perpendicular, tan- Taxonomic remarks
gential, or paratangential to the surface, and acanthostyles, which
may typically occur in a larger size class, which is often partly Microscleres. These are the usual arcuate chelae and sigmas,
smooth in the pointed half, and a smaller size, which is usually either one or both of which may be absent. In some genera interest-
entirely spined. Diactinal spicules forming thinner or thicker ing modifications of the arcuate isochelae occur. Bipocilli-like
plumose or plumo-reticulate tracts or columns in the choanosome, microscleres were reported from a genus Lissopocillon, but these are
with only a thin coat of spongin, or without optically visible spon- merely malformed arcuate chelas and the genus is considered a jun-
gin. Columns usually consist of spicules identical to those in the ior synonym of Phorbas. Somewhat similar chelae were reported in
ectosome, but sometimes these are replaced by acanthostyles. They a genus Bipocillopsis, but they are also obviously reduced chelae,
mayor may not be echinated by 1-2 sizes of acanthostyles. At the and the genus is a junior synonym of Hymedesmia. True bipocilli
base of the sponge there is usually a spongin plate closely adhering are a synapomorphy for the genus Iophon (suborder Microcionina,
the substrate and echinated by single acanthostyles or groups or family Acamidae). Chiasto sigmas occur occasionally.
bundles of acanthostyles. Microscleres may include arcuate Genera excluded. A number of genera, which have tradition-
isochelae and sigmas. ally been assigned to Hymedesmiidae based on their hymedesmioid

575
576 Porifera' Demospongiae • Poecilosclerida • Myxillina • Hymedesmiidae

architecture, are here referred to other families on account of the H. japycina Topsent, 1927b) is a Myxillidae, genus Hymenancora.
characters of their microscleres. Holorodesmia Topsent, 1928c Lissomyxilla Hanitsch, 1894a (type Tethea spinosa Bowerbank,
(type Hymedesmia flaccida Topsent, 1927b) is a Clathria 1874b) is a junior synonym of Tethyspira Topsent, 1890b (see de
(Microciona) (type slide examined in MNHN). Hymedesanisochela Laubenfels, I 936a), order Halichondrida, family Dictyonellidae.
a Bakus, 1966b (typeH. rayae Bakus, 1966b) is an Acamidae, prob- Dendoricella Lundbeck, 1905 (type D. rhopalum Lundbeck, 1905)
ably Iophon but it lacks bipocillae (paratype specimen examined in is assigned to a separate family Dendoricellidae uniting plumose
BMNH). Quindesmia de Laubenfels, 1936a (type Hymedesmia sponges with only oxeas and arcuate chelae. Xytopsaga Burton,
inflata Bowerbank, 1866) is a Halicnemia, as its type is a junior 1932b (type P. myxilloides Burton, 1932b) is a junior synonym of
synonym of H. patera) (slide of type examined in BMNH). genus Fibulia, family Dendoricellidae. Fibulia Carter, 1886c (type
Chiastosia Topsent, 1928c (type Myxilla pecqueryi Topsent, F. camosa Carter, 1886c) goes to a separate family Dendoricellidae.
1892a) is a Myxillidae, genus Hymenancora. Arndtanchora de Sister family. The relationship with the family Crellidae is
Laubenfels, 1936a (type Leptosia sirventi Topsent, 1927b) is a close, because of many shared features. The two families are kept
Myxillidae, genus Hymenancora. Hymesigmia Topsent, 1927b (type apart because of the unique crellid ectosomal skeleton.

KEY TO GENERA

(1) Spicules exclusively smooth styles and strongyles, no microscleres or acanthose spicules ........................................... Hemimycale
Microscleres and/or spined spicules present ...................................................................................................................................... 2
(2) Microscleres are profusely ornamented chelae or derivates .............................................................................................................. 3
Microscleres if present are smooth-shafted ........................ ... .......... ....... ... ... ..... .............. .... ........... ............ ....... .......... ..... ................. 4
(3) Skeleton structure hymedesmioid, i.e., with single acanthostyles erect on the substratum ......................................... Acanthancora
Skeleton consistst of columns of tomotes without acanthostyles ...................................................................... Pseudohalichondria
(4) All spicules smooth, spined spicules absent ....................................................................................................................... Hamigera
Lightly or more heavily spined spicules present .............................................. ,............... ...... .......................... ..................... ...... ...... 5
(5) Acanthostrongyles present forming a basal mass or reticulation ................................................................................. Plocamionida
No acanthostrongyles or basal reticulation ........................................................................................................................................ 6
(6) Skeleton hymedesmioid, i.e., with acanthostyles singly erect on the substratum .......................................................... Hymedesmia
Skeleton plumose or plumoreticulate, acanthostyles, if present, echinating the choanosomal tracts, incorporated in the skeletal
columns or strewn in the interior, not singly erect on the substrate .................................................................................................. 7
(7) Accessory acanthose spicules (acanthostyles, acanthoxeas) present, strewn in the interior, not localized; choanosomal styles smooth
or microspined ..................................................................................................................... ,.... ........ .... ....... ................. .... Spanioplon
Acanthostyles localized in echinating position, or incorporated in the skeletal tracts ........................................................................ 8
(8) Acanthostyles only in skeletal columns or tracts .............................................................................................................................. 9
Acanthostyles echinating the columns or tracts or strewn randomly .............................................................................................. 10
(9) A single category of acanthostyles ........................................................................................................................................ Phorbas
Two distinct categories of acanthostyles ........................................................................................................................... Myxodoryx
(10) Acanthostyles in echinating position ..................................................................................................................................... Phorbas
Acanthostyles in the columns and strewn in the interior, not echinating ...................................................................... Kirkpatrickia

ACANTHANCORA TOPSENT, 1927 composed of acanthostyles erect on the substrate; microscleres are
isochelae ornamented with smaller or larger spines, derived from
Synonymy arcuate isochelae; further microscleres may include chiastosigmas.
Half a dozen species.
Acanthancora Topsent, 1927b: 12. Hymenamphiastra de
Laubenfels, 1930: 27. Ferrerhemandezia de Laubenfels, 1936a: 87. Previous reviews

Type species Topsent, 1928c, Van Soest, 1984b.

Leptosia schmidti Topsent, 1898b: 250 (by original Description of type species
designation).
Acanthancora schmidti (Topsent, 1898b) (Fig. lA).
Definition Synonymy. Leptosia schmidti Topsent, 1898b: 250, fig. 2b;
Topsent, 1904b: 189, pI. XV fig. 9; Acanthancora schmidti;
Hymedesmiidae with modified spined-tuberculate chelae and Topsent, 1927b: 12; Topsent, 1928c: 274.
hymedesmioid structure. Material examined. Holotype: MOM (not seen); MNHN
DT961- slide labeled 'PA 1897,48' made from holotype.
Diagnosis Description (based on Topsent, 1898b, 1904b). Thinly
encrusting, smooth, irregular. Colour white. Skeleton hymedesmioid,
Encrusting; ectosomal skeleton composed of smooth tomotes acanthostyles are arranged in bouquets with heads embedded in the
or anisotomotes, some with polytylote bases; choanosomal skeleton spongin adhering the substrate. Tomotes are concentrated in the
Porifera· Demospongiae • Poecilosclerida • Myxillina • Hymedesmiidae 577

..
1
E
F
Fig. 1. Acanthancora. A, Acanthancora schmidt; (Topsent, 1898b as Leptos;a), drawing of spicules reproduced from Topsent (1904b, pI. XV fig. 9) (sizes
see text). B-E, Acanthancora cyanocrypta (de Laubenfels, 1932 as Hymenamphiastra). B, SEM photo made from a paratype showing arrangement of the
skeleton with groups of acanthostyles (scale 100 /-Lm). C, details of apices of tomote (scale I /-Lm). D, spined and deformed chela (scale 10 /-Lm). E, draw-
ing of skeleton and spicules made from a slide of a paratype. F, Acanthancora aenigma (Lundbeck, 1910 as Hymedesmia), type of Ferrerhernandezia de
Laubenfels (l936a), drawing of spicules reproduced from Lundbeck (1910, pI. IX fig. 1) (sizes see text).

periphery, scattered singly among the numerous microscleres. Acanthancora, A. clavatancora, which seems quite close to A.
Spicules, ectosomal tomotes, strongyle-like, inequiended, with schmidti, including the chiastosigmas. It is quite possible that the two
a rounded and a mucronate end, 200--220 X 3 fLm; choanosomal belong to the same species. Burton (1959a) described A. stylifera
acanthostyles, of widely different sizes but not clearly differenti- from the Indian Ocean and Van Soest (1984b) A. coralliophila from
ated in structural and echinating spicules, all entirely spined, the Caribbean. This genus is similar to Hymedesmia, differing in the
112-260 X 7-15 fLm. Microscleres strongly deformed and spined, peculiar microscleres, which are shared by non-hymedesmioid
looking somewhat like diplasters, but presumably are derived from Pseudohalichondria (cf. below).
(arcuate) chelae, 12-14 fLm. Chiastosigmas, 7-8 fLm. Distribution The genus Hymenamphiastra de Laubenfels, 1930 was
and ecology. Azores; deep-water, 349 m. erected for type species (monotypy) H. cyanocrypta de Laubenfels
Remarks. The slide of the holotype examined contains large (1930: 27) (cf. also de Laubenfels, 1932: 87, fig. 51). A paratype,
amounts of curved smooth oxeas, but these are obviously not BMNH 1929.8.22.18, including two slides, was re-examined.
proper. Also, the chiastosigmas mentioned by Topsent (1904b) are The skeleton (Fig. lB, E) of this blue thinly encrusting species has
not found in the slide, which could mean that they may also not be single large acanthostyles of 250-280 X 13 fLm erect on the substrate
proper. Topsent described a second Azoran deep-water species of surrounded and echinated by short acanthostyles of 75-90 X 10 fLm
578 Porifera' Demospongiae • Poecilosclerida • Myxillina • Hymedesmiidae

to form distinct tree-like bouquets. Ectosomal tomotes are thin and distributed tangentially and intercrossing, but they are arranged
oxea-like with lanceolate apices (Fig. lC,E), 160-170 X 3 /-tm. in parallel sheets in the wall of the areolae. The areolate surface
Microscleres are strongly deformed bluntly spined reduced chelae contains mostly microscleres, strewn around the pores.
(Fig. lD, E) of 10-15 /-tm.1t conforms closely to Acanthancora. Choanosomal skeleton consists of plumose bundles of strongyles
The genus Ferrerhernandezia de Laubenfels, 1936a: 87 was echinated by smooth styles. Spicules (Fig. 2C-E), tomotes in
erected for type species (by original designation) Hymedesmia the form of strongyles to tylotes (Fig. 2C), smooth with faintly
aenigma Lundbeck (1910: 84, pI. IX fig. 1, here reproduced in swollen endings, 230-320 X 3-7/-tm; styles to subtylostyles
Fig. IF) (not examined). This is a thinly encrusting sponge with (Fig. 2D), often rather abruptly curved in the second half,
vertically placed acanthostyles, 130-340 X 14-21 /-tm, and bundles 240-320 X 5-9/-tm; arcuate isochelae (Fig. 2E), 15-22/-tm.
of ectosomal polytylote oxeote tomotes, 320-430 X 6-7/-tm. Distribution and ecology. Mediterranean; on rocks and under
Microscleres are 'asteroid' chelae, 28-35 /-tm. This combination of overhangs, down to 23 m.
characters conforms closely to Acanthancora. Re1lUlrks. Hamigera hamigera was originally described in
the genus [Cribrella] Schmidt, 1862 which was preoccupied by
Cribrella Agassiz, 1835 (Echinodermata). The present species
was fixed as the type species of [Cribrella] Schmidt, 1862
HAMIGERA GRAY, 1867 by Desqueyroux-Faundez & Stone (1992). Subsequently, Gray
(1867a) erected Hamigera with the same species, C. hamigera as
Synonymy the type. Amoibodictya Zahn et al. (1977: 105) is a synonym,
because the type species (by monotypy) A. Jorsteri from the
[Cribrella] Schmidt, 1862: 70 (preocc.). Hamigera Gray, Adriatic is an obvious junior synonym of Hamigera hamigera.
1867a. Amoibodictya Zahn et al., 1977: 105. This genus is distinguished from Phorbas and Hymedesmia
by the absence of acanthostyles, instead of which smooth styles are
Type species found in echinating position. About half a dozen Hamigera species
have been reported from Indo-Pacific and Australian-New Zealand
Cribrella hamigera Schmidt, 1862: 70 (by original waters.
designation).

Definition HEMIMYCALE BURTON, 1934

Hymedesmiidae without acanthostyles; with strongyles or Synonymy

tylotes as tomotes; with coring and echinating smooth styles or
subtylostyles; arcuate chelae. Hemimycale Burton, 1934a: 556.

Diagnosis Type species

Encrusting to massive sponges; oscules slightly elevated, Desmacidon columella Bowerbank, 1874b: 243 (by original
rounded, pores in characteristic and conspicuous areolate pore- designation).
fields; ectosomal diactinal spicules form fans around areolae;
choanosomal skeleton composed of plumose tracts of smooth Definition
strongyles or strongylote tomotes, which may be mixed with
smooth styles or subtylostyles, with echinating smooth styles or Hymedesmiidae without acanthostyles; without microscleres;
subtylostyles; microscleres are arcuate isochelae. Half a dozen megascleres strongyles and styles, not divisible into ectosomal or
species. choanosomal spiCUles.

Previous review Diagnosis

Bergquist & Fromont, 1988: 73. Thickly encrusting to massive, with striking and characteristic
areolated porefields; skeleton of plumose columns of mixed styles
Description of type species and slightly inequiended strongyles, which are probably derived
from tomotes; the choanosomal columns fan out near the surface
Hamigera hamigera (Schmidt, 1862) (Fig. 2A-E). and strengthen the raised porefields; no microscleres. One definite
Synonymy. Cribrella hamigera Schmidt, 1862: 70, pI. VI valid species, but others may exist.
fig. 13; Desqueyroux-Faundez & Stone, 1992: 10, pI. XXIV
fig. 138. AmoibodictyaJorsteri Zahn et aI., 1977: 106, figs 1-6. Previous review
Material examined. Holotype: LMJG 15359 - Zara Canal.
Schizotype: BMNH 1867.3.11.70 - slide of Schmidt #59, Adriatic. Forster, 1955.
Description. Encrusting to massive lobose (Fig. 2A-B), up
to 2-3 cm thick, up to 10 cm in widest expansion. Smooth surface Remarks
with conspicuous slightly elevated, oval areolae (Fig. 2B), with
oscules on similar elevations. Consistency fleshy. Colour orange- Assignment of Hemimycale to this family is controversial,
red. Skeleton of the ectosome consists of strongyles or tylotes because of the uniform spiculation of smooth tomote-like spicules
Porifera· Demospongiae • Poecilosclerida • Myxillina • Hymedesmiidae 579

Fig. 2. A-E, Hamigera hamigera (Schmidt, 1862 as Cribrella). A, photo of holotype reproduced from Desqueyroux-Faundez & Stone (1992, pl. XXIV
fig. 138) (scale 1 cm). B, Schmidt's drawing of the type and microscleres (sizes see text). C-E, SEM photos of spicules made from the holotype (scale
1 fLm). F-I, Hemimycale columella (Bowerbank, 1874b as Desmacidon). F-G, SEM photos of spicules made from the holotype (scale 100 fLm). H, photo
of live specimen made by Bernard Picton. I, photo of type specimen (scale 1 cm).

without clear differentiation and lack of microscleres. Several from Hemimycale, a logical and defendable action. Ulosa pos-
authors assigned it to the Halichondriidae, along with such genera sesses exclusive styles. It has been misinterpreted for Scopalina
as Batzella and Ulosa. The presence of stylote and strongylote (Halichondrida: Dictyonellidae) and is now assigned to Mycalina,
derivatives of apparently the same spicule type would allow inclu- family Esperiopsidae.
sion of this genus in the family Chondropsidae. However, the
genus is here assigned to Hymedesmiidae corroborated by the con- Description of type species
spicuous presence of surface areolae and skeletal architecture of
strengthening of the elevated areolae by parallel arrangement of the Hemimycale columella (Bowerbank, 1874b) (Fig. 2F-I).
spicules in the wall. These constitute compelling similarities with Synonymy. Desmacidon columella Bowerbank, 1874b:
Hamigera, Phorbas and Hymedesmia. Batzella does not possess 243, pI. LXXVIII figs 6-8; Stylinos columella; Topsent, 1892a:
these traits, and moreover has spicules only in a single type, i.e., 136; Stylotella columella; Topsent, 1894b: 26; Prianos columella;
strongy1es, which makes its assignment to Chondropsidae, away de Laubenfels, 1932: 62 (also as Prianos columellus sic);
580 Porifera· Demospongiae • Poecilosclerida • Myxillina • Hymedesmiidae

Hemimycale columella; Burton, 1934a: 556, fig. 10; Forster, 1955: bundles in the choanosome; choanosomal basal skeleton
553, figs 1-3. hymedesmioid, composed of one or two sizes of acanthostyles
Material examined. Holotype BMNH 30.7.3.434, Exmouth, standing singly erect on the substrate; microscleres are arcuate
Bowerbank collection; four unregistered slides, BMNH 'Bk 818', isochelae and/or sigmas. Many species.
Bowerbank collection.
Description. Encrusting; several mm in thickness, lateral Remarks
expansion up to 5 cm or more (Fig. 21). Surface closely covered in
numerous large rounded areolate porefields (Fig. 2H), which are Hymedesmia is close to Phorbas in most spicular characters,
conspicuous because they have elevated rims lighter coloured than but the two are separated by the hymedesmioid arrangement of the
the pore areas. Consistency soft, slippery. Colour bright red to basal acanthostyles in Hymedesmia against the echinating and/or
pale orange or pink. No definite ectosomal skeleton, although columnar arrangement of the acanthostyles in Phorbas. This dis-
scattered spicules occur tangentially in the skin; the walls of the tinction is problematical in individual species and may not be ten-
areolae are supported by sheets of spicules. There is no differentia- able eventually. In this family species with profusely ornamented
tion in shape of the spicules into ectosomal and choanosomal spined chelae may show typical hymedesmioid (Acanthancora
spicules. Choanosomal skeleton consists of plumose bundles of schmidti, Hymedesmia crux) or Phorbas-like arrangement
spicules, branching and anastomosing infrequently. In many speci- (Pseudohalichondria clavilobata), indicating the low phylogenetic
mens a secondary skeleton of calcareous nodules is developed significance of these skeletal arrangements. In view of the large
(see Vacelet et al., 1987); it is unclear whether this has taxonomic number of Hymedesmia and Phorbas species it is practical to
significance. Spicules. Probably only a single type, but their maintain them as separate genera. In line with this decision it is
endings are either rounded at both ends, thus technically strongyles also proposed to maintain two genera for Hymedesmiidae with
(Fig. 2F), or inequiended, occasionally one end mucronate, spined chelae, i.e., Acanthancora for those with hymedesmioid
technically styles (Fig. 2G) or strongylosty1es, 330-465 X 2-8 fLm. structure, and Pseudohalichondria for those with a more Phorbas-
Reproduction. Viviparous, April to October. Distribution and like structure (cf. above and below). Because the number of
ecology. North East Atlantic, Mediterranean, Azores; on rocks, Hymedesmia species is very large, it is proposed to employ two
shallow water. subgenera for convenience, Hymedesmia s.s. for species with
Remarks. So far this remains a monotypical genus, although microscleres (usually chelae, to which may be added sigmas), and
Sara (1960) and Sara & Siribelli (1960) described two additional Stylopus Fristedt, 1885 for species without microscleres. It is not
species from the Mediterranean, viz., Hemimycale ambigua and likely that loss of chelae indicates close relationship, and the dis-
H. brevicuspis, both probable synonyms of Crambe crambe (see tinction is very likely to be artificial. Dolichacantha Hentschel,
chapter on family Crambeidae). Vacelet et al. (1987: 260) recorded 1914 was erected for type species D. macrodon Hentschel (1914)
an unnamed species from the Red Sea. This assignment is chal- from the Antarctic. The description of Hentschel (1914) records an
lenged by Van Soest et al. (1996: 93), because its chemistry con- extraordinary combination of spicules, including very long smooth
forms to Crambe and Monanchora. A further record, although not tylotomotes acanthostyles of two distinct size classes, peculiar
corroborated by a description, is from Brazil (Muricy et al., 1991). acanthostrongyles with numerous spined 'cladi', palmate isochelae
The calcareous nodules described by Vacelet et al. (1987) were also and sigmas. Some of the spicules are said to be 'not numerous'
recorded by Bowerbank in his plate LXXVm as 'gemmules'. (sigmas) and 'rare' (palmate isochelae). The shape and architecture
is hymedesmioid, but there are long projections, and the acan-
thostyles are said to be mixed in with the tomotes. The peculiar
HYMEDESMIA BOWERBANK, 1864 acanthostrongyles are apparently arranged in bundles on the sur-
face, making the sponge very hispid. It is difficult to decide which
Synonymy of the spicules are proper to the sponge, and thus the affinities of
the sponge remained obscure. Koltun (1970) described a second
Hymedesmia Bowerbank, 1864: 190. Sclerilla Schmidt, 1868: species with the peculiar long acanthostrongyles with excessively
13. Stegxella Bowerbank, 1874b: xi. Stegilla Bowerbank, 1874b: long spines D. shikotani Koltun (1970). This apparently lacks the
xii. Stylopus Fristedt, 1885: 28. [Leptosia] Topsent, 1892b: xxii. sigmas but the chelae are clearly pictured and stated to be palmate.
Tylosigma Topsent, 1894a: 32. Ectyodesmia Topsent, 1928c: 55. Until further information is provided, it is assumed that sigmas are
Ectyostylopus Topsent, 1928c: 56. Poecilochela de Laubenfels, foreign, and that Dolichacantha is a deviating member of the
1936a: 87. HymoxeniaAlander, 1938: 72. microcionine family Acarnidae (see chapter on that family by
Hooper).
Type species
Previous review
Hymedesmia zetlandica Bowerbank, 1864 (by monotypy).
Lundbeck,191O.
Definition

Hymedesmiidae with hymedesmioid skeleton. SUBGENUS HYMEDESMIA BOWERBANK, 1864

Diagnosis Synonymy

Persistently thinly encrusting; ectosomal skeleton composed Sclerilla Schmidt, 1868: 13. Stegxella Bowerbank, 1874b: xi.
of smooth ectosomal tomotes strewn at the surface and forming Stegilla Bowerbank, 1874b: xii. Tylosigma Topsent, 1894a: 32.
Porifera • Demospongiae • Poecilosclerida • Myxillina • Hymedesmiidae 581

Key to subgenera of Hymedesmia

(1) Chelae present ........................ ...... .......... ........................... ............................ ..................... ..................................... ......... Hymedesmia
No chelae ................................................................... ........................................................ ...................................................... Stylopus

Ectyodesmia Topsent, 1928c: 55. Poecilochela de Laubenfels, Description of type species

1936a: 87. Hymoxenia Alander, 1938: 72.
Hymedesmia (Hymedesmia) zetlandica Bowerbank, 1864
(Fig. 3A-F).
Definition Synonymy. Hymedesmia zetlandica Bowerbank, 1864: 190;
Bowerbank, 1874b: 73, pis XXIX figs 1-7; Leptosia zetlandica
Hymedesmia with chelae. Topsent, 1894a: 31.

c.
F
I
" G

Fig.3. Hymedesmia. A-F, Hymedesmia (Hymedesmia) zetlandica Bowerbank (1866). A-E, SEM photos of spicules made from the holotype (scale A left,
Band E, 10 f.Lm; A right, C, and D, 1 f.Lm). F, drawing of skeleton and spicules made from a slide of the holotype. G,Hymedesmia (Hymedesmia) inflata
(Alander, 1935 as Hymoxenia), reproduced from Alander (1935 : fig. 1) (sizes see text).
582 Porifera' Demospongiae • PoeciloscIerida • Myxillina • Hymedesmiidae

Material examined. Holotype: BMNH unregistered - dry decepta Kirkpatrick (1907a: 278, not illustrated here). The holo-
specimen and slide, Bk.467, Shetland. type and slides BMNH 1908.2.5.160(b+f) were examined. This
Description. Thinly encrusting on pebbles. Lateral expan- material is scanty, thinly encrusting on a branching bryozoan.
sion up to 2 cm. Colour yellow or brownish yellow. Surface Megascleres are single acanthostyles, erect on the substrate, larger
smooth, no oscules recorded. Skeleton (Fig. 3F) hymedesmioid, ones, 423-517 X 14-20 flm surrounded by a circle of smaller acan-
with single acanthostyles erect on the substrate, head embedded in thostyles, 172-205 X 9-12flm. Ectosomal strongyles, inequiended,
a basal plate of spongin. Bundles of tomotes, 3-5 spicules in cross one end mucronate, 234-288 X 3-4.5 flm. Microscleres consist of
section, traverse the thin choanosome and fan out at the surface. a majority of normal arcuate isochelae, 19-23 flm, and reduced
Spicules (Fig. 3A-F), ectosomal and subectosomal symmetrical unguiferate and malformed ('bipocillae') presumed derivates
tylotes (Fig. 3A), 225-255 X 5-8 flm; acanthostyles (Fig. 3B), thereof, 8-15 flm. These do not constitute enough evidence for
robust, with well-developed head, fully and heavily spined, in separate generic status of this single species.
a wide size range of 80-150 X 10-20 flm, possibly divisible in two The genus Hymoxenia Alander, 1938 was erected for
categories; arcuate isochelae of two distinct morphological types, Norwegian type species (by monotypy) H. inflata Alander (1938. 72,
larger normal shaped (Fig. 3C), 20-30 flm, smaller sigma- fig. 1, here reproduced as Fig. 3G), type stated to be in the
like reduced chelae (Fig. 3D), with only two sharp teeth, shaft Gothenburg Museum. This is an encrusting species conforming
undulate or bluntly spined, 10-15 flm; sigmas (Fig. 3E), wide and entirely to Hymedesmia, including single acanthostyles erect on the
thin, in sigmodragmas, 40-70 flm. Distribution and ecology. substrate, in two size classes, 650-1400 X 20-25 flm and 150-
Shetland, S and W coast of Ireland; from deeper water, 65 m and 200 X 8-10 flm, ectosomal tylotes, 320-385 X 3-5 flm, and arcuate
downward. chelae, 50-60 flm, but having in addition an accessory spicule
Remarks. The type of Hymedesmia is rather atypical for the type of centrotylote acanthoxeas. These spicules, 290-400 X
large assemblage of species united traditionally under this genus 8-lOflm, are very probably merely 'double' versions of the
name. Neither the second category of small reduced and spined small acanthostyles, and accordingly the genus is considered a junior
chelae, nor the sigmas arranged in dragmata are found in any synonym.
known Hymedesmia, while the possession of symmetrical tylote The genus Tylosigma Topsent, 1894a: 32 was erected for type
tomotes is quite infrequent. This combination is more reminiscent species (by original designation) Hymedesmia campechiana
of Lissodendoryx (Ectyodoryx), than of mainstream Hymedesmia. Topsent (1889: 30, fig. 8C) from the Caribbean coast of Mexico.
It needs to be kept in mind that the genus characters of A slide of the type, MNHN DT1844, was examined. The species is
Hymedesmia are obviously linked with the thin growth form, and an atypical Hymedesmia by lacking proper chelae and diactinal
this may prove to be an unreliable set of characters for determining tomotes. The microscleres are large sigmas arranged in dragmas and
phylogenetic relationships. small sigmas, which are possibly reduced chelae, not unlike those
Topsent (1928c: 55) erected in a casual way a subgenus of H. zetlandica. The smaller tylostyles may be considered as
Hymedesmia (Ectyodesmia) for Hymedesmia in which the tomotes. In view of the similarities with H. zetlandica it is proposed
acanthostyles could be divided into larger main megascleres to consider Tylosigma a junior synonym of Hymedesmia
and smaller echinating acanthostyles. Since the type species (Hymedesmia).
of Hymedesmia has a smaller and a larger acanthostyle type, it
would fall to Topsent's subgenus Ectyodesmia. However, the type
species automatically belongs to the nominal subgenus, and this
makes Ectyodesmia an objective synonym of Hymedesmia SUBGENUS STYLOPUS FRISTEDT, 1885
(Hymedesmia).
The genus Stegxella Bowerbank, 1874b: xi was erected for Synonymy
type species (by original designatiom) S. saccea ascribed to
Schmidt (1864), but no such combination is found in that publica- Stylopus Fristedt, 1885: 28. [Leptosial Topsent, 1892b: xxii
tion. On the following page (p. xii) Bowerbank spelled the name (preocc.). Ectyostylopus Topsent, 1928c: 56.
as Stegilla, again not a name used in Schmidt's publication.
Technically, Stegilla is a nomen nudum. De Laubenfels (1936a: 85) Type species
declared Stegxella to be a synonym of Myxilla. It is conceivable
that both Bowerbank names are misspellings for Sclerilla Schmidt, Stylopus coriaceus Fristedt, 1885: 28 (by monotypy).
1868: 13, which de Laubenfels (1936a: 85) also declared to be
a synonym of Myxilla, but of which the type species has not been Definition
fixed yet. This is here fixed as Sclerillafilans Schmidt (1868: 13),
with lectotype specimen (dry) in MNHN, from Algiers. A Schmidt Hymedesmia without microscleres.
slide from Algiers, BMNH 1868.3.2.20, was examined, and
Topsent (1938a) provided a redescription. Spicules include Description of type species
polytylote inequiended smooth tomotes, 150-175 X 3-4 flm,
acanthostyles, 115-120 X 3 flm. The BMNH slide did not Hymedesmia (Stylopus) coriacea (Fristedt, 1885) (Fig. 4A-B).
contain proper microscleres (a few broken sigmas), but Topsent Synonymy. Hymeniacidon dujardini Bowerbank, 1866:
recorded arcuate chelae, 21-24 flm and sigmas, 25-33 flm. 244. Not Halisarca dujardini Johnston, 1842. Leptosia dujardini;
This conforms to Hymedesmia, and it is proposed to consider Topsent, 1894b. 8. Hymedesmia broendstedi Burton, 1930b: 497.
Stegxella and Stegilla along with Sclerilla as synonyms of Stylopus coriaceus Fristedt, 1885: 28, pI. II, fig. 8. Hymedesmia
Hymedesmia. coriacea; Van Soest, 1987b: 20.
The genus Poecilochela de Laubenfels, 1936a: 87 was erected Material examined. Syntype: BMNH 1927.2.22.3 -
for type species (by original designation) the Antarctic Myxilla Bohuslan, Sweden, colI. S. Loven.
584 Porifera· Demospongiae • PoeciloscIerida • Myxillina • Hymedesmiidae

Despite its unfortunate choice, de Laubenfels' type species desig- Consistency soft and fleshy. Colour orange-red, pale brown in
nation appears valid, and accordingly Hymeniacidon dujardini alcohol. Skeleton (Fig. 4D) of the ectosome consists of spiCUles
sensu Bowerbank is the type species. Bowerbank's specimen arranged in bouquets, fanning out and becoming tangentially
BMNH 1910.1.1.124 and five unregistered slides, Bk 536,537 and arranged at the surface. Walls of the areolae strengthened by verti-
539, from Strangford Lough, N Ireland, were examined. By its pos- cal palissades. Choanosomal skeleton plumoreticulate, with longi-
session of 2 sizes of acanthostyles and strongylote tomotes, this is tudinal spicule tracts connected irregularly by thinnner spiCUle
confirmed to be a clear junior synonym of Hymedesmia (Stylopus) tracts. Spicules (Fig. 4D), ectosomal tomotes, strongylote, up to
coriacea (cf. also Van Soest, 1987b). Since [Leptosia] Topsent, 260 X 6.5 fLm. Choanosomal styles smooth, with occasional
1892b was already preoccupied by Leptosia Hiibner, 1818 spines, about 400 X 13 fLm. Distribution and ecology. Antarctic and
(Lepidoptera), Stylopus remains the valid subgenus name. Subantarctic Islands; shallow-water.
Topsent, 1928c: 56 erected a subgenus Stylopus (Ectyostylopus) Remarks. The genus has been associated with Tedaniidae,
for Stylopus with two categories of acanthostyles. The type species but it lacks onychaetes, and the possession of areolated porefields
is Hymedesmia dermatus Lundbeck, 1910 (monotypy). This dis- is a strong indication it is related to Phorbas and Hymedesmia. It
tinction is considered superfluous and Ectyostylopus is considered differs from these genera in the absence of microscleres and the
a junior synonym of Hymedesmia (Stylopus). plumoreticulate architecture. It is also close to Myxodoryx but this
genus possesses echinating acanthostyles. Three other species have
been assigned to Kirkpatrickia, viz., Tedania coulmani Kirkpatrick
KIRKPATRICKIA TOPSENT, 1912 (1907a), K. spiculophila Burton & Rao (1932), and K. borealis
Koltun (1970). They differ from the type in having properly spined
Synonymy styles and in the shape of the tomotes.

Kirkpatrickia Topsent, 1912: 3.

MYXODORYX BURTON, 1929
Type species
Synonymy
Tedania variolosa Kirkpatrick, 1907a: 279 (by original
designation). Myxodoryx Burton, 1929a: 438.

Definition Type species

Hymedesmiidae with strongylote tomotes and smooth or Lissomyxilla hanitschi Kirkpatrick, 1907a: 275 (by original
spined styles in plumose but anastomosing bundles; no microscleres. designation).

Diagnosis Definition

Flabellate or digitate; pores grouped in sieve-plates (areolae). Hymedesmiidae with strongylote tomotes and lightly (spined)
Ectosomal skeleton composed of ectosomal strongylote tomotes or styles in plumose but anastomosing bundles; with a separate
strongyles in tangential tracts and whispy brushes on the surface (smaller) category of non-echinating acanthostyles with spines
(the latter around pores). Choanosomal skeleton composed only on the lower (pointed) half; no microscleres.
of loosely agglomerated primary and secondary fibres cored
by choanosomal styles, curving out towards the surface in the Diagnosis
peripheral skeleton. Megascleres choanosomal styles with
smooth or spined bases microscleres are absent. Three or four Massive growth form; ectosomal skeleton composed of
species. smooth tylotes with mucronate bases forming tangential and
oblique tracts at the surface; choanosomal skeleton confused retic-
Previous review ulate, with branching fibres cored by smooth choanosomal styles
and echinated by acanthostyles with verticillate spination;
Koltun, 1964b. microscleres are absent. A single species.

Description of type species Description of type species

Kirkpatrickia variolosa (Kirkpatrick, 1907a) (Fig. 4C-D). Myxodoryx hanitschi (Kirkpatrick, 1907a) (Fig. 5A-C).
Synonymy. Tedania variolosa Kirkpatrick, 1907a: 279; Synonymy. Lissomyxilla hanitschi Kirkpatrick, 1907a: 275;
Kirkpatrick, 1908c: 32, pI. XXI fig. I , pI. XXV fig. I; Kirkpatrickia Kirkpatrick, 1908c: 26, pI. XXII fig. 7, pI. XXVI fig. 4; Myxodoryx
variolosa; Topsent, 1912: 3. hanitschi; Burton, 1929a: 438.
Material examined. Holotype: BMNH 1908.2.5.162. Para- Material examined. 'Cotype': BMNH 1908.2.5.132 (includ-
type: BMNH 1908.2.5.164 - from Winter Quarters, Antarctica. ing 2 slides BMNH 1908.2.5.132 c+e) - 'Discovery' collection,
Paratype: BMNH 1908.2.5.163 (not examined). Antarctic. The type material consists of three specimens, two of
Description. Thickly flabellate or digitate fronds, up to 18 which are from the same location, Coulman Island, the other,
cm high, 13 cm wide, ending in sphinctrate oscules (Fig. 4C). smaller specimen is from Barrier. The latter specimen is here cho-
Surface covered with slightly raised areolate porefields. sen as the lectotype, because it has a complete ectosome.
Porifera· Demospongiae • Poecilosclerida • Myxillina • Hymedesmiidae 583

- -- - --<200

B
150

D
I
Fig. 4. A-B, Hymedesmia (Sty/opus) coriacea (Fristedt, 1885). A,drawing of shape, skeleton and spicules of the type material, reproduced from Fristedt
(1885, pI. II fig . 8) (sizes see text). B, drawing of skeleton and spicules made from a slide of one of the syntypes. C-D, Kirkpatrickia variolosa
(Kirkpatrick, 1907a as Tedan ia). C, drawing of the holotype (reproduced from Kirkpatrick's pI. XXI fig . 1; size see text). D, drawing of skeleton and
spicules made from a slide of the holotype.

Description. Thinly encrusting on other organisms (barna- latter was ascribed to Johnston, 1842. Bowerbank misinterpreted
cles, brachiopods, tubeworms (see type of Fristedt, reproduced Halisarca dujardini Johnston, 1842 for the present species, so the
here as Fig. 4A), algae, corals). Surface smooth, mucous. name dujardini cannot be used for it.
Consistency firm. Skeleton (Fig. 4A-B) of the ectosome consists The genus [Leptosia] Topsent, 1892b: xxii was originally
of brushes of tomotes fanning out to form tangential single erected for a group of three species consisting of Hymedesmia
spicules; choanosomal skeleton consists of single acanthostyles zetlandica, Hymeniacidon dujardini Bowerbank, 1866 (not Halisarca
erect on the subtstrate, from which arise ill-defined bundles of dujardini Johnston, 1842), and Leptosia exilis Topsent, 1892b
tomotes. Spicules (Fig. 4A-B), strongylote or anisotylote tomotes (=Forcepia luciensis Topsent, 1890b). De Laubenfels (1936a: 88),
about 200 X 2.5 f.Lm (range given by Van Soest, 1987b: 140-280 X for unexplained reasons, designated H. dujardini as type species,
2-4 f.Lm). Acanthostyles in a wide size range, probably divisible in whereas Topsent himself (e.g., 1892a, 1904b, 1928c) expressly
two categories, 120-180 X 5-8 f.Lm (range given by Van Soest, mentioned chelae in his definitions of [Leptosia] . In 1928, Topsent,
1987b: 56-312 X 1.7-8 f.Lm). Distribution and ecology. Sweden when confronted with the fact that Hymedesmia was a valid genus
(Fristedt), E Atlantic S to Mediterranean and W Africa, on stones, covering more-or-less his concept of [Leptosia], restricted the use
shells and corals, shallow-water down to 100 m. of it to species with anchorate chelae (a concept here covered by
Remarks. The reason for preferring the name coriacea the genus Hymenancora), but this is impossible because the origi-
Fristedt, 1885 over dujardini sensu Bowerbank, 1866, is that the nal series of sponges did not contain species with anchorate chelae.
Porifera· Demospongiae • Poecilosclerida • Myxillina • Hymedesmiidae 585

A _-_.50 I
c 00

B
Fig. 5. Myxodoryx hanitschi (Kirkpatrick, 1907a as Lissomyxilla). A, drawing of habit reproduced from Kirkpatrick (l908c, pI. XXII fig. 7) (sizes see
text). B, drawing of spicules reproduced from Kirkpatrick (1908c, pI. XXVI fig. 4) (sizes see text). C, drawing of skeleton and spicUles made from a slide
of the holotype.

Description. Lobate sponges (Fig. 5A), with transparent Type species

dermal membrane roofing over extensive subdermal openings.
Surface areolae are not conspicuously developed. Colour whitish Phorbas amaranthus Duchassaing & Michelotti, 1864 (by
grey. Skeleton (Fig. 5C) of the periphery consists of brushes subsequent designation; de Laubenfels, 1936a: 63).
of ectosomal spicules fanning out at the surface and becoming
tangentially arranged. Choanosomal skeleton plumose, confused, Definition
with longitudinal tracts echinated by acanthostyles. Spicules
(Fig. 5B-C), ectosomal tomotes tylote-like with pointed Hymedesmiidae with plumose tracts of tomotes and/or acan-
mucronate endings, 280--(317.4)-350 X 6-(7.6)-10 /-Lm. Styles thostyles, echinated by acanthostyles (but these may be absent);
smooth, curved near the blunt end, sharp-pointed, 385-(449.2)- microscleres normal-shaped chelae and sigmas, which may be
503 X 15-(19.3) - 22 /-Lm. Echinating acanthostyles, with long absent.
spines at the head and near the pointed end, but virtually smooth
inbetween, 210--(260.3)-288 x 12-(13.6)-18 /-Lm. Distribution and Diagnosis
ecology. Antarctic; deep-water.
Remarks. The species was originally assigned to Thickly encrusting, massive or ramose sponges. Pores grouped
Lissomyxilla Hanitsch, 1894a, which is a junior synonym of in sieve-plates (areolae); ectosomal skeleton with a crust of
Tethyspira Topsent, 1890b (Halichondrida: Dictyonellidae). So far isochelae and diactinal spicules which form fans disposed at right
the genus remains mono typical, and also somewhat isolated from angles and tangential to the surface; heavily spined acanthostyles
other Hymedesmiidae. No closely related genera are apparent. core plumose or plumo-reticulate choanosomal tracts, or these may
be replaced by smooth diactina1 tomotes; echinating acanthostyles
also heavily spined; microscleres are arcuate isochelae and sigmas,
PHORBAS DUCHASSAING & MICHELOTTI, 1864 both of which may be absent. About 70 species.

Synonymy Taxonomic remarks

Phorbas Duchassaing & Michelotti, 1864: 92. Anchinoe Gray, Voultsiadou-Koukoura & Van Soest (1991b) demonstrated
1867a: 535. [Pronax) Gray, 1867a: 536. Suberotelites Schmidt, 1868: that the alleged distinction between Phorbas, Anchinoe and
12. Plumohalichondria Carter, 1876: 236. Clathrissa Lendenfeld, Stylostichon (viz., skeletal columns consisting of tomotes only or
1888: 217. Stylostichon Topsent, 1892a: 111. Lissopocillon Ferrer- of acanthostyles only) intergrades in the various species assignable
Hernandez, 1916. 32. Grayax de Laubenfels, 1936a: 63. Merriamium to these genera. Consequently, it was proposed to merge these gen-
de Laubenfels, 1936a: 83. Pronaxella Burton, 1931d: 168. era and this is followed here. Perhaps a subdivision of the now
Bipocillopsis Koltun, 1964b: 79. Podotuberculum Bakus, 1966b: 505. enlarged genus Phorbas is possible based on the possession of a
586 Porifera· Demospongiae • Poecilosclerida • Myxillina • Hymedesmiidae

c
Fig. 6. Phorbas. A-B, Phorbas amaranthus Duchassaing & Michelotti (1864). A, habit of holotype (photo K. Riitzler). B, drawing of skeleton and
spicules reproduced from Van Soest (l984b, text-fig. 34). C, microphoto of spicules reproduced from de Laubenfels (1936a, pI. II fig. 1 as Merriamium
tortuganensis). D-F, Phorbas perarmatus (Bowerbank, 1866 as Hymeniacidon), type of Anchinoe Gray, 1867a, SEM photos of spicules made from the
type (scale 10 f,Lm; except D, 100 f,Lm). G, Phorbas nexus (Koltun, 1964b as Bipocillopsis), drawing of habit and spicules reproduced from Koltun's fig. 19
(sizes see text).

single acanthostyle size category (Phorbas s.s.) or two size Description of type species
categories, a larger acanthostyle coring the choanosomal bundles,
a smaller echinating the columns (Suberotelites). This is best Phorbas amaranthus Duchassaing & Michelotti, 1864
postponed until the species content of Phorbas s.l. is properly (Fig. 6A-C).
revised. The genus Acanthodoryx Levi, 1951, on paper at least, Synonymy. Phorbas amaranth us Duchassaing &
appears to be close to Phorbas. However, the type species Michelotti, 1864. 92, pI. XXI fig. 1; Van Soest, 1984b: 86, pI. VI
A. fibrosa Levi, 1951 lacks the surface areolae, and the skeleton is fig. 7, text-fig. 34. Merriamium tortugasense de Laubenfels,
plumoreticulate rather than plumose; echinating acanthostyles are 1936a: 83, pI. 11, fig. 1.
lacking. For these reasons, Acanthodoryx is assigned to the family Material examined. Mus. Torino POR. 59 - St. Thomas
Coelosphaeridae, and considered a subgenus of Lissodendoryx (see (cf. Fig. 5A). Other material. Many additional specimens from
chapter Coelosphaeridae). various Caribbean localities.
Porifera • Demospongiae • Poecilosclerida • Myxillina • Hymedesmiidae 587

Description. Encrusting, several mm in thickness, with dis- the type of Phorbas, and accordingly Clathrissa is considered
tinctive areolate surface patterns. Consistency soft, easily dam- a junior synonym.
aged. Gives off a red exudate when squeezed. Skeleton (Fig. 6B) of The genus Lissopocillon Ferrer-Hernandez, 1916 was erected
the ectosome a tangential arrangement of loose tomotes; walls of for type species (by monotypy) L.dendyi Ferrer-Hernandez, 1916:
areolae strengthened by parallelly arranged tomotes; microscleres 32, fig. 10 (here reproduced as Fig. 7E), a lobate-mammillate
surround the pores. Choanosomal skeleton of thick (ca. 100 jJ-m in sponge of 15 X 12 X ?cm. Surface provided with areolated pore
diameter) columns of tomotes mixed with increasing numbers of fields and oscules of the same size. Colour probably reddish.
acanthostyles towards the base; columns lightly echinated by acan- Skeleton of diactinal oxeote tomotes, 300-320 X 6-8 jJ-m, arranged
thostyles. Spicules (Fig. 6B), robust smooth oxeote tomotes, in the ectosome and in the choanosomal tracts, in the
straight with hastately pointed apices, 235-266 X 3.5-5.5 jJ-m. bundles partly replaced by large acanthostyles, averaging 256 X
Large acanthostyles with occasional strongylote modifications, 12-14jJ-m. The bundles are echinated by shorter acanthostyles,
129-182 X 5-8 jJ-m. Shorter echinating acanthostyles, 91-112 X averaging 128 X 6-12 jJ-m. Chelae in two size categories, normal
4.5-7.5 jJ-m. Robust arcuate isochelae with short alae, 20-24 jJ-m. arcuate isochelae, about 26 jJ-m, and smaller malformed chelae, up
Distribution and ecology. Caribbean; encrusting the base of corals to 20 jJ-m. The latter were called 'pocillos' by Ferrer-Hernandez
in the reef environment, 6-33 m. and constitute the generic character. Such malformed chelae are
Remarks. A common genus with cosmopolitan distribution. not unusual in Myxillina and may not even be a species character,
Anchinoe Gray, 1867a: 535 was erected for type species (by mono- let alone a genus character. In most features L. dendyi conforms
typy) Hymeniacidon perarmatus Bowerbank, 1866 from Shetland. to Phorbas lieberkuehni Burton (1930c: 532, as Hymedesmia),
The holotype, BMNH 1910.1.1.144 was re-examined. This is a a replacement name for Halichondria Jasciculata sensu Lieberktihn,
thin crust on a stone. Surface provided with conspicuous areolate 1859: 522. The consequence of reassignment of Lissopocillon
porefields. The skeleton consists of an ectosomal tangential skele- dendyi to Phorbas is that it will have to be given a new name, since
ton of intercrossing tomotes of 445-510 X 4-9 jJ-m, with the walls the combination Phorbas dendyi is already reserved for the species
of the areolae supported by parallel bands of tomotes. Ectosomal originally described as Stylostichon dendyi Topsent, 1913. It is here
skeleton is carried by choanosomal plumose bundles of tomotes proposed to rename Lissopocillon dendyi as Phorbas Jerrerhernan-
(Fig. 6D) with a low number of long acanthostyles (Fig. 6E), dezi n.sp., although it may tum out to be a junior synonym of
283-305 X 4-8 jJ-m, mixed in. The bundles of 60-100 jJ-m in diam- Phorbas lieberkuehni Burton when type material is re-examined.
eter are profusely echinated by short acanthostyles (Fig. 6E), The genus Merriamium de Laubenfels, 1936a was erected
142-165 X 4-7 jJ-m. Arcuate chelae (Fig. 6F) in two size cate- for type species (by monotypy) Merriamium tortuganensis de
gories, 62-92 and 28-44 jJ-m. This sponge conforms closely to the Laubenfels (1936a: 83, pI. 11 fig. 1, here reproduced as Fig. 6C).
type species of Phorbas. The type specimen, USNM 22418 from Florida, was reexamined.
The genus Bipocillopsis Koltun, 1964b: 79 was erected for It has surface areolae and a skeleton consisting of bundles of
type species (by monotypy) the Antarctic B. nexus Koltun (1964b: oxeote tomotes gradually converted to bundles of acanthostyles;
80, fig. 19, here reproduced in Fig. 6G). No material was examined microscleres are in a single size category of arcuate chelae. This is
(stated to be housed in the Zoological Institute of the Russian an obvious junior synonym of Phorbas amaranth us (see also
Academy of Sciences). This is a mass of branches with irregular Van Soest, 1984b: 88), making Merriamium an objective synonym
surface provided with porefields (clearly pictured by Koitun, but of Phorbas as both share the same type species. De Laubenfels'
not mentioned in his description). Tomotes are stylote, 370-480 X (1939) monograph of Merriamium contains a mixture of
5-7 jJ-m, and form anastomosing bundles, echinated by acan- Hymedesmia, Phorbas and Lissodendoryx (Ectyodoryx) species.
thostyles, spined lightly, 520-720 X 15-25 jJ-m. Chelae, 16-20 jJ-m, The genus Plumohalichondria Carter, 1876 was erected
are peculiar, but likely to originate from arcuate chelae in view of for type speces (by monotypy) Plumohalichondria microcionides
their resemblance to the reduced chelae of Fibulia. The drawing Carter (1876: 236, pI. XII fig. 11 pI. XV fig. 30). The type speci-
and the description of Koltun do not entirely match, and it is possi- men, BMNH 1882.7.28.47, labelled 'Porcupine coIl. stat. 51, 440
ble that there is a second smaller category of acanthostyles pictured fms, between Scotland and Faroe Islands' was re-examined (cf.
in his fig. 19 (here reproduced), but not mentioned in the text. If the Fig. 7F). It is a lobe with irregular mammillated surface and firm
choanosomal tracts are indeed made up of stylotomotes, as is consistency. Skeleton (Fig. 7G) consisting of plumose bundles of
apparently what Koltun describes, then the peculiar reduced chelae sharply pointed oxeote tomotes, 430-630 X 10-13 jJ-m, towards
are the only difference with Phorbas, and accordingly the genus is the interior replaced by long club-shaped acanthosty!es, spined
considered a junior synonym. profusely on the head but virtually smooth along the shaft,
The genus Clathrissa Lendenfeld, 1888 was erected for type 695-1140 X 20-28 jJ-m. The bundles are echinated by short acan-
species C. arbuscula Lendenfeld, (1888: 217, pI. V fig. 2) (by sub- thostyles, spined all-over, 235-300 X 12-14 jJ-m. Chelae (Fig. 7G)
sequent designation; Hallmann, 1912: 146). The following type are arcuate, and occur in a large size range, probably divisible in
and other material was examined, BMNH 1886.12.12.22 (labeled two categories, 65-125 and 42-63 jJ-m. The species is only remark-
'Herkunft Port Jackson 1140'), slide BMNH 1954.2.10.73 and able for the large size of acanthostyles and chelae.
slide 1954.2.12.52 (both Dendy collection). All that material is The genus Podotuberculum Bakus, 1966b was erected for
identical and concerns a mass of papillar projections, possibly type species (by monotypy) P. hoffmanni Bakus (1966b: 505, fig.
constituting fused branches, size 10 cm high, 8 cm in diameter 23, here reproduced as Fig. 8A). A paratype, BMNH 1964.12.1.4,
(Fig. 7A). The skeleton consists of plumose bundles of oxeote from San Juan Island, W coast of North America, was re-examined.
tomotes (Fig. 7B), 192-230 X 3-5 jJ-m, echinated by acanthostyles This is encrusting and its surface bears 'tubercles', i.e., areolae.
(Fig. 6C), 100-126 X 3-8 jJ-m, towards the interior grading into Spicule sizes and shape conform closely to those of Myxilla versi-
bundles of long acanthosty!es 180-207 X 7-9 jJ-m. Chelae are the color califomiana de Laubenfels (1932:81), with strongylote
usual arcuate type (Fig. 7D), 20-30 jJ-m. This conforms closely to tomotes of about 250 X 4-8 jJ-m, styles with spined heads, about
588 Porifera' Demospongiae • Poecilosclerida • Myxillina • Hymedesmiidae

G
Fig. 7. Phorbas species. A-D, Phorbas arbuscula (Lendenfeld, 1888 as Clathrissa). A, habit ofholotypereproduced from Lendenfeld's pI. V fig. 2 (sizes
see text). B-D, SEM photos of spicules made from the type (scale 10 J.Lm). E, Phorbas Jerrerhernandezi n.sp., replacement name for Phorbas dendyi
(Ferrer-Hernandez, 1916 as Lissopocillon), drawing of spicules reproduced from Ferrer-Hernandez's fig. 10 (sizes see text). F-G, Phorbas microcionides
(Carter, 1876 as Plumohalichondria). F, fragment of type specimen (scale I cm). G, drawing of skeleton and spicules made from a slide of the type.

250 X 12 fLm, some of which take up an echinating position while Montagu's Type Spec. Hab.?' , were examined. The cuts and
other are mixed with the tomotes coring the choanosomal bundles. spicules conform to Bowerbank's description. It is a massively
No microscleres. On paper this species comes close to encrusting orange sponge, with a tuberculate surface, but the areo-
Kirkpatrickia, but in that genus the choanosomal tracts are con- lae are very small and indistinct. Ectosomal skeleton a tangential
nected by secondary tracts, and no echinatig spicules are found. It mass of intercrossing oxeote tomotes (Fig. 8C), 140-165 X 3-6 fLm.
conforms to Phorbas. Choanosomal skeleton (Fig. 8B) is a system of plumose bundles
The genus [Pronax] Gray, 1867a: 536 was erected for type of acanthostyles (Fig. 8D), 160-200 X 8-12 fLm, anastomosing
species (by monotypy) Hymeniacidon plumosa Bowerbank, 1866: at sharp angles, and profusely echinated by shorter acanthostyles
195 (=Spongia plumosa Montagu, 1818). The specimen here pro- (Fig. 8D), 70-100 X 4-7 fLm. Spongin well-developed. Arcuate
posed as lectotype, BMNH 1930.7.3.484 (certainly Johnston's isochelae (Fig. 8E) of 15-20 fLm. The bundles of acanthostyles
specimen, possibly Montagu's specimen) and slides of Montagu's contain a core of oxeas in their peripheral parts, thus testifying of
specimen consisting of 5 unregistered slides, three of which the artificiality of distinguishing a separate genus for Phorbas-like
are labeled 'Spongia plumosa Mont, portions from various parts species with a predominance of acanthostyles in the choanosomal
of type with Dr Grant. Devon Coast. Bk 944, Bk945 , , and two tracts. The name [Pronax] is preoccupied by Pronax Gray, 1867a:
are labeled 'Spongia plumosa Mont. (Microciona plumosa Bk) 526 (Clionaidae).
Porifera· Demospongiae • Poecilosclerida • Myxillina • Hymedesmiidae 589

,.
= A
B

I
..,

G -~.I~

Fig.8. Phorbas species. A, Phorbas hoffmanni (Bakus, 1966b as Podotuberculum), dmwing of skeleton and spicules reproduced from Bakus' text-fig. 23
(sizes see text). B-E, Phorbas plumosus (Montagu, 1818 as Spongia), type of Pronax Gray (1867a) and Pronaxella Burton (1931d). B, drawing of skele-
ton and spicules made from a slide of the type. C-E, SEM photos of spicules made from the type (scale 1 fLm; except D, 10 fLm). F-G, Phorbas dendyi
(Topsent, 1892a as Stylostichon). F, drawing of halo type reproduced from Topsent (1 892a. pI. II fig.6) (size see text). G, drawing of skeleton and spicules
reproduced from Topsent (1 892a. pI. XI figs 9-10) (sizes see text).

The genus Pronaxella Burton, 1931d: 168 was erected as soft consistency. Topsent maintains the skeletal columns are
a replacement name with the same species Spongia plumosa entirely filled with acanthostyles, but Norwegian specimens in
Montagu (1818) as type (by original designation). Since Pronax ZMA similar to the Azoran type specimen demonstrate the
plumosus is now assigned to Phorbas, Pronaxella likewise presence of mixtures of acanthostyles and tomotes in peripheral
becomes a junior synonym. De Laubenfels (I 936a: 63) was parts of the skeletal columns. Columns profusely echinated by
apparently unaware of Burton's proposed replacement name, and acanthostyles (Fig. 8G) of the same size range as those in the
he proposed Grayax as a second replacement name, likewise columns. Spiculation. Strongylote tomotes (Fig. 8G), 210-300 X
a junior synonym of Phorbas. 4-8/-Lm. Acanthostyles (Fig. 8G), about 180-350 X 10-18 /-Lm.
The genus Stylostichon Topsent, 1892a: 111 was erected for Arcuate isochelae (Fig. 8G), about 15-56 /-Lm.
type species (by subsequent designation; Lundbeck, 1909) The genus Suberotelites Schmidt, 1868 was erected for type
S. dendyi Topsent (1892a: 112, pI. II fig. 6, pI. XI figs 9-10, here species S. mercator Schmidt (1868: 12, pI. II fig. 15) (by mono-
reproduced as Fig. 8F-G) (see Voultsiadou-Koukoura & Van Soest typy). A fragment of Schmidt's type was re-examined in MNHN.
(1991b) for a justification of assuming S. dendyi as the type). The species was also redescribed by Topsent (1938a: 12), and
Stylostichon dendyi is a lobate to arborescent sponge (Fig. 8F) of Pulitzer-Fin ali (1983: 565 as Anchinoe, fig. 65, here reproduced
590 Porifera' Demospongiae • PoeciloscIerida • Myxillina • Hymedesmiidae

B I.0

Fig. 9. Phorbas mercator (Schmidt, 1868 as Suberotelites). A, photo of habit, reproduced from Pulitzer-Finali (1983 fig. 65) (size see text). B, drawing of
skeleton and spicules made from a fragment of the holotype.

as Fig. 9A). The sponge is a greyish crust, with some digitate pro- clustered around the bases of the long spicules; microscleres are
jections of 1.5 cm long. The skeleton (Fig. 9B) consists of poly- arcuate isochelae. Three or four species.
spicular bundles of 50-100 fLm diameter with a coating of spongin,
anastomosing irregularly but frequently. The bundles are predomi- Previous review
nantly cored by anistrongylote tomotes (Fig. 9B), 200-250 X
2-4 fLm, but occasionally acanthostyles (Fig. 9B), 190-205 X Topsent, 1928c.
7.5-8 fLm are mixed in. The bundles are echinated sparingly
by shorter acanthostyles (Fig. 9B), 80-160 X 4.5-6 fLm. No Description of type species
microscleres. The lack of chelae is considered a species character
and accordingly Suberotelites is a junior synonym of Phorbas. Plocamionida ambigua (Bowerbank, 1866) (Fig. lOA-F).
Synonymy. Microciona ambigua Bowerbank, 1866: 136;
Bowerbank, 1874b: 65, pI. XXXV, figs 1-9; Hastatus ambiguus.
PLOCAMIONlDA TOPSENT, 1927 Fristedt, 1885: 31, pI. II, fig. 1; Plocamia ambigua; Topsent, 1894b:
8; Plocamionida ambigua; Topsent, 1927b: 16; Topsent, 1928c: 303.
Synonymy Material examined. Lectotype (designation herein):
BMNH 1910.1.1.65 (dry) - Shetland, encrusting shell.
Plocamionida Topsent, 1927b: 16. Hymendectyon Bakus, Paralectotype: BMNH 1930.7.3.227 - Shetland, encrusting Lima
1966b: 476. shell, both Norman collection; two unregistered slides, Bk 435.
Description. Thin sheets, up to ca. 20cm across (the lecto-
Type species type specimen is now a little piece of shell, 2.5 X 1.5 X 0.5 cm
with a crust on it). Surface even, hispid; reported as tuberculate
Microciona ambigua Bowerbank, 1866: 136 (by subsequent in Mediterranean specimens. Oscules numerous and small.
designation; de Laubenfels, 1936a: 77). Consistency soft, but difficult to determine because very thin.
Colour pale red, orange, yellow or ochre. Grey-brown in alcohol,
Definition light brown dried. Skeleton (Fig. IOF) a basal layer of acan-
thostrongyles lying tangentially to the substratum or forming an
Hymedesmiidae with plocamiid basal skeleton. Megascleres irregular thin basal reticulation ('plocamiid' skeleton) with short,
mucronate tomotes, long acanthostyles, and basal acantho- entirely spined acanthostyles oriented more-or-less at right angles
strongyles. Microscleres arcuate chelae. to the substratum with their heads towards the base. Larger acan-
thostyles and much larger almost smooth styles project from this
Diagnosis basal layer, the latter protruding through the ectosomal region.
There is a surface layer of tomotes arranged as bouquets or some-
Encrusting. Ectosomal skeleton contains paratangential what tangentially. Chelae are scattered sparsely in a layer near the
brushes of tomotes. Choanosomal skeleton with hymedesmioid surface. Spicules (Fig. lOA-G), straight, very slender tomotes
architecture, with long smooth styles erect on the substrate, singly or (Fig. lOA), in the type 235-(252.7)-274 X 2.5-3 fLm, ca. 200-
in bundles, and protruding through the surface. Long acanthostyles 420 fLm in length over the range of the species. They may have
and short acanthostrongyles forming a reticulate basal skeleton, short and simple mucrons at their ends. The principal spicules of the
and also short echinating acanthostyles erect on the substrate and main skeleton are large styles (Fig. lOB), lightly acanthose at the
Porifera' Demospongiae • PoeciioscIerida • Myxillina • Hymedesmiidae 591

1 G
F
Fig. 10. Plocamionida. A-F, Plocamionida ambigua (Bowerbank, 1866 as Microciona). A-E, SEM photos of spicules made from the lectotype (scales A left
and right, Bright, D left, 10 jim; scale B left, Cleft, 100 jim; scale C right, D right, E, 1 jim). F, drawing of skeleton and spicules made from a slide of the
lectotype. G, Plocamionida lyoni (Balms, 1966b as Hymendectyon), drawing of skeleton and spicules reproduced from Bakus' text-fig. 16 (sizes see text).

head but otherwise smooth, measuring 500-960 X 6--14 J..Lm in up near the San Juan Archipelago, W coast of North America.
the type, up to 2000 X 28 J..Lm in other specimens. Auxiliary spicules Skeleton consists of a dense plocarniid skeleton of small acan-
are smaller, entirely spined acanthostyles (Fig. 1OC), in two distinct thostyles, spined all over, 100-128 X 7-9 J..Lm, upon which longer
categories, the larger surrounding the heads of the main styles, acanthostyles, with spined only in the upper half, 160-345 X
311-(353.0)-432 X 7-(8.2)-9 J..Lm in the type, the smaller echinat- 12-16J..Lm, are arranged in an erect outwardly projecting position.
ing the nodes of the basal skeleton, 149-(190.4)-235 X 5-(5.8)-7 Their heads are surrounded by a bouquet of smaller acanthostyles.
J..Lm in the type; overall acanthostyle sizes over the range of the Ectosomal spicules are smooth oxeote tomotes, 133-153 X 4 J..Lm.
species about 155-450 J..Lm. Acanthostrongyles (Fig. IOD) entirely Chelae are arcuate, 25-35 J..Lm. Although several discrepancies with
spined, curved, with often unequally swollen ends, may be distorted P. ambigua are apparent (small size of the acanthostyles, absence
and variable; spines have a crown of smaller spines, of acanthostrongyles), the plocamiid skeleton in combination with
98-(102.8)-110 X 3.5-(4.2)-5 J..Lm; 70-180 J..Lm in other specimens. the full spicule complement, make this species a likely member
Microscleres arcuate chelae (Fig. 10E) with curved shafts, of Plocamionida, and accordingly Hymendectyon is considered
20-(23.0)-26 J..Lm in the type, 19-37 J..Lm in other specimens. a junior synonym.
Distribution and ecology. NE Atlantic, Mediterranean; on bivalve
shells, stones, bedrock and saddle oysters, 40-360 m.
Remarks. A further N Atlantic species of Plocamionida
is P. microcionides (Carter, 1876, as Hymeraphia). The genus PSEUDOHALICHONDRIA CARTER, 1886
Hymendectyon Bakus, 1966b was erected for type species (by mono-
typy) Hymedectyon lyoni Bakus (1966b: 476, pI. ITa, text-fig. 16, Synonymy
here reproduced as Fig. lOG) (no material examined). According
to Bakus, this is a 'persimmon' -coloured encrustation dredged Pseudohalichondria Carter, 1886c: 454.
592 Porifera· Demospongiae • Poecilosclerida • Myxillina • Hymedesmiidae

...-
It: [1 ' fI ..

---...
C r
••

II,~ 111 !1I/~

.-.. ~ •.-.~..
I V. (.e,
~\_; ~~ ~ ... I ~
III/
II".
.1.0
I
"" v
I
\~ ';t==~::l ~ • "
fir. ~~ftJit i 'I'
\tTl) 1------,... . vu D E
F
Fig. 11. A-D, Pseudohalichondria clavilobata Carter, 1886c. A, habit of holotype (scale 1 cm). B-C, SEM photos of spicules made from the holotype
(scale IIJ-m). D, drawing of skeleton and spicules made from a slide of the holotype. E, Spaniop/on annaturum (Bowerbank, 1866 as Hymeniacidon),
drawing of skeleton and spicules made from a slide of the holotype. F, Spaniop/on werthi (Hentschel, 1914 as Acanthoxa) drawing of spicules reproduced
from Hentschel's pI. VII fig. 4 (sizes see text).

Type species Previous reviews

Pseudohalichondria clavilobata Carter, 1886c: 454 (by Carter, 1886c; de Laubenfels, 1936a.
monotypy).
Description of type species
Definition
Pseudohalichondria clavilobata Carter, 1886c (Fig. I lA-D).
Hymedesmiidae with plumose columns of smooth tomotes, Synonymy. Pseudohalichondria clavilobata Carter, 1886c:
without acanthostyles, with modified spined chelae. 454, pI. X figs 6-9.
Material examined. Holotype: BMNH 1886.12.15.81
Diagnosis (including 2 slides) - Port Philip Heads.
Description. Massive-Iobose (Fig. lOA), substipitate, size
Massive, flabellate; ectosomal skeleton with subtylostyles; 15 X 10 X IOcm. Surface smooth with very slightly raised small
choanosomal skeleton composed of thick fibres cored with sand pore-areas. Compressible, but compact consistency, fleshy. Colour
and subtylostyles; microscleres are spined arcuate isochelae. yellowish-white in alcohol. Skeleton (Fig. lID) of the ectosome
Porifera' Demospongiae • Poecilosclerida • Myxillina • Hymedesmiidae 593

a single-spicule palisade of megascleres strengthening the porefields Description of type species

and ranging individually slightly above the surface. Microscleres
evenly distributed around the ostia. Choanosomal skeleton feebly Spanioplon armaturum (Bowerbank, 1866) (Fig. lIE).
developed consisting of single widely spaced spongin fibres with a Synonymy. Hymeniacidon armatura Bowerbank, 1866:
core of megascleres and sand grains, 50-300 fLm thick depending of 183; Bowerbank, 1874b: 88, pI. XXXIV figs 12-15; Spanioplon
the amount of sand, running straight from the interior to the surface. armaturum; Topsent, 1890b: 31. Hymedesmia stephensi Burton,
Few interconnecting fibres of about the same thickness. Distance 1930b: 494.
between the fibres may be > 1000 fLm or more. Spicules, smooth Material examined. Holotype: BMNH 1930.7.3.125 -
styles (Fig. liB), interpreted as tomotes, found singly at the surface Strangford Lough, thin crust on Pecten shell. Schizotypes (slides):
and filling the fibres, 200-250 X 1.5-4 fLm. Sigmoid, strongly BMNH unregistered-labeled 'Hymeniacidon armaturum
warty-undulating microscleres (Fig. 11C) assumed to be derived Strangford Lough, Dr Dickie, #Bk. 509 (R1835 and R1836)'.
from arcuate chelae, 12-18 fLm. Distribution. South Australia. Description. Thinly encrusting, up to 1 cm thick and 9 cm in
lateral expansion. Surface smooth with obvious exhalant canals,
Remarks which may become grooves when the sponge is taken out of the
water. In places the surface is conulose. Few oscules, and no obvi-
This genus shares the peculiar chela-modification with ous areo1ate porefields. Consistency soft. Colour greyish yellow.
Acanthancora but is otherwise radically different in structure, Skeleton (Fig. 11E) consists of 1umose bundles of styles rising up
possessing elaborate shape and lacking acanthostyles. Even the from a basal plate of spongin. They integrade towards the periph-
chelae, though both technically 'spined', are morphologically dis- ery with bundles of tomotes which fan out when reaching the sur-
tinct. A second species, P. fibrosa was described by Whitelegge face, forming a tangential skeleton. Loose styles and many
(1901) from the same geographic area; its skeleton is a bit tighter- acanthostyles are scattered singly in the choanosome. The acan-
meshed, but the main difference with P. clavi/obata is the growth thostyles are not echinating and are simulating microscleres.
form which is distinctly arborescent. Spicules (Fig. lIE), tomotes, irregularly bent or curved,
inequiended, ends tylote or mucronate, 186-(221.3)-242 X
2.5-(3.85)-5 fLm. Styles, in majority smooth and with a faint tyle,
SPANIOPLON TOPSENT, 1890 occasionally with a spined head, 258-(269.7)-282 X 5-(7.65)-
10 fLm. Acanthostyles thin, straight, long-spined except in the
Synonymy middle part, 42-(50.6)-61 X 2-(2.95)-3.5 fLm. Distribution and
ecology. European coasts from Norway to the Mediterranean, on
Spanioplon Topsent, 1890b: 31. Acanthoxa Hentschel, 1914: 96. bivalve shells and stones, down to 50m.
Remarks. A second species, S. fertile Topsent (1892a: 117,
Type species pI. XI fig. 11) differs in spicule sizes and in possessing centroty-
lote oxeas as tomotes, which is unusual in Myxillina. The genus
Hymeniacidon armatura Bowerbank, 1866: 183 (by original name was also used for species with chelae. S. cheliferum
designation). Hentschel (1912: 368; type specimen SMF 1571 re-examined),
which is assigned to Clathria (cf. also Hooper, 1996a), and S. oscu-
Definition losum (Topsent, 1892b: xxiii, as Amphilectus), which conforms to
Phorbas. De Laubenfels (1936a) considered Spanioplon to be a
Hymedesmiidae with non-echinating accessory acanthostyles junior synonym of Stylopus, but this has echinating acanthostyles
or acanthoxeas. No chelae. different from the arrangement in S. armaturum.
The genus Acanthoxa Hentschel, 1914 was erected for type
Diagnosis species (by monotypy) the Antarctic A. werthi Hentschel (1914:
96, pI. VII fig. 4, here reproduced as Fig. UF), a massive species
Encrusting. Ectosomal skeleton composed of smooth aniso- of 3 X 2 X 2cm. It has ectosomal stylote tomotes, 312-360 X
tomotes, smooth or microspined choanosomal styles, small acan- 7-8 fLm, carried by plumose tracts of acanthosty1es, 456-504 X
thostyles or acanthoxeas dispersed (possibly homologues of 11-15 fLm, apparently becoming arranged in a network near the sub-
echinating spicules or true microscleres), chelae absent. Three strate. The accessory spicules are acanthoxeas, 120-144 X 2.5 fLm,
species. scattered in the choanosome. The species shares the absence of
echinatinglbasal acanthosty1es, accessory acanthose spicules, and
Previous review the absence of microscleres with Spanioplon. Boury-Esnault &
Van Beveren (1982: 100, pI. XVII fig. 65, text-fig. 29) proposed to
Van Soest, 1987b. synonymize Acanthoxa with Spanioplon and this is here endorsed.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Iotrochotidae Dendy, 1922

Rob W.M. Van Soest

Zoological Museum, University of Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, Netherlands. (soest@science.uva.nl)

Iotrochotidae Dendy (Demospongiae, Poecilosclerida) is revived in Myxillina with the distinguishing feature being possession of birotula
microscleres. Resemblance of birotulas to polydentate chelae of some Myxillidae make it likely that birotulas are derived from anchorate
chelae by proliferation of the alae. This is supported by the occurrence of true anchorate isochelae in a few members of the family. From
these facts a close relationship between Myxillidae and Iotrochotidae may be concluded. The shape, skeletal architecture and further
spicule complement in this family is quite diverse, paralleling that of the larger families of Myxillina. Six genera are considered valid.
Keywords: Porifera; Demospongiae; Poecilosclerida; Myxillina; Iotrochotidae; Amphiastrella; Hymetrochota; Iotroata; Iotrochopsamma;
Iotrochota; Rotuloplocamia.

DEFINITION, DIAGNOSIS, SCOPE History and biology

Synonymy Dendy's section Iotrochoteae so far has been largely ignored

by later authors. Genera united by the possession of the peculiar
Iotrochoteae Dendy, 1922b: 96. birotulas were assigned to a variety offamilies (e.g., Desmacididae
(Iotrochota), Coelosphaeridae (Amphiastrella), Myxillidae
Definition (Iotrochota. Iotroata), Hymedesmiidae (Hymetrochota), and
Microcionidae (Rotuloplocamia». The formal recognition of this
Myxillina with birotula microscleres. group at the family level is made here for the first time. The family
is represented in most areas of the world in shallow as well as
Diagnosis deeper waters.

Encrusting, massive, fistular, ramose or flabellate sponges;

ectosomal skeleton variously consisting of perpendicular or tan-
gentially arranged tomotes or absent; many genera demonstrate Taxonomic remarks
a lack of differentiation of megascleres. Choanosomal skeleton
reticulate or hymedesmioid. Choanosomal megascleres styles, The previous assignments of genera cited above, here consid-
strongyles, or tylotes, smooth or acanthose. Microscleres birotulas, ered members of Iotrochotidae, were based usually on features of
occasionally also anchorate isochelae and sigmas. skeletal architecture. In other families employed here, skeletal
architecture is quite diverse, and the possession of the unique biro-
Scope tulas is considered to be a stronger and clearer indication of shared
history than broadly similar skeletal features, which are probably
Eleven nominal genera, six of which are considered valid: strongly related to shape and size of the sponge, and indirectly to
Amphiastrella, Hymetrochota. Iotroata, Iotrochopsamma. Iotrochota habitat and ecology.
and Rotuloplocamia

KEY TO GENERA

(I) No megascleres; the skeleton is a reticulation of sand grains ................................................................................... Iotrochopsamma

At least one category of megascleres ..... ............... ................. ............ ............... ................ ................ ...... ..................... ................ ....... 2
(2) Fistular growth form with parchment-like surface made up of intercrossing strongylote or tylote spicules. Choanosomal
skeleton vestigial ..... .......... ........... ................................................ ............................ ............... ........... ...... ............... ....... Amphiastrella
Choanosomal skeleton elaborate, reticulated, plocamiid or hymedesmioid ....................................................................................... 3
(3) Acanthose megascleres present ........................................................................................................................................................... 4
Acanthose megascleres absent .... .......... .... ............. ........ ........ ...... ...... .... ........... ........... ...... .......................... ........... ............................ 6
(4) Among the acanthose megascleres there are acanthose strongyles and tylotes; skeleton arrangement is plocamiid ... Rotuloplocamia
No acanthostrongyles or -tylotes; skeleton hymedesmioid or reticulate ............................................................................................. 5
(5) Thin encrustations. Skeleton hymedesmioid (i.e., with acanthostyles erect on the substrate, points outwards, with shorter acan-
thostyles in groups at the base ....................................................................................................................................... Hymetrochota
Massive or lamellar. Skeleton reticulate, no smaller echinating acanthostyles ....................................................................... Iotroata
(6) Rectangular reticulation of polyspicular tracts; differentiation in shorter/fatter and longer/thinner spicules which may be either styles
or strongyles or both, but these are not functionally localized ............................................................................................ Iotrochota
Reticulation of single spicules or paucispicular tracts; differentiation into ectosomal tomotes and choanosomal styles ....... Iotroata

594
Porifera· Demospongiae • Poecilosclerida • Myxillina • Iotrochotidae 595

Fig. 1. A-D, Amphiastrella birotulifera (Carter, 1886c as Phloeodictyum). A, holotype (scale 1 cm). B-C, SEM photos of spicules made from the holotype
(scales 10 j-Lm). D, drawing of skeleton and spicules made from a slide of the holotype. E, Hymetrochota rotula Topsent (l904b), drawing of birotulas
reproduced from Topsent's plate XIV fig. 6 (size see text). F, Hymetrochota topsenti (Burton, 1930 as Hymenotrocha), drawing of spicules reproduced from
Burton's fig. 4 (encadred chela assumed to be foreign, sizes not recorded).

AMPHIASTRELLA DENDY, 1896 Diagnosis

Synonymy Massive, erect, fistulose growth form; choanosomal skeleton

consists of diactinal megascleres which are strongyles or tylotes
Amphiastrella Dendy, 1896: 27. forming an irregular mesh; ectosomal megascleres same as in
choanosome, arranged tangential to surface forming a compact
dermal layer; microscleres are birotulas and in one species sigmas.
Type species Two species, apparently restricted to South Australia and New
Zealand.
Phloeodictyon birotuliferum Carter, 1886c: 447 (by original
designation). Previous review

Bergquist & Fromont (1988).

Definition
Description of type species
Iotrochotidae with fistulose hollow shape and tylotes as
megascleres. Amphiastrella birotulifera (Carter, 1886c) (Fig. lA-D).
596 Porifera· Demospongiae • Poecilosclerida • MyxilIina • Iotrochotidae

Synonymy. Phloeodictyon birotuliferum Carter, 1886c: 447, Definition

pI. X figs 1-5; Amphiastrella birotulifera; Dendy, 1896: 27.
Material examined. Syntypes: BMNH 1887.7.11.12 - Iotrochotidae with hymedesmioid arrangement of mega-
Westernport Bay, Victoria, Bracebridge Wilson Collection, and 2 scleres, including echinating acanthostyles erect on the substrate.
slides, BMNH 1887.7 .11.12a, and unregistered.
Description. Hollow mass (Fig. lA) with branching fistules
on top and rooting processes at the base (the latter observation Diagnosis
is from Dendy, 1896, from an additional specimen). Length of
fistular tubes ca. 9 cm, diameter 0.5 cm. Some of the branches of Encrusting sponges with hispid surface. Skeleton hymedes-
the fistules are open-ended and presumably function as oscules. mioid. Ectosomal tomotes smooth. Choanosomal spicules are
Surface smooth. Consistency parchment-like, firm. Colour brown- spined styles; auxiliary acanthostyles echinate the substrate.
ish in alcohol. Skeleton (Fig. ID) of the ectosome a tangential Microscleres are birotulas. Two species, apparently restricted to
reticulation of spongin enforced spicule tracts, 3-5 spicules in North Atlantic deep water.
cross section, making rather neat triangular meshes. Choanosomal
skeleton virtually absent: a few thin spicule lines traverse the wide
internal space, branching infrequently and irregularly. Tylotes are Previous review
the only megascleres. Spicules (Fig. lB-D), tylotes, straight,
robust, with clearly developed, elongate tyles, smooth, 400-700 X Topsent (1904b).
7-20/-Lm (Carter mentions a second thinner category with less
developed tyles, but it seems they are part of a continuous series);
birotulas with relatively thick shaft often constricted in the middle, Description of type species
which may bear an occasional spine, variable in length, probably
divisible in two size categories, 17-35 and 51-56/-Lm; there are a Hymetrochota rotula Topsent, 1904b (Fig. IE).
few sigmas of about 40/-Lm in the slide (also recorded by Dendy), Synonymy. Hymetrochota rotula Topsent, 1904b: 168,
but these are probably foreign. Distribution. Victoria. pI. XIV fig. 6.
Remarks. A second species of this genus is A. kirkpatricki Material examined. Holotype (not seen): MOM. Fragment
Dendy (1924: 371) from New Zealand, differing primarily in the size of holotype: MNHN D.T. 999 (slide) - labeled 'PA 1897, 152',
of the birotulas which are in a single small size category of 17-34 Azores, Princesse Alice Cruise 1897, 200 m depth.
/-Lm (see Bergquist & Fromont (1988: 50) for a subsequent record of Description. A thin hispid crust epibiotic on a fistule of
that species). These authors considered Xytopsene de Laubenfels Oceanapia. Brownish in alcohol. Skeleton. Hymedesmioid, i.e.,
(1936a) a junior synonym of Amphiastrella, but this was based on large acanthostyles stand erect on the substrate surrounded by
the statement of de Laubenfels that Cornulum novaezealandiae small acanthostyles. Tomotes scattered singly or occasionally in
Brpndsted (1924b: 473, fig. 26) should be assigned to Xytopsene. bundles near the surface. Microscleres abundant in all parts of the
Brpndsted's species is a junior synonym of Amphiastrella kirk- sponge. Spicules, tomotes smooth, style-like, with unequally
patricki Dendy (1924). However, the type species of Xytopsene is not swollen mucronate ends, frequently polytylote, 350-380 X 3/-Lm;
Brpndsted's species, but Suberitesjistulatus Carter (1880b: 53, pI. V choanosomal acanthostyles, slightly curved, heavily spined with
fig. 22), a clear Damiria, making Xystopsene a junior synonym of truncate spines on the head, less densely and more finely spined
that genus and not of the present one. along the shaft, 800-1000 X 10-15 fLm; echinating acanthostyles
Levi & Levi (1983b: 966) assigned Amphiastrella to their fam- straight, entirely spined, 80-180 fLm. Microscleres birotulas
ily Comulidae (subsequently named Iophonidae for priority reasons, (Fig. IE) with 20, relatively short, teeth, 20-23 /-Lm, umbrella 7 /-Lm
cf. Van Soest et al., 1994, but now calledAcarnidae, again for priority (growth stages with reduced teeth and prominent falces).
reasons), while Bergquist & Fromont (1988) maintained it in the fam- Distribution and ecology. Azores, collected at 200m.
ily Coelosphaeridae. Neither assignment is here followed because of Remarks. The distinction of this genus from the closely
the possession of the peculiar birotulas shared with the other mem- related Iotroata rests on the combination of a hymedesmioid skele-
bers of this family. The fistular shape and skeletal organization, ton (as opposed to a reticulate one in Iotroata), and echinating
although common in the families Coelosphaeridae and Acamidae, acanthostyles (absent in Iotroata). Both characters are not consid-
is found in many unrelated sponges. It is considered a functional ered of phylogenetic significance if occurring singly, but in combi-
adaptation to life in sedimented environments. nation they are provisionally considered to unite a monophyletic
group. A second species, H. topsenti (Burton, 1930c: 506, text-
fig. 4) from Stavanger (Norway) is here assigned to Hymetrochota.
HYMETROCHOTA TOPSENT, 1904 Burton described it under the name Hymenotrocha, without indi-
cating that he intended it to be a new genus and in his comments
Synonymy indicated there were 'other species of the genus'. Our assignment
of Burton's material to Hymetrochota is conditional on the correct-
Hymetrochota Topsent, 1904b: 168. Hymenotrocha Burton, ness of our assumption that his description includes a mixture of
1930c: 506. spicules of a Hymetrochota and a Hymedesmia, tylotes, strongyles,
smooth styles, two categories of acanthostyles, arcuate chelae and
Type species two sizes of birotulas. His figure is here reproduced (Fig. IF). No
material of this species was found in the BMNH London (although
Hymetrochota rotula Topsent, 1904b: 168 (by original Burton mentions the reg. no. 1910.1.1.2626), so the assignment to
designation). Hymetrochota remains tentative.
Porifera· Demospongiae • Poecilosclerida • Myxillina • Iotrochotidae 597

IOTROATA DE LAUBENFELS, 1936 De Laubenfels (1936a), after having erected Iotroata on p. 87

for thinly encrusting species combining the presence of birotulas
Synonymy and anchorate chelae, some pages further on (p. 95) proposed a fur-
ther new genus Iotaota for species with a reticulate skeleton,
Iotroata de Laubenfels, 1936a: 87. Iotaota de Laubenfels, tylotes as ectosomal megascleres and birotulas and unguiferate
1936a: 95. Sigmarotula Bergquist & Fromont, 1988: 94. chelae as microsclere. The type was indicated (by original designa-
tion) to be Halichondria abyssi Carter (1874a); in this genus he
Type species included several species already assigned to Iotroata, including its
type I. acanthostylifera, apparently having forgotten his actions
Iotrochota acanthostylifera Stephens, 1916: 236 (by original on p. 87. It is clear that the two 'genera' are synonyms.
designation). The name Iotroata was chosen in preference over Iotaota
because of page prority. The holotype of Halichondria abyssi
Definition Carter (1874a: 245), BMNH 1890.4.10.4 with slide BMNH
1910.1.1.1393, was examined (Fig. 2B-G). The skeleton consists
Iotrochotidae with an isotropic skeleton; microscleres may of smooth oxeas (Fig. 2B, E) and styles (Fig. 2D) for structural
include anchorate isochelae. megascleres, but is otherwise similar to acanthostylifera in having
ectosomal tylotes (Fig. 2B) of ca. 250 fLm, poly dentate chelae of
Diagnosis ca. 65 fLm (Fig. 2F) and birotulas (Fig. 2G) of ca. 17 fLm. Carter
reported toxas, but these are here assumed to be contamination
Encrusting, massive or flabellate; ectosomal skeleton com- because the type slide examined here did not contain them.
posed of smooth ectosomal tylotes with mucronate or spined ends, Likewise, Lundbeck (1905: 195) re-examined the type and did not
mostly standing erect, sometimes paratangentially, in bundles on find them. The genus Iotroata has been largely ignored so far, with
the surface; choanosomal skeleton composed of smooth or slightly the exception of a record by Van Soest (1987b: 18) of Iotroata spin-
acanthose styles, sometimes oxeote or absent, forming an isotropic osa (Lundbeck, 1905: 195), ZMA POR. 4708. This species differs
reticulation; fibres are poor but there may be abundant collagen; from I. acanthostylifera in being thickly flabelliform and in the
microscleres are birotulas, anchorate (unguiferous) isochelae and absence of true chelae, instead of which it has two size categories
sigmas. About 10-11 species, occurring in northern and southern (33 and 18 fLm) of birotulas (Fig. 2H). The choanosomal styles
oceans, but so far not recorded in the tropics. are entirely spined and measure 450-500 X 12-15 fLm; they
are arranged in a loose reticulation. Further species of Iotroata
Previous review appear to be I. affinis (Lundbeck, 1905: 194), I. oxeata (Lundbeck,
1905: 186), I. polydentata (Lundbeck, 1905: 192), I. rotulancora
Van Soest (l987b). (Lundbeck, 1905: 191, with junior synonym I. dubia Lundbeck,
1905: 189), I. varidens (Lundbeck, 1905:185), and I. somovi
Description of type species (Koltun, 1964b: 52), all described originally as Iotrochota.
The genus Sigmarotula Bergquist & Fromont, 1988 was
Iotroata acanthostylifera (Stephens, 1916) (Fig. 2A). erected (by monotypy) for type species S. lamellata Bergquist &
Synonymy. Iotrochota acanthostylifera Stephens, 1916: Fromont (1988: 94, pI. 44 figs D-F, pI. 45 figs A-D). This was
236; Stephens, 1920: 29, pI. II fig. 5; Iotroata acanthostylifera; de excellently described by these authors; it is an orange-red, lamellate
Laubenfels, 1936a: 87. sponge, with ectosomal tylotes of 210-225 X 4-5 fLm, choanosomal
Material examined. Holotype (slide): BMNH 1953.11.1.89- smooth styles of 360-425 X 7.5-20 fLm, birotulas of 12-16.5 fLm
from W coast of Ireland. and sigmas of 36-50 fLm. Bergquist & Fromont state that their new
Description (from Stephens, 1916). Massive encrustation genus is quite distinct from other birotula-possessing genera, but it
on deep-water corals, up to 2 cm thick. No further data. Skeleton. is clearly closely related and synonymous with Iotroata because the
Choanosomal skeleton is an irregular reticulation of acanthostyles. only difference with several Iotroata species, such as I. polydentata,
Ectosomal skeleton consists of tangential bundles of tomotes, is the possession of sigmas. In other Myxillina genera
which may form longer fibres. Spicules (Fig. 2A), ectosomal (Amphiastrella, Myxilla, Phorbas) the presence or absence of sig-
tylotes, ends often unequal, one being more elongate, 300-330 X mas is not considered to unite monophyletic groups (the presence of
5 fLm; choanosomal acanthostyles, straight or slightly curved, with sigmas is considered an ancestral Myxillina character).
the head very slightly swollen and thickly spined; a few small
spines are scattered along the shaft, 320-350 X 8 fLm. Microscleres
are polydentate unguiferate chelae with 8-10 teeth at either end, IOTROCHOTA RIDLEY 1884
45-50 fLm and birotulas with 15 teeth, about 15 fLm. Distribution
and ecology. W coast of Ireland, encrusting hard substrates at Synonymy
450-1300m.
Remarks. The genus is peculiar because the type species Iotrochota Ridley, l884a. Hiattrochota de Laubenfels, 1950b:
and several other forms (e.g., Iotroata somovi (Koltun, 1964b as 20. Iotrochostyla de Laubenfels, 1954: 150.
Iotrochota) combine the possession of birotulas with unguiferate-
anchorate chelae, thus providing evidence of the likely origin of Type species
birotulas from anchorate chelae. The genus is closely related to
Hymetrochota with which it shares presence and shape of tomotes Halichondria birotulata Higgin, 1877: 296 (by subsequent
and spined styles. It differs in having an isotropic reticulation. designation; de Laubenfels, 1936a: 48).
598 Porifera· Demospongiae • Poecilosclerida • Myxillina • Iotrochotidae

'00
1

Fig. 2. Iotroata spp. A, Iotroata acanthostylifera (Stephens, 1916 as Iotrochota) , drawing of skeleton and spicules made from a slide of the holotype.
B-G, Iotroata abyssi (Carter, 1874a as Iotrochota), type of Iotaota. B, drawing of skeleton and spicules made from a slide of the holotype. C-G, SEM
photos of spicules made from the holotype (scales: C-E, 100 ILm; F, 10 ILm; G, llLm). H, Iotroata spinosa (Lundbeck, 1905 as Iotrochota), SEM photos
of two size categories of birotulas (scale llLm) made from a ZMA specimen.

Definition choanosomal styles, or oxeas, or only strongyles inside fibres (all

about the same size, without definite localization of strongyles
Iotrochotidae with smooth megascleres normally differenti- in the ectosomal skeleton); microscleres are birotulas. About
ated in two categories, longer and thinner monactinal (styles) or 10--15 species, predominantly recorded from tropical waters, but
diactinal spicules (usually strongyles) and shorter and thicker at least one species is from the Arctic.
styles or diactinal spicules, which do not have distinct localization
in ectosomal and choanosomal regions. Previous reviews

Ridley (1884a: 433), Wiedenmayer (1977b: 138), Van Soest

Diagnosis (1984b: 38).

Encrusting, massive, digitate or bushy growth forms, often Description of type species
black colouration with a purple stained mucus; ectosomal skeleton
thick, difficult to detach, composed of single spicules or poorly Iotrochota birotulata (Riggin, 1877) (Fig. 3A-D).
defined short tracts of spicules; choanosomal skeleton a regular Synonymy. Hyrtios musciformis Duchassaing & Michelotti,
reticulation of multispicular tracts; spongin maybe well-developed 1864: 75, pI. 14 fig. 3; Van Soest, 1984b: 38 (senior synonym
and is usually darkly pigmented; megascleres are smooth proposed for suppression by Wiedenmayer, 1977b: table 48).
Porifera· Demospongiae • Poecilosclerida • Myxillina • Iotrochotidae 599

I 1

Fig.3. A-D, Iotrochota birotulata (Higgin, 1877 as Halichondria). A, drawing of habit and spicules copied from Higgin, 1877 (sizes see text). B-D, SEM
photos of spicules made from the lectotype (scales: B-C, 10 !Lm; D, 1 !Lm). E, [otrochopsamma arbuscula (Whitelegge, 1906 as Iotrochota), habit, copied
from Whitelegge (1906) (size see text). F-K, Rotuloplocamia octoradiata Levi (1952). F-J, SEM photos of the spicules made from a ZMA specimen
(scale 10 !Lm; except E, l!Lm). K, drawing of skeleton and spicules made from the holotype.

Halichondria birotulata Riggin, 1877: 296, pI. 14 figs 11-15. squeezed, it gives off a dark exudate. Many specimens are infested
Iotrochota birotulata; Ridley, 1884a: 433; Van Soest, 1984b: 38, by yellow or orange zoanthids. Ectosomal skeleton an organic crust
pI. V, figs 1-2, text-fig. 11. Hircinia atra Whitfield, 1901: 49, pI. 5. with scattered loose megascleres. Choanosomal skeleton a rectan-
? Iotrochota imminuta Pulitzer-Finali, 1986: 141, figs 62-63. gular reticulation of thick spicule tracts, diameter 35-100/-Lm,
Material examined. Lectotype: BMNH 1877.3.9.1 - dry mesh size 100-700 /-Lm. Strongy les and styles are mixed in the
specimen from Jamaica. Paralectotype: BMNH 1877.3.9.2 - 2 tracts. Many loose scattered megascleres. Spicules (Fig. 3A-D)
slides from BMNR collection. Other material. Many specimens two types of megascleres are present (Fig. 3B-C), occurring both
from the ZMA collection (see list in Van Soest, 1984b). in a continuous range from relatively long and thin to relatively
Description. Encrusting, massive-lobate, with repent short and fat; strongyles, 140-230 X 3-5 /-Lm, and styles,
branches, or large branching bushes (Fig. 3A). Surface conu10se 140-240 X 3-6 /-Lm; birotulas (Fig. 3A,D), 10-15 /-Lm. Distribution
(sharp conules in dry specimens), but may be smooth for extended and ecology. Caribbean, reefs down to 90 m, lagoons, mangroves.
areas. Length of branches up to 50 cm or more, diameter 1-5 cm; Remarks. Localization of styles and strongyles reported by
encrustations may be as thin as 1 mm. Consistency tough and Bergquist (1965) cannot be confirmed. Strongyles (presumably
resilient, barely compressible. Colour black, typically mottled with derived from tomotes) and styles (presumably the choanosomal
light green patches; occasionally entirely light yellow-green. When megascleres) are intermingled throughout the sponge. Hyrtios
600 Porifera' Demospongiae • Poecilosclerida • Myxillina • Iotrochotidae

muscifonnis Duchassaing & Michelotti (1864), type specimen network of heavy spongin fibres, with long ascending primary
ZMA POR. 1427, redescribed by Van Soest (1984b: 38) is a clear fibres and shorter transverse secondary fibres both cored with sand
senior synonym of Halichondria birotulata, but the proposal by grains and foreign spicule fragments; megascleres are absent;
Wiedenmayer (1977b: table 48, footnote f) to suppress this name microscleres are birotulas. A single species from South Australia.
under ICZN Article 79c in favour of stability is here endorsed.
Iotrochota imminuta Pulitzer-Finali (1986) was erected for speci- Previous review
mens resembling I. birotulata in most aspects, but are lacking
birotulas and styles. Wiedenmayer (1977b) also discussed such De Laubenfels (1954).
specimens and concluded that they are merely destitute representa-
tives of I. birotulata. That is also endorsed here. However, Pulitzer- Description of type species
Finali's material could also belong to chondropsid genera as
Chondropsis or Strongylacidon. Iotrochopsamma arbuscula (Whitelegge, 1906) (Fig. 3E).
Lundbeck's North Atlantic deep water species described as Synonymy. Iotrochota arbuscula Whitelegge, 1906: 482,
Iotrochota are all referred to Iotroata, due to the fact they have pI. XLIV fig. 18. Iotrochopsamma arbuscula; de Laubenfels,
ectosomal tomotes morphologically distinct from megascleres of 1954: 127.
the choanosome. Material examined. None. Holotype: AM - Thetis'
The genus Hiattrochota de Laubenfels, 1950b was erected for Collection, stat. 41, (fide Hooper & Wiedenmayer, 1994).
H. protea de Laubenfels (1950b: 20, fig. 12) (original designation). Description (from Whitelegge, 1906). Dichotomously
The type specimen USNM 22750 was re-examined. It has both ramose sponge with a stem of 7.5 cm and dividing branches of up
styles and strongyles, as well as birotulas and it is entirely unclear to 24cm long. Stem and branches are 0.7-3cm in diameter. Colour
why this was considered distinct from Iotrochota. Also the wide- grey. Skeleton. Rectangular reticulation of spongin fibres cored by
spread Iotrochota baculifera Ridley, 1884a (type specimen BMNH sand grains, no proper megascleres. Fibres 80-150 fLm in diameter,
1882.2.23.315 re-examined) was assigned to this genus. meshes 250-500 fLm. Spicules exclusively birotulas, 20 fLm.
Subsequently H. ditrochota de Laubenfels (1954: 122), H. hiatti de Distribution and ecology. New South Wales coast, South Australia,
Laubenfels (1954: 125), and H. mystile de Laubenfels (1954: 126) trawled from some depth.
were added. The genus allegedly differs from Iotrochota proper in Remarks. The complete absence of megascleres is here con-
having no size distinction between monactinal and diactinal sidered as sufficient evidence for a separate status of this monotyp-
spicules. This is considered of minor importance. ical genus. If similar specimens or species with sand columns, but
Likewise, Iotrochostyla de Laubenfels, 1954 was erected for including proper megascleres, would be found the genus may be
type species (by original designation) I. iota de Laubenfels (1954: merged with Iotrochota, with which it shares the rectangular retic-
150, fig. 97), and the Alaskan species Iotrochota magna Lambe ulation. A striking similarity is apparent between myxillid
(1894: 120, pI. ill fig. 2), based on the lack of diactinal megascle- Psammochela chalinifonnis Carter (1885) and iotrochotid
res. This is also considered a minor feature. Both I. iota (type spec- Iotrochopsamma in the absence of megascleres and the small
imen USNM 22909) and l. magna indeed appear to have a majority polydentate microscleres.
of styles as megascleres (125 X 4 fLm in the type, 438 X 24 and
240-333 X 6-9 fLm in I. magna), but these are clearly differenti-
ated in a longer-thinner and shorter-thicker category. Moreover, the ROTULOPLOCAMIA LEVI, 1952
two species differ considerably in appearance and spicule sizes,
and appear not at all closely related. If more than one species of Synonymy
Iotrochota would be extant with exclusively styles, Iotrochostyla
may be conveniently recognized as a subgeneric unit. Rotuloplocamia Levi, 1952: 51.

Type species
IOTROCHOPSAMMA DE LAUBENFELS, 1954
Rotuloplocamia octoradiata Levi, 1952: 51 (by monotypy).
Synonymy
Definition
Iotrochopsamma de Laubenfels, 1954: 127.
Iotrochotidae with plocamiid skeletal structure, including
Type species basal acanthostrongyles and echinating acanthostyles.

Iotrochota arbuscula Whitelegge, 1906: 482 (by original Diagnosis

designation).
Thin encrustations with basal reticulation of acanthostrongyles
Definition on the nodes of which long styles stand erect surrounded by short
acanthostyles. Tomotes are anisostrongyles or subtylotes arranged
Iotrochotidae lacking megascleres. in erect bundles. Microscleres birotulas. A single species from the
Eastern Atlantic.
Diagnosis
Previous reviews
Ramose; ectosomal skeleton finely reticulate, porous, arena-
ceous; choanosomal skeleton composed of an open reticulate Desqueyroux-Faundez & Van Soest (1996).
Porifera· Demospongiae • Poecilosclerida • Myxillina • Iotrochotidae 601

Description of type species dumbell-type acanthostrongyles (Fig. 31, K) with swollen and
rugose heads and only a few spines on the shaft, 40-120 X
Rotuloplocamia octoradiata Levi, 1952 (Fig. 3F-K). 8-12lJ-m. Microscleres birotulas (Fig. 3J, K) with 8 teeth on each
Synonymy. Rotuloplocamia octoradiata Levi, 1952: 51, umbrella, probably in two size categories, 5-12 f.Lm. Distribution
figs. 15-16; Van Soest, 1993a: 213, table 5. Plocamiancora spec. and ecology. West Africa, Atlantic coast of Spain, cryptic habitats
Solorzano et al., 1991: 182, fig. 4. in shallow water down to 30 m depth.
Material examined. Holotype: MNHN DCl. 1271 - Remarks. There are a few discrepancies between Levi's
Senegal. Other material. ZMA 6688 - Mauritania Expedition, stn. (1952) and the ZMA material on the one hand, and the North Spain
49/5. ZMA 6706 - Mauritania Expedition, stn. 49/18. material described as Plocamiancora spec. by Solorzano et al.
Description. Extremely thin, encrusting the undersurface of (1991) in the shape of the tomotes and the size of the large styles.
stones or insides of dead shells, not easily detached; several square However, the tomotes are variable in shape in the West African
cm in lateral expansion. Surface optically smooth, slightly hispid. specimens, so these differences are considered infraspecific varia-
Consistency firm. Colour yellow-orange or reddish orange. Skeleton tion. Rotuloplocamia octoradiata has the plocamiid skeletal struc-
(Fig. 3K). acanthostrongyles form a basal, somewhat obscured ture considered in the past (Topsent, 1928c; Burton, 1935a) a
isotropic reticulation. Upon it large acanthostyles are standing singly synapomorphy for a separate and polyphyletic family Plocamiidae.
and erect, surrounded by groups of shorter acanthostyles. The However, the diverse spiculation of the various forms has caused
ectosomal tomotes occur in dense bundles perpendicular and their current assignment to several unrelated Myxillina families, as
paratangential near the surface forming a more-or-Iess dense crust. well as to some Microcionina families. The genus is considered to
Spicules (Fig. 3F-K). Megascleres ectosomal tomotes in the form be valid despite its monotypical status because it differs from other
of styles or subtylotes (subtylostyles with mucronate pointed end, birotule-bearing genera in the possession of dumbell-type acan-
Fig. 3F, K). smooth, 130-220 X 4-5 f.Lm; large styles (Fig. 3G, K) thostrongyles forming the isotropic basal skeleton. This type of
with strongly rugose heads and smooth curved shafts, 360-740 X skeleton is considered ancestral (cf. above) to Myxillina, and also
13-20 f.Lm; short, echinating acanthostyles (Fig. 3H, K), with heav- represented in Microcionina families Acarnidae, Microcionidae
ily rugose heads and lightly spined shafts, 120-275 X 9-16f.Lm; and Raspailiidae.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademicIPlenum Publishers, New York, 2002

Family Myxillidae Dendy, 1922

Rob W.M. Van Soest

Zoological Museum, University of Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, Netherlands. (soest@science.uva.nl)

Myxillidae Dendy (Poecilosclerida: Myxillina) is revised and narrowed down to a new concept including only those genera which
combine the possession of anchorate chelae (or polydentate derivations) with diactinal ectosomal tomotes and choanosomal styles in a
reticulate arrangement. Two genera deviating from this general concept are included here: i.e., Hymenancora with hymedesmioid skele-
ton, and Psammochela with only a single megasclere type, because of chelae similarities with other included genera. Other families with
anchorate chelae are Desmacididae (oxeas as the only megascleres), Iotrochotidae (with birotulas) and Crambeidae (monactinal ectoso-
mal tomotes and choanosomal styles). Thirty-three nominal genera answer to the new concept of Myxillidae and of these eight are con-
sidered valid: Damiriopsis, Ectyonopsis, Hymenancora, Melonanchora, Myxilla, Plocamiancora, Psammochela, Stelodoryx. In the large
genus Myxilla four subgenera have been defined to alleviate classing the large numbers of species: Myxilla (Myxilla), Myxilla
(Burtonanchora), Myxilla (Ectyomyxilla), and Myxilla (Styloptilon).
Keywords: Porifera; Demospongiae; Poecilosc1erida; Myxillina; Damiriopsis; Ectyonopsis; Hymenancora; Melonanchora; Myxilla
(Myxilla); Myxilla (Burtonanchora); Myxilla (Ectyomyxilla); Myxilla (Styloptilon); Plocamiancora; Psammochela; Stelodoryx.
- - - - _ ...- .. - - -- --"- -- - - - - -

DEFINITION, DIAGNOSIS, SCOPE History and biology

Synonymy Ridley & Dendy (1887) included sponges now considered

members of Myxillidae into a larger subfamily Ectyoninae which
Myxilleae Dendy, 1922b: 85; Myxillidae Topsent, 1928c: 237. was based on the possession of echinating acanthostyles (thus
covering the present concepts of Myxillina + Microcionina).
Hentschel, 1923 defined the family to include sponges with chelae,
Definition diactinal ectosomal megascleres and monactinal (often spined)
choanosomal megascleres. He included hymedesmiids in the
Myxillina with anchorate or unguiferate chelae, choanosomal family, but kept Tedaniidae, Dendoricellidae, Coelosphaeridae and
styles and diactinal ectosomal tomotes (oxeote and tylote). Crellidae as separate families. Topsent (1928c) employed a broad
concept of Myxillidae including subfamilies Myxiliina, Tedaniinae
and Phoriospongiinae, based on the possession of reticulate skele-
Diagnosis ton, diactinal ectosomal megascleres and monactinal choanosomal
megascleres. A major difference of Topsent's subfamily Myxillinae
Encrusting, massive, lobate, fan-shaped or branching sponges. with the presently employed contents of the family Myxillidae
Specialised ectosomal skeleton composed of tomotes with smooth is the transfer of genera with arcuate chelae and a reticulate
or apically spined bases, arranged as bouquets or lying paratangen- choanosomal skeleton to the family Coelosphaeridae, whereas
tial or perpendicular to the surface. This is occasionally absent. Topsent restricted Coelosphaeridae to genera with hollow-bladder
Choanosomal skeleton composed of isotropic, anisotropic or shape. De Laubenfels (1936a) also employed a large concept of
plumoreticulate tracts of smooth or partially spined monactinal or Myxillidae, but included crellids and hymedesmiids in it. He
diactinal choanosomal megascleres, sometimes echinated by small subdivided this large group into two subfamilies, Myxillinae with
acanthose styles. Hymedesmioid arrangement occurs in thinly smooth ectosomal megascleres, and Grayellinae with spined
encrusting forms. Spongin development variable, usually consisting ectosomal megascleres. Within Myxillinae, he distinguished
of light spongin cementing spicules together at their nodes, but 'Myxillinae I' which more or less covered Topsent's Myxillinae,
sometimes with heavy fibres. Microscleres include one or more size and 'Myxillinae II' which covered Topsent's Hymedesmiidae. Levi
and form categories of anchorate isochelae and/or derivatives (spat- (1973) followed Topsent's (1928c) classification, as did most other
ulate, unguiferous isochelae, sometimes anisochelate, occasionally authors prior to Hajdu et al. (1994a). These authors restricted
polydentate), and one or two size categories of smooth sigmas. Myxillidae to genera possessing as chelae exclusively anchorate
forms or derivates thereof, in combination with diactinal tomotes.
This was followed by Desqueyroux-Faundez & Van Soest (1996).
Scope Here, Myxillidae is further restricted by exclusion from the family
of genera Desmacidon and Desmapsamma, which go to a separate
Thirty three nominal genera are included in this family, of family Desmacididae based on the loss of choanosomal styles, and
which eight are recognised here as valid: Damiriopsis, Ectyo- of genera possessing birotula microscleres, which go to a separate
nopsis, Hymenancora, Melonanchora, Myxilla, Plocamiancora, family Iotrochotidae. Myxillidae tend to be encrusting to massive
Psammochela, Stelodoryx. One genus (Myxilla) has four recog- sponges occurring on many different substrates, including rocks,
nized subgenera. mangrove roots, and sand. They may occur at considerable depths.

602
Porifera' Demospongiae • Poecilosclerida • Myxillina • Myxillidae 603

The majority of the species and genera have been reported from chapter on suborder Myxillina) are common in this family,
temperate and cold waters. but in thinly encrusting or bladder-like sponges this may be
obscured or absent, or replaced by an emphasis of zone II skele-
Taxonomic remarks tons. The anchorate chelae may be tridentate, unguiferate or
polydentate.
The concept of M yxillidae is here narrowed down further than
any previous concept and clearly focused on the possession of Previous reviews
anchorate or anchorate-derived chelae in combination with differ-
entiated ectosomal and choanosomal megascleres. Reti- Topsent (1928c), Hajdu et ai. (1994a), Desqueyroux-Faundez &
culate skeletons (presumably by emphasis of skeletal zone I; see Van Soest (1996).

KEY TO GENERA

(1) Choanosomal skeleton includes smooth or spined strongyles ............................................................................................................ 2

Choanosomal skeleton consists of styles only ..................................................................................................................................... 4
(2) Strongyles large and thick; there are no zone II (cf. Fig. of Myxillina chapter) long styles ............................................................... 3
Skeleton is plocamiid, i.e. there are long styles erect on a basal reticulation of strongyles ......................................... Plocamiancora
(3) Strongyles spined all over .................................................................................................................................................. Ectyonopsis
Strongyles smooth except the heads which may have light spination ............................................................................... Damiriopsis
(4) Microscleres include sphaerancorae ............................................................................................................................. Melonanchora
No sphaerancorae ................................................................................................................................................................................. 5
(5) Chelae include polydentate chelae ...................................................................................................................................................... 6
Chelae are spatulate or unguiferate with not more than 3 teeth ................................................................................................ Myxilla
(6) Skeleton hymedesmioid ................................................................................................................................................. Hymenancora
Skeleton reticulate ................................................................................................................................................................ ............... 7
(7) Tomotes present next to the choanosomal styles ................................................................................................................. Stelodoryx
No tomotes, a single megasclere type or no megascleres at all ...................................................................................... Psammochela

DAMIRIOPSIS BURTON, 1928 Material examined. Holotype: BMNH 1926.10.1.20 (slide

BMNH 1926.10.1.20a) - both labeled with unpublished species
Synonymy name 'dendyi'.
Description. Small, irregularly massive, size 2 X I X 0.5 cm.
Damiriopsis Burton, 1928a: 124. Surface minutely hispid. Conspicuous subdermal cavities visible
through a transparent surface membrane (Fig. IA). Oscules 1.5mm
Type species in diameter. Colour greyish-yellow (alcohol). Skeleton (Fig. 4A).
Ectosomal skeleton consists of loose bouquets of tomotes, not form-
Damiriopsis brondstedi Burton, 1928a: 124 (by monotypy). ing a continuous skeleton. Choanosomal skeleton a rectangular,
somewhat irregular reticulation of bundles of 2-4 strongyles thick.
Dermition Spicules. Ectosomal tomotes are tylotes with spined apices (Figs lB,
4A), 264--357 X 6-9 f.Lm. Choanosomal spicules are thick strongyles
Myxillidae with choanosomal strongyles in an isotropic with faintly rugose apices (Figs 1C, 2A), 625-1053 X 36-70 f.Lm.
reticulation; no sigmas. Microscleres. Anchorate chelae in two distinct size categories
(Figs 10, 2A), 33-44 and 54--97 f.Lm. Distribution and ecology. Known
Diagnosis only from the Andaman Islands, from considerable depth, 234--522m.
Remarks. Burton recorded the chelae as being anchorate as
Thickly encrusting. Surface rough. Ectosomal skeleton con- well as palmate. In light microscopy, the smaller chelae may super-
sisting of tylotes with spined apices. Choanosomal skeleton a ficially look like palmate, but under SEM they are clearly anchor-
close-meshed reticulation of thick strongyles rugose apices. Micro- ate, similar to the larger category. This single species is related to
scleres anchorate chelae. No sigmas, no echinating acanthostyles. the genera Ectyonopsis and Stelotrochota - here united under
One species. Ectyonopsis - because they share a choanosomal reticulation of
thick strongyles. This is nevertheless retained as a separate genus
Previous review because the choanosomal spicules are entirely spined.

Burton (1928a).
ECTYONOPSIS CARTER, 1883
Description of type species
Synonymy
Damiriopsis brondstedi Burton, 1928a (Figs 1, 4A).
Synonymy. Damiriopsis brondstedi Burton, 1928a: 124, Ectyonopsis Carter, 1883b: 315. Ectyonancora Levi, 1963:
pI. I figs 3-4, text-fig. 6. 36. Stelotrochota Bakus, 1966b: 501.
604 Porifera' Demospongiae • PoeciloscIerida • Myxillina • MyxiUidae

Fig. 1. Damiriopsis brondstedi Burton, 1928a. A, fragments of holotype (scale I cm). B-D, SEM photos of spicules of holotype (scale 10 fLln).

Type species (Fig. 2A) consist of three thin-bladed fragments of which the
largest is 2.5 X 1.5 X 0.4 mm. Consistency firm, colour brown in
Ectyonopsis ramosa Carter, 1883b: 315 (by monotypy). dry state. Surface finely grained, even. The ectosomal skeleton is
represented by scattered tomotes. The skeleton (Fig. 2C-E) is an
Definition isodictyal-anisotropic reticulation of acanthostrongyles encased in
spongin, echinated by similar but slightly longer acanthostyles
Myxillidae with smooth tomotes with hastate/mucronate (arrows in Fig. 2D). Spicules, acanthostrongyles (Fig. 2B, G) and
apices and an isotropic reticulation of thick acanthostrongyles, acanthostyles (Fig. 2B, F), 177-270 X 20--24/-Lm, ectosomal
echinated by thick acanthostyles; microscleres are robust spatulif- tomotes (Fig. 2H), 165-188 X 4-6 /-Lm, smooth with swollen,
erous anchorate isochelae. mucronate ends, more or less symmetrical. Anchorate isochelae
(Fig. 2I) rather elongate-narrow, in a single size category, 21-30 /-Lm.
Diagnosis Distribution. South Australia, no further data.
Remarks. The genus is here revived. It was never reported
Flabellate; ectosomal tomotes are entirely smooth with after its original description, which was incomplete, as Carter did
mucronate apices; choanosomal strongyles heavily spined all-over; not mention the anchorate isochelae.
echinating acanthostyles thick, heavily spined all-over; chelae The genus Ectyonancora Levi, 1963 was erected for type
spatulate anchorate. Four species. species Ectyonancorajlabellata Levi, 1963: 36, pI. VIB fig. 40 (by
subsequent designation; Desqueyroux-Fatindez & Van Soest, 1996:
Previous reviews 47). The syntypes (MNHN LBIM DCL. 548 & 549 from South
Africa, stat. FAL 330), were re-examined. These are erect, flabel-
Levi (1963), Desqueyroux-Fatindez & Van Soest (1996), both late, lamellar sponges (Fig. 3A), 7.5X 10X0.4 cm. Numerous small
as Ectyonancora. oscules of 0.5 mm diameter cover both sides. Brownish or ochre in
alcohol. Skeleton (Fig. 4B). Loose tomotes and chelae are abun-
Description of the type species dant in the detachable surface membrane. The choanosomal skele-
ton consists of thick spicule tracts enclosing isotropic meshes.
Ectyonopsis ramosa (Carter, 1883b) (Fig. 2A-I). Spicules. Ectosomal tomotes (Figs 3B, 4B), smooth with lanceo-
Synonymy. Ectyonopsis ramosa Carter, 1883b: 315, late apices, 145-180 X 7/-Lm. Choanosomal acanthostrongyles
pI. XIII fig. 5. (Figs 3C, 4B), thick, heavily spined, 220--250 X 22-25 /-Lm.
Material examined. Holotype (fragment): BMNH Acanthostyles (Figs 3D, 4B), heavily spined (seem to be the same
1884.4.14.6 -labeled 'From Type, South Australia'. spicule type), 225-300 X 20--25 /-Lm. Anchorate isochelae (Figs 3E,
Description (partly from Carter, 1883b). A thick cluster of 4B) in two size classes, 27-32 and 40--57/-Lm. Distribution and
branches, up to 6 cm wide and 0.4 cm thick, rising from a hard con- ecology. Cape Town area, South Africa, on sandy rocky bottom
tracted peduncular base. The sponge may form a clathrate blade with shells, 14-51 m. From this description it is clear that
by anastomosis. The remnants now representing the holotype Ectyonancora Jlabellata is almost identical with Ectyonopsis
Porifera· Demospongiae • Poecilosclerida • Myxillina • Myxillidae 605

Fig. 2. Ectyonopsis ramosa Carter, l883b, A, lectotype BMNH 1884.4.14.6 (scale 1 em). B, drawing of megascleres reproduced from Carter's, l883b:
pI. XIII fig. 5 (sizes see text). C-E, microphotos of skeleton of lectotype (scale 100 /-Lm) (arrows in D indicate echinating acanthostyles, in E isochelae).
F-I, SEM images of spicules of lectotype (scale 10 /-Lm).

ramosa, the only difference of substance being the presence of some are unguiferate (Fig. 3H) with reduced short teeth, no clear
a second size category of isochelae. Several species have been size categories, 30-51 f1m. Skeleton (Fig. 4C). Ectosomal spicules
described in this genus: E. pluridentata Levi (1963), E. panis loosely arranged, for the most part tangential; choanosomal skele-
Boury-Esnault & van Beveren (1982) and E. walvisensis Driz ton somewhat irregular isotropic reticulation. This species is
(1989), all from Southern Ocean localities. The latter is not an included in Ectyonopsis because of the overall close similarity in
Ectyonopsis on account of its possession of long subectosomal skeletal structure. The lack of echinating acanthostyles and the
styles. It conforms to Plocamiancora (cf. below). presence of spines on the tornote apices is judged to be of specific
The genus Stelotrochota Bakus, 1966b was erected (by mono- value, since these are common enough features of many individual
typy) for Stelotrochota hartmani Bakus (1966b: 501, pi. 2b). A myxillid species.
paratype (Fig. 3F) was examined, BMNH 1964.12.1.3, including
slide 1964.12.1.3a, from San Juan Archipelago, Washington State,
USA. This has ectosomal tylotes (Figs 3G, 4C), with one end globu- HYMENANCORA LUNDBECK, 1910
lar and the other more elongate, both microspined on the head,
139-166 X 7-9 f1m. Acanthostrongyles (Figs 31, 4C), with swollen Synonymy
ends which are slightly unequal, entirely heavily spined, especially
on the swollen ends, 185-211 X 19-22 f1m. Chelae (Fig. 4C) robust, Hymenancora Lundbeck, 1910. Leptosiopsis Topsent, 1927b:
strongly curved, some are anchorate spatulate with well-developed 13. Hymesigmia Topsent, 1927b: 14. Leptosia sensu Topsent,
alae, others are polydentate, likewise with 5 well-developed teeth, 1928c: 56 (not Topsent, 1892b: xxii). Chiastosia Topsent, 1928c:
606 Porifera· Demospongiae • Poecilosclerida • Myxillina • Myxillidae

Fig. 3. A-E, Ectyonopsis jlabellata (Levi, 1963 as Ectyonancora). A, holotype (scale I cm). B-E, SEM photos made from spicules of the holotype
(scale 10 /Lm). F-I, Ectyonopsis hartmani (Bakus, 1966b as Stelotrochota). F, habitus of paratype (scale I cm). 0-1, SEM photos of spicules from paratype
(scale 10 /Lm).

276. Arndtanchora de Laubenfels, 1936a: 110. Jelissima de Previous reviews

Laubenfels, 1936a: 79. Herceus Koltun, 1959: 173.
Lundbeck (1910: 116), Desqueyroux-Faundez & Van Soest
Type species (1996: 46).

Leptosia umbellifera Topsent, 1904b: 192 (by subsequent Description of type species
designation; de Laubenfels, 1936a: 87).
Hymenancora umbellifera (Topsent, 1904b) (Fig. 5A).
Definition Synonymy. Leptosia umbellifera Topsent, 1904b: 192,
pI. XV fig. 4; Hymenancora umbellifera; Lundbeck, 1910: 116.
Myxillidae with hymedesmioid arrangement of acanthostyles; Material examined. Holotype: MOM (not seen).
microscleres include polydentate anchorate chelae. Schizotype: MNHN DT 1039 - '#40, Prince Albert 1895'.
Description (mostly from Topsent, 1904b). Smooth crust on
Diagnosis Siphonidium, 0.5 mm in thickness. Colour white. Skeleton.
Choanosomal skeleton hymedesmioid, with acanthostyles erect on
Encrusting, with smooth or papillate surface; ectosomal skele- the substrate, heads down; bundles of tomotes go to the surface,
ton with smooth ectosomal strongy1es or styles, occasionally with where they are arranged tangentially. Spicules. Ectosomal polyty-
polytylote tyles; choanosomal skeleton hymedesmioid with acan- 10te tomotes, somewhat anisotylote, smooth, 345 X 4.5 /-Lm.
thostyles erect on the substrate, often in two functional categories, Acanthostyles in a large size range, 130-450 X 7-12 /-Lm. Chelae
structural and echinating; microscleres are isochelae - occasionally anchorate or unguiferate, po1ydentate, with on the average 5 teeth
anisochelae - often with polydentate alae, and sigmas, which are but may be up to 7-9 teeth, 26-30/-Lm. Distribution and ecology.
often absent. Approximately 20 species. Azores, deep water.
Porifera· Demospongiae • Poecilosclerida • Myxillina • Myxillidae 607

_ _~ ooo

A B

----- ,,0

1 I ~~
.; 1
.0
[
...
C
D
Fig. 4. A, Damiriopsis brondstedi Burton, 1928a, drawing of skeleton and spicules. B, Ectyonopsisflabellata (Levi, 1963 as Ectyonancora), drawing of
skeleton and spicules made from a slide of the holotype. C, Ectyonopsis hartmani (Bakus, 1966b as Stelotrochota), drawing of skeleton and spicules made
from a slide of the BMNH paratype. D, Hymenancora tenuissima (Thiele, 1905 as Hymedesmia), drawing of skeleton and spicules (all four drawings made
from slides of the respective type specimens).

Remarks. Most species of the genus are thinly encrusting this genus, but not all of these assignments have been followed by
deep-water species. The genus unites those former Hymedesmiidae authors after Lundbeck. A 14th record is that of Hymenancora
that have anchorate, unguiferate or polydentate chelae. Because lundbecki Hentschel, 1912.
hymedesmioid architecture is a suspect character, it is possibly an Topsent's (1927b) genus Leptosiopsis was erected (by mono-
artifactual group, which may need to be split up and distributed typy) for L. inaequalis (1927b: 13). Four type slides were exam-
over other myxilline genera based on similarities of spicules. The ined, MNHN DT 1083. This is a thinly encrusting species from
type species designation by de Laubenfels could be challenged, deep water (2165 m) off the Azores. The genus character is the
since he can hardly be considered a 'first reviser' in the sense of the peculiar polydentate anchorate anisochela, which resembles those
ICZN. His choice of Topsent's species Leptosia umbellifera is of Hymenancora in their 'normal' half (Fig. 5C). The skeletal
rather random, and it might have been more logical to indicate one architecture is hymedesmioid with two size categories of acan-
of Lundbeck's extensively described species. Still, Lundbeck thostyles, 217-245 X 20 J.Lm, and 120-135 X 15 J.Lm; ectosomal
indeed named this species, as third in a list of literature records spicules are stylote, poIytyIote anisotomotes of 180-210 X 2 J.Lm.
considered by him to be members of his new genus. Lundbeck The genus Hymesigmia was erected by Topsent (1927b) for
assigned 13 species earlier described in the genus Hymedesmia to H. japycina Topsent (1927b: 14) (by monotypy). A type slide
608 Porifera' Demospongiae • Poecilosclerida • Myxillina • Myxillidae

D B c
".
"
... .:

'.
I I
I I

E F
Fig. 5. Hymenancora species. A, Hymenancora umbellifera (Topsent, 1904b as Leptosia), drawing of chelae, reproduced from Topsent's plate XV fig. 4.
B, Hymenancorajapycina Topsent (1927b, as Hymesigma), drawing of spicules reproduced from Topsent's (I928c, plate IX fig. 26). C, Hymenancora
inaequalis Topsent (1927b, as Leptosiopsis), drawing of spicules, reproduced from Topsent's (I928c, pI. IX figs 18-19). D, Hymenancora pecqueryi
Topsent (l892a, as Myxilla) , type of Chiastosia Topsent, 1928c, drawing of spicules reproduced from Topsent's plate IX fig . 16. E, Hymenancora
sirventi Topsent (I927b as Leptosia), type of Amdtanchora, drawing of spicules reproduced from Topsent's (l928c) plate IX fig. 17. F, Hymenancora
orienta lis Koltun (1959, as Herceus), drawing of spicules reproduced from Koltun's fig. 134 (scales for all drawings see text).

MNHN DT 1085 was examined. This thinly encrusting Hymesigmiaa junior synonym of Hymenancora despite the absence
hymedesmioid specimen was collected at the same deep-water sta- of chelae in its type species.
tion (2165 m) off the Azores as Leptosiopsis inaequalis. The slide Topsent (1928c) restricted the use of his genus Leptosia
preparations of both have many peculiarly incurved sigmas 70 /-Lm Topsent (l892b: xxii) to Hymedesmia-like species with anchorate
high and 85 /-Lm wide (Fig. 5B), which were described only for H. chelae, thus making it implicitly a possible senior synonym of
japycina by Topsent (they are the genus character for Hymesigmia), Hymenancora. However, since no anchorate-bearing species were
but appear so common in the slide of H. inaequalisas to be an assigned to Leptosia when originally erected, this appears to be
unlikely contamination. The two specimens differ in the absence of illegal. Also, Leptosia is preoccupied by Leptosia Hubner, 1818
the peculiar anchorate anisochelae in H. japycina, in the larger size (Lepidoptera), see elsewhere in the chapter on Hymedesmiidae
of the acanthostyles (up to 660/-Lm) and also in the stylote nature of under Hymedesmia (Sty/opus).
the ectosomal spicules (which may be up to 315 /-Lm). It is possible Topsent's (1928c: 276) genus Chiastosia was erected for type
they are closely related, and thus it is proposed to consider species (original designation) Myxilla pecqueryi Topsent, 1892a
Porifera· Demospongiae • Poecilosclerida • Myxillina • Myxillidae 609

(including var. minima). A type slide was examined (MNHN DT Definition

940), as well as a slide of the var. minima (MNHN DT 1179). The
genus was founded on the peculiar chiastosigmas (Fig. 5D), con- Myxillidae with smooth ectosoma1 tylotes. Microscleres
sisting of two normal sigmas apparently attached in their centre include spherancoras.
and thus forming spider-like spicules. Such spicules occur also in
unrelated Acanthancora clavatancora Topsent (1927b), and Diagnosis
Guitarra cf. indica sensu Van Soest (1988). With its hymedesmioid
arrangement of strongylote tomotes, 470 X 6 fLm, acanthostyles Fistular growth form, with paper-like thin ectosome com-
in two size classes of 420 and 120 fLm, and 5-teethed anchorate posed of smooth ectosomal tylotes, smooth choanosomal styles,
chelae, this deep water (1250-2460m) species from the Azores microscleres are spherancoras (also known as me10nche1as) and
conforms to the present genus; its peculiar choastosigmas are anchorate chelae. Four species all from the Northern Hemisphere,
considered a specific, but not a generic character. concentrated in cold deep water.
De Laubenfels (1936a: 110) erected the genus Arndtanchora
for type species Leptosia sirventi Topsent (1927b: 12) (by original Previous review
designation). A type slide MNHN DT 1218 was examined. This is
a deep water (2460 m) encrustation from the Azores. It has acan- Desqueyroux-Faundez & Van Soest (1996).
thostyles of two sizes, 840 X 33 and 160-210 X 10 fLm, smooth
ectosomal anisostrongyles of 230-385 X 6.5 fLm, and two categories Description of type species
of chelae (Fig. 5E) one of which is the usual anchorate
type of 50--60 fLm, with on average about 4 teeth, and the other Melonanchora elliptica Carter, 1874a (Fig. 6).
is peculiar in being cleistochelate, approximately 20 fLm; it is Synonymy. Melonanchora elliptica Carter, 1874a: 212,
here assumed that these are derived from anchorates. Thin sigmas pI. XIII figs 6-12, pI. XV fig. 35a-b; Vosmaer, 1885a: 31, pI. I
occur in low numbers and may be foreign. A second species assigned fig. 14, pI. V figs 69-70.
to Amtanchora is Hymedesmia exigua Kirkpatrick (1907a: 273); it is Material examined. Ho10type: BMNH 1882.7.28.54 - N
likewise a clear Hymenancora by its possession of anchorate chelae Atlantic, 'Porcupine' Exped., 1869. Other material. ZMA POR.
with 5 teeth (or 6, see Kirkpatrick, 1908c: pI. XXVI fig. 2c1). 1548-Arctic, Willem Barents Exped., stat. 4 (Vosmaer's material).
The genus lelissima de Laubenfels (1936a: 79) was erected Description. Globular, surface corrugated, with projecting
for type species (by original designation) Hymedesmia tenuissima tubercules. Skin stiff, bladderlike. Size up to 10 cm diameter.
Thiele (1905: 454, fig. 70). This was assigned to Ectyomyxilla Fragile, surface parchment-like. Colour grey. Skeleton. Ectosomal
by Desqueyroux-Faundez & Van Soest (1996: 41), but subse- skeleton is a well-developed skin-like crust of tangential tylotes.
quent examination of 'co-type' (=paratype) material BMNH Choanosomal skeleton ill-defined, vague tracts of styles. The inte-
1931.3.27.13 (with slide BMNH 1930.11.28.6, ex Berlin Museum rior is mostly a soft organic mass without spicule tracts. Spicules.
collection) revealed that Lundbeck was right in assigning this to Ectosomal tomotes (Fig. 6A) smooth, tylote or subtylote, with
Hymenancora. It has an ectosomal skeleton consisting of brushes unequal ends, one being swollen, the other evenly rounded,
of tomotes, 200 X 8 fLm, fanning out to form a tangential crust 450-615 X 13-15 fLm. Choanosomal smooth styles (Fig. 6B),
(Fig. 4D); choanosomal skeleton composed of single acan- 650-860 X 13-15 fLm. Anchorate isochelae (Fig. 6D), relatively
thostyles, possibly in two size categories,l00-290 X 7-15 fLm, elongate and slim, with spatulate alae and well developed fimbriae,
erect on the substratum. Microscleres anchorate isochelae, 10-30 in two size categories, 24-44 and 60 fLm. Spherancoras (mel-
fLm, of which the larger have 5 teeth, and sigmas, 45-48 fLm. The onchelas, Fig. 6C), 48-68 fLm. Distribution and ecology. Arctic,
tomotes are peculiar in being oxeotes with small spines scattered North Atlantic (including Caribbean?); deep water, 135 m.
over the whole shaft, but no similarity to crellid arrangement exists. Remarks. This is a distinct genus on account of its peculiar
Koltun's (1959) genus Herceus erected (by monotypy) for microscleres, which are derived from anchorate isochelas by fusion
North Pacific Herceus orientalis Koltun (1959: 173, fig. 134, here and proliferation of the alae (cf. Fig. 6C). Schmidt (1880b)
reproduced as Fig. 5F) (holotype quoted to be in the Zoological reported this species also from the Caribbean, but that record needs
Institute of the Academy SSSR), differs from the type species in to be confirmed, since Schmidt only compared the microscleres. A
having stylote tomotes, but is otherwise similar in skeleton. It has a second North Atlantic species was described by Schmidt (1875),
hymedesmioid arrangement of acanthostyles of a wide size range, viz., M. emphysema. It has larger and thicker tomotes (900 X
176-416 X 12-21 fLm. Tomotes, 290-416 X 8-10 fLm, arranged in 25 fLm), and lacks styles. Koltun (1959, 1970) described two
tufts or fibres. Chelae polydentate and unguiferate, 23-28 fLm. The further species M. kobjakovae and M. tetradentifera from the
species was collected at 28-63 m depth. North Pacific.

MELONANCHORA CARTER, 1874 MYXILLA SCHMIDT, 1862

Synonymy Synonymy

Melonanchora Carter, 1874a: 212. Myxilla Schmidt, 1862. Dendoryx Gray, 1867a: 535.
Emplocus Gray, 1867a: 535. Tereus Gray, 1867a: 537. Hastatus
Type species Vosmaer, 1880: 136. Ectyomyxilla Hentschel, 1914. Crellomyxilla
Dendy, 1924: 364. Burtonanchora de Laubenfels, 1936a: 94.
Melonanchora elliptica Carter, 1874a: 212 (by monotypy). Styloptilon Cabioch, 1968b: 236.
610 Porifera· Demospongiae • PoeciioscIerida • Myxillina • Myxillidae

Fig. 6. Melonanchora elliptica Carter (1 874a). SEM photos of spicules. A. tylote-strongyle (scale IOO/Lm). B. details of tylote-strongyle and style (scale
10 /Lm). C-D. microscIeres (scale 10 /Lm). C. growth stages of spherancoras. D. two size categories of anchorate isochelae.

Type species Previous review

Halichondria rosacea Lieberkiihn. 1859: 521 (by subsequent Desqueyroux-Faundez & Van Soest (1996).
designation; de Laubenfels. 1936a: 85).
Remarks
Definition
In accordance with proposals made by Desqueyroux-Faundez &
Myxillidae with ectosomal tylote tomotes with spined apices Van Soest (1996: 26) a wide definition of Myxilla is here
and a choanosomal reticulation of spined or smooth styles; employed. Three of the genera synonymized here with Myxilla
microscleres include anchorate chelae with only 3 teeth. were previously considered valid by various authors. They differ
from Myxilla s.s. in characters presumed artificial: presence of
Diagnosis a second. smaller category of acanthostyles (Ectyomyxilla)
possession of smooth instead of spined styles (Bunonanchora),
Encrusting. massive, stipitate or branching sponges. Surface and plumose rather than reticulate arrangement of the skeleton
irregular to conulose. The ectosomal tylotes have variable termina- (Styloptilon). Such characters, though hardly likely to be of phy-
tions: mucronate. frequently with 1 or few prominent spines, or logenetic significance, still may serve to separate groups among
exceptionally entirely smooth. forming both a tangential skeleton the large number of species answering to a wide definition of
and vertical spicule brushes; choanosomal spicules stylote, usually Myxilla employed here. These groups are used here as con-venient
styles with or without spined bases. forming a tight meshed subgenera. In contrast with Desqueyroux-Faundez & Van Soest
isotropic reticulation or looser isodictyal skeleton; echinating acan- (1996) the genus Stelodoryx (with synonyms Onychomyxilla and
thostyles often present but sometimes lost; microscleres are anchor- Pseudomyxilla) is retained as valid, based on the possession of
ate spatulate or unguiferate isochelae with 3 teeth. often of several polydentate chelae (5 or more complete alae). The use of Myxilla is
sizes, and sigmas (the latter sometimes lost). About 85 species. limited to species possessing chelae with only 3 complete alae.

Key to subgenera of Myxilla

(1) Megascleres forming the main skeleton entirely smooth ............................................................................. Myxilla (Burtonanchora)
Megascleres of the main skeleton spined on the heads or entirely spined ........................................................................................... 2
(2) Skeleton an isotropic reticulation ........................................................................................................................................................ 3
Skeleton plumose ................................................................................................................................................. Myxilla (Styloptilon)
(3) A second smaller category of spined megascleres present .............................................................................. Myxilla (Ectyomyxilla)
Only a single category of spined megascleres .......................................................................................................... Myxilla (Myxilla)
Porifera· Demospongiae • Poecilosclerida • Myxillina • Myxillidae 611

SUBGENUS MYXILLA SCHMIDT, 1862 becoming branched - erect in deeper waters (Lieberktihn's type
was described as finger-thick anastomosing branches). It may
Synonymy form upright growths around hydroid stems. Surface has a loose,
porous appearance (Fig. 7B), similar to that of M. incrustans.
Myxilla Schmidt, 1862. Dendoryx Gray, 1867a: 535. Emplocus Slightly hispid. The oscules are scattered across the animal, and
Gray, 1867a: 535. Tereus Gray, 1867a: 537. Hastatus Vosmaer, have raised edges. Exhalant channels run up to the oscules; not
1880: 136. apparent when preserved. Considerable quantities of slime are
exuded. Consistency loose, somewhat crumbly, elastic. Colour
Type species yellow, or dirty rose-red, or brownish orange when alive. Skeleton
(Fig. 8A). ectosomal tomotes are arranged in vertical brushes at
Halichondria rosacea Lieberkiihn, 1859: 521 (by subsequent the surface. The choanosomal skeleton is an isotropic paucispicu-
designation; de Laubenfels, 1936a: 85). lar reticulation of acanthostyles, without echinating spicules.
Microscleres are present throughout the sponge body. Spongin is
Definition scarce. Spicules (Figs 7A, 7 C-H, 8A). surface tomotes bear three
or four small spines at the ends (Figs 7C-D, 8A) and are 140--170 X
Myxilla with isotropic skeleton made up of acanthostyles in 2-3 j.Lm long (Lieberktihn: 170 j.Lm). Megascleres of the choanoso-
a single size category. mal skeleton are acanthostyles whose spines are few but strong
(Fig. 7E-F, 8A), and present throughout the length of the spicule,
Diagnosis 140--200 X 3-6 j.Lm (Lieberktihn pictured 2 acanthostyles of
170--200 X 5 j.Lm). The microscleres are spatuliferous anchorate
Predominantly massive to lobate sponges with punctate surface chelae (Figs 7G, 8A) of two sizes, 20--25 j.Lm, and 12-15 j.Lm, and
and often slimy when handled out of the water. Oscules usually promi- small sigmata (Figs 7H, 8A), ca. 14-35 j.Lm (Lieberkiihn pictured
nent on apex of lobes or along ridges. Consistency firm but crumbly. 4 sigmas of 14-35 j.Lm, see Fig. 7A). Distribution and ecology.
Ectosomal megascleres usually short diactines, often with spined or Mediterranean, Atlantic coasts of France and Spain, British Isles,
mucronate endings. Choanosomal megascleres a single size category Faroes, Arctic; exposed sites, on rock, on shells, from the shore
of short acanthostyles. Skeleton close-meshed isotropic, with 2 or down to 300 m.
more acanthostyles forming the sides. Microscleres abundant anchor- Remarks. The type is presumably lost and Schmidt's mate-
ate isochelae in one or more size categories and usually sigmas, often rial from Triest (Fig. liB) is topotypical, thus it is proposed to
also in more than one size category. Myxilla (Myxilla) species have adopt this as neotype. In Lieberktihn's original description there is
been reported from Northern Hemisphere and tropical waters, but no mention of chelae, nor are they figured (cf. Fig. 7A) and the
appear rare or absent in Southern Oceans. description of the skeleton is also atypical: "Die Nadelztige beste-
hen aus vielen nebeneinanderliegenden Nadeln von zweierlei
Remarks Form ... ". Schmidt (1862) at first just repeated this description but
later (Schmidt, 1868: 27) acknowledged his oversight and reported
The separation of Myxilla species with only a single acan- the occurrence of 'Doppelhaken' in the Triest specimens of
thostyle category is probably artificial, because the skeletal struc- M. rosacea. Super-ficially, M. rosacea can be very similar to
ture of Myxilla species with a second category is often very similar. M. incrustans, but it does not seem to form the raised, oscule-
Still it allows easier identification among the many Myxilla species bearing ridges of the latter species. Examination of the tomote ends
described. The distinction between arcuate and anchorate chelae at high power is necessary to confirm identification. Typical mem-
was not made until Levinsen (1893b) and Lundbeck (1905, 1910) bers of this subgenus next to the type species are M. incrustans
discussed the issue and pointed out the subtle but consistent differ- (Johnston, 1842), M. Jimbriata (Bowerbank, 1882), and
ences between these chela types. Several Myxilla species described M. mexicensis (Dickinson, 1945), but a comprehensive enumera-
previous to Lundbeck, notably those of Ridley & Dendy (1887) tion is beyond the scope of the present revision.
possess arcuate chelae and need to be referred to coelosphaerid The genus Dendoryx Gray (1867a: 535) was erected for type
genera such as Lissodendoryx and Ectyodoryx. species (by original designation) Halichondria incrustans sensu
Bowerbank, 1866: 249 (=H. incrustans Johnston, 1842). The
Description of type species type specimen of this species could not be found in the BMNH
collection, accordingly it is proposed to designate the figured
Myxilla (Myxilla) rosacea (Lieberkiihn, 1859) (Figs 7, 8A). specimen in Bowerbank (1874b, pI. XLIV fig. 7), BMNH
Synonymy. Halichondria rosacea Lieberkiihn, 1859: 521, 1930.7.3.271, Hastings, Bk 2013, as neotype. For a description of
pI. XI fig. 2; Myxilla rosacea; Schmidt, 1862: 71; Vosmaer, 1880: this species one is referred to Ackers et al. (1992) and Van Soest
123; Ackers et al., 1992: 124, fig. 209. Myxilla fasciculata; et at. (1999).
Schmidt, 1862: 71 (not: Lieberktihn, 1869); Topsent, 1920a: 14. The genus Emplocus Gray (1867a: 535) was erected (by
Myxilla tridens Schmidt, 1864: 36, pI. IV fig. 5. original designation) for type species Myxilla tridens Schmidt
Material examined. Neotype (designated herein): LMJG (1864: 36) which has been synonymized with M. rosacea by
15293 - Schmidt collection, Triest (cf. Desqueyroux-Faundez & Vosmaer (1880: 123) and Burton (1930c: 533), thus making
Stone, 1992: 14, fig. 141). Other material. ZMA POR. 00265 - Emplocus an objective junior synonym. Examination of a specimen
Baie de Morlaix, NW France, 10m. of Schmidt, LMJG 15291, and a slide of Schmidt from
Description (mostly from Ackers et al., 1992). Cushions the Adriatic, BMNH 1868.3.11.98, confirmed the previous
0.5-1 cm thick (Fig. 7B), forming patches up to 15 cm across; conclusions (cf. Fig. 7C-H).
612 Porifera· Demospongiae • Poecilosclerida • MyxiUina • Myxillidae

'==
\

A B

Fig. 7. Myxilla rosacea (Lieberkiihn, 1859 as Halichondria), type of Myxilla. A, drawing of spicules reproduced from Lieberkiihn's pI. XI fig. 2 (size see
text). B, Schmidt's (1862) fragments of the Venice specimen of Myxilla rosacea, proposed as neotype material, reproduced from Desqueyroux-Faundez &
Stone's (1992, fig. 141) (scale 1 em). C-H, SEM photos of spicules made of one of Schmidt's type specimens of Myxilla tridens, LMJG 15291 (considered
a junior synonym of M. rosacea) (scales IO j-Lffi, except D, 1 j-Lffi).

The genus Tereus Gray (1867a: 537) was erected (by original SUBGENUS BURTONANCHORA DE LAUBENFELS, 1936
designation) for type species Isodictya fimbriata (Bowerbank,
1866: 337), a species quite similar to M. rosacea, differing only in Synonymy
life appearance (more orangy, no slime) and in details of tomote
apices; sigmas tend to be rare. The dry type from Shetland and Burtonanchora de Laubenfels, 1936a: 94.
three unregistered slides, BMNH #Bk 759, were examined and
found to conform to descriptions in, for example Ackers et al., Type species
1992 and Van Soest et al., 1999; sigmas were not found in the type
slide. Myxilla cruci/era Wilson, 1925: 430 (by original designation).
The genus Hastatus Vosmaer (1880: 136) was erected
(by original designation) for Isodictya lurida Bowerbank (1866: Definition
336), a synonym of Myxillafimbriata (Bowerbank, 1866), which
makes this genus an objective synonym of Tereus Gray (l867a), Myxilla with choanosomal styles smooth, never spined.
which in its tum is doubtlessly a synonym of Myxilla (Myxilla).
Slides of the type, BMNH unregistered, Bk 758, of unknown Diagnosis
origin, bearing the indication 'Type' and of a second specimen,
here assigned paralectotype #Bk 757, from the coast of Devon. Flabelliform or massive sponges. Skeleton a reticulation of
They conform to Myxilla fimbriata, and likewise do not contain smooth styles. Ectosomal skeleton bundles of tomotes fanning out
sigmas. to form a tangential skeleton at the surface. Microscleres tridentate
The genus Stegxella Bowerbank (1874b) was synonymized anchorate isochelae and sigmas. Species answering to the defini-
with Myxilla by de Laubenfels (1936a: 85), but it is here reassigned tion of Burtonanchora have been reported from tropical as well as
as a synonym of Hymedesmia (q.v.). polar areas, and also in deep water.
Porifera· Demospongiae • Poecilosclerida • Myxillina • Myxillidae 613

I
"

A c

----,..

Fig. 8. A, Myxilla (Myxilla) rosacea (Lieberkiihn. 1859), drawing of skeleton and spicules made from a representative specimen in the ZMA collection
(ZMA POR. 00265). B, Myxilla (Burtonanchora) crucifera Wilson (1925), habitus and spicules, reproduced from Wilson's pIs 42 fig. 5 and 49 figs 3-4
(size see text). C, Myxi/la (Ectyomyxilla) kerguelensis (Hentschel, 1914), drawing of skeleton and spicules made from the type of Crellomyxilla interme-
dia Dendy (1924), considered a junior synonym. D, Myxilla (Styloptilon) ancoratum (Cabioch, 1968b), drawing of skeleton and spicules made from a slide
of the holotype.

Remarks Description. (from Wilson, 1925). Flabelliform (Fig. 8B),

about 4 X 5 X 0.8 cm. The convex surface bears regularly distributed
The smooth styles may seem a clear distinction from the oscules of 1-2mm in diameter. The concave surface is irregular
spined styles found in most Myxilla species. However, some and bears closely set pores. Firm consistency. Light brown (alco-
species are known to have only a few spines or very faint spination, hol). Skeleton. ectosomal tomotes form brushes at the surface
making this distinction dubious. which fan out to form a tangential layer. The main skeleton is
isotropic in the interior but may become somewhat anisotropic at
the periphery. Meshes are enclosed by 2-6 spicules. Spicules
Description of type species (Fig. 8B). Tomotes oxeote, smooth straight, 165-200 X 6-7 f.Lm;
styles, short, fat, entirely smooth, 270-300 X 18-21f.Lm; anchorate
Myxilla (Burtonanchora) cruci/era Wilson, 1925 (Fig. 8B). isochelae in two size categories, 40 and lOf.Lm; sigmas in two size
Synonymy. Myxilla cruci/era Wilson, 1925: 430, pI. 42 fig. 5, categories, 50-70 and 10 f.Lm. Distribution. Philippines.
pI. 49 figs 3-4; Burtonanchora crucifera; de Laubenfels, 1936a: 94. ReTlUlrks. Several other species were assigned to
Material examined. None. Holotype: USNM 21359- Burtonanchora by de Laubenfels (1936a): Myxilla lacunosa
'Albatross' Philippine Expedition, stat. D5168 (not examined). Lambe (1892: 70), M. hastata Ridley & Dendy (1886: 472),
614 Porifera· Demospongiae • Poecilosclerida • Myxillina • Myxillidae

M. magna Topsent (1916: 168), M. pedunculata Lundbeck (1905: Description of type species
148), M. pistillaris Topsent (1916: 169), M. spongiosa Ridley &
Dendy (1886: 471) and Amphilectus osculosus Topsent (1893c). Myxilla (Ectyomyxilla) kerguelensis (Hentschel, 1914)
Levi (1960a, 1963, 1965) added Burtonanchora myxilloides (West (Figs 8C, 9).
Africa), B. sigmatifera (South Africa) and B. gracilis (Red Sea). Synonymy. Ectyomyxilla kerguelensis Hentschel, 1914:
Desqueyroux-Faundez & Van Soest (1996: 34) added a further 103, pI. IV fig. 10, pI. VII fig. 10. Crellomyxilla intermedia Dendy,
species, Myxilla (Burtonanchora) asymmetrica to this group, but 1924: 364, pI. XV figs 16-21.
reassigned (p. 31) M. spongiosa to the subgenus Myxilla (Myxilla) Material examined. No type material of Hentschel was
(as junior synonym of Myxilla (Myxilla) moWs Ridley & Dendy reexamined. Holotype of Crellomyxilla intermedia Dendy: BMNH
(1886: 471) on account of the slight spination of the styles. 1923.10.1.145 - 'Terra Nova' Exped. stat. 90, Three Kings islands,
New Zealand, 180m (this is considered a junior synonym).
Description. Elongated, subcylindrical; branches of 7 cm
SUBGENUS ECTYOMYXILLA DENDY, 1924 diameter, 11.5 cm high. Surface uneven, granular. Dendy's frag-
ments (Fig. 9A) are black in alcohol, but apparently this was
Synonymy induced by osmic acid; other smaller fragments are white in alco-
hol; Hentschel's material (Fig. 9G) was greyish brown. Skeleton.
Ectyomyxilla Hentschel, 1914: 103 (not Lundbeck, 1909: 445 Ectosomal skeleton (Fig. 8C) consists of two layers: an outer layer
=nomen nudum). Crellomyxilla Dendy, 1924: 364. of single acanthostyles perpendicular to the surface, points out-
wards, carried by partly tangentially arranged bouquets of tomotes.
Type species Choanosomal skeleton (Fig. 8C) isotropic, with triangular meshes
in the peripheral skeleton, becoming quadrangular deeper in the
Ectyomyxilla kerguelensis Hentschel, 1914: 103 (by original choanosome; meshes made up of 1-4 spicules. Hentschel records
designation). rare echinating acanthostyles. Spicules. Ectosomal acanthostyles
(Figs 8C, 9C, J), short and thick, entirely spined, 56-100 X
Definition 5-12/-lm; ectosomal tomotes (Figs 8C, 9D, H) with swollen
mucronate apices, 152-210 X 5-7/-lm; choanosomal styles (Figs
Myxilla with a smaller category of spined styles in addition to 8C, 9B, I), entirely acanthose, with thick spines, 176-225 X
the main (acantho-)styles. 14-17 /-lm). Microscleres. anchorate chelae (Figs 8C, 9E, K) in two
size categories, 28 and 16 /-lm (Hentschel's material: only a single
Diagnosis category, 12.5-19/-lm); sigmas (Figs 8C, 9F, L), 17-36/-lm.
Distribution and ecology. Kerguelen, New Zealand; deeper water.
Encrusting, massive, flabellate or branching sponges with Remo:rks. Crellomyxilla intermedia Dendy, 1924 shares
irregular surface. Skeleton isotropic paucispicular, which may most characters, including the smaller acanthostyles concentrated
become loosely anisotropic in some species. Megascleres include in the ectosome, with Myxilla (Ectyomyxilla) kerguelensis and
two size categories of acanthostyles, one of which may form a sur- is generally considered synonymous (compare Fig. 8A-F with
face palisade or is echinating the nodes of the skeletal meshes. 8G-L). A further species sharing this character is M. (E.) chilensis
Tomotes smooth or lightly spined on the apices. Microscleres (Thiele, 1905: 443, figs 22, 62). Species answering to the defini-
anchorate isochelae and sigmas. Many Myxilla (Ectyomyxilla) tion of Myxilla, but possessing echinating acanthostyles next
have been described from cold or deep water. to choanosomal styles are: M. arena ria Dendy (1905: 169),
M. (Ectyomyxilla) dracula Desqueyroux-Faundez & Van Soest
Remarks (1996: 37), Ectyomyxilla hentscheli Burton (1929a: 438), M. mariana
Ridley & Dendy (1886: 472), M. massa Ridley & Dendy (1886:
Contrary to what Desqueyroux-Faundez & Van Soest (1996: 473), M. mariana tylacantha Hentschel (1914), E. methanophila
27) maintained, the name Ectyomyxilla is a nomen nudum, because Maldonado & Young (1998a: 798), M. nobilis patagonica Ridley &
Lundbeck (1909: 445) did not mention any species when erecting Dendy (1886: 473), Myxilla parasitica Lambe (1893: 31), and
this genus. The subsequent use of the name by Hentschel (1914: E. ramosa Bergquist & Fromont (1988: 87).
103) does not satisfy Article 12.2.5 of the ICZN (Anonymous, The genus lelissima de Laubenfels, 1936a (type species
1999) as type species designation. However, the name may still be Hymedesmia tenuissima Thiele, 1905) was assigned to
used in the sense of Hentschel (1914), and it takes his authorship. Ectyomyxilla by Desqueyroux-Faundez & Van Soest (1996), but
Thus, its type species is Ectyomyxilla kerguelensis Hentschel, 1914 subsequent examination of 'co-type' material revealed that it
by monotypy. This group unites the Myxilla-like species with a sec- answers to the definition of Hymenancora (cf. above).
ond smaller category of entirely spined styles. In the type species
(cf. below) the smaller spined styles occur as an ectosomal pal-
isade. In other species considered to belong to the subgenus SUBGENUS STYLOPTILON CABIOCH, 1968
Ectyomyxilla, these spicules may occur in the usual echinating
position at the nodes of the choanosomal reticulation. Synonymy

Previous reviews Styloptilon Cabioch, 1968b: 236.

Lundbeck (1909), Hentschel (1914), Dendy (1924), Type species

Desqueyroux-Faundez & Van Soest (1996), Maldonado & Young
(1998a). Styloptilon ancoratum Cabioch, 1968b: 236 (by monotypy).
Porifera' Demospongiae • Poecilosclerida • Myxillina • Myxillidae 615

Fig. 9. Myxilla (Ectyomyxilla) kerguelensis (Hentschel, 1914 as Ectyomyxilla). A, two type fragments of Crellomyxilla intermedia Dendy (1924),
considered a junior synonym (scale 1 em). B-F, SEM photos of spicules made from the type material of Crellomyxilla intermedia (scales 10 f,lm).
G-L, Hentschel's drawings of habit and spicules reproduced from his pis. IV fig. 10 and pI. VII fig. 10 (sizes see text).

Definition Description (partly after Cabioch, J968b). Thinly encrusting,

about 1 mm in thickness, agglutinating shell debris. The surface
Myxilla with plumose skeleton. is finely hispid. Colour yellowish white in alcohol. Skeleton
(Fig. 8D). the dermal membrane is charged with microscleres and
Diagnosis carried by bouquets of tomotes, which also form short tangential
bundles. The bouquets of tomotes are in tum erected on the peri-
Encrusting; with a plumose skeleton of bundles of larger acan- pheral ends of the choanosomal spicule tracts, which consist of
thostyles echinated by similar acanthostyles. Tomotes anisoxeote. acanthostyles. The skeletal columns are thick and form a plumoretic-
Microscleres anchorate chelae and sigmas. So far only a single ulate skeleton, in which the longitudinal elements dominate. Spicules
species has been assigned to this subgenus and its status as a sepa- (Fig. 8D). Tomotes smooth with pointed, but unequal ends, one being
rate group is thus uncertain. Still, its plumose skeletal architecture slightly swollen; they may be somewhat polytylote or have an irregu-
prevents its inclusion in Myxilla (Myxilla). lar outline, 140--200 X 2.5-5 /-lm; acanthostyles are entirely spined
and perhaps divisible in two size categories, 100--135 X 5-8 /-lm and
Description of type species 190--280 X 5-10 /-lm, but they do not constitute two functional cate-
gories (main and echinating). Microscleres spatulate anchorate
Myxilla (Styioptilon) ancoratum (Cabioch, 1968b) (Fig. 8D). isochelae in two size categories, 35-48 /-lm and 11-15 /-lm; sigmas in
Synonymy. Styioptilon ancoratum Cabioch, 1968b: 236, a single variable size category, 32-50/-lm. Distribution and ecology.
fig. lOB; Sol6rzano & Urgorri, 1993: 64. Atlantic coasts of France and Spain; on pebbles down to 85 m.
Material examined. Holotype: specimen from the Roscoff Remarks. The species remains ill-known. Its characters are
area, incorporated in the collections of the Station Biologique de in between those of Phorbas (plumose columns, pointed tomotes,
Roscoff. two sizes of acanthostyles) and Myxilla (anchorate chelae).
616 Porifera· Demospongiae • Poecilosclerida • Myxillina • Myxillidae

- ,..

c
D
Fig. to. A, Plocamiancora denticulata Topsent (l927b), drawing of spicules reproduced from Topsent (1928c, pI. X fig. 17) (sizes see text). B, Psammochela
elegans Dendy (1916a) drawing of skeleton and spicules made from the slide of the holotype. C, Psammochela chaliniformis (Carter, 1885b as Dysidea),
type of Psammodoryx Burton (1934a), drawing of skeleton and polydentate chelae made from a slide of the holotype. D, Psammochela psammodes
(Hentschel, 1911 as Desmacidon), type of Anomomyxilla Burton (1934a), drawing of spicules reproduced from Hentschel (1911, fig. 19) (sizes see text).

PLOCAMIANCORA TOPSENT, 1927 Diagnosis

Synonymy Encrusting; smooth ectosomal tomotes; choanosomal skele-

ton plocamiid with an isotropic basal reticulation of single acan-
Plocamiancora Topsent, 1927b: 17. Plocamissa Burton, thostrongyles echinated by long microspined styles in ascending
1935c: 401. plumose axial columns; short acanthostyles echinating skeletal
tracts; microscIeres are unguiferate or polydentate anchorate
isochelae. Three species.
Type species
Previous review
Plocamiancora denticulata Topsent, 1927b: 17 (by original
designation). Topsent (1928c).

Definition Description of type species

Myxillidae with plocamiid skeletal structure. Plocamiancora denticulata Topsent, 1927b (Fig. lOA).
Porifera· Demospongiae • Poecilosclerida • Myxillina • Myxillidae 617

Synonymy Type species

Plocamiancora denticulata Topsent, 1927b: 17; Topsent, Psammochela elegans Dendy, 1916a: 126 (by monotypy).
1928c: 307, pI. X fig. 17. ? Plocamiancora arndti Alander, 1942:
54, pI. 12 fig. 16. Definition
Material examined. Lectotype (slide): MNHN DT. 1164 -
labeled 'St. 1420, 1902'. Paralectotypes (2 slides): MNHN DT. Myxillidae with a single megasclere category consisting of
1081 -labeled 'St. 1116, 1901'. smooth styles or strongyles. The skeleton consists of a reticulation
Description (from Topsent, J928c). Extended crusts, all on of megascleres and sand grains. Microscleres include anchorate
stony or gorgonian corals, with hispid surface. Colour greyish (in chelae, and/or small polydentates; sigmas may be present.
alcohol). Skeleton. plocarniid, i.e., with a basal isotropic reticulation
of acanthostrongyles, singly or in bundles of two or three, upon which Diagnosis
single long acanthostyles are erected, at their base echinated by
groups of smaller acanthostyles. Spicules (Fig. lOA). Smooth oxeote Massively encrusting. Ectosomal skeleton with a fine-meshed
anisotomotes with mucronate endings, 308-350 X 4--5 /-Lm; long reticulation of small sand grains; choanosomal skeleton irregularly
acanthostyles, somewhat curved at the base, spined only on the blunt reticulate, composed of thin fibres with poor spongin, cored with
end, 600-850 X 20--30/-Lm; smaller echinating acanthostyles, entirely sand and styles, and numerous sand particles and spicules dis-
spined, 224--490 X 10--12/-Lm; acanthostrongyles, strongly curved, persed within the mesohyl; megascleres are styles; microscleres are
spined predominantly at both ends, lighter inbetween, 140--160 X anchorate and polydentate isochelae and sigmas.
10 /-Lm; large unguiferate-spatulate chelae with 3-5 teeth, 73-85 /-Lm;
smaller unguiferate-polydentate chelae with 5-9 teeth, 38-52/-Lm. Remarks
Distribution and ecology. Azores; deep water, 1740--2460 m.
Remarks. This genus in many aspects represents the model The three species united here under Psammochela share the
Myxillina, as it contains all three skeletal zones found in the presence of sand columns replacing the choanosomal skeleton, and
various genera and families of this group, and resembles certain anchorate chelae or derived forms. In other respects they show
Microcionina genera such as Antho and Lithoplocamia. considerable differences: P. elegans has styles as megascleres,
Plocamiancora amdti Alander (1942) is probably a synonym of P. psammodes has strongyles, and P. chaliniformis lacks megascle-
P. denticulata, but was distinguished on the lack of polydentate res entirely. Their close relationship is uncertain.
(5-teethed) chelae. This is subject to variation. Driz (1987)
reported P. denticulata from 260 m off the coast of SW Africa, but Previous reviews
later (1989) assigned this to a new species of Ectyonancora,
E. walvisensis Driz (1989: 73). However, this is not an Ectyonopsis Dendy (1916a), Burton (1934a), Wiedenmayer (1989: 67).
(senior synonym of Ectyonancora) because of its possession of
long styles erect on the isotropic reticulation of acanthostrongyles. Description of type species
It conforms to the present definition of Plocamiancora, and
accordingly is renamed Plocamiancora walvisensis. Solorzano Psammochela elegans Dendy, 1916a (Figs lOB, 11F-J).
et al. (1991: 182, fig. 4) recorded a Plocamiancora spec. from the N Synonymy. Psammochela elegans Dendy, 1916a: 126, pI. I
coast of Spain. This clearly conforms to Rotuloplocamia octoradiata fig. 6, pI. III fig. 22.
Levi (1952) (see chapter on family Iotrochotidae). Material examined. Lectotype: BMNH 1920.12.9.36.
The type species (original designation) of Plocamissa Burton Paralectotype: BMNH 1920.12.9.37 - 2 unregistered slides CN
(1935c: 401), i.e., Plocamia igzo de Laubenfels, 1932: 102 (type XVIII.3.
DSNM 22058 from California re-examined), in spite of de DescriptWn. Irregular lamellar or digitate sponges (Fig. IIF),
Laubenfels' inadequate description, conforms in all important respects often forming long slender processes, up to 9 cm long 0.3-1 cm in
to Plocamiancora denticulata except for the smaller size of the poly- diameter. Surface irregularly rugose or finely conulose, but smooth
dentate chelae. The ectosomal skeleton contains strongly developed in between. Oscules small and scattered. No colour recorded.
bouquets of anisotomotes many of which have one mucronate and one Skeleton (Fig. lOB). Ectosomal skeleton consists of a tight-meshed
microspined rounded end (Fig. lIA). Because some of them are reticulation of fine sand grains, meshed 50--80 /-Lm in diameter, with
smooth (Fig. llA right) they were not distinguished by de Laubenfels few megascleres and rnicroscleres. Choanosomal skeleton an
from the choanosomal styles. The structural styles (Fig. lIB) are isotropical to irregular reticulation of spiCUle tracts mixed with sand
united in bundles and have spined heads, size 350 X 35 /-Lm; these are grains. Meshes 100--200/-Lm, tracts with up to 5 spicules in cross
echinated at the base by rare, thin acanthostyles (Fig. 11C) of approx- section. Spicules (Figs lOB, llG-J). No ectosomal megascleres;
imately 190 X 11 /-Lm; the basal/central isotropic reticulation is made choanosomal styles (Fig. 11 G), smooth straight, relatively robust,
up of dumbell-shaped acanthostrongyles (Fig. lID) of 130 X 13 /-Lm. rather uniformly about 160 X 5-7/-Lm; chelae in two categories, one
The chelae are small polydentate anchorates (Fig. liE) of 14 /-Lm. commonplace tridentate anchorate chelae (Fig. 111) of 24--25/-Lm
(these are not abundant and may possibly be foreign), and abundant
small polydentate chelae (Fig. 11H) of about 12 /-Lm, with 9 or more
PSAMMOCHELA DENDY, 1916 alae; sigmas (Fig. 111) in two size categories, 50/-Lm and 12/-Lm
(Dendy gives 33 /-Lm and a much smaller category). Distribution and
Synonymy ecology. India, Indonesia; rocky bottom, 5-7 m.
Remarks. The type species is one of the sand sponges which
Psammochela Dendy, 1916a: 126. Psammodoryx Burton, so frequently demonstrate reduced skeleton and spicule complement.
1934a: 556. Anomomyxilla Burton, 1934a: 555.
618 Porifera· Demospongiae • PoeciioscIerida • Myxillina • Myxillidae

Fig. 11. A-E, Plocamiancora igzo (de Laubenfels, 1932 as Plocamia), type of Plocamissa Burton (l935c), SEM photos of spicules made from the type
(scales 10 fLm; except E, 1 fLm). F-J, Psammochela elegans Dendy (1916a). F, fragments of the holotype (scale I cm). G-J, SEM photos of spicules made
from the type (scales 10 fLm; except H, 1 fLm). K-L, Psammochela chalinifonnis (Carter, 1885b as Dysidea), type of Psammodoryx Burton (l934a).
K, fragments ofholotype (scale I cm). L, SEM photos of poly dentate chelae (scale I fLm).

Burton (1934a) synonymized it with Phoriospongiafibrosa Ridley forming loose dermal brushes supporting a close-meshed tangen-
(1884a), but justification for this is thin, since this species has tial skeleton of sand and foreign spicules (sandy crust); choanoso-
oxeas as megascleres and these are quoted as much thicker than the mal skeleton consists of an irregular network of fibres
styles of Dendy's type; furthermore fibrosa has tridentate, rather incorporating sand and foreign spicules, and with plumose tracts of
than polydentate chelae. Burton's (1934a) record of P. fibrosa is strongyles; microscleres are rare spatulate anchorate isochelae,
dubious because he could not find any microscleres. Several other 15-17 /-Lm, and unguiferous isochelae, 9-10 /-Lm, and sigmas in two
species have been assigned to this genus: Psammochela rigida size categories, 25-32 and 10-12 /-Lm.
(Bowerbank, 1875b: 289, as Halichondria) from Singapore is close The genus Psammodoryx Burton, 1934a: 556 was erected (by
to P. elegans and a possible senior synonym; however, there is original designation) for type species Dysidea chaliniformis Carter
apparently no sand reticulation at the surface, which is a distinct (1885b: 217) from South Australia, collected from a depth of 35 m.
difference. P. recife and P. tylota Boury-Esnault (1973: 283) Syntypes BMNH 1886.12.15.341 (with slide) and 343 were re-
have tylotes as megascleres and arcuate isochelae. They do not examined. Since these represent two distinct species, a lectotype
conform to Psammochela and are very likely coelosphaerids, is here chosen: BMNH 1886.12.15.341 (see remarks below).
probably members of the subgenus Anomodoryx of the genus Originally the type specimen (Fig. 11K) was a compressed bunch
Lissodendoryx. of solid cylindrical stalks rising from a common stem; size about
The genus Anomomyxilla Burton (1934a: 555) was erected 30 X 12 X 2.5 cm. Branches small and 0.3-2 cm in diameter. Only
(by original designation) for type species Desmacidon psammodes a few fragments have survived. Oscules in rows on the branches.
Hentschel (1911: 322, fig. 19), type specimen not examined. This Consistency fragile (dry). Colour reddish brown. Skeleton
has an ectosomal skeleton composed of strongyles (Fig. lOD), (Fig. 1DC) composed of a neat isodictyal reticulation of sand grains
136-166 X 2-3 /-Lm, of the same form as found in the choanosome, without proper megascleres. Ectosomal skeleton a reticulation of
Porifera • Demospongiae • Poecilosclerida • MyxilUna • MyxilUdae 619

rounded meshes of ca. 100 fJ.m in diameter. The only spicules present choanosomal skeleton. The choanosomal megascleres are smooth
are polydentate unguiferate isochelae (Figs 1Oe, IlL) of 10-12 fJ.m. or spined styles, and a second category of smaller echinating acan-
There is confusion among the BMNH specimens labeled 'Dysidea thostyles may be present. Microscleres include polydentate (5 or
chaliniformis', due to the fact that two distinctly different species more alae) anchorate or unguiferate isochelae and sigmas.
have received identical or similar labeling. Burton (1934a) omitted Occasionally tridentate anchorate chelae may be also present.
to check the type specimens when he indicated this species as the About 10 species. Most representatives of this genus originate
type of his new genus Psammodoryx. BMNH 1886.12.15.341 con- from deep water or from boreal or Antarctic waters.
forms to the genus definition of Burton ('reduced Myxilleae (?)
with skeleton composed of a reticulation of horny fibres filled Remarks
with sand and foreign spicules; microscleres isochelae unguiferae')
and accordingly this is here chosen as the lectotype. The labels The name of the genus suggests stalked growth form such as
with this specimen in Miss S.M. Stone's handwriting say: found in the type species as a genus character, but growth form is
'?Chalinopsilla arborea var. micropora Lendenfeld, 1889, p. 149, not likely to constitute a valid generic character. Other growth
South coast of Australia J.B. Wilson Coli.', and an older label, pos- forms are also included and accordingly the genera Onychomyxilla
sibly in Carter's handwriting says "Dysidia chaliniformis fragment and Pseudomyxilla are here synonymized with Stelodoryx.
of main mass which is net (1790) Australia" (information provided
by Clare Valentine). The fragment of the type redescribed by Previous review
Dendy (1896) was also taken from this specimen. Confusion with
Dendy'S Chondropsis chaliniformis (attributed to Lendenfeld, Burton (1932).
1889a, because this author described specimens with strongyles
and sigmas not conforming to Dysidea chaliniformis) may well Description of type species
have occurred among the various curators of the BMNH collection.
The other presumed type specimen, BMNH 1886.12.15.343, Stelodoryx procera Topsent, 1904b (Fig. 12A).
though labeled as the holotype (in Miss S.M. Stone's handwriting),
and later reassigned to Psammochela by Burton, appears to belong Synonymy
to Echinoclathria, probably E. subhispida (Carter). It has proper
styles and lacks chelae. Possibly as a result of this long-time confu- Stelodoryx procera Topsent, 1904b: 174, pI. ill fig. 5, pI. XIV
sion, Burton (1934a) also recorded Great Barrier Reef material fig. 17, pI. xvrn fig. 2.
as Chondropsis chaliniformis (Carter), while quoting Dysidea Material examined. Holotype (slides): MNHN DT. 990 -
chaliniformis Carter, 1885b as a synonym, without referring to his labeled 'PA 1895 53'. Slides of topotypical specimens: MNHN
action of a few pages further, viz., narning this same Dysidea DT. 989 - labeled 'PA st. 1349'. DT. 991 - labeled
chaliniformis Carter, 1885b as type of his new genus Psammodoryx. 'PA 1897, 53'.
Burton (1934a) also included Desmacidon arenosa Whitelegge Description. A stalked species (Fig. 12A), 2-6 cm high;
(1906: 478) from South East Australia in Psammodoryx. On paper main body flattened, softly spongy, with grooved irregular surface.
this sounds very similar to P. chaliniformis, and this was also Colour pale brown. Skeleton. Ectosomal skeleton of brushes of
realized by Whitelegge. Differences he noted were a massive shape tornotes; the main body has a typical Myxilla-like isotropic reticu-
(7 X 4.5 X 3 cm), less clearly developed spongin fibres and a larger lation of 1-3 spicules per side; the stalk has a plumoreticulate
size of the chelae (15 fJ.m). arrangement oflong styles forming thick longitudinal bundles con-
nected here and there by short cross-bundles. Spicules. Ectosomal
anisotornotes, with a thicker more heavily spined end, and a thin-
STEWDORYX TOPSENT, 1904 ner more lightly spined end, 235-300 X 5 fJ.m; choanosomal styles
of the main body, entirely smooth, blunt-ending, 350-650 X
Synonymy 12 fJ.m; styles of the stalk, entirely smooth, pointed, 990-1300 X
20-23 fJ.m. Microscleres. Anchorate, 5-teethed chelae in two mor-
Stelodoryx Topsent, 1904b: 174. Onychomyxilla Topsent, phologically distinct forms, with spatulate, long teeth 35-55 fJ.m
1927b: 12. Pseudomyxilla Koltun, 1955b: 14. (possibly in two size categories), and thin, elongate chelae with
short spiky teeth (50-55 fJ.m). Distribution and ecology. Azores;
Type species deep water, on rocks or on stalks of gorgonians, 350-1200 m.
Remarks. The type species differs from other species
Stelodoryx procera Topsent, 1904b: 174 (by monotypy). described in this genus by its possession of clearly localized
choanosomal megasclere categories, probably correlated with the
Definition stalked shape. However, if more than one species would exist with
that differentiation, a separate (sub)genus may need to be recog-
Myxillidae with reticulate skeleton and a full complement nized for stalked species.
of megascleres; microscleres include polydentate anchorate Koltun (1955b) erected the genus Pseudomyxilla for P. vitjazi
isochelae. Koltun (1955b: 14, by monotypy) emphasizing the reduced unguifer-
ate teeth of the isochelae. However, S. procera possesses chelae
Diagnosis with reduced teeth next to spatulate chelae. The difference with
Pseudomyxilla thus becomes void. P. vitiazi (Fig. 12C) has tomotes
Stalked, lobate, massive or encrusting sponges. Resembling with microspined apices of 190-290 X 4-7 fJ.m, choanosomal acan-
Myxilla in most respects, including ectosomal tomotes and reticulate thostyles entirely spined, 435-520 X 20-29 fJ.m, and polydentate
620 Porifera· Demospongiae • Poecilosclerida • Myxillina • MyxilJidae

, ,

c
Fig. 12. A, Stelodoryx procera Topsent (1904b), drawings of habit and spicules reproduced from Topsent (1904b, pI. ill fig. 5, pI. XIV fig. 17, pI. XVill
fig. 2). B, Stelodoryx pectinata Topsent (1892a as Dendoryx), type of Onychomyxilla Topsent (1927b), drawing of spicules reproduced from Topsent (1892a,
pI. X fig. 6). C, Stelodoryx vitiazi (Koitun, 1955b as Pseudomyxilla), drawing of spicules reproduced from Koitun (1959, fig. 78) (all sizes see text).

unguiferate isochelae of 26-40 /-Lm. It is a deep water species Hymetrochota (cf. chapter on family Iotrochotidae) which has
(ll5-820m) from the NE Pacific. failed to develop its birotulas. Its characters are apparently stable
The genus Onychomyxilla Topsent (1927b: 12) was erected since several specimens were obtained from different localities off
(by original designation) for type species Dendoryx pectinata the Azores. The species is allocated to Stelodoryx because true
Topsent (1892a: 100, pI. X fig. 6). A MNHN slide (DT. 995) of the birotulas are absent.
holotype was examined, labeled 'No. 78. Camp. Hirondelle, 1888, Several Stelodoryx and Pseudomyxilla species have been
125', from the Azores area, 861-2460m. Its skeleton is basically a described from various parts of the world, most of them from cold
paucispicular reticulation of acanthostyles, with echinating acan- or deep water: S. chlorophylla Levi (1993: 49), S. cribrigera
thostyles at the nodes. Ectosomal skeleton of bundles of tylotes, (Ridley & Dendy, 1887: 472, as Myxilla), S. jlabellata Koltun
which are smooth, with well-developed tyles, 415-500 X 4-5 /-Lm; (1959: 119) S. lissostyla (Koltun, 1959: 124, as Pseudomyxilla),
choanosomal acanthostyles (Fig. 24B), curved, mostly smooth S. multidentata (Boury-Esnault & van Beveren, 1982: 93, as
with few big spines near the blunt end and a few small spines near Pseudomyxilla), S. phyllomorpha Levi (1993: 51), S. pluridentata
the pointed end, 448-504 X 13-16 /-Lm; smaller echinating acan- (Lundbeck, 1905: 151, as Myxilla), and S. toporoki Koltun (1958:
thostyles, similarly spined, 224-266 X 8/-Lm. Microscleres are 57). Koltun considered Chondrocladia alascensis Lambe (1894:
polydentate isochelae (Fig. 12B), sometimes slightly anisochelate, 119) a member of Stelodoryx, but this conforms clearly to
with numerous short sharp teeth, in two size categories, 36--60 /-Lm Monanchora. However, his own specimen recorded as Stelodoryx
and 18-22/-Lm. This species could be regarded as a deviating alascensis appears to be a true Stelodoryx.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademicIPlenum Publishers, New York, 2002

Family Phellodermidae ram. nov.

Rob W.M. Van SoestI & Eduardo Hajdu2

1 Zoological Museum, University of Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, Netherlands. (soest@science.uva.nl)
2Museu Nacional, Departamento de Invertebrados, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, sin, 20940-040,
Rio de Janeiro, RJ, Brazil & Centro de Biologia Marinha, Universidade de Sao Paulo, Sao Sebastiao, SP, Brazil. (hajdu@acd.ufrj.br)

Phellodermidae fam. nov. (Demospongiae, Poecilosclerida) is newly erected in the suborder Myxillina to accomodate sponges with arcu-
ate chelae or derivates thereof lacking special ectosomal diactinal megascleres. The only megascleres present are styles, which may occur
in size categories but are of uniform shape in all parts of the sponge. There are only few sponges with such a combination of characters,
and these are assigned to two genera, Echinostylinos and Phellodenna, differing in the possession of normal or asymmetrically twisted
chelae in the former, and peculiar symmetrically modified chelae, called abyssochelae, in the latter. They mostly occur in deep and cold
waters. Phellodermidae are similar to Dendoricellidae and Chondropsidae in having only a single megasclere type in combination with
arcuate chelae. However in these families the megascleres are respectively oxeas and strongyles, which are probably tomote-derived indi-
cating loss of choanosomal megascleres. In contrast, Phellodermidae seem to lack tomotes and retained their choanosomal megascleres.
Keywords: Porifera; Demospongiae; Poecilosclerida; Myxillina; Phellodermidae fam. nov.; Echinostylinos; Phellodenna.

DEFINITION, DIAGNOSIS, SCOPE Remarks

Definition The family is newly erected to unite sponges with the combi-
nation of styles and arcuate isochelae. The family differs from
Myxillina with exclusively styles as megascleres and arcuate other Myxillina sponges that contain arcuate chelae, such as
or modified arcuate chelae, and if present, sigmas. Coelosphaeridae, Hymedesmiidae and Crellidae, in lacking differ-
entiated ectosomal and choanosomal megascleres. Sponges of the
Diagnosis family Dendoricellidae and Chondropsidae appear similar in like-
wise lacking differentiated ectosomal and choanosomal megascle-
Semiglobular, stalked or thinly branching sponges. Surface res in combination with arcuate chelae, but the megascleres
slightly hispid. Skeleton plumose or radiating, without special in those family are diactinal (respectively oxeas and strongyles).
ectosomal skeleton. Spicules exclusively smooth styles, no dif- The lack of megasclere differentiation in these families is not con-
ferentiation into ectosomal and choanosomal types, although sidered an indication of sister-group relationships, as the diactinal
a smaller size may occur at the surface. Microscleres arcuate megascleres are probably tomote-derived, whereas Phellodermidae
isochelae, which may be modified into unique forms by twisting or fam. nov. appears to lack these entirely, having retained
fusion of the alae, and sigmas, which may be absent. the choanosomal styles. Other poecilosclerid families, e.g.,
Desmacididae and certain Mycalina, may appear similar, but these
Scope have different chela-types, anchorate or palmate.

Two out of five nominal genera are considered valid,

Echinostylinos and Phellodenna, comprising about ten species
reported from cold deeper waters.

KEY TO GENERA

(1) Chelae include 'abyssochelae', with alae meeting or fusing in the frontal plane ............................................................ Phelloderma
Chelae normal or asymmetrically modified, no fusion of alae ..................................................................................... Echinostylinos

ECHINOSTYLINOS TOPSENT, 1927 Definition

Synonymy Phellodermidae with ectosomal megascleres conslstmg of

a smaller size of styles or subtylostyles; choanosomal skeleton consist
Echinostylinos Topsent, 1927b: 8. Camptisocale Topsent, of tracts of larger styles; microscleres arcuate isochelae and sigmas.
1927b: 8. Ysila de Laubenfels, 1936a: 126.
Diagnosis
Type species
Anastomosing branches, erect bushes or semiglobular masses.
Echinostylinos reticulatus Topsent, 1927b: 8 (by monotypy). Surface irregular or conulose. Skeleton of smaller ectosomal

621
622 Porifera· Demospongiae • Poecilosclerida • Myxillina • PheUodermidae

D F

E
Fig. 1. A-C, Echinostylinos reticulatus Topsent (1927b). A, habit, reproduced from Topsent's pI. II fig . 21 (scale 1 em). B, skeleton of branch, reproduced
from his pI. VII fig. 15 (scale see text). C, spicules, from his pI. VII fig. 16 (sizes see text) . D, Echinostylinos glomeris (Topsent, 1904b as Esperiopsis),
type of Camptisocaie Topsent, 1927b, drawing of chelae, from Topsent, 1904b: pI. XVII fig. 3(scale 10 !Lm). E-F, Echinostylinos schmidti (Arnesen, 1903
as Esperiopsis), type of Ysila de Laubenfels. E, photo of holotype, reproduced from Arnesen, 1903 pI. I fig. 4 (size unknown). F, drawing of spicules, from
Arnesen, 1903 pI. IV fig . 4 (sizes see text).

subtylostyles assuming a partly erect and partly tangential position Material examined. Holotype: MOM (not seen). Schizotypes:
and larger choanosomal styles forming an axially condensed mass MNHN DT1293 - TWo microscopic slides, labeled 'n.g. n.sp. st.
or a vague reticulation of tracts and single spicules, Microscleres 3144' , made from the holotype.
arcuate chelae and sigmas. Approximately eight species. Description (from Topsent, 1928c). Clathrate mass
(Fig. lA) of thin twisted anastomosed branches with hispid sur-
Previous reviews face. Branches ca. 1.5 mm in diameter, total mass of branches
about 10 cm long, 8 cm wide. Consistency tough, not easily broken.
Topsent, 1928c; Bergquist & Fromont, 1988; Van Soest, 1990. Colour white in alcohol. The branches contain an axis of aligned
thick styles (Fig. lB) occasionally interconnected to form a tight
Description of type species spongin-enforced reticulation, and echinated profusely with thick
styles at the periphery, forming thus an extra-axial skeleton which
Echinostylinos reticulatus Topsent, 1927b (Fig. lA-C). carries the surface skeleton of small subtylostyles. The echinating
Synonymy Esperiopsis polymorpha var. Topsent, 1892a: 91, styles protrude through the surface to form hispid conules. The
pI. VI fig. 10. Echinostylinos reticulatus Topsent, 1927b: 8; Topsent, ectosomal skeleton contains a loose arrangement of bundles of thin
1928c: 207, pI. IT fig. 21, pI. VIT figs 15-16; ? Bergquist & subtylostyles and solitary subtylostyles. Spicules (Fig. lC), subty-
Fromont, 1988: 45, pI. 17 figs A-D. lostyles, smooth, straight, largely isodiametric, with a short point,
Porifera· Demospongiae • Poecilosclerida • Myxillina • Phellodermidae 623

245-280 X 3-4 fLm; styles robust, smooth, somewhat curved, architecture, would have to be considered plumose. It seems evi-
500-560 X 26--30 fLm. Microscleres, arcuate chelae, with thick, dent that de Laubenfels misinterpreted the skeleton of E. schmidti
curved shaft and short teeth, 27-49 fLm; sigmas of normal shape, and thus the genus Ysila was erected for nought.
15-22 fLm, possibly in two size/form categories. Distribution and
ecology. Azores, ? New Zealand, deep water, 900 m (?55-110 m).
Remarks. The New Zealand material shows similar spicule PHELLODERMA RIDLEY & DENDY, 1886
sizes and for that reason was assigned to E. reticulatus. However,
the shape is more lamellate or lobately ramose, different from the Synonymy
thin branches in the North Atlantic specimens. In view of the large
geographic separation, it is likely that both are different at the Phelloderma Ridley & Dendy, 1886: 347. Abyssocladia Levi,
species level. Echinostylinos has been associated with Monanchora 1964a: 78.
and Crambe of the family Crambeidae (e.g., by Van Soest, 1990),
on account of shared smaller and larger (subtylo-)styles and seem- Type species
ingly similar microsclere complement. However, the chelae of
Echinostylinos are clearly arcuate and the sigmas are true sigmas Phelloderma radiatum Ridley & Dendy, 1886: 347 (by
and not the reduced chelae that pass for sigmas in several monotypy).
Crambeidae. Further species are E. gorgonopsis Levi (1993: 49),
E. hirsutus Koltun (1970: 202), E. mycaloides Koltun (1970: 200), Diagnosis
E. shimushirensis Koltun (1970: 200), and E. stylophora (Levi &
Levi, 1983b: 960, as Lissodendoryx). Echinostylinos unguiferus de Phellodermidae with large styles in a single size category, and
Laubenfels (1953a: 528, fig. 6) is a junior synonym of Monanchora abyssochelae (stout isochelae with shaft markedly curved on
arbuscula (Duchassaing & Michelotti, 1864). profile view and frontal teeth touching or fused together).
The genus Camptisocale Topsent (1927b: 8) was erected for
the type species (by original designation) Esperiopsis glomeris Description of type species
Topsent, 1904b: 213, pI. XVII fig. 3. A MNHN slide (DT. 1000)
labeled "108 PA 1897" of the MOM type was examined. Phelloderma radiatum Ridley & Dendy, 1886 (Fig. 2A).
The chelae are definitely arcuate rather than modified palmates as Synonymy. Phelloderma radiatum Ridley & Dendy, 1886:
Topsent (1904b) maintained. The habit consisted of a damaged thin 347; Ridley & Dendy, 1887: 113, pI. XXIII fig. 8.
branch, with a main skeleton of radially arranged larger styles,
1400-1500 X 17-20 fLm; at the ectosome there is a partially tan-
gentially arranged layer of polytylote smaller subtylostyles,
900-1000 X 7-8 fLm. Peculiarly shaped arcuate isochelae
A B
(Fig. 1D) up to 60 fLm in length are abundant below the surface.
In the combination of stylote ectosomal spicules, smooth
choanosomal styles and arcuate isochelae, this species appears to
be close to Echinostylinos, and only the peculiar shape of the
chelae is a clear difference. This is judged to be of specific value
only, and accordingly Camptisocale is considered a synonym.
Camptisocale and Echinostylinos were erected on the same page,
but priority is given here to the name Echinostylinos as this has
been used much more frequently than Camptisocale.
The genus Ysila de Laubenfels (1936a: 126) was erected for
the type species (by original designation) Esperiopsis schmidti
Arnesen, 1903: 9. This is a semiglobular sponge (Fig. 1D, size not
known) with a confused-plumose skeleton of styles and large num-
bers of sigmata and arcuate isochelae (Fig. IF). It has not been
redescribed since its introduction and remains ill-known.

(J
Lundbeck (1905) noted its odd combination of spicules, but
retained the species in Esperiopsis (family Mycalidae). Arnesen
(1903) stated that the sigmas are vey large - and illustrates them
like that - but quotes a size of 20 fLm, whereas the chelae are said
to be 40 fLm, but are illustrated at less than a quarter the size of the
sigmas. The legend to the figures states that all spicules are illus-
trated at the same scale. Apart from this inconsistency, if the
description by Arnesen will prove to be otherwise correct, the
species is better assigned to Echinostylinos and it is proposed here
to consider Ysila a junior synonym of this genus. De Laubenfels
also noted the resemblance to Echinostylinos, but nevertheless
Fig. 2. A, Phelloderma radiatum Ridley & Dendy (1886), drawing
erected Ysila because Echinostylinos would have a plumose skele- of spicules reproduced from their pI. XXIII fig. 8 (sizes see text).
ton lacking in Esperiopsis schmidti. However, the skeletal structure B, Phelloderma bruuni (Levi, 1964a as Abyssocladia), drawing of spicules
described and figured by Arnesen, if it does present a recognizable reproduced from his fig. 30 (sizes see text).
624 Porifera· Demospongiae • Poecilosclerida • Myxillina • Phellodermidae

Material examined. Holotype: BMNH 1887:5:2:167 (wet)- shape of their isochelae, termed here 'abyssochelae'. Levi (l964a)
Rio de la Plata (fragment in 2MB, fide Hooper & Wiedenmayer, used the name thaumatoche1ae in referring to the isoche1ae of
1994:169). Abyssocladia, but it is suggested here that its usage should be
Description (adapted from Ridley & Dendy, 1887: 113). restricted to chelae similar to those of Mycale thaumatochela
Sponge subglobular, with a concave base of attachment, about 13 mm Lundbeck, 1909, viz. palmate anisochelae with caliciform feet encir-
in diameter. A cork-like cortex (ca. 240 fLm thick) is found underneath cling the frontal ala of the head. The type of A. bruuni (presumably
the easily peeled off dermal membrane. A few dubious scattered in ZMUC, a slide of it MNHN DC11404 was examined) from the
oscula on small papillae. Skeleton of radiating bundles of megasc1e- Kermadec Trench is a deep sea (>5000 m) pedunculate sponge, with
res with points directed outwards and embedded in the dense cortex. stalk of 10 X 1 mm, ending in an umbrella-like main body of 13 mm
Large amount of sand scattered throughout. Megasc1eres styles, at the widest. It has fusiform, slightly polytylote styles with a smaller
straight, smooth, verging upon tylostyles; fairly gradually and sharply head, 1300--1650 X 25-28 fLm, abyssochelae of 70-75 fLm, and
pointed, often subpolytylote, 650 X 12.6jl.m. Microscleres stout relatively thick sigmas of 30 fLm. As both Phellodenna and
isochelae with shafts markedly bent on profile view and frontal teeth Abyssocladia are known from a single species, it is difficult to objec-
which may fuse together, ca. 44 fLm high. Distribution and ecology. tively evaluate the significance of their apomorphic characters (pres-
Rio de la Plata, Argentina, 1080m, bottom green sand. ence of a 'cortex' in Phelloderma, and of centrally thickened sigmas
Remarks. The genus Abyssocladia Levi, 1964a, with type in Abyssocladia). It is proposed here to stress the similarities (virtu-
species A. bruuni Levi, 1964a: 78 (by monotypy) is considered ajun- ally identical abyssochelae and polytylote styles) and accordingly
ior synonym of Phellodenna, on account of the similar and peculiar Abyssocladia is considered a synonym of Phelloderma.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademicIPlenum Publishers, New York, 2002

Family Tedaniidae Ridley & Dendy, 1886

Rob W.M. Van Soest

Zoological Museum, University of Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, Netherlands. (soest@science.uva.nl)

Tedaniidae Ridley & Dendy (Demospongiae, Poecilosclerida) is characterized by the possession of onychaetes, thin finely acanthose
monactinal microscleres. Three genera are recognized based on the shape and differentiation of megascleres. One genus, Tedania, is
subdivided in three subgenera, based on the morphology of ectosomal tomotes and size of choanosomal megascleres. The use of the
nominal genus Tedanione is abandoned.
Keywords: Porifera; Demospongiae; Poecilosclerida; Myxillina; Tedaniidae; Hemitedania; Strongylamma; Tedania (Tedania); Tedania
(Tedaniopsis); Tedania (Trachytedania).
- - - - - - - _. ---_._.- - - - - - - - - - - - - - - - - -

DEFINITION, DIAGNOSIS, SCOPE (1928c) recognized a subfamily Tedaniinae (with content closely
similar to what is used here) within a wider concept of Myxillidae.
Synonymy De Laubenfels (1936a) retained separate families Tedaniidae and
Myxillidae, but compared with Topsent's classification mixed their
Tedaniina Ridley & Dendy, 1886: 335. Tedaniae Burton, contents on the basis of smooth vs. spined choanosomal megascle-
1932b: 345. res, thus including genera like Lissodendoryx and Acarnus within
Tedaniidae. Levi (1973), Wiedenmayer (1977b) and Van Soest
Definition (1984b) did not distinguish between subfamilies and employed a
broad assemblage named variously Myxillidae (including both
Myxillina with onychaetes, without chelae or sigmas. Tedania and Myxilla) or Tedaniidae (ibidem). Finally, Bergquist &
Fromont (1988) restricted the use of Tedaniidae to sponges pos-
Diagnosis sessing onychaetes, and Desqueyroux-Faundez & Van Soest (1996)
followed this. Tedaniids are soft slimy sponges with irregular
Encrusting, massive or digitate poecilosclerid sponges. shape. They are found in many different habitats, predominantly
Choanosomal skeleton predominantly plumo-reticulate or favouring sedimented environments: e.g., mangroves and stones
plumose, composed of tracts of smooth or spined monactinal among sand and mud, from shallow to deep waters. Reproduction
megascleres, or smooth diactinal megascleres, enclosed within is viviparous. Mature larvae have a bare posterior pole and are
light or moderate spongin fibres, or with no visible fibres and 300-600 11m diameter (Wapstra & Van Soest, 1987).
spicules merely cemented together at their nodes. Occasionally
fibres contain sand particles partly replacing spicules. Ectosomal Taxonomic remarks
spicules (tomotes) diactinal, tylotes or strongyles, with smooth or
spined bases, lying tangential, paratangential or erect on the sur- Tedaniidae is well delimited by the possession of unique ony-
face, although usually not in bundles. Microscleres onychaetes; chaete microscleres. Their position within the Poecilosclerida, and
chelae or sigmas absent. its suborder Myxillina, is less well established due to reduced spic-
ulation. Despite the lack of the major poecilosclerid synapomorphy
Scope (chelae), the family can nevertheless be assigned to this order on
the basis of its possession of tomotes, a feature shared with
Nine nominal genera. Genera and subgenera considered two Poecilosclerida suborders. The lack of tridentate chelae and
valid are Hemitedania, Strongylamma, Tedania (Tedania), Tedania sigmas is shared with Microcionina (which have palmate chelae),
(Tedaniopsis), Tedania (Trachytedania). whereas the lack of toxas is shared with Myxillina. Diversity
of tomote shapes and apical ornamentations are more often
History and biology found in Myxillina families than in Microcionina families.
Similarly, the loose plumoreticulate structure of most tedaniids is
Ridley & Dendy (1887) erected the family (as a subfamily) for more often observed in Myxillina. For these (perhaps) inconclusive
only two genera, viz., Tedania and Trachytedania. Tedaniinae was reasons Tedaniidae is assigned to Myxillina rather than to
considered related to and united with sigma-bearing Haplosclerida Microcionina.
(their subfamilies Gelliinae and Phloeodictyiinae) and lipochelate
Poecilosclerida (Desmacellinae and Hamacanthinae) within a larger Previous reviews
family Heterorhaphidae. Myxillinae was classed further away in
the chela-bearing family Desmacidonidae. Since that time various Bergquist & Fromont (1988), Desqueyroux-Faundez & Van
authors assigned a wide variety of genera to tedaniids. Topsent Soest (1996).

625
626 Porifera· Demospongiae • PoeciloscIerida • Myxillina • Tedaniidae

KEY TO GENERA

(1) Megascleres are oxeas or strongyles of one type only ......................................................................................................................... 2

Megascleres are differentiated in ectosomal (diactinal) and choanosomal spicules (usually monactinal, but in any case differently
shaped from those of the ectosome) ............ ......... ........................ ................ ................ ........................... .......... ...... .......... ....... Tedania
(2) Megascleres oxeas ............................................................................................................................................................ Hemitedania
Megascleres thin strongyles ........................................................................................................................................... Strongyiamma

HEMITEDANIA HALLMANN, 1914 (1996) erroneously considered the type species to be one of the
'sand sponges' (i.e., sponges which have replaced part or whole of
Synonymy their skeletal spicule tracts by sand tracts), but this was based on
examination of one of Dendy's subsequently named specimens,
Hemitedania Hallmann, 1914: 431. not on Carter's original specimens. The type specimens do not con-
tain any appreciable build-up of sand tracts and it is dubious
Type species whether such specimens belong to H. anonyma at all.
Desqueyroux-Faundez & Van Soest (1996) suggested that the
Amorphina anonyma Carter, 1886b: 49 (by original genus Tedaniopsamma Burton, 1934a may be synonymous with
designation). Hemitedania. However, after examination of the type of
Tedaniopsamma (Hircinia) Jlabellopalmata (Carter, 1885b) it is
Definition clear that this lacks the onychaetes and consequently is referred to
Chondropsidae.
Tedaniidae with exclusively oxea megascleres.
Previous reviews
Diagnosis
Hallmann (1914), Hooper & Wiedenmayer (1994),
Massive; ectosomal skeleton reticulate but without any spe- Desqueyroux-Faundez & Van Soest (1996).
cialised spicules; choanosomal skeleton composed of plumo-
reticulate fibres cored by smooth choanosomal oxeas, occasionally Description of type species
with an arenaceous skeleton; oxeas sharply pointed, of a single cat-
egory; microscleres one or more size categories of onychaetes. Hemitedania anonyma (Carter, 1886b) (Fig. lA-E).
Distribution. New South Wales and Victoria, two species. Synonymy. Amorphina anonyma Carter, 1886b: 49;
Rhaphisia anonyma; Dendy, 1895: 256; Hemitedania anonyma;
Remarks Hallmann, 1914: 431, pI. 18 fig. 4, pI. 19 figs. 1-5, pI. 24 figs. 3-5,
text-fig. 20; Hooper & Wiedenmayer, 1994: 414. Reniera pandaea
Hallmann (1914) based his description of Hemitedania Lendenfeld, 1888: 79; Rhaphisia pandaea; Whitelegge, 1902: 281.
anonyma on 'some twenty specimens' from Port Jackson and Halichondria rubra Lendenfeld, 1888: 81. Halichondria rubra dig-
neighbouring localities, and furthennore on a specimen from Port itata Lendenfeld, 1888: 81; Rhaphisia rubra; Whitelegge, 1901: 77.
Phillip Bay and a slide preparation presented to the Australian Material examined. Lectotype (designated herein): BMNH
Museum by Dendy. It is likely that Hallmann did not examine the 1886.12.15.119 - preserved in alcohol, Port Phillip Heads,
original Carter specimens. The BMNH collection holds one alco- Bracebridge Wilson collection (slide of the holotype labeled:
hol specimen (BMNH 1886.12.15.119) and two dry specimens Dendy, colI. 1954.2.12.291 (Fig. 1A». Paralectotype: BMNH
(BMNH 1886.12.15.390,466), all labeled 'Amorphina anonyma, 1886.12.15.390 - dry specimen, same data as the lectotype
Bracebridge Wilson Collection, Port Phillip Heads'. The speci- (Fig. 1B).
mens 390 and 466 appeared to have switched labels (inspected Description. Massive, lobate (Fig. lA-B) to thickly ramose,
March 1999) and moreover are not conspecific, one (466) appeared laterally compressed. Size up to 20 X 10 X 10 cm. Surface smooth.
to be a Xestospongia, lacking the onychaetes. The other (390) con- Numerous oscules on the upper parts and at the apex of lobes and
forms to the description of Carter. It seemed the best course to branches. Colour possibly orange (based on remarks made by
select the alcohol material as the type, although Carter's descrip- Dendy); in alcohol it is yellow-brown; in the dry state it is beige.
tion is probably based more on the dry material. The BMNH col- Skeleton (Fig. IE) of the ectosome formed by the brushed endings
lection also holds several further specimens and slides labeled of the choanosomal tracts, with spicules fanning out and partly
"Rhaphisia anonyma" from the same area described by Dendy becoming tangential. Choanosomal skeleton plumoreticulate, with
(1895). These are not type specimens and have not all been re- emphasis on main ascending tracts, which are 30-70 /-Lm in diame-
examined for the present study. Van Soest (1984b) included in this ter, made up of 4-20 spicules in cross section. Interconnecting
genus species with thin strongyles answering to Strongylamma tracts irregularly branching off from the main tracts at right angles,
based on the shared possession of eccentric tyles on the onychaetes thinner and with less coring spicules. Spongin binding the spicules.
(emphasized by Hallmann in his definition of the genus Between the tracts there are many loose single megascleres and
Hemitedania). This feature is of much wider occurrence in the family microscleres. Spicules (Fig. 1B-E), oxeas, 242-(257.3)-318 X
and in view of the asymmetric nature of onychaetes is thought to be 4-8/-Lm (Carter gives a size of 317 X 1O.5/-Lm); onychaetes in two
without value at the genus level. Desqueyroux-Faundez & Van Soest size categories, 155-188 and 60-65/-Lm. Distribution and ecology.
Porifera· Demospongiae • Poecilosclerida • Myxillina • Tedaniidae 627

..1 H -----. ...

Fig. 1. A-E, Hemitedania anonyma (Carter, 1886b as Amorphina), type material. A, lectotype (wet, scale 1 cm), B, paralectotype (dry, scale 1 cm).
C, SEM photo of onychaete (right, scale 10 11m) and detail (scale 111m). D, SEM photo of oxea (right, scale 10 11m) and detail (scale 111m). E, drawing of
skeleton and spicules made from a slide of the holotype. F-H, Strongylamma carteri (Dendy, 1895 as Chondropsis). F, SEM photo of strongyle. G, SEM
photo of onychaete (scales 111m), both made from topotypical specimen. H, drawing of skeleton and spicules made from a slide of the type.

SE Australia, Port Phillip Heads, Port Jackson, Port Stephens, Type species
Green Point, Shark Reef, Bass Strait, 7-57 m.
Remarks. In addition to the type species a further SE Chondropsis carteri Dendy, 1895: 256 (by original
Australian species may exist, H. ramosa (Whitelegge, 1906: 463, designation).
as Rhaphisia). Hallmann (1917c), who assigned Whitelegge's
material provisionally to Hemitedania, thought it would need a Definition
genus of its own, but judging from Hallmann's redescription the
spicule characters (oxeas and onychaetes) perfectly fit with Tedaniidae with smooth strongyles as the only megascleres
Hemitedania. For type specimen information on the various forming a loosely plumose skeleton.
Lendenfeld synonyms see Hooper & Wiedenmayer (1994: 414).
Diagnosis

STRONGYLAMMA HALLMANN, 1917 Encrusting, cryptic, or more massive growth forms. Skeleton
of smooth strongyles as the only megascleres forming a loosely
Synonymy plumose skeleton. In one species the spicule bundles are reinforced
by sand grains. No ectosomal specialization. Little spongin.
Strongylamma Hallmann, 19l7c: 643. Tedanione sensu Microscleres onychaetes in single or 2 size classes, with or without
Dendy, 1922b: 100 (not Wilson, 1894: 338). Hemitedania sensu distinct tyles. Distribution. Victoria, Tropical W Atlantic, W Africa,
Van Soest, 1984b: 52 (not Hallmann, 1914: 431). and Indian Ocean. Three species.
628 Porifera· Demospongiae • Poecilosclerida • Myxillina • Tedaniidae

Remarks two species from the general area appear likely candidates as junior
synonyms, H. poa de Laubenfels (1947) and H. corrugata Dfaz et
Emphasis is placed on the lack of differentiation of the at. (1993). Until Wilson's material has been traced, it is not possible
megascleres and feebly-developed spicule bundles. Hemitedania to draw a firm conclusion about this synonymy. West African waters
also has only a single category of megasclere, but these are oxeas. harbour specimens of Strongylamma baki indistinguishable from
Moreover, the type species and a second species assigned to those of the Caribbean (cf. Van Soest, 1993a, as Tedanione), so it is
Hemitedania are elaborate sponges with reticulate skeletons, assumed to be the same species. In addition to S. carteri and S. baki
unlike the thin crusts of Strongylamma. Tedanione sensu Dendy, a further species belonging to this genus is the Western Indian
1922b is a clear synonym, but the type of Tedanione (i.e., T. foetida Ocean S. wilsoni Dendy (1922b: 101, as Tedanione). Tedanione
Wilson, 1894) is a Halichondria (cf. below). connectens Brfilndsted, 1924 (see also Bergquist & Fromont, 1988)
and T. obscurata de Laubenfels, 1930 do not conform to the present
Previous review definition of the genus Strongylamma and are referred to Tedania
(Tedania).
Desqueyroux-Faundez & Van Soest (1996 as Tedanione).

Description of type species TEDANIA GRAY, 1867

Strongylamma carteri (Dendy, 1895) (Fig. IF-H). Synonymy

Synonymy. Chondropsis carteri Dendy, 1895: 256;
Strongylamma carteri; Hallmann, 1917c: 643. Tedania Gray, 1867a: 520. Trachytedania Ridley, 1881: 122.
Material examined. Holotype (not seen): NMVG2322. Tedaniopsis Dendy, 1924: 367. Paratedania Burton, 1929a: 411.
Fragment of holotype: BMNH 1902.10.18.166 - slide made from Xytopsihis de Laubenfels, 1936a: 54. Oxytedania Sara, 1978: 61.
holotype, Port Phillip Heads, South Australia. Other material.
BMNH 1933.8.1.1 - specimen and slide, from Port Phillip Heads. Type species
Description. Small, massive, rounded, constricted below
and somewhat flattened above. Oscules small and grouped on the Reniera digitata Schmidt, 1862: 73 (by original desig-
upper part. Surface smooth but uneven. Consistency compressible. nation) (=junior synonym of Halichondria anhelans Lieberkiihn,
Colour (alcohol) pale yellowish grey. Skeleton (Fig. IH) of the 1859: 521).
ectosome with tangential megascleres and microscleres strewn in
confusion. Choanosomal skeleton of wispy loose tracts of Definition
strongyles containing also some sand grains. The tracts have a
width of ca. 200 /-Lm and are arranged at distances of 100-400 /-Lm. Tedaniidae with differentiated ectosomal and choanosomal
Spicules, strongyles, straight and thin (Fig. IF-H): megascleres.
165-(253.5)-303 X 2.5-(2.9)-3.5 /-Lm. Microscleres, two size cate-
gories of onychaetes (Fig. 1G-H) , tyles not visibly developed: Diagnosis
84-90.8-99 and 42-47.6-54 /-Lm. Distribution. Victoria.
Remarks. Dendy's description does not mention onychaetes, Massive to lobate; ectosomal skeleton composed of tylotes or
but merely 'long hair-like rhaphides'. but Hallmann (1917c: 643) tomotes with microspined bases forming tangential or paratangential
discovered that these were onychaetes when he re-examined the surface tracts; choanosomal skeleton composed of styles with
type of Dendy. Under SEM these are clear and unequivocal. This smooth or microspined bases, producing reticulate, plumo-reticulate,
genus is known under the aliases Tedanione and Hemitedania. Type plumose or dendritic architecture; microscleres onychaetes.
material of Hemitedania baki (ZMA POR. 4766-4768; Curac;ao, Distribution cosmopolitan. Approximately 65 species.
encrusting on corals, 20--35 m), which answers in all aspects to
Strongylamma, was compared to Wilson's (1894) description of Previous reviews
Tedanione foetida from the Bahamas; no original Wilson material
has been unearthed so far. Since Tedanione sensu Dendy, 1922b is Bergquist & Fromont (1988), Desqueyroux-Faundez & Van
clearly congeneric with H. baki it seemed a fair assumption that the Soest (1996).
latter could be a junior synonym of Wilson's species as they are both
from the same broad locality. However, not only are Tedanione Remarks
foetida and H. baki clearly different in morphology and spicule size,
it has also become clear that Tedanione is an unlikely tedaniid. The Three subgenera have been recognized by Desqueyroux-
description and figures of Wilson allow the conclusion that the only Faundez & Van Soest (1996): Tedania s.s. with basically the
spicules likely to be proper to the sponge are oxeas 350 /-Lm long and spicule complement and arrangement of T. anhelans, including
'microscleres (oxeas) of varying length' , and also that the larvae are microspined ectosomal tylotes and small choanosomal styles
uniformly ciliated. Tylotes were also mentioned, but only 'very few' <350/-Lm; Trachytedania Ridley, 1881: 122 (type species
were observed. The description fits perfectly with sponges of the T. spinata Ridley, 1881) with mucronate ectosomal tomotes, and
genus Halichondria and not with tedaniids on two important main spicules in the same size range as Tedania s.s.; and
counts: Halichondria species often have a smaller and a larger cate- Tedaniopsis Dendy, 1924: 366 (type species T. turbinata
gory of oxeas whereas tedaniids have differentiated spicules includ- Dendy, 1924) with variably shaped ectosomal tomotes, and
ing onychaetes, and Halichondria larvae are typically ciliated styles (occasionally stylote or oxeote modifications) larger than
all-over, whereas tedaniid larvae have a bare posterior pole. At least 350/-Lm.
Porifera' Demospongiae • Poecilosclerida • Myxillina • Tedaniidae 629

Key to subgenera of Tedania

(1) Ectosomal megasc1ere microspined tylotes ............................................................................................................. Tedania (Tedania)

Ectosomal megasc1eres mucronate tornotes or smooth tylotes ............................................................................................................ 2
(2) Choanosomal megasc1eres >350 /-Lm ................................................................................................................ Tedania (Tedaniopsis)
Choanosomal megasc1eres <350 ftm ............................................................................................................ Tedania (Trachytedania)

SUBGENUS TEDANIA GRAY, 1867 reticulation of styles, spongin indistinct, with many loose spicules.
Spicules, ectosomal tylotes (Fig. 2A) with finely spined apices,
Synonymy 140--302 X 2-7 ftm; styles (Fig. 2B) smooth, slightly curved:
173-280 X 5-12 ftm; onychaetes (Fig. 2C), finely rugose, without
Tedania Gray, 1867a: 520. Xytopsihis de Laubenfels, 1936a: 54. a distinctly swollen asymmetrical tyle, in a single but variable size
category: 40--220 X 0.5-2 ftm (Lieberktihn pictured 2 styles of
Type species 240--270 X 7-12 ftm and an onychaete of 180 ftm). Distribution
and ecology. Western and Eastern Mediterranean, NE Atlantic
Reniera digitata Schmidt, 1862: 73 (by original designa- from Bretagne down to Senegal, Azores; shallow-water, on rocks
tion) (=junior synonym of Halichondria anhelans Lieberktihn, and stones, littoral to 100 m.
1859: 521). Remarks. Lieberktihn (1859) described and figured only
styles and onychaetes (Fig. 2F), but Schmidt (1862: 72) found tylotes
Diagnosis in Lieberktihn's material. It is remarkable that on the next page, when
describing Reniera digitata, Schmidt did not refer at all to his descrip-
Smooth, relatively small styles, occasionally strongylote tion on the previous page of Myxilla anhelans for which he described
styles as structural megascleres and microspined tylotes as ectoso- exactly the same spicules. The species has been reported from allover
mal megascleres. Distribution: predominantly in tropical and the world, and it is clear that closely similar populations exist over the
warm-temperate waters of all three ocean systems. whole Tethyan region. Recent insights in biogeography and ecology
favour the recognition of a series of allopatric (sibling) species (e.g.,
Description of type species the Caribbean representatives are known under the name T. ignis
(Duchassaing & Michelotti, 1864 as Thalysias), cf. Fig. 2G), rather
Tedania (Tedania) anhelans (Lieberktihn, 1859) (Fig. 2A-F). than a single cosmopolitan species (cf. Van Soest, 1987b).
Synonymy. ? Spongia anhelans Vio in Olivi, 1792: ix-xxxi The genus Xytopsihis de Laubenfels, 1936a was erected for
(fide Schmidt, 1862); Halichondria anhelans Lieberktihn, 1859: 521, type species Halichondria aspera Bowerbank, 1875b: 287) from
pI. XI fig. 6; Myxilla anhelans; Schmidt, 1862: 72; Tedania anhelans: the Straits of Malacca (by original designation). Bowerbank's
Levi, 1952: 48; Levi, 1959: 130, fig. 21; Boury-Esnault, 1971b: 312; description is misleading in giving the spicule combination of
Sara, 1972: 80; Pulitzer-Finali, 1978: 57; Sol6rzano & Babio, 1979: tylotes, oxeas, styles, 2 size categories of chelae, sigmas. Not sur-
55, fig. 11; Pulitzer-Finali, 1983: 561; Van Soest, 1987b: 13, figs 1-4. prisingly, Hofman & Van Soest (1992: 87) misinterpreted the nature
Reniera digitata Schmidt, 1862: 73, pI. VII fig. 11; Desqueyroux- of the type specimen. These authors referred it to the genus
Faundez & Stone, 1992: 16, figs 175-177; Tedania digitata; Gray, Lissodendoryx (as L. aspera), and this was made the senior syn-
1867a: 520; Topsent, 1920a: 16; Babic, 1922: 245; Topsent, 1932b: 4; onym of a number of species variously known as Crella schmidti
Topsent, 1936: 23. Reniera nigrescens Schmidt, 1862: 74; Ridley (1884a: 432), Zetekispongia zonea de Laubenfels (1936b:
Desqueyroux-Faundez & Stone, 1992: 17, figs. 195-196. Reniera 446) and Damiriana hawaiiana de Laubenfels (1950b: 14). The
muggiana Schmidt, 1868: 28; Desqueyroux-FaUndez & Stone, 1992: type specimen and two microscopic slides are kept in the BMNH
17, fig. 193. Tedania chevreuxi Topsent, 1891b: 13, pI. II figs 1-2. London, unregistered. It is an irregularly lobate sponge of 8 X 6 X
Tedania suctoria, sensu Arndt, 1941: 12 (not Schmidt, 1875). 3 cm, with an irregularly pitted surface. Attached at its undersurface
Material examined. Lectotype (designation herein): LMJG there is a smaller cushion-shaped sponge, size 3 X 2 X 0.5 cm, with
15344 - labeled 'Reniera digitata, Venedig' (Venice), alcohol a smooth surface. The larger sponge is a Tedania, probably match-
preserved specimen (cf. Desqueyroux-Faundez & Stone, 1992 ing the description of T. dirhaphis Hentschel (1912) which would
pI. XXIX fig. 175). Paralectotype: LMJG 15335 - same locality then become a junior synonym of T. aspera. It has microspined
(cf. Desqueyroux-Faundez & Stone, 1992 pI. XXIX fig. 176). tylotes of 224--(247.4)-276 X 4--(4.3)-5 ftm, thick smooth styles
Other material. ZMA POR. 4863 - Chateau du Taureau, Baie de of 253-(279.1)-302 X 11-(13.5)-15 ftm, and two size categories
Modaix, Bretagne, France. of onychaetes, viz., 221-(303.6)-352 X 2.5-5.5 ftm and 50--144 X
Description. Massive-ramose (Fig. 2D-E), with irregular <0.5 ftm (cf. Fig. 2H). No chelae or sigmas were found in this spec-
branches and flattened fistule-like projections issuing from a thick imen. However, the smaller sponge is a Lissodendoryx, identified as
base. Size up to 10 cm in lateral expansion and elevation. Surface L. tematensis (Thiele, 1903a) using Hofman & Van Soesfs key to
irregular, corrugated. Oscules of ca. 5 mm diameter on the main the Lissodendoryx of the Indo-Malayan Archipelago (it has tylotes
body as well as on the branches and at the end tubular elevations. of ca. 180 X 4 ftm, styles of ca. 180--200 X 6 ftm, two categories of
Consistency soft, compressible, easily damaged. Colour variably arcuate isochelae, ca. 25 and 15 ftm, and two categories of sigmas,
brownish, orange-green, reddish brown and 'blue'. Skeleton of the 35 and 18 ftm). Since the Tedania makes up the bulk of the material
ectosome consists of bundles of tylotes, partly tangential, partly of Halichondria aspera and was obviously the material intended for
perpendicular or in surface brushes; individual tylotes are strewn it by Bowerbank, we are forced to accept that Tedania aspera is the
throughout the peripheral regions. Choanosomal multispicular type of Xytopsihis and this genus is an obvious junior synonym of
630 Porifera· Demospongiae • Poecilosclerida • MyxiUina • Tedaniidae

H I I

Fig. 2. A-F, Tedania (Tedania) anhelans (Lieberkiihn, 1859 as Halichondria). A-C, spicules of a type specimen of the type of Tedania, i.e. , Reniera digitata
Sclunidt (1862), considered a junior synonym of T. anhelans. A, microspined tylote (left, scale 10 /Lm) with detail, (scale l/Lm). B, style (left, scale 10 /Lm) with
detail (scale I fLm) . C, onychaetes (left, scale 10 /Lm) with detail (scale l/Lm). D, Schmidt's drawing of Reniera digitata. E, photo of the lectotype of Reniera
digitata from Desqueyroux-Faundez & Stone, 1992 (scale 1 cm). F, Lieberkiihn's (1869) drawing of the spicules of Halichondria anhelans (sizes see text).
G, Tedania (Tedania) ignis (Duchassaing & Michelotti, 1864 as Thalysias), drawing of the skeleton and spicules (reproduced from Van Soest, I 984b). H, Tedania
(Tedania) aspera (Bowerbank, 1875b as Halichondria), type of Xytopsihis de Laubenfels (l936a), drawing of skeleton and spicules made from the type.

Tedania (Tedania). Hofman & Van Soest's (1992) use of the species Diagnosis
name for Lissodendoryx material with oxeas as the structural
megascleres is thus incorrect. The name of that species now Relatively long (>350 j.l.m), thick, smooth styles, occasion-
becomes Lissodendoryx (Waldoschmittia) schmidti (Ridley, 1884a). ally modified to anisostrongyles, anisotylotes or anisoxeas as struc-
tural megascleres, ectosomal megascleres are mucronate or
tylostrongylote tornotes occasionally with one or more vestigial
spines. Distribution. Predominantly known from the southern
SUBGENUS TEDANIOPSIS DENDY, 1924 oceans.

Synonymy Description of type species

Tedaniopsis Dendy, 1924: 367. Paratedania Burton, 1929a: 441. Tedania (Tedaniopsis) turbinata (Dendy, 1924) (Fig. 3A-D).
Synonymy. Tedaniopsis turbinata Dendy, 1924: 367, pI. XI
Type species figs 2-3, pI. XIV figs 31-35.
Material examined. Holotype: BMNH 1923.10.1.148 -
Tedaniopsis turbinata Dendy, 1924: 367 (by monotypy). Three King Islands, New Zealand.
Porifera· Demospongiae • Poecilosclerida • Myxillina • Tedaniidae 631

ftl
o
a
-- I.. H
D i
Fig. 3. A-D, Tedania (Tedaniopsis) turbinata (Dendy, 1924). A--C, SEM photos of spicules from the holotype (scales 10 ILm; except Bright, llLm).
D, drawings of skeleton and spicules made from a slide of the holotype. E-H, Tedania (Trachytedania) spinata Ridley (1881). E-G, SEM photos of
spicules from the holotype (scales 10 ILm; except D left, E and Fright. llLm). H, drawings of skeleton and spicules made from a slide of the holotype.

Description. Large stipitate sponge fonning a thick-walled Remarks. Tedaniopsis was erected because of the replace-
shallow cup, up to 20cm high, 14cm wide. Surface smooth, ment of choanosomal styles by strongylotylotes. However, no sim-
grooved on the sides. Oscules scattered over inner surface. Colour ilar species with comparable spiculation has been described and
in alcohol pale yellowish greyish. Skeleton (Fig. 3D) of the surface thus the generic status is questionable. Nevertheless, the large sizes
consists of brushes of tomotes, forming a fringe around each of the of the spicules and the irregular skeletal architecture unite the pres-
oscules. Choanosomal skeleton an irregular reticulation of spicule ent species with several others, and accordingly Desqueyroux-
tracts. Spicules, ectosomal tylotomotes (Fig. 3B, D), inequiended, Fat1ndez & Van Soest (1996) proposed to recognize a distinct
mostly provided with two or three spines at one end: 340 X 6 f.1m; subgenus for them. Species assigned to this subgenus are: Tedania
choanosomal strongylotylotes (Fig. 3A, D), large and thick, with massa Ridley & Dendy, 1886 (with junior synonyms T. actiniformis
oval heads: 500 X 25 f.1m; onychaetes (Fig. 3C-D), finely spined, Ridley & Dendy, 1886 and Tedania tantula (Kirkpartick, 1907a, as
in two categories, the larger with a clear asymmetrical tyle: 600 X Oceanapia) (Antarctica, Magellan province); Tedania charcoti
4 f.1m and 180 X 2 f.1m. Distribution and ecology. New Zealand; Topsent, 1917 (Antarctica, Magellan province, SE Pacific);
dredged up from 180 m. Tedania infundibuliformis Ridley & Dendy, 1886 (SE Pacific);
632 Porifera· Demospongiae • Poecilosclerida • Myxillina • Tedaniidae

Tedania tenuicapitata Ridley, 1881 (Magellan province, SE the choanosome. Oscules, 2-3 mm diameter, irregularly scattered.
Pacific); Tedania oxeata Topsent, 1907 (Antarctica); Tedania Consistency soft, compressible. Colour of surface dark gray, inter-
vanhoeffeni Hentschel, 1914 (Antarctica); and Tedania phacellina nally brownish to grayish. Skeleton (Fig. 3H) of the ectosome
Topsent, 1928c (deep water North Atlantic). a dense palisade of tornotes reinforced by the terminal part of
The genus Paratedania Burton, 1929a, with type species styles and abundant onychaetes. Choanosomal skeleton irregular to
Oceanapia tantula Kirkpatrick, 1907a (by monotypy) (which is isotropic reticulation of spicule tracts. Spicules, ectosomal smooth
probably a junior synonym of Tedania massa Ridley & Dendy, mucronate tornotes (Fig. 3F) abruptly pointed: 120--280 X 3-6 fLm;
1886) was erected because of a densely packed layer of tangential choanosomal styles (Fig. 3E) with frequently a few spines on along
megascleres, as opposed to the bouquet-type arrangement of all the upper half: 160--280 X 3-10 fLm; onychaetes (Fig. 3G) straight,
other Tedaniidae. Oceanapia tantula (persistently named tarantula strongly spined, in two size categories: 90--230 X 2 fLm and
by Burton, 1929a, 1932b), of which the type specimen BMNH 30--100 X 0.5 fLm length. Distribution and ecology. SE Pacific,
1908.2.5.198 was examined, is a lobate to erect sponge with 39°-S0oS; SW Atlantic, SOo-S4°S, Falkland Islands; encrusting
smooth anisostrongyles as structural megascleres and tylotornotes shells, pebble and sand bottom, 10-69 m.
with an occasional mucron at one of the apices. The tangential Remarks. Desqueyroux-Faundez & Van Soest (1996: 62)
arrangement at the surface, however, was probably entirely reviewed this species extensively. Ridley (1881) emphasized the
induced by the periostracum of mollusc shells on which the speci- presence of spines on the heads of part of the styles when describ-
mens were found to be attached, and thus must be considered ing T. spinata. The styles are essentially smooth, but many have a
artifactual. Burton (1932b) withdrew this genus himself, and sug- few spines along the shaft. They are not homologous to echinating
gested that the type was a junior synonym of Tedania massa. acanthostyles. The variability and vestigial presence of the
Examination of type material of both revealed some differences in spination is paralleled by that of Lissodendoryx isodictyalis
the shape of tornote endings (hastate in T. massa), but in view of (Coelosphaeridae). Ridley considered the smooth styles as a sepa-
the variability of this character, conspecificity of both is certainly rate spicule type, claiming they were also morphologically distinct
possible. Both are also similar to Tedania turbinata, and accord- in having a tapering or mucronate head. Examination of the type as
ingly Paratedania is considered a junior synonym of the subgenus well as redescriptions of topotypical specimens (Topsent, 1929b;
Tedaniopsis. Desqueyroux-Faundez & Van Soest, 1996) did not confirm
Ridley's claims. The 'styles' with pointed or mucronate heads are
tornotes, which are in the same size range as the styles. The type
SUBGENUS TRACHYTEDANIA RIDLEY, 1881 specimens of T. murdochi Topsent, 1904a, T. corticata Sara, 1978
and T. laminariae Sara, 1978 were re-examined and could not be
Synonymy separated from T. spinata, on skeletal structure, spicule sizes and
form, so their conspecificity is probable. Further species assigned
Trachytedania Ridley, 1881: 122. Oxytedania Sara, 1978: 61. to the subgenus by Desqueyroux-Faundez & Van Soest (1996) are
Tedania mucosa Thiele, 1905 (with synonyms T. excavata Thiele,
Type species 1905, T. pectinicola Thiele, 1905 and T. juegiensis Thiele, 1905),
T. patagonica Ridley & Dendy, 1886, T. gurjanovae Koltun,
Trachytedania spinata Ridley, 1881: 122 (by monotypy). 1958, T. microrhaphidiophora Burton, 1935c, and T. toxicalis
de Laubenfels, 1930.
Diagnosis The genus Oxytedania Sara (1978), with type species
O. bifaria Sara, 1978: 61 (by monotypy), was erected for a species
Structural spicules are relatively small styles <350 fLm, with the spicule combination of typical Tedania in addition to
which may show a few spines; ectosomal megascleres are a reticulate skeleton of oxeas. The type specimen studied by
mucronate or oxeote tornotes. Distribution. Predominantly known Desqueyroux-Fa6ndez & Van Soest (1996) did not contain any ofthe
from the southern oceans, but there are also records from tedaniid spicules described by Sara. The skeletal architecture was
California and from the Arctic. typically that of the haplosclerid family Niphatidae with multispicu-
lar tracts of oxeas in a tight reticulation. It is assumed that the genus
Description of type species is based on a contamination of Tedania spinata spicules (including
styles with a few spines) in a niphatid specimen. Indications for this
Tedania (Trachytedania) spinata Ridley, 1881 (Fig. 3E-H). conclusion are not only Desqueyroux-Faundez & Van Soest's failure
Synonymy. Trachytedania spinata Ridley, 1881: 122, pI. X to find any Tedania-like spicules, but also the fact that Sara not only
fig. 10; Tedania spinata: Topsent, 1929b: 282, fig. 67; Tedania pictured the oxeas as structural megascleres, but also styles of quite
(Trachytedania) spinata: Desqueyroux-Faundez & Van Soest, dissimilar size and form as a second category of structural megascle-
1996: 62, figs 129-134. Tedania murdochi Topsent, 1904a: 63; res. TWo such different structural megasclere categories together in
Topsent, 1913b: 629, pI. V fig. 5. Tedania corticata Sara, 1978: 56, a single sponge are unlikely to occur in a tedaniid. Accordingly
figs 34-35. Tedania laminariae Sara, 1978: 54, figs 32-33. this genus is pronounced unrecognizable c.q. a junior synonym
Oxytedania bifaria Sara, 1978: 61, figs. 38-39. of Tedania (Trachytedania). Recent, Cristobo & Urgorri (2001)
Material examined. Holotype: BMNH 1879.12.27.9 - described a Trachytedania species with hymedesmioid skeleton (two
Portland Bay, Chile. Other material. See Desqueyroux-Faundez & size categories of fully spined acanthostyles with heads embedded in
Van Soest (1996: 62). a basal spongin plate), which is quite unlike Tedania (Trachytedania)
Description. Massive sponge, size 8-10 X 3-8 cm. Surface spinata in architecture and spiculation. This appears to belong to an
smooth, covered by a translucent membrane, which is darker than undescribed genus of Tedaniidae.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Suborder Mycalina Hajdu, Van Soest & Hooper, 1994

Rob W.M. Van SoestI & Eduardo Hajdu2

I Zoological Museum, University of Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, Netherlands. (soest@science.uva.nl)
2Museu Nacional, Departamento de Invertebrados, Universidade do Brasil, Quinta da Boa Vista, sIn, 20940-040, Rio de Janeiro,
RJ, Brazil & Centro de Biologia Marinha, Universidade de Sao Paulo, Sao Sebastiao, SP, Brazil. (hajdu@acd.ufij.br)

Suborder Mycalina Hajdu et al. (Demospongiae, Poecilosclerida) contains sponges characterized by possession of palmate chelae or
derivates thereof and a single megasclere type, usually smooth styles with a slight constriction subapically (dubbed 'mycalostyles').
Several families possess diancistras or derivates thereof and along with the mycalostyles these are interpreted as synapomorphies of the
suborder. In some families chelae are absent, presumed lost, and in rare cases megascleres are strongyles or oxeas. Chelae may be modi-
fied to become anisochelate (families Mycalidae and Cladorhizidae), anchorate (genus Chondrocladia), polydentate (subgenus Mycale
(Grapelia», or strongly modified (family Guitarridae). These chela types are presumed to be derived from ordinary palmate isochelae.
Microscleres that may be present in addition to chelae and diancistras are normal sigmas, toxas (which are occasionally spined), micro-
xeas (which may be rugose, spined or amphiaster-like), trichodragmas and commata. Skeletal structure is mostly plumose or plumoreti-
culate, occasionally isodictyal-anisotropic, with tangential ectosomal skeleton frequently developed, but occasionally absent. One family
contains coralline sponges with a limestone basal skeleton. Nine families are distinguished based on the presence of microscleres,
morphology and skeletal architecture, distributed worldwide and in all marine habitats.
Keywords: Porifera; Demospongiae; Poecilosclerida; Mycalina; Cladorhizidae; Desmacellidae; Esperiopsidae; Guitarridae;
Hamacanthidae; Isodictyidae; Merliidae; Mycalidae; Podospongiidae.

DEFINITION, DIAGNOSIS, SCOPE derivates occurring in several families across the suborder. With
suborder Microcionina they share the possession of palmate chelae
Synonymy and toxas (sometimes lost); with suborder Myxillina they share
the possession of sigmas (also sometimes lost). Microscleres are
Mycalina Hajdu, Van Soest & Hooper, 1994a. diverse: palmate isochelae, palmate anisochelae, palmate ungui-
ferate anisochelae, anomochelae, naviculichelae, anchorate ungui-
Definition ferate isochelae, spined isochelae, placochelae, biplacochelae,
tetropocilli, smooth sigmas, spined sigmas, sigmancistras, diancis-
Poecilosclerida possessing palmate chelae or derivates thereof tras, clavidiscs, commata, toxas, spined toxas, microxeas, rugose
(occasionally absent) and smooth megascleres of a single type; microxeas, spined microxeas, spinorhabds, raphides, trichodragmas,
tridentate chelae and toxas absent; sigmas or derivates thereof and rarely, forceps.
usually present. Skeleton plumose or plumoreticulate.
Scope
Diagnosis
In the extended concept of Mycalina presented here, nine fam-
Sponges with encrusting, massive, lobate, repent, branching, ilies are included: Cladorhizidae, Desmacellidae, Esperiopsidae,
tubular, stipitate or flabellate shape. Consistency usually soft and Guitarridae, Hamacanthidae, Isodictyidae, Merliidae, Mycalidae,
compressible, but may be firm to hard exceptionally. One family has and Podospongiidae. Together these comprise a complement of
a coralline basal skeleton. Spicule skeleton structure is basically sponges occurring all over the world in all marine habitats. Apart
plumose, with bundles of spicules fanning out towards the surface; from family Mycalidae, the individual families have a modest
in various Mycalina, these bundles may be consolidated by spongin species diversity not exceeding a few dozen species.
and interconnected by secondary bundles. At the surface, crusts of Compared to Hajdu et al.'s (1994a) original content of
tangential spicules or a reticulated skeleton may be present. Within Mycalina, the family Isodictyidae (previously assigned to hap-
Poecilosclerida, the Mycalina are unique in lacking a differentiation losclerids, see Hajdu et aI., 1994b) is added based on the study
of megasclere types, all megascleres being of a single smooth type, of Samaai et al. (1999). Podospongiidae is also included in
usually characteristic styles with a slight subapical constriction, Mycalina - previously considered part of Latrunculiidae and gener-
dubbed mycalostyles, but occasionally megascleres are strongyles, ally assigned to Hadromerida (see e.g., Bergquist, 1978), based on
oxeas or tylostyles (derived from styles as indicated by the existence studies of Kelly & Vacelet (1995) and Kelly & Samaai (this
of intermediate forms in some taxa). However, some differentiation volume). Nevertheless, the family is not 'typical' of other mycalin-
in size between ectosome and choanosome may exist and occasion- ids and is therefore included as incertae sedis (see below). The fam-
ally this is also expressed in modification of the megasclere mor- ilies Esperiopsidae (previously assigned to family Mycalidae) and
phology, but no division into main and auxiliary megascleres is Merliidae (previously assigned to Demacellidae or Hamacanthidae)
found, unlike suborders Microcionina and Myxillina. Mycalina are are revived for reasons explained elsewhere in this volume
also unique in the possession of diancistras, knife-edged sigma (see chapters by Van Soest & Hajdu).

633
634 Porifera· Demospongiae • Poecilosclerida • Mycalina

TAXONOMIC HISTORY as synapomorphy for the entire suborder Mycalina is not entirely
convincing, although holding true for this central group of
The suborder Mycalina Hajdu et al. (1994a: 126) was only Mycalina. The mycalostyles are more widespread, but appear to
recently erected from a re-evaluation of all poecilosclerid charac- be absent or rare in families Desmacellidae, Isodictyidae and
ters from the type species of most nominal genera. SEM examina- Podospongiidae. It may be significant that two of the families
tion of microsclere microstructure, especially the structure of lacking these synapomorphies also lack palmate chelae. In the
chelae and spination on megascleres, was instrumental in revising past these groups were indeed assigned to other orders,
the systematics of this group (and other poecilosclerids), and not e.g., Desmacellidae (also Biemnidae) were allocated to Axinellida;
reliant solely on light microscopy as many earlier studies. Podospongiidae to Hadromerida. In the case of Desmacellidae, the
Previously employed poecilosclerid taxa, especially the families possession of sigmas is considered compelling evidence for
Desmacidonidae (or Desmacididae) and Coelosphaeridae s.s., Mycaline membership. Affinities of family Podospongiidae are
were based on muddled combinations of megasclere shape and more obscure, with spinose microrhabd (spinorhabd) microscleres,
features of the habit, and ignored conflicting distributions of having suggested affinities with the Mycalidae based on possession
microscleres. Until Hajdu et al.'s (1994a) study, no authoritative of a sigmoid protorhabd with presumed homology to diancistra
unified classification of Poecilosclerida existed, with each major derivatives (Kelly & Samaai, this volume). For a discussion of
author employing his or her own insights, and with their analyses membership of Podospongiidae, see below.
sometimes biased by examination of limited comparative material A minority of individual species of the other families have
or based only on restricted regional faunas. Useful reviews of these oxeas or strongyles. These are considered derived from styles,
schemes are found in Bergquist & Fromont (1988) and Hajdu which is occasionally made clear from a series of intermediate
(1995), and are summarized in the various family chapters con- spicule modifications found in the same individual.
tained within this volume. The SEM studies referred to above The lack of differentiated ectosomal and choanosomal megas-
revealed that fine structures of chelae coincide with broad distribu- cleres is also not shared by all species; notable exceptions are
tions of other microsclere types such as toxas and sigmas, which Coelodischela (two species), which has strongyles and tylotes
led to the recognition of three suborders (a fourth suborder incertae which appear distinct. In Podospongiidae, megasclere diversity is
sedis has now been added in this volume, see chapter on Order also notable. What remains as a universal feature is the fact that all
Poecilosclerida). The suborder Mycalina differs from other megascleres are smooth, but many individual myxilline and micro-
Poecilosclerida by (1) the lack of differentiated main and auxiliary cionine species and genera also have smooth megascleres.
acanthostyles, (2) combined possession of palmate chelae, sigmas
and toxas, and (3) lack of spined megascleres. Inclusion of Podospongiidae

This family is revived and included in Mycalina for two com-

REMARKS pelling reasons: (I) spinorhabd microscleres derived from sigmas
or sigmoid microscleres, traceable ontogenetically, and (2) shared
Synapomorphies chemistry (norsesterterpene peroxides) of several podospongiid
genera and several Mycale species. More subtle similarities are the
In the original definition emphasis was placed on: (1) styles plumoreticulate skeleton. The spinorhabd microscleres are similar
with subapical constriction, and (2) the occurrence of sigmancis- to those of Phlyctaenopora (Barbozia), which is assigned to the
tras and derivates over the families originally united in this subor- family Mycalidae on account of the possession of palmate
der: Mycalidae, Hamacanthidae, Desmacellidae, Cladorhizidae anisochelae. Nevertheless, the position of Podospongiidae in
and Guitarridae. However, sigmancistras are absent from Mycalina is not completely clear and presently considered incertae
Desmacellidae, Isodictyidae and Podospongiidae, and only dubi- sedis. The affinities of Podospongiidae with Mycalidae and the
ously present in a single Mycalidae species (Mycale simonis) and a discrepancies with Latrunculiidae are discussed elsewhere in this
single Guitarridae species (Euchelipluma pristina). Thus, its status volume (see chapters by Kelly & Samaai and Samaai & Kelly).

KEY TO FAMILIES

(I) Microscleres entirely lacking .................................................................................................. Esperiopsidae (Ulosa, Semisuberites)

Some form of microscleres present ... .......... ................. .......... ...... ................ ................ ........................... ..................................... ....... 2
(2) Microscleres include chelae or derivates (placochelae, tetrapocilli, etc.) ........................................................................................... 5
No chelae or derivates ......................................................................................................................................................................... 3
(3) Microscleres exclusively spinorhabds ........................................................................................................................ Podospongiidae
No spinorhabds .................................................................................................................................................................................... 4
(4) Microscleres include clavidiscs ............................................................................................................................................ Merliidae
Microscleres diancistras or sigmancistras ... .......... .................. ............... ...... .......... ........................... ........... ...... ........ Hamacanthidae
Microscleres sigmas, and/or trichodragmas, no clavidiscs or sigmancistras ................................................................ Desmacellidae
(5) Placochelae or derivates (tetrapocilli, coelodiscs) present (these are strongly modified chelae in which the
frontal ala forms a flat plate attached to the rhabd by a large number of fimbriae) ......... ........... ................ ................. .... Guitarridae
No placochelae or derivatives .............................................................................................................................................................. 6
(6) Deep-sea sponges with root system, stalk and feather-like or thinly ramose shape ..................................................... Cladorhizidae
Shape may be ramose, but not featherlike ... ...... .......... ...... .......... ... ......... ............... ........... ..... ........... ........... .......... ...... ...... .... ............. 7
Porifera· Demospongiae • Poecilosclerida • Mycalioa 635

(7) Anisochelae .. .... ... ................................................... ................ ................ ................ ................ ........... ................. .......... ....................... 8
Exclusively isochelae ........................................................................................................................................................................... 9
(8) Skeleton an isodictyal-anistropic reticulation of paucispicular bundles of styles; no other microscleres .............................................. .
...................................................................................................................................................... Esperiopsidae (Amphilectus lobata)
Skeleton plumose or plumoreticulate; usually sigmas and/or other microscleres present .................................................. Mycalidae
(9) Megascleres diactinal (oxeas and/or strongyles) .............................................................................................................. Isodictyidae
Megascleres (mycalo-)styles ............................................................................................ Esperiopsidae (Amphilectus, Esperiopsis)
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Cladorhizidae Dendy, 1922

Eduardo Hajdu! & Jean Vacelet2

1 Museu Nacional, Departamento de Invertebrados, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, sIn, 20940-040, Rio de
Janeiro, RJ, Brazil & Centro de Biologia Marinha, Universidade de Slio Paulo, Slio Sebastilio, SP, Brazil. (hajdu@acd.ufrj.br)
2Centre d'Oceanologie de Marseille, Universite de la Mediterranee - CNRS UMR 6540, Station Marine d'Endoume, 13007 Marseille,
France. (jvacelet@sme.com.univ.mrs.fr)

Cladorhizidae Dendy (Demospongiae, Poecilosclerida, Mycalina) are predominantly deep water taxa, encompassing fifteen nominal genera
of which only three genera are considered to be valid. Sponges usually small, symmetrical, possessing diagonal, radiating supporting
processes and basal root adaptations for living in soft sediments, with the axial skeleton composed of monactinal or diactinal megascleres
from which radiating extra-axial tracts diverge to the ectosome. Microscleres include (an)isochelae, sigmas, forceps or micro(subtylo)styles
(microspined, spear-shaped in a few cases). However, the family lacks any remarkable synapomorphy, with characters such as the presence
of forceps, unguiferate chelae, sigmancistras, stipitate growth fonn, deep-water habitat, carnivorous habit and/or lack of an aquiferous sys-
tem not universal amongst all genera (or at least not yet demonstrated to be so), nor are these features unique to the family. These genera
are retained within this family as understood by previous authors.
Keywords: Porifera; Demospongiae; Poecilosclerida; Mycalina; Cladorhizidae; Asbestopluma; Chondrocladia; Cladorhiza.

DEFINITION, DIAGNOSIS, SCOPE isochelae and mycalostyles in E. desmophora, typical of the

Esperiopsdae - but see alternative interpretation in Hajdu
Synonymy (Guitarridae, Esperiopsdae, this volume), and complex placochelae
in E. pristina, typical of the Guitarridae).
Cladorhizeae Dendy, 1922: 58. Cladorhizidae de Laubenfels, There is still doubt concerning authorship of family
1936a: 122. Cladorhizidae. Dendy (1922) erected Cladorhizeae assigning only
Amphilectus (?) unguiculatus to it, with the only reference to inclu-
Diagnosis sion of Cladorhiza being the proposed family name. He made
no other comments on the taxon. De Laubenfels (1936a) was much
Sponges usually small, symmetrical, mostly in deep water, more thorough, which is probably why some contemporary authors
with diagonal, radiating supporting processes and basal root adap- have attributed the authorship to him. Nevertheless, ICZN Article
tations for living in soft sediments. Axial skeleton composed 11.7 (Anon., 1999) supports recognition of Dendy's authorship
of monactinal or diactinal megascleres, from which radiating with the inclusion of Cladorhiza in the family 'inferred from
extra-axial tracts diverge to the ectosome. Microscleres include the stem'.
(an)isochelae, sigmas, forceps or micro(subtylo)styles (micro-
spined, spear-shaped in a few cases). Considerable reduction to Scope
complete loss of the choanocyte layer is known for some species,
being associated with an adaptation to carnivory, preying on A great deal of attention has been focused on cladorhizids
relatively large food items. over the past decade due to the observation of some remarkable
biological traits in this mostly deep-water group (Vacelet & Boury-
Remarks Esnau1t, 1995a,b, 1996; Vace1et et al., 1995, 1996). In part, these
discoveries were made possible as a consequence of the discovery
A strong synapomorphy is missing for the Cladorhizidae. of a thriving shallow-water population of the carnivorous
Characters such as the presence of forceps, unguiferate chelae, sig- Asbestopluma hypogea Vacelet & Boury-Esnault, 1996. The most
mancistras, stipitate growth fonn, deep-water habitat, carnivorous interesting aspect of this discovery is the total absence of
habit and/or lack of an aquiferous system, are not shared by all choanocyte chambers associated with a carnivorous existence
members, or at least have not yet been demonstrated to be so; nor (presumed secondary loss), with choanocytes being amongst the
are these characters confined to the family. On the basis of strongest synapomorphy for the entire Porifera.
morphological evidence alone we prefer to keep the family intact, De Laubenfels (1936a) emphasised the symmetrical, mostly
as understood by previous authors (e.g., Levi, 1973; Hooper, stalked habit, and deep-water habitat as diagnostic features of the
1997) who assigned it to the vicinity of the Mycalidae and allied Cladorhizidae, although already recognising convergent evolution
families. The decision to keep Asbestopluma, Chondrocladia and of similar traits on other deep-water sponges. Microscleres, espe-
Cladorhiza within a single family appears to be the most parsimo- cially the chelae, were considered diagnostic at the genus level, of
nious at present, although their affinities to sponges such as which he considered nine genera to be valid. Fifteen nominal
Esperiopsis desmophora Hooper & Levi, 1989 (Esperiopsidae) and genera were considered here, of which only three are considered
Euchelipluma pristina Topsent, 1909 (Guitarridae) remain unclear. valid. One is transferred to the Crambeidae. Taxonomic decisions
These sponges share some of the characters referred to above, but made here follow the rationale proposed by Van Soest & Hajdu
additional features strongly point to distinct assignments (palmate (Mycalina, this volume).

636
Porifera· Demospongiae • Poecilosclerida • Mycalina • Cladorhizidae 637

KEY TO GENERA

(1) Anisochelae ......................................................................................................................................................................................... 2

Unguiferate isochelae ....... ................ ...... .......... .......... .... ....... ........... ................ .......... ............ ...... ........... ... .................. Chondrocladia
(2) Palmate anisochelae ....................................................................................................................................................... Asbestopluma
Anchorate or unguiferate anisochelae ......... ........... ................ ................ ...... ...... ..................... .......... ........... ........... ...... ...... Cladorhiza

ASBESTOPLUMA TOPSENT, 1901 Remarks. A canal system is lacking in Asbestopluma

hypogea Vacelet & Boury-Esnault, 1996 (Vacelet & Boury-
Synonymy Esnault, 1995a; Figs IH-J), and it is likely that this feature may
be widespread in the genus. The subgenera of Lundbeck (1905),
[Cometella] Schmidt, 1870: 49 (nomen oblitum). [Asbesto- viz., Asbestopluma, Cotylina [type species established here as
pluma] Lankester, 1882: 478 [nomen nudum]. Asbestopluma being Esperella infundibulum Levinsen, 1887 (=Asbestopluma
Topsent, 1901d: 23. Lycopodina Lundbeck, 1905: 58. Cotylina infundibulum), no material reexamined] and Lycopodina [type
Lundbeck, 1905: 68. Helophloeina Topsent, 1928d: 8. species Esperella cupressiformis var. lycopodium Levinsen, 1886
(=Asbestopluma lycopodia); by subsequent designation; de
Laubenfels, 1936a), no material re-examined] were defined mostly
Type species on the basis of habit, which has subsequently been shown to be an
unstable character (Hentschel, 1914; Vacelet & Boury-Esnault,
Cladorhiza pennatula Schmidt, 1875 (by subsequent 1996). The use of these subgenera should thus be avoided. De
designation; Topsent, 1901d). Laubenfels' (1936a) suggestion of raising Lycopodina to full
generic rank is not accepted. Conversely, Helophloeina (type
Diagnosis species H. stylivarians Topsent, 1928d; by monotypy, no material
re-examined), was erected by Topsent (1928d) on the basis of
Cladorhizidae with palmate anisochelae. a unique combination of spicules. Hooper & Levi (1989) and
Hooper (1997) suggested it may be synonymous to Meliiderma
Description of type species Ridley & Dendy, 1887, which is not accepted here. Meliiderma
(syntypes re-examined, see below) is a likely synonym of
Asbestopluma pennatula (Schmidt, 1875) (Fig. 1). Chondrocladia Thomson, 1873, well distinguished from
Synonymy. Cladorhiza pennatula Schmidt, 1875: 119, pI. I Asbestopluma. Chondrocladia and Helophloeina share small
figs 14--16. Cladorhiza bihamatifera Vosmaer, 1882a: 47, pI. I fusiform anisochelae which is here considered as strong evidence
figs 105-112. Esperia bihamatifera; Hansen, 1885: 15, pI. ill fig. 7, for their close relationship. Despite the fact that the short/stout
pI. IV, fig. 2, pI. VII figs 5, 14. Cladorhiza nordenskiOldii Fristedt, micro(tylost)(strong)yles found in both Helophloeina and
1887: 455, pI. 25 figs 56--59, pI. 31, fig. 25. Asbestopluma pennatula; Meliiderma are indeed similar in overall shape, their distinct
Topsent, 1901d: 24, 28, pI. ill figs 9a-d. Esperella plumosa dimensions (11-12 f.1m vs. 60-100 f.1m, respectively), coupled with
Arnesen, 1903: 11, pI. II fig. 1, pI. IV fig. 3, pI. VI fig. 7 (Not the presence of tylost(rong)yles in the ectosome of Asbestopluma
Esperia plumosa Carter, 1882a). Taxonomic decision for syn- and Chondrocladia, suggest the plausibility of the alternative inter-
onymy: Lundbeck (1905). pretation given above. As argued elsewhere (Hajdu, 1999), the use
Material examined. Holotype: Not seen. Other material. of derived monotypic genera (e.g., Helophloeina with desmas)
MNRJ 1236 - don. O.S. Tendal, 'Ingolf' Stn. 67, 61°30'N, clearly pertaining to more universal plesiomorphic genera (e.g.,
22°30'W (sample from ZMK). MNRJ 1237 - don. O.S. Tendal, Asbestopluma without desmas) is undesirable for it highlights the
'Ingolf' Stn. 68, 62°06'N, 22°30'W (sample from ZMK). MNRJ amount of apomorphy in a single species, at the cost of making
1238 - don. O.S. Tendal' 'Ingolf' Stn. 78, 60 37'N-27°52'W 0
paraphyletic an assemblage where many species are classified.
(sample from ZMK). In parallel with decisions taken for the Mycalidae (Hajdu, 1999
Description (adapted/rom Lundbeck, 1905). Slender, pen- and this volume), Asbestopluma and Helophloeina could be given
niform, reaching over 18 cm high; axis slightly compressed and the subgeneric rank within a larger, supposedly monophyletic genus
variably long lateral branches inserted in the sides (Fig. lA). Asbestopluma. It should be made clear though that for phyloge-
Oscula and pores apparently lacking. Axial skeleton with a rela- netic purposes genus Asbestopluma would mean absolutely the
tively powerful, densely spiculated exterior layer. Microscleres same as subgenus Asbestopluma, the latter being thus redundant
abound in the layer coating the branches. Megascleres: styles in the and useless. The same reasoning applies to Helophloeina, be it a
central portion of the main axis, 680-1000 f.,Lm long, 15-32 f.,Lm genus or a subgenus, and species H. stylivarians Topsent, 1928d.
thick (Fig. lB); subtylostyles in the branches and in the outer layer The genus Cometella Schmidt, 1870: 49 was erected for type
of the axis, 520-750f.,Lm long, 9-17f.,Lm thick (Fig. lC). species CometeUa gracilior Schmidt, 1870: 49 pI. IV fig. 9 (by
Microscleres: subtylostrongyles, irregularly sinuous, microspined, subsequent designation; De Laubenfels, 1936a: 151). Schmidt's
in the coating of the stalk, 50-137 f.1m long, 1-2.8 f.1m thick drawing shows a pedunculate sponge with a small ovate main
(Fig. ID); large palmate anisochelae, with the lower end polylobate, body. A slide of the lectotype BMNH 70.5.3.97 was reexamined.
48-63 f.1m in total height (Fig. I G); small palmate anisochelae, This contains sections and many dissociated spicules. The spicule
fusiform, 10-11.4f.1m in total height (Fig. IF); sigmancistras, complement conforms to Asbestopluma, with megascleres styles
21-24 f.1m long (Fig. IE). with blunt endings, 900-1400 X 20-25 f.,Lm, smooth echinating
638 Porifera· Demospongiae • Poecilosclerida • Mycalina • Cladorhizidae

I
I I E

B c o G

Fig. 1. A, Asbestopluma pennatula Schmidt (as Cladorhiza) (reproduced from Schmidt, 1875, pI. I, fig. 14). B-F, Asbestopluma pennatula, drawings of
spicules (reproduced from Lundbeck 1905, pI. 10, fig. 4). B, styles from the axis (scale 80 fj-m). C, subtylostyle from the lateral branches (scale 75 fj-m).
D, microspined subtylostrongyles from the stalk-coating (scale 17.5 fj-m). E, sigmancistra (scale 10 fj-m). F, small palmate anisochelae (scale 3.2 fj-m).
G, large palmate anisochelae (scale 12 fj-m). H-J, Asbestopluma hypogea Vacelet & Boury-Esnault, 1996. H, living specimen (scale 2.5 min). I, ho1otype
preserved in formalin (scale 3.1 mm). J, SEM photo of a palmate anisochelae (scale 2 fLm).

subtylostyles, 300--600 X 2-4/-Lm, irregularly sinuous microspined Asbestopluma has prevailing usage in the sense ofICZN article 23.9.
subtylostrongyles, 90-120 X 1-3/-Lm, sigmancistras 20-25/-Lm, and [Cometella 1is a nomen oblitum in favour of Asbestopluma.
palmate anisochelae 10-12 /-Lm. The genus Cometella was not recog-
nized so far as a genus in Cladorhizidae, as Schmidt himself used it CHONDROCIADIA THOMSON, 1873
for sponges from a variety of families, Cometella stellata Schmidt,
1870 (= Halicometes in Tethyidae) and Cometella spermatozoon Synonymy
Schmidt, 1875 (=Suberites in Suberitidae), whereas Carter used it
for sponges of yet another series of families: Cometella simplex Chondrocladia Thomson, 1873: 188. Crinorhiza Schmidt,
Carter, 1876 (=Stylocordyla in Stylocordylidae), and Cometella 1880b: 83. Meliiderma Ridley & Dendy, 1887: 102, text-fig. 6,
pyrula Carter, 1876 (=Crella in Crellidae). More importantly, pI. XX fig. 11, pI. XXI figs 6a-c, 14. Neocladia Koltun, 1970: 193,
the name has not been used after 1876 and the junior name text-figs 18.1-6, pI. VI fig. 6.
Porifera· Demospongiae • Poecilosclerida • Mycalina • Cladorhizidae 639

o
A

Fig. 2. A, Chondrocladia virgata Thomson (reproduced from Thomson, 1873: 188, 20-40cm high), B-D, Chondrocladia virgata, drawings of spicules
(reproduced from Carter, 1874a, pI. 14 fig, 20-21 and pI. IS fig. 38). B, anchorate isochelae (scale 1O.6IJom). C, sigma (scale 10 IJom). D, style (scale
230 IJom). E, a living specimen of Chondrocladia sp. photographed in situ in the North Pacific, 5320 m depth (from Tendal et ai., 1993, photo Volodin).
F, same specimen after collection, preserved in alcohol (scale 3.5 mm). G, SEM photo of an anchorate isochelae of an unidentified Chondrocladia from the
deep North Pacific (scale 30 IJom).

Type species syntypes) - Kyska Harbour, 'Unalashka', U.S.A. Chondrocladia

gigantea (Hansen, 1885): MNRJ 1243 - don. O.S. Tendal,
Chondrocladia virgata Thomson, 1873 (by monotypy). 'Michael Sars', 62°53'N, 04°14'E, det. W. Lundbeck (sample from
ZMK). MNRJ 1244 - don. O.S. Tendal, 'Godthab', stn. 162,
Diagnosis 62°48'N, 60 46'W (sample from ZMK). MNRJ 1245 - don. O.S.
0

Tendal, 'Bioice', stn. 2085, 67°15.67'N, 17°26.41'W, det. O.S.

Cladorhizidae with isochelae. Tendal (sample from ZMK). Chondrocladia pulchra Lambe, 1894:
CMNIl9002824 (fragment from syntype) - Chika Island, Alaska,
Description of type species U.S.A. Syntypes of Meliiderma stipitata (Ridley & Dendy, 1886):
BMNH 1887.5.2.191 and BMNH 1891.10.3.52a.
Chondrocladia virgata Thomson, 1873 (Fig. 2). Description (adapted from Carter, 1874a; Thomson, 1875).
Synonymy. Chondrocladia virgata Thomson, 1873: 188. Erect, branching scantily and dichotomously, 20-40 cm high. With
Carter, 1874a: 217, pI. XIV figs 20-21, pI. XV fig. 38. divergent rooting processes, short straight stalk (ca. 6mm wide),
Material examined. Holotype: Not seen. Other material. long axis bearing abundant conical processes (ca. 25 mm long,
Chondrocladia alaskensis Lambe, 1894: CMNI1900-2828 - 4 mm across at the base) arranged around it in a confused spiral
Chika Island, Alaska, U.S.A. CMNl1900-2869 (fragments from manner (Fig. 2A). Choanosomal skeleton a central axis formed by
640 Porifera· Demospongiae • Poecilosclerida • Mycalioa • Cladorbizidae

long subtylostyles, from which transverse bundles built by the Type species
same spicules radiate to form the conical processes. Ectosomal
skeleton with large amounts of microscleres, and smaller megas- Cladorhiza abyssicola Sars, 1872 (by monotypy).
cleres piercing the sponge surface. Axis and spicules diminishing
in size upwards from the base. Megascleres: (subtylo)styles, Diagnosis
smooth, slightly curved, fusiform, head narrower than the shaft, ca.
1530/Lm long (average largest size) and 28/Lm across (greatest Cladorhizidae with anchorate/unguiferate anisochelae (Fig. 3f).
diameter, Fig. 2D). Microscleres: anchorate isochelae, with seven
teeth at each end, curved shaft, ca. 67/Lm long (average largest Description of type species
size) and 21/Lm across (breadth of head, Fig. 2B). Sigmas, smooth,
ca. 25/Lm long (average largest size, Fig. 2C). Cladorhiza abyssicola Sars, 1872 (Fig. 3).
Remarks. In some preserved specimens of Chondrocladia Synonymy. Cladorhiza abyssicola Sars, 1872: 65, pI. VI,
the conical processes bear irregular swellings which appear to be figs 16-34; Lundbeck, 1905: 79, pI. XII, fig. 2.
inflated, translucent spheres in the living state (Tendal et al., 1993; Material examined. Holotype: Not seen. Other material.
KUbler & Barthel, 1999; Fig. 2E-G). The aquiferous system Syntype ofAxoniderma mirabile (Ridley & Dendy, 1886): BMNH
consists of rows of large choanocyte chambers running along the 1887.5.2.141. Cladorhiza corticocancellata Carter, 1876: MNRJ
main axis of the sponge. Crinorhiza (no material re-examined 1246 - don. O.S. Tendal, 'Ingolf', stn. 143, 62°58'N, 07°09'W, det.
here) was erected for cladorhizids with branches condensed around W. Lundbeck (sample from ZMK). Cladorhiza geUda Lundbeck,
a flattened stalked body (cf. Topsent, 1930), the branches without 1905: MNRJ 1240 (syntype) - don. O.S. Tendal, 'Ingolf', stn. 113,
terminal swellings, and a special category of ectosomal microsub- 69°31'N, 07°06'W (sample from ZMK, syntype); MNRJ 1241 -
tylostyles missing. Ridley & Dendy (1887) did not attach much don. O.S. Tendal, 'Michael Sars', 600 19'N, 05°39'E (sample from
confidence to these diagnostic characters, and used the name ZMK). Cladorhiza tenuisigma Lundbeck, 1905: MNRJ 1242 (syn-
'Crinorhiza' to denote a special habit only. Topsent's (1930) sub- type) - don. O.S. Tendal, 'Ingolf', stn. 105, 65°34'N, 07°31'W
generic arrangement for Chondrocladia should be abandoned. For (sample from ZMK).
instance, forms of 'Crinorhiza' were described for Cladorhiza too, Description (adapted from Sars, 1872; Lundbeck, 1905).
thus rendering any group of species recognized on the basis of the Slender, unbranched (young specimens), or with relatively long
diagnostic characters for Crinorhiza very suspect in phylogenetic branches issuing from a central axis; below a richly branched root.
terms. The type species of Crinorhiza, C. amphactis Schmidt, Total length may exceed 7 cm. Secondary processes, thin conical
1880b, possesses the characteristic spicule complement of a pointed and slightly curved, strongly compressed laterally, issue
Chondrocladia, and our opinion is that both genera are synonyms. circularly from the stem and the side branches. A variably long sec-
This contrasts with opinions implicit in the recent literature (e.g., tion of the stem, just above the roots, may be devoid of secondary
Levi, 1973; Hooper, 1997). Chondrocladia (Amorphocladia) processes (Fig. 3A). Live-colour yellowish transparent to reddish-
Topsent, 1930, was erected for Chondrocladia alaskensis Lambe, yellow. The skeleton is closely dependent on the form; it consists in
1894 and C. pulchra Lambe, 1894, but these species are best axes and branches of a powerful, polyspicular fibre, often of great
referred to Crambe Vosmaer, 1880b/Monanchora Carter, 1883c thickness, where the fibres supporting the branchlets are inserted.
(see Crambeidae, this volume). The above explanation leaves In the axial fibres spongin is found. Spicules: megascleres - styles,
only Chondrocladia (Chondrocladia) as a potentially valid, but slightly fusiform or not, slightly bent or straight, 390-730 /Lm
certainly superfluous subgeneric assemblage. Meliiderma (type (Fig. 3B). Microscleres - anchorate/unguiferate anisochelae with
species Chondrocladia stipitata Ridley & Dendy, 1886; type spec- five teeth on each termination, 21-25 /-Lm (Fig. 3E); sigmata, large,
imen re-examined) is considered a synonym of Chondrocladia in typical, 78-100 /Lm (Fig. 3C); sigmancistras, 40-42 /-Lm (Fig. 3D).
view of its possession of typical anchorate isochelae, this being the Remarks. The smaller sigmas of C. abyssicola are treated
original interpretation by Ridley & Dendy too (Ridley & Dendy, here as sigmancistras - Lundbeck (1905) leaves no doubt in this
1886). The possession of micro(tylost)(strong)yles in the ectosome respect. Nevertheless, a definitive decision can only be made after Sars'
of Meliiderma sets the species apart, but does not warrant the erec- (1872) specimens have been re-examined. Trochoderma Ridley &
tion of a monotypic higher taxon. Neocladia (no material exam- Dendy (1886) is a junior homonym of Trochoderma Theel (1877,
ined), on the contrary, is set apart by the peculiar shape of its Echinodermata), and was replaced by Axoniderma Ridley & Dendy.
forcepslhirotule-like isochelae, but again it is considered here The latter genus (type species Trochoderma mirabile Ridley &
that the erection of a monotypic taxon is unwarranted (refer to the Dendy, 1886), comprises a single species, distinguished by the pecu-
discussion of Asbestoplumal Helophloeina above). liar shape of its 'pseudoamphiasters', which are indeed birotule-like,
modified anchorate isochelae. In spite of arguing for this character's
possible low classificatory/phylogenetic significance, Ridley &
CLADORHlZA SARS, 1872 Dendy (1887) erected the genus Axoniderma, which differs
from Cladorhiza mainly by the possession of these 'pseudoamphi-
Synonymy asters'. Exaxinata (no material re-examined; type species
C. oxeata Lundbeck, 1905), is another monotypic genus, erected
Cladorhiza Sars, 1872: 65, pI. VI figs 16-34. [Trochoderma] on account of its possession of oxeas instead of the typical styles in
Ridley & Dendy, 1886: 344 (preocc. by Trochoderma Theel, most cladorhizids. Raoa (no material re-examined; type species
1877, Echinodermata). Axoniderma Ridley & Dendy, 1887: 96, C. tridentata Ridley & Dendy, 1886), is yet another monotypical
text-fig. 5, pI. :xx fig. 5, pI. XXI figs 8-10 (replacement name genus, set apart by its anchorate anisochelae with only three teeth
for Trochoderma Ridley & Dendy). Exaxinata de Laubenfels, and hollow cup-shaped sponge habit. It is postulated here that
1936a: 122. Raoa de Laubenfels, 1936a: 123. A. mirabile, C. oxeata and C. tridentata belong to a clade together
Porifera· Demospongiae • PoeciloscIerida • Mycalina • Cladorhizidae 641

Fig. 3. A, Cladorhiza abyssicola Sars 1872, a ramified specimen (from Sars, 1872: fig. 17). B-E, C. abyssicola, drawings of spicules (reproduced from
Sars, 1872). B, style from the axis (scale 50 !Lm). C, sigma (scale 20 !Lm). D, sigmancistras (scale 20 !Lm). E, anchorate anisochelae (scale 6.S !Lm). P, SEM
view of an anisochelae from an undescribed species from Barbados (scale 4.4!Lm).

with C. abyssicola, to which neither species of Asbestopluma or deeply thanked for the provision of scientific literature. The fol-
Chondrocladia belong. In the absence of a phylogeny for the species lowing curators are gratefully acknowledged for the provision of
of Cladorhiza, Axoniderma, Exaxinata and Raoa are considered here type specimens and/or other specimens on loan: Judith Fournier
SUbjective synonyms of Cladorhiza. Otherwise, Cladorhiza may (CMN) Rob W.M. Van Soest (ZMA), Ole S. Tendal (ZMK), Clare
quite possibly be made paraphyletic. A canal system is apparently Valentine (BMNH). Rob W.M. Van Soest is further acknowledged
lacking, as verified in Cladorhiza Metanophila by Vacelet & Boury- for the provision of SEM facilities during EH's PhD sabatical in
Esnault, 2002. Amsterdam, when most of his morphological studies on the
cladorhizids were conducted and from bringing Schmidt's
ACKNOWLEDGEMENTS Cometella to our attention. Financial support to EH to attend
the 'Systema Porifera' Workshops and to construct the means for
We want to express our sincere thanks to Dr John N.A. writing this revision was obtained from CAPES, CNPq, FAPERJ,
Hooper (Queensland Museum) and Dr Rob W.M. Van Soest and FAPESP, from Brazil, and the organisers of the International
(University of Amsterdam), for inviting us to take part in the Conference on Sponge Science (Otsu, Japan). N thanks K.
'Systema Porifera' project, and also for valuable discussions Tabachnik for providing in situ photograph and specimen of
throughout the preparation of this manuscript. Claude Levi is Chondrocladia sp., and C. Marschal for his help with illustration.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Desmacellidae Ridley & Dendy, 1886

Eduardo Hajdu· & Rob W.M. Van Soest2

1 Museu Nacional, Departamento de Invertebrados, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, sin, 20940-040, Rio de
Janeiro, RJ, Brazil & Centro de Biologia Marinha, Universidade de Sao Paulo, Sao Sebastiao, SP, Brazil. (hajdu@acd.ufrj.br)
2 Zoological Museum, University of Amsterdam, P.O. Box 94766,1090 GT Amsterdam, Netherlands. (soest@science.uva.nl)

Desmacellidae (Demospongiae, Poecilosclerida) are poecilosclerids lacking chelae, a feature shared with the closely related families
Hamacanthidae and Merliidae. Desmacellidae are distinguished from these families by the lack of diancistra-like microscleres.
Noticeable in this group is a preponderance of toxic or dermatitis-producing sponges. Six valid genera are included in the Desmacellidae,
viz., Biemna, Desmacella, Dragmatella, Microtylostylifer, Neofibularia and Sigmaxinella. Nominal genera Allantophora and Toxemna are
considered junior synonyms of Biemna. Tylodesma, Tylosigma, Sigmotylotella and Sigmatoxella are junior synonyms of Desmacella.
Hallmannia is a junior synonym of Microtylostylifer. [Fibula ria] is a senior synonym of Neofibularia, but is preoccupied. Sigmaxia is
considered a junior synonym of Sigmaxinella.
Keywords: Porifera; Demospongiae; Poecilosclerida; Mycalina; Desmacellidae; Biemna; Desmacella; Dragmatella; Microtylostylifer;
Neofibularia; Sigmaxinella.
--.---.--------.---------------------~

DEFINITION, DIAGNOSIS, SCOPE Merliidae. Hooper (1984b) revised some of the conflicting ideas
existing in the literature regarding the concept of the Desmacellidae.
Synonymy Of special concern here is the hypothesis put forward by Hallmann
(1917 c) that Desmacella Schmidt, 1870 (and Desmacellidae) would
Desmacellina Ridley & Dendy, 1886: 336. Desmacellinae be close to Axinellidae Carter, 1875b; while Biemna Gray, 1867a
Ridley & Dendy, 1887: 58. Biemnidae Hentschel, 1923: 406. (and a separate family Biemnidae) would be closer to the Mycalidae
Sigmaxinellidae Levi, 1955: 84. according to a.o. Burton (1930c) and Levi (1957). However, follow-
ing the majority of authors (e.g., Van Soest, 1984b; Bergquist &
Definition Fromont, 1988; Hooper et al., 1991), we prefer to assign both
Biemna and Desmacella to the Poecilosclerida and to the same fam-
Mycalina lacking chelae and diancistra-like microscleres. ily. Since Desmacellidae has priority over Biemnidae, the latter is
Microscleres include sigmas, microxeas, commata and raphides. designated a junior synonym. Desmacellids are lipochelous poe-
cilosclerids which seemingly attained this condition through the
Diagnosis loss of their ancestral chelae (following Van Soest, 1984b as
Biemnidae). Other lipochelous poecilosclerid genera, in general,
Encrusting, massive, cup-shaped, fan-shaped and branching are confidently assigned to various families/suborders on the basis
sponges. Megascleres are usually styles, occasionally oxeas or of possession of one or more synapomorphies shared with these
strongyles. Spicules are typically enclosed within plumose, reticu- groups (e.g., acanthostyles, or rosettes of microscleres, etc.).
late, halichondroid-reticulate or compressed axial fibres. Micro- However, apart from the likely loss of chelae and presumably also
scleres are diverse, usually consisting of sigmas (one genus the loss of sigmancistra-like microscleres, desmacellids do not
excepted), and often including microxeas of several sizes, raphides share additional positive traits, so the monophyletic status of this
in trichodragmata or occurring singly, commata, some of which group and its affinities with other families in the Mycalina, remain
may be microspined or rugose. Species belonging to this family weakly established. Few reproductive data have been published on
may occur in all oceans and at all depths. desmacellid genera to date. Brien (1973a: 304) records incubated
larvae in Tylodesma (=Desmacella) annexa, inferring viviparity.
Scope Alternatively, Neofibularia nolitangere is an oviparous sponge with
spectacular synchronous gamete release in reef populations
Six genera out of fifteen nominal genera are considered valid, (Reiswig, 1970a; Hoppe & Reichert, 1987). Although not an exclu-
Biemna, Desmacelia, Dragmatella, Microtylostylifer, Neofibularia, sive character for Desmacellidae, it is nevertheless striking that
and Sigmaxinelia. many species are the cause of severe dermatitis when touched.

Taxonomic remarks Reviews

A strong synapomorphy for Desmacellidae is lacking. There Hallmann (1917c), Burton (1930c), Hartman (1967), Hooper
are no chelae, nor any diancistra-like microscleres. Toxiform (1984b), Van Soest (1984b), Hooper & Wiedenmayer (1994),
microxeas and commata are common, but are shared with Hajdu (1995, 1999).

642
Porifera· Demospongiae • Poecilosclerida • Mycalina • Desmacellidae 643

KEY TO GENERA

(1) Two widely divergent sizes of subtylostyles, the smaller may be considered rnicroscleres ...................................... Microtylostylifer
Subtylostyles may be present, but no distinct microsubtylostyles ....................................................................................................... 2
(2) Fistular sponge with parchment-like surface enclosing a bladder-like body; styles and trichodragmas, no sigmas ........ Dragmatella
Sigmas present ............ ................ ................ ................ ...................................... .................... ....................... ..... ............ ............... ........ 3
(3) Skeleton strongly compressed axially .............................................................................................................................. Sigmaxinella
No axial condensation; skeletons plumose, plumo-reticulate, reticulate or vaguely halichondrioid ................................................... 4
(4) Megascleres tylostyles, occasionally styles; microscleres limited to sigmas and raphides ................................................ Desmacella
Megascleres (subtylo-)styles, oxeas or strongyles; in addition to sigmas and raphides there are microxeas and
often also commata; frequently producing dermatitis ........................................................................................................................... 5
(5) Ectosomal skeleton arranged brush-like; choanosomal skeletons mostly plumose or plumoreticulated ................................... Biemna
Ectosomal skeletons tangential, choanosomal skeletons neatly reticulated, spongin mostly abundant ........................... Neoflbularia

BIEMNA GRAY, 1867 Material examined. Holotype: BMNH 1877.5.21.2112 -

Unregistered slide of Halichondria variantia, labeled as
Synonymy 'Hymeniacidon variantia, St Katherine's Rocks, Tenby, Mrs Brett.
1854'. Other material. Several specimens from Ireland and
Biemna Gray, 1867a: 538. Allantophora Whitelegge, 1907: Norway, a.o. ZMA POR. 4288 - Sherkin Island, Ireland. The type
505. Toxemna Hallmann, 1917c: 673. [Toxemmal de Laubenfels, material of Desmacidon peachi could not be found in a recent
1936a: 124 (/apsus). search of the BMNH collections (Ms C. Valentine, pers. comm.).
Description. Encrusting, becoming a thick cushion (e.g.,
Type species 1-1.5 cm thick by 7 cm diameter; thickest at the centre and tapering
to the edge). It can grow into cup-like or lamellate forms
Desmacidon peachi Bowerbank, 1866: 349 (by original when mature. Colour beige, occasionally yellow. Surface uneven
designation). This is generally considered a junior synonym of (Fig. 1C) with conulose projections, due to the ends of spicule
Halichondria variantia Bowerbank, 1861: 69. fibres supporting the surface. Larger specimens may have a dis-
tinctly 'lumpy' or shaggy appearance due to development of sur-
Definition face projections. Several oscules, scattered across the upper surface
of the sponge, seen as dark holes in surface. Inhalant pore areas are
Demacellidae with stylote or exceptionally oxeote megascle- obvious between conulose projections when seen alive.
res arranged in plumoreticulate fashion; ectosomal skeleton con- Consistency very soft and friable. Skeleton plumose. Stout multi-
sists of the brushed endings of choanosomal tracts; microscleres spicular fibres (of styles), branching and anastomosing, ascend
include sigmas, rnicroxeas, commata and raphides. through the sponge and support the surface. Tissue is packed with
abundant microscleres. Minimal amounts of spongin are present.
Diagnosis Megascleres, styles, 360-(580)-700 p.,m, abruptly bent near the
rounded end. Microscleres, sigmas of two size categories, ca.
Massive, cup-shaped or tubular sponges, with uneven surface. 50-70 p.,m and ca. 10-20 p.,m; microxeas and raphides, grading into
Plumose or plumoreticulate choanosomal skeleton, with variable each other, ca. 63-165 p.,m, gathered loosely into trichodragmata;
development of spongin fibres cored by (subtylo-)styles of a single commata, 10-20 p.,m, looking like small, halved sigmas. These
size, occasionally replaced by oxeote spicules; ectosomal skeleton microscleres are very numerous. Ecology and distribution. On ver-
made up of brushes of megascleres making the surface often tical surfaces with moderately strong water movement, on under-
shaggy; microscleres include sigmas, raphides, microxeas, com- side of boulders; on mud, gravel, stones. Tolerant of brackish
mata, rnicrostrongyles and spheres. Most species cause a dermatitis- conditions. Arctic-boreal in distribution, south to ca. 40oN. Mostly
like reaction when in touch with bare skin. About 55 species occur recorded from deep water (down to 1608 m), although type locality
in all oceans, in a large depth range. (Tenby, S England) was intertidal. Recorded recently from Ireland,
Wales, Scotland, Guernsey. If synonymy is correct then it occurs
Description of type species also off the coast of East Greenland, down to 250 m.
Remarks. In the present concept of the species high arctic
Biemna variantia (Bowerbank, 1861) (Fig. lA, C). specimens which tend to have styles up to 1500 p.,m long are tenta-
Synonymy. Halichondria variantia Bowerbank, 1861: 69 tively included (various 'species' of Lundbeck, 1902 are thus con-
(see Burton, 1930c: 522 for full synonymy). Desmacidon peachi sidered synonyms). This species has had a chequered taxonomic
Bowerbank, 1866: 349; Desmacella peachi var. stellifera Fristedt, history and a revision may decide that its name has been too
1885: 29: pI. II fig. 10; Desmacella peachi var. groenlandica severely compromised and has to be changed. A tropical represen-
Fristedt, 1887: 441, pI. XXIV figs 38-45, pI. XXVIII fig. 14. tative of Biemna showing a full complement of spicules is Biemna
Desmacidon koreni Schmidt, 1875: 117, pI. I fig. 7. Gellius saucia Hooper et al., 1991 (Fig. lB).
capilliferus Levinsen, 1887: 357, pI. XXX figs 7-10. Raphiodesma The genus Allantophora Whitelegge, 1907: 505 (by mono-
aculeatum Topsent, 1888: 152. Desmacella hamifera Lundbeck, typy, no material re-examined here) was erected for Allantophora
1902: 93, pI. VII figs 4-6, pI. XVII fig. 1. pUcata Whitelegge, 1907: 505, pI. XLV fig. 28 (holotype is AMS
644 Porifera· Demospongiae • Poecilosclerida • Mycalina • DesmacelIidae

__ ~B

,
F

Fig. 1. A-B, Biemna. A, habit (scale I cm) and spicules (scales 50 /Lm) of Desmacidon peachi Bowerbank, 1866 (=Biemna variantia) (after Bowerbank,
1874, pI. LXIII figs 1-7). B, Biemna saucia Hooper et al., 1991, skeleton (scale 500 /Lm), megascIeres (scale 50 /Lm) and spicules (scale 25 /Lm) (copied
from Hooper et al., 1991, figs 1-9). C, Photo of live specimen of Biemna variantia, Ireland (courtesy B. Picton). D-E, Desmacella. D, skeleton (scale 500
/Lm) and spicules (scale 50/Lm) of Desmacella pumilio Schmidt, 1870 (copied from Van Soest, 1984b: fig. 53). E, photo of specimen of Desmacella
pumilio (copied from de Laubenfels, 1936a: pI. 22 fig. 1 (approx. life size). F-G, Dragmatella. F, habit of Desmacella aberrans Topsent, 1892a (copied
from his pI. II fig. 7). G, spicules of the same (copied from his pI. IX, fig. 10).

G4345, cf. Hooper & Wiedenmayer, 1994: 157), on the basis of its Type species
possession of 'echinating' megascleres. Hallmann (1916b) was
hesitant about the status of Allantophora, which was used both as Desmacella pumilio Schmidt, 1870: 53 (by subsequent
a separate genus and as a subgenus of Biemna. Hooper (1984b), designation; Dendy, 1922b; invalid subsequent designation of
Hooper et al. (1991) and Hooper & Wiedenmayer (1994) consid- D. annexa, by Topsent, 1925c: 704).
ered Allantophora a subjective junior synonym of Biemna
(contra Burton, 1930c), and this view is shared here, stressing Dermition
the similar spicule complement of the type species of both. The
genus Toxemna Hallmann, 1917c: 673 was erected (by original Desmacellidae with monactinal megascleres, arranged in
designation) for type species Desmacella tubulata Dendy, 1905: plumose bundles; microscleres sigmas and raphides; the latter may
155, pI. IX fig. 4 (type in BMNH) on the supposed occurrence be absent.
of toxas in its type species. The spicules described by Dendy as
toxas, organized in dragmata, are commata instead (Van Soest, Diagnosis
1984b). The relationships of Biemna to Neofibularia are discussed
below. Massive or thinly encrusting sponges with hispid surface.
Reticulate, plumo-reticulate and vaguely halichondrioid choanosomal
skeleton; poorly developed spongin fibres; megascleres tylostyles or
DESMACELIA SCHMIDT, 1870 styles of one or more sizes; microsc1eres sigmas and raphides. About
40 species distributed over all major areas of the oceans, predomi-
Synonymy nantly in deeper waters.

Desmacella Schmidt, 1870: 53. Tylodesma Thiele, 1903a: Description of type species
944. Sigmotylotella Burton, 1932b: 295. Sigmatoxella de
Laubenfels, 1936a: 121. Desmacella pumilio Schmidt, 1870 (Fig. ID-E).
Porifera· Demospongiae • PoeciioscIerida • Mycalina • Desmacellidae 645

Synonymy. Desmacella pumilio Schmidt, 1870: 53, pl. V, with a curved shaft and well-developed spherical heads, arranged
fig. 14; de Laubenfels, 1936a: 114, pl. 22, fig. 1; Van Soest, 1984b: in a confused reticulation; and smaller ectosomal tylostyles (600 X
136, pl. IX, fig. 2, text-fig. 53; Desmacodes pumilio; Vosmaer, 1880: 12 /-lm), similar in shape to the choanosomal ones, forming a dense
108; Tylodesma pumilio; Hallmann, 1916b: 519; Burton, 1930c: 525. palisade. Microscleres are sigmas, 28 /-lm. Hooper (1984b) already
Material examined. Holotype: Unknown. Schizotype: expressed doubts over the validity of this genus, arguing that tuber-
BMNH 1870.5.3.27 - Slide of Schmidt's specimen, locality not culate heads may be found in some Desmacella species, such as D.
mentioned, depth 592.5m. ZMA 4761 - near Jamaica, 18°20'N grimaldii (Topsent, 1892a) and D. ithystela Hooper, 1984b. What is
77°20'W, described in Van Soest, 1984b. left as a seemingly unique feature of Sigmotylotella suberitoides is
Description. The properties of the type species remain elu- its ectosomal palisade, which is not common in mycaline sponges.
sive, because the original description is minimal, and subsequent Ectosomal brushes, on the other hand, are very common, and these
records of additional specimens show some discrepancies. Schmidt may be arranged in a dense side-by-side manner, which is often
mentions there are two small, club-shaped, tubular sponges, with difficult to distinguish from a true palisade.
apical oscula, with tylostyles of varying sizes lying partly in fibres, The genus Sigmatoxella de Laubenfels, 1936a: 121, with type
and sigmas. It would also appear that there is no dermal skeleton species Desmacella annexa Schmidt, 1870: 53 (by original desig-
and that the outermost spicules of the main skeleton project slightly nation), was erected on account of the addition of toxiform
at the surface. De Laubenfels' specimen (Fig. IE) is 'amorphous', raphides to an otherwise typical Desmacella set of spicules. We
8 X 3 cm, and there are groups of oscules, whereas Van Soest's re-examined a slide of the type in the Natural History Museum,
specimen is a thin encrustation. Spicules (Fig. 1D), megascleres London, BMNH 1870.5.3.29; this appears to be the only extant
tylostyles, size given by de Laubenfels: 530-1400 X 9-17 /-lm, type material (Desqueyroux-Faundez & Stone, 1992). The species
Van Soest gives 320-(480)-800 X 9-(11)-15 /-lm. Microscleres. is described as massive or more thinly encrusting, occasionally
Schmidt and de Laubenfels give a single size of sigmas, 38-78 /-lm forming erect masses, with irregular hispid surface. Size may
and 30-40/-lm respectively, whereas Van Soest records two sizes, exceed 5 cm. Size of oscules not recorded. Consistency fragile.
larger sigmata, 45/-lm, smaller sigmata, 15/-lm. Apparently there Skeleton plumose, with ill-defined bundles and scattered single
are no raphides. This is a Caribbean deep-water species found spicules generally directed towards the surface; no special ectoso-
between Cuba and Florida, and near Jamaica. mal skeleton. Megascleres, tylostyles with prominent tyles, quite
Remarks. The genus Tylodesma was erected by Thiele variable in size, 220-1050 X 2.5-14 /-lm. Microscleres, sigmas in
(1903a: 944) who believed that Desmacella was a synonym of two size classes: 25-42 and 11-15/-lm; toxiform raphides, with
Hamacantha Gray, 1867a, because the type species of the latter, a bend in the middle with a slight counter-bend on both sides:
Hymedesmia johnsoni Bowerbank, 1866, had been included by 50-115 X 0.5-2 /-lm. The species was originally described from
Schmidt (1870) in his list of species assigned to Desmacella. 350 m deep off the coast of Florida, with a tantalizingly short
Dendy (1922b: 56) established the type species of Desmacella, as description. The assignment of European specimens to Schmidt's
D. pumilio. Thiele's (1903a) view was in agreement with an earlier species is tentative and conspecificity with specimens from the
opinion expressed by Vosmaer (1887), and also based on the recog- Indian Ocean likewise needs further corroboration. Sigmatoxella
nition that the remaining species assigned to Desmacella seemed to is considered a subjective junior synonym of Desmacella, in
belong to a natural group. Tylodesma had its type species estab- agreement with opinions expressed by Hooper (1984b), Van
lished by Hallmann (1916b: 518), as Halichondria inornata Soest (1984b) and Hooper et al. (1991).
Bowerbank, 1866: 271, reluctantly, because the properties of
Schmidt's (1870) Desmacella species did not appear to him suffi-
ciently known. Bowerbank's species is represented by slides in
BMNH made from the type, bearing the number 'Bk 654'. A recent DRAGMATELLA HALLMANN, 1917
search of the collections did not reveal any extant specimens (Ms
C. Valentine, pers. comm.). This is a greyish white, massive, irreg- Synonymy
ular species, occasionally with creeping branches. Thickness up to
7 cm. Surface irregular. Oscules few and rather large. Consistency Dragmatella Hallmann, 1917c: 640.
fragile. Ectosomal skeleton a tangential layer of single megascleres
or a reticulation of bundles, or a confused mass. Choanosomal Type species
skeleton multi spicular bundles arranged in irregular reticulation.
Megascleres tylostyles of great size variation, 190-1000 X Desmacella aberrans Topsent, 1892a: 85 (by original
6-18/-lm. Microscleres are sigmas, 20-45/-lm, no raphides. Deep designation).
water, 100-270 m, recorded from Shetland, W coasts of Ireland,
Norway, Portugal. Wiedenmayer (1977b) followed by many Definition
authors (e.g., Hooper, 1984b; Van Soest, 1984b; Bergquist &
Fromont, 1988), argued that Halichondria inornata is very close Desmacellidae with fistular habit; spicules styles and
to Desmacella pumilio in terms of spiculation, so that Tylodesma trichodragmata.
is better regarded as a junior synonym of Desmacella. This view
is shared here. Diagnosis
The genus Sigmotylotella was erected on the fact that the bases
of the tylostyles are often tuberculate. Its type species (by mono- Encrusting sponges with hollow bladder-like habit, pointed fis-
typy), Sigmotylotella suberitoides Burton, 1932b: 295, tules and parchment-like skin. Ectosomal skeleton a feltwork of tan-
fig. 20 (holotype in BMNH 1928.2.15.400, labeled 'Discovery Exp. gentially arranged styles, choanosomal skeleton scanty bundles of
Stn. 6.'), possesses large, choanosomal tylostyles (1000 X 18 /-lm), styles. Microscleres exclusively long raphides arranged in sinuous
646 Porifera· Demospongiae • Poecilosclerida • Mycalina • Desmacellidae

trichodragmata; no sigmata. The type species is from deep water in arranged rosette-like. Main skeleton plumoreticulate, consisting
the North Atlantic; a second species from the South Pacific remains of large subtylostyles (mycalostyles). Microscleres very common
to be described. in the type species, consisting of microstyles/raphides occurring
singly or arranged in trichodragmas. Sigmas are definitely present
Previous reviews but uncommon. Two species known, one from New Zealand, the
other from Indonesia.
Hallmann, 1917c, Topsent, 1928c.

Description of type species Description of type species

Dragmatella aberrans (Topsent, 1892a) (Fig. IF-G). Microtylostylifer anomalus Dendy, 1924 (Fig. 2A-G).
Synonymy. Desmacella aberrans Topsent, 1892a: 85, pI. II Synonymy. Microtylostylifer anomalus Dendy, 1924: 382,
fig. 7, pI. IX fig. 10; Dragmatella aberrans; Hallmann, 1917c: 640. pI. XV figs 46-49.
Material examined. Holotype: Presumably in MOM (not Material examined. Holotype: BMNH 1923.10.1.162 (wet),
seen). Other material. MNHN DT1187 (slide) - labelled 'PA st. 1923.10.1.162a (slides) - Three Kings Islands, New Zealand,
475' (not from type). 180m depth.
Description (from Topsent, 1892a, 1928c). Small hollow Description (modifiedfrom Dendy, 1924 and re-examination
sponge encrusting on stones or corals. No size quoted. Colour grey of the holotype). Small, massive sponge (fragment) of 1.5 cm
(preserved). Surface smooth, but provided with long thin, pointed diameter. Consistency crumbly but harsh. No apparent oscules.
fistules (Fig. IF). Skeleton of the ectosome a tangential crust of Colour in alcohol light yellowish grey. Ectosomal skeleton (Fig.
intercrossing styles. Choanosomal skeleton consisting of thick 2D) easily detachable, transparent, consisting of a reticulation of
spongin enforced bundles, mostly erect and occasionally branch- bundles of megascleres and single spicules, with large numbers of
ing; choanosome cavernous. Spicules (Fig. 1G) styles, somewhat microsubtylostyles scattered in between. Choanosomal skeleton
curved, well-rounded, very slightly fusiform, and sharply pointed, (Fig. 2E) an irregular reticulation of bundles of mycalostyles and
600-800 X 9-11.5 fLm; raphides in very thick trichodragmas, single spicules, with scattered but dense single microsubtylostyles.
70-200 X 12-20 fLm. Distribution. Azores, deep-water. Megascleres mycalostyles to styles, constricted below the rounded
Remarks. Topsent (1892a) originally considered this end (Fig. 2A, F). sharply pointed at the other end, up to 750 X
species to be a Desmacella. Hallman, 1917c assigned the new 27 fLm. Microsubtylostyles (Fig. 2B-C), of the same shape but
genus to his concept of the family Axinellidae near to Biemna, much smaller, and definitely classified as microscleres, uniformly
which was included in Axinellidae by him. Later, Topsent (1928c: 64 X 1.6 fLm. Sigmas (Fig. 2G) of several sizes are present
224) believed this was a reduced coelosphaerid, on account of the throughout the skeleton but are not common, and it is assumed that
fistular habit with parchment-like skin. Both views have merit, they are native to the sponge.
although neither proposal is well-founded. The combination of Remarks. Dendy (1924) assigned the genus to Suberitidae
styles and trichodragmata is not conclusive, but the fistular habit and compared it to Polymastiidae. However, there seems to be little
is also certainly polyphyletic. By assigning Dragmatella to in common with that group. Assignment of this genus to
Desmacellidae, we assume implicitly that sigmata have become Desmacellidae is based on the shape of the megascleres, which are
lost. The second, undescribed species from the SE Pacific is very clearly mycalostyles, and the reticulate choanosomal skeleton with
similar in spicule sizes and form. tangential ectosomal skeleton, which is shared with some Biemna
and Neofibularia. The peculiar microsubtylostyles in such large
quan-tities is a unique marker and the genus is considered valid
MICROTYLOSTYLIFER DENDY, 1924 with confidence.
The genus Hallmannia Burton, 1930c: 519 was erected for
Synonymy type species Biemna aruensis Hentschel, 1912: 352, pI. XV fig. 9,
pI. XIX fig. 22 (by original designation) from Indonesia (not:
Microtylostylifer Dendy, 1924: 381. Hallmannia Burton, [Hallmannia] Burton, 1931: 352, a junior homonym for type
1930c: 519. species [Hallmannia] spirophora Burton, 1931 = Rhabderemia
spirophora). The holotype, SMF 958 was re-examined. This sponge
Type species is similar to Microtylostylifer anomalus in having a (plumo)reticu-
late choanosomal skeleton (see Fig. 3A), and a complement of large
Microtylostyliferanomalus Dendy, 1924: 382 (by monotypy). subtylostyles (Fig. 3D), 400-720 X 12-15 fLm, and very much
smaller subtylostyles (Fig. 3B-C, E-F), 88-136 fLm (up to 3 fLm in
Definition thickness), many of which are arranged in tight bundles (Fig. 3C).
The latter are curved and were considered as homologous to com-
Desmacellidae with a small category of (subtylo-)styles con- mata by Hentschel, but this is highly debatable. As with M. anom-
centrated at the surface; microscleres are microstyles, microxeas or alus, we observed uncommon sigmas (Fig. 3G). Hallman (1916b:
raphides, uncommon sigmas. 501) considered this species to be a polymastiid, which in tum
induced Burton (1930c) to erect a separate genus Hallmannia for it.
Diagnosis However, Hentschel's assignment to Desmacellidae appears cor-
rect, although the homology of the small subtylostyles to commata
Massive but small sponges with smooth surface. Ectosomal seems far-fetched. Hallmannia aruensis conforms in all major
skeleton tangential, consisting of bundles of small subtylostyles, details to Microtylostylifer and accordingly it is a junior synonym.
Porifera· Demospongiae • Poecilosclerida • Mycalina • Desmacellidae 647

)(
B

I
c

I
A

Fig. 2. A-G, Microtylostylifer anomalus Dendy, 1924. A-C, drawings of spicules reproduced from Dendy's PI. V figs 46-49. A, large subtylostyles (scale:
100 f-Lm). B, small subtylostyles (scale see A). C, the same enlarged (scale: 10 f-Lm). D-G, photographs of sections and spicules made by J.N.A. Hooper.
D, ectosomal skeleton (scale: 100 f-Lm). E, perpendicular section (scale: 100 f-Lm). F, detail of head of subtylostyle (scale: 10 f-Lm). G, sigma (scale: 10 f-Lm).

Fig. 3. A-G, Microtylostylifer aruensis (Hentschel, 1912 as Biemna), type of Hallmannia Burton, 1930c, photos made by J.N.A. Hooper. A, cross section
showing reticulate choanosomal skeleton (scale: 500 f-Lm). B, surface 'bouquets' of microsubtylostyles (scale: 100 f-Lm). C, bundles of microsubtylostyles
(scale: SO f-Lm). D, large subtylostyles (scale: 100 f-Lm). E-F, microsubtylostyles (scale: 50 f-Lm). G, sigma (scale: 50 f-Lm).
648 Porifera· Demospongiae • Poecilosclerida • Mycalina • Desmacellidae

I
__
r=)
~==~~==~~_A======

--=-=-====-----=======
~---------------
~~------------
J

Fig. 4. A---{i, Neofibularia Hechtel, 1965. A,photo of holotype of Fibularia massa Carter, 1882a (scale 5 cm) (copied from Hartman, 1967: pI. I, fig. 2).
B, drawing of Amphimedon nolitangere Duchassaing & Michelotti, 1864 (copied from their plate XV, fig. 3). C, photo of type specimen of the latter,
Museum Torino nr. 42 (scale 5 cm) (copied from Hartman, 1967: pI. I, fig. I). D, drawing of spicules of holotype of Fibularia massa (scales: megasc1eres,
50 !Lm; microsc1eres, 10 !Lm) (copied from Hartman, 1967: fig. 2). E-F, drawings of cross section of choanosomal skeleton and tangential view of surface
of a Cura~ao specimen of Neofibularia nolitangere (scale 500 !Lm) (copied from Van Soest, 1984b: fig. 55). G, drawing of spicules of holotype of
Amphimedon nolitangere (scales: megascieres, 50 !Lm; microscieres, 10 !Lm) (copied from Hartman, 1967: fig . I). H-J, Sigmaxinella (for scales see text).
H, habits of S. australiana Dendy, 1897 (copied from Hallmann, 1916b, pI. XXXIII, figs 1- 3). I, cross section of a branch of the same (copied from
Hallmann, 1916b, p. XXXIV, fig. I). J, drawing of spicules of the same (copied from Hallmann, 1916b, text-fig. 12).

NEOFIBULARIA HECHTEL, 1965 Description of type species

Synonymy Neofibularia nolitangere (Duchassaing & Michelotti, 1864)

(Fig.4A-G).
Neofibularia Hechtel, 1965: 23; [Fibularia] Carter, 1882a: Synonymy. Amphimedon nolitangere Duchassaing &
282 (preocc. by Fibularia Lamarck, 1816, Echinodennata). Michelotti, 1864: 82, pI. XV, fig. 3; Fibulia nolitangere; de
Laubenfels, 1936a: 51; Neofibularia nolitangere; Hartman, 1967:
Type species 7, fig . 1, pI. I, fig. 1. Fibularia massa Carter, 1882a: 282; Gellius
massa; Arndt, 1927: 151, pI. II fig. 5; Neofibularia massa; Hechtel,
Fibularia massa Carter, 1882a: 282 (by subsequent designa- 1965: 23.
tion; de Laubenfels, 1936a: 54). This is generally considered Material examined. Holotype: Mus. Torino 42
a junior synonym of Amphimedon nolitangere Duchassaing & St. Thomas (photo in Hartman, 1967: pI. I, fig. 1, here reproduced
Michelotti, 1864: 82. in Fig. 4C). There are fragments (schizolectotypes) in MNHN
(D.NEE 1062) and BMNH (1928.11.12.34). The holotype of
Definition Fibularia massa (photo in Hartman, 1967: pI. I fig. 2, here repro-
duced in Fig. 4A), from Long Cay, Bahamas, is kept in the City of
Desmacellidae with diactinal megascleres, exceptionally sty- Liverpool Museums, according to Hartman (1967), under nr.
lote, and reticulate choanosomal skeleton; microscleres include CLMI08. An additional Liverpool Museum reg. nr. is LIVCM:ZI
sigmas, raphides, microxeas and commata. 69 (Mr Ian Wallace, pers. comm.). A slide of the type is kept in
BMNH (1939.3 .24.47) (Ms C. Valentine, pers. comm.).
Diagnosis Description. Massive to cake-shaped in smaller specimens,
developing an apical depression and growing out into groups of
Massive or tubular sponges with irregular surface. All species wide-vented tubes (Fig. 4A-C). Colour dark red or brown. Surface
are toxic sponges causing severe dennatitis. Irregular isodictyal optically smooth, hispid to the touch, occasionally shaggy. Size up
choanosomal skeleton. Megascleres diactinal (strongyles, oxeas) or to 30 cm high and wide. The ectosomal skeleton (Fig. 4F) may be
sometimes styles; microscleres sigmas, raphides, microxeas and traversed at regular intervals by vertical choanosomal spongin
commata. Four species have been described from tropical Atlantic fibres cored by strongyles, which may project beyond the sponge
and West Pacific locations. surface giving it a hispid appearance. Alternatively, a regular retic-
ulate pattern of flattened spongin fibres may occur at the surface.
Previous review Where this reticulation meets the tenninations of the vertical
choanosomal fibres, a few spicules may protrude through the ecto-
Hartman, 1967. some, but in these cases the sponge surface is rather smooth. Both
Porifera· Demospongiae • Poecilosclerida • Mycalina • Desmacellidae 649

types of surface may occur in a single specimen. The choanosomal About a dozen species have been described from South Australia,
skeleton (Fig. 4E) consists of an irregular reticulation of flattened S New Zealand and a few Indian Ocean locations.
spongin fibres cored by strongyles. Most strongyles are disposed in
rows parallel to the fibres, but some occur diagonally, piercing the Previous reviews
fibres in all directions. Microscleres occur throughout the sponge.
All types may be found at the surface, but sigmas are more abun- Hallmann, 1916b, Hooper, 1984b.
dant. Regularly arranged groups of microxeas do not occur near the
surface. All types of microscleres may be found associated with Description of type species
the surface of the spongin fibres. Oscules may open individually at
the surface, at times at the summit of an upright lobe of the sponge, Sigmaxinella australiana Dendy, 1897 (Fig. 4H-I).
or they may open into wide cloacas. Spicules (Fig. 4D,G). Sizes Synonymy. Sigmaxinella australiana Dendy, 1897: 240;
based on measurements of holotype of Fibularia massa provided Hallmann, 1916b: 535, pI. xxxm figs 1-3, pI. XXXIV fig. 1,
by Hartman (1967: 25): strongyles, often gently curved, seldom text-fig. 12.
straight, a few sinuous, 226-(316)-348 X 6.1-(7.2)-9.2 fLm. Material examined. BMNH 1902.10.18.408-409 (slides).
Microscleres, sigmas, with roughened ends, 18-(20)-22 fLm X Ho!otype and paratypes: NMV G2292 -fide Hooper (1984b: 32).
1.4-1.6 fLm; microxeas, faintly roughened, occurring singly or in Description (adaptedfrom Dendy, 1897 and Hooper, 1984b).
dragmas, 104-(112)--124 X 1.8-(2.0)-2.2 fLm thick; raphides, Erect, stipitate, ramose, bushy sponge. Branches cylindrical-
occurring singly or in loosely arranged goups, 82-(95)-110 X compressed, dichotomously branching in two planes. Size up to 18 cm
0.8-1.0 fLm; commata, 7 X 0.8 fLm. Distribution and ecology. This high, branches 3-8 mm in diameter. Oscules are shallow stelliform
is a widespread species in Caribbean reefs. depressions, 1-2 mm in diameter, arranged in rows along the
Remarks. The distinction of Neofibularia from Biemna is branches. Colour brownish red to rufous orange. Surface minutely
subtle (e.g., Hooper, 1996b). Hooper & Levi (1993a) and Hooper hispid, generally smooth. Choanosomal skeleton an axially con-
(1996b) suggested that species of Neofibularia and Biemna share densed reticulation of spicule tracts enclosed in visible spongin;
the dermatitis-production, and may be considered closely related extra-axial skeleton plumose ending in peripheral spicule brushes;
also for that reason. Currently, both genera are distinguished no special ectosomal skeleton. Mesh size 100-250 fLm, spicule
essentially by their skeletal arrangement. Neofibularia, known tracts 40-100 fLm in the axial region, with a core of 4-5
from only a few species, has a rather uniform skeletal architecture; spicules in cross section. Connecting fibers may be devoid of
while Biemna, on the contrary, is quite diverse in that respect, spicules. Megascleres styles, and some strongylote modifications,
ranging from halichondrioid-reticulate to hymedesmioid, with 120-450 X 2-17 fLm. Microscleres sigmas in two size categories,
sub-renieroid reticulate and plumose examples (Hooper, 1996b). 25-45 fLm and 9-16 fLm; microxeas singly or in trichodragmas,
The distinction of monactinal vs. diactinal megascleres appears to 20-45 fLm. Distribution. South Australia.
transgress the respective genus boundaries, with both genera show- Remarks. The importance of the axial condensation of the
ing species with either of those megascleres. skeletal architecture has been intensely debated lately (e.g., Hooper,
1984b; Bergquist & Fromont, 1988; Hooper et al., 1991; Van Soest,
1991). In spite of the currently widespread (although not unani-
mous) acceptance of the artificiality of taxa such as the Axinellida,
SIGMAXINELLA DENDY, 1897 the discussion has not settled down, but merely changed its level of
universality. The axial compression of the choanosomal skeleton is
Synonymy now seen as either a likely synapomorphy for several genera, or a
species-specific marker (e.g., Esperiopsis desmophora Hooper &
Sigmaxinella Dendy, 1897: 240. Sigmaxia Hallmann, 1916b. Levi, 1989; Mycale quadripartita Boury-Esnault, 1973). The recog-
nition of Sigmaxinella depends on the synapomorphic value of the
Type species axial compression of the skeleton in sponges which otherwise are
similar to Biemna. The possession of commata and microxeas in the
Sigmaxinella australiana Dendy, 1897: 240 (by subsequent latter may be regarded as further discriminating features.
designation; Hallmann, 1916b: 535). The genus Sigmaxia Hallmann (1916b: 535, pI. xxxm fig. 5,
pI. XXXVI figs 2-3) was erected for type species Axinella
Definition jlabellata Carter, 1885d: 361 (by original designation), types
BMNH 1886.12.15.471 (dry, here designated paralectotype),
Desmacellidae with axially compressed reticulate skeleton 1886.12.15.473 (wet, here designated lectotype). Dendy (1897:
and extra-axially plumose skeleton; spicules styles, sigmas and 241) assigned this species to Sigmaxinella, but Hallmann decided
microxeas. the skeleton was less clearly axially condensed and thus could not
fit into Sigmaxinella. Moreover, S. jlabellata has trichodragmata
Diagnosis as a further character distinguishing it from S. australiana. Hooper
(1984b: 6) maintained Sigmaxia as distinct from SigmaxineUa,
Arborescent or stipitate growth form. Cylindrical branches, but subsequently acknowledged that they are probably synonyms
rarely fused to form a flabellate sponge, with optically smooth (Hooper & Wiedenmayer, 1994: 154). As pointed out above, recent
surface. Condensed axial skeleton, plumose or plumo-reticulate discussions (e.g., Van Soest, 1991) resulted in a lower emphasis of
to plumose extra-axial skeleton, well-developed spongin fibres; axially condensed skeletons as a character for genus or family dis-
plumose, erect ectosomal skeleton; megascleres coring styles or tinction due to the obvious adaptive value of a reinforced axis
anisostyles of a single size; microscleres are sigmas and microxeas. in branching sponges. Raphides in trichodragmata are widespread
650 Porifera' Demospongiae • Poecilosclerida • Mycalina • Desmacellidae

in desmacellids and obviously not of generic value. The properties Mr Ian Wallace (CLM) provided information on the type
of Axinella flabellata largely overlap with Sigmaxinella and it is specimens of Fibularia massa. John N.A. Hooper provided
proposed to consider Sigmaxia a junior synonym. illustrations of Microtylostylifer. EH is further thankful to
CAPES, CNPq, FAPERJ and FAPESP, from Brazil, and the
ACKNOWLEDGEMENTS organizers of the International Conference on Sponge Science
(Otsu, Japan) for financial support to attend the 'Systema
Clare Valentine (BMNH) is thanked most sincerely for the Porifera' Workshops and to construct the means for writing this
provision of type specimens and/or other specimens on loan. revision.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademicIPlenum Publishers, New York, 2002

Family Guitarridae Dendy, 1924

Eduardo Hajdu l & Clea Lemer2

1 Museu Nacional, Departamento de Invertebrados, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, sIn, 20940-040, Rio de
Janeiro, RJ, Brazil & Centro de Biologia Marinha, Universidade de Sao Paulo, Sao Sebastiao, SP, Brazil. (hajdu@acd.ufrj.br)
2Museu de Ciencias Naturais, Funda9ao ZoobotfuIica do Rio Grande do SuI, Salvador Fran9a, 1427, 90690-000, Porto Alegre, RS,
Brazil & Instituto de Biociencias, Universidade de Sao Paulo, SP, Brazil. (cblemer@portoweb.com.br)

Guitarridae Dendy (Demospongiae, Poecilosclerida, Mycalina) includes species with typical mycalostyles and sigmancistras, lacking
tridentate chelae, and having restricted architectural specialization shown by their megascleres (e.g., absence of echinating megascleres).
Microscleres consist of placochelae-derivatives (placochelae, biplacochelae, dischelae and/or tetrapocillas), uniting genera irrespective of
their disparate skeletal arrangements. The family includes seven nominal genera of which four are valid. A key for the valid genera is given.
Keywords: Porifera; Demospongiae; Poecilosclerida; Mycalina; Guitarridae; Coelodischela; Euchelipluma; Guitarra; Tetrapocillon.
-----------------------------------------~---~-

DEFINITION, SCOPE clustering together all the taxa bearing the so-called placochelae-
derivatives (placochela, biplacochela, dischela and tetrapocilla;
Synonymy Van Soest, 1988; Hajdu et al., 1994a). These taxa were spread over
different families within the Poecilosclerida, on account of their
Guitarreae Dendy, 1924: 336. Not Gray, 1867a (erroneous diversified skeletal architectures and complement of megascleres
authorship attributed by Carballo & Uriz, 1998: 809). Not Burton, (Coelosphaeridae, Cladorhizidae, Desmacididae). Their classifica-
1929a (erroneous authorship attributed by Hajdu et al., 1994a: 126). tion in a single, possibly monophyletic family is presented here.
Van Soest (1988) recognized the shared affinities of Coelodischela,
Definition Guitarra and Tetrapocillon, to which Hajdu (1994) and Hajdu et al.
(1994a) added Euchelipluma. These sponges form a uniform
Mycalina with microscleres including placochelae-derivatives clade in terms of their shared possession of complex placochelae-
(placochelae, biplacochelae, dischelae and/or tetrapocillas). derived microscleres, irrespective of their disparate skeletal
arrangements (cf. Hajdu & Van Soest, 1996). Their classification
Taxonomic remarks within the Mycalina is supported on the grounds of their posses-
sion of typical mycalostyles and sigmancistras, their lack of triden-
Guitarridae Dendy, 1924 was informally proposed as a tate chelae, and on the restricted architectural specialization shown
Section within the Mycalinae (as Esperellinae; Dendy, 1924), orig- by their megascleres (e.g., absence of echinating megascleres, viz.,
inally comprising only Guitarra Carter, I 874a. Subsequent revi- acanthostyles). Taxonomic decisions taken herein follow the
sionary studies on the Poecilosclerida argued for the necessity of rationale outlined by Van Soest & Hajdu (Mycalina, this work).

KEY TO GENERA

(1) With placochelae ................................................................................................................................................................................. 2

With placochelae derivatives, but no true placochelae ........................................................................................................................ 3
(2) Erect, with sigmancistras ............................................................................................................................................... Euchelipluma
Other habits; palmate isochelae, when present, spiny; no sigmancistras ................................................................................ Guitarra
(3) With tetrapocilli ............................................................................................................................................................... Tetrapocillon
With dischelas ................................................................................................................................................................. Coelodischela

COELODISCHEIA VACELET, VASSEUR & LEVI, 1976 Definition

Synonymy Guitarridae with dischelae.

Coelodischela Vacelet, Vasseur & Levi, 1976: 57. Description of type species

Type species Coelodischela diatomorpha Vacelet, Vasseur & Levi, 1976

(Fig. 1).
Coelodischela diatomorpha Vacelet, Vasseur & Levi, 1976 Synonymy. Coelodischela diatomorpha Vacelet, Vasseur &
(by monotypy). Levi, 1976: 57, fig. 36, PI. VI, figs. d-e).

651
652 Porifera· Demospongiae • Poecilosclerida • Mycalina • Guitarridae

C D

Fig. 1. Coeiodischeia. A-K, C. diatomorpha Vacelet, Vasseur & Levi, 1976. A, holotype (scale I mm) (adapted from Vacelet et ai., 1976, Pl.VId). B-K,
drawing of the spicule complement (adapted from Vacelet et aI., 1976, fig. 36). B, tylote. C-D, strongyles. E-G, dischelae (scales 20 ILm). H-K, sigmas
(scale 50 !Lm) . L-N, C. massa Levi & Levi, 1983b. L, dische1a, paratype re-examined (MNHN D CL 2860), top oblique view. M-N, dischelae (adapted
from Van Soest, 1988, PI. 3, figs. 2, 4) (scales 20 ILm). M, side view. N, broken ala viewed from below.

Material examined: Holotype: Not seen. Comparative Levi & Levi, 1983b). Sigmas, typical or slightly tlagelliform,
material. Coelodischela massa Levi & Levi, 1983b: Paratype 17.5-110/-Lm wide by 0.5-2.5/-Lm thick. The larger ones (tlagelli-
MNHN 2860 - from Grand Recif, New Caledonia. form) have their endings suddenly bent 900 (Fig. IH-K).
Description (adapted from Vacelet et aI., 1976: 57). Small Remarks. Coelodischela is atypical within the family due
tubes attached directly to the substrate, 10 mm long, 1 mm diameter, to its possession of tylotes with well formed rounded tyles, as
with a few annular thickenings; the extremities are open and slightly are common within the Coelosphaeridae (sensu Hajdu et al.,
wider (Fig. lA). The walls of the tubes, rather thin, are reinforced 1994a; Van Soest, this volume). Furthermore, it possesses two cat-
by tangential strongyles, in disorder or sometimes arranged in loose egories of megascleres, which is paralleled only here-and-there
groups with 4-5 spicules, and by uncommon, dispersed tylotes. within the Mycalina, being a character relatively more widespread
Dischelae may be grouped irregularly. Megascleres: Tylotes, with in the Microcionina and Myxillina. Nevertheless, the denticulated
rounded tyles, 155-170/-Lm long and 4-5/-Lm thick, tyles 7.5 /-Lm inner-surface of Coelodischela's dischelas (cf. Levi & Levi, 1983b;
wide (Fig. IB). Strongyles, 140-160/-Lm long and 9-10/-Lm thick Hajdu et aI., 1994a) is undeniably akin to the microstructure
(Fig. lC-D). Microscleres: Dischelae, (placo)chelae-derived observed in the typical placochelae of Guitarra. In fact the
microscleres with two opposed disks bearing densely denticulated coelodiscs appear closer to a placochelae-derivative found in some
inner-surfaces, connected by two shafts on opposed sides of the Guitarra, the biplacochelae (e.g., G. abboti Lee, 1987; G. isabellae
spicules' borderline; both disks are slightly concave, the concavity Lee, 1987), differing only by the possession of a second fusion
being more conspicuous along the diameter which crosses both con- (shaft) between both disks, and of two symmetrical holes near
nection shafts; they are pierced centrally by two semicircles each. the central portion of each of both disks. The bridge separa-
Dimensions: 19.2-31.7/-Lm wide by 13.3-16.7 /-Lm high (calculated ting both holes is probably homologous to the falx of typical
from Vacelet et al.'s, 1976, fig. 36) (Fig. IE-G, L-N from C. massa chelae.
Porifera· Demospongiae • PoeciioscIerida • Mycalina • Guitarridae 653

C 0 E F G

Fig. 2. Euchelipluma pristina Topsent, 1909. A-B, holotype (10M 04 1041). A, bundle of subtylostyles witb projecting isochelae. B, tangential view of
bundle of subtylostyles showing projecting placochelae (scale 100 !Lm). C-N, drawing of the spicule complement (adapted from Topsent, 1909, PI. II,
fig. 6). C-D, subtylostyle of the main shaft. E-G, subtylostyles of the spines (scale 20 !Lm). H,subtylostyle of the peduncle. l-K, st(rong)yles of the base
(scale 100 !Lm). L, sigmancistras. M, isochela. N, placochela (scale 20 !Lm). O-P, SEM photographs, holotype re-examined (10M 04 1041).0, placochela
(scale 20 !Lm). P, sigmancistras-l and -II (scale to !Lm).

EUCHEllPLUMA TOPSENT, 1909 base, and divided in several parallel fibres upwards. The fibres are
twisted at the peduncle. Tufts of spicules are inserted on the spicular
Synonymy fibres at regular intervals, symetrically, giving thus origin to the
spines. Microscleres are situated around the shaft, the larger sigman-
Euchelipluma Topsent, 1909: 18. Desmatiderma Topsent, cistras standing perpendicularly. Chelae are also localized around
1928d: 308. the megasclere bundles (Fig. 2A-B), supposedly participating in
the capture of prey (i.e., camivory). In the widened portion of the
Type species shaft, the ascending spicular-fibres spread, surrounding yellow,
subspherical or ovoid, mostly compressed fleshy masses around
Euchelipluma pristina Topsent, 1909 (by monotypy). 200 fLm in diameter. Megascleres: Subtylostyles of the main shaft,
typically straight, slightly fusiform, with short obtuse points, and
Diagnosis bases which taper very gradually; tyles mostly inconspicuous,
heads as thick as necks, around 1000 fLm long, 30 fLm thick near
Guitarridae with narrow placochelas in face view, smooth the middle portion, and 13 fLm thick at the base (Fig. 2C-D).
palmate isochelae, sigmancistras and an erect habit coupled to Subtylostyles of the spines, straight, with a long inconspicuous
axially compressed architecture. tyle, sometimes polytylote, short apex, 400-600 fLm long, ca. 11
fLm thick (Fig. 2E-G). Subtylostyles of the peduncle, fusiform,
Description of type species curved, ca. 600 fLm long and 20 fLm thick (Fig. 2H). Styles of the
base, sometimes strongyles, flexuous, 240-370 fLm long, 10-14
Euchelipluma pristina Topsent, 1909 (Fig. 2). fLm thick (Fig. 2I-K). Microscleres: Palmate isochelae, slightly
Synonymy. Euchelipluma pristina Topsent, 1909: 19, PI. I, curved, 80-100 fLm long, shaft 6 fLm across, frontal alae 28 fLm
fig. 4, PI. 11, fig. 6. apart (Fig. 2M). Placochelae, with denticulated inner surfaces of
Material examined. Syntypes: MOM 04 1041 - 'Princesse the alae, 60-73 fLm long, shaft 6 fLm across, alae 28 fLm high and
Alice', stn. 1203 (15°54'N, 25°15'W). BMNH 1930.7.1.19 - same 15 fLm wide (Fig. 2N-O). Sigmancistras-I, 22-24 fLm long, gener-
locality (Prince of Monaco Collection, microscopy slide, ally contorted, with a conspicuous narrowing in the middle portion
schizosyntype ). of its inner fringe (Fig. 2L, P). Sigmancistras-II, ca. 12 fLm long
Description (adapted from Topsent, 1909: 19). Small (Fig. 2L,P).
sponges 14-22mm high, with a slight basal widening, a short, Remarks. The genus is clearly recognizable as the only gui-
naked, constricted peduncle, and a slightly widened and flattened tarrid with sigmancistras. There are two well characterized species,
shaft, omated with short paired spines. One specimen bears three E. pristina and E. arbuscula (Topsent, 1928d), and a third in need of
rows of spines, instead. The sponge is firm near the base, but flexible revised study, Euchelipluma congeri de Laubenfels, 1936a, as no
towards the apex; specimens are more-or-Iess flexuous. The skeleton illustration of skeletal architecture and/or of the spicule complement
consists of a spicular axis of styles (Fig. 2A), condensed near the was originally provided.
654 Porifera· Demospongiae • Poecilosclerida • Mycalina • Guitarridae

GUITARRA CARTER, 1874 Guitarra sigmatifera Topsent, 1916: ZMA 11540 - Antarctica, det.
R.W.M. Van Soest.
Synonymy Description (adapted from Carter 18740: 210, Pl. XIII,
figs. 2-5, Pl. Xv, fig. 34; Burton, 1929a: 427). Conical sponge
Guitarra Carter, 1874a: 210, PI. XIII, figs. 2-5, PI. XV, fig. 34. (13 mm high and 8 mm wide); hispid surface, with a perioscular
Hoplakithara Kirkpatrick, 1907a: 285. Pocilloguitarra Topsent, tubular spicular fringe. Large apical oscule, smaller ones scattered
1928c: 45. over the surface. Megascleres: oxeas, smooth, fusiform, nearly
straight, finely pointed, up to 375 J.Lm long and 7 J.Lm thick (average
Type species largest size) (Fig. 3A). Oxeas of the perioscular fringe up to 1042
J.Lm long. Microscleres: placochelae with denticulated inner sur-
Guitarrafimbriata Carter, 1874a (by monotypy). faces, a clear central trapezoidal area devoid of denticles occurs in
the inner face of the shaft, up to 67 J.Lm long, 25 J.Lm wide in its
Definition widest part, and 13 J.Lm wide in its central constricted portion (Fig.
3B-C). Burton (1929a) gives the following spicule dimensions
Guitarridae with placochelae, and palmate acanthoisochelae after restudy of the type material: 'Tornostrongyla' (=oxeas),
or bipocilla-like isochelae. 310 X 5 J.Lm; large placochelae, 90-100 J.Lm; small placochelae,
40-50 J.Lm; spiny isochelae, 10-11 J.Lm (Fig. 3D).
Description of type species Remarks. There has been considerable discussion on the
status of Guitarra fimbriata (Burton, 1929a; Lee, 1987; Boury-
Guitarrafimbriata Carter, 1874a (by monotypy) (Fig. 3). Esnault et al., 1993; Carballo & Uriz, 1998), but the following dis-
Synonymy. Guitarrafimbriata Carter, 1874a: 4. cussion will be concerned with higher taxa only. Starting with
Material examined. Holotype: BMNH 1954.3.9.319. Hoplakithara (type species H. dendyi Kirkpatrick, 1907a), erected
Comparative material. Guitarra fimbriata: MNRJ 1492 (fragment on account of its possession of exotyles. The type species is the
from Cabioch's Private Collection 336) - Roscoff, Brittany, only known species, and it is treated here as a Guitarra with
France, det. L. Cabioch. Guitarra abboti Lee, 1987: CASIZ exotyles instead. Pocilloguitarra was erected by Topsent, 1928c for
060483 (holotype) - stn. 821009, Cordell Bank, Marin County, G. bipocillifera Brjiindsted, 1924b, on account of its bipocilla-like
California. Guitarra indica Dendy, 1916: ZMA 1158 - Aru I., microscleres ('spiny isochelae with a characteristic spoon-like
'Siboga' Exp., Indonesia, det. M. Burton. ZMA 10440, 10578, shape' sensu Carballo & Uriz, 1998). It is clear from recent revi-
10719, 10815, 12549 - Seychelles, det. R.W.M. Van Soest. sions that a complete series of shapes exists from the so-called
spiny isochelae of G. laplani Boury-Esnault et al., 1993 to the
'bipocilla' of G. bipocillifera Brjiindsted, 1924b (Carballo & Uriz,
1998: fig. 5), so that the recognition of a higher taxon on the
basis of these microscleres would be premature. We consider here
A Pocilloguitarra as a junior synonym of Guitarra.

TETRAPOCILLON BR0NDSTED, 1924

Synonymy

Tetrapocillon Brjiindsted, 1924b: 456.

Type species

Tetrapocillon novaezelandiae Brjiindsted, 1924b

(by monotypy).

Definition

Guitarridae with tetrapocilla.

Description of type species

I
Fig. 3. Guitarra fimbriata Carter, 1874a. A, drawing of the oxea (adapted
Tetrapocillon novaezelandiae Brjiindsted, 1924b (Fig. 4).
Synonymy. Tetrapocillon
1924b: 457.
novaezelandiae

Material examined: Holotype: Not seen. Comparative

Brjiindsted,

from Carter, 1874a, PI. XV, fig. 34) (scale 50 /-Lm). B-D, SEM photographs
material. Tetrapocillon atlanticus Van Soest, 1988: ZMA 6226
of the spicule complement of the holotype (BMNH 1954.3.9.319 - adapted
from Boury-Esnault et aI., 1993: 369, fig. I). B, placochela. C, bottom view (holotype) - 'CANCAP VIT' Stn. 081, W of Boavista, Cape Verde
of broken ala (scales 10 /-Lm). D, spiny isochela (scale 5 /-Lm). Islands.
Porifera' Demospongiae • Poecilosclerida • Mycalina • Guitarridae 655

BCD

-
Fig. 4. Tetrapocillon novaezelandiae Br~ndsted, 1924b. A-H, J-K, drawing of the spicule complement (adapted from Br~ndsted , 1924b: 457, fig . 15).
A-D, mycalostyles. E-H, tetrapocilli. E, fac e view. F, side view. G, view from the back. H, young microsclere viewed from the back. I, SEMphotograph
oftetrapocilla (adapted from Bergquist & Fromont, 1988: 158, PI. 18b). l-K, drawing of palmate isochelae. J,face view. K, side view (scale 20f1m).

Description (adaptedfrom Brfindsted, 1924b: 457, Bergquist tetrapocilli Tetrapocillon would be classified close to genera such as
& Fromont, 1988: 46). Thinkly encrusting, ca. 3 mm thick, 30 X Batzella Topsent, 1893c and Strongylacidon Lendenfeld, 1897c, at
25 mm in area, consistency as felt, finely granular surface (vel- the time postulated to be close to the myxillids. The view of Hajdu
vety). Live colour is black outside, and bright orange-yellow (1994) and Hajdu et al. (1994a) is followed here. Tetrapocillon is
inside. Oscules in live specimens flush with the surface, 0.5-0.8 mm considered to be a true guitarrid, as foreseen by Van Soest (1988), but
long and 0.2-0.4 mm wide. They tend to disappear upon fixation. the latter taxon is better assigned to the Mycalina as argued above.
Dermal membrane packed with pigment cells. No special ectoso-
mal skeleton. Choanosomal skeleton mostly a spread of single
spicules, but a few tracts (ca. 60 fLm across) are seen, arranged in ACKNOWLEDGEMENTS
an irregular reticulation. Primary tracts branch at the surface, each
forming two or three spicule brushes. Tetrapocilli are abundant The authors are thankful to Dr John N.A. Hooper
throughout the sponge. Megascleres: (subtylo)styles (=mycalostyles), (Queensland Museum) and Dr Rob W.M. Van Soest (University of
straight or slightly undulating, generally slightly fusiform, apices Amsterdam), for the invitation to take part in the 'Systema
sharp or blunt, 260-325 fLm long and 10 fLm thick (holotype); and Porifera' project, and also for valuable discussions throughout the
230-330 fLm long and 5-8.5 fLm thick (other specimens from New preparation of this manuscript. Louis Caboich (SBR) Christien
Zealand) (Fig. 4A-D). Microscleres: tetrapocilli, with denticulated Carpine (MOM), Rob W.M. Van Soest (ZMA) and Clare Valentine
inner surfaces, 40-80 fLm long (holotype); 6--55 fLm long (other (BMNH) are acknowledged for sending specimens on loan. Welton
specimens from New Zealand) (Fig. 4E-I). (Spiny?) palmate L. Lee (Lee & Associates, California, U.S.A.) is thanked for allow-
isochelae, 15 fLm long (holotype) (Fig. 4J-K); 9-18 fLm long (other ing me access to his superb guitarrid SEM portfolio. Financial sup-
specimens from New Zealand). port to attend the 'Systema Porifera' Workshops and to construct
Remarks. Tetrapocillon is clearly recognizable by the pres- the means for writing this revision was obtained from CAPES,
ence of tetrapocilli, which are undisputable placochelae-derivatives. CNPq, FAPERJ, and FAPESP, from Brazil, and the organisers of
Van Soest (1988) pointed to the fact that was it not for their the International Conference on Sponge Science (Otsu, Japan).
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Esperiopsidae Hentschel, 1923

Rob W.M. Van Soestl & Eduardo Hajdu2

1 Zoological Museum, University of Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, Netherlands. (soest@science.uva.nl)
2 Museu Nacional, Departamento de Invertebrados, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, sIn, 20940-040, Rio de
Janeiro, RJ, Brazil & Centro de Biologia Marinha, Universidade de Sao Paulo, Sao Sebastiao, SP, Brazil. (hajdu@acd.ufrj.br)

Esperiopsidae Hentschel (Demospongiae, Poecilosclerida) is revived and limited to sponges with the combination of possession of
mycalostyles and lack of an ectosomal skeleton. The styles are united in tracts which form an isodictyal reticulation or an irregular anas-
tomosing system. Microscleres if present include palmate isochelae, exceptionally anisochelae, and sigmas. The possession of forceps is
reported but considered dubiously proper. The family is probably closest to Mycalidae, with which it is usually associated, and which
shares the palmate chelae and mycalostyles. However, most Mycalidae have an elaborate ectosomal skeleton and most chelae are
anisochelate. The family is predominantly recorded from temperate and colder waters.
Keywords: Porifera; Demospongiae; Poecilosclerida; Mycalina; Esperiopsidae; Amphilectus; Esperiopsis; Semisuberites; Ulosa.

DEFINITION, DIAGNOSIS, SCOPE with isochelae. He did not distinguish between different chela types
and included, amongst others, Phelloderma (Myxillina:
Synonymy Phellodennidae), Artemisina (Microcionina: Microcionidae) and
Chondrocladia (Mycalina: Cladorhizidae). Topsent (1928c) likewise
Esperiopsidae Hentschel, 1923. Amphilectidae de Laubenfels, did not use chela morphology as a character at the family level, but
1936a: 123. Semisuberitidae de Laubenfels, 1936a: 135. went further in combining Esperiopsis and Mycale and relatives
into a large subfamily Mycalinae. De Laubenfels (1936a) assigned
Esperiopsis to his huge family Ophlitaspongiidae (which contained a
Definition broad assemblage of poecilosclerid sponges), whereas Amphilectus
and Ulosa were put in a separate family Amphilectidae, (which
Mycalina with a choanosomal skeleton consisting of a reticu- again contained a broad assemblage of microcionine and mycaline
lation of tracts of styles, without a special ectosomal skeleton. genera). Semisuberites was assigned to a separate family
Microscleres if present palmate isochelae and sigmas. Semisuberitidae, which contained a remarkable assemblage of
genera such as Cladocroce (now Haplosclerida: Chalinidae),
Dragmatella (now Desmacellidae), Rhaphisia (now Chalinidae),
Diagnosis and Pachaxinella (a junior synonym of the suberitid Homaxinella).
His classification has no serious followers.
EnclUsting, massive, lobate, flabellate or cup-shaped sponges.
Surface finely conulose or microhispid due to projecting spicule
blUshes. Skeleton a reticulation of tracts of styles ending at the sur- Relationship with MycaJidae
face in blUshes or single spicules, no special ectosomal stlUctures.
Exclusively styles as megascleres although in one species there Prior to Hentschel's (1923) distinction of a separate family
are desmas of unknown derivation in addition to the styles. Esperiopsidae, Esperiopsis was usually assigned to a broader assem-
Microscleres palmate isochelae and sigmas, often absent or rare, blage including Mycalidae (e.g., Topsent, 1928c), and this tendency
but may be abundant in one genus. still finds followers, such as Bergquist & Fromont (1988) and Hajdu
et al. (1994a). It is clear that Esperiopsis and Amphilectus show sim-
ilarities in megasclere and microsclere shape with Mycale, and it is
Scope likely that Mycalidae and Esperiopsidae are sister families. A recent
cladistic analysis of the major Mycale subgenera and various
Four out of seven nominal genera are considered valid mem- Esperiopsis species demonstrated the monophyly of the Mycalidae,
bers of this family, Amphilectus, Esperiopsis, Semisuberites and but showed the Esperiopsis representatives were in a paraphyletic
Ulosa. Only Esperiopsis is widespread and diverse, the other gen- position. Nevertheless, the synapomorphic characters repsonsible for
era have only few species. Most representatives typically occur in this result may need to be studied further using a larger complement
high latitudes or deeper waters. of Esperiopsidae and additional characters. On a more practical
level, it is evident that a group comprising all the various Mycale
subgenera and Esperiopsis c.s. cannot be defined as a single family
History without also including Desmacellidae, for example. We prefer to
retain two separate families Mycalidae and Esperiopsidae, which
Hentschel's (1923) concept of Esperiopsidae was based on pos- clearly differ in the reticulate structure, lack of surface specialization
session of styles, lack of diactinal ectosomal spicules, in combination of the latter and absence of isochelae in the former.

656
Porifera· Demospongiae • Poecilosclerida • Mycalina • Esperiopsidae 657

Inclusion of Ulosa and Semisuberites Previous reviews

These genera lack microscleres and their exclusive possession of Hentschel (1923), Levi (1973), Van Soest (1984b), Hooper &
styles has made them subject of widely divergent family assignments Levi (1989), Hooper & Wiedenmayer (1994).
(see below). The esperiopsid nature of these small genera is tenta-
tive and defended on the basis of similarity of style shape and a
reticulate skeleton.

KEY TO GENERA

(I) Chelae and/or sigmas present .............................................................................................................................................................. 2

No microscleres ................................................................................................................. .................................................................. 3
(2) Skeleton of irregularly anastomosing spicule tracts, denser in the interior; styles usually over 400 /Lm; sigmas usually
present, chelae usually in several size categories ............................................................................................................... Esperiopsis
Skeleton regularly isodictyal, anisotropic; megascleres usually not exceeding 400 /Lm; only a single category of
isochelae or derivates; sigmas usually absent ................................................................................................................... Amphilectus
(3) Choanosomal skeleton consists of spongin encased polyspicular bundles making a rectangular reticulation;
styles in a single size category ..................................................................................................................................................... Ulosa
Choanosomal skeleton loosely reticulate, with many spicules in confusion; styles in two size categories ................... Semisuberites

AMPHILECTUS VOSMAER, 1880 priority of the genus name Amphilectus over Esperiopsis. The two
genera differ in the structure of the skeleton which is irregular, and
Synonymy more condensed in the interior in Esperiopsis, size of megasc1eres
which is usually over 400 /Lm in Esperiopsis, and the diversity of
[Corybas] Gray, 1867a: 537 (preocc.). Amphilectus Vosmaer, microsc1eres, which is usually greater in Esperiopsis.
1880: 109. Brondstedia Burton, 1929a: 428.
Description of type species
Type species
Amphilectus fucorum (Esper, 1794) (Fig. 1).
Isodictya gracilis Bowerbank, 1866: 331 (by subsequent des- Synonymy (restricted). Spongia fucorum Esper, 1794: 278,
ignation; Dendy, 1922b: 58). This is generally considered a junior pI. 49 figs 1-2; Halichondriafucorum; Johnston, 1842: 112, pis 9,
synonym of Amphilectus fucorum (Esper, 1794: 278). 12 fig. 2; Isodictyafucorum; Bowerbank, 1866: 322; Amphilectus
fucorum; Vosmaer, 1880: 117; Esperiopsis fucorum; Topsent,
Definition 1890a: 201. Spongia parasitica Montagu, 1818: 34. Isodictya gra-
cilis Bowerbank, 1866: 331; Bowerbank, I 874b: 149, pI. LVIII
Esperiopsidae with ladder-like skeleton of ascending and figs 23-26. Refer to Burton (1929a), Arndt (1935) and Van Soest
interconnecting spicule tracts; microsc1eres only small palmate et al. (2000) for suggested additional synonymy.
isochelae (occasionally slightly anisochelate), usually no sigmas. Material examined. Holotype of Isodictya gracilis (not
seen): BMNH 1930.7.3.357 (dry), and I slide BMNH
Diagnosis 1877.5.21.754 - labelled 'from type' (Hooper & Wiedenmayer,
1994 claim there are 3 syntypes but these were not found in a
Encrusting, cushion-shaped, lobate, occasionally branching recent search, Ms C. Valentine, pers. comm.). Other material. ZMA
sponges with velvety to finely hispid surface. Skeleton an isod- 4302,4400,4532,5925,5927,8795 - Ireland. ZMA 4723,4832,
ictyal or subisodictyal reticulation of short styles. Microscleres are 5913,9668,9703,14102, 14141-France. ZMA 6775,6796,6843-
small palmate isochelae or exceptionally anisochelae or sigmas. France. MNRJ 1166, 1184 - France.
There are at least two species, both North Atlantic, but several oth- Description. Extremely polymorphic and fast-growing,
ers may be hiding under the name Esperiopsis. changing shape in a few weeks. Encrusting as thin sheets or cush-
ions (Fig. lB), massive-Iobose or branched (Fig. lA, C). Size may
Remarks range from a few cm 2 to several dm 2, up to 15 em high. The oscules
may be only slightly raised from the surface, or be characteristi-
Despite widespread use of the name Esperiopsis for the cally at the top of volcano-like growths, and may have 'tassel'
species fucorum, Amphilectus is here recognized as a valid genus, growths originating near the margin of the oscules. The oscules do
in accordance with available phylogenetic evidence, which not contract. They are inconspicuous in encrusting forms, and large
assumes an independent origin for species allied to Esperiopsis or crateriform in massive forms; they are often arranged in linear
villosa Carter, 1874a and Amphilectus fucorum Esper, 1794 (Hajdu & series in the branching forms. Surface even, minutely hispid, fairly
Desqueyroux-Faundez, 1994), as Esperiopsis-I and -II, respec- open in appearance. It has a characteristic and unpleasant strong
tively. Previously, Bergquist & Fromont (1988: 25) distinguished smell, not dissimilar to that of Halichondria panicea. Consistency
sponges here assigned to Amphilectus as 'Esperiopsis sub-group soft and easily tom. Colour an intense reddish orange. Skeleton
two'. Authors who accept A. fucorum as the type of Amphilectus, irregularly reticulate. The main skeleton (Fig. ID) is an isodictyal
but did not consider it distinct from Esperiopsis overlooked the to sub-isodictyal reticulation of multi spicular fibres, reinforced
658 Porifera· Demospongiae • Poecilosclerida • Mycalina • Esperiopsidae

Fig. 1. Amphilectus fucorum (Esper, 1794). A, habit of ramose specimen (photo B. Picton, size see text). B, habit of encrusting specimen (photo R.W.M.
Van Soest, size see text). C, drawings ofho1otype lsodictya gracilis Bowerbank, 1866, reproduced from Bowerbank, 1874b: pI. LVIII figs. 23-26 (sizes see
text). D, drawing of skeleton of lsodictya hispida Bowerbank, 1882, considered a junior synonym, reproduced from Bowerbank, 1882: pI. XII fig. 3 (sizes
see text). E, SEM photo of style of Irish specimen ZMA 4832 (scale 10 ILm). F, SEM photo of palmate isochela of same (scale 10 ""m).

with variable amounts of spongin (depending on the age of the derived characters of each of the three mentioned genera would
sponge). There is no special ectosomal skeleton. Spicules smooth, have developed (Vosmaer, 1880). Vosmaer (1887) pointed to
usually curved, styles (Fig. lC, E), size variable: up to 150-400 X Microciona armata Bowerbank, 1866 as an example ('Beispiel')
3-19 flm. Microscleres small palmate isochelae (Fig. IF), which of what he intended to classify in Amphilectus (one of two species
may be rare, 14-28 flm. Reproduction August (Levi, 1956a). actually available to him, out of 42 species assigned to Amphilectus
Distribution and ecology. Atlantic coasts of Europe from Norway by Vosmaer). Levi (1960a) took this as a type species designation.
and the Faroes to the Mediterranean (latter record is by Driz, In spite of the fact that in other quoted genera the 'Beispiel'
1983); on rock and under stones; amongst hydroids (it will grow up species coincide with the type species as currently understood,
Tubularia stems) and has even been found on ascidian tests. It is we submit that Vosmaer did not formally make a type species
often found growing on/in Laminaria holdfasts on the lower shore designation. Accordingly, we take Dendy's (1922b), followed
and shallow sublittoral, down to 108 m. by Burton (1929a), as the objective designation, and recognize
Remarks. The species is polymorphic, both in habit and in Isodictya gracilis (=Amphilectus fucorum) as the type species
skeletal characters, and there are a large number of Bowerbank of Amphilectus. It is worth mentioning that Isodictya gracilis
synonyms (at least 16, see Van Soest et aZ., 2000) still to be (as Amphilectus g.) was the first species listed by Vosmaer (1880),
checked which may result in a 'splitting' of the entity A. fucorum as the remaining 41 lacking an alphabetical or a chronological order.
now understood. The genus [Corybas] Gray, 1867a: 537 was erected for type
AmphiZectus was erected by Vosmaer (1880: 109) for sponges species Isodictya Zobata sensu Bowerbank, 1866 (by original des-
which showed characters of dubious assignment to either of three ignation), which is based on Spongia Zobata Montagu (1818).
genera, viz., Desmacidon Bowerbank, 1862b; Mycale Gray, 1867a A type slide is extant in BMNH (unregistered, bearing number Bk.
and Myxilla Schmidt, 1862. Forty-two species were assigned to 952), labeled 'spicula from the Type with Dr. Grant. Devon
Amphilectus, believed to be the ancestral stock, from which the Coast' (not examined). Synonyms include Chalinula ovulum
Porifera· Demospongiae • Poecilosclerida • Mycalina • Esperiopsidae 659

Schmidt (1870: 38), type in ZMUC with slides in 2MB and ESPERIOPSIS CARTER, 1882
BMNH (not examined) and Esperia lanugo Schmidt (1875: 118),
no type material located (Desqueyroux-Faundez & Stone, 1992). Synonymy
Specimens ZMA 5388 (from the Netherlands) and ZMA 8453,
8454 (from Northern Ireland) were examined. This is a pale yellow, Esperiopsis Carter, 1882a: 296. Mycalopsis Topsent, 1927b: 7.
beige sponge forming round, egg-shaped, or flattened cushions
(Fig. 2A) growing on brown algae, hydroids or erect bryozoans, Type species
4-15 mm in diameter. Larger specimens are cylindrical or rather
irregular in shape; larger specimens may have oscules in rows Esperia villosa Carter, 1874a: 213 (by original designation).
along a ridge, or somewhat scattered. Consistency slightly elastic,
very soft. Skeleton of primary polyspicular bundles, cross-linked Definition
here and there by single spicules, running from the substratum to
the surface. Spongin is not obviously abundant, and is colourless. Esperiopsidae with anastomosing skeleton of thick spicule
Megascleres (Fig. 2C) more or less curved styles, 145-(190)-310 X tracts condensed in the interior and running upwards to the surface,
of 2-11 /-Lm. Microscleres (Fig. 2B) anisochelae variable in size, occasionally desma-like spicules may occur; microscleres chelae
from 15-23/-Lm in length. They also vary greatly in the relative and sigmas.
sizes of the chelate ends. In some specimens the difference
between the ends is so slight as to suggest isochelae. Distribution. Diagnosis
North Atlantic and North Sea, down to 200m. The species is very
close to A. jucorum but the anisochelate condition usually makes Massive, pedunculate or flabellate sponges with plumoreticu-
distinction easy. [Corybas] Gray is considered a synonym of late choanosomal skeleton, composed of anastomosing columns of
Amphilectus and would have taken priority if it had not been preoc- large subtylostyles (mycalostyles), condensed in the interior and
cupied by Corybas Boisduval in Westwood, 1850 (Lepidoptera). thinning out towards the surface. In species with desmas, these are
The genus Brondstedia Burton, 1929a: 428 was erected for concentrated in the interior forming an axial column (Hooper &
the New Zealand type species (by original designation) Esperiopsis Levi, 1989). Microscleres include up to three categories of palmate
glaber Brondsted, 1924a: 141, fig. 20. This was described as hav- isochelae, which may occur in rosettes, and up to two categories
ing styles of 200-370 /-Lm, palmate isochelae of 18 /-Lm and sigmas of sigmas. About 50 species have been recorded but these need
of 14-35 /-Lm. Brondsted also mentioned a thick tangential ectoso- revision. Most occur in higher latitudes or in deeper water.
mal crust and a reticulate choanosomal skeleton. Bergquist &
Fromont (1988: 26, pI. 6B-C) redescribed this material but were Previous reviews
unable to find any microscleres or the ectosomal skeleton; the
choanosome appeared to be confused with very thick spicule tracts. Bergquist & Fromont (1988), Hooper & Levi (1989), Hooper &
They returned the species to Esperiopsis, but in view of the spicule Wiedenmayer (1994).
size it seems better to consider it a species of Amphilectus,
although it is by no means typical for this genus. If the microscle- Description of type species
res described by Brondsted (1924a) are indeed foreign, is not
impossible that it will have to be assigned to Semisuberites. Esperiopsis villosa (Carter, 1874a) (Fig. 3A-B).
Synonymy. Esperia villosa Carter, 1874a: 213, pI. XIII
figs 13-15, pI. XV fig. 36; unknown sponge; Carter, 1879a: 502,
pI. XVlIa, figs 12a-c; Esperiopsis villosa; Carter, 1882a: 296.
Material examined. Holotype: BMNH 1882.7.28.55a -
'Porcupine' Exp. 'between the N coast of Scotland and the Faroe
Islands; det. H.J. Carter'; slide from holotype registered as BMNH
1954.2.12.158.
Description (adapted from Lundbeck, 1905; spicules and
architecture reevaluated from microscopic preparations of type).
Erect, more or less leaf-shaped (Fig. 3A-B). Surface finely
shaggy from protruding megascleres. Oscula mostly on top of
small conical projections situated on the upper edge of the
sponge. No specialized ectosomal skeleton. Dermal membrane
rests on diverging tufts of megascleres from terminating ascend-
ing choanosomal tracts of megascleres. Choanosomal skeleton
with ascending multispicular fibres which diverge and anasto-
mose along the way, slendering when approaching the surface.
Megascleres (Fig. 3A-B) are robust, slightly fusiform, smooth
mycalostyles, 560-(619)-670 X 1O-20/-Lm. Microscleres
(Fig. 3A-B) are palmate isochelae of three size categories: I,
Fig. 2. Amphilectus [obata (Montagu, 1818 as Spongia), type of [Corybas] 83-113 /-Lm, II, 50-83 /-Lm and III, 23-28 /-Lm. Categories I and II
Gray, 1867a. A, habit in situ (photo B. Picton, size see text). B, SEM image may overlap, but the conspicuous narrow face and straight profile
of palmate anisochela (scale 10 f,lm). C, detail of head of style (scale 1 f,lm). views of category I clearly set them apart. Sigmas are 50-193 /-Lm
660 Porifera' Demospongiae • Poecilosclerida • Mycalina • Esperiopsidae

Fig. 3. A-B, Esperiopsis villosa (Carter, 1874a as Esperia). A, drawing of habit and spicules reproduced from his pI. XIII figs 13-15, and pI. XV fig. 36.
B, drawing of habit and spicules reproduced from Lundbeck, 1905, pI. I fig. 4, and pI. VIII fig. l. C, Esperiopsis radiata (Topsent, 1927b as Mycaiopsis) ,
drawings of spicules and photo of skeleton reproduced from Topsent, 1928c: pI. VII figs 8-9.

long. Sigmas 50-100 f.Lm long are much more frequent (80%) the skeleton is reticulate rather than plumose, it is more delicate,
than larger ones (20%), but no clear gap exists in their dimen- the spicules are shorter, the diversity of microscleres is limited to
sions. Distribution and ecology. North Atlantic and Arctic Ocean, one small size category of chelae to which one size category of
also reported from Antarctic waters (Kirkpatrick, 1907a), sigmas may be added.
650-200m. The genus Mycalopsis Topsent, 1927b was erected for type
Remarks. Esperiopsis is employed here in the sense of species Mycalopsis radiata Topsent, 1927b: 7 (monotypy). See
Burton (1929a), for a restricted group of species conforming to also Topsent, 1928c: pI. vn figs 8-9, here reproduced as Fig. 3C.
Esperia villosa. These possess a plumo-reticulate skeleton of tracts A slide of the MOM holotype was examined, MNHN DT. 1336.
of large mycalostyles, and microscleres generally including more This is a blackish encrustation from 1331 m off the Azores. It has
than one category of isochelae and sigmas. These species were col- a skeleton 'similar to Carmia', which may be translated as having
lectively termed Esperiopsis-I in Hajdu & Desqueyroux-Faundez polyspicular bundles rising up to the surface and lacking an ectoso-
(1994) and Hajdu (1999). Further species included in this concept mal skeleton. Spicules (Fig. 3C) consist of straight, somewhat
of Esperiopsis are E. flagellum Lundbeck, 1905, and E. typichela polytylote styles of 385-455 X 7-8 f.Lm, and three size categories
Lundbeck, 1905, but a revision of original specimens is necessary of palmate isochelae, 80-95 f.Lm, 40-60 f.Lm and 18-20 f.Lm, of
to be certain of the genus membership. The genus Amphilectus is which the largest occur in rosettes. This combination of characters
closely related, as is demonstrated above. Many contemporary conforms to Esperiopsis as here defined and accordingly
authors consider Amphilectus to be a synonym, but the structure of Mycalopsis is considered a synonym.
Porifera' Demospongiae • Poecilosclerida • Mycalina • Esperiopsidae 661

SEMISUBERITES CARTER, 1877 shape are also included. Surface even, velvety. Oscules only on the
inside of tubes orin the depressed apex of branches (Fig. 4A, D, F, G).
Synonymy Consistency delicate, easily damaged. Ectosomal skeleton not
clearly separated from the choanosomal skeleton, consisting of
Semisuberites Carter, 1877a: 39. Stylaxia Topsent, 1913a: 52. brushes or loose palisades, which are the ends of loose choanosomal
Siphonocalypta Burton, 1931c: 140. tracts fanning out at the surface (Fig. 4C). Some small open spaces
are apparent subdermally between the ectosomal brushes. The
Type species choanosomal skeleton (Fig. 4C) consists of a system of loose longi-
tudinal spicule tracts parallel to the surface and tracts running to the
Semisuberites arctica Carter, 1877a: 39 (by monotypy). This surface at right angles to these longitudinal tracts; many single
is here considered a junior synonym of Semisuberites crib rosa spicules strewn in between. Spicules (Fig. 4C, E) exclusively styles
(Miklucho-Maclay, 1870: 6 as Veluspapolymorpha var. cribrosa). in what could perhaps be two size categories 400-500 and
200-300 /-Lm; many styles show a faint swollen head, and are similar
Definition to mycalostyles. Distribution and ecology. Arctic, down to 100m.
Remarks. This genus has been largely ignored by modem
Esperopsidae with skeleton consisting of loose brushes at the authors. Topsent (1928c: 37) assigned it to the Suberitidae, but most
surface and a system of loose tracts parallel to the surface inter- suberitids are compact and firm, with a high density of spicules,
crossed by polyspicular tracts ending in the surface brushes. which are also usually true tylostyles. Semisuberites cribrosa is a
Spicules styles with slightly swollen heads. No microscleres. fragile sponge with a loose, confusedly arranged skeleton. It shows
some similarities with Hymeniacidon in spicule size and shape, but
Diagnosis lacks the tangential surface skeleton of that genus. The synonymy of
Veluspa polymorpha var. cribrosa is based on Koltun's treatment of
Trumpet-shaped sponges with velvety outer surface, caused this species (as Phakettia cribrosa), no original specimens could be
by brushes of spicules ending at the surface; choanosomal skeleton studied. Still, there are no other matching records of Arctic species,
with spicules arranged in loose tracts perpendicular to the surface and the synonymy with Semisuberites arctica appears well-founded.
crossing long tracts running parallel to the surface, with many Burton (1932a: 196) also suggested the synonymy of Semisuberites
loose single spicules; megascleres exclusively styles in one or arctica and Miklucho-Maclay's species but confined himself to
more size categories, many with faintly swollen heads. A single adding a questionmark. The genus Veluspa Miklucho-Maclay is a syn-
species? onym of Haliclona (see chapter by De Weerdt on Haplosclerida:
Chalinidae).
Previous review The genus Stylaxia Topsent, 1913a: 52 was erected for
type species Cribrochalina variabilis Vosmaer, 1882a: 36, pI. I
Burton (1932a). figs 16-17 (here reproduced as Fig. 4D), pI. ill figs 67419 (here
reproduced as Fig. 4E), pI. IV figs 145-147 (by original designa-
Description of the type species tion). This is considered a junior synonym of Semisuberites
cribrosa. The type specimens of Cribrochalina variabilis (incl.
Semisuberites cribrosa (Miklucho-Maclay, 1870) (Figs 4-5). varieties crassa and salpingoides), RMNH 725-727, and the very
Synonymy. Veluspa polymorpha var. crib rosa Miklucho- similar Cribrochalina sluiteri Vosmaer, 1882a, RMNH 724, as well
Maclay 1870: 6, pI. I figs 12-16. Semisuberites arctica Carter, as 4 specimens of C. sluiteri subsequently collected in the same
1877a: 39; Burton, 1932a: 196, figs 1-2, pI. VII figs 1-2. area and kept in ZMA, were re-examined and they conform in all
Cribrochalina sluiteri Vosmaer, 1882a: 36; Vosmaer, 1885a: 22, details to Semisuberites crib rosa.
figs 10-11; Levinsen, 1887: 352, pI. XXIX figs 6-9, pI. XXX The genus Siphonocalypta Burton, 1931c: 140, fig. 1 (here
fig. 6; Stylaxia sluiteri Topsent, 1913a: 54, pI. ill fig. 1. Cribrochalina reproduced as Fig. 5B) was erected for type species (original des-
variabilis Vosmaer, 1882a: 36, pI. I figs 16-17, pI. ill figs 67419, ignation) Auletta elegans Vosmaer, 1882a: 40, pI. I fig. 20 (here
pI. IV figs 145-147 (incl. var. crassa and var. salpingoides); reproduced as Fig. 5A), pI. ill fig. 70, pI. IV fig. 152. The lectotype
Fristedt, 1887: 418, pI. 26 fig. 4; Stylaxia variabilis; Topsent, 1913a: ZMA 448, from northern Norway, collected by the Willem Barents
52. Auletta elegans Vosmaer, 1882a: 40, pI. I fig. 20, pI. ill fig. 70, Exped. 1881, was studied. It was originally a tube shaped sponge
pI. IV fig. 152; Siphonocalypta elegans; Burton, 1931c: 140, fig. 1. of 5 cm high, 1 cm diameter, which is now disintegrated. With the
Material examined. Lectotype of Semisuberites arctica help ofVosmaer's description, however, the properties of this spec-
Carter (in alcohol): BMNH 1877.7.3.1 - 'Smith's Sound, Cape imen can be reconstructed. Despite Burton's subsequent (1932a;
Napoleon, 50 fthm, presented by the Lords of the Treasury'. 197) defensive statements, this appears to be a clear junior syn-
Paralectotype (dry): BMNH 1877.7.3.5 - 'Smith Sound 50 fthm onym of Semisuberites arctica, combining a tubular habit and
Aug.1876, H.W. Jeilden, Lords of the Treasury'; including 3 sec- curved styles of 300-500 /-Lm, in two size categories, arranged in an
tions and 1 spicule mount. Holotype of Cribrochalina sluiteri irregular loose reticulation.
Vosmaer: RMNH 724 - Willem Barents Exped. 1878/79; ZMA
818, 819, Wi\lem Barents Expedition 1881. Lectotype of Auletta
elegans: ZMA 448 - Willem Barents Expedition 1881. ULOSA DE LAUBENFELS, 1936
Description. Trumpet-shaped (Fig. 4B, D, F), or bushes of
flaring tubes (Fig. 4A, G) or hollow branches (Fig. 4A, F), with a Synonymy
thin stalk, greatest height 14cm, widest expansion of individual
tubeslbranches 3 cm in diameter. If synonymy of Cribrochalina Ulosa de Laubenfels, 1936a: 126. Stylaxinella Vacelet,
sluiteri is accepted, then flabelliform sponges with widely expanded 1960a: 264.
662 Porifera· Demospongiae • Poecilosclerida • Mycalina • Esperiopsidae

Fig. 4. ~emisuberites cribrosa (Miklucho-Maciay, 1870). A, drawings of habit reproduced from Miklucho-Maciay, 1870: pI. I figs 12-13 (sizes see text).
B, photo of lectotype of Semisuberites arctica Carter, 1877a, BMNH 1877:7:3:1 (scale I cm). C,drawing of skeleton and spicules made from a slide of the
lectotype. D, drawings of habits of Cribrochalina variabilis, considered junior synonyms, reproduced from Vosmaer, 1882a: pI. I figs 16--17 (sizes see
text). E, styles of Cribrochalina variabilis and Aueletta elegans, both considered junior synonyms, reproduced from Vosmaer, 1882a: pI. III figs 69-70
(scale 50 j.Lm). F, drawing of habits of specimens recorded as Cribrochalina sluiteri by Levinsen, 1887: pI. XXIX figs 6--9 (sizes see text). G. drawing of
habit of specimen recorded as Cribrochalina variabilis by Fristedt, 1887: pI. 26 fig. 4 (size see text).

B
Fig. 5. A, drawing of Auletta elegans Vosmaer, 1885a, type of Siphonocalypta Burton, 1931c, considered a junior synonym of Semisuberites cribrosa,
reproduced from Vosmaer, 1882a: pI. I fig. 20. B, drawings of spicules, habit and skeleton of Siphonocalypta elegans Burton, 1931c, considered a junior
synonym of Semisuberites cribrosa, reproduced from Burton, 1931c: fig . 1(sizes see text).
Porifera· Demospongiae • Poecilosclerida • Mycalina • Esperiopsidae 663

Fig. 6. Ulosa spp. A, photo of detail of surface of MNHN lectotype of Ulosa angulosa (Lamarck, 1814 as Spongia) (size see text). B, water colour of
Ulosa stuposa (Esper, 1794 as Spongia), reproduced from his plate X (size see text). C, habit in situ of Roscoff specimen of Ulosa stuposa (photo MJ. de
Kluijver). D, drawings of habit, skeleton and spicules of Chalina inornata Bowerbank, 1874b considered a junior synonym of Ulosa stuposa, reproduced
from Bowerbank, 1874b: pI. LXXXIII figs 12-16 (sizes see text).

Type species Description of type species

Spongia angulosa Lamarck, 1814: 376 (by original designation). Ulosa angulosa (Lamarck, 1814) (Fig. 6).
Synonymy. Spongia angulosa var. B Lamarck, 1814; 376;
Definition Cacochalina angulosa; Topsent, 1930: 27, pI. III fig. 2; Ulosa
angulosa; de Laubenfels, 1936a: 126.
Esperiopsidae with a skeleton of poly spicular spicule fibres Material examined. Holotype: MNHN DT. 549 - from
encased in spongin making a rectangular or polygonal reticulation. King Island, S Australia. Other material. ZMA 3042 - Indonesia.
Megascleres styles of uniform size. No microscleres. Description (from Topsent, 1930). Lobate sponge
(Fig. 6A). The type specimen is bilobate, 8 cm high with lobes
Diagnosis of 3.5 cm in diameter and its surface appears macerated. There
are regularly distributed rounded holes, 1.5-2 mm in diameter,
Encrusting, lobate, and branching sponges with conulose sur- which are considered oscules by Topsent. Surface conulose due
face due to projecting fibers; no ectosomal skeleton. Choanosomal to projecting fibres. Skeleton a reticulation of primary fibers of
rectangular or polygonal reticulation of spongin encased spicule 55-80 fLm in diameter, interconnected by secondary fibres of
tracts. Megascleres short styles of a single size, occasionally modi- 30-60 fLm in diameter, forming rectangular meshes of 100-
fied to oxeas. Microscleres absent. Three or four species from 350 fLm, narrowing to only 30 fLm near the periphery. The
Atlantic-Mediterranean and South Pacific regions. primary fibres have a continuous core of aligned spicules with a
thin spongin border, the secondary fibres have fewer coring
Previous review spicules. Spicules are styles, somewhat curved, isodiametrical,
with well-rounded head and conical point, 125-180 X 5 fLm; a
Van Soest (1987). small minority of the spicules are oxeote, with the rounded end
664 Porifera· Demospongiae • PoeciioscIerida • Mycalina • Esperiopsidae

conically pointed. Distribution and ecology. King Island, N of Esper, 1794: pI. V (not Spongia stuposa Ellis & Sol ander, 1785:
Tasmania; ? Indonesia. 186 = Stelligera stuposa), with junior synonyms Chalina digitata
Remarks. The Indonesian material in ZMA answers in most Schmidt, 1866a, Chalina inornata Bowerbank, 1874b and
aspects to Topsent's redescription, but fibres are thicker and Desmacidon pannosus Bowerbank, 1874b. Esper's original speci-
meshes are wider, hence its assignment to this species with hesita- mens are considered lost, but his figure (here reproduced as
tion, including its disjunct distribution from the type locality. Fig. 6B) of this common European species is recognizable. It forms
This genus was associated with Stylinos Topsent, 1891a: 535 by digitate cushions to ramose forms, up to 8 cm thick, branches up to
some authors (e.g., Hooper & Wiedenmayer, 1994). This genus was 17 cm long and with a diameter up to 1.5 cm. Irregular in outline.
erected for type species lsodictya uniformis Bowerbank, 1866: 329 Surface finely conulose (Fig. 6C), in places honeycombed, some-
(by subsequent designation; Hooper & Wiedenmayer, 1994: 295). A what slimy. Consistency softly spongy, compressible, easily tom or
microscopical slide containing cross sections of the type specimen damaged. Colour pinky orange, salmon pink or yellowish in life,
labeled 'Isodictya uniformis Bk. from Type', BMNH 1877.5.21.751, pale yellow in alcohol. The organic ectosome is carried by the
bearing the number 'Bk.751' and with engraved number 'R2698, endings of choanosomal fibres, giving a conulose surface.
was re-examined. The skeleton is confused with vague spicule bun- Choanosomal skeleton is an irregular rectangular fibre system
dles without visible spongin running to the surface following a wavy (Fig. 6D) with relatively few soft parts and large open spaces; fre-
course. Many interstitial spicules in confusion. At the surface there is quently digitations and branches are partly hollow. Fibre reticulation
a crust of tangential and paratangential spicules intercrossing at all consisting of barely distinguishable main and secondary fibres,
angles. The spicules are smooth styles, 220-310 X 3-6/l-m, match- which connect at right angles. Fibres heavily cored, but spongin coat
ing those of Hymeniacidon perlevis as do the other features. All of normally well-developed. Meshes of the reticulum vary in size
these observations match the description by Bowerbank. Stylinos is between 100 and 1000 /l-m, fibre diameter in the different specimens
an obvious synonym of Hymeniacidon by virtue of the perhaps varies between 15 and 70/l-m, coring extent between 3 and 20
unfortunate type species designation by Hooper & Wiedenmayer. spicules per cross section. Spicules (Fig. 6D) predominantly styles,
Topsent, in subsequent publications (Topsent, 1892a: 135, 137) made but oxeote and strongylote modifications are common. Size rather
it clear he intended this genus to be a poecilosclerid, as he erected variable, especially the thickness, both within a single specimen and
Stylinos jullieni in 1892a: 137 as 'type' of Stylinos. However, since among different specimens, 115-260 by 2-10 /l-m. The species
S. jullieni was not among the species mentioned in Topsent's (1891a) occurs on rocks and surrounding algae in the deeper sublittoral,
publication, it cannot be the type of Stylinos. This species is very 15-55 m. Distribution. Ireland, S England, Atlantic coasts of France
probably a member what is now to be called Ulosa de Laubenfels, and Spain, Western Mediterranean, probably also North West Africa.
1936a, and consequently Hooper & Wiedenmayer attempted to sink The characters of this species match those of Ulosa angulosa and
Ulosa into synonymy of Stylinos. Hooper & Wiedenmayer (1994) thus Stylaxinella is a clear junior synonym of Ulosa.
were apparently unaware of the true nature of lsodictya uniformis Related forms (u. tenellula and U. tubulosa) have been
when they designated this the type of Stylinos. recently described from the Mediterranean by Pulitzer-Finali
Furthermore some doubt about the Ulosa nature of Stylinos (1983). All these appear to conform to the definition of Ulosa
jullieni was raised by Topsent himself (1928c: 14), because he based on Lamarck's type.
found spines on the styles and a dubious presence of tylotes. He Until recently, the genus name Ulosa was used for quite
suggested that S. jullieni is close to Kirkpatrickia (now in different sponges in recent West Indian literature. Particularly,
Hymedesmiidae), but left it incertae sedis. the extremely common reef form 'Ulosa' ruetzleri Wiedenmayer
A further complication arose when Vacelet, 1960a: 264 - (1977b), is widely divergent from U. angulosa and U. stuposa.
accepting that Stylinos sensu Topsent, 1892a, 1928c could not be 'Ulosa' ruetzleri is an incrusting sponge, fleshy-conulose, with a
what Topsent originally intended, and apparently unaware of the high content of organic material and a very loose system of
properties of Ulosa - erected a replacement genus Stylaxinella. dendritic spongin-fibres cored by rather few, long, sharply pointed
He chose as type species (original designation) Spongia stuposa styles. It conforms to Scopalina (Halichondrida: Dictyonellidae).
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Hamacanthidae Gray, 1872

Eduardo Hajdu

Museu Nacional, Departamento de Invertebrados, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, sin, 20940-040,
Rio de Janeiro, RJ, Brazil & Centro de Biologia Marinha, Universidade de Sao Paulo, Sao Sebastiao, SP, Brazil. (hajdu@acd.ufrj.br)

Hamacanthidae Gray (Demospongiae, Poecilosclerida, Mycalina), including Vomerulinae Ridley & Dendy, includes eight nominal
genera of which only two genera and two subgenera are considered valid. The family is defined by its possession of mycalostyles and
diancistra derivatives (diancistras and cyrtancistras), c-shaped, flat or contorted, and placed near MycaJidae by their lack of tridentate
chelae and restricted architectural specialization (e.g., lack of acanthostyles).
Keywords: Porifera; Demospongiae; Poecilosclerida; Mycalina; Hamacanthidae; Hamacantha; Hamacantha (Hamacantha); Hamacantha
(Vomerula); Pozziella.

DEFINITION, DIAGNOSIS, SCOPE which cannot be entirely ruled out on the basis of available morpho-
logical evidence, would classify Hamacantha Gray, 1867a and
Synonymy Merlia Kirkpatrick, 1908c (Merliidae) closer together, because of
the striking similarity between diancistras and clavidiscs (e.g.,
Hamacanthidae Gray, 1872a: 449. Vomerulinae Ridley & Topsent, 1928c; Wiedenmayer, 1994; Lehnert & Van Soest, 1998).
Dendy, 1886: 337. Despite our fragmentary knowledge of poriferan biodiversity during
geological time, the diagnostic spicules that bear on the issues
Definition raised here are known from the paleontological literature.
Diancistras and clavidiscs are known since the lower Jurassic
Mycalina with diancistra-derivatives (diancistras or (Mostler, 1990), while anisochelae are only confidently known
cyrtancistras) which are always c-shaped, flat or contorted. from the lower Cretaceous (Gruber & Reitner, 1991; Wiedenmayer,
1994), which supports the second hypothesis outlined above. It has
Remarks been considered here though, that affinities among every sigmancis-
tra derivative (sigmancistras, diancistras, cyrtancistras and cla-
The hamacanthids are a well-characterized group of about 25 vidiscs) are probably plesiomorphic. Accordingly, the classification
species distributed worldwide. The classification of Hamacanthidae proposed reflects instead, the undeniable similarities between
within the Mycalina is considered well corroborated by its posses- Hamacantha and Mycale Gray, 1867a (Mycalidae; easily detach-
sion of mycalostyles and likely sigmancistra derivatives (diancistras able reticulated ectosomal skeletons, plumo-reticulated choanoso-
and cyrtancistras), their lack of tridentate chelae, and on the mal construction, microscleres in rosettes, sigmas disposed
restricted architectural specialization produced by their megascleres alongside megasclere bundles with hooks pointing outward), and
(e.g., absence of echinating acanthostyles; Hajdu, 1994). On the those of Merlia (Merliidae) and Biemna Gray, 1867a1Desmacelia
other hand, their closer affinity to the Mycalidae still warrants fur- Schmidt, 1870 (Desmacellidae; tylostyles, commas). Taxonomic
ther investigation, perhaps through the assessment of complimen- decisions made here follow the rationale proposed by Hajdu (1999,
tary data sets (e.g., cytology, DNA, etc.). A rivalling arrangement, and present work, see Mycalina).

KEY TO GENERA AND SUBGENERA

(1) With diancistras .................................................................................................................................................................................. 2

With cyrtancistras ................................................................................................................................................................... Pozziella
(2) With oxeas .............................................................................................................................................. Hamacantha (Hamacantha)
With styles ................................................................................................................................................... Hamacantha (Vomerula)

HAMACANTHA GRAY, 1867 Definition

Synonymy Hamacanthidae with diancistras.

Refer to subgenera. Description of type species

Type species See nominotypical subgenus.

Hymedesmia johnsoni Bowerbank, 1864 (mistakenly emended

to Halichondriajohnsoniby Gray, 1867a; ICZN 67d) (by monotypy).

665
666 Porifera' Demospongiae • Poecilosclerida • Mycalina • Hamacanthidae

B c

c
Fig. 1. A, Hamacantha (Hamacantha) johnsoni (Bowerbank, 1864), drawing of ectosomal architecture (scale 500 11m) and larger category of diancistra
(scale IOOl1m) (reproduced from Bowerbank, 1864, figs 112, 293). B, drawing of the spicule complement of Hamacantha (Vomerula) popana (de
Laubenfels, 1935a) (reproduced from Hajdu, 1994, fig. I) (scale 100 11m). C, drawing of the spicule complement of Pozziella clavisaepta Topsent, 1896b
(reproduced from Topsent, 1896b, fig. I) (scale 100 11m).

Remarks Synonymy. Hymedesmia johnsoni Bowerbank, 1864: 35,

127, figs 112, 293. Halichondria johnsoni Gray, 1867a (objective
Part of the abundant synonymy of Ha11U1cantha may be attrib- replacement name for Bowerbank's (1864), Hymedesmiajohnsoni).
uted to the conflicting interpretations historically put forward for Ha11U1cantha complanata Topsent, 1892: 86, pI. II, fig. 9, pI. VII,
the diagnostic characters of its type species. The situation has been fig. 5f (as H. johnsoni var. complanata). Ha11U1cantha
made clear by Stephens (1921), who, after re-examining a type schmidti (Carter, 1882a) (=Desmacella johnsoni Schmidt, 1870
slide, established that the spiculation of H. johnsoni consisted of from Portugal). Not D. johnsoni Schmidt, 1870, from Florida
oxeas, two categories of diancistras and sigmas. (=H. schmidti).
Material examined. Holotype: Not seen. Comparative mate-
rial examined: Ha11U1cantha schmidti (Carter, 1882a): BMNH
SUBGENUS HAMACANTHA GRAY, 1867 1870.5.3.28 (holotype) - Florida (277 m depth). Hamacantha sim-
plex Burton, 1959a: BMNH 1936.3.4.339a and 1936.3.4.412a
Synonymy (holotype) - 'Murray' Exp. (stn. 157, Maldive area; det. M. Burton).
Description (adapted from Ridley & Dendy, 1887; Topsent,
Hamacantha Gray, 1867a: 538. Athnacama de Laubenfels, 1904b; Stephens, 1921). Deep-water sponge, encrusting or
1936a: 52. variably-thick cushion-like, with easily detachable, neatly reticu-
lated surface peel (Fig. lA). Megascleres are oxeas (frequently
Type species st(rong)yloid), ca. 450-600/-Lm long by 5-10 /-Lm thick.
Microscleres are diancistras in two size classes, 100-160 /-Lm and
Hymedesmiajohnsoni Bowerbank, 1864 (by monotypy). ca. 30-35 /-Lm long; and sigmas 20-25 /-Lm long.
Remarks. Hamacantha (Hamacantha) is established here
Definition for those species with diactinal megascleres, which are customarily
seggregated from others with monactines in sponge classification.
Hamacantha with diactinal megascleres. Athnacama de Laubenfels (type species Ha11U1cantha lundbecki
Topsent, 1904b), was erected on the basis of its possession of oxeas
Description of type species as sole megascleres, thus being a clear junior synonym of
Ha11U1cantha (Ha11U1cantha). De Laubenfels (1936a) considered
Hamacantha (Ha11U1cantha) johnsoni (Bowerbank, 1864) Ha11U1cantha to be a likely synonym of Vomerula or perhaps even
(Fig.lA). unrecognizable, which explains his establishment of Athnacama.
Porifera· Demospongiae • Poecilosclerida • Mycalina • Hamacanthidae 667

SUBGENUS VOMERULA SCHMIDT, 1880 other hamacanthid has yet been found with similar megascleres,
suggests a more likely interpretation of the character as a likely
Synonymy apomorphy instead, and Zygherpe is maintained in the synonymy
of Hamacantha in agreement with an earlier proposal (Hajdu,
Vomerula Schmidt, 1880b: 82. Zygherpe de Laubenfels, 1994). Hypsispongia was based on the study of a contaminated
1932: 65. Hypsispongia de Laubenfels, 1935a: 5 (in part). specimen. Hajdu (1994) has shown that its type species was a typi-
Crellancistra de Laubenfels, 1936a: 112. Evomerula de cal (valid) species of Hamacantha, contaminated by abundant
Laubenfels, 1936a: 125. anisochelae of Mycale bellabellensis (Lambe, 1905). Crellancistra
(type species Hamacantha integra Topsent, 1904b), was established
Type species for its simple exotyles (although described as acanthostyles by
de Laubenfels, 1936a) and diancistras with open-hooks (forming
Vomerula tenda Schmidt, 1880b (by subsequent designation, a more obtuse angle with the shaft). These features are understood
present work, following recommendations by Ridley & Dendy, as apomorphies here, and the genus is transferred into synonymy
1887 that it was a 'typical species of the genus'). with Hamacantha. Evomerula (type species Hamacantha agassizi
Topsent, 1920a), was established on the basis of its association of
Definition styles to diancistras and sigmas. This genus is clearly synonymous
with Hamacantha (Vomerula).
Hamacantha with monactinal megascleres.

Description of type species POZZIELLA TOPSENT, 1896

Hamacantha (Vomerula) tenda (Schmidt, 1880b) (Fig. lB). Synonymy

Synonymy. Vomerula tenda Schmidt, 1880b: 82.
Material examined. Holotype: Not seen. Comparative mate- Pozziella Topsent, 1896b: 147.
rial examined. Hamacantha esperioides (Ridley & Dendy, 1886):
BMNH 1887.5.2.157ii (syntype) - 'Challenger' Exp. (stn. 142, Type species
Agulhas Bank, SW Atlantic). MNRJ 1194 (fragment from CEAB
6B-92) - 'Benguela VII' Exp. (P-56, Namibia; det. MJ. Uriz). Pozziella clavisaepta Topsent, 1896b (by monotypy).
Hamacanthafalcula (Bowerbank, 1874): MNRJ 870 (fragment from
ZMA 4944) - Vaagegrund (Bergen, Norway; det. R.W.M. Van Diagnosis
Soest). Hamacantha hyaloderma (de Laubenfels, 1932): USNM
22060 (holotype) - California (U.S.A.). Hamacantha mindanaensis Hamacanthidae with cyrtancistras.
Wilson, 1925: BMNH 1936.3.4.186 - 'Murray' Exp. (stn. 152,
Indian Ocean; det. M. Burton). Hamacantha popana (de Laubenfels, Description of type species
1935a): AMNH 262 (holotype) - Lower California (Mexico).
Description (adapted from Topsent, 1920a). Choanosome Pozziella clavisaepta Topsent, 1896b (Fig. lC).
reduced (cavernous) to some skeletal tracts on which the dermal Synonymy. Pozziella clavisaepta Topsent, 1896b: 147, fig. 1.
membrane rests. There are two layers of spicules in this membrane. Material examined. None.
The superficial one is made of tangential styles, densely distrib- Description (adapted from Topsent, 1896b). Encrusting
uted, criss-crossed in some parts, arranged in bundles in others, but sponge with easily detachable dermal membrane and cavernous
mostly directed toward a central papilla, and keeping this arrange- choanosome (strongly reminiscent of Hamacantha johnsoni).
ment until the apex of the papilla. The inner layer is less dense and Spicules (Fig. IC): Megascleres are styles, 470-500 t-Lm long
more irregular. The criss-crossing of styles establishes some nodes, (strongly resembling Hamacantha falcula - Topsent (1928c: 198)
which coincide with the termination of the ascending choanosomal remarked that his 1904 record of H. johnsoni was to be considered
tracts. The papilla of two of the specimens possesses an areolated H. falcula instead), and exotyles, ca. 210 fLm long, with a styloid
pore field just below its apex. There are rosettes of up to 30 dian- termination on one side, and an acanthotylote on the other
cistras around the choanosomal bundles of megascleres. Spicules. (15-20 t-Lm thick). The exotyles stand on the ectosome with their
Megascleres: Styles, straight or slightly bent, slightly fusiform, acanthotylote terminations projecting away from the sponge,
with a slightly prominent head (mycalostyles ?), apex short and forming thus a characteristic crust. Microscleres are cyrtancistras
obtuse, 450--760 fLm long and 11-18 fLm thick. Microscleres: (very large diancistras without most of their inner fringe/edge),
Diancistras, 250--265 fLm long. Toxas, acanthose, with large, ca. 450 fLm long, occuring in loose dragmas alongside choanoso-
thin spines, implanted variably perpendicularly, sparsely spread mal bundles of megascleres, and variably shaped (from regular to
over the length of the spicule; absent from the terminations; flagelliform) and sized sigmas, 25-120 fLm long.
56--103 fLm long. Remarks. Topsent (1920a) described Hamacantha clav-
Remarks. Figure 1B shows the spicule complement isaepta var. aperta from the West Indies. This is clearly a Pozziella,
of Hamacantha (Vomerula) popana de Laubenfels, 1935a. and is considered here a valid species, Pozziella aperta, on account
Hamacantha (Vomerula) is established here for those species with of diagnostic morphological features and likely biogeographical
monactinal megascleres, which are customarily seggregated from isolation. It differs from the type species by the possession of a
others with diactines in the sponge classification. Zygherpe was second category of much smaller cyrtancistras (larger ones up
erected on the basis of its possession of (sub)ty10styles. This char- to 473 fLm, smaller ones up to 167 t-Lm). Pozziella is retained as a
acter alone is seen as a weak diagnostic feature. The fact that no valid genus on the basis of its rather peculiar set of spicules, thus
668 Porifera' Demospongiae • Poecilosclerida • MycaJina • Hamacanthidae

opposing Topsent's own change of mind (Topsent, 1904b). Wiedenmayer (1994) identified a lower Cretaceous fossil spicule
Cyrtancistras are undisputably related to diancistras, and diancis- drawn by RUst (1885) as a possible cyrtancistra. He was already
tras such as the smaller ones in Hamacantha acerata Levi, 1993 hesitating in this respect, a suspicion shared here, so that no asser-
could be seen as an evolutionary stepping-stone (=character trans- tion can still be made confidently on the paleontological record of
formation series) from typical forms, as in H. johnsoni, to the cyr- Pozziella.
tancistras, where most of the inner edge is reduced. Hajdu (1994)
postulated a character transformation series to explain the relation
among the sigmancistra derivatives. Cyrtancistras are also conspic- ACKNOWLEDGEMENTS
uous by their very large length (over 400 f.Lm), the largest diancis-
tras known to date being those of H. megancistra Pulitzer-Finali, The author is grateful to Dr John N.A. Hooper (Queensland
1978, which are under 300 f.Lm in length. Known fossil diancistras Museum) and Dr Rob W.M. Van Soest (University of Amsterdam),
also are smaller than 300 f.Lm (e.g., Wiedenmayer, 1994). The for the invitation to take part on the 'Systema Porifera' project, and
exotyles of Pozziella also set it apart as they do not appear closer to also for valuable discussions throughout the preparation of this man-
those of H. integra (=Crellancistra, sensu de Laubenfels, 1936a), uscript. Klaus RUtzler and Kate Smith (USNM), Rob W.M. Van Soest
than they appear to those of clear mycalids such as Mycale (ZMA), Clare Valentine (BMNH), G.w. Thurmann and J.E. Winston
(Rhaphidoteca) ItUlrshallhalli Kent, 1870a (originally as (AMNH) are thanked for sending specimens on loan. Financial
Rhaphidoteca). It is quite possible that Pozziella would sit at a basal support to attend the 'Systema Porifera' Workshops and to construct
position within the hamacanthid clade, although no formal phyloge- the means for writing this revision was obtained from CAPES,
netic analysis has yet been attempted, an outcome which would not CNPq, FAPERJ, and FAPESP, from Brazil, and the organisers of the
imply the acceptance of either genus as non-monophyletic. International Conference on Sponge Science (Otsu, Japan).
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N,A, Hooper and Rob WM. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Mycalidae Lundbeck, 1905

Rob W.M. Van SoestI & Ednardo Hajdu2

1 Zoological Museum, University of Amsterdam, P.O. Box 94766,1090 GT Amsterdam, Netherlands. (soest@science.uva.nJ)
2Museu Nacional, Departamento de Invertebrados, Universidade do Brasil, Quinta da Boa Vista, sIn, 20940-040, Rio de Janeiro, RJ,
Brazil & Centro de Biologia Marinha, Universidade de Sao Paulo, Sao Sebastiao, SP, Brazil. (hajdu@acd.ufrj.br)

Mycalidae Lundbeck (Demospongiae: Poecilosclerida) is restricted to taxa with the combination of palmate anisochelae and a tangential
surface skeleton. It consists of two genera, Mycale s.l. and Phlyctaenopora, which includes Barbozia. The large genus Mycale is subdi-
vided in 11 subgenera, viz., Aegogropila, Anomomycale, Arenochalina, Carmia, Grapelia, Mycale, Naviculina, Oxymycale, Paresperella,
Rhaphidotheca and Zygomycale. These subgenera are probably artificial and some 'overlap' in characters, but they may serve to differen-
tiate the 150 or more species assigned to Mycale. The genera Raphioderma, Rhaphiodesma, Protoesperia and Mycalecarmia are consid-
ered junior synonyms of Mycale (Mycale). The genus Esperia and its replacement name Esperella, as well as Pellinula, are considered
junior synonyms of subgenus Aegogropila. The genera Acamas, Acamasina, Kerasemna and Parisociella are junior synonyms of
Arenochalina. The genus Oxycarmia is considered a junior synonym of Carmia. The genus Pseudoesperia is considered a junior synonym
of Grapelia. The genera Gomphostegia and Sceptrospongia are junior synonyms of Rhaphidotheca. Keys to the genera and subgenera are
provided.
Keywords: Porifera; Demospongiae; Poecilosclerida; Mycalina; Mycalidae; Mycale (Aegogropila); Mycale (Anomomycale); Mycale
(Arenochalina); Mycale (Carmia); Mycale (Grapelia); Mycale (Mycale); Mycale (Naviculina); Mycale (Oxymycale); Mycale
(Paresperella); Mycale (Rhaphidotheca); Mycale (Zygomycale); Phlyctaenopora (Phlyctaenopora); Phlyctaenopora (Barbozia).
- - - - - - - - - - - - - - - - - - - - - -

DEFINITION, DIAGNOSIS, SCOPE Scope

Synonymy Two genera comprising 13 subgenera out of 25 nominal (sub-)

genera are recognized as valid, Mycale Gray, 1867a: 533 and
Esperiadae Gray, 1867a: 531. Esperellinae Ridley & Dendy, Phlyctaenopora Topsent, 1904b: 198. The first is subdivided in
1887: 62. Arenochalininae Lendenfeld, 1888. Mycalinae eleven subgenera, the second in two subgenera.
Lundbeck,1905.
Remarks
Dermition
Nomenclatorial remarks, The name Mycalidae and author-
Mycalina with 'mycalostyles' (styles with slightly constricted ship of Lundbeck is to be maintained over the older names
neck), occasionally modified to oxeas or strongyles, arranged in Esperiadae, Esperellinae and Arenochalininae because the syn-
a plumose, plumoreticulate or confused manner; microscleres onymy of Esperia, Esperella and Arenochalina with Mycale was
include palmate chelae, sigmas (may be serrated), toxas (may established before 1961 and Lundbeck's replacement of the family
be spined), spined microxeas, and raphides, all of which may name was likewise made before 1961. Mycalidae has been used
be absent; no diancistras or derivates, no commata, no spined hundreds of times in many different publications by many different
micro(subtylo-)styles. authors, and thus satisfies the condition of ICZN article 40.2 as
'prevailing usage' (Anon., 1999).
Diagnosis History and biology. Gray (1867a: 531) used the name
Esperiadae for a Jarge group of genera covering more-or-Jess what
Encrusting, massive, fan-shaped and branching growth forms. is now considered the order Poecilosclerida. Ridley & Dendy, 1887:
Subectosomal sculpturing, grooves and ridges often found on the 62 assigned to their Esperellinae all poecilosclerids lacking echinat-
surface, within which are usually found the ostia. Skeleton ing acanthostyles, thus more-or-less covering what is now under-
plumose or plumoreticulate, composed of styles or rarely oxeas stood as the suborder Mycalina. However, they also included
enclosed in spongin fibres or without visible spongin. Ectosomal Iophon (now Microcionina), and Desmacidon and Phelloderma
tangential skeleton usually present. Ectosomal spicules may differ (now Myxillina). Lendenfeld's 'group' Arenochalininae was
from the choanosomal ones by smaller size. Megascleres most assigned to a larger subfamily Chalininae, and he was obviously
often subtylostyles (mycalostyles), but (aniso)strongyles, oxeas unaware of the mycalid nature of Arenochalina. Lundbeck (1905: 7)
and exotyles occur occasionally. Microscleres characteristically erected Mycalinae to replace Esperellinae and included not only
palmate anisochelae, but also palmate isochelae, sigmas, toxas, Ridley & Dendy's concept, but also most Myxillina; however, he
spined microxeas and raphides may occur. The family is quite subdivided the group into Mycaleae and Myxilleae, which covered
diverse with about two hundred species described occurring in what is now understood as Mycalina and Myxillina. Topsent (1928:
all parts of the world oceans, and possibly many more hundreds 46) returued to Ridley & Dendy's concept, but continued the use
awaiting discovery. of the name Mycalinae. De Laubenfels (1936a: 112) assigned Mycale

669
670 Porifera· Demospongiae • Poecilosclerida • Mycalina • Mycalidae

and its relatives to a family Ophlitaspongiidae, defined as having the of a restricted Mycalidae may be considered monophyletic based on
fibres echinated by smooth rather than acanthose styles. Mycale was the shared possession of palmate anisochelae. However, one species
defined by him (p. 116) as having 'fibers cored and echinated by of the genus Amphilectus, viz., A. lobata (Bowerbank, 1866), here
smooth monaxons that are frequently subtylostyles'. He assigned assigned to the family Esperiopsidae, also possesses palmate
a wide range of unrelated genera in Ophlitaspongiidae which are anisochelae. Further character analysis is necessary to arrive at
now spread over three suborders. Levi (1973: 609) was the first to a more robust phylogeny of the families of Mycalina.
restrict the contents of the family to Mycale and its immediate rela- Phlyctaenopora, although not usually included in this group,
tives, all sharing the palmate anisochelae. Many authors followed shares anisochelae and a tangential surface crust with Mycale,
his proposal. Bergquist & Fromont (1988: 17) expanded Levi's con- which conforms to the present definition of the family. It differs in
cept with the inclusion of Esperiopsis, which is likely to be closely having strongyles and oxeas as megascleres instead of the usual
related, but has palmate isochelae. Their proposal was followed by mycalostyles. Barbozia has similar structure and spiculation, but in
Hajdu & Desqueyroux-Faundez (1994). They assigned a number of addition possesses peculiar spined microxeas. It is here considered
incertae sedis genera to the family (e.g., Ulosa, Phelloderma), but a subgenus of Phlyctaenopora.
pointed out that a monophyletic Mycalidae was not established Mycalidae are numerous and common members in all seas
convincingly. If the mycaline families are indeed related as suggested and all marine habitats. Reproduction is often seasonal. The larvae
by Hajdu & Desqueyroux-Faundez (1994), viz., «Cladorhizidae, have a bare posterior pole and diverse larval spiculation. Gemmule-
Guitarridae) (Desmacellidae (Hamacanthidae, Mycalidae»), like resting stages consisting of undifferentiated globular masses of
Mycalidae s.l. (which for them included the family here separated archaeocytes are reported from Mycale species from temperate
off as Esperiopsidae) could share the possession of chelae, suppos- waters.
edly lost two nodes down the tree, by the ancestor of the
Desmacellidae-Hamacanthidae-Mycalidae s.l. clade. Another possi-
ble synapomorphy, although underlying, is the presence of three Previous reviews
categories of chelae. However, these characters do not appear to
establish the monophyly very strongly. Further discussions on the Hallmann (1914), Topsent (1924), de Laubenfels (1936a),
phylogenetic relationships of Mycalidae and Esperiopsidae genera Van Soest (1984b), Bergquist & Fromont (1988), Hajdu &
and subgenera may be found in Hajdu (1999). The present concept Desqueyroux-Faundez (1994), Hajdu (1999).

KEY TO GENERA

(1) Only mycalostyles present ........................................................................................................................................................ Mycale

Megascleres oxeas and/or strongyles .................................................................................................................................................. 2
(2) Megascleres exclusively oxeas ............................................................................................................................ Mycale (Oxymycale)
Megascleres oxeas and strongyles ............................................................................................................................... Phlyctaenopora

MYCALE GRAY, 1867 Definition

Synonymy Mycalidae with megascleres in a single category of shape.

There may be size categories.
[Esperia] Nardo, 1833: 522 (preocc.) (not Esperia HUbner,
1816: Lepidoptera). [A camas] Duchassaing & Michelotti, 1864: 94 Diagnosis
(preocc.) (not Acamas Montfort, 1808, Mollusca). Mycale Gray,
1867a: 533. Aegogropila Gray, 1867a: 533. Grapelia Gray, 1867a: Encrusting, lobate, repent, ramose, branching erect or tubular
534. Carmia Gray, 1867a: 537. Naviculina Gray, 1867a: 538. sponges. Surface irregular or smooth, occasionally grooved.
Esperella Vosmaer, 1887: 353 (misprinted as [Esperiella] on p. 348 Consistency fibrous, compressible or soft, occasionally firm or
(lapsus». Raphioderma Bowerbank in Norman, 1869b: 333. hard. Skeleton of discrete bundles of spicules arranged in plumose
Rhaphidotheca Saville Kent, 1870a: 219. Raphiodesma fashion, with occasional anastomosing, rarely reticulate. Ectosomal
Bowerbank, 1874b: 235. Pseudoesperia Carter, 1886c: 455. skeleton usually a well-developed tangential crust of intercrossing
Arenochalina Lendenfeld, 1887c: 821. Protoesperia Czemiavsky, bundles or single megascleres, occasionally absent or consisting
1879: 248. Gomphostegia Topsent, 1896b: 149. Paresperella of the brushed endings of choanosomal tracts. Spicules usually
Dendy, 1905: 162. Anomomycale Topsent, 1924: 116. exclusively mycalostyles, rarely replaced by oxeas; size categories
Sceptrospongia Dendy, 1926: 6. Oxymycale Hentschel, 1929: 932. are common. Microscleres palmate anisochelae (exceptionally
Zygomycale Topsent, 1930: 431. Acamasina de Laubenfels, 1936a: isochelae), the larger ones frequently in rosettes, sigmas, toxas,
117. Mycalecarmia de Laubenfels, 1936a: 120. Parisociella raphides; rarely micracanthoxeas which appear to be toxa-derived;
Burton, 1952: 169. Oxycarmia de Laubenfels, 1954: 155. size categories common for anisochelae (up to four have been
Kerasemna Pulitzer-Finali, 1981: 105. recorded) and sigmas, occasionally also trichodragmas and toxas.

Type species Scope

Hymeniacidon lingua Bowerbank, 1866: 187 (by subsequent The genus comprises some two hundred species, subdivided
designation; Thiele, 1903a: 949). among eleven subgenera, Mycale (Mycale), Mycale (Aegogropila),
Porifera· Demospongiae • Poecilosclerida • Mycalina • Mycalidae 671

Fig. 1. Skeletal structures of Mycale subgenera. A-B, Mycale (Mycale), cross section (A) and tangential section (B) to show ectosomal crust of inter-
crossing single spicules. C-D, Mycale (Aegogropila), ditto. E, Mycale (Carmia), cross section. F, Mycale (Arenochalina), cross section (all redrawn from
Van Soest, 1984b: fig. I).

Mycale (Anomomycale), Mycale (Arenochalina), Mycale (Carmia), of Aegogropila. Bergquist & Fromont (1988: 7) went further and
Mycale (Grapelia), Mycale (Naviculina), Mycale (Oxymycale), proposed to raise Topsent's subgenera to genera. Hajdu (1995,
Mycale (Paresperella), Mycale (Rhaphidotheca), Mycale 1999) revived some additional (sub)genera, e.g., Grapelia and
(Zygomycale). Overall distribution cosmopolitan. Naviculina. Hajdu (1995, 1999) and Carballo & Hajdu (1998)
pointed out that characters of several of the (sub-)genera over-
Remarks lapped with other (sub-genera), e.g., Paresperella and Zygomycale
have a typical Aegogropila structure (Fig. lC-D), Grapelia,
The idea that several, relatively homogeneous groups can be Anomomycale and Oxymycale have a typical Mycale s.s. structure
recognised within the genus Mycale is a concept as old as the genus (Fig. lA-B). Thus, several of these taxa appear to be paraphyletic
itself, as Gray (1867a) erected five different genera for sponges or overlapping, as they are based on different features of the skele-
sharing a comparable set of spicules (viz. Mycale, Aegogropila, ton and spicules. It is here decided to keep the species groups pos-
Grapelia, Carmia and Corybas) and distinguished from each other sessing a single megasclere type assigned to Mycalidae, all at the
in minor morphological features only (cf. Hajdu & Desqueyroux- level of subgenera of Mycale instead of genera. In this way we
Faundez, 1994). Hallmann (1914) stressed the need for a division avoid some of the paraphyly problems, but it is certainly not ideal
of the large genus, into subgenera Aegogropila, Paresperella, and must be considered a provisional, convenient system to man-
Grapelia, Arenochalina and Mycale. He was doubtful about age large groups of taxa. When hypotheses of phylogenetic rela-
Protoesperia. Topsent (1924) followed his example and elaborated tionships for these taxa, attempted by Carballo & Hajdu (1998) and
and extended the subgenera, recogmzmg Aegogropila, Hajdu (1999) (amongst others) become more robust, it may turn
Anomomycale, Carmia and Mycale. However, he restricted his pro- out to be preferable to have several more genera in Mycalidae (see
posals to species occurring in Western Europe, and thus avoided further comments in the remarks for each subgenus).
genera like Paresperella, Arenochalina and Rhaphidotheca. These
proposals found few contemporaneous followers, but they were Previous reviews
revived by Van Soest (1984b: 10), who constructed diagrammatic
drawings of the major subgeneric skeletal features (here repro- Van Soest (1984b), Bergquist & Fromont (1988), Hajdu &
duced in Fig. lA-F). He also added Acamasina = Arenochalina to Desqueyroux-Faundez (1994), Hajdu (1995), Hajdu & Riitzler
the list of subgenera, but assigned Zygomycale to the synonymy (1998).

Key to subgenera of Mycale

(1) Megascleres exclusively oxeas (Fig. lOB) ..................................................................................................................................... Oxymycale

Megascleres include exotyles (Figs 12-14), styles with swollen or proliferated ends) piercing the ectosome, next to
normal mycalostyles ............................................................................................................ ........................................Rhaphidotheca
Megascleres exclusively subtylostyles (mycalostyles) ........................................................................................................................ 2
(2) Microscleres include naviculichelae (Figs 8-9: complete or near fusion of both frontal alae, falx markedly expanded
along the shaft, lateral alae of the head project backward and upward) ............................................................................. Naviculina
672 Porifera· Demospongiae • PoeciioscIerida • Mycalina • Mycalidae

Sigmas are serrated (Fig. 11) ............................................................................................................................................ Paresperella

Microscleres include anomochelae (Fig. 4: palmate anisochelae with a slightly curved shaft on face view; and frontal ala of the foot
considerably expanded, approaching or even surpassing the dimensions of the usually larger frontal ala ofthe head, contorted,
sometimes with a serrated upper border) ....................................................................................................................... Anomomycale
Microscleres include small isochelae (Fig. 15B) ............................................................................................................... Zygomycale
Microscleres include polydentate unguiferate chelae (Fig. 7); rosettes in two size categories ............................................... Grapelia
Microscleres consist of normal shaped sigmas and palmate anisochelae; toxas, raphides and micracanthoxeas may be present .......... 3
(3) Ectosomal tangential skeleton present ................................................................................................................................................. 4
Coherent ectosomal skeleton lacking ................................................................................................................................................... 5
(4) Ectosomal skeleton consisting of intercrossing bundles of megascleres making triangular or polygonal
meshes (Fig. ID) ............................................................................................................................................................... Aegogropila
Ectosomal skeleton consisting of a felted mass of intercrossing megascleres without clear meshes or bundles (Fig. IB) ...... Mycale
(5) Choanosomal skeleton consisting of thick spongin enforced spicule tracts arranged in a rectangular reticulation; fibres cored by
foreign objects (such as algae or sand); sponges extremely slimy when lifted out of the water (Fig. IF) ..................... Arenochalina
Choanosomal skeleton consisting of wispy plumose bundles of megascleres showing little or no cohesion; sponges usually soft and
encrusting (Fig. IE) ............................ ....................................................................................................................................... Carmia

SUBGENUS MYCALE GRAY, 1867 described as being 'roundish', 'resembling the tongue of a sheep',
or 'an inverted triangular pyramid'. It is apparent that some speci-
Synonymy mens tend to grow in an erect, flattened form, with the attached
base narrower than the apex. Grows up to 30 cm in height, with
Mycale Gray, 1867a: 533. Raphiodenna Bowerbank in Norman, variable width and depth. Surface sulcate (Fig. 2A), the furrows
1869b: 333. Raphiodesma Bowerbank, 1874b: 235. ? Protoesperia forming an irregular reticulation over the surface. Within the fur-
Czemiavsky, 1879: 248. Mycalecarmia de Laubenfels, 1936a: 120. rows, the dermal membrane is smooth, elsewhere it is rather
shaggy in appearance. Colour grey, pale buff; in alcohol pale yel-
Type species low, whitish yellow or gray. Oscules are borne on shallow, trans-
parent cones of height 2-3 mm, and have a diameter of 4-10 mm.
Hymeniacidon lingua Bowerbank, 1864: 48 (by subsequent The inhalant pores lie along the surface of the furrows, and can
designation; Thiele, 1903a: 949). only be seen with the aid of a microscope in preserved material
but are obvious in living specimens. The sulcate cracks close
Definition on collection. Consistency rather tough, very fibrous, but easily
compressed. Ectosomal skeleton a mass of tangential spicules.
Mycale with a confused tangential ectosomal skeleton (many Choanosomal skeleton plumoreticulate, consisting of ascending
with pore-grooves, and/or three categories of anisochelae, and/or multispicular fibres of styles, with small amounts of spongin bind-
basally-spurred anisochelae-III, and/or raphides in two categories). ing the fibres. Towards the surface, some fibres run parallel to
the surface, and some become splayed like a brush and penetrate
Description of type species the surface. Megascleres are styles (Fig. 2B-C), usually straight,
occasionally slightly curved: 460-1150 X 13-20 J.1m. Their ends
Mycale (Mycale) lingua (Bowerbank, 1866) (Fig. 2A-C). are variable in shape. The head may be that of a typical style, or
Synonymy. Hymeniacidon lingua Bowerbank, 1864: 48, may tend to be that of a subtylostyle, or may be constricted like a
pI. VI fig. 147; Hymeniacidon lingua Bowerbank, 1866: 187; handle ('mycalostyles'). The points may be abrupt or long.
Mycale lingua; Gray, 1867a: 533; Raphiodesma lingua; Microscleres (Fig. 2B-C) are palmate anisochelae, sigmata and
Bowerbank, 1874b: 235, pI. XLVII fig. 8, pI. LXXVII figs 1-6; trichodragmas. The anisochelae fall into two distinct size classes,
Mycale (Mycale) lingua; Topsent, 1924: 86, fig. 1. Desmacidon 35 J.1m and 70 J.1m, the larger ones being found in rosettes, mainly
constrictus Bowerbank, 1866: 350; Bowerbank, 1874b, 181, near the dermal membrane; sigmas, 16-32 J.1m; trichodragmas,
pI. LXXI figs 3-10. Raphioderma coacervata Bowerbank in 42-78 X 11-14 J.1m. Distribution and ecology. Widely distributed
Norman, 1869b. Esperia lucifera Schmidt, 1873: 148. Esperia pla- over the north Atlantic, from the Azores to Spitzbergen. Also
coides Carter, 1876: 316; Mycale placoides; Stephens, 1912: 34. known from the Mediterranean; found at depths between 30 and
Esperella vosmaeri Levinsen, 1887: 20. Esperella massa; Arnesen, 2460m.
1903: 9, pI. I fig. 5, pI. IV fig. 2 (not massa Schmidt, 1862). Remarks. Gray (1867a: 533) misquotes Bowerbank's type
Material examined: Lectotype (here designated): BMNH species as Hymeniacidon lingula.
1930.7.3.235 - label in Bowerbank's handwriting reads The genus Raphioderma Bowerbank in Norman, 1869b: 333
'Raphiodesma lingua/Loch Fyne. McNab. Bwk Coll. Vol. 2. p. 187, was erected for type species (monotypy) Raphioderma coacervata
type of Genus & species 1852'. (Loch Fyne is on the Kintyre penin- Bowerbank in Norman, 1869b. No material of this sponge
sula, Argyll, Scotland). Paralectotype: BMNH 1877.5.21.2019 - was examined, but the species and the genus name were ignored
Shetland; also registered as BMNH 1930.7.3.233, dry; as by Bowerbank (1874b), and assigned to the synonymy of
Raphiodesma. Other material. BMNH 1882.7.28.98 - syntype of Hymeniacidon lingua in Bowerbank (1882: 162). It is assumed to
Esperia placoides Carter, 1876, 'Porcupine' Exped., Shetland. have been a mistake, later to be replaced by Raphiodesma lingua
Description (based on Ackers et al., 1992). Encrusting to in Bowerbank (1874b: 237). If that is correct, then technically
massive lobose (Fig. 2A-B), to massive fig-shaped. Variously speaking, Raphioderma is a senior synonym of Raphiodesma.
Porifera· Demospongiae • Poecilosclerida • Mycalina • Mycalidae 673

(CC

Fig. 2. Mycale (Mycale) lingua (Bowerbank, 1864). A, In situ photo from Ireland showing sulcate grooves (photo B. Picton). B, habit and spicules of type
specimen, reproduced from Bowerbank, 1874b: pI. LXXVII (size see text). C, spicules reproduced from Lundbeck, 1905: pI. IX fig. 6 (sizes see text).

The genus Raphiodesma Bowerbank. 1874b: 235 (earlier also 1887.5.2.160 and additional material. This is a massive sponge
introduced on p. vi) was erected for type species Hymeniacidion from Southern Argentina, with a skeletal arrangement consisting of
lingua (subsequent designation by Hallmann, 1914: 399). Since thick spicule bundles ending at the surface in dense ectosomal
this is also the type species of Mycale, the genus Raphiodesma is tufts. This is not a typical Mycale (Mycale) ectosome, which
an objective junior synonym of Mycale (Mycale).1t is also likely to has tangentially intercrossing spicules, but is sufficiently different
be a junior synonym of Raphioderma, and thus by inference, that from the ectosome of typical Carmia, that assignment to Mycale
genus is also a junior synonym of Mycale (Mycale). (Mycale) is justified. The spicules include a single category oflong
The genus Protoesperia Czerniavsky, 1879: 248 was erected robust mycalostyles, 500-900 X 15-20l1m, three size categories
for type species (designation herein) Protoesperia simplex of anisochelae, 75-95I1m, 45-70 11m, and 30-45 11m, and tri-
Czerniavsky, 1879: 249, pI. II fig. 13, pI. III fig. 18, from the Black chodragmas 45-88 11m; some of the studied specimens also con-
Sea. The type was stated to be located in the museum of the tained sigmas, but these were lacking in the holotype.
University of Charkov, Ukrain, but is now presumably lost (Koltun, De Laubenfels (1936a: 116) restricted Mycale to species hav-
in litt.). This is a possible synonym of Mycale (Mycale), but the ing anisochelae and sigmas as microscleres, differentiated from
description is ambiguous and the thickly encrusting species has not other genera like Carmia in the lack of toxas. This simplified and
been rediscovered. The megascleres are said to consist of three cat- artificial restriction found only few followers.
egories of subtylostyles (400 X 4.5 11m, 420-480 X 14 11m, and The most recent phylogenetic analysis of the genus Mycale by
270-500 X 3-18 11m, which differ only in the shape of the malfor- Hajdu (1999) found the confused ectosomal architecture to be
mations) and a predominance of oxeas of 135-578 X 7-16 11m. synapomorphic for a cluster of species-groups, among which is
The author also recorded the presence of anisochelae. It is Mycale (Mycale). Proper classification principles demand unique
here assumed that the oxeas are not proper. This species needs characters for taxa, and these appear to be lacking for Mycale
redescription. (Mycale). However, strict application of these principles would
The genus Mycalecarmia de Laubenfels, 1936a: 120 was have led to the inclusion within Mycale (Mycale) of easily recog-
erected for type species (by original designation) Esperella nizable subgenera as Grapelia and Rhaphidoteca. We prefer to
lapidiformis Ridley & Dendy (1886: 338). Hajdu & Desqueyroux- keep them here as provisional separate units without pretending
Faundez (1994: 574) redescribed the holotype BMNH phylogenetic soundness.
674 Porifera' Demospongiae • Poecilosclerida • Mycalina • Mycalidae

Fig. 3. Mycale (Aegogropila) contareni (Martens, 1824). A, In situ photo from Ireland (photo B. Picton). B-E. SEM images of spicules of ZMA POR.
4287 from Sherkin Island, Co. Cork, Ireland (scale bars 10 j.Lm). B, detail of mycalostyle. C, two size categories of sigmas. D, three size categories of
anisochelae. E, toxa.

SUBGENUS AEGOGROPlLA GRAY, 1867 are palmate anisochelae in one to four size categories, the larger
ones forming rosettes, sigmas, raphides, toxas and eventually
Synonymy micracanthoxeas.

[Esperia] Nardo, 1833: 522 (preocc.) (not Esperia Hiibner, Description of type species
1816: Lepidoptera). Aegogropila Gray, 1867a: 533. Pellinula
Czerniavsky, 1879: 125. Esperella Vosmaer, 1887: 353 (misprinted Mycale (Aegogropila) contareni (Martens, 1824) (Fig. 3A-E).
as Esperiella on p. 348). Synonymy. Spongia contarenii Von Martens, 1824: 455,
580. Esperia typica Nardo, 1833: 522 (fide Schmidt, 1862: 54).
Type species Halichondria aegogropila Johnston, 1842: 119, pI. XI fig. 1;
Mycale aegagropila; Stephens, 1912: 32. Esperia foraminosa
Halichondria aegagropila Johnston, 1842: 119 (by Schmidt, 1862: 54, pI. V fig. 3. Esperia bauriana Schmidt, 1862:
subsequent designation; Dendy, 1922b: 55). 55. Esperia modesta Schmidt, 1862: 57. Esperia bacillaria
Schmidt, 1864: 34. Aegogropila varians Gray, 1867a: 533.
Definition Desmacidon copiosus Bowerbank in Norman, 1869a: 299. Mycale
microchaela Ferrer-Hernandez, 1922: 9. Mycale contareni;
Mycale with surface visibly reticulate to the naked eye; Topsent, 1924: 91, fig. 3.
choanosomal skeleton plumose or plumoreticulated made of multi- Material examined. Lectotype of Halichondria aegagropila:
spicular tracts of megascleres of one category only; ectosomal BMNH 1847.9.7.39. Esperia contarenii (sensu Schmidt, 1862):
skeleton a dense tangential reticulation of megascleres isolated or in LMJG 15364. Syntypes of Esperia foraminosa Schmidt, 1862:
tracts, megascleres are subtylostyles (mycalostyles); microscleres LMJG 15361, 15372. Syntypes of Esperia bauriana Schmidt,
Porifera· Demospongiae • Poecilosclerida • Mycalina • Mycalidae 675

1862: LMJG 15370, 15950, SMF 48? - Bay of Muggia, Adriatic. pI. I (=pl. V) fig. 8), which has page priority, because a question
Other material. ZMA 4187 - Roscoff, France. MNRJ 1081, 1149, mark was added to the name by Czerniavsky, possibly indicating
1153, 1157, 1167, 1173, 1175 - Roscoff, France. he considered this species to be a dubious Pellinula. The descrip-
Description (from Ackers et ai., 1992). The shape varies tion of P. schmidtii in Latin is not very informative, but from
from sheets or cushions of unequal thickness to massive-lobose the recorded spicules, it seems a clear Mycale s.l., as styles of
(Fig. 3A). The lobes may become elongate and anastomose, lead- up to 220 fLm, sigmas of 7-9 fLm and chelae of 10-32 fLm are
ing to a superficially ramose appearance. Specimens can grow to described. If the description is understood well, there also was an
a considerable size, at least 10 cm diameter and thickness. Surface Aegogropila-type of ectosomal skeleton. Pellinula is here tenta-
typically covered by small conules, raised up by the skeletal fibres, tively assigned to Mycale (Aegogropila) as a junior synonym. Its
giving the surface a somewhat reticulate appearance. Oscules few, type material is presumably lost (Koltun, in litt.).
large when open, found at summits of the lobes or scattered and Vosmaer (1887: 353) erected Esperella for type species
carried on membraneous fistules on encrusting forms. Colour yel- (by monotypy) Spongia contareni Martens, 1824 (to replace the
low, yellow green, ochrous yellow; preserved white to light brown. preoccupied Esperia), which he also indicated as senior synonym
Consistency firm to soft, even, compressible. Smell quite marked, of Esperia typica Nardo, 1833. He ignored the fact that Gray
sweetish-pungent. Contraction slight. Skeleton plumoreticulate. (1867a) already had created a number of genera covering the char-
The ectosomal skeleton is a tangential, triangular or polygonal acters of Esperia, among which Mycale and Aegogropila. He even
reticulation of multispicular fibres. The choanosomal skeleton is mentioned Gray's Aegogropila and Bowerbank's Raphiodesma
an irregular system of anastomosing stout multi spicular fibres. as synonyms of Esperella. By sharing the same type species
Spongin reinforces the skeletal fibres to a varying degree. Aegogropila and Esperella are objective synonyms.
Megascleres (Fig. 3B) are slightly curved (sub)tylostyles (so-called De Laubenfels (1936a: 122) declared Aegogropila a synonym
mycalostyles), thicker towards the pointed end, with a more or less of Carmia without explanation.
pronounced head, 220-357 X 7-11 fLm, varying between individu- Aegogropila is not diagnosable on the basis of known mor-
als. Microscleres include palmate isochelae in three size groups phological characters, yet. Traditionally recognized on the basis of
(Fig. 3D), the largest are grouped into rosettes, 31-47 fLm; the mid- its reticulated ectosomal skeleton, Aegogropila was discovered to
dle size are solitary and variable in number (and may even be be paraphyletic by the recognition of more exclusive assemblages
absent), 12-25 fLm; the smallest are solitary, often abundant, with also bearing this character, such as Naviculina, Paresperella and
one ala characteristically standing out from the shaft giving the Zygomycale, which bear good synapomorphies each. Thus, like
spicule a noticeable angular profile, 13-15 fLm. Sigmas of two Mycale (Mycale) the present subgenus is one of convenience, with
sizes (Fig. 3C) are present, the larger are numerous, robust and little phylogenetic basis.
somewhat twisted, sizes generally in the range 40-65 fLm; the The studied specimens mentioned above assigned to Mycale
smaller (if present) are thinly scattered and very slender and conse- contareni presented some variation in the abundance of microscle-
quently sometimes hard to find, 16-22 fLm. Toxas (Fig. 3E) are res, which is considered here to be of minor importance, especially
either single and scattered or, nearly always, in small bundles form- as artefacts from preparation cannot be completely ruled out.
ing toxodragmata; they are small and fine, with a widely extended Nevertheless, there has also been detected some variation in
central flexion; they are always present but sometimes rare and dif- the shape of the anisochelae-ill. Both specimens of Esperia
ficult to detect; size 20-70 fLm. Sexual reproduction in summer. foraminosa (syntypes) possess anisochelae-ill which are more
Asexual reproduction also in summer by budding, buds located at straight-shafted, and with more obtuse head and foot, approaching
summits of the conules. Distribution and ecology. British Isles, closer an isochela condition. As intermediary forms were not seen,
Atlantic coast of France and Spain, Mediterranean, Azores; littoral it appears to us that the suggested synonymy between Esperia
to 37 m, growing on stones, shells, algae, sessile coelenterates and foraminosa and Mycale contarenii (sensu Topsent, 1924) needs to
scallop shells, boulders, cobbles, at sheltered sites with moderate be checked further.
tidal streams.
Remarks. Hajdu & Riitzler (1998) argued for acceptance
of Gray's (1867a) Aegogropila varians as type species of
Aegogropila. However, Dendy (1922b: 55) indicates Halichondria SUBGENUS ANOMOMYCALE TOPSENT, 1924
aegagropila Johnston, 1842 as the type of Aegogropila. Both
A. varians and H. aegogropila were mentioned by Gray, 1867a, Synonymy
and moreover no description of A. varians Gray, 1867a was
offered, so there is ample reason to accept Dendy's designation as Anomomycale Topsent, 1924: 116.
valid. This decision was again confirmed by Van Soest (1984b: 12)
and Bergquist & Fromont (1988: 21). Hajdu & Riitzler (1998) Type species
hypothesized about the reasons for Gray to propose A. varians (it is
likely that Gray intended the name A. varians to replace A. Desmacidon titubans Schmidt, 1870: 55 (by monotypy).
aegogropila in order to avoid a tautologous name), but this is not
certain from re-examined specimens or slides. Definition
The genus Pellinula Czerniavsky, 1879: 125 (1880: 60)
was erected for type species (here designated) Pellinula schmidtii Mycale with anomochelae (palmate anisochelae with a slightly
Czerniavsky, 1879: 126, pI. I (=pl. V) figs 9-10 (1880: 62, curved shaft on face view; and frontal ala of the foot considerably
pI. I (=pl. V) figs 9-10), from the northern shores of the Black Sea expanded, approaching or even surpassing the dimensions of the
at a depth of 10-15 m. This designation foregoes the choice of usually larger frontal ala of the head, contorted, sometimes with
P. cribrosa Czerniavsky, 1879: 125, pI. I (=pl. V) fig. 8 (1880: 60, a serrated upper border).
676 Porifera· Demospongiae • Poecilosclerida • Mycalina • Mycalidae

A
B

c D
Fig. 4. Mycale (Anomomycale) titubans (Schmidt, 1870). A, drawing of anomochelae reproduced from Schmidt, 1870: pI. V fig. 18 (sizes see text).
B-D, drawings of spicules reproduced from Lundbeck, 1905: pI. X fig. 3 (sizes see text). B, subtylostyles. C, sigmas. D, anomochelae.

Description of type species same species as Schmidt's. This is a deep-water species recorded
from the Atlantic as well as from the Southern Oceans.
Mycale (Anomomycale) titubans Schmidt, 1870 (Fig. 4A-D). Remarks. We examined further material belonging to this
Synonymy. Desmacidon titubans Schmidt, 1870: 55, pI. V subgenus from Antarctica (USNM 43481, 43482, 43483,
fig. 18; Mycale titubans; Lundbeck, 1905: 41, pI. X fig. 3; Mycale Antarctic), identified as Anomomycale sp. by V.M. Koltun. These
(Anomomycale) titubans; Topsent, 1924: 116, fig. 16. Antarctic specimens have sigmas often 400 fLm long. Likewise,
Material examined. Lectotype (here designated): BMNH material recorded by Boury-Esnault & Van Beveren, 1982 from
1870.5.3.41 - slide made by Schmidt, labeled 'Florida 176 fms, Kerguelen has overall larger dimensions than the Atlantic material.
nr. 79'. Lectotype designation is made for this material since no
further material of the type species is known to exist
(Desqueyroux-Faundez & Stone, 1992: 41).
DescriptWn (based on Schmidt, 1870). Shapeless mass. SUBGENUS ARENOCHAUNA LENDENFELD, 1887
Skeleton consisting of an irregular system of finn spicule tracts built
from large thick megascleres making an axial structure from which Synonymy
issue at oblique angles thinner tracts made up of smaller megascleres.
The megascleres are mycalostyles in two size classes. Microscleres [Acamasl Duchassaing & Michelotti, 1864: 94 (preocc.) (not
anomochelae (Fig. 4A) of about 30 fLm long and sigmas in a large Acamas Montfort, 1808, Mollusca). Arenochalina Lendenfeld,
size range, divisible into two sizes, larger ones up to 250 fLm and 1887c: 821. Acamasina de Laubenfels, 1936a: 117. Parisociella
smaller of ca. 30 fLm (Schmidt says 3 fLm, but that is clearly a mis- Burton, 1952: 169. Kerasemna Pulitzer-Finali, 1981: 105.
print), the latter occurring in sigmodragmata. North Atlantic material
described by Lundbeck (1905: 41) has mycalostyles (Fig. 4B) in two Type species
size categories, 590-900 fLm and 320-400 fLm long. Microscleres
anomochelae (Fig. 4A, D) 24-52 fLm long, and sigmas (Fig. 4C), Arenochalina mirabilis Lendenfeld, 1887c: 821 (by
50-140 fLm long. It is uncertain whether this material belongs to the monotypy).
Porifera· Demospongiae • Poecilosclerida • Mycalioa • Mycalidae 677

Fig.S. Subgenus Mycale (Arenochalina). A, photo ofholotype of Arenochalina mirabilis reproduced from Lendenfeld, 1887c: pI. XXVI fig. 70 (size see text).
B, drawing of cross section of skeleton reproduced from Lendenfeld, 1887c: pI. XXVII fig. 28 (size see text). C, drawing of spicules reproduced from
Wiedenmayer, 1989: text-fig. 56 (sizes see text). D, holotype of Acamas laxissima Duchassaing & Michelotti, 1864 (type of [Acamas] and Acamasina (photo
L.A. van der Laan) (size see text). E, in situ photo of Jamaican specimen of Mycale (Arenochalina) laxissima (photo H. Lehnert). F-G, SEM images of spicules
of M (A.) laxissima (photos H. Lehnert). F, anisochela (scale bar 1 fLm). G, sigmas (scale bar 10 fLm). H-I. Kerasemna tenuityla Pulitzer-Finali, 1981. H. photo of
holotype reproduced from Pulitzer-Finali, 1981: fig. 16 (scale bar I cm). I, drawing of spicules reproduced from Pulitzer-Finali, 1981: fig. 17 (scale bar 10 fLm).

Definition Synonymy. Arenochalina mirabilis Lendenfeld, 1887c:

821, pIs XXVI fig. 70, XXVII fig. 28; Pulitzer-Finali, 1981: 100,
Mycale with choanosomal skeleton conslstmg of a stout, fig. 101; Mycale (Arenochalina) mirabilis; Wiedenmayer, 1989: 84,
mostly quadrangular reticulation of spongin fibres cored by foreign pI. 9 figs 8-10, text-figs 53-56. Esperella spongiosa Dendy, 1896:
material and/or filamentous algae and few proper megascleres; 16. Mycale jistulata Hentschel, 1911: 292.
ectosomal skeleton absent; few scattered choanosomal megascleres Material examined. Holotype: BMNH 1886.8.27.587 -
and microscleres; megascleres mycalosty1es of one category only; Torres Straits, Nortlt Australia.
microscleres anisochelae of one category only, often absent or rare, Description (adapted from Wiedenmayer, 1989: 84).
and sigmas, occasionally absent or rare. Variably lobate, massive, contorted, erect or repent, sometimes
with pedicel. The holotype (Fig. 5A) is finger-shaped, 10 X 2.5 X
Previous reviews 1.5 cm. Oscules conspicuous, apical on lobes, 4-8 mm in diameter
in the holotype, other specimens often with much smaller, scattered
Van Soest (1984b as Mycale (Acamasina)), Wiedenmayer ones. Surface conulose to bristly, with commonly blunt primary
(1989), Riitzler (1990), Hajdu & Riitzler (1998). fibre tips piercing dermis. Some specimens with coarser, widely
spaced fibres, with longer conules and tapering fibre tips. Other
Description of type species specimens fleshier, with thicker, more continuous dermis, surface
then smoother. Softly spongy, very compressible. Much slime upon
Mycale (Arenochalina) mirabilis Lendenfeld, 1887c collecting. Dermis and choanosome mostly yellowish cream to
(Fig. 5A-C). beige in life, with some pink and greenish. Colour recorded as
678 Porifera' Demospongiae • PoeciioscIerida • Mycalina • Mycalidae

beige, orange red, cream; in shallow waters off Heron Island (south- (Fig. 51) and, according to Pulitzer-Finali, thin toxas, which were not
ern Great Barrier Reef) the species is conspicuously blue, with observed by us. Instead we noticed many juvenile megascIeres, which
dark-red to purple-coloured fibres in life. Fibrous skeleton mostly often take a sinuous shape and may have been mistaken for toxas.
regUlar. In the holotype meshes are rectangular and 400-800 fLm Pulitzer-Finali (1981) assigned his genus to Desmacellidae because
wide; in other specimens mesh size may be around 1 mm. Primary of the lack of anisochelae. However, this is not unusual in species of
fibres (Fig. 5B) usually cored by foreign debris. Meshwork in some Mycale (Arenochalina) and in view of all other compelling similari-
specimens lax, less regular, primaries then with few foreign inclu- ties we consider Kerasemna a junior synonym of Arenochalina.
sions. Most fibres contain filamentous algae. Thickness of fibres in
the holotype averages 50 fLm. Megascleres mycalostyles (Fig. 5C),
variably thick, mostly slender, in the holotype they measure SUBGENUS CARMIA GRAY, 1867
155-165 X 4.8--6 fLm. These were not noted by Lendenfeld, who
quotes oxeas of 20 X 4 fLm as megascleres. Other records (Dendy, Synonymy
1896; Wiedenmayer, 1989) give an overall size range of 155-268 X
1.9-9.5 fLm. Microscleres anisochelae and sigmas, but they are very Carmia Gray, 1867a: 537. Oxycarmia de Laubenfels, 1954: 155.
rare in the holotype, only a single palmate anisochela of 35 fLm was
found. Other descriptions (Dendy, 1896; Wiedenmayer, 1989) give Type species
an overall range of anisochelae of 19.~30 fLm. Sigmas (Fig. 5C),
54 fLm in the holotype, with recorded range of 24--86 fLm. Hymedesmia macilenta Bowerbank, 1866 (by subsequent
Distribution. North, East and South East Coasts of Australia. monotypy).
Remarks. Duchassaing & Michelotti, 1864: 95 erected a
genus Acamas for type species (subsequent designation by de Definition
Laubenfels, 1936a: 117) Acamas laxissima. The genus name
Acamas is preoccupied by Acamas Montfort, 1808 (Mollusca), Mycale with plumose or plumoreticulated choanosomal skele-
which induced de Laubenfels (1936a: 117) to erect a new genus ton; ectosomal skeleton absent, or only a few scattered megascIeres
name Acamasina with the same species A. laxissima as the type. lying tangentially, as well as dispersed microscIeres; megascIeres
The lectotype (designation by Van Soest et al., 1983), Mus. Torino are subtylostyles (mycalostyles) in one category only; microscIeres
Por. 34 (with fragments deposited in BMNH, MNHN and USNM) are palmate anisochelae in one or more size-categories, the larger
was re-examined (Fig. 5D). It is a macerated mass of crooked- of which may form rosettes, as well as a variable complement of
laminated fibres, similar to those of A. mirabilis, filled usually sigmas, toxas, raphides and micracanthoxeas.
with algal strands and a core of subtylostyle megascleres. Some
sigmas and anisochelae of similar size to A. mirabilis are present. Previous reviews
Wiedenmayer (1977, as Thorecta horrida) described also foreign
spicules inside the fibres. The amount and nature of coring Van Soest (1984b), Bergquist & Fromont (1988), Carballo &
with foreign material is here interpreted as species-specific and we Hajdu (1998).
follow here Wiedenmayer's (1989) suggestion that Arenochalina
and Acamasina are synonyms. Full-grown specimens of M. (A.) Description of type species
laxissima assume the shape of a cluster of tubes (Fig. 5E), but
retain the sparing presence of anisochelae (Fig. 5F) and sigmas Mycale (Carmia) macilenta (Bowerbank, 1866) (Fig. 6A-B).
(Fig.5G). Synonymy. Hymeniacidon macilenta Bowerbank, 1866: 176;
The genus Parisociella Burton, 1952: 169 was erected for Carmia macilenta; Gray, 1867a: 537; Esperella macilenta; Topsent,
type species Esperiopsis anomala Ridley & Dendy, 1886: 341 (see 1894: 8; Mycale macilenta; Stephens, 1912: 33. Hymeniacidon
also Ridley & Dendy, 1887: 84) collected near Honolulu and floreum Bowerbank, 1866: 190; Bowerbank, 1874b: pI. XXXIII
recorded from the Red Sea by Burton (1952), with type BMNH figs 7-13; Raphiodesma floreum; Bowerbank, 1874b: 94.
1887.5.2.165, wet and slides (re-examined here). There are no pub- Raphiodesma fallaciosum Bowerbank, 1882: 163, pI. XVII
lished illustrations of this species. This is an irregularly ramose, figs 7-12. Esperella hamata Topsent, 1892b: xxi.
digitate sponge, length 12.5 cm, diameter of the branches 0.6 cm. Material examined. Holotype (re-examined in Carballo &
Colour light or dark grey-yellow, consistency soft, fibrous. The Hajdu, 2001): BMNH 1910.1.1.135 - dry, Norman collection.
skeleton is a rectangularly meshed system of stout spongin fibres Other material. MNRJ 1013 - La Tortue, Roscoff, France.
cored by relatively few styles/tylostyles of 250 X 5 fLm, vestigial ZMA 5918 - Sherkin Island, Ireland. ZMA 7218 - Pregui~a,
chelae scarce, minute, 10 fLm. Ridley & Dendy (1887) call these sao Nicol1io, Cape Verde Islands.
chelae 'isochelae' whereas Burton (1952) refers to them as palmate Description (after Ackers et al., 1992). Encrusting, sheet or
anisochelae. They are apparently vestigial and may be difficult to cushion, up to 10cm across. Surface smooth. Oscules are clearly
classify as either type. The remaining features are in accordance visible in living sponges, on erect tubes, but are not very numerous.
with characters of Arenochalina and it is proposed here to Pores small, scattered over the surface. Alive, the sponge is pale
synonymize Parisociella with the present subgenus. yellow. When preserved it is yellow or brown. Consistency soft.
The genus Kerasemna Pulitzer-Finali, 1981: 105 was erected for Skeleton plumose; ascending multi spicular tracts of subtylostyles
type species (by monotypy) Kerasemna tenuityla Pulitzer-Finali, (which do not divide or anastomose), ending in loose brushes at the
1981: 106, figs 1~17. The type specimen MSNG 46937 was re- surface, with single megascIeres lying scattered in between.
examined. This is a clathrate, macerated lobate mass of stout spongin Spongin is minimal. There is no specialised ectosomal skeleton. The
fibres (Fig. 5H) cored by subtylostyles (Fig. 5I) in the same size range surface is supported by the widely spaced, terminal brushes of
as those of Arenochalina mirabilis. Microscleres are thin sigmas the main skeleton, with a few single megascleres (sometimes these
Porifera' Demospongiae • Poecilosclerida • Mycalina • Mycalidae 679

A B

c-=========~~ r TO

Fig. 6. Subgenus Carmia. A-B. Mycale (Carmia) macilenta. A, habit of specimen in situ (photo B. Picton). B, type specimens and spicules reproduced
from Bowerbank (l874b: pI. XXXIII figs 7-13) (sizes see text). C, spicules of Oxycarmia confundata reproduced from de Laubenfels, 1954: fig. 101 (sizes
see text). D-E, Mycale (Carmia) micracanthoxea Buizer & Van Soest, 1977. D, SEM overview of spicules, showing masses of micracanthoxeas (arrow)
(scale bar 10 jJ.m). E, SEM image of micracanthoxeas (scale bar 1 jJ.m).

are absent altogether) scattered in the large interstices. The megas- (see below), instead, but in that subgenus these replace the
cleres are generally straight-shafted, slightly fusiform, subty- mycalostyles. In this case it is thought more likely that the oxeas
lostyles, with barely formed elliptical heads, between 200 and 300 may be foreign. Carmia was given a quite different emphasis by
/-Lm long. Microscleres include palmate anisochelae of three sizes. de Laubenfels (1936a: 118), who defined it as an independent
The largest are grouped into rosettes in the ectosome, 33-59/-Lm. genus similar to Mycale but differing from it through the posses-
The middle size are solitary and scarce, 17-24/-Lm. Those of the sion of toxas.
smallest size are very abundant and widely dispersed throughout A remarkable species of Carmia is Mycale (Carmia)
the body, 11-15/-Lm. They have a distinctive lower tooth which is micracanthoxea Buizer & Van Soest, 1977, based on its possession
very short and curved in towards the shaft. The sigmas fall into two of a special microsclere type, the micracanthoxeas (Fig. 6D-E).
size categories, 65-115 /-Lm and 21-28 /-Lm. Toxas are variable in These tiny microscleres were subsequently found in a Mycale of
length, 60-250/-Lm. Larvae are found between August and mid- the Straits of Gibraltar (Carballo & Garcia-Gomez, 1994) assigned
October. Distribution and ecology. Common along the Channel to the same species, M. micracanthoxea, but also in a NE Pacific
coasts of the British Isles and France; Belgium; Atlantic coasts species (M. bamfieldensis Reiswig & Kaiser, 1989, as bamfieldense)
of France, Spain, Mediterranean, North Africa; littoral to 27 m, and a South American species (M. escarlatei Hajdu et al., 1995).
encrusting mainly on Pecten and Chlamys. The latter two species are assignable to subgenus Aegogropila on
Remarks. The suggested synonymy of Oxycarmia de the basis of their reticulate tangential skeleton. Carballo & Hajdu
Laubenfels, 1954: 155 (type species (by monotypy) O. confundata (1998) described another two species with micracanthoxeas,
de Laubenfels, 1954: 155, fig. 101; here reproduced in Fig. 6C), viz. M. (Aegogropi/a) lilianae Carballo & Hajdu, 1998 and
needs to be confirmed by re-examination of the holotype. The M. (Carmia) urizae Carballo & Hajdu, 1998.
genus was founded on the presence of large numbers of oxeas The observation that at least one species of Aegogropi/a
of 685 X 15 /-Lm, apparently arranged perpendicular to the surface may lack the ectosomal skeleton (cf. Hajdu & Rtietzler, 1998), and
and criss-cross in the choanosome. The remaining skeleton is typi- that micracanthoxeas are shared by some species of Carmia-like
cally that of Carmia; it consists of the usual bundles of thin subty- sponges and some Aegogropi/a-like sponges, is a strong indication
lostyles of 225 X 1 /-Lm, palmate anisochelae, 20/-Lm, two size that Carmia is most likely polyphyletic. The assemblage is kept
categories of sigmas, 70/-Lm and 20/-Lm, toxas of 130-235/-Lm for convenience only, and should preferably not be given full
and trichodragmas of 60-120/-Lm long. The oxeas would seem generic status (as did Bergquist & Fromont, 1988), until a more
to indicate its closest affinities to Oxymycale Hentschel, 1929 comprehensive revision of its species is undertaken.
680 Porifera· Demospongiae • Poecilosclerida • Mycalina • Mycalidae

A further remarkable species of Carmia is an unnamed strengthened by inclusion of variable amounts of sand grains, in a
species forming a close symbiotic relationship with the octocoral tight pack, spread, or in patches. Spicular density varies inversely.
Tubipora musica, reported by Van Soest & Verseveldt, 1987, from Architecture confused; sometimes reticulated, with megascleres
Indonesia. The normally semiglobular colonies of Tubipora are not organized in well defined bundles. Projecting terminations of
forced into clusters of tubes with apical vents by the cohabitation subectosomal brushes often well apparent, along with few tangen-
with the MycaZe species. tially displaced megascleres. Rosettes of anisochelae-I (ca. 100/-Lm
in diameter) uncommon. Rosettes of anisochelae-II (ca. 50/-Lm in
diameter) very rare, not seen in some specimens, to common in
SUBGENUS GRAPEUA GRAY, 1867 ZMA 10712. Apparent absence could be an artefact due to the dry
(=contracted) state of the specimens and/or abundance of sand-
Synonymy grains, which renders (uncommon) spicular characters difficult to
assess in thick-sections. Anisochelae-III may be common among
Grapelia Gray, 1867a: 534. Pseudoesperia Carter, 1886c: 455. subtylostyles. Choanosomal skeleton often bearing a remarkable
number of sand grains, with megascleres merely filling-in inter-
Type species stices. Sand-cored fibres more than 1 mm wide in the subectosomal
area, where they diverge in profusion to support the ectosomal
Grapelia australis Gray, 1867a: 534 (by monotypy). layer. Fibres no more than 400 /-Lm wide, where subtylostyles pre-
dominate. Spongin present, more apparent in sand-cored fibres.
Definition Overall pattern observed is plumo-reticulated, ascending fibres
diverging and anastomosing frequently, spreading into brushes
MycaZe with a confused tangential ectosomal skeleton, and in the subectosomal area. One specimen from Tasmania (ZMA
with three categories of anisochelae one or more of which have 10712) possesses a much more spicular, much less sandy skeleton,
unguiferate alae or alae reduced to a series of spines. with fibres equally stout. Spongin scarce, which seems compen-
sated by a spread pattern shown by subtylostyles in places where
Remarks tracts intersect. Megascleres: Subtylostyles, straight, slightly sinu-
ous, gently or markedly curved, slender with elliptic head or stout
The definition may be extended with the following details: with oval head, sharp apex (228-382 X 3-12 X 3-15/-Lm, total
anisochelae-I with a curved shaft in profile view, ratio height of length X width of head X width of shaft, 254-(291.5)-313 X
the head/total height of the spicule <25%, alae of the foot projecting 4-(5.4)-7 X 5-(6.1)-7/-Lm in neotype). Microscleres (Fig. 7C),
downward forming a pore, and rosettes built both by anisochelae-I anisochelae-I, shaft markedly curved on profile view, unguiferate
and -II; mostly with unguiferate anisochelae-I, acanthose head, lateral alae narrow and sharp, often longer than frontal alae,
anisochelae-II, and basally-spurred anisochelae-ill). Hajdu (1995) frontal alae bifid, height of the head 15-20% of total length;
assigned one species apparently lacking unguiferate or spined chelae palmate foot, top border of frontal ala often wavy, or with a single,
to Grapelia, but the only available material, a microscopic slide, pre- variably sharp projection, or small spines (29-61 X 4-12/-Lm,
vented a clear determination of the fine structure of the alae. total height X height of head, 36-(41.2)-50 /-Lm in neotype);
anisochelae-II, shaft markedly curved on profile view, variably
Description of type species stout; acanthose head with crown of spines, alae not distinguish-
able, larger-primary (up to 8-9) and smaller-secondary spines
MycaZe (Grapelia) australis Gray, 1867a (Fig. 7A-C). (20-30 or more) often recognizable; palmate foot, height 25-35%
Synonymy. Hymeniacidon spec. Bowerbank, 1864: fig. 135. of total length, spiny top border of the alae (15-22/-Lm,
Grapelia australis Gray, 1867a: 534. Esperia parasitica Carter, 15-(17.6)-20/-Lm in neotype); anisochelae-ill, palmate, shaft gen-
1885a: 108. Pseudoesperia enigmatica Carter, 1886c: 455. MycaZe tly curved on profile view, height of the head 30-50% of total
parasitica var. arenosa Hentschel, 1911: 311. length, lateral alae longer than frontal one; reduced foot with vari-
Material examined. Neotype (designated by Hajdu et aZ., ably vestigial alae, basal spur-like projection (15-21/-Lm,
1995: 9): BMNH 1886.12.15.467 (=holotype of Pseudoesperia 15-(17.0)-19/-Lm in neotype).
enigmatica Carter, 1886c). Other material. Dendy's (1896) mate- Remarks. Grapelia is one of the most easily diagnosed
rial of Esperella enigmatica: NMV F 65700-65704. Hentschel's subgenera of MycaZe. Extensive taxonomic remarks on Grapelia
(1911) material of MycaZe parasitica var. arenosa: ZMH 1666. are provided by Hajdu (1995) and Hajdu et aZ. (1995). The sub-
Material of M. (G.) parasitica var. arenosa (sensu Shaw, 1927): genus has representatives in all tropical and warm-temperate
BMNH 1925.11.1.729; ZMA 10712 - Tasmania, colI. M. Carpay. regions of the world oceans.
Description (adapted/rom Hajdu, 1995). Specimens mas- By choosing the holotype of Pseudoesperia enigmatica
sive, lobose, or spherical, with or without conical protuberances Carter, 1886c as neotype of Grapelia australis, the genus
(up to 5 cm high). Dimensions of the neotype (Fig. 7A) 15 X 15 X Pseudoesperia Carter, 1886c was made an objective synonym
10 cm. Surface mostly reticulated to the naked eye, occasionally of Grapelia. A recently described species from the Colombian
wrinkled or conulose, easily removed. Oscula large, up to 7 mm in Caribbean, MycaZe (GrapeUa) unguifera Hajdu et aZ., 1995,
dry neotype, in a row, or on top of protuberances, sometimes not although differing greatly in habit (thin slimy crusts) exhibits
seen. Consistency elastic, soft to firm, dry specimen is hard. Colour strikingly similar anisochelae, both in shape and size (Fig. 7D-E).
of the specimens from Port Phillip Heads, dull orange-yellow Differences are the slightly larger subtylostyles and the presence of
or ochre-yellow (Dendy, 1896). Specimen from Stockyard sigmas in M. (G.) unguifera. Other species of GrapeUa likewise are
Point beige to mustard-yellow in life. Ectosomal skeleton often closely similar to the type species in spicular characters.
Porifera· Demospongiae • Poecilosclerida • Mycalina • Mycalidae 681

Fig. 7. Subgenus Grapelia. A-C, neotype of Mycale (Grape/ia) australis (Gray, 1867a) =holotype specimen of Esperia parasitica Carter, 1885a, repro-
duced from Hajdu (1995; Figs 6.3, 6.9-12). A, habit photographed from damaged (under-)side to show skeletal structure (scale bar I em). B, subtylostyle
(scale bar 10 fLm). C, anisochelae I (top, scale bar IOfLm), II (middle) and III (bottom) (bottom scale bar 2 fLm). D-E, Mycale (Grapelia) unguifera,
anisochelae I (D, scale bar 10 fLm) and II (E) (scale bar 2 fLm), reproduced from Hajdu et aI., 1995; figs 24, 27-28.

SUBGENUS NAV/CUliNA GRAY, 1867 Description of type species

Synonymy Mycale (Naviculina) cliftoni Gray, 1867a (Fig, 8A-C).

Synonymy. Hymedesmia spec. Bowerbank, 1864: 252,
Naviculina Gray, 1867a: 538. fig. 152. Naviculina cliftoni Gray, 1867a: 538; Hooper &
Wiedenmayer, 1994: 293; Mycale (Naviculina) cliftoni; Hajdu,
Type species 1999: 227, figs 1-2.
Material examined. Holotype: BMNH 1877.5.21.270 -
Naviculina cliftoni Gray, 1867a: 538 (by monotypy). slide, Freemantle, Western Australia.
Description (from Hajdu, 1999: 227). One single thick-
Definition section preparation remains. It contains a perfectly preserved
fragment of the specimen's surface peel, from which it is possible
Mycale with a reticulated tangential ectosomal skeleton and to gather the whole series of spicules in Mycale. This peel contains
naviculichelae (complete or near fusion of both frontal alae, falx an ectosomal skeleton characterized by a neat reticulation
markedly expanded along the shaft, lateral alae of the head project (Fig. 8A) of megasclere bundles (2-6 spicules across) or single
backward and upward). megascleres, forming meshes which are mostly triangular (from
40 X 70 to 240 X 350 /-Lm across), and inside which pores are
Previous review clearly visible (60/-Lm across). Naviculichelae abound inside the
meshes (Fig. 8B), especially around the bundles of megascleres.
Hajdu (1999). Megascleres mycalostyles, smooth, mostly straight, slightly
682 Porifera· Demospongiae • Poecilosclerida • Mycalina • Mycalidae

c
Fig. 8. Mycale (Naviculina) cliftoni (Gray, 1867a). A, tangential view of surface skeleton (scale bar 100 ILm). B, detail to show spicule bundles and navi-
culichelae (scale bar 100 ILm). C, naviculichelae photographed from various angles and details (scale bars 51Lm) (all reproduced from Hajdu, 1999: figs 1-2).

fusifonn, with elliptic or oval heads, and points which taper more- complement of sigmas (Fig. 9A-F). In order to avoid confusion
or-less gradually, 330-(357.4)-388 (N = 20) X 4.8-8.4 (head, with the isochelae of Clathria cleistochela Topsent, 1925, Hajdu
N = 10) X 6-9.6/l-m (shaft, N = 10). Microscleres naviculichelae (1999) proposed to name naviculichelae, the so-called 'cleis-
(Fig. 8C), head 60-70% the total spicule length, with narrowing or tochelae' of Mycale.
complete regression of the frontal alae of the head, which may
touch the one of the foot, lateral alae of the head projecting back-
ward and slightly upward, downward expansion of the upper falx SUBGENUS OXYMYCALE HENTSCHEL, 1929
along the shaft, 12-(17.3)-21.6 (N= 100). Sigma (proper ?),
slender, smooth, sharp endings, 14.4/l-m (N = 1). Synonymy
Remarks. Naviculina is diagnosed in accordance with
recent assessment of its characters (see Hajdu, 1999). Hooper & Oxymycale Hentschel, 1929: 932.
Wiedenmayer (1994: 293) employed Naviculina as a separate
genus in Mycalidae and apparently emphasized different aspects of Type species
Naviculina. They made Arenochalina a junior synonym, but made
no statements to support this. We fail to detect similarities or Esperia intermedia Schmidt, 1874: 433 (by monotypy).
shared features between Naviculina cliftoni and Arenochalina
mirabilis Lendenfeld, 1888 (see above). Hajdu (1999) proposed to Definition
assign all Mycale species with so-called 'cleistochelae' (in which
the frontal alae touch and the shaft expands towards them), proba- Mycale with exclusively oxeas as megascleres.
bly about half a dozen in total, to Naviculina, since these all share
a reticulated ectosomal skeleton. Examples are Mycale diversisig- Previous review
mata Van Soest, 1984b (Fig. 9A-C) and Mycale cleistochela
flagellifera Vacelet & Vasseur, 1971 (Fig. 9D-F). By this feature Hentschel (1929).
Naviculina belongs to the Aegogropi/a-like Mycale, but differs
from that subgenus in having the peculiar chelae. The type Description of type species
species M. (N.) cliftoni lacks nonnal chelae, but M. diversisigmata
and M. cleistochela flagellifera possess them, as well as a varied Mycale (Oxymycale) intermedia (Schmidt, 1874) (Fig. lOA-B).
Porifera· Demospongiae • Poecilosclerida • Mycalina • Mycalidae 683

Fig. 9. Mycale with naviculichelae, assigned to Naviculina, A-C, Mycale diversisigmata Van Soest, 1984b from Curac;ao, ZMA 4783. A, tangential view
of reticulate ectosomal skeleton (scale bar 100 fLm). B, SEM image of naviculichela (scale bar 10 fLm). C, flagellated sigma (scale bar 10 fLm).
D-F, Mycale cleistochelaflagellifera Vacelet & Vasseur, 1971, SEM images of spicules of ZMA 4597 from Indonesia. D, naviculichelate anisochela II
(scale bar 10 fLm). E, naviculichelate anisochela III (scale bar 1 fLm). F, normal anisochela I (scale bar 10 fLm).

Synonymy. Esperia intermedia Schmidt, 1874: 433, fig. 10; Remarks. Other species may be used to complete the diag-
Mycale intermedia; Thiele, 1903b: 381, fig. 12; Oxymycale inter- nosis of this subgenus. We examined material of Mycale
media; Hentschel, 1929: 932. Oxymycale wagini Koltun, 1952: 127. (Oxymycale) acerata Kirkpatrick, 1907a (BMNH 1933.6.10.128,
Material examined. None. Type material has not been South Georgia; MNRJ 1212, Chilean Antarctic Territory; MNRJ
recovered so far. Thiele (1903b) described a slide made by Weltner, 1254, off Elephant Island, Antarctica). Some of the North Pacific
which could be in 2MB. species described by Sim et al. (1988) also appear to be true mem-
Description (adapted from Thiele, 1903b: 381 and Koitun, bers of this subgenus. Oxymycale comprises massive or lobate
1959: 71). The type apparently was an insignificant fragment. sponges with skeleton of spicule bundles, which may be coarsely
Koltun describes the species as cushion-shaped, 'lumpy', up to reticulate. Surface skeleton usually developed consisting of inter-
5 cm in height, often in the form of an elongated oval encrustation crossing spiCUle bundles. Megascleres exclusively sharp-pointed
on the branches of hydroids, polychaete tubes, algae etc. There is oxeas. Microscleres the usual complement of several size cate-
apparently an ectosomal skeleton, but no details are recorded. The gories of anisochelae, sigmas may be present but are lacking in the
skeleton consists of weakly developed bundles of megascleres type species. A couple of species from cold water mostly.
which are not cemented together. Megascleres curved, sharply
pointed oxeas (Fig. lOB) of 300-550 X 10-16/-Lm. Microscleres
anisochelae I (Fig. lOA-B), 50-74/-Lm, and II (Fig. lOB), 8-27/-Lm. SUBGENUS PARESPERELLA DENDY, 1905
No further microscleres are recorded. Distribution and ecology.
E coast of Greenland (type locality), no depth known; subsequent Synonymy
specimens have been reported from the Barents Sea, Laptev Sea
and the sea NW of Greenland; depth 17-325 m. Paresperella Dendy, 1905: 162.
684 Porifera· Demospongiae • Poecilosclerida • Mycalina • Mycalidae

the Liverpool Free Museum). Spicules (Fig. 11A) consisting of

fusiform subtylostyles, 175 X 6 J.Lm; microscleres large robust ser-
rated sigmas, 100 J.Lm (thickness 6 J.Lm), palmate anisochelae,
occurring singly or in rosettes, 17 J.Lm, toxas arranged in toxodrag-
mas (not shown in Fig. 11A), 50 J.Lm. Distribution and ecology.
Gulf of Manaar, off Galle, Sri Lanka. If Dendy's subsequent record
is correct, then it is a deep water species.
Remarks. Dendy's (1905) subsequent record of Paresperelia
serratohamata likewise was a minute specimen encrusting calcare-
ous algae, so it may well be the usual shape of this species. Dendy
(1905: 163) recorded a second species from Sri Lanka with a more
elaborate shape allowing description of details of the skeletal struc-
ture. The ectosomal skeleton consists of tangential spicule tracts
making a wide-meshed reticulation. The choanosomal skeleton con-
sists of multi spicular columns and many loose single spicules.
These skeletal details match the ZMA Indonesia specimen studied
here (Fig. lIB-F), so it is likely that all members of this subgenus
A have an Aegogropila-type of ectosomal skeleton. The ZMA speci-
men studied here, which bears an unpublished name given by
M. Burton, is peculiar in having the pointed ends of many megascle-

~ B
res formed into an exotyle-like crown of spines or lobes (Fig. liD),
quite similar to those described for the subgenus Sceptrospongia
(see below). Another comparable occurrence is the bifid termination
Fig. 10. Mycale (Oxymyca/e) intermedia (Schmidt, 1874). A, drawing of in the megascleres of M. (Paresperelia) bidentata (Dendy, 1905).
anisochela I reproduced from Schmidt, 1874: fig. 10 (size see text).
B, spicules of the type reproduced from Thiele's (1903b: fig. 12) redescription
(sizes see text).
SUBGENUS RHAPHIDOTECA SAVILLE KENT, 1870

Type species Synonymy

Esperia serratohamata Carter, 1880b: 49 (by original Rhaphidotheca Saville Kent, 1870a: 219. Gomphostegia
designation). Topsent, 1896b: 149. Sceptrospongia Dendy, 1926: 7.

Definition Type species

Mycale with choanosomal skeleton plumose or plumoreticu- Rhaphidotheca marshallhalli Saville Kent, 1870a: 219 (by
lated, made of multi spicular tracts of megascleres; ectosomal monotypy).
skeleton often a tangential reticulation of megascleres; megascleres
subtylostyles (mycalostyles) often of one size category only; Definition
microscleres are palmate anisochelae of one or more categories,
the larger ones may form rosettes, serrated sigmas, and occasion- Mycale with ectosomal skeleton pierced by club-shaped
ally toxas. exotyles. Next to these there are the usual mycalostyles,
anisochelae, sigmas and raphides.
Previous reviews
Previous reviews
Dendy (1905), Hajdu & Riitzler (1998).
Thiele (1903b), Hajdu (1995).
Description of type species
Description of type species
Mycale (Paresperelia) serratohamata (Carter, 1880b)
(Fig. 11A). Mycale (Rhaphidotheca) marshalihalii (Saville Kent, 1870a)
Synonymy. Esperia serratohamata Carter, 1880b: 49, pI. V (Fig. 12A-E).
fig. 20; Paresperella serratohamata; Dendy, 1905: 162, pI. XI fig. 2. Synonymy. Rhaphidotheca marshallhalli Saville Kent,
Material examined. Type material: not in BMNH 1870a: 219, pI. XV figs 1-7. Esperia rhopalophora Schmidt, 1875:
(Ms C. Valentine, pers. comm.). Other material. ZMA 1739 - 118, pI. I fig. 12; Rhaphidotheca rhopalophora; Thiele, 1903b:
Paresperella spec., Indonesia, Siboga Exped. stat. 318, 06°36'S 383, fig. 14. Rhaphidotheca affinis; Carter, 1879c: 496.
114°55'E, depth 88m. Material examined. Holotype (not examined): BMNH
Description (from Carter, 1880b). The type apparently was 1882.4.27.5 - 1 slide labelled 'from type', from Saville Kent's type
a minute encrustation, 0.4 cm in horizontal expansion, on a calcare- specimen. Presented by Dr J. Millar FRS. Cambridge House,
ous alga. There was enough material to make a slide and still keep Bethnal Green'. Other material. ZMA 5017, 5018 - from Bergen
a dry piece (which was probably destroyed during the bombing of area, Norway, depth 100-600m.
Porifera' Demospongiae • Poecilosclerida • Mycalina • Mycalidae 685

f
,~

I
C
A

Fig. 11. Subgenus Paresperella. A. Mycale (Paresperella) serratohamata Carter, l882a, drawing of spicules of the holotype reproduced from Dendy,
1922b: pI. XI fig. 2 (sizes see text). B-F, SEM images of spicules of Indonesian specimen of Paresperella, ZMA 1739. B-C, subtylostyle (scale bars: B,
100 f,Lm; C. 10 f,Lm). D, detail of crown of lobes of 'pointed' end of subtylostyles (scale bar 10 f,Lm). E. serrated sigma (scale bar 10 f,Lm). F. anisochela II
(scale bar 10 f,Lm).

Description (after Saville Kent, 1870a and Thiele, 1903b). the original descriptions were incomplete; these (cf. above syn-
Cushions or thick encrustations, 1-3cm wide, 1 cm high. Surface onymy) are here considered a single species.
hispid ('bristling') due to projecting exotyles. Colour beige. The genus Gomphostegia Topsent, 1896b was erected for
Consistency soft and fragile, Skeleton of strongly developed type species (by monotypy) Gomphostegia loricata Topsent,
plumose spicule tracts, up to 150/-Lm in thickness, spreading out 1896b: 149, fig. 2, from 845 m near the Azores. A schizotype of the
near the surface to carry a mass of tangential megascleres. Beyond MOM holotype was studied (BMNH 1930.7.1.18). This species is
that surface skeleton, individual exotyles protrude some distance. very similar to R. marshallhalli, but the exotyles have a flattened,
Megascleres styles (Fig, 12B), slightly constricted near the base almost cup-shaped tyle (Fig. 13), which is not found in marshall-
('mycalostyles'), fusiform shaft (hence they were termed 'acerate' haW, although Hajdu (1995: 103) reports variability in this feature.
by Saville Kent), 500-1000 X 16 /-Lm; exotyles (Fig. 12A-D) In any case, the similarities are sufficiently great to consider
with a large but variably shaped and sized elongated tyle the Gomphostegia a clear junior synonym; this was already acknowl-
head of which is rugose (Fig. 12D), protruding beyond the surface edged by Topsent (1904b: 202, pI. XIV fig. 15) himself.
of the sponge tyles outward, 800-1400 X 15-25 /-Lm, tyles 35-55 The genus Sceptrospongia Dendy, 1926: 6 (see also Burton,
/-Lm. Microscleres palmate anisochelae (Fig. 12B, F) in two size I 928a: 121) was erected for type species (by monotypy)
categories, 75-90/-Lm (arranged in rosettes), and 25-40/-Lm; sig- Sceptrospongia coronata Dendy, 1926: 6, pI. I figs 1-8 (see Burton,
mas (Fig. 12B, not recorded by Saville Kent), 13-20/-Lm; tri- 1928a: 121, text-figs 4-5, here reproduced in Fig. 14). The type mate-
chodragmas (Fig. 12B, mistaken for 'acerates' by Saville Kent), rial of the 'Investigator' collection from the Arabian Gulf and
60-80 fLm. Distribution and ecology. Off Spain and Portugal, Andaman Sea is in the Indian Museum Calcutta, 1M P308 and 309.
W coast of Ireland, between Scotland and the Faroes, Norway; on These are subspherical sponges with an ectosomal skeleton of tan-
rocks and dead corals in deep water, from 75 m downwards (type gential intercrossing mycalostyles of 360 X 13 fLm (Fig. 14A). Single
from 900m). exotyles (called 'stephanotyles' by the author), of similar size as
Remarks. Several North Atlantic deep water species the mycalostyles, are erected at right angles, in a regular loose pal-
belonging to this subgenus have been described, due to the fact that isade, heads embedded in the ectosomal skeleton and swollen ends
686 Porifera' Demospongiae • Poecilosclerida • Mycalina • MycaJidae

Fig. 12. Mycale (Rhaphidolheca) marshallhalli (Saville Kent, 1870a). A-B, Esperia rhopalophora Schmidt (1875) considered a junior synonym.
A, drawing of exotyles reproduced from Schmidt, 1875: pI. I fig. 12 (size see text). B, drawing of spicules of Esperia rhopalophora reproduced from
Thiele's (1903b: fig. 14) redescription of Schmidt's type. C-E, SEM images of spicules of ZMA 5017, from Bergen, Norway. C, overview of spicules
(scale bar 100 J.I.m). 0, detail of exotyle (scale bar 10 J.I.m). E, anisochela I (scale bar 10 J.I.m).

A 0" fl
.' ~

..' . G
,,
~ .'
.. ...

.'
;.
.
·. .,-
~ :

. ,,'
\
.

~c========================~~~
B

Fig. 13. Mycale (RhaphiuUlIIt:LU) WltCUtu \~Vp.)t::Ill, lO::;rUU uuml'HUOl

ct;') fig. 2(size see
text). B. photos of sections reproducedfrom Topsent (1928: pI. VII figs 11-12) (size see text).
Porifera' Demospongiae • Poecilosclerida • Mycalina • Mycalidae 687

mm
c
A B
Fig. 14. Mycale (Rhaphidotheca) coronata (Dendy, 1926 as Sceptrospongia), drawings of spicules reproduced from Burton, 1928a: text-figs 4-5, A-B,
megasc1eres consisting of normal subtylostyles and exotyles with a crown of spines, C, microsc1eres. D, cross section of peripheral skeleton showing
arrangement ofthe exotyles (sizes see text).

projecting outwards (Fig. 14D). The swollen ends are peculiar in only; ectosomal skeleton a dense tangential reticulation of megas-
being flattened and crowned with a circle of spines (Fig. 14B), which cleres, singly or in tracts; megascleres mycalostyles; microscleres
are the result of proliferation of the axial canal. Microscleres palmate anisochelae in one or more size categories, the larger ones
(Fig. 14C) consist of the usual complement of anisochelae, I: 75 11m, forming rosettes, small palmate isochelae, sigmas, raphides and
II: 27 11m and III: 15 11m, sigmas I: 45 11m, II: 18 11m, and trichodrag- toxas, the latter two often in bundles (dragmas).
mataI: 45 X 9 11m, II: 15 X 9 11m. The exotyles of this species are per-
haps peculiar, but not fundamentally different from those of M. (R.) Remarks
marshallhalli, and accordingly Sceptrospongia is considered a junior
synonym. We also found megascleres of similar shape in an unidenti- This subgenus is based on the possession of small palmate
fied specimen ZMA 1739, from Indonesia, possessing also serrated isochelae in addition to the usual anisochelae. Most recent authors
sigmas (see Fig. lID). Since these megascleres do not occupy a per- considered these to be plesiomorphic and in view of the fact that
pendicular ectosomal position in the Indonesian specimen, they are other features are shared with Aegogropila synonymized it with
not functional exotyles, although they are morphologically quite sim- that subgenus. Nevertheless, there are at least two species with
ilar. By showing these overlapping characters it is demonstrated that small isochelae and thus it is convenient to distinguish these from
subgenera Paresperella and Rhaphidotheca are likely to be artificial. Aegogropila proper.

Description of type species

SUBGENUS ZYGOMYCALE TOPSENT, 1929
Mycale (Zygomycale) parishi (Bowerbank, 1875b)
Synonymy (Fig. 15A-D).
Synonymy. Raphiodesma parishi Bowerbank, 1875b: 283;
Zygomycale Topsent, 1930: 431. Amphilectus parishi; Vosmaer, 1880: 119; Esperella parishi;
Ridley & Dendy, 1887: 65; Zygomycale parishi; Topsent, 1930: 431.
Type species Material examined. Holotype (not examined): BMNH
1877.5.21.2113 (dry) and 4 slides 1877.5.21.1358 -labeled 'from
Raphiodesma parishi Bowerbank, 1875b: 283 (by original type. Str. of Malacca' (Ms C. Valentine, pers. comm.). Bowerbank
designation). (1875b) noted there were 6 specimens in the original collection
made by Commander Parish. Other material. ZMA 1606-1609 -
Definition all from Indonesia, Siboga collection. ZMA 9590 - Pulau Rantu,
Singapore, colI. H. Moll.
Mycale with plumose or plumoreticulated choanosomal skele- Description (based on Bowerbank, 1875b). Thinlyencrust-
ton made of multi spicular tracts of megascleres of one category ing, up to 8 X 4.5 X 0.3 cm in the original type series; mostly
688 Porifera' Demospongiae • Poecilosclerida • Mycalina • Mycalidae

PHLYCTAENOPORA TOPSENT, 1904

Synonymy

Phlyctaenopora Topsent, 1904b: 199. Barbozia Dendy,

1922b: 131.

Type species

Phlyctaenopora bitorquis Topsent, 1904b: 199 (by

monotypy).

Definition

Fig. 15. Mycale (Zygomycale) parishi (Bowerbank, 1875b), SEM images

Mycalidae with a megasclere complement of oxeas and
of spicules made from ZMA 1606 from Indonesia. A, anisochela I (scale
bar 10 /-Lm). B, isochela (scale bar I/-Lm). C, sigma I (scale bar 10 /-Lm). strongyles in combination with a microsclere complement which
D, trichodragma (scale bar 10 /-Lm). may include peculiar amphiaster-like spined microxeas and
anisochelae, to which sigmas may be added.

similar thin encrustations in reliably identified subsequent speci- Diagnosis

mens. Colour grey-brown alive, cream-white in the dried state.
Surface optically smooth, microscopically hispid due to projecting Massive, smooth or with erect fistules; confused compact
choanosomal spicule tracts. Consistency firm. Ectosomal skeleton choanosomal skeleton of smooth oxeas and scattered bundles
consists of intercrossing tracts of megascleres. Choanosomal skeleton of isolated strongyles; ectosomal skeleton a crust of smooth
an irregular system of anastomosing spicule tracts, with little spongin. strongyles; microscleres include palmate anisochelae which may
Megascleres subtylostyles (mycalostyles) up to 350 X 10 fLm; be basally spurred, sigmas and spiny rhabds, the latter two of
microscleres anisochelae I (Fig. 15A), arranged in rosettes, 55 fLm, which may be absent.
anisochelae III, 17 fLm, palmate isochelae (Fig. 15B), 12 fLm,
sigmas I (Fig. 15C), 82 fLm, sigmas II, 17 fLm, and toxas, Scope
82 fLm. Apparently there were also raphides in trichodragmas
(Fig. 15D), 30-160 fLm, in the type (subsequent observation by The genus is divided in two subgenera, Phlyctaenopora
Ridley, 1884: 436), and in many subsequently recorded specimens. (Phlyctaenopora) and Phlyctaenopora (Barbozia), both with two
Distribution and ecology. Indo-West Pacific, widespread, also species from Atlantic and Pacific localities. The subgenera differ in
reported from the East Pacific; shallow-water. the presence or absence of spined microxeas (dubbed 'oxydis-
Remarks. The species has also been reported from the West corhabds' by various authors).
Indies and Brazil (e.g., de Laubenfels, 1956 and Hechtel, 1965),
but this concerns Mycale (Zygomycale) angulosa (Duchassaing & Previous reviews
Michelotti, 1864 as Pandaros), a blueish ramose sponge, clearly
different from M. (z.) parishi (see e.g., Van Soest, 1984b: 16). Levi & Levi (1983b), Van Soest & Stentoft (1988).

Key to subgenera of Phlyctaenopora

(1) Spined microxeas present ....................................................................................................................................................... Barbozia

Spined microxeas absent .............................................................................................................................................. Phlyctaenopora

SUBGENUS PHLYCTAENOPORA TOPSENT, 1904 Description of type species

Synonymy Phlyctaenopora (Phlyctaenopora) bitorquis Topsent, 1904b

(Fig. 16A-B).
Phlyctaenopora Topsent, 1904b: 199. Synonymy. Phlyctaenopora bitorquis Topsent, 1904b: 199,
pI. V fig. 24, pI. XIV fig. 18.
Type species Material examined. Holotype: MOM (not seen) - 599 m off
the Azores. Fragment examined: MNHN DT 1022 - microscopic
Phlyctaenopora bitorquis Topsent, 1904b: 199 (by mono- slide made from the holotype.
typy). Description (from Topsent, 1904b). Convex crust
(Fig. 16A), tom at the underside (forcibly removed from the sub-
Definition stratum), 4 X 5 X 0.6 cm. Cream-coloured. Surface shiny-smooth,
bearing numerous unequal papillae of about 5 mm high. One large
Phlyctaenopora lacking spined microxeas. elevated oscule, with exhalant canals leading up to it. Ectosomal
Porifera· Demospongiae • Poecilosclerida • Mycalina • Mycalidae 689

Fig. 16. Subgenus Phlyctaenopora (Phlyctaenopora). A-B, Phlyctaenopora (Phlyctaenopora) bitorquis Topsent, 1904b. A, habit ofholotype reproduced
from Topsent, 1904b: pI. V fig. 24 (size see text). B, spicules of holotype reproduced from Topsent, 1904b: pI. XIV fig. 18 (sizes see text).
C, Phlyctaenopoa (Phlyctaenopora) halichondrioides Van Soest & Stentoft, 1988, SEM images of spicules of holotype (scale bars: anisochelae, 1 J.Lm;
megasc\eres, 10 J.Lm).

crust firm, 250-300 f.Lm in thicknes, made up of a tightly massed Type species
tangential spicule skeleton. Most ectosomal spicules are oxeas.
Choanosomal skeleton largely confused, with strongyles tending to Barbozia primitiva Dendy, 1922b: 132 (by subsequent
form ill-defined tracts, whereas oxeas are intercrossing at all designation herein).
angles. In the walls of the papillae the strongyles are arranged par-
allel to the axis of the papilla. Megascleres (Fig. 16B) oxeas, angu- Definition
larly curved twice, 300 X 10-12 f.Lm; strongyles slightly curved,
slightly thinner at the rounded ends, 360 X 12 f.Lm. Microscleres Phlyctaenopora with spined microxeas.
(Fig. 16B) anisochelae, usually spurred, 20-27 f.Lm; sigmas, abun-
dant, 38 f.Lm. Distribution and ecology. Azores, deep water. Description of type species
Remarks. The habit and ectosomal reinforcement indicate
that the sponge probably lives buried in the sediment with the Barbozia primitiva Dendy, 1922b (Fig. 17 A).
papillae and oscules raised above it. A second species of this sub- Synonymy. Barbozia primitiva Dendy, 1922b: 132, pI. VIII
genus, P. (P.) halichondrioides was described by Van Soest & fig. 9, pI. 18 fig. 1.
Stentoft (1988: 119, fig. 59), from Barbados. It differs from Material examined. None. Holotype (not examined):
P. (P.) bitorquis in lacking sigmas and having smaller sized oxeas BMNH 1921.11.7.111 - wet & slide, 'Sealark' Expedition, Indian
(Fig. 16C). Ocean RN.LXXXII.1.
Description (from Dendy, 1922b). Hemispherical sponge of
7.5 cm diameter and a height of 4.5 cm, with truncated fistules of
SUBGENUS BARBOZIA DENDY, 1922 4 mm on the upper surface. Oscules likewise conically elevated.
The ectosomal skeleton is a thick crust (up to 350 f.Lm in thickness)
Synonymy of mostly tangentially arranged megascleres. This is carried by
the peripheral columns of spicules of the choanosomal skeleton,
Barbozia Dendy, 1922b: 132. which consists of an irregular system of very thick spicule tracts.
690 Porifera· Demospongiae • Poecilosclerida • MycaJina • MycaJidae

Fig. 17. Subgenus Phlyctaenopora (Barbozia). A, Phlyctaenopora (Barbozia) primitiva (Dendy, 1922b), drawing of spicules reproduced from Dendy's
pI. 18 fig . I (sizes see text). B-D, Phlyctaenopora (Barbozia) bocagei Levi & Levi (l983b), SEM images of spicules of the type. B,SEM overview of
megasc1eres (scale bar 100 fLm). C, chela (scale bar 10 fLm), D, amphiaster-like microxeas (scale bar 10 fLm).

Megascleres (Fig. 17A), strongyles, slightly curved, 350 X 13 f..l.m, ACKNOWLEDGEMENTS

oxeas, often biangulate, some with proliferations along the
shaft, 266 X 10 f..l.m. Microscleres (Fig. 17A), palmate anisochelae, The following curators and collection managers helped locat-
spurred, numerous, 20 f..l.m; amphiaster-like microxeas, 36-57 f..l.m. ing specimens, sending them out on loan at our request, or supplied
Distribution and ecology. Amirante Islands, W Indian Ocean, collection numbers: Clare Valentine, Mary Spencer-Jones, Shirley
dredged at a depth of 40-153 m. Stone (BMNH), Judith Fournier (CMN), Ruth Desqueyroux-
Remarks. Levi & Levi (1983b: 955) described Faiindez (MHNG), Claude Levi (MNHN), Klaus Rtitzler and Kate
Phlyctaenopora bocagei from New Caledonia (here illustrated in Smith (USNM), Phil Alderslade (NTM), Alan Beu (NZGS),
Fig. 17B-D), differing from P. (Barbozia) primitiva in the lack of Manfred Grasshoff (SMF), Prof. Dzwillo (ZMH), Gerald Bakus
true strongyles, instead of which there are blunt-ended styles, and (USC). Financial support to EH to attend the Systema Porifera
in possessing two size categories of oxeas (Fig. 17B), the larger of Workshops was obtained from CAPES, CNPq, FAPERJ, and
which is 1050 X 80/Lm. Subsequently, Levi (1993: 58) reported the FAPESP, from Brazil, and the organisers of the International
same species again from New Caledonia, with one of the speci- Conference on Sponge Science (Otsu, Japan).
mens possessing 'serpentiform' oxeas. Levi (1993) suggested that
Phlyctaenopora and Barbozia are synonyms, which is here
partially confirmed by recognizing both at the subgenus level.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Merliidae Kirkpatrick, 1908

Eduardo Hajdu! & Rob W.M. Van Soest2

Merliidae Kirkpatrick (Demospongiae, Mycalina) is a monogeneric family of poecilosclerids with a 'sclerosponge' body plan. The cal-
careous basal skeleton of the single genus Merlia Kirkpatrick, 1908d has a distinct 'chaetetid' architecture and the mineral composition is
high magnesium calcite. The poecilosclerid affinity is evident from the siliceous spicules, particularly the unique microscleres called cla-
vidiscs, which bear structural resemblance to hamacanthid diancistras. Other spicular features, such as the possession of tylostyles and
commata-like raphides indicate relationships with desmacellid sponges. Thus, the family is assigned to the poecilosclerid suborder
Mycalina. Five species of Merlia have been named, including one fossil. Some species lack either the calcareous basal skeleton or the
clavidiscs.
Keywords: Porifera; Demospongiae; Poecilosclerida; Mycalina; Merliidae; Merlia.

DEFINITION, DIAGNOSIS, SCOPE by Van Soest (1984a) and Vacelet (1985), who pointed out that
the calcareous basal skeleton is facultative and certainly not of
Definition phylogenetic significance at the ordinal or class levels. Vacelet
(1980a, 1985) proposed assignment of an order Merliida to the sub-
Mycalina with calcareous basal skeleton and clavidisc class Tetractinomorpha, but Van Soest (1984a), Hoshino (1990),
microscleres. Reitner (1992) and Hajdu (1995) emphasized similarities of clavi-
discs and diancistras, and the commata-like raphides, and returned
Diagnosis Merlia to Poecilosclerida, either as a member of Hamacanthidae or
BiemnidaelDesmacellidae. In view of the fact that Merlia shows
Thin crusts consisting of a chaetetid calcareous basal skeleton affinities to both Hamacanthidae and Desmacellidae, and because the
(i.e., formed of a layered system of calcareous chambers), the outer calcareous basal skeleton is unique in Mycalina, it is proposed here
layer of which is filled with sponge tissue and siliceous spicules. to follow Kirkpatrick in assigning Merlia to a family of its own.
The basal skeleton is made up of high magnesium calcite and the
microstructure is of the 'water-jet' type. The spiculation consists of
thin tylostyles arranged in wispy plumose bundles, unique key-
hole type microscleres called clavidiscs, rugose rap hides, and MERLIA KIRKPATRICK, 1908
small commata-like spicules appearing monactinal.
Synonymy
Scope
Merlia Kirkpatrick, 1908d: 510. Noronha Kirkpatrick, 1909: 48.
A single genus, Merlia, with about five species.
Type species
Taxonomic history
Merlia normani Kirkpatrick, 1908d (by monotypy).
Kirkpatrick (1908d) discovered this remarkable organism and
immediately recognized its sponge nature, but misinterpreted it as Definition
a 'pharetronid' sponge because he attributed some foreign tuning-
fork spicules to it. In a follow-up study (Kirkpatrick, 1909) he As for definition of family.
assumed that the siliceous spicules were from an independent
organism, naming the latter Noronha sealariformis. However, after Diagnosis
extensive fieldwork and the study of many specimens he reached
the conclusion (Kirkpatrick, 1911) that the limestone base is Thin crusts with smooth surface, covering usually a calcare-
secreted by the siliceous sponge, and that the complex should be ous basal skeleton consisting of a system of layered chambers. The
classed as a sister-group to Desmacellinae. Subsequently, the sys- chambers of the outer layer are filled with choanosomal tissue and
tematic position of Merlia varied among authors (see Hoshino, spicules, the inner chambers are filled with undifferentiated cell
1990 for a table of assignments). Hartman & Goreau (1970) masses. The calcareous basal mass may be absent. Spicules consist
assigned Merlia to their class Sclerospongiae, erected for sponges usually of thin tylostyles, clavidiscs, raphides, and finely spined
with calcareous basal skeletons. This was subsequently contested monactines (commata). Clavidiscs may be absent.

691
692 Porifera· Demospongiae • PoeciloscIerida • Mycalina • Merliidae

D E
F)

Fig. 1. Merlia normani Kirkpatrick, 1908d. A, SEM image of cross section of calcareous basal skeleton (scale 100 ""m) (from Vace1et & Uriz, 1991).
B, SEM image of upper chamber layer (scale 100 ""m) (from Vacelet & Uriz, 1991). C-F, drawing of spicules (scale 15 ""m) (from Van Soest, 1984a).
G-H, SEM images of spicules. G, tylostyle (scale 10 ""m). H, clavidisc (scale I ""m) (from Gautret et al., 1991). I-J, rugose commata (scale 5 ""m) (I, from
Gautret et al., 1991; J, from Van Soest, 1984a).

Recent reviews of the blocks. Meshes are IS0-220 /-Lm in diameter, the cavities
being 120-150/-Lm across, and the walls 40-60/-Lm thick. Youngest
Vacelet, 19S0a: 227; Van Soest, 19S4a: 212; Hoshino, 1990; specimens with no crypts at all, the calcareous skeleton made of
Vacelet & Uriz, 1991: 171; Reitner, 1992: 236. slender bars of a wide-meshed polygonal reticulation. The number of
layers diminishes from centre to the edges of the sponge, where there
Description of type species may be a region devoid of the basal calcareous skeleton. Meshes sur-
rounded by 4-S tubercles. Tubercles are 75 /-Lm high and 75/-Lm
Merlia normani Kirkpatrick, 1905d (Fig. lA-J). broad, and bear sharp-pointed conu1es, about 10 /-Lm high and 16/-Lm
Synonymy. Merlia normani Kirkpatrick, 1905d: 510, broad at the base. The basal skeleton and its outer layer of chambers
pI. XV, figs 1001S; Noronha scalariformis Kirkpatrick, 1909: 4S. is filled with choanosomal tissue (choanocyte chambers, canal sys-
Material examined. Holotype: BMNH 1911.4.7.10 - tem) and covered by organic ectosome. The inner chambers are filled
Canon Norman's original type specimen, on rock, Porto Santo. with masses of archaeocytes which are assumed to have a function
Other material. BMNH 1911.4.7.2-9 (including the 1911 figured similar to that of gemmules. Siliceous skeleton. Upright bundles of
specimen 1911.4.7.7), BMNH 1954.2.21.115-116, 1954.6.3.2 - tylostyles with points outwards, along with a few raphides, forming
specimens and slides, all from Porto Santo. more or less vertical pillars of support. Megasclere bundles may be
Description (adopted from Kirkpatrick, 1911: 661). 'The seen lying transversely on the floor of an upper open crypt. Only
great majority of the specimens, of which there are about 1000, were very rarely spicules are seen in the lower crypts. Clavidiscs occur
obtained from sixty fathoms off Porto Santo Island.' Small incrus- everywhere, but are more abundant on the surface. Commata occur
tations, ca. 1 cm2, 1 mm thick, growing on shells, octocorals in the choanosome, especially close to the choanocyte chambers.
('branches of corallines'), foraminiferans and worm-tubes; bright- Spicules (Fig. lC-J). Tylostyles (Fig. IC, G), slender, commonly
vermillion in colour, smooth surface. Calcareous basal skeleton curved at the distal end, sometimes nearly straight, ca. 140 X I.S /-Lm
(Fig. lA-B) more-or-Iess regularly honeycombed, 'with blocks of thick, with tyles 5 X 2.2 11m. Clavidiscs (Fig. lD, H), ca. 45 X 30 11m,
reddish orange-coloured jelly filling in the spaces', with three or four, with a rim beveled inwards to a thin edge, ca. 3 11m thick. A key-hole
rarely five layers from edge to centre of the sponge. Blocks (crypts, shaped sinus or notch is present on the inner margin at each end of
chambers) in fairly regular horizontal and vertical rows; occasionally the long axis. Rugose raphides (Fig. IE), in trichodragmata ca.
some blocks of double breadth are found; regularity of horizontal SO 11m long. Monactine curved commata-like spicules (Fig. 1I-J),
rows frequently broken due to shorter or longer than average size rugose or spined, occuring singly or in bundles, ca. 15/-Lm.
Porifera· Demospongiae • Poecilosclerida • Mycalina • Merliidae 693

Remarks. Merlia is well distinguished from other mycaline forms may occur. The often markedly lobate head of the tylostyles
genera based on either the basal skeleton or the clavidiscs or both. in Terpios (e.g., Riitzler & Smith, 1993) should prevent such erro-
Noronha is a clear objective synonym as it was based on the mis- neous identifications.
taken assumption that the soft parts pertained to a different sponge
than the hard basal calcareous skeleton. Five species of Merlia
have been named, including one fossil, M. morlandi (Hinde & ACKNOWLEDGEMENTS
Holmes, 1892). Some species lack the generic synapomorphies: the
calcareous basal skeleton is lacking in M. deficiens Vacelet, 1980a The authors are thankful to Dr. Jean Vacelet (SMB) for valuable
and M. tenuis Hoshino, 1990, and the clavidiscs are lacking in discussions throughout the preparation of this manuscipt. EH is fur-
M. lipoclavidisca Vacelet & Uriz, 1991. The latter authors ther thankful to CAPES, CNPq, FAPERJ and FAPESP, from Brazil,
expressed concern for the possible inclusion of deficientllipocla- and the organizers of the International Conference of Sponge Science
vidisc species of Merlia in genera such as Terpios, but this could (Otsu, Japan), for financial support to attend the 'Systema Porifera'
apply to many other cases in the Poecilosclerida where reduced Workshops and to construct the means for writing this revision.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Podospongiidae De Laubenfels, 1936

Michelle Kellyl & Toufiek Samaai2

1 National Institute of Water & Atmospheric Research (NIWA) Ltd, Private Bag 109-695, Newmarket, Auckland, New Zealand.
(m.kelly@niwa.cri.nz)
2 Zoology Department, University of the Western Cape, Private Bag Xl7, Bellville, 7535, Cape Town, South Africa. (tsamaai@uwc.ac.za)

Podospongiidae de Laubenfels (Demospongiae, Poecilosclerida) is resurrected to include taxa previously referred to Latrunculiidae
Topsent based on the possession of spinose microrhabds (spinorhabds), as compared to acanthodiscorhabds, or 'chessman' spicules, in the
latter. The family includes Podospongia, Sigmosceptrella, Diacarnus and Negombata, defined as having spinorhabds in one or two size
classes disposed in a dense crust or scattered in a band within the ectosome. The family is included in Mycalina as incertae sedis based on
an interpretation that the protorhabd of spinorhabds is sigmoid and potentially a sigmancistra derivative. The family also has clearly dif-
ferentiated ectosomal and choanosomal skeletons; an extremely collagenous texture; where known, a viviparous reproductive strategy
incubating large conspicuous bright yellow larvae; oxeas, strongyles, and variants of these megascleres; and a range of distinctive
chemistries including norsesterterpene peroxides, to which cytotoxic latrunculins in Negombata are added, supporting postulated affini-
ties with Mycalina and clearly differentiating them from Latrunculiidae. Species of Podospongiidae are more common in tropical and
subtropical waters, occasionally also found in temperate and cold deep waters.
Keywords: Porifera; Demospongiae; Poecilosclerida; Podospongiidae; Podospongia; Sigmosceptrella; Diacarnus; Negombata.

DEFINITION, DIAGNOSIS, SCOPE incorporated into clear fibres (in Negombata). Microscleres are spin-
orhabds, often in two size categories, with two whorls of spines
Synonymy arranged serially around the spicule, between a proximal and distal
whorl of spines, one of which may be longer than the other. The pro-
Podospongiidae de Laubenfels, 1936a: 165. torhabd is sigmoid in Sigmosceptrella, Negombata, and Podospongia.
Microscleres are typically arranged in a compact crust or are scattered
Definition throughout the ectosomal region, with smaller categories also present
in the choanosome. Where known reproduction is viviparous, produc-
Mycalina incertae sedis with a plumoreticulate radiating ing huge bright yellow-orange larvae throughout the year that are
skeleton in which the primary fibres dominate and the secondary clearly visible to the unaided eye. Diacarnus and Sigmosceptrella are
fibres are reduced or absent. The ectosomal skeleton can be very known to produce the biologically active compounds norsesterter-
elaborate, forming dendritic umbelliform tracts that emanate from pene peroxides, and Negombata produces latrunculins.
the apices of primary fibres. Megascleres are generally strongyl-
oxeas but also include strongylote, tomote, stylote and oxeote mod- Scope
ifications. Spinorhabd microscleres in two size categories form a
crust or are scattered throughout the ectosome and choanosome. There are four valid genera, Podospongia, Sigmosceptrella,
Diacarnus, and Negombata.
Diagnosis
Distribution
Thickly encrusting, digitate branching, massive tubular, vasi-
form to stipitate sponges with microscopically smooth conulose Found throughout tropical to subtropical waters, occasionally
surface, and membranous oscules. Texture is tough, rubbery (almost in temperate to cold deep waters.
cartilaginous in Diacarnus), compressible and fleshy to the touch.
Colour in life is uniformly tan with surface pigmentation of maroon History and biology
and oak brown (Sigmosceptrella, Diacarnus), brilliant orange red
(Negombata), or creamy to orange (Podospongia). Structural Sigmosceptrella Dendy, 1922b, Podospongia du Bocage,
megascleres include strongyloxeas (often vestigial in Diacarnus), 1869 and Latrunculia du Bocage, 1869 were initially included in
robust curved anisostrongyles (Negombata) or more typically vari- a single family Latrunculiidae by Topsent (1922), defined as contain-
ants of styles, strongyles and oxeas (Podospongia). Ectosomal ing either spinose or discate microscleres and oxeote or stylote
skeleton with characteristic plumose 'starbursts' of spicule tracts, megascleres. Prior to the review of Kelly-Borges & Vacelet (1995)
with fine, dendritic individual tracts arising from the ascending Latrunculiidae was considered to be monophyletic based on the
choanosomal fibres and expanding into tiny brushes in the ectoso- common possession of these characteristic microscleres, even
mal membrane (most elaborate and umbelliform in arrangement in though these taxa represented at least three significantly different
Diacarnus and Sigmosceptrella). Choanosomal skeletal architecture skeletal architectures and many other forms of microscleres. Kelly-
dominated by huge thick hollow primary fibres interconnected by Borges & Vacelet (1995) recognised the non-homologous nature of
occasional short secondary fibres forming extremely elongate these various microscleres and illustrated the clear differences
narrow meshes (in Diacarnus and Sigmosceptrella), or fibroreticu- in the arrangements of their megasclere skeletons in each taxon,
late, forming a tight-meshed square reticulation of anisostrongyles suggesting that their combined presence in a single family was

694
Porifera· Demospongiae • Poecilosclerida • Mycalina • Podospongiidae 695

probably erroneous - although no alternative classification was and one species of Negombata have aciculospinorhabds with elon-
proposed at that time. gated spines on both apical whorls. There are usually two size cate-
This prior concept of Latrunculiidae has now been revised gories of spinorhabds. The protorhabd is sigmoid in Sigmosceptrella,
(Samaai & Kelly, this volume) to include only those taxa containing Negombata, and Podospongia, and straight in Diacarnus (Fig. 3).
discate acanthose micromabds (acanthodiscorhabds), that develop from These microscleres show no tendency to a vertical orientation as they
a straight protorhabd, and are located in a tightly packed palisade in the do in some Latrunculiidae, except in the case of Podospongia in
ectosomal membrane. In Latrunculiidae megascleres are anisostyles which the asymmetrical spinorhabds are vertically arranged in a pal-
arranged in a wispy reticulation and an ectosomal skeleton containing isade in the ectosome. The sigmoid tendency of the protorhabd
tangentially orientated megascleres. Latrunculiids also form thick is hypothesized as possible evidence for a close affinity to the
encrusting masses and all have areolate porefields and raised oscular Mycalina, as a potential sigmancistra derivative. Nevertheless,
fistules. Based on the observation of larvae in wtrunculia citharistae, Podospongiidae remains incertae sedis within Mycalina, in particu-
by Vacelet (1969), we conclude that the Latrunculiidae are viviparous. lar, and Poecilosclerida in general, whereas the family taxon is
They also contain bioactive compounds, the discorhabdins and their clearly differentiated from Latrunculiidae (as revised here).
derivatives, and pyroloquinoline alkaloids. These characters directly
contrast with those of Podospongiidae, in particular the possession of a Megascleres & skeletal architecture
sigmoid protomabd of the spinomabds, indicative of potential affinities
to Mycalina. Latrunculids also have a centre of diversity in the Southern All Podospongiidae have strongyloxeas as megascleres, and
Ocean, contrasting with the distribution of Podospongiidae found pre- Negombata and Podospongia have additional forms that include
dominantly in tropical and subtropical waters. stout curved anisostrongyles in the former, and strongyloxeas
with various stylote, tornote or oxeote modifications in latter.
Anisostrongyles in Negombata are restricted to the fibrous network
REVISION OF PODOSPONGIIDAE of the 'axial' region and they are embedded in spongin. The four
podospongiid genera are characterised by the common possession
Prior concepts of Latrunculiidae included genera now allo- of a fibroreticulate plumose skeleton, except in the case of
cated to Podospongiidae, based on the misinterpreted homology of Podospongia, which has tracts that radiate from a centrum into the
microrhabd microscleres (disc ate versus spinose microrhabds, globular mass and stalk. In Diacarnus huge spongin-reinforced
respectively), which are fundamentally different in their ontoge- fibres form elaborate dendritic umbelliform arrangements in the
netic or developmental stages (see Latrunculiidae chapter). ectosome, while in Sigmosceptrella, these are less elaborate. In
Podospongia (Topsent, 1928c; Boury-Esnault et al., 1994b; Levi, both these genera the primary fibres are huge and hollow with only
1993) Diacarnus, Sigmosceptrella and Negombata (Kelly-Borges minor secondary connections, whereas in Negombata the primary
& Vacelet, 1995) have tracts of strongyloxeas or variants of this and secondary fibres are not readily distinguishable. Podospongia
form of megascleres, and spinorhabds in the ectosomal region. The has a radiating stipitate morphology, and Negombata has an elabo-
spinorhabds are very similar across the group, with a tendency rate 'axial' fibroreticulation with 'extra-axial' radiating dendritic
towards asymmetry in Podospongia and Sigmosceptrella. Unlike tracts that diverge from the fibroreticulation towards the ectosome.
the latter three genera, which have an elaborate plumoreticulate
skeleton, Podospongia has very simple tracts of spicules that radi- Gross morphology
ate from a centrum in globular apex of the body (Fig. lA). These
tracts extend through the stalk to the base of the sponge where they Sigmosceptrella and Diacarnus form digitate lobate rambling
attach the sponge to the surface in a root-like structure. Further encrustations, large tubes and huge barrels in the case of Diacarnus.
comparisons between the families are provided in the chapter on Negombata species are digitate or foliose. An as-yet undescribed
Latrunculiidae (Samaai & Kelly, this volume). species from Indonesia is pedunculate. Podospongia is stipitate and
most species are tiny, being less than 60mm in length, but mostly less
Microsclere ontogenetic development, than 10 mm. None of the genera have any special surface features,
morphology and disposition although Podospongia has an apical oscule that is fringed by a palisade
of megascleres. These sponges are flexible, compressible and very
Microscleres of Podospongiidae are termed spinorhabds. These elastic due to their rich collagenous mesohyl matrix. The texture is
spicules dominate the ectosome in a compact crust, and are scattered smooth and flesh-like as the sponge ectosome is also charged with
in the choanosome. They are generally rod-shaped with four whorls abundant collagen.
of spines arranged serially along the spicule. Spinorhabds of
Negombata range from very regular and whorled as in Diacarnus, to Reproduction
irregular and spiraster-like (Fig. 3A-B). In Sigmosceptrella the spin-
orhabds are more-Of-less symmetrical, but the central whorls are sep- In several species of Sigmosceptrella, Diacarnus, and
arated by a much greater distance than they are from the apical and Negombata reproduction is viviparous, with characteristically huge
proximal whorls, giving the spicule a dumbbell shape (Fig. 3C-D). (2-3 mm diameter) bright yellow-orange larvae visible to the
In addition, one of these end groups of whorls is often expanded, unaided eye produced throughout the year (Fig. 2D). No reproduc-
forming an asymmetry along the spicule. In Diacarnus the spicules tive products have been observed in Podospongia.
are very regular with the four whorls separated equidistant along the
shaft, with the two distal whorls forming a spinose crown Pigmentation
(Fig. 3E-F). Podospongia, and one species of Negombata have
spinorhabds and asymmetrical aciculospinorhabds, which have an Podospongia, Sigmosceptrella and Diacarnus have a cream basal
elongated apical spine (Fig. lB--C). Some species of Podospongia pigmentation and a mottled maroon or oak brown pigmentation in
696 Porifera' Demospongiae • Poecilosclerida • Mycalina • Podospongiidae

the ectosomal membrane where they are exposed to light. Negomhata group characterised by the possession of spinorhabds, as opposed
ItUlgnifica and N. corticata are a brilliant orange red that is persistent to acanthodiscorhabds in Latrunculiidae (Kelly-Borges & Vacelet,
in ethanol preservative for some time (Kelly-Borges & Vacelet, 1995). 1995; Samaai & Kelly, this volume). This is considered to repre-
Podospongia spp are invariably described as cream or white in life. sent a fundamental difference in the ontogenetic development
of these microscleres, and not merely a morphometic variation
Biochemistry (see chapter on Latrunculiidae). In all genera except Diacamus the
protorhabd of the spinorhabd is sigmoid, or a derivative of a
Sigmosceptrella and Diacamus contain biologically active sigmoid shape. This ontogenetic development of microscleres,
norsesterterpene peroxides (see Urban et aZ., 2000; Kelly-Borges & coupled with the predominance of a pseudoaxial plumoreticulate
Vacelet, 1995). These include muqubilins recorded from a species skeleton, strongyloxeas and anisostrongyles with various tornote,
of Diacamus and specimens from the Red Sea identified as oxeote and stylote modifications and vivipary, indicate greater
'Prianos'. This latter identification is highly suspect and it is likely affinity with poecilosclerid sponges than with hadromerids
that this material belongs to Diacamus, for which species exist that (to which 'Latrunculiidae' of authors, i.e., Latrunculiidae +
have incomplete spicule complements, possessing only vestigial Podospongiidae, was previously assigned; see Samaai & Kelly, this
strongyloxeas similar to typical species of Prianos (containing volume). We consider that Podospongiidae has a greater affinity
only strongyles) (Urban et aZ., 2000). Cytotoxic latrunculins have with the poecilosclerid suborder Mycalina than with Latrunculiidae,
been described from Red Sea species of Negombata (Gillor et aZ., the latter assigned to its own suborder (Latrunculina subord. nov.)
2000). The chemistry of Podospongia is unknown. incertae sedis within Poecilosclerida. Closely related genera
PhZyctaenopora Topsent and Barbozia Dendy were also previously
Ecology, bathymetric and biogeographical distribution included within Latrunculiidae, but are now assigned incertae
sedis to the Mycalidae due to their possession of anisochelae.
Podospongia is predominantly a deep-water sponge collected While Podospongiidae do not have chelae microscleres or any
down to 600 m, typically found around 100 m or more. Species are other type of microsclere that could be construed as directly (or
known from New Caledonia, Natal coast of South Africa, north- obviously) linking them to the Mycalidae, their plumose, semi
eastern and central Atlantic Ocean. Three species of Negombata axial/extra-axial architecture of spongin-bound stylote or oxeote
are known from the shallow tropical western Indian Ocean and megascleres, and possession of a sigmoid protorhabd, suggest their
southeast Asian waters, Sigmosceptrella overlaps this distribution affinities lie most clearly with the Mycalina (particularly with
and extends from the Western Indian Ocean to cool temperate Cladhorizidae, Mycalidae).
southwest and eastern coasts of Australia. Diacamus is common This conclusion is supported by non-morphological evidence.
throughout southeast Asia and the central west Pacific, extending Norsesterterpene peroxides have also been found in a species
east to Fiji and south along the southwest, east and northeast coasts of MycaZe (Capon & Macleod, 1987a) and the suggested poecilo-
of Australia. Most species live in typical coral reef environments, sclerid affinity of Podospongiidae is strengthened by molecular
on dead coral substrate, down to depths of about 50 m. evidence. An 18S rDNA sequence of Negombata corticata
was consistently contained within the clade containing two
Phylogenetic affinities species of Clathria (Microcionidae, Poecilosclerida), rather
than Acanthochaetetes (Spirastrellidae, Hadromerida) (Adams
The four genera included in Podospongiidae (Podospongia, et aZ., 1999). Refer to further discussion in the chapter on
Diacamus, Sigmosceptrella and Negombata) form a cohesive Latrunculiidae.

KEY TO GENERA

(I) Choanosomal tracts radiating from a centrum, stipitate body .......................................................................................... Podospongia
Plumoreticulation ................................................................................................................................................................................. 2
(2) Megascleres styles in addition to strongyloxeas ................................................................................................................. Negombata
Megascleres strongyloxeas with various tornote, oxeote or stylote modifications ............................................................................. 3
(3) Huge hollow primary fibres with umbelliform ectosomal skeleton, protorhabds of microscleres straight ......................... Diacarnus
Thick dense primary tracts with robust ectosomal tracts, surface crust of spinorhabds, protorhabds of microscleres
sigmoid ............. .......... ....... ............... ........... ...... ................ .... ....... .......... ................. ... ....... ...... ............... ...... .......... ..... Sigmosceptrella

PODOSPONGIA DU BOCAGE, 1869

Synonymy Definition

[Lovenia) du Bocage, 1868 (preocc.). Podospongia du Podospongiidae with tracts of oxeas with various strongylote,
Bocage, 1869: 160. AZcyospongia de Laubenfels, 1934: 18. stylote, tornote and oxeote modifications radiating along the stalk
and within the elliptical expansion above the stalk; spinorhabds
Type species and aciculospinorhabds are packed in a vertical arrangement in the
ectosome and scattered throughout choanosome, protorhabds are
Podospongia loveni du Bocage, 1869: 160 (by monotypy). sigmoid.
Porifera· Demospongiae • Poecilosclerida • Mycalina • Podospongiidae 697

Fig. 1. Podospongia (A-C) and Negombata (D-F). A-B, Podospongia loveni Bocage (redrawn from du Bocage, 1869, pI. 10, fig. 1). A, whole sponge
(upper left) and vertical section showing radiating tracts of megascleres that emerge to form a solid stalk, and crypt-like apical oscule (upper right) (scale
10 mm). Acicu10spinorhabd (lower left) and spinorhabd (lower right) microsc1eres that form a crust in the ectosome and scattered throughout the
choanosome (scale 30 fLm). B, (paratype MNHN LBIM.DT 1115) aciculospinorhabds and spinorhabds (scale 40 fLm). C, Alcyospongia india de Laubenfels,
1934 (holotype USNM 22365), acicu10spinorhabds with prominent apical spine (scale 40 fLm). D-E, Negombata corticata (Carter) (holotype BMNH
1840.6.56-58; after Kelly-Borges & Vacelet, 1995). D, ho!otype (scale 50 mm). E, ectosomal and choanosomal skeleton (scale 600 fLm). F, Negombata mag-
nifica (Keller) (schizotype BMNH 1908.9.24.118, after Kelly-Borges & Vace1et, 1995), ectosomal and choanosoma1 skeleton (scale 60 fLm).

Diagnosis Description of type species

Predominantly stipitate with an elliptical head, usually less Podospongia loveni du Bocage, 1869 (Fig. lA-B).
than 6cm in total length, typically less than 20mm long, apical Synonymy. [Lovenial borealis du Bocage, 1868: 37 [nom-
oscule often with a fringe of megascleres, possibly rarely thinly enD blituml. Podospongia loveni du Bocage, 1869: 160.
encrusting. Megascleres are modified oxeas radiating from a cen- Material examined. Holotype: Lost.
trum in uniform tracts extending into the stalk where they extend to Description (modified from du Bocage, 1869). Sponge
root-like processes for attachment. Microscleres are smooth composed of a curved vertical stem and an elliptical head, strongly
straight spinorhabds of a variety of forms including asymmetrical attached to the substratum by a large base not divided into root-like
and symmetrical forms, and aciculospinorhabds; reproduction and processes (Fig. IA). There is a single apical oscule. The surface of
biochemistry unknown; colour in life uniformly cream to white. the elliptical portion has numerous depressions and is uneven and
bristling with the emergent tips of the megasclere tracts. The ecto-
Previous reviews somal region is distinctive with a layer of spinorhabd microscleres
that are aligned vertically in the ectosome. The skeleton consists of
Topsent (1922, 1928c); de Laubenfels (1936a); Boury-Esnault radiating tracts of megascleres (which appear from the illustrations
et al. (1994b). to be a mixture of oxeas, strongyles and strongyloxeas, 400-500 f.1m
698 Porifera' Demospongiae • Poecilosclerida • Mycalina • Podospongiidae

long) which extend into the axis of the sponge (Fig. lA). Type species
Microscleres are scattered in the choanosome as well (Fig. lA).
Remarks. Podospongia was erected by du Bocage (1869) to Latrunculia corticata Carter, 1879b: 298 (by original
replace the genus [Lovenia], which he had earlier erected for Lovenia designation).
borealis du Bocage (du Bocage, 1868). The genus name was preoc-
cupied by Lovenia Desor, 1847, an echinoderm. Theoretically the Definition
species name P. borealis has priority over the name P. loveni.
However, the former has not been used after 1899 and therefore Podospongiidae with a small square-meshed reticulation of
prevailing usage is maintained (ICZN Art. 23.9; Anon., 1999).' strongyloxeas and robust styles bound into clear spongin, dendritic
The holotype has since been confirmed destroyed. plumose tracts arise from this 'axial' fibroreticulation in an 'extra-
It is not necessary to emend the genus diagnosis as the origi- axial' region, microscleres are irregular spinorhabds, these are
nal description and subsequent descriptions of Podospongia loveni packed in the outer ectosome and are abundant throughout the
(Topsent, 1928c; Boury-Esnault et al., 1994b) and other species choanosome, protorhabds pseudo-sigmoid (recurved).
(Levi, 1993; Kirkpatrick, 1903) are adequate to provide a firm con-
cept of the species, and consequently also of the genus. A slide of Diagnosis
spicules of a paratype specimen of Podospongia loveni, from
Portugal, supplied by Professor Claude Levi, conforms closely to Erect, foliose, pedunculate, digitate, branching. Compressible,
du Bocage's original description (Fig. lB). The genus is charac- with an extremely smooth surface through which large pores are
terised by its stipitate gross morphology, with unusual modified clearly visible. Megasclere skeleton consists of uniform, elongate,
oxeote spicules radiating in tracts from a centrum in the centre of to square-meshed reticulation of clear spongin cored fairly irregu-
the globular head. Spicule tracts are aligned longitudinally within larly with thick, slightly curved oxea with strongylote ends, and
the stalk or peduncle. In some species, the base of the peduncle a clear hollow axial canal. Ectosome with thin fusiform wavy
forms fine rhizoid-like structures for attachment to the substrate. oxeas in tracts that fan within a thick collagenous ectosome, ulti-
Aciculospinorhabds form a crust within the ectosome. mately forming surface brushes. Microscleres, irregular spin-
De Laubenfels (1934) considered Alcyospongia (type species A. orhabds in two sizes, largest very irregular, thick, without a distinct
india de Laubenfels, 1934: 18) to belong to Latrunculiidae because of shaft, smaller, with a distinct straight or curved shaft with spines of
its apparent relationship to the genus Podospongia. This West Indies uneven length arranged roughly in two central whorls and two ter-
specimen (USNM 22365) was differentiated from Podospongia by its minal bunches, terminally commonly double. Immature microscle-
possession of 'straight streptasters', and the fact that the centrum from res, straight rods in which one or more spines from apices are
which the tracts radiated was at the base of the globular mass, rather strongly recurved, disposed predominantly on the extreme external
than at its centre, as in Podospongia. In all other characters, the sponge surface of the ectosome, but can also be very dense within the
is similar to species of Podospongia. Examination of the type speci- choanosome (modified from Kelly-Borges & Vacelet, 1995).
men revealed that the centrum was depressed in the globular region,
but we consider that it could easily be the result of dehydration as an Previous reviews
artifact of preservation. A new spicule preparation revealed that the
'straight streptasters' were foreign, and very similar to those that De Laubenfels (1936a); Hooper (1986); Kelly-Borges &
would occur in astrophorid families, and that the sponge had asym- Vacelet (1995).
metrical spinorhabds as in other species of Podospongia (Fig. Ie).
Sigmoid protorhabds were also present in the new spiCUle preparation. Description of type species
De Laubenfels (1934) regarded the new genus assignment as doubt-
ful, considering that the position of point of radiation of the tracts may Negombata corticata (Carter, 1879b) (Fig. ID-E).
not be that important phylogenetically. Given that the streptasters are Synonymy. Latrunculia corticata Carter, 1879b: 298.
foreign inclusions or may have been from another sponge at the time Latrunculia tarentina Pulitzer-Finali, 1983: 513.
of collection, we concur with de Laubenfels' doubts and hereby for- Material examined. Holotype: BMNH 1840.5.6.56-58.
mally relegate Alcyospongia into synonymy with Podospongia. Description (modified from Kelly-Borges & Vacelet, 1995).
Sponge (in the dried state) is erect and composed of short, curved,
Distribution flanged, narrow fans of different lengths with a restricted base of
attachment (Fig. ID). Surface texture of dry sponge is fibrous and
Podospongia is generally found in deeper water down to 6OOm, roughened; Carter (1879b) described the fresh sponge as 'chon-
from the Natal coast of South Africa, New Caledonia and the droid' and 'smooth' with pores 20/-Lm diameter and 74/-Lm apart.
central and north Atlantic. Latrunculia normani Stephens is also most Colour yellowish white in dry sponge. The choanosomal skeleton
probably a species of Podospongia but it is thinly encrusting, the only consists of a central axis of rectangular meshes formed by spongin
character that does not conform to the diagnosis of the genus. fibres, 300-600/-Lm diameter, not clearly differentiated into pri-
mary and secondary tracts (Fig. IE). Megascleres are not arranged
uniformly within fibres, but rather, are scattered singly or in
groups, and oblique or occasionally perpendicular to the fibre axis.
NEGOMBATA DE LAUBENFELS, 1936 An ectosomal skeleton of wavy oxeas is present, but this has col-
lapsed in the holotype, rendering the arrangement difficult to deter-
Synonymy mine. The holotype of Negombata magnifica (Keller) shows this
feature more clearly (Fig. IF). Microscleres are densely packed on
Negombata de Laubenfels, 1936a: 159. the sponge surface and also in the choanosome, almost obscuring
Porifera· Demospongiae • Poecilosclerida • Mycalina • Podospongiidae 699

fibres (Kelly-Borges & Vacelet, 1995). Megascleres are oxeas, apical with low fleshy raised collars. Texture extremely tough,
being thick, straight or slightly curved with rounded strongylote almost cartilaginous, fleshy, elastic. Colour in life cream base with
ends, 346(317-384) X 10-12fLm, and thin with ends fusiform, pale purple-pink, and mottled with deep reddish brown and yellow
422(394-451) X 2.4-6 fLm. Microscleres are spinorhabds of 2 size brown. Skeletal arrangement plumoreticulate with huge thick hol-
classes, having slightly curved or spiraled spines of uneven lengths, low fibres optically visible, connected sparsely by thin secondary
26-35 /-Lm in length (Fig. 3A-B). fibres. Ectosome dense, highly collagenous, distinct from the
Remarks. Negombata is clearly differentiated from underlying choanosome, with layers of collencytes and collagen
Latrunculia (Latrunculiidae) by its regular axial fibrous reticula- fibrils parallel to the surface. Megascleres are strongyloxeas, with a
tion, a dendritic-plumose extra-axial skeleton, and irregular spino- faint subterminal swelling at the proximal end, with the distal end
rhabds (Kelly-Borges & Vacelet, 1995). Negombata also differs oxeote or more typically strongylote. Microscleres are two size
from Diacamus, Sigmosceptrella and Podospongia in having two categories of spinorhabds. Sponges incubate huge bright orange-
forms of strongyloxea megascleres, one of which is robust, curved, yellow larvae.
anisostrongyle which is always embedded in the spongin of the
axis. In N. kenyensis these spicules have stylote modifications to Previous reviews
one end rendering their appearance style-like. Microscleres of
Negombata are spinorhabds and are densely packed in the sponge Carter (1 879b); Hooper (1986); nan (1995); Kelly-Borges &
ectosome, but they are irregular and the spines are spiraled rather Vacelet (1995).
than serially arranged as in the other three genera. A particular
characteristic of this genus is the faintly translucent glass-smooth Description of type species
surface, noted by Keller (1889), when he described N. magnifica.
Diacamus spinipoculum (Carter) (Fig. 2A-C).
Distribution Synonymy. Axos spinipoculum Carter, 1879b: 286; Diacamus
spinipoculum; Burton, 1934: 549; Latrunculia spinipoculum; Hooper,
Negombata has a similar distribution to Sigmosceptrella (see 1986: 181. Latrunculia hallmanni Wiedenmayer, 1989: 41.
below), although species do not extend into the West Central Material examined. Holotype: BMNH 1846.10.14.174.
Pacific or into Australia. Negombata is represented so far by four Description (modified from Kelly-Borges & Vacelet, 1995).
species, three of which have a brilliant orange-red colouration and Large spherical to oval barrel-shaped sponge with a deep central
a digitate or leafy gross morphology: Negombata magnifica and atrium, oscules, scattered on interior walls of atrium (Fig. 2A).
N. corticata from the Red Sea; Negombata kenyensis Pulitzer-Finali Surface heavily tuberculate to mammillate, rubbery and micro-
from Zanzibar; and a new pedunculate species still undescribed scopically smooth to touch. Colour in life, mahogany brown, mot-
from Indonesia, with a cream-orange colouration. tled with pale pinkish red, rim of apical atrium and atrium walls
cream-white. Cream in ethanol. The choanosomal skeleton con-
sists of large primary fibres, 500-900 /-Lm diameter, which radiate
towards the surface, connected by rare short secondary fibres,
DIACARNUS BURTON, 1934 250-300/-Lm diameter, or anastomosing with adjacent primary
fibres in the deeper choanosome (Fig. 2B). Primary fibre tip
Synonymy divides into an umbel of numerous slender dendritic spicule tracts,
38-70/-Lm, where the primary fibres passes through ectosome-
Diacamus Burton, 1934: 549. choanosome boundary. Dendritic spicule tracts occasionally
branch below surface and diverge and ramify within ectosome,
Type species where they form slightly fanned brushes (see Fig. 2C). Ectosomal
skeleton, 1200-1500 fLm deep, extremely dense with parallel colla-
Axos spinipoculum Carter, 1879b: 286 (by original designation). gen fibrils, easily distinguished from underlying choanosome
which is dense and soft. A plumose ectosomal skeleton of wavy
Definition oxeas is present, but this has collapsed in the holotype, rendering
the arrangement difficult to determine. Larger microscleres are
Podospongiidae with strongyloxeas arranged in huge hollow scattered predominantly around edges of exhalant canals in
spongin-bound primary fibres, connected by sparse thin secondary choanosome, around ectosomal lacunae, and just below lower
fibres, umbelliform dendritic arrangements of thin tracts emerge boundary of ectosome where smaller thinner spinorhabds are also
from the apex of each primary fibre. Microscleres are regular spino- found. Megascleres are strongyloxeas with slight subterminal
rhabds with whorls of spines arranged equidistant along the shaft swellings, 57(53-65) X 2(1-5)/-Lm. Microscleres are spinorhabds
of the spicule, forming a crust at the surface of the sponge or in a of 2 size classes, 46-57 fLm in length (Fig. 3E-F).
mid-ectosomal band, and are scattered throughout the choanosome, Remarks. In a revision of Axos Gray (Hemiasterellidae,
protorhabds straight. Hadromerida) Hooper (1986) transferred Axos spinipoculum to
Latrunculia in recognition of the superficial similarity of the gen-
Diagnosis (modified from Kelly-Borges & Vaceiet, 1995) eral skeletal and spicule features to those of Latrunculia sensu lato.
Kelly-Borges & Vacelet (1995) progressed this idea further by
Massive, barrel-shaped, tubular or lobate-digitate sponges. reassigning this species to a revised concept of Burton's
Surface with single or multiple conules or blunt broad tubercles or Diacamus. They also noted that the skeletal arrangements,
mounds, microscopically smooth, slippery and rubbery. Ostia radi- microsclere morphology and arrangement, colouration and texture
ate in stellate formation in shallow rounded depressions, oscules differed from all known species of Latrunculia, and that Diacamus
700 Porifera· Demospongiae • PoeciloscIerida • MycaIina • Podospongiidae

Fig. 2. Diacarnus (A-D) and Sigmosceptrella (E-F). A-B, Diacarnus spinipoculum (Carter) (holotype BMNH 1846.10.14.174, after Kelly-Borges &
Vacelet, 1995). A, holotype (scale SOmm). B, ectosomal and upper choanosomal skeleton (scale 130 J..Lm). C, Diacarnus bellae Kelly-Borges & Vacelet
(holotype BMNH 1994.S.22.1I, after Kelly-Borges & Vacelet, 1995), ectosomal and upper choanosomal skeleton (scale 150J..Lm). D, Diacarnus
megaspinorhabdosa Kelly-Borges & Vacelet (specimen QMG30S013, after Kelly-Borges & Vacelet, 1995), incubated larva (scale 2 mm).
E, Sigmosceptrella quadrilobata Dendy (holotype BMNH 1925.11.1.1641), skeletal arrangement (scale 60 J..Lm). F, Sigmosceptrella fibrosa Dendy
(holotype BMNH 1925.11.1.717) (scale 50mm).

spinipoculum possibly represented a family separate from are difficult to differentiate, unlike Sigmosceptrella and
Latrunculiidae. Species of Diacarnus are particularly common Negombata that are represented by discrete, easily recognisable
across the Indo-Pacific and it is surprising that the genus had not species - a phenomenon that has been noted for Western Indian
been differentiated from typicallatrunculiids prior to the review of Ocean Porifera in general (Kelly-Borges & Valentine, 1995). This
Kelly-Borges & Vacelet (1995). These sponges are very characteris in complete contrast to the poriferan fauna of Southeast Asia,
istic as they produce huge yolk-like yellow-orange larvae which Australasia, Micronesia, and the West Central Pacific where, apart
are 2-3 mm in diameter, these appear to be present year round from the presence of the same very discrete, easily recognisable
(Fig. 2D) (see Ilan, 1995). Diacamus have an extremely collagenous species, many genera have numerous sibling species which are dif-
texture and feel and look somewhat like uncooked chicken in life, ficult to differentiate based solely on skeletal features. This is par-
complete with sinew-like cartilagenous fibres. Most species of ticularly evident for genera of Chalinidae, Niphatidae, Petrosiidae,
Diacamus were assigned to Latrunculia prior to the review of Callyspongiidae, Halichondriidae, Microcionidae and others.
Kelly-Borges & Vacelet (1995). Diacamus differs from other The genus Sceptrintus Topsent, 1898b (Topsent, 1898b: 239;
members of Podospongiidae in that it has numerous species that 1904b: 117, pI. 1, fig. 11, pI. 12, fig 4) from the Azores was originally
Porifera· Demospongiae • Poecilosclerida • Mycalina • Podospongiidae 701

Fig. 3. Spinorhabds and immature protorhabds of Negombata, Sigmosceptrella and Diacamus. A, Negombata magnifica (Keller), mature spinorhabd.
B, Negombata magnifica (Keller), semi-sigmoid protorhabd. C, Sigmosceptrella quadrilobata Dendy, mature spinorhabd. D, Sigmosceptrella quadrilobata
Dendy, sigmoid protorhabd. E, Diacamus ardoulwbae Kelly-Borges & Vaceiet, mature spinorhabd. F, Diacamus ardoukobae Kelly-Borges & Vacelet,
mature spinorhabd (scale 8 j-Lm).

assigned to the Spirastrellidae, but is transferred here to SIGMOSCEPTRELLA DENDY, 1922

Podospongiidae with some reservation. The type species,
Sceptrintus richardi Topsent (by monotypy), has styles that are Synonymy
lightly spined, being occasionally verticillate, and spinorhabd-like
microscleres of 2 sizes. One of the microscleres is huge, being as Sigmosceptrella Dendy, 1922b: 136.
large, or larger than the megascleres. The choanosome is domi-
nated by discasters of all sizes scattered abundantly without orien- Type species
tation between loose tracts of styles. There are no visible fibres,
rendering the texture of the sponge firm but friable. At the surface Spirastrellajibrosa Dendy, 1897: 254 (by original designation).
of the sponge discasters and styles appear to be tangential and
paratangential, the styles being grouped into small bundles that do Definition
not project through the surface. Discasters are smaller at the sur-
face. The sponge is massive and very smooth with no visible Podospongiidae with well-defined plumose tracts of strongy-
oscules or papillae, reminding Topsent of the genus Latrunculia. loxeas, no secondary fibres are apparent, radiating and diverging
The colouration appears to be mottled yellow. The proposed tenta- from a spongin base, in one case in the form of a candelabra.
tive affinity of Sceptrintus with Podospongiidae is based upon the Microscleres are asymmetrical spinorhabds with sigmoid pro-
similarity of the microscleres and their protorhabds, with the spin- torhabds, forming a thick crust at the surface of the sponge and
orhabd form of Diacamus in particular. The surface morphology densely scattered within the choanosome.
and colouration seem to be similar as well. However, the styles are
quite different as they are lightly spined and the microscleres are Diagnosis
enormous compared to those of Diacamus species. The skeletal
details are also quite different from Diacamus, Sigmosceptrella Thickly encrusting, lobate, semi-digitate sponge with a smooth
and Negombata. A more definite assignment may be arrived at to faintly conulose surface. The choanosomal skeleton consists of
through examination of fresh material. large diverging dendritic or plumose tracts that are not joined by
secondary fibres, diverging to form brushes in the ectosome.
Distribution Megascleres are strongyloxeas. The ectosome is packed with abun-
dant spinorhabds which are characteristically dumbbell shaped and
Diacamus contains numerous species, some of which are rea- often asymmetrical.
sonably easy to differentiate in the field based on their gross mor-
phology. However, for most species, a combination of the gross Previous reviews
morphology, architecture of the ectosomal region, morphology of
microscleres and their disposition within the ectosome, provide Dendy (1921); Dendy (1922b); Kelly-Borges & Vacelet (1995).
the clearest indication of species (Kelly-Borges & Vacelet,
1995). Species diversity is highest in Papua New Guinea and the Description of type species
Philippines, with different species recorded from the southwest
Pacific (New Caledonia) to the Great Barrier Reef, and Micronesia. Spirastrellajibrosa Dendy, 1897: 254 (Fig. 2F).
702 Porifera· Demospongiae • Poecilosclerida • MycaIina • Podospongiidae

Synonymy. Spirastrella fibrosa Dendy, 1897: 254. Sigmosceptrella clearly fits within the revised concept of
Latrunculia conulosa Hallmann, 1912: 126. Podospongiidae, and is probably most closely related to Diacamus
Materinl examined. Holotype: BMNH 1925.11.1.717. in that it has plumose tracts of megascleres that diverge to form
Description. Sponge forms a thick lobate mass, with dendritic fibres in the ectosome and brushes at the surface. While
a conulose surface, the texture is rubbery (Fig. 2F). The choanoso- the choanosomal tracts are less elaborate than those in Diacamus
mal skeleton is composed of strongyloxeas which form dense well- spp., and the spinorhabds are dumbbell-shaped and occasionally
defined fibres which spring from the base of the sponge, diverging aciculate, the general pattern of architecture is very similar within
to form brushes, or candelabra, below the ectosome. The ectosome the family as a whole. The protorhabds are also sigmoid.
is well-defined as it is packed with spinorhabds and dense with
collagen. The ectosome is not detachable. Distribution
Remarks. Sigmosceptrella was first introduced by Dendy
(1921), in a description of characteristic spicules but without men- Sigmosceptrella are less diverse and have more discrete
tion or a description of the type species (and thus a nomen nudum), species distributions than Diacamus, widely dispersed in cool
and so the genus was not formally described until Dendy (I922b). temperate waters of New South Wales and western Australia
In describing Sigmosceptrella quadrilobata Dendy, 1922b, from (S.fibrosa) and the Western Indian Ocean (S. quadrilobata).
Mauritius, Dendy stated that he considered his 'Spirastrella' fibrosa
Dendy, 1897, from Port Phillip Heads, Australia, was the true
holotype of Sigmosceptrella. Figure 2E illustrates the skeletal ACKNOWLEDGEMENTS
organisation of the holotype of Sigmosceptrella quadrilobata, an
encrusting species. The skeletal arrangement of the type species, We are indebted to The Royal Society London and The
Sigmosceptrella fibrosa, is very similar but branching of the pri- National Research Foundation, South Africa, for provision of
mary fibre occurs more towards the apex of the fibre, giving the a PhD scholarship for TS, and for general funding under their joint
appearance of an unruly candelabra. Examination of the type Capacity-Building Programme (1996-2001). We also thank the
species Sigmosceptrella fibrosa and S. quadrilobata reveals charac- Natural History Museum, London for field and laboratory support,
teristics that clearly differentiate Sigmosceptrella from Diacamus when the bulk of this research was carried out. The National
and Negombata. The largest microscleres of Sigmosceptrella are Institute of Water & Atmospheric Research (NIWA), New Zealand,
very similar to those of Diacamus and Negombata, but the spicules has supported the preparation of this contribution. We are indebted
are dumbbell-shaped and one end of the spinorhabd is enlarged and to The Coral Reef Research Foundation, Micronesia, for collection
thus asymmetrical (Fig. 3 C-D). These microscleres are also packed of new sponge material that has advanced our knowledge of
densely in the ectosome and rather than occurring in a scattered Podospongiidae diversity across the Indo-Pacific. This is a contri-
band within the ectosome (like Diacamus and Negombata). bution from the Coral Reef Research Foundation, Micronesia.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob w.M. Van Soest
© Kluwer AcademicIPlenum Publishers, New York, 2002

Family Isodictyidae Dendy, 1924

Eduardo Hajdu l & Gisele Lobo-Hajdu2

1 Museu Nacional, Departamento de Invertebrados, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, sIn, 20940-040, Rio de
Janeiro, RJ, Brazil & Centro de Biologia Marinha, Universidade de Slio Paulo, Slio Sebastilio, SP, Brazil. - (hajdu@acd.ufij.br)
2 Departamento de Biologia Celular e Genetica (DBCG), Instituto de Biologia Roberto Alcantara Gomes (mRAG), Universidade do
Estado do Rio de Janeiro, Rio de Janeiro, RJ, Brazil. (glhajdu@uerj.br)

Isodictyidae Dendy, (Demospongiae, Poecilosclerida) contains two genera, Coelocarteria and Isodictya, allocated to suborder Mycalina.
Ichnodonax is considered ajunior synonym of Coelocarteria. Cercidochela. Homoeodictya. Neoesperiopsis. Textiliforma and Valentis are
considered junior synonyms of Isodictya.
Keywords: Porifera; Demospongiae; Poecilosclerida; Mycalina; Isodictyidae; Coelocarteria; Isodictya.

DIAGNOSIS, SCOPE Taxonomic remarks

Synonymy Isodictya has been the focus of some recent ping-pong assign-
ments in the poriferan classification. Conflicting hypotheses were
Isodictyeae Dendy, 1924: 334. proposed by Hajdu et al. (1994a) and Samaai et al. (1999), postu-
lating niphatid (Niphatidae, Haplosclerida) or mycalid (Mycalidae.
Diagnosis Poecilosclerida) affinities, respectively. As Samaai et al. (1999)
undertook a most rigorous formal analysis on the affinities of
Mycalina with (mostly) diactinal megascleres arranged in a Isodictya their results are followed here. Accordingly, Isodictya
(p1umo)reticulate skeletal architecture (niphatid- or phloeodictyid- and allied taxa are included within the Mycalina. Their suggestion
like) and palmate isochelae. of closer relationships between Isodictya and mycalid genera,
such as Esperiopsis and Mycale, needs further investigation, and
Scope consequently Isodictya has not been accommodated within the
Mycalidae but referred to Dendy's (1924) 'section' Isodictyeae,
The family contains eight nominal and two valid genera, which is here promoted to family-level status. Refer to the remarks
Coelocarteria and Isodictya, with Ichnodonax included as a junior on Isodictya for further comments on the interpretation of the
synonym of Coelocarteria, and Cercidochela. Homoeodictya. phylogenetic affinities of the genus.
Neoesperiopsis. Textiliforma and Valentis included in Isodictya.

KEY TO GENERA

(1) Reticulated choanosomal skeleton with thick tracts of oxeas, ectosomal skeleton a dense tangential reticulation of strongyles,
Fistulose .......................................................................................................................................................................... Coelocarteria
Reticulated choanosomal skeleton with variably thick tracts of oxeas (rarely styles), ectosomal skeleton with tufts of the same
spicules, digitiforrnlflabellate forms are common .................................................................................................................. Isodictya

COELOCARTERIA BURTON, 1934 Description of type species

Synonymy Coelocarteria singaporensis (Carter, 1883b) (Fig. 1).

Synonymy. Phloeodictyon singaporense Carter, 1883b:
Coelocarteria Burton, 1934a: 563. Ichnodonax de 326. Histoderma vesiculatum Dendy, 1905: 166. Ichnodonax
Laubenfels, 1954: 112. kapne de Laubenfels, 1954: 112.
Material examined. Holotype: BMNH (not seen). Other
Type species material. ZMA 790-807, 810 - 'Siboga' Exped., Indonesia, det.
M. Burton. ZMA 6501, 6511, 8556, 8594, 8625, 8690, 8840,
Phloeodictyon singaporense Carter, 1883b: 326 (by original 9279 - 'Snellius II' Exped., Indonesia, det. R.W.M. Van Soest.
designation). ZMA 11417 -Madang, Papua New Guinea, det. R.W.M. Van Soest.
Description (adapted/rom Dendy, 1905: 166, as Histoderma
Diagnosis vesiculatum; Bergquist, 1965: 161, compilation o/literature data
and own observation on p. 162). Body of varied morphology;
Isodictyidae with fistules; a phloeodictyid-like architecture subhemispherical, tubular, barrel-shaped, oval, cushion-like; with
with a dense, but neat reticulation of plurispicular tracts of fistules up to 6.5 cm high (Fig. 1A). Colour alive, yellow to brown,
strongyles and/or oxeas. turning to dark-brown, or purplish color in spirit. Ectosomal

703
704 Porifera· Demospongiae • Poecilosclerida • Mycalina • Isodictyidae

Fig. 1. A-G, Coelocarteria singaporensis (Carter, 1883b as Phloeodictyan). A, live-specimen in the intertidal at Sumba (Indonesia, 'Snellius II' Exped.,
photo & del. RW.M. Van Soest, ZMA 6501) (scale 5 em). B-C, stout reticulation of multi spicular tracts of megascleres (ZMA 8840, del. RW.M. Van
Soest) (scale 500 fLm). D-E, drawings of megasc1eres (from Ridley, 1884a, pI. XLI fig. s, as Rhizachalina singaporensis) (scale ca. 50 fLm). F, palmate
isoche1a (ZMA 8840, det. R.W.M. Van Soest) (scale ). G, toxa (ZMA 8840, del. RW.M. Van Soest) (scale 10 fLm).

skeleton a single tangential layer of strongyles packed together diagnosis to a reticulate choanosomal architecture in addition to
with a few larger oxeas. Choanosomal skeleton mainly a dense arcuate chelae, in view of the occurrence of smooth ectosomal
phloeodictyid reticulation of plurispicular tracts up to ca. 300/Lm tylotes in the type species of Hymedesmia Bowerbank, 1864, viz.,
thick (Fig. lB-C). Megascleres: Oxeas, 300/Lm long and 17 /Lm H. zetlandica Bowerbank, 1864. Consequently, Coelocarteria had
thick in the holotype, 237-312/Lm long and 3.5-16/Lm thick in to be transferred from this redefined taxon. Its possession of
other records (Fig. ID). Strongyles, 40-80 /Lm long and 4/Lm thick palmate chelae, as well as absence of acanthostyles or any acces-
in the holotype, 30-120 /Lm long and 3.5-12 /Lm thick in other sory megascleres, suggested inclusion within the Mycalina, more
specimens (Fig. IE). Microscleres: palmate isochelae, 14-17.5 /Lm precisely, within one of three available ill-defined families, viz.,
long in specimens other than the type. Desmacellidae Ridley & Dendy, 1886, Isodictyidae or Mycalidae
Remarks. The specimen ZMA 8840 studied under SEM Lundbeck, 1905. The reticulated choanosomal archi-tecture of
by Hajdu et al. (1994b) had palmate isochelae with a few spines on diactinal megascleres and possession of palmate isochelae suggest
the inner side of their shafts (Fig. IF), as well as rare toxas, about a closer affinity with Isodictya than to any of these other taxa.
50/Lm long (Fig. 1G). Coelocarteria has been assigned to the
Coelosphaeridae by recent authors (e.g., Bergquist, 1965; Van
Soest, 1984) on the basis of its fistular habit. Hajdu et al. (1994a,b) ISODICTYA BOWERBANK, 1864
argued against a synapomorphic value for the fistulae in sponge
phylogeny, suggesting an emphasis on anatomical features Synonymy
when higher taxa are circumscribed within the Poecilosclerida
and Haplosclerida. Accordingly, a new interpretation of the Isodictya Bowerbank, 1864: 197. Homoeodictya Ehlers, 1870:
Coelosphaeridae was advanced, where the diagnosis was based on 32. Textiliforma Carter, 1885d: 288. Cercidochela Kirkpatrick,
the combined possession of smooth ectosomal tylotes and arcuate 1907: 284. Valentis de Laubenfels, 1936: 96. Neoesperiopsis de
chelae (Hajdu et al., 1994a). Van Soest (this volume), shifted the Laubenfels, 1949a: 15.
Porifera· Demospongiae • Poecilosc1erida • Mycalina • Isodictyidae 705

A C

-
G

H
I
Fig. 2. A-H, Isodictya palmata (Ellis & Solander, 1786 as Spongia). A, Ellis & Solander's (1786) specimen (from Ellis & Solander, 1786, pI. 58 fig. 6).
B-C, BMNH 1930.7.3.381 (Orkney Isles, from Bowerbank, 1874, pI. LII fig. 2). B, drawing of choanosomal architecture (scale 200 !Lm). C, drawing of
palmate isochelae on side (upper three) and face views (scale 10 !Lm). D-H, ZMA 3280, off Iceland (det. R.W.M. Van Soest). D, tangential view of the surface
of the specimen showing the projecting terminations of the choanosomal fibres (scale 500 !Lm). E, transverse section through a branch showing the plumo-
reticulated organization of the skeleton with large subectosomal spaces (scale 5 mm). F, detail of the isodictyal reticulation showing primary and secondary
connecting tracts of megascleres cemented by spongin (scale 500 !Lm). G, fusiform oxea (scale 50 !Lm). H, palmate isochela on side view (scale 10 !Lm).

Type species Description of type species

Spongia palmata Ellis & Solander, 1786 (by subsequent Spongia palmata Ellis & Solander, 1786 (Fig. 2).
designation; Dendy, 1924). Synonymy. Spongia palmata Ellis & Solander, 1786: 189.
Spongia digitata Esper, 1797: 190. Halichondria palmata
Diagnosis Johnston, 1842: 92. Isodictya palmata Bowerbank, 1866: 31l.
Pachychalina compressa Schmidt, 1870: 37.
Isodictyidae of flabellate/digitate growth forms; choanosomal Material examined. 'Holotype' of Halichondria palmata
skeleton reticulate or plumoreticulate (niphatid-like); megascleres Johnston, 1842: BMNH 1847.9.7.1 - Berwickshire coast. Other
are mostly diactinal, usually oxeas; microscleres, palmate material. BMNH 191O.l.l.2197 - NE Atlantic, Norman Collection.
isochelae, frequently with plate-like inner extensions of the falxes. ZMA 3280 - off Iceland, 64°48'N, 12°45'W, det. R.W,M. Van Soest.
706 Porifera· Demospongiae • PoeciloscIerida • Mycalina • Isodictyidae

Description (adapted from Lundbeck, 1905: 121). Erect, South Africa by Levi (1963), but none exactly match the descrip-
digitate with variably compressed branches which may coalesce tion of Carter. Nevertheless, the synonymy of Textiliforma with
into plate-shaped areas (Fig. 2A). Surface is finely and densely Isodictya is clear. Valentis de Laubenfels, 1936 was erected for
shaggy from the projecting ends of the skeletal fibres (Fig. 2D), type species Desmacidon lentus Vosmaer, 1880: 131 from the
connected by thin organic ectosome draped over the fibre ends. 'coast of France' (by original designation). The dry holotype
Oscula up to 5 mm in diameter (in spirit), mostly along the edges of (RMNH 261), was re-examined. The label reads 'Desmacidon
the compressed branches, but also scattered on the plates formed lentus Vosm. type, coli. Persoon, ? Kust van Frankrijk', which indi-
by the fusion of some branches (Fig. 2A). Reticulated skeleton cates that it is not certainly from the coast of France. In view of its
composed of primary ascending tracts connected by thinner characters, it appears unlikely to be from Western Europe. It is a
secondary tracts (Fig. 2B, E-F). Skeletal meshes larger, the closer thinly flabelliform sponge with a distinct oscular and poral surface,
to the surface, in a pattern resembling that of niphatids (Fig. 2E). size 21 X 11 X 0.8 cm. The skeleton is a reticulation of spongin-
Megascleres are cigar-shaped oxeas (116-277,...,m long and cemented thick spicule tracts forming elongate meshes, primary
4.5-16.8,...,m thick; 199-(206.5)-216,...,m long and 1O-11,...,m thick tracts up to 250,...,m in diameter, interconnecting tracts 50-150 ,...,m,
in the holotype, Fig. 2G), occasionally styles or strongyles. meshes 200-800 X 200-450 ,...,m. No special ectosomal skeleton,
Microscleres are palmate isochelae (23-35,...,m in length, 23.8,...,m the tracts ending at the surface in spreading brushes of megascle-
in the holotype, Fig. 2C, H). res. Spicules consist of thick, strongly curved oxeas, 250-300 X
Remarks. Comparative material examined: Isodictya deich- 24-30 ,...,m, and palmate isochelae 20-25 ,...,m. Vosmaer (1880) men-
mannae (de Laubenfels, 1949a) - ZMA 6186, east Cape Cod tioned several other spicule types, but these were not observed;
(NW Atlantic, det. R.WM. Van Soest). Samaai et al. (1999) con- presumably they were foreign. This is undoubtedly a species of the
ducted a formal phylogenetic analysis in order to test the contro- genus Isodictya, but it differs from the European Isodictya palmata
versial allocation of Isodictya (and its synonym Cercidochela) to in several distinct traits: thinly flabelliform, smaller oscules,
the haplosclerids proposed by Hajdu et al. (1994b), as well as that thicker spicules. Since the origin of the specimen is unknown, it is
of Coelocarteria, as discussed above. Their results were taken here here provisionally named Isodictya lenta (Vosmaer, 1880). On the
as the state of the art of our knowledge regarding the affinities of basis ofVosmaer's partly faulty description, de Laubenfels (1936)
Isodictya, but it is of special concern that neither haplosclerids with made a genus definition which does not cover the properties of
strict isodictyal skeletons (= Isodictya palmata-like), nor poe- Desmacidon lentus. The genus Valentis is here assigned to the syn-
cilosclerids devoid of chelae were considered in their analysis. onymy of Isodictya based on the characters of the type specimen.
Additionally, too much synapomorphic value was attached to
details of skeletal anatomy (7 out of 12 characters), which may
either be interpreted differently by other authors working on the ACKNOWLEDGEMENTS
same species (there are no strict definitions of skeletal architecture
terminology - terms are mainly broad descriptors), or vary widely The authors are thankful to Drs John N.A. Hooper (QM) and
when additional species are added to the analysis. The only chemi- Rob WM. Van Soest (ZMA), for the invitation to take part on the
cal character considered by Samaai et al. (1999), viz., manzamines, Systema Porifera project and for valuable discussions during the
coded synapomorphic for the haplosclerids, is known to occur in preparation of this manuscript. Dr. Wallie H. de Weerdt (ZMA) is
poecilosclerids too (Harper, in litteris; in Van Soest & Braekman, especially thanked for valuable discussions during the assessment
1999). of the classification whereabouts of Isodictya and related genera.
Two other genera are also included here in synonymy with We are most grateful to Rob Van Soest and Clare Valentine
Isodictya, based on recent re-examination of their respective type (BMNH) for providing access to type specimens and other mate-
material (courtesy of Rob Van Soest). Textiliforma Carter, 1885d rial. Rob Van Soest is also gratefully acknowledged for providing
was erected for type species Textiliforma joliata Carter, 1885d: 288 access to SEM facilities during my PhD candidature in
from Cape of Good Hope, South Africa (by monotypy). The type Amsterdam, where most of the morphological studies on Isodictya
series (not examined) contains thinly flabelliform sponges, with and allied genera were conducted, and for re-examination and
groups of oscules scattered over the surface. The skeleton is a re-description of the type species of Textiliforma and Valentis.
ladder-like reticulation of spicule tracts embedded in spongin. Financial support to attend the Systema Porifera Workshops and
Spicules are oxeas of 120 X lO,...,m and isochelae of 25 ,...,m. From to construct the means for writing this revision was obtained
Carter's characterization of the latter ('navicular, with obtuse from CAPES, CNPq, FAPERJ, and FAPESP, from Brazil, and the
ends') these are presumed to be the palmate type peculiar organisers of the International Conference on Sponge Science
to Isodictya. Several Isodictya species have been described from (Otsu, Japan).
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Suborder Latrunculina subord. nov. incertae sedis

Michelle Kelly! & Toufiek Samaai2

1 National Institute of Water & Atmospheric Research (NIWA) Ltd, Private Bag 109-695, Newmarket, Auckland New Zealand.
(m.kelly@niwa.cri.nz) 2Zoology Department, University of the Western Cape, Private Bag X17, Bellville, 7535, Cape Town South
Africa. (tsamaai@uwc.ac.za)

Latrunculiidae Topsent (Demospongiae, Poecilosclerida) is assigned as a monophyletic taxon to suborder Latrunculina subord. nov.,
incertae sedis within Poecilosclerida. Latrunculiidae lacks any of the major apomorphies of this order but has certain other characters
indicative of closer affinities with Poecilosclerida than with Hadromerida, to which it has previously been assigned. Latrunculiidae are
unique amongst Poecilosclerida in having discate canthose microrhabd microscleres (acanthodiscorhabds, or 'chessman' spicules)
arranged vertically in a palisade on the exterior surface forming an ectosomal crust. The inferred homology between acanthodiscorhabds
and spirasters of Hadromerida is rejected, as is their homology with spinose microrhabds (spinorhabds) found in Podospongiidae.
Keywords: Porifera; Demospongiae; Poecilosclerida; Latrunculina subord. nov. incertae sedis; Latrunculiidae.

DEFINITION, DIAGNOSIS, SCOPE areolate porefields, fistular oscuies, and biochemical similarities
(sterols and free amino acids; Bergquist et ai., 1969, 1980). (2)
Definition Arguing for its allocation to Poecilosclerida Levi (1973), Vacelet
(1969), Boury-Esnault & Lopes (1985), Hooper (1986), and Kelly-
Poecilosclerida incertae sedis with acanthodiscorhabds form- Borges & Vacelet (1995) also cite evidence associated with megas-
ing an ectosomal skeleton; without chelae or sigma microscleres. clere geometry, skeletal arrangement, ontogenetic development
of microscleres and their localization to the ectosome, etc.
Diagnosis (see chapter on Latrunculiidae). (3) Van Soest (1984b) considered
all these arguments were weak and included the family incertae
As for family. sedis within Porifera. (4) Many aspects of this debate are now
redundant given the revised concept of 'Latrunculiidae' into two
Scope families, Latrunculiidae and Podospongiidae. In both families
the ontogenetic development of acanthodiscorhabds and spin-
A single family, Latrunculiidae, and four valid genera. orhabds, respectively, differ from those of spirasters of
Hadromerida (Spirastrellidae), supporting their allocation within
Remarks Poecilosclerida. The two families differ in the ontogenetic develop-
ment of their microscleres, with the protorhabd of acanthodis-
Latrunculiidae was initially allocated to Poecilosclerida corhabds in Latrunculiidae being straight, and those of
(Topsent, 1922, 1928c), but has since alternated between this order Podospongiidae being sigmoid, with the consequence that the latter
and the tetractinomorph Hadromerida. (1) In support of the latter is postulated to have affinities with sigmancistra derivatives of
assignment Bergquist (1968, 1978), Hartman (1982) and Mycalina. The former has no obvious affinities with other
Wiedenmayer (1989) cited evidence based on megasclere geometry, Poecilosclerida families or suborders, and is incertae sedis within
skeletal arrangement, ectosomal microsclere crust, special this order.

707
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcadernicIPlenurn Publishers, New York, 2002

Family Latrunculiidae Topsent, 1922

Toufiek Samaajl & Michelle Kelly2

1 ZoologyDepartment, University of the Western Cape, Private Bag X17, Bellville, Cape Town, South Africa. (tsamaai@uwc.ac.za)
2 NationalInstitute of Water & Atmospheric Research (NIWA), Private Bag 109-695, Newmarket, Auckland, New Zealand.
(m.kelly@niwa.cri.nz)

Latrunculiidae Topsent (Demospongiae, Poecilosclerida incertae sedis) is revised substantially from prior concepts, recognizing two fam-
ily taxa (Latrunculiidae and Podospongiidae) differentiated primarily by the morphology of microrhabd microscleres that reflect major
ontogenetic differences in their formation. Latrunculiidae has discate acanthose microrhabd microscleres (acanthodiscorhabds or 'chess-
man' spicules), that form a dense perpendicular palisade in the outer ectosome, areolate porefields, and short fistular oscules on the
sponge surface. Four genera are recognised as valid (Latrunculia, Sceptrella, Strongylodes1rW and Tsitsikam1rW gen. nov.), with suspected
affinities to Poecilosclerida but no obvious close relatives within any of the suborders, and hence is allocated to Latrunculina subord. nov.
incertae sedis in Poecilosclerida pending a more thorough phylogenetic analysis.
Keywords: Porifera; Demospongiae; Poecilosclerida; incertae sedis; Latrunculina; subord. nov.; Latrunculiidae; Latrunculia; Sceptrella;
Strongylodesma; Tsitsikamma gen. nov.
- - - - - - - - - _... _ .._ . - _ ..__. _ - - _ ._ _ .

DEFINITION, DIAGNOSIS, SCOPE regions for Latrunculia and Tsitsikamma, southern and northern
warm temperate to tropical regions for Strongylodesma, and north-
Synonymy ern hemisphere temperate regions for Sceptrella.

Latrunculiidae Topsent, 1922. Scope

Definition There are 12 nominal genera, four of which are considered to

be valid Latrunculiidae, and include Latrunculia, Sceptrella, and
Wispy wide-meshed reticulate to reinforced reticulate Strongylodes1rW, the latter transferred from the poecilosclerid
choanosomal skeleton of anisostyles or strongyles, ectosomal family Chondropsidae, and Tsitsikam1rW gen. nov. erected here
skeleton compact tangential, microscleres are acanthodiscorhabds, (See Fig. 1). Negombo (with synonym Oxylatrunculia) is trans-
or 'chessman' spicules, typically arranged vertically in a palisade ferred to Desmoxyidae (Halichondrida), and Sigmosceptrella,
on outside of ectosome. Diacarnus, Negombata, Podospongia (with synonym Alcyospongia),
are transferred to Podospongiidae (Poecilosclerida, with suspected
Diagnosis affinities to Mycalina - see Kelly & Samaai, this volume).
Phlyctaenopora Topsent and Barbozia Dendy, presently assigned
Massive encrusting, hemispherical, spherical or pedunculate to the poecilosclerid family Crellidae, are transferred incertae sedis
sponges with areolate porefields and raised fistular oscules; surface to Mycalidae.
smooth and velvety between fistules; texture leathery and barely
compressible when in preservative, whole animal generally very History and biology
dense with heavy uniform body mass. Colour in life typically
brownish black with forest-green or blue tinges, liquorice brown, Latrunculiidae was erected by Topsent (1922) to include gen-
fawn, or forest green (Latrunculia, Strongylodes1rW, Tsitsikam1rW), era with either spinose or discate microrhabds and anisostyle
or pale beige to white (Sceptrella). Structural megascleres are megascleres. Topsent initially included three genera in the family,
anisostyles or strongyles, rarely oxeas, these are frequently slightly Sigmosceptrella Dendy, 1922b, Podospongia du Bocage, 1869 and
irregular, sinuous, forming a compact tangential layer under the Latrunculia du Bocage, 1869. As for many other families within
ectosome, and a wide-meshed reticulation in the choanosome. the Demospongiae Latrunculiidae was, and still is, the subject of
Microscleres are acanthodiscorhabds bearing apical and basal considerable controversy. Until the review of Kelly-Borges &
whorls of spines (apical whorl and manubrium) and several crenu- Vacelet (1995) it was considered to be monophyletic (Dendy,
late discs (subsidiary and median whorls) in between, typically I 922b; Topsent, 1922, 1928c; Bergquist, 1968, 1978; Vacelet,
arranged in a compact palisade of single perpendicular spicules, 1969, Levi, 1973; Wiedenmayer, 1977b; Boury-Esnault & van
their bases buried in the ectosomal membrane. Based on the obser- Beveren, 1984; Wiedenmayer, 1989; Hooper & Wiedenmayer,
vation of larvae in Latrunculia citharistae by Vacelet (1969), we 1994; Hooper, 1986, 1997), and included Didiscus Dendy, 1922b
conclude that the Latrunculiidae are viviparous. in addition to the latter three genera. Negombata de Laubenfels,
Latrunculia, Strongylodes1rW, and Tsitsikamma are known to 1936a, Diacarnus Burton, 1934a, Oxylatrunculia Hoshino, 1981,
contain biologically active discorhabdins and their derivatives. Sanidastra Volkmer-Ribeiro & Watanabe, 1983, and Alcyospongia
Bathymetric distribution is shallow sublittoral to bathybenthic. de Laubenfels, 1934, were subsequently added to the family,
Biogeographic distribution is polar to warm temperate with centres although the affinities between these genera were not always com-
of diversity and abundance in southern hemisphere temperate pletely clear. Sanidastra is now clearly accepted as belonging to

708
Porifera· Demospongiae • Poecilosclerida • Latrunculina • Latrunculiidae 709

is presented here, although it remains incomplete given there are

many new data awaiting publication (Samaai, unpublished data).
Latrunculiidae was first established within the Order
Poecilosclerida (Topsent, 1922, 1928c) but Bergquist (1968, 1978),
Hartman (1982), and Wiedenmayer (1989) subsequently consid-
ered the family to have greater affinities with Hadromerida. This
action was based on: the collagenous nature of the mesohyl (in
some genera at least); the similarity of the areolate porefields to
those in Cliona; similarity of the radial skeleton between Tethya,
Stylocordyla, and Podospongia; presence of fistular oscules in
Polymastia; presence of a microsclere crust in Spirastrellidae; and
the similarity of sterol (Bergquist et al., 1980) and free amino acid
patterns (Bergquist et al., 1969) with the hadromerids. Van Soest
(1984b), on the other hand, stated that arguments are weak for
placement in either Order, and included the family incertae sedis
within the Porifera. As it is currently perceived, the family is het-
erogeneous, sharing features with both hadromerid sponges
Fig. 1. Discorhabd microsclere morphologies used to define genera (Subclass Tetractinomorpha) and poecilosclerid sponges (Subclass
of Latrunculiidae, and discorhabd-like microscleres found in other Ceractinomorpha). Nevertheless, although there is indeed a sug-
groups of sponges as a comparison. A-B, Latrunculia, anisodiscorhabd
and aciculodiscorhabd. C, Sceptrella, isoconicodiscorhabd ('sceptre'). gested common ancestry to hadromerids the vast majority of char-
D, Tsitsikamma, isochiadiscorhabd. E-F, Sigmosceptrella, spinorhabds. acters are seen to be shared with poeciloscerids (Table I). These
G-H, Negombata, spinorhabds. I-J, Barbozia, oxydiscorhabds. K-L, include the type of megascleres, their arrangement within the
Diacamus, spinorhabds. M, Didiscus, didiscorhabds. N, Negombo, sponge, the ontogenetic development of the microscleres, and their
sanidaster. O-P, Oxylatrunculia, sanidasters. Q-R, Podospongia, spin-
surface position (Carter, 1879b; Ridley & Dendy, 1887; Dendy,
orhabds (podorhabds). S, Sanidastra, sanidasters.
1921; Topsent, 1922, 1928c; Bergquist, 1968, 1978; Levi, 1973;
Vacelet, 1969; Boury-Esnault, 1985; Hooper, 1986; Wiedenmayer,
1989; Kelly-Borges & Vace1et, 1995), all of which support the
the freshwater Spongillidae by Manconi & Pronzato (1996). Until inclusion of Latrunculiidae within Poecilosclerida, but its relation-
the work of Kelly-Borges & Vacelet (1995) no distinction was ships within the order remain incertae sedis.
made between discate and spined forms (i.e., with the prior
assumption that these microrhabds were homologous), leading to
considerable instability in the family. REVISION OF LATRUNCULIIDAE
Recent work on several genera within Latrunculiidae (Kelly-
Borges & Vacelet, 1995; Van Soest, 1987; Van Soest et at., 1990) The major character that defines the genera Latrunculia
indicated that it might be polyphyletic, and these authors began the and Sceptrella (Latrunculiidae) and Podospongia, Negombata,
task of ascertaining the phylogenetic affinities of the various taxa. Diacamus and Sigmosceptrella (Podospongiidae) is possession of
However, none of these studies could be construed as comprehensive acanthose microrhabds, traditionally used to unite them historically
revisions of the family, and thus such a revision was necessary and into a single family. Characters such as the form and disposition of

Table 1. Characters shared between Poecilosclerida and Hadromerida for the prior concept of family Latrunculiidae
(Latrunculiidae + Podospongiidae).

Character Poecilosclerida Hadromerida

- - - - - - - - - -_ .. _ - - - _.._ - - - - - - - _ . -----------

Microsclere disposition ectosomal palisade, erect surface spicules throughout ectosome + crust (armoured surface)
Microsclere morphology Sigmoid protorhabd
(Podospongiidae) Terminally spined strongyles
Megasclere morphology anisostyles, styles, polytylotes, polytylote features strongyloxeas
terminally spined styles erect palisade
Ectosomal architecture tangential, plumoreticulate
(dendritic), axial fibro-reticulation,
Choanosomal architecture reticulate, fibroreticulate radial (Podospongia)
Gross morphology massive encrusting, digitate branching massive encrusting
Surface features areolate pore fields & fistulose oscules areolate porefields & fistulose oscules
Colour deep brown or green, bright colouration red (Spirastrellidae)
(Zyzzya), bright red (Negombata)
Histology highly collagenous (Diacarnus)
Bioactive chemistry pyrroloquinoline alkaloids + norsesterterpene sterol and free amino acid patterns
peroxides + latrunculins
710 Porifera· Demospongiae • PoeciloscIerida • Latrunculina • Latrunculiidae

Apex

~
~t
•
Apical whorl (a.w.)

• C?
Subsidiary whorl (s.w.)
•
t..,c::
Median whorl (m.w.)

~(
.....l

Shaft Aciculodiscorhabd

iFf
Manubriwn (rna.)
•
Anisodiscorhabd
Fig. 2. Schemalic representalion of picu le lerminology u cd 10 de cribe the di corhabd of LalTllllculia.

these microscleres, the skeletal arrangement of the megascleres Tsitsikamma gen. nov. These microscleres differ consider-
and the sponges' general morphology, have been used to distin- ably from those in Latrunculia and Sceptrella as they are
guish genera from one another. However, Kelly-Borges & Vacelet isochiadiscorhabds with a stout straight shaft, bearing three whorls
(1995) demonstrated that acanthose microrhabds were non- of truncate tubercles (prefix chia- means trucate - see Bergquist &
homologous between certain groups of genera, correlated with Kelly-Borges, 1991, for microsclere terminology), which are ter-
clear differences in the arrangement of their megasclere skeletons minally acanthose (Figs 1, 6E-F). In these microscleres the sub-
within each taxon, leading them to the hypothesis that the prior sidiary whorl is absent, the medium whorl is midway between
concept of 'Latrunculiidae' was polyphyletic. manubrium and apical whorl.
Ontogenetic development of acanthomicrorhabds.
Microsclere ontogenetic development, morphology and Although the adult form of the microscleres in all of the above gen-
disposition era are superficially similar, it is their differences in developmental
stages that gives a clear indication of their separate origins. Dendy
Latrunculia. Species have anisodiscorhabds as microscle- (1921) referred to these developmental stages as reflecting their
res (Figs 1-2). These have a straight shaft bearing four disc-like ontogenetic development, but it is equally tenable that these also
whorls of spines that are differentiated in size and shape. The apex reflect a transformational homology of character states. The
of the spicule usually consists of a rosette or crown of spines and anisodiscorhabds of Latrunculia follow a particular ontogenetic
the base of the spicule, which is embedded in the ectosomal mem- developmental pattern that starts with a straight protorhabd, as
brane, is called the manubrium (ma.). The shape of this part of the indicated by Dendy (1921, pI. 15 fig. lOa) (see Fig. 4G). In con-
microsclere is typically bushy and spinose. Immediately beneath trast, this developmental process, as suggested by Topsent (1922),
the apex of spines, is the apical whorl (a.w.), which has a uniform clearly differs from the early ontogenetic stage of Podospongiidae
(non-partitioned) margin. Between the apical whorl and the (Sigmosceptrella, Negombata , and Podospongia) in that the pro-
manubrium are subsidiary (s.w.) and median whorls (m.w.) torhabd is sigmoid in the later species (see also Dendy. 1921;
arranged more-or-less at equal distances along the shaft, although Wiedenmayer, 1989; Kelly-Borges & Vacelet, 1995). Although the
in some species these discs are closer to the apical end of the protospinorhabds of Diacamus are faintly reminiscent of the pro-
spicule. These central whorls are usually disc-shaped with crenu- toanisodiscorhabds of Latrunculia and Sceptrella (Fig. 5C), in that
late margins, parallel with each other, and partitioned into several the protorhabd shaft is straight, the development of spines on the
leaf-like flanges . Several species of Latrunculia also contain acicu- shaft in Diacamus protospinorhabds is in complete contrast
lodiscorhabds (Figs 1-2) which have an elongated apical spine. to the development of the discs in anisodiscorhabds. The pro-
Sceptrella. The microscleres are isoconicodiscorhabds or toisochiarhabd in Tsitsikamma gen. nov. (see Fig. 6G) is straight,
'sceptres' (Figs 1, 5A-C) with a stout straight shaft bearing an api- and spines appear simultaneously on the distal curved ends of the
cal whorl and manubrium that is not differentiated in size or form shaft as in Latrunculia and Sceptrella. Although there are substan-
(prefix conico- means conical - see Boury-Esnault & Riitzler, tial differences in the ontogenetic development of microscleres
1997, for terminology). The apical whorl differs from those in between Latrunculia, Sceptrella and Tsitsikamma, the develop-
Latrunculia S.s. in that it is fused with the apex and consists of a ment of discs completely contrasts with the development of spines
rosette of crenulated spines. They also bear furcate or conical in spinorhabds of the Podospongiidae. The lack of microscleres in
spines (subsidiary and median whorls) which are often acanthose Strongylodesma is viewed here as a secondary loss during the evo-
and arranged at equal distances along the shaft. lution of the family; in all other characters there are remarkable
Porifera· Demospongiae • Poecilosclerida • Latrunculina • LatruncuIiidae 711

similarities between Strongyiodesma, Latrunculia, Tsitsikamma known reproduction in Podospongiidae is viviparous (nan, 1995;
and Sceptrella. Kelly-Borges & Vacelet, 1995).
Disposition within the skeleton. In Latrunculia the anisodis-
corhabds are typically arranged in a single layer of perpendicular Base pigmentation
spicules forming a tightly packed palisade on the outside of the
sponge, with the bases of the spicules buried in the ectosomal mem- Latrunculia, Strongyiodesma, and Tsitsikamma have a charac-
brane. In Sceptrella and Tsitsikamma the arrangement is not as teristic base pigmentation of deep brownish black that is often and
regular but the isodiscorhabds are also arranged in a single layer variously tinged with forest green and deep blue. These specimens
on the outside of the sponge. In contrast, the disposition exude a dark brownish or greenish black pigment; the ethanol pre-
of the microscleres in the Podospongiidae genera Diacarnus, servative is always oily-looking and deeply pigmented. Preserved
Sigmosceptrella, Podospongia, and Negombata is quite different specimens always retain their dark pigmentation. Some species are
with the spinorhabds forming a thick crust within the ectosome and a lighter oak brown throughout. As we have not seen Sceptrella
frequently throughout the distinct cortical region. These microscle- in life their living colouration is uncertain, however, ethanol-
res also show no tendency towards any vertical orientation. preserved specimens are uniformly cream to tan. It is interesting to
note that specimens of Tsitsikamma without discorhabdins are a
Megascleres and skeletal architecture lighter brown than their discorhabdin-containing siblings. Variation
in pigmentation has been associated with increased levels of cyto-
Latrunculia, Tsitsikamma and Sceptrella have large often poly- toxicity in the genus Latrunculia, i.e., the most cytotoxic extracts
tylote anisostyles, and Strongyiodesma has strongyles as megascle- of discorhabdins have been found from heavily pigmented green
res. Another so-far undescribed species of Latrunculia from the and brown sponges of Latrunculia (Perry et ai., 1986; Perry et ai.,
Sea of Okhotsk possesses terminally spined anisostyles (Samaai, 1988b; Copp et ai., 1994; Lill et ai., 1995).
unpublished). By comparison, Podospongiidae all have strongylox-
eas, to which robust curved styles are added in Negombata, with the Biochemistry
latter restricted to the choanosomal 'axial' fibroreticulation.
The general skeletal architecture that typifies Latrunculiidae Latrunculia spp. (Perry et al., 1986; Perry et ai., 1988b; Kita
is a large-meshed polygonal reticulation formed by wispy tracts of et ai., 1989; Copp et ai., 1994; LiB et al., 1995; Yang & Baker,
spicules with no spongin reinforcement. The ectosomal skeleton is 1995; Beukes, 2000; Beukes et ai., in press), and Strongyiodesma
a tangential layer of the same megascleres. This skeletal organisa- spp. (Beukes, 2000; Beukes et al., in press) contain biologically
tion is reminiscent of the genus Phorbas (Hymedesmiidae). active discorhabdins and their derivatives. A new pyrroloimino-
Although the reticulation in Sceptrella is irregular and wispy, the quinone alkaloid tsitsikammamine, isolated from Tsitsikamma
structure at the base is slightly radial. The major differentiating together with brominated discorhabdin derivatives, is structurally
characteristic of Tsitsikamma is the massive reinforcement of these related to the makaluvarnine compounds found in the poeciloscle-
meshes to form huge thick tracts that are visible with the unaided rid genus Zyzzya (Hooper et ai., 1996; Beukes, 2000; Urban et al.,
eye. The overall gross structure is honeycombed with convoluted 2000). Interestingly, Tsitsikamma gen. nov. also appears superfi-
layers, similar to poecilosclerid genera such as Inflatella and cially to be morphologically intermediate between Latrunculia and
Coelocarteria. In contrast, Podospongiidae are characterised by Zyzzya, at least in terms of their general choanosomal skeletal
the possession of a fibroreticulate plumose skeleton. arrangement (Table 2). The apparent relatedness of these com-
pounds (discorhabdins, tsitsikammamines and makaluvamines)
Gross morphology and isolation of derivatives (i.e., discorhabdins in Zyzzya spp.,
makaluvamines in Latrunculia sp. and 'hybrid' makaluvarninesl
Latrunculiidae present a range of growth forms extending discorhabdins in Tsitsikamma) is a promising target for future
from relatively thin encrustations to thick hemispherical masses. investigation. Determination of whether pyrroloquinoline alkaloids
All taxa have areolate porefields and raised trumpet-like oscular are a potential chemotaxonomic marker for these genera would
fistules, and are velvety to the touch. The texture of Latrunculia, also require investigation of the homology of their respective
Strongyiodesma and Sceptrella is very cakey, dense and compress- biosynthetic pathways.
ible, and not very elastic. Tsitsikamma is extraordinarily tough A further group of compounds, the prianosins, were described
and leathery due to the sponge's huge internal spicule tracts. from a Japanese sponge identified as Prianos meianos de
Latrunculiidae are thus superficially similar to the genera Zyzzya Laubenfels, 1954, (Kobayashi et ai., 1987; Cheng et ai., 1988;
(Acarnidae) and Phorbas (Hymedesmiidae). By comparison, Kobayashi et ai., 1991; Van Soest et al., 1996a). These are in fact
growth forms of Podospongiidae are typically digitate and branch- discorhabdins; prianosin A is identical to discorhabdin A and pri-
ing, flexible, compressible, very elastic due to rich collagen anosin D to discorhabdin D (Perry et al., 1988a,b; Lill et al., 1995).
mesohyl, and lacking any special surface features. Prianosin Band prianosin C are new pyrroloquinoline alkaloids
(Van Soest et al., 1996a), similar to discorhabdins. It is likely,
Reproduction however, that this record of P. meianos is a misidentification for a
Strongyiodesma, since both taxa have depauperate morphological
Oocytes have been observed in southern hemisphere speci- characteristics, including only strongyles as megascleres, lacking
mens of Latrunculia, Strongylodesma, and Tsitsikamma (Kelly microscleres and dark brownish pigmentation (Urban et ai., 2000).
unpubl. data), but Vacelet (1969) provides the only report of the Van Soest et ai. (1996a) reported on the discovery of two
presence of parenchymella larvae in Latrunculia citharistae groups of biogenetic ally unrelated alkaloids, cyclic guanidine alka-
Vacelet, indicating that the group are probably viviparous. No loids and pyrroloquinoline alkaloids called batzellins and iso-
reproductive products have been observed in Sceptrella. Where batzellins, in sponges that had been assigned to Batzella (i.e., based
712 Porifera· Demospongiae • PoeciloscIerida • Latrunculina • Latrunculiidae

Tab le 2. Po ible affinitie between Lammclliia and Tsilsikamma. and Zy;:;:ya

Tsil ikalllllW Zyzzya

Character LalrlmclIiia
gen. nov.

massive encrusti ng, emi pherical. thicklyencru ting

External morphology
pedu ncu late
brownish or greeni h b al ck. pink browni h black
Base pigmentalion browni h or grceni h b ack
l
areolatc p refield + Ii tular 0 cu le arcolate porefield + Ii tular 0 cule
Surface feature areolate porelield + listular 0 cu les

Skeletal architecture

tangential mega clerc + micro clere tangential mega c leres + tangen tial megasclere
Ectosomal architccture
pali ade microsclere pali ade

on their possession of a simple complement of strongyles in a wispy subsequently been found on southeast coast of South Africa. The
reticulation). The sponge with cyclic guanidine alkaloids, originally genus is also known from New Caledonia (Vacelet & Kelly in
identified as Batzella (ZMA Por 8788), is now considered to be Levi, 1998) and Tonga (Kelly, unpublished). The seamount and the
Monanchora arbuscula (Crambeidae), and the genus Batzella, like shallow subtidal South African species grow on hard rocky sub-
Prianos, is a dustbin taxon. Batzellin and iso-batzellin-containing strates, whereas the subtropical species grow attached to dead coral
sponges from Harbar Branch Oceanographic Institution, also identi- rubble in sandy lagoon environments (Vacelet & Kelly in
fied as Batzella (HBOM 003:00050, HBOM 003:00051) are identi- Levi, 1998). Species previously identified as Batzella from deep-
cal in all respects to Strongylodesma, which also produces water in the west central Atlantic region (Van Soest, 1996;
pyrroloquinoline alkaloids. These highly functionalised pyrrolo- Gunasekera et aI., 1999) are now recognised as Strongylodesma.
quinoline alkaloids are very closely related to the discorhabdins of Tsitsikamma is only known from the unique, warm temperate
Latrunculia species and the makaluvarnines of Zyzzya. These alka- region off the southeast coast of South Africa, where they are found
loids thus become a strong consistent chemical marker for Zyzzya, in relative abundance in sheltered coastal rocky reefs down to 25 m
Latrunculia and Strongylodesma (Urban et al., 2000). depth.

Ecology, bathymetric and biogeographical distribution Phylogenetic affinities

Latrunculiidae are widespread, found predominantly in cold Latrunculia, Strongylodesma, Tsitsikamma and Sceptrella are
water with centres of diversity in the southern and northern hemi- unequivocally differentiated at a higher taxonomic level from
spheres (Fig. 3). Species of Latrunculia are predominantly sublit- Podospongiidae based on fundamental differences in having differ-
toral and often a dominant organism in the benthic fauna. In all ent development stages of their microrhabd microscleres.
except Antarctica, the subantarctic and subtropical environments The presumed homology of spicules and skeletal characters of
these sponges are found in relative abundance in exposed coastal Latrunculiidae (and Podospongiidae) have been diversely inter-
environments attached to hard rocky substrate, often on vertical preted, resulting in the switching back and forth of the groups
walls or rocky reef surfaces down to 30 m depth. In Antarctica between the tetractinomorph Order Hadromerida and the ceractin-
Latrunculia apicalis is found under the sea-ice and grows on a sed- omorph Order Poecilosclerida. However, whereas Podospongiidae
imentary substrate of predominantly sponge spicules. Although the has more obvious affinities with the Poecilosclerida Mycalidae
predominant centre of diversity for these genera appears to be in (see Kelly & Samaai, this volume), those of Latrunculiidae remain
the southern hemisphere, the genus has an anti boreal distribution more obscure, and hence remains incertae sedis within the
(Plate, Antarctica, subantarctic, South Africa, New Zealand, south- Poecilosclerida subordinal classification.
west Australia, Tasmania), extending into subtropical water (New
Caledonia, Tonga, Southern Japan) and the western Boreal Pacific.
There are only a few known species of Sceptrella and these appear Levels of speciation
to be uncommon in the environments in which they have been
found. Species of Sceptrella are predominantly deepwater with Latrunculiidae, as revised here, is a small but well-defined
a centre of diversity in the northern hemisphere, in the Atlantic family with four easily differentiated genera. Latrunculia, found
Ocean. Sceptrella was first described off the coast of Florida but predominantly in southern ocean waters, is characterised by abun-
species have also been found off the Azores, the coast of Gibraltar, dant and what appears to have been rapid speciation. In New
and the north coast of Norway down to a depth of 2460 m. Zealand, as in the Polymastiidae (Kelly-Borges & Bergquist,
Strongylodesma was first described from an oceanic seamount 1997), species of Latrunculia are numerous and relatively difficult
in the southern Atlantic Ocean, and a shallow subtidal species has to differentiate and distinction is based upon genetic and small
Porifera· Demospongiae • Poecilosclerida • Latrunculina • Latrunculiidae 713

X •

•
] .'
• Latrllnclllia
X Sceplrella
• •
*
• Irongylodesma
Tsilsikall//lla •
Fig. 3. Known di tribution of Latrunculiidac. e LaIrlIllCIIlill; . Seep/rel/a: . /rollgylodesl/la: * Tsilsik(ll/lll/(1 gen. no\.
morphological and morphometric differences in spiculation Previous reviews
(Alvarez et al., 2002). In South Africa however, species are
discrete and easily differentiated. Sceptrella and Strongylodesma Carter (I 879b); Ridley & Dendy (1887); Dendy (1921);
are characterised by having only a few discrete species that are Topsent (1922, 1928c); de Laubenfels (1934); Bergquist (1967,
geographically isolated. Tsitsikamma is known from two South 1968, 1978); Hooper (1986); Wiedenmayer (1989); Kelly-Borges &
African species. Vacelet (1995); Urban et al. (2000); Alvarez et al. (2002).

KEY TO GENERA

(1) Megascleres anisostyles or styles, microscleres present ...................................................................................................................... 2

Megascleres strongyles, microscleres absent ............................................................................................................. Strongylodesma
(2) Microscleres anisodiscorhabds with spined apical whorl and manubrium, and
disc-shaped median and subsidiary whorls .................................................................. .........................................................lAtrunculia
Microscleres are isoconicodiscorhabds with crenulated apical whorl, and conical or furcate whorls of spines .................... Sceptrella
Microscleres are isochiadiscorhabds with cylindrical terminally acanthose truncate tubercles ....................................... Tsitsikamma

LATRUNCUUA DU BOCAGE, 1869 Diagnosis

Synonymy Encrusting to massive oval-shaped sponge with raised trumpet-

like or mammiform oscular fistules and areolate porefields; surface
Latruneulia du Bocage, 1869. velvety to the touch; texture is very cake-like, dense but compress-
ible. Colour in life deep brownish-black or dark green, sometimes
Type species tinged with deep blue, in preservative specimens always retain
their dark pigmentation. The choanosomal architecture consists of
Latrunculia eratera du Bocage, 1869 (by monotypy). monactinal or rarely diactinal spicules arranged in an irregular,
polygonal, large-meshed reticulation formed by wispy tracts of
Definition spicules, which lack spongin reinforcement. There is no distinction
between primary and secondary tracts. The ectosomal skeleton is
Latrunculiidae with polygonal large-meshed reticulation a tangential layer of choanosomal megascleres, being somewhat
formed by wispy tracts of anisostyles. Megascleres are typically plumose at the base of the ectosome. Megascleres are typically
smooth and sinuous, occasionally polytylote anisostyles to which smooth and sinuous, occasionally polytylote anisostyles to which
terminally spined styles or diactinal spicules may be added. Micro- terminally spined styles or diactinal spicules may be added.
scleres are anisodiscorhabds. Aciculodiscorhabds, amphiaster-like Microscleres are anisodiscorhabds. Aciculodiscorhabds, amphi-
microscleres, and acanthomicroxeas are present in one or other of aster-like microscleres, and acanthomicroxeas are present in one or
two species. Microscleres are disposed in a palisade in the outer other of two species. Microscleres are disposed in a palisade in the
ectosome. outer ectosome. Where known sponges are viviparous. lAtruneulia
714 Porifera· Demospongiae • Poecilosclerida • Latrunculina • Latrunculiidae

spp. produce pyrroloquinoline alkaloids discorhabdins and their cylindrical oscules, -5 mm height, and mammiform areolate pore-
derivatives. fields. Ectosome thick and leathery not separable from underlying
choanosome. Colour in life dark brown; in preservative pale yel-
Previous reviews low. The choanosomal skeleton is a very irregular polygonal-
meshed reticulation formed by wispy tracts of smooth styles
Du Bocage (1869); Carter (1879b); Topsent (1922, 1928c); (Fig. 4B). These tracts range in width from 91-120 /-Lm in thickness,
Ridley & Dendy (1887); Dendy (1921); de Laubenfels (1934); forming meshes that are 200 /-Lm wide. Towards the surface these
Burton (1934a); Hooper (1986); Wiedenmayer (1989); Kelly-Borges & spicules tend to be vertically arranged. The surface of the ectosome
Vacelet (1995); Urban et aZ. (2000); Alvarez et aZ. (2000b). is aligned with an erect layer of single discorhabds (Fig. 4C).
Beneath the discorhabds in the ectosome is a thick paratangential
Description of type species layer of densely interlocking megascleres, approximately 700/-Lm
wide. Megascleres (Fig. 4D): anisostyles are smooth, centrally
Latrunculia cratera du Bocage, 1869. thickened, fusiform and faintly sinuous, 510 (455-547) X
Synonymy. Latrunculia cratera du Bocage, 1869: 161, 16(16-18) /-Lm diameter. Microscleres (Fig. 4E-G): anisodiscorhabds,
pI. 11 fig. 2. the manubrium is an expanded spinose base armoured with two hori-
Material examined. Holotype: Lost. zontally arranged whorls of spines followed by a smooth slender
Description. Small thickly encrusting Latrunculiidae, with cylindrical shaft 20 /-Lm long and 7 /-Lm wide. Medium whorl is mid-
smooth, microscopically hispid surface. Surface usually with way between manubrium and apical whorl, this is circular, flat and
volcano-shaped or cylindrical oscules, together with mammiform to horizontally arranged, 30 /-Lm in diameter, similar in diameter to the
crater-shaped areolate porefields. Colour in life and in preservative subsidiary and apical whorl, the subsidiary whorl is more or less per-
unknown, (after du Bocage, 1869). The choanosomal skeleton con- pendicular to the shaft. The spines of the apical whorl are slanted
sists of a large-meshed polygonal reticulation formed by wispy tracts slightly upwards ending in a crown-like tuft of blunt terminally
of spicules with no spongin reinforcement. The ectosomal skeleton spined projections. The whorls are deeply notched along the rim and
is a tangential layer of megascleres. The surface of the sponge has divided into four segments each segment possessing a denticulate
a distinct palisade of anisodiscorhabds (modified from du Bocage, margin or 7-8 spines, 66(57-71) /-Lm in length.
no measurements given). Megascleres are smooth or polytylote Remarks. Although du Bocage (1869) failed to assign
anisostyles 160-190 X 6 /-Lm. Microscleres are anisodiscorhabds LatruncuZia to a family or order, he described the sponge L. cratera
40 /-Lm in length (after Burton, unpublished catalogue, BMNH). in conjunction with Podospongia Zoveni du Bocage (1869). Carter
Remarks. Du Bocage (1869) originally described (1879b) was the first to indicate affinity by placing Latrunculia
Latrunculia cratera from Cape Verde in the Atlantic Ocean, distin- purpurea in the Order Holorrhaphidota and Latrunculia
guishing it from Podospongia Zoveni by the presence of a 'distinct (=Negombata) corticata in the Family Ectyonida (Order
dermis of thorny microscleres' forming a 'complete envelope Echinonemata). His placement of the two species in different fam-
around the sponge'. He named the species Latrunculia cratera but ilies was based on the differing structure of the 'flesh' spicules.
did not give a generic diagnosis nor any indication of its phyloge- Ridley & Dendy (1886, 1887) included Latrunculia with
netic affinity. The original type material consisted of only one Spirastrella in Spirastrellidae on the basis that the discorhabds
specimen (a holotype) and was deposited in the Museu Bocage, were thought to be related to spinispires forming an ectosomal
Lisbon, Portugal. Unfortunately, du Bocage's sponge collection crust as do the spirasters of Spirastrella. Various other authors
was lost after the museum burnt down on in 1978. Alleged schizo- (Kirkpatrick, 1900a, 1903b, 1908c; Stephens, 1915a; Hentschel,
types of Latrunculia cratera have been reported in the Norman and 1914; Dendy, 1921) followed Ridley & Dendy's (1887) classifica-
Lervig collections in the BMNH, all of which have been examined tion for the placement of Latrunculia within Spirastrellidae, but
and all transferred to other genera. No other record of L. cratera Topsent (1922) placed the genus in the newly defined family
has been reported since its initial description by du Bocage, and Latrunculiidae together with Podospongia and Sigmosceptrella,
with the destruction of the holotype the species is clearly incertae thus separating the genus Latrunculia from Spirastrellidae. Various
sedis. There is no provision under the ICZN to nominate a new minor reorganisations in the taxonomic placement of Latrunculia
type species for a genus once this is fixed (Anon., 1999, ICZN were proposed, such as with the Family Choanitidae by de
Articles 66-70). Consequently, the concept of Latrunculia bocagei Laubenfels (1936a) and Bergquist (196Ia), but Levi (1973)
Ridley & Dendy is used here to emend the genus diagnosis as adopted Topsent's (1922) system of classification in placing the
a 'representative species' that exemplifies of Latrunculia given that genus with the Latrunculiidae in Poecilosclerida.
the holotype of L. cratera is confirmed destroyed and the original Twenty-seven species have been historically allocated to
description and diagnosis are completely inadequate to provide Latrunculia, giving the impression of a global distribution.
a firm concept for the genus. According to redefined criteria for Latrunculia in this work, how-
ever, only eleven of the originally recognised species now conform
Description of representative species to the strict definition of LatruncuZia. The remainders are assigned
to other genera, both within Latrunculiidae and elsewhere. An
Latrunculia bocagei Ridley and Dendy, 1886 (Fig. 4). exhaustive inspection of world-wide museum material (Samaai,
Synonymy. Latrunculia bocagei Ridley and Dendy, 1886: unpublished), has revealed the existence of a further nine previ-
492; 1887: 238-239, pI. 44 fig. 1, pI. 45 fig. 8. ously undescribed species which are now also assigned to the
Material examined. Holotype: BMNH 1887.5.2.237. genus. This brings the total number of species within Latrunculia
Holotype of variety: BMNH 1887.5.2.238 - Kerguelen. to 20 (Samaai, unpublished).
Description. Massive, hemispherical sponge, 3 X 2.5 X The genus MicrostyZifer Vacelet, 1969 (Vacelet, 1969: 179)
3.8cm diameter (Fig. 4A). Surface smooth with volcano-shaped or was originally described as having styles of three sizes with
Porifera· Demospongiae • Poecilosclerida • Latrunculina • Latrunculiidae 71S

Fig. 4. A-G, Latrunculia bocagei du Bocage (1869). A, holotype (after Ridley and Dendy, 1887: pI. 44 fig. 1). B-C, photomicrograph of skeletal archi-
tecture made from a slide of the holotype. E, drawing of discorhabds made from a slide of the holotype. D, F-G, SEM photos of spicule made from the
holotype (scales D, 150 f,Lm; F, 20 f,Lm). G, developmental stages of discorhabds (scale 20 f,Lm).

microspined heads, the smallest microstyles forming a palisade have been overlooked by Vacelet (1969) when initially describing
in the ectosome and present in the fascicles of the megascleres. this species because if they were definitely present they would
Vacelet (1969) considered the genus to be an axinellid, but had have been visible in the ectodermal palisade. Moreover, the species
reservations about this assignation. Re-examination of a micro- was described as hispid with projecting styles, whereas no
scope slide of the holotype of the type species, Microstylifer Latrunculiidae are hispid as they have a tangential ectosomal
rugosus Vacelet (MNHN DNlO) (by monotypy), courtesy of Rob megasclere skeleton. The only evidence for affinity of Microstylifer
Van Soest, revealed that the megascleres had constricted necks, fre- with Latrunculiidae, apart from the discorhabds in the slide, is the
quently malformed heads, and they were occasionally polytylote. 'vague tracts of megascleres' in the choanosome and the oblique
The distal ends were frequently blunt. The spicule slide contained layer of megascleres at the ectosome. However, above the oblique
only a few spicules, including foreign sigmas and several latrun- layer there is a palisade of microstyles. If discorhabds were native to
culid discorhabds, indicating a potential affinity with Latrunculiidae. this species they would be in this layer, and no Latrunculiidae in our
One species of Latrunculia is also known to possess micros pined knowledge has microstyles. Until fresh material is examined, or
styles, but nevertheless, it is very unlikely that discorhabds would a histological section of the holotype can be examined, this species
716 Porifera· Demospongiae • PoeciloscIerida • Latrunculina • Latrunculiidae

Fig.5. A-C, Sceptrella regalis Schmidt, 1870. A, drawing of discorhabd made from a slide of the holotype. B-C, photomicrograph of microscleres made
from a slide of the holotype. B, mature sceptres. C, developing isoconicodiscorhabd. D-G, Strongylodesnw areolata Levi (1969). D, holotype (diameter
25 X l3 X 30 mm) (after Levi, 1969: 956, pI. 2 fig. 5). E-F, photomicrograph of skeletal architecture made from a slide of the holotype. G-H, drawing and
SEM photo of strongyle from the holotype.

is a very questionable Latrunculiidae, and presently Diagnosis

Poecilosc1erida incertae sedis.
Encrusting sponges with mammiform papillae and areolate
porefields; surface even, non-hispid. Colour in life and in preserva-
SCEPTRELLA SCHMIDT, 1870 tive rose-tan to grey. Microsc1eres are isoconicodiscorhabds with
furcate whorls of spines, on a straight axial rod, megasc1eres are
Synonymy smooth styles or anisostyles. Choanosomal architecture consists of
anisostyles arranged in wispy irregular polygonal meshes, ectoso-
Sceptrella Schmidt, 1870: 58. mal skeleton consists of a tangential layer of anisostyles, with an
erect palisade of microsc1eres.
Type species
Previous review
Sceptrella regalis Schmidt, 1870 (by monotypy).
None.
Definition
Description of type species
Latrunculiidae with anisostyles arranged in wispy irregular
polygonal meshes, ectosomal skeleton consists of a tangential layer Sceptrella regalis Schmidt, 1870 (Fig. 5).
of anisostyles, with an erect palisade of microsc1eres, microsc1eres Synonymy. Sceptrella regalis, Schmidt, 1870: 58, pI. 5
are isoconicodiscorhabds ('sceptres') with furcate whorls of fig. 24; Latrunculia regalis; Van Soest, 1984b: 149; Wiedenmayer,
spines, on a straight axial rod. 1994: 73.
Porifera· Demospongiae • PoeciloscIerida • Latrunculina • Latrunculiidae 717

Material examined. Holotype (fragment): BMNH Definition

1870.S.3.22 (spicule slide) - Rorida, 262 fathoms.
Description (after Schmidt, 1870). Sponge encrusting, Latrunculiidae with wispy polygonal reticulation of
bearing oscular processes, surface even, non-hispid. Colour in life strongyles with no distinction between primary and secondary
unknown; in preservative grey, texture unknown. The choanosomal tracts. The ectosomal skeleton consists of a clear thick band of col-
architecture consists of anisostyles arranged in wispy irregular lagenous mesohyl, the base of which has a thin band of paratan-
polygonal meshes, the base of the choanosome is somewhat radial. gential strongyles. Megascleres are smooth or terminally spined
The ectosomal skeleton consists of a tangential layer of anisostyles strongyles in one size category.
with isoconicodiscorhabds scattered in the choanosome and form a
palisade at the surface. The megascleres are smooth centrally thick- Diagnosis
ened fusiform anisostyles 410 X ll/-Lm (Fig. SB). Microscleres are
isoconicodiscorhabds, or 'sceptres' (Fig. SA, C). These spicules Massive spherical or hemispherical sponges with elevated
have a stout shaft, the manubrium is surrounded by a ring of volcano-shaped or cylindrical oscules and fungiform or concave
obliquely directed spines, and at equal distances along the shaft are elliptical areolate porefields. Colour brown to reddish brown. The
two whorls of projecting similar spines, which are armed with choanosomal skeleton consists of a wispy polygonal reticulation
secondary spines. The apical whorl is oblique and arranged in a of tracts of strongyles with no distinction between primary and
crown-like structure. There are three size categories: 50 /-Lm; 70 /-Lm; secondary tracts. The ectosomal skeleton consists of a clear thick
127/-Lm in length. band of collagenous mesohyl, the base of which has a thin band
Remarks. Schmidt (1870) presumed that the type species of of paratangential strongyles. The thickness of this paratangential
Sceptrella, S. regalis, collected off the Florida coast, was closely layer differs considerably between species. Most species have
related to Latrunculia as it had ani so styles and a similar skeletal a very thick layer. Strongyles support the raised surface structures
architecture, although he recognised that the microsclere was spin- and protrude beyond the surface. Megascleres are smooth, or
ose and bore little resemblance to the lobed discorhabd microscle- terminally spined strongyles in one size category. Microscleres
res characteristic of Latrunculia. When Schmidt (187S) described absent. Strongylodesma spp. contain pyrroloquinoline alkaloids
Sceptrella triloba from Norway he synonymised Sceptrella such as batzellins, isobatzellins, discorhabdins and their
with Latrunculia on the basis of identical skeletal and spicule derivatives.
morphology. Thiele (1905) and Burton (unpublished catalogue in
the BMNH), transferred the species Sceptrella triloba to Previous review
Latrunculia.
However, Carter (1879b: 358-359, pI. 29 figs 14-15) clearly None.
illustrated the difference between the microscleres of Sceptrella
and Latrunculia, stating that 'the spines may be microspined as in Description of type species
Sceptrella regalis Schmidt, or the groups on the shaft may be trans-
formed into circular plates with serrated margins, as in Latrunculia Strongylodesma areolata Levi, 1968 (Fig. 5).
cratera'. Examination of the spicule slide of holotype Sceptrella Synonymy. Strongylodesma areolata Levi, 1969: 959, pI. 2
regalis reveals that although the microscleres of this genus arise fig.4c.
from a straight protorhabd as in Latrunculia, the post development Material examined. Holotype: MNHN LBIM DCL 1425-
of spicules differs. The whorls of spinose projections develop Vema Seamount.
simultaneously in Sceptrella in contrast to the asynchronised Description. Sponge massive encrusting, 2S X 13 X 30 mm
development of the different whorls in Latrunculia, and the spines in diameter (Fig. 5D); surface smooth with flat circular or elliptical
develop directly from the shaft and are smooth or entirely areolate porefields which are 0.5-3.5mm in diameter. Oscules are
microspined. Sceptrella is upheld here as a valid genus of raised and membranous. Colour in life reddish-brown, beige in pre-
Latrunculiidae. servative. Texture firm and compressible. Choanosomal architec-
Several specimens identified as Latrunculia have been trans- ture consists of an irregular polygonal-meshed reticulation of
ferred to Sceptrella. Presently we recognise six valid species of wispy tracts of faintly microspined strongyles (Fig. SE). These
Sceptrella, including S. regalis Schmidt, S. triloba Schmidt, tracts range in width from 9-100 /-Lm in thickness, forming meshes
S. biannulata Topsent, S. insignis Topsent, and S. insignis var. that are 120/-Lm wide. The ectosomal skeleton consists of a clear
regularis. At least one more species of Sceptrella is known from thick band of collagenous mesohyl 227-270/-Lm thick, the base of
Icelandic waters, but remains unnamed (Samaai, unpublished). which is a thin band of paratangential strongyles (20/-Lm deep)
(Fig. 5F). Strongyles protrude beyond the surface (Fig. 5F) in a
haphazard manner. Megascleres are faintly terminally microspined
strongyles, 299(282-319) X 6(5-7) /-Lm in length (Fig. 5G-H).
STRONGYLODESMA LEVI, 1969 Microscleres absent.
Remarks. The holotype of Strongylodesma areolata was
Synonymy first described by Levi (1968) from the Vema Seamount in the
south Atlantic Ocean, and included in Spirastrellidae. Conversely,
Strongylodesma Levi, 1969: 959. an unnamed species of Strongylodesma described by Battershill
(in Levi 1998) was placed in Phoriospongiidae (=Chondropsidae)
Type species (Poecilosclerida) on the basis that some other taxa of
phoriospongiids also possessed areolate porefields, and no doubt
Strongylodesma areolata Levi, 1969 (by monotypy). also because many have strongyles as their primary megascleres
718 Porifera· Demospongiae • Poecilosclerida • Latrunculina • Latrunculiidae

(although these are usually coupled with chelae microscleres, Description of type species
e.g., Strongylacidan etc.). If other characters are taken into
account, however, including skeletal architecture, base pigmenta- Tsitsikammafavus gen. nov., sp. nov. (Fig. 6).
tion, and biochemistry, the majority are shared with Latrunculia. Material examined. Holotype: BMNHI997.7.3.2
Species of Strongyladesma differ primarily from those of Rheeders Bay, Tsitsikamma National Park, 34°lO'S, 23°54'E,
Latrunculia and Sceptrella in that they lack microscleres and that South Africa, depth 22m, colI. P. Coetzee, 1995. Paratypes:
the megascleres are strongyles. BMNH - several specimens from Rheeders Bay, Knoll reef
Seven species of Strongyladesma are now recognised, including Tsitsikamma National Park, and Algoa bay, Port Elizabeth, South
three species from South Africa, a new species from New Caledonia Africa. Habitat. Depth range 18-22m on rocky substrate, associ-
(Vacelet & Kelly in Levi, 1998), and one from Tonga (Kelly, unpub- ated with soft and hard coral and other invertebrates.
lished). A further two Caribbean species are now recognised amongst Description. Sponge is massive, ovo-spherical, sometimes
the Harbor Branch Oceanographic Institution collections (HBOM thickly encrusting, 1-8cm high and IO-l5cm wide (Fig. 6A).
003:00922) and (HBOM 003:00050, HBOM 003:00051). Texture is hard and only slightly compressible, resilient and leath-
ery; surface smooth and crowded with large single to multicham-
bered cylindrical lance-shaped oscules, 1-4mm wide, 8mm high
TSITSlKAMMA GEN. NOV. and many pedunculate cauliform areolate porefields, 3 mm wide,
5-6 mm high, with no membrane; colour in life is dark brown or
Type species liver brown and between the cauliform structures dark turquoise, in
alcohol, ectosome dark turquoise, choanosome dark brown. The
Tsitsikammafavus gen. nov., sp. nov. choanosome is divided into honeycomb-like chambers and convo-
luted layers by very thick reinforced tracts of anisostyles, these
Definition tracts range in width from 1000-1700 f..lm thick, forming meshes
that are elliptical in shape and 5800 f..lm wide. These tracts are lined
Latrunculiidae with elliptical honeycomb-like chambers with microscleres, suggesting that they may represent an early
(sensu stricto), or a single sac-like chamber (sensu lata), chamber ectosermal surface, and that the sponge grows in size and volume
walls being very thick tracts of style megascleres. Microscleres are by putting on new chambers, as in many species of Petrosia and
isochiarhabds. Caelocarteria. Within and between the chambers and convoluted
layers the skeleton consists of an ill-formed, irregular reticulation of
Diagnosis small anisostyles, these tracts range in width from 100-150 f..lm (Fig.
6B). Microscleres are isochiadiscorhabds and these are abundant
Massive encrusting, hemispherical or pedunculate sponges throughout the choanosome. The ectosome is composed of a thick
with a smooth undulating surface, covered with large robust cylin- dense feltwork of tangential and paratangential anisostyles approxi-
drical or volcano-shaped oscules and raised cauliform areolate mately 900 f..lm wide. This layer is present in the fistulae with
porefields. Colour in life is dark liver brown to dark turquoise, or anisostyles disposed in a compact regular vertical to oblique arrange-
pinkish brown. The texture is extraordinarily tough and leathery. ment supporting the cauliform areolate structures. A single layer of
Megascleres are styles, microscleres are isochiarhabds. Elliptical erect isochiadiscorhabds (48 f..lm wide) lines the surface of the ecto-
honeycomb-like chambers, or a single sac-like chamber surround- some (Fig. 6C). Megascleres in two categories of anisostyles, (1)
ing a soft mesohyl with wispy tracts characterise the genus. The slightly curved and thickened centrally, 621(537-700) X 14(14) f..lm,
walls of the chambers or sac are extremely thick, up to 1500 f..lm, n = 20; (2) thinner slightly curved centrally: 530(480-566) X
and visible with the unaided eye. The chambered holotype is 9.6(9.6)f..lm (Fig. 6D). Microscleres are isochiadiscorhabds with
incompressible. Microscleres are present in an irregular palisade three whorls of cylindrical, conical tubercles, the apex of each tuber-
on the surface ectosome and lining the internal tracts. Chemistry cular projection is acanthose: 48(41-60) X 9(7.2-9.6)f..lm (Fig.
includes pyrroloquinoline alkaloids (tsitsikammamines), and the 6E-G). Reproduction: oocytes present in specimens but no further
discorhabdin derivatives 14-Bromo-3-dihydrodiscorhabdin C and indication of reproductive mode yet corroborated.
14-Bromodiscorhabdin C. Etymology. favus = honeycomb (L.).
Remarks. The huge spicule tracts that compartmentalise the
Etymology body of Tsitsikammafavus are remarkable and clearly differentiate
it from Latrunculia, Strongylodesma and Sceptrella, and other
Tsitsikamma means 'clear water' in Khoi language, and the related demosponges. The characters that link the genera are the
genus has been named for the type locality, a unique area of south- base pigmentation, chemistry, the surface palisade of microscleres
east Africa. and the soft skeletal parts. A second pendunculate sac-like species
is known from Algoa Bay, Port Elizabeth, South Africa.
Previous reviews

Pertaining only to the chemistry literature: Urban et al. ACKNOWLEDGEMENTS

(2000), Beukes (2000), Beukes et al. (in press).
We are indebted to The Royal Society London and The
Geographic distribution National Research Foundation, South Africa, for provision of
a PhD scholarship for TS, and for general funding under their joint
Rheeders Bay, Tsitsikamma National Park, East Coast, South Capacity-Building Programme (1996-2001). We also thank the
Africa; Algoa Bay, Port Elizabeth, East Coast, South Africa. Lottery Science Grants Board of New Zealand and UNITEC
Porifera· Demospongiae • Poecilosclerida • Latrunculina • Latrunculiidae 719

E G

Fig. 6. A-G, Tsitsikammafavus gen. nov. sp. nov. A, holotype. B-C, photomicrograph of skeletal architecture made from a slide ofholotype. D-F, megas-
clere and microsclere drawing made from the holotype slide. G-I, SEM photos of isochiasdiscorhabd and terminations (scales G, 20 fLm; H, 3.8 fLm; I, 10
fLm). J, developmental stages of isochiadiscorhabds (scale 20 fLm).

Institute of Technology, Auckland, for laboratory support in 1991 collection of new sponge material that has advanced our knowl-
when the bulk of this research was carried out. The National edge of Latrunculiidae diversity in South Africa and New Zealand.
Institute of Water & Atmospheric Research (NIWA), New Zealand, We are very grateful to Professor Mike Davies-Coleman, Rhodes
has supported the preparation of this contribution and, with University, Grahamstown, South Africa, for assisting our under-
the National Museum of New Zealand Te Papa Tongarewa, standing of latrunculiid chemotaxonomy. This is a contribution
Wellington, provided specimens for general reference. We are from the Coral Reef Research Foundation, Micronesia, and Harbor
indebted to The Coral Reef Research Foundation, Micronesia, for Branch Oceanographic Institution Contribution No. 1441.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Order Halichondrida Gray, 1867

Rob W.M. Van SoestI & John N.A. Hooper2

I Zoological Museum, University of Amsterdam, P.O. Box 94766, 1090 GT, Amsterdam, Netherlands. (soest@science.uva.nl)
2 Queensland Museum, P.O. Box 3300, South Brisbane, Qld, 4101, Australia. (JohnH@qm.qld.gov.au)

Order Halichondrida Gray (Demospongiae) is here employed in the redefined sense of Van Soest et al. (1990; with subsequent published
adjustments), and includes the former much more restricted Halichondrida and part of the former Axinellida as proposed in earlier classi-
fications (e.g., Levi, 1973; Bergquist, 1978; Hartman, 1982). The order is defined to include sponges with smooth styles, oxeas or
strongyles, occurring often in tandem within the same sponge, but usually without distinct localization, and lacking microscleres other
than spined microxeas and raphides. Flexuous, sinuously curved, or contorted megascleres are relatively common. Skeletal architecture is
basically plumoreticulate, but in various groups this is modified to dendritic columns of megascleres or confused bundles with many loose
single megascleres. Ectosomal skeleton ranging from specialized, tangentially intercrossing or obliquely arranged palisades or bouquets
of mega- and microscleres, to absent or undifferentiated, or replaced by thick organic skin, or rarely with sand grains. Five families are
recognized, based on features of the ectosomal and choanosomal skeleton. Axinellidae, Bubaridae and Dictyonellidae lack a specialized
ectosomal skeleton, whereas Halichondriidae and Desmoxyidae with few exceptions have an ectosomal skeleton consisting of spicules
arranged tangentially or in the form of palisades/bouquets. Axinellidae differ from Bubaridae and Dictyonellidae in having a well-developed
dense reticulation of megascleres, often showing axial and extra-axial differentiation, and their surface is velvety or microhispid due to
projecting choanosomal spicule bundles. Bubaridae are thinly encrusting sponges with monactinal megascleres erect on a basal mass of
bent or flexuous diactinal spicules. Dictyonellidae have a fleshy organic skin, and lack a neat reticulation, megascleres usually forming
dendritic columns or a vague confused reticulation of styles, rarely oxeas. Desmoxyidae have spined microscleres or smaller megascleres
at the surface, either tangentially or perpendicularly arranged. Halichondriidae have only smooth megascleres, and with few exceptions
have a tangential crust of intercrossing oxeas, rarely styles. Where known most Halichondrida are oviparous, but Halichondria,
Hymeniacidon (Halichondriidae), Scopalina and Svenzea (Dictyonellidae) are viviparous; the latter have very large ovoid larvae.
Halichondrida are common inhabitants of all marine habitats in all oceans and seas.
Keywords: Porifera; Demospongiae; Halichondrida; Axinellidae; Bubaridae; Dictyonellidae; Desmoxyidae; Halichondriidae.

DEFINITION, DIAGNOSIS, SCOPE absent or vaguely developed, resulting in dendritic skeletal colunms
or entirely confused skeletons. Spongin and interstitial collagen may
Synonymy be present in considerable quantities, also binding the spicule bun-
dles, but may also be vestigial, resulting in lax or crumbly texture.
Halichondriadae Gray, 1867a: 518. Halichondrina Vosmaer, Megascleres styles, oxeas and strongyles, often intergrading into each
1887: 335. Axinellides Levi, 1953a: 3 (in part). Axinellida Levi, other with apices ranging from sharply and gradually tapering, to
1957b: 181 (in part). Halichondrides Levi, 1953a: 3. Clavaxinellides bluntly rounded, often with characteristic stair-stepped appearance.
Levi, 1956a: 167 (in part). Clavaxinellida Levi, 1957b: 181 (in part). Styles may occasionally have a slightly developed subterminal tyle.
[Claraxinellida] Levi, 1957b (Iapsus, p. 183). Halichondriida The respective megasclere types frequently occur together in a single
Wiedenmayer, 1977: 148 (nomen correctum). sponge, but may also be confined to exclusively oxeas or styles;
strongyles always occur together with one of the other megasclere
Definition types. Diactinal spicules (oxeas, strongyles) in some genera and
species may be flexuous, sinuously curved or contorted. Microscleres
Ceractinomorpha Demospongiae with styles, oxeas, strongyles except trichodragmata are absent, but some genera have the smallest
or intermediate spicules, of widely diverging sizes, and not func- category of oxeas or styles ornamented with spines or, in one genus,
tionally localized; skeleton plumoreticulate, dendritic or confused; disc-like outgrowths; such spicules may be conveniently named
microscleres if present microxeas and/or trichodragmas. microscleres but the homology with true microsc1eres is debatable.

Diagnosis Scope

Encrusting, massive, lobate, digitate, fistular, ramose, tubular or Five families are recognized in the order, Axinellidae Carter,
flabellate sponges. Surface smooth, microhispid, conulose or 1875, Bubaridae Topsent, 1894, Desmoxyidae Hallmann, 1916,
grooved. Ectosomal skeleton consisting of a crust of intercrossing Dictyonellidae Van Soest et aI., 1990 and Halichondriidae Gray,
megascleres, or bouquets/palisades of small megascleres c.q. microx- 1867a. Together they contain many hundreds of species occurring
eas, but also frequently absent or thickly organic. Choanosomal in all marine habitats and in all geographic regions. Axinellidae
skeleton basically plumoreticulate, consisting of plumose sheets or and Halichondriidae are particularly speciose, Bubaridae and
bundles of megascleres obliquely or squarely interconnected by Dictyonellidae are less diverse, with Desmoxyidae intermediate in
shorter plumose bundles; frequently the interconnecting bundles are diversity.

721
722 Porifera· Demospongiae • Halichondrida

Remarks cycle information, discovered broad group-related modes of repro-

duction. In large groups of sponges, like tetractinellids and
The plumoreticulate skeletal architecture may be a synapomor- hadromerids, all species investigated were found to be oviparous,
phy for a much wider group of Demospongiae (see Van Soest, whereas other groups, like haplosclerids, poecilosclerids and 'ker-
1991). This character is progressively lost in the families atose' sponges appeared to be always brooding their larvae. He
Dictyonellidae and Halichondriidae. However, a basic plumorectic- later proposed (Levi, 1953a, 1956a, 1957b) to subdivide the
ulate or plumose architecture is found even in these taxa with few Demospongiae in two subclasses, which were based on ovipary
exceptions. Although the combination of styles and oxeas is also (Tetractinomorpha) and vivipary (Ceractinomorpha). Levi (I.c.)
found in a few poecilosclerid Microcionina and Myxillina, in these demonstrated that within the Halichondrida some families were
groups they are localized to particular regions and/or have several oviparous (Axinellidae sensu de Laubenfels, 1936a), other families
geometric categories. For instance, in Myxillina stylote choanoso- were viviparous (Halichondriidae sensu de Laubenfels). This led
mal spicules are combined with ectosomal diactines; in the ras- him to propose a subdivision of the group, with orders Axinellida
pailiid Echinodictyum choanosomal oxeas are arranged into tracts (families Axinellidae, Bubaridae, Desmoxyidae, Euryponidae,
which are echinated by (acantho-)styles. Indiscriminate occurrence Hemiasterellidae, Raspailiidae, Rhabderemiidae, Sigmaxinellidae
of styles and oxeas is not found. Localized spicules in Didiscus, and Trachyc1adidae), assigned to Tetractinomorpha, and
Mynnekiodenna and Higginsia are conveniently considered Halichondrida s.s. (families Halichondriidae and Hymeniacidonidae)
microscleres. Sinuous longer or shorter diactinal spicules are found assigned to Ceractinomorpha. Axinellida was further underbuilt
in various groups distributed throughout the order: e.g., Axinella by emphasizing the frequent occurrence of axially condensed
cannabina, Phakellia, Auletta (Axinellidae), Bubaris (Bubaridae), skeletons in members of this group (see Levi, 1973 for a compre-
Acanthella, Dactylella (Dictyonellidae), some Axinyssa and some hensive treatment). The combination of axial skeletons and asters
Topsentia (Halichondriidae). This distribution may be interpreted in Hemiasterellidae and axial skeletons and spinispirae in
as retention of ancestral spicules, possibly related to 'lithistid' Trachycladidae led Levi (1956a: 167) to propose a formal higher
spicules, through Monocrepidium (Bubaridae) and fossil genera taxon including both Hadromerida and Axinellida, the superorder
such as Cephalorhaphidites and Megaloraphium, and thus consti- Clavaxinellides, which he later (Levi, 1957b: 181) corrected to
tute an underlying synapomorphy of the order. Clavaxinellida (and misspelled as Claraxinellida on p. 183). These
combined proposals were received with approval by most contem-
Taxonomic history porary authors (e.g., Vacelet, 1969; Bergquist, 1978; Hartman,
1982), and much effort was directed towards obtaining additional
Gray (1867a: 518) erected a family Halichondriadae which support for this scheme. From 1985 onwards, again the consensus
contained a mixture of Haplosclerida, Poecilosclerida and over these groups was broken, by independent studies of Hooper
Hadromerida, and indeed included Halichondriidae. Later Vosmaer (1990b) on the relationship of oviparous ('tetractinomorph')
(1887: 335) used the suborder name Halichondrina for a similarly Raspailiidae and viviparous ('ceractinomorph') Microcionidae,
wide group of Ceractinomorpha (dubbed Comacuspongida by and Van Soest et al. (1990) and Van Soest (1987a, 1991) on the
him), basically only excluding the 'Keratosa'. Ridley & Dendy relationships ofAxinellidae and Halichondriidae. Criticism of the
(1887: Iviii) employed a suborder Halichondrina in the order Levi system amounted to the conclusion that too much emphasis
Monaxonida Sollas, 1883, which covered the same groups. was placed on only a few characters and conflicting character dis-
Halichondrida were narrowed down by Topsent (1928c: 37) to a tributions were ignored. These heavy-weight characters (vivipary,
group of four families (Axinellidae, Astraxinellidae, Bubaridae, axial condensation of the choanosomal skeleton) were moreover
and Heteroxyida), which together cover most of the present con- given as much weight for their presence as for their (primitive)
cept of Halichondrida. Remarkably, the family Halichondriidae absence, a philosophically unsound character treatment: one can-
was merged with Axinellidae, and some genera now considered not define a group of organisms on the fact that it does not possess
Halichondriidae like Spongosorites, were excluded. The most a certain feature, unless one assumes it was lost. Both Hooper (I.c.)
important difference with present content of Halichondrida is the and Van Soest (I.c.) brought forward contrasting evidence and the
inclusion of Astraxinellidae (= Hemiasterellidae), which are here latter suggested to abandon the (Clav-)Axinellida sensu Levi, in
considered to belong to Hadromerida on account of the possession favour of a more conservative classification. Halichondrida sensu
of asters. De Laubenfels (1936a) distributed the latter group over Topsent and de Laubenfels, with necessary emendations, contain-
several orders and families, and retained the remaining groups in a ing core families Axinellidae and Halichondriidae, along with
Halichondrida of five (the apparently preferred number of this Bubaridae, Dictyonellidae and Desmoxyidae, fit better with the
author) families: Axinellidae, Halichondriidae, a new family available morphological characters. An initial 28S rRNA sequence
Hymeniacidonidae (containing in addition to Hymeniacidon, analysis performed with a 27 species sample (Alvarezet al.,
many genera now united in Dictyonellidae), a new family 2000a) generally confirmed the likely close relationships of these
Semisuberitidae (containing an odd assemblage of poecilosclerid families. Chemosystematic evidence (Van Soest & Braekman,
and haplosclerid genera), and a new family Monanthidae (for 'sub- 1999) also points at close relationships with two families of com-
lithistid' genera). By this subsequent building on previous classifi- pounds (pyrrole-2-carboxylic derivatives and cyanoditerpene-
cation schemes, and by increasing the focus of the group by derivatives) distributed over species and genera ofAxinellidae,
removing outlying families and genera, a continuity and consensus Bubaridae, Dictyonellidae and Halichondriidae. The pyrroles
was achieved, which allowed some form of stable use of the classi- also occur in Agelasida (in both families Agelasidae and
fication. This stability was significantly disrupted by new, non- Astroscleridae), which may be evidence along with 28S rRNA
skeletal evidence produced by Levi (e.g., 1951, 1953a, 1956a, sequence data (Chombard et al., 1997) to support a possible future
1957b and following). This author, after a long and careful study of merger of Agelasida and (part of) Halichondrida. An abstract of a
sponges in the field, giving him access to previously unknown life study by Chombard et al. (1999) reports the finding of 28S rRNA
Porifera' Demospongiae • Halichondrida 723

sequence similarity between Suberitidae, Polymastiidae and and (parts of) Halichondrida, or a merger of Hadromerida and
Halichondriidae, resulting even in a proposal for a new suborder (parts of) Halichondrida. Future studies with larger samples of rep-
Suberitina comprising these families, to be assigned to resentative taxa and using more than a single gene are necessary to
Hadromerida. Whereas such studies and results indicate that the enable confmnation or refutation of the present concept, which is
present concept of Halichondrida may be subject to future change, based on morphological analysis of all type specimens and species
it is premature to follow suggestions such as a merger of Agelasida and a large sample of additional species.

KEY TO FAMILIES

(1) Ectosomal specialization present, either in the fonn of a tangential crust or single spicule layer, a palisade of smaller and/or larger
megascleres or erect or scattered microscleres; surface smooth, but may be wrinkled or thrown up into folds and depressions ...... 2
No ectosomal specialization, surface velvety, or hispid due to single projecting spicules, or fleshy-organic, without
tangential spicules ................................... ...... .... ...... ...... ..................... .......... ... ... ..... ................ ................ ............ .... ...... .......... ............. 3
(2) At the surface there is a tangential layer or a palisade of spiny smaller oxeas or strongyles ......................................... Desmoxyidae
No spiny megascleres or microscleres; surface skeleton consists either of a tangential crust or tangential single spicules, or palisades
c.q. bouquets of smaller spicules mingled with larger choanosomal spicules ........................................................... Halichondriidae
(3) Surface velvety or hispid, due to projecting single spicules or choanosomal spicule tracts; sponges firmly resilient ........................ 4
Surface fleshy-conulose, with organic skin thrown up into conules by choanosomal fibres or spicules; comes off in flakes when
attempted to obtain a surface peel................ ... ....... ...... ....... .... ....... ... ................. ... ............ ...... .......... ........... ................. Dictyonellidae
(4) Thinly encrusting, not exceeding 1 cm thickness, strongly hispid, due to erect monactinal megascleres, with heads embedded in
a basal layer of interlacing megascleres ...... ........................... .............. ....... .......... ............ ...... ............... ............ .......... ....... Bubaridae
Elaborate growth fonns; surface microhispid, often velvety ............................................................................................................... 5
(5) Choanosomal skeleton reticulate, often with axial and extra-axial differentiation ............................................................ Axinellidae
Choanosomal skeleton confused ................................................................................................... Halichondriidae (genus Axinyssa)
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademicIPlenum Publishers, New York, 2002

Family Axinellidae Carter, 1875

Belinda Alvarez l & John N.A. Hoope.-2

1 Divisionof Botany and Zoology, Australian National University, Canberra, ACT 0200. Present address: Museum and Art Gallery of the
Northern Territory, P.O. Box 4646, Darwin, NT 0801, Australia. (belinda.glasby@nt.gov.au)
2 Queensland Museum, P.O. Box 3300, South Brisbane, Qld, 4101, Australia. (JohnH@qm.qld.gov.au)

Axinellidae Carter (Demospongiae, Halichondrida) contains 10 valid genera (of 90 nominal genera) and approximately 300 species
described so far worldwide. The family has a wide distribution from shallow to deep waters (down to 1800m depth). Species are gener-
ally erect, branching, fan-shaped or tubular, but encrusting and massive forms also occur. Species have no specialised ectosomal skeleton.
The choanosomal skeleton is generally differentiated in axial and extra-axial regions with two basic types of skeletal architecture:
(1) ascending and anastomosing spicule tracts, spongin fibres cored with spicules or fibrofascicles radiating to the periphery and ending
in surface processes; (2) plumose to plumoreticulated. The axial skeleton of some species is either compressed or absent. Megascleres are
styles, oxeas (and transitional forms) and sinuous strongyles in any combination. Bubaridae is excluded from synonymy with Axinellidae
and reinstated as a valid taxon in Halichondrida.
Keywords: Porifera; Demospongiae; Halichondrida; Axinellidae; Auletta; Axinella; Cymbastela; Dragmacidon; Dragmaxia;
Pararhaphoxya; Phakellia; Phycopsis; Ptilocaulis; Reniochalina.

DEFINITION, DIAGNOSIS, SCOPE trichodragmata, generally difficult to observe and located near the
periphery.
Synonymy
Scope
[Phakellidael Gray, 1867a: 503, 518; Gray, 1872a: 447
(nomen oblitum, Article 23.9 ICZN (Anon., 1999». Axinellidae A literature review revealed a total of 92 nominal genera that
Carter, 1875c; Lendenfeld, 1889a: 903 (in part); de Laubenfels, have been erected in, or transferred to, the family at one time or
1936a: 127 (in part). another. Of these, only 23 genera belong in the Axinellidae with 10
valid genera (Le., Auletta, Axinella, Cymbastela, Dragmacidon,
Definition Dragmaxia, Pararhaphoxya, Phakellia, Phycopsis, Ptilocaulis,
Reniochalina) and the rest considered junior synonyms. The total
Halichondrida without specialised ectosomal skeleton; with number of species in the family according to Hooper & Levi
velvety or microhispid surface; with choanosomal skeleton of (1994) is approximately 300 worldwide, but given the uncertainties
ascending spiculo-fibres connected irregularly by loose spicules in the taxonomy of this family the actual number may differ
and short tracts, or plumoreticulated with ascending plumose tracts substantially above or below that figure.
connected by thinner ones or single spicules. Megascleres are
oxeas, anisoxeas, styles, sinuous strongyles in any combination. Distribution
Microscleres are raphides, single or in trichodragmata.
According to Van Soest (1994), the Axinellidae has a cosmo-
Diagnosis politan distribution without any obvious or distinct centre of diver-
sity. Regional revisions record 15 species for New Caledonia
Encrusting, massive, branching, fan-shaped and tubular; gen- (Hooper & Levi, 1993b), 11 for New Zealand (Bergquist, 1970),
erally red, orange or yellow sponges; surface is usually velvety to 21 (in 7 genera) from the Central-West Atlantic region (Alvarez
hispid due to the projection of choanosomal spicules and without et al., 1998) and 17 for the NE Atlantic (Van Soest et al., 2000).
specialised ectosomal skeleton. Choanosomal skeleton is formed Other estimates are: 57 species described for Australia (Hooper &
by ascending spiculo-fibres that radiate to the periphery, connected Wiedenmayer, 1994); 130 species from the Indo-west Pacific
irregularly by loose spicules and short tracts, and ending in skeletal region, of which 45% are represented in Australian waters (Hooper &
projections at the surface, or plumoreticulate with ascending Levi, 1994); 63 species for the South China Sea (Hooper et aI.,
plumose tracts of oxeas and styles connected regularly by pau- 2000b) and 30 for the Mediterranean (Van Soest, unpublished
cispicular or multispicular tracts, or reticulated with main spicule data).
tracts of sinuous or vermiform strongyles, or anisoxeas, plumo-
echinated by single spicules. In some species the choanosomal History and biology
skeleton is divided into an axial region, where it often becomes
compressed, and an extra-axial region with plumoreticulated skele- The Axinellidae was initially erected by Carter (1875c: 133,
ton. Megascleres are mainly oxeas, anisoxeas with or without tele- 145) as 'Axinellida' under the order Echinonemata, to include
scoped tips or microspines, straight or sinuous styles, strongyles sponges with horny fibres 'echinated with proper spicules project-
are always sinuous. Microscleres are raphides, single or in ing from the interior of the fibre'. Carter (1883b: 316) recognised

724
Porifera' Demospongiae • Halichondrida • AxinelIidae 725

the simplicity of his previous diagnosis and extended the definition of these genera are polythetic, in that they cannot be defined by
of the family as: sponges with an axial and extra-axial skeleton; a single set of diagnostic characters. This makes allocation of
generally with sinuous oxeas, located in the core of the fibres and species problematical, and raises the likelihood that genera contain
echinated by styles of two size categories; with both types of heterogeneous assemblages of species.
spicules occurring in the axial skeleton. He recognised that the type The higher taxonomic position of the family Axinellidae has
and location of the spicules could vary among species, but empha- been very unstable and subjected to different higher taxonomic
sised that the presence of spicules echinating the fibres charac- assignments. Topsent (1894c; 1928c) and de Laubenfels (1936a;
terised all species and therefore were diagnostic for the family. 1950a) considered it belonged to the Halichondrina (Vosmaer
He also suggested that Axinella Schmidt, 1862 and Phakellia 1885b), a suborder of the Monaxonida, but later elevated to full
Bowerbank, 1862b were typical representatives. Carter (1885d: order status. Dendy (1916a; 1922b) had a different view, consider-
348) further expanded the definition ofAxinellidae to include one ing it closely related to the family Desmacidonidae (currently in
of the most controversial characters in contemporary sponge taxon- the order Poecilosclerida), and tentatively included it as a subfam-
omy, 'the compression of the axial skeleton'. He also grouped ily of this group (which he considered to be polyphyletic).
axinellid sponges into several assemblages - branching, stipitate, But one of the most remarkable changes influencing the
flabelliform, vasiform and massive shapes - however, he recog- taxonomic position of the Axinellidae was the subdivision of
nised that form was a character with poor informative value. the Demospongiae into the subclasses Ceractinomorpha and
Ridley & Dendy (1886; 1887) described the family again Tetractinomorpha by Levi (1953a). As consequence of this subdi-
under the old order Monaxonida. Their definition included sponges vision the Axinellidae was allocated to the former order Axinellida
having a 'skeleton typically non-reticulate; consisting of ascending Levi, 1955, along with other groups allegedly having in common
axes of fibres from which arise subsidiary fibres radiating to the an axially compressed skeleton and an oviparous mode of repro-
surface; fibres typically plumose; megascleres chiefly styles to duction. This scheme was criticised by Van Soest et al. (1990) and
which oxeas and (or) strongyles may be added; microscleres rarely Van Soest (1991) who returned the family to the order
present, never chelas'. The differences between the definitions of Halichondrida in an emended form.
Carter and Ridley & Dendy are insignificant. Ridley & Dendy These higher taxonomic changes and the inclusion of other
(1886, 1887) did not mention Carter as author of the original genera with condensed axial skeletons, plumose or plumoreticulate
description of the Axinellidae and since then, with some exceptions extra-axial skeletons and microscleres by Hallmann (1916a,
(Wiedenmayer, 1977b; Hooper & Wiedenmayer, 1994), the author- 1916b, 1917c) and de Laubenfels (1936a) have contributed to the
ity of the family has been incorrectly attributed to these authors large number of nominal genera (92) included in the family at one
instead of Carter (1875c). time or another.
The definitions of Carter (1875c, 1883b), Ridley & Dendy The phylogenetic relationships of the family Axinellidae in
(1886, 1887), and the subsequent amendments of Topsent (1894c, relation to other demosponge families were explored by Van Soest
1904b), Dendy (1905; 1916a), Hallmann (1917c), de Laubenfels et al. (1990) using cladistic and parsimony methods based on mor-
(1950a), Levi (1973), Wiedenmayer (1977b); Bergquist (1970; phological characters. In their view, the Axinellidae belongs to
1978) and Hartman (1982) reflect the lack of diagnostic characters a redefined order Halichondrida, and is placed as the sister clade of
to define the family. Hooper & Wiedenmayer (1994: 69) were a group of taxa that includes the families Desmoxyidae,
the last authors to give a comprehensive definition of the family as Dictyonellidae and Halichondriidae. The synapomorphy proposed
follows: to define the Axinellidae within this order is the presence of an
'Encrusting, massive, branching, fan shaped, and tubular; axially condensed and extra-axial plumoreticulate choanosomal
surface usually hispid due to projecting spicules; megascleres are skeleton. However, some genera, such as Dragmacidon (i.e.,
styles, oxeas, strongyles (sometimes sinuous) in all combinations, Pseudaxinella of contemporary authors; see below) and Ptilocaulis
usually smooth, sometimes tuberculate, spined, flexuous or vermi- have (secondarily?) lost the axial skeleton, while others like
form; skeleton typically divided into axial and extra-axial compo- Auletta and Phakellia lack a clear axial component, which renders
nents; main skeletal tracts, both spicules and spongin fibres, the strict definition untenable. Although axial condensation of
typically condensed in axial portion of skeleton, with extra-axial the skeleton has been traditionally used to link taxa within the
plumose or plumoreticulate tracts running to surface; encrusting Axinellidae, it is obviously a very weak synapomorphy for the
species having condensed basal skeleton usually with basal flexu- family, as it is also present in other unrelated families such as
ous or vermiform strongyles, and perpendicular extra-axial skele- Raspailiidae (e.g., Ceratopsion), Hemiasterellidae (e.g., Axos),
ton of styles (straight or rhabdose), or oxeas. Microscleres usually Desmacidinae (e.g., Desmacidon) and some Microcionidae
absent, although few genera having raphides or microraphides in (e.g., Clathria (Axosuberites)) Suberitidae (e.g., Homaxinella,
groups (trichodragmata) or alone'. Plicatellopsis, Rhizaxinella).
This definition agrees with previous ones and is used in this The phylogenetic relationships of some members of the fam-
revision as an initial criterion to select the valid genera of ily Axinellidae has been studied by Alvarez & Crisp (1994) using
Axinellidae. Hooper & Levi (1993b) and Hooper & Wiedenmayer a group ofaxinellid species from the Central-West Atlantic, and
(1994) recognised the poor value of the characters included in this more recently by Alvarez et al. (2000a). In the latter study, a group
definition, the occurrence of many exceptions to the general pattern of 27 species ofAxinellidae and related families (Dictyonellidae,
and the poor understanding of the affinities of many of the included Halichondriidae and Agelasidae) were selected with the aim of
species. testing the monophyly ofAxinellidae and to investigate their phy-
The paucity of reliable diagnostic characters creates a prob- logenetic relationships using morphological and molecular data.
lem at the infra-family level. Many of the nominal genera assigned The results of this work based on molecular data and some chemi-
to the Axinellidae have skeletal structures different to the axinellids cal evidence, showed that the family might not be monophyletic;
in the sense of Hooper & Wiedenmayer (1994). Furthermore, most but the analysis based on morphological data indicated however,
726 Porifera' Demospongiae • Halichondrida • Axinellidae

that the family is monophyletic. Therefore, at this stage there is not (e.g., Acanthella, Stylissa and Rhaphoxya) have skeletal arrange-
sufficient conclusive evidence to establish the monophyly of the ment similar to some genera ofAxinellidae (e.g., Axinella) but the
Axinellidae or to clearly define the phylogenetic position of the surface features in those cases are very characteristic of
family in the higher classification. Dictyonellidae (see chapter in this volume).
A great number of novel chemical compounds have been iso- The definition of the family is also emended from Hooper &
lated from members of this family (e.g., Rudi et aI., 1997; Mancini Wiedenmayer (1994) to exclude encrusting forms (i.e., Bubaris,
et al., 1997; Li et al., 1998b; Urban et aI., 1999) some with bioac- Bubaropsis, Cerbaris, Hymerhabdia, Lithobubaris, MOllocrepidium,
tive properties (e.g., Newbold et al., 1999) and others that function Rhabdobaris, Rhabdoploca, Skeizia and Uplexoa). These were
as chemical defence (e.g., Wilson et aI., 1999). The use of these included in Axinellidae at one time or another based on the similar-
compounds as chemotaxonomic markers has also been discussed ities between the axial condensation of the skeleton ofAxinellidae
(Van Soest & Braekman, 1999). and the 'basally condensed' skeleton of encrusting forms such as
Some members of the family Axinellidae are oviparous Bubaris (see Bergquist, 1970; Van Soest et al., 1990). They are
(Siribelli, 1962), but it is true to say that the reproductive biology transferred here to Bubaridae sensu Topsent, 1928c and
of the family remains poorly known, and the importance of this Desmanthidae (see respective chapters in this volume).
character in defining higher taxa is largely speculative - despite
extensive interpretation of these few data as 'rules' for the family Previous reviews
and for the nominal order 'Axinellida' in general (e.g., Levi, 1973;
Bergquist, 1978; Hartman, 1982). Carter, 1883b: 316; Ridley, 1884a: 462; Carter, 1885d: 347;
Ridley & Dendy, 1886: 477; Ridley & Dendy, 1887: 166; Topsent,
Remarks 1894c: 24--25; Dendy, 1897: 231; Wilson, 1902: 399; Topsent,
1904b: 137; Dendy, 1905: 181; Row, 1911: 354; Hentschel, 1912:
The definition given above restricts Axinellidae to genera 113; Dendy, 1916a: 96; Hallmann, 1917c: 673-674; Dendy, 1922b:
without specialised ectosomal skeleton and velvety or microhispid 111; Wilson, 1925: 444; de Laubenfels, 1936a: 127; de Laubenfels,
surface; with choanosomal skeleton predominantly plumose to 1950a: 87; Bergquist, 1970; Levi, 1973: 605; Wiedenmayer,
plumoreticulate, mainly formed by spiCUle tracts of sinuous 1977b: 151-152; Bergquist, 1978: 167; Hartman, 1982; Tanita &
strongyles, oxeas, anisoxeas or styles and without microscleres Hoshino, 1989: 84; Hooper & Levi, 1993b: 1396; Hooper &
other than raphides and trichodragmata. Some dictyoneIlid genera Wiedenmayer, 1994: 69; Alvarez et al., 2000a.

KEY TO GENERA

(1) Choanosomal skeleton with ascending and anastomosing tracts radiating to periphery; spicule tracts are coring spongin
fibres, and/or fibrofascicles .................................................................................................................................................................. 2
Choanosomal skeleton with spicule tracts of sinuous strongyles plumo-echinated (radiating obliquely and perpendicularly)
by styles or oxeas ................................................................................................................................................................................. 4
Choanosomal skeleton plumoreticulate ............................................................................................................................................... 5
Choanosomal skeleton consisting of plumose axes of spicules, dendritic ally branching and rarely interconnected, with
peripheral spicules curving outward. Megascleres are only styles, some bent near the base. Microscleres are long
sinuous trichodragmata and single raphides ornamented with fine spines at terminal points ...... .... ...... ...... ...................... Dragmaxia
(2) Flat spatula-shaped scopiform processes at surface ............................................................................................................................ 3
Long and filamentous processes at surface ....... ... ... .... ................ ........... ........... ................. .... ........... ................... ........ ...... ... Phycopsis
(3) Megascleres are anisoxeas with or without spined tips ................................................................................................... Reniochalina
Megascleres are styles in two size categories ...................................................................................................................... Ptilocaulis
(4) Body generally fan-shaped and stalked; with ascending choanosomal tracts of strongyles interconnected and
plumo-echinated by single or short tracts of styles. Main tracts often imprinting the surface as 'veins' .............................. Phakellia
Body generally tube-shaped and stalked; with a layer of sinuous strongyles lining the inner wall of the tube and
a reticulation, nearly isotropic of choanosomal tracts of strongyles plumo-echinated by styles. Surface free of
skeletal projections but with single spicules ............................................................................................................................... Auletta
Body generally dichotomously branching and stalked, with a main axial core of interwoven strongyles echinated
by single oxeas or short plumose tracts of oxeas ......... ....... ... ...... ........ .............. .... .... ...... ................. ................. .......... Pararhaphoxya
(5) With styles and oxeas. Microscleres, if present, are trichodragmata ................................................................................................... 6
Body generally cup-shaped and stalked; no styles ............................................................................................................. Cymbastela
(6) Massive or massive encrusting forms; plumose tracts are generally thick and multispicular ......................................... Dragmacidon
Body branching, arborescent or bushy; plumose tracts are pauci-multispicular, connected at regular intervals by
uni-paucispicular secondary tracts ........................................................................................................................................... Axinella
Porifera' Demospongiae • Halichondrida • Axinellidae 727

AULETTA SCHMIDT, 1870 Synonymy. Auletta sycinularia Schmidt, 1870: 45; Ridley &
Dendy, 1887: 187; Topsent, 1904b: 143; Van Soest & Stentoft, 1988:
Synonymy 105; Alvarez et al., 1998: 38.
Material examined. Lectotype (here designated): BMNH
Auletta Schmidt, 1870: 45. 1939.2.10.36 - Alligator Reef, Florida, 210m. (based on data pub-
lished by Desqueyroux-Faundez & Stone, 1992: 25).
Type species Description. Cluster of 6 tubes, 1-3 cm long approximately;
on thin stalk approximately 1 cm long; surface hispid; oscules api-
Auletta sycinularia Schmidt, 1870: 45 (by monotypy). cal, with sphincters; pores, 60-80 mm in diameter, within atrial
cavity of tubes; colour unknown when alive, beige in alcohol;
Definition consistency fragile and soft (Fig. IA). Specialised ectosomal skele-
ton absent; choanosomal skeleton with thick and longitudinal
Axinellidae of tubular habit with a layer of sinuous strongyles strongyle tracts, 50-180 !-lm, plumo-echinated or connected at
lining the inner wall of the tube and a reticulation, nearly right angles by single or plumose tracts of two size categories of
isotropic of choanosomal tracts of strongyles plumo-echinated by styles. Inner tube wall reinforced by masses of sinuous strongyles
styles. (Figs lB-D, 2A). Sinuous strongyles and strongyloxeas, length
300-770!-lm (465.8 ± 133.8), width 13-25!-lm (18.4 ± 3.1)
Diagnosis (n = 24); styles of two size categories: (a) length 450-950!-lm
(657.9 ± 136.6), width 10-20!-lm (15.7 ± 3.2) (n = 13), (b) length
Tubular, erect on peduncle or narrow base. Surface smooth 140-280!-lm (206.25 ± 57.6), width 8-1O!-lm (9.5 ± 1) (n = 4)
with choanosomal spicules projecting slightly; ectosome without (Fig. 2B). Additional size category of styles found in specimens
specialised skeleton. Choanosomal skeleton plumoreticulate, with from Gulf of Mexico (Alvarez et al., 1998).
longitudinally strongyle tracts, connected by single styles or Remarks. This species seems to be variable in its shape, with
plumose tracts of styles; masses of sinuous strongyles reinforcing some specimens having the tubes fused, resulting in a flabelliform
inner tube wall. Megascleres sinuous strongyles, always coring shape (Alvarez et al., 1998). Auletta sycinularia is similar to
main spicule tracts and inner tube walls; styles and or oxeas, A. sessilis Topsent, 1904b, a species recorded from the Azores but
plumo-echinating and connecting main tracts. Microscleres absent. with a slightly different habit in which the tubes are fused to
a broad base. Auletta tube rosa Alvarez et al., 1998, recorded from
Remarks the Gulf of Mexico, differs from A. sycinularia in shape, surface
features and in having oxeas, in addition to styles and strongyles.
The organisation of the skeleton of Auletta is comparable to that Habitat and distribution. This species has been found in the
of Phakellia and Pararhaphoxya. In Phakellia the primary skeleton Gulf of Mexico, Florida and Barbados at depths of 70-200 m;
is reticulated, with axes of sinuous strongyles plumo-echinated by substrate unknown.
styles, single or in bundles connected by secondary spicule tracts,
generally styles; in Pararhaphoxya styles have been replaced by
oxeas. The three genera have distinctive habit and surface features. AX/NELLA SCHMIDT, 1862
Auletta has been referred previously to Cornacuspongidae:
Halichondrina: Halichondridae (Vosmaer, 1885b), Monaxonida: Synonymy
Halichondrina: Axinellidae (Topsent, 1894c; 1904b: 137; de
Laubenfels, 1936a), Axinellida: Axinellidae (Levi, 1973) and Axinella Schmidt, 1862: 60. Axidragma Hallmann, 1917c: 637
Halichondrida: Axinellidae (Van Soest et al., 1990: 18; Alvarez (new synonym). Axinosia Hallmann, 1914: 349 (new synonym).
et al., 1998). Chalinissa Lendenfeld, 1887c: 771. Pseudaxinella Schmidt, 1875:
120. (new synonym; Not Pseudaxinella Schmidt sensu de
Distribution Laubenfels, 1936a: 131; Bergquist, 1970: 20; Levi, 1973;
Wiedenmayer, 1977b: 152,155; Wiedenmayer, 1989: 47; Hooper &
A total of 17 species of Auletta are recorded in the literature Levi, 1993b: 1436; Alvarez et al., 1998 =Dragmacidon, see
from Central-West Atlantic, East Atlantic, Indian Ocean, NE below). Querciclona de Laubenfels, 1936a: 46. (new synonym).
Australia, Central-West Pacific and Japan. Teichaxinella de Laubenfels, 1936a: 128 Tragosia Gray, 1867a:
513. [Astrospongia] Gray, 1867a: 514 (objective synonym).
Previous reviews
Type species
Topsent (1894c: 25); Dendy (1889: 93); Topsent (1904b: 137,
143); Dendy (1905: 194); (Dendy 1916a: 119); Dendy (1922b: Axinella polypoides, Schmidt, 1862: 60 (by subsequent desig-
121); Burton (1931c: 142); de Laubenfels (1936a: 129); nation; de Laubenfels, 1936a).
de Laubenfels (1954: 170); Levi & Vacelet (1958: 233); Levi
(1973: 605); Van Soest & Stentoft (1988: 105); Alvarez et al. Definition
(1998: 32).
Axinellidae with choanosomal skeleton differentiated in
Description of type species axial (compressed or vaguely reticulated) and extra-axial
(plumoreticulated) region. Megascleres are styles and oxeas.
Auletta sycinularia Schmidt, 1870 (Figs 1-2). Microscleres, if present, are microraphides and trichodragmata.
728 Porifera· Demospongiae • Halichondrida • Axinellidae

Fig. 1. Auletta sycinularia Schmidt. A, lectotype, BMNH 1939.2.10.36. B, longitudinal section through the skeleton. C, SEM of a longitudinal skeletal
section. D, SEM of a transverse section through the skeleton (scales A, 1 cm; B, 200 /Lm).

Diagnosis recognised as typical for the genus (i.e., branching habit, skeleton
axially condensed and extra-axial skeleton with a reticulation of
Ramose, bushy or lamellate habit. Surface generally smooth plumose tracts, styles and oxeas). However, many other species
with choanosomal spicules projecting slightly; oscules, when visi- presently included in Axinella lack some of these features;
ble, with stellate morphology (i.e., superficial canals leading to additionally, some of the characters can be variable at the intra-
opening 'imprinted' in superficial skeleton); ectosome without spe- specific level (see Pansini, 1982-1983; Alvarez et ai., 1998). The
cialised skeleton. Choanosomal skeleton differentiated in axial genus currently includes a heterogeneous assemblage of species,
and extra-axial regions; axial skeleton compressed or vaguely some of which might be better allocated to Acanthella or Phakellia
reticulated; extra-axial skeleton plumose or plumoreticulate. based on the revised diagnoses provided in this work. A revision of
Megascleres styles, or styles and oxeas, or oxeas; when both pres- the nominal species ofAxinella, Phakellia and Acanthella is recom-
ent, one type may be rare; modifications of megascleres common mended here, probably best undertaken on a regional basis due the
in several species. Microscleres, if present, microraphides and large number of species currently assigned to each of these genera.
raphides, mostly in tightly packed trichodragmata. Axidragma (type species Axinella padina Topsent, 1896a:
431) is considered here to be a junior synonym of Axin ella, as sug-
Remarks gested by Cabioch (1968b). It was erected in the Axinellidae by
Hallmann (1917c) to include Axinella padina, a species considered
Axinelia is difficult to define due to the great morphological plas- to be a junior synonym of Axinella jlustra by Topsent (1904b).
ticity exhibited by species. The type species has features generally According to Topsent (1904b), A. padina differs from A. jlustra
Porifera· Demospongiae • Halichondrida • Axinellidae 729

Fig. 2. Auletta sycinularia Schmidt, BMNH 1939.2.10.36. A, diagram of the skeleton. B, style. C, sinuous strongyle (scales A, 200j.Lm; B-C, 100 j.Lm).

only in the thickness of the oxeas which seem to be smaller in Chalinissa (type species Isodictya dissimilis Bowerbank,
A. padina, a feature that could vary at intraspecific level. 1866: 318) was erected in the Homorrhaphidae: Chalininae
Axidragma in the sense of Hallmann shared the presence of tri- (Lendenfeld, 1889a) and agrees with Axinella (Burton, 1927: 502).
chodragamata with Dragmaxia Hallmann and Dragmacidon Other species described under Chalinissa (i.e., C. communis
Hallmann, but is different in terms of skeletal architecture. Neither Lendenfeld, 1887c: 772; C. communis var. flabellum Lendenfeld,
the type material of Axin ella padina or A. flustra could be located 1887c: 772; C. elegans Lendenfeld, 1887c: 773; C. elongata
but, according to the descriptions given by Topsent (1892a: 122; Lendenfeld, 1887c: 774; C. macropora Lendenfeld, 1887c: 772;
1904b: 139; 1928c: 177) and Cabioch (1968b), and based on the C. ramosa Lendenfeld, 1887c: 774; C. rigida Lendenfeld, 1887c:
examination of a specimen of A. flustra from Banyuls (see below) 773; c. serpens Lendenfeld, 1887c: 773; C. tenuifibris Lendenfeld,
and the published descriptions, the species can be confidently 1887c: 773) are currently allocated to Callyspongia serpentina
placed in its original genus. Dendy (1922a) and de Laubenfels (Lamarck) (Hooper & Wiedenmayer 1994).
(1936a: 132) regarded Axidragma as a junior synonym of Tragosia, Pseudaxinella (type species Pseudaxinella sulcata Schmidt,
but this genus is considered also a synonym of Axinella in this 1875: 120 by monotypy) is included here as a junior synonym of
work. Axinella based on recent examination of its type material (ZMUC
Axinosia (type species Axinella symbiotica White1egge, 1907: DEM-241, lectotype, here designated; ZMUC DEM-242; ZMUC
508) was erected in the Axinellidae by Hallmann (1914: 269) DEM-243, ZMUC DEM-244, paralectotypes, Fig. 5A, wet, SW
to include a group of species that he considered congeneric vom Bukenfjord, 106 fathoms). The genus was erected by Schmidt
(i.e., Axinella symbiotica, Stylotella irregularis Kirkpatrick, 1900a, (1875) for Pseudaxinella sulcata, from the North Sea to allocate
Phakellia multiformis Whitelegge, 1907: 508 and A. arborescens those Axinella-like species without a clear compression of the
Ridley & Dendy, 1886). It was related to Teichaxinella by axial skeleton, some of which were included in Dictyocylindrus
Wiedenmayer (1989: 48-49) and to Reniochalina and Ptilocaulis by and Isodictya by Bowerbank (1866) but are currently allocated
Hooper & Levi (1993b: 1404). The genus is considered here ajunior to Axinella, e.g., A. dissimilis (Bowerbank, 1866: 318) and
synonym of Axin ella based on the re-examination of the type material A. infundibuliformis (Linnaeus, 1791: 1296). Since de Laubenfels
of A. symbiotica, which agrees with the definition of Axinella given (1936a) however, the genus has been reserved for species of mas-
above. Other species reported under Axinosia are: A. incrustans sive, subspherical, cushion-shaped, unbranched or lobate growth
Burton, 1930e: 333 from the South Pacific and A. xutha Laubenfels, form, with a skeleton typically plumoreticulate and without axial
1954: 175 from the Central-West Pacific. The identity of these compression or differentiation between axial and extra-axial
species, and those transferred to Axinosia by Hallmann (1914) needs regions. Although the concept of Pseudaxinella by contemporary
to be confirmed as they might belong to genera other than Axinella. authors (Bergquist, 1970: 20; Levi, 1973; Wiedenmayer, 1977b:
730 Porifera· Demospongiae • Halichondrida • Axinellidae

152, 155; Wiedenmayer,1989: 47; Hooper & Levi, 1993b: 1436; Halichondrina: Halichondridae (Vosmaer, 1885b); Monaxonida:
Alvarez et aI., 1998) has been unequivocal, it does not match either Halichondrina: Axinellidae (Ridley & Dendy, 1886; Topsent,
the description of Schmidt, or the redescription of Thiele (1903b: 1894c; 1896a; 1904b; de Laubenfels, 1936a), Sigmatomonaxonida:
378), or the observations made from the lectotype of Pseudaxinella Axinellidae (Dendy, 1905; Topsent, 1928c); Axinellida: Axinellidae
sulcata which agrees with Axinella. Species assigned to (Bergquist, 1970; Levi, 1973) and Halichondrida: Axinellidae
Pseudaxinella in the past are better placed under Dragmacidon (Van Soest et al., 1990; Alvarez et aI., 1998).
Hallmann, 1917c (see below).
Querciclona (type species Antherochalina quercifolia Keller, Distribution
1889: 338, by original designation) was synonymised with
Phakellia by Hooper & Levi (1993b: 1413). Hooper (1996a) The genus has a widespread distribution (Hooper &
redescribed the type material (BMNH 08.9.24.179) and referred it Wiedenmayer, 1994; Van Soest, 1994) and approximately 60 valid
to Phakellia; he further suggested thatA. quercifolia might be a syn- species (Alvarez, 1998).
onym of Phakellia donnani (Bowerbank, 1873c as [sodictya). The
type specimen of A. quercifolia was re-examined here and nothing Previous reviews
additional needs to be added to the description of Hooper (1996a).
However, the material, is considered to be in agreement with Gray (1867a: 513); Ridley & Dendy (1886: 479; 1887: 178);
Axinella and not with Phakellia. It is very similar to other species Topsent (1894c); Lendenfeld (1888: 234); Dendy (1897: 231);
of Axinella such as A. symbiotica Whiteiegge, 1907: 508 from Wilson (1902: 400); Topsent (1904b: 137); Dendy (1905: 188);
Australia, A. waltonsmithi (de Laubenfels, 1953a: 533) from Vosmaer (1912: 308); George & Wilson (1919: 158); Dendy
the Central-West Atlantic, and A. damicomis sensu Pansini, (1922b: 114); Topsent (1928c: 173); Vosmaer (1932-35: 721); de
1982-1983, from the Mediterranean. The oxeas in these species, Laubenfels (1936a: 130); Levi & Vacelet (1958: 233); Vacelet
including the type species of Querciclona, are rare or absent; the (1969: 177); Levi (1973: 605); Bergquist (1970: 14); Donadey et al.
choanosomal skeleton is clearly plumoreticulate, with paucispicular (1990: 259); Hooper & Levi (1993b: 1407); Alvarez et al. (1998).
primary tracts ending in brushes that protrude through the surface,
and connected by uni-paucispicular secondary tracts generally one Description of type species
spicule long. The skeleton is slightly compressed in the axial region.
Teichaxinella (type species Teichaxinella shoemakeri de Axinella polypoides Schmidt, 1862 (Figs 3-4).
Laubenfels, 1936a: 129, by original designation) was established for Synonymy. Axinella polypoides Schmidt, 1862: 60; Gray,
some species of Axinella that lack axial condensation because of 1867a: 514; Schmidt, 1868: 9; (not Schmidt, 1870: 60 = Axinella
their external morphology (de Laubenfels, 1936a: 128-129; polycapella de Laubenfels, 1953a: 530); Carter, 1884b: 205;
Wiedenmayer, 1977b: 154). Detailed examination of several speci- Topsent, 1902: 348; Babic, 1922: 238; Topsent, 1928c: 173;
mens of the type species, including the holotype (see Alvarez et al., Vosmaer, 1932-35: 723; Topsent, 1934a: 34; Sara, 1960a: 32;
1998) showed that the extra-axial skeleton consists of single spicules Vacelet, 1969: 175; Boury-Esnault, 1971b: 303; Pulitzer-Finali,
and short plumose spicule tracts and therefore should be included 1983: 516; Uriz, 1982: 59 (with additional records). Tragosia
in Axinella. As in other species of Axinella, e.g., A. polypoides polypoides; Hanitsch, 1894a: 179.
Schmidt, 1862: 60, A. dissimilis (Bowerbank, 1866: 318) and Material examined. Lectotype (here designated): BMNH
A. infundibuliformis (Linnaeus, 1791: 1296), the type species of 1867.7.26.81, dry - Schmidt's specimen from type locality,
Teichaxinella has also trichodragmata which supports further Adriatic. Paralectotype (here designated): BMNH 1929.10.11.1
synonymy with Axinella. Other species reported under Teichaxinella and BMNH 1877.5.2.335 (registered twice according to the
are: Phakellia jlabellata Carter, 1885d: 363 and Axinella vermiculata BMNH records) - fragment of Schmidt's specimen from type
Whitelegge, 1907: 511 (Hooper & Wiedenmayer, 1994: 85), locality.
Axinella labyrinthica Dendy, 1889: 88 (Hooper & Levi, 1993b: Description. Branching-ramose, 30 cm high; dichotomous
1410) and T. morchella Wiedenmayer, 1977b: 154 (junior synonym branches, 1-2 cm in diameter; surface smooth-velvety when moist;
of Axinella corrugata (George & Wilson, 1919: 161». The generic oscules areolate star-shape, 0.5-5 mm diameter approximately,
placement of these species needs to be re-examined. uniformly distributed; colour orange or yellow-orange in life, dark-
Tragosia (type species Spongia infundibuliformis Linnaeus, brown in dry state; consistency firm but flexible (description of
1791: 1296, by original designation) (Fig. 5B) was revised by external features taken from Uriz, 1982) (Fig. 3A). Specialised
Dendy (1922a) and differentiated from Axinella based on the pres- ectosomal skeleton absent; choanosomal skeleton differentiated in
ence of trichodragmata. This character is also present in other axial and extra-axial regions; extra-axial skeleton plumoreticulate
species of Axinella including the type species (i.e., A. dissimilis, with primary tracts, plumose, pauci-, pluri- or multi-spicular,
A. jlustra, A. polypoides. A. shoemakeri) and also in other genera of 20-100 f.Lm thick, radiating from the axis towards the surface,
Axinellidae (e.g., Dragmacidon Hallmann, 1917c, Dragmaxia ending in single spicules or brushes projecting slightly through
Hallmann, 1916b) and is considered here of little value to differen- ectosome, connected at more-or-less regular intervals by secondary
tiate the genus. Cabioch (l968b) considered Tragosia as a sub- tracts of 1-3 spicules, generally no more than one spicule long;
genus ofAxinella and included A. dissimilis and A. jlustra. Some axial skeleton condensed to reticulate, present both in central col-
species currently assigned to Tragosia such as, T. alba Descatoire, umn of specimens and in branches (Figs 3C-D, 4A). Mega-
1969: 188, T. fibrosa Topsent, 1890b: 65, T. hirondellei Topsent, scleres are styles, length 210-500f.Lm (345::!:: 118.6), width 8-12
1892a: 120, should be revised as they might not belong in Axinella. f.Lm (9.4::!:: 1.3) (n = 10) and oxeas, length 270-420 f.Lm (324::!::
Axinella has been referred previously to Halichondriae 45.3) (n = 10) 5-12 f.Lm (8.7::!:: 1.9) (n = 10) (Fig. 4B-C).
(Schmidt, 1862), Keratospongia: Chalinidae (Gray, 1867a), Microscleres are trichodragmata, up to 30 f.Lm in length, difficult to
Echinonema: Axinellida (Carter, 1885d), Cornucuspongidae: observe (Fig. 3B).
Porifera' Demospongiae • Halichondrida • Axinellidae 731

Fig. 3. Axinella poJypoides Schmidt, 1862. A, lectotype, BMNH 1867.7.26.81. B, trichodragmata. C, SEM of the choanosomal skeleton. D, detail of the
choanosomal skeleton (scales A, 10 cm; B, 10 /-Lm; C, 1 mm; D, 500 /-Lm).

Remarks. Axinella polypoides is very similar in general associated to soft or hard substrates, protected vertical walls or
morphology, skeletal architecture and spicule dimensions to shallow and dark habitats (Uriz, 1982).
A. polycapella de Laubenfels, 1953a, from the Central-West
Atlantic and A. australiensis Bergquist, 1970, from New Zealand.
In the material examined trichodragmata are not very abundant, CYMBASTELA HOOPER & BERGQUIST, 1992
difficult to detect and generally located near the surface; they do
not appear to be homologous to the microraphides detected by Synonymy
Donadey et al. (1990). Axinella polypoides shares the presence of
trichodragmata with other species of AxineUa, such as A. flustra Cymbastela Hooper & Bergquist, 1992: 103.
Topsent, 1892a, A. dissimilis (Bowerbank, 1866), A. infundibuliformis
(Linnaeus, 1791) and A. shoemakeri (de Laubenfels, 1936a). Type species
Habitat and distribution. The species is restricted to
the Mediterranean and East Atlantic. It is found from shallow water Pseudaxinyssa stipitata Bergquist & Tizard, 1967: 189 (by
to 367 m depth but is generally abundant at depths of 30-40 m original designation).
732 Porifera' Demospongiae • Halichondrida • Axinellidae

Fig. 4. A.tillel/a {Jo/Y{Joities chmidl. 1862. A. diagram f lhe h an omal kelelon. B. ly le. C. oxea ( calc 50 I.Lln).

Fig. S. A, Axinella sulcata (Schmidt, 1875), lectotype, ZMUC DEM-24l (indicated with an arrow) and paralectotypes, ZMUC DEM-242-244.
B. Axillella infundibuliformis (Linneaeus, 1791), BMNH 1930.7.3.370 (scales A, 1 em; B, 2.5 em).

Definition (1992). One of its species however, C. coralliophilla Hooper &

Bergquist, possesses a specialised ectosomal skeleton which is
Axinellidae with choanosomal skeleton plumoreticulated and not typical of other Axinellidae. Phylogenetic analysis using partial
made exclusively of oxeas. 28S rDNA sequences also showed that C. coralliophilla is not
closely related to other Cymbastela species, such as C. stipitata and
Diagnosis C. vespertina (Alvarez et al., 2000a). Future studies are required
to confirm the generic position of C. coralliophila. Cymbastela
Cup-shaped, thinly lamellate or lamellate-flattened-encrusting hooperi Van Soest et al., 1996b reported from the Great Barrier
habit. Surface smooth with choanosomal or ectosomal spicules pro- Reef, is distinguished from other species of Cymbastela by its
jecting slightly; with or without specialised skeleton of smaller growth form, which is stalkless and flattened on the substratum,
oxeas. Choanosomal skeleton generally differentiated in axial and yellow coloured with a loose skeletal morphology of vaguely
extra-axial regions; axial skeleton condensed or vaguely reticulated; plumose and wavy ascending tracts (Van Soest et al., 1996b).
extra-axial skeleton plumoreticulate with plumose spicule tracts Amphimedon terpenensis Fromont, 1993 is now considered a
enveloped with thin sheets of spongin or well formed spongin species of Cymbastela based on skeletal and chemical (i.e., presence
fibres. Megascieres are oxeas often with telescoped points of di-isocyano-adociane) similarities with C. hooperi (Van Soest
(emended from Hooper & Bergquist, 1992; Van Soest et al. , 1996b). et at. , 1996b).
Other species of Cymbastela are: C. cantharella (Levi, 1983)
Remarks (as Pseudaxinyssa) (Hooper & Levi, 1993b: 1397); C. concentrica
(Lendenfeld, 1887c) (as Antherochalina); C. coralliophila Hooper &
Cymbastela is considered here to have close affinities with Bergquist, 1992: 120; c. hooperi Van Soest et at., 1996b: 104;
species of Axinella having similar growth forms (i.e., flabellate- C. marshae Hooper & Bergquist, 1992: 127; C. notiaina Hooper &
lamellate) and skeleton as suggested by Hooper & Bergquist Bergquist, 1992: 128; C. vespertina Hooper & Bergquist, 1992: 110.
Porifera· Demospongiae • Halichondrida • Axinellidae 733

Distribution Material examined. Holotype: AM Z3101, wet - Dudley

Point Reef, Darwin, Northern Territory, colI. E. Pope & J. Boase,
The genus has been reported from Australia (Hooper & 13. x.1965.
Bergquist, 1992; Hooper & Wiedenmayer, 1994) and New Description. Cup-shaped, approximately 8 cm wide and
Caledonia (Hooper & Levi, 1993b) but is also present in New 7 cm tall; on stalk 2 cm in diameter; surface smooth with rounded
Zealand (Alvarez, unpublished data). and digitate projections, microscopically hispid due to projection
of spicules; pores 1-1.8 mm in diameter, evenly distributed; colour
Description of type species green-brown; consistency firm but flexible (Fig. 6A). Specialised
ectosomal skeleton absent. Choanosomal skeleton plumoreticulate
Cymbastela stipitata (Bergquist & Tizard, 1967) (Figs 6-7). with plumose primary tracts, 50--150 fLm thick, ending in brushes
Synonymy. Pseudaxinyssa stipitata Bergquist & Tizard, 1967: at surface; secondary tracts (no more than one spicule long), often
189-191. Cymbastela stipitata; Hooper & Bergquist, 1992: 106. paucispicular but some multispicular, with spicules stacked, con-
necting primary tracts at more or less regular intervals. (Figs
6B-D, 7A). Spicules are oxeas, length 250--320 fLm (290::':: 22.0),
width 5-20 fLm (13.4 ::':: 3.9) (n = 10) (Fig. 7B-D).
Remarks. Cymbastela stipitata has been extensively
redescribed and compared to other species of Cymbastela by
Hooper and Bergquist (1992). Its closest relative is C. vespertina
Hooper & Bergquist, 1992, another northwestern Australian species,
that differs from C. stipitata in habitat, colour and in the thickness of
the lamella (Hooper & Bergquist, 1992). Phylogenetic analyses based
on molecular and morphological data, also point out to the closest
relationships between these two species (Alvarez et al., 2000a).

B o

Fig. 6. Cymbastela stipitata (Bergquist, 1967). A, holotype, AM Z3101.

B, SEM of the skeleton. C, longitudinal section through the periphery. Fig. 7. Cymbastela stipitata (Bergquist, 1967). A, diagram of the
D, detail of spicule tracts (scales A, 2 cm; B, 500 fLm; C, 200 fLm; D, 100 fLm). choanosomal skeleton. B, oxeas (scales A, 200 fLm; B-D, 50 fLm).
734 Porifera· Demospongiae • Halichondrida • Axinellidae

Habitat and distribution. The species is restricted to north- The closest genera to Dragmacidon are Axinella and
ern Australia. It is found from shallow waters to 19 m depth; mostly Cymbastela. They all share a plumoreticulate choanosomal skele-
associated to hard substrates and turbid waters (Hooper & ton of ascending plumose tracts but Dragmacidon differs from
Bergquist, 1992). them in growth form, surface features and in the absence of an
axial skeleton. Dragmacidon also shares with Axinella the spiCUle
composition (i.e., oxeas, styles and trichodragmata in similar size
categories). These three genera are likely to constitute a mono-
DRAGMACIDON HALLMANN, 1917 phyletic group that could be classified as a subfamily within
Axinellidae. Phylogenetic analyses based on morphological
Synonymy characters support this view (Alvarez, 1998).

Dragmacidon Hallmann, 1917c: 639. Pseudaxinella sensu de Distribution

Laubenfels, 1936a: 131; Bergquist, 1970: 20; Levi, 1973;
Wiedenmayer, 1977b: 152,155; Wiedenmayer, 1989: 47; Hooper & The genus has a widespread distribution according to Van Soest
Levi, 1993b: 1436; Alvarez et al., 1998; Not Pseudaxinella (1994) and contains approximately 17 species, including those
Schmidt, 1875: 120; Thiele, 1903b: 378 (=Axinella). assigned to Pseudaxinella by previous authors (Alvarez, 1998).

Type species

Thrinacophora agariciformis Dendy, 1905: 186 (by original

designation).

Definition

Axinellidae with a plumoreticulate choanosomal skeleton and

undifferentiated axial and extra-axial regions. Megascleres are oxeas
and styles. Microscleres if present, are trichodragmata.

Diagnosis

Unbranched, club-shaped, lobate, shrub-like, thickly encrust-

ing or massive habit. Surface more or less smooth with short
conules or tubercles; oscules circular, flush or slightly elevated,
sometimes with superficial canals leading to opening. Ectosome
without specialised skeleton. Choanosomal skeleton plumoreticu-
late with ascending plumose tracts, anastomosing or interconnected
by secondary multispicular tracts; not differentiated into axial or
extra-axial regions. Megascleres oxeas and/or styles generally in
similar proportions and dimensions. Microscleres if present,
raphides in tightly packed trichodragmata.

Remarks

Dragmacidon was erected by Hallmann (1917c) to include

two species of Thrinacophora that lack the special dermal megas-
cleres (i.e., without a typical raspailiid ectosome). The genus was
reserved for more or less massive species with skeleton of plumose
columns of styles and oxeas, and with raphides, single or in tri-
chodragmata as the only type of microscleres. Other species
included in Dragmacidon by Hallmann (1917c) were T. durissima
Dendy, 1905, Halichondria clathriformis Lendenfeld, 1888 and
Ciocalypta incrustans Whitelegge, 1897. Hallmann's diagnosis of
Dragmacidon corresponds to the concept of Pseudaxinella by
recent authors (Bergquist, 1970: 20; Levi, 1973; Wiedenmayer,
1977b: 152, 155; Wiedenmayer, 1989: 47; Hooper & Levi, 1993b:
1436; Alvarez et al., 1998). Given that Pseudaxinella s.s is ajunior
synonym of Axinella (see above under remarks of Axinella),
Dragmacidon becomes a suitable genus to locate those species that Fig. 8. Dragmacidon agariciformis (Dendy). A, holotype, BMNH
have been referred to Pseudaxinella by most authors. The above 1907.2.1.74. B,longitudinal section through the periphery. C, SEM of lon-
gitudinal section through skeleton. D, trichodragmata. E, detail of spicule
diagnosis is based on the type species of Dragmacidon and the tracts. F, SEM of skeleton and trichodragmata near surface (scales A, 120mm;
descriptions of many of those Pseudaxinella spp. B; 300 fLm; E, 50 fLm).
Porifera· Demospongiae • Halichondrida • Axinellidae 735

c
B

Fig. 9. Dragmacidon. A, Dragmacidon reticulata (Ridley & Dendy, 1886), diagram of the choanosomal skeleton. B-E, Dragmacidon agariciformis
(Dendy), spicules (scales B-D, 50 fLm; E, 10 fLm).

Previous reviews Dragmacidon durissima is closely related, both genetically and

morphologically to other species of Dragmacidon previously
Topsent (1928c: 37); de Laubenfels, 1936a: 131; Bergquist, referred to Pseudaxinella, i.e., D. australis from the South Pacific
1970: 20; Levi, 1973; Wiedenmayer, 1977b: 152, 155; and North coast of Australia (Hooper & Wiedenmayer, 1994;
Wiedenmayer, 1989: 47; Hooper & Levi, 1993b: 1436; Alvarez Alvarez et al., 2000a) and D. reticulata from the Central-West
et al., 1998. Atlantic (Alvarez et al., 1998).
Habitat and distribution. The species is restricted to the
Description of type species Indian Ocean at depths of 38m (Dendy, 1922b; Burton, 1959a).

Dragmacidon agaricijormis (Dendy, 1905) (Fig. 8).

Synonymy. Dragmacidon agaricijormis Hallmann, 1917c: DRAGMAXIA HALLMANN, 1916
639; Axinella agaricijormis Burton, 1959a: 259.
Material examined. Holotype: BMNH 1907.2.1.74 - Ceylon. Synonymy
Description. Massive, cushion shaped, approximately 4
cm in diameter; on broad base; surface conulose, corrugated Dragmaxia Hallmann, 1916b: 543.
with rounded to irregular depressions, covered by a thin trans-
parent membrane; colour yellow-orange in alcohol; consistency Type species
firm. (Fig. 8A). Specialised ectosomal skeleton absent;
choanosomal skeleton plumoreticulate, with ascending plumose Spongosorites variabilis Whitelegge, 1907: 513 (by original
tracts, 60-150 J.Lm loaded with broken spicules and cemented by designation).
spongin, connected by uni-multispicular tracts, up to 100 j.Lm, in
all directions.(Figs 8B-C, E-F, 9A). Styles in two size cate- Definition
gories: (a) length 650-670 j.Lm (656.7 ± 11.6), width 5-8 j.Lm
(6 ± 1.7) (n = 3) very rare; (b) length 165-335 j.Lm (212.6 ± Axinellidae (?) with styles/rhabdostyles forming plumose
56.4), width 5-10 j.Lm (6.5 ± 2.1) (n = 11). Oxeas, with axes and pointing outwards in the periphery. Microscleres are long
telescoped tips, length 185-298 J.Lm (224.5 ± 34.9), width 3-10 sinuous trichodragmata and raphides. Terminal ends of raphides
j.Lm (7.8 ± 2.3) (n = 10). (Fig. 9B-E). Trichodragmata, length ornamented with fine spines.
20-70 J.Lm (43.3 ± 25.2), width 6-35 j.Lm (15.7 ± 16.7) (n = 3),
perhaps in two categories, generally located below the surface
(Figs 8D, F, 9E). Diagnosis
Remarks. Dragmacidon agaricijormis was referred to
Axinella by Burton (1959a) without comment and its transfer here Lamellate, flabellate, cup-shaped or massive habit; surface
to Dragmacidon was suggested by Hallmann (l917c). It is very marked with small ridges; ectosome without specialised ectoso-
similar to D. durissima (Dendy, 1905) from the same locality, dif- mal skeleton. Choanosomal skeleton composed of ascending and
fering slightly in habit and in the dimensions of the megascleres. dendritically branched axes of spicules, with peripheral individual
736 Porifera· Demospongiae • Halichondrida • AxinelIidae

spicules curving outward toward surface and rarely intercon- Material examined. Holotype: AM, missing. Other mate-
nected. Megascleres styles only, straight or bent near the base, in rial. AM GlO173 - dry, offWata Mooli, NSW, Australia, coli. R.V.
widely different sizes. Microscleres long, sinuous trichodrag- 'Thetis', stn 41. AM Z5103-Z 5109 - slides, Off Wata Mooli,
mata and single raphides ornamented by fine spines at terminal NSW, Australia.
points. Description. Flabelliform, approximately 7 cm high and
9 cm wide; on small peduncle; surface marked by a reticulum of
Remarks spicule tracts; colour brown-orange in dry state. Specialised
ectosomal skeleton absent; choanosomal skeleton with ascend-
Dragmaxia was erected by Hallmann (19l6b) in Axinellidae ing plumose tracts, 300-600 f.Lm thick, with spicules or brushes
to include Spongosorites variabilis, a species characterised by a of spicules of same kind, curved outward to surface; generally
skeleton of plumose axes of styles and by the presence of long dendritic but might anastomose or be joined by spicules of
raphides (trichites in Hallmann's terminology) that were inter- adjacent tracts (Fig. lOA-B). Subtylostyles-styles-rhabdostyles,
preted by Whitelegge (1907) as microxeas and sinuous trichodrag- length 354-709 f.Lm (549.4 ± 87.8), width 10-26 f.Lm (16.8 ±
mata. The definition of Hallmann was expanded by Alvarez et al. 4.2) (n = 25) (Fig. lOD-E), some sinuous; raphides, single and
(1998) to include D. undata Alvarez et aI., 1998: 26, a massive as trichodragmata (Fig. lOF), ornamented with extremely fine
encrusting species with the same type of long raphides and tri- spines (Fig. 11), length 60-110 f.Lm (90.8 ± 17.4).
chodragmata, but without clear spicules bundles, as the ones Remarks. Although the holotype of the species is missing,
present in the type species. The raphides, in both D. variabilis the species can be recognised from the descriptions given by
and D. undata are ornamented with extremely fine spines in arrow Whitelegge (1907) and Hallmann (19l6b) and from the exam-
shape, which seems to be a diagnostic feature for the genus ination of additional specimens collected in the same area and
(Fig. 11). expedition.
Dragmaxia was considered a synonym of Stylissa by Habitat and distribution. The species is known only from
Hooper & Levi (1993b) following de Laubenfels (1936a). According the type locality (SE coast, New South Wales, Australia). It is
to these authors, the differences between the two genera were not found at depths of 42m (Hooper & Wiedenmayer, 1994).
substantial to keep them separated; however, after examination of
the type material of Stylissa it is concluded that the two genera,
should be kept separate. The skeleton in Stylissa (see chapter on
DictyoneIiidae) is less organised, nearly halichondrioid as in other PARARHAPHOXYA BURTON, 1934
Dictyonellidae such as Dictyonella, Scopalina and Rhaphoxya
(transferred to Dictyonellidae in this work). The skeleton of Synonymy
Dragmaxia consists of dendritic axes with spicules pointing
outward toward the surface. Pararhaphoxya Burton, 1934a: 565.
Dragmaxia is provisionally kept in Axinellidae as it lacks a
clear plumose or plumoreticulate skeleton. Further, some of the Type species
megascleres are rhabdostyle-like, which resembles a feature present
in raspailiid sponges and the raphides with ornamented ends do not Pararhaphoxya tenuiramosa Burton, 1934a: 565 (by original
seem to be homologous with those present in species ofAxinella or designation; junior synonym of Sigmaxinella pulchra Br!llndsted,
Dragmacidon. 1924a).

Distribution Definition

The genus is represented only by two species with a disjunct Axinellidae with an axial core of interwoven strongyles and
distribution: Dragmaxia variabilis, from the East coast of Australia extra-axial skeleton made exclusively by oxeas.
and D. undata Alvarez et al., 1998 from the Caribbean. Tuberella
ciliata Wilson, 1925 from the Philippines, was redescribed by de Diagnosis
Laubenfels (1935a) under Dragmaxia. The descriptions of Wilson
(1925) and de Laubenfels (1935a), however, do not agree with the Stalked and branching habit. Surface smooth; oscules, when
concept of Dragmaxia given in this work. That species seems to visible, form stellate patterns on the branches; consistency
belong to the genus Aaptos (Suberitidae). cartilaginous-like when alive; ectosome membranous without spe-
cialised skeleton. Choanosomal skeleton differentiated in axial and
Previous reviews extra-axial regions; axial skeleton with thick core of interwoven
strongyles dividing dichotomously along the branches; extra-axial
Hallmann (19l6b): 543; de Laubenfels, 1936a: 130; Hooper & skeleton with oxeas, single or in plumose tracts orientated at right
Levi, (1993b): 1422. Alvarez et al. (1998). angles to the axis. Megascleres sinuous strongyles, always located
at the axial skeleton and oxeas. Microscleres absent.
Description of type species
Remarks
Dragmaxia variabilis (Hallmann, 19l6b) (Figs 10-11).
Synonymy. Spongosorites variabilis Whitelegge, 1907: Sigmaxinella pulchra was considered conspecific with
513. Dragmaxia variabilis; Hallmann, 1916b: 543. Stylissa Pararhaphoxya tenuiramosa by Bergquist (1970) based on her
variabilis; Hooper & Wiedenmayer, 1994: 84. examination of the type material. Br!llndsted's specimen was
Porifera· Demospongiae • Halichondrida • Axinellidae 737

E
o

Fig. 10. Dragmaxia variabilis (WhiteJegge, 1907). A-B, possible syntype, AM GI0173. C, diagram of the skeleton. E-D, styles. P, raphides (scales
A, 1 em; B-E, 100 j..Lm; F, 50 j..Lm).

contaminated with spicules from the species Biemna stylotata and

therefore overlooked by Burton when he erected Pararhaphoxya.
Pararhaphoxya is a typical axinellid, well defined by the fea-
tures of the extra-axial skeleton. It shares features with Axinella
(Le., branching habit, surface imprinted by stellate oscules; skele-
ton differentiated in axial and extra-axial regions, extra-axial skele-
ton plumoreticulated and oxeas) and Phakellia and Auletta (i.e.,
sinuous strongyles in the main axis).
Pararhaphoxya has been previously referred to the
Axinellidae (Burton, 1934a), Halichondrina: Axinellidae:
Axinellinae (de Laubenfels, 1936a) and Axinellida: Axinellidae
(Bergquist, 1970).

Distribution

Fig. 11. Dragmaxia. SEMs of spiny raphides. A, Dragmaxia variabilis, The genus is restricted to Australia, New Zealand and W
AM GlO173. B-C, Dragmaxia undata, USNM 39451 (scales 1 j..Lm). Pacific Ocean and currently represented only by the type species. A
738 Porifera' Demospongiae • Halichondrida • Axinellidae

second species from northern New Zealand (Battershill & Bergquist, Diagnosis
1990: 68) remains to be formally described. Other material from
Western Australia examined by one of us (BA) also belongs to this Thin-plated or fan-shaped growth form, generally on peduncle.
genus. Surface hispid, with points of large choanosomal styles projecting
through ectosome; ectosome without specialised skeleton.
Previous reviews Choanosomal skeleton plumoreticulate, with multiple axes differ-
entiated into primary and secondary tracts. Primary tracts, thick,
De Laubenfels, 1936a: 130; Bergquist, 1970: 16; Hooper & visible as 'veins' or stout lines strengthening fan; formed by sinu-
Levi, 1993b: 1413. ous strongyles or strongyloxeas; plumo-echinated by styles and
connected by secondary tracts of styles, strongyles or strongylox-
Description of type species eas. Megascleres, sinuous or vermiform strongyles, strongyloxeas,
and styles sometimes in several size categories; occasionally oxeas.
Pararhaphoxya puZchra (Br\'indsted, 1924a) (Fig. 12). Microscleres absent.
Synonymy, Sigmaxinella puZchra Br\'indsted, 1924a: 151.
Pararhaphoxya puZchra Bergquist, 1970: 16. Pararhaphoxya Remarks
tenuiramosa Burton, 1934a: 565; de Laubenfels, 1954: 173;
Bergquist, 1970: 16. Some authors (Vosmaer, 1912; Bergquist, 1970; Hooper &
Material examined. Holotype (of Pararhaphoxya Levi, 1993b) employ a broad definition of Phakellia, which over-
tenuiramosa): BMNH 1930.8.13.145 - Great Barrier Reef. laps with what is considered Acanthella by others (Van Soest
Description. Erect on stalk; branches dichotomous, slender et aZ., 1990; Alvarez et aZ., 1998). The lack of clear definition and
and cylindrical (3-7 mm in diameter), generally with bifid ends; differentiation between the two genera has created confusion and
surface hispid; colour orange alive, grey-white in alcohol; consis- consequently they contain a heterogenous complex of species. We
tency cartilaginous (description of external characters completed emphasise here the reticulate (Phakellia) versus the dendritic
from Burton, 1934a; Bergquist, 1970) (Fig. 12A). Specialised ecto- (Acanthella) peripheral skeletons, and the flabelliform (Phakellia)
somal skeleton absent; choanosomal skeleton with axial core of versus the irregularly ramose-bushy form (Acanthella) as distinc-
interwoven strongyles, dividing dichotomously along the branches, tive characters to define both genera and avoid further confusion
plumo-echinated by oxeas single, or in bundles (Fig. 12B). Sinuous over the assignment of species. Regional revisions of species are
strongyles, length 950-2050/-lm (1314 ± 401), width 5-10 /-lm required to allocate taxa to either Acanthella or Phakellia.
(6.1 ± 1.9) (n = 10) some with subterminal knobs; oxeas, length
340-610 /-lm (492 ± 76.6), width 5-10 /-lm (7.2 ± 2.0) (n = 10) c 0
with telescoped, blunted tips. Occasionally styles (see Bergquist,
1970: 16) (Fig. 12C-D).
Remarks. Pararhaphoxya puZchra is so far the only species
described for the genus. It is related to an undescribed common
northern New Zealand species recorded by Battershill & Bergquist
(1990). Both are arborescent species with dichotomous branches.
In the latter, however, the extra-axial skeleton is well defined,
plumoreticulate and formed exclusively by oxeas.
Habitat and distribution. The species has been recorded
from the Great Barrier Reef, New Zealand and western Pacific, at
depth range 4-81 m.

PHAKELUA BOWERBANK, 1862

Synonymy

Phakellia Bowerbank, 1862b: 1108. Adreissa Topsent, 1932a:

63 (new synonym).

Type species

Halichondria ventilabrum (sic) (=Spongia ventilabra

Linnaeus, 1767: 1296) (by original designation).

Definition
Fig. 12. Pararhaphoxya tenuiramosa Burton. A, holotype, BMNH
Axinellidae of planar habit with mUltiple axes of sinuous 1930.8.13.145. B, SEM of the choanosomal skeleton. C-D, spicules (scale
strongyles connected by secondary tracts of single spicules. 100 fLm).
Porifera· Demospongiae • Halichondrida • Axinellidae 739

Adreissa (type species Spongia labellum Lamarck, 1814, strongyles, 60--200 f.1m thick (Figs 13C, 14A). Spicules are sinuous
Lamarck'species No. 56, two specimens, MNHN LBIM DT607 and strongyles, length 630--1060 f.1m (897 ± 155.1), width 13-18 f.1m
DT3385; locality, Madagascar [?]) is considered here to be a syn- (14.1 ± 1.7) (n = 10) and styles, length 360--710 f.1m (505 ± 93.8),
onym of Phakellia. The species is a typical Phakellia in growth form width 10--15 f.1m (11.7 ± 1.9) (n = 10), some flexuous and gener-
and surface features, with vein-like tracts enforcing the fan, and with ally bent near the head (Fig. 14B-D).
two size categories of styles. It differs from other species in that Description of species. Flabelliform or cup-shaped to fla-
strongyles are reduced to very thin and sinuous styles-strongyles. belliform; on small peduncle; surface marked by a reticulation of
Phakellia has been previously referred to Silicea (Bowerbank, 'vein-like' lines; oscules pores uniformly distributed on one or both
1862b: 1095), Coralliospongia: Phakelliadae (Gray, 1867a), sides of the lamella; colour grey or pale-yellow-green; grey- beige
Echinonema: Axinellida (Carter, 1885d), Axinellidae: Axinellinae in alcohol; consistency firm, elastic (external features taken from
(Lendenfeld, 1889a), Monaxonida: Halichondrina: Axinellidae Bowerbank 1866; Ackers et al., 1992) (Fig. 13A-B). Specialised
(Ridley & Dendy, 1886; Topsent, 1894c; Topsent, 1896a; Topsent, ectosomal skeleton absent; choanosomal skeleton consists of an
1904b; Topsent, 1928c) and Halichondrida: Axinellidae (Van Soest ascending reticulation of thick primary tracts (up to 7 mm), mark-
et al., 1990; Alvarez et al., 1998). ing surface, densely packed with sinuous strongyles, plumo-
echinated by styles single or in bundles and connected by lines or
Distribution secondary tracts of sinuous strongyles or styles; axial skeleton
compressed and only distinguishable at peduncle (Figs 13D, 14A).
According to Van Soest (1994), the genus has an Arctic- Sinuous strongyles, length 460--1060 f.1m (758 ± 189.1) (n = 20);
Boreal distribution and is restricted to deep and/or colder waters. width 10--18 f.1m (13.4 ± 2.3) (n = 20); styles, length 340--710 f.1m
On the other hand Hooper & Wiedenmayer (1994) reported a wider (458 ± 91.0) (n = 20); width 8-15 /-Lm (11.2 ± 1.8) (n = 20).
distribution for the genus, which includes the amphi-Atlantic, Remarks. The description of Spongia ventilabra of
Mediterranean Sea, Red Sea, Indian Ocean, Indo-Malayan region, Linnaeus refers to an illustration given by Seba (1758: 184, tab. 95
Japan and New Zealand. Species of Phakellia are also present in fig. 8) and also to Spongia strigosa Pallas, 1766: 397. The descrip-
the Central-West Atlantic (Alvarez et al., 1998). Differences in the tion of Pallas' species also mentions Seba's illustration. Vosmaer
patterns of distribution are a consequence of the different interpre- (1912: 311), following Esper (1794), suggested on this basis that
tations of this genus by various authors and the fact that many both species were synonyms and that the type of Phakellia should
species allocated to Phakellia probably belong to other genera. The be called Phakellia strigosa (Pallas). However, no author except
genus contains approximately 34 valid species (Alvarez, 1998). Bergquist (1970) who merely listed the name P. strigosa has fol-
lowed this synonym.
Previous reviews An application to suppress the name of Spongia strigosa
Pallas, 1766 has been submitted to the International Commission on
Bowerbank, 1864: 186; Bowerbank, 1866: 122; Gray, 1867a: Zoological Nomenclature (Alvarez & Willan, in press). This appli-
516; Schmidt, 1880b: 81; Ridley & Dendy, 1886: 478; Ridley & cation (Case number 3216) seeks also to preserve the spelling of
Dendy, 1887: 169; Dendy, 1897: 235; Thiele, 1898: 51; Topsent, Linneaus' species as ventilabrum as this is the name in prevailing
1894c; Wilson, 1902: 399; Topsent, 1904b: 137; Dendy, 1905: 190; use, and to designate the specimen BMNH 10.1.1.2687 as the neo-
Vosmaer, 1912: 310; Ferrer-Hernandez, 1914b: 26; Dendy, 1922b: type of the species, given that there is good evidence that the origi-
116; de Laubenfels, 1936a: 130; Levi, 1973: 605; Bergquist, 1970: nal specimen of Spongia ventilabra is no longer extant.
17; Hooper & Levi, 1993b: 1413. Habitat and distribution. The species is distributed in the
Arctic, North and East Atlantic and Mediterranean. It is found at
Description of type species depths of 10--1863m on sheltered locations and rock ridges
(Ackers et al., 1992; Boury-Esnault et aI., 1994b).
Phakellia ventilabra (Linnaeus, 1767) (Figs 13-14).
Synonymy. Spongia ventilabra Linnaeus, 1767: 1296.
Halichondria ventilabrum (sic); Johnston, 1842: 107. Phakellia
ventilabrum (sic); Bowerbank, 1866: 122; Carter, 1876: 240; PHYCOPSIS CARTER, 1883
Fristedt, 1885: 47; Vosmaer, 1885a: 23; Ridley & Dendy, 1887:
170; Topsent, 1891a: 529; Vosmaer, 1912: 316; Dendy 1922b: 118; Synonymy
Topsent, 1928c: 174; Cabioch, 1968b: 221; Vacelet, 1969: 178;
Ackers et al., 1992: 85; Boury-Esnault et al., 1994b: 89. Phycopsis Carter, 1883b: 319. Ketosus de Laubenfels, 1936a:
Material examined. Neotype (proposed): BMNH 130 (new synonym).
10.1.1.2687, dry - near Lervig, Norway. Other material. Nine spec-
imens, BMNH 77.5.21.141, wet - Haff Banks, Shetland, coll. Type species
Humphreys, 1852.
Description of neotype (Fig. 13A). Flabellate specimen, Phycopsis hirsuta Carter, 1883b: 319 (by subsequent
dry, approximately 10--23cm high by 10--20cm wide; on short designation; Thiele, 1899: 11).
peduncle, approximately 1 cm in diameter; surface hispid, marked
by reticule of spicule tracts, some ascending ones thick (5-7mm) Definition
like veins. Specialised ectosomal skeleton absent; choanosomal
skeleton reticulated, with primary tracts of sinuous strongyles, Axinellidae with ascending multispicular tracts, radiating
300--600 f.1m thick, plumo-echinated by styles or connected by sec- from central axis and ending in long and filamentous surface
ondary unispicular or paucispicular tracts of styles, and sometimes processes.
740 Porifera· Demospongiae • Halichondrida • Axinellidae

Fig. 13. Phakellia ventilabra (Linneaus). A, neotype, BMNH 10.1.1. 2687. B, 4 of 9 specimens from the Shetlands, BMNH 77.5.21.141. C, BMNH
10.1.1.2687, section through the choanosomal skeleton at the periphery. D, BMNH 77.5.21.141, SEM of the choanosomal skeleton (scales B, 3cm;
C, 400 j.Lm; D, 200 j.Lm).

Diagnosis spicule tracts sheathed by spongin, that end as surface processes

(distinct for each of the genera).
Branching-bushy-stipitate and stalked growth form. Surface Ketosus is a monotypic genus erected by de Laubenfels
villous with long processes as filaments; ectosome without spe- (1936a) for Dictyocylindrus setosus Bowerbank, 1873b: 325, with
cialised skeleton. Choanosomal skeleton differentiated into axial a brief diagnosis that overlaps significantly with several genera
and extra-axial regions. Axial skeleton compressed and vaguely ofAxinellidae (i.e., Acanthella, Phakellia, Pararhaphoxya). The
reticulated with close-set of spongin fibres and spicules. Extra- features of the type species Phycopsis setosa (Bowerbank, 1873b)
axial skeleton with thick spongin fibres cored by ascending multi- (new combination and lectotype designated here, BMNH
spicular tracts, radiating from central axis and ending in surface 1877.5.21.1310, slide, 'Bowerbank collection, Bere Regis,
processes. Megascleres strongyles, oxeas-anisoxeas and styles Devonshire (?)') as well as the original description given by
(including sinuous forms), not localised in any particular area of Bowerbank (1873b), agrees with the diagnosis of Phycopsis as
the skeleton. Microscleres absent. presently understood. Phycopsis setosa has surface projections in
the shape of 'setae', considered here homologous with the long fil-
Remarks amentous processes present in other species the genus. It also
shares with other species the skeletal organisation (Fig. 15E) and
Phycopsis is related to Reniochalina and Ptilocaulis. The three the type of spicules (strongyles, oxeas-anisoxeas and styles; Fig.
genera have similar spicule composition (with only Reniochalina 16D-E). It, however, differs in growth form, which is fan-shaped
having oxeas-anisoxeas with spined tips) and a skeleton of ascending instead branching or arborescent as seen in other Phycopsis spp.
Porifera· Demospongiae • Halichondrida • Axinellidae 741

Previous reviews
c
Carter, 1883b: 319; Carter, 1884a: 130; Thiele, 1899: 11;
Topsent, 1904b: 137; Vosmaer, 1932-35: 721,722; de Laubenfels,
1936a: 131; Tanita & Hoshino, 1989: 88.

Description of type species

Phycopsis hirsuta Carter, 1883b (Figs 15-16).

Synonymy. Phycopsis hirsuta Carter, 1883b: 319; Hooper &
Wiedenmayer, 1994: 79.
Material examined. Holotype: BMNH 84.4.14.1, dry -
Australia, 'D6 KKl , Deditam Dickie, Presd. by H.I. Carter'.
Description. Branching, 20 cm high and 15 cm wide on
broad stalk, 0.5 cm in diameter; branches cylindrical, generally
dichotomously divided, rigid, tapering towards ends, some anasto-
mosing; surface with thin and long processes 1-2 mm long; colour
brown in dry condition; consistency hard (Fig. 15A). Specialised
ectosomal skeleton absent. Choanosomal skeleton differentiated in
axial and extra-axial regions. Axial skeleton condensed with close-
set spongin fibres and spicules forming a vague reticulation; extra-
axial skeleton with light spongin fibres 60-100/-Lm, cored by
spicule tracts, radiating from central axis and ending in surface
processes. Single spicules dispersed among spongin fibres or
arranged in a halichondrioid-like skeleton. Spicules are not
localised in any particular area of the skeleton (Figs 15B-E, 16A).
Oxeas-anisoxeas, length 310-410 /-Lm (366 ± 34.7), width 5-12 /-Lm
(9.9 ± 2.3) (n = 10), fusiform or telescoped, straight, slightly bent
or slightly sinuous; strongyles, length 300-690/-Lm (502 ±
Fig. 14. Phakellia ventilabra (Linneaus), BMNH 10.1.1.2687. A, diagram
of the skeleton. B, strongyle. C-D, styles (scale 100 fLm).
136.45), width 5-13 /-Lm (8.8 ± 2.1) (n = 10), sinuous or straight;
styles-styloids, length 310-410 /-Lm (355 ± 34.7), width 5-12 /-Lm
(9.1 ± 2.1) (n = IO)(Fig. 16B-E).
The type locality of P. setosa is ambiguous, given the possibility Remarks. The species has not been recorded since its origi-
that the locality originally assigned by the collector is erroneous nal description. The type material is well preserved and corresponds
(Bowerbank, 1873b). Indeed a species like this one has not been partially with the description given by Carter (1883b) in which only
recorded yet from the well-known fauna of the NE Atlantic and one type of spicules (i.e., oxeas-anisoxeas) was reported. The
therefore this record is definitely not reliable to expand the distribu- strongyles and the styles found in the type material were not men-
tion of Phycopsis which, is currently limited to southern latitudes. tioned in the original description of Carter. According to Carter
Both Phycopsis hirsuta and P. setosa closely resemble species (1883b), the species resembles Phycopsisfruticulosa recorded from
of Ptilocaulis reported from New Caledonia, viz., P. fusiformis Tasmania, differing only in the length of the surface processes and
Levi. 1967a, P. epakros Hooper & Levi, 1993b and P. papillatus the diameter of the axis. No strongyles or styles were reported for
Hooper & Levi, 1993b. Phycopsis might be a more appropriate this species either. Phycopsis hispidula (Ridley, 1884a), another
genus for the New Caledonian species; they all share with species from the north coast of Australia, was originally described
Phycopsis hirsuta the long surface projections of the choanosomal under the genus Protoschmidtia Czerniavsky, 1879 (=Haliclona)
skeleton and have similar spicule composition and habit. Close and referred to Phycopsis by Hooper & Wiedenmayer (1994). The
phylogenetic relationships among these species were also shown identity of this species needs confirmation as its original description
by Hooper & Levi (1994). Other species of Phycopsis are: P. fru- disagrees with the diagnosis of Phycopsis given here.
ticulosa Carter, 1883b: 320 and P. hispidula (Ridley, 1884a), and P. Habitat and distribution. Only known from its type locality
terpnis de Laubenfels, 1954: 176. Axinyssa aculeata Wilson, 1925 (South Australia).
from Hawaii was referred to Phycopsis by de Laubenfels (l936a:
131; 1951 : 263) and Tanita & Hoshino (1989: 89), but the species
is currently assigned to its original genus (Van Soest et al., 1990; PTILOCAUllS CARTER, 1883
Hooper & Bergquist, 1992).
Phycopsis has been previously referred to Echinonemata: Synonymy
Axinellida (Carter, 1883b; Carter, 1884a,b, 1885d), Halichondrina:
Axinellidae (Topsent, 1904b; Topsent, 1928c) and Raspailiidae Ptilocaulis Carter, 1883b: 321.
(Van Soest et al., 1990).
Type species
Distribution
Ptilocaulis gracilis Carter, 1883b: 321 (by subsequent desig-
Tasmania, South Australia, Indo-Malaysia, W Pacific nation; de Laubenfels 1936a: 127) (junior synonym of Pandaros
(Hooper & Wiedenmayer, 1994), New Caledonia. walpersi Duchassaing & Michelotti, 1864).
742 Porifera· Demospongiae • Halichondrida • Axinellidae

Fig. 15. Phycopsis. A-D, Phycopsis hirsuta Carter. A, holotype BMNH 84.4.14.1. B, SEM of a longitudinal section through the skeleton. C, detail of
skeletal projection at surface. D, SEM of a transverse section through the skeleton. E, Phycopsis setosa (Bowerbank), longitudinal section through the
lectotype BMNH 1877.5.21.1310 (scales A, 2.5cm; B, 500 !Lm; C, 30 !Lm; D, 500 !Lm; E, 600 !Lm).

Definition Choanosomal skeleton differentiated in axial and extra-axial

regions; axial skeleton vaguely reticulated; extra-axial skeleton
Axinellidae with axial skeleton vaguely reticulated and extra- irregularly reticulated with ascending fibrofascicIes cored with
axial skeleton formed by ascending fibrofascicIes cored with styles styles and ending in surface processes. Megascleres are styles gen-
and ending in surface processes. erally in two size categories; occasionally oxeas or anisoxeas.
Microscleres absent.
Diagnosis
Remarks
Clavate, branching, bushy or lamellate growth form. Surface
covered by scopiform (i.e., flattened and spatula-shaped (Carter, The flattened surface processes, also referred to as 'scopi-
1833» processes; ectosome without specialised skeleton. form' (Carter, 1883b), 'surface tufts' (Ridley, 1884a), papilliform
Porifera· Demospongiae • HaIichondrida • Axinellidae 743

Distribution

The genus has relatively well-recognised species with disjunct

distributions (Hooper & Levi, 1994). These are: Ptilocaulis rigidus
Carter, 1883b: 322 from Australia, P. marquezi (Duchassaing &
B Michelotti, 1864: 40 and P. walpersi (Duchassaing & Michelotti,
1864) from the central West Atlantic (Alvarez et al., 1998); P. digi-
tatus Topsent, 1928c: 172 from the central North Atlantic; P. spi-
culifera (Lamarck, 1813-1814: 449) from the Indo West Pacific
(Ridley, 1884a: 617; Dendy, 1922b: 115; Burton, 1959a: 266;
Pulitzer-Finali, 1993: 289); P. epakros Hooper & Levi, 1993b; P.
fusiformis Levi, 1967a and P. papillatus Hooper & Levi, 1993b
from New Caledonia (Hooper & Levi, 1993b). As mentioned
above, the species of Ptilocaulis reported from New Caledonia by

I Hooper & Levi (1993b) have affinities with Phycopsis and are
probably better placed in this genus. Ptilocaulis echidnaeus
(Lamarck, 1813) (fide Topsent, 1932a: 108) is recorded for the
coast of Africa (?) by Lamarck (1813), and thus the type locality of
this species remains ambiguous.

Previous reviews

Carter, 1884a: 130; Topsent, 1928c: 37, 172; de Laubenfels,

1936a: 127; Levi, 1973: 605; Wiedenmayer, 1977b: 152; Hooper &
Levi, 1993b: 1426; Alvarez et al., 1998: 22.

Description of type species

Ptilocaulis walpersi (Duchassaing & MicheIIoti, 1864)

(Figs 17-18).
Synonymy. Pandaros walpersi Duchassaing & Michelloti,
1864: 90. Ptilocaulis gracilis Carter, 1883b: 321. Ptilocaulis
Fig. 16. Phycopsis. A-C, Phycopsis hirsuta Carter, holotype BMNH walpersi; Zea 1987; Alvarez et al. 1998 (with additional synonyms).
84.4.14.1. A, diagram of the skeleton. B, strongyle. C, oxea. D-E, Material examined. Lectotype: MT Por. 56 (USNM 31025
Phycopsis serosa (Bowerbank), lectotype BMNH 1877.5.21.1310. D,
slide) - St. Thomas, Virgin Islands, Caribbean. Holotype of
strongyle. E, oxea (scale 100 !Lm).
Ptilocaulis gracilis: BMNH 1845.12.30.1, dry - Caribbean.
Description. Encrusting, irregularly massive, plate-shaped,
projections (Hooper & Levi, 1993b) are distinctive for species of fan-shaped, or erect finger-shaped; dichotomously branching (as in
Ptilocaulis but cannot be used as a diagnostic character to define the lectotype of Ptilocaulis gracilis) or fonning brushes of 10 or
the taxon. The genus was included in the Desmoxyidae by Van more closely-cropped branches. Surface uniformly covered with
Soest et al. (1990) based on the similarities in surface characteris- scopifonn flattened processes, 2 mm long and approximately
tics with other species of Desmoxyidae, such as Higginsia. Alvarez 5-lOmm apart and minute pores, less than 1 mm in diameter;
et al. (1998) showed that similar surface processes were present in colour orange-red when alive; white-beige in alcohol; brown in dry
some species of Axinelia and on this bases the genus was trans- state (Fig. 17A-B). Specialised ectosomal skeleton absent;
ferred to its original family. Similar surface processes are found choanosomal skeleton with developed fascicles of ascending and
also in Phycopsis and Reniochalina. anastomosing primary fibres (styles and spongin), radiating to the
The organisation of the choanosomal skeleton, although periphery and ending in flattened surface processes. Main fibres
differentiated in axial and extra-axial regions, differs slightly from interconnected irregularly by single fibres, some unevenly core
other axinel\id genera. The extra-axial skeleton of Ptilocaulis, instead with single spicules, and/or short and thin paucispicular tracts (Figs
of plumo-reticulated (e.g., Axinella, Cymbastela. Dragmacidon) is 17C-D, 18A). Styles of two sizes: (1) length, 260-340 f.Lm (295.6
fonned by fibres that merge or intricately interconnect (fibrofasci- ± 18.0), width 15-10 f.Lm (13.3 ± 1.9) (n = 25); (2) length,
cles) in an irregular reticulation. Similar architecture is also seen in 860-490 f.Lm (715.6 ± 81.0), width 10-5 f.Lm (7.2 ± 2.2) (n = 25);
Phycopsis and Reniochalina. longest projecting from scopifonn processes and less frequent than
The similarities and close phylogenetic relationships among shorter ones of the choanosomal skeleton (Fig. 18B-C).
species of Ptilocaulis and Reniochalina has been well established Remarks. The species is common in the Caribbean and due
by Hooper & Levi (1993b; 1994) and Alvarez et al. (2000a). to its variable growth fonn has often been confused with other
The genus has been previously referred to Echinonemata: axinellids (see list of synonyms and extended remarks in Alvarez
Axinellida (Carter, 1884a,b; 1885d), Halichondrina: AxineIIidae et al., 1998). Ptilocaulis walpersi is similar to P. marquezi in its
(Topsent, 1904b; 1928c), Halichondrida: Desmoxyidae growth fonn, skeletal organisation and distribution, but differs in
(Van Soest et aI., 1990); Axinellida: Axinellidae (Hooper & Levi, spiculation. Ptilocaulis marquezi has styles transitional to styloids,
1993b) and Halichondrida: Axinellidae (Alvarez et al., 1998). in only one size category, and oxeas transitional to strongyloxeas.
744 Porifera· Demospongiae • Halichondrida • Axinellidae

Fig. 17. Ptilocaulis gracilis Carter. A, holotype BMNH 1845.12.30.1, dry. B, detail of scopiform processes. C, detail of spiculo-fibres. D, longitudinal
section through the choanosomal skeleton (scales A, 5 cm; C, 50 /-Lm; D, 300/-Lm).

C Definition
\
B
Axinellidae with extra-axial spongin fibres projecting
into surface processes and cored with paucispicular tracts of
oxeas, anisoxeas and styles. Oxeas with tips sunnounted by
microspines.

Diagnosis

Branching or fan-shaped, occasionally lobate, growth fonn.

Surface covered by flattened (spatulate) long processes. Ectosome
without specialised skeleton. Choanosomal skeleton differentiated
into axial and extra-axial regions, projecting into surface processes.
Extra-axial skeleton reticulated with ascending spongin fibres,
Fig. 18. Ptilocaulis. A, diagram of the skeleton of the type species. interconnected at all angles by single spicules or short fibres, or
B-C, styles ofUSNM 31025 (spicule preparation made from the lectotype anastomosing and fonning oval to round meshes; spongin fibres
MT POR. 56 of Ptilocaulis waipersi (Duchassaing & Michelotti)) (scale
cored with paucispicular spicular tracts, sometimes plumose;
100/-Lm).
axial skeleton slightly condensed and reticulated as in extra-axial
region. Megascleres are oxeas-anisoxeas with tips sunnounted by
microspines, which are rudimentary or absent; occasionally styles
Habitat and distribution. Widely distributed in the or styloids. Microscleres absent.
Caribbean, from Florida to Colombia and off Bennuda. It is found
from shallow water to 35 m depth on coral reef bottoms. Remarks

The morphological and genetic affinities among species

RENIOCHALINA LENDENFELD, 1888 of Ptilocaulis and Reniochalina have been well established
(Hooper & Levi, 1993b, 1994; Alvarez et al., 2000a). These genera
Synonymy together with Phycopsis have a similar choanosomal skeleton of
ascending spiculo-fibres ending in conspicuous processes at the
Reniochalina Lendenfeld, 1888: 82. Axiamon Hallmann, surface. They differ in spicule composition and shape of the surface
1914: 440 (objective synonym). processes (long and flattened in Reniochalina, spatula-shaped
in Ptilocaulis and filamentous in Phycopsis. These three genera
Type species are likely to constitute a monophyletic group that could be
classified as a subfamily within Axinellidae. Phylogenetic analyses
Reniochalina stalagmitis Lendenfeld, 1888: 82 (by based on morphological characters support this view (Alvarez,
subsequent designation; Hallmann, 1914). 1998).
Porifera' Demospongiae • Halichondrida • Axinellidae 745

Fig. 19. Reniochalina staiagmitis Lendenfeld. A, lectotype BMNH 1887.4.27.122. B-C, SEM of the choanosomal skeleton (scale A, 2 cm).

The synonymy ofAxiamon with Reniochalina was established and New Zealand (Alvarez, unpublished data). It is represented so
by Wiedenmayer (1989) and confirmed here by comparison far by three other species besides the type: Reniochalina sectilis
of their respective type specimens (see Hooper & Levi, 1993b for Wiedenmayer, 1989: 49, from South Australia, R. condylia
further comment in this synonymy). Hooper & Levi, 1993b: 1405, and R. plumosa (Levi & Levi,
The genus has been previously referred to Halichondrina: 1983b: 945) from New Caledonia (Hooper & Levi, 1993b; 1994).
Haploscleridae: Renierinae (Topsent, 1894c), Homorrhaphidae: The descriptions of these species, however, do not fully agree
Renierinae (Lendenfeld, 1889a), Haploscleridae (de Laubenfels, with the diagnosis of the genus proposed here and their identities
1936a) and Axinellida: Axinellidae (Wiedenmayer, 1989; Hooper & need to be confirmed.
Levi, 1993b).
Previous reviews
Distribution
Whitelegge, 1902: 283; Hallmann, 1914: 346; de Laubenfels,
Reniochalina seems to be restricted to Australia, Indo-west 1936a: 47, 130; Wiedenmayer, 1989: 48; Hooper & Levi, 1993b:
Pacific region (Hooper & Wiedenmayer, 1994; Van Soest, 1994) 1403.
746 Porifera· Demospongiae • Halichondrida • Axinellidae

Fig. 20. Reniochalina stalagmitis Lendenfeld. A, holotype of Axiamon folium Hallmann, AM 09004. B, SEM of the skeleton. C-D, tips of anisoxea
(scales A, IOem; B, 200 IJ.m; C, 21J.m; D,5IJ.m).

Description of type species consistency firm but flexible (Figs 19A, 20A). Specialised ectosomal
skeleton absent; choanosomal skeleton differentiated in axial and
Reniochalina stalagmitis Lendenfeld, 1888 (Figs 19-21). extra-axial regions. Extra-axial skeleton reticulated with ascending
Synonymy. Reniochalina stalagmitis Lendenfeld, 1888: 83; spongin fibres, 50-100 f.1m interconnected at all angles by single
Whitelegge, 1902: 283; Hallmann, 1914: 346; Hooper & spicules or short fibres, or anastomosing and forming oval to round
Wiedenmayer, 1994: 81 (with additional synonyms). meshes up to 200 f.1m in diameter. Spongin fibres slightly developed
Material examined. Lectotype: BMNH 1887.4.27.122 - and cored with paucispicular tracts, sometimes plumose. Axial
Western Australia. Holotype of Axiamon folium: AM G9004, wet - skeleton slightly condensed and reticulated as in extra-axial region
Western Australia. 'Syntype' (?): AM B5478, wet - no locality data. (Figs 19B-C, 20B, 21A). Oxeas or anisoxeas, straight, bent or
Description. Branching to fan-shape, flat branches, dichoto- slightly sinuous; with tips surmounted by microspines, which might
mous or fused; surface with long surface processes and parallel be rudimentary or absent; microspines at one end might be half size
channels, 1 mm apart, 1 mm depth, running longitudinally with pro- to those at other end; length 190-315 (240.2 ± 37.6) f.1m, width 8-15
jecting spicules; colour orange-red alive, dark brown in alcohol; (11.9 ± 2.8)f.1m (n = 10) (Figs 20C-D, 21B-D).
Porifera· Demospongiae • Halichondrida • Axinellidae 747

B
o

Fig. 21. Reniochalina stalagmitis Lendenfeld: A, diagram of the choanosomal skeleton; B-D. different classes of anisoxeas (scale 100 ILm).

Remarks. According to Wiedenmayer (1989) the species is Additional financial and logistic support was also provided by
similar to Reniochalina sectilis but we consider that the latter has Division of Botany and Zoology, Canberra; Queensland Museum,
affinities with the Raspailiidae (see below). Reniochalina lamella Brisbane; Electron Microscopy Unit, Research School of
Lendenfeld, 1888 was synonymised by Wiedenmayer (1989) with Biological Sciences (EMU-ANU); National Institute of Water and
R. stalagmitis. The two species seem to differ slightly in habit (see Atmospheric Research, Wellington and Museum and Art Gallery
Wiedenmayer, 1989: plate 4.2, and additional remarks therein). of the Northern Territory, Darwin. We are especially grateful: to
Reniochalina stalagmitis is genetically and morphologically John Kennedy, Steve Cook and Lisa Goudy (QM) for assisting
similar to an undescribed species of Reniochalina from the Great with the preparation of the material; to Ms Joane Moore for the
Barrier Reef, Australia, recorded in Alvarez et al. (2000a). This line drawings. Rob Van Soest (ZMA Amsterdam) and Richard
species differs from R. stalagmitis in habit, which is branching- Willan (NTM, Darwin) made valuable suggestions this chapter.
erect; with cylindrical branches, semi-buried in the substrate, We are also grateful to the following museum curators/collection
dichotomously divided near the tips. It differs also in the size of the managers who made available the type material required for the
oxea microspines, which are much smaller and only visible under taxonomic revision: Penny Berents (AM); Clare Valentine
high magnification (e.g., SEM analysis). (BMNH); Claude Levi (MNHN); e. Carpine (MOM), Manfred
Habitat and distribution. The species seems to be widely Grasshoff (Natur-Museum und Forschungsinstitut Senckenberg,
distributed throughout Australia: W coast, Western Australia, Great Frankfurt); Dieter Kuehlmann (Zoologisches Museum fUr
Barrier Reef, Queensland coast, Torres Strait (see Hooper & Naturkunde an der Humboldt-Universitat zu Berlin); J.e. den
Wiedenmayer, 1994: 82). It is found in coralline substrate between Hartog, National Natural History Museum, Leiden; Klaus Riitzier
10-20m depth. and Kate Smith (USNM); Ole Tendal (ZMUC DEN). JH also
acknowledges a post-doctoral fellowship (1988) at the MNHN
ACKNOWLEDGEMENTS Paris under Claude Levi, and a Churchill Fellowship at BMNH
London (1988), from which many of the raw data concerning
BA's studies on Axinellidae and Bubaridae were made possi- types and the true characteristics ofAxinellidaelBubaridae taxa
ble by an Australian National University (ANU) scholarship. were initially gathered.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John NA. Hooper and Rob WM, Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Bubaridae Topsent, 1894

Belinda Alvarez! & Rob W.M. Van Soest2

1 Museum and Art Gallery of the Northern Territory, P.O. Box 4646, Darwin, NT 0801, Australia. (belinda.g1asby@nt.gov.au)
2Zoological Museum, Faculty of Science, University of Amsterdam, P.O. Box 94766, 1090 GT, Amsterdam, The Netherlands.
(soest@science.uva.nl)

Bubaridae Topsent (Demospongiae, Halichondrida) is used here in the restricted sense of Topsent (1928c) to group exclusively encrusting
forms that were related to the family Axinellidae at one time or another. The family includes four valid genera and approximately 30
species, characterised by a basal layer of interlacing spicules, generally diactines, and erect monactines spicules, with bases embedded in
the basal layer, projecting perpendicularly to the substratum. Most species of the family are found in deep waters down to 1300 m but
shallow water species are also known.
Keywords: Demospongiae; Halichondrida; Bubaridae; Bubaris; Cerbaris; Hymerhabdia; Monocrepidium.
- - - - - - ~ . - - - - . - - -... - - - -

DEFINITION, DIAGNOSIS, SCOPE Distribution

Synonymy Most genera are found only in the East Atlantic and
Mediterranean, and generally in deep waters (down to 1300 m
Bubarinae Topsent, 1894c: 20 (in part); Bubaridae sensu depth). However, the genus Bubaris has an allegedly wider distri-
Hentschel, 1923: 407 (in part); Topsent 1928c: 42; Vacelet, 1969: bution including species records from the Arctic, Indian Ocean,
180 (in part); Levi, 1973. Indonesia, Japan, New Zealand and Antarctica.

Definition History and biology

Halichondrida of encrusting habit and skeleton differentiated Topsent (1894c: 20; 1898b: 247) erected the subfamily
into a basal layer of interlacing spicules and a perpendicular layer Bubarinae (under Poeciloscleridae: Halichondrina) to include
of monactines with bases embedded in the basal skeleton. Plocamia Schmidt, Suberotelites Schmidt, Bubaris Gray, Cerbaris
Topsent, Rhabderemia Topsent and Hymerhabdia Topsent.
Diagnosis Cerbaris, Plocamia, Rhabderemia, and Suberotelites were later
transferred by Topsent (1904b) to a subfamily Ectyoninae.
Encrusting sponges generally with hispid surface. Ectosomal Hentschel (1923) defined the family Bubaridae as 'with spe-
skeleton absent. Choanosomal skeleton differentiated into a basal cial diactinal spicules of remarkable form, which are placed
layer of interlacing diactines and, in some cases additional styles; in encrusting sponges at the base, in erect sponges in the centre,
and perpendicular to this layer longer monactines with bases and in branching sponges in the axis. If these spicules are absent
embedded in the basal skeleton. Megascleres in the basal skeleton there may be walking-stick (cane) spicules (rhabdostyles). Main
are diactines of different kinds (strongyles, oxeas, anisoxeas, megascleres almost always monactinal. Microscleres can be
angulated-centrotylote oxeas, anisoxeas, ceroxas, i.e., m-shape chelae or sigmas, or can be absent'. Genera included were
diactines), different shapes (sinuous or vermicular, bent, angulate Plocamiopsis, Lithoplocamia, Crambe, Monocrepidium (erected
or centrotylote) and different textures (smooth, evenly or unevenly under Desmanthidae by Topsent, 1898b: 229), Rhabdoploca
acanthose, tuberculate or annulate). Rhabdostyles or styles with (erected under Ectyoninae by Topsent, 1904b: 157) in addition to
contorted or reflexed bases are also present. Longer styles (some- those assigned by Topsent (1894c, 1898b) to Bubarinae.
times in two size categories) or tylostyles are located perpendicu- Topsent (1928c) recognised that Bubaridae as defined by
larly to substrata. Raphides occasionally present. Hentschel (1923) was a heterogeneous assemblage of genera and
proposed a new arrangement to include only those genera with a
Scope basal layer of spicules and perpendicular monactines (i.e., Bubaris,
Cerbaris, Hymerhabdia, Monocrepidium and Rhabdoploca). Other
The family includes nine nominal genera, but only four are con- genera included in Bubaridae subsequent to the arrangement pro-
sidered valid under the definition of the family employed here. Other posed by Topsent were: Bubaropsis Levi and Vacelet, Lithobubaris
genera of the nominal Bubaridae are currently included in other Vacelet, Skeizia Cabioch, Rhabdobaris Pulitzer-Finali and Uplexoa
families (i.e., Suberotelites Schmidt (= Phorbas) in Hymedesmii- de Laubenfels.
dae; Plocamia Schmidt (=Antho (Acarnia)) in Microcionidae; In some axinellid genera (e.g., Phakellia, Auletta and Acanthella)
Plocamiopsis Topsent (=Antho (Acarnia)) in Microcionidae; the axial skeleton is compressed and formed by interlaced strongyles
Crambe Vosmaer in Crambeidae; Lithobubaris Vacelet in from which longer monactines project perpendicularly. A similar
Desmanthidae; Lithoplocamia Dendy in Raspailiidae; Rhabderemia architecture is seen in members of Bubaridae, as initially suggested by
Topsent in Rhabderemiidae (see respective chapters in this volume). Hentschel (1923), and led some authors to include some bubarid

748
Porifera· Demospongiae • Halichondrida • Bubaridae 749

genera in AxinelJidae (e.g., Bergquist, 1970; Van Soest et aI., 1990). similarities of the skeletal organisation seen in both groups (see
The resemblance in skeletal structure between some AxinelJidae and Desmanthidae in this volume).
Bubaridae suggests a close relationship between these two families. At Bubaridae is restricted here to encrusting forms with similar
this stage, however, is not possible to establish whether or not this sim- skeletal architecture (i.e., a basal layer of spicules with perpendicu-
ilarity can be interpreted as a homology to relate both families, or if it lar monactines). Detailed studies of bub arid species must be under-
is a condition independently acquired in both groups. taken to establish whether or not this type of skeletal arrangement
Relationships among some members of Desmanthidae (i.e represents a good synapomorphy to unite the members of
Lithobubaris) and Bubaridae have also been proposed based on Bubaridae.

KEY TO GENERA

(1) Basal skeleton formed only by diactines ............................................................................................................................................. 2

Basal skeleton includes monactines (rhabdostyles). Diactines bent or angulate oxeas .................................................. Hymerhabdia
(2) Diactines of basal skeleton always smooth, typically vermicular or sinuous. Spicules projecting perpendicular to
substrate are long and smooth styles ........................................................................................................................................ Bubaris
Diactines (including strongyles, toxostrongyles, oxeas and ceroxas) evenly or unevenly acanthose, but smooth ones
occasionally present. Spicules projecting perpendicular to substrate are styles and rhabdostyles sometimes in two size
categories ....................................................................................................................................................... .......................... Cerbaris
Diactines are tuberculate or annulate but smooth ones occasionally present. Spicules projecting perpendicUlar to
substrate are styles and tylostyles ................................................................................................................................ Monocrepidium

BUBARIS GRAY, 1867 Distribution

Synonymy Approximately twenty species of Bubaris have been reported

in the literature (Alvarez, 1998), mainly from deep waters, but a
Bubaris Gray, 1867a: 521. revision is necessary as some of these species may be better allo-
cated to Phakellia or Acanthella. The genus has an allegedly wide
Type species distribution including records from the Arctic, amphi-Atlantic,
S Atlantic Ocean, Mediterranean Sea, Indian Ocean, Japan,
Hymeraphia vermiculata Bowerbank, 1866: 141 (by original New Zealand, Antarctica (Hooper & Wiedenmayer, 1994).
designation).
Previous reviews
Definition
Gray (1867a: 521); Carter (1880b: 46); Topsent (1891a: 546);
Bubaridae with basal skeleton formed by smooth sinuous or Topsent (1894c: 22); Thiele (1903b: 379); Topsent (1904b: 137,
vermicular strongyles or strongyloxeas. 145); Dendy (1922b: 62); Topsent (1928c: 189); Burton (1928a:
128); de Laubenfels (1936a: 131); Bergquist (1970: 19); Hechtel
Diagnosis (1969: 25); Boury-Esnault & Van Beveren (1982: 48); Boury-
Esnault & Lopes (1985: 171).
Encrusting. Surface hispid. Choanosomal skeleton formed by
bundles or individual styles projecting perpendicularly to substrata Description of type species
with heads embedded in a basal layer of vermicular or sinuous
strongyles or strongyloxeas. Bubaris vermiculata (Bowerbank, 1866) (Figs 1-2).
Synonymy. Hymeraphia vermiculata Bowerbank, 1866:
Remarks 141. Bubaris vermiculata; Topsent, 1891a: 545; Topsent, 1894c.
Material examined. Lectotype (here designated):
The skeleton of Bubaris resembles the one found in some BMNH 30.7.3.86 - Shetland, Bowerbank collection, colI. Barlee,
axinellid genera such as Phakellia, Acanthella and Auletta Peach & Gregor (Fig. lA). Paralectotypes (here designated): BMNH
(i.e., axes of sinuous strongyles with projecting styles); but 30.7.3.85 - Shetland, Bowerbank collection (Fig. lB); BMNH
as stated above, this homology is considered insufficient to 10.1.1.89, dry - Shetland, 100 fathoms (Fig. lC). Other material.
relate Bubaris to Axinellidae. It is also possible that some BMNH not registered - 2 slides, Bowerbank collection No. 453;
species of Bubaris are actually juvenile forms of Phakellia or BMNH 1910.1.1.564 - Lervig, Norway, Norman collection.
Acanthella, but currently there is no empirical evidence to support Description. Encrusting, 0.5-1 cm diameter, on pebbles or
this hypothesis. bivalve shells; surface microhispid; red in life (Boury-Esnault &
The genus has been previously referred to Leiospongia Lopes, 1985) (Fig. 2); grey in dry state. Specialised ectosomal
(Halichondriadae) (Gray, 1867a), Halichondrina: Poeciloscleridae: skeleton absent; choanosomal skeleton basal layer of sinuous-
Bubarinae (Topsent, 1894c), Halichondrina: Bubarinae (Topsent, vermiform strongyles or strongyloxeas echinated by long
1896a), Halichondrina: Bubaridae (Topsent, 1928c), Axinellida: styles perpendicular to substrate with heads embedded in basal
Bubaridae (Levi, 1973); Halichondrida: Axinellidae (Van Soest layer (Fig. 10). Megascleres styles (Fig. IE), length 230-
et al., 1990). 2200 p..m (681::': 572.5), width 13-30 p..m (21.7::': 5.8) (n= 10),
750 Porifera· DeIDospongiae • Halichondrida • Bubaridae

Fig. 1. Bubaris vermiculata (Bowerbank). A, lectotype, BMNH 30.7.3.86. B, paralectotype BMNH 30.7.3.85. C, paralectotype, BMNH 10.1.1.89.
D-G, BMNH 1910.1.1.564. D, SEM ofthe skeleton. E-G, styles and vermicular anisoxeas (scales A-C, I cm; E, 100 fLm; F-G, 50 fLm).

strongyloxeas (Fig. IF)- anisoxeas (Fig. 10), sinuous to vermi-

form, length 360-510 /LID (438 ± 45.2), width 8-13 /Lm (10.5 ±
1.9) (n = 10).
Remarks. Other material examined included possible syn-
types of Hymeraphia vermiculata var. erecta (Carter, 1876): 307
(BMNH 1982.7.28.30, W of Shetlands Is., sta. 82, 312 fms, colI.
HMS 'Porcupine'; BMNH 1882.7.28.34, West of Shetland Is.,
624m depth, sta.82, HMS 'Porcupine'(?». This species does not
agree with the concept of Bubaris under this revision. It has close
affinities with species of Acanthella such as A. flagelliformis (Van
Soest & Stentoff, 1988) and A. mastophora (Schmidt, 1870) as
suggested by Carter (1876: 308).
Habitat and distribution. East Atlantic, 9-1360 m depth. The
species has been also recorded for the Mediterranean (Babic, 1922:
238; Sara, 1960a: 452; Sara & Siribelli, 1962: 30; Pulitzer-Finali,
1977: 36; Pulitzer-Finali, 1983: 523), Indonesia (Topsent, 1897a;
Desqueyroux-Faundez, 1981), Kerguelen Island, Tristan da Cunha
Fig. 2. Bubaris vermiculata, in situ, Ireland (photo Bernard Picton). (Boury-Esnault & Van Beveren, 1982: 48) and New Zealand (Dendy,
Porifera' Demospongiae • Halichondrida • Bubaridae 751

1924: 351; Bergquist, 1970: 19) - but all these latter records might initially erected in Bubaridae and described as a small encrusting
represent different species of Bubaris. fragment with disorganised skeleton of spicUles. The megascleres
included oxeas of variable sizes, straight, curved and angulated
(600-1600/-lm X 20-55 /-lm) toxiform oxeas (110-150/-lm) and
CERBARIS TOPSENT, 1898 strongyles slightly sinuous (300-450 /-lm X 22 /-lm).Trichodragmata
(11 0-130 /-lm) and small acanthostrongyles were located at the
Synonymy base (75 X 12 /-lm) (Levi & Vacelet, 1958). The type slide examined
at MNHN (DCL741L) has all the spicules originally described; but
Cerbaris Topsent 1898b: 247; Topsent 1904b: 160. it also contains long styles and a mixture of foreign spicules some
Rhabdoploca Topsent, 1904b: 157 (new synonym). Bubaropsis of which could be assigned to haplosclerid genera such as Petrosia.
Levi & Vacelet, 1958: 235 (new synonym). Rhabdobaris Pulitzer- A second species, Bubaropsis alborani Boury-Esnault et al.,
Finali, 1983: 526 (new synonym). 1994b: 82, has been recently described. The species differs from
B. curvisclera in the absence of raphides, trichodragmata and
Type species toxiform oxeas, and in the size of the acanthostrongyles.
Rhabdobaris Pulitzer-Fin ali, 1983: 526 (type species
Cerbaris torquatus Topsent, 1898b: 248 (by monotypy). Rhabdobaris implicata, by monotypy) was erected for a very small
specimen (18 mrn high X 2 mrn diameter) with a spicule composi-
Definition tion similar to species included previously in Rhabdoploca and
Bubaropsis (i.e., styles-rhabdostyles in two size categories, oxeas
Bubaridae with evenly or unevenly acanthose diactines in the and strongyles sinuous or centrotylote, smooth or acanthose and
basal skeleton. raphides). The synonym of Rhabdobaris with Cerbaris is estab-
lished here with hesitation. The habit of the type species was
Diagnosis described as stipitate, which is not typical of Bubaridae, and there

Encrusting. Surface hispid. Choanosomal skeleton formed by

a basal layer of acanthose and smooth diactines with monactines

rt4i V"'f
projecting perpendicularly to substrata. Megascleres in the basal
skeleton are evenly or unevenly acanthose ceroxas, i.e., m-shaped A

~~
oxeas, toxostrongyles or strongy1es; smooth oxeas or strongyles .l \ ;:s:) iJ
are also present. Styles, subtylostyles or rhabdostyles, sometimes
in two size categories located perpendicularly to the basal layer. .J U ' ~,

Raphides may be present.

f!?~1P~
Remarks
B
The definition of Cerbaris has been expanded here to include
bubarid-like species that share the presence of acanthose diactines
in the basal skeleton and that were previously assigned to
Rhabdoploca Topsent, 1904b, Bubaropsis Levi & Vacelet, 1958
and Rhabdobaris Pulitzer-Finali, 1983.
Rhabdoploca Topsent, 1904b: 157 (type species Microciona
curvispiculifera Carter, 1880b: 43) is considered here a junior syn-
onym of Cerbaris. The type material is confirmed destroyed during
an air raid in 1941 (S. Stone, pers. com.), but the species is recog-
nisable from Carter's description who described it as: encrusting
with surface hispid, colour cream, basal skeleton of toxostrongyles
(i.e., strongyles bent in the centre) smooth or fully spined; perpen-
dicular styles, smooth, in two size categories, with bases fixed
in the basal layer (Fig. 3B). Topsent (l904b) initially erected
Rhabdoploca in Ectyonidae and transferred it later to Bubaridae
(Topsent, 1928c). As indicated by Topsent (1928c), the genus
shares the presence of a basal layer of spicules with perpendicular
monactine spicules with other members of Bubaridae. Hooper
(1996a: 578), suggested that the type species of Rhabdoploca
might be a Clathria (Clathria) but this is less likely. The generic ,
assignment of another species previously assigned to Rhabdoploca \ .
(i.e., R. topsenti Hentschel 1912: 43; Vacelet & Vasseur 1965: 21)
needs to be re-evaluated.
Fig. 3. A, styles and ceroxas found in Cerbaris torquatus Topsent 1898b
Bubaropsis Levi & Vacelet, 1958 (type species Bubaropsis
(reproduced from Topsent, 1904b, pI. XIII fig . 18). B, styles and tox-
curvisclera Levi & Vacelet, 1958: 235, by original designation) ostrongyles found in C. curvispiculifera (reproduced from Carter, 1880b,
agrees also with the definition of Cerbaris given above. It was pI. IV fig. 6).
752 Porifera' Demospongiae • Halichondrida • Bubaridae

is no indication in the original description or in the extant material H. papillosa). We reject this proposal and suggest that such species
(a spicule slide) on the type of skeletal organisation. are better placed in Axinyssa (see Family Halichondriidae, this
volume).
Distribution Uplexoa de Laubenfels, 1936a: 131 (type species Bubaris
oxeata Dendy, 1924: 349, by original designation) is referred here
East Atlantic, Mediterranean, Gulf of Manaar, 121-599 m depth. to Hymerhabdia, as suggested previously by Bergquist (1970). The
type material of Uplexoa (BMNH 23.10.1.124, slide, Terra Nova'
Description of type species Expedition) was examined and shown to be an encrusting speci-
men with a thin dermal membrane. The skeleton is formed by
Cerbaris torquatus Topsent, 1898b (Fig. 3A). styles (250-1000 X 20---45 J.tm) projecting perpendicularly to the
Synonymy. Cerbaris torquatus Topsent, 1898b: 248; surface with bases embedded in a basal layer of angulate oxeas
Topsent, 1904b: 160. (180-310 X 5-10 J.tm) (Fig. 5). Smaller and less common rhab-
Material examined. Holotype: MOM (?) - 1897; Azores; dostyles, some located in the basal layer and some projecting at an
station 866, 599m (not found). Slide from the holotype: MNHN acute angle ('echinating'), are also present. The genus was erected
LBIMDT943. in Axinellidae by de Laubenfels (1936a) to include Bubaris oxeata
Description. Encrusting. Surface hispid. Colour grey. and B. elegans Dendy, 1924, two species that share the presence of
Skeleton formed by long subtylostyles (1500-2000 J.tm), smooth bent oxeas. The description of B. elegans given by Dendy (1924:
with bases slightly constricted, embedded in a basal layer of 350) is not in agreement with the concept of Bubaridae adopted
'ceroxas' and projecting perpendicularly through surface. Ceroxas here. This species is clearly related to Axinellidae and should be
unevenly acanthose or smooth with two parallel conical branches referred to Axinella.
(120 J.tm long, 17 J.tm wide) connected transversely, sometimes Other species assigned to Hymerhabdia are H. contracta
forming a spiral; spines abundant at terminal ends. Sara & Siribelli, 1960: 30, H. diversicolor Carballo, 2000: 580;
H. intermedia Sara & Siribelli, 1960: 48; H. oxytrunca Topsent,
1904b; H. papillosa Sara & Siribelli, 1960: 31, H. pori Tsurnamal,
HYMERHABDIA TOPSENT, 1892 1969: 346; H. reichi Tsurnamal, 1969: 348, all from the
Mediterranean and H. topsenti Levi, 1952: 56 from Senegal.
Synonymy The genus has been previously referred to Monaxonida:
Ectyoninae (Topsent, 1892b); Halichondrina: Poeciloscleridae: sub-
Hymerhabdia Topsent, 1892b: 25; Topsent, 1894c: 23; family Bubarinae (Topsent, 1894c), Halichondrina: Axinellidae
Topsent, 1904b: 159; Topsent, 1928c: 42; de Laubenfels, 1936a: (Topsent, 1896a), Halichondrina: Bubaridae (Topsent, 1928c),
151; Bergquist, 1970: 20. Uplexoa de Laubenfels, 1936a: 131. Hadromerina: Suberitidae (de Laubenfels, 1936a) and
Halichondrida: Axinellidae (Van Soest et al., 1990; Carballo, 2000).
Type species
Distribution
Hymerhabdia typica Topsent, 1892b: 26 (by monotypy).
East Atlantic, Mediterranean, North Africa.
Definition
Description of type species
Bubaridae with a basal skeleton of interlacing rhabdostyles
and/or bent angulate oxeas. Hymerhabdia typica Topsent, 1892b (Fig. 4).
Synonymy. Hymerhabdia typica Topsent, 1892b: 26; Topsent,
Diagnosis 1894c: 24; Topsent, 1934a: 38; Pulitzer-Finali, 1977: 37.
Material examined. Holotype: MNHN LBIM DT3075
Encrusting. Surface hispid. Choanosoma1 skeleton formed by (not found). Slide made from holotype MNHN DT2359 - Banyu1s.
individual or bundles of long styloid spicules with heads embedded Description. Encrusting. Surface hispid. Skeleton formed
in a basal layer of smaller rhabdostyles and angulate or bent oxeas. by a basal layer of interlacing rhabdostyles with long tylostyles and
Megascleres are styles, subtylostyles and tylostyles, smooth, styles projecting perpendicularly to substrata and with bases
straight or slightly bent; rhabdostyles, some with bases contorted embedded in the layer. Styles and tylostyles, smooth, straight or
or reflexed, or oxeas angulate, centrotylote or irregularly bent. slightly bent, sometimes with prominent bases, 650-800 X 10 J.tm.
Microscleres absent. Rhabdostyles, some with bases contorted or reflexed, 80-120 X
8-IOJ.tm.
Remarks Remarks. Hymerhabdia oxytrunca Topsent, 1904b: 15 is
considered a junior synonym of H. typica by Carballo (2000),
Hymerhabdia has been related to other bubarid-like genera. based on the similarities of types and dimensions of spicules. Both
The skeletal architecture is similar to Bubaris, but spicules located species bear, in particular, rhabdostyles and centrotylote oxeas. The
in the basal layer are rhabdostyles and angulate oxeas instead of presence of centrotylote oxeas were also recorded in H. typica by
sinuous or vermiform strongyles and anisoxeas, and there are Topsent (1934a). It seems, however, that this might be a variable
tylostyles in addition to the styles, projecting perpendicularly to the character within the species as it has not been recorded in other
substratum. descriptions.
Carballo (2000) expanded the definition of Hymerhabdia Habitat and distribution. East Atlantic and Mediterranean;
to include species with fistules (i.e., H. diversicolor and 15-120m.
Porifera· Demospongiae • Halichondrida • Bubaridae 753

Fig. 4. Hymerhabdia typica (Topsent). A, photo in situ, Ireland (photo B. Picton). B, diagram of skeleton of Hymerhabdia typica as observed in the slide
made from holotype (MNHN DT2359). C, drawing of spicules (reproduced from Topsent, 1904b, pI. XIII, fig . 16).

B "

c
'. '.:

Fig. 5. Hymerhabdia oxeata (Dendy). A ,BMNH 23.10.1.124, slide (labelled as 'type'); B, style. C, oxea (scale 100 fLm).

MONOCREPIDIUM TOPSENT, 1898 Diagnosis

Synonymy Encrusting. Surface hispid. Choanosomal skeleton fonned by

styloids single or in bundles, projecting perpendicularly to the sub-
Monocrepidium Topsent, 1898b: 229; Topsent, 1904b: 137, strata and through the ectosome with heads embedded in a basal
147; Topsent, 1928c: 28; de Laubenfels, 1936a: 103; Vacelet, 1969: layer of vermicular diactines some of which are tuberculate or
183. Skeizia Cabioch, 1968b: 220; Van Soest et al., 2000. annulate. Megascleres are long smooth styles, slightly curved near
the bases, subtylostyles or tylostyles. Diactines of the basal skele-
Type species ton are strongyles or strongyloxeas, stout, vermiculate, tuberculate
or annulate.
Monocrepidium vermiculatum Topsent, 1898b: 229 (by
original designation). Remarks

Definition Monocrepidium has skeletal architecture typical of Bubaridae.

It was initially referred to Desmanthidae by Topsent (1898b) based
Bubaridae with basal skeleton that includes stout diactines, on the resemblance of the tuberculate diactines with monocrepid
smooth tuberculate or annulate. desmas. That view was later rejected by Topsent (1904b) based on
754 Porifera· Demospongiae • Halichondrida • Bubaridae

B h
?\

~JfRfij
'?'i
Fig. 6. Monocrepidium vermiculatum Topsent. A, SEM of the surface and basal layer of tuberculate spicules (photo J. Vacelet; scale 100 fLm). B, drawing
of spicules (reproduced from Topsent, 1904b, pI. XIII fig. 9).

the presence of an axial canal observed in the diactines of both Distribution. East Atlantic, Mediterranean and Indian Ocean.
Bubaris and Monocrepidium, which he considered to be homolo-
gous. The genus is related mainly to Bubaris and Hymerhabdia, Description of type species
both of which have smooth diactine spicules in the basal skeleton.
Hymeraphia eruca Carter, 1880b: 46 was referred to Monocrepidium vermiculatum Topsent, 1898b (Fig. 6).
Monocrepidium by Topsent (1928c: 41) and to Laonoenia Hallmann, Synonymy. Monocrepidium vermiculatum Topsent, 1898b:
1917c: 675 by de Laubenfels (1936a:I44). The description of Carter 229; Topsent, 1904b: 148; Topsent, 1928c: 41; Vacelet, 1969: 183;
agrees with the definition of Monocrepidium adopted here, and there- Pulitzer-Finali, 1977: 36; Pulitzer-Finali, 1983: 526.
fore this species is the second representative of the genus. Material examined. Holotype (fragment): MNHN LBIM
Skeizia (type species Skeizia minuta Cabioch, 1968b: 220; DT 887, slide.
by original designation) is here considered to be a synonym Description. Encrusting. surface hispid. Colour, grey.
of Monocrepidium. The type species has all the features of Skeleton formed by a basal layer of interlacing diactines with
Monocrepidium but differs in having tylostyles instead of styles. styles embedded by their bases in the layer and projecting perpen-
We consider that this is not a valid reason to erect a new monotypic dicular to the substrate (Fig. 6A). Styles or subtylostyles smooth,
genus, and consequently, we have expanded the definition of slightly bent near the base (170-1400 X 5-25 /-lm). Strongyles and
Monocrepidium here to include species with tylostyles. strongyloxeas, thick (15/-lm), sinuous, with tubercles roughly
The genus has been previously referred to Lithistida: arranged in a spiral, located in the basal skeleton. Smooth, thinner
Desmanthidae (Topsent, 1898b), Halichondrina: Axinellidae (Topsent, and sinuous oxeas also present (Fig. 6B).
1904b), Halichondrina: Bubaridae (Topsent, 1928c), Myxilliformes: Habitat and distribution. The species seems to be restricted
Arnphibleptulidae (de Laubenfels, 1936a), Axinellida: Bubaridae to deep waters (121-600m depth) across the Mediterranean and
(Levi, 1973) and Halichondrida: Axinellidae (Van Soest et aI., 1990). East Atlantic.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcaderniclPlenurn Publishers, New York, 2002

Family Desmoxyidae Hallmann, 1917

John N.A. Hooper

Queensland Museum, P.O. Box 3300, South Brisbane, Qld, 4101, Australia. (JohnH@qm.qld.gov.au)

Desmoxyidae Hallmann (Demospongiae, Halichondrida) contains 19 genera, 10 of which are recognised here, and probably less than
100 species worldwide. Desmoxyids are widely distributed and live mainly in shallow coastal waters but a few have also been recorded
from 1300m depth. Major morphological characters diagnostic for the family include spined or smooth diactinal microscleres (oxeas or
strongyles), often centrangulate or strongly bent centrally, localised in the ectosomal region; choanosomal skeleton with monactinal
(styles or tylostyles) and/or diactinal spicules forming confused reticulate tracts or widely spaced, irregular muitispicular bundles bound
together by sparse collagen, or enclosed in poorly developed fibres; microscleres also sometimes include raphides occurring singly or in
bundles (trichodragmata), or acanthose microrhabds (sanidaster-like), acanthose cladotoxa and birotules.
Keywords: Porifera; Demospongiae; Halichondrida; Desmoxyidae; Acanthoclada; Didiscus; Halicnemia; Heteroxya; Higginsia; lulavis;
Microxistyla; Myrmekioderma; Negombo; Parahigginsia.
--------------

DEFINITION, DIAGNOSIS, SCOPE them were not always completely clear. Topsent (1928c) subse-
quently removed most of these spurious genera to Hemiasterellidae,
Synonymy but Wiedenmayer (1977b) suggested that although some were hemi-
asterellids, others were more appropriately included in Hadromerida
Heteroxyinae Dendy, 1905 (nomen oblitum). Desmoxyinae (e.g., Paratimea Hallmann) or Astrophorida (e.g., Holoxea Topsent,
Hallmann, 1917c. Desmoxyidae Bergquist, 1970. Higginsiinae de 1892a). Hooper & Levi (1993b) discuss the affinities of other genera
Laubenfels, 1936a. included in Desmoxyidae at one time or another.
Several desmoxyid genera have received recent attention
DefInition in isolation, although prior to the present work there has been
no comprehensive synthesis or revision of the entire family.
Halichondrida with smooth or spined, usually centran- Contemporary reviews include: Bergquist (1965) and Hooper &
gulate microxea microscleres or modified forms forming an ectoso- Levi (1993b) for Myrmekioderma and Higginsia; Van Soest et al.
mal skeleton. (1990), Hiemstra & Van Soest (1991) and Diaz et al. (1993) for
Myrmekioderma and Didiscus; Pansini & Pesce (1998) for
Diagnosis Higginsia; and Van Soest & Lehnert (1997) for Julavis, also
providing the most recent overview of the family.
Encrusting, massive or ramose sponges often with hispid Together these reviews have moved closer towards a more
or conulose surfaces bearing smooth or spined microxea micro- homogeneous definition and scope of Desmoxyidae, although
scleres, often centrangulate or strongly bent centrally, sometimes there is still controversy as to the true affinities of Myrmekioderma
acanthose and rhabdose, usually found localised on the ectosome, and Didiscus.
sometimes also with raphides occurring singly or in bundles Bergquist (1965) provides a good historical overview on the
(trichodragmata), or acanthose cladotoxa and birotules in one chequered history of taxonomic placement of Myrmekioderma,
genus; megascleres monactinal (styles or tylostyles), diactinal including: subfamily Heteroxyinae (Dendy, 1905) in the family
(oxeas, strongyles) or both; choanosomal skeleton a confused Haploscleridae; section Heteroxyeae, subfamily Axinellidae,
arrangement of single spicules or widely spaced reticulate bundles family Desmacidonidae (Dendy, 1922b); family Jaspidae,
of multispicular fibres, with little spongin, with poorly developed order Epipolasida (Wilson, 1925); family Heteroxyidae (Topsent,
or no axial compression, and a relatively poorly differentiated 1928c); family Phorbasidae (de Laubenfels, 1936a); section
extra-axial skeleton (disorganised-plumose). Raspelieae, family Axinellidae (Burton, 1937).
Of contemporary authors, Hooper & Levi (1993b) included
Scope Myrmekioderma in Desmoxyidae, following Bergquist (1965),
based on its morphological similarities with Higginsia (e.g., spined
Of 19 nominal genera only 10 are considered to be valid and ectosomal spicules), with empirical support from chemotaxonomic
presently included in Desmoxyidae. studies of both genera and outgroups. Conversely, Van Soest et al.
(1990), Hiemstra & Van Soest (1991) and Diaz et al. (1993) referred
History and biology Myrmekioderma to Halichondriidae based on the halichondroid
nature of its more-or-Iess confused choanosomal skeleton, posses-
The family was established initially for five genera, of which sion of a detachable ectosomal crust, and the fact that the acanthose
only three were correctly assigned (Desmoxya, Higginsia and nature of the ectosomal spicules was not as prominent for the genus
Halicnemia). Many other genera were subsequently added or associ- as previous authors had indicated. Didiscus was also included in the
ated with the family by various authors, although affinities between Halichondriidae based on its very close morphological resemblance

755
756 Porifera' Demospongiae • Halichondrida • Desmoxyidae

to Myrmekioderma (external features, skeletal architecture). spicules, and closer affinities to other halichondrids. On that basis
However, Van Soest & Lehnert (1997) subsequently agreed that it could also be argued that several genera currently assigned to
both Myrmekioderma and Didiscus should be returned to the Bubaridae (e.g., Cerbaris and junior synonyms Rhabdoploca,
Desmoxyidae, indicating further that lulavis represented a close Bubaropsis) also have inferred affinities with the Desmoxyidae,
link between Higginsia and Myrmekioderma, possessing both having acanthose or smooth oxeote megascleres, but these also have
coarsely spined diactinal surface spicules (like Higginsia) and quite different skeletal structures and megasclere geometries (mon-
wispy trichodragmata (like Myrmekioderma). actinal spicules with heads embedded in a basal layer of interlacing
Of other genera, Hooper & Levi (1993b) retained Holoxea vermiform diactinal spicules). The presence of smooth or spined
Topsent in the Desmoxyidae following Hallmann (1917c), which oxeas, generally associated with the ectosomal skeleton, is charac-
had sanidaster-like microxeas associated with the ectosome, in teristic of, and apomorphic for, desmoxyid genera. These spicules
addition to raphides in trichodragmata. However, re-examination are traditionally classed as microscleres (e.g., Hallmann, 1917c;
of the type species (H. furtiva Topsent, 1892a; MNHN DT2414) Wiedenmayer, 1977b), although in comparison to many other taxa
confirms that these are not true microxeas but more similar to the they are generally much too large to be considered as such.
spiny microrhabds seen in Melophlus Thiele, and the genus is more Negombo Dendy is transferred to Desmoxyidae in this
appropriately referred to Ancorinidae (Astrophorida) (see chapter work following Kelly (pers. comm.), with suspected affinities
by Uriz, this volume). to Didiscus, although this relationship is not completely clear
and requires further corroboratory evidence from other sources
Differences with similar families (e.g., molecular data).

Although Topsent (1928c) implied close affinities between Previous reviews

desmoxyids and hemiasterellids, more recent concepts of the family
(e.g., Hooper, 1986; Voultsiadou-Koukoura & Van Soest, 1991a) Bergquist (1965), Wiedenmayer (1977b), Van Soest et al.
restrict the latter to forms with asterose microscleres (in (1990), Hiemstra & Van Soest (1991), Diaz et al. (1993), Hooper &
Hadromerida), whereas desmoxyids have only microxeote Levi (1993b), Van Soest & Lehnert (1997), Pansini & Pesce (1998).

KEY TO GENERA

(1) Microscleres (acantho-)microxeas ...................................................................................................................................................... 2

Microscleres discorhabd-like microxeas ................................................................................................................................. Didiscus
Microscleres sanidaster-like acanthose microrhabds ............................................................................................................. Negombo
Microscleres raphides ....... ....... .... ................ ........... ................. ............... ................ ................ ................ ............ ..................... ... ... ...... 4
Microscleres acanthose cladotoxas and birotules .... .......................... ...... .......... ...... ................ .......... ........... .................. Acanthoclada
Microscleres absent .............................................................................................................................................................. Heteroxya
(2) Cortex absent, without microscleres or with microscleres scattered but not confined to the surface ................................... Higginsia
Cortex moderate with scattered microscleres concentrated near surface ....................................................................... Parahigginsia
Cortex well-developed, with an erect palisade of microscleres perpendicular to the surface ............................................................. 3
(3) Extra-axial megascleres centrangulate oxeas ..................................................................................................................... Halicnemia
Extra-axial megascleres absent .... ...... .......... ........... ........... .......... ...... .......... ............ ................ ................ ........... .... ...... .... Microxistyla
(4) Ectosomal megascleres acanthose oxeas ..... ........... ........... ................. ......... ....... .......... ...... ... ............ ................. ........ Myrmekioderma
Ectosomal megascleres raphidiform oxeas .. ,... ....... ........... ..... ...... ...... ............. ............. ...... ... ....... ..... ................. ........... ........ Higginsia
Ectosomal megascleres (acantho-)strongyles ............................................................................................................................. lulavis

ACANTHOCLADA BERGQUIST, 1970 Diagnosis

Synonymy Thickly encrusting; choanosomal skeleton lax, fibrous, con-

sisting of ascending fibres and tracts reinforced with small quanti-
Acanthoclada Bergquist, 1970: 22. ties of collagen, without distinctive axis; basal skeleton composed
of tangled bundles of spicules containing rhabdostyles in plumose
Type species or hymedesmioid tracts erect on the substrate bases of larger
choanosomal styles embedded in these bundles, and abundant
Acanthoclada prostrata Bergquist, 1970 (by monotypy). microscleres lying on basal spongin; ascending fibres cored by
smooth styles forming multi spicular tracts, with smooth rhabdo-
Definition styles scattered within and protruding from fibres. Long styles also
protrude through fibres and abundant microscleres are scattered
Desmoxyidae with a plumose or hymedesmioid basal skeleton throughout the mesohyl producing a lax halichondroid skeleton.
composed of rhabdostyles and bases of choanosomal styles perpendi- Subectosomal fibres terminate in brushes of long centrangulate
cular to the substrate; choanosomal skeleton lax, composed of oxeas which form conulose surface projections. Ectosome is also
ascending tracts of long smooth styles extending into the periphery, packed with acanthose microscleres. Megascleres include smooth
terminating in brushes of long centrangulate oxeas; microscleres styles, rhabdostyles and oxeas which are usually centrangulate or
acanthose cladotoxas and birotules forming a dense ectosomal cortex. toxiform. Microscleres acanthose cladotoxas and birotules.
Porifera' Demospongiae • Halichondrida • Desmoxyidae 757

IA CI
G

Fig. 1. Acanthoclada prostrata Bergquist (holotype). A, base and point of choanosomal style (scale 100 JLm). B, centrangulate oxea (scale 200 JLm).
C, enlarged apex of oxea (scale 100 JLm). D, rhabdostyles (scale 50 JLm). E, birotules (scale 15 JLm). F, cladotoxas (scale 15 JLm). G, idealised reconstruction
of skeleton. H, ectosomal skeleton (scale 200 JLm). J, subectosomal skeletal tracts (scale 200 JLm). J, basal skeleton (scale 200 JLm). K, holotype (from
Bergquist, 1970: pI. 5B).

Description of type species conules) or forming paratangential tracts near the surface, and the
surface is also densely packed with spined birotule-like microscle-
Acanthoclada prostrata Bergquist, 1970 (Fig. 1). res (52-72 X 3-4 11m) and 'cladotoxas' (80-96 X 5-6 11m);
Synonymy. Acanthoclada prostrata Bergquist, 1970: 22. choanosomal skeleton with two more-or-less distinct components:
Material examined. Holotype: NMNZ Por.27 - Takatu basal skeleton consists of entirely smooth rhabdostyles with a
Point, New Zealand. prominent basal rhabd (213-600 X 6-8 11m) forming dense
Description. Thick, slimy, encrusting on a bivalve, firm tex- plumose or hymedesmioid bundles erect on the the substrate, and
ture, granular surface raised into conules; ectosomal skeleton con- also with the bases of choanosomal styles embedded, usually in
sists of bundles of centrangulate oxeas (960-1320 X 1-9 11m) bundles, within this tangled basal spicular network and standing
embedded in the ends of choanosomal skeletal tracts, protruding perpendicular to the substrate; ascending component of skeleton
through the surface in erect bundles (mainly at the ends of surface consists of thick, more-or-less discrete multispicuiar tracts coring
758 Porifera· Demospongiae • Halichondrida • Desmoxyidae

light spongin fibres composed of long straight or slightly curved Type species
styles with evenly rounded, slightly subtylote, or occasionally
subterminal tylote swellings (677-1850 X 9-18 fJ.m) standing per- Didiscus ptacospongioides Dendy, 1922b (by monotypy).
pendicular to the substrate and ascending to, but not protruding
through the surface; longer, more curved styles also project out Definition
from skeletal tracts and tracts are also sparsely 'echinated' with
rhabdostyles (although the latter are mostly concentrated at the Desmoxyidae with ectosomal crust of 'discorhabd' -like
base of the skeleton); mesohyl also contains individual styles, microxea microscleres.
bundles of oxeas erect or running transversely through the
skeleton, and abundant microscleres are scattered throughout Diagnosis
the skeleton together producing a lax halichondroid skeletal
structure. Massive-amorphous to lobate growth forms; spicules strewn in
Remarks. Bergquist (1970) assigned Acanthoclada to confusion and sporadically grouped in spongin-reinforced direction-
Desmoxyidae because the overall skeletal pattern was similar to less tracts; ectosomal skeleton is a gradation from a perpendicular
Higginsia (lax and lacking any axial compression), differing from palisade to a tangential or paratangential arrangement of spicules with
the latter in having echinating rhabdostyles and a special category 'discorhabd' -like microscleres arranged perpendicular to the surface;
of microscleres termed 'cladotoxas' (curved rhabds with large ter- surface with sculptured grooves and subdermal drainage canals.
minal clads and mostly smooth shafts, or with few larger hook-like
spines at the centre) somewhat resembling the cladotylotes of Description of type species
Acamus (Acamidae, Poecilosclerida), and 'birotules' (of similar
shape but with evenly spined shaft and small terminal hooks). Didiscus ptacospongioides Dendy, 1922b (Fig. 2).
These microscleres appear to be unique amongst the demosponges Synonymy. Didiscus ptacospongioides Dendy, 1922b: 135.
and as such they are of uncertain derivation and affinity within Material examined. Holotype: BMNH 1921.11.7.112 -
Desmoxyidae. Unlike the cladotylotes of Acamus, which comprise Cargados Carajos, W Indian Ocean.
a special category of megasclere, these cladotoxas in Acanthoclada Description. Massively encrusting growth form; surface
are much smaller relative to its style and oxea megascleres and strongly grooved with angular striations forming polygonal plates;
clearly represent a special category of microsclere (and thus the plates contractile with oscula in between; ectosomal skeleton a thick
affinities between these two genera appear to be merely superfi- tough crust composed of generally smaller oxeas forming discrete
cial). Bergquist (1970) recorded that rhabdostyles were 'echinat- bundles in a continuous palisade, erect, paratangential or tangential
ing', but this may be a slightly misleading comparison with to the surface; discorhabd microscleres generally arranged perpen-
raspailiids and microcionids. Certainly in the basal part of the dicular to the surface, and concentrated near the surface singly or in
skeleton rhabdostyles form plumose or hymedesmioid brushes thick tracts; choanosomal skeleton with confused radial tracts of
erect on ('echinating') the surface, whereas they do not appear to generally larger oxeas, the largest at the centre running mainly lon-
'echinate' the ascending choanosomal spicule tracts but appear to gitudinally, with medium and smaller sized oxeas becoming
be scattered, embedded or protrude from these tracts at all angles increasingly transverse, eventually carrying the ectosomal crust;
and directions. Consequently, the term 'echinating' is not used spongin moderately developed; megascleres are oxeas more-or-less
in this context. in two size categories, the smaller sometimes centrotylote, and occa-
This genus is somewhat enigmatic within Desmoxyidae (and sionally with tylote modifications (400-1400 X 11-20 fJ.m);
indeed Halichondrida). Microsclere morphology and derivation is microscleres rugose or finely spined microxeas, sharply pointed
not typical of other desmoxyids, although the birotules are vaguely with 'frayed' or ragged points, possessing two unequally sized discs
rhabd-like and are conceivably derived from the rhabd-like microx- asymmetrically aligned along the shaft, superficially resembling
eas typical of the family. The possession of rhabdostyles is discorhabds of the Latrunculiidae (70-90 X 2-5 fJ.m).
also unusual amongst halichondrids. Morphologically these Remarks. There is little doubt that Didiscus and
spicules are very similar to those rhabdose genera of Raspailiidae Myrmekioderma are closely related and belong to the same family,
(Autospongus, Raspailia (Raspaxilla)) , although the two groups despite the former previously being included with the
have few other affinities. The genus is retained in Desmoxyidae, Hadromerida. Part of the confusion regarding the family placement
following Bergquist (1970), being most similar to Higginsia based of Didiscus centred on the origin and nature of the discorhabd
largely on their affinities in skeletal structure, whereas this assign- microxeas, which up until the review of Van Soest et at. (1990)
ment is still not certain. were considered to be homologous with the true discorhabds of the
latrunculids (formerly Hadromerida, now Poecilosclerida). These
Distribution authors correctly noted that the lateral disks on microxeas of
Didiscus were merely highly modified and exaggerated spines on
Monotypic, and known only from New Zealand waters. centrotylote microxeas, not related to the streptaster/spiraster ori-
gin of true discorhabds. In those examples of microxeas that lacked
lateral disks, or where they were rudimentary, there are clear simi-
larities to those of Myrmekioderma, with tapering points, granular
DIDISCUS DENDY, 1922 spination and slightly centrangulated at their centres (Fig. 2C).
Further clues on its affinities can be seen in skeletal architec-
Synonymy ture, in which both genera have an ectosomal skeleton where
smaller oxeas are arranged tangentially, paratangentially or in erect
Didiscus Dendy, 1922b: 135; Van Soest et at., 1990: 33; brushes (discorhabd-like microxeas and smaller oxeas in Didiscus;
Hiemstra & Van Soest, 1991: 45; Diaz et at., 1993: 145. smaller oxeas in Myrmekioderma). This surface crust is carried by
Porifera' Demospongiae • Halichondrida • Desmoxyidae 759

B
A I
I
c
I

Fig. 2. Didiscus placospongioides Dendy (holotype). A. larger oxeas (scale 150 f.Lm). B, smaller oxeas (scale 100/-Lm). C, discorhabds (scale 25 /-Lm) .
D, idealised reconstruction of skeleton. E, ectosomal skeleton (scale 200 /-Lm). F, choanosomal skeleton (scale 200 /-Lm) . G, holotype (from Dendy. 1922b:
pI. 5 fig. 10).

irregular radiating tracts of larger spicules in both genera (Hiemstra HAUCNEMIA BOWERBANK, 1864
& Van Soest, 1991). Diaz et at. (1993) further remarked on the sim-
ilarities between the two genera, citing examples of two Caribbean Synonymy
species, Didiscus oxeata and Myrmekioderma styx, in which exter-
nal sculpturing of the surface and skeletal structure were so similar Halicnemia Bowerbank, 1864: 184, 1866: 96; Topsent,
that they were indistinguishable in the field without subsequent 1897b: 235; Hallmann, 1917c: 675; Topsent, 1928c: 39 (in part);
microscopic examination. Consequently, Didiscus differs substan- Descatoire, 1966: 239; Van Soest, 1987b: 24; Desqueyroux-
tially only from Myrmekioderma in possessing discorhabd-like Faundez & Van Soest, 1997: 429. Alloscleria Topsent, 1927: 6;
microxeas and lacking trichodragmata. Didiscus and 1928c: 160. Quindesmia de Laubenfels, 1950. (Taxonomic deci-
Myrmekioderma also consistently produce the same marine natural sion for synonymy: Hooper, this work.)
products chemistry (M.K. Harper, pers. comm.).

Distribution
Type species
Tropical, subtropical; Brazil, Caribbean, Mediterranean, NE
Atlantic, W Indian Ocean, Indonesia. Halicnemia patera Bowerbank, 1864: 184 (by monotypy).
760 Porifera· Demospongiae • Halichondrida • Desmoxyidae

D
I
A Be

Fig. 3. Halicnemia patera Bowerbank (syntypes). A, larger tylostyle (scale 400 fLm). B, smaller tylostyle (scale 200 fLm). C, centrotylote oxea (scale
400 fLm). D, acanthoxeas (scale 50 fLm). E, peripheral skeleton (scale 100 fLm). F, choanosomal skeleton (scale 250 fLm). G, spicule fringe around margin
of disk (scale 250 fLm). H, idealised reconstruction of skeleton. I, syntype (from Bowerbank, 1874b: pI. 15 fig. 31).

Definition type species) and protruding choanosomal megascleres; megascleres

of one or two categories, styles/tylostyles with or without polytylote
Desmoxyidae with acanthose centrangulate microxeas forming smooth oxeas; microscleres are centrangulate spined microxeas, usu-
erect ectosomal brushes. ally with abrupt angular central curve, sometimes straight.

Diagnosis Description of type species

Encrusting or disk-shaped growth forms; surface smooth or Halicnemia patera Bowerbank, 1862b (Fig. 3).
minutely conulose and usually very hispid; choanosome reduced to Synonymy. Halicnemia patera Bowerbank, 1862b, pI. 10,
a compressed basal layer of paratangential tracts on the substrate pI. 32; 1864: 184; 1874b, pI. 15. Crella inflata Arndt, 1935: 76.
including a halichondroid reticulation of styles with or without Bubaris gallica Topsent, 1893c: 35.
polytylote smooth oxeas; styles/tylostyles embedded and erect on Material examined. Syntypes: BMNH 1910.1.1.2459-60,
the basal layer, protruding through and tangential to the surface; 1930.7.3.25 - Shetland Is, N Atlantic.
specialised ectosomal spicules consist of erect brushes of acan- Description. Thinly encrusting; surface conulose, fibrous,
those, centrangulate or straight microxeas (best developed in the with margins prominently hispid and shaggy from protruding
Porifera· Demospongiae • Halichondrida • Desmoxyidae 761

longer tylostyle megascleres; choanosomal skeleton a basal layer in A. tenuispinosa. This species has long smooth style megas-
of scattered tylostyles arranged paratangential to the substrate, long cleres (lacking a swollen basal tyle as characteristic of the
centrotylote oxeas forming dense bundles ascending to surface, type species) and two categories of centrotylote acanthoxea
and a dense mass of small acanthoxeas forming a crust on the microscleres - the smaller being more heavily spined of the two.
surface and subectosomal skeletons; ectosomal skeletons with Its acanthoxeas do not form a prominently erect surface, as in
paratangential crust of microxeas and longer tylostyles protruding the type species, but are reportedly paratangential and scat-
through the surface; megascleres include a long tylostyles with tered throughout the mesohyl, and its choanosomal skeleton con-
a pronounced basal swelling and constriction, entirely smooth, sists of parallel tracts of styles forming a halichondroid skeleton.
fusiform points (1400-2750 X 35-55 /-Lm), smaller tylostyles of Given these features it is perplexing why Topsent (1928c) did not
similar morphology (255-790 X 15-32 /-Lm), and long, thin, centrally associate the genus as having affinities to Halicnemia and
bent and centrotylote oxeas (1200--1750 X 10--15/-Lm); microscleres Higginsia.
are acanthoxeas with centrotylote swelling, straight or with a central Quindesmia de Laubenfels, 1950 (type Hymedesmia inflata
bend, spines range from vestigial to prominent, mostly erect Bowerbank, 1866) is an objective synonym of Halicnemia, as its
(125-215 X 6-12 /-Lm). type is ajunior synonym of H. patera (Rob Van Soest, pers. comm.,
Remarks. There is some confusion surrounding the genus from re-examination of a slide of the type in the BMNH).
Halicnemia, arising from Topsent's (1928c: 39) revision of species.
Topsent included a diverse assemblage of taxa, including Distribution
hadromerids, hemiasterellids and true desmoxyids into the genus.
Aster-bearing species (e.g., Bubaris constellata Topsent), and North Atlantic (Shetland Islands, Ireland, France,
those with acanthose or smooth microxeas (e.g., H. patera Mediterranean), southeast Pacific (Galapagos Islands, S Chile) and
Bowerbank, H. gallica (Topsent». Halicnemia (sensu Topsent) do NE Pacific (British Columbia).
not form a natural assemblage of taxa, contrary to his (1928c: 39)
strong opinions otherwise, although the presence of certain shared
characters seems to cut across systematic boundaries (e.g., posses-
sion of microxea microscleres). Nevertheless, aster-bearing species HETEROXYA TOPSENT, 1898
are referred to Stelligera Gray (family Hemiasterellidae; e.g.,
Hooper, 1986), those with asters and microxea are probably Synonymy
hadromerids (e.g., Diplastrella, Spirastrella), whereas Halicnemia
has only acanthose ectosomal microxeas, very similar in geometry Heteroxya Topsent, 1898b: 231; Topsent, 1904b: 133; de
to those seen in Higginsia. Laubenfels, 1936a: 72; Bergquist, 1965: 182.
Following the removal of most of Topsent's (1928c) species
included in Halicnemia, leaving only the type species in it, Type species
Desqueyroux-Faundez & Van Soest (1997) subsequently added
several other species. They also briefly review the genus and Heteroxya corticata Topsent, 1898b (by monotypy).
included in it a new species from the Galapagos Is, SE Pacific, and
Higginsia papillosa Thiele, 1905 (from S Chile). They also noted Definition
that several species currently included in Higginsia may eventually
be transferred to Halicnemia. Van Soest (1987b) also remarked on Desmoxyidae lacking microscleres but with two sizes of oxea
this close similarity, suggesting that the only significant differences megascleres both with apical spination, the smaller forming a
relate mainly to reticulate versus radial skeletons in Higginsia nearly continuous palisade perpendicular to the ectosome and the
and Halicnemia, respectively. This argument was again raised by larger forming both a halichondroid choanosomal skeleton and
Desqueyroux-Faundez & Van Soest (1997), citing observed simi- protruding through the ectosome.
larities between a Galapagos species of Halicnemia and other
Higginsia, as evidence for a close relationship between the genera, Diagnosis
and between Desmoxyidae and some poecilosclerids (Raspailiidae,
Rhabderemiidae). They suggested that the similarity of the angu- Thickly encrusting growth form; surface lobate, folded to the
lated acanthoxeas of both genera was supported by a grade in contours of the substrate, highly hispid; choanosome with a con-
skeletal architecture halfway between a strictly Eurypon-like archi- densed basal layer of spongin lying on the substrate, without a
tecture in Halicnemia patera and a strictly reticulate architecture in complete mineral skeletal structure, containing only acanthoxeas
Higginsia coraUoides. The Galapagos Halicnemia had long styles distributed without appreciable order on basal spongin and strewn
surrounded by shorter styles and centrotylote oxeas, also observed throughout the mesohyl; subectosomal skeleton consists of oxeas
in H. patera, plumose spiCUle bundles also seen in some Higginsia or tomotes perpendicular to the ectosome, protruding through the
and surface bouquets of special oxeas reminiscent of a raspailiid surface, but not embedded in basal spongin; ectosomal skeleton
surface structure. They suggested further that Desmoxyidae and with a perpendicular palisade of smaller acanthoxeas, through
Raspailiidae might be conceivably merged in future revisions - an which the larger subectosomal oxeas protrude; megascleres curved
action not presently contemplated. to sinuous smooth or spined hastate oxeas!tomotes, and curved
Alloscleria Topsent (type species A. tenuispinosa Topsent, acanthoxeas; microscleres absent.
1927, by original designation, schizotypes MNHN DT1190) is also
referred here to Halicnemia, with skeletal structure and spicule Description of type species
geometry similar to the type species but with some differences in
its ectosomal skeleton and absence of polytylote oxea megascleres Heteroxya corticata Topsent, 1898b (Fig. 4).
762 Porifera· Demospongiae • Halichondrida • Desmoxyidae

B
I
Fig. 4. Heteroxya corticata Topsent (slides from holotype). A,larger acanthoxea (scale 250 f,l.m). B, smaller acanthoxeas (scale 50 f,l.m) . C, apex of smaller
acanthoxea (scale 20 f,l.m) . D,enlarged view of cortical ectosomal bundle (scale 100 f,l.m) . E,cortical ectosomal bundles (scale 250 f,l.m) . F,idealised recon-
struction of skeleton. G, holotype (from Topsent, 1904b: pI. 1 fig . 15).

Synonymy. Heteroxya corticata Topsent, 1898b: 231 ; smooth) than the smaller (235--420 X 12-23 f,Lm), with spination
1904b: 133, pI. 1 fig.15. pI. 12 figs 22-23. predominantly at apices of spicules; microscleres absent.
Material examined. Holotype: MOM (not seen); slide made Remarks. Topsent (1898b, 1904b, 1928c) remarked on
from holotype - MNHN DT905, BMNH 1930.7.1.46 - Azores, the resemblance between H. corticata and Trachya (=Aaptos,
N Atlantic. Suberitidae), particularly in surface features, but in other respects
Description. Thinly encrusting plaque, hispid smface; their affinities are purely superficial. The higher systematic place-
choanosome nearly completely lacking skeletal structure, with only ment of Heteroxya has altered with each successive description in
acanthoxeas scattered indiscriminantly throughout the mesohyl; cor- the literature: Aciculida (Topsent, 1898b: 233), Tethyidae (Topsent,
tical ectosome well developed, thick, very spiculose, with two sizes of 1904b: 134), Axinellida (Topsent, 1928c: 189) and Coelosphaeridae
acanthoxeas, the larger extending into the choanosomal mesohyl inte- (de Laubenfels, 1936a: 72), but the genus appears to have closest
rior and protruding a long way through the smface, perpendicular to affinities with the Desmoxyidae. Van Soest et ai. (1990) included
it, and the smaller forming a nearly continuous perpendicular palisade Heteroxya as a synonym of Myrmekioderma, with question, based
on or just below the smface; megascleres two sizes of oxeas, both pre- on Topsent's description of H. corticata, but re-examination of the
dominantly acanthose, with pronounced central curvature, the larger holotype does not indicate such a close relationship as inferred by
(1600--2000 X 20--35 f,Lm) less spined (or sometimes completely Topsent. The genus is retained here in Desmoxyidae.
Porifera' Demospongiae • Halichondrida • Desmoxyidae 763

Distribution Synonymy. Higginsia coralloides Higgin, 1877: 291.

Material examined. Holotype: BMNH 1886.12.15.92 -
N Atlantic. Grenada (Bowerbank slide). Other material. BMNH
1902.10.18.416 - Dendy's specimen RN891.
Description. Erect, lamellate, massive growth form; surface
HIGGINSIA HIGGIN, 1877 conulose, striated; ectosomal skeleton with slender extra-axial
oxeas (occasionally styles) protruding through the surface in sparse
Synonymy plumose bundles, and centrangulate microxeas tangential and
below the surface (not forming a crust); extra-axial skeleton with
Higginsia Higgin, 1877: 291; Hallmann, 1917c: 655; Dendy, radial tracts of very long, slender, sinuous styles surmounted by the
1922b: 126; Topsent, 1928c: 39; de Laubenfels, 1936a: 132; oxeas protruding through the surface, arising from the axis in
Burton, 1959a: 255; Wiedenmayer, 1977b: 156; Hooper & Levi, sparse plumose tracts; choanosomal skeleton halichondroid with a
1993b: 1452. Dendropsis Ridley and Dendy, 1886: 483, 1887: 191; partially compressed, reticulate axis composed of robust oxeas
Hallmann, 1917c: 693; de Laubenfels, 1936a:132; Levi, 1973: 606. without any obvious fibre component but mesohyl heavily invested
AllantellaHallmann, 1917c: 675. Desmoxya Hallmann, 1917c: 649; with collagen, and abundant centrangulate microxeas scattered
de Laubenfels, 1936a: 132; Van Soest et al., 1990: 18. Taxonomic throughout; ectosomal oxeas/styles raphidiform, widely curved or
decision for synonymy: Hooper & Levi (1993b) and this work. sinuous (430-620 X 3-5 !-Lm); extra-axial styles long, slender, sinu-
ous, bases evenly rounded (1100-1650 X 14-22 !-Lm); choanoso-
Type species mal oxeas robust, thick, usually symmetrical, with fusiform points
(580-740 X 11-32 !-Lm); microscleres spined, centrangulate, curved
Higginsia coralloides Riggin, 1877 (by subsequent designation, or straight microxeas (65-240 X 3-6 !-Lm).
Hallmann, 1917c), (holotype BMNH 1886.12.15.92), possible Remarks. The nominal genera Higginsia, Dendropsis and
junior synonym of Spongia strigilata Lamarck, 1814: 450 Desmoxya differ essentially in skeletal construction (reticulate or
(Wiedenmayer, 1977b) (holotype MNHN DT637). plumo-reticulate; with a compressed axis and radial extra-axis; and
plumose-halichondroid, with meandering, occasionally reticulate
Definition skeletal tracts, respectively). Most species lack definite axial com-
pression of the skeleton (except Dendropsis bidentifera Ridley &
Desmoxyidae with sinuous raphidiform ectosomal oxeasl Dendy, 1886 (holotype BMNH 1887.5.2.59), the type species, by
styles protruding through the surface in sparse plumose bundles, original designation, of Dendropsis), having instead a halichon-
supported by long sinuous extra-axial styles in sparse plumose droid, vaguely reticulate axis, and in most species there is often
tracts, embedded in a choanosomal skeleton of variable structure some differentiation of axial and extra-axial skeletons (except
(halichondroid to axially compressed) composed of choanosomal Desmoxya lunata), suggesting some sort of affinity with the con-
oxeas in bundles, with coarsely spined centrangulate microxeas cept ofAxinellida. All three nominal genera share the apomorphy
tangential and below the surface (not forming a crust). of spined microxeas, but it is not entirely convincing that the major
differences in their skeletal structures can be ignored, particularly
Diagnosis in the case of Dendropsis which has a nearly 'classical axinellid'
architecture, in recognising these taxa merely as synonyms of
Growth forms erect, lamellate, massive, vasiform or lobate; Higginsia (e.g., Hallmann, 1917c; Van Soest et al., 1990).
surface conulose, papillose, often silt covered or membraneous. It is possible that resurrection of Desmoxya (with type species
Choanosomal skeleton predominantly with an elaborate system of Higginsia lunata Carter, 1885d (by original designation) holotype
bundles of megascleres, although skeletal structure ranges from BMNH 1886.12.15.138) is required to accomodate Higginsia-like
halichondroid with a partially compressed, reticulate axis, and species that lack any evidence of axial compression (as seen in
an irregularly plumo-reticulate extra-axial region (Higginsia), a both H. lunata and H. anfractuosa Hooper & Levi, 1993b), having
compressed axis and a radial, non-plumose extra-axial region instead a simply halichondroid, meandering reticulation of
(Dendropsis), to a lax plumose or plumo-reticulate structure, choanosomal tracts as seen in other axinellid genera such as
without axial compression or regional differentiation of the skeleton Rhaphoxya. These species also differ from Higginsia sensu stricto
(Desmoxya); spongin fibres usually poorly developed although only in having one size category of megaScleres, raphides in addi-
heavy collagen forms the mesohyl, usually with numerous megas- tion to spined microxeas, as well as a more lax skeletal architec-
cleres and microscleres scattered between main skeletal tracts; all ture. However, intermediate forms of architecture between
skeletal tracts cored by robust oxeas and/or styles, or sometimes Higginsia and Desmoxya are present in some species of Higginsia
modified further to strongylote megascleres with bifurcated points. (e.g., H. massalis), and this condition is interpreted here as being
Ectosome with a protruding extra-axial skeleton composed of merely a reduced form of the skeleton that is typical for Higginsia
longer oxeas and/or styles, and smaller oxeas/styles forming sparse (Van Soest et al., 1990). Allantella Hallmann (with type species
brushes or tangential layers on the surface. Megascleres oxeas, Trachytedania arborea Keller, 1889) has spined microstrongyles
strongyles and/or styles of 1-3 sizes. Microscleres spined, centran- and is clearly synonymous with Higginsia (confirmed by examina-
gulate curved or straight microxeas, and sometimes also raphides tion of specimen 2MB 444 courtesy of R. WM. Van Soest).
occurring singly or in bundles (trichodragmata). Hallmann (1917 c) provided a comprehensive review of the
genus and the diagnostic features of species included at that time,
Description of type species although many more species have since been described and all now
are in need of detailed revision. Only a few type specimens have
Higginsia corallo ides Higgin, 1877 (Fig. 5). yet been located and seen, so it is not possible to undertake
764 Porifera· Demospongiae • Halichondrida • Desmoxyidae

I
A B CD

Fig. 5. Higginsia coral/oides Higgin (holotype and Dendy specimen). A, basal end of extra-axial style (scale 100 fLm). B, choanosomal oxea (scale
100 fLm) . C, ectosomal oxea (scale 100 fLm). D, microspined microxea (scale 10 fLm). E, peripheral skeleton (XS) (scale 250 fLm). F, ectosomal skeleton
(TS) (scale 250 fLm). G, choanosomal skeleton (scale 250 fLm). H, idealised reconstruction of skeleton. I, holotype of Spongia strigilata Lamarck (MNHN
DT637), senior synonym of H. coralloides.

a review of these species at the present time. Twenty species are Distribution
thought to be valid: H. anfractuosa Hooper & Levi, 1993b; H.
bidentifera (Ridley and Dendy); H. ciccaresei Pansini & Pesce, Worldwide, but most common in tropical and temperate Indo-
1998; H, coralloides Higgin, with varieties H. c. liberiensis Pacific Ocean - Chile, Caribbean, Azores, Ireland, Iceland,
Higgin, 1877, and H. c. arcuata Higgin, 1877 [possible synonym Mediterranean, South Africa, West Africa, W Indian Ocean,
of H. strigilata (Lamarck)]; H. higgini Dendy, 1922b; H. lunata Seychelles, Red Sea, Gulf of Aden, Indonesia, Palau, SE, NW and
Carter; H. massalis Carter, 1885d; H. mediterranea Pulitzer-Finali, NE Australia, New Caledonia.
1977; H. mixta Hentschel, 1912; H. natalensis Carter, 1885d;
H. petrosioides Dendy, 1922b; H. pumila (Keller, 1889);
H. robusta Burton, 1959a; H. scabra Whitelegge, 1907; H. strigi- JULA VIS DE LAUBENFELS, 1936
lata (Lamarck); H. tanekea Hooper & Levi, 1993b; H. tethyoides
Levi, 1960a; and H. thielei Topsent, 1898b (Van Soest, 1987b; Synonymy
Hooper & Levi, 1993b; Pansini & Pesce, 1998), with H. papillosa
Thiele, 1905 transferred to Halicnemia (Desqueyroux-Faundez & lulavis de Laubenfels, 1936a: 79; Van Soest & Lehnert,
Van Soest, 1997). 1997: 502.
Porifera· Demospongiae • Halichondrida • Desmoxyidae 765

Fig. 6. lulavis levis (Kirkpatrick) (holotype). A, choanosomal acanthostrongyles (scale 50 IJ..m). B, ectosomal acanthostrongyles (scale 50 fLm). C, basal
end of smooth strongyle (scale 100 fLm) . D, smooth style (scale 100 fLm) . E, raphide (scale 10 fLm). F, peripheral skeleton (XS) (scale 250 fLm). G, periph-
eral skeleton (XS) (scale 100 fLm). H, choanosomal skeleton (scale 100 fLm) . I, idealised reconstruction of skeleton.

Type species Description of type species

Tedania levis Kirkpatrick, 1900b (by original designation). lulavis levis (Kirkpatrick, 1900b) (Fig. 6).
Synonymy. Tedania levis Kirkpatrick, 1900b: 355, pI. 14,
Definition fig. 4.
Material examined. Holotype: BMNH 1900.10.19.16 -
Desmoxyidae with spiny strongyles forming a tangential ecto- Funafuti (Ellice Islands) (=Tuvalu), central S Pacific.
somal skeleton and raphides in dragmata. Description. Thinly encrusting on dead coral, with smooth
surface; ectosomal skeleton a tangential, criss-cross layer of thinner,
Diagnosis curved, smooth, granular or vestigially microspined strongyles;
choanosomal skeleton halichondroid reticulate, composed of loose
Thinly encrusting growth form; choanosomal skeleton tracts of larger, curved or sinuous, occasionally vermiform, promi-
reduced, consisting of a halichondroid-reticulate arrangement of nently acanthose strongyles with poorly developed spongin fibres
acanthose strongyles forming a basal skeleton, and longer smooth but abundant granular collagen dispersed throughout the mesohyl;
strongyles and styles oriented perpendicularly to the surface, single skeletal reticulation denser near the periphery than towards the
or in bundles; ectosomal skeleton with smaller spined strongyles basal skeleton, with some peripheral tracts forming distinct spicule
lying tangential to the surface; megascleres are long, thick, curved bundles; few larger, smooth strongyles and styles ascend to
or vermiform acanthose strongyles, and longer smooth strongylesl the surface from the basal skeleton to the periphery in more-or-less
styles; microscleres are raphides in trichodragmata. plumose or radial tracts or as single spicules; megascleres
766 Porifera· Demospongiae • Halichondrida • Desmoxyidae

acanthose strongyles of 2 sizes, both curved, sinuous or sometimes Description of type species
vermiform, the smaller with vestigial spination, the larger promi-
nently spined (250-300 X 3-12 f,Lm), and long smooth, curved or Microxistyla petrina Topsent, 1928c (Fig. 7).
sinuous strongyles or styles (245-1380 X 4--12 f,Lm); microscleres Synonymy. Microxistyla petrina Topsent, 1928c: 179, pI. 6
raphides in trichodragmata (150-200 X 6-12 f,Lm). fig. 18.
Remarks. De Laubenfels (1936a) erected lulavis for Material examined. Holotype: MOM (not seeen), schizotypes
Kirkpatrick's (1900b) species, based on a single encrusting speci- MNHNDTlI41, BMNH 1930.7.1.42-CapeVerdels, NEAtiantic.
men containing two distinct categories of spiny strongyles and Description. Encrusting-massive, irregular surface perfo-
raphide microscleres. De Laubenfels overlooked the presence of rated by small oscula, stony consistency; ectosomal skeleton with
the longer smooth strongyles and styles that arise from the basal erect, plumose bundles of microxeas arising from subectosomal
skeleton, although admittedly these are uncommon in the holotype tracts forming a nearly continuous palisade on the surface; subecto-
slide preparation (BMNH 1900.10.19.16a), and also mentioned by somal skeleton plumose, cavernous, with multispicular bundles
Kirkpatrick in the original description. Kirkpatrick initially of styles; choanosomal skeleton riddled with aquiferous canals,
assigned his species to Tedania based on the possession of raphide between which is an irregularly halichondroid reticulate skeleton
microscleres as well as diactinal ectosomal megascleres but these composed of robust styles poorly bound with sparse collagen;
characters are not truly homologous to the onychaetes and ectoso- megascleres curved thick styles (400-575 X 20-30 f,Lm); microscle-
mal tylotes of Tedania; moreover T. levis contains spined diactinal res smooth centrangulate or straight microxeas (60-160 X 2-4 f,Lm).
choanosomal megascleres and differentiated axial and extra-axial Remarks. Topsent (1928c) remarked that this monotypic
spicules. De Laubenfels subsequently erected the genus lulavis to genus showed affinities with the genera Axinyssa (Halichondriidae)
his admittedly 'unsatisfactory family' Acarniidae (= Microcionidae and Ceratopsion (Raspailiidae) in having oxeote microscleres, but
in the contemporary sense) based on possession of spiny spicules this similarity is probably superficial given that microxeas found
(megascleres), but the genus appears to have closest resemblance in either of these genera are not centrangulate. Similarly, in texture
to the Desmoxyidae having a halichondroid reticulate choanoso- and skeletal architecture Topsent (1928c) suggests that M. petrina
mal skeleton and more organised tangential ectosomal skeleton, shows similarities to the stony haplosclerids (Petrosia, Strongy-
including spined diactinal spicules and raphide microscleres. The lophora), emphasized by the paucity of spongin fibres or collagen
genus contains only two species so far, and still relatively poorly within the choanosome, but this is again only superficial. The genus
known, although the recent description of a new species and is more obviously related to the desmoxyids (Higginsia, Desmoxya)
redescription of the type species by Van Soest & Lehnert (1997) with centrotylote microscleres, and with similarities to some hali-
has made the genus now perfectly recognisable. chondriids (like Spongosorites dendyi (Topsent) and S. placenta
Topsent), which have centrotylote modifications to some spicules
Distribution (although in the Halichondriidae these are classed as megascleres
and not microscleres as in Desmoxyidae). Topsent omitted to men-
Tuvalu, central Pacific, and Jamaica, Caribbean. tion that the species has a specialised ectosomal skeleton in which
plumose bundles of microxeas project from peripheral subectoso-
mal tracts. However, the holotype is dry and this may have become
MICROXISTYIA TOPSENT, 1928 detached in his material.

Synonymy Distribution

Microxistyla Topsent, 1928c: 179. Monotypic, known only from the NE Atlantic and Sea of
Okhotsk.
Type species
MYRMEKlODERMA EHLERS, 1870
Microxistyla petrina Topsent, 1928c (by monotypy).
Synonymy
Definition
Myrmekioderma Ehlers, 1870: 28; Bergquist, 1965: 177.
Desmoxyidae with plumose bundles of smooth centrangulate Acanthoxifer Dendy, 1905: 156; Dendy, 1922b:129; Bergquist,
or straight microxeas forming a nearly continuous palisade on the 1965: 177. Anacanthaea Row, 1911: 329; Van Soest et al., 1990:
surface, supported by plumose extra-fibre tracts of choanosomal 31. Callites Schmidt, 1868: 16; Van Soest et al., 1990: 31.
styles and a halichondroid reticulation of the same styles below. Neoprosypa de Laubenfels, 1954: 189. Taxonomic decision for
synonymy. Van Soest et al. (1990: 31).
Diagnosis
Type species
Encrusting-massive growth form, stony texture; ectosomal
skeleton with erect, plumose bundles of smooth microxeas forming Alcyonium granulatum Esper, 1830 (by monotypy).
a nearly continuous palisade on the surface; extra-axial region cav-
ernous with plumose tracts of choanosomal styles; choanosomal Definition
skeleton a confused halichondroid reticulation of smooth robust
styles without fibres but bonded together with minimal collagen; Desmoxyidae with a detachable ectosomal skeleton consisting
microscleres smooth straight or centrangulate microxeas. of smaller (acanth-)oxeas in brushes perpendicular or paratangential
Porifera· Demospongiae • Halichondrida • Desmoxyidae 767

I
A B

Fig. 7. Microxistyla petrina Topsent (schizotypes). A, styles (scale 150 /-Lm). B, microxeas (scale 10/-Lm). C, peripheral skeleton (scale 250 /-Lm). D, ecto-
somal peel (scale 200 /-Lm). E, choanosomal spicule tract (scale 400 /-Lm). F, idealised reconstruction of skeleton.

to the surface, supported by larger choanosomal (acanth-)oxeas, species but not all). Subectosomal skeleton composed of ascending
strongyles or styles forming a slightly compressed halichondroid tracts of the same spicules running radially or obliquely to the
reticulation in the central portion of the choanosome and a more surface, with larger choanosomal megascleres protruding, and fre-
cavernous peripheral skeleton of oblique spicule tracts, with wispy quently cavernous towards the periphery. Ectosomal skeleton without
trichodragmata of raphides in one or two size classes dispersed specialised spiculation although smaller (acanth-)oxeas protrude
throughout the skeleton. from the subectosomal region, forming a dense 'peel' of closely adja-
cent brushes perpendicular or paratangential. Megascleres consist of
Diagnosis two distinct categories of oxeas, strongyles, or more rarely styles,
sometimes sinuous and centrally flexed, either entirely smooth or
Massive or encrusting growth forms. Surface hispid, with with even spination, the larger choanosomal spicules more variable in
characteristic excavating meandering, sinuous or straight canals their terminations than the smaller ectosomal spicules which are typ-
and grooves sometimes forming polygonal tuberculate plates. ically oxeas or acanthoxeas. Microscleres raphides in trichodrag-
Choanosomal skeleton collagenous, with compressed central portion mata, in one or more categories, the larger usually sinuous or curved.
composed of a confused halichondroid mass of larger choanosomal
megascleres ranging from acanthoxeas and oxeas, strongyles or less Description of type species
frequently styles, forming irregular, ascending, multispicular tracts
bound together with sparse collagen (spongin fibres present in some Mynnekiodenna granulata (Esper, 1830) (Fig. 8).
768 Porifera· Demospongiae • HaIichondrida • Desmoxyidae

Fig. 8. Mynnekiodenna granulata (Esper) (specimen QMG303135). A, choanosomal oxeas (smooth or vestigial spination) (scale 100 j.1m). B, ectosomal
acanthoxeas (scale 50 j.1m). C, trichodragmata (scale 50 j.1m). D. peripheral skeleton (scale 250 j.1m). E, subectosomal skeleton (scale 250 j.1m).
F, choanosomal skeleton (scale 250 j.1m). G, idealised reconstruction of skeleton. H, specimen in situ (photo author).

Synonymy. Alcyonium granulatum Esper, 1830: 71; of smaller ectosomal (acanth-)oxeas arising from the subectosomal
Mynnekiodenna granulata; Ehlers, 1870: 28. Acanthoxifer ceylonen- region (305-708 X 3-12ll-m) forming a continuous paratangential
sis Dendy, 1905: 157. Mynnekiodenna tylota de Laubenfels, 1954: palisade, supported below by long, pillar-like tracts of larger
119. Neoprosypa atina de Laubenfels, 1954: 190. Acanthoxifer four- choanosomal (acanth-)oxeas and styles producing a cavernous sub-
manoiri Levi, 1956c: 5. dermal region; choanosome cavernous, with a vaguely halichon-
Material examined. Holotype: Unknown - Indonesia. droid reticulate skeletal structure, sometimes with a compressed
Other material. Numerous specimens, widespread tropical central portion, and with differentiated primary and secondary
Australasia (see Hooper & Levi, 1993b: 1449; Hooper et al., 1997: spongin fibres and spicule tracts; primary, ascending, fibres very
8). Holotypes of junior synonyms (see below). large, with up to 50 spicules per tract but diminishing in size
Description. Massive, bulbous, partially burrowing, light towards the periphery, more-or-Iess regularly spaced, intercon-
orange-brown to bright orange exterior; surface convoluted with nected by secondary, transverse or oblique, pauci-, uni- or aspicular
large conules or rounded or polygonal plates, each separated by fibres; fibres cored by larger choanosomal (acanth-)oxeas (495-
shallow but distinct grooves, excavated channels containing large 950 X 8-22Il-m). Two categories of megascleres differing only in
oscula; ectosomal skeleton a detachable 'peel', with a distinct crust size and distribution within skeleton, both straight or slightly curved
Porifera· Demospongiae • Halichondrida • Desmoxyidae 769

at centre, rarely asymmetrical, with variable terminations, usually retained this genus in the family Desmoxyidae, supporting their
tapering to sharp fusiform points but also with telescoped, decision with chemotaxonomic evidence based on an inferred close
strongylote, styloid modifications and other terminal deformities, relationship between Higginsia and Myrmekioderma (Hooper
and both categories of spicules may be entirely smooth or lightly et al., 1992). However, morphological support for homology
and evenly acanthose; ectosomal spicules tend to be mostly between the occasionally acanthose ectosomal oxeas in
oxeas whereas choanosomal megascleres have greater geometric Myrmekioderma and the true ectosomal acanthoxeas in Higginsia
variability. Microscleres raphides mostly in trichodragmata is still equivocal, with the consequence that Hooper et aZ. (1997)
(110-155 X 10-15 fLm). returned it to Halichondriidae. The overall similarity between the
Remarks. Bergquist (1965) meticulously redescribed the ectosomal structure, choanosomal organisation and spicule compo-
type species of Myrmekioderma from a number of specimens from sition (apart from the possession of microxeas or discorhabds,
a range of geographic localities. She showed conclusively that the respectively) largely supports the inferred close relationship
type species of Acanthoxifer (A. ceyZonensis Dendy, 1905 (by between Myrmekioderma and Didiscus, of which the latter had
monotypy) holotype BMNH 1907.2.1.54,55) and Neoprosypa been previously included in Halichondriidae (e.g., Van Soest et al.,
(N. atina de Laubenfels, 1954 (by monotypy) holotype USNM 1990). Conversely, structural similarities in the erect palisade of
22974) were conspecific and also identical to M. granulata. ectosomal spicules is remarkable between Myrmekioderma and
Bergquist also noted that M. granulata had a wide geographical dis- Higginsia (see illustrations in Hooper & Levi, 1993b). As a conse-
tribution with a corresponding high degree of skeletal variability, quence of these discrepancies both Myrmekioderma and Didiscus
particularly in the presence, absence and size of certain spicule cat- are now referred to the Desmoxyidae, to acknowledge their closer
egories, and the presence or absence of spination on megascleres. affinities to genera such as Higginsia. This decision is supported by
Bergquist (1965: 182) also discussed at length the relation- the findings of Van Soest & Lehnert (1997) who recently
ships between Myrmekioderma, Acanthoxifer, Anacanthaea (type redescribed and revived the genus lulavis, previously incertae
species A. nivea Row, 1911 (by monotypy) holotype BMNH sedis. They showed that luZavis had obvious morphological links
1912.2.1.54), Spongosorites and Heteroxya, and she concluded that with Myrmekioderma (with wispy trichodragmata) and Higginsia
Myrmekioderma, Anacanthaea and Heteroxya were good genera, (coarsely spined diactinal spicules), and on this basis they
and most closely related to Higginsia. Conversely, Van Soest et aZ. re-assigned all these genera to Desmoxyidae.
(1990) included Anacanthaea in synonymy with Myrmekioderma. Van Soest et aZ. (1990) included six species in
Two species were initially assigned to Anacanthaea, the type Myrmekioderma, of which only two live in the Indo-west Pacific,
species and A. rea de Laubenfels (1934: 11). Apart from and a seventh species should also be included: M. dendyi (Burton,
megasclere size and live colouration both species are identical. 1959a) from the south Arabian coast and Indonesia; M. granulata
In external morphology, distribution of pores and ornamentation widespread throughout the Indo-west Pacific; M. nivea (Row,
of pore grooves A. nivea and A. rea are also very similar to 1911) from the Red Sea; M. rea de Laubenfels, 1934 from the
Myrmekioderma granulata (Esper), which by contrast is bright vicinity of Puerto Rico; M. spelaea (Pulitzer-Finali, 1983) from the
orange whereas the two Anacanthaea species are white and dark Mediterranean; M. styx de Laubenfels, 1953a from the Gulf of
pinkish-grey, respectively. In addition, M. granuZata has acanthose Mexico; and M. tulearensis (Vacelet et al., 1976) from SW
microxeas, choanosomal oxeas, and a separate category of Madagascar.
ectosomal oxeas, whereas Anacanthaea has only oxeas. However,
several regional populations of M. granulata have their spiculation Distribution
reduced to only smooth oxeas, in addition to microxeas (e.g.,
Hooper et al., 1997), and in this regard Anacanthaea fits comfort- Widely distributed in shallow tropical and subtropical oceans
ably within Myrmekioderma. Van Soest et al. (1990) commented throughout the Indo-west Pacific: Mediterranean, Madagascar,
further on the generic synonymy of Myrmekioderma, and included Aldabra, Seychelles, Gulf of Manaar, Indonesia, NW Australia,
Callites (type species C. lacazii Schmidt, 1868 (by monotypy) northern Great Barrier Reef, central western Pacific - Ponape,
fragment of holotype BMNH 1868.3.2.3) into this synonymy. Truk, Ebon Atoll, Palau, Ifaluk, Chesterfield Islands, New
Subsequent to Bergquist'S (1965) work the genus has been Caledonia.
closely examined by Van Soest et al. (1990: 31), Diaz et al. (1991),
Hooper & Levi (1993b: 1448) and Hooper et al. (1997: 6).
Together these studies demonstrate that Myrmekioderma, Didiscus NEGOMBO DENDY, 1905
and Higginsia are closely related and most appropriately assigned
to the same family. Myrmekioderma and Didiscus are superficially Synonymy
very similar and both easily recognisable in the field by their
deeply sculptured sinuous and plate-like grooves on the surface, Negombo Dendy, 1905: 127. Oxylatrunculia Hoshino,
having an ectosome composed of smaller spicules forming an erect 1981: 221.
palisade over tangential larger spicules, and a similar basic
choanosomal skeletal organisation (Van Soest et aI., 1990; Diaz Type species
et al., 1991). Myrmekioderma lacks discorhabd microscleres, char-
acteristic of Didiscus, but has instead bundles of sinuous raphides Negombo tenuistellata Dendy, 1905 (by monotypy).
(trichodragmata). The genus has been traditionally defined with
lightly acanthose ectosomal oxeas (e.g., Bergquist, 1965; Hooper & Definition
Levi, 1993b), but Diaz et al. (1991) correctly note that these ecto-
somal spicules are as frequently smooth as they are acanthose, and Desmoxyidae with a halichondroid skeleton of styles and/or
they emend the definition accordingly. Hooper & Levi (1993b) oxeas and with sanidaster-like acanthose microrhabds.
770 Porifera· Demospongiae • Halichondrida • Desmoxyidae

A IB
Fig. 9. Negomho keUyae sp. nov. (holotype OCDN3701-I). A, choanosomal oxeas and styles (scale 300 fLm) . B, acanthose sanidaster-like microrhabds
(scale 10 fLm). C, peripheral skeleton illustrating disposition of microrhabds on surface (scale 100 fLm). D, halichondroid choanosomal skeleton (scale
500 fLm). E, idealised reconstruction of skeleton.

Diagnosis inclusions, hard and slightly incompressible. Oscula and pores not
observed. Texture soft and flexible. Ectosomal skeleton with a
Massive fistular, with a tangential surface layer of sanidaster- layer of sanidasters lying tangentially or irregularly scattered on
like acanthose microrhabds which are also scattered within the the surface, together with abundant foreign debris. Choanosomal
mesohyl lining the canals. Choanosomal skeleton with halichon- skeleton with very large gently curved styles and oxeas forming
droid tracts of large styles and/or oxeas forming longitudinal or confused longitudinal halichondroid tracts, apparently without
radial, compact tracts. spongin fibres, and with sanidasters lining the aquiferous system.
Megascleres are smooth styles and oxeas 540-1100 X 12-17 f.Lm.
Description of type species Microscleres are acanthose microrhabds (sanidasters) with irregu-
lar whorls of spines along the shaft, 12-15 f.Lm in length.
Negombo tenuistellata Dendy, 1905. Description of Negombo keUyae sp. nov. (Fig. 9). Massive
Synonymy. Negombo tenuistellata Dendy, 1905: 127-128, lump, slightly hemispherical. Texture firm, spiculose, sticky, tears,
pI. 5, fig. 8. encrusted by other sponges and detritus. Large central oscule.
Material examined. Holotype (not seen): BMNH Black alive, with 'sweet, pleasing' odour. Ectosome heavily
1907.2.1.28 - Sri Lanka. Comparative material. Holotype of encrusted with coral debris, diatoms, and foreign spicules, with
Negombo kellyae sp.nov.: BMNH 2000.9.15 .1 (OCDN3701-I) - thin tangential or paratangential layer of sanidasters 2-5 deep
Pemba I., W Tanzania, 5°0.60'S, 39°40.14'E, coIl. CRRF, 26.i.96. forming sparse crust on the surface. Choanosome with a sparse
Description of type species (from Dendy, 1905). Sponge halichondroid reticulate skeleton composed of large oxeas and
massive with fistular outgrowths. Surface roughened with foreign styles, the latter less common than the former, in two intergrading
Porifera· Demospongiae • Halichondrida • Desmoxyidae 771

II
AB

Fig. 10. Parahigginsia phakellioides Dendy (holotype). A, oxea (scale 150 11m). B, microxeas (scale 10 11m). C, peripheral skeleton (scale 1 mm). D, ide-
alised reconstruction of skeleton. E, holotype (from Dendy, 1924: pI. 11, fig. 4).

size classes. Reticulation open-meshed, with meshes up to 500,...m microrhabds suggest that the genus might be best assigned to
wide, bounded by uni- or paucispicular tracts. No spongin fibres Desmoxyidae, closest to Didiscus. Microrhabds in Negombo
observed. Spicule tracts near the surface slightly diverging, paucispic- and Didiscus appear to be modified microxeas, whereas those of
ular, not protruding through the surface. Mesohyl heavily collagenous Latrunculia (acanthodiscorhabds), and Diacamus and Negombata
containing abundant cyanobacteria. Choanocyte chambers lined by (spinorhabds) have different geometry and are not considered to be
sparse sanidasters, fewer than at the surface. Few interstitial spicules homologous (M. Kelly, pers. comm.) (see chapters on Podospongiidae
present. Megascleres oxeas and styles with approximately the same and Latrunculiidae in this work).
geometry, very long and robust, slightly curved at the centre, long Oxylatrunculia Hoshino, 1981 (type species Oxylatrunculia
tapering points (1325-(1814)-2240 X 37-(51)-66,...m). Smaller acanthosanidastra Hoshino, 1981) is virtually identical to N.
oxeas similar in shape but much less robust (765-(922)-1245 X tenuistellata but having a more radiate skeletal architecture, much
22-(27)-34,...m). Microscleres acanthose microrhabds resembling larger spiCUlation with exclusively oxeote megascleres.1t too clearly
sanidasters, 18-26,...m long, with large spines (up to 3,...m long) scat- belongs to Negombo making Oxylatrunculia a junior synonym.
tered over shaft, not in whirls and not discoid in shape. Etymology. The genus presently contains three species: Negombo
Named for Michelle Kelly for discovering this species. tenuistellata Dendy, N. acanthosanidastra (Hoshino) and N. kellyae
Remarks. Hooper (1986) and Wiedenmayer (1989) regarded sp.nov. described here. The new species differs from N. tenuistellata
Negombo tenuistellata as a synonym of Latrunculia, while de in having much larger megascleres and microscleres, having pre-
Laubenfels (1936a) considered the possibility that Negombata was dominantly oxeas in the skeleton (whereas the type species has pre-
synonymous with Negombo. Examination of Negombo tenuistellata dominant styles), having a much more lax and sparse choanosomal
Dendy by Kelly-Borges & Vacelet (1995) upheld the generic status of reticulation (the type species has long wispy halichondroid tracts),
Negombo and concluded that it was distinct from Latrunculia, and in its live specimen characteristics (the type species is fistulose,
Diacamus and Negombata based on differences in skeletal structure, tubular, yellowish grey). In spicule dimensions the present species
gross morphology and spiculation. The large oxea and styloid megas- is more similar to N. acanthosanidastra but megascleres consist of
cleres, the halichondroid skeletal architecture and the geometry of both oxeas and styles (only oxeas in the Japanese species), sanidasters
772 Porifera· Demospongiae • Halichondrida • Desmoxyidae

have several large spines but is not microspined (unlike the latter), without distinct fibres or obvious collagenous spongin; megascle-
and live characteristics (massive, sepia coloured). res smooth oxeas of a single category; microscleres curved acan-
those microxeas.
Distribution
Description of type species
Japan, Sri Lanka, Tanzania.
Parahigginsia phakellioides Dendy, 1924 (Fig. 10).
Synonymy. Parahigginsia phakellioides Dendy, 1924: 375,
PARAHIGGINSIA DENDY, 1924 pI. 11 fig. 4, pI. 15 figs 32-33.
Material examined. Holotype: BMNH 1923.10.1.156 -
Synonymy North Cape, New Zealand.
Description. Branching and anastomosing, curved, thinly
Parahigginsia Dendy, 1924: 375; de Laubenfels, 1936a: 159; lamellate growth form; surface smooth, regular, with distinct
Bergquist, 1970: 22. osculiferous inhalant and porous exhalant surfaces; texture hard,
brittle; ectosomal skeleton with membraneous skin-like surface
Type species bearing scattered tangential microxea microscleres on the surface,
multispicular tracts of choanosomal oxea megascleres forming the
Parahigginsia phakellioides Dendy, 1924 (by monotypy). dense tangential crust just below the surface, up to 5 spicules thick;
ascending choanosomal spicule tracts do not protrude through the
Definition surface; choanosomal skeleton a dense petrosiid-like isotropic
reticulation of robust oxeas forming thick irregular tracts, 2-8
Desmoxyidae with a glabrous ectosomal skeleton containing spicules thick, without spongin fibres but bound together with
only curved acanthose microxeas lying tangentially, supported sparse collagen, and with large oval meshes up to 500 f.Lm diame-
below by a dense peripheral crust of smooth choanosomal oxeas ter; microxea microscleres also scattered within the mesohyl;
becoming isotropic and more compressed towards the axis. megascleres short, stout, slightly curved hastate oxeas or some-
times strongyloxeas (280-355 X 18-23 f.Lm); microscleres slightly
Diagnosis curved, minutely spined microxeas with evenly dispersed spines
(78-95 X 1-2f.Lm).
Erect, lamellate, lobate growth forms; surface smooth, not Remarks. Parahigginsia is closely related to Higginsia in
hispid; ectosome glabrous, containing only spongin and scattered skeletal construction and spiculation, differing only by its Phakellia-
acanthoxeas lying tangential to the surface; extra-axial skeleton not like lamellate growth form, smooth ectosome, and absence of long
clearly delineated from axis, but consisting of irregularly anasto- slender megascleres forming the extra-axial skeleton.
mosing and ascending primary and secondary tracts of choanoso-
mal oxeas, which do not protrude through the surface but become Distribution
more ordered and compact near the periphery, forming a dense
crust below the ectosome; choanosomal skeleton with a condensed Monotypic, known only from New Zealand.
axial reticulation of oxeas forming irregular isotropic tracts,
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Dictyonellidae Van Soest, Diaz & Pomponi, 1990

Rob W.M. Van Soestl, Dirk Erpenbeckl & Belinda Alvarez2

1 ZoologicalMuseum, University of Amsterdam, P.O. Box 94766, 1090 GT, Amsterdam, Netherlands. (soest@science.uva.nl,
erpenbeck@science.uva.nl)
2 Museum and Art Gallery of the Northern Territory Museum, P.O. Box 4646, Darwin NT 0801, Australia. (belinda.glasby@nt.gov.au)

Dictyonellidae Van Soest et ai. (Demospongiae, Halichondrida) are sponges with fleshy appearance, lacking a special surface skeleton of
spicules. The spicular density is often relatively low and skeletons consist of ill-defined bundles following a dendritic pattern or making
an irregular reticulation. Spongin may be important as binding substance. Spicules comprise usually long, often somewhat flexuous
styles, occasionally strongyles, oxeas or oxeote derivations, but shorter straight styles are not uncommon. Characteristically, spicule end-
ings are variable and often telescoped. Dictyonellidae are cosmopolitan in occurrence, but are more common in warmer waters. Compared
with the original contents, the family has been here extended to include Phakettia, Stylissa and Svenzea, and now comprises ten valid
genera (out of 17 nominal genera).
Keywords: Porifera; Demospongiae; Halichondrida; Dictyonellidae; Acanthella; Dictyonella; Liosina; Lipastrotethya; Phakettia;
Rhaphoxya; Scopalina; Stylissa; Svenzea; Tethyspira.
- - - - - - ~---- --~----~-------~-------~---~---~. -----

DEFINITION, DIAGNOSIS, SCOPE Scope

Synonymy Ten genera out of eighteen nominal genera are recognized as

valid: Acanthella Schmidt, 1862: 65; Dictyonella Schmidt, 1868:
[Acanthellidae1Carter, 1885d: 364 as Acanthellina (to be sup- 10; Liosina Thiele, 1899: 17; Lipastrotethya de Laubenfels, 1954:
pressed under ICZN Article 23.9.1 as an unused name, not having 235; Phakettia de Laubenfels, 1936a: 130; Rhaphoxya Hallmann,
been used after 1899 and the junior name in frequent use in the last 1917c: 641; Scopalina Schmidt, 1862: 79; Stylissa Hallmann, 1914:
50 years and longer than 10 years prior to this publication). There 349; Svenzea Alvarez et al., 2002: in press; Tethyspira Topsent,
is also a genus [Acanthellinal Carter, 1885d which is considered a 1890a: 195.
nomen oblitum (cf. below), which is a further reason not to use
Carter's family name. Dictyonellidae Van Soest, Diaz & Pomponi,
1990: 20. TAXONOMIC HISTORY AND BIOLOGY

Definition The family was only recently erected in an attempt to

rearrange the genera assigned to the former orders Axinellida
Halichondrida without ectosomal skeleton, but with dense and Halichondrida sensu Levi, 1973, which was necessary because
organic ectosomal layer, giving the sponge a fleshy appearance; of the increased awareness that these groups as then understood
choanosomal skeleton is predominantly built of styles, occasion- were clearly polyphyletic (Van Soest et ai., 1990; Diaz et al., 1991;
ally oxeas or strongyles occur. Pomponi et ai., 1991; Diaz et al., 1993; Alvarez, 1998).
Abandonment of a separate order Axinellida and revision of the
Diagnosis families Halichondriidae and Hymeniacidonidae yielded a pro-
posal for an order Halichondrida sensu lato comprising four fami-
Encrusting, massive, lobate, digitate, bushy or colurnnar- lies, Axinellidae, Desmoxyidae, Halichondriidae and a new family
tubular sponges, usually with a conulose surface. Between conules, Dictyonellidae (Van Soest et al., 1990). Since these families were
the surface is shiny-smooth and distinctly fleshy. Ectosomal skele- newly defined, the generic content has been under constant debate
ton is entirely absent. If attempts are made to make a tangential and rearrangements have been proposed in the recent literature up
section, the surface comes off in flakes. Choanosomal tracts and to and including this volume. Further challenges of the adopted
individual spicules directed towards the surface, often collected in classification of the order Halichondrida and its families may be
vague tracts or more definitely in spongin enforced spiculofibre. expected from the newest DNA sequence results (Chombard et al.,
Irregularly reticulate or confused skeletal arrangements are not 1999; Alvarez et al., 2000a). However, until sponge DNA data have
uncommon. Spicular density is relatively low compared to other gained sufficient powers of conviction (e.g., multi-gene analyses),
Halichondrida. Spicules predominantly styles, often of consider- we propose to leave the family classification intact. Dictyonellidae
able length and then flexuous or curved. Occasionally oxeas or are clearly differentiated from Halichondriidae in the absence of
strongyles occur. Telescoped endings are common. Ten genera are a surface skeleton and the presence of some degree of organization
considered valid of seventeen nominal genera. The family occurs of the choanosomal skeleton (lacking in Halichondriidae). From
in all three ocean basins, but appears absent from the colder Axinellidae it is differentiated by the fleshy surface appearance
regions. (velvety in Axinellidae produced by the close-meshed endings

773
774 Porifera· Demospongiae • Halichondrida • Dictyonellidae

of the choanosomal skeleton). From Desmoxyidae they are differen- contents of the family, the genera Phakettia, Stylissa, and Svenzea
tiated by the absence of a surface skeleton of (spined) spicules and have been added in this current revision; [Dactylella] was found to
also the fleshy surface appearance. Compared to Van Soest et al.'s be preoccupied and is replaced by its junior synonym Lipastrotethya.

KEY TO GENERA

(1) Surface muddy-sandy with fine grooves making a polygonal surface pattern .......................................................................... Liosina
Surface not muddy and grooved ........................................................................................................................................................... 2
(2) Spicules include sinuously bent strongyles or strongyloxeas .............................................................................................. Acanthella
Spicules may be flexuous but are not bent twice in opposite directions ............................................................................................... 3
(3) Spicules include oxeas or oxeotes ........................................................................................................................................................ 4
Spicules exclusively styles ................................................................................................................................................................... 5
(4) Skeleton lax, low spicular density, spicule length less than 650 fLm .................................................................................. Rhaphoxya
Skeleton dense, with high spicular density and, spicules over 650 fLm .................................................................. ....... Lipastrotethya
(5) Next to long thin smooth styles there are (often rare) short spined styles ............................................................................ Tethyspira
No spined styles ................................................................................................................................................................................... 6
(6) Encrusting with styles arranged in dendritic fibres originating from a basal spongin plate ................................................. Scopalina
Growth form massive or erect. Styles differently arranged .................................................................................................................. 7
(7) Styles arranged in sheets or columns with considerable spongin ........................................................................................................ 8
Styles arranged in a reticulation or intercrossing in confusion ............................................................................................................ 9
(8) Surface smooth, glistening, bluntly conulose, consistency fairly soft ................................................................................ Dictyonella
Surface bristly, consistency hard ............................................................................................................................................ Phakettia
(9) Reticulation unispicular, regular ............................................................................................................................................... Svenzea
Reticulation vague, often somewhat confused ........................................................................................................................... Stylissa

ACANTHELLA SCHMIDT, 1862 Remarks

Synonymy Acanthella has been referred to Axinellidae by contemporary

authors (e.g., Bergquist, 1970; Levi, 1973; Hooper & Levi, 1993b;
Acanthella Schmidt, 1862: 65. Alvarez et al., 1998; Alvarez et al., 2000a) who adopted vague
definitions of the genus with elements such as the presence of an
Type species axial skeleton, typical of members of the family Axinellidae.
Consequently, Acanthella has been confused with other axinellid
Acanthella acuta Schmidt, 1862: 65 (by subsequent designa- genera such as Phakellia, and contains a heterogeneous complex of
tion; Vosmaer, 1912: 312). species. Acanthella sensu stricto does not fit in the revised defini-
tion ofAxinellidae (see chapter on Axinellidae) and therefore it is
Definition allocated to Dictyonellidae, as suggested by Van Soest et al.
(1990). The habit and skeletal organisation of the type species is
Dictyonellidae with choanosomal skeleton of dendritic tracts quite distinctive, sharing with other dictyonellid genera such as
cored by interwoven sinuously bent strongyles or strongyloxeas Dictyonella and Scopalina, the dendritic nature of the choanosomal
and echinated by straight styles, oxeas or anisoxeas. skeleton, with a central axial column producing dendritic ally
branching spicule tracts and thinning toward the periphery, and the
Diagnosis fleshy appearance of the surface.

Erect, bushy, ramose, lamellate or lobate habit. Surface gener- Previous reviews
ally coarsely conulose with protruding choanosomal spicules.
Ectosome membranous, glistening in appearance, without specialised Vosmaer (1912); de Laubenfels (1936a); Levi (1973); Van
skeleton. Consistency cartilaginous-like when alive. Choanosomal Soest et al. (1990); Hooper & Levi (1993b).
skeleton with thick tracts or axes of interwoven strongyles or strongy-
loxeas, sometimes condensed in the axial region, dendritically Description of type species
branching toward the periphery and ending in conules at surface;
reduced to single spicules in some thinly branching forms. Acanthella acuta Schmidt, 1862 (Fig. lA-E).
Megascleres doubly flexed and sinuously curved strongyles, some- Material examined. Lectotype: BMNH 1867.7.26.66 -
times strongyloxeas, always coring main spicules tracts; straight Sebenico, Adriatic. Other material. ZMA POR. 14589 - NW
styles, oxeas or anisoxeas in any combination, echinating main tracts; Spain, SYMBIOSPONGE 98/SP/APR121RG/024, depth 16m,
microscleres absent. Distribution. According to Van Soest (1994) the colI. R. Gomez.
genus has a widespread distribution. A total of 22 species of Description. Bush-shaped, type specimen approximately
Acanthella are recorded in the literature from the following areas: 12 cm wide and 13 cm high; on short peduncle; surface conulose;
Central-West Atlantic, Mediterranean Sea, Indian Ocean, Australia, marked with thick spicule tracts and covered with tough and
New Zealand, New Caledonia and Japan, (Alvarez 1998). smooth skin-like collagenous membrane. Oscules not seen; colour
Porifera· Demospongiae • Halichondrida • Dictyonellidae 775

Fig. 1. Acanthella acuta Schmidt, 1862. A, lectotype BMNH 1867.7.26.66 (scale 1 cm). B, In situ photo of ZMA POR. 14589 from NW Spain (photo
MJ. de Kluijver). C, drawing of skeletal structure (scale 1 mm). D, SEM photos of sinuous strongyle (scale 10 fLm) and detail of apices (scale 1 fLm) from
ZMA POR. 14589. E, ditto of style (scale 10 fLm) and details of apices (scale I fLm).

dark orange alive, brown-orange in dry state; consistency firm Type species
(cartilage-like when alive) (Fig. lA-B). Specialised ectosomal
skeleton absent; choanosomal skeleton with ascending, thick (up to Dictyonella cactus Schmidt, 1868: 10 (by subsequent desig-
500/-Lm) and dendritic axes of sinuous strongyles plumo-echinated nation; de Laubenfels, 1936a: 139) (=junior synonym of
by styles, becoming thinner towards periphery; axial and extra- Acanthella obtusa Schmidt, 1862: 65).
axial skeleton not differentiated (Fig. I C). Sinuous strongyles
(Fig. ID), 640-1490 X 5-13 /-Lm; flexuous and straight styles, Definition
970-2150 X 8-15 /-Lm (Fig. IE). Distribution and ecology. The
species is restricted to the Mediterranean, found often between Dictyonellidae with long styles in anastomosing and diverg-
40-100 m depth, in shallow and dark habitats, on sandy or coralline ing bundles progressively thinning out towards the surface.
substrate (Uriz, 1982).
Diagnosis

Thick-lobed bushes or irregular massive sponges with shiny

DICTYONELLA SCHMIDT, 1868 smooth surface covered with blunt ridges and conules. Ectosomal
skeleton absent, choanosomal skeleton consisting of anastomosing
Synonymy and diverging spicule bundles arranged lengthwise in the lobes and
branches. Spicules long, often curved or flexuous styles. Little
Dictyonella Schmidt, 1868: 10. Phacanthina Vosmaer, 1912: spongin. Approximately 15 species spread over the warmer seas of
313. Perisinella Topsent, 1928c: 171. the world.
776 Porifera· Demospongiae • Halichondrida • Dictyonellidae

Fig. 2. Dictyonella spp. A-F. Dictyonella obtusa (Schmidt, 1862), A, watercolour of type specimen, reproduced from Schmidt's, 1862 pI. VI fig. 8 (scale
I cm). B, habits of specimens from Naples, reproduced from Topsent (1925: pI. VIII figs 2-4 as Stylotella plicata) (scale I cm). C, drawing of skeleton
(scale I mm). D, drawing of spicules (scale 100 !Lm). E, SEM photo of spicule apices made from the type specimen of Dictyonella cactus (junior synonym
of D. obtusa) (scale l!Lm). F, SEM photo of skeletal bundles made from ditto (scale 250 !Lm). G-H, Dictyonella madeirensis Topsent, 1928c, type of
Perisinella. G, habit reproduced from Topsent, 1928c: pI. II fig. 12 (scale I cm). H, spicule and variation of apices (scale 100 !Lm).

Remarks madeirensis Topsent (1928c: 170, pI. II fig. 12, pI. VI fig. 13). This
East Atlantic species forms a yellowish mass of digitations on a coni-
Due to Schmidt's poor definition and description, this genus cal basis, 12 cm high, 11 cm in diameter (Fig. 2G). The skeleton con-
was not employed until Topsent (l938a) redescribed the type. Prior sists of spongin-encased dendritic spicule tracts forming an irregular
to him, Vosmaer (1887, 1935) gave an erroneous interpretation to it meshwork. No ectosomal skeleton. Spicules (Fig. 2H) are a mixture
and few authors used the name. Of the five specimens of the original of styles, subtylostyles and occasional tylostyles, 400--700 X 2-3 /-Lm.
Dictyonella collection from Algiers, only the specimen labeled Carballo et al. (1996) assigned D. madeirensis to the genus
'no. 10' appeared with certainty to conform to Schmidt's descrip- Scopalina, based on the abundant spongin, shared with Scopalina
tion. The second species of Dictyonella described by Schmidt lophyropoda (cf. below). However, in view of the thinly encrusting
(1868) along with D. cactus, viz., Dictyonella labyrinthica, is a habit and solitary, non-anastomosing fibres of Scopalina this is
specimen of Myxilla prouhoi according to Topsent (1938a). In addi- considered erroneous.
tion to the type specimen mentioned above, there is a microscope The genus Stylotella Lendenfeld (1888) was used for sponges
slide in MNHN registered as DT. 940 (Desqueyroux-Faundez & answering to the present concept of Dictyonella by many authors
Stone, 1992). A slide in BMNH 1867.3.11.77 labeled 'Acanthella prior to Topsent's (1938a) redescription and redefinition of the
obtusa 49 Adriatic' clearly contains spicules of Acanthella acuta. genus. Stylotella is a junior synonym of Hymeniacidon (Halichon-
Type material proper of Acanthella obtusa appears to be lost. It is driidae) as it possesses a tangential skeleton and the spicules are
proposed here to assign above mentioned lectotype specimen of small styles (see chapter on Halichondriidae).
Dictyonella as the lectotype of Acanthella obtusa. Several members of Dictyonella, e.g., D. incisa (Schmidt,
The genus Phacanthina Vosmaer (1912: 313, pI. 16 figs 1-2) 1880a), D. marsilii (Topsent, 1893c), were assigned to the genus
was erected for type species Acanthella obtusa Schmidt, 1862 Rhaphidostyla Burton (1935b), but the type species R. kitchingi
(original designation). Since this is considered the senior synonym Burton, 1935b is a Hymeniacidon (see chapter on Halichondriidae).
of the type species of Dictyonella, Phacanthina becomes an
objective synonym. Previous review
The genus Perisinella Topsent (1928c: 171) was erected as a
subgenus of Stylotella for type species (by monotypy) Stylotella Topsent (1938a).
Porifera • Demospongiae • Halichondrida • Dictyonellidae 777

Description of type species of this genus remains tentative in the absence of any definite
synapomorphies with either Halichondrida and Poecilosclerida.
Dictyonella obtusa (Schmidt, 1862) (Fig. 2A-F). The possession of iregularly arranged oxeotes in a low density and
Synonymy. Acanthella obtusa Schmidt, 1862: 65, pI. VI the occurrence of monactinal spicules in a species showing strong
fig. 8. Dictyonella cactus Schmidt, 1868: 10. Phakellia pUcata overall similarities with the type species both point towards
Schmidt, l880a: 282. Halichondrida, and within that order, to Dictyonellidae. Future
Material examined. Lectotype (designated by Topsent, studies using non-morphological characters need to confirm this.
1938a: 9): MNHN unregistered - labeled 'No. 10, Exp. Algerie'. The genus [Migas] Sollas, 1908 was erected for type species (by
BMNH slide 1868.3.2.27 - labeled 'Algiers' (this contains just monotypy) Migas porphyrion Sollas (1908: 395, figs 1-3) from
a few broken spicules, probably from fairly long styles). Mozambique. The type material could not be found, but from the
Description. Lobate or leafy bushes (Fig. 2A-B) of up to description it is clear that it belongs to Liosina. With the surface orna-
7 cm high (lectotype is 5 cm high); smaller specimens may be mentation and foreign material inclusion, and oxeas of 960 X l6/Lm,
massive-lobate. Colour red alive, pale violet in alcohol. There is a it is very close and probably identical to Liosina paradoxa. Thus,
distinct thick organic 'skin', stretched over the endings of the [Migas] Sollas is a junior synonym of Liosina (see also de Laubenfels,
choanosomal spicule tracts. This comes off in flakes (Schmidt 1936a: 54). De Laubenfels, 1954, however, discovered that [Migas]
states 'Oberhaut lasst sich in Fetzen abziehen'). The choanosomal Sollas, 1908 was preoccupied by Migas Koch, 1873 (Arachnida). He
skeleton consists of bundles of megascleres (Fig. 2C, F) that are had apparently forgotten that he synonymized Migas Sollas with
generally directed upwards toward the periphery, but they anasto- Liosina in 1936, and erected the genus Milene de Laubenfels, 1954:
mose and diverge along the way in a loose manner. The bundles 116 with Migas porphyrion as the type species (by original designa-
thin out towards the surface where they may form low conules. tion). Milene, thus becomes an objective synonym of Liosina.
Binding spongin is visibly present especially in the interior.
Spicules (Fig. 2D-E) long flexuous styles, occasionally with Description of type species
strongylote or telescoped endings, up to 1700 X 17 /Lm. Distribu-
tion and ecology. Mediterranean, shallow-water. Liosina paradoxa Thiele, 1899 (Fig. 3A-E).
Synonymy. Liosina paradoxa Thiele, 1899: 17, pI. 2 fig. 5,
pI. 4 fig. 4, pI. 5 fig. 9. Migas porphyrion Sollas, 1908: 395; Milene
UOSINA THIELE, 1899 porphyrion; de Laubenfels, 1954: 116.
Material examined. Holotype: NMB 22 - Celebes, Sarassin
Synonymy collection. Other material. Many topotypical specimens, including:
ZMA POR. 14478, SYMBIOSPONGE 98/SS/APR291BHl031 -
Liosina Thiele, 1899: 17. [Migas] Sollas, 1908: 395 (preocc.). off Langkai Island, SW Sulawesi, 4m, coIl. B.W. Hoeksema.
Milene de Laubenfels, 1954: 116. Description. The holotype (Fig. 3A) is an upright mass
(which is likely to be part of a tube), topotypical and other speci-
Type species mens are clusters of thick-walled tubes with narrow apical open-
ings. The surface has a characteristic polygonal pattern of grooves
Liosina paradoxa Thiele, 1899: 17 (by monotypy). and ridges (Fig. 3B). Grooves consist of thin lines connecting
small rounded openings. Colour muddy grey or whitish grey, with
Definition an orange interior shining through the grooves and openings.
Consistency firm, but easily damaged. Ectosomal region a crust of
Dictyonellidae with a sandy-muddy surface divided into polyg- reddish brown, peculiar-looking mud particles enclosed in sponge
onal areas; skeleton loose bundles of oxeote or stylote spicules. tissue membranes. The ectosomal mud particles form a dense layer
of about 1 mm thickness through which may protrude bundles of
Diagnosis spicules or individual spiCUles (Fig. 3E). No special ectosomal
skeleton. The choanosome likewise has a high density of mud par-
Tube-shaped, massive or encrusting sponges with a character- ticles, partly covering and engulfing irregular bundles of spicules,
istic muddy surface and polygonal surface pattern caused by a sys- which are bound by considerable spongin. Bundles vague in out-
tem of thin-walled shallow grooves. Mud particles enclosed in line and of indefinite width. Loose spicules abound, but spiculation
sponge tissue form a dense surface cover and also penetrate into has an overall low density. Spicules (Fig. 3C-D) with mostly
the interior. Spicules relatively low in density, irregularly arranged rounded ends, basically oxeas, but stylote forms not infrequent,
into bundles which are bound by variable amount of spongin. in a wide size range: 360-900 X 8-20 /Lm. No microscleres.
Spicules may be both monactinal and diactinal. So far, only two Distribution and ecology. Indo-West Pacific, widespread, common;
species have been assigned to this genus, both widespread in the shallow-water, on reefs and in lagoons and bays.
Indo-West Pacific.

Remarks UPASTROTETHYA DE LAUBENFELS, 1954: 235

The muddy appearance and the mud particles form an easily Synonymy
recognizable feature of this species, and they are also found in the
species originally described as Laxosuberites arenosus by Vacelet & [Dactylella] Thiele, 1898: 56 (preocc. by [Dactylella] Gray,
Vasseur (1971), which however has tylostyles. Nevertheless, this 1873: 461, Tetillidae, nomen oblitum). Lipastrotethya de
species is reassigned to Liosina. The family and class assignment Laubenfels, 1954: 235.
778 Porifera· Demospongiae • Halichondrida • Dictyonellidae

Fig. 3. Liosina paradoxa Thiele, 1899. A, habit of holotype 5MB 22 reproduced from Thiele's, 1899 pI. 2 fig . 5 (size see text). B, photo of freshly col-
lected specimen from the reefs of the Seychelles. C, oxea (scale 100 fLm). D, details of oxea apices (scale 10 fLm). E,SEM image of section of peripheral
region to show mud particles penetrated by spicule bundles (scale 100 fLm).

Type species Halichondriidae), but in that genus the spicules are confusedly
arranged, lacking a vertical orientation.
Lipastrotethya ana de Laubenfels, 1954: 235 (by monotypy). The characters of this sponge conform to [Dactylella] Thiele
and Lipastrotethya would seem a junior synonym. The type species
Definition (by monotypy) is Dactylella hilgendorfi Thiele, 1898: 56, pI. 4
fig. 8, pI. 5 fig. 25, pI. 8 fig. 41 a-b. A fragment of the holotype
Dictyonellidae with exclusively oxeas or derivates thereof. 2MB 920a from Hokodate, Japan and a further fragment of holo-
type BMNH 1908.9.24.143 were reexamined. From a shared base
Diagnosis arise 4-5 finger-shaped projections (Fig. 4B), about 4cm long,
1.5 cm diameter. The fingers may have lateral lobes. The ends of
Digitate sponges with microhispid, finely grooved surface. the projections are distinctly grooved, and grooves converge into
Ectosomal skeleton absent. Choanosomal skeleton a bundled mass an apical oscule. Surface slightly rugose, microhispid due to pro-
of oxeas. No microscleres. jecting spicule bundles. The colour is pale brown in preserved con-
dition. Consistency firm. No ectosomal skeleton. Choanosomal
Remarks skeleton (Fig. 4D) a mass of bundles of aligned spicules traversing
the centre of the projections and diverging outwards. High spicular
This species sounds close to Dictyonella on paper, but in density, little spongin. Spicules (Fig. 4C) oxeas, straight or slightly
that genus the spicules are invariably styles or monactinal curved, 1000-1600 X 20-25 flm. Thiele states they are not infre-
derivatives. Also, spongin is usually abundantly present. On quently flexuous, but in the fragment examined this is hardly the
paper it also sounds close to Axinyssa Lendenfeld, 1897 (family case. Also the proportion of stylotes is negligible. A second record
Porifera· Demospongiae • Halichondrida • Dictyonellidae 779

A c
Fig. 4. Lipastrotethya. A, Lipastrotethya ana de Laubenfels, 1954, drawing of spicules, reproduced from de Laubenfels, 1954: fig. 162.
B-D, Lipastrotethya hilgendorfi (Thiele, 1898 as [Dactylella]). B, drawing of ho1otype reproduced from Thiele's, 1898 pI. 4 fig. 8. C, drawing of spicules
from the same, pI. 8 fig. 4Ia-b. D, photo of cross section from the same pI. 5 fig. 25 (for sizes see the text).

of Dactylella is from the Caribbean, D. rhaphoxea de Laubenfels, ill-defined bundles of oxeas arranged perpendicular to the surface
1934. It conforms in some detail to D. hilgendorji, but has a much and ending in the surface tubercles. The ectosomal region is
larger range of spicule sizes, including small ectosomal oxeas. Its described as corticate by de Laubenfels, but this is not a cortex in
identity should be further studied. the usual sense; rather it is an ectosomal layer containing special
This genus [Dactylella] Thiele is preoccupied by [Dactylella] granular cells and ?calcareous nodules. The spicules (Fig. 4A) are
Gray, 1873: 461, which was erected for type species Tethya symmetrical oxeas, although some of them are blunt-ending at both
dactyloidea Carter, 1869a: 15, figs 1-4 from the SE coast of Arabia sides; they are curved, and occasionally flexuous; in a wide size
(by original designation). This is undoubtedly a member of range, up to 850 X 25 fJ-m. They are clearly not strongyloxeas as
Tetillidae, because Carter described a radiate structure and oxeas de Laubenfels maintained, and the resemblance to Tethya megas-
and protriaenes as megascleres. No certainty exists about the pres- cleres he made is absent. Distribution. Chuuk, Central Pacific,
ence of sigmaspires, because Carter had 'given away the specimen' shallow water.
(to Dr Bowerbank, cf. Carter, 1871: 103). In view of the absence of
clear porocalyces and apparent lack of a cortical skeleton, this is
likely to be a Tetilla. Since the name Dactylella predates Tetilla it PHAKETTlA DE LAUBENFELS, 1936
needs to be suppressed as an unused name in the sense of ICZN
article 23.9. The name has not been used for a tetillid sponge after Synonymy
its original proposal, although subsequently, [Dactylella] was also
used by Thiele, 1898 for a sponge in the family Dictyonellidae Phakettia de Laubenfels, 1936a: 130.
(now considered preoccupied, and replaced by Lipastrotethya de
Laubenfels, 1954). The name Tetilla has been frequently used in Type species
the past 50 years by many authors, and thus fulfills the demands for
continued usage under ICZN Article 23.9. But, the subsequent use Phakellia cactoides Burton, 1928b: 127 (by original
by Thiele of [Dactylella] for a dictyonellid sponge is nevertheless designation).
against ICZN rules and it has to be replaced by the present
genus name. Definition

Previous review Dictyonellidae with a central region of long styles aggregated

in a confused and dense mass, from which they radiate, some-
Thiele, 1898 (as [Dactylella]). times in thick bundles, to the periphery and project through the
surface.
Description of type species
Diagnosis
Lipastrotethya ana de Laubenfels, 1954 (Fig. 4A).
Synonymy. Lipastrotethya ana de Laubenfels, 1954: 235, Erect on broad base or short stems, bushy or with coalescent
fig. 162. lobes. Surface bristly. Ectosomal skeleton absent. Choanosomal
Material examined. Holotype: USNM 23094 (fragment) - skeleton with a central region of styles aggregated in a confused
Truk Atoll (Chuuk State), Federated States of Micronesia. and dense mass from which other styles, sometimes in thick bun-
Description (after de Laubenfels, 1954). The species is dles radiate to the periphery. Megascleres are only long styles.
described as an irregularly rounded mass of up to 6 X 8 X 16cm, According to de Laubenfels (1936a) it is widespread, but species
golden brown in colour with a slightly greenish, cartilaginous assignments need to be verified. The type species is from the
subcutaneous layer. Surface tuberculate. Skeleton consists of Indian Ocean.
780 Porifera· Demospongiae • Halichondrida • Dictyonellidae

Fig. 5. Phakettia cactoides (Burton, 1928b). A, ho1otype BMNH 1926.10.1.38 (left specimen) and paratypes, at least one of which is presumably located
in the Indian Museum, Calcutta, photo reproduced from Burton, 1928b: pI. I figs 6-8 (scale 5DOfLm). B, skeleton (scale 250fLm). C, drawing of style
(scale 1DO fLm).

Remarks RHAPHOXYA HALLMANN, 1917

Several species of Phakellia (i.e., Phakellia beringensis Synonymy

Hentschel, 1929: 975, P. bowerbankiVosmaer, 1885a: 24, P. crassa
Carter, 1885d: 363, P. conulosa Dendy, 1922b: 116, P. datli Lambe, [Acanthellina] Carter, 1885d: 365 (nomen oblitum).
1894: 125, P.fusca Thiele, 1898: 51, P. papyracea Ridley & Dendy Rhaphoxya Hallmann, 1917c: 643.
1886: 478, P. ridleyi Dendy, 1887: 159; P. tumida Dendy, 1897)
were referred to Phakettia by de Laubenfels (1936a). The assign- Type species
ment of these species to Phakettia however, must be revised, as in
most cases, they do not fully agree with the definition proposed Rhaphoxya typica Hallmann, 1917c (by original designation).
here. Hooper & Levi (1993b) included Phakettia in synonymy with
Rhaphoxya, but this is not accepted here, with the two respective Definition
type species showing significant differences in their spicule geom-
etry (long styles vs. anisoxeas and styles) and structure (confused Dictyonellidae with anisoxeas and styles, single or in loose
central mass with radial periphery vs. halichondrioid with loose wavy and anastomosing bundles, ascending towards the surface
wavy anastomosing tracts ascending to the periphery). and reinfOlced with spongin.

Previous reviews Diagnosis

De Laubenfels (1936a); Wiedenmayer (1989). Massive to thickly flabellate; surface shiny and conulose.
Ectosome fleshy without special skeleton. Choanosome cavernous with
Description of type species a lax skeleton of wavy and anastomosing bundles of spicules reinforced
with spongin. Spicules are anisoxeas and styles with telescoped ends.
Phakettia cactoides (Burton, 1928b) (Fig. 5A-C). Raphides and trichodragmata may be present. Distribution of the genus
Synonymy. Phakellia cactoides Burton, 1928b: 127, pI. I exclusively Australia. Seven nominal species.
figs 6-8.
Material examined. Holotype: BMNH 1926.10.1.38 - wet. Remarks
Paratype: BMNH 1926.10.1.33 -labeled 'Co-type, RN. VIl (Pars),
donated by the Indian Museum'. The genus has been referred to Axinellidae by other authors
Description. Bushy, lamellate or lobate on a broad base. (e.g., Bergquist, 1970; Wiedenmayer, 1989; Hooper & Levi,
The type series apparently consisted of three specimens (Fig. 5A). 1993b); however, as Hooper and Levi (1993b) admitted, the genus
The holotype (Fig. 5A, left specimen) is 2.5 cm high. Colour in life is atypical for that family. Based on its close similarity to dictyonel-
unknown, beige in alcohol. Surface hispid, due to the projection of lid genera such as Dictyonella and Scopalina, it is transferred here
single spicules or spicule tracts from the choanosomal skeleton to the Dictyonellidae. Rhaphoxya shares with other dictyonellids
with smooth depressions as channels running longitudinally the type of spicules and the presence of bundles of spicules rein-
through the length of the specimen. Consistency hard; fleshy forced with spongin or, spongin fibres cored with spicules, single or
appearance. Ectosomal skeleton absent. Choanosomal skeleton in loose tracts. The presence of long raphides (or thin oxeas) with
formed by long styles aggregated in a confused and dense mass in rudimentary spination in the type of Rhaphoxya may be peculiar to
the central region ofthe skeleton (Fig. 5B). From the central skele- the type species and not a diagnostic feature to define the genus.
ton single styles or thick bundles radiate in all directions to the [Acanthellina] Carter, 1885d was erected for type species
periphery and project through the surface. Megascleres exclusively (by monotypy) Acanthellina rugolineata Carter, 1885d: 365
smooth styles (Fig. 5C), 1850-2800 X 10-33/J-m. Distribution and (Fig. 6D-F) from South Australia. The genus was considered a
ecology. Bay of Bengal, Indian Ocean, 174m. junior synonym of Acanthella by de Laubenfels (1936a). Bergquist
Porifera· Demospongiae • Halichondrida • Dictyonellidae 781

I
F

Fig. 6. Rhaphoxya spp. A-C, Rhaphoxya typica Hallmann, 1917c, largest portion of the fragmented holotype AM Zl595 (scale I em). B, photo of thick
section of peripheral region of the holotype (scale 500 j.Lm). C, spicules consisting of anisoxea, details of apices and raphide (scale 100 j.Lm).
D-F, Rhaphoxea rugolineata (Carter, 1885d as Acanthellina). D, photo of holotype, reproduced from Hooper & Levi (1993b: fig. 32). E, photo of thick
section of peripheral skeleton, reproduced from ditto (scale 500 j.Lm). F, spicules (scale 100 j.Lm).

(1970: 18), however, considered its type species to belong in Description. Massive; holotype fragmented into nine
Rhaphoxya. The genus was later declared a nomen oblitum under pieces, 2-10 cm of maximum diameter (Fig. 6A); surface with
the 50 year rule of the ICZN by Wiedenmayer (1989: 50) (also in shallow (3.5 cm deep) and narrow (4.5 cm wide) grooves and scat-
Hooper & Levi, 1993b: 1441; Hooper & Wiedenmayer, 1994: 82), tered conuli-papillae 1 mm high. Ectosome membranous, detach-
and we propose here to follow this course (ICZN article 23.9.1; able, reticulated with circular-ovoid meshes, 100-200/-Lm in
Anon., 1999). The skeleton (Fig. 6E) and other features are in diameter; colour brown in alcohol; consistency firm. Ectosomal
agreement with the definition of Rhaphoxya as employed here. The skeleton absent. Choanosomal skeleton halichondrioid (Fig. 6B)
genus includes seven species (i.e., Rhaphoxya cactifonnis Carter, with single spicules, in bundles or in vague tracts ascending
1885a: 114, R. felina Wiedenmayer, 1989: 52, Rhaphoxya pallida towards surface or irregularly positioned, enveloped by sheaths
Dendy, 1897: 257, R. rugolineata Carter, 1885d: 365 (junior or slightly developed fibres of spongin. Megascleres (Fig. 6C)
synonym of R. cactifonnis according to Bergquist (1970) and are flexuous anisoxeas with telescoped ends of different shapes,
Wiedenmayer (1989», R. solida Carter, 1885d: 362 and R. typica. and less frequent styles; length 320-620 /-Lm, width 5 /-Lm. Also
raphides, single or in trichodragmata (Fig. 6C, right), smooth or
Previous reviews microspined, some with crenulate ridges and small tubercles.
Distribution and ecology. Eastern and southern Australian waters
Hallmann (1917c); de Laubenfels (1936a); Bergquist (1970); (Hooper & Wiedenmayer, 1994).
Wiedenmayer (1989); Hooper & Levi (1993b).

Description of type species

SCOPAUNA SCHMIDT, 1862
Rhaphoxya typica Hallmann, 1917c (Fig. 6A-C).
Synonymy. Rhaphoxya typica Hallmann, 1917c: 643. fig. 17, Synonymy
pI. 29 fig. 3, pI. 38 figs 8-9, pI. 39 fig. 5, pI. 42 figs 1-2.
Material examined. Holotype: AM Z1595 - wet, Port Scopalina Schmidt, 1862: 79. Hoplochalina Lendenfeld,
Phillip, Victoria, Australia. 1887c: 822.
782 Porifera· Demospongiae • Halichondrida • Dictyonellidae

Fig. 7. Scopalina lophyropoda Schmidt, 1862. A-B, In situ photos from Marseille (photo B. Picton) and NW Spain (photo M.J. de Kluijver). C, SEM
photo of details of style (scale 10 iJ.m). D, SEM image of peripheral skeleton to show sheet-like spicule-fibre made from ZMA POR. 14434 (scale
100 iJ.m). E, Schmidt's (1862) drawing of the skeletal fibers, reproduced from his pI. VII fig. 18 (size see text).

Type species and the Indo-Pacific S. australiensis (Pulitzer-Finali, 1981), and

thus may be diagnostic.
Scopalina lophyropoda Schmidt, 1862: 79 (by monotypy). The genus Hoplochalina Lendenfeld, 1887c: 822 was
erected for type species H. incrustans Lendenfeld, 1887c: 823
Definition (subsequent designation by Van Soest, 1980: 118; confirmed by
Hooper & Wiedenmayer, 1994). The type species designation is
Encrusting Dictyonellidae with conulose surface caused by based on the fact that it is the first species mentioned by
prominent dendritic spongin fibres cored by long styles. Basal Lendenfeld, but it is admittedly rather unfortunate, as the genus
spongin plate. Consistency soft. definition given by Lendenfeld and the other species described
by him (H. dendrilla, H. fenella, and H. renieroides) are unlike
Diagnosis H. incrustans. De Laubenfels (l936a: 138) clearly adopted a dif-
ferent view of Hoplochalina, emphasizing the possession of
Encrusting sheets with surface covered in sharp conules. Soft oxeas, rather than styles. However, de Laubenfels did not make a
consistency. Ectosomal skeleton absent. Choanosome characteris- type species designation, so his remarks are invalid. The type of
tically grainy in appearance. Choanosomal skeleton consisting of Hoplochalina incrustans was described as a conulose rosy-red
bundles of thin styles entirely enclosed in spongin. Fibres dendritic crust, 4 mm in height, with conules of 1.5 mm high. A slide
rising up from a basal spongin plate. Low spicular density and soft of the type of H. incrustans BMNH 1886.8.27.521 from Port
consistency. Half a dozen species distributed over all warmer parts Jackson was re-examined and found to conform closely to
of the world oceans. Scopalina, including dendritic spongin fibres of approximately
200/-Lm diameter, containing tracts of long styles, 600 X 10 /-Lm,
Remarks and the grainy appearance of the choanosome. It may be close,
and even con specific with Dictyonelia australiensis Pulitzer-
Scopalina lophyropoda is a characteristic species and similar Finali, 1981.
forms are common in all three ocean basins. In the Mediterranean-
Atlantic area, at least three species appear to exist: S. lophyropoda, Description of type species
S. azurea Bibiloni, 1993 and an unnamed species from
Canary Islands (Cruz, 1986, pers. comm.). Carballo et al. (1996) Scopalina lophyropoda Schmidt, 1862 (Fig. 7A-E).
erroneously assigned Dictyonella madeirensis to Scopalina Synonymy. Scopalina lophyropoda Schmidt, 1862: 79,
(see above). The common West Indian Scopalina ruetzleri pI. VII fig. 18.
(Wiedenmayer, I 977b) was originally assigned to Ulosa de Material examined. Lectotype: LMJG 15117 - Sebenico,
Laubenfels, 1936a, but that mycalid genus has a reticulate skeleton Adriatic. Other material. Several topotypical specimens including:
and is quite unlike Scopalina. The grainy appearance of the ZMA POR. 14434 - NW Spain, SYMBIOSPONGE 98/SP/APRI2/
choanosome is characteristic of at least S. lophyropoda, S. ruetzleri, MKl028, 9m, coil. M.J. de Kluijver.
Porifera· Demospongiae • Halichondrida • Dictyonellidae 783

Description. Thinly encrusting with characteristic sharp Topsent, 1897a or Stylotella aurantium Kelly-Borges & Bergquist,
conules. In life the orange to dark red sponges appear fleshy, 1988), and S. variabilis (Whitelegge, 1907 as Spongosorites). A fur-
pumped up, but in preserved or dry condition the sponge collapses ther prominent species of this genus is Stylissa carteri (Dendy,
to form pale beige, fleshy crusts. Surface layer without spicules, 1889), a.k.a. Acanthella carteri or Axinella carteri (with junior syn-
thickly organic with characteristic grainy appearance caused by onyms Acanthella ehrenbergi Keller, 1889 and Acanthella auranti-
clusters (20-40 fLm in diameter) of ? pigment granules. aca Keller, 1889). Hooper & Levi (1993b) assigned to the type
Choanosomal skeleton consists of spongin fibres rising up from species specimens from New Caledonia which appear to belong to
a basal spongin plate covering the surface. The fibres consist of Stylissa carteri. This differs from S. flabelliformis in having
layered spongin with a central core of 1-4 spicules. Fibre system a stalked thickly lobate to flabellate growth form with strongly
dendritic, i.e., they are singly rising towards the periphery or are sub- papillate-conulose surface. Hooper & Levi (1993b) proposed that
dividing, but do not anastomose. Length of fibres at least 1 mm, fre- the genus Stylissa be considered a senior synonym of Dragmaxia,
quently longer when subdividing, lying at distances of 20~500 fLm. but that genus has long sinuous trichodragmata.
Thickness of fibres 5~80 fLm at the base tapering to 30-40 fLm at
the periphery, where they often end with single spicules sticking Description of type species
out of the fibres. Spicules styles, flexuous, often curved in the
lower half, 950-1200 X 12-15 fLm. Distribution and ecology. Stylissaflabelliformis Hentschel, 1912 (Fig. 8A-F).
Mediterranean-Atlantic, possibly extending as far south as the Synonymy. Stylotella flabelliformis Hentschel, 1912: 355,
Cape Verde Islands. pI. XIX fig. 26; Stylissa flabelliformis; Hallmann, 1914: 349
(not Hooper & Levi, 1993b: 1422 =Stylissa carteri).
Material examined. Lectotype (designation by Hooper &
STYUSSA HALLMANN, 1914 Levi, 1993b: 1422, as holotype): SMF 1587; schizotype fragment:
MNHN DCl. 2214); paratypes: SMF 1527, 1534, 1538, 1689, 1690-
Synonymy all from Am Islands, Indonesia. Other material. ZMA POR. 14486-
SW Sulawesi, Indonesia, SYMBIOSPONGE 98/SSIMAYOll
Stylissa Hallmann, 1914: 349. BHl039, off Kappoposang Island, 31 m, vertical wall, colI. B.W.
Hoeksema.
Type species Description. Thinly flabellate, with side-blades in the larger
specimens. The lectotype (Fig. 8A) measures 10 cm high and 5 mm
Stylotella flabelliformis Hentschel, 1912: 355 (by original thick. It is probably a fragment of an originally much larger speci-
designation). men, because Hentschel mentions a specimen of 20 em high, and
recent specimens may be even larger (Fig. 8B). There is no genuine
Definition stalk although the sponge narrows towards the base, but distinct
thickened 'ribbed veins' consisting of thick spicule tracts traverse
Dictyonellidae with a skeleton consisting of styles arranged the body radially from base and subdivide and anastomose periph-
in a confused plumose reticulation, with many single spicules in erally. These ribs often bear conulose projections. In between the
confusion. Styles in a single size category. ribs, the surface is smooth and bears a few oscules. The edges of
the blade are incised and bear projections. Colour dark orange-
Diagnosis brown, beige in alcohol. Skeleton consisting of thicker and thinner
spicule tracts, bound by spongin, and confusedly plumoreticulate
Erect, flabellate, or compressed-lobate sponges with irregu- arranged single spicules (Fig. 8C-D). Spicules curved styles
larly conulose and/or ridged surface. Conules blunt. Surface (Fig. 8E-F), 45~(478)-550 X 1~(17.7)-25 fLm. Distribution and
smooth between conules often with a slight colour difference ecology. Occurring in many Eastern Indonesian reefs.
between smooth and conulose parts. Colours usually red, orange or
yellowish. Skeleton confused, but some plumose reticulation
usually recognizable. In the interior and in the stem of erect SVENZEA ALVAREZ ET AL., 2002
forms there is axial condensation. Styles curved, usually stout,
relatively short and of a single size category. Several species are Type species
common in the Indo-West Pacific, one is recorded from the
Caribbean. Pseudaxinella zeai Alvarez et al., 1998: 20 (by original
designation).
Remarks
Definition
Hallmann (1914) erected this genus because Stylotella
flabelliformis did not agree with his definition of Stylotella. The latter Dictyonellidae with a neat uni- paucispicular reticulation of
genus is now considered a junior synonym of Hymeniacidon (family styles.
Halichondriidae), as its type species has a tangential ectosomal
skeleton. De Laubenfels (1936a: 130) related Stylissa to Diagnosis
Homaxinella based only on the presence of styles. Homaxinella is
now considered a suberitid. Stylissa was revived by Hooper & Levi, Massive, cake shaped or more thinly encrusting. Surface opti-
1993b for the type species S. flabelliformis, S. massa (Carter, 1881 b, cally smooth, slightly microhispid. Soft consistency. No ectosomal
as Axinella virgultosa massa, a.k.a. Hymeniacidon conulosa skeleton. Choanosomal skeleton uni- paucispicular reticulation,
784 Porifera· Demospongiae • Halichondrida • Dictyonellidae

Fig. 8. Stylissa flabelliformis (Hentschel, 1912). A, photo of lectotype, SMF 1587 (scale 1 cm). B, ZMA POR. 14486, photographed in situ in SW
Sulawesi reefs (photo B.W. Hoeksema). C, SEM photo of choana somal skeleton (scale 500 /Lm). D, SEM photo of surface (scale 100 /Lm). E, SEM photo
of details of style (scale I/Lm). F, drawing of styles reproduced from Hentschel's, 1912 pI. XIX fig. 26 (scale 10 /Lm).

Fig.9. Svenzea zeai (Alvarez et aI., 1998). A., photo ofZMA POR. 14539 in situ on the reef of Cura~ao (photo M.J. de Klnijver) (scale I cm). B-C, draw-
ing of skeleton and spicule, reproduced from Alvarez et aI., 1998 fig. II (scales 500 /Lm and 100 /Lm). D, SEM photo of detail of style (scale I/Lm).

with some spongin at the nodes. Characteristic grainy soft parts Remarks
(similar to Scopalina). Spicules short styles and less frequent
oxeas. Reproduction viviparous, larvae huge, up to 5 mm or more, The type species was tentatively assigned to the genus
oval, entirely ciliated. Three species from the tropical Atlantic and Pseudaxinella (now Dragmacidon, see chapter on Axinellidae) by
Indo-Pacific. the original authors (Alvarez et al., 1998), but it was already noted
Porifera· Demospongiae • HaIichondrida • DictyoneIlidae 785

Fig. 10. Tethyspira spinosa (Bowerbank, 1874b). A, drawings of habit and spicules, reproduced from Bowerbank's, 1874b pI. LXXXIII figs 17-22 (sizes
see text). B, specimen photographed in situ (photo B. Picton). C-E, SEM photos made from ZMA POR. 5975. C, section of peripheral skeleton (scale
1 mm). D, spined microtylostyles (scale 10 f.Lm). E, details of styles (scale 10 f.Lm).

that it did not fit into Pseudaxinella because of the thin spicular Material examined. Holotype: USNM 39361 - Islas del
tracts and soft consistency. With the discovery of two other species Rosario, Colombia. Paratypes: ZMA 8876 - U.S. Virgin Islands.
with similar properties as P. zeai sufficient evidence was assembled USNM 42805 - Little Tobago, Trinidad & Tobago. Other material.
for the erection of a new genus Svenzea, named after the original ZMA POR. 14359 - Cura9ao, SYMBIOSPONGE 98/CUIMAY23/
discoverer of the type specimen, Prof. Sven Zea. The reassignment RG/017, 33.4m, coil. R. Gomez.
of the genus from Axinellidae to Dictyonellidae is made largely on Description. Spreading thick masses with lobes (Fig. 9A),
the circumstantial evidence presented by the characteristic mass of up to 6 cm thick and indefinite lateral expansion. Oscules chimney-
granular cells found in all three species and shared with Scopalina like, 1 cm in diameter. Surface smooth, punctate. Consistency
species. In most other respects, induding its peculiar haplosderid- crumbly, soft. Colour brown with yellow interior. Skeleton
type skeleton, Svenzea is not typical of the family Dictyonellidae. (Fig. 9B) a reticulation of spicule tracts of 1-3 spicules intercon-
nected by single spicules. In the peripheral parts of the sponge,
Description of type species extending deep into the choanosome, there is a mass of dark gran-
ular cells, obscuring the spicules and the skeleton. This is similar
Svenzea zeai (Alvarez et al., 1998) (Fig. 9A-D). to the condition observed in Scopalina. Spicules (Fig. 9C-D): short
Synonymy. Hymeniacidon amphilecta sensu Pulitzer-Finali, styles, occasionally with oxeote endings, 190-330 X 5-12.5 /-lm
1986: 117 (not de Laubenfels, 1936a). Calyx podatypa sensu less frequent oxeas with similar range. Distribution and ecology.
Humann, 1992: 33 (not de Laubenfels, 1934). Pseudaxinella (?) zeai Colombia, Bahamas, Belize, Jamaica, Cura9ao, deeper parts of
Alvarez et al., 1998: 20, fig. 11; Svenzea zeai; Alvarez et aZ., 2002. the reefs.
786 Porifera· Demospongiae • Halichondrida • Dictyonellidae

TETHYSPlRA TOPSENT, 1890 an apomorphy and are not homologous to echinating megascleres
common to the poecilosclerids.
Synonymy The genus Lissomyxilla Hanitsch, 1894a is an objective junior
synonym of Tethyspira as it shares the same type species (by
Tethyspira Topsent, 1890a: 195. Lissomyxilla Hanitsch, original designation).
1894a: 194.
Previous review
Type species
Topsent, 1900: 257.
Tethea spinosa Bowerbank, 1874b: 279 (by original
designation). Description of type species

Definition Tethyspira spinosa (Bowerbank, 1874b) (Fig. lOA-E).

Synonymy. Tethea spinosa Bowerbank, 1874b: 279,
Dictyonellidae with auxiliary spined styles in addition to long pI. LXXXIII figs 17-22; Tethyspira spinosa; Topsent, 1890a: 195;
choanosomal styles. Topsent, 1900: 257, pI. VII fig. 7; Lissomyxilla spinosa; Hanitsch,
1894a: 194. Rhaphidostyla incisa sensu Van Soest & Weinberg,
Diagnosis 1980: 3, fig. 4 (not Schmidt, 1880a).
Material examined. Holotype: BMNH unregistered
Massive cushion-shaped sponges with slippery conulose Bk.1081 - dry, Fowey Harbour. BMNH unregistered - BMNH
appearance. No ectosomal skeleton. Choanosomal skeleton con- slide labeled 'from same specimens as R.1628. Fowey Harbour.
sisting of bundles of long styles and single styles; at the base there Bk 394'. Other material. ZMA POR. 5975 - East Calf Island,
are short long-spined (tylo-)styles. A single species from the Roaringwater Bay, Ireland, August 7, 1982, depth 5 m.
Eastern Atlantic. Description. Low cushion (Fig. lOA-B) with tall sides and
rugose upper surface. Larger specimens may become massive-
Remarks globose, to ca 15 cm across, and are often angular with ridges and
crests. Surface slippery, conulose. Oscules translucent chimneys (Fig.
The record of Rhaphidostyla incisa by Van Soest & Weinberg lOB). Consistency firm, slightly compressible, elastic. Greyish-white
(1980) from Sherkin Island, Ireland, was found on re-examination or orangish. Choanosomal skeleton consists of parallel bundles of
to be Tethyspira spinosa (Van Soest, 1987). This genus so far long megascleres cemented to some extent by spongin (Fig. 1OC).
remains only known from a single, rather enigmatic species. Some spicules penetrate the surface. There is a scattered basal layer of
Topsent (1900) assigned it to his family Mesapidae in the order small acanthose styles. These are easily missed as the basal layer of
Clavulida (=Hadromerida), interpreting the spined spicules as massive sponges is not always collected or sectioned, and also the
microrhabds. Hooper (1991) assigned it to Raspailiidae as incertae acanthostyles may be scarce or absent. The main structural megascleres
sedis on the basis of resemblance of the spined styles to those of are straight or somewhat curved styles or subtylostyles (Fig. lOA, E):
Endectyon, for example. However, with regard to general appear- up to 1-2mmX3-12fJ.m. Long-spined microtylostyles (Fig. lOA,
ance and structure of the skeleton, T. spinosa fits better in D): ca. 70-120 X 4--6 fJ.m. Distribution and ecology. British Isles,
Dictyonellidae. The peculiar spined styles are here interPreted as Brittany; on wave exposed rocks, from 6m to at least ca. 60 m.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Halichondriidae Gray, 1867

Dirk Erpenbeck & Rob W.M. Van Soest

Zoological Museum, University of Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, The Netherlands. (erpenbeck@science.uva.nl;
soest@science.uva.nl)

Halichondriidae Gray (Demospongiae, Halichondrida) contains 11 valid genera (of 47 nominal genera) and probably around 180 species.
Halichondriidae are widely distributed and live mainly in shallow coastal waters but a few have also been recorded from 500 m depth. The
major morphological characters distinguishing this group from related sponges is the thoroughly confused arrangement of the choanoso-
mal megascleres (oxeas, styles and derivates thereof) usually coupled with a differentiated ectosomal skeleton, consisting of a tangential
or concentrated arrangement of spicules mostly smaller sized than those of the choanosome. All spicules are smooth. Except for rhaphides
(trichodragmas) there are no microscleres.
Keywords: Porifera; Demospongiae; Halichondrida; Halichondriidae; Amorphinopsis; Axinyssa; Ciocalapata; Ciocalypta; Epipolasis;
Halichondria (Halichondria); Halichondria (Eumastia); Hymeniacidon; Laminospongia; Spongosorites; Topsentia; Vosmaeria.

DEFINITION, DIAGNOSIS, SCOPE of the megascleres seemingly randomly strewn and often with rem-
nants of main spicule tracts and vague interconnecting spicules or
Synonymy tracts. Binding spongin is rare. Choanosomal organic parts show a
distinct lack of collagenous matter in many genera. Spicule density
Halichondriadae Gray, 1867a: 518. Halichondridae Vosmaer, may be quite high, and in combination often with large sizes of
1887: 335. Stylotellinae Lendenfeld, 1888: 185. Ciocalyptidae the megascleres the consistency may become hard and brittle.
Hentschel, 1923: 408. Spongosoritidae Topsent, 1928c: 35. 24. However, Halichondriidae with smaller spicules are often quite
Halichondriidae de Laubenfels, 1936a: 133. Hymeniacidonidae de compressible and easily damaged due to lack of skeletal organi-
Laubenfels, 1936a: 136. zation. Some genera are 'fleshy' due do relatively high collagen
content and low spicule density. Halichondriidae contain 47 nomi-
Definition nal genera, 11 of which are considered valid. In one genus
Halichondria, two valid subgenera are recognized.
Halichondrida with a confused arrangement of smooth oxeas
and/or styles in the choanosome and usually an organized special Scope
ectosomal skeleton consisting of tangentially arranged or densely
confusedly arranged crust of oxeas and/or styles of sizes similar to Genera considered valid members of the family:
or smaller than those of the choanosome. Amorphinopsis Carter, 1887, Axinyssa Lendenfeld, 1897c,
Ciocalapata de Laubenfels, 1936a, Ciocalypta Bowerbank, 1863,
Diagnosis Epipolasis de Laubenfels, 1936a, Halichondria Fleming, 1828, with
subgenera Halichondria and Eumastia, Hymeniacidon Bowerbank,
Halichondrida with oxeotes and stylotes, i.e., spicules taking 1862, Laminospongia Pulitzer-Finali, 1983, Spongosorites Topsent,
variously the shape of oxeas, strongyloxeas, oxeas with blunt ends, 1896a, Topsentia Berg, 1889, Vosmaeria Fristedt, 1885.
and true styles, occasionally with slightly expanded tyle near one
of the endings or more central. Spicules invariably smooth, often
elongate fusiform, but also equidiametrical over much of their TAXONOMIC HISTORY AND BIOLOGY
length, pointed apices normally elongated and gradually narrowing
to a sharp point, usually slightly curved, occasionally straight or History
rarely crooked or doubly angulated. Individual genera and species
may be recognizable on possessing either exclusively distinct This family was established by Gray (1867a) in his overview
oxeas or styles, but there is a strong tendency for species and gen- of sponge taxa known in his time. The type genus Halichondria was
era to have these extreme shapes intergraded with intermediate used in a very general sense by him and his contemporaries, so a
forms. Spicules are typically of a wide size range, with not infre- wide variety of sponges was included in the family along with the
quently distinct size categories without a clear overlap; ifthat is the type species Spongia panicea Pallas, 1766. Subsequently, the con-
case, then the smaller sizes are often concentrated at the surface. tents of the family were restricted by various authors, often in a
There is a distinct ectosomal skeletal differentiation with either a widely diverging manner. To date, no consensus has become appar-
tangential crust of intercrossing spicules, which is often easily ent mainly due to the poorly differentiated characters of the type
detached due to underlying subdermal spaces, or a confusedly species of Halichondria and its assumed allies. The assignment of
paratangential or palisade-like arrangement, in which case the sur- genera employed here is again slightly different from the latest revi-
face skeleton is not easily detached and comes off in 'flakes'. The sion of the family by Van Soest et al., 1990. Molecular data
choanosomal skeleton is characterized as confused, with many (Alvarez et al., 2000a; Chombard et a!., 1999) indicate we may

787
788 Porifera· Demospongiae • Halichondrida • Halichondriidae

expect in future further instability of the family contents. It is Phakellia), Batzella (now considered a valid genus of
beyond the scope of this chapter to enumerate all the genera at one Chondropsidae), Collocalypta (now considered a junior synonym of
time or another assigned by various authors under this family the Halichondriidae genus Ciocalypta), Densa (now considered a
name, but concepts of the family content used by a few of the junior synonym of the haplosclerid genus Xestospongia),
major authors are here briefly summarized. Gray, 1867a assigned Ectyobatzella (now considered a junior synonym of the
Halichondriidae (as Halichondriadae) to his subsection Poecilosclerid genus Monanchora), Hoplochalina (now considered
Spiculospongiae, order Leiospongia (sponge spicules of one kind a junior synonym of Scopalina), Oxeostilon, Prianos (now consid-
only). He defined the family as having a skeleton of 'fusiform' or ered a junior synonym of Haliclona), Rhaphidostyla, Viles,
'pin-shaped' spicules variously fasciculated together or rarely Dictyonella (now considered a valid genus of Dictyonellidae), and
united by a small quantity of spongin. Genera assigned were subdi- Hemimycale (now considered a valid genus of Hymedesmiidae).
vided into two groups, one with fusiform or 'needle-like' spicules, Levi (1973: 614) considered Halichondrida to be unrelated
perhaps considered as Halichondriidae sensu stricto: Reniera, to Axinellidae and assigned both to two different sub-
Halichondria, Dictyocylindrus, Aaptos, Halisarca, Lieberkuehnia, classes (Axinellidae to Tetractinomorpha, Halichondrida to
Tedania, Oro idea, Prianos, Schmidtia, Crella, Sophax, Epicles, Ceractinomorpha), on the basis of the possession of brooded larvae
Eurypon, Bubaris, Ciocalypta, Rasalia, Adocia, Philotia, and the in Halichondria and Hymeniacidon. He considered the
other with 'pin-shaped' spicules: Abila, Suberites, Ficulina, Halichondrida an artificial group, based on negative characters
Raspalia, Raphiophora, Spinularia, Antho, and Pitalia. The (absence of microscleres and organization of the skeleton), with two
family as a whole is a heterogeneous mixture of Haplosclerida, families, Halichondriidae (predominantly with oxeas) and
Poecilosclerida and Hadromerida, but included in the Hymeniacidonidae (predominantly with styles). He admits this divi-
Halichondriidae s.s. were Halichondria and Ciocalypta. Schmidt, sion was not absolute as Hymeniacidonidae might sometimes pos-
1870 did not employ the genus name Halichondria, nor a family sess oxeas and Halichondriidae might have styles. Of the genera
Halichondriidae, but he maintained a strict distinction between assigned to his Halichondriidae, Rhaphisia is now considered to
sponges with oxea- or strongyle-shape (family Renierinae) and belong to Haplosclerida. Most of the genera assigned by him to
those with 'Stecknadeln' (family Suberitidinae). Halichondria Hymeniacidonidae - with the exception of the type genus
panicea was considered a member of his genus Amorphina and thus Hymeniacidon - are here considered members of the family
Halichondria by inference was assigned to Renierinae. In this group Dictyonellidae based on the absence of an ectosomal skeleton.
he distinguished the following genera: Reniera, Amorphina, Pellina, Bergquist (1978) and Hartman (1982) essentially followed Levi's
Eumastia, Foliolina, Tedania, Schmidtia, Plicatella, and Auletta. scheme of the two families Halichondriidae and Hymeniacidonidae,
This is a mixture of modem Halichondrida and Haplosclerida, with but only few genera were assigned. Hartman mentions Rhaphisia as
Tedania as the odd Poecilosclerida. Topsent (1928c) assigned most an example of Halichondriidae but this is now considered
of the modem Halichondriidae (Halichondria, Ciocalypta, a Haliclona (see chapter on Chalinidae by de Weerdt), and Ulosa as
Coelocalypta, Topsentia, Hymeniacidon) to a widely employed an example of Hymeniacidonidae, but this is now considered a
family Axinellidae, of which he did not present a formal definition. member of Esperiopsidae (see chapter on Mycalidae by Van Soest &
One notable exception is Spongosorites, which is assigned to Hajdu). Van Soest et al. (1990) in a revision of the types of all
Hadromerida along with Aponastra and Alloscleria as a family of genera reestablished the close relationship of Halichondriidae and
its own, Spongosoritidae. In his comments he indicates nevertheless Axinellidae. This scheme was followed substantially by Hooper
a close, presumably ancestral, relationship with Topsentia and et al. (1997). Conversely, Hooper & Bergquist (1992) and Carballo
Halichondria. Of all the older authors, Topsent's classification is et al. (1996) proposed slight rearrangements of the contents of
probably closest to what is now perceived as a natural classification: Halichondriidae and Axinellidae described by Van Soest et al.
close kinship ofAxinellidae, Desmoxyidae, Dictyonellidae and (1990), but these latter proposals are not followed here because they
Halichondriidae. Topsent also believed this group to stem from merely represented a reinterpretation of characters of only a few
hadromerid stock, a notion which may still prove to be of value con- selected genera, and did not explore the consequences of the pro-
sidering some results of molecular studies. De Laubenfels (1936a: posed changes for the overall generic contents of the family. No new
126-139) employed an order Halichondrina with five families, viz., data or observations were offered to support the proposals. For the
Axinellidae, Halichondriidae, Semisuberitidae, Hymeniacidonidae present revision we revisited all the types and additional specimens
and Monanthidae. Halichondriidae were defined as having a spicu- and could add considerable further material based on visits to the
lation of smooth oxeas, a special dermal skeleton and lacking musea of Paris (LSF grant to DE) and London (LSF grant to RVS).
a conspicuously isodictyal reticulation. The family was assumed This revision supports most of the conclusions of Van Soest et al.
to contain the genera Halichondria, Apatospongia, Ciocalypta, (1999), but the following changes are proposed: (1) genera with
Ciocalapata, Dactylella (now considered a genus of Dictyonellidae), spined microxeas (Myrmekioderma and Didiscus) are reassigned to
Halichondriella, Leucophloeus, Trachyopsilla, and Trachyopsis. Desmoxyidae following e.g., Hooper & Levi (1993b), Van Soest &
He assigned Coelocalypta as junior synonym to Apatospongia, Lehnert (1997), (see chapter on Desmoxyidae by Hooper), whereas
and Halispongia and Spuma to Halichondria. The family Van Soest et al. (1990), followed by Hooper et al. (1997) retained it
Hymeniacidonidae de Laubenfels, 1932 was defined by having a within Halichondriidae. (2) Nailondria de Laubenfels, 1954 is con-
fleshy ectosome not profusely echinated by spicules which are usu- sidered ajunior synonym of Amorphinopsis (instead of ajunior syn-
ally tangentially placed, with the choanosomal structure plumose to onym of Hymeniacidon). (3) Ciocalapata de Laubenfells, 1936a is
confused and spiculation of oxeas, styles or strongyles, or combina- revived as a valid genus (previously considered ajunior synonym of
tions of these. The family was erected for Hymeniacidon (with Halichondria). (4) Collocalypta Dendy, 1905 is proposed to be a
assigned synonyms Amorphilla and Stylohalina), Acanthella junior synonym of Ciocalypta Bowerbank, 1861 (previously con-
(with assigned synonym Acanthellina; now considered a genus sidered a valid genus). (5) Coelocalypta Topsent, 1927b is proposed
of Dictyonellidae), Adreissa (now considered a junior synonym of to be a junior synonym of Ciocalypta Bowerbank, 1861 (previously
Porifera' Demospongiae • Halichondrida • Halichondriidae 789

considered a junior synonym of Topsentia). (6) Eumastia Schmidt, Taxonomic remarks

1870 is proposed to be a valid subgenus of Halichondria instead
of a mere junior synonym. (7) Pellina Schmidt, 1870 is proposed The choanosomal (lack of) arrangement of the megascleres in
to be a junior synonym of Halichondria (Halichondria). a confused manner constitutes the major synapomorphy for the
(8) Laminospongia Pulitzer-Finali, 1983 is revived as a valid genus family, although it is difficult to describe or illustrate this in precise
(previously considered a junior synonym of Topsentia). terms or images. There is a lack of visible binding spongin and
(9) Alloscleria Topsent, 1927b is removed from the family (previ- bundles or tracts of megascleres are not interconnected, and often
ously considered a junior synonym of Topsentia) and assigned lack a definite orientation. Vague tracts are found in many hali-
to Desmoxyidae (see Hooper's chapter on Desmoxyidae). chondrids, especially in the subectosomal region, where they tend
(10) Oxeostilon Ferrer Hernandez, 1922 is considered a to fan out and carry a special tangential skeleton or become part of
junior synonym of Spongosorites (rather than of Topsentia). it. The surface skeletal arrangement is a major discriminating char-
(11) Oxeosarcodea de Laubenfels, 1954, previously not considered acter at the genus level. Some members of the family lack a definite
as a member of Halichondriidae, is included in the synonymy of surface skeleton, and the surface is often somewhat conulose due to
Axinyssa. (12) Stylinos Topsent, 1891a, previously not considered projecting choanosomal tracts. Such sponges clearly bridge the gap
as a member of Halichondriidae, is now included in the synonymy with families Axinellidae and Dictyonellidae, which likewise lack
of Hymeniacidon. (13) Vosmaeria Fristedt, 1885, previously not a surface skeleton. Family membership of Axinyssa has been con-
considered as a member of Halichondriidae is now included as a tested (e.g., by Hooper & Bergquist, 1992, but later reincluded in
valid genus. (14) Astromimus Lendenfeld, 1897a, previously not the family by Hooper et aI., 1997), and it is proposed here to leave
considered as a member of Halichondriidae, is included in in the it in Halichondriidae based on the confused arrangement of the
synonymy ofAxinyssa. (15) Petromica Topsent, 1898b (with junior skeleton of most members of Axinyssa. Several Halichondriidae
synonym Monanthus Kirkpatrick, 1903a) is returned to 'Lithistida' species show a burrowing habit, typically with conical fistules
(family Desmanthidae) because of the complement of monocrepid sticking up from the sandy substrate from a massive basis buried
desmas. For this volume the 'Lithistida' are maintained as a poly- underneath the sand. This habit does not agree fully with details of
phyletic higher taxon, awaiting molecular and other characters for skeletal architecture and spicule shapes and sizes. Moreover, over-
assignment to existing orders and families of the Demospongiae. all similar habits are also found in definitely unrelated orders and
We predict that Petromica will eventually be found a valid member families (e.g., Poecilosclerida: Coelosphaeridae, Haplosclerida:
of Halichondrida, probably of Halichondriidae. Phloeodictyidae). In the classification of the genera of this family
Arguments for all these proposed changes are given at the such a rather distinctive habit thus will be assumed to have been
appropriate locations below. developed in several genera independently, notably in Ciocalypta
Finally, the genus Cryptax de Laubenfels, 1954: 214, was and Topsentia. Only in the case of Ciocalypta this habit is part of
erected for type species Cryptax orygmi de Laubenfels, 1954: 214, the discriminating characters for the genus in combination with a
fig. 146 (by monotypy) from Majuro Atoll, S Pacific. It was found particular arrangement of the skeleton and transparence of the
occupying cavities of up to 4 mm diameter within corals which in fistules. The exclusive occurrence of oxeas is so widespread in
tum were covered by a Spirastrella. Spiculation was reported to the family that it is not a discriminating character for genera;
consist exclusively of fusiform tylostyles with rounded endings however, the exclusive occurrence of styles of a particular shape
('tylostrongyles'), 600-700 X 2-12 /-Lm. De Laubenfels (1954) was (faint tyle near the blunt end) is one of the synapomorphies for the
not sure that it was actually boring and assigned it to Suberitidae, genus Hymeniacidon. Size categories of a single megascleres or
but it is also possible it might be a clionaid lacking spiraster of styles and oxeas combined in the same sponge may constitute
microscleres (see chapter on Clionaidae). However, re-examination additional generic synapomorphies. Secondary metabolites
of the holotype (USNM 22966), consisting of a spicule slide pre- reported from members of this family are abundant and often quite
pared by de Laubenfels, and some coral crumbs from which was characteristic. Van Soest & Braekman (1999) record isocyanoter-
peeled bits of the sponge coating them (prepared and examined penes (shared with Axinellidae and Dictyonellidae), sulfated
courtesy of Klaus Rlitzler), revealed only long oxeas with rounded sterols and macrolides (both shared with unrelated sponges).
tips, or with one tip rounded and the other one sharply pointed, or Various species of one genus, Spongosorites, produce characteris-
with a few thinner ones both pointed. There was no sign of a tylote tic topsentins.
modification reported by de Laubenfels (he may have mixed in the
Spirastrella, but there were no tylostyles in his slide preparation Biology
either). The oxeas are 650-850 X 5-12 /-Lm, agreeing with sizes of
'tylostrongyles' reported by de Laubenfels, with some bent at the Two genera (Halichondria and Hymeniacidon) are viviparous,
center, some off-center, some slightly, some sharply. The peel and their larvae are wholly ciliated. Those of Halichondria have a
shows criss-cross orientation parallel to the substrate, indicating characteristic tuft of long cilia at the posterior pole (Wapstra & Van
possible affinities with the halichondriids, but owing to the discrep- Soest, 1987). Larvae of other genera so far remain unknown, these
ancies between the original description and the holotype, the genus may be likely oviparous. Halichondriidae occur in all areas of the
Cryptax and its type species are declared here incertae sedis and world oceans, from the intertidal down to continental slope habi-
unassignable to any known halichondriid. tats. Many species are specialists of sedimented habitats.

KEY TO GENERA

(1) Trichodragmas (hair-like spicules packed in wispy bundles) present .................................................................................. Epipolasis
No trichodragmas ................................................................................................................................................................................ 2
790 Porifera· Demospongiae • HaIichondrida • HaIichondriidae

(2) Shape a massive base from which rise long conical semitransparent fistules strengthened by an internal spicule axis and extra-axial
spicule bundles at right angles to the surface .. ........... ................ ................. ................ ........... .... ... ..................................... Ciocalypta
Various shapes, may include short fistules or papilllae, but no large semitransparent conical fistules ............ .......... ...... ................. 3
(3) Shape a thin blade, with scattered small oscules ....................................................................................................... Laminospongia
No thin blade ..................................................................................................................................................................................... 4
(4) Finely conulose surface, no special surface skeleton, choanosome collagenous, spicule density relatively low ................ Axinyssa
Surface smooth, surface strengthened by special tangential or paratangential skeleton, spongin not visibly present ....... ,............... 5
(5) Spicules exclusively short styles (many with a subterminal swelling) ....................................................................... Hymeniacidon
Spicules oxeas, or a mixture of oxeas and (sub/tylo-)styles ............................................................................................... ,.............. 6
(6) Surface skeleton easily detachable without taking away part of the choanosome ............................................................. ,.............. 7
Surface skeleton only detachable as flakes with part of the choanosome attached ......................................................................... 10
(7) Spicules exclusively oxeas in a wide size range ........................................................................................................... Halichondria
Spicules oxeas and (sub/tylo-)styles .................................................................................................................................................. 8
(8) Stylote spicules have a distinct tyle and are in fact (sub-)tylostyles ...................................................... ............................ Vosmaeria
True styles, without tyle .................................................................................................................................................................... 9
(9) Styles only in a small size category (200-400 /-Lm), next to larger oxeas .................................................................. Amorphinopsis
Styles only in a large size category (>800 /-Lm), next to smaller and larger oxeas .......................................................... Ciocalapata
(10) Consistency stony hard, crumbly, surface rough to the touch, choanosome a dense mass of single spicules ..................... Topsentia
Consistency dense but compressible, surface smooth, choanosome with tracts of spicules; spicules often angular or doubly bent;
most species show aerophobic reaction when collected ....... ................ ................. .......... ............................ .... ...... ....... Spongosorites

AMORPHINOPSIS CARTER, 1887 Synonymy. Amorphinopsis excavans Carter, 1887: 77, pI. V
figs 12-15; Annandale, 1915b: 467, figs 4-5; Hooper et aZ., 1997:
Synonymy 25, figs 15-16.
Material examined. Holotype (slide): BMNH 1981.10.14.3-
Amorphinopsis Carter, 1887: 77. Prostylissa Topsent, 1925b: a slide of Carter's type specimen labelled "Mergui Archipelago, fro
208. Tumata de Laubenfels, 1936a: 77. Nailondria de Laubenfels, Anderson, 28 Dec. 82, don. by Linn. Soc. via Wyn. Wheeler, 1981,
1954: 182. '3', '74', sponge surface excavating sp". The holotype (not exam-
ined), is apparently located in the Indian Museum, Calcutta IM
Type species 659717 ZEV (cf. Annandale, 1915b).
Description. This is based on Annandale's (1915b)
Amorphinopsis excavans Carter, 1887: 77 (by monotypy). redescription of a presumed fragment of the type in the Indian
Museum Calcutta. The BMNH slide which is - based on the text on
Definition the label - quite certainly made from the type is a dissociated
spicule mount. Shape of the type specimen is encrusting on a piece
Halichondriidae with ectosomal tangential skeleton of inter- of dead coral, size 6 X 3 cm. The surface is smooth but displays
crossing megascleres single or in tracts. Spicules differentiated into some conules in the dry state (Fig. IC). No visible openings. The
larger oxeas and derived forms, smaller oxeas and small styles, dry sponge is firm. Annandale's subsequent specimens were digi-
concentrated at the surface. Choanosomal skeleton confused with tate (Fig. ID). Ectosomal skeleton (Fig. lA) tangential, consisting
high spicular density and little or no spongin. of thick intercrossing tracts of larger oxeas, the spaces in between
filled with loose oxeas of all sizes as well as small styles.
Diagnosis Choanosomal skeleton confused, with thick but vaguely delimited
tracts and many loose individual spicules. Megascleres predomi-
Encrusting to massive, occasionally with irregular branches nantly oxeas in a large size variation and small styles concentrated
issuing from a massive base. Firm, hard, but fragile sponges. at the surface (Fig. IB). Long oxeas, fat, fusiform, curved,
Ectosomal skeleton tangential, with intercrossing larger oxeas and 520-700 X 18-20 /-Lm. Shorter oxeas, fusiform, 200-375 X 8-
derived forms, often forming intercrossing bundles. Small oxeas 12/-Lm. Styles (Fig. IB), thickest in the middle, blunt end tapering
and styles concentrated at the surface. Choanosomal skeleton con- 120 X 6 /-Lm. Distribution and ecology. Mergui Archipelago, off the
fused, with irregular tracts of large oxeas. Approximately ten coast of Birma, coral reefs.
species, mostly from the Indian Ocean and West Pacific, but the Remarks. Carter's description is misleading in its emphasis
genus is also recorded from Brazil (Muricy et al., 1991). on a star-shaped surface appearance (very probably caused by des-
iccation) and 'excavating' habit. Annandale's notes make it clear
Previous reviews that the surface is merely slightly conulose due to the ectosome
being lifted by thicker choanosomal tracts. The excavating habit is
Annandale (1915b), Van Soest et al. (1990), Hooper et aZ. simulated by penetration of portions of the sponge into bore holes
(1997). made by and subsequently left vacant by clionids. Amorphinopsis
excavans is not really an excavating sponge and subsequent records
Description of type species of it as such (e.g., Thomas, 1973) must be considered erroneous.
New material of the type species was described from NW Australia
Amorphinopsis excavans Carter, 1887 (Fig. lA-D). by Hooper et aZ. (1997).
Porifera' Demospongiae • Halichondrida • Halichondriidae 791

Fig. 1. A-D, Amorphinopsis excavans Carter, 1887. A, tangential view of the surface skeleton. B, large megasderes and small styles (scale 100 !Lm).
C, habit of the holotype reproduced from Carter, 1887 (scale 1 mm). D, longitudinal section through a digitiform process reproduced from Annandale,
1915. E-G, Amorphinopsis maza (de Laubenfels, 1954 as Nailondria) spicules (scales: E-F, 100 !Lm; G, 10 !Lm). H-L, Amorphinopsis siamensis (Top sent,
1925b as Prostylissa). H-J, spicules (scales: H-I, 100 !Lm; J, 10 !Lm). K, surface view (scale 100 !Lm). L, view on peripheral skeleton (scale 1000 !Lm).

The genus Prostylissa Topsent (1925b: 208) was erected it is a related but separate species, to be named Amorphinopsis
for Prostylissa siamensis Topsent, 1925b: 208, figs 1-2 (by mono- siamensis.
typy). The holotype of Prostylissa siamensis, MNHN DT. 3453, The genus Tumata de Laubenfels, 1936a: 77 was erected for
labeled "Golfe de Siam, A. Krempffleg. 1921", was reexamined. It type species Reniera megarrhaphea Lendenfeld, 1888: 79 from
consists of a mass of irregular thick anastoming branches, up to Port Jackson, East Australia (by original designation). The type
2 em in diameter. The surface is smooth but slightly conulose, and series (not examined) is rather confusedly treated by subsequent
it displays a few large openings, presumably oscules. Consistency authors: Whitelegge (1902: 277, 280) redescribed the 'type' given
finn but friable. The ectosomal skeleton (Fig. lK) is detachable as No. 385, presumably from the Australian Museum and from
due to some limited subectosomal vestibules (Fig. lL). It consists Port Jackson but this is not expressly mentioned and can be only
of tangentially intercrossing tracts of large oxeas surrounded by circumstantially deduced. Hallmann (1914: 330, pI. XVII figs 5-6,
small spicules, which are strewn irregularly or appear to 'echinate' text-fig. 3) gave a full description with some figures of the type, but
the tracts. Choanosomal skeleton consisting of thick tracts of large failed to mention a specimen number; presumably it was the same
oxeas and many single spicules. Smaller spicules concentrated at specimen referred to as type by Whitelegge, 1902. Hooper &
the periphery. Large fusifonn oxeas (Fig. IH), 980-1050 X 40 /-lm. Wiedenmayer (1994: 205) list a series of eight specimens as syn-
Smaller similar oxeas (Fig. 1I), 300 X 7-10 /-lm. Small fusifonn types, two from AMS and six from BMNH, with slides in AMS
styles (Fig. lJ), 180-300 X 7-10 /-lm. Tbese features make it clear of two of the BMNH specimens, and a slide in 2MB from one of
that Prostylissa is a synonym of Amorphinopsis. Prostylissa the BMNH specimens. However, this series contains specimens
siamensis was considered an elaborate specimen of the same from Mauritius and New Zealand, which cannot be part of the type
species A. excavans by Van Soest et at. (1990) for example, but series since these localities were not mentioned in Lendenfeld's
in view of the clearly larger and thicker size of the large oxea original description. The specimens that are from the type locality,
category in combination with the ramose habit, it is likely that presumably BMNH 1887.4.27.84 (with slide in AMS G3338),
792 Porifera· Demospongiae • Halichondrida • Halichondriidae

BMNH 1887.4.27.14 (slide AMS G3377), BMNH 1887.4.27.6, spicule bundles. Choanosomal skeleton disorganized with spicules
AMS G9016 and AMS Z2015, may contain the original type speci- strewn in confusion and/or composed of vaguely ascending, widely
men described by Lendenfeld, but this remains undetermined. Both spaced vertical tracts of large oxeas or strongyloxeas, forming
Whitelegge describing the 'type No. 385' and Hallmann noted that loose bundles. Choanosome with poor or moderate spongin devel-
their specimen was quite different in shape from what Lendenfeld opment, but heavy interspicular collagen; spicule density relatively
described. In the absence of a connection between Whitelegge's low. Spicules oxeas, strongyloxeas or stylote modifications, usu-
No. 385 and the type series mentioned by Hooper & Wiedenmayer, ally of only one size class. About 15 species, distributed over the
it is decided here to indicate AMS G9016 as the lectotype. Reasons warmer waters of the world oceans.
are twofold: the first author to redescribe the species referred to a
specimen from AMS rather than from BMNH, and the only other Previous reviews
AMS specimen is a mere fragment. The descriptions of Whitelegge
and Hallmann leave little doubt that Reniera megarrhaphea is an Van Soest et al. (1990), Hooper & Bergquist (1992), Hooper
Amorphinopsis, with an ectosomal skeleton of intercrossing oxeas et al. (1997).
and small styles and a confused choanosomal skeleton of oxeas.
Sizes of oxeas 220-950 X 6-31 fLm, of styles 160-250 X 6-9 fLm Description of type species
(the original description of Lendenfeld did not mention styles, but
only small oxeas of 20 fLm (presumably a misprint for 200) X 8 fLm. Axinyssa topsenti Lendenfeld, 1897c (Fig. 2A-F).
The genus Tumata is a junior synonym of Amorphinopsis. Synonymy. Axinyssa topsenti Lendenfeld, l897c: 116,
The genus Nailondria de Laubenfels (1954: 182) was erected pI. 10 figs 134-144.
(by monotypy) for type species Nailondria maza de Laubenfels Material examined. Holotype: 2MB 2971 - from Zanzibar.
(1954: 182, fig. 121). The type specimen, USNM 23083, from the Schizotype: BMNH 1908.9.4.145, including slide 1897.3.25.70.
West Central Pacific, was reexamined. It is described as an amor- Description. Lobate mass of tubes, 10 cm in largest expan-
phous soft sponge with the consistency of soggy bread, diameter sion. Individual lobes (Fig. 2A) 2 cm high, 3 cm diameter at the
15 cm, yellow. The surface is smooth, detachable, with conspicu- base, tapering to 0.5 cm. Lobes tube-like with apical oscules pene-
ous oscules of 5 mm diameter, 3 cm apart. The ectosomal skeleton trating deep into the interior of the sponge. Surface finely but regu-
is a dense tangential crust of intercrossing single spicules. The larly conulose. Consistency firm. Colour dark green. Ectosomal
choanosome is cavernous, with indistinct spicule tracts 40-100 fLm region (Fig. 2D-E) traversed by protruding choanosomal bundles
in diameter directed to and carrying the surface crust. Spicules: interconnected more or less tangentially by irregular bundles of
strongyloxeas (Fig. IE-F), 460-690 X 10-18 fLm, and styles 1-2 spicules enclosing large pore-fields. Protruding bundles at the
(Fig. IG), 220-288 X 5-8 fLm. Van Soest et al. (1990) assigned this surface consist of small categories of megascleres, whereas the tan-
species to Hymeniacidon, because both sizes of spicules could be gential spicules are of a larger category. The subectosomal region
interpreted as styles. However, the strongyloxeas are definitely contains large open spaces (Fig. 2F) of 300-500 fLm diameter sep-
oxeote in origin. The soft consistency is unusual, but the other arated by bundles of spicules 50-200 fLm in diameter. In between
characters match well with Amorphinopsis, thus we propose this are masses of loose direction-less spicules, which may become
species to be named Amorphinopsis maza. longitudinally arranged (Fig. 2B) along the inner wall of the
oscular cavity. Spicules (Fig. 2C) are large oxeas, 550-740 X
14-18 fLm, some with stylote endings and small oxeas located
AX/NYSSA LENDENFELD, 1897 at the conules, 165-250 X 4-5 fLm, many with one end rounded
but clearly of oxeote origin. Distribution and ecology. Zanzibar,
Synonymy shallow-water.
Remarks. The type species has never been reliably
Axinyssa Lendenfeld, 1897c: 116. Astromimus Lendenfeld, described since its original description (Stephens', 1915b record is
1897a: 148. Pseudaxinyssa Burton, 1931a: 350. Axinomimus de dubious, as the description differs substantially from the type), and
Laubenfels, 1936a: 163. Oxeosarcodea de Laubenfels, 1954: 230. it appears to be unique in its possession of the tube-shaped habit
and distinct size categories of the megascleres. Subsequent use of
Type species Axinyssa was rare until Burton (1931a: 350) described the genus
Pseudaxinyssa as distinct from Axinyssa in possessing a single size
Axinyssa topsenti Lendenfeld, 1897c: 116 (by monotypy). category of megascleres. Pseudaxinyssa was erected for type
species (original designation) Axinyssa tethyoides Kirkpatrick
Definition (1903a: 245, fig. 18). The type of this species, BMNH
1902.11.16.25, was redescribed by Van Soest et al. (1990) and
Halichondriidae lacking an ectosomal tangential skeleton. Hooper & Bergquist (1992). It is a globular black sponge (Fig. 3A)
Choanosomal skeleton largely disorganized, but at the periphery with a papillate surface, which is detachable but organic, and it has
the spicules are arranged in bundles at right angles to and protrud- a black colour. The skeleton is an irregular dendritic system of bun-
ing slightly beyond the surface causing a fine conulation. dles and fibres of oxeas (Fig. 3B), 500-1000 fLm in diameter, an
ectosomal skeleton is lacking entirely. There are many loose
Diagnosis spicules. The oxeas are fusiform and thick, oxeas, stylote modifica-
tions, 420-700 X 8-34 f.Lm. There is little resemblance to the type
Massive, lobate or tubular sponges with conulose surface. specimen ofAxinyssa topsenti, but like that it appears to be unique
Ectosomal region lacking a distinct surface skeleton, largely in its unusual shape, surface and skeleton. Burton (1931a)
organic, tough, with sparsely scattered spicules and protruding described a further species of Pseudaxinyssa. P. tenuispiculata
Porifera· Demospongiae • Halichondrida • Halichondriidae 793

Fig. 2. Axinyssa topsenti Lendenfeld 1897. A, BMNH schizotype. B, longitudinal section made from schizotype (size see text). C, spicules. D, surface
view (both reproduced from Lendenfeld, 1897: pI. 10 figs 134-144, size see text). E, cross section (scale 100 fLm). F, surface view (scale 100 fLm).

Burton (1931a). This was reexamined, but the similarity with of a detachable ectosomal skeleton in combination with the surface-
A. tethyoides and A. topsenti is low. oriented vague choanosomal tracts in this species conform to the
The genus Astromimus Lendenfeld, 1897a: 148 was erected characters of Axinyssa and accordingly it is assigned to the syn-
for type species Astromimus luteus Lendenfeld, 1897a: 148, pI. V onymy of that genus. Both names were erected in 1897 (although
fig. 44, pI. VII fig. 80, pI. XII figs 217-221, 224, 225 (monotypy) the title page of 'Die Clavulina' gives 1896, the last page
from Lesina, in the Adriatic. The holotype (wet, reexamined) is announced that it was in fact printed in 1897). In view of the pre-
kept in the Natural History Museum of Vienna and now consists of vailing usage ofAxinyssa it is proposed to continue its use and con-
half the specimen depicted in Lendenfeld's pi. V fig. 44. This is sider Astromimus a junior synonym (if necessary under ICZN
a semiglobular mass of 6cm diameter and 7 cm height, the live article 23.9, because Astromimus subsequent to its original descrip-
colour quoted as an intense yellow. The surface is smooth to tion was only mentioned in an uncritical species list of Pulitzer-
slightly irregular-hispid, somewhat shaggy in appearance, punc- Finali, 1983). Axinyssa luteus is very likely a specimen of what is
tate, even seemingly reticulate to the naked eye. Oscules with a generally known as Halichondria aurantiaca (Schmidt, 1864: 38),
raised rim, 4-6 mm diameter. The skeleton is loosely radiate, with and accordingly the valid name for it is Axinyssa aurantiaca
vague bundles of oxeas and single spicules oriented generally (Schmidt, 1864).
towards the surface, where they spread out to form a rather con- The genus Axinomimus de Laubenfels (1936a: 163) was
densed skeleton compared to that of the interior. In cross section of erected for type species Axinella paradoxa Ridley & Dendy (1886:
the peripheral skeleton a few paratangentially arranged bundles 482). The type specimen, BMNH 1887.5.2.68, was reexamined.
and loose spicules are apparent, which cause the reticulated aspect It is a massive lobate sponge (Fig. 3C) of 2.5 cm high and 4 cm
of the surface, but there is no easily detachable ectosomal skeleton. diameter. Oscules arranged in small groups on top of the lobes.
Spicules oxeas, curved, often blunt-ending, in a wide size range, It has a conulose surface and rubbery-fibrose consistency. The
but without clear size categories, 300-1000 X 8-13 f..lm. The lack ectosomal region has bundles of spicules protruding beyond the
794 Porifera· Demospongiae • Halichondrida • Halichondriidae

Fig.3. A-B, Axinyssa tethyoides Kirkpatrick (1903a) (= type of Pseudaxinyssa Burton 1931 a). A, holotype specimen (scale I cm). B, SEM image of sur-
face showing bundles of oxeas (scale 100 j.Lm). C-E, Axinyssa paradoxa (Ridley & Dendy, 1886 as Axinella) (=type of Axin om in us de Laubenfels 1936a).
C, holotype (scale I cm). D, cross-section, reproduced from Van Soest et al. (1990) (scale 500 j.Lm). E, spicules (scale 100 j.Lm).

organic skin to form low conules, but it lacks tangential spicules. of partly tangential spicules of the same size as those in the
Between the conules, there are prominent porefields, The interior. Oxeosarcodea is considered a junior synonym of
choanosomal skeleton (Fig, 3D) consists of large oxeote spicules Axinyssa.
forming stout fibres running vertically upwards; numerous irregu- The surface characters ofAxinyssa are atypical in the family
larly scattered spicules between the fibres, Spicules (Fig, 3E) Halichondriidae, but through the presence of a large complement
smooth large oxeas, 480-640 X 14-18 fJ-m. of loose irregularly scattered oxeas in the choanosome the genus
It is proposed to distinguish the properties of a genus Axinyssa Axinyssa is more naturally placed in this family than any other
by combining the overlapping characters of the three above family of the order Halichondrida.
described species (Axinyssa topsenti, A. tethyoides and A. para-
doxa). They share the absence of a clear ectosomal skeleton
(although the type species still has a tangential complement), a CIOCALAPATA DE LAUBENFELS, 1936
conulose surface due to protruding choanosomal spicule tracts, and
a fibrous choanosome with low spicular density. Synonymy
The genus Oxeosarcodea de Laubenfels, 1954: 230, which was
erected for type (by monotypy) Oxeosarcodea oinops de Laubenfels, Ciocalapata de Laubenfels, 1936a: 134.
1954: 230, fig. 158, from Ebon Atoll, Central Pacific, shares
these characters. Slides of the holotype, USNM 22982, were Type species
reexamined. This is a red, massive, microconulose sponge, with
a confused skeleton of oxeas and stylote/strongylote modifica- Ciocalypta amorphosa Ridley & Dendy, 1886: 479 (by original
tions, 420-660 X 4-12 fJ-m. The surface has an incomplete cover designation).
Porifera· Demospongiae • Halichondrida • Halichondriidae 795

Fig. 4. Ciocalapata amorphosa Ridley & Dendy, 1886. A, paralectotype specimen (scale 1 cm). B, spicules (from Ridley & Dendy, 1886, size see text).
e, drawing of spicules and cross section of paralectotype (from Van Soest et al., 1990: fig. 73) (scales: spicules, 100 .... m; section, 1 mm). D, SEM image of
surface of paralectotype (scale 100 .... m).

Definition Synonymy. Ciocalypta amorphosa Ridley & Dendy, 1886:

479; Ridley & Dendy, 1887: 175, pI. XL fig. 9; Ciocalapata
Halichondriidae with detachable tangential ectosomal skele- amorphosa; de Laubenfels, 1936a: 134; Halichondria amorphosa;
ton of intercrossing bundles of spicules and a trabecular choanoso- Van Soest et al., 1990: 47, fig. 73.
mal skeleton of thick spicule tracts. Spicules oxeas (in two size Material examined. Lectotype: BMNH 1887.5.2.56 -
categories in the type species) and a separate category of styles. largest of 5-6 fragments, 'Challenger' expedition, Stn. 320.
Paralectotypes: 4-5 remaining fragments registered under the same
Diagnosis number.
Description. Massive, sprawling, riddled with rounded
Massive, fragile, soggy sponges, with a thin detachable ectoso- holes (small paralectotype fragment shown in Fig. 4A).
mal skeleton roofing over a system of large choanosomal holes in Consistency soggy. Surface skeleton (Fig. 4D) where still present,
the type species. Choanosomal thick spicules tracts fonning a tra- easily detached; elsewhere the surface is rough and tufted.
becular system enclosing the holes. Spicules include both oxeas and Choanosomal cavities 1-2mm in diameter. Ectosomal skeleton
genuine styles. A single species from deep water in the SW Atlantic. consisting of intercrossing bundles of megascleres (Fig. 4C),
smaller oxeas dominant but larger oxeas and styles also present.
Previous review Choanosomal spicule bundles of 200-300 J.Lm in diameter fonn a
trabecular system surrounding the holes, but in addition there are
Van Soest et al. (1990, as a junior synonym of Halichondria). many megascleres in confusion. Spicules (Fig. 4B) oxeas in two
distinct size categories, 900-1500 X 18-25 J.Lm and 515-680 X
Description of type species 8-10 J.Lm, and styles in a single category, 940-1150 X 30 J.Lm.
Strongylote modifications of the styles occur in a low frequency.
Ciocalapata amorphosa (Ridley & Dendy, 1886) (Fig. 4A-D). Distribution and ecology. SW Atlantic, 600 m.
796 Porifera· Demospongiae • Halichondrida • HaIichondriidae

Remarks. The possession of a special category of large true to cream-yellow to grey. The surface of the fistule has a glassy,
styles is not shared with the type species of Halichondria and other translucent appearance, through which vertical spicule fibres can
related species. Amorphinopsis has only small styles and is also be seen with the unaided eye. Oscules are on top of some of the fis-
otherwise distinct. For that reason we prefer to retain Ciocalapata tules. Consistency firm. The ectosomal skeleton (Fig. 5C) is a tan-
as a valid genus. Van Soest et al.'s (1990) suggestion that it may gential reticulation of intercrossing bundles, thickness 45-110 /Lm.
be a member of the genus Biemna without microscleres is Spaces in between bundles 100-250/Lm with smaller spicules
refuted here, as skeletal architecture is clearly halichondrioid. predominant. Subectosomal spaces present. The main skeleton is
A second species has been ascribed recently to Ciocalapata, viz., an irregular, confused reticulation consisting mostly of larger
Ciocalapata almae Carballo et al., 1996. The shape, spicule size, spicules, with a tendency to form ascending fibres. In the fistules
skeletal architecture, and habitat of this species deviate strongly there is a central condensation consisting of a column of 3-4
from C. amorphosa, so this assignment would stretch the genus aligned thick tracts (Fig. 5D), together about 800/Lm diameter,
characters considerably. The styles described from C. almae appear with radially arranged ascending fibres ending at right angles at the
to be modified oxeas rather than true monactinal spicules, and surface and lifting it up in preserved or dried specimens. Distance
membership of Halichondria appears more likely. of these supporting tracts 1500-1600 /Lm, space between has a low
spicular density and carries mostly only single spicules in confu-
CIOCALYPTA BOWERBANK, 1862 sion. The spiculation consists of large and small, slender styles and
occasional oxeas. These are incompletely differentiated into two
Synonymy size categories. Long styles, 600-630 X 12-18 /Lm. Small styles,
340-390 X 5-10 /Lm. Small oxeas, 200-260 X 5/Lm. The spicules
Ciocalypta Bowerbank, 1862: 1105. Leucophloeus Carter, frequently have telescoped or distorted extremities, and it is possi-
1883b: 323. Collocalypta Dendy, 1905: 199. Coelocalypta ble that the oxeas are none other than modified styles. Distribution
Topsent, 1928c: 167. Pseudohymeniacidon Carballo, 2001: 258. and ecology. European seaboard of Atlantic from Helgoland south
to Spain, Portugal Mediterranean, in clear water.
Type species Remarks. By retaining the fistule growth form as an integral
feature of the genus definition, we create a possible confusion in the
Ciocalypta penicillus Bowerbank, 1864: 179 (by monotypy). family Halichondriidae, because several halichondrids with
fistular growth form appear closely related to non-fistular forms.
Definition Collocalypta digitata Dendy (1905) (cf. below) is convincingly
similar in shape to Ciocalypta penicillUS, but shares an important
Halichondrids with finger-shaped fistu1es possessing a feature with Axinyssa, i.e., the complete absence of an ectosomal
central spicular axis, with strong secondary tracts supporting skeleton. These facts cast doubt over the validity of the fistular
the ectosome. Spicules styles and/or oxeas, usully in two size growth form as a character uniting a monophyletic group of
categories. sponges. Emphasis should here be placed on features of the skeleton
in addition to the fistular habit: the possession of a central column
Diagnosis and extra-axial thick supporting tracts, may serve to differentiate the
species of Ciocalypta from other fistular halichondrids. Van Soest
Basal mass usually buried in the soft sediment with conically et al. (1990) emphasized possession of styles, thus excluding
tapering erect finger-shaped fistules rising above the sediment. species like Coelocalypta porrecta Topsent, 1928c, Collocalypta
Surface smooth, usually somewhat transparent, without visible digitata, Ciocalypta tyleri Bowerbank, 1875b and C. gibbsi Wells
oscules. Ectosomal skeleton usually present, but occasionally lack- et al., 1960 from Ciocalypta. In contrast, we submit here that the
ing. If present it is tangential, carried by strong spicule tracts issu- possession of exclusively oxeas in an otherwise closely similar
ing from a thick central spicular axis. Skeleton of basal mass species such as found in these mentioned above is insufficient rea-
largely confused. About 15 species occuring mostly in temperate son to exclude them from Ciocalypta, as a minor complement of
and subtropical waters of the Atlantic and the South Pacific. oxeas is found even in its type species. Topsent (1921) also took the
view that oxeas and styles may assume a variable dominance in
Description of type species species of Ciocalypta, and it is advocated here to follow this.
The genus Leucophloeus Carter (1883b) was erected (by
Ciocalypta penicillus Bowerbank, 1864 (Fig. 5A-D). monotypy) for Leucophloeus massalis Carter (1883b: 323, pI. XIV
Synonymy. Ciocalypta penicillus 1864: 179, pI. XXX fig. 15A-B). The type material cannot be unambiguously identified
fig. 360; Bowerbank, 1874b: 33, pI. XIII figs 2-4. Ciocalypta leei from well-labelled specimens, but Van Soest et al. (1990) report the
Bowerbank, 1874b: 295, pI. LXXXVI figs 1-4. existence of a specimen labelled Leucophloeus massalis in the
Material examined. Lectotype: BMNH 1877.5.21.1069 (with BMNH 'Bowerbank collection' bearing the numbers 706 and 31,
additional number 1930.7.3.28) - from Hastings, SE England. further data and registration lacking, possessing several of the char-
Paralectotype: BMNH 1830.4.3.29. acters described by Carter (1883b). It is a piece of a conical fistule
Description. Basal cushion of up to 10 cm in diameter which has a smooth furrowed surface, a thick white surface crust of
buried in sand or gravel, from which project large conical, translu- spicules over a beige coloured interior of spicule tracts forming
cent, ridged, thick walled, non-contractile fistules (Fig. 5B), nor- a central axis and radiating spicule tracts. Spicules (Fig. 7C)
mally up to about 5-9 cm high, 0.5-0.6 mm in diameter. In the are styles in two sizes. It is proposed to adopt this material as
lectotype (Fig. 5A) there are 18 individual fistules, each about the lectotype of Leucophloeus massalis. Its properties allow the
2 cm diameter at the base, 0.5 cm at the apex. Colour white-cream synonymization of Leucophloeus with Ciocalypta.
Porifera' Demospongiae • Halichondrida • Halichondriidae 797

Fig. 5. Ciocaiypta penicillus Bowerbank, 1864. A, lectotype specimen and spicules (from Bowerbank, 1874b: pI. LXXXVI figs 1-3) (sizes see text).
B, specimens photographed in situ (photo B. Picton). C, tangential view of surface (from Van Soest et ai., 1990: fig. 60) (scale 500 /Lm). D, SEM image of
cross-section (scale 1000/Lm).

The genus Collocalypta Dendy (1905: 199) was erected precisely these features (Ciocalypta-like habit without ectosomal
(by monotypy) for Collocalypta digitata Dendy (1905: 199, pI. VII skeleton), it is assumed to be autapomorphic. If further species
fig. 6, pI. XllI figs 1-2). The holotype BMNH 1907.2.6.89 was reex- of Ciocalypta without ectosomal skeleton are found to exist
amined. It resembles Ciocalypta in habitus (Fig. 6A), including the Collocalypta may be raised to subgenus level. Carballo (2000)
transparent fistules which are furrowed and conulose in preserved revived Collocalypta as a valid genus in Axinellidae, and associ-
condition. The basal mass is flattened, covered with shells and ated it with Hymerhabdia.
sand, and the fistules are relatively widely apart, up to 5 cm high. The genus Coelocalypta Topsent (1928c) with type species
No visible oscules. Consistency tough. The skeleton of the basal (by monotypy) Coelocalypta porrecta Topsent (1928c: 167, pI. II
crust consists of stout, erect, plumose columns of spicule tracts fig. 6, pI. VI fig. 4) is now considered a member of Ciocalypta,
cemented by spongin. The fistules have a central column of aligned not of Topsentia as Van Soest et al. (1990) proposed. Most of the
spicules, from which diverge thick loose bundles of spicules features (Fig. 7A-B) of this subtropical East Atlantic species con-
towards the surface (Fig. 6B, 6D), lifting this up into conules. form to Ciocalypta, induding a central column and supporting
Oxeas (Fig. 6C), in various shapes, shorter and thicker, slightly spicule tracts in the fistules and an ectosomal skeleton. The tan-
curved, irregularly ended 450-630 X 20-30 /Lm, and long and gential tracts in the ectosome are absent, instead of which there
slender, slightly curved, 760-930 X 20-30 /Lm. A noteworthy are single tangential spicules overlying a dense crust of paratan-
difference with Ciocalypta penicillus is the absence of an ectoso- gential brushes of oxeas. Nevertheless, the ectosomal skeleton
mal tangential skeleton (Fig. 6E), instead of which there is is detachable like in Ciocalypta penicillus because of the presence
a thick organic skin. This absence was considered to be of generic of a regular system of subectosomal vestibules, between the
significance by previous authors (e.g., Van Soest et at., 1990), supporting tracts. Carballo (2001) revived Coelocalypta as a valid
but in view of the fact there is only a single species with genus.
798 Porifera· Demospongiae • Halichondrida • Halichondriidae

A B c

Fig. 6. Ciocalypta digitata Dendy (1905, as Collocalypta). A, type specimen. B, cross-section. C, oxea (all reproduced from Dendy, 1905: pI. VI fig. 6 &
pI. XIII figs 1-2 (sizes see text)). D, SEM image of cross-section made from the type (scale 500 /-Lm). E, surface view of same (scale 100 /-Lm).

The genus Pseudohymeniacidon Carballo, 2001: 258 was EPIPOLASIS DE LAUBENFELS, 1936
erected for type species Coelocalypta aderma Levi & Vacelet,
1958: 236, fig. 18 from the Azores (original designation). The Synonymy
definition given by Carballo is rather enigmatic, but it is evident
from the characters of the type species, that it is intended for Epipolasis de Laubenfels, 1936a: 162.
Ciocalypta- or Collocalypta-like fistule-bearing sponges with
oxeas as megascleres and lacking a tangential ectosomal skeleton. Type species
Previously, Carballo (2000) assigned Collocalypta to Axinellidae
and associated it with Hymerhabdia. We have problems under- Spongosorites suluensis Wilson, 1925: 331 (by original
standing the arguments and classification proposed by Carballo designation).
(2000, 2001) for a group of sponges that all seem closely related
to us. In his proposals, these must apparently be assigned to four Definition
genera of two families (Ciocalypta. Collocalypta. Coelocalypta
and Pseudohymeniacidon, families Halichondriidae and Halichondriidae with trichodragmas. Ectosomal skeleton con-
Axinellidae). We admit that it is possible to rearrange genera of sisting of an ectosomal crust of intercrossing spicules; choanoso-
Halichondrida and redistribute them among families. because char- mal skeleton a confused mass of single spicUles.
acters are not clear-cut. However, we claim to have studied this group
of difficult sponges in its entirety arriving at the present 'state-of- Diagnosis
the-art' classificaton. We propose to consider the above mentioned
genera synonymous with Ciocalypta s.1. until independent support Massive-amorphous to flabellate, with parchment-like ectoso-
is obtained for generic and familial distinctness of these forms. mal tangential skeleton, without a collagenous choanosome and
Porifera' Demospongiae • Halichondrida • Halichondriidae 799

B c
Fig. 7. Ciocalypta porrecta (Topsent 1928c as Coelocalypta). A, type specimen reproduced from Topsent, 1928c: pI. II fig. 6 (size see text). B, spicules
reproduced from Topsent 1928c: pI. VI fig. 4 (size see text). C, Ciocalypta massalis Carter (1883b as Leucophloeus), spicules reproduced from Carter
1883b: pI. XIV fig. 15 (size see text).

without any spongin, resulting in an utterly confused arrangement HALICHONDRIA FLEMING, 1828
of spicules. The spicules include large numbers of trichodragmas
(raphides) which may be flexuous or sinuous in appearence. Two Synonymy
species, both from tropical waters.
Halichondria Fleming, 1828: 520. Raspaigella Schmidt,
Previous review 1868: 25. Amorphina Schmidt, 1870: 40. Eumastia Schmidt, 1870:
42. Pellina Schmidt, 1870: 41. Spuma Miklucho-Maciay, 1870: 13.
Van Soest et al. (1990). ? Tedanione Wilson, 1894: 338. Menanetia Topsent, 1896a: 115.
Halichondriella Burton, 1931 b: 137. Trachyopsilla Burton, 1931 b:
Description of type species 138. Cioxeamastia de Laubenfels, 1942: 265.

Epipolasis suluensis (Wilson, 1925) (Fig. 8A-D). Type species

Synonymy. Spongosorites suluensis Wilson, 1925: 331,
pl. 38 fig. 8, pl. 48 fig. 3; Epipolasis suluensis; de Laubenfels, Spongia panicea Pallas, 1766: 388 (by original designation).
1936a: 162; Van Soest et al., 1990: 41, figs 56, 65.
Material examined. Lectotype and paralectotype: USNM Definition
21297 - Philippines.
Description. Shape lamellate (Fig. 8A). The lectotype is the Halichondriidae with tangential ectosomal skeleton carried by
larger of both original specimens, measuring 8 X 5.5 cm with subectosomal spicule tracts or brushes separated by subdermal
a thickness of 6 mm. One side is the poral side without any visible spaces. Megascleres exclusively oxeas or derivates in a wide size
openings, the other is the oscular side, with abundant 2mm diameter range.
oscules scattered at distances of 3 mm of each other. The surface of
both sides is smooth. The ectosomal skeleton at the oscular side is a Diagnosis
tangential, parchment-like crust of tightly intercrossing spicules,
about 100 IJ..m in thickness, that of the poral side (Fig. 8B) consists of Encrusting, massive, occasionally irregularly branching, or
close-set brushes of spicules forming a peripheral layer of about 400 digitate sponges with smooth or papillate surface. Oscules often on
IJ..m in thickness. The choanosomal skeleton is largely an irregular conical elevations. Surface skeleton well-developed with tangential
mass of single spicules with a few vague tracts, spongin very scarce. bundles of spicules and single spicules intercrossing to form a
Spicules (Fig. 8C) oxeas in a large size range, in several size cate- lighter or heavier built surface crust. Subectosomal spaces usually
gories, in the ectosomal region they measure 140-450 X 7-16IJ..m, in well-developed causing the surface crust to be often rather inde-
the choanosome the range is 140-1350 X 17-32IJ..m. Trichodragmas pendent of the main skeleton and easily peeled off. Choanosomal
(Fig. 8D) straight or more often curved, flexuous or sinuous, 100-228 X skeleton of rather ill-defined bundles of spicules, which at the sur-
5-10 IJ..m. Distribution and ecology: Philippines, Indonesia, reefs. face become oriented perpendicular to the surface crust. They often
Remarks. In many of its characters this genus approaches fan out and carry the surface crust. Many single spicules distributed
Topsentia, but the possession of trichodragmas in the two species randomly. Spongin not visibly present. Spicules oxeas with gradu-
of the genus (also in Epipolasis profunda Diaz et al., 1993) ally tapering sharp points, in a wide size range, often seemingly
makes it likely that Epipolasis is a valid genus. Trichodragmas divisible into a smaller and a larger category but overlap is
are common in other families of Halichondrida (Axinellidae, extensive. Occasionally style-like modifications occur at a low fre-
Desmoxyidae), but within Halichondriidae they are diagnostic. quency. About 110 species distributed over all regions and habitats.
800 Porifera' Demospongiae • Halichondrida • Halichondriidae

Fig. 8. Epipolasis suluensis Wilson, 1925. A, photo of holotype (scale 1 cm). B, SEM image of surface of type (scale 100 !Lm). C, spicules of type
(scale 100!Lm). D, trichodragmas of type (scale lO!Lm).

Remarks appears to be Eumastia Schmidt (1870). Van Soest et al. (1990)

simply considered these genera junior synonyms of Halichondria,
Several species from high latitudes with Halichondria-like but the shared habitus may indicate common ancestry. For that
skeleton architecture share a surface covered in short papillae. For reason, we propose to employ Eumastia as a subgenus within
these several generic names are available, the most senior of which Halichondria s.l.

Key to subgenera of Halichondria

(I) Surface densely covered with short conical papillae ............................................................................................................ Eumastia
Surface smooth or digitate, no continuous cover with conical papillae .......................................................................... Halichondria

SUBGENUS HAUCHONDRIA FLEMING, 1828 1894: 338. Menanetia Topsent, 1896a: 115. Halichondriella
Burton, 1931b: 137. Trachyopsilla Burton, 1931b: 138.
Synonymy

Halichondria Fleming, 1828: 520. Halina de Blainville, 1830:

497 (fide Neave, 1940). Seriatula Gray, 1867a: 515. Raspaigella Type species
Schmidt, 1868: 25. Amorphina Schmidt, 1870: 40. Pellina Schmidt,
1870: 41. ? Apatospongia Marshall, 1892: 16. ? Tedanione Wilson, Spongia panicea Pallas, 1766: 388 (by original designation).
Porifera· Demospongiae • Halichondrida • Halichondriidae 801

Fig. 9. Halichondria panicea Pallas (1766. as Spongia). A, original specimen reproduced from Ellis, 1755: plate XVI fig. D. B, spicules reproduced from
Van Soest et al., 1990: fig. 69 (scale 100 j.Lm). C, neotype BMNH 1964.6.8.6 (scale 1 cm). D-E, SEM images of cross sections of different ZMA specimens
(scale 100 j.Lm). F, tangential ectosomal skeleton in surface view (scale 100 j.Lm).

Definition the typical volcanoe-shaped chimneys up to 4 or 5 cm high.

Oscules are relatively large, conspicuous, 2-4 mm in diameter.
Halichondria with smooth or digitate surface. Sponge body may be up to 25 cm thick and 60 cm across, but much
smaller specimens are the rule. In localities with strong currents
Description of type species specimens may grow out to form longer oscular chimneys closely
adhering and verging toward palmate forms. In deeper or stagnant
Halichondria panicea (Pallas 1766) (Fig. 9A-E). waters, with absence of wave surge, specimens may form masses
Synonymy (restricted). Spongia panicea Pallas, 1766: 388; of anastomosing branches, with oscular chimneys here and there
Linnaeus (in Gmelin), 1791: 3823; Halichondria panicea; on the branches. Consistency is firm, compressible, easily tom.
Fleming, 1828: 520; Johnston, 1842: 114, pIs 10-11. Spongia Colour basically light orange-yellow or pale yellowish green.
tomentosa Linnaeus, 1767: 1299. Spongia papillaris Linnaeus (in However, intertidal specimens exposed to the light may be dark
Gmelin), 1791: 3824; Esper, 1794: pI. Spong. II. Spongia seriata greyish green, presumably due to microsymbionts. Ectosomal
Grant, 1826d: 11. skeleton (Fig. 9D-E) tangential (as is usual for the genus), with
Material examined. Neotype (here designated): BMNH spicules arranged disorderly or in tight bundles of 10-30 f.1m dia-
1964.6.8.6 - Burnham-on-Crouch, Essex (Fig. 9B). Other material. meter, leaving very little open space (usually only up to 100 f.1m in
Numerous freshly collected specimens incorporated in ZMA. diameter) for ostia. Specimens in exposed localities have thicker
Description. Quite variable in shape, related to exposure and more organized ectosomal skeletons. Thickness of the ectoso-
to water movement. Specimens growing in intertidal localities mal spicule crust (Fig. 9C) 200-300 f.1m. Choanosomal skeleton
exposed to the full oceanic surf may be entirely smooth with barely (Fig. 9C) largely confused, but near the surface some organization
visible oscular chimneys. More intermediate environments show into tracts of ca. 50 f.1m diameter is observed in many specimens.
802 Porifera· Demospongiae • Halichondrida • Halichondriidae

These tracts carry the ectosomal reticulation, leaving characteristic (1989). Riisgard et a!. (1993) measured the rate of suspension feed-
subectosomal spaces of up to 100 fLm diameter between them. ing of this species and determined its energy costs at different
There is very little visible spongin. Spicules (Fig. 9B) oxeas only; temperatures. Forester (1979) studied the association between this
size in specimens from Western Europe: 124-482 X 2-15 fLm. species and scallops (Chlamys varia). The scallops allow bread-
Average sizes are: 300 X 7 fLm (Holland), 280 X 8 fLm (France), crumb sponges to overgrow their shells in order to escape predation
360 X 9 fLm (Ireland). Several authors report a subtle but statisti- by starfishes; the sponge receives increased nutrient supply from
cally significant difference in spicule lengths between ectosomal the inhalant current created by the bivalve. Other associated
(shorter) and choanosomal (longer) spicules, but no definite size fauna include endosymbiont nematodes (Bongers, 1983), annelids,
categories can be distinguished due to large overlap. Distribution crustaceans, pycnogonids, echinoderms and fish (Frith, 1976).
and ecology. Northern Atlantic, both along the European coasts Distribution is the entire Northern Atlantic, both along the European
and those of America; from intertidal down to more than 500 m. coasts and those of America. High arctic occurrence has not been
Remarks. Proposal for neotype designation. Pallas (1766) established with certainty, but the species is certainly found in north-
did not describe specimens in his hands, but instead erected new ern Norway, Iceland and northern Canada. Similar sponges occur in
species for pictures and descriptions of pre-Linnean authors. In the the North Pacific and their conspecificity with Atlantic populations is
case of Spongia panicea Pallas, 1766: 388 he refers to drawings of likely. Southward, the species reaches New England and the
Ellis, (1755), and Seba (1758), and also to a description of Ray Mediterranean, although it is uncommon in the Mediterranean itself.
(1724). The specimens depicted in Seba (pI. 96 fig. 4, and pI. 99 Records from the southern ocean (e.g., New Zealand) are doubtful.
fig. 3) are atypical, probably beachworn specimens, from England A separate species H. reticulata (Bowerbank, 1866) (with junior
and North America (no further indication), whereas Ellis pictured an synonym H. topsenti de Laubenfels, 1936a: 133) has been distin-
obvious living specimen (his plate XVI fig. D, here reproduced as guished for species from exposed habitats showing thick ectosomal
Fig. 9A) freshly collected off the shore, somewhere in England, intercrossing tracts and strongly developed surface crust.
probably off the mouth of the Thames. This location is based on Halichondria topsenti is not considered valid for intergrading of
Ellis' (p. 30) remarks on a Tubularia depicted in the same plate specimens with lesser and stronger developed surface skeletons
growing apparently in close contact with the sponge (of which no are observed when a larger series of specimens from various locali-
locality is mentioned by Ellis). An obvious choice of the lectotype ties is studied (Van Soest et a!., 1999).
would have been Ellis' specimen, but this material is no longer Synonymy of Halichondria (Halichondria). The genus
extant. Halichondria panicea has been recorded from allover the Halina De Blainville, 1830: 497 is apparently a junior synonym of
world, at all depths. To ascertain stability in this group of sponges Halichondria, intended to replace Fleming's name without justifi-
with rather vague and variable characters, a neotype needs to be cation and for the same type species. A later genus [Halina]
established. We propose to adopt the specimen BMNH 1964.6.8.6 Bowerbank, 1858 was employed for type species Halina bucklandi
(Fig. 9C) from Burnham-on-Crouch, Essex, about 20 kms from Bowerbank, 1858, which was renamed Dercitus by Gray, 1867a
mouth of Thames. Labelled "28.9.1963, off Power Station, tacking (see Maldonado's chapter on Pachastrellidae).
from bank to bank, Stn. 2, Otter trawl, 12-30ft. Rich yellow-brown The genus Seriatula Gray, 1867a: 515 was erected for
in colour. Quite prolific & found with a number of Polycladi. ColI. type species Spongia seriata Grant, 1826d: 11. A neotype for
Miss S.M. Stone & Mr. J.E Castle" (courtesy Ms Clare Valentine) as this species was designated by Hooper, 1966a: 62 as BMNH
the neotype of Spongia panicea. A photo of the neotype is given in 1847.9.7.14, Johnston's specimen of Halichondria seriata. However,
Fig.9B. Howson & Chambers (1999: 612) discovered that slides of the Grant
Biology and variability. This is a viviparous species; larvae collection are still extant in the University College of London.
have been observed by many authors (e.g., Topsent, 1911; Among these are several slides numbered UCLZ B73, all individu-
Hartman, 1958b; Wapstra & Van Soest, 1987 (SEM photos». They ally wrapped in paper, and on one of the paper wraps it is written
are orange-yellow, oval to oblong, ciliated uniformly, but with a "Spongia seriata Gr variety of S. papillaris" (quote from Howson &
posterior tuft of longer cilia; size up to 600 X 18 fLm. Conflicting Chambers). This slide is here designated the lectotype of Spongia
observations have been made about the sexual cycle of this species. seriata Grant, 1826d. The identity of this material was established as
Sponges in Holland were found to be hermaphroditic (Wapstra & Halichondria panicea (Howson & Chambers, 1999: 612) and
Van Soest, 1987), those in the Baltic gonochoric (Witte & Barthel, accordingly, Seriatula becomes a junior synonym of Halichondria.
1994). Reproductive periods are April to September, with larvae The genus Raspaigella Schmidt (1868: 25) was erected for
being released from June to September. Sometimes parasitized by type species (by monotypy) Raspaigella brunnea Schmidt, 1868:
the alga Tribonema endozooticum (Wille) (Borojevic et al., 1968c). 25. The lectotype LMJG 15330 (designation herein) from Triest,
This species occurs in the intertidal region as well as in the sublit- the paralectotype LMJG 15524 from an unknown locality, and
toral down to more than 500 m. In the intertidal region it occurs on a slide BMNH 68.3.2.29, probably of one of the type specimens,
upper, lateral and undersides of boulders and holdfasts of brown were examined. This sponge forms a mass of irregularly anasto-
algae. It is the commonest intertidal sponge of the Eastern Atlantic. mosing limp branches of 11-16 cm length and 5 mm diameter
In the sublittoral it is likewise common, especially in northern (Fig. lOA). Consistency in the dry condition: fragile, easily broken.
areas, e.g., the coasts of the North Sea. Vethaak et al. (1982) estab- Surface crust easily removable (Fig. lOB). Ectosomal skeleton
lished that its tolerance towards siltation is lower than its sympatric (Fig. 10C) of irregular thin short bundles of oxeas, 2-3 spicules in
sister species H. bowerbanki Burton, 1930c, preferring somewhat cross section, with a high concentration of smaller spiCUles.
more exposed habitats. However, the ecological range is broad and Choanosomal tracts wispy, following mostly the length of the
overlaps considerably with that of H. bowerbanki. Ecophysiological branches, many single spicules distributed randomly. Oxeas in a
aspects such as substrate specificity, growth, biomass, production wide size range, seemingly in two categories, 150-600 X 3-10 fLm
and energy budget were studied by Barthel (1986; 1988; 1991). (Fig. IOD). These characters conform to Halichondria and accord-
Production of faecal pellets was measured by Wolfrath & Barthel ingly Raspaigella is considered a junior synonym.
Porifera· Demospongiae • Halichondrida • Halichondriidae 803

Fig. 10. Halichondria brunnea Schmidt (1868, as RaspaigeUa). A, photo of LMJG lectotype (scale 1 cm). B, SEM image of cross section (scale 100 fLm).
C. ditto of surface (scale 100 j.Lm). D. spicules drawn from slide of type (scale 100 j.Lm).

The genus Amorphina Schmidt, 1870: 40 was erected for who made a subsequent designation of Reniera grossa as type, but
Reniera grossa Schmidt, 1864, Halichondria panicea sensu this designation could not be found in Higgin's paper. Topsent
Bowerbank (1866), Reniera compacta Schmidt, 1864 and Reniera (1925b) suggested that Schmidt himself indicated Reniera grossa
aurantiaca Schmidt, 1864. Several authors (Topsent, 1925b; as the type of Amorphina, but this is strictly speaking not true:
Burton, 1934a; de Laubenfels, 1936a) assumed Reniera grossa to Schmidt only started his list of assigned species with R. grossa, but
be the type of Amorphina. The type specimens of R. grossa, LMJG did not indicate it as type. Burton (1934a) reluctantly accepted that
15329 from Lesina and LMJG 15189, from unknown locality were Reniera grossa is the type based on Topsent's suggestion, although
examined. Since LMJG 15329 (figured in Desqueyroux-Faundez he emphasized that Schmidt intended the genus to be for sponges
& Stone, 1992: fig. 186) is the only specimen with the type locality like Halichondria panicea. This is certainly obvious from
on its label, it is here proposed as the lectotype, and the remaining Schmidt's definition and some of the species formally described by
series of specimens in LMJG are thus paralectotypes. Several other him as Amorphina, e.g., A. genitrix Schmidt, 1870 which is a
specimens are in existence with Schmidt's labels (Strassbourg, Spongosorites (cf. below). Since no formal (correct) type species
Berlin), but their type status is doubtful. Both reexamined speci- designation, (i.e., a species expressly stated to be the type) appears
mens are members of what is now considered Haliclona s.l., to have been made (Hooper & Wiedenmayer's erroneous reference
probably of De Weerdt's Haliclona (Rhizoniera) (De Weerdt, cannot be counted as such), it is proposed to select Halichondria
pers. comm.), with Haliclona-type oxeas of about 160 X 5 f.1m. panicea sensu Bowerbank, 1866 - which conforms to the neotype
The skeletons are irregularly reticulate, and in places somewhat of Spongia panicea Pallas, 1766 selected above - as the type
confused, hence Schmidt's association of this species with species, which makes Amorphina an objective synonym of
Halichondria panicea. The status of Reniera grossa as type of the Halichondria Fleming.
genus Amorphina is, however, a matter of discussion. Hooper & The genus Pellina Schmidt, 1870: 41 was erected for type
Wiedenmayer (1994) indicate Riggin (1877) as the first revisor species (by original designation) Reniera semitubulosa Schmidt,
804 Porifera· Demospongiae • Halichondrida • Halichondriidae

Fig. 11. A-B, Halichondria semitubulosa (Schmidt 1862 as Reniera) (=type of Pellina Schmidt, 1870). A, photo of type specimen (reproduced from
Desqueyroux-Faundez & Stone, 1992) (scale I em). B, SEM image of cross section made from the lectotype (scale 100 j.Lm). C-D, Halichondria minchini
(Topsent, 1896a as Menanetia) (=junior synonym of H. panicea). C, photo of MNHN type specimen (scale I em). D, cross-section made from a slide of
the type (scale I mm). E, Halichondria corticata (Burton 1931b as Halichondriella) (=junior synonym of H. panicea), cross-section reproduced from
Burton (1931 b: fig, 2) (scale 1 mm).

1862, which mayor may not be a junior synonym of Spongia rounded holes of 250 11m diameter, separated by thin tracts of
semitubulosa Lamarck (1813: 455; 1816: 380). Since the Lamarck spicules. The choanosomal skeleton (Fig. lIB) is a largely con-
material is no longer extant, it is impossible to ascertain what the fused mass of single spicules, very few tracts. Spicules oxeas, in
exact properties of this species are. However, Schmidt's material of a large size range but undifferentiated categories, smaller spicules
Reniera semitubulosa is extant in LMJG, and the properties of concentrated at the surface, 250-400 X 3-8 11m. These characters
these specimens objectively determine the nature of Pellina (see conform to Halichondria and it is possible that the species
also De Weerdt, 1986: 114). The type material of Reniera semitubulosa is identical to brunnea described above. The genus
semitubulosa consists of 7 samples (cf. Desqueyroux-Faundez & Pellina has been employed as a Haplosclerida genus by many
Stone, 1992: 17), but only one bears a label with the type locality authors, but this is obviously erroneous.
('Venedig'), LMJG 15320, figured in Desqueyroux-Faundez & The genus Apatospongia Marshall, 1892: 16 was apparently
Stone, 1992: fig. 203 (here reproduced as Fig. lIA). It is here cho- erected for type species Apatospongia fallax Marshall, 1892: 16
sen as the lectotype, while the other specimens all from unknown (monotypy). We have been unable to find the paper, as the volume
localities (LMJG 15616, 15617, 15990, 15614, 15615 and 15991) it is supposed to be published in does not seem to contain it.
are assigned paralectotype status. The lectotype and paralectotype De Laubenfels (1936a: 134) assigned it to the synonymy of
LMJG 15991 were reexamined. This species forms an irregular Halichondria, and we maintain this on his authority.
mass of lobes, surface smooth, consistency firm. The ectosomal The genus Menanetia Topsent (1896a: 115) was erected (by
skeleton is detachable and consists of a fairly light tangential skele- monotypy) for type species Menanetia minchini Topsent (1896a:
ton, mostly of single spicules, with very few tracts, surrounding 115). A type specimen MNHN DT. 2099 from Roscoff, including a
rounded porefields. Subectosomally there is a system of fairly large slide was examined. It is massively encrusting (Fig. lIe), with
Porifera· Demospongiae • Halichondrida • Halichondriidae 805

smooth surface, large oscules, firm consistency. The ectosomal ciliated. Tylotes were also mentioned, but only "very few" were
skeleton is made up of heavily spiculated bundles of 50-70 fLm observed. The description fits perfectly with sponges of the genus
diameter intercrossing to form a tight reticulation with triangular to Halichondria and not with tedaniids on two important counts:
rounded meshes of 50-120 fLm diameter. The thickness of the Halichondria species often have a smaller and a larger category of
ectosomal-subectosomal region is 300-400 fLm. In the choanosome oxeas whereas tedaniids have differentiated spicules including ony-
(Fig. liD) there are wavy bundles of spicules, 30-40 fLm in diame- chaetes, and Halichondria larvae are typically ciliated all-over,
ter following an erratic course with frequent anastomes. In whereas tedaniid larvae have a bare posterior pole. Until Wilson's
between, many single spicules lie in confusion. Oxeas relatively material has been traced, it is not possible to draw a firm conclu-
small, seemingly divisible in two sizes but with considerable over- sion about this synonymy.
lap, 100-280 fLm. This is a specimen similar to Halichondria Van Soest et aZ. (1990) suggested as further synonyms of
reticuZata (cf. above), which is considered the exposed form of Halichondria: CiocaZapata, Cioxeamastia, Eumastia, PyZoderma
H. panicea. Accordingly, Menanetia is considered ajunior synonym and Spuma. Eumastia with junior synonyms Cioxeamastia and
of Halichondria. Spuma is proposed as a valid subgenus of Halichondria, rather than
The genus Halichondriella Burton (1931b: 137) was erected a junior synonym, because of the possession of a mass of hollow
for type species (by monotypy) Halichondriella corticata Burton papillae in combination with a Halichondria type skeletal arrange-
(1931b: 137, fig. 2). A portion of the holotype BMNH ment in several distinct species (cf. below). CiocaZapata possesses
1931.10.28.25 from the wall of the Tromsoe aquarium (Norway) a complement of genuine styles in addition to the Halichondria-
was examined, as well as a fragment BMNH 1938.8.18.2 subse- like skeleton of oxeas, and for that reason is considered valid.
quently collected and incorporated in the Oslo Museum identified PyZoderma has been found to conform to the Poecilosclerida
by Arnesen as H. corticata. This is a cushion-shaped sponge, with- family Dendoricellidae (see Van Soest's chapter on that family)
out visible oscules, with an undulating mammillate surface. The because of compelling similarity of its type species, P. Zatrun-
ectosomal skeleton is detachable with difficulty and consists of culioides with chela-bearing P. demonstrans.
randomly strewn single spicules. Thickness of the ectosomal crust
ca. 40 fLm. The subectosomal lacunae are small, about 50 mm,
separated by spicule tracts of 50-60 fLm fanning out to carry the SUBGENUS EUMASTIA SCHMIDT, 1870
ectosomal crust. The choanosomal skeleton (Fig. liE) is organized
in bundles 20-50 fLm in diameter with 6-8 oxeas in cross section, Synonymy
oriented perpendicular to the surface. Many loose single spicules.
Oxeas relatively thin and short, 160-300 X 1-4 fLm. Since this is an Eumastia Schmidt, 1870: 42. Spuma Miklucho-Maclay, 1870:
aquarium specimen it is to be expected that it would show adapta- 13. Cioxeamastia de Laubenfels, 1942: 265.
tion of shape and poor silica supply. Other features in combination
are in general agreement with Halichondria and the fact that Type species
Halichondria panicea is a dominant sponge in the Tromsoe
environment, make it quite likely that this is a rather delicate form Eumastia sitiens Schmidt, 1870: 42 (by monotypy).
of H. panicea.
The genus Trachyopsilla Burton (1931b: 138) was erected for Definition
type species (by monotypy) Trachyopsilla gZaberrima Burton
(1931b: 138, figs 3-4) in the same paper as the Halichondriella. Halichondriidae with surface covered by short conical fis-
A portion of the holotype from the Trondheim fjord (Norway), tules. Ectosomal detachable tangential skeleton, choanosomal con-
BMNH 1931.10.28.77) was examined. This is a cushion-shaped fused skeleton. Spicules exclusively oxeas.
sponge (Fig. 12A) without visible oscules. The surface is smooth,
consistency friable, soft. The ectosomal skeleton is a loose and thin Previous review
tangential layer of mostly single spicules intercrossing randomly.
The subectosomal skeleton shows subdermal spaces (Fig. 12B) of Van Soest et aZ. (1990, as junior synonym of Halichondria).
200-300 fLm separated by brushes of spicules carrying the ecto-
some. Choanosomal skeleton of ill-defined bundles of ca. 50 fLm Description of type species
diameter, with many confusedly arranged single spicules. Oxeas,
240-340 X 4-8 fLm. Burton emphasized the brushes or palisades at Eumastia sitiens Schmidt, 1870 (Figs 12C-D, 13A-C).
the surface as peculiar to his new genus, but such structures are Synonymy. Eumastia sitiens Schmidt, 1870: 42, pI. V
also found in many Halichondria spp. including H. panicea. Again, fig. 12; Halichondria sitiens; Van Soest et al., 1990: 49, fig. 79.
it is our firm belief that Trachyopsilla is a mere extreme form of Material examined. Holotype: ZMUC - Egdesminde,
H. panicea and accordingly falls into synonymy of Halichondria. W Greenland, 56-100 m, colI. Olrik. Microscope slide: BMNH
The genus Tedanione Wilson (1894) was erected (by mono- 1870.5.3.112 -labelled "Gronland 50". Other material. Specimens:
typy) for T.foetida Wilson, 1894: 338, figs 95-113. This genus was ZMA POR. 1014 and 4707 -Arctic waters.
misinterpreted by Dendy (1922b) as a tedaniid genus (now known Description. Semi-globular cushion (Fig. 12D), yellowish
as Strongylamma, see chapter on Tedaniidae elsewhere in this inside, with white surface, densely covered with cone-shaped papil-
volume). The description and figures of Wilson (no original Wilson lae (Fig. 12C-D) which may be branched and may have an oscule.
material has been unearthed so far, it is not in USNM, K. Riitzler Size up to 15 cm across. Surface smooth, papillae hollow, conical,
in litteris) allow the conclusion that the only spicules likely to be stubby at the summit, or more tapering if they bear a terminal
proper to the sponge are oxeas 350 fLm long and "microscleres oscule. Consistency rather firm. Ectosomal skeleton a tangential
(oxeas) of varying length", and also that the larvae are uniformly 'skin' of intercrossing bundles of oxeas (Fig. 13B); subdermal
806 Porifera· Demospongiae • Halichondrida • Halichondriidae

Fig. 12. A-B, Halichondria glaberrima (Burton 1931 bas TrachyopsiUa). A, photo of BMNH schizotype (scale 1 cm). B. SEM image of peripheral skele-
ton of type (scale 100 /-Lm). C-D, Halichondria (Eumastia) sitiens (Schmidt, 1870 as Eumastia). C, specimen photographed in situ (photo B. Picton).
D, drawing of type specimen (reproduced from Schmidt, 1870: pI. V fig. 12) (scale 1 cm).

lacunae make the ectosome easily detachable. Choanosomal skele- drawing of its habit (Fig. 13D), an encrusting cushion covered with
ton of ill-defined spicule tracts and many loose single spicules; long papillae, resembles Eumastia sitiens. The var. convoluta,
spicule tracts may reach a diameter of ~90 /Lm. In the papillae tube rosa and velamentosa are unmistakably Halichondria panicea.
longitudinal supporting fibres (Fig. 13A) may be as thick as The paper of Miklucho-Maclay is dated 1869 but in fact has not
1500/Lm. Spicules (Fig. 13C) exclusively oxeas, gradually taper- appeared earlier than 1870, thus simultaneously with Eumastia
ing and sharply pointed, in a single but variable size category: Schmidt, 1870. For reasons of stability Eumastia has been chosen
360-1020 X 7-20/Lm. Distribution and ecology. Boreo-Arctic, as the senior of the two names.
down to 100 m. The genus Cioxeamastia de Laubenfels (1942) was erected
Remarks. A second species of Eumastia, E. attenuata (by monotypy) for Cioxeamastia polycalypta de Laubenfels (1942:
Topsent, 1915b, was reported from the Falkland Islands, Southern 265). The holotype USNM 22691 from the northern Hudson Bay,
Atlantic. N Canada, was reexamined. It is a subspherical sponge of 3 cm
The genus Spuma Miklucho-Maclay (1870) was erected diameter, with surface covered by more than 50 short papillae of
(by monotypy) for Spuma borealis Miklucho-Maclay (1870: 13, 4 mm high and 1-3 mm in diameter. Consistency spongy. Ectosomal
pI. II figs 23-30) from the White Sea. No material of this species tangential skeleton of intercrossing spicules, carried by subecto-
was examined. The type material is reported lost by Koltun (1959), somal bundles of aligned spicules, up to 15 spicules in cross sec-
who had access to the collections of the St. Petersburg Museum. tion. Choanosomal skeleton of single spicules in confusion.
The species is described in four varieties, papillosa, convoluta, Spicules exclusively oxeas, 200-700 X 4-12 /Lm. In view of the
tube rosa, and velamentosa, which appear to belong to two different close similarity to Eumastia sitiens this is a likely junior synonym
species. Spuma borealis papillosa is mentioned first and the of this apparent circum-arctic species.
Porifera· Demospongiae • Halichondrida • Halichondriidae 807

Fig. 13. A-C, Halichondria (Eumastia) sitiens (Schmidt, 1870 as Eumastia). A, SEM image of fistule skeleton of ZMA specimen (scale 100/-Lm).
B, surface view of same (scale 100 /-Lm). C, spicules of same (scale 100 /-Lm). D, Halichondria (Eumastia) borealis (Miklucho-Maclay, 1870 as Spuma)
(=junior synonym of H. (E.) sitiens), drawing of type specimen reproduced from Miklucho-Maclay, 1870 (no size data).

HYMENIAClDON BOWERBANK, 1859 Ectosomal tangential skeleton of intercrossing bundles end single
megascleres. Choanosomal skeleton with ascending vague bundles
Synonymy and many loose, confusedly arranged megascleres.

Hymeniacidon Bowerbank, 1859: 286. Stylotella Lendenfeld,

1888: 185. Stylinos Topsent, 1891a: 535. Amorphilla Thiele, 1898:
44. Stylohalina Kirk, 1909: 539. Thieleia Burton, 1932b: 329. Diagnosis
Uritaia Burton, 1932a: 199. Rhaphoxiella Burton, 1934a: 544.
Rhaphidostyla Burton, 1935b: 651. Encrusting to massive, lobate or occasionally irregularly
branching growth forms. Surface irregularly papillate, grooved,
Type species corrugated or smooth. Ectosomal skeleton is a thin tangential or
paratangential crust, often incomplete, membranous, detachable or
Hymeniacidon caruncula Bowerbank, 1859: 286 (by subse- coming off in flakes, containing styles or stylotes. Choanosomal
quent designation; Bowerbank, 1864: 191) (this is considered a skeleton varies from truly confused to disorganized with wispy,
junior synonym of Spongia perlevis Montagu, 1818: 86). slightly plumose ascending spicule tracts of styles or stylotes. Most
species have a characteristic fleshy consistency; oxeas have been
Definition lost completely and megascleres consist only of styles or stylotes.
Several dozens of species, predominantly from temperate and
Halichondriidae with encrusting or massively lobate shape colder waters, but several tropical species have been reliably
and exclusively small styles (>500/-Lm) for megascleres. reported.
808 Porifera· Demospongiae • Halichondrida • Halichondriidae

Fig. 14. A-C. Hymeniacidon perlevis (Montagu, 1818 as Spongia). A, type specimen of Hymeniacidon caruncuncuia Sowerbank, 1859 (=junior syn-
onym of H. perlevis) (scale I cm). S, tangential view of ectosomal skeleton reproduced from Van Soest ef ai., 1990: fig. 87 (scale 100 J.1m). C, blunt end of
apices of styles from ditto (scale lOJ.1m). D-E, Hymeniacidon agminata Ridley, 1884a (=type species of Styiofella). D, SEM image of cross section of
holotype (scale 100 J.1m). E, surface view of same (scale 100 J.1m).

Previous reviews covered with irregularly shaped short papillae and small digita-
tions. Consistency fragile, crumbly. Fresh specimens of this
Van Soest et al. (1990); Hooper et al. (1997). species (Ackers et al., 1992; Van Soest et al., 1999) are orange to
blood-red and have a fleshy consistency. The ectosomal skeleton
Description of type species (Fig. 14B) is a dense crust, with spicules and spicule bundles or
brushes in paratangential (at some angle to the surface), rather than
Hymeniacidon perlevis (Montagu, 1818) (Fig. 14A-C). strictly tangential arrangement. The choanosomal skeleton is less
Synonymy (restricted). Spongia perlevis Montagu, 1818: dense, with wispy bundles following a wavy course to the surface,
86; Hymeniacidon perleve; Burton, 1957: 31 ; Hymeniacidon and many single spicules in confusion. Spicules exclusively styles,
perlevis; Van Soest, I 977a: 270, fig. 5, pI. 3D. Spongia aurea slightly curved, often with a faint tylote swelling subterminally
Montagu, 1818: 87. Spongia sanguinea Grant, I 826d: 203, pI. 2 (Fig. 14C), 152-475 X 3-12 f.lm. Distribution and ecology. NE
fig. 9. Hymeniacidon caruncula Bowerbank, 1859: 286; Bowerbank, Atlantic, from Norway (as Hymeniacidon fallax) and the British
1864: 191, pI. XXXV fig. 372; Bowerbank, 1866: 179; Bowerbank, Isles along most European coasts, penetrating the Mediterranean
1874b: pI. XXXII fig. 1. ]sodictya uni/ormis Bowerbank, 1866: and reaching south and west to Macaronesian islands, intertidal to
329; Bowerbank, 1874b: 139, pI. LV figs 8-10. shallow subtidal, on rocks, Laminaria holdfasts, and also buried in
Material examined. Lectotype of Hymeniacidon caruncula: the sand (then developing relatively long papillae). This is one of
BMNH 1930.7.3.1290 (including spicule slide) - Tenby, Wales. the most common species of Europe.
Description. A rounded thickly encrusting mass 5 X Remarks. The gender of Hymeniacidon remains con-
4.5 X 1.5 cm, orange-brown in dry condition, bearing some resem- tentious. It is true that most names ending in the Greek suffix '-on'
blance to Bowerbank's (1874b) drawing (Fig. 14A). Surface are of neuter gender, but the combination 'Hymeniacidon' is not
Porifera· Demospongiae • Halichondrida • Halichondriidae 809

a known noun in the Greek language, so there is room for doubt. It bearing the number "Bk.751" and with engraved number "R2698"
is also customary to derive the gender of a genus name from the use was reexamined. The skeleton is confused with vague spicule bun-
by its author, and it is clear from Bowerbank's use of the genus dles without visible spongin running to the surface following
name that he considered Hymeniacidon to be female (e.g., a wavy course. Many interstitial spicules in confusion. At the sur-
H. caruncula, H. aurea, H. mammeata, H. perlevis, H. sanguinea). face there is a crust of tangential and paratangential spicules inter-
We propose here to settle this matter by establishing the gender crossing at all angles. The spicules are smooth styles, 220-310 X
as female. Although the type specimen(s) of Spongia perlevis 3-6 f.1m, matching those of Hymeniacidon perlevis as do the other
Montagu (1818) are not with certainty identified in the collec- features. All of these observations match the description
tions of BMNH, there are two unregistered slides labelled by Bowerbank. Stylinos is an obvious synonym of Hymeniacidon
"Hymeniacidon perlevis Montagu (Halichondria Montagu) from by virtue of the perhaps unfortunate type species designation
type 'Devon'" which may represent Montagu's material. The by Hooper & Wiedenmayer. Topsent, in subsequent publications
slides conform closely to Hymeniacidon caruncula. The affinities (Topsent, 1892a; 1892b) made clear that he intended this genus to
of Hymeniacidon are controversial. The genus was made the type be a poecilosclerid, as he erected Stylinos jullieni in 1892a: 137 as
of a family Hymeniacidonidae de Laubenfels, 1936a (see above), type of Stylinos. This species is very probably a member what is
which was subsequently used by many authors for genera with now to be called Ulosa de Laubenfels, 1936a, and consequently
exclusively styles as megascleres and a dendritic or confused Hooper & Wiedenmayer attempted to sink Ulosa into synonymy of
architecture. Genera with such characters are now distributed Stylinos. However, since S. jullieni was not among the species
over various families (Dictyonellidae and Axinellidae), whereas mentioned in Topsent's 1891a publication, it cannot be the type of
Hymeniacidon because of its possession of a (para-) tangential Stylinos. Hooper & Wiedenmayer (1994) were apparently unaware
surface crust is assigned to Halichondriidae, as a genus with exclu- of the true nature of Isodictya uniformis when they designated this
sively truly monactinal megascleres (styles of the other genera are the type of Stylinos.
suspected diactinal modifications). The genus Amorphilla Thiele (1898: 44) was erected for type
The genus Stylotella Lendenfeld (1888) was erected for type species (by original designation) Hymeniacidon sanguinea Grant
species Stylotella digitata Lendenfeld (1888: 185) (by subsequent (1826d: 203, pI. 2 fig. 9). No type material could be found in
designation; Hallmann, 1914: 349) from N Australia. A portion BMNH. It is described as encrusting on the surface of rocks, 6.5 cm
of the type specimen is kept in the collections of BMNH under diameter, more than 1.5 cm thick. Surface flat, but with small,
an unpublished name 'Truncatella micropora'. We agree with round elevations and depressions, colour deep red, numerous small
Hallmann (1914), that this is an obvious junior synonym of oscules. Slimy when tom. Spicules depicted as styles, curved,
Hymeniacidion agminata Ridley (1884a: 466, pI. XLI fig. E, equally thick throughout, but no size given. The described charac-
pI. LIII fig. f), of which the type specimen, BMNH 1881.10.21.347, ters conform to those of Hymeniacidon perlevis and we consider
was reexamined. This is a mass of erect, limp cylindrical branches H. sanguinea a junior synonym of it, thus making Amorphilla an
and digitations tapering to a thin apex. The surface is smooth, small objective junior synonym of Hymeniacidon.
inconspicuous oscules are scattered in low numbers over the The genus Stylohalina Kirk, 1909 was erected for type species
branches. The ectosomal skeleton is a tangential crust of inter- (by monotypy) Stylohalina conica Kirk (1909: 539, pI. 25) from
crossing spicule tracts (Fig. 14E), very much like that found in New Zealand. The type material is probably lost (Barraclough Fell,
Halichondria, overlying a more open choanosomal skeleton with 1950), but the description is clear enough to be able to establish
few tracts and many loose confusedly arranged megascleres (Fig. that this is an obvious junior synonym of Hymeniacidon. The type
14D). Spicules slightly curved styles, 182-271 X 4-8 f.1m. These is described as a flattened, conical mass, with a single large oscules
characters approach Hymeniacidon to such an extent that we pro- at its apex. The surface has minute elevations, small, round or oval
pose to consider Stylotella a junior synonym. The genus name has pores, it is glossy and of dense texture. The ectosomal skeleton is
been used for a variety of sponges different from the type in lack- quite thin, and minutely conulose through projecting spicules, but
ing a surface skeleton, e.g., Stylotella aurantium Kelly & Bergquist they are parallel to the surface. Choanosomal skeleton consists of
(1988). This and similar sponges assigned to Stylotella are irregularly scattered spicules, in low density, sometimes arranged
better placed in the Dictyonellidae genus Stylissa, which has in small irregular bundles. Spicules styles, with slight swelling at
a rather confused choanosomal skeleton of thick styles, but lacks stylote end, slightly curved, average 360 X 10 f.1m.
any surface skeleton. In fact, the species 'Stylotella' aurantium The genus Thieleia Burton (1932b: 329) was erected for type
conforms closely to Stylissa massa (Carter, 1887, as Axinella species (by original designation) Hymeniacidon rubiginosa Thiele
virgultosa massa). Wiedenmayer (1989) employed a subfamily (1905: 421, fig. 44), from the SE Pacific. A paralectotype BMNH
Stylotellinae for poecilosclerid sponges lacking microscleres, 1908.9.24.133 was reexamined. The species is a flat crust
possessing thin styles or strongyles as megascleres, and having (Fig. 15A) with undulating but optically smooth surface, yellowish
their skeletal tracts frequently partly or wholly replaced by sand in alcohol. The ectosomal skeleton is paratangential (Fig. 15B),
columns. Such sponges are now united in the family consisting of brushes diverging at all angles, no tangential spicules
Chondropsidae (see Van Soest's revision of that family elsewhere are apparent. The choanosomal skeleton consists of thin bundles
in this volume), but the genus Stylotella by virtue of the properties running parallel to the surface, in between which there is a con-
of its type species is excluded from this family, and assigned here fused mass of single megascleres. Spicules styles, 325 X 7 f.1m.
to Halichondriidae. This species conforms to Hymeniacidon but for its lack of tangen-
The genus Stylinos Topsent, 1891a: 535 was erected for type tial spicules. Since so far only one such species has been described,
species Isodictya uniformis Bowerbank, 1866: 329 (by subsequent we propose to consider Thieleia a junior synonym.
designation; Hooper & Wiedenmayer, 1994: 295). A microscopical The genus Uritaia Burton, 1932a was erected for type species
slide containing cross sections of the type specimen labelled (by monotypy) Uritaia halichondrioides Burton (1932a: 199, pI. 7
"Isodictya uniformis Bk... from Type, BMNH 1877.5.21.751", figs 3--4, text-fig. 4). The type is not in BMNH, but may still be
810 Porifera· Demospongiae • Halichondrida • Halichondriidae

~======--=-======---
D

Fig. 15. A-B, Hymeniacidon rubiginosa (Thiele, 1905) (=type of Thieleia Burton, 1932a). A, BMNH paralectotype specimen (scale I cm). B, SEM
image of ectosomal skeleton of paralectotype (scale 100 fLm). C-D, Hymeniacidon halichondrioides (Burton, 1932 as Uritaia). C, type specimen repro-
duced from Burton, 1932: pI. 7 fig. 3 (scale I cm). D, spicules from Burton, 1932, text fig. 4 (scale 100 /-Lm). E-F, Hymeniacidon corticata (Thiele, 1905
as Batzella) (=type of Rhaphoxiella Burton, I 934a). E, photo of BMNH schizotype (scale I cm). F, SEM image of cross section of schizotype (scale
250/-Lm). G-I, Hymeniacidon kitchingi (Burton, 1935b as Rhaphidostyla). G-H, spicules of holotype (scale 10 /-Lm). I, SEM image of cross-section of
holotype (scale 100/-Lm).

extant in the Tohoku Imperial University, Japan. It is described and reexamined. This is a rather distinct species forming a flat thick
figured as an irregularly massive sponge (Fig. 15C). The surface crust (Fig. 15E) with smooth but somewhat irregular surface. It has
bears irregular ridges or tubercles; few, small, inconspicuous a thick, dense ectosomal crust consisting of a mass of intercrossing
oscules level with the surface. The ectosomal skeleton is described single spicules (Fig. 15F), overlying a choanosome with large open
and figured as a close-set palisade of brushes of spicules with single spaces and low spicular density. Spicules include all forms
spicules arranged tangentially. The choanosomal skeleton is loose between strongyles and oxeas, up to 430 X 7 /..lm in size.
and irregular, with many spicules in confusion. Spicules styles in Burton erected this genus because of the possession of strongyles
two size categories (Fig. 15D), the smaller concentrated at the sur- in combination with styles. For the time being it is proposed to
face, 420 X 14/..lm and 260 X 7/..lm. The only character in which it class this ill-known species in the genus Hymeniacidon, but should
differs from Hymeniacidon proper is the possession of two distinct other similar species be found, then the genus may be revived.
size categories of styles rather than a single overlapping size range. The genus Rhaphidostyla Burton (1935b: 651) was erected for
In view of the fact that the sizes of both category fall within the size type species (by original designation) Rhaphidostyla kitchingi
range of the styles of the type species of Hymeniacidon (cf. above), Burton (1935b: 652) from Scotland. The holotype BMNH
we propose to consider this a junior synonym. 1934.9.26.79 and four paratypes 1934.9.26.80-83 were reexam-
The genus Rhaphoxiella Burton (1934a: 544) was erected for ined. The holotype is an irregularly massive sponge of 2 X 3 X 2 em,
type species (by original designation) Batzella corticata Thiele with glabrous, uneven surface, yellow/yellow-green in colour.
(1905: 438, fig. 58) from Juan Fernandez Island in the SE Pacific. Fresh specimens (Ackers et al., 1992; Van Soest et al., 1999) are
A type fragment (2 XI cm) and slide, BMNH 08.9.24.151, were pinkish yellow and may have long limp digitations. Oscules rather
Porifera· Demospongiae • Halichondrida • Halichondriidae 811

conspicuous with raised rim. The consistency is soft, compressible. (Fig. 16B). Spicules (Fig. 16D) oxeas to styles, not separable
The ectosomal skeleton is light, with spicule cover rather patchy. in categories, among which a few strongyles with unequal
The choanosomal skeleton (Fig. 151) is distinct in consisting of a ends, 350--1300 X 9-19 /Lm. Distribution and ecology: Italian
confused mass of wispy bundles, each 20--30/Lm in diameter and Mediterranean, 137 m.
following an erratic course, to end eventually at the surface where Remarks. Van Soest et al. (1990) assigned this species to
the spicules fan out to form bundles of more or less tangential ecto- Topsentia, but there is little evidence to support that assignment.
somal megascleres. Spicules slender styles (Fig. 15G-H), relatively The lack of a clearly detachable surface crust as well as a large size
few with telescopic ends, thin, rather uniformly 220--230 X 2--6/Lm. range in spicules are shared with Topsentia, but there the agree-
Burton emphasized the telescopic ends of the spicules and included ment ends. Laminospongia subtilis has the bladed form and indis-
for that reason several species with dendritic spicules tracts and very criminate occurrence of styles and oxeas of some Axinellidae
much longer styles in Rhaphidostyla. These other species are obvi- genera (Axinella, Cymbastela, Phakellia), but these have a well-
ous members of Dictyonella (family Dictyonellidae). The type of organized reticulate choanosomal skeleton entirely lacking in
Rhaphidostyla conforms to Hymeniacidon, but differs from Laminospongia. It is proposed here to leave this distinctive species
H. perlevis in colour, smaller and thinner spicules, and wispy tracts. as a valid genus of Halichondriidae until further evidence for its
Van Soest et al. (1990) suggested that Nailondria systematic position has been obtained.
de Laubenfels (1954) was a junior synonym of Hymeniacidon, but
subsequent study of the type material, revealed that it conforms to
Amorphinopsis (cf. above). SPONGOSORITES TOPSENT, 1896

Synonymy
LAMINOSPONGIA PULITZER-FINAL!, 1983
Spongosorites Topsent, 1896a: 117. Aponastra Topsent,
Synonymy 1927b: 5. Oxeostilon Ferrer Hernandez, 1922: 9.

Laminospongia Pulitzer-Finali, 1983: 546. Type species

Type species Spongosorites placenta Topsent, 1896a: 117 (by monotypy).

Laminospongia subtilis Pulitzer-Finali, 1983: 546 (by Definition

monotypy).
Halichondriidae with smooth surface caused by dense ecto-
Definition somal crust of smaller megascleres; choanosomal skeleton a densely
confused mass of smaller and larger megascleres traversed by
Halichondriidae (?) without special ectosomal skeleton, with directionless vaguely delimited multi spicular tracts. Megascleres
confused choanosomal skeleton; megascleres variable in size and oxeas in several size categories.
shape.
Diagnosis
Diagnosis
Massive-amorphous to subspherical. Surface optically
Laminar shape, with evenly distributed small oscules at one smooth. Oscules inconspicuous. Ectosomal skeleton a thick flaky
of the sides. Skeleton of confusedly arranged single spicules, no crust of paratangentially arranged, relatively thin and small
visible spongin. Spicules oxeas and styles in a large size range. A spicules; choanosomal skeleton with spicules strewn in confusion,
single species which is not certainly a member of this family and occasionally grouped in spongin-enforced tracts, directed mainly
is considered valid only because it does not conform clearly to parallel or oblique to the surface. Many species with aerophobic
any described genus. pigments causing a discolouration to dark or black tinges after col-
lection. Megascleres are oxeas of various sizes, sometimes with
Description of type species their points modified to strongyles or styloids, often centrangulate.
Approximately ten species distributed in deeper waters of all
Laminospongia subtilis Pulitzer-Finali, 1983 (Fig. 16A-D). oceans.
Synonymy. Laminospongia subtilis Pulitzer-Finali, 1983:
546, fig. 50; Topsentia subtilis; Van Soest et al., 1990: 39, fig. 50. Recent reviews
Material examined. Holotype: MCSNG 47175 - Calvi,
Italy. Van Soest et al. (1990), Pomponi et al. (1991), Hooper et al.
Description. The holotype now consists of several frag- (1997).
ments of a single laminar specimen of 120 cm 2 in lateral expansion
and max 3 mm thick (Fig. 16A). The surfaces are both optically Description of type species
smooth but differ in the occurrence of numerous scattered small
oscules, 0.2-0.3 mm in diameter. The ectosome is not detachable. Spongosorites placenta Topsent (1896a) (Fig. 17A-B).
Consistency firm, lamella breaks, when bent. The skeleton lacks Synonymy. Spongosorites placenta Topsent, 1896a: 117.
an ectosomal cover or other specialization (Fig. 16C), the Material examined. Holotype: MNHN DT. 2409
choanosome is halichondrioid, with single spicules in confusion Concarneau, W France.
812 Porifera· Demospongiae • Halichondrida • Halichondriidae

Fig. 16. A-D, Laminospongia subtilis Pulitzer-Finali 1983. A, type specimen reproduced from Pulitzer-Finali, 1983: fig. 59 (scale 1 cm). B, SEM image
of cross section of fragment of the holotype (scale 1 mm). C, surface view of ditto (scale 1 mm). D, spicules reproduced from Pulitzer-Finali 1983: fig. 59b
(scale 50 !Lm).

Description. Shape a flat massive, thick crust. The holotype clearly in excess (up to 600 f.Lm) of those of the type specimen of
in MNHN consists of two small, globular, fragments (Fig. 17A). S. placenta, which may indicate specific distinctness.
Surface smooth, no visible apertures. Consistency firm but friable. The genus Aponastra Topsent (1927b: 5) was erected (by
The ectosomal skeleton is a dense mass of smaller spicules, monotypy) for type species Aponastra dendyi Topsent (1927b: 5,
arranged in all directions (Fig. 17B). It is detachable only as a see also Topsent, 1928c: 159, pI. ill fig. 24, pI. V fig. 12) from 91 m
thicker flake. Choanosome cavernous, but densely spiculated off the Cape Verde Islands. A slide of the type, MNHN DT. 2865
between the cavities, tracts vaguely delimited and not oriented from stat. 1203, was reexamined. This sponge (Fig. 17C) has an
towards the surface. Spicules. smooth oxeas double curve, sharply irregular massive base from which rises a short branch of black
pointed at the endings, large size range and probably at least two colour with smooth surface. The ectosomal skeleton consists of a
size categories, in ectosome a shorter category of approximately layer of tangential smaller spicules, deposited on an inner layer of
70 X 5 f.Lm, in the choanosome a relatively long and thin category tangential larger spicules. The interior is cavernous. The choanoso-
300-360 X 6 f.Lm predominates. Distribution and ecology. W coast mal skeleton consists of single, irregularly arranged spicules.
of France, probably also along most other coasts of Europe, Spicules (Fig. 17D) oxeas, the smaller of which are termed
dredged from deeper waters. 'microrhabds' by Topsent, because of their position and frequent
Remarks. The type was presumed to be lost by Van Soest possession of centrotylote swelling. We believe there are at least
et al. (1990), but a subsequent visit to MNHN revealed that it still two categories, 500 X 16--20 f.Lm and 45-145 X 2-7 f.Lm, but
exists. Van Soest et al. (1999) synonymized Spongosorites pla- common intermediate sizes constitute probably a third category.
centa with Amorphina genitrix Schmidt, 1870. We reexamined The shape and arrangement of the spicules conforms to
some of Schmidt's type specimens from the Copenhagen Museum. Spongosorites and accordingly we propose to consider Aponastra
It is definitely a Spongosorites, but it has the larger spicule sizes a junior synonym.
Porifera· Demospongiae • Halichondrida • Halichondriidae 813

\\ D

Fig. 17. A-B. Spongosorites placenta Topsent, 1896. A, holotype specimen. B, peripheral skeleton of holotype (scale 100/Lm). C-D Spongosorites
dendyi (Topsent, 1927 as Aponastra). C type specimen. D, spicules (both from Topsen!, 1928).

The genus Oxeostilon Ferrer Hernandez (1922: 9) was erected Type species
(by monotypy) for Oxeostilon annandalei Ferrer Hernandez (1922:
9, figs 8-9). No material could be traced. It is described as a thick Anisoxya glabra Topsent, l898b: 234 (by original designation).
mass with undulating shiny surface. The skeleton is confused with
some thicker tracts developed. Spicules are oxeas and strongylote Definition
modifications, often curved abruptly, in a large size range of
320-800 J.Lm, possibly in size categories. Van Soest et al., 1990 Halichondriidae with an ectosomal skeleton conslstmg of
proposed to consider this genus a junior synonym of Topsentia on a crust-like partly tangential or paratangential arrangement of
the grounds that the large spicule size conformed to Topsentia spicules grading into the densely confused choanosomal skeleton.
rather than to Spongosorites. This decision is reversed here, as Ectosome detachable only in flakes due to lack of subdermal cavi-
spicule sizes overlap with those of Spongosorites and also the sur- ties, but choanosome cavernous. Spicules oxeas or modifications,
face characters of S. annandalei are more reminiscent of that in a large size range, including smaller spicules concentrated at the
genus. The species remains ill-known. surface.

TOPSENTIA BERG, 1899 Diagnosis

Synonymy Massive, lobate, or wall-shaped sponges, with brittle and

rough texture; bright colours typically absent; ectosomal skeleton
[Anisoxya] Topsent, l898b: 234 (preocc. by Anisoxya composed of crust of compact, smaller ectosomal oxeas lying tan-
Mulsant, 1856, Coleoptera). Topsentia Berg, 1899: 77. Trachyopsis gentially or paratangentially producing a microhispid surface (but
Dendy, 1905: 147. Viles de Laubenfels, 1934: 13. optically smooth); choanosomal skeleton with very little collagen,
814 Porifera· Demospongiae • Halichondrida • Halichondriidae

I E F I

Fig. 18. A-C, Genus Topsentia. A, Topsentia glabra (Topsent, 1898b as Anisoxya), spicules reproduced from Topsent, 1904b: pI. XIII fig. 4
(scale 100 j,Lm). B-C, Topsentia halichondrioides (Dendy, 1905 as Trachyopsis). B, photo ofBMNH holotype (scale I cm). C, SEM image of cross section
of holotype (scale I mm). D-G, Vosmaeria crustacea Fristedt, 1885. D, drawing of habit, reproduced from Fristedt, 1885, pi .11 fig. 5 (scale 1 cm). E, draw-
ing of spicules from ditto (scale 100 j,Lm). F, watercolour ofVosmaer's (1885a) Inflatella spec. considered a synonym of V. crustacea, reproduced from his
pI. II fig. 5 (scale I cm). G, spicules from ditto (scale 100 j,Lm).

no spongin-cemented fibres, resulting in an utterly confused, direc- Material examined. Lectotype: MOM (not seen). Slide:
tionless arrangement of spicules packed around canals and cavities; MNHN DT. 1169 - Prince de Monaco Expedition, stn. 589,
megascleres are oxeas of a wide size range, usually in 2-3 size Azores.
classes; twisted, bent or doubly-bent spiCUles sometimes present; Description. Massive flat yellowish crust; thickness 2-5 cm.
no raphide rnicroscleres. A dozen species recorded from all three Surface smooth, but rough to the touch, no visible oscules.
oceans, mostly at lower latitudes. Consistency fragile, due to cavernous interior. Ectosome easily
detachable, but comes off in flakes. The ectosomal skeleton con-
sists of an abundance of larger oxeas tangential to the surface, with
Previous reviews spaces in between these filled by irregularly arranged small oxeas
intercrossing at all angles. Choanosome cavernous, skeleton a solid
Van Soest et al. (1990), Hooper et al. (1997). halichondrioid mass of spicules. No spongin, no fibres or tracts.
Spicules (Fig. 18A) oxeas, occasionally stylote modifications, in
three size categories, large oxeas, smooth, fusiform sharply
Description of type species pointed, curved in the middle, 500-1000 X 13-30 fLm; intermedi-
ate oxeas, smooth, fusiform, sharply pointed, slightly curved,
Topsentia glabra (Topsent, 1898b) (Fig. 18A). 200-350 X 5-7 fLm; small oxeas, fusiform, sharply pointed,
Synonymy. Anisoxya glabra Topsent, 1898b: 234; Topsentia slightly curved, concentrated at the surface, 50-120 X 1-4 fLm.
glabra; Berg, 1899: 77; Topsent, 1904b: 131, pI. xm fig. 4. Distribution and ecology. Azores, deep water.
Porifera· Demospongiae • Halichondrida • HaIichondriidae 815

Remarks. The discriminating characters of this genus con- Definition

cern the brittle consistency based on a high spicule density in the
absence of tracts and spongin, and the occurrence of a large size Halichondriidae with tylostyles and oxeas as megascleres, and
range of fusiform oxeas, the largest of which are 600 J.1m long tangential ectosomal skeleton.
or more.
The genus Trachyopsis Dendy (1905: 147) was erected (by Diagnosis
monotypy) for type species Trachyopsis halichondrioides Dendy,
1905: 147, pl. X fig. 10. The holotype BMNH 1907.2.1.44 was Encrusting sponges; type species with long thin papillae.
reexamined. This is a massive sponge with volcanoe-shaped oscules Surface smooth. Tangential ectosomal skeleton of oxeas carried by
(Fig. 18B). Surface rough, consistency hard. Skeleton an extremely bundles of tylostyles. Three species from the cold or temperate
dense confused mass of oxeas (Fig. 18C), of which the larger are Atlantic and Pacific.
600--700 X 30--35 J.1m; small oxeas rare. Although the spicule size
categories are not obvious, the remaining characters of the species Description of type species
conform quite closely to Topsentia and accordingly this genus is
considered a junior synonym. Most other species assigned to Vosmaeria crustacea Fristedt, 1885 (Fig. 18D-G).
Trachyopsis belong to Axinyssa rather than to Topsentia. Van Soest Synonymy. Vosmaeria crustacea Fristedt, 1885: 24, pl. II
et al. (1990) considered the genus Alloscleria Topsent, 1927b to be fig. 5; Burton, 1930c: 672; Arndt, 1935: 37, fig. 60; Alander, 1942:
a junior synonym of Topsentia, ignoring the description of spined 76; Koltun, 1966: 91, pl. XXIV figs 5-6, text-fig. 62. Inflatella
oxeas among the megascleres. Reexamination of a type slide in spec. Vosmaer, 1885a: 24, pl. II fig. 5; Inflatella crustacea;
MNHN revealed that these are indeed present in the slide, and for Levinsen, 1893a: 414. Inflatella robusta Swartschevsky, 1906:
that reason we reassign this genus to the family Desmoxyidae (see 320, pl. 14.
chapter on Desmoxyidae by Hooper in this volume). Material examined. None. The type material is presumed to
The genus Viles de Laubenfels, 1934: 13 was erected for type be in the Stockholm museum.
species Viles ophiraphidites de Laubenfels, 1934: 13 from Puerto Description. Encrusting with numerous thin papillae
Rico (monotypy). The holotype USNM 22334 was reexamined. This (Fig. 18D, F), lateral size 10 X 25 mm, papillae 2-7 mm long.
is greyish pink, massive sponge, 4 cm in diameter, 2 cm high. The Surface smooth, oscules on apex of papillae. Colour white. The
skeleton is confused, and at the surface there is some notable strength- ectosomal skeleton is a detachable membrane entirely composed of
ening in the ectosomal region, with a concentration of smaller megas- intercrossing oxeas. The choanosomal skeleton consists of oxeas
cleres. Megascleres oxeas in a large size range, 300--1000 X and tylostyles in confusion. The skeleton on of the papillae is exclu-
10--20 J.1m. Irregularly curved or twisted forms are not uncommon sively built by tylostyles. Spicules (Fig. 18D-E, G). Oxeas, smooth,
among the smaller oxeas. Characters of Viles ophiraphidites conform curved, sharply pointed, 520--850 J.1m (Arndt, 1935) or 440--1100 X
to those of Topsentia (see also Diaz et aI., 1993: the species is 18-38 J.1m (Koltun, 1966); (sub-)tylostyles, straight, with tyle often
widespread in Caribbean reefs), and accordingly the genus is ajunior subterminal, occasionally true styles, pointed end often blunt or
synonym. rounded, 850--910 J.1m (Arndt, 1935) or 600--1000 X 12-22 J.1m.
Distribution and ecology. Scandinavia, Arctic Ocean, 13-311 m.
Remarks. There are two other species assigned to
VOSMAERIA FRISTEDT, 1885 Vosmaeria, V. laevigata Topsent, 1896a from Roscoff, and V. retic-
ulosa Thiele, 1905 from Iquique, Northern Chile. Both are devoid
Synonymy of papillae, but share the combination of a tangential ectosomal
skeleton of oxeas and vertical bundles of ty10sty1es. The genus
Vosmaeria Fristedt, 1885: 24. Vosmaeria has been assigned variously to Suberitidae (Fristedt),
Po1ymastiidae (Arndt, Koltun) and Axinellidae (Topsent).
Type species However, it fits in Halichondriidae on account of its tangential
skeleton of intercrossing oxeas. The papillate habit and combina-
Vosmaeria crustacea Fristedt, 1885: 24 (by monotypy). tion of diactines and monactines reminds of Ciocalypta.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcaderniclPlenurn Publishers, New York, 2002

Order AgeJasida Hartman, 1980

Rob W.M. Van Soestl & John N.A. Hooper

Agelasida Hartman (Demospongiae) is a small order characterized by the possession of verticillately spined styles (occasionally with
oxeote derivatives). Sponges of the two families belonging to this order are otherwise quite different in their structure. Agelasidae have
elaborate habits, are compressible, and possess a reticulate skeleton of spongin fibres, whereas Astroscleridae are coralline sponges hav-
ing an inorganic limestone basal skeleton upon which is draped a thin organic veneer as the only living tissue. Agelasidae comprise a
single species-rich circumtropical genus. Astroscleridae were formerly joined with other such 'sclerosponges' in a separate class
Sclerospongiae Hartman & Goreau, but subsequent studies (e.g., Vacelet, 1985), on living and fossil coralline sponges clearly demonstrate
them to be polyphyletic, and assignable to various orders of the Demospongiae. Astroscleridae have low species and ecological diversity,
confined to cryptic reef habitats, but are circumtropical in distribution.
Keyword: Porifera; Demospongiae; Agelasida; Agelasidae; Astroscleridae.

DEFINITION, DIAGNOSIS, SCOPE History and biology

Synonymy The two families of this order have had a chequered history.
Until quite recently Agelasidae (partly as Ectyonidae) was consid-
Agelasida Hartman, 1980a: 29. Sclerospongiae Hartman & ered a member of Poecilosclerida (or equivalent earlier higher
Goreau, 1970: 228 (in part). Ceratoporellida Hartman & Goreau, taxon) by most authors, and in the older literature it was simply
1972: 145. defined to contain Poecilosclerida with echinating spicules.
Topsent (1928c) subdivided the former widely defined Ectyoninae
into a number of families including Microcionidae (as Clathriinae),
Definition Myxillinae, Hymedesmiinae, etc., and was the first to restrict
Ectyonidae to Ectyon (at the same time casting doubt over the
Demospongiae with verticillately spined monactine synonymy with Agelas) and Ectyonopsis Carter (1883b: 315). The
megascleres. latter genus is not a member of Agelasidae as the spines of its
spicules are not arranged in verticils, it is here considered a
member of Poecilosclerida: Myxillidae. Up to and including Levi
Diagnosis (1973) all major authors assigned Agelasidae to Poecilosclerida.
However, Bergquist (1978) was the first to realize that this was not
Encrusting, semiglobular, lobate, repent-ramose, branching- consistent with life history and biochemical data.
erect, tubular, globular, of flabellate sponges. Surface optically Bergquist (1978: 168) assigned Agelasidae to the order
smooth. Hypercalcified (basal limestone) skeleton present in one Axinellida on the basis of free amino acid patterns, sterol composi-
of the two families. Skeleton of siliceous spicules with binding tion and on the apparent oviparity. Although this was not formally
spongin or a system of anastomosing spongin fibres, cored and followed by many authors, possibly because Axinellida sensu Levi
echinated by spicules. In hyperca1cified sponges the spicules may (1973) and Bergquist (1978) was obviously polyphyletic, close
echinate the limestone chamber walls, blunt ends embedded in it relationship of Agelas and Axinella was nevertheless generally
and progressively becoming entrapped in the limestone walls. In accepted. Later, additional evidence for this close relationship was
sponges with spongin fibre system, the fibres are echinated by obtained by Chombard et al. (1997) using 28S rRNA sequence
spicules, blunt ends embedded in the fibre, with main fibres also data. In the same study they also found a sister group relationship
having a core of spicules. Megascleres verticillately spined styles, for Agelasidae and Astroscleridae. Hartman (1980a: 29) tentatively
occasionally oxeas, no microscleres. Representatives of both fami- erected the order Agelasida, admitting that there were biochemical
lies of the order produce similar pyrrole-2-carboxylic compounds, affinities with Axinellida, but arguing that in other features
characteristically with a bromine addition. , ... these groups do not seem especially close.'
The family Astroscleridae was first discovered by Lister (1900),
but its sponge nature was contested due to the similarity of the basal
Scope skeleton to that of cnidarians. Likewise, the next discovered represen-
tative of the fanIily, Ceratoporella nicholsoni (Hickson, 1911) was
Two families are recognized, Agelasidae with a single genus ignored by spongologists because it was wrongly classified
and several dozen species, and Astroscleridae with five, mostly in Cnidaria until its rediscovery (Hartman & Goreau, 1966).
monotypical, genera. The order is predominantly circumtropical, Astroscleridae are the main representatives of the calcified demo-
with a few species occurring in the Mediterranean and in subtropi- sponges that were classified in a special class of sponges,
cal waters of the NW Pacific. Sclerospongiae (Hartman & Goreau, 1970), which have subsequently

817
818 Porifera· Demospongiae • Agelasida

been shown to be polyphyletic (e.g., Vacelet, 1985; Van Soest, Hispidopetra in Agelasida, family Astroscleridae because of over-
1984a; Reitner, 1992), and now with corroboratory molecular data all similarity with genera of that family (see Vacelet's chapter on
(Chombard et aZ., 1997). The family has affinities with extinct reef Astroscleridae, this volume). We assume here, that the spines of
builders such as chaetetids and stromatoporoids (Hartman & Goreau, Hispidopetra have become lost.
1972). The monophyly of the family, however, is not well estab- Agelas species are usually large, bright-coloured, conspicuous
lished. The five genera included in Astroscleridae share the basal cal- inhabitants of shallow-water reefs and other clear-water envi-
careous skeleton made of aragonitic sclerodermites. However, the ronments. Greatest recorded depth is 150m. Where known, sexual
basal skeleton of Astrosclera is spherulitic, the spherules being of reproduction is oviparous. Astrosclera willeyana incubates
intracellular origin, whereas the other genera have a clinogonal embryos of the parenchymella type and is thus viviparous.
structure, with extracellularly secreted sclerodermites. Astroscleridae occur predominantly in submarine caves and are
The synapomorphy for the order, viz., the verticillately spined only found in the open at greater depths.
styles, is lacking in one genus of Astroscleridae, Hispidopetra.
For this reason, Reitner (1992: 195) assigned Hispidopetra to Recent reviews
the halichondrid family Hymeniacidonidae (now included in
Halichondriidae). The similarity with non-coralline style-bearing Reitner (1992), Hooper & Wiedenmayer (1994), Chombard
halichondrids is not great, and it is here preferred to retain et aZ. (1997).

KEY TO FAMILIES

(1) Calcareous basal skeleton with soft parts only a thin veneer ....................................................................................... Astroscleridae
Compressible sponge without calcareous skeleton ............................................................................................................. Agelasidae
Systema Porifera: A Guide to the Classification of Sponges. Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Agelasidae Verrill, 1907

Rob W.M. Van Soest

Zoological Museum, University of Amsterdam, P,O, Box 94766,1090 GT Amsterdam, Netherlands, (soest@science,uva,nl)

Agelasidae Verrill (Demospongiae, Agelasida) (nomen conservandum pro Chalinopsididae and Ectyonidae) is a monotypical family with
a characteristic skeleton of spongin fibres cored and echinated by verticillate styles, occasionally oxeas, The spongin fibres are unique and
morphologically quite distinct among the fibres of Demospongiae, showing arched columns and transverse bundles of collagen instead
of the usually arrangement of collagen bundles aligned with the axis of the fibres. The spicules are uniquely shared with sclerosponges of
the family Astroscleridae, and together the two families comprise the order Agelasida. In Agelas 34 nominal species have been described
predominantly from the tropical Atlantic and Indo-West Pacific, with a single Mediterranean and a single South Australian species.
Keywords: Porifera; Demospongiae; Agelasida; Agelasidae; Agelas.

DEFINITION, DIAGNOSIS, SCOPE (l875c: 133, as Ectyonida) and Chalinopsididae Schmidt (1870: 59,
as Chalinopsidinae). The latter name has been used subsequently only
Synonymy by Schmidt himself (in 1880b: 80) and by Fristedt (1885: 44; 1887:
458). Vosmaer (1887: 222) discussed the name Chalinopsididae but
Chalinopsididae Schmidt, 1870: 59. Ectyonida Carter, 1875c: did not adopt it. Article 23.9.1.2 of the ICZN (Anon., 1999) clearly
133. Agelasidae Verrill, 1907: 333. states the conditions under which senior unused names may be sup-
pressed in favour of junior names in general use. The junior name
Definition must have been used at least 25 times by different authors in the past
50 years, while the senior name must not have been used since at least
Agelasida with reticulation of spongin fibres, made up of 1899. The latter rule precludes suppression ofEctyonidae in favour of
transversely arranged collagen bundles; fibres cored and echinated Agelasidae since Ectyonidae was in general use before 1925.
by verticillate styles, occasionally including verticillate oxeas Ectyonidae may be conserved as a generally used junior synonym by
(Fig. 1). suppression of the unused Chalinopsididae Schmidt, 1870 (not used
after 1899, whereas Ectyonidae was used by at least 20 authors in at
Diagnosis least 40 publications (albeit a non-exhaustive count) between 1882
and 1925). Since Chalinopsididae can be suppressed, but Ectyonidae
Growth form is variable, ramose, lamellate, tubular or mas- cannot, we have to conserve the name Agelasidae under ICZN article
sive. The colour is frequently orange or reddish brown and texture 40.2. Agelas was discovered to be a senior synonym of Ectyon so
extremely tough but compressible. The skeleton is reticulate, very Verrill (1907) replaced Ectyonidae by Agelasidae, and since this hap-
fibrous, with a well-developed system of profusely echinated spon- pened well-before 1961, and the name Agelasidae is the prevailing
gin fibres, of which the main or ascending fibres usually contain name, the conditions for article 40.2 are satisfied.
coring spicules. The fibres are peculiar and unique in having
arched transversely arranged collagen bundles. Spicules are styles Taxonomic history
with unique verticillated spines. In some species coring spicules
differ geometrically from those echinating. Oxeas or strongyles Until quite recently, Agelasidae (partly as Ectyonidae) was con-
may also occur along with the styles in some species. The homo- sidered a member of Poecilosclerida (or equivalent earlier higher
geneity of skeletal characteristics makes the external growth form taxa) by most authors, and in the older literature it was simply
of great importance and utility in identification. defined to contain Poecilosclerida with echinating spicules. Fristedt
(1885, 1887) assigned axinellid-type sponges to Chalinopsidinae,
Scope but presumably this was done in ignorance of the true nature of
Chalinopsis and Schmidt's use of the family. Verrill's (1907) deci-
Of six nominal genera only a single valid genus is recognized, sion to erect Agelasidae as a replacement name for Ectyonidae was
Agelas Duchassaing & Michelotti, 1864. done without clarification and too casually to have any impact on
the established use of Ectyonidae by mainstream taxonomists of the
Remarks on synonymy time. Topsent (1928) subdivided the former widely defined
Ectyoninae into a number of families including Microcionidae (as
ICZN Code Article 40.2 (Anon., 1999) has to be invoked in Clathriinae), Myxillinae, Hymedesmiinae, etc., and was the first to
order to be able to continue the use of the well-established family restrict Ectyonidae to Ectyon (at the same time casting doubt over the
name Agelasidae Verrill, 1907: 333. The name Agelasidae was rather synonymy with Agelas) and Ectyonopsis Carter (1883b: 315). The
casually introduced by Verrill in a list of families. It is a clear junior latter genus is not a member of Agelasidae as the spines of its
synonym of two previously proposed names, Ectyonidae Carter spicules are not arranged in verticils, it is here considered a member

819
820 Porifera· Demospongiae • Agelasida • Agelasiidae

1870: 59. Pachychalinopsis Schmidt, 1880b: 80. Siphonochalinopsis

Schmidt, 1880b: 80.

Type species

Agelas dispar Duchassaing & Michelotti, 1864: 76 (by subse-

quent designation; Burton & Rao, 1932: 355).

Definition

Agelasidae with skeleton of spongin fibres cored and

echinated by verticillate megascleres.

Diagnosis

Massive-lobate, encrusting, tubular, branching or flabellate

sponges, often of considerable size, with smooth surface provided
with small rounded and/or key-hole shaped apertures. Colour usu-
ally orange or brownish orange. Consistency toughly compressible,
firm. No ectosomal specialization. Choanosomal skeleton is an
isotropic or anisotropic, occasionally irregular, system of spongin
fibres. The main fibres are distinguishable by a core of megascle-
res, both main and interconnecting fibres are profusely echinated
Fig. 1. Agelasidae characters. A, SEM photo of skeleton showing coring by megascleres. Spicules verticillate styles or oxeas. 34 nominal
and echinating verticillate styles (X 160). B, TEM photo of spongin fibre species distributed over most tropical waters; a single species
showing transversely arranged collagen bundles (X7520) (photos repro- occurs in the Mediterranean. The genus has not been recorded from
duced from De Vos et al., 1992). the tropical East Pacific and tropical West Africa.

of Poecilosclerida: Myxillidae. Burton & Rao (1932: 355) subse- Previous reviews
quently established the synonymy of Ectyon with Agelas, and this was
followed by most contemporary authors. The affinities of Ageias were Wiedenmayer (1977b), Assmann et al. (2001).
left undecided by Burton & Rao (1932). De Laubenfels (1936a: 73)
assigned Agelas to Verrill's Agelasidae, in turn assigned to his large Description of type species
assemblage of 'Phorbasiformes', and included Acanthostylotella (now
Poecilosclerida: Raspailiidae), Scopalina (now Halichondrida: Agelas dispar Duchassaing & Michelotti, 1864 (Fig. 2A-F).
Dictyonellidae) and Ophistospongia Gray, 1867a (now Poecilo- Synonymy. Agelas dispar Duchassaing & Michelotti, 1864:
sclerida: Microcionidae) in the family. The definition provided by 76, pI. XV fig. I.
de Laubenfels (1936a) was erroneous: he emphasized the echinating Material examined. Holotype: ZMA POR. 607 - original
spicules and stated that coring spicules were absent. Levi (1973) label 'Agelas Dispar/St. Martin' (cf. Fig. 2C).
simply repeated this definition. Bergquist (1978: 168), on the basis Description. The holotype (only extant type) is an erect lobe
of chemical and reproductive data, concluded that Agelasidae of 9 cm high and 5 cm in diameter, with an apical depression of
(stripped to a single genus Agelas) belonged in the order Axinellida. 2.5 cm in diameter (Fig. 2B-C). The colour according to Duchassaing
On similar grounds, Hartman (1980a: 29) subsequently erected a & Michelotti's description and plate, is pale orange-brown. The dry
separate order Agelasida for Agelas. holotype is greyish brown. Surface smooth, with scattered small
rounded and keyhole-shaped apertures. Larger rounded openings
Biology are concentrated in the apical depression. Interior strongly
cavernous-clathrate. Skeleton a tight-meshed reticulation of spongin
Agelas species are usually large, bright-coloured, conspicuous fibres (Fig. 2D). Main fibres of 70-110 fJ..m in thickness, cored by
inhabitants of shallow-water reefs and other clear-water envi~n 0-8 spicules, lie at distances of 180-350 fJ..m. Interconnecting fibre
ments. Greatest recorded depth is 150m (Van Soest & St7ritoft, of 20-70 fJ..m in thickness are uncored and are 200-350 fJ..m apart
1988). Where known, sexual reproduction is oviparous. Agelas (Fig. 2F). Both main and interconnecting fibres are echinated by
species have unique secondary metabolites (bromopyrroles), which 1-4 spicules over 500 fJ..m fibre length (Fig. 2E). Spicules verticil-
belong to a wider distributed group of molecules named pyrrole-2- late styles 70-150 X 2.5-7.5 fJ..m with 7-12 whorls of spines
carboxylic derivatives (Van Soest & Braekman, 1999). (Fig. 2G). Distribution and ecology. West Indian region, shallow-
water reefs.
Remarks. The holotype is the only specimen that can be
AGELAS DUCHASSAING & MICHELOTTI, 1864 attributed with certainty to this species, most other records need ver-
ification in view of the confusion over the extent to which the shape
Synonymy and colour of the species is variable. Duchassaing & Michelotti's
water colour of this species, though crude, is obviously not a repre-
Agelas Duchassaing & Michelotti, 1864: 76. Ectyon sentation of the holotype (here reproduced as Fig. 2A), so we must
Gray, 1867a: 515. Oro idea Gray, 1867a: 520. Chalinopsis Schmidt, assume some of the type material has become lost.
Porifera· Demospongiae • Agelasidae • Agelasiidae 821

Fig. 2. Agelas dispar. A, Duchassaing & Michelotti's plate XV fig. I. B-C, holotype ZMA Por. 607 photographed from two sides (scale I em). D-E, SEM
images of skeleton of holotype (scales: D, 500 !Lm; E, 100 !Lm). F, photomicrograph of skeleton of holotype showing coring of ascending fibres (scale
100 !Lm). G, verticillate style of holotype (scale 10 !Lm).

The genus Ectyon Gray, 1867a: 515 was erected for the (1977b) considered E. sparsus a junior synonym of Agelas dispar,
Caribbean Ectyon sparsus Gray, 1867a: 515 (by subsequent desig- but it could also quite possibly be a specimen of Agelas dilatata
nation; de Laubenfels, 1936a: 74). The type specimen (BMNH Duchassaing & Michelotti (1864: 77, pI. II fig. F, pI. XIV fig. 1),
1845.5.12.123) was re-examined, consisting of two halves of an the original material of which is lost. A recent record (Assmann
extensive specimen of approximately 15 X 12X4 cm each. The et al., 2001) of A. dilatata confirmed the specific distinctness of this
skeleton is tight-meshed, anisotropic at the surface with main species. In any case, Ectyon is a clear junior synonym of Agelas as it
fibres of 50-90 11m thickness cored by 2-3 spicules and secondary shares with Age/as dispar the possession of verticillate styles coring
fibres of 30-50 11m thickness uncored. The fibre system gradually and echinating a skeleton of spongin fibres.
becomes isotropic in the interior, with little distinction of primary The genus Oroidea Gray, 1867a: 520 was erected for type
and secondary fibres. Rounded meshes measure 150-300 11m, species (by monotypy) Oro idea adriatica Gray, 1867a: 520.
echination of fibres is sparse with 0-5 spicules over a distance The definition provided by Gray leaves no doubt that he was
of 500 11m. Verticillate styles 120-180 X 7-11 11m, with 11-13 referring to the species now known as Agelas oroides (Schmidt,
whorls of spines. This material has been described and 1864: 35, pI. IV figs 1-2, as Clathria). Clathria oroides (holo-
figured by Carter (187lc: 270, pi. XVII figs 1-3). Wiedenmayer type LMJG 15957, here reproduced in Fig. 3E, copied from
822 Porifera· Demospongiae • Agelasida • Agelasiidae

Fig. 3. Agelas type specimens. A, lectotype of Chalinopsis clathrodes Schmidt, 1870 from ZMUC (scale 1 cm). B, lectotype of Chalinopsis conifera
Schmidt, 1870 from ZMUC (scale 100 ILm). C, lectotype of Spongia arborescens Lamarck, 1814 (reproduced from Topsent, 1932a). D, lectotype of
Alcyonium sceptrum Lamarck, 1814 (reproduced from Topsent, 1933, scale 1 cm). E, lectotype of Clathria oro ides Schmidt, 1864 (reproduced from
Desqueyroux-Paundez & Stone, 1992, scale 1 cm). P, photomicrograph of skeleton of a Jamaican specimen of Agelas conifera (ZMA Por 12778) to show
coring and echination of spongin fibres (scale 250 ILm).

Desqueyroux-Faundez & Stone (1992); there is also a slide in BMNH 60-700/-Lm. Verticillate styles 93-163 X 5-10 /-Lm, with 10-14
1868.3.2.22, from Algiers) shares the possession of verticillate styles whorls of spines.
coring and echinating a skeleton of spongin fibres with Agelas dis- The genus Chalinopsis Schmidt, 1870: 59 was erected for
par. It is a lobate sponge with irregular surface. The interior is Chalinopsis cervicornis Schmidt (1870; 60, pI. V fig. 2a), subse-
densely organic with very few larger canals or cavities. The skele- quently designated type species by de Laubenfels 1936a: 7, and
ton is an irregular reticulation of spongin fibres, without a clear dis- C. clathrodes Schmidt (1870: 60) and C. conifera Schmidt (1870: 60,
tinction between primary ascending and secondary interconnecting pI. V fig. 2b). There is a junior homonym, Chalinopsis Lendenfeld,
fibres, all fibres being 30-70/-Lm in diameter. Some are cored by 1887c, which belongs to Callyspongiidae (Haplosc1erida). The
1-5 spicules, so presumably are primary fibres. Echination promi- type material of Schmidt's type species appears to be lost
nent and uniformly 3-4 spicules over 500 /-Lm fibre length. Meshes (Desqueyroux-Faundez & Stone, 1992), the figure of Schmidt is
Porifera· Demospongiae • Agelasidae • Agelasiidae 823

a non-informative drawing of a spicule, and the description is too Verticillate styles 120-164 X 9-10 /-Lm, with 11-13 whorls of
short to be certain of its identity in modem context. Schmidt's type spines. Topsent noted that the description of Ectyon cylindricus
is generally considered a junior synonym of Spongia arborescens Carter, 1883b: 314 is very similar. Examination of Carter's type,
sensu Lamarck, 1814 (not Pallas, 1766). Topsent (1932a: 104, BMNH 1884.5.26.6, confirmed this both macroscopically and
pi. V fig. 2, as Ectyon cervicornis, here reproduced as Fig. 3C) microscopically. It appears likely that Agelas sceptrum (long single
redescribed Lamarck's type specimen. It has an antler-like growth branches and low number of verticils) and Agelas cervicornis
form with flattened smooth branches which become plate-like (ramose, flattened, with high number of verticils) are distinct
forms lower down though fusion. Oscules are regularly distributed species. Further proof that Schmidt's Chalinopsis is indeed a junior
and flush. The overall resemblance with Schmidt's description of synonym of Agelas are the lectotype specimens (designation herein)
C. cervicornis is compelling. The verticillate styles are stated to of both Chalinopsis clathrodes and C. conifera, which are still
be 90-210 X 4.5-9/-Lm, with 19-23 whorls of spines. Zea (1987: extant in ZMUC, and both contain the typical features of Agelas.
215), quoting a personal communication from Dr Klaus Riitzler, They are here illustrated in Fig. 3A (c. clathrodes) and Fig. 3B
synonymized Chalinopsis cervicornis with Alcyonium sceptrum (c. conifera). A representative photomicrograph taken from the
Lamarck (1814: 163). The type of this was redescribed by Topsent skeleton of a recent specimen of Agelas conifera is shown in Fig. 3F.
(1933: 33, pi. II fig. 5, as Ectyon, here reproduced as Fig. 3D). It is The genera Pachychalinopsis Schmidt, l880b: 80 and
a single long branch of 25 cm long and 2.5 em diameter. Oscules Siphonochalinopsis Schmidt, 1880b: 80 were erected without
are numerous and flush with the surface. Thickest fibres are 70 /-Lm. naming any species and thus are nomina nuda.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Astroscleridae Lister, 1900

Jean Vacelet

Centre d'Oceanologie de Marseille (CNRS-Universite de la Mediterranee, UMR 6540), Station Marine d'Endoume, 13007 Marseille,
France. (jvacelet@sme.com.univ-mrs.fr)

Astroscleridae Lister (Demospongiae, Agelasida) contains five nominal genera all considered valid, and six described species from coral
reef caves and deep-fore reefs from tropical Indo-Pacific and West Atlantic regions. The relationship of the family with Agelasidae is indi-
cated by a spiculation of verticillately spined styles and by biochemical characters. The family is characterized by the presence of a basal
calcareous skeleton made of spherulitic or clinogonal sclerodermites in aragonite, more or less similar to the skeleton of various fossil
taxa such as stromatoporoids and chaetetids.
Keywords: Porifera; Demospongiae; Agelasida; Astroscleridae; Astrosclera; Ceratoporella; Goreauiella; Hispidopetra;
Stromatospongia.

DEFINITION, DIAGNOSIS, SCOPE to that of cnidarians. Another representative of the family,

Ceratoporella nicholsoni (Hickson, 1911) has long been ignored
Synonymy by spongologists because it was wrongly classified in Cnidaria
(Hickson, 1911) until its rediscovery (Hartman & Goreau, 1966).
Astroscleridae Lister, 1900. Ceratoporellidae Hartman & Astroscleridae are the main representatives of the calcified demo-
Goreau, 1972. sponges that were classified in a special class of sponges,
Sclerospongiae (Hartman & Goreau, 1972), which has been shown
Definition to be polyphyletic (Vacelet, 1985). The family has affinities with
extinct reef builders such as chaetetids and stromatoporoids
Agelasida with verticillately spined styles, occasionally smooth (Hartman & Goreau, 1972). An exact correspondence with fossil
styles, and a basal calcareous skeleton made of spherulitic or clinog- genera or families, however, is difficult to establish. The inclusion
onal sclerodermites in aragonite. of possible fossil representatives is not attempted in the present
paper. It appears that true Chaetetidae are more closely related to
Diagnosis the calcified demosponges of the genus Merlia (Gautret et al.,
1991). Astrosclera willeyana incubates embryos of parenchymella-
Encrusting or massive subspherical growth forms. Basal type. The reproduction of the other genera has not been observed.
skeleton composed of a mass of aragonitic sclerodermites, which The family has a circumtropical distribution, with Astrosclera
are either spherulitic and intracellularly secreted, or clinogonal and largely distributed in the Indo-Pacific area and the other genera
extracellularly secreted. Living tissue organized as in typical predominantly Caribbean. These calcified demosponges are local-
Demospongiae, forming a veneer on the surface of the basal skele- ized in submarine caves and epibathyal cliffs, where they could
ton and extending down into calicular skeletal units, cylindrical, replace corals as main reef builders.
meandroid or irregular in form. No tabulae between the tissue-
filled external skeleton and the interior interskeletal spaces, which Taxonomic remarks
are generally filled in solidly with secondary calcium carbonate
deposits. Siliceous spicules styles or strongyles with verticillate The affinity of Astrosclera with Agelasidae is clearly con-
spines, more rarely smooth styles, sometimes secondarily lost, firmed by chemo-taxonomic studies (review in Worheide, 1998),
secondarily entrapped into the calcareous skeleton. cytological characters and molecular sequences (Chombard et aI.,
1997). The monophyly of the family, however, is presently not well
Scope established. The five genera here included in Astroscleridae share
the verticillately spined styles and a basal calcareous skeleton
Astroscleridae contains five nominal genera, all considered made of aragonitic sclerodermites. However, the basal skeleton of
valid: Astrosclera, Ceratoporella, Goreauiella, Hispidopetra and Astrosclera is spherulitic, the spherules being of intracellular ori-
Stromatospongia. gin, whereas the other genera have a clinogonal structure, with
extracellularly secreted sclerodermites. Astrosclera is viviparous,
History and biology and its embryos have been frequently observed, whereas the
embryology of the other genera is unknown (with a faint indication
Astrosclera willeyana is the first discovered Recent demo- of viviparity in Stromatospongia), which could mean that they are
sponge with a basal calcareous skeleton. Although the first descrip- oviparous as Agelasidae. It is thus possible that future studies will
tion was very accurate (Lister, 1900), its sponge nature has been indicate that the family has to be split in Astroscleridae, with
contested (Weltner, 1910) due to the similarity of the basal skeleton Astrosclera, and Ceratoporellidae, with the four other genera.

824
Porifera· Demospongiae • Agelasida • Astroscleridae 825

KEY TO GENERA

(1) Calcareous skeleton with spherulitic microstructure .......................................................................................................... Astrosclera

Calcareous skeleton with clinogonal microstructure ........................................................................................................................... 2
(2) Calcareous skeleton with regular calicular units ............................................................................................................ Ceratoporella
Calcareous skeleton irregular or meandriform .................................. :................................................................................................. 3
(3) Siliceous spicules smooth styles ...................................................................................................................................... Hispidopetra
Siliceous spicules verticillately spined styles or strongyles ................................................................................................................ 4
(4) Encrusting on serpulid tubes ....... ..... ...... ...... ............... ... ........... ... .......... ................ ................ ........... ................. ....... Stromatospongia
Pedunculate and auriculiform, non associated with serpulid ............................................................................................. Goreauiella

ASTROSCLERA LISTER, 1900 skeleton. Living tissue located at the surface and inside the irregu-
larly reticulated lacunae of the superficial (1-8 mm thick) calcare-
Synonymy ous skeleton. No special ectosomal differentiation. Choanocyte
chambers small, ca. 15 fLm in diameter. Exhalant canals bearing
Astrosclera Lister, 1900. a valvule near the aphodus. Tissue containing a high density of
morphologically diverse intercellular bacteria. No well defined
Type species spherulous cells. Basal calcareous skeleton in aragonite, alveolar in
surface and with an organisation more or less similar to that of
Astrosclera willeyana Lister, 1900 (by monotypy). the fossil stromatoporoids, solid in the backfilled central parts.
Tabulae absent. Basal parts externally covered by an epitheca.
Definition Microstructure spherulitic, built up of sclerodermites 10-60 fLm in
diameter, with crystal fibres, 1-3 fLm in diameter, arranged in a
Astroscleridae with a reticulate secondary skeleton composed radiate structure. Sclerodermites intracellularly secreted as gran-
of spherulitic sclerodermites in aragonite and marked with an ules which pass through a spheraster-like stage and are incorpo-
astrorhizal system. rated into the superficial parts of the solid skeleton when their
size is 20-25 fLID. Siliceous styles dispersed in the living tissue.
Diagnosis In some specimens, styles entrapped by their basis within the cal-
careous skeleton, obliquely echinating the basal skeleton. Styles
Massive growth form. Meandroid or reticulate calcareous usually with a swelling in the basal third, but highly variable in
skeleton in aragonite, composed of spherulitic sclerodermites of shape, size and abundance according to the geographic distribution.
intracellular origin and further epitaxial growth. Living tissue In the Indian Ocean, styles 42-87 X 2.5-11 fLm, with verticillated
inside the irregular lacunae of the superficial calcareous skeleton. spines generally well developed; spines absent in the Red Sea.
Superficial canals etching an astrorhizal system on the surface of In West Pacific, acanthostyles 47-164 X 1.7-10.5 fLm, some-
the calcareous skeleton. Siliceous styles of variable shape, often times vestigial or absent, with an irregular spination. Siliceous
echinating the calcareous skeleton, sometimes absent. spicules absent in Central Pacific. Reproduction viviparous, by
incubated parenchymella. Growth rate of the skeleton: ca. 0.2 mmly
Previous review (Worheide, 1998).
Distribution and ecology. Indo-Pacific tropical area:
Worheide, 1998. Red Sea, Indian Ocean (Madagascar, Seychelles, Comoro
islands, Christmas island, Mascarene islands), Pacific (Philippines,
Description of type species Indonesia, Great Barrier Reef, New Caledonia, Guam, Enewetak,
French Polynesia, Japan). Common on undersurfaces of coral
Astrosclera willeyana Lister, 1900 (Fig. 1). rubble, reef cavities, caves, deep cliffs, 1-185 m depth.
Synonymy. Astrosclera willeyana Lister, 1900: 459. Remarks. In the recent species of Astroscleridae, the affini-
Material examined. Holotype: BMNH 1900.10.19.1. Other ties with the Agelasidae indicated by the spicules are confirmed
material. Numerous specimens from Red Sea (Djibouti), by the chemistry of secondary metabolites and by cytology.
Madagascar (Tulear), Europa Island, Comoro Islands, La Reunion, Agelasidae differ, however, by a well developed spongin skeleton
Philippines (Cebu), New Caledonia, French Polynesia. and by oviparity. A similar spherulitic structure of aragonitic
Description. Massive, globular, bulbous, cushion-shaped or sclerodermites is observed in Permo-Triassic fossils belonging to
cylindrical. Young specimens encrusting, growing upwards into various grades of organization (chaetetid, stromatoporoid, sphinc-
a cylindrical structure, with a dead stalk and a living "head" progres- tozoid and other), indicating possible affinities (Gautret, 1986).
sively becoming rounded and larger than the stalk (bulbous shape). A triassic sponge, Astrosclera cuiji Worheide, 1998 has been
In largest specimens, shape subspherical with a stalk hidden by the described as a fossil representative of the genus (Worheide, 1998).
head margins growing down towards substratum. Growth rings vis- There are questionable affinities with Middle Ordovician genus
ible, but without any trace of regeneration or budding. Bathyal Lapidipanis Paquette et aZ., 1983 in which the spherules are consid-
specimens mostly cylindrical. Size usually 0.5-2.5 cm, up to 25 cm erably larger (Paquette et aZ., 1983). Specimens from various
in diameter. Texture stony. Colour orange. Surface smooth, irregu- geographical locations differ by the shape, spination, size and
larly mammillate in some large specimens. Oscules small, abundance of the siliceous spicules, showing six different
2-5/cm2, in the centre of an astrorhizal system etched into the geographical groups. Although an initial genetic study based on
826 Porifera· Demospongiae • AgeJasida • AstroscIeridae

Fig. 1. Astrosclera willeyana Lister, 1900. A, specimen from Philippines (scale 4.4 mm). B, SEM photo of the surface of the skeleton showing astrorhizal
processes, specimen from Philippines (scale 720/-Lm). C, SEM photo of a fracture in the skeleton, with echinating verticillately spined styles, specimen
from Philippines (scale 140/-Lm). D, section through the skeleton and the living tissue, including intracellular calcareous spherules, specimen from
Glorieuses Islands (Indian Ocean) (scale 100 /-Lm). E, SEM photos of verticillately spined styles: a, specimen from Indonesia (scale 10 /-Lm); b, specimen
from Palau (scale 8.3 /-Lm); c, specimen from Philippines (scale II /-Lm); d, specimen from Glorieuses Islands (scale 6.5 /-Lm); a, b & c from Worheide, 1998.

RFLP-PCR ofITS rDNA did not detect any significant differences Definition
between some of these populations (Worheide, 1998), a more
detailed examination using ITS rDNA sequences provided strong Astroscleridae with basal skeleton superficially organized in
support for the presence of at least two additional cryptic species, polygonal calicles and verticillately spined styles.
in addition to Astrosclera willeyana s.s. (Worheide et al., 2002).
Diagnosis
Distribution
Massive growth form; basal skeleton made of sclerodermites
As for type species. with a clinogonal microstructure, superficially organized in polyg-
onal calicular units; styles with regular whorls of spines free in the
CERATOPORELLA mCKSON, 1911 tissue or obliquely echinating the calicle walls.

Synonymy Previous reviews

[Ceratopora] Hickson, 1911. Hartman & Goreau, 1970, 1972; Willenz & Hartman, 1989.

Type species Description of type species

Ceratopora nicholsonii Hickson, 1911 (by monotypy). Ceratoporella nicholsoni (Hickson, 1911) (Fig. 2).
Porifera· Demospongiae • Agelasida • Astroscleridae 827

Fig. 2. Ceratoporella nicholsoni (Hickson, 1911), specimens from Jamaica. A, underwater photograph of a specimen about 15 cm in diameter. B, SEM
photo of the skeleton (scale 84 fLm). C, section through the skeleton and the living tissue; arrows: siliceous styles; sk: calcareous skeleton; ca: canal;
It: living tissue (scale 54 fLm).

Synonymy. Ceratopora nicholsonii Hickson, 1911: 200. regular whorls of spines, free in the living tissue or obliquely
Material examined. Holotype: not seen, Caribbean. Other entrapped in the calicle wall by their basis surrounded by collagen
material. Numerous specimens from Jamaica and Bahamas. microfibrils. Entrapped siliceous spicules progressively dissolved
Description. Massive, mound-shaped when mature, with reg- within the basal calcareous skeleton. Living tissue forming a thin
ularly spaced mamelons; young individuals cone-shaped or pedun- veneer at the surface of the calcareous skeleton and extending down-
culate. Colour orange in life. Mound-shaped specimens up to 100 cm wards into each calicle; total thickness of the living tissue 1.5 mm.
in diameter and 50 cm in thickness on deep fore-reef, extraordinarily Each calicular unit corresponds to a single inhalant and exhalant
tough and heavy. Basal and lateral surfaces of the skeletal mass canal. Canals bearing special valvules. Choanocyte chambers small,
covered by an epitheca showing growth lines. Superficial parts of 20.7 fLm in mean diameter. Exhalant collecting system leaving stel-
the basal calcareous skeleton marked by closely spaced calicles, late depressions (astrorhizae) on the surface of the skeleton.
0.2-O.4mm across and 1-1.2mm deep, irregularly polygonal, rang- Intercellular symbiotic bacteria abundant and highly diverse in
ing from 150 fLm-30O-400 fLm in diameter. The lumen of the cali- shape. Growth rate of the skeleton 0.21-O.23rrunJy in Jamaican
cles are filled in with aragonite in the inner part of the basal skeleton. caves (Willenz & Hartman, 1999). Depth range: 8-184 m.
Basal skeleton made of sclerodermites with a clinogonal microstruc- Remarks. The regular calicular structure of the skeleton is
ture, consisting of closely packed crystalline units that diverge at a reminiscent of the calcified demosponges Merlia Kirkpatrick,
low angle. Siliceous spicules styles, 206-298 x 3.1-4 fLm, bearing 1908, family Merliidae and Acanthochaetetes Fischer, 1970,
828 Porifera· Demospongiae • Agelasida • Astroscleridae

family Spirastrellidae. The genus, however, is easily distinguished Synonymy. Goreauiella auriculata Hartman, 1969: 17.
from these by the microstructure and composition of the skeleton Material examined. Holotype (not seen): YPM 6858 -
and by the spicule complement. Caribbean. Other material. One specimen from Jamaica.
Description. (from Hartman, 1969; Hartman & Goreau,
Distribution 1970 and new material). Auriculate or saucer-like form, with the
edges upturned or curled downward, attached by a broad peduncle.
Monotypic, endemic to Caribbean. Colour light yellow in life. Size up to 16 cm in diameter and 3 mm
in thickness. Basal skeleton thin, made up of aragonitic scleroder-
mites, with numerous arborescent, spinose processes, 0.5-1.1 mm
GOREAUIELLA HARTMAN, 1969 in height, and multibranching patterns of higher processes drain-
ing to the edge of the skeleton. Siliceous spicules strongyles
Synonymy bearing whorls of spines, 35-124 X 1.3-1.9/-tm (mean 60-68 X
2.3-2.7 /-tm, becoming embedded in the basal skeleton fIrst by one
Goreauiella Hartman, 1969. end provided with rounded knobs. Living tissue forming a thin
veneer fIlling the space between the superficial processes of the
Type species basal skeleton. Oscules 300 /-tm in diameter, opening out along the
edge of the sponge. Eurypylous choanocyte chamber 18-20 /-tm in
Goreauiella auriculata Hartman, 1969 (by monotypy). diameter. Occasionally associated with zoanthids that induce the
formation of processes in the calcareous skeleton. Depth range:
Definition 8-70 m in caves and deep fore-reef.
Distribution. Monotypic, endemic to Caribbean.
Pedunculate, auriculiform Astroscleridae with verticillately
spined strongyles and with basal skeleton superficially covered
with arborescent processes. HISPIDOPETRA HARTMAN, 1969

Diagnosis Synonymy

Auriculate or saucer-like form, attached to the substratum by Hispidopetra Hartman, 1969.

a broad peduncle. Siliceous spicules with verticillate spines,
strongyles or truncated styles. Aragonitic basal skeleton with Type species
superficial arborescent processes, with raised patterns forming
astrorhizae running to the edge of the skeleton. Hispidopetra miniana Hartman, 1969 (by monotypy).

Description of type species

Goreauiella auriculata Hartman, 1969 (Fig. 3).

Fig. 3. Goreauiella auriculata Hartman, 1969. Siliceous strongyles Fig. 4. Hispidopetra mmwna Hartman, 1969. Siliceous styles (scale
(scale 12.3 fLm), from Hartman (1969). 62.5 fLm), from Hartman (1969).
Porifera· Demospongiae • Agelasida • Astroscleridae 829

~:: .••••• , < • :.~

c
~~.~ : . ..
C
~=::: ....:=.

o
Fig. 5. Stromatospongia vermicola Hartman, 1969. A,holotype YPM 6376 (scale 16.5 mm), from Hartman (1969). B , polished section through a specimen
from Jamaica; It: living tissue; se: tube of serpulid worms; sk: calcareous skeleton with siliceous verticillately spined styles (scale 3l2!Lm).
C, acanthostyles (scale 30 !Lm), from Hartman (1969). D,three dimensional schematic representation; Ar: aragonitic skeleton; c: choanosome; DM: dermal
membrane; EC: exhalant canal; IS: inhalant space or vestibule; 0: osculum; S:spicule (scale 1 mm), from Willenz & Hartman (1989) .

Definition Synonymy. Hispidopetra miniana Hartman, 1969: 12.

Material examined. Holotype (not seen): YPM 6853 -
Encrusting to massive Astroscleridae with smooth styles and Caribbean. Other material. One specimen from Jamaica.
basal skeleton bearing long superficial processes. Description. (from Hartman, 1969; Hartman & Goreau,
1970 and new material) . Encrusting to massive. Colour carmine
Diagnosis to vermilion in life. Size reaching 15 cm in diameter and 3 cm in
height. Basal skeleton made up of aragonitic sclerodermites, with
Encrusting to dome-shaped or massive. Siliceous spicules superficial arborescent processes, up to 7 mm high, with rounded
smooth styles, partially or totally embedded in the calcareous spines and protruding siliceous spicules, without astrorhizae.
skeleton. Basal skeleton devoid of astrorhizae, with surface Spicules smooth styles, range of means 269-301 X 5.4-7.4 /Lm,
covered with conspicuous spinose processes up to 7 mm high. with an overall range 125-818 X 1.3-10.4 /Lm, partially (by the
head) or totally entrapped in the basal skeleton. Eurypylous
Description of type species choanocyte chambers, 6-18 /Lm in diameter. Frequently overgrown
by serpulid worms. Depth range: 10-95m in caves and deep
Hispidopetra miniana Hartman, 1969 (Fig. 4). fore-reef.
830 Porifera· Demospongiae • AgeJasida • Astroscleridae

Remarks. Well characterized among other astrosc1erids by Description of type species

its vermilion colour, smooth styles and well developed superficial
processes. Stromatospongia vermicola Hartman, 1969 (Fig. 5).
Synonymy. Stromatospongia vermicola Hartman, 1969: 3.
Distribution Material examined. Holotype (not seen): YPM 6376 -
Caribbean. Other material. Several specimens from Jamaica.
Monotypic, endemic to Caribbean. Description (from Hartman, 1969; Hartman & Gareau,
1970; Willenz & Hartman, 1989 and new material). Encrusting,
always growing in association with tangled masses of serpulid
worms. Colour apricot to light salmon pink in life. Size up to 40cm
STROMATOSPONGIA HARTMAN, 1969 in diameter and 10 cm in height. Basal skeleton intimately mixed
with serpulid worm tubes, superficially marked by numerous
Synonymy upright, multibranched processes, 1.5-2mm high, with the living
tissue extending down into irregular spaces left between the
Stromatospongia Hartman, 1969. processes. Sc1erodermites with aragonite crystals radiating in
all directions from centers of calcification usually located around
Type species spicule heads. Siliceous verticillately spined styles, 75-519 X
3.3-13 /-Lm with a mean of 165-187 X 6.2-8 /-Lm, with whorls of
Stromatospongia vermicola Hartman, 1969 (by original spines on the shaft, more or less completely overgrown by the arag-
designation). onite. Living tissue forming a thin veneer between the processes of
the surface of the basal skeleton. Eurypylous choanocyte chambers,
Definition 16-20 /-Lm in diameter. The association with serpulids appears oblig-
atory. Depth range: 10--95 m under overhangs of deep fore-reefs.
Encrusting to massive Astrosc1eridae with verticillately Remarks. The association with serpulids, which appears to
spined styles, associated with calcareous tubes of serpulid worms. be a good distinctive character for the Caribbean species, is less
characterized for the Pacific representative (Hartman & Goreau,
Diagnosis 1976), and the genus is possibly synonym with Ceratoporella
(Hartman & Goreau, 1972; Willenz & Hartman, 1989). Its superfi-
Encrusting to massive, with an aragonitic basal skeleton asso- cial skeleton, however, is devoid of the regular calic1es highly
ciated with the calcareous tubes of serpulid worms. Siliceous characteristic of Ceratoporella.
spicules with verticillately spined styles more or less completely
entrapped in the basal skeleton. Surface of the basal skeleton Distribution
ornamented with processes 0.8 to 2 mm high. Living tissue forming
a thin veneer in the spaces between the processes of the basal Mostly Caribbean. One species described from the Central
skeleton. Pacific (Hartman & Goreau, 1976).
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Order Haplosclerida Topsent, 1928

Rob W.M. Van Soest1 & John N.A. Hoope.-2

Haplosclerida Topsent (Demospongiae) is characterized by possession of an isodictyal skeleton of diactinal megascleres. The megascleres
are relatively short fusiform oxeas, or short compact strongyles. Microscleres are restricted to microxeas/strongyles, sigmas, toxas, raphides
and amphidiscs. The sigmas and toxas may be interpreted as a trait shared with the order Poecilosclerida. A large number of common shal-
low-water marine sponges and all sponges occurring in freshwater are considered members of this order. To accommodate the high diver-
sity of groups and habitats, and in acknowledgement of the continuing debate about monophyly of the order, three suborders are now
recognized: Haplosclerina with families: Callyspongiidae, Chalinidae and Niphatidae; Petrosina with families Calcifibrospongiidae
Petrosiidae and Phloeodictyidae; and Spongillina with families Spongillidae, Malawispongiidae, Metaniidae, Metschnikowiidae,
Palaeospongillidae, Potamolepiidae and Lubomirskiidae. Haplosclerina and Petrosina appear closely related morphologically and are con-
troversial higher taxa. They may be conveniently separated by the absence of a clear anisotropic skeletal structure and the apparent
oviparous reproduction in Petrosina. Spongillina appear more distantly related and are characterized by absence of a tangential ectosomal
skeleton, possession of spined megascleres and unique amphidisc microscleres, and gemmules with an elaborate anatomy and physiology.
Keywords: Porifera; Demospongiae; Haplosclerida; Haplosclerina; Petrosina; Spongillina subord.nov.

DEFINITION, DIAGNOSIS, SCOPE dubbed microxeas or microstrongyles. These are usually concen-
trated in the ectosome. Megascleres are usually smooth, but in one
Definition suborder are frequently finely spined. Microxeas occurring in that
same group are also invariably spined. Further microscleres in the
Demospongiae in which the main skeleton is partially or order are sigmas, toxas and raphides. In one suborder there is
entirely composed of an isodictyal anisotropic or isotropic, occa- a unique microsclere type, the amphidiscs, associated with the gem-
sionally alveolate reticulation of spongin fibres andlor spicules, mule resting stage. Sigmas and toxas are often characteristically
with uni- to multi spicular tracts of diactinal spicules forming trian- angularly bent, making them distinct from such microscleres in other
gular, rectangular or polygonal meshes. Megascleres are exclusively orders. Two of the three suborders share secondary metabolites of
oxeote or strongylote, bonded together with collagenous spongin the pyridine and acetylene compound types. One monotypical genus
or enclosed within spongin fibres; microscleres, if present, may has a sclerosponge basal skeleton.
include sigmas andlor smooth toxas (both frequently centrangu-
late), microxeas or microstrongyles, and in one group amphidiscs. Scope

Diagnosis Three suborders and 13 families are recognized. Together

these sponges occur in all habitats, including freshwater, in all seas
Encrusting, massive, lobate, tubular, arborescent, flabellate or and on all continents. The order comprises the highest biodiversity
excavating sponges. Habit, colour and oscular features broadly vari- of all sponges in terms of species and habitats.
able. Consistency generally compressible to soft, brittle or hard and
incompressible in one group. Ectosomal skeleton usually unispicu- Taxonomic history
lar, tangential, regular reticulation of single spicules, bundles or
spongin enforced fibres. It may be frequently absent or occasionally Synonymy. Haplosclerina Topsent, 1928c. Haplosclerida
developed into a thick impenetrable crust. Choanosomal skeleton de Laubenfels, 1955b. Nepheliospongida Bergquist, 1980.
a regular isodictyal reticulation of megascleres encased in variable Petrosida Boury-Esnault & Van Beveren, 1982.
amounts of spongin. The reticulation may take the form of a strictly History of Haplosclerida. An extensive historical account
unispicular-isotropic skeleton. Frequently it is anisotropic with of ideas of classification of marine sponges now united in this
thicker or thinner ascending tracts interconnected by single order may be found in De Weerdt (1985: 16). For the freshwater
spicules or thin tracts. In some groups the skeletal tracts are sponges, a survey of ideas on classification are presented elsewhere
arranged in thickly alveolar fashion in which meshes may become (Manconi & Pronzato, this volume). It is beyond the present chap-
rounded and result in a disoriented reticulation. Spicules usually ter to repeat these, and only a few major ideas are here elaborated.
are relatively short fusiform sharply pointed oxeas ('cigar- The marine sponges were originally distributed among two groups,
shaped'), which in general may be easily differentiated from oxeas chalinid sponges with spongin as a major component of the skele-
in other orders. Strongyles are also common and it is not infrequent ton, and renierid sponges without visible spongin. Such a subdivi-
that they derive from juvenile oxeote stages. Oxeas and strongyles sion was already employed by Schmidt (1870), and was also used
may also occur together, then often in different size categories, the by Vosmaer (1887) and Ridley & Dendy (1887), for example,
smallest of which may be functional microscleres and then are Renierid sponges were frequently associated with the genus

831
832 Porifera' Demospongiae • Haplosclerida

Halichondria and some other halichondrid genera. Topsent postulated to incubate their larvae, the second, Petrosida was pos-
(1928c) was the first to include both types of sponges into a single tulated to be oviparous. So far this distinction is upheld in various
group, and to disassociate them from halichondrids. This was fol- studies (e.g., Wapstra & Van Soest, 1987; Han & Loya, 1990;
lowed by most major authors since then. De Laubenfels (1936a) Fromont, 1994; Fromont & Bergquist, 1994). In addition, the sec-
attempted to compromise the integrity of Haplosclerida by adding ond group would have unique chemistry by exclusively possessing
a group of chelae-bearing poecilosclerid genera into it, which he sterols with a cyclic sidebranch. Subsequent studies have failed to
called 'Desmacidonidae'. The contents of his Desmacidonidae are substantiate the chemical distinctness of the Petrosida (cf. Fromont
at present distributed over many different suborders and families of et aI., 1994), and other chemical characters (notably 3-alkyl piperi-
Poecilosclerida and all have been excluded from Haplosclerida dine derivatives, cf., Andersen et al., 1996 and straight-chain
long since. A recent attempt (Hajdu et al., 1994b) to reintroduce acetylenes, cf. Van Soest et al., 1998) have been found to occur
a chela-bearing family (Isodictyidae) into Haplosclerida likewise over both groups, re-establishing their close relationship.
was refuted (Samaai et al., 1999) and is not adopted in the present Nevertheless, the two groups are recognizable and definable on the
volume. Chalinid and renierid sponges continued to be recognized basis of skeletal architecture in addition to the difference in repro-
within Haplosclerida as basal stemgroups representing two diverg- ductive strategy. Since both show many similarities in spicule form
ing lines of evolution (cf. Griessinger, 1971; Levi, 1973). They and size, and share unique chemistry, they are here recognized as
were employed as 'families' Haliclonidae and Renieridae, with suborders, along with the freshwater sponges. The sclerosponge
a third family Gelliidae for taxa with microscleres. Independently, genus Calcifibrospongia is newly assigned to Petrosina in a family
a fourth overlapping family Adociidae was employed by some of its own, Calcifibrospongiidae.
authors. Subsequently the classification was refined and diversified Biology. Marine Haplosclerida are typical inhabitants of
(Van Soest, 1980; Bergquist & Warne, 1980; Bergquist, 1980; shallow-water and intertidal habitats, reefs and mangroves, where
De Weerdt, 1985). Van Soest, followed by De Weerdt, recognized they form a colourful and striking element of the filter-feeding
five families among marine haplosclerids, three of which may be community. Several species may reach a large size and presumably
regarded to belong in the 'chalinid' group, and two in the 'renierid' these represent long-lived microhabitats for a large variety of
group. However, continuation of the terms chalinid and renierid for symbionts. One genus is excavating limestone substrates (corals,
the two groups was demonstrated to be impossible, as it was made shells) and several groups may live buried in the sediment. Larvae
clear that Reniera itself belonged to the chalinid group. Bergquist of at least one suborder have a characteristic ciliation pattern
(1980) proposed to subdivide the order into two distinct (and consisting of a 'skirt' of longer cilia surrounding a bare posterior
according to her unrelated) orders, viz., Hap10sclerida s.s. and pole. Freshwater sponges with few exceptions have gemmules as
Nepheliospongida, again more or less covering the chalinid vs. the a resting stage to tide them over adverse conditions (ice, drought).
renierid lines. The name Nepheliospongida was later replaced by The gemmules also enable them to cross large areas of dry land by
Petrosida (see Boury-Esnault & Van Beveren, 1982; Hartman, means of wind transport or bird's feet.
1982), because the fossil genus Nepheliospongia could not be Previous reviews. Griessinger (1971), Bergquist & Warne
linked with the recent members of the group on convincing evi- (1980), Van Soest (1980), Desqueyroux-Faundez (1984, 1987),
dence (amongst others, the presence of spicules in Nepheliospongia de Weerdt (1985, 1986), Fromont (1991, 1993), Penney & Racek
could not be demonstrated). The first order, Haplosclerida s.s. was (1968).

KEYTOTHESUBORDERSOFHAPLOSCLERIDA

(1) Marine sponges .................................................................................................................................................................................... 2

Freshwater sponges ..................................................................................................................................................... ,...... Spongillina
(2) Skeleton anisotropic, i.e., consists of clearly recognizable ascending spicule tracts or fibres interconnected at regular distances by
secondary spicule tracts or fibres. Larvae are incubated ............................ ................................................................... Haplosclerina
Skeleton isotropic, i.e., consists of a reticulation without a clear orientation and without distinction in primary ascending and secondary
interconnecting spicule tracts or fibres, skeleton densely confused. Larvae unknown, presumably oviparous .................... Petrosina
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademicIPlenum Publishers, New York, 2002

Suborder Haplosclerina Topsent, 1928

Rob W.M. Van Soest1 & John N.A. Hooper2

Haplosclerina Topsent (Porifera, Demospongiae) are here newly defined in a new content, although this follows earlier proposals for
a subdivision of Haplosclerida at the ordinal level made by Bergquist (1980a). In the present concept, Haplosclerina Topsent, 1928c are
marine Haplosclerida characterized by the possession of a regular anisotropic skeleton, in which primary ascending spicule tracts or fibres
are invariably recognizable, even in unispicular skeletons. In this respect they differ from the suborder Petrosina, which has the skeleton
isotropic ally organized. Spicules are smooth oxeas or strongyles in a single size category (Petrosina frequently have differentiated size
categories). A further difference, which appears to be valid after exhaustive testing, is the viviparous reproduction (Petrosina are probably
all oviparous). Haplosclerina differ from freshwater sponges that show some similarities, here united in the suborder Spongillina
Manconi & Pronzato, in having a tangential ectosomal skeleton (with few exceptions), and by the lack of special microscleres protecting
the gemmules. Also, spicules in Spongillina are frequently spined. Three families, Chalinidae, Callyspongiidae and Niphatidae, are
recognized, differing in the organization of the skeleton. Chalinidae have a unispicular ectosomal skeleton and a lightly built paucispicu-
lar choanosomal skeleton; Callyspongiidae have a two-dimensional, often double- or triple-meshed ectosomal skeleton and usually a
rectangular-meshed spongin-rich choanosomal skeleton; Niphatidae have a three-dimensional, often irregular ectosomal skeleton, and
a pauci- or more often multispicular choanosomal skeleton, with strong primary fibres or tracts.
Keywords: Porifera; Demospongiae; Haplosclerida; Haplosclerina; Callyspongiidae; Chalinidae; Niphatidae.

DEFINITION, DIAGNOSIS, SCOPE rectangular. Niphatids usually have multi spicular tracts consoli-
dated by a less obvious spongin sheath and the primaries often
Synonymy dominate the skeleton causing a more irregular and coarse skeleton
than that of callyspongiids. Spicules consist of smooth diactinal
Haplosclerina Topsent, 1928c. megascleres (oxeas or strongyles), and an erratic and presumably
vestigial presence of microscleres, usually sigmas and/or toxas,
Definition rarely microxeas.

Marine Haplosclerida with an anisotropic choanosomal skele- Scope

ton; spicules smooth oxeas or strongyles of a single size category.
In the concept of Haplosclerina presented here, three fami-
Diagnosis lies are included: Chalinidae Gray, 1867, Callyspongiidae
de Laubenfels, 1936 and Niphatidae Van Soest, 1980. Together these
Sponges with thinly or thickly encrusting, cushion-shaped, comprise a complement of sponges occurring all over the world in
massive growth forms, commonly with oscular chimneys or all marine habitats. All three families are highly speciose and diverse
mounds; or ramose, vase-shaped, tubular, lamellate, fan-shapes. in form and size. Tropical shallow-water marine habitats such as
Consistency often soft, compressible, resilient, but sometimes may reefs and mangroves harbour many haplosclerine sponges, but they
be firm, stiff, or brittle to hard. Ectosomal skeleton may be absent, are also frequently observed in temperate and cold water, especially
strictly unispicular-tangential, multispicular-tangential or three- Chalinidae. A few are known to occur in deep water.
dimensional ('paratangential'), single-meshed, double-meshed or
triple-meshed, with the larger primary meshes formed by pauci- or
multispicular fibres or spicule tracts, and the secondary or tertiary TAXONOMIC HISTORY
meshes unispicular. Also, in some niphatids dense multi spicular
brushes of spicules obscure such ectosomal meshes. The choanoso- For a more extensive history of Haplosclerina we refer to the
mal skeleton is always anisotropic, with pauci- or multi spicular chapter on the order Haplosclerida (Van Soest & Hooper, this vol-
primary or ascending fibres or tracts interconnected by uni-, pauci- ume). In summary, from the relatively diffuse earlier classifications,
or multi spicular secondary tracts, which may in tum be intercon- the existence among sponges now classed in Haplosclerida two
nected by tertiary tracts. Chalinids usually have paucispicular types of skeletal structure emerged: 'renierid' and 'chalinid'.
primary tracts and unispicular interconnecting spicules, but even in These are now formalized at the subordinal level. The suborder
some entirely unispicular chalinids the primary lines are subtly dis- Haplosclerina is newly erected in this volume, but since its contents
tinct from the interconnecting lines. Callyspongiids tend to have largely cover Topsent's (1928c) Haplosclerina, and in fact its name
fibres with a large amount of spongin and a relatively weakly is a homonym of Topsent's group, it is considered unnecessary to
developed core of spicules, which may be frequently thin or vestig- claim authorship by the present authors. Bergquist (1980a), on the
ial, occasionally entirely absent. These skeletons are often neatly basis of assumed vivipary and distinct chemistry, distinguished

833
834 Porifera' Demospongiae • HaploscIerida • HaploscIerina

a restricted order Haplosclerida, covering more-or-less the same of skeleton is the more ancestral type for Haplosclerida, and that the
groups as proposed here. isodictyal or alveolar skeleton of many Petrosina is derived from it.
As such, the anisotropic skeleton cannot serve as a synapomorphy
of Haplosclerina. Megascleres occurring in a single category is
REMARKS shared with some Petrosina (e.g., Xestospongia) and Spongillina.
Thus, we are left with the ectosomal skeleton which according
Synapomorpbies to De Weerdt (1989) is 'organized', presumably meaning it is retic-
ulate, whereas that of Petrosina is usually crust-like with tangential
The vivipary of all Haplosclerina was confirmed by a series of spicules intercrossing. In Spongillina it is usually absent.
studies (e.g., Wapstra & Van Soest, 1987; Fromont, 1994; nan & De Weerdt's (1989) character analysis of the five families
Loya, 1990, etc.), but the chemical distinctness could not be con- of marine Haplosclerida (Spongillina were excluded) suggested
firmed (e.g., Fromont et al., 1994; Andersen et al., 1996; Van Soest that two distinct clades exist: one comprising Chalinidae,
et al., 1998; Van Soest & Braekman, 1999). Vivipary is shared with Callyspongiidae and Niphatidae; and the other comprising
many poecilosclerid families and most dictyoceratids and dendrocer- Petrosiidae and Phloeodictyidae. Thus, a division of the order
atids, and thus this makes a poor synapomorphy. The known chem- Haplosclerida into suborders Haplosclerina and Petrosina, as pro-
istry so far provides insufficient evidence that typical Haplosclerina posed here, has support from a formal character analysis even
compounds may exist (Van Soest & Braekman, 1999). though morphological apomorphies are vague.
The anisotropic skeleton is shared with closely related
Spongillina and several more distantly related sponge groups in Previous reviews
Poecilosclerida (e.g., Esperiopsidae, Isodictyidae), and a few
Petros ina species appear to have an underlying anisotropic structure Bergquist, 1980a; Van Soest, 1980; Desqueyroux-Faundez,
(e.g., Neopetrosia subtriangularis). It seems likely that this type 1984; De Weerdt, 1985, 1989; Fromont, 1993; Fromont et aI., 1994.

KEY TO FAMILIES

(1) Ectosomal skeleton a unispicular reticulation (sometimes absent); choanosomal skeleton with uni-, pauci- or multispicular primary
fibres or tracts and unispicular interconnecting fibres or tracts ... ................. ........... .... ........... ................ ........... ..... ...... ...... Cbalinidae
Ectosomal skeleton a pauci- or multispicular reticulation, or double-meshed with meshes formed by pauci- or multi spicular fibres or
tracts subdivided by unispicular fibres or lines; choanosomal skeleton always with multispicular primary fibres or tracts .............. 2
(2) Ectosomal skeleton a neat tangential two-dimensional reticulation of paucispicular or multi spicular fibres or tracts, usually double- or
triple-meshed; choanosomal skeleton usually a rectangularly meshed reticulation, usually with a core of spicules encased in ample
spongin ................. ............ ....... .... ...... ........... .......... ................. ......... ....... .......... ........... ...................... ........... ........ ..... Callyspongiidae
Ectosomal skeleton a three-dimensional, often somewhat irregular reticulation of pauci- or multi spicular fibres or tracts; choanosomal
skeleton anisotropic, but not neatly rectangularly meshed ................................................................................................. Nipbatidae
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademicIPlenum Publishers, New York, 2002

Family Callyspongiidae de Laubenfels, 1936

R. Desqueyroux-Faundez1 & c. Valentine2

1 Museum d'histoire Naturelle, P.O. Box 6434, 1211 Geneva 6, Switzerland. (ruth.faundez@mhn.ville-ge.ch)
2 The Natural History Museum, Cromwell Road, London SW7 5BD UK. (c.valentine@nhm.ac.uk)

Callyspongiidae de Laubenfels (Demospongiae, Haplosclerida) contains four valid genera, four subgenera and a large number of nominal
species with a worldwide distribution in tropical, temperate and cold waters. Species live principally in shallow waters. Major taxonomic
characters for differentiating genera are type of ectosomal mesh, morphology of subectosomal fibres, and presence, absence and type of
choanosomal fibres.
Keywords: Porifera; Haplosclerida; Callyspongiidae; Callyspongia; Callyspongia (Callyspongia); Callyspongia (Cladochalina);
Callyspongia (Euplacella); CaUyspongia (Toxochalina); Arenosclera; Dactylia; Siphonochalina.

DEFINITION, DIAGNOSIS, SCOPE History and biology

Synonymy A number of early names for several taxa have recently been
discovered (J.N.A. Hooper & R.W.M. Van Soest, in litteris) which
[Cavochalinidae] Carter, 1875c (nomen nudum). supposedly have priority. [Cavochalinidae] Carter, l875c is a senior
[Tubulodigitidae] Carter, 1875c (nomen nudum). Callyspongiidae synonym of Callyspongiidae. However, Carter did not nominate a
de Laubenfels, 1936a: 55. type species, instead it was 'named after a typical genus, but where
no typical species has yet been described, was named adjectively
Definition after its most striking characters'. This name is unavailable (Article
11.7.1.1, ICZN; Anon., 1999), in lacking a valid generic name
Haplosclerida with two dimensional ectosomal skeleton of when first published. Similarly, Carter (1875c: 141) erected a
primary, secondary and sometimes tertiary spongin fibres, fibres suprageneric taxon [Tubulodigitata], providing a short diagnosis
are cored by oxeas or strongyles, occasionally uncored or cored but again without mentioning a type genus. This suprageneric group
with foreign material. could also be interpreted as an available family taxon,
'Tubulodigitidae', although the genus Tubulodigitus was erected for
Diagnosis T. communis several years later by Carter (188lb: 367), and hence
these taxa of the suprageneric level were both nomina nuda (erected
Encrusting, massive, vase-shaped, tubular, lamellate, fan- without naming a type genus or species; Article 11.7.1.1, ICZN).
shaped or branching growth form. Sponge texture smooth to firm, De Laubenfels (1936a) erected Callyspongiidae to include
resilient. Surface smooth or characteristically sculptured into 'Haplosclerida possessing a characteristic surface reticulation of
conules, ridges or spines and fibre-reticulation tangential to the smaller fibres delimiting regular smaller meshes, with minute
surface. Ectosomal skeleton is a two dimensional tangential reticu- oxeas as the rule and simple microscleres toxas, present'. For
lation of primary, secondary and sometimes tertiary spongin fibres, de Laubenfels (1936a: 55) this family was differentiated from
fibres are cored by oxeas or strongyles, occasionally uncored or Haliclonidae ( = Chalinidae), and closer to his concept of
cored with foreign material. Choanosomal skeleton forms a net- Poecilosclerida than to Haplosclerida. However, the absence of spe-
work of primary principal or longitudinal, aspicular, pauci- to mul- cial spicules localised to any particular region of the sponge induced
tispicular spongin fibres, and secondary or connecting spongin de Laubenfels to leave Callyspongiidae in the Haplosclerida. The
fibres, uni-, bi- or aspicular. Choanosomal mesh typically regular presence of a secondary ectosomal reticulation composed of smaller
or rectangular with some exceptions, e.g., subgenus Cladochalina fibres with smaller meshes is good reason to consider the two fami-
with an irregular confused choanosomal mesh. Spongin is typically lies to be taxonomically different. For Wiedenmayer (1977b: 78) the
abundant. Megascleres: oxeas to strongyles in variable number, concept of continuous tangential reticulation or distinct ectosomal
generally small sizes, may be vestigial, completely absent or skeleton was apparently unsuitable as a diagnostic character, and
replaced by sand grains and/or foreign material. Microscleres: if having considered de Laubenfels' restriction of the family to the
present, toxas. type species he abandoned its use. Van Soest (1980) subsequently
redefined and resurrected Callyspongiidae based primarily on West
Scope Indian species. Bergquist & Warne (1980) revised the family and
described New Zealand species in three genera: Caliyspongia,
TWenty-three nominal genera and many hundreds of species are Chalinopsilla and Dactylia. Their interpretation was based strictly
included in this family worldwide, of which the following are consid- on de Laubenfels' (1936a) diagnosis ofthe family, and they insisted
ered valid: Arenosclera, Callyspongia, Callyspongia (Callyspongia), on the importance of surface reticulation as a distinguishing charac-
Callyspongia (Cladochalina), Callyspongia (Euplacella), ter to discriminate between genera and species of Callyspongii-
CaUyspongia (Toxochalina), Dactylia and Siphonochalina. dae. Desqueyroux-Faundez (1984) described New Caledonian

835
836 Porifera· Demospongiae • Haplosclerida • Haplosclerina • Callyspongiidae

Haplosclerida, including Callyspongiidae, 'introducing total con- Distribution

tent of silica in the skeleton' as a new diagnostic character, includ-
ing in that study the genera Callyspongia, Toxochalina (here Species have been collected from most tropical, temperate
considered as a subgenus of Callyspongia) and Arenosclera, with and cold waters. They are commonly represented along the East
silica content varying from 1-30% of dry weight. Wiedenmayer Pacific region from the Californian, Chile-Peru and Magellan
(1989) subsequently accepted Callyspongiidae, providing an regions; East South America, Brasilian and West Indies regions,
expanded synonymy of Callyspongia to include Spinosella (based West and South Africa and Mediterranean Atlantic region;
on features of the ectosomal and choanosomal skeleton, anatomy of Indo-Malaysian, Japan-China; South and North East and West
fibres, spicules and morphological variations). Fromont (1993) pro- Australian and New Zealand regions. No species have yet been
vided a diagnosis of Callyspongiidae modified from Wiedenmayer recorded from the Boreal East and West Pacific, Atlantic regions or
(1989), and described several species from the Great Barrier Reef. from Antarctica.

KEY TO GENERA AND SUBGENERA

(I) Single meshed ectosomal skeleton of foreign material, spicule tracts or single spicules, or a double meshed ectosomal reticulation of
fibres or tracts ..................................................................................................................................................................................... 2
Three sizes of regular ectosomal mesh ................................................................................................................................................ 7
(2) One size of confused ectosomal mesh, with abundant foreign material .......................................................... ................................... 3
Single or double, triangular to polygonal ectosomal mesh, without foreign material......................................................................... 4
(3) Tangential ectosomal mesh with no proper fibres, only fragments of foreign debris and spicules. Choanosomal irregular, fragmentary
fibres with foreign material proper spicules or both .................................. .............. ......................................................... Arenosclera
Tangential ectosomal network with proper fine aspicular fibres and foreign material. Choanosomal, continuous, aspicular fibres, with
only foreign material ................................................................................................................................................................ Dactylia
(4) Ectosomal strong echinated layer with continuous, strong palisade of brushes of spicules, or free spicules,without peripheral
condensation ........... ........... .......... ...... ...... .... ........... ...... .......... ................. ................................ .... ............ ...................... ................ ...... 5
Ectosomal non-echinated layer, peripheral condensation present ....................................................................................................... 6
(5) Ectosomal mesh echinated by strong palisade of spicular brushes. Choanosomal multi spicular primary fibres well cored, narrow
spongin sheath. One single surface layer ................................................................................................ Callyspongia (Cavochalina)
Ectosomal mesh echinated by free spicules. Choanosomal paucispicular primary fibres, with large spongin sheath. Three
surface layers .............................................................................................................................................. Callyspongia (Euplacella)
(6) Ectosomal mesh single, small, triangular to rectangular, unispicular fibres or single spicules. Choanosomal primary longitudinal fibres
paucispicular, subectosomal ends abundantly ramified, horizontally orientated and divided by connecting fibres .... Siphonochalina
Ectosomal mesh double, large, rounded to polygonal, unispicular fibres. Choanosomal primary longitudinal multispicular fibres,
parallel, secondary fibres subdividing subectosmal meshes parallel to the principal fibres ................... Callyspongia (Callyspongia)
(7) Surface strongly conulose. Choanosomal multispicular primary fibres compact, abundantly divided, non-fasciculated, abundantly
ramified triangular to irregular mesh. Scarce spongin. Microscleres toxas ............................................. Callyspongia (Toxochalina)
Surface conulose to finely conulose or spiny. Choanosomal primary longitudinal fasciculated fibres, abundantly ramified, tertiary
network always present. spongin variable amount, always visible. Microscleres absent ...................... Callyspongia (Cladochalina)

CALLYSPONGIA DUCHASSAING & MICHELOTTI, 1864 Definition

Synonymy Callyspongiidae with a regular ectosomal tangential reticula-

tion of primary, secondary and sometimes tertiary spiculo-fibres,
Callyspongia Duchassaing & Michelotti, 1864: 57. [Tuba] ectosomal morphology: one single size or three sizes of rounded to
Duchassaing & Michelotti, 1864: 44 (preocc. by Tuba Renier, 1804, irregular, or triangular to rectangular ectosomal mesh. Spongin
Polychaeta). Dactylochalina Lendenfeld, 1886c: 570. Ceraochalina abundant. Microscleres toxas may be present (e.g., Callyspongia
Lendenfeld, 1887c: 740, 778. Chalinella Lendenfeld, 1887c: 770, (Toxochalina) multiformis Pulitzer-Finali, 1986).
pI. 27, fig. 20. Chalinopora Lendenfeld, 1887c: 764. Euchalina
Lendenfeld, 1887c: 742, 816. Patuloscula Carter, 1882a: 365. Diagnosis
Cavochalina Carter, 1885c: 287; Topsent, 1932: 68. Placochalina
Lendenfeld, 1887c: 741, 790. Cladochalina Schmidt, 1870: 35. Varying greatly, from massive to ramose, lamellate, folia-
Spinosella Vosmaer, 1887: 342. Euplacella Lendenfeld, 1887c: 741, ceous to infundibuliform, tubular or lobate, repent or erect growth
789. Toxochalina Ridley, 1884a: 402. Platychalina sensu forms. Ectosomal skeleton a tangential network formed by second-
Wiedenrnayer in Hooper & Wiedenrnayer, 1994: 87 (Not ary and sometimes finer tertiary fibres (triple mesh ectosomal
Platychalina Ehlers, 1870: 21, 30). layer) or less ramified and with regular size of mesh (single mesh
ectosomallayer). Choanosomal skeleton a well-developed network
Type species of primary longitudinal fibres, fasciculated or non-fasciculated,
spongin sheath always present. Primary fibres ramified to form
Callyspongia fallax Duchassaing & Michelotti, 1864 (by secondary and tertiary fine fibres and tertiary choanosomal net-
subsequent designation; Burton, 1934a). work always present; or non-ramified, parallel and connected by
Porifera· Demospongiae • Haplosclerida • Haplosclerina • Callyspongiidae 837

short, parallel, non-ramified, secondary fibres, there may be several different ways: (a) by the presence of more than one surface
peripheral condensation of the choanosomal skeleton. layer of the ectosome (e.g., in Euplacella we observed three suc-
cessive layers, each growing over a previous surface layer 'growth
Previous reviews lines') (Fig. 5C); (b) by subectosomal smaller mesh obtained by
vertical subdivision of primary fibres and modification of subecto-
Burton (1934a: 539), Wiedenmayer (1977b: 90), Bergquist & somal connective fibres (e.g., Ceraochalina, Fig. lA-D); or
Warne (1980: 24), Van Soest (1980: 46), Desqueyroux-Fatindez (c) simply by transversal condensation of the subectosomal second-
(1984: 785), Wiedenmayer (1989: 103), Hooper & Wiedenmayer ary fibres, occurring between terminal ends of parallel primary
(1994: 87). fibres (in Siphonochalina, Fig. 9D). In C. fallax (D. & M., 1864)
peripheral condensation is variably present, whereas it is always
Remarks present in C. typica (Lendenfeld, 1887c). The latter is the type
species of Ceraochalina (by indication; Burton's (l934a: 540) sub-
Wiedenmayer (1977b) considered Callyspongia and Spinosella sequent designation of Cladochalina nuda Ridley, 1884a as the
as two distinct genera, with their principal differences being the type species is invalid), which is included in this work as synonym
absence of fasciculation and a much denser surface fibro-reticulation of Callyspongia. Further comment is required on the generic syn-
or peripheral layer, often discontinuous and not two-dimensional, but onymy of Callyspongia. Chalinella, with type species Chalinella
rather a condensation of the main skeleton in Callyspongia. In rnacropora (by subsequent designation, Burton, 1927: 296), is a
Spinosella (pro Cladochalina), Wiedenmayer (1977b) emphasized subjective synonym of Ceraochalina (taxonomic decision this
the special two-dimensional surface network, smaller angular mesh, work) and a synonym of Callyspongia (taxonomic decision
fasciculated choanosomal fibres and the presence of conules, spines Wiedenmayer in Hooper & Wiedenmayer, 1994). Chalinopora is
and laciniae to a variable degree. Wiedenmayer's (l977b) concept of included in synonymy with Callyspongia following Wiedenmayer
Callyspongia included: [in part] Siphonochalina, Sclerochalina, [?] (1989: 104) (type species Chalinopora typica Lendenfeld, 1887c:
Tubulodigitus, Patuloscula and Polysiphonia. Spinosella sensu 765, by indication; Burton's (1927: 293) subsequent designation of
Wiedenmayer (1977b) included as synonyms, Tuba, Cladochalina, Acervochalina clavifonnis Carter, 1886 as the type species is
but also Callyspongia (sensu Burton, 1932b) and [in part] invalid, and Burton's (1934a: 535) record of Chalinopora
Siphonochalina, Dactylochalina, Ceraochalina, Dasychalina, clavifonnis refers to a typical Adocia). Euchalina (with type
Siphonella (fide Burton, 1934a) and Euchalina (fide Burton, 1934a). species Euchalina typica Lendenfeld, 1887c, by indication), is a
Bergquist & Warne's (1980) concept of Callyspongia included subjective synonym of Chalinopora (taxonomic decision this
Siphonochalina, Toxochalina, Ceraochalina, Dactylochalina and work), and a synonym of Callyspongia (taxonomic decision
Spinosella. Van Soest (1980) used these names as subgenera and Wiedenmayer in Hooper & Wiedenmayer, 1994), with Burton's
concluded that Callyspongia exhibited a high proportion of spongin, (1934a: 540) subsequent designation of Euchalina rigida as the
variable presence of peripheral condensation (e.g., in C. fallax and type species being invalid (Wiedenmayer, 1989: 104).
C. eschrichti Duchassaing & Michelotti, 1864), presence or absence Wiedenmayer, in Hooper & Wiedenmayer, (1994), included
of fasciculated primary fibres (e.g., C. vaginalis (Lamarck) and Platychalina Ehlers, 1870 (type species Platychalina foliacea
C. plicifera (Lamarck». Van Soest (1980: 55) placed emphasis on the (Esper, 1797) Ehlers, 1870, by monotypy) in synonymy with
rectangular choanosomal mesh in CaUyspongia, as opposed to Callyspongia, whereas other authors have produced contradictory
Spinosella (pro Cladochalina) which has an irregular, confused decisions. The first author to 'tackle this mess', Ridley (1884a),
choanosomal mesh. We observed that this character is largely deter- doubted the chalinid nature of the genus at all. He studied material
mined by the type of fibres present: non-fasciculated or fasciculated. The from H.M.S. 'Alert' which, in his opinion, strongly agreed with
secondary, discriminatory taxonomic character in subgenera of Esper's figure and description with the exception of small morpho-
CaUyspongia is a special dermal skeleton with thinner secondary or even logical details described by Ehlers, 1870 (such as the single
tertiary fibres, associated or not with the presence of ectosomal conules occurrence of isolated spicules, or different disposition of 'pores')
of different size. Over the years the taxon Callyspongia has become Ridley identified his specimen as Euspongia foliacea. Conversely,
quite unmanageable given the high diversity of species and the subtle several other authors treated P. foliacea in very different ways.
characters differentiating it from other allied genera (or subgenera). None of these subsequent authors, however, have studied the type
The classification proposed here is largely a 'convenient one', material given that it is lost (Wiedenmayer, 1989: 104): "not repre-
with five of the fifteen generic names included in synonymy with sented in the Australian region" (Lendenfeld, 1887c: 792); syn-
Callyspongia by Wiedenmayer (in Hooper & Wiedenmayer, 1994) onym of lsodictya Bowerbank (de Laubenfels, 1936a: 54);
treated here as subgenera: Euplacella; Toxochalina; Cladochalina Platychalina foliacea, unnecessary designation of type species,
(including Spinosella); Cavochalina (including Placochalina). lipochelous form of Homeodictya elastica, and a pre-emptive of
This present classification is based on an earlier morphological Homoeodictya Ehlers, 1870 (Burton, 1936: 143); Platychalina,
analysis of Callyspongiidae and other haplosclerids (Desqueyroux- might replace Spinosella as subgenus of Callyspongia if the
Fatindez, 1999) using structural characters more so than previous specific identity of S. vaginalis Lamarck and S. foliacea could
classifications. One of the difficulties encountered in the recogni- be established (Van Soest, 1980: 60). But as the debate cannot be
tion of reliable characters to separate taxa of Callyspongiidae is the resolved without re-examination of Ehlers' missing type, if it still
irregular occurrence of microscleres in some genera, where toxas exists, in this work, we follow Wiedenmayer in Hooper &
are sometimes present (Toxochalina), not present (Callyspongia Wiedenmayer (1994) , and we consider here, Platychalina as ajun-
s.s.), or only fine skeletal details appear to differentiate taxa at even ior synonym of Callyspongia.
the species levels. Another taxon that is potentially associated with Callyspongia is
A similar case is observed for other skeletal characters. For [Cacochalina] Schmidt, 1868, but this is a nomen nudum, following
example, peripheral condensation of the fibres may be produced in ICZN Article 5.1. as Schmidt (1868) did not name a type species.
838 Porifera· Demospongiae • Haplosclerida • Haplosclerina • CaUyspongiidae

Fig. 1. Callyspongia (Callyspongia) typica (Lendenfeld. 1887c as Ceraochalina) (holotype BMNH 1886.8.27.439). A, holotype. B. enlarged view of the
surface. C, ectosomal skeleton. one size of mesh, hispid surface, small conules (scale 100 ILm). D, ectosomal and choanosomal skeleton, longitudinal view,
subectosomal smaller mesh obtained by vertical subdivision of primary fibres and modification of subectosomal connective fibres (peripheral condensation).

However, Article 11.4.1. also specifies that a genus group name with- Definition
out associated nominal species could be accepted consistent with the
Principle of Binomial Nomenclature in the absence of evidence to the Callyspongia with one size of ectosomal mesh. Single ectoso-
contrary. Cacochalina Schmidt, 1870 was also declared unrecognis- mal non-hispid, layer. Choanosomal fibres multispicular, non-
able (Burton, 1927; de Laubenfels, 1936a), because the two species fasciculated, spongin sheath present. Well defined, connecting
assigned by Schmidt (1870) (c. subtilis, Florida and C. rubiginosa, fibres, rectangular mesh without free spicules.
Antilles), are unrecognizably described. Van Soest (in litteris) studied
slides of Schmidt in London (BMNH 1870.5.3.135 Florida, 9 fath- Diagnosis
oms) and in Copenhagen (ZMUC), and in his opinion these slides
were so poor that no genus characters can be extracted from them. Lobate repent encrusting to massive or short anastomosed
Consequently, we propose here to declare [Cacochalina] Schmidt a upright tubes with apical oscule. Surface smooth. Single ectosomal
nomen nudum (taxonomic decision, this work). non-hispid, layer and regular size of single, rounded to polygonal
mesh. Choanosomal a well-developed network with rectangular
empty mesh, primary choanosomal fibres multispicular, non-
SUBGENUS CALLYSPONGIA DUCHASSAING & fasciculated, non ramified, spongin sheath well-defined. There
MICHELOTTI, 1864 may be peripheral condensation of the choanosomal skeleton.
Microscleres, toxas may be present.
Synonymy
Distribution
Callyspongia Duchassaing & Michelotti, 1864: 57.
Ceraochalina Lendenfeld, 1887c: 740, 778. Chalinella Lendenfeld, Cosmopolitan (Hooper & Wiedenmayer, 1994).
1887c: 770, pI. 27, fig. 20. Chalinopora Lendenfeld, 1887c: 764.
Euchalina Lendenfeld, 1887c: 742, 816. Anatoxius Lendenfeld, Previous reviews
1887c: 798. Patuloscula Carter, 1882a: 365.
Burton (1934a: 539), Wiedenmayer (1977b: 90), Van Soest
(1980: 46).
Type species
Description of type species
Callyspongiafallax Duchassaing & Michelotti, 1864 (subse-
quent designation; Burton, 1934a: 539). Callyspongiafallax Duchassaing & Michelotti, 1864 (Fig. 2).
Porifera· Demospongiae • Haplosclerida • Haplosclerina • Callyspongiidae 839

Fig. 2. Callyspongia (Callyspongia) fallax Duchassaing & Michelotti, 1864 (Lectotype: MT Por 84). A, lectotype. Ectosomal skeleton a typical
tangential network of rectangular to polygonal mesh, formed around the apex of a primary longitudinal fibre that ramifies on surface (scale 200 /-Lm).
B, choanosomal skeleton a regular network of rectangular to roundish mesh, formed by primary, longitudinal paucispicular, connected at right angle by
secondary unispicular fibres. C, Patuloscula procumbens Carter, 1882a, ectosome uniform, membranous layer with one size of rounded mesh, limited
by one single spicule by side (scale 500 /-Lm). D, C. fallax, primary paucispicular and connecting unispicular fibres, regular choanosomal skeleton.

Synonymy. Callyspongia fallax Duchassaing & Michelotti, Sometimes primary fibres are fused andlor subdivided, but not fas-
1864: 57; Wiedenmayer, 1977b: 92 (in part); Van Soest, 1980: 47. ciculated. If fused, two central axes of coring spicules are visible.
Material examined. Lectotype (designated by Van Soest, Van Soest (1980) described subdermal peripheral condensation of
1980: 47): MT Por 84 (fragments BMNH 1928.11.12.5, MNHN choanosomal skeleton by intercalated vertical secondary fibres,
NBE 1067, MHNG 22213) - St Thomas. Comparative material. which we did not observe. Spicules: strongyloxeas, 38-(46)--80 f..Lm
Lectotype of CaUyspongia eschrichtii Duchassaing & Michelotti, length and 0.5-2 f..Lm in diameter.
1864: 56: ZMA POR 620 - St Thomas. Paralectotypes: MZUT Distribution. Caribbean: St Thomas, Cura9ao, Bonaire,
POR 44, 45, ZMA POR 623, (fragment MHNG 14839) - Viecques, Florida, Grenada, Tortola, Bahamas. Occurring only on
St Thomas. Holotype of Dactylochalina australis Lendenfeld: reefs. From secondary source: Van Soest (1980).
BMNH 1886.8.27.466a (slide); BMNH 1886.8.27.469, wet sub-
sample. Dactylochalina reticulata Lendenfeld, 1886c: BMNH Remarks
1886.8.27.467a - slide, W coast of Australia.
Description. Repent mass of anastomosing lobes, measuring Van Soest (1980) considered C. eschrichti an available species
12-14cm in length by 2-3cm in diameter, with terminal oscule, and different from C. fallax, contrary to Burton (1934a) and
10--12mm in diameter, and a short aquiferous canal. Surface Wiedenmayer (1977b) who both considered C. eschrichti to be a
smooth. Consistency firm, tough to barely compressible in dry state. synonym of C.fallax or a growth form of Callyspongia (Spinosella
Colour yellowish, amber-like to beige, with purple spots. In dry pro Cladochalina) vaginalis. We support Van Soest's contention
state, light brown. Ectosomal skeleton a typical tangential network that C. eschrichti is a valid species but closest to C. fallax because
of rectangular to polygonal mesh, formed around the apex of a pri- of its typical Callyspongia (Callyspongia) type of skeleton, but
mary longitudinal fibre that ramifies on the surface. Mesh delimited without peripheral condensation. They also have different habits
by primary fibres, 107-(193)-350 f..Lm large, mesh delimited by and spicule micrometries.
finer fibres, 50--74 f..Lm, in diameter. Primary fibres 30--50 f..Lm, in Subgenus CaUyspongia differs from other subgenera by the
diameter. Secondary and tertiary fibres, 7-26 f..Lm and 4-6 f..Lm in absence of choanosomal fibro-fascicles and the presence of con-
diameter respectively. Choanosomal skeleton a regular network of stantly regular, rectangular mesh. Ectosomal network is formed by
rectangular to roundish mesh 246-(297)--360 f..Lm large, formed by ramified ends of primary longitudinal fibres, spread on the surface
primary, longitudinal paucispicular fibres (5-7 divergent spicules and delimiting regular triangular to rounded or polygonal ectoso-
in a central row), 53-(78)--114 f..Lm in diameter, connected at right mal mesh. Burton (1934a) considered Patuloscula Carter, 1882a as
angle by secondary unispicular fibres, 12-(23)--33 f..Lm in diameter. synonym of Callyspongia, since P. procumbens its type species
840 Porifera· Demospongiae • Haplosclerida • Haplosclerioa • Callyspongiidae

was synonym of C. fallax (Fig. 2A-D). De Laubenfels (1936a) rugous, inner surface smooth, microhispid, concentrically lined,
remarked that Patuloscula was 'very similar by its external form, strongly porous, with regularly distributed oscula. Consistency
with Callyspongia, except by the possession of toxiform spicules' . tough, rubbery. Skeleton: ectosomal small mesh echinated by erect
Several syntypes of P. procumbens studied by Carter (1882a) spicule brushes and unispicular fibres. Choanosomal (Fig. 3B)
and Dendy (189Oa) illustrate the large morphological variability of regular network with longitudinal parallel paucispicular fibres and
the type species of Patuloscula, and consequently the difficulties of strong central core and spongin sheath. Connective secondary
discriminating between this genus and Callyspongia, to the extent fibres, short, non-ramified, paucispicular. Spicules: small oxeas.
that Wiedenmayer (1977b) also included P. procumbens in syn-
onymy with C. fallax. Dendy (1890a) provided details of the ecto- Previous reviews
somal skeleton of Patuloscula (as Siphonochalina) procumbens,
describing a polygonal reticulation of mesh delimited by sparse Topsent (1932: 68; in part, as Cribrochalina bilamellata),
uni-serially distributed spicules. The structure of the ectosome we Wiedenmayer (1989: 106).
observed in the type specimens of P. procumbens, surface mem-
brane pierced by one size of ectosomal mesh delimited by unispic- Description of type species
ular fibres is, in our opinion, the only difference between
P. procumbens and C. fallax and following Wiedenmayer (1977b) Callyspongia (Cavochalina) bilamellata (Carter, 1885d)
and Van Soest (in litteris), in this work we consider Patuloscula (Fig. 3).
a junior subjective synonym of Callyspongia (Callyspongia). Synonymy. Spongia bilamellata Lamarck, 1813-14: 484;
Dactylochalina Lendenfeld, 1886c (type species Cribrochalina bilamellata Topsent, 1932: 68; Callyspongia bil-
Dactylochalina australis Lendenfeld, 1887c, by subsequent desig- amellata Wiedenmayer, 1977b: 118; 1989: 106; Placochalina
nation; Burton, 1934a), is considered here a junior synonym of pedunculata var. dura Lendenfeld, 1887c: 791.
Callyspongia (Callyspongia). The type of skeleton in both genera Material examined. Holotype: MNHN LBIM DT 602 -
is similarly formed by fibres with abundant spongin with a central King I., Bass Strait. Holotype of Cavochalina bilamellata Carter,
row of spicules, unispicular in secondary fibres and a paucispicular 1885d (Fig. 3D): BMNH 1886.12.15.182 - S coast Australia.
in primary fibres. Callyspongia and Dactylochalina are different Comparative material. Holotype of Placochalina: BMNH
only by their spicule sizes, which in any case are vestigial 1886.12.15.182 - Port Phillip Heads, S coast Australia. Syntype
as Lendenfeld (1886c) mentions. Skeletal meshes are empty of of Cavochalina bilamellata (Carter's specimen): BMNH
fibres and foreign material is absent. Spicule micrometries in 1886.8.27.561 - Port Jackson. Syntypes of Placochalina pedun-
D. cylindrica Lendenfeld, 1886c, are: oxeas 48-61 X 4.8-6.4 J.Lm. cula var. dura Lendenfeld, 1887c: BMNH 1886.8.27.563, 564,
Longitudinal primary fibres are 92 J.Lm in diameter, connecting sec- 565 - Torres Straits, Port Jackson and Dlawarra. Holotype of
ondary fibres 42 J.Lm in diameter, meshes of the surface are 60 J.Lm Placochalina peduncula var. poculum Lendenfeld, 1887c: BMNH
diameter and choanosomal meshes are 170-176 J.Lm diameter. 1886.8.27.562 - South coast of Gulf St Vincent. Holotype of
Anatoxius Lendenfeld, 1887c (type species Phylosiphonia Placochalina peduncula var. mollis Lendenfeld, 1887c: BMNH
stalagmites Lendenfeld, 1887c, here designated, syntypeAM Z618 1886.8.27.562 - South coast of Gulf StVincent.
not seen), was erected as a subgenus of Phylosiphonia, which is a Description. Pedicellate sponge, 28 cm high, bilamellate,
Chalinula (Chalinidae) by virtue of Burton's type species assign- 15-16cm in length, doubled up like a bivalve shell with a long,
ment, but most if not all Phylosiphonia (Anatoxius) species are hard stem, ending in a root-like expansion, vasiform, compressed,
Callyspongia, including the type species here designated. infundibular toward terminal part. Consistency leathery. Colour
pale pinkish brown. Outer surface irregular, corrugated and wrin-
kled, reticulated with numerous microconules. Internal surface
SUBGENUS CAVOCHALINA CARTER, 1885 even, smooth, microhispid, concentrically lined, small oscula regu-
larly scattered over internal surface. Skeleton: Ectosome with
Synonymy numerous erect spicule brushes from terminal part of strongly rami-
fied primary fibres, one size of rounded mesh. Choanosome with
Cavochalina Carter, 1885d: 287; Topsent, 1932: 68. regularly parallel-branched primary longitudinal fibres, 35-140 J.Lm
Placochalina Lendenfeld, 1887c: 741, 790. in diameter. Secondary fibres, short connected, some of them
slightly branched, similar in diameter to primary fibres. Rounded
Type species mesh. Spongin abundant, except on surface where spicules are
more important, ends of fibres ramified to form bundles with
Cavochalina bilamellata (Lamarck, 1814) Carter, 1885d (by spicule brushes, resembling ectosomal skeleton of Cribrochalina.
subsequent designation; Topsent, 1932). Spicules: fusiform oxea, slightly bent, 37-43-51 X 2-3 J.Lm, small
free vestigial oxeas and strongyles.
Definition Remarks. Topsent (1932) affirmed that C. bilamellata is a
Haplosclerida but close to Cribrochalina given the possession of an
Callyspongia with two different types of surfaces. External ectosomallayer in both. Wiedenmayer (1977b) remarked that this
surface very rough (Fig. 3A). Internal surface (Fig. 3C), smooth, species was much like Cribrochalina vasculum from the West
concentrically lined, extremely porous. Ectosomal tangential net- Indies, a species he also associated with Lendenfeld's varieties of
work with strong fibres and rounded mesh of one size. Placochalina, with spicule-dimensions being the only significant
difference between them. We have also remarked on similar charac-
Diagnosis ters between Cribrochalina infundibulum (type species of
Cribrochalina) and C. bilamellata, but in addition to differences in
Stalked, bilaterally compressed, bilamellate to funneliform spicule micrometries they have a different type of fibres and a dif-
growth form. Outer surface strongly rough, coarse, reticulate, ferent type of ectosomallayer. Cribrochalina infundibulum has larger
Porifera· Demospongiae • Haplosclerida • Haplosclerina • CaUyspongiidae 841

Fig.3. Callyspongia (Cavochalina) bilamellata Carter, 1885 (holotype MNHN LBIM DT602). A, holotype, external surface view. B, choanosomal skele-
ton, regular network with longitudinal parallel paucispicular fibres and strong central core, large spongin sheath. C, internal surface of the holotype.
D, Cavochalina bilamellata, Carter's specimen BMNH 1886.12.15.182.

spicules (strongylote oxeas: 108-(134)-164 X 3-4 /-Lm) and stronger, Lendenfeld; Cribrochalina infundibulum Schmidt as P. pedunculata
compact multispicular primary fibres (160-180/-Lm in diameter), poculum; Cribrochalina variabilis vars. crassa Vosmaer and salp-
arising to the surlace to form strong brushes of spicules in a palisade ingoides Vosmaer as P. variabilis vars. variabilis and salpingoides;
or crust. Isodictya infundibuliformis Bowerbank as P. infundibuliformis).
Cavochalina was introduced by Carter (I 882a: 277) for S. bil- We confirm here, from re-examination of type material, that
amellata (Lamarck) but without nominating it as the type species. Placochalina is a junior objective synonym of Cavochalina.
We consider Cavochalina to be an available genus, based on
Article 11.5 of the ICZN 'by bibliographic reference' through
Topsent's (1932) revision of Lamarck's collection in the MNHN SUBGENUS CLADOCHAUNA SCHMIDT, 1870
Paris. Topsent (1932) confirmed Spongia bilamellata Lamarck,
1814 typical form, as the species described by Carter as Synonymy
Cavochalina bilamellata 'a typical chalinine' that he named
Cribrochalina bilamellata. Topsent (1932) therefore considered Cladochalina Schmidt, 1870: 35. Spinosella Vosmaer,
Cavochalina and Placochalina to be objective synonyms (both 1887: 342.
genera share the same type species), and closer to Cribrochalina
(Niphatidae) (both sharing a similar type of ectosomallayer). Only Type species
two species of Cavochalina have been described, both by Carter
(l882a; 1885d), and both from the S coast of Australia. Tuba armigera Duchassaing & Michelotti, 1864 (by subsequent
Placochalina, a genus of doubtful status, with the type species designation; Burton, 1927).
Cavochalina bilamellata (Lamarck) Carter, 1885d (by subsequent
designation de Laubenfels, 1936a) was mixed up in Cavochalina Definition
bilamellata and Cribrochalina Schmidt, 1870 by Lendenfeld
(1887c). Lendenfeld (1887c: 790) included several species of Carter, Callyspongia with ectosomal network with three sizes of mesh
Schmidt, Vosmaer and Bowerbank in his new genus Placochalina around terminal ends of primary fibres largely spread on the sur-
(viz., Cavochalina bilamellata Carter, as Placochalina pedunculata face. Multispicular primary fibres well defined, always fasciculated,
var. dura; Cavochalina digitata var. arenosa Carter, 1882a, as and ramified to form a choanosomal tertiary network of fine fibres
P. arenosa; Cribrochalina cretacea Schmidt as P. cretacea always present. Spongin sheath clearly visible. Conulose surface.
842 Porifera· Demospongiae • HapJoscierida • HapJoscierina • CaUyspongiidae

Fig.4. A-B, Callyspongia (Cladochalina) armigera (Duchassaing & Michelotti, 1864) (lectotype MUS. TURIN POR 75). A, lectotype, ectosomal skele-
ton network, strictly tangential, primary-secondary and tertiary fibres in one plan, determining three sizes of irregular mesh. B, longitudinal view of conule,
fasciculated primary fibre, apex at bottom of the figure. C-D, Spinosella sororia (Duchassaing & Michelotti, 1864) (lecto- and paralectotypes MT Por 126,
118). C, paralectotype, choanosomal skeleton strong, irregular, with primary longitudinal fasciculated fibres, abundantly ramified to form free secondary
and tertiary fibres with rounded to elongate always subdivided mesh. D, lectotype, longitudinal view of choanosomal skeleton, Spongin always present.
Spicules: oxeas to strongyloxeas well developed or vestigial.

Diagnosis Spinosella vaginalis forma armigera; Wiedenmayer, 1977b: 104;

Callyspongia (SpinoseUa) vaginalis forma armigera; Van Soest,
Branched, tubuliform or vase-like growth form, with tendency 1980: 60.
to form coalescent tubes or masses of branches, with a common Material examined. Lectotype (designated by Van Soest,
base, with large apical vents, or irregularly distributed oscula. 1980: 60): BMNH 1870.5.3.136, MT Por 75 (Schmidt's original
Surface strongly to finely conulose or spiny. Texture elastic, strong slides) - St Thomas (Fig. 6A, B). Paralectotype: ZMA POR 2407
or fragile and breakable, in dry state. Ectosoma1 skeleton network, (fragments. BMNH 1928.11.12.93, 94, MHNG INVE POR
strictly tangential, primary secondary and tertiary fibres in one 980.176) - Florida, Antilles.
plane, determining three sizes of irregular mesh (Fig. 4A). Several Description (emendedfrom Van Soest, 1980: 60). Ramose,
end branches of one primary longitudinal fibre form the skeletal branches 12 cm long by 1.5 cm in diameter, somewhat compressed.
axis of each surface conule or spine (Fig. 4B). Choanosomal skele- Oscula scattered over branches, 3-6 mm in diameter. Surface
ton strong, irregular, with primary longitudinal fasciculated fibres, conulose, smooth in between. Ectosomal skeleton, large mesh
abundantly ramified to form free secondary and tertiary fibres with 250 f.Lm in diameter, small mesh 60-100 f.Lm in diameter, second-
rounded to elongate, always subdivided mesh (Fig. 4C, D). Spongin ary fibres 8-12 f.Lm in diameter. Choanosomal skeleton: fibro-
always visible present in variable amounts. Spicules: oxeas to fascicles 200 f.Lm in diameter, primary, paucispicular fibres (1-4
strongyloxeas well developed or vestigial. spicules), 40-70 f.Lm in diameter; secondary paucispicular fibres
(1-3 spicules), 25-35 f.Lm in diameter, tertiary, aspicular to unispic-
Previous reviews ular, fibres, 6-15 f.Lm in diameter (Fig. 4D). Spicules: oxeas,
48-(88)-99 X 1.5-(3)-4.5 f.Lm in diameter.
Burton (1934a: 540), Wiedenmayer (1977b: 100; 1989: 104, Remarks. Further comment is required on the synonymy
105), Van Soest (1980: 110). outlined above. [Tuba] Duchassaing & Michelotti, 1864 (type
species T. sororia D. & M., by subsequent designation) (Burton,
Description of type species 1934a: 540), Topsent (1932: 75), is junior homonym of Tuba
Renier, 1864 (Polychaeta), [Tuba] Oken, 1815 (Nemertina), [Tuba]
Callyspongia (Cladochalina) armigera (Duchassaing & Fabricius, 1823 (Mollusca), [Tuba] Lea, 1833 (Mollusca) and
Michelotti, 1864) (Fig. 4). [Tuba] Barrande, 1848 (Mollusca). Vosmaer (1887) proposed
Synonymy. Tuba armigera Duchassaing & Michelotti, Spinosella as a replacement name (with type species Tuba sororia
1864: 48, pI. 8, fig. 3; Callyspongia armigera; Burton, 1934a: 540; D. & M., 1864, by original designation). Burton (1934a: 540) and
Porifera· Demospongiae • Haplosclerida • Haplosclerina • Callyspongiidae 843

Bergquist & Warne (1980: 24) considered it to be a synonym of Subectosomal region of upper surface exhibits three (in type speci-
Callyspongia; Dendy (1890a: 357) suggested it was close to men) (Fig. 5C) successive ectosomallayers or 'growth lines or lamel-
Siphonochalina; and Van Soest (1980: 56, 64) used Spinosella as a lae', Inner surface with single ectosomallayer. Choanosomal skeleton
subgenus, including redescription of the lectotypes of T. megas- (Fig. 5D) of parallel paucispicular primary longitudinal fibres, non-
toma, T. irregularis and T. sororia, synonymising them in ramified, strongly cored with large spongin sheath, rounded to rectan-
Callyspongia (Spinosella) vagina lis, therefore making it a subjec- gular large empty choanosomal mesh, and short, non-ramified
tive synonym of Callyspongia. connecting unispicular fibres. Spicules, small, slightly bent oxeas.
Burton (1927) designated T. armigera (Fig. 4A, B) the type
species of Cladochalina, differing from C. fallax in having three sizes Previous reviews
of mesh in the ectosomal skeleton, but he also concluded that this
difference was not significant at the generic level, and included Van Soest (1980: 110), Wiedenmayer (1989: 104, 105),
Cladochalina in synonymy with CaUyspongia. Burton (1934a) con- Wiedenmayer in Hooper & Wiedenmayer (1994: 87).
sidered that T. sororia, the type species of SpinoseUa (Fig. 4C, D),
was a synonym of C. faUax. Consequently, both Cladochalina and Description of type species
SpinoseUa became synonyms of CaUyspongia. The present study
confirms their synonymy from re-examination of their respective type Callyspongia (Euplacella) australis (Lendenfeld, 1887c)
species, T. armigera and T. sororia. Wiederunayer (1989: 103, 105) (Fig. 5).
proposed the use of the name Cladochalina to replace Spinosella as Synonymy. EuplaceUa australis Lendenfeld, 1887c: 741,789;
a subgenus of CaUyspongia based on seniority of the two taxa. Burton, 1934a: 597; Wiedenmyaer, 189: 106.
Cladochalina presents a skeleton similar to Toxochalina Material examined. Holotype (fragment): BMNH
Ridley, these genera are included here as subgenus of 1886.8.27.591 (dry) - Torres Straits. Comparative material. Slide
CaUyspongia. Both exhibit three sizes of ectosomal mesh, irregular of Lendenfeld as Euplacella bilamellata (Carter): BMNH
choanosomal mesh and abundantly ramified, confuse principal 1887.8.27.591a. Syntype of Euplacella frondosa (Lendenfeld,
fibres. Their principal difference being the presence of toxa in 1887c): BMNH 1886.8.27.424 - Port Jackson. Syntypes of
Toxochalina. This has been considered in contradictory ways: EuplaceUafrondosa (Lendenfeld, 1887c): BMNH 1886.8.27.453,
Burton (1934a) considered Desmacidon folioides had a skeleton 453a - Port Jackson. Syntype slide of Euplacella frondosa
like that of Cladochalina armigera with toxas for microscleres, (Lendenfeld, 1887c): BMNH 1954.2.12.88 - Port Jackson.
but toxas, could be also present in CaUyspongia (e.g., C. fallax), Holotype of Euplacella mollissima (Lendenfeld, 1887c): BMNH
therefore, Toxochalina was synonym of Callyspongia. 1886.8.27.423 (dry) - Westernport Bay, Victoria.
Description. Holotype of Euplacella australis (Fig. 5A-D)
Distribution is represented by a small flattened rectangular fragment of frond,
lamella or thin plate, with two different surfaces. The original
West Indies: St Thomas, Viecques (Duchassaing & Michelotti, description stated: 'A pedunculated plate, undulated and curved,
1864), Florida, Antilles, Jamaica (Lehnert & Van Soest, 1999a). diameter 150mm, thickness 3 mm, margin continuous.
Consistency strong and hard' (Lendenfeld, 1887c). Outer surface
finely hispid, with numerous, irregularly distributed oscules. Inner
SUBGENUS EUPLACELLA LENDENFELD, 1887 surface with one single surface layer. Consistency tough, hard, in
dry state. Ectosomal skeleton with one size of small rounded mesh,
Synonymy with erect brushes of spicules, present in the outer surface.
Choanosomal skeleton, with primary longitudinal parallel pau-
Euplacella Lendenfeld, 1887c: 741,789. cispicular fibres, 33-(55)-66 f-Lm in diameter with central core of
spicules and large spongin sheath, connected by uni- to trispicular,
Type species short, non-ramified fibres, 25-(33Hl f-Lm in diameter. Large
rounded to rectangular regular mesh 82-(120)-148 f-Lm in diame-
Euplacella australis Lendenfeld, 1887c: 782 (by subsequent ter, without or with a few free spicules. Mesh near the surface,
designation; Burton, 1934a: 597). 5-(6)-7 f-Lm in diameter. Longitudinal fibres of triple layers,
16--(22)-25 f-Lm in diameter, ending at the surface by perpendicular
Definition palisade of spicules delimiting ectosomal mesh Spicules: small
oxeas, 48-(56)-65 X 3-4 f-Lm, or even smaller (Lendenfeld, 1887c
Callyspongia with three distinct surface layers and one size of cites oxeas as 67 X 3.3 f-Lm).
close polygonal-triangular mesh delimited by echinated secondary Remarks. Burton (1934a) designated E. australis the type
fibres. Choanosomal parallel, strong longitudinal paucispicular fibres. species of Euplacella (making de Laubenfels' (1936a: 59) subse-
quent designation of E. mollisima Lendenfeld, 1887c, invalid), and
Diagnosis gave a diagnosis that emphasised the typical ectosomal network
characteristic of this genus, which he considered important enough
Thin lamellae or thin plates, foliate, with two different surface to warrant its recognition as a valid genus of Callyspongiidae.
layers: upper surface smooth, regular, with numerous, evenly distrib- Burton (1934a) also included Placochalina in synonomy with
uted oscules (Fig. 5A). Ectosomal skeleton a tangential layer of upper Euplacella, and Van Soest (1980: 110) redefined the genus based
surface with one size regular small mesh ends of primary fibres echi- on Pachychalina elongata Ridley & Dendy, 1886, including
nated by free spicules (Fig. 5B, D). Inner surface irregularly undu- Placochalina as a junior synonym of EuplaceUa, whereas we have
lated, ectosomallayer with small rounded mesh, without hispidation. demonstrated here that it is a synonym of Cavochalina Carter.
844 Porifera· Demospongiae • Haplosclerida • Haplosclerina • CaUyspongiidae

Fig. 5. Callyspongia (Euplacella) australis (Lendenfeld. 1887c) (holotype bmnh 1886.8.27.591). A, holotype, small flattened rectangular fragment of
lamella, with numerous small oscula irregularly distributed on the outer, finely hispid surface. B, ectosomal skeleton with one size of small rounded mesh,
underlined by abundant free spicules. Unispicular fibres, tangential view (scale 50 /Lm). C, subectosomal region as triple surface layer or 'growth lines or
lamellae'. D, echinated ends of primary fibres.

Wiedenmayer (1989: 104-5) incorrectly proposed that mesh. In Cavochalina bilamellata surface hispidation is also made
Ceraochalina Lendenfeld (type species C. typical was a senior of isolated spicule brushes, but they are more evident at the ends
synonym of Euplacella. Both are lamellate with a smooth, regular of the primary fibres, of the choanosomal skeleton. Primary fibres
porous surface, and numerous small oscula, but their external mor- in Cavochalina are abundantly cored, while secondary fibres are
phology appears to be the only common character whereas there sparsely cored by small oxeas, representing a CaUyspongia type
are many skeletal differences between the two. These differences of skeleton. In Cribrochalina (Niphatidae) surface hispidation is
are clear at the level of subectosomal and ectosomal regions. apparently similar, consisting of a strong continuous palisade of
Ceraochalina exhibits subectosomal peripheral condensation. The single oxeas, issuing from the surface ends of primary fibres, like
hispid surface of Ceraochalina (Fig. IB), results from isolated a crust. In addition, the compact multi spicular longitudinal fibres
spicules echinating connecting fibres around mesh. The ends of cored by sinuous, larger spicules produce more obvious differences
primary fibres form small hispid conules around which ectosomal between these callyspongiids and Cribrochalina. Euplacella
small one size mesh are distributed. In contrast, in Euplacella lon- Lendenfeld is also close, by its morphology and skeleton, to
gitudinal fibres of subectosomal region form three succesive sur- Cavochalina. But the three surface layers in Euplacella is signifi-
face layers (Fig. 5C). The ends of primary fibres form three dense cantly different between both genera.
ectosomallayers with small mesh of a single size without conules
(a hispid surface is produced by echinated ends of primary and Distribution
connecting fibres, appearing as palisade of isolated brushes of
spicules (Fig. 5D). The small size of conules in Ceraochalina and North coast of Australia, Torres Strait.
the regular size of the ectosomal mesh in both genera make their
surfaces appear the same type. Therefore, we consider here that
Ceraochalina is a synonym of Callyspongia (CaUyspongia). SUBGENUS TOXOCHAUNA RIDLEY, 1884
Surface hispidation is an important character of Euplacella,
Cribrochalina and Cavochalina bilamellata (see Wiedenmayer, Synonymy
1989: 107). In Euplacella surface hispidation is visible as palisade
of spicule brushes especially on the secondary fibres of surface Toxochalina Ridley, 1884a: 402.
Porifera· Demospongiae • Haplosclerida • Haplosclerina • Callyspongiidae 845

Fig. 6. Cal/yspongia (Toxochalina) folioides (Bowerbank, 1875) (Holotype and and Ridley, 1884a specimen, BMNH 1877.5.21.2034, 1882.2.23.311).
A, holotype. B, Ridley, l884a specimen, enlarged view of the surface. C, ectosomallayer, central conule, three sizes of mesh (scale 100 !Lm). D, choanoso-
mal strong network of primary irregularly distributed multispicular fibres, abundantly ramified to form connective, ramified smaller network; ectosomal
network on top (scale 500 !Lm). E, choanosomal fibres, abundantly cored, multi spicular without spongin sheath (scale 20 !Lm).

Type species Description of type species

Desmacidon folioides Bowerbank, 1875 (by subsequent des- Callyspongia (Toxochalina) folioides (Bowerbank, 1875)
ignation; Burton, 1934a: 597). (Fig. 6).
Synonymy. Desmacidon folioides Bowerbank, 1875: 295;
Definition Ridley, 1884a: 402.
Material examined. Holotype: BMNH 1877.5.21.2034 -
CaUyspongia with three sizes of ectosomal mesh and conspic- Straits of Malacca (Bowerbank collection). Specimens of Toxochalina
uous surface conules. Choanosomal strong, irregular network with folioides Ridley, 1884a: BMNH 1882.2.23.311, 550 - Prince of
compact multi spicular fibres abundantly divided to form tangled, Wales Channel, Torres Strait. H.M.S. 'Alert'.
irregular connective fibres and large triangular to irregular Description. Ramose, sponge issued from a basal, enlarged
choanosomal mesh. Scarce spongin. portion. Surface conulose but smooth in between, with small ostia
and oscula, irregularly distributed, covered by a conspicuous
Diagnosis membrane minutely reticulated with abundant toxa and oxeas.
Ectosomal fibres strong and ramified to form irregular large mesh.
Compact ramified subcylindrical branched growth form Choanosomal strong network of multi spicular primary fibres abun-
(Fig. 6 A, B), issuing from an enlarged basal portion. Surface conulose dantly divided to form secondary and tertiary fibres of a tangled
with a strong ectosomal network and strong fibres, numerous small network. Spicules: oxeas (1) 70-(78)-88 X 4/-Lm in diameter;
conules. Oscula irregularly distributed. Consistency hard, strong. oxeas (2) 62-(69)-67 X 2/-Lm in diameter. Microscleres, if present,
Ectosomal irregular network, three sizes of mesh around compact toxas, 64-(81)-115 X 2/-Lm in diameter.
small conules, strong multi spicular primary fibres, abundantly Remarks. The most distinctive character of Toxochalina
ramified (Fig. 6C). Mesh with abundant free oxeas and toxas. folioides is its surface. It has a strong surface membrane with
Choanosomal (Fig. 6D) strong network of compact multi spicular strongly ramified fibres, clearly visible under the surface mem-
primary fibres, irregularly distributed (Fig. 6E), abundantly rami- brane. The ectosomal network with large mesh is subdivided by
fied to form connective paucispicular fibres, as smaller network of fine fibres with small spicules. Toxochalina exhibits a compact and
tertiary uni to aspicular small fibres, similar to the surface network. ramified ectosomal skeleton, close to that seen in Callyspongia,
Spongin scarce, present only to bind spicules. Spicules: straight and these two genera are considered by most authors to be syn-
oxeas, microscleres toxas. onyms. We use Toxochalina as a subgenus of Callyspongia,
because in addition to possessing toxa microscleres its skeletal
Previous reviews structure is much more compact and irregular, with tangled tertiary
fibres, than Callyspongia sensu stricto, whereas additional taxo-
Ridley & Dendy (1887: 49), Dendy (1905: 139), Topsent nomic characters are required to accurately establish its exact taxo-
(1928: 66), de Laubenfels (1936a: 59), Bergquist & Wame (1980: nomic status. De Laubenfels (1936a) suggested that it was a
24), Van Soest (1980: 110), Desqueyroux-Faundez (1984: 811). synonym of Patuloscula, a review rejected here, whereas Van Soest
846 Porifera· Demospongiae • HapJoscIerida • HapJoscIerina • Callyspongiidae

Fig. 7. Arenosclera heroni Pulitzer-Finali, 1981 (holotype MSNG 46949 her 32, specimen MHNG 22920). A, New Caledonian specimen. B, Great
Barrier Reef holotype, ectosomal tangential network, no proper fibres, roundish to irregular mesh, foreign debris and spicules, slightly amount of spongin
(scale 200 J..Lm). C, choanosomal skeleton, irregular, primary and secondary fibres not clearly distinguishable. D, thicker tracts cored by foreign debris,
proper spicules or both (scale 20 J..Lm).

(1980) and Bergquist & Warne (1980) both suggested it was a syn- Diagnosis
onym of CaUyspongia.
Distribution. N coast of Australia, Great Barrier Reef, Massive, irregularly lobate and repent growth form. Oscula
Torres Strait, Prince of Wales Channel, Thursday Island, Indo- unique at end of lobes, or numerous and scattered, volcano-shaped
Malayan region, New Zealand (Bergquist & Warne, 1980); (Fig. 7A). Surface uneven, smooth, fine reticulate sandy mem-
Auckland islands (Br~ndsted, 1923), New Caledonia, Ambon, brane, aquiferous system clearly visible. Consistency hard, firm,
Indo-Pacific region (Desqueyroux-Faundez, 1984). resilient. Ectosomal tangential network, no proper fibres, roundish
to irregular mesh, foreign debris and spicules, with a small amount
of spongin. Choanosomal skeleton intricate, irregular, primary and
ARENOSCLERA PULITZER-FINALI, 1981 secondary fibres not clearly distinguishable, thicker tracts cored by
foreign debris, proper spicules or both. Thinner tracts irregularly
Synonymy distributed, sparse spicules or uncored. Spongin scarce. Micro-
scleres absent.
Arenosclera Pulitzer-Finali, 1981: 123.
Previous reviews
Type species
Pulitzer-Finali (1981: 123), Desqueyroux-Faundez (1984: 817).
Arenosclera heroni Pulitzer-Finali, 1981, by original designation.
Description of type species
Definition
Arenosclera heroni Pulitzer-Finali, 1981 (Fig. 7).
Callyspongiidae with ectosomal skeleton of sand and foreign Synonymy. Arenosclera heroni Pulitzer-Finali, 1981: 123,
debris cemented by scarce spongin, rounded mesh. Choanosomal figs 33-35; Desqueyroux-Faundez, 1984: 817, figs 27, 201-206.
fibres irregular, discontinuous, with foreign debris, proper spicules Material examined. Holotype: MSNG 46949 HER 32 -
or both. Heron Island, S Great Barrier Reef, Australia. Other material.
Porifera· Demospongiae • HapJosclerida • HapJosclerina • CalJyspongiidae 847

Arenosclera heroni. MHNG 22920 - Ile Daougae, New Caledonia. Type species
Holotype of A. rosacea Desqueyroux-Faundez, 1984: MHNG
978.437 - Ile Ndu, New Caledonia. Dactylia chaliniformis Carter, 1885c: 309 (by subsequent des-
Description. Massive, irregularly lobate, repent, issued ignation; de Laubenfels, 1936a).
from a basal encrusting plate (Fig. 7A). Each lobe 5-25 mm high,
and 5-15 mm in diameter. Apical oscules, 3-12 mm in diameter, Definition
at each lobe, deep aquiferous cavity pierced by numerous pores.
Surface smooth, uneven, with low conules, 1-2 mm length, cov- Callyspongiidae without spicules, ectosomal skeleton of fine
ered by the sandy reticulate membrane, under which the aquiferous network with rounded mesh, well differentiated aspicular fibres
system is clearly visible. Colour: grey to violet. Consistency hard, cored by abundant foreign material. Choanosomal principal fibres
firm, resilient. Ectosomal, a tangential irregular network, formed sand cored.
by foreign debris and spicules agglutinated by scarce spongin,
60-70 f.Lm in diameter with rectangular to polygonal mesh 80-250 Diagnosis
f.Lm wide (Fig. 7B). Very rare, unispiculate, fragmentary, fine
fibres, 10-15 f.Lm in diameter, irregularly present. The choanoso- Arborescent growth forms with solid digitate branches of
mal skeleton (Fig. 7C, D) is an irregular, confused network irregular diameter, dichotomously divided and swollen terminal
of undifferentiated fibres. Thicker irregular tracts, 50-100 f.lm parts (Fig. 8A). Surface even, smooth and pierced by numerous
in diameter, alternatively cored by foreign debris and by ostia. Oscula numerous, irregularly scattered, or linearly distrib-
spicules; rounded irregular mesh, 90-164-295 f.lm large, subdi- uted, tubuliform. Surface minutely reticulated. Texture elastic.
vided by uni- or bispicular fine fragmentary, secondary fibres, Ectosomal skeleton (Fig. 8B) irregular, fine network with rounded
10-30 f.lm in diameter, free of debris, that form a fine isodictyal mesh, formed by fine aspicular fibres, cored by abundant foreign
network of one spicule size. Spicules: fine oxeas, slightly bent, material. Choanosomal skeleton a network of principal longitudi-
51-(63)-90 X 1-2 f.Lm. nal divergent aspicular fibres, abundantly cored by sand and for-
Remarks. Arenosclera is atypical of Callyspongiidae in eign spicules. Connecting fine secondary fibres, free of inclusions.
having an irregular and disorganised fibre skeleton, and by its sur- Foreign inclusions present in andlor outside the fibres make it dif-
face reticulate skeleton which is only partially visible in the type ficult to observe the skeletal framework. No proper spicules.
species, and is formed by very rare, fragmentary, fine fibres and Spongin present.
abundant foreign material. We do not follow Wiedenmayer (1989:
105) who included Arenosclera as a synonym of Callyspongia, Previous reviews
considering that presence or absence of spicules and of foreign
material correspond to poorly correlated traits and represent only De Laubenfels (1936a: 59), Bergquist & Warne (1980: 34),
variations of degree within the Callyspongiidae. Arenosclera has Wiedenmayer (1989: 105).
some resemblance to Dactylia, both genera exhibiting an arena-
ceous ectosomal skeleton and abundant foreign debris. But the Description of type species
skeletal network of each is different. In Arenosclera the ectosomal
spongin is present between foreign debris but it is only fragmen- Dactylia chaliniformis Carter, 1885c (Fig. 8A-D).
tary, which is different from the well developed aspicular ectoso- Synonymy. Dactylia chaliniformis Carter, 1885c: 309;
mal fibres of Dactylia. The choanosomal skeleton of Arenosclera is Lendenfeld, 1888: 110 (as Chalinopsilla arborea var. macropora);
irregular and confused. Proper fibres are not clearly distinguish- Lendenfeld, 1888: 111 (as Chalinopsilla arborea var. micropora).
able, and it is irregularly cored by foreign debris and spicules, with Material examined. Lectotype: BMNH 1912.15.196 -
scarce spongin. In contrast, Dactylia has well differentiated S coast of Australia, Port Phillip Heads (JB Wilson collection;
choanosomal primary and connecting fibres cored only by foreign labelled '? Phoriospongia chaliniformis (Carter, 1885c) (Lendenfe1d,
debris, without spicules. Growth forms of both genera are also dif- 1889a: 200) as Dactylia chaliniformis = Spongelia varia (Gray).
ferent, with Arenosclera containing encrusting, repent and shortly Det. Burton'). Slide. BMNH 12.15.196a - slide of Carter. Compara-
lobate growth forms, whereas Dactylia chaliniformis, the type tive material. Holotype (fragment) of Chalinopsis dichotoma
species of Dactylia, was described by Carter (1885d) to be a dis- Lendenfeld, 1886c: AM G8960 (MNHN LBIM DCL2061). Holotype
tinctly 'digito-chalina-sponge'. (fragment) of Chalinopsilla elegans Lendenfeld, 1886c: BMNH
1886.8.27.62 (MNHN LBIM DCL2116).
Distribution Description. Arborescent, caulescent, with solid, digitiform
branches of irregular diameter, enlarged at their terminal parts,
There are only two species of Arenosclera described so "fig-like" (Carter, 1885c). Surface even, smooth, covered by a fine
far, apparently endemic to the southwest Pacific (Australia ectosomal network minutely reticulated, with small rounded
(Pulitzer-Finali, 1981) and New Caledonia (Desqueyroux- mesh, 90-(137)-213 f.Lm diameter, and abundant tubuliform ostia
Faundez, 1984». (Fig. 8C), masked by foreign material. Fine ectosomal aspicular
fibres irregularly distributed, 12-(19)-33 f.lm in diameter.
Choanosomal skeleton (Fig. 8D) an irregular network of primary
DACTYUA CARTER 1885 ascending fibres, 41-(60)-80 f.lm in diameter. These are radiating
to irregularly divergent, not ramified, aspicular but abundantly
Synonymy cored by foreign material. Short connecting secondary aspicular
fibres, 12-16f.Lm in diameter. Foreign inclusions, present inside
Dactylia Carter, 1885c: 309. [Chalinopsisj Lendenfeld, andlor outside the fibres, make it difficult to observe the skeleton
1886c: 569. Chalinopsilla Lendenfeld, 1888: 104; 1889a: 133. or skeletal framework.
848 Porifera· Demospongiae • HapJosclerida • HapJosclerina • CaIJyspongiidae

Fig. 8. Dactylia chaliniformis Carter, 1885. (lectotype BMNH 1886.12.15.196). A, lectotype. B, ectosomal irregular, fine network with rounded meshes,
abundantly cored by foreign material. C, surface tangential view, even, minutely reticulated, tubuliform pores (scale 20 f.Lm). D, choanosomal network with
primary aspicular fibres, abundantly cored by sand and foreign spicules.

Remarks. De Laubenfels (1936a: 59) included Dactylia in imitated species. Lendenfeld (1889a) also considered species
Callyspongiidae based on the characters of its principal skeleton, of Paraspongia and Psammoclema as species of Chalinopsilla.
despite the absence of spicules, and that it had a typical structure as Lendenfeld (1888, 1889a) assigned most of Carter's new species
'belonging to one of the orders properly having spicules' . In addition, to his new form, to give new concepts to Carter's genera.
de Laubenfels synonyomised several genera ([ Chalinopsis] Wiedenmayer (1989: 105) provided and extensive discussion of the
Lendenfeld, 1886c (preocc. by Chalinopsis Schmidt, 1870), taxonomic problems between these genera. Pulitzer-Finali (1981)
Chalinopsilla Lendenfeld, 1888, a replacement name for the preoccu- transferred Chalinopsilla arborea Lendenfeld, 1888 to his recently
pied Chalinopsis Lendenfeld, Paraspongia Carter and Psammoclema described genus Arenosclera.
Marshall) with Dactylia, of which only Chalinopsilla is now How then to correlate the presence of sand and foreign material
accepted in synonymy. with the absence of spiCUles? The absence of a strict definition of
The genus has a tortuous taxonomic history. Lendenfeld Callyspongiidae based on types of skeletons makes these charac-
(1888) described Dactylia chalinifonnis as Chalinopsilla arborea ters very subjective. Bergquist & Warne (1980) considered
var. macropora Duchassaing & Michelotti (sic), and var. micropora Dactylia and Chalinopsilla as valid genera, based on the presence
Carter (sic). To justify his changes he inferred the 'imitating power' or absence of foreign material, respectively, however this discrimi-
of some of his new species. He considered Chalinopsilla arborea nation is incorrect. Chalinopsilla sensu Bergquist & Warne, 1980,
was able to include spicules taken from decayed specimens of the lacks foreign material, whereas Wiedenmayer (1989) showed that
Porifera· Demospongiae • Haplosclerida • Haplosclerina • CaUyspongiidae 849

its type species, Chalinopsilla dichotoma, contains abundant foreign Description of type species
material, which we also confirm from re-examination of type mate-
rial, especially in the primary fibres. Wiedenmayer (1989: 105) con- Siphonochalina coriacea Schmidt, 1868 (Fig. 9).
sidered the presence or absence of spicules versus foreign material to Synonymy. Schmidt 1868: 7; Topsent, 1927a [file card],
be poorly correlated traits, with differences often only in abundance figs 1-4.
of foreign material coring the Callyspongiidae skeleton between Material examined. Holotype: MNHN LBIM DCL
taxa. Consequently, we consider that the absence of spicules and the (Schmidt spec. 77) (fragment BMNH 1868.3.2.13) - La Calle,
presence of foreign material could be two independent characters to Algeria. Other material. Holotype (fragment) of Siphonochalina
differentiate Dactylia from CaUyspongia as there are different gen- typica var. levis Lendenfeld, 1887c: AM G8976 (MNHN LBIM
era of Callyspongia-like sponges with spicules replaced by sand. We DCL2077) - Port Jackson, NSW, Australia. Holotype (fragment)
have conserved Dactylia Carter, 1885c (which has priority over of Siphonochalina osculata var. macropora Lendenfeld, 1887c:
Chalinopsilla Lendenfeld, 1886c), for these sponges which have lost AM G8993 (MNHN LBIM DCL2082) - Western Australia.
spicules and replaced them, at least in the ascending primary fibres, Holotype of Siphonochalina axialis Lendenfeld, 1887c: AM G
by sand and other foreign material. unreg. (fragment MNHN LBIM DCL1956) - Port Jackson, NSW,
Australia. Holotype of Sclerochalina asterigena Schmidt, 1868:
Distribution MNHN LBIM DCL89/11 - Algeria. Neotype of Tubulodigitus
(=Siphonochalina) communis Dendy, 1889: BMNH 1889.1.21.1.
Species of Dactylia are known from Australia (Carter, 1885c; Specimen (Fig. 9B) and slide of Siphonochalina (Tubulodigitus)
Lendenfeld, 1886c, 1889a; Hooper & Wiedenmayer, 1994); West communis Cater, 1881b: BMNH 1881.10.21.338, 339a - H.M.S.
Indies (Van Soest, 1980); New Zealand (Lendenfeld, 1888; 'Alert', Port Jackson. BMNH 1882.1.5.2 - Kurrachee.
Bergquist & Warne, 1980). Description. Tubular sponge (Fig. 9A), divided into several
tubes from the basal part, with a large, interiorly smooth aquiferous
cavity of uniform diameter, ending in a central oscule of the same
SIPHONOCHAUNA SCHMIDT, 1868 diameter. Surface, smooth covered by a fine ectosomal network
with numerous ostia. Texture fragile, resilient. Ectosomal skeleton
Synonymy (Fig. 9C) formed by fine, irregular network of unispicular fibres or
single spicules. Longitudinal view of the surface (Fig. 9D) exhibits
Siphonochalina Schmidt, 1868: 7. Sclerochalina Schmidt, small peripheral condensed mesh of the subectosomal region, pro-
1868: 8. [Siphonellal Lendenfeld, 1887c: 808 (preoccupied). duced by terminal part of choanosomal fibres, abundantly ramified
Tubulodigitus Carter, 1881b: 367. and by connecting fibres (Fig. 9D). Principal fibres paucispicular,
2-3 spicules, longitudinally divergent, with a large spongin sheath,
Type species 26-40 J..lm in diameter, connecting or secondary fibres, unispicular
parallel, 12-20 J..lm or more in diameter, often of the same diameter
Siphonochalina coriacea Schmidt, 1868 (by monotypy). as primary fibres. Spicules: slightly bent oxeas, 96-(111)-128X
3.5-(4)-4.5 J..lm in diameter.
Definition Remarks. Topsent (192530 1927a, 1938b) redescribed,
defined and erected new species of this genus originally created for a
Callyspongiidae with an ectosomal network with one size of single briefly (but exactly) described species by Schmidt (1870). It
clearly defined mesh, unispicular fibres. Choanosomal, isolated, has been included as a synonym of Callyspongia for a long time by
scarcely cored primary longitudinal fibres. Peripheral subectoso- authors (e.g., Burton, 1932b, 1934a; de Laubenfels, 1936a) who
mal condensation present. were not familiar with the type species. Based on Topsent's (1938b)
descriptions of Sclerochalina asterigena Wiedenmayer (1977b: 91)
Diagnosis considered 'peripheral condensation' of the main skeleton to be the
main diagnostic criterion differentiating Siphonochalina from other
Tubular (Fig. 9A, B), solid, branching, creeping or erect growth callyspongiids. Dendy (1890a) described several West Indian species
form. Surface even, smooth, minutely reticulated and pierced by recognising the peripheral condensation of the ectosomal skeleton as
numerous ostia. Oscula unique, at the end of tubes and of the same a 'proliferation of new fibres between the old ones'. In spite of this
diameter. Texture elastic, resistant. Ectosomal skeleton a fine irregu- statement he subsequently confused the genera Siphonochalina,
lar tangential network, triangular mesh, formed by unispicular fibres Callyspongia and Patuloscula when placing some of the West Indian
or single spicules. Subectosomal skeleton with ends of longitudinal species (Dendy 1890a). At the same time he considered
fibres profusely divided and connected by short, secondary unispicu- Siphonochalina, Callyspongia and Spinosella (pro Cladochalina) as
lar fibres, peripheral condensation. Choanosomal skeleton with iso- genera exhibiting very similar morphologies that could result in a
lated, parallel, strong longitudinal primary fibres with large spongin wrong identification and that peripheral condensation of the ectoso-
sheath. Connecting secondary fibres, parallel, regularly distributed. mal skeleton is an important character to differentiate them.
Spongin and spicules abundant. Our study of type specimens showed clearly the different
types of ectosomal skeletons in Siphonochalina, Callyspongia
Previous reviews (CaUyspongia) and CaUyspongia (Cladochalina). Siphonochalina
(including the junior subjective synonyms Sclerochalina,
Schmidt (1868: 7), Ridley (1884a: 401), Topsent (1892a: 66; Siphonella and Tubulodigitus) presents a dense ectosomal tangen-
1925a: 236; 1938b: 4) (as Sclerochalina), Van Soest (1980: 113), tial network with small mesh, varying from triangular-rectangular to
Wiedenmayer (1989: 104). polygonal, which is in no way divisible into primary or secondary
850 Porifera· Demospongiae • Haplosclerida • Haplosclerina • Callyspongiidae

Fig. 9. Siphonochalina coriacea Schmidt, 1868 (holotype MNHN DT77), A, holotype. B, neotype of Siphonochalina communis Dendy, 1889. C, ectoso-
mal skeleton formed by fine, irregular network of unispicular fibres or single spicules over choanosomal isolated, parallel, strong longitudinal primary
fibres with large spongin sheath, longitudinal view. Connecting secondary fibres, parallel, regularly distributed. Spongin and spicules abundant. D, periph-
eral condensation occurring by proliferation of the subectosomal ends of parallel primary fibres, connected by fine unispicular fibres, smaller mesh.

mesh. Longitudinal section through the subectosomal region surface to form connecting fibres parallel to the surface, or badly
(Fig. 9D) shows a condensed region of rectangular small mesh differentiated fibres, replaced by single spicules. In Callyspongia
and abundant, parallel secondary fibres, produced by ramification (Callyspongia) and Callyspongia (Cladochalina) primary longitu-
of subectosomal ends of primary longitudinal fibres, in contrast dinal and connecting fibres are well differentiated, fasciculated or
to the larger ectosomal polygonal meshes of Callyspongia non-fasciculated, forming regular, short connecting fibres and
(Cladochalina). Primary longitudinal and connecting fibres of both rectangular scalariform in Callyspongia (Callyspongia) or tangled,
genera are also different. In Siphonochalina primary. fibres are irregular meshes in Callyspongia (Cladochalina). In addition, ter-
parallel-radiating, non-fasciculated, abundantly ramified near the minal ends of primary fibres in CaUyspongia (Cladochalina) form
Porifera· Demospongiae • Haplosclerida • Haplosclerina • Callyspongiidae 851

conspicuous or inconspicuous conules. Siphonochalina exhibits in Hooper & Wiedenmayer (1994: 88) synonymised it, without
a smooth, non-conulous surface. justification, with Callyspongia (following Wiedenmayer, 1989:
Several nominal genera are included here in synonymy with 104). Tubulodigitus (type species Tubulodigitus communis Carter,
Siphonochalina and require further comment. The inclusion of 1881b, by monotypy) was referred to Adocia by Burton (1934a:
Sclerochalina Schmidt (type species Sclerochalina asterigena 536), with the type specimen lost and Dendy's (1889: 81) specimen
Schmidt, 1868, by monotypy) follows Topsent (1938b). of Siphonochalina communis nominated as neotype. De Laubenfels
[Siphonella] Lendenfeld (type species Siphonochalina fortis (1936a: 59) considered that in this genus 'Callyspongia and
Ridley, 1881, by subsequent designation; Burton, 1934a: 540) is callyspongid tendencies are so pronounced that it would seem to be
a junior homonym of Siphonella Macquart, 1835 (Diptera), decidedly worthy of separate generic rank'. This opinion is not
[Siphonella] IsseI, 1869 (Mollusca) and [Siphonella] Hagenow, adopted here and Tubulodigitus is retained in synonymy with
1850 (Polyzoa). The subsequent designation of Siphonella commu- Siphonochalina.
nis Lendenfeld, 1887c as the type species by Van Soest (1980: 118) Distribution. Coast of Algeria, East coast of Australia,
is therefore invalid, who in any case included it in his monograph Western Australia, New Zealand (Wiedenmayer in Hooper &
in a section of 'generic names of uncertain identity'. Wiedenmayer Wiedenmayer, 1994), Gulf of Manaar, Kurrakhee (Carter, 1881 b).
Systema Porifera: A Guide to the Classification of Sponges, Edited by John NA. Hooper and Rob WM. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Chalinidae Gray, 1867

Wallie H. De Weerdt

Zoological Museum, University of Amsterdam, PO. Box 94766, 1090 GT Amsterdam, Netherlands. (weerdt@science.uva.nl)

Chalinidae Gray (Demospongiae, Haplosclerida), including synonyms Renieridae Ridley, Gelliidae Ridley, Haliclonidae de Laubenfels
and Adociidae de Laubenfels, belongs to the suborder Haplosclerina together with Callyspongiidae de Laubenfels and Niphatidae
Van Soest. Chalinidae have a world-wide distribution with possibly hundreds of extant species living in all parts of the oceans. Twenty-
seven nominal genera have been described, but only four genera are presently considered valid: Chalinula, Cladocroce, Dendroxea and
Haliclona, the latter containing the subgenera Gellius, Halichoclona, Haliclona, Reniera, Rhizoniera and Soestella. Chalinidae differ
from other haplosclerid families by the possession of a unispicular tangential ectosomal skeleton and a choanosomal skeleton with
unispicular secondary lines. Characters in common with other marine haplosclerid families are smooth, diactinal megascleres and
microscleres, if present, in the form of sigmas, toxas, raphides and microxeas. Some species exhibit extremely high variability in mor-
phological characters, inhibiting their reliable identification by the non-specialist, whilst others are remarkably constant in their features.
Keywords: Porifera; Demospongiae; Haplosclerida; Haplosclerina; Chalinidae; Chalinula; Cladocroce; Dendroxea; Haliclona (Gellius);
Haliclona (Halichoclona); Haliclona (Haliclona); Haliclona (Reniera); Haliclona (Rhizoniera); Haliclona (Soestella).

DEFINITION, DIAGNOSIS, SCOPE with a 'skeleton formed of regular, reticulated, anastomosing,

horny fibres, which have one or more series of regular small
Synonymy siliceous spicules in the central lines' . Gray (1867a) assigned nine
genera to the family, including [sodictya, Acanthella, Clathria, and
Chalinidae Gray, 1867a: 511. Renierinae Schmidt, 1870: 39. Astrospongia, but only the first mentioned genus, Chalina
Gelliadae Gray, 1872a: 449. Renieridae Ridley, 1884a: 406. Gelliidae Bowerbank (which is a junior synonym of Haliclona Grant, 1836),
Ridley, 1884a. Gelliinae Ridley & Dendy, 1887: 37. Haliclonidae de is a 'true' chalinid as presently understood. That Gray's definition
Laubenfels, 1932: Ill. Adociidae de Laubenfels, 1936a: 65. was far too wide is also obvious from the work of Lendenfeld
(1887c) who took the liberty to add many more genera to the fam-
Definition ily, most of which are incompatible with the currently accepted
concept of Chalinidae. Three more families were later proposed by
Haplosclerina with a delicate reticulated choanosomal skele- different authors, all three comprising a wide range of genera, of
ton of uni-, pauci- or multi spicular primary lines, which are regu- which the type genera are closely related to Haliclona. Renieridae
larly connected by unispicular secondary lines. Ectosomal skeleton, Schmidt (1870), was established without a clear definition, for
if present, a regularly hexagonal, unispicular, tangential reticulation. Reniera (subgenus of Haliclona), Amorphina, Pellina, Eumastia
(all Halichondriidae), Foliolina (Phloeodictyidae), Tedania
Diagnosis (Tedaniidae), Schmidtia (Petrosiidae), Plicatella (Microcionidae),
and Auletta (Axinellidae). Gelliadae was erected by Gray (1872a)
Thickly encrusting, cushion-shaped, ramose or tubular growth as part of the order Hamispongia ('Sponge horny or fleshy,
forms, cushion-shaped sponges commonly with oscular chimneys strengthened with fusiform or needle-like spicules, interspersed
or mounds. Consistency soft to rather firm, also spongy. Colour with anchorate or bihamate spicules.'), for sponges with 'defensive
purple, violet, pink, brown, blue or green, occasionally white. spicules filiform' (Gellius (subgenus of Haliclona), Biemna
Megascleres smooth oxeas or strongyles, microscleres, if present, (Desmacellidae), Asychis (Gellius, subgenus of Haliclona) and
sigmas, toxas, raphides or microxeas. Oceanapia (Phloeodictyidae» and sponges with 'defensive
spicules clavate at the end' (Dymnus (Damo)). In 1936 de
Scope Laubenfels erected the family Adociidae, as part of the order
Poecilosclerida for sponges with a 'special dermal skeleton' and
Twenty-seven nominal genera have been described, four of included, apart from the type genus Adocia (Haliclona, subgenus
which are considered valid: Chalinula, Cladocroce, Dendroxea of Haliclona), such widely divergent genera like Melophlus
and Haliclona (with the subgenera Haliclona, Reniera, (Ancorinidae), Menanetia (Halichondriidae), and Protoschmidtia
Halichoclona, Soestella, Gellius, and Rhizoniera). (Chalinidae, incertae sedis). Topsent (1928c) included the first three
families, as subfamilies of the family Haploscleridae, in a newly
History and biology created order Haplosclerida. Although he accepted various kinds of
choanosomal skeletal architectures, he made the 'Haploscleridae'
The taxonomic history of Chalinidae is long and complicated, [name corrected to Haliclonidae by de Laubenfels, 1932, which is
and only the main developments can be dealt with here. The name a junior synonym of Chalinidae1 much more comprehensible by
Chalinidae was proposed by Gray in 1867 for a group of highly allowing only diactinal spicules as megascleres ('mostly oxeas,
unrelated sponges of the order Keratospongia ('horny sponges') sometimes strongyles' [translated]), only sigmas, toxas, microxeas

852
Porifera· Demospongiae • Haplosclerida • Haplosclerina • Chalinidae 853

and raphides as microscleres, and excluding sponges with echinat- Toxadocia. Van Soest's new concept of the family was followed
ing spicules. He used the subfamily Gelliinae for sponges with by De Weerdt (1986) who, after a taxonomic revision and phylo-
microscleres, the Renierinae for sponges with spicule-reinforced genetic analysis of North East Atlantic Chalinidae, went a step
skeletons, and the Chalininae for sponges with spongin-reinforced further by abandoning most of the nominal genera and recognizing
skeletons. Since the addition of the family Adociidae by de only eight monophyletic 'species groups' (De Weerdt, 1989).
Laubenfels (1936a) for sponges with a special dermal skeleton, Recently, De Weerdt (2000) published a monograph of Caribbean
two general trends can be observed (cf. Van Soest, 1980 for Chalinidae, in which the species groups have been given proper
a detailed review): one of spongologists following the line of names, based on study of all the available type species of the nom-
Topsent (e.g., Levi, 1973; Griessinger, 1971), the other following inal genera. Chalinidae occur in all seas, from polar regions to the
de Laubenfels (e.g., Hechte1, 1965; Bergquist & Warne, 1980). Van tropics. Although they can be found at all depths, they are most
Soest (1980) made clear that neither of these classifications were common in shallower parts of both tropical reef environments and
based on characters reflecting phylogenetic relationships. He rede- colder, subtidal habitats. They may form substantial components of
fined the family Haliclonidae with emphasis on the choanosomal mangrove communities. Chalinid sponges usually do not reach
skeletal architecture (' ... choanosomal skeleton consisting of uni- large sizes, but under certain conditions, like mangrove habitats,
or paucispicular primary (ascending) tracts or fibres, intercon- they may form proliferations of more than 1 m long. The reproduc-
nected by single spicules at right angles .. .'), and included both tion is so far known to be exclusively viviparous, with ciliated lar-
genera without a dermal skeleton and without microscleres, like vae of 140 X 470 /-Lm (cf. Wapstra & Van Soest, 1987). Like other
Haliclona, with a dermal skeleton and without microscleres, like haplosclerids, some species produce toxic chemical compounds,
Adocia, with microscleres and a dermal skeleton, like Sigmadocia, like cyclic alkylpiperidines (Van Soest & Braekman, 1999) and
and other microsclere bearing genera like Gellius, Orina and amino alcohols (Devijver et aI., 2000).

KEY TO GENERA

(1) At the basis of the sponge there is a densely reticulated mass of spicula, choanosomal skeleton consisting of almost plumose,
branching spicular tracts, which thin out towards the surface ............................................................................................. Dendroxea
No densely reticulated mass of spicules at the basis of the sponge, choanosomal skeleton a rather delicate, iso- or anisotropic,
ladder-like reticulation or consisting of multi spicular fibre tracts with a rather dense subisotropic reticulation in between ............. 2
(2) Secondary lines ofthe choanosomal skeleton more than one spicule long .......................................................................... Chalinula
Secondary lines of the choanosomal skeleton one spicule long ...................................................................................... .......... .......... 3
(3) Skeleton consisting of multi spicular fibre tracts with a rather dense subisotropic reticulation in between ....................... Cladocroce
Skeleton iso-, subiso- or anisotropic, but without multi spicular fibre tracts throughout the sponge .................................... Haliclona

CHAUNULA SCHMIDT, 1868 Previous review

Synonymy De Weerdt, 2000.

Chalinula Schmidt, 1868: 7. Acervochalina Ridley, 1884a: Description of type species

398. Phylosiphonia Lendenfeld, 1887c: 796. Chalinodendron
Lendenfeld, 1887c: 819. Nara de Laubenfels, 1954: 76. Katiba de Chalinula renieroides Schmidt, 1868 (Fig. 1).
Laubenfels, 1954: 186. Synonymy. Chalinula renieroides Schmidt, 1868: 7.
Chalinula fertilis Keller, 1879: 318, pis. 18-20. Acervochalina
Type species fertilis; De Weerdt & Van Soest, 1986: 27, figs 19,20. Taxonomic
decision for synonymy: De Weerdt, 2000.
Chalinula renieroides Schmidt, 1868 (by subsequent Material examined. Holotype (slide): MNHN D.T 748 -
designation; de Laubenfels, 1936a: 47). two microscopic slides made by Topsent from Schmidt's specimen,
labeled 'Chalinula renieroides O. Schmidt, Type, Museum, Exp.
Definition sc.de l' Algerie'.
Description. Thin crusts to laterally spreading cushions of
Choanosomal skeleton with secondary lines more than one equal thickness (up to 0.5 cm), with numerous, regularly distrib-
spicule length. No ectosomal skeleton. uted oscula, flush with the surface and on slightly raised mounds,
0.8-1.5 mm in diameter (Fig. 1A). Surface even, slightly hispid.
Diagnosis Consistency soft, somewhat resilient, compressible. Colour light
brown. Choanosomal skeleton (Fig. lB) a regular reticulation of
Thinly encrusting, cushion-shaped or irregularly digitate unispicular (occasionally paucispicular) primary lines which are, at
growth forms. Consistency spongy, elastic, or very soft and limp. unequal distances but regularly, connected by secondary lines of
Colour brown or purplish. Oxeas short, ranging from vestigial to one spicule thick and two-four spicules in length. Most meshes
cigar-shaped; a high variation in form and size may occur within between the lines are rectangular, 133-195/-Lm in diameter.
the same species. Spongin usually abundant, with high variation Spongin moderate, completely enveloping the skeletal lines.
within one species. No microscleres. Spicules (Fig. 1C) short, straight or only very slightly curved oxeas
854 Porifera· Demospongiae • Haplosclerida • Haplosclerina • Chalinidae

Fig. 1. Chalinula renieroides Schmidt, 1868. A, habit of preserved specimen, ZMA Por. 3043 (scale 0.5 cm). B, perpendicular section of choanosome,
Topsent's slide of Chalinula renieroides, MNHN DT. 748 (scale 200/Lm). C, SEM photo of spicules made from ZMA Por. 3043 (scale 20 /Lm). D, per-
pendicular section of choanosome, Topsent's slide of Chalinulafertilis Keller, 1879, MNHN DT. 2279 (scale 250 /Lm).

with a pronounced fusifonn shape and sharp points, 81-93.9-105 X leaves no doubt about their identity. For further synonymy of
3.6--5.1--6.6 f.Lm. Frequently the sponges produce many embryos Chalinula limbata see De Weerdt, 1986 (as Acervochalina).
(Fig. ID). Distribution and ecology. Mediterranean, Azores, Phylosiphonia was erected by Lendenfeld (1887c: 796) for
Caribbean. Growing on and under stones in shallow sandy bottom. Chalinula fertilis Keller (1879), subsequently designated as type
Remarks. The genus Acervochalina Ridley (1884a: 398) species by Burton (1934a: 531). For the conspecifity of Chalinula
was erected (by original designation) for North Atlantic Chalina fertilis and C. renieroides see De Weerdt, 2000.
limbata (Bowerbank, 1866: 373). This species was first described Chalinodendron was erected by Lendenfeld (1887c: 819) for
by Montagu (1818: 111, pI. XV figs 2, 3) as Spongia limbata Chalina gracilenta Bowerbank (1866), subsequently designated
(Fig. 2A), then by Johnston (1842: 168, pI. XIX figs 3-5) under the as type species by Burton (1934a: 530). This species is a junior
same name (Fig. 2B-D). Bowerbank (1866) transferred it to synonym of Spongia limbata Montagu (1818). Both the holotype
Chalina (Fig. 2E). The earliest available material of Spongia lim- of Chalina gracilenta (BMNH 1877.5.21.2089, Torquay, figured
bata is that of Johnston in the BMNH collection. BMNH 47.9.7.88 specimen of Bowerbank, 1874, pI. LXVII fig. 4, designated as the
(,RabT) consists of two sponges, one of which is the specimen of type by Bowerbank on p. 171 of the same volume) (Fig. 2G-I), and
Johnston, 1842, pI. XIX fig. 5 (Fig. 2D). It is a finn, cushion- the other specimen of Bowerbank (BMNR 1877.5.21.2090,
shaped sponge attached to an alga, with a very fibrous structure, Scarborough) are cushion-shaped with circular oscula, with a very
and hispid surface. The skeleton is a reticulation of primary and spongy and fibrous structure. Because of the fragility of this mate-
secondary spongin fibres (Fig. 2J), the first 18-76f.Lm thick, cored rial no slides were made, but the BMNH holds specimens identi-
by 4--20 spicules. The secondary fibres are 1--6 spicules long and fied by Nonnan as Chalina gracilenta (BMNR 1910.1.1.382,
3-7 spicules thick. The diameter of the meshes is 170-400 f.Lm. North Devon; 1910.1.1.384, Cumbra), which were studied instead.
The oxeas are vestigial, fusifonn, 52.8--65.1-74.4 X 1-1.5-2.2 f.Lm. The skeleton of BMNH 10.1.1.383 (Fig. 2K) is a reticulation
This specimen, as well as Johnston's other BMNH material of the of spongin fibres, 9.5-38 f.Lm thick, cored by 2-20 spicules. The
species (BMNH 47.9.7.89 ('Rab?'), 47.9.7.91 ('Britain'), and secondary fibres are 3-5 spicules in length and 1-5 spicules thick.
47.9.7.92 (,Devonshire'» confonns to Montagu's description and The meshes between the fibres are 114--460 f.Lm in diameter.
figures. Bowerbank's material of Chalina limbata is in the BMNH The oxeas are vestigial, 60--100.8 X 1-4.3 f.Lm. There is no doubt
represented by, among others, BMNH 1877.5.21.2086 (Cornwall, about the conspecifity of Bowerbank's Chalina gracilenta with
figured specimens of Bowerbank, 1874, pI. LXVII figs 7-10) Montagu's Spongia limbata and therefore about the synonymy of
(Fig. 2E-F), 1877.5.21.2087 (Cornwall, figured specimen of Chalinodendron with Chalinula.
Bowerbank, 1874, pI. LXVII fig. 11) and 1877.5.21.2088 Nara was erected by de Laubenfels (1954: 76, fig. 46; by
(St. Andrews). No slides of these specimens were made, but their original designation) for Nara nematifera de Laubenfels, 1954.
external appearance (all of them being small, fibrous cushions) The holotype, USNM 22980 (Ebon Lagoon, Marshall) is a thinly
Porifera' Demospongiae • Haplosclerida • Haplosclerina • Chalinidae 855

Fig. 2. Chalinufa fimbata (Montagu, 1818 as Spongia) . A, habit, reproduced from Montagu, 1818 (scale I cm). B-D, habit, reproduced from Johnston
(1842, as Spongia). E, habit, reproduced from Bowerbank (1874, as Chafina) (scale 0.5cm). F, oxea, reproduced from Bowerbank (1874, as Chafina)
(scale 25 fLm). G-H, habit, reproduced from Bowerbank (1874, as Chafina g racifenta) (scale I cm). I, oxea, reproduced from Bowerbank (1874,
as Chafina gracifenta) (scale 25 fLm). J, perpendicular section of choanosome made from Johnston 's specimen of Spongia fimbata , BMNH 1847.9.7.88
(scale 300 j.Lm). K, perpendicular section of choanosome made from BMNH 1910.1.1.383 (Chafina gracifenta) (scale 300 j.Lm).

Fig. 3. A, perpendicular section of choanosome made from the holotype of Nara nematifera de Laubenfels (1954) (scale 300 fLm). B, perpendicular sec-
tion of choanosome made from the holotype of Katiba milnei de Laubenfels (1954) (scale 150 j.Lm).

encrusting, very soft sponge, completely covering a piece of coral Katiba was erected by de Laubenfels, (1954: 186, fig. 124; by
of 3.3 X 2 X 1.8 cm. Oscules were not observed. The skeleton monotypy) for Katiba milnei de Laubenfels, 1954. The holotype,
(Fig. 3A) is a delicate reticulation of slightly wavy primary lines of USNM 22948 (Ebon Atoll, Marshall Islands) consists of two thin,
1-2 spicules thick, loosely connected by secondary lines of 1-2 rather soft crusts on coral rubble of ca. 2 X 1 cm, with a smooth
spicules long and 1 spicule thick. Spongin is not abundant, but surface. The skeleton (Fig. 3B) is an extremely delicate reticulation
clearly present at the nodes of the spicules. The oxeas are very of very small, vestigial, somewhat crooked strongyloxeas of
slender, straight or slightly curved, 85.5-99 X 0.9-2.3 j.Lm. The 55.1-63.9-77.9 X 0.4--0.6--0.8 j.Lm, without a clear distinction
enigmatic 'threads' mentioned by de Laubenfels (1954) is a symbi- between primary and secondary lines. Most oxeas are encased in
otic fungus (Dr w.F. Prud'homme van Reine, pers. comm.). This a thin sheath of spongin. The abundant trichodragmata mentioned
species is extremely common throughout the tropical Indo-west by de Laubenfels could not be found. The species clearly belongs
Pacific, encrusting live coral in particular. It has a distinctive to Chalinula . Similar to Nara nematifera this species is also very
vibrant mauve colouration with the wavy 'surface threads' pale or common throughout the Indo-west Pacific, occurring as extremely
white, and with very small oscules scattered over the surface. It is thin crusts on dead coral, khaki green in live colouration, and once
mucusy, with a slimy texture that mostly disintegrates if removed again nearly impossible to remove intact from the substratum
from the substratum (J.N.A. Hooper, pers. comm.). (J.N.A. Hooper, pers. comm.).
856 Porifera· Demospongiae • HapJosc1erida • HapJosc1erina • Chalinidae

Fig. 4. A, Cladocrocefibrosa Topsent, l892a. Habit, reproduced from Topsent, 1892a. B, Cladocroce incurvata Levi & Levi, 1983b. Tangential view of
ectosomal skeleton made from the holotype (scale 500 fLm). C, Cladocroce aculeata Pulitzer-Finali, 1982. Perpendicular section of choanosome made
from a preserved specimen in ZMA (48!III/OIA) (scale 500 fLm).

CLADOCROCE TOPSENT, 1892 paucispicular reticulation. Ectosomal skeleton, if present, a tangen-

tial, uni- or paucispicular, iso- or subisotropic reticulation.
Synonymy Microscleres, if present, toxas.

Cladocroce Topsent, 1892a: 72. Toxiclona de Laubenfels,

1954: 73. Diagnosis

Type species Sponges lamellate or tube-shaped. Colour grey, salmon pink,

mauve or ochre. Surface smooth, sometimes conulose. Consistency
Cladocrocefibrosa Topsent, 1892a (by monotypy). firm but elastic, compressible. In the lamellate forms the oscules
are circular, flush with the surface, rather small, not abundant and
Definition (modified from Fromont, 1993) they may appear at both sides of the lamellae.

Chalinidae in which the choanosomal skeleton is reinforced Previous review

by multispicular primary tracts that occasionally anastomose. In
between these tracts the skeleton is a rather dense, subisotropic, Fromont, 1993.
Porifera' Demospongiae • Haplosclerida • Haplosclerina • Chalinidae 857

:1f
E/,J ~
Fig.5. Cladocroce gaussiana (Hentschel, 1914, as Siphonochalina). A, habit of preserved specimen in the USNM (scale 1 cm). B, perpendicular section
of choanosome reproduced from Hentschel (1914) (scale 150 j,Lm). C, oxea, reproduced from Hentschel (1914). D, perpendicular section of choanosome
made from preserved specimen in the NMHN (scale 500 j,Lm). E, oxeas and toxas (scale 250 j,Lm).

Description of type species by De Weerdt (2000) to be a synonym of the subgenus Haliclona,

but this viewpoint cannot be sustained. The ZMA has several
Cladocrocefibrosa Topsent, 1892a: 72 (Fig. 4). specimens from the Antarctic, on loan from the USNM, which
Synonymy. Cladocroce fibrosa Topsent, 1892a: 72, pl. III conform entirely to Hentschel's S. gaussiana, and which exhibit
figs 1,2. all the characters of Cladocroce. The sponges are fan-shaped
Material examined. None. (Fig. 5A), with a firm but flexible consistency, without apparent
Description (after Topsent, 1892a). Vase-shaped sponge oscula or with small, circular oscula either at the outer or on the
(Fig. 4A) of grey colour, 15 cm high, 11 cm in diameter, thickness inner side of the fan. The choanosomal skeleton consists of multi-
of the wall 0.6 cm. Inner side of the wall finely hispid. Both sides spicular tracts, with a rather dense, subisotropic reticulation in
of the wall with openings ('without distinction between pores or between (Fig. 5B, D). The ectosomal skeleton is a tangential,
oscula') of about 0.6 mm. From the base of the sponge arise several subisotropic reticulation. Spongin moderate. The oxeas are robust,
strong, white fibres which branch frequently and diminish in size with slightly mucronated, conical points, 207-255.5-278 X
towards the upper part of the sponge. They consist of oxeas running 9.3-14.4-16.8 /-Lm. Microsc1eres toxas with slightly recurved apices,
parallel and bound together by spongin. In between these fibres 93-134.1-165 X 1.5-3.2-4.5 /-Lm. Siphonochalina gaussiana and
the skeleton is that of 'Reniera'. Oxeas robust, 600 X 18 /-Lm. the genus Toxiclona may evidently be assigned to Cladocroce.
Distribution and ecology. Known only from the type locality,
Azores (between Pico and Sao Jorge), 1300m.
Remarks. Cladocroce is a rare, hitherto somewhat ill-known DENDROXEA GRIESSINGER, 1971
genus, with mainly deep-water representatives. These are
C. incurvata Levi & Levi (1983b: 974, pl. IV figs 4, 35, New Synonymy
Caledonia), C. osculosa (Topsent, 1927b: 19, Portugal, 749-31Om),
and C. spathiformis Topsent (1904: 242, pl. III fig. 1, pl. XVII Dendroxea Griessinger, 1971: 152. Dendrectilla Pulitzer-
fig. 17, Azores, 1165 m). C. aculeata Pulitzer-Finali (1982: 109, Finali, 1983: 593.
figs 20, 21) was described from shallow-water from the Great
Barrier Reef (see also Fromont, 1993). The holotype of C. incurvata Type species
is MNHN DCL. 2942; a microscope slide made from this specimen
is present in the ZMA and contains a section of the ectosomal Reniera [enis Topsent, 1892b (by monotypy).
skeleton (Fig. 4B). It clearly shows the multispicular tracts and the
rather dense, subisotropic skeleton. The oxeas (after Levi & Levi, Definition
1983b) are 180-220 X 8-10 /-Lm. Cladocroce aculeata is repre-
sented in the ZMA by several specimens, which all agree with the Chalinidae with a basal densely reticulated mass of spicules,
description of Pulitzer-Finali (1982) and Fromont (1993) (Fig. 4C). from which arise multi spicular, plumose, branching spicular tracts,
Toxiclona was erected by de Laubenfels (1954: 73, by original which thin out towards the surface. Between the primary lines
designation) for Siphonochalina (?) gaussiana Hentschel (1914: there is a reticulation of single spicules, with many spicules
136, pl. IV fig. 17, pl. VIII fig. 17). Toxiclona was first considered irregularly scattered in between.
858 Porifera· Demospongiae • Haplosclerida • Haplosclerina • Chalinidae

Fig. 6. Dendroxea lenis (Topsent, 1892b, as Reniera). A, habit of preserved specimen, reproduced from De Weerdt & Van Soest (1986) (scale 1 cm).
B, oxeas, reproduced from De Weerdt & Van Soest (1986) (scale 50 f.Lm). C, perpendicular section of choanosome of ZMA Por. 5462, Banyuls (scale 500 f.Lm) .

Diagnosis densely reticulated mass of spicules. The tracts are connected

by transverse single spicules, but there also many loose, disori-
Soft, thinly encrusting sponges, to 0.5 cm thick, with an even ented spicules scattered in between. Spongin very scarce. Oxeas
or somewhat irregular surface and greyish-white colour. Oscula (Fig. 6B) thin, slightly curved, with tapering points, 118-165 X
indistinct. No ectosomal skeleton. Spongin scarce. Oxeas thin, ca. 3-3.5 fLm. Distribution and ecology. Mediterranean, Azores,
slightly curved, with tapering points, 118-165 X ca. 3.5 fLm. Canary Islands. On rocks and in cavities, sublittoral.
Remarks. The single species known from the genus
Previous review Dendroxea, D. lenis, is so far only found in the Mediterranean,
Azores and Canary Islands. De Weerdt & Van Soest (1986) men-
De Weerdt, 2000. tioned the possibility that Reniera carmabi Van Soest (1980) could
also be a Dendroxea, but De Weerdt (2000) has shown that this
Description of type species species is a synonym of Chalinula molitba (de Laubenfels).
Despite the fact that the genus contains only one species, it has
Dendroxea lenis (Topsent, 1892b) (Fig. 6). such characteristic features, different from all the other chalinid
Synonymy. Reniera lenis Topsent, 1892b: XIX. Dendroxea genera and subgenera, that it is considered a valid genus.
len is; Griessinger, 1971: 152. Dendrectilla was erected by Pulitzer-Finali (1983: 593, fig. 81;
Material examined. Holotype (not examined): Laboratoire by monotypy) for Dendrectilla tremitensis Pulitzer-Finali, 1983. From
Arago de Banyuls-sur-Mer. Other material. NNM-CANCAP 5 his description it is clear that his species is conspecific with D. lenis.
stat. 5, Azores (cf. De Weerdt & Van Soest, 1986). ZMA Por. 5464-
Banyuls, France (cf. Van Lent & De Weerdt, 1987). HAliCLONA GRANT, 1836
Description. Thinly encrusting (Fig. 6A), overall size
ca. 3 X 1 X 0.5 cm. Surface smooth, even. Oscula indistinct. Synonymy
Consistency soft, somewhat viscous. Colour greyish-white. No
ectosomal skeleton. Choanosomal skeleton (Fig. 6C) consisting of Haliclona Grant, 1836. [Chalina] Grant, 1861 (nomen nudum).
branching, multi spicular tracts, rising up from a common, basal, Reniera Schmidt, 1862. Diplodemia Bowerbank, 1864. Gellius Gray,
Porifera· Demospongiae • Haplosclerida • Haplosclerina • Chalinidae 859

1867a. Orina Gray, 1867a. Asychis Gray, 1867a. Adocia Gray, 1867a. to firm, elastic, brittle or corky. Surface rather smooth and even,
Philotia Gray, 1867a. Veluspa Miklucho-Maclay, 1870. Euchalinopsis slightly to rather strongly punctate, sometimes slightly hispid.
Lendenfeld, 1887c. Rhaphisia Topsent, 1892b. Halichoclona de Oscula usually circular, flush with the surface or at the top of oscu-
Laubenfels, 1932. Toxadocia de Laubenfels, 1936a. Neoadocia lar mounds or chimneys. Ectosomal skeleton, if present, either a
de Laubenfels, 1950b. Reniclona de Laubenfels, 1954. Kallypilidion very regular, tangential, unispicular, isotropic reticulation, with
de Laubenfels, 1954. Rhizoniera Griessinger, 1971. three- to six-sided meshes, or less regular, subisotropic and rather
dense or consisting of a discontinuous, rather open reticulation due
Type species to many rounded meshes. Choanosomal skeleton a regular, fre-
quently ladder-like reticulation of uni-, pauci- or multi spicular pri-
Spongia oculata Pallas, 1766 (by original designation). mary lines, regularly connected by unispicular secondary lines; it
may also be a rather dense, subisotropic reticulation with many
Definition subdermal and choanosomal spaces. Spongin is nearly always
present, confined to the nodes of spicules or more abundant, some-
Chalinidae with unispicular secondary lines. times forming the dominant part of the skeleton. Megascleres
smooth diactines, oxeas or strongyles, length usually between
Diagnosis 80-250 f..I.,m, with a thickness of ca. 5-10 f..I.,m, rarely larger, up to
370 X 15 f..I.,m. Microscleres, if present, sigmas, toxas, raphides or
Cushion-shaped, frequently with oscular mounds or chim- microxeas.
neys, branching, tube-shaped, repent ramose, rarely thinly encrust-
ing sponges. Colour purple, violet, pink, brown, yellowish, green, Recent reviews
blue, black, sometimes white, orange or red. Some species show a
two-colour combination of a whitish ectosome and darker, purple Griessinger, 1971; Van Soest, 1980; De Weerdt, 1986, 1989,
or brownish choanosome. Consistency varying from soft, fragile, 2000.

KEY TO SUBGENERA OF HAliCLONA

(1) Choanosomal skeleton an anisotropic, ladder-like reticulation of uni- to paucispicular, ascending primary lines, regularly or less
regularly connected by unispicular secondary lines ............................................................................................................................ 2
Choanosomal skeleton an iso- or subisotropic reticulation without a clear distinction between primary and secondary lines ... ....... 4
(2) Skeleton very regular, with straight, uni- to paucispicular ascending primary lines, regularly connected by unispicular secondary
lines, spongin always present, at the nodes of the spicules or more copious, oxeas usually short, relatively thick, cigar-shaped,
microscleres, if present, toxas ............................................................................................................................ Haliclona (Haliclona)
Skeleton less regular, with pauci- to multi spicular primary lines, irregularly connected by unispicular secondary lines, with many
spicules in confusion ......... ................ ...... .......... ...... .................. ... .................................... ....... ........... ...... ....... ... ..... ...... ...... .......... ... .... 3
(3) Primary lines pauci- to multispicular, somewhat wavy, spongin scarce or absent, oxeas usually slender, with long, hastate points, no
microscleres ...................................................................................................................................................... Haliclona (Rhizoniera)
Skeleton towards the inner parts of the sponge becoming a confused, subhalichondroid reticulation, oxeas usually long, stout, with
hastate points, microscleres, if present, toxas, sigmas or raphides or a combination of these ............................... Haliclona (Gellius)
(4) Both the ectosomal and choanosomal skeleton show a tendency to form rounded meshes, choanosomal skeleton consisting of ill-
defined paucispicular primary lines, irregularly connected by unispicular secondary lines, with many choanosomal spaces, spongin
scarce to more abundant, oxeas slender, microscleres, if present, toxas or raphides .......................................... Haliclona (Soestella)
Both the ectosomal and choanosomal skeleton a regular, iso- subisotropic reticulation ...... ............................................................... 5
(5) Ectosomal and choanosomal skeleton a very regular, delicate, unispicular, isotropic reticulation, spongin at the nodes of the spicula,
oxeas slender, frequently blunt-pointed to strongylote, microscleres, if present, toxas and sigmas, sponges usually soft, compressible
but fragile .............................................................................................................................................................. Haliclona (Reniera)
Ectosomal and choanosomal skeleton forming a subisotropic, paucispicular reticulation, ectosome overlaying the choanosome very
loosely with many subectosomal spaces, no spongin, oxeas usually rather stout, microscleres, if present, sigmas, sponges usually
brittle, incompressible, fragile ...................................................................................................................... Haliclona (Halichoclona)

SUBGENUS GELUUS GRAY, 1867 Definition

Synonymy Chalinidae with a choanosomal skeleton consisting of a rather

confused, subhalichondroid reticulation of pauci- to multi spicular
Gellius Gray, 1867a: 538. Orina Gray, 1867a: 539. Asychis primary lines, irregularly connected by unispicular secondary
Gray, 1867a: 539. Rhaphisia Topsent, 1892b: XX. Sigmadocia de lines. Ectosomal skeleton, if present, either a regular, tangential,
Laubenfels, 1936a: 69. The 'angulata' group of De Weerdt (1986, unispicular, isotropic reticulation, or consisting of irregularly
1989). strewn, tangentially orientated spicules.

Type species Diagnosis

Isodictya jugosa Bowerbank, 1866 (original designation) Cushion-shaped or submassive sponges, sometimes with
(senior synonym of Renierafibulata Schmidt, 1862: 73). a tendency to form fistular outgrowths. Consistency soft, fragile,
860 Porifera· Demospongiae • HaploscIerida • HaploscIerina • Chalinidae

I
I

A
Fig. 7. Haliclona (Gellius) jibulata (Schmidt, 1862, as Reniera). A, habit, reproduced from Bowerbank (1874, as Isodictya jugosa) (scale 0.5 em).
B, oxeas, reproduced from Bowerbank (scale 50 !Lm). C, sigma, reproduced from Bowerbank (scale 5 !Lm). D, habit of preserved specimen ZMA Por. 6107
(scale I cm). E, oxea and sigma, reproduced from Schmidt (scale 50 !Lm). F, tangential view of ectosomal skeleton, reproduced from De Weerdt 1986
(scale 250 !Lm). G, perpendicular section of choanosomal skeleton made from the holotype of Isodictyajugosa (scale 300 !Lm). H, sigmas (scale 25 !Lm).

sometimes slightly brittle. Spongin scarce or absent. Oxeas com- a rather dense and irregular reticulation of paucispicular primary
monly robust needles of fairly large size, up to 370 X 15 f.1m. lines, irregularly connected by unispicular secondary lines, with
Microscleres are usually present, and may be toxas, sigmas or many spicules in confusion. Spongin scarce, at the nodes of the
raphides or a combination of these. spicules. Spicules: straight or slightly curved oxeas with a long and
sharp point, 160-270 X 5-11 f.1m (Fig. 7B); microscleres irregularly,
Recent review weakly curved sigmas (Fig. 7C, H), 9.6-37 X 1-2.4 f.1m. Distribution
and ecology. Mediterranean-Atlantic. In the infralittoral, to ca. 60m,
De Weerdt, 2000. on stones, conglomorates of shells, calcareous algae, sponges, etc.
Remarks. Haliclona (Gellius) jibulata is well characterized
Description of type species by the combination of its cushion-shaped fonn, pink colour,
strongly reticulated surface and irregularly, weakly curved sigmas.
Haliclona (Gellius)jibulata (Schmidt, 1862) (Fig. 7). The species is without doubt con specific with the later described
Synonymy (restricted). Reniera jibulata Schmidt, 1862: 73, lsodictya jugosa Bowerbank, 1866. Although Schmidt's original
pI. vn fig. 9. Gelliusjibulatus; Topsent, 1890a: 201. Gelliodesjibula- material could not be studied and his description and figures of the
tus; Babic, 1922: 268. lsodictya jugosa Bowerbank, 1866. Gellius spicules are rather uninfonnative (see Fig. 7E), the identity of the
jugosus; Gray, 1867a: 538. Halichondria couchii Bowerbank, 1874: species has been made clear by Topsent (1890a: 201; 1925c: 706,
203, pI. Lxxm figs 12-15. Amorphina couchii; Topsent, 1888: 145. pI. VIn fig. 6) and his material in the MNHN (for exact spicule
? Gellius pyrrhi Hanitsch, 1895: 211. ? Gellius arcticus Hentschel, sizes of this material see De Weerdt, 1986). The holotype of
1916: 12, fig. 2. Taxonomic decision for synonymy: De Weerdt, 2000. lsodictya jugosa, BMNH 1910.1.1.294 (Fig. 7A), is a small cush-
Material examined. Holotype: not seen. Other material. ion of rather finn consistency, growing on a piece of conglomer-
Specimens of Gellius jibulatus: MNHN DT. 2699 - Luc (one micro- ated bryozoans, calcareous algae and tube wonns. Oscula were not
scope slide containing a cross-section, det. Topsent, cf. Topsent, observed and there is no ectosomal skeleton. The choanosomal
1890a: 201). MNHN DT. 3204 - Mediterranean (fragment, det. skeleton is a subhalichondroid reticulation (Fig. 7E) of paucispicu-
E. Topsent). MNHN DT. 3069 - Saint-Jean-de-Luz (fragments, det. lar primary lines, irregularly connected by unispicular secondaries.
E. Topsent). Holotype of lsodictyajugosa: BMNH 1910.1.1.294- The oxeas (Fig. 7B) are straight or slightly curved, with long and
Shetland, figured specimen of Bowerbank, 1874, pI. L fig. 11 (des- sharp points, ca. 260 X 8.2 f.1m. Microscleres sigmas (Fig. 7C, H),
ignated as the type by Bowerbank on p. 128 of the same volume). irregularly curved; ca.--22 X 1.4 f.1m. It confonns in every respect to
Description. Cushion-shaped to somewhat laterally spreading Haliclona (Gellius) jibulata. For further synonymy of the species
sponges (Fig. 7A, D), occasionally with fistular processes. Size up to see De Weerdt (1986).
10 cm in diameter. Oscula not numerous, circular, 1-2 mm, flush with Orina was erected by Gray (1867a: 539, by original designa-
the surface. Surface smooth, strongly reticulated. Colour pink to dark tion) for Halichondria angulata Bowerbank, 1866: 233. The
pink. Ectosomal skeleton (Fig. 7F) an irregular and rather confused, holotype is BMNH 1910.1.1.173, Guernsey, figured specimen
somewhat subisotropic reticulation. Choanosomal skeleton (Fig. 7G) of Bowerbank, 1874, pI. XLI fig. 4, designated as the type by
Porifera· Demospongiae • Haplosclerida • Haplosclerina • Cbalinidae 861

D
Fig. 8. Haliclona (Gel/ius) angulata (Bowerbank, 1866 as Halichondria). A, holotype, reproduced from Bowerbank, 1874 (scale 1 em). B, oxeas, repro-
duced from Bowerbank, 1874 (scale 50 11m). C, toxa, reproduced from Bowerbank, 1874 (scale 25 11m). D, tangential view of ectosomal skeleton, repro-
duced from Bowerbank, 1874 (scale 100 11m). E, perpendicular section of choanosome made from the holotype (scale 300 11m). F, close-up of sigmas and
toxas made from the holotype (scale 50 11m).

Bowerbank on p. 101 in the same volume (Fig. 8A). It is an irregu- the Azores, 200 and 523 m, as Thrinacophora ? spissa Topsent
larly shaped, rather firm (in the dried state) clump of 2.5 X (1892a: 124, pI. VI fig. 12, pI. IX fig. 9), which he later transferred
1 X 1 cm. The ectosome is for the greater part lost, but is still pres- to Rhaphisia (Topsent, 1904: 233, pI. I fig. 8, pI. XVII fig. 18)
ent in a few places. No oscula were observed. The ectosomal skele- (Fig. 9A-E). Material of this species is in the MNHN represented
ton (Fig. 8D) is a regular, tangential reticulation of intercrossing by MNHN DT. 1051, one microscope slide containing a cross-
oxeas. The choanosomal skeleton (Fig. 8E) is a somewhat con- section. The skeletal architecture (Fig. 9F) is a somewhat con-
fused, subhalichondroid reticulation of paucispicular primary lines, fused, subhalichondroid reticulation formed by strongyles of
irregularly connected by single secondary spicules. The oxeas 300.3-785.4 X 15.4-30.8 /-Lm (Topsent mentioned strongyles in three
(Fig. 8B) are long, rather thin, with long and sharp points, size categories: 1000 X 40/-Lm, 400 X 15 /-Lm and 300-400 X 5-8
237.5-277-307.8 X 6-7.7-9.1 /-Lm. There are numerous sigmas /-Lm). In addition there are numerous trichodragmas (Topsent, 1904:
(not mentioned by Bowerbank) and toxas, both in the ectosome three size categories, viz., 11 0-160 X 10-12 fLm, 70 x 30 fLm, and
and choanosome. The sigmas (Fig. 8F) are very regular in shape 20-30 X 7-8 fLm) and rather sharply curved toxas of 115.5-207.1 x
and size, slightly centrangulated, strongly curved, 9.6-12-14.4 X 4.6-6.9/-Lm (Fig. 9G). The skeletal architecture of Topsent's
0.7-O.9-1.2/-Lm. The toxas (Fig. 8C, F) are sharply bent, with Rhaphisia conforms in every respect to the subgenus Gellius.
recurved apices, also very regular in shape and size, 43.2-55.6- Sigmadocia was erected by de Laubenfels (1936a: 69, by
63.6 X 1-1.2-1.4/-Lm. Haliclona (Gellius) angulata is a valid, original designation) for Gellius textapatina de Laubenfels, 1926:
Mediterranean-Atlantic species, entirely conforming to the subgenus 567. The holotype (USNM 21446, examined) is a thick crust,
Gellius. For further synonymy see De Weerdt (1986). rather firm, brittle, 5 X 3.5 X 0.6 cm, with a smooth, even surface,
Asychis was erected by Gray (1867a: 539, by original designa- and numerous circular and oval oscula, ca. 1 mm in diameter;
tion) for Reniera jibulata Schmidt (1862), which needs no further ectosome easily detachable. The ectosomal skeleton is a rather
discussion. confused, tangential reticulation. The choanosomal skeleton is
Rhaphisia was erected by Topsent (1892b: XX, by monotypy) a somewhat confused reticulation, but with tracts showing a ten-
for Rhaphisia laxa Topsent, 1892b. He established the genus for dency to form rounded meshes approaching those of Xestospongia
'Renierinae' without a regular skeleton, with oxeas as megascleres de Laubenfels (1932). The megascleres are robust, sharply-pointed
and trichodragmas as microscleres. The species is represented in oxeas, 415.8-446.8 X 19.3-25.4 /-Lm. Microscleres are sigmas of
the MNHN by MNHN DT. 300 (one microscope slide, labeled very variable size and shape, with sharply recurved apices, some of
Rhaphisia laxa, 'Type', Banyuls), and MNHN DT. 299 (two them flagellated, 53.9-123.2 X 2.3-6.2 /-Lm. It is with somewhat
microscope slides, Banyuls), but the material of these slides is too hesitation considered a synonym of Gellius.
scanty to enable a reliable redescription of the skeleton and The subgenus Gellius is not very common, and it may have
spicules. In another work Topsent (1892a) described a species from a preference for deeper water. There are only three shallow water
862 Porifera· Demospongiae • Haplosclerida • Haplosclerina • ChaIinidae

Fig. 9. Haliclona (Gellius) spissa (Topsent, 1892a, as Thrinacophora ?). A, habit, reproduced from Topsent (1 892a) (no size given). B, strongyle, repro-
duced from Topsent (1892a) (scale 100/Lm). C, trichodragmas, reproduced from Topsent (l892a) (scale 100 /Lm). D, habit, reproduced from Topsent
(1904, as Rhaphisia) (no size given). E,toxas, reproduced from Topsent (1904) (scale 50/Lm). F, perpendicular section of choanosome (from microscope
slide MNHN DT. 1051) (scale 300 /Lm). G, close-up of trichodragmas and toxas (scale 50/Lm).

species known from the North East Atlantic region: H. (G.) angulata orange choanosome. Consistency relatively crisp and brittle, only
Bowerbank, H. (G.) fibulata Schmidt, and H. (G.) rava Stephens, and slightly compressible. Spongin absent or very scarce, at the nodes of
only one in the Caribbean: H. (G). tenerrirrm Burton. It is presently the spicules. Ectosomal skeleton of the same structure as the
unknown how many species occur in the Pacific and Antarctic, but in choanosome, usually very loosely overlaying the choanosome, from
the deeper parts of the Caribbean it is represented by H. (G.) calcinea which it may be separated by extensive subectosomal spaces.
Burton and H. (G.) megasclera Lehnert & Van Soest (1996). Choanosomal skeleton a subisotropic, somewhat confused reticula-
tion, commonly intercepted by many choanosomal spaces.
MegascIeres usually acerate or hastate oxeas, reaching substantial
SUBGENUS HAUCHOCLONA DE LAUBENFELS, 1932 sizes in some species. Microscleres, if present, microxeas or sigmas.

Synonymy Previous review

Halichoclona de Laubenfels, 1932: 114. Neoadocia de De Weerdt, 2000.

Laubenfels, 1950b: 15. The 'jistulosa' group of De Weerdt (1986,
1989; De Weerdt et aI., 1991). Pellinella Thiele, 1905: 471. Description of type species

Type species Haliclona (Halichoclona) gellindra de Laubenfels, 1932

(Fig. 10).
Halichoclona gellindra de Laubenfels, 1932 (by monotypy). Synonymy. Halichoclona gellindra de Laubenfels, 1932:
114, fig. 68.
Definition Material examined. Holotype: USNM 22063 - Carmel,
California.
Chalinidae with a choanosomal skeleton consisting of Description (partly after de Laubenfels, 1932). The holo-
a subisotropic, somewhat confused reticulation, commonly inter- type consists of very small fragments of indefinite shape (Fig. lOA).
cepted by many choanosomal spaces. Ectosomal skeleton of the De Laubenfels (1932) described the species as: 'Encrusting sponges,
same structure as the choanosome, usually very loosely overlaying 2 to 4 mm thick, somewhat more than 4 cm2 in area.' Consistency
the choanosome, from which it may be separated by extensive fragile. Colour in life pale lavender, in spirit cream. Oscules few,
subectosomal spaces. Spongin absent or very scarce, at the nodes irregular in shape, about 1 mm in diameter, often with raised collars
of the spicules. MegascIeres usually acerate or hastate oxeas. nearly 1 mm high. Surface superficially smooth. Ectosomal skele-
MicroscIeres, if present, microxeas or sigmas. Sponges commonly ton ('specialization') a crust of tangentially placed oxeas; easily
relatively crisp and brittle, only slightly compressible. removed 'in flakes'. Choanosomal structure a very regular isodictyal
reticulation. The ectosomal skeleton is not present anymore, but this
Diagnosis may have disappeared upon preservation. The choanosomal skele-
ton (Fig. lOB) is a rather dense, confused, subisotropic reticulation;
Massive, tubiform, cushion-shaped or encrusting sponges, it is denser and less regular than described by de Laubenfels. No
with a strong tendency in several species to form fistular out- visible spongin. The oxeas are rather small, hastate, with very sharp,
growths. Oscula usually large, not numerous. Colour orange, pink, long points, 105-111.6-121.5 X 5.6-6.2-6.9 f.Lm. Distribution and
purplish or white; in some species there is a two-colour combination ecology. Known only from the type locality, Carmel, California,
of a whitish transparent ectosome and a darker coloured, purple or intertidal.
Porifera· Demospongiae • Haplosclerida • Haplosclerina • Chalinidae 863

Fig. 10. Haliclona (Halichoclona) gellindra (de Laubenfels, 1932, as Halichoclona). A, holotype (fragmented) (scale 0.5cm). B, perpendicular section of
choanosome made from holotype (scale 500 jJ.m). C, Haliclona (Halichoclona) jistulosa (Bowerbank, 1866), perpendicular section of choanosome of
preserved specimen ZMA Par. 5656.

Pellinella was erected by Thiele (1905: 471) for Pellinella slender, with long, rather sharp points, 129-135.7-144 X 6-6.2--6.9
conica Thiele, 1905 (by monotypy). From his description and f..Lm. Raphides, as mentioned by de Laubenfels, could not be detected.
especially drawings it is clear that this is a species of Haliclona
(Halichoclona). SUBGENUS HAliCLONA GRANT, 1836
Remarks. De Laubenfels described the choanosomal skele-
ton as a very regular, isodictyal structure, but the slide made from Synonymy
the holotype clearly shows a rather dense, subisotropic reticulation.
Though scanty, the information from de Laubenfels' description Haliclona Grant, 1836: 6. [Chafina] Grant, 1861 (nomen
and the holotype is just sufficient to get a picture of characters nudum). Diplodemia Bowerbank, 1864: 202. Adocia Gray, 1867a:
shared by several chalinid species, for which the name Pellina has 552. Veluspa Miklucho-Maclay, 1870: 5. Lessepsia Keller, 1883:
been applied by authors. This starts with Griessinger (1971), who 19. Euchafinopsis Lendenfeld, 1887c: 744. The 'oculata' group of
re-introduced the name Pellina Schmidt, 1870 for chalinid sponges De Weerdt (1986, 1989).
with a crisp, fragile consistency, large ectosomal spaces and a
dense, irregularly meshed choanosome. He placed Haliclona Type species
(Halichoclona) semitubulosa Lieberktihn (1859, as Spongia) in
this genus, together with the Mediterranean-Atlantic Haliclona Haliclona oculata (Pallas, 1766) (by original designation;
(Halichoclona) Jistulosa Bowerbank (1866, as lsodictya), Grant, 1836: 6).
Haliclona (H.) magna Vacelet (1969, as Reniera) and Haliclona
(H.) parietalis Topsent (1893c, as Reniera). Van Soest (1980) Definition (modified from De Weerdt, 2(00)
transferred Pellina to his new family Oceanapiidae and used the
genus for fistule bearing sponges with a simple spicule comple- Choanosomal skeleton consisting of a very regular, ladder-like
ment of small oxeote megascleres (cf. also Van Soest & Sass, reticulation of uni - to paucispicular primary lines, regularly con-
1981). De Weerdt 1986 reviewed the history of the genus Pellina nected by unispicular secondary lines. Ectosomal skeleton, if pres-
and concluded that it is a synonym of Halichondria Fleming, 1828. ent, a unispicular, tangential, isotropic reticulation; occasionally with
Halichoclona is the only available name for these crisp, fragile the oxeas 'intercrossing'. Oxeas short, rather robust, fusiform or with
sponges with a subisotropic skeleton. Though frequently devel- acerated points. Spongin moderate to abundant. No microscleres.
oped, fistules do not necessarily have to be present.
Neoadocia was erected by de Laubenfels (l950b: 15) for Diagnosis
Neoadocia mokuoloea de Laubenfels (1950b: 15, fig. 8). The holo-
type, USNM 22745, from Moku 0 Loe, Kaneohe Bay, Hawaii, is a Sponges branched, repent ramose or thinly encrusting.
cushion of 3 X2X 1 cm, without visible oscula, very soft and fragile. Consistency varying from soft, flexible, to firm, corky. Oscula in the
Both the ectosomal and choanosomal skeleton (Fig. 11A-B) are branched and ramose forms circular, small, but clearly visible; in the
subisotropic, entirely conforming to Halichoclona. The oxeas are branched form they may be arranged alongside the narrower sides of
864 Porifera· Demospongiae • Haplosclerida • Haplosclerina • Chalinidae

Fig.n. Haliclona (Halichoclona) mokuoloea (de Laubenfels, 1950b, as Neoadocia). A, tangential view of ectosomal skeleton made from holotype (scale
500 fLm). B, perpendicular section of choanosome (scale 500 fLm).

Fig. 12. Haliclona (Haliclona) oculata (Pallas, 1766 as Spongia). A, habit, reproduced from Bowerbank (1874, as Chalina). B, perpendicular section of
choanosome made from preserved specimen ZMA Por. 5557 (scale 250 fLm). C, oxeas, reproduced from Bowerbank (1874) (scale 25 fLm).

the branches; in encrusting sponges they are very small, with a slight Description of type species
tendency to be surrounded by star-shaped aquiferous canals.
Haliclona oculata (Pallas, 1766) (Fig. 12).
Previous review Synonymy (restricted). Spongia oculata Pallas, 1766: 390.
Haliclona oculata; Grant, 1841: 5. Halichondria oculata;
De Weerdt, 1986; 2000. Johnston, 1842: 94, pI. ill figs 1,2. Clwlina oculata; Bowerbank,
Porifera' Demospongiae • Haplosclerida • Haplosclerina • Chalinidae 865

1864: 208, pI. XIII fig. 262; 1874: 169, pI. LXVI figs 1-3. skeleton (Fig. 12B) a regular, ladder-like reticulation of uni- to
Chalinula oculata; Schmidt, 1870: 77. Euchalinopsis oculata; paucispicular primary lines, regularly connected by unispicular
Lendenfeld, 1887c: 744. Haliclona oculata tavaresi Arndt, 1941: secondary lines. Spongin moderate to abundant, especially towards
25. Spongia dichotoma Linnaeus, 1767: 1299. Halichondria the base of the sponge. Spicules: short, fat, fusiform oxeas with
cervicornis Johnston, 1842: 96, pI. VI figs 1, 2, pI. V figs 1, 2. short, sharp points (Fig. 12C), stylote and strongylote modifi-
Chalina cervicornis; Bowerbank, 1866: 364; 1874: 171, pI. LXVII cations rather common, 80-145 X 5-12 JJ.-m. Distribution and ecol-
figs 1-3. Diplodemia vesicula Bowerbank, 1864 pI. XIV fig. 273, ogy. Arctic-Boreal. Infralittoral, to ca. 100m, on rocky and sandy
pI. XXIII fig. 324, pI. XXXVI fig. 377; 1866: 357; 1874: 178, bottom, attached to stones, Mytilus, etc. The species can tolerate
pI. LXX figs 12-14. Isodictya pygmaea Bowerbank, 1866: 313; low salinity and turbid water with suspended silt.
1874: 141, pI. LVI figs 6-10. Chalina grantii Bowerbank 1866: Remarks. The large number of synonyms of Haliclona
375; 1874: 173, pI. LXVIII figs 6, 7. Pachychalina grantii; Ferrer- (Haliclona) oculata is easily explained by its variable growth form
Hernandez, 1922: 18. Veluspa polymorpha var. gracilis Miklucho- and common occurrence, and the tendency of earlier authors to
Maclay, 1870: 5, pI. I figs 1,2, pI. II figs 22, 3. Veluspa polymorpha describe every growth form as a variety or a new species. Contrary
var. digitata Miklucho-Maclay, 1870: 5, pI. I figs 3, 4. Veluspa to its variable growth form, the skeletal architecture and size and
polymorpha var. arctica Miklucho-Maclay, 1870: 5, pI. I fig. 11, shape of the oxeas is fairly consistent and characteristic, enabling
pI. II figs 22, 2. Chalina arbuscula Verrill, 1873: 742. Isodictya reliable identification of the species. For a detailed treatment of the
varians Bowerbank, 1874: 307, pI. LXXXVIII figs 1-6. Chalina synonyms one is referred to De Weerdt (1986); here only the type
flemingii (pars) Bowerbank, 1874: 357. Reniera gracilis; species of the nominal genera which are synonymous with the sub-
Dybowsky, 1880: 47, pI. III fig. 3, pI. IV fig. 16. Haliclona gra- genus Haliclona will be dealt with.
cilis; Koltun, 1959: 216, fig. 175. ? Pachychalina caulifera Diplodemia was erected (by monotypy) by Bowerbank (1864:
Vosmaer, 1882a: 33, pI. I fig. 14, pI. III figs. 64-66. Taxonomic 202) for Diplodemia vesicula Bowerbank. BMNH 1877.5.21.2082
decision for synonymy: De Weerdt, 1986. is the holotype, figured specimen of Bowerbank, 1874, pI. LXX
Materio:l examined. ? Holotype: BMNH 1841.1.13.46 - fig. 12, designated as the type on p. 178 of the same volume. See
two dried specimens, connected by a small rope, with a label (prob- also Bowerbank, 1864, pI. XIV fig. 273, pI. XXIII fig. 324 [not fig.
ably by Ridley): 'Spongia oculata, 106,4/3.45', and another label: 234 as erroneously printed], and pI. XXXVI fig. 377. Bowerbank
'Chalina oculata (Pallas), Type? (History to be investigated),. erected the genus for this species because of the combination of
Other material. BMNH 1847.9.7.2 - Chalina oculata, Johnston a fibrous skeleton and 'membranous and spiculous ovaries'. The
Collection (JC), Firth of Fourth. ZMA - several specimens identi- holotype consists of a small cluster of gemmules, attached to a
fied as Haliclona oculata. Halichondria cervicornis: BMNH 'valve of a pecten' (Fig. 13A), held together by spongin fibres of
1847.9.7.144 - figured specimen of Johnston, 1842, pI. V 26.4-43.2 JJ.-m thick, cored by oxeas of ca. 93.6-124.8 X 2.9-6 JJ.-m
fig. 2. Holotype of Diplodemia vesicula: BMNH 1877.5.21.2082- (Fig. 13B-D). The oxeas within the gemmules measure 100.8-
Shetlands, figured specimen of Bowerbank, 1874, pI. LXX 111.4-117.6 X 4.3-4.8-5.3 JJ.-m. The gemmules (Fig. BE-F) are
fig. 12. Isodictya pygmaea Bowerbank: BMNH 1932.1.5.10 - ca. 950 JJ.-m in diameter. There is no doubt that the species belongs
Scarborough, figured specimens of Bowerbank, 1874, pI. LVI to Haliclona (Haliclona) oculata, in which the development of
figs 6-8 (the specimen of his fig. 6 designated lectotype by De gemmules is a common feature.
Weerdt, 1986). Holotype of Chalina grantii Bowerbank: BMNH Adocia was erected by Gray (1867a: 552, original designa-
1932.1.5.12 - figured specimen of Bowerbank, 1874, pI. LXVIII tion) for Isodictya simulans Bowerbank (1866: 308). There are sev-
fig. 6. Microscope slide of Chalina arbuscula Verrill: BMNH eral species described by Bowerbank and other authors which are
25.11.1.775a- Dendy Collection, Wood's Hole Museum.lsodictya synonymous with Haliclona (Haliclona) simulans, for which one
varians Bowerbank: BMNH 1877.5.21.2083 - Mersey. Chalina is referred to De Weerdt (1986). The species was first described by
flemingii Bowerbank: BMNH 1877.5.21.2079 - Peterhead, Johnston (1842: 109, pI. VIII figs 1-6) as Halichondria simulans,
described by Bowerbank, 1874: 357. Chalina oculata: BMNH initially transferred by Bowerbank (1861: 71) to Haliclona, then
1847.9.7.144, BMNH 1847.9.7.2 - Johnston Collection, Firth of to Chalina (Bowerbank, 1864: 101), and then to Isodictya
Fourth. Halichondria cervicornis: figured specimen of Johnston, (Bowerbank, 1866: 101). Johnston's specimen of his plate VIII fig.
1842, pI. V fig. 1. Chalina cervicornis: BMNH 1877.5.21.2080- 5 is catalogued as BMNH 1847.9.7.9, and designated lectotype by
Bowerbank Collection, Torquay, figured specimen of Bowerbank, De Weerdt (1986: 98). Isodictya simulans is in the BMNH repre-
1874, pI. LXVII fig. 1. sented by several specimens. BMNH 1932.1.5.2a, from Hastings,
Description. Solid branches, growing from a common stalk is the specimen figured by Bowerbank (1874, pI. LI fig. 5), desig-
which is attached to the substratum with a pedicel (Fig. 12A). nated as 'genolectotype' of Adocia by Burton, 1934a: 534
There is a considerable variation in the number and degree of fus- (Fig. 14A-B). He kept Adocia for species with a dermal skeleton.
ing of the branches. The branches may be thin and delicate (diame- It has, in the dried state, a light yellowish-brown colour and fragile,
ter 0.2-0.4 cm), entirely isolated from each other, but they may also though somewhat corky consistency. The sections made from this
grow much thicker and broader, and fused to such a degree that the specimen show a skeleton clearly damaged by the handling;
sponge becomes almost flabelliform. Usually the branches are the ectosomal skeleton is a somewhat subisotropic, tangential,
somewhat laterally compressed. The oscula are always small and basically unispicular reticulation with intercrossing oxea; the
circular, 1-3 mm, mainly arranged along the narrower sides of the choanosome consists of paucispicular primary lines, irregularly
branches. Height of the sponge up to 30 cm. Surface smooth, vel- connected by unispicular secondaries. There is a moderate to con-
vety, slightly hispid. Consistency rather soft, but elastic, compress- siderable amount of spongin at the nodes of the spicula. The oxeas
ible, not fragile. Colour light brown. Skeleton: ectosomal skeleton are short, fat, cigar-shaped, with rather short and sharp points,
usually absent, occasionally consisting of an irregular, incoherent 108-119.7-141.6 X 5-6.6-9.1 JJ.-m. See Fig. 14C for a drawing of
reticulation of some tangentially orientated spicula. Choanosomal the choanosomal skeleton, and Fig. 14D for the ectosomal skeleton
866 Porifera· Demospongiae • Haplosclerida • Haplosclerina • ChaIinidae

I'II/
E F-

B
Fig. 13. Haliclona (Haliclona) oculata (Pallas, 1766, as Spongia). A, gemmules attached to the 'valve of a pecten', reproduced from Bowerbank (1874,
as Diplodemia vesicula) (scale I cm). B, spongin fibres cored by oxeas, reproduced from Bowerbank, 1864 (scale 100 /lorn). C, spongin fibres, microscope
slide made from the holotype of Diplodemia vesicula (scale 100 /lorn). D, oxeas reproduced from Bowerbank, 1874 (scale 50/lom). E, gemmules,
reproduced from Bowerbank, 1864 (scale 100/lom). F, gemmule microscope slide made from the holotype of Diplodemia vesicula (scale 100 /lorn).

D
Fig. 14. Haliclona (Haliclona) simulans (Johnston, as Halichondria). A, habit, reproduced from Bowerbank (1874, as Isodictya) (scale 1.5 em). B, oxeas,
reproduced from Bowerbank (scale 50/lom). C, perpendicular section of choanosornal skeleton made from preserved specimen in ZMA (scale 500/lom).
D, tangential view of ectosomal skeleton (scale 500 /lorn) (after De Weerdt, 1986).

of a preserved ZMA specimen of Haliclona (Haliclona) simulans. Miklucho-Maclay (1870: 5, figs 1-2) (Fig. 15). Although the orig-
The conspecifity of Johnston's and Bowerbank's simulans is obvi- inal material has not been studied, the description and figures leave
ous. The presence of a distinct ectosomal skeleton is considered no doubt about the con specificity of this species with Haliclona
a primitive character within the Haplosclerida (see De Weerdt, (Haliclona) oculata.
1989), therefore there is no reason to keep Adocia separate from Lessepsia was erected (by monotypy) by Keller (1883: 19) for
Haliclona. Lessepsia violacea Keller (1882: 19, pi. I figs 1-2). The holotype
Veluspa was erected (by original designation) by of L. violacea could not be examined (it is not present in the
Miklucho-Maclay (1870) for Veluspa polymorpha var. gracilis Museum flir Naturkunde der Humboldt-Universitat, Berlin), but
Porifera· Demospongiae • Haplosclerida • Haplosclerina • Chalinidae 867

Keller gave a good description of the species. He described it as Type species

a very delicate, cushion-shaped sponge, with one large osculum of
5 mm diameter, and a few smaller ones, with a vivid violet colour, Reniera aquaeductus Schmidt, 1862: 73 (by subsequent
growing between shells of the mussel Mytilus variabilis. The skele- designation; de Laubenfels, 1932: 61).
ton a delicate, irregular reticulation with three to four sided
meshes, with oxeas completely embedded in spongin. Primary Definition
lines uni- to paucispicular, with only very few spicula scattered in
between the lines. The oxeas with both ends pointed or rounded, Chalinidae with a choanosoma1 skeleton consisting of a deli-
100--120 /Lm. Although Keller described the skeleton as 'irregular', cate, regular, unispicular, isotropic reticulation. Ectosomal skele-
his figure (I.c., fig. 2) shows a rather regular structure, conforming ton, if present, also a tangential, unispicular, isotropic, very regular
to the subgenus Haliclona. Keller considered the species and new and continuous reticulation. Spongin always present at the nodes of
genus to be placed between 'Chalinula' and 'Reniera'. It is here spicules, but never abundant. Oxeas frequently blunt-pointed or
placed in the synonymy of the subgenus Haliclona. strongylote. Microscleres, if present, toxas and sigmas. Sponges
Euchalinopsis was erected by Lendenfeld (1887c: 744), for commonly soft and fragile.
Chalina oculata (Bowerbank, 1864), subsequently designated as
type species by Burton (1934a: 529). Diagnosis
Toxiclona de Laubenfels (1954), type species Siphonochalina
gaussiana Hentschel (1914), was first considered by De Weerdt Delicate sponges with a cushion-shaped or encrusting, later-
(2000) to be a synonym of the subgenus Haliclona. This viewpoint ally spreading growth form, also masses of anastomosing slender
cannot be upheld anymore, and the species is now transferred to branches intermittently and loosely attached to the substratum, or
Cladocroce (see above). clusters of tubes; some species have a tendency to form long, thin,
blind-ending proliferations. Oscula at the end of oscular chimneys
or mounds, or flush with the surface. Consistency varying from soft,
SUBGENUS RENIERA SCHMIDT, 1862 compressible but fragile to moderately firm. Several species pro-
duce slime strands when tom apart. Colour commonly bright, vary-
Synonymy ing from purple, violet to orange and yellow. Surface smooth, even.
Ectosomal skeleton, if present, a regular, tangential, unispicular,
Reniera Schmidt, 1862: 73. Prianos Gray, l867a: 520. isotropic, reticulation. Choanosomal skeleton also a delicate, regu-
Philotia Gray, 1867a: 522. Toxadocia de Laubenfels, 1936a: 69. lar, unispicular, isotropic reticulation. Spongin always present at the
Reniclona de Laubenfels, 1954: 64. Kallypilidion de Laubenfels, nodes of the spicula, but never abundant. Oxeas frequently blunt-
1954: 110. The 'aquaeductus' group of De Weerdt (1986, 1989; De pointed or strongylote. Microscleres, if present, toxas and sigmas.
Weerdt et al., 1991).
Previous reviews

Griessinger, 1971; Van Soest, 1980; De Weerdt, 1989,2000.

Description of type species

Haliclona (Reniera) aquaeductus Schmidt, 1862 (Fig. 16).

Synonymy. Reniera aquaeductus Schmidt, 1862: 73, pI. VII
figs 6, 6a, 6b; Griessinger, 1971: 126, fig. 4a; Boury-Esnault &
Lopes, 1986: 199, fig. 48, pI. ill, 12.
Material examined. Holotype: Not seen. Topotypical speci-
men: BMNH 1867.7.26.47a (dry) - one of Schmidt's specimens of
Reniera aquaeductus from the type locality, Sebenico, Adriatic.
Description (partly after Boury-Esnault & Lopes, 1986).
Laterally spreading masses up to 15 cm in diameter and 0.5-1.5 cm
thick, from which arise fistule-like outgrowths of 5-7 mm high and
other elevations. Oscules at the top of these elevations, with
a diameter of 1-3 mm (Fig. 16A). Colour light purple or pink, con-
sistency soft, very fragile. Surface smooth and even. Ectosomal
skeleton (Fig. 16C) a very regular, tangential, unispicular and
isotropic reticulation. Choanosomal skeleton (Fig. 16D) also a
very regular, delicate, unispicular, isotropic reticulation, without
clear distinction between primary and secondary lines. Oxeas
(Fig. 16B) blunt-pointed or strongylote, ca. 145-175 X 6-8/Lm.
Spongin moderate, at the nodes of the spicula. Distribution and
Fig. 15. Haliclona (Haliclona) oculata (Pallas, 1766). Habit, repro-
ecology. Mediterranean, Azores, on hard substratum, 5-35 m.
duced from Mikluco-Maclay (1870 as Veluspa polymorpha var. gracilis) Remarks. Prianos was erected by Gray (1867a: 520, origi-
(scale 2cm). nal designation) for Reniera amorpha Schmidt, 1864: 38. Holotype
868 Porifera· Demospongiae • Haplosclerida • Haplosclerina • Chalinidae

Fig. 16. Haliclona (Reniera) aquaeductus (Schmidt, 1862, as Reniera). A, habit, reproduced from Schmidt (1862) (scale I em). B, oxeas, reproduced from
Schmidt (scale 50 fLm). C, tangential view of ectosomal skeleton made from specimen in BMNH identified by Schmidt (scale 250 fLm). D, perpendicular
section of choanosomal skeleton made from specimen in BMNH identified by Schmidt (scale 250 fLm).

LMJG 15628 (not examined). Topsent (1925c: 710) considered it specimens in the BMNH identified by Bowerbank as I. varians
a junior synonym of Reniera cratera Schmidt, 1862: 73, which is all belong to Haliclona (Haliclona) oculata (Pallas) (e.g.,
followed here. BMNH 1019.1.1.349, figured specimen of Bowerbank, 1874,
Philotia was erected by Gray (1867a: 522, by original desig- pI. LXXXVIII, fig. 1). See Figs 17D-E for slides of Johnston's
nation) for Isodictya varians Bowerbank, 1866: 281. This species specimens of Halichondria cinerea marked '17e' .
was first established for a single specimen, illustrated in Toxadocia was erected by de Laubenfels (1936a: 69, by origi-
Bowerbank, 1874, pI. XLVIII figs 14-16 and designated as type on nal designation) for Gellius abbreviatus Topsent (1918: 538, San
p. 124 of the same volume. This specimen forms part of four spec- Thome). It was described as 'very small specimens' of 10 X 6 X
imens in the BMNH, identified by Johnston as Halichondria 1-2 mm, white, with a loose structure, slightly hispid, without dis-
cinerea (cf. Johnston, 1842: 111, pI. IV figs 3 and 4), all dried and tinct oscules. Skeleton of a simple structure, forming multi spicular
mounted on a piece of carton. They have the numbers BMNH lines towards the periphery, with rather abundant spongin at the
1847.9.7.41 (Johnston's card '17d'), 1847.9.7.42 (Johnston's card nodes of the spicules. Megascleres slightly curved strongyles,
'17e'); 1847.9.7.71 (left, larger specimen on Johnston's card '17c') 100-110 X 5.5-6 /-Lm. Microscleres toxas of variable size, the
and 1847.9.7.156 (right, smaller specimen on Johnston's card larger ones 56-70 X 1.7-2.4 /-Lm, the smaller ones 20 X 0.8 /-Lm,
'17c)'. Of these, only the sponge on card '17d' was considered by with intermediates. Topsent's material is represented in the MNHN
Bowerbank to belong to 'cinerea' (ef. Bowerbank, 1866: 274--279), by two microscope slides, MNHN DT. 2057, labeled: "Iype, Praia
the others were re-identified by him as Isodictya varians (1847. das Conchas, 3.8.1906, Det. E.T., Descr. E.T., Arch. Zool. 1918,
9.7.42), I. peachii (1847.9.7.71), and I. permollis (1847.9.7.156) 57, p. 538-540', one of which contains a cross-section. The skele-
respectively. Having reexamined these specimens there is no doubt ton is a very regular, unispicular, isotropic reticulation (Fig. 18A);
that they all belong to Haliclona (Reniera) cinerea (Grant, 1826d) multi spicular lines could not be detected, and the spongin is less
(cf. De Weerdt, 1986 and De Weerdt & Stone, in De Weerdt, 1987). abundant than mentioned by Topsent. The strongyles (Fig. 18B)
The holotype of I. varians is a light greyish-brown, fragile cushion measure 96-117 X 4.5-6 /-Lm; the toxas (Fig. 18C), abundant,
of 1 X 2.5 X 1 cm (Fig. 17A), without visible oscula, with a smooth measure 24-66.9 X 0.9-2.4 /-Lm. As already mentioned by Topsent,
surface. The ectosomal skeleton is a very regular, tangential, the species is very similar to the Arctic Haliclona (Reniera)
unispicular, isotropic reticulation. The choanosomal skeleton primitiva (Lundbeck, 1902: 69, pI. XIII fig. 11, as Gellius; ef. De
(Fig. 17C) has the same structure, very regular and isotropic. There Weerdt, 1986). Both species have the regular, unispicular, isotropic
is a cons'iderable amount of spongin at the nodes of the oxeas. skeletal architecture typical of the subgenus Reniera.
These are short, rather fat, cigar-shaped, with many stylote modi- Reniclona was erected by de Laubenfels (1954: 64, by origi-
fications (Fig. 17B), 76.8-100.3-112.8 X 4.8-7.6-10.3 /-Lm. The nal designation) for Isodictya permollis Bowerbank, 1866, which
species clearly belongs to Haliclona (Reniera) cinerea. The other is ajunior synonym of Haliclona (Reniera) cinerea (Grant, 1826d).
Porifera· Demospongiae • Haplosclerida • Haplosclerina • Chalinidae 869

Fig. 17. Haliclona (Reniera) cinerea (Grant, 1826d, as Spongia). A, habit, reproduced from Bowerbank (1874, holotype Isodictya varians) (scale 0.5 em).
B, oxeas, reproduced from Bowerbank (scale 25 ILm). C, perpendicular section of choanosomal skeleton made from holotype of l. varians (scale 250 ILm).
D, perpendicular section of choanosomal skeleton made from Johnston's specimen' l7e' (as Halichondria cinerea) (scale 250 ILm). E, tangential view of
ectosomal skeleton made from Johnston's specimen' 17e' (scale 250 ILm).

Fig. 18. Haliclona (Reniera) abbreviata (Topsent, 1918 as Gellius). A, perpendicular section of choanosomal skeleton made from Topsent's slide
(scale 50 ILm). B, toxas made from Topsent's slide (scale 50 ILm).

The lectotype of I. permollis (designated by De Weerdt & Stone, Laubenfels at Ponape (USNM 23035 and 23060) and identified
in De Weerdt, 1987) is BMNH 1932.1.5.11, Scarborough, the by him as Reniclona permollis (de Laubenfels, 1954: 67) still
specimen figured by Bowerbank, 1874, pI. XLVIII fig. 9. It is a flat needs to be established, but they certainly do not belong to the
crust of 5 X 3.5 X 0.2 cm (Fig. 19A), growing on a stone, of a very eastern Atlantic Haliclona (Reniera) cinerea (cf. De Weerdt,
fragile consistency, light grey with yellowish tinges, with about 1986, 1989).
five circular oscules flush with the surface, and a diameter of 1-2 Kallypilidion was established by de Laubenfels (1954: 110,
mm. There is a clearly visible ectosomal skeleton. The ectosomal by monotypy) for Kallypilidion poseidon de Laubenfels (1954:
skeleton (Fig. 19C) is a very regular, tangential, unispicular, 110, fig. 70, pI. IX fig. a; Palau Islands), which is a junior synonym
isotropic reticulation. The choanosomal skeleton (Fig. 19D) is also of Siphonochalinajascigera Hentschel (1912: 398, pI. XVI fig. 3)
a very regular, unispicular, isotropic reticulation. There is some (Fig. 20A-B). De Laubenfels erected the genus for the single
spongin at the nodes of the spicules. The oxeas (Fig. 19B) are species on basis of the external shape: 'It is characterized sharply
slightly curved, with acerated, short points, 100.8-112.2-122.4 X by its external shape, which is regularly that of a very thin-walled
5-6.2-7.7 f.Lm. The identity of the sponges collected by de bowl or vase. It is easily detached from the substratum and, when
870 Porifera· Demospongiae • Haplosclerida • Haplosclerina • Chalinidae

Fig. 19. Haliclona (Reniera) cinerea (Grant, 1826d, as Spongia). A, habit, reproduced from Bowerbank (1874, as lsodictya permollis) (scale 2cm).
B, oxeas, reproduced from Bowerbank (scale 50 J.Lm). C, tangential view of ectosomal skeleton made from Bowerbank's specimen (scale 250 J.Lm).
D, perpendicular section of choanosomal skeleton made from Bowerbank's specimen (scale 250 J.Lm).

Fig. 20. Haliclona (Reniera)fascigera (Hentschel, 1912 as Siphonochalina). A, habit of preserved specimen, ZMA Por. 15268 (scale 2cm). B, spongin
fibre cored by oxeas, reproduced from Hentschel (no size given). C, tangential view of ectosomal skeleton made from holotype of Kallypilidion poseidon
de Laubenfels (1954) (scale 250 J.Lm). D, perpendicular section of choanosomal skeleton made from holotype of Kallypilidion poseidon.
Porifera' Demospongiae • Haplosclerida • Haplosclerina • Chalinidae 871

turned upside down, strikingly resembles a brimless cap, such as Previous review
a man might wear. The name is derived from the Greek for 'beautiful
hat', or 'cap' (which was possibly used as a head gear by Poseidon; De Weerdt, 2000.
de Laubenfels, 1954: 111). He described the colour as a vivid light
blue, very slightly verging toward violet. Consistency flexible, Description of type species
'like that of wet paper.' Surface mostly very smooth, with here
and there a few lumps. The ectosomal skeleton a tangential, iso- Haliclona (Rhizoniera) rhizophora Vacelet, 1969 (Fig. 21).
dictyal reticulation, 'as characteristic of the Adociidae', and the Synonymy. Reniera rhizophora Vacelet, 1969: 212, fig. 53,
endosome 'also strikingly isodictyal in nature'. The oxeas united pI. IV fig. 3; Rhizoniera rhizophora; Griessinger, 1971: 152, fig. 21.
by spongin, measuring 64-77 X 2-3.5/-lm. In addition to the basal Material examined. Holotype: MNHN JV 68-13 -
isodictyal reticulation there are tracts of fibres containing a moder- Mediterranean.
ate amount of spongin. The holotype, USNM 23121 has been Description (partly after Vacelet, 1969). Club-shaped
examined and consists of a tube, 6 cm high, 2 cm thick at the base, sponges (Fig. 21A), with a stalk attached to the substratum, to 8 cm
gradually increasing to 2.6 cm at the top. The top part of the tube is high; main body 2-4 cm high, 1.5-2.5 cm thick. Colour grey in
recurved. Consistency soft, surface smooth, punctate. The ectoso- spirit. Consistency rather soft, stalk not very rigid. Oscula apical,
mal skeleton (Fig. 20C) a very regular, tangential, unispicular, 2-3 rum in diameter. No ectosomal skeleton. Choanosomal skele-
isotropic reticulation. The choanosomal skeleton (Fig. 20D) also a ton (Fig. 21B) a somewhat irregular reticulation of paucispicular
regular, unispicular, reticulation. A moderate amount of spongin primary lines, irregularly and loosely connected by unispicular sec-
unites the spicules at the nodes. The oxeas are slender, slightly ondary lines, with many loose spicules scattered in between.
curved, somewhat fusiform, 72-77.9-81.9 X 3-3.~.2/-lm. In Throughout the skeleton run multi spicular tracts, arising from the
all respects the species fits within the subgenus Reniera. The stalk, 30-85 /-lm in diameter. Spongin very scarce, at the nodes
original material of Siphonochalina jascigera Hentschel (1912, of the spicula. Oxeas slender, hastate, 220-330 X 5-7.5/-lm.
Banda Sea) could not be studied, but Hentschel's description Distribution and ecology. Mediterranean, 130-150 m.
(see also Fig. 20B) leaves no doubt about the conspecifity of his Remarks. The type species differs from all other species
species with Kallypilidion poseidon. The ZMA holds several spec- assigned to Rhizoniera by the possession of a stalk and the rein-
imens of Haliclona (Reniera) jascigera from Indonesia, and forcement throughout the sponge of extensions of the multi spicular
Hooper (pers. corum.) reports that the species is commonly tracts originating from the stalk. Because of this, the regular
observed in many of the western Pacific island and Great Barrier anisotropic skeleton of pauci- to multi spicular primary lines con-
Reef faunas. nected by unispicular secondaries, as found in all other members
assigned to the subgenus, is somewhat obscured in the type
species. See for comparison Fig. 21C, which is the choanosomal
skeleton of Haliclona (R.) rosea (Bowerbank, 1866, as Isodictya).
SUBGENUS RHIZONIERA GRIESSINGER, 1971 Presently, it does not seem appropriate to keep Rhizoniera as
a separate genus or subgenus, or to erect a new subgenus for
Synonymy species with a better developed anisotropic skeleton, because
R. rhizophora has a stalk and occurs in deeper water. Though
Rhizoniera Griessinger, 1971: 152. The 'rosea' group of perhaps not the most appropriate, Rhizoniera is, however, the only
De Weerdt (1986, 1989). available name for species with this type of skeleton.

Type species
SUBGENUS SOESTELLA DE WEERDT, 2000
Reniera rhizophora Vacelet, 1969 (by monotypy).
Synonymy
Definition
Haliclona (Soestella) De Weerdt, 2000: 7.
Chalinidae with a regular anisotropic, ladder-like choanoso-
mal skeleton consisting of pauci- to multispicular ascending pri- Type species
mary lines, connected by unispicular secondary lines. Ectosomal
skeleton usually absent, if present, consisting only of some Reniera mamillata Griessinger, 1971 (by original designation).
vaguely strewn tangentially orientated oxeas. Spongin scarce or
absent. Definition

Diagnosis Chalinidae with a subanisotropic choanosomal skeleton con-

sisting of ill-defined paucispicular primary lines, irregularly con-
Sponges thickly encrusting, cushion-shaped with oscular nected by paucispicular secondary lines. There is a slight but
chimneys or mounds, or massive, rarely stalked. Consistency soft consistent tendency of the spicules to form rounded meshes.
to moderately firm, sometimes viscous. Surface frequently slightly Ectosomal skeleton a discontinuous, tangential, rather open reticu-
hispid through projecting spicules of the primary lines. Colour lation, due to many rounded meshes framed by spicules in lines of
brown, pink, purple or bluish-grey. Megasc1eres usually slender 2-5 spicules thick. Spongin always present at the nodes of spicules,
oxeas with acerated points. No microsc1eres. Consistency soft to but never abundant. Oxeas usually slender. Microsc1eres, if present,
moderately firm. sigmas, toxas or raphides.
872 Porifera' Demospongiae • Haplosclerida • Haplosclerina • Chalinidae

A
c
Fig. 21. Haliclona (Rhizoniera) rhizophora (Vacelet, 1969 as Reniera). A, habit, reproduced from Griessinger (1971, as Rhizoniera) (scale 1 cm). B, per-
pendicular section of choanosomal skeleton made from the holotype (scale 500 ILm). C, perpendicular section of choanosomal skeleton of preserved spec-
imen of Haliclona (Rhizoniera) rosea (Bowerbank, 1866 as [sodictya), ZMA Por. 4182 (scale 500 ILm).

Diagnosis Material examined. Holotype: MNHN DJMG 4 -

Dubrovnik, Adriatic. Other material. ZMA Por 5460 - Cap
Growth form irregularly massive, lhickly encrusting, l' Abeille, 32 m, Banyuls.
lobate/cushion-shaped, or digitate. Oscula at the end of oscular DeSCription (after Griessinger, 1971 and Van Lent &
chimneys or mounds, alongside the branches of digitate forms, De Weerdt, 1987). Massive base (25 mm thick) from which arise
flush with lhe surface in encrusting forms. Consistency varying differently shaped (mammiform, vulcano-shaped) osculiferous
from soft to moderately firm. Surface even or somewhat irregular, elevations, up to 7 cm high (Fig. 22A-B). Colour pink-orange,
slightly to rather strongly punctate. Colour varying from light sometimes with yellowish tinges at lhe base. Consistency moder-
to dark brown, black, green to dark red. Ectosomal skeleton a ately soft, compressible, somewhat viscous. The ectosomal
discontinuous, tangential, rather open reticulation, due to many skeleton (Fig. 22C, F) is a rather irregular, tangential, discontinu-
rounded meshes framed by spicule lines of 2-5 spicules lhick. ous reticulation of spicules forming rounded meshes. The
Choanosomal skeleton a subanisotropic reticulation consisting of choanosomal skeleton (Fig. 22E) also a somewhat irregular reticu-
ill-defined paucispicular primary lines, irregulary connected by lation of ill-defined primary lines, irregularly connected by
paucispicular secondary lines. Spongin always present at the unispicular secondary lines, wilh a vague tendency to form rounded
nodes of spicules, but never abundant. Oxeas usually slender. meshes. Spongin scarce, at the nodes of the spicUla. Oxeas
Microscleres, if present, sigmas, toxas or raphides. (Fig. 22D) wilh tapering, ralher short points, 105-180 X 2.5-6.8 fLm.
Remarks. The subgenus Soestella was erected by
Previous review De Weerdt (2000) for a group of sponges named 'du type Reniera
arenata' by Griessinger (1971: 130). All the species assigned
De Weerdt, 2000. by Griessinger to this group, as well as most species of the group
'du type R. fulva' show a high resemblance in skeletal architecture,
Description of type species conforming to the above given definition of Soestella. Van Soest
(1980: 12) based his redefinition of 'Reniera' on this group of
Haliclona (Soestella) mammilata Griessinger, 1971 (Fig. 22). sponges, but without having studied the type species of Reniera,
Synonymy. Reniera mammilata Griessinger, 1971: 132, Reniera aquaeductus Schmidt (1862). This species, wilh its
figs 5b, 6c, j, pI. I fig. 3; Haliclona mamillata; Van Lent & entirely unispicular, isotropic skeleton, is clearly different from lhe
De Weerdt, 1987: 135. above mentioned group of species.
Porifera· Demospongiae • Haplosclerida • Haplosclerina • Chalinidae 873

Fig. 22. Haliclona (Soestella) mamillata (Griessinger, 1971 as Reniera). A, habit of preserved specimen, ZMA Por. 5460 (scale I cm). B, different shaped
osculiferous elevations, reproduced from Griessinger. C, tangential view of ectosomal skeleton, reproduced from Griessinger (scale 250 fLm). D, oxeas,
reproduced from Griessinger (scale 50 fLm). E, perpendicular section of choanosomal skeleton made from holotype (scale 500 fLm). F, tangential view of
ectosomal skeleton made from holotype (scale 500 fLm).

GENUS INCERTAE SEDIS 'short with acute endings' . Together this conveys an image of
a Haliclona-type of sponge. Three other species of Protoschmidtia
The genus Protoschmidtia Czerniavsky, 1878: 392 (1879: 95; were described in the same publication, and of only one
1880: 30) was erected for Protoschmidtia simplex Czerniavsky, (Protoschmidtia (sic)foraminosa Czerniavsky, 1878: 393 (1879: 98;
1878: 392 (1879: 95; 1880: 31) (here designated type species), 1880: 33), pI. I (=pl. V) figs 6--7) data on spicule size (140-210 X
from the northern shores of the Black Sea, from the midshore. The 7-12 fLm) and shape are provided. This is consistent with an
description in Latin is not very informative. The shape is encrust- assumed synonymy with Haliclona. Several other authors (Ridley,
ing with tubular elevations, and was compared with 'Reniera 1884a; Thiele, 1903; Burton, 1928a) employed Protoschmidtia
palmata' and 'Amorphina grossa'. Spicules are described as mostly for species belonging to the order Haplosc1erida.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Nipbatidae Van Soest, 1980

Ruth Desqueyroux-Faundez1 & Clare Valentine2

1 Museum d'histoire naturelle, P.O. Box 6434, 1211 Geneva 6, Switzerland. (ruth.faundez@mhn.ville-ge.ch)
2 The Natural History Museum, Cromwell Road, London SW7 5BD, UK. (c.valentine@nhm.ac.uk)

Niphatidae Van Soest (Demospongiae, Haplosclerida) contains nine valid genera and a number of nominal species with a worldwide dis-
tribution. Species live principally in shallow tropical waters, although species of Amphimedon and Pachychalina are known from Chile,
Peru and Magellan regions, and Gelliodes, Microxina and Haliclonissa are also represented in Antarctica. Major taxonomic characters for
differentiating genera include: the type of ectosomal skeleton, which is formed by brushes of spicules, sometimes complemented by
a fibrous network; morphology of choanosomal fibres, longitudinal, parallel-radially plumose or divergent, often fasciculated or largely
ramified; presence of interstitial spicules in some species to form a free network and spicule micrometries, usually larger than in
Callyspongiidae, in general, between 100--600 f.Lm length.
Keywords: Porifera; Demospongiae; Haplosclerida; Niphatidae; Amphimedon; Cribrochalina; Dasychalina; Gelliodes; Haliclonissa;
Hemigellius; Microxina; Niphates; Pachychalina.

DEFINITION, DIAGNOSIS, SCOPE Scope

Synonymy Twelve nominal genera and possibly more than 100 species
are included in this family worldwide. Of the nominal genera nine
Niphatidae Van Soest, 1980: 25. are considered to be valid genera (taxonomic decision, this work):
Amphimedon, Cribrochalina, Dasychalina, Gelliodes, Haliclonissa,
Definition Microxina, Hemigellius, Niphates and Pachychalina.

Haplosclerida with three dimensional ectosomal skeleton of ffistory and biology

multispicular fibres. Choanosomal skeleton of multispicular fibres,
cored by oxeas, often strongylote or stylote. Microscleres if pres- The family was erected by Van Soest (1980) to include
ent, sigmas or microxeas. Haplosclerida with an ectosomal paratangential multispicular
reticulation of fibres or tracts, and a choanosomal reticulation of
Diagnosis multispicular fibres or tracts. Microscleres, if present, include
only sigmata or microxeas. Van Soest (1980) remarked this family
Encrusting, massive, fan-shaped, flabelliform, vase-shaped was specifically created to receive chalinid sponges which had
or branching growth forms, often bearing chimney-like oscular multispicular fibres, and in this work described many Caribbean
processes. Sponge texture stiff or resilient, to hard and brittle. species of Amphimedon, Niphates and Cribrochalina.
Surface smooth or rough, partially hispid, tuberculate, ridged, or The definition of Niphatidae has evolved from several subse-
strongly spiny. Ectosomal skeleton a dense multispicular, tangential quent reviews of the family. Hartman (1982) based his revision
(or paratangential) reticulation of brushes of spicules, representing principally on the nature of the ecto- and choanosomal skeleton, the
the protruding terminal parts of the choanosomal primary longitudi- type of mega- and microscleres, and their morphologies. Pulitzer-
nal fibres, supporting an ectosomal network of rectangular to Finali (1981) included two new species of Gelliodes and
rounded meshes, made of secondary fibres, characteristic at the sur- Amphimedon from Australia. Desqueyroux-Fa11ndez (1984) empha-
face of some species. This structure sometimes may be absent, sised the nature of the ectosomal skeleton as the principal taxo-
weakly developed or obscured by terminal brushes of primary nomic character, and describing species of Niphates, Amphimedon
fibres. Interstitial spicules are present in some species and form a and Gelliodes from New Caledonia. De Weerdt & Van Soest (1986)
free network. Choanosomal skeleton is a dense network of primary did not report any Niphatidae from the North Atlantic fauna
multispicular, longitudinal fibres, parallel-radially plumose or although mentioning species of Gelliodes previously recorded from
divergent, often fasciculated or largely ramified, cored by multispic- this region by authors. These authors included Aka de Laubenfels,
ular bundles of oxeas or strongyles. Primary fibres are transversally 1936a in Niphatidae despite similarities with Siphonodictyon (both
connected by short, well developed or irregularly ramified tracts to now in Phloeodictyidae). Kelly Borges & Bergquist (1988) reported
form confused secondary or connecting fibres. Spongin typically on species of Niphates, Gelliodes and Siphonodictyon from Papua
abundant, but may be scarce or just present to unite spicules (e.g., New Guinea. Wiedenmayer (1989) described only a single species
Microxina). Megascleres oxeas or strongyles of variable sizes, from Bass Strait but provided an analysis of the proportion of
longer and stronger than in Callyspongiidae (usually between spicules to spongin in massive-lobose species of Gelliodes with
100--600 f.Lm length and 3-29 f.Lm in diameter). Free spicules often Indo-West Pacific records. Fromont (1993) noted the practical diffi-
abundant. Microscleres if present, sigmas or microxeas. culty of using paratangential three dimensional ectosomal structure

874
Porifera· Demospongiae • Haplosclerida • Haplosclerina • Nipbatidae 875

as a principal character to define the family, suggesting instead that the structure of the surface and the strong fibres virtually without
the strong development of a primary and secondary multi spicular spongin in Gelliodes, Indo-Pacific Gelliodes is not congeneric with
skeleton was more useful to differentiate it from Chalinidae, and West-Indian Niphates.
described species of Amphimedon, Niphates, Gelliodes and Aka
from the Great Barrier Reef (Aka subsequently referred to Distribution
Phloeodictyidae). Hooper & Wiedenmayer (1994) recognised six
valid genera of Niphatidae: Amphimedon, Cribrochalina, Gelliodes, Species are generally found in shallow waters, although
Microxina, Niphates and Hoplochalina, five with Australian records. Hartman (1982) stated that genera other than Amphimedon occur
Hoplochalina was included in the family for the first time, whereas from tropical shallow-waters to depths of 2400m, and Koltun
previously Van Soest (1980) listed it as a Haplosclerids of uncertain (1964b) recorded species of Dasychalina and Microxina from
identity. We have examined the type specimen (H. incrustans, BMNH 256 and 540m depth. Genera are widely distributed (Van Soest,
1886.8.27.521), and conclude that it does not belong to Niphatidae. personal communication): Chile, Peru and Magellan Regions
Actually, there is no concensus about valid genera of this family. Van (Amphimedon, Haliclonissa, Hemigellius, Microxina, Niphates and
Soest (1980) mentioned seven genera of Niphatidae: Amphimedon, Pachychalina), Antarctica (Amphimedon, Gelliodes, Haliclonissa,
Niphates, Siphonodictyon, Cribrochalina, Hemigellius, Haliclonissa, Hemigellius, Microxina and Niphates), East South America and
and probably also Microxina, but no representatives of Gelliodes Brazilian Regions (Amphimedon, Haliclonissa, Niphates), Caribbean
and Siphonodictyon were found in his West Indies collection. Islands and West Indies (Amphimedon, Gelliodes, Niphates),
Contradictory opinions about the availability and synonymy of two West Africa (Amphimedon, Gelliodes, Hemigellius, Niphates),
genera often included in this family: Aka de Laubenfels, 1936a and Mediterranean-Atlantic, West Mediterranean (Amphimedon,
Siphonodictyon Bergquist, 1965, are discussed below. In Van Soest's Gelliodes, Hemigellius), Boreal West Atlantic (Hemigellius,
(1980) opinion the identity of some genera remain uncertain. For Pachychalina), Boreal East Pacific (Pachychalina), Japan-China
example, Gelliodes is only barely differentiated from Niphates, with (Amphimedon, Gelliodes, Hemigellius), Red Sea (Amphimedon,
alleged differences such as the amount of spongin and morphology of Gelliodes), central Indian Ocean (Amphimedon, Gelliodes, Niphates),
interconnecting fibres being difficult to quantify. Notwithstanding the Australia (Amphimedon, Cribrochalina, Gelliodes, Microxina,
opinion of Wi edenmayer (1977b), who considered characters such as Niphates), New Zealand (Gelliodes).

KEY TO GENERA

(1) Microscleres (sigmas, microxeas, rarely toxas) present ...................................................................................................................... 2

No microscleres ................................................................................................................................................................................... 4
(2) Surface skeleton a paratangential (three-dimensional) network of spongin enclosed paucispicular tracts, surface uneven, maybe
conulose, or stiffly spiny but not hispid ....................................................... ........................................................................................ 3
Surface skeleton consisting of spicules brushes, surface finely or coarsely hispid, no (para-)tangential spicule tracts,
spongin not visible ............................................................................................................................................................. Hemigellius
(3) Surface uneven, microconulose, but not stiffly spiny, choanosomal skeleton regularly anisotropic, microscleres
vestigial ... ........... .......................... ... .................. ................. ..... ................................. .......... ................ ........... ..................... ..... Niphates
Surface stiffly spiny, choanosomal skeleton with reduced secondary fibres, microscleres numerous .................................. Gelliodes
(4) Surface optically smooth, microconulose or bumpy, but not stiffly conulose, spiny or coarsely hispid ............................................. 5
Surface stiffly conulose, spiny or coarsely hispid ............................................................................................................................... 8
(5) Ectosomal skeleton a paratangential or three-dimensional network of spicule tracts ......................................................................... 6
Ectosomal skeleton lacking a network, consisting of the closely set brushed endings of the choanosomal skeleton making a crust-like
ectosome ............ ...... ........... ................ .......... ................................ ................ ................. ........................... ...................... Cribrochalina
(6) Spongin absent or invisible, choanosomal skeleton with emphasis on primary tracts, many loose interstitial spicules .... Pachychalina
Spongin clearly present enclosing ectosomal and choanosomal spicule tracts, choanosomal skeleton regularly anisotropic ........... 7
(7) Surface more or less smooth, but may be uneven or bumpy, ectosomal network tangential .......................................... Amphimedon
Surface microconulose, ectosomal network paratangential (three-dimensional) ................................................................... Niphates
(8) Surface spiny 'aculeate' (with stiff conules) ..................................................................................................................... Dasychalina
Surface coarsely hispid, irregular ..................................................................................................................................... Haliclonissa

AMPHIMEDON DUCHASSAING & MICHELOTTI, 1864 Definition

Synonymy Niphatidae with an optically smooth surface, regular tangen-

tial ectosomal network with rounded meshes of a single size. Ends
Amphimedon Duchassaing & Michelotti, 1864: 78; de of choanosomal longitudinal primary fibres barely protruding.
Laubenfels, 1936a: 45. Hemihaliclona Burton, 1937: 18, pI. i, fig. 7. Spongin abundant. Microscleres absent.

Type species Diagnosis

Amphimedon compressa Duchassaing & Michelotti, 1864 (by Irregularly massive, lamellate, flabelliform, branching growth
subsequent designation, de Laubenfels, 1936a). forms, with numerous oscules linearly distributed on rims or
876 Porifera· Demospongiae • HapJosclerida • HapJosclerina • Nipbatidae

Fig. 1. Amphimedon Duchassaing & Michelotti, 1864. A-D, type species Amphimedon compressa Duchassaing & Michelotti, 1864. Lectotype ZMA POR
863. A, lectotype (reproduced from Duchassaing & Michelotti, 1864) (scale I cm). B, diffuse to radially plumose choanosomal skeleton (scale 50 ILm).
C, ascending, longitudinal multispicular primary fibres (scale 50 ILm). D, primary fibres grouped in bundles and irregular interconnections (scale 50 ILm).

scattered on branches. Ectosomal skeleton a tridimensional tangential (Fig. lA). Consistency in dry state, hard. Ectosomal tangential net-
network of secondary fibres with rounded meshes, covered by a fine work incomplete, irregular, with uniform rounded meshes, 100-200
membrane. Choanosomal skeleton irregular, diffuse, radially fLm, abundant free spicules, formed by secondary fibres, 20-90 fLm
plumose network of primary, ramified multispicular fibres, irregu- in diameter. Choanosomal skeleton a diffuse to radially plumose net-
larly connected by secondary multi spicular fibres. Abundant spongin. work (Fig. lB) with irregular meshes, 90-300 fLm in diameter, with
Megascleres oxeas often with modified or strongylote apices. ascending, longitudinal multispicular primary fibres, rich in spongin
Microscleres absent. (Fig. lC), 41-63-120 fLm in diameter cored by 15 or more spicules.
Primary fiures, grouped in bundles, are quite irregularly intercon-
Previous reviews nected (Fig. ID), by secondary multispicular fibres, 8-21-33 fLm in
diameter. Megascleres: oxeas, slightly bent, regular in diameter, with
De Laubenfels, 1936a: 213; Burton, 1937: 18; Hartman, 1955: multi telescoped or strongylote apices, 106-140-168 fLm length and
167; Wiedenmayer, 1977b: 82; Van Soest, 1980: 26; Desqueyroux- 3--4.7-5 fLm in diameter. Microscleres absent.
Faundez, 1984: 777; Fromont, 1993: 21. Distribution. West Indies; east coast of South America;
Australia; New Caledonia; Antarctica.
Description of type species Remarks. De Laubenfels (1936a: 45) included Amphimedon
in his new family Haliclonidae (= Chalinidae) for sponges contain-
Amphimedon compressa Duchassaing & Michelotti, 1864 ing predominantly diactinal spicules distributed more-or-Iess
(Fig. 1). homogeneously throughout the skeleton, and lacking any ectoso-
Synonymy. Amphimedon compressa Duchassaing & mal specialization. He suggested that it differed from Haliclona
Michelotti, 1864: 78, pI. 17, fig. 2. principally by the possession of longitudinal fibres projecting
Material examined. Lectotype ZMA POR. 863 - St. through the surface. Burton (1937) erected Hemihaliclona, desig-
Thomas (designation by Wiedenmayer, 1977b): Not examined. nating Amphimedon viridis as the type species. Wiedenmayer
Paralectotype: MT Por. 35 - St. Croix. Schizoparalectotype: (1977b: 82) considered Hemihaliclona as a subjective junior syn-
BMNH 1928.11.12.42, as Haliclona (Amphimedon) compressa. onym of Amphimedon, and assigned it to a subgenus of Haliclona
Other comparative material. Holotype of Amphimedon cristata based on its possession of a halichondroid-like skeleton with abun-
Pulitzer-Finali, 1996: MSNG 48708 - Laing I., PNG. Holotype of dant spongin enclosing spicules, and a surface network of hispid
Amphimedon conferta Pulitzer-Finali, 1996: MSNG 48709 - Laing fibres partially or well developed. Van Soest (1980) considered
I., PNG. Holotype of Amphimedon rudis Pulitzer-Fin ali, 1996: that the presence of a special tangential ectosomal reticulation
MSNG 48711 - Laing I., PNG. was justification to restore Amphimedon to generic rank, and
Description (emended from Van Soest, 1980). Massive to to remove it from Haliclonidae (=Chalinidae) to his new family
flabelliform, up to 13 cm high, cylindrical branches 7 cm in diameter, Niphatidae.
with rows of oscules flush with the surface, 2-8 mm in diameter, Van Soest (1980) included the poorly known type species of
scattered over the the upper part of branches or concentrated on side Pachychalina Schmidt into synonymy with Amphimedon, whereas
rims. Surface smooth on the flattened parts, tuberculate and ridged we suggest that there are significant skeletal differences between
on the rims. Colour, bright red, alive, ochre to brown in dry state their respective type species to recognise both genera as valid.
Porifera· Demospongiae • Haplosclerida • Haplosclerina • Nipbatidae 877

Fig.2. Cribrochalina Schmidt, 1870. A-D, type species Cribrochalina infundibulum, lectotype BMNH 1870.5.3.165. A, ectosomal palisade of oxeas and
protruding longitudinal primary fibres (scale 200 11m). B, strong surface crust, transverse section (scale 100 11m). C, longitudinal section through
choanosomal skeleton showing a regular network of multispicular longitudinal primary fibres and elongate mesh with abundant free spicules (scale
50 11m). D, choanosomal fibres, nearly parallel, with large spongin sheath (scale 50 11m).

Desqueyroux-Faiindez (1984) followed Van Soest's (1980) concept nearly parallel to radiating strong fibres, with a large spongin
of Amphimedon by introducing a new character (percentage of sil- sheath. Meshes are elongated to rectangular.
ica in the skeleton) to define species. These values were found to be
very variable (between 8.6-71.0 mgllOO g), but this character now Diagnosis
appears to be poorly informative for systematics of this group.
Fromont (1993) also included Pachychalina into synonymy with Lamellate, folded, fan, vase, or cup-shaped, thin-walled
Amphimedon, and also redefined Amphimedon to include the pres- growth forms, attached directly to the substrate, with a basal por-
ence of sigma microscleres as a good specific character. tion burrowed into the sediment. Oscules generally not distinguish-
able externally from numerous holes puncturing the surface.
Surface optically smooth, with an ectosomal network consisting of
a palisade of spicule-brushes covered by a fine membrane (crust).
CRIBROCHALINA SCHMIDT, 1870 Choanosomal skeleton with strong, ascending longitudinal multispic-
ular primary fibres radially distributed to produce an anisotropical
Synonymy skeleton with regularly elongated meshes. Spongin abundant.
Microscleres absent.
Cribrochalina Schmidt, 1870: 36, pI. 4, fig. 3; 1880: 77.
Topsent, 1920a: 6. Distribution

Type species Thirteen nominal species of Cribrochalina are known from

West Indies and West Central Pacific (Van Soest, pers. comm.).
Cribrochalina infundibulum Schmidt, 1870 (by subsequent
designation, de Laubenfels, 1936a: 46). Previous reviews

Definition Topsent, 1920a: 6; de Laubenfels, 1936a: 46; Wiedenmayer,

1977b: 118; Van Soest, 1980: 42.
Niphatidae with ectosomal network masked by a palisade
of spicule brushes produced by the ends of primary longitudinal Description of type species
fibres and appearing as a hard, strongly hispid surface crust.
Choanosomal skeleton composed of multi spicular ascending, Cribrochalina infundibulum Schmidt, 1870 (Fig. 2).
878 Porifera' Demospongiae • HaploscIerida • HaploscIerina • Nipbatidae

Synonymy. Cribrochalina infundibulum Schmidt, 1870: 36. and named Spongia bilamellata as Cribrochalina bilamellata
Material examined. Lectotype (here designated): BMNH (Lamarck). Re-examination of the Lamarck specimen confirms its
1870.5.3.165 (fragment) -Antilles. Paralectotypes: MZUS POR 123 suspected affinities with Cribrochalina of the West Indies,
(fragment 25, 47) - West Indies (labelled 'type' by Schmidt, fide although in Cribrochalina spicule dimensions are larger, the sur-
Topsent, 1920a). BMNH 1870.5.3.134 - teased fragment. Other com- face is a crust, and fibre structure is more compact - suggesting
parative material. Holotype of Spongia bilamellata Lamarck: MNHN that Cavochalina and Cribrochalina are different and probably
LBIM DT 602 - King 1., Bass Strait. Carter's specimen of valid taxa, allocated to two different families: Callyspongiidae and
Cavochalina bilamellata: BMNH 1886.12.15.182 - S coast Australia. Niphatidae, respectively. Wiedenmayer (1977b: 118) also included
Cribrochalina cretacea Schmidt, 1870: MCZ 9135 (orig. 6633w) - Cavochalina Carter, 1885 and Placochalina Lendenfeld, 1887c in
Florida. MCZ 6090 - 'Blake Expedition', stn 200. off Martinique. synonymy with Cribrochalina (following Topsent, 1932a).
Description. Lamellate, funnel-shaped, thin-walled. Surface Wiedenmayer (1977b) also associated Cribrochalina (in part)
smooth, outer surface irregularly undulated, inner surface concen- with Petrosia and Xestospongia, as members of the family
trically undulated. Under the fine membrane covering the surface NepheJiospongiidae, which he revived. In Van Soest's (1980)
(appearing as imperforate), are very small oscules and pores scat- opinion the structure of Cribrochalina with strong primary fibres,
tered over the surface. The strong surface crust produces a hard rich in spongin and radially distributed to form a crust, is very dif-
consistency, stiff and brittle in the dry state. Colour alive purple to ferent from the isotropical meshes of Petrosia and from the skeletal
purple-brown. Ectosomal skeleton is a continuous palisade of oxea structure of Xestospongia. Van Soest (1980: 108, fig. 37) refuted
brushes produced by the diverging surface ends of multi spicular the proposition of Wiedenmayer (1977b) but acknowledged the
longitudinal primary fibres, to form a strong crust (Fig. 2A-B). strong affinity of Cribrochalina with Amphimedon and Niphates,
Choanosomal fibres are nearly parallel, with a large spongin except that Cribrochalina had a much finer skeletal mesh and had
sheath, forming a regular network of multi spicular longitudinal an ectosomal crust of spicule brushes. In Wiedenmayer's (1977b)
fibres, 85-120 f.Lm in diameter, with elongate meshes 300-400 f.Lm concept Cribrochalina also corresponded, in part, to Pachychalina
in diameter and abundant free spicules (Fig. 2C-D). Diverging (e.g., Pachychalina aurantiaca (Lendenfeld) var. dura Wilson,
terminations of the ascending primary fibres under the surface are 1902, from the Virgin Islands), which is probably a species of
prolifically divided to produce an irregular system of subectosomal Petrosia (Van Soest, pers. comm.).
cavities and peripherically condensed meshes, 120-180 f.Lm in Finally, Wiedenmayer (1977b: 119) considered Cribrochalina
largest diameter. Megasc1eres: strongylote oxeas, strongly bent or infundibulum was con specific with C. vasculum (Lamarck). We
flexuous. Topsent (1920a) provides the following dimensions, observed in both species the same types of spiCUles (slightly longer
180-215 X 3.5-5 f.Lm to 250 X 5.5 f.Lm Wiedenmayer (1997: 121) in C. vasculum), with fibres having the same structure (with the
records micrometries as 180-215 X 3.5-5 f.Lm. mesh of C. vasculum irregular and largerin size), and with a subec-
Remarks. Cribrochalina Schmidt, 1870 was redescribed tosomal condensation (less marked than in C. infundibulum).
by Topsent (1920a) based on two fragments (MZUS 25 and 47, Wiedenmayer (1977b) suggested that these differences were
here considered paralectotypes) of Cribrochalina infundibulum. intraspecific, despite Topsent's (1920a) assertion that Lamarck's
Schmidt's material in ZMUS was also examined but the real status specimen was Calyx vasculum. Van Soest (pers. comm.) suggested
of this material is completely uncertain, with the consequence that that it was not certain that Spongia vasculum was West Indian,
the fragment BMNH 1870.5.3.165 was designated as lectotype. irrespective of the alleged similarity between C. infundibulum and
This fragment has the exact wording on the label 'Antillen', con- C. vasculum, with the consequence that the argument remains
forming exactly to Schmidt's (1870) description, whereas the speculative. Wiedenmayer (1977b: 119) noted that two potential
locality of other material is less specific. In our diagnosis here we syntypes of Cribrochalina infundibulum exist in the Museum of
discuss the properties of this fragment. Comparative Zoology, Harvard University. Re-examination of this
Topsent (1920a) remarked that Cribrochalina resembled material (courtesy of Ardis B. Johnston, Assistant Curator,
Pachychalina in the number of spicules and their arrangement Department of Invertebrate of this Museum) indicates that there is
within fibres, with abundant long, fine, bent and sinuous oxeas. But no justification to consider this material as representing valid types,
based on our re-examination of the lectotype we conclude that with all labelled 'non type'.
there are important differences between the type material of both
genera, which is evidence for recognising them as distinct genera.
Fibre structure is similar in both genera, but in Cribrochalina fibres DASYCHAUNA RIDLEY & DENDY, 1886
are stronger, nearly parallel, with a large spongin sheath and their
umbelliform ends and free spicules form a strong surface crust. In Synonymy
Pachychalina fibres are vaguely parallel, without visible spongin,
and meshes are confused, without properly connecting fibres. Dasychalina Ridley & Dendy, 1886: 329; 1887: 19.
A surface crust is not present in Pachychalina, but its ectosomal
skeleton is three-dimensional, irregular and confused, and it is not Type species
recognizable as distinct from the main skeleton.
Topsent (1920a) had a clear concept of Cribrochalina, but Dasychalina fragilis Ridley & Dendy, 1886: 330 (by subse-
later (Topsent, 1932a) included Spongia bilamellata (typical form) quent designation, Burton, 1932b: 278).
Lamarck in the genus, recognising the same specimen redescribed
by Carter (1885; as Cavochalina bilamellata, a callyspongiid), and Definition
by Lendenfeld (1887c; as Placochalina pedunculata). Topsent
(1932a) considered Cavochalina Carter, 1885 and Placochalina Niphatidae with rugged and spinous or conulose surface.
Lendenfeld, 1887c as junior objective synonyms of Cribrochalina, Ectosomal skeleton an irregular network of stout fibres and free
Porifera • Demospongiae • Haplosclerida • Haplosclerina • Niphatidae 879

Fig. 3. Dasychalina Ridley & Dendy, 1886. A--C, type species Dasychalinafragilis Ridley & Dendy, 1886, lectotype BMNH 1887.5.2.170. A, ectosomal
skeleton, an irregular reticulation of isolated spicules supported by an additional stronger reticulation (scale 50 ILm). B, apices of the ectosomal conules
(scale 500 ILm). C, compact multispicular principal fibres without spongin sheath (scale 500 ILm). D, Gelliodes jibulatus (Carter) Ridley, 1884a specimen
from Torres Strait, BMNH 1882.2.23.202 (scale 1 cm).

spicules, reinforced by a secondary subectosomal reticulation, form- Description of type species

ing surface conules. Choanosomal skeletal network a compact retic-
ulation of stout fibres and scattered spicules. Spongin not visible. Dasychalinafragilis Ridley & Dendy, 1886 (Fig. 3).
Synonymy. Ridley & Dendy, 1886: 330; 1887: 20, pI. 4,
Diagnosis figs 2, 2a.
Material examined. Lectotype (here designated): BMNH
Massive, irregularly branched, ramose to flabellate growth 1887.5.2.170 - 'Challenger' expedition, stn. 208, Philippines.
forms . Surface strongly aculate, conulose, uneven and striated, with Other comparative material. Holotype of Dasychalina melior
striae radiating from the summit of conules (Ridley & Dendy, 1887: Ridley & Dendy, 1886: BMNH 1887.5.2.178 - 'Challenger'
20). Texture strong, hard and brittle. Oscules large, irregularly scat- Expedition, stn. 208, Philippines. Holotype of Dasychalina validis-
tered on upper side of branches. Ectosomal skeleton, (Fig. 3A) sima (Thiele, 1905): 2MB 2234 - Plate Collection, Calbuco, Chile.
formed by stout, multispicular fibres, and free spicules in a dense Holotype of Dasychalina magellanica Thiele, 1905: 2MB 3327 -
irregular network with irregular mesh, under which a subectosomal Plate Collection, Bahia Parke, Cockburn Canal, Punta Arenas,
network of stout fibres is visible. Aculeated conules or spines on the Chile.
surface (Fig. 3B) are formed by central branches at ends of primary Description (emended from Ridley & Dendy, 1886: 330).
longitudinal fibres. Choanosomal skeleton an irregular network of Irregularly ramose, subcylindrical branches about 18 mm in diame-
primary fibres, without clear orientation, strong, abundantly rami- ter, covered with strong conules or spines (aculeated). Colour in
fied, and with no visible spongin sheath (Fig. 3C). spirit light greyish or brownish yellow. Texture hard and brittle.
Surface uneven, delicately striated, with striations radiating from
Distribution ends of conules. Oscules large, about 4 mm in diameter, almost
entirely scattered and restricted to one side of each branch.
Philippines (Ridley & Dendy, 1886); South Georgia, Falkland Ectosomal skeleton (Fig. 3A), an irregular reticulation of isolated
Islands; South coast of Chile: Calbuco, Magellan Strait; South spicules supported by an additional stronger reticulation of stouter
Georgia; Antarctica (Burton, 1932b, 1934a, 1940; Thiele, 1905; fibres the ends of which form the apices of the ectosomal conules
Desqueyroux, 1975, 1976; Koltun, 1964b). (Fig. 3B). Choanosomal skeleton an irregular reticulation of very
stout, compact multispicular principal fibres, 35-50 JJ..m in diame-
Previous reviews ter, without spongin sheath, spongin present but only cementing
spicules (Fig. 3C) in the fibres and in between fibres, with
Burton, 1932b: 278; 1934a: 15; [non] de Laubenfels, 1936a: abundantly scattered free spicules. Megascleres: oxeas 350-420-
45; Wiedenmayer, I977b: 53,95, 100. 500 JJ..m in long by 15-20 JJ..m in diameter.
880 Porifera· Demospongiae • Haplosclerida • Haplosclerina • Nipbatidae

Remarks. Ridley & Dendy (1886: 329) described Type species

Dasychalinafragilis as 'solid, coarsely spined on surface; skeleton-
fibres stout, spicules polyserial; amount of spongin variable, never Gelliodes fibulata Ridley, 1884a (by subsequent designation;
very great', further indicating that the solid, coarsely spined surface Hooper (1986: 187), this is generally considered a junior synonym
was the main character differentiating it from Pachychalina. of Axosfibulata Carter, 1881) (by monotypy).
However, Ridley & Dendy (1887: 19) realised that the type species
of Pachychalina (P. rustica Schmidt, 1868), may have a spinous Definition
('struppig') surface, and consequently, if both Dasychalina and
Pachychalina had a spinose surface they should be included in a Niphatidae in which the tangential ectosomal layer of fibres
single genus. Thus, Dasychalina became a junior subjective syn- with an irregular mesh is interrupted by protruding ends of
onym of Pachychalina, although several authors subsequently con- choanosomal longitudinal fibres. Choanosomal skeletal network
tinued to use the name Dasychalina (e.g., Burton, 1934a: 15, is composed of compact multispicular longitudinal fibres.
Dasychalina validissima (Thiele, 1905». Burton (1932b: 278) was
the first author to comment significantly on the genus, with the Diagnosis
type species (designated by him as Dasychalinafragilis) exhibiting
a different skeletal structure from his concept of Pachychalina, Thickly incrusting to massive, tubular growth form, intri-
although he admitted that Pachychalina was unknown to him. cately branching, long cylindrical stems irregularly ramified and
Burton's (1932b) definition of Dasychalina was confined to anastomosing at points of contact (single branches attain a length
sponges like Adocia and Callyspongia (= Pachychalina of of about 100mm), rampant or erect, arising from a common basal
authors), defined as 'haplosclerids with ectosomal skeleton irregu- portion. Oscules usually numerous, unevenly scattered over the
lar multispicular reticulation with primary and secondary mesh'. surface and often conspicuous. Surface uneven, membranous,
Wiedenmayer (1977b: 100) included Dasychalina into synonymy strongly aculeated at intervals of about 2-5 mm, sustained by
with Spinosella (pro Cladochalina), considering that Burton's strong, slender, sharp ramified spines, 2-3 mm long (Fig. 3D) sur-
(1932b) concept of Dasychalina corresponded to sponges like face may be also ridged or tuberculate or smooth, and finely hispid
Spinosella with little or no spongin, and with oxeas packed in tracts or velvety. Texture very hard. Ectosomal skeleton is a tangential
(e.g., Callyspongia (Cladochalina) ridleyi Burton, 1934a). Van network of secondary fibres, free oxeas and abundant sigmas
Soest (1980: 106) rejected Ridley & Dendy's (1886) and (Fig. 4A), often interrupted by the ends of the strong primary lon-
Wiedenmayer's (1977b) concepts of Dasychalina and gitudinal fibres protruding from the choanosomal skeleton to form
Pachychalina, considering the genus to be a subjective junior syn- the spines. Choanosomal skeleton composed of primary longitudi-
onym of Niphates. As we see, however, Dasychalina has been nal-radial multispicular and ramified primary fibres, distinct and
compared with genera containing a very different type of skeleton very compact (Fig. 4B). Primary fibres (Fig. 4C) form rectangular
(i.e., C. (Cladochalina), Pachychalina and Niphates), in spite of to rounded meshes, subdivided irregularly by secondary fibres, and
the presence of a different type of surface (aculeate) and the large mesh containing abundant free spicules. Megascleres (Fig. 4D)
size of its spicules (more than 400 11m). Skeletal differences consist of robust oxeas with sharp apices. Microscleres are abun-
between these genera, confirmed from study of their respective dant sigmas.
type species, are important and thus we consider here that these
three genera are different and valid. Pachychalina has a very irreg- Distribution
ular surface, abundantly perforated by aquiferous openings, no
aculeations, no clear ectosomal tangential network; there is no der- Indian Ocean, West Pacific Ocean, Torres Strait, Northern
mal membrane, fibres are multispicular, of irregular diameter and Territory and Western and N coast of Australia, SE coast of
without a visible spongin sheath. Niphates exhibits a conulose to Australia (Port Jackson), Amboina, Indo-Malayan region, New
spiny surface with numerous oscules; its irregularly distributed Guinea and New Caledonia (Carter, 1881; Lendenfeld, 1887c;
fibres are pauci- to multispicular, but spongin is always dominant. Topsent, 1897a; Thiele, 1903; Dendy, 1905; Van Soest, 1980;
Callyspongia (Cladochalina), in spite of the aculeated surface sim- Hooper, 1986; Desqueyroux-Fatindez, 1984).
ilar to that of Dasychalina, exhibits an ectosomal network with
three mesh sizes, all smaller than those of Dasychalina, and pos- Previous reviews
session of fasciculated primary fibres. In addition, in all three gen-
era spicules are much smaller than in Dasychalina, which appears Carter, 1881, 383, pI. 18, fig. 4; 1882: 288; Ridley, 1884a:
to be an important character to differentiate species of Dasychalina 427, pI. 39, fig. 1, pI. 41, fig. bb-bbii; Ridley & Dendy, 1887: 47,
from sponges with an aculeated surface and containing small pI. 12, fig. 2; Topsent, 1897a: 470; 1932a: 114; Hentschel, 1912:
spicules, resembling Dasychalina (which Van Soest (in litteris) 393; Burton, 1934a: 549; Wiedenmayer, 1977b: 95; 1989: 117; Van
suggests are common in Indonesia and neighbouring areas, but in Soest, 1980: 106; Desqueyroux-Fatindez, 1981: 744, figs 56, 117;
reality are species of Callyspongia (Cladochalina». 1984: 780, figs 6, 51, 52, 58--61; Hooper, 1986: 184, figs 11, 12,
pI. 1, fig. E; Hooper & Wiedenmayer, 1994: 318.

Description of type species

GEUIODES RIDLEY, 1884
Gelliodesfibulatus Ridley, 1884 (Fig. 4).
Synonymy Synonymy. ?Axos fibulata Carter, 1881: 383, pI. 18, fig. 4.
Gelliodes fibulata Ridley, 1884a: 427, pI. 39, fig. 1, pI. 41,
Gelliodes Ridley, 1884a: 426. figs bb-bbff.
Porifera· Demospongiae • Haplosclerida • Haplosclerina • Nipbatidae 881

Fig. 4. Gelliodes Ridley, 1884a. A-D, Gelliodesfibulata Ridley, I 884a, lectotype, BMNH 1882.2.23.226. A, ectosomal skeleton a tangential network of
secondary fibres, free oxeas and abundant sigmas (scale 200 ~m). B, choanosomal skeleton with irregularly connected primary fibres and scarce spongin
binding spicules (scale 50 ~m). C, choanosomal spicules, robust oxeas with sharp apices (scale 20 ~m). D, sigma microscleres (scale 2 ~m) and oxea
megasc1eres (scale 5 ~m).

Material examined. Lectotype (here designated): BMNH region but are rare in the choanosomal region. Megascleres: oxeas
1882.2.23.226 - Prince of Wales channel, Torres Straits, 7-9 fath., straight to slightly curved, hastate, 135-182-250/-Lm long by
bottom sand, colI. Dr Coppinger, 9/1881. Other material. BMNH 3-6-16/-Lm in diameter. Microscleres: sigmas, c-shaped, slightly
1882.2.23.193 - Thursday Island, Torres Straits. BMNH centrangulate, 10-12-16 /-Lm long.
1882.2.23.202 (dry), 202a (slide) - Prince of Wales channel, Torres Remarks. The holotype ofAxos jibulata Carter, 1881 was
Straits. BMNH 1882.2.23.240 (spirit), 240 (slide) - Torres Straits, confirmed as lost (Burton, 1934a) precluding a definite conclusion
bottom mud, 10/1881. BMNH 1882.2.23.547, 548, 549 - Thursday that it is a senior synonym of Gelliodes jibulatus Ridley, 1884.
Island, Torres Straits, 3-5 fath., bottom mud and sand. Hooper (1986) revised the genus Axos Gray, 1867 from old and
Description. Long, cylindrical, repent branches issued at recent material collected from Northern Territory and Western
different levels, from a common base, branches anastomosing at Australia and compared this with Gelliodes jibulatus Ridley,
points of contact. Surface covered by a transparent membrane, over 1884a. Hooper confirmed that this species was distinct and clearly
which protrude the ends of long primary fibres, as long and short differentiated from other species of Axos (Hooper, 1986: 187).
spines, making the surface extremely spinous and aculeated at Gelliodes jibulatus has been accurately and frequently described
intervals of about 2-5 mm. Longer spines, 2-3 mm long, are rami- by contemporary authors as it is very easily recognisable by its
fied to form secondary spines. Choanosomal skeleton cavernous, distinctive arborescent growth form, its strongly spiny surface
formed by abundant primary multispicular longitudinal fibres, the (with its outer surface 'beset with pointed eminences') and its spic-
spines issue from the ends of primary fibres. Intermediate surface ulation (Ridley, 1884a: 426), whereas other species subsequently
between the spines is cavernous and covered by a transparent mem- assigned to Gelliodes are not consistently spiny, including G.
brane. Colour in life grey to pale blue. Ectosomal skeleton is a tan- carnosa Dendy, 1889, G. incrustans Dendy, 1905 and G. fragilis
gential reticulation of multi spicular fibres and triangular to Desqueyroux-Faundez, 1984 which exhibit a smooth surface.
rectangular meshes. Choanosomal skeleton is cavernous, formed Hooper (1986) remarked on the possession of 'choanosomal fibres
by abundant primary longitudinal, multi spicular fibres ascending, are heavily invested by spongin' in the type species, whereas
radiating, strong, 180-450-647/-Lm in diameter, abundantly Wiedenmayer (1989) noted the highly variable proportion of spon-
ramified to form anastomosing secondary connecting fibres, gin in the skeleton, the presence of secondary fibres that may be
82-191-295/-Lm in diameter as irregular branches of primary largely reduced, the fibrous skeleton that may be isodictyal, and
fibres. Choanosomal mesh rectangular, 451-670-902 /-Lm in diam- that the surface in some species (other than the type species) may
eter, with abundant free oxeas and a few sigmas. Aquiferous spaces be almost smooth. Variations between species were confirmed by
are visible, especially on the surface, sigmas are abundant in this an analysis of the silica content of New Caledonian specimens
882 Porifera' Demospongiae • Haplosclerida • Haplosclerina • Niphatidae

(Desqueyroux-Faundez, 1984), ranging from 8.6% (in G. incrus- Previous reviews

tans, with a smooth surface) to 71.0% (in G.fibulatus), confirming
an apparent level of heterogeneity amongst taxa. Authors have De Laubenfels, 1936a: 46; Burton, 1940: 100; Koltun, 1964b:
commented on the resemblance between some nominal species of 102; Hooper & Wiedenmayer, 1994: 320.
Gelliodes and Niphates, possibly representing misidentifications of
either or both taxa given the emphasis on the presence of sigmas in Description of type species
both genera - regular in Gelliodes or irregularly in Niphates.
Wiedenmayer (1977b) remarked that characters such as the struc- Haliclonissa verrucosa Burton, 1932b (Fig. 5).
ture of the surface and the strong fibres, the latter virtually without Synonymy. Haliclonissa verrucosa Burton, 1932b: 270,
spongin, confirm that Indo-Pacific Gelliodes are not congeneric pI. 51, fig. 3, text-fig. 8.
with West-Indian Niphates. Both genera are also different in the Material examined. Holotype: BMNH 1928.2.15.473 -
distribution and structure of their fibres, where in Gelliodes fibres Palmer Archipelago, stn. 180, 'Discovery' Expedition, 160m depth.
are compact, sharply delimited, distinct, multispicular and with Description. Holotype measures 13-15 cm long and 5cm in
scarce, or no visible, spongin. In contrast, in Niphates the fibres are diameter, composed of a stout irregularly cylindrical branch with a
pauci- to multi spicular, abundantly ramified to form fascicles, few conspicuous oscula connected to a central cavity, 10 mm in
spongin is dominant, and always visible. diameter, as a shallow, large atrium. Texture soft, friable. Surface
densely spined, conulose or verrucose, irregular. Ectosome with-
out a surface membrane, with protruding ends of strong primary
longitudinal fibres protruding through the surface at different lev-
HALICLONISSA BURTON, 1932 els, forming the spines or aculeations, and in between these spines
there is an irregular nework of abundant free spicules; there is no
Synonymy special surface skeleton. At the surface the primary longitudinal
fibres are clearly visible on the outer side of branches, as strong
Haliclonissa Burton, 1932b: 270. tracts.(Fig. 5A). Choanosomal skeleton (Fig. 5B-C) is an irregular
network of diffuse, multispicular, primary longitudinal fibres, with
Type species irregular secondary fibres in between, vaguely present or diffuse
and forming an irregular mesh, with abundant free spicules.
Haliclonissa verrucosa Burton, 1932b (by original designation). Spongin scarce. Megascleres: oxeas (Fig. 5D) 369-395-422/-lm in
length and 12-14-16 /-lm in diameter. A second category of oxeas,
Definition 254-290-344/-lm in length, 4-8/-lm in diameter, was observed
although not mentioned by Burton (1932b). Stylote oxeas,
Niphatidae with a densely hispid surface composed of isolated 312-377/-lm long, 1-15-16/-lm in diameter, are also present.
brushes of spicules formed at the ends of primary longitudinal Remarks. Burton (1932b) erected Haliclonissa for the
fibres. Choanosomal skeleton with primary longitudinal multi- type species which he considered an intermediate between
spicular fibres connected to diffused secondary fibres, to form irreg- Haliclona and Microxina, and cited as a principal diagnostic char-
ular elongated meshes with numerous free spicules. Spongin scarce. acter its surface, which was verrucose, and the structure of its
skeleton composed of diffuse fibres forming an irregular network
Diagnosis of isolated spicules. The protruding ends of primary fibres form a
special surface skeleton corresponding to the Niphatidae model,
Stout, tubular growth form with a deep or a shallow atrium although Burton (1932b) defined Haliclonissa as lacking a surface
and cylindrical branches, irregularly ramified. Sponge fixed skeleton.
directly to the substrate. Oscules conspicuous, connected to the
atrium, irregularly scattered over the entire sponge. Surface
strongly irregular, spiny, with dense and short spicule projections,
produced by the terminal part of primary fibres, with numerous HEMIGELLIUS BURTON, 1932
free spicules in between forming an irregular, confused tangential
network without a clear mesh. Consistency is soft, easily com- Synonymy
pressible, friable. Ectosomal skeleton consists of perpendicular
spicule brushes protruding from the ends of primary longitudinal Hemigellius Burton, 1932b: 272. ? Plumocolumetta
multi spicular fibres, producing a densely spiny surface. In between de Laubenfels, 1936a: 54. Calyxadocia de Laubenfels, 1936a: 70.
brushes of spicules there is an irregular network with abundant
free spicules. Choanosomal skeleton composed of pauci- to multi- Type species
spicular primary fibres, longitudinally parallel to oblique, abun-
dantly ramified, anastomosing. Interconnecting diffuse secondary HemigeUius rudis Burton, 1932b (=Gellius rudis sensu
fibres pauci- to multi spicular. Irregular mesh with numerous free Kirkpatrick, 1908c, a synonym of Gellius fimbriatus Kirkpatrick,
spicules. Spongin scarce. Megascleres are robust oxeas. 1908c; not Gellius rudis Topsent, 1901a) (by original designation).

Distribution Definition

Antarctica, Palmer Archipelago; Falkland Islands; Atlantic Niphatidae with a finely hispid surface produced by the
coast of South America. (Burton, 1932b; 1940; Koltun, 1964b). divergent paniculated (sensu Kirkpatrick, 1908c) ends of primary
Porifera· Demospongiae • Haplosclerida • Haplosclerina • Nipbatidae 883

Fig. 5. Haliclonissa Burton, 1932b. A-D, type species Haliclonissa verrucosa Burton, 1932b, holotype BMNH 1928.2.15.473. A, tennina1 part of tube
with longitudinal fibres clearly visible on the outer side of the sponge (scale 1cm). B, internal view of a longitudinal section of the central cavity (scale
I mm). C, choanosomal skeleton an irregular network of multi spicular primary longitudinal fibres and secondary fibres in between fonning irregular
meshes with abundant free spicules (scale 50 fLm). D, oxea megasc1eres (scale 20 fLm).

longitudinal fibres arising from the subectosomal region. Description of type species
Choanosomal skeleton is composed of confused multi spicular par-
allel diverging primary fibres, without visible spongin. Hemigellius rudis Burton, 1932b (Fig. 6).
Synonymy. Gellius rudis sensu Kirkpatrick, 1905c: 45, pI. 17,
Diagnosis figs I, la, pI. 24, fig. la. Gelliusfimbriatus Kirkpatrick, 1905c: 46,
pI. 17, figs 2, 2a, 24, fig. 2a-b. Not Gellius rudis Topsent, 1901a.
Massive to lobate and branching growth forms (Fig. 6A). Hemigellius rudis Burton, 1932b: 272.
Oscules large, with raised rims, irregularly distributed. Surface Material examined. Specimen of Gellius rudis sensu
smooth when covered by the surface membrane, finely hispid if the Kirkpatrick, 1905c: BMNH 1905.2.5.197A-B - Mc Murdo Bay,
membrane has been rubbed off. Ectosomal skeleton forms a hispid Winter Quarters, Antarctic Expedition. Holotype of G. fimbriatus
surface by a slight palisade of free spicules issuing from the ends of Kirkpatrick, 1905c: BMNH 1905.2.5.192C - Winter Quarters, off
primary longitudinal fibres. Subectosomal region with numerous Hut Point, 12-20 fms. Other material. Lectotype of Hemigellius
(embryonic, sensu Kirkpatrick, 1908c) holes formed by uni- to rudis Burton, 1932b: BMNH 1928.2.15.128, stn. 39 (RN
multi spicular connecting secondary fibres and by the bifurcated CIX.I.197-235), 25.03.1926, 'Discovery'. BMNH 1928.2.15.95 -
ends of primary longitudinal fibres (Fig. 6B). Choanosomal same locality data.
skeleton composed of primary longitudinal multispicular fibres Description (emended from Kirkpatrick, 1908c).
(Fig. 6C) whose divergent terminal parts in the subectosomal region Massively lobate or thick subcylindrical unbranched or branched
are divided in a paniculate fashion to strengthen the surface and to fragments of sponges. The largest fragments are 18 X 5 X 2.5 cm
form the fine hispidation that characterises the genus. Embryonic and 14 X 5 X 2.5 cm. Several fragments exhibit oscules 7 mm in
holes are well developed (Fig. 6D). Megascleres: Oxeas, slightly diameter, with circular raised rims. The surface, where the ecto-
bent. Microscleres: sigmas with angulate and curved ends. some exists, is quite smooth (Fig. 6A), and where the ectosome has
rubbed off is finely hispid. Ectosomal skeleton is a finely hispid
Distribution network formed by the terminal ends of primary fibres (Fig. 6B).
Choanosomal skeleton(Fig. 6C) is formed by multi spicular longi-
Antarctica: off Winter Quarters, off Coulman Island and tudinal, irregular, confused, discontinuous fibres or tracts, without
Mc Murdo Bay. (Kirkpatrick, 1905c). visible spongin and curved outwards; some of them with more than
10 spicules, I11S-3160 /-Lm in diameter. Primary fibres are
Previous reviews subsequently divided in paniculated branches (Fig. 6D), and each
terminal branch is SO-I71-380 /-Lm in diameter, ending below the
Burton, 1932b; Burton, 1938; Vacelet & Arnaud, 1972; ectosomal membrane, and producing an ectosomal palisade
Boury Esnault & van Beveren, 19S2; Van Soest, 19S0; Hooper & formed by the central oxeas of each end of the branches. This
Wiedenmayer,1994. network is visible as the support for the ectosomal membrane.
884 Porifera· Demospongiae • Haplosclerida • Haplosclerina • Niphatidae

Fig. 6. Hemigellius Burton, 1932b. A-C, type species, Gellius rudis sensu Kirkpatrick, 1908c. A, specimen BMNH 1908.2.5.197B (scale I cm). B, longi-
tudinal view of ectosomal hispid surface and subectosomal rounded cavity (scale 500 !Lm). C, choanosomal skeleton of primary longitudinal multispicular
fibres with robust oxeas (scale 200!Lm). D, Hemigellius rudis, Burton's lectotype BMNH 1928.2.15.128. Longitudinal section of subectosomal region,
with embryonic holes, multispicular primary fibres without visible spongin (scale I mm).

Immediately under the surface network there are several subecto- Antarctica (70023'S, 82°47'W), and G. rudis collected by
somal rounded cavities, 600-1556-2000 fLm in diameter, formed Kirkpatrick (l908c), from Mc Murdo Bay, especially concerning
by paniculated curved branches of principal fibres (Fig. 6D), which the subectosomal ends of primary longitudinal fibres in both: pan-
finally expand to form the ectosomal network. These cavities iculated in G. rudis Topsent collected by Kirkpatrick, and non pan-
are defined by Kirkpatrick (1908c) as embryo-containing cavities iculated in the type of G. rudis Topsent, 1901a. At the same time
(although in our studies of specimens these cavities were empty). Vacelet & Arnaud (1972) remarked that G. Jimbriatus and G. rudis
Remarks. Burton (1932b) erected Hemigellius, and desig- sensu Kirkpatrick, 1907a were subjective synonyms. The character
nated Gellius rudis Topsent, 1901a as the type species. However, 'paniculated ends' of primary fibres in Kirkpatrick material, in
Burton misinterpreted Topsent's (1901a) description of Gellius their opinion, justified the erection of the new genus Hemigellius
rudis (1929a) and thus Hemigellius is based on Kirkpatrick's by Burton (1932b). Burton (1932b: 272) considered his genus
(l908c) specimens of Gellius rudis, which Burton named Hemigellius to be closely allied to Haliclona, but later (Burton,
Hemigellius, and which are certainly different from Topsent's spec- 1938: 7), observing the variation in the skeletal arrangement of
imen (=Haliclona (Gellius); see Vacelet & Arnaud, 1972: 20). H. pachyderma, he considered it was doubtful whether the species
Burton (l932b) remarked that the original description of Topsent was distinct from the type species of his genus, and indeed if the
(1901a) gave little information regarding the structure of the skele- genus Hemigellius itself was distinct from Haliclona. These argu-
ton, but the surface was said to be finely hispid, and in Burton's ments are rejected here and Hemigellius clearly belongs to
opinion specimens of Gellius rudis Topsent described by Niphatidae as presently interpreted.
Kirkpatrick (1908c) allegedly resembled the holotype of Topsent Van Soest and de Weerdt (in litteris) remarked on the genus
closely enough to justify his diagnosis of Hemigellius (i.e., based Plumocolumetta de Laubenfels, 1936a: 54 (type species Gellius
on Kirkpatrick's specimens of G. rudis). Vacelet & Arnaud bidens Topsent, 1901a: 2 from Antarctica, by original designation).
(1972: 20) remarked on the differences exhibited between G. rudis For a full description see Topsent (l901d: 14, pI. II fig. 8, pI. III
Topsent, collected by the 'Belgica' Expedition, 1897-99, from fig. 7). The holotype (not examined) is in IRSNB no. POR032,
Porifera· Demospongiae • Haplosclerida • Haplosclerina • Nipbatidae 885

cf. Willenz, 1994: 8). A subsequently collected specimen from Diagnosis

MNHN (Jean Charcot Expedition 1905, No. 47) was examined and
this was found to conform to Topsent's original description. It is a Tubular and branched, to globular with cylindrical branching
hollow cylindrical species with a finely hispid, punctate surface. growth forms. Tubes or branches are issued from a common base,
The choanosomal skeleton consists of very loose longitudinal have thin walls, and the sponge has a deep central exhalant cavity.
spicule tracts and many single species arranged in confusion. Several scattered oscules, terminal if singular. Surface is uneven,
At the surface there is a more-or-Iess tangential crust of disorderly membraneous and strongly spined. Ectosomal skeleton forms
arranged spicules carried by the endings of the choanosomal spicule a tangential irregular network of oxeas between the spines, and
tracts, which also protrude beyond the surface for slight distance. appears obscured by them. Choanosomal skeleton exhibits longitu-
The megascleres are robust curved oxeas, 660-700 X 17-20 fLm. dinal, compact, multi spicular, fasciculate and anastomosing pri-
Microscleres are sigmas with unique bifid apices, curved outwards mary fibres, issuing from the base of the sponge. Between the
and downwards, rather resembling a snake's tongue. Its oxeas and fibres, large, irregular meshes occur with abundant free spiCUles.
surface skeleton are quite different from sponges of Haliclona No visible spongin. Microscleres microxeas.
(Gellius), and in combination with the peculiar sigmas, this induced
de Laubenfels (1936a) to erect a separate genus for this species. Previous reviews
However, apart from the uniquely ornamented sigmas, this sponge
conforms to Hemigellius. Until further species with such sigmas are Topsent, 1916, 1917; Hooper & Wiedenmayer, 1994.
found, we prefer to consider Gellius bidens a member of this genus
and accordingly Plumocolumetta is a subjective junior synonym of Description of type species
Hemigellius (taxonomic decision this work).
Calyxadocia de Laubenfels, 1936a was considered to be Microxina charcoti Topsent, 1916 (Fig. 7).
a junior synonym of Microxina by Wiedenmayer (in Hooper & Synonymy. Microxina charcoti Topsent, 1916: 170; 1917:
Wiedenmayer, 1994), but its allocation there is debatable. The type 73, pI. 1, fig. 3, pI. 2, fig. 3, pI. 6, fig. 17; [in part] Burton, 1932b:
species, Gellius calyx Ridley & Dendy, 1887 (by original designa- 271, pI. 50, fig. 1 (under M. benedeni (Topsent, 1901a».
tion), has an external morphology close to that of Calyx (a thin Material examined. Lectotype (here designated): MNHN
walled hollow body), but the surface skeleton is closer to that of LBIM DT 226 - Antarctica, near Terre Alexandre, Baie Marguerite,
Haliclona (a loose, single layer of oxeas). Ridley & Dendy (1887) 200-297m. Paralectotypes: MNHN LBIM DT 692, 216, 217, 223,
describe it as having 'loose main bundles of spicules running in a 316, 317 (BMNH 1926.10.26.338 slides) - stn 8, Man-O-War
direction vertical to the surface of the sponge. At the surface these Anchorage, Funchal, Madeira. 0-25 m, 'Terra Nova' collection.
spicula project freely, giving it a characteristic hirsute appearance' . Other comparative material. Holotype of Microxina benedeni
The main skeleton resembles Gellius (a loose reticulation and some (Topsent, 1901a) (Fig. 7D): IRSNB 24 POR MOD I1- Antarctica,
loose bands of big oxeas, connected by free spicules, projecting 700 20'S, 83°23'W, 29.11.1898, 'Belgica' 1897-99, no. 725.
freely in a hispid surface), and microscleres are abundant sigmas. Description. Tubular and ramified from a narrow fibrous
Burton (1959), describing a specimen from Iceland as Haliclona base (Fig. 7 A), with a deep central aquiferous cavity ending in a
calyx (Ridley & Dendy, 1887), said that it coincided exactly with large oscule of the same diameter. Surface strongly spined by long
the holotype of Gellius calyx, except that it lacked sigmas. An alter- surface spines up to 10mm long (Fig. 7B). Ectosomal skeleton
native view, that Calyxadocia may belong to the subgenus Gellius is a tangential network of oxeas interrupted by the terminal tufts of
of genus Haliclona, is rejected because in the skeleton is more con- longitudinal primary fibres subdivided to form the surface spines
fused, subhalichondroid, and on this basis de Weerdt (pers. comm.) (Fig. 7B). Choanosomal skeleton with large and irregular meshes
suggests that it is closer to Hemigellius. The large oxeas of Gellius delimited by fasciculate, longitudinal primary multi spicular fibres,
calyx is also another clue as to its affinities with Niphatidae, where containing more than 15 spicules, packed compactly (Fig. 7C) and
these huge oxeas are never found in Chalinidae. Calyxadocia is con- issued from the base of the sponge, 296-426-800 fLm in diameter.
sidered a synonym of Hemigellius (taxonomic decision this work). Primary fibres may be irregularly split to form secondary fibres,
41-90-148 fLm in diameter. Meshes contain abundant microscleres
and free oxeas. No visible spongin. Megascleres: oxeas 1, strong,
MICROXINA TOPSENT, 1916 slightly curved with acerate points, 459-512-600 fLm long X
20-25-29 fLm in diameter; oxeas 2, 262-360-459 fLm long X
Synonymy 4-7-12fLm in diameter. Microscleres: microxeas, 45-73-95fLm
long X 3-4 fLm in diameter.
Microxina Topsent, 1916: 170; 1917: 72. ? Sigmaxinyssa Remarks. Topsent (1917) included his new genus Microxina
Kirkpatrick, 1907a: 272. in the Gelliinae, and made a comparison between Microxina and
Strongylophora. This genus was included in Gelliinae by Dendy
Type species (1905), and Topsent (1917) considered that both genera were close
due to the common possession of microxeas, and because of the
Microxina charcoti Topsent, 1916 (by monotypy). fibrous character of their skeletons. At the same time he also
remarked that both genera should probably be included with the
Definition axinellids. Later, in the same work, he gave up this idea because
Strongylophora was (in his opinion) less fibrous than Microxina, and
Niphatidae with strongly spined surface formed by the termi- because of the existence of other Gelliinae with a spined surface
nal tufts of primary fibres, with a tangential layer of free oxeas (e.g., Gelliodes). We consider that both genera are sufficiently differ-
between the spines, and microxeas are present. ent in their skeletal structures to be valid. Burton (1932b) considered
886 Porifera· Demospongiae • Haplosclerida • Haplosclerina • Nipbatidae

Fig. 7. Microxina Topsent, 1917. A-C, type species Microxina charcoti Topsent, 1916. A, tubular branched fragments of lectotype MNHN LBIM DT 226
and 216, with strongly spined surface and spines up IOmm long (scale I cm). B, terminal tufts of longitudinal primary fibres subdivided to form the sur-
face spines (scale 500 I-\-m). C,primary longitudinal choanosomal fibres with no visible spongin (scale 200 I-\-m). D, Microxina benedeni (Topsent, 190Ia),
holotype IRSNB 24 POR MOD II (scale 1 cm).

that Gelliodes benedeni and Microxina charcoti were subjective syn- Type species
onyms, due to their external form and spiculation, with the only
notable difference being that G. benedeni had only sigmas (and addi- Niphates erecta Duchassaing & Michelotti, 1864 (by subse-
tional raphides in var. fonior). By comparison, Microxina had only quent designation; Wiedenmayer, 1977b).
microxeas although Burton (1932b) also observed raphides.
Wiedenmayer (1989) regarded Hemigellius as a subjective synonym Definition
of Microxina. We also compared both genera and observed signifi-
cant differences between their respective type species, such as the Niphatidae with a paratangential ectosomal reticulation of
presence (in Hemigellius rudis, sensu Kirkpatrick, 1908c) of panicu- fibres or tracts, obscured by the conulose surface produced by the
late primary fibres and subectosomal 'embryonary holes', and differ- ends of primary longitudinal fibres.
ences in the continuous fibres or tracts observed in Microxina
charcoti. In addition, we found that the structure of the choanosomal Diagnosis
skeleton was compact and disorganised in Hemigellius rudis, but
clearly fibrous in Microxina charcoti. The external morphology, con- Massive-vasiform, funnel-shaped to tubular or ramose growth
sistency and surface skeleton are also different: Hemigellius is mas- forms (Fig. SA). Surface conulose to spiny, irregularly smooth to
sively lobate or thickly subcylindrical, whereas Microxina is tubular, rough, produced by primary longitudinal fibres ending on surface.
to massive or globular (e.g., M. benedeni; Fig. 7D). Hemigellius has Oscules numerous, irregularly distributed, flush with the surface
a clear surface palisade or a finely hispid surface, whereas in or slightly raised. Consistency spongy to rigid, tough, firm.
Microxina there is a fine membrane between the solid surface spines Ectosomal skeleton is a tangential network of secondary fibres,
producing a strongly spiny surface. obscured by protruding tufts of primary fibres. Microscleres are
rare sigmas. Choanosomal skeleton an irregularly diverging reticu-
lum of pauci- to multi spicular primary longitudinal fibres, abun-
NIPHATES DUCHASSAING & MICHELOTTI, 1864 dantly ramified, (Fig. SB) to form fibrofascicles and irregular
branches ascending to the surface. Interconnecting secondary
Synonymy fibres pauci- to multispiculate, well developed to form rounded
to irregular meshes. Spongin dominant, abundant and covering
Niphates Duchassaing & Michelotti, 1864: 93; de Laubenfels, the spicules (Fig. SC). Megascleres oxeas (Fig. SD). Microscleres
1936a (as unrecognisable). sigmata, present or absent.
Porifera· Demospongiae • Haplosclerida • Haplosclerina • Niphatidae 887

Fig. 8. Niphates Duchassaing & Michelotti, 1864. A-D, type species Niphates erecta Duchassaing & Michelotti, 1864. A, lectotype (fragments) ZMA
POR 1633, MT Por 51 , two branches with conulose surface and scattered oscules (scales 1cm). B, choanosomal skeleton with an irregularly diverging
reticulum of pauci- to multi spicular primary longitudinal fibres (scale 50 Ilm). C, ectosomal skeleton with polygonal meshes and multi spicular fibres
(scale 200 Ilm). D, oxeote to strongylote megascleres and abundant spongin (scale 20 Ilm).

Distribution Description (emendedfrom Van Soest, 1980). Ramose with

branches 15 cm long and 1.5-2 cm in diameter. Branches undivided
Caribbean: Bimini, Florida, St Thomas, Cura~ao, Bonaire; over most of their length. Oscules numerous, 4-7 mm, irregularly
Colombia; Barbados, Cuba (as Gelliodes ramosa), Tortola, scattered over branches. Surface smooth to conulose-spinous.
Viecques, Mexico, Jamaica (as Gelliodes aerolata). Saba, Puerto Consistency fragile, spongy, tough. Ectosomal skeleton an irregu-
Rico. Indo-west Pacific: New Caledonia; Papua New Guinea; lar to polygonal network of superficial multi spicular fibres extend-
Australia (decision for synonymy Van Soest, 1980). ing between the protruding ends of primary fibres. This network is
covered by protruding and ramified ends of primary longitudinal
Previous reviews fibres producing a conulose to spiny surface. Choanosomal skele-
ton an irregular reticulation with rounded to angular meshes
Duchassaing & Michelotti, 1864; Wiedenmayer, 1977b: 95; formed by radiating and fasciculate ascending longitudinal, multi-
Van Soest, 1980: 35; Desqueyroux-Faundez, 1984: 50; Fromont, spicular fibres. Meshes average between 139-435 f.Lm in largest
1993: 17. diameter. Primary fibres are 55-100 f.Lm in diameter, cored by 6-20
spicules. At the surface they are 200-500 f.Lm apart. Secondary
Description of type species or inteconnecting fibres are 25-45-90 f.Lm in diameter, cored by
4-15 spicules. Primary fibrofascicules 361-515-615 f.Lm in diame-
Niphates erecta Duchassaing & Michelotti, 1864 (Fig. 8). ter. Megascleres: Oxeas, slightly curved, oxeote and strongylote
Synonymy. Niphates erecta Duchassaing & Michelotti, forms are present, 99-139-154 f.Lm long, 2-5-6 f.Lm in diameter.
1864: 93 pI. 21. fig 3; Wiedenmayer, 1977b: 96, pI. 20, figs 7, 8, Microscleres: if present, sigmata, 12-15 f.Lm.
pI. 21, figs 1-4, text-fig. 119; Zea, 1987 (with additional Remarks. The precise species content of this genus, and
synonyms). indeed the family, remains uncertain. For example, many species
Material examined. Lectotype (designation Weidenmayer, presently included in Pachychalina have been referred to
1977b: 96, 271): ZMA POR. 1633 - St. Thomas. Paralectotype MT Amphimedon, with some originally described as Pachychalina of
Por 51 - St. Thomas. Schizoparalectotype: USNM 31018. Other authors more closely related to Callyspongia, e.g., Pachychalina
material. ZMA 3305 - Cura~ao, W of Piscadera. ZMA 3518 - near fibrosa Ridley & Dendy, 1887. Similarly, some Amphimedon
Carmabi. ZMA 3642 - outside of Jan Thiel Bay, Cura~ao. and Niphates may be better referred to Pachychalina, although
888 Porifera· Demospongiae • Haplosclerida • Haplosclerina • Nipbatidae

precisely which ones requires a more thorough revision. The pres- Description of type species
ence or absence of sigma microscleres in Niphates is considered an
unreliable generic character (see Van Soest, 1980; Fromont, 1993). Pachychalina rustica Schmidt, 1868 (Fig. 9).
Synonymy. Pachychalina rustica Schmidt, 1868: 8, pI. 2,
fig. 6; Topsent, 1901b: 357, pI. 13, fig. 4; 1938a: 6; 1938b: 31.
Material examined. Lectotype (here designated): MNHN
PACHYCHALINA SCHMIDT, 1868 LBIM DT47 - La Calle, Algeria, Expedition Lacaze-Duthier.
Paralectotype: MNHN LBIM DT25.
Synonymy Description. Lobate, clavi form, compact branches, growing
on a fragment of mollusc shell or on a calcareous pebble, small,
Pachychalina Schmidt, 1868: 8; Ridley & Dendy, 1887: 19. 2-3 cm high, 1-2 cm wide. Surface with numerous orifices, conu-
lose or strongly spiny in appearence (Fig. 9A). Without surface
Type species membrane. Consistency elastic, compressible, due to its strongly
developed aquiferous system with large irregular canals, limited by
Pachychalina rustica Schmidt, 1868: 8 (by monotypy). spongin plates. Ectosomal skeleton not clearly visible; in the type
material it contains free spicules and fragments of fine tracts, pro-
Definition ducing a tridimensional network (Fig. 9B). Choanosomal skeleton
irregular and confused, formed by strong multi spicular fibres with-
Niphatidae with a paratangential ectosomal reticulation out preferent orientation in the skeleton, large and repeatedly
of fibres or tracts obscured by the irregularly, conulose to spiny longitudinally divided, ultimately forming isolated and finer rami-
surface, pierced by abundant aquiferous orifices. fications (Fig. 9C). All fibres are multi spicular, with more than
15 spicules, no visible spongin sheath. Diameter of primary fibres
Diagnosis 100-272-440 fLm, secondary fibres are not connecting fibres
but free within the choanosomal skeleton, 40-60-80 fLm, tertiary
Columnar, clavate, subcylindrical growth forms (Fig. 9A), fibres paucispicular, free in the terminal parts of secondary tracts,
having a well-developed aquiferous system with large canals 20-30 fLm diameter. The interfibre spaces contain few free
rendering the texture highly compressible. Numerous small rounded spicules. Megascleres: from type material, 138-171-195 fLm long,
aquiferous openings not clearly differentiated into oscules or pores. 3-5-6 fLm wide. Topsent (1938b), 190-240 X 4-7 fLm, with most
Some of these openings are lengthened and divided by a fine wall 215-225 X 5-6 fLm (see Table 1).
with irregular edges, especially at the high part of the sponge Re11Ulrks. Lendenfeld (1887c: 775) provided an emended
where pores are very close to one another. At this location the sur- diagnosis of Pachychalina and described ten new species from
face has a very irregular character due to the numerous aquiferous Australia, none of which has been re-examined. Dendy (1905)
orifices and because the surface lacks a membrane. Ectosomal included Pachychalina in the subfamily Chalininae, but recognised
skeleton (Fig. 9B) appears to be undifferentiated from the that genera included in this group (viz., Pachychalina, Chalina,
choanosomal skeleton, and is extremely irregular because of the Ceraochalina and Siphonochalina), were impossible to define
numerous aquiferous orifices at the surface. No ectosomal mem- sharply. Some of these species are actually Callyspongia, such as
brane is present. Ectosomal skeleton with rounded meshes and free Pachychalina (pro Callyspongia) spinilamella Dendy, 1889. Burton
spicules, three-dimensional, irregular and confused, between the (1929a) considered Pachychalina incertae sedis as he was uncertain
ends of primary fibres and aquiferous canals. Choanosomal skele- as to what 'Schmidt intended it to include'. The type specimen, in
ton is confused, ascending, slightly parallel multispicular primary Burton's (1928) opinion, was 'for the moment, apparently irretriev-
fibres of irregular diameter, with compact spicules and no visible ably lost, and not available for re-examination' . Nevertheless, Topsent
spongin binding them. Primary fibres are obliquely branched to (1901b) found in MNHN collections, two small specimens, which are
form secondary lines, with decreasing diameter towards the high probably the specimens we have studied in this work, but he did not
part of the sponge (Fig. 9C). The choanosomal skeleton is highly describe them providing only an illustration (Topesent, 1901b: pI. 13,
lacunar due to the aquiferous canals, with lacunae as large as the fig. 4) and spicule dimensions (which are similar to those observed in
narrow spongin walls between them. Spongin not visible. the type material re-examined). Wiedenmayer (1977b) included with
Megascleres: oxeas, irregularly bent, with short apices and regular doubt Pachychalina in synonymy with Niphates. He remarked that
diameter (Fig. 9D). Microscleres: absent. the surface of Pachychalina, illustrated by Topsent (1901 b), was very
similar to that of West Indian Niphates but without a developed sur-
Distribution face reticulum. Pachychalina, was subsequently included as a subjec-
tive junior synonym of Amphimedon by Van Soest (1980: 34) and
Mediterranean; Australia (many recorded as Cribrochalina followed by Hooper & Wiedenmayer (1994). Van Soest (1980) sup-
and Amphimedon, in Hooper & Wiedenmayer, 1994). Chile-Peru ported his decision by the fact that Pachychalina has probably been
and Magellan region; West Indies, Bahamas, St Thomas, compromised in the older literature, and consequently a doubtful
Guadeloupe, New Providence, (as Amphimedon [part]), Boreal taxon although a well-established genus in the sense of the ICZN. The
East Pacific, Red Sea, Indian Ocean, Tasmania. (Van Soest, person- knowledge of the real taxonomic features of Pachychalina sensu
nal communication, in part.) stricto, from re-examination of the type material, reveals several
distinctive features and indicates that the genus is valid within
Previous reviews Niphatidae. Future work is required to determine the species and geo-
graphic scope of the genus, requiring re-examination of all nominal
Topsent, 1901b: 357; 1938a: 6; 1938b: 31; Burton, 1929a: 505. species assigned to the various niphatid genera.
Porifera· Demospongiae • Haplosclerida • Haplosclerina • Nipbatidae 889

Fig. 9. Pachychalina Schmidt, 1868. A-D, type species Pachychalina rustica Schmidt, 1868. A, lectotype MNHN LBIM DT 747 (scale I cm). B, ectoso-
mal skeleton extremely irregular undifferentiated from choanosome (scale 50 !Lm). C, choanosomal skeleton with an irregular network of confused ascend-
ing, slightly parallel multi spicular primary fibres obliquely branched to form secondary lines (scale 50 !Lm). D, oxeas (scale 5 !-Lm).

Pachychalina differs from Amphimedon, Gelliodes and Pachychalina possesses a three-dimensional surface skeleton
Niphates as follows. largely undifferentiated from the main skeleton, and with no surface
It differs from Amphimedon as it lacks a clear ectosomal membranous organic layer distinguishable from the subdermal por-
tangential network or membraneous ectosomal skeleton. In tion. In Gelliodes there is a light organic surface layer and the ecto-
Amphimedon there is a regular tangential skeleton of secondary somal skeleton is a tangential system of multi spicular fibres through
fibres with regular rounded meshes, overlying the choanosomal which the protruding ends of primary fibres are clearly observed.
canals and covered by a fine membranous organic layer. In addi- Niphates is also similar to Pachychalina but has a more regu-
tion, the choanosomal skeletal reticulation of Pachychalina is lar skeleton with secondary fibres well-developed to form a regular
peculiar in having an almost tertiary structure, with thick primary rectangular reticulation. Their surface skeletons are also similar,
tracts, thinner irregular very thin interconnecting tracts, and with being three-dimensional and not well-separated from the choanoso-
loose spicules in between. By comparison, in Amphimedon the mal skeleton in Pachychalina, compared to Niphates which has a
choanosomal skeleton has a regular system of main and secondary surface that is conulose to spiny, and irregularly smooth to rough
fibres which do not differ much in diameter. In Pachychalina the produced by primary longitudinal fibres ending at the surface, and
whole skeleton is rather confused, with the choanosomal skeleton well differentiated from the choanosomal skeleton. Consistency in
irregular, diffuse, and occurring as a radially plumose network of both genera is also different. Niphates is spongy to rigid, tough,
primary, ramified, multi spicular fibres, irregularly connected by no firm, whereas Pachychalina has an extremely compressible texture.
proper secondary multi spicular fibres and no visible spongin. Choanosomal skeletons in Pachychalina and Niphates differ where
The skeletons of Pachychalina and Gelliodes have both the former has a skeleton containing irregular fibres, without a vis-
similarities and differences, with the former having an emphasis ible spongin sheath, and highly lacunar, and Niphates has pauci- to
on the main fibres, producing a coarse skeleton with secondary multispicular fasciculated primary and interconnecting fibres,
tracts thinner and ill-defined and with loose spicules, and lacking which are well-developed and spongin is dominant, abundant, and
sigmas which are present in Gelliodes (Fig. 9C). In addition, covers the spicules.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Suborder Petrosina Boury-Esnault & Van Beveren, 1982

Rob W.M. Van Soest1 & John N.A. Hooper

Petrosina Boury-Esnault & Van Beveren (Porifera, Demospongiae) are here defined in new content following earlier proposals for a
subdivision of Haplosclerida at the ordinal level suggested by Bergquist (1980a). In the present concept, Petrosina Boury-Esnault &
Van Beveren, 1982 are marine Haplosclerida characterized by the possession of an isotropic skeleton, in which spicule tracts or fibres lack
orientation. In this respect they differ from the suborder Haplosclerina, which has the skeleton organized in primary ascending and sec-
ondary interconnecting tracts or fibres. Spicules are frequently (but not exclusively) smooth oxeas and/or strongyles, in several size cate-
gories (Haplosclerina have only a single size category). A further difference which appears to be valid is demonstration of oviparous
reproduction (Haplosclerina are probably all viviparous). Petrosina differ from freshwater sponges, here united in the suborder
Spongillina Manconi & Pronzato, in having a tangential ectosomal skeleton which frequently takes the form of a crust of intercrossing
spicules or a reticulation of spicules of smaller size than those of the choanosomal skeleton. Petrosina differ also from Spongillina in lack-
ing gemmules and special microscleres protecting these asexual resting bodies. Also, spicules in Spongillina are frequently spined. Three
families are recognised: Calcifibrospongiidae, Petrosiidae and Phloeodictyidae, differing in their habit and in the organization of the
skeleton. Calcifibrospongiidae contains a single species of coralline sponges with a unique aragonitic limestone basal skeleton and a loose
skeleton of thin strongyles. Petrosiidae include sponges with an alveolar choanosomal skeleton made by usually thick spicule tracts.
Phloeodictyidae have a choanosomal reticulation of single spicules strengthened by independent thick and long tracts of spicules.
Keywords: Porifera; Demospongiae; Haplosclerida; Petrosina; Calcifibrospongiidae; Petrosiidae; Phloeodictyidae.
-~- --- - -~- - -~ -- -- - - - - - -

DEFINITION, DIAGNOSIS, SCOPE (Le., rounded meshes made by thick spicule tracts), are especially
common in some petrosiid genera. In phloeodictyids, the
Synonymy choanosomal skeleton consisting of an isodictyal uni- or paucispic-
ular reticulation is strengthened by apparently independent long
Nepheliospongida Bergquist, 1980a; Petrosida Boury- and thick spicule tracts forming their own anastomosing system, or
Esnault & Van Beveren, 1982; Hartman, 1982. remain clear of other such spicule tracts over large distances. These
spicule tracts also run subdermally in fistules as an independent
Definition supporting structure. Spicules consist of smooth diactinal mega-
scleres (oxeas or strongyles), and an erratic and presumably vestig-
Marine Haplosclerida with an isotropic choanosomal skeleton ial presence of microscleres, usually sigmas and/or toxas. In this
lacking differentiation in primary and secondary fibres or tracts; suborder, microxeas or microstrongyles are recognized as a special
spicules smooth oxeas and/or strongyles in a single, or up to five microsclere, although they may in fact be part of a series of
size categories. increasingly smaller megasclere categories.

Diagnosis Scope

Sponges with thickly encrusting, cushion-shaped, massive In the concept of Petrosina presented here, three families are
repent-ramose, erectly ramose, vase-shaped, tubular, lamellate, included: Calcifibrospongiidae Hartman, 1979, Petrosiidae Van
fan-shaped or turnip-shaped with main body buried in the substrate Soest, 1980 and Phloeodictyidae Carter, 1882. Together these com-
and often long fistules sticking out above the sediment. One genus prise a complement of sponges occurring all over the world in all
has a basal limestone skeleton, and two (?) genera are excavating, marine habitats. Calcifibrospongiidae are monotypical, the other
with often long epilithic papillae or tubes. Consistency usually two families are speciose. Tropical shallow-water marine habitats
hard or firm, barely compressible, but crumbly, brittle and fragile such as reefs harbour the majority of petrosine sponges, but
sponges are not uncommon. Surface usually smooth, but may be phloeodictyids especially are also frequently observed in sediment-
ridged or honeycombed occasionally. Ectosomal skeleton uni- or rich environments, including cold- and deep-water habitats.
polyspicular, crust-like, (lacking a well-organized triangular reticu-
lation of single spicules or neat polygonal meshes, such as found in
Haplosclerina). The choanosomal skeleton is isotropic, which TAXONOMIC HISTORY
means there is no orientation of the reticulation, with pauci- or
multispicular primary or ascending fibres or tracts forming square, For a more extensive history of Haplosclerina we refer to the
rounded or irregular polygonal meshes. In some petrosines, an chapter on the order Petrosina (Van Soest & Hooper, this volume). In
irregular, vague anisotropic arrangement appears to be super- summary, two types of skeletal structure have emerged ('renierids'
imposed on the isotropic reticulation. Alveolar skeletons and 'chalinids') from the diffuse earlier classifications of sponges

891
892 Porifera· Demospongiae • HapJosclerida • Petrosina

now classed in Haplosclerida. These are now formalized at the sub- petrosins may be part of the radiate-accretive (sensu Wiedenmayer,
ordinal level, with Petrosina largely comprising the former 1977a) growth pattern, which in petrosins may be secondarily
'renierids'. Since Reniera is a subgenus of Haliclona (see contribu- (derived) obscured by further silicification, resulting in the isotropic
tion in this volume on Chalinidae by De Weerdt), any reference to pattern. Alternatively, the isotropic choanosomal skeleton may be
this in the name of the present suborder needs to be avoided. derived from a more widespread anisotropic skeletal arrangement,
Bergquist (l980a), on the basis of assumed ovipary and distinct evidenced by its vaguely detected presence in species of Neopetrosia.
chemistry, distinguished a restricted order Nepheliospongida, cov- The ectosomal skeleton according to De Weerdt (1989) is usu-
ering more-or-less the same groups. It was demonstrated subse- ally crust-like with tangential spicules intercrossing. It is rather dif-
quently that the fossil sponge Nepheliospongia, which formed the ficult to define the ectosomal skeleton of petrosine sponges due to
basis for the order Nepheliospongida, lacked spicules and could not the large variation observed. The neatly meshed ectosomal skele-
be grouped together with the recent genera Petrosia and Calyx, orig- tons ofhaplosclerine sponges do not occur in petrosines, but this is
inally assigned to this order. Boury-Esnault & Van Beveren (1982), not operational as a distinct synapomorphy.
and independently Hartman (1982), proposed to rename the order The occurrence of several distinct size categories of megascle-
Petrosida to avoid confusion with Nepheliospongia. Since Hartman res in various genera of this suborder is unique among the
(1982) excluded Phloeodictyidae (as Oceanapiidae) from Petrosida, Haplosclerida, but unfortunately it is not of general occurrence as
authorship for the present concept of Petrosina is given to Boury- many petrosine genera possess only a single category (e.g.,
Esnault & Van Beveren (1982). Neopetrosia, Xestospongia, Aka, many species of Oceanapia).
Microscleres have the same haphazard distribution in individ-
ual species as found in Haplosclerina. They are considered to be
REMARKS remnants from an ancestral broad parentage including poeciloscle-
rids, and their occurrence does not make them useful markers for
Synapomorphies taxa above the species level.
In spite of the paucity of genuine synapomorphies, De
The ovipary of Petros ina was upheld in a recent study Weerdt's (1989) character analysis of the five families of marine
(Fromont & Bergquist, 1994), but the chemical distinctness (pos- Haplosclerida (Spongillina were excluded) suggested that two dis-
session of sterols with a cyclic sidebranch) could not be confirmed, tinct clades exist, one comprising Chalinidae, Callyspongiidae and
since many species appear to lack them and several non-petrosines Niphatidae, the other Petrosiidae and Phloeodictyidae. Thus, a
possess structurally similar molecules (e.g., Fromont et al., 1994; division of the order Haplosclerida into suborders Haplosclerina
Van Soest & Braekman, 1999). Ovipary is shared with many and Petrosina, as proposed here, is supported by a formal character
tetractinomorph families and with verongids, and thus this makes a analysis but the morphological synapomorphies that support these
poor synapomorphy. The known chemistry so far provides evi- clades remain vague and partially elusive.
dence of the existence of genus markers (e.g., for Petrosia) but
Petrosina-wide occurrence of the same compound family has not Previous reviews
(yet) been demonstrated.
The anisotropic skeletal arrangement seen in haplosclerines, Bergquist, 1980a; Van Soest, 1980; Desqueyroux-Faiindez,
and in the periphery (e.g., tops of tubes, tips of branches) of some 1987; De Weerdt, 1985, 1989; Fromont, 1991; Fromont etal., 1994.

KEY TO FAMILIES

(1) Coralline sponge with basal limestone skeleton and a reticulation of thin strongyles ....................................... Calci6brospongiidae
No basal limestone skeleton ................................................................................................................................................................. 2
(2) Choanosomal skeleton an isotropic unispicular skeleton, causing a 'pulpy' macroscopical texture, strengthened by long and
thick spicule tracts running independently through the choanosome and forming an anastomosing subdermal supporting
system ................ .......... ................................ ................... ........ ................... ........ ................................ ...... .......... ......... Phloeodictyidae
Choanosomal skeleton an isotropic reticulation of pauci- or multi spicular tracts or fibres making squarish or rounded meshes and
lacking a differentiation in primary ascending and secondary interconnecting tracts ........................................................ Petrosiidae
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Phloeodictyidae Carter, 1882

R. Desqueyroux-Faundez 1 & C. Valentine2

I Museum d'histoire naturelle P.O. Box 6434, CH-1211 Geneva 6, Switzerland. (ruth.faundez@mhn.ville-ge.ch)
2 The Natural History Museum, Cromwell Road, London SW7 5BD, UK. (c.valentine@nhm.ac.uk)

Phloeodictyidae Carter (Demospongiae, Haplosclerida), including Akaidae Alander, Oceanapiidae Van Soest, is distinguished from other
haplosclerids primarily in having a detachable ectosomal crust, an isotrophic reticulation of single spicules, and a concentric laminated
internal structure. Species commonly have fistules and tubes, an adaptation to their common habitat of burrowing or being buried in the
substrate. Megascleres are oxeas or strongyles, microscleres, if present, are sigmas and/or toxas. Species are distributed in shallow and
deeper tropical waters; Calyx and Pachypellina have been also recorded from the North Atlantic, Antarctica and subantarctic regions, and
Oceanapia are known worldwide.
Keywords: Porifera; Demospongiae; Haplosclerida; Phloeodictyidae; Aka; Calyx; Oceanapia; Pachypellina; Tabulocalyx.

DEFINITION, DIAGNOSIS, SCOPE with pustular eminences only inside. Structure essentially laminated
and concentric; laminae of two distinct kinds, which may alternate
Synonymy with each other in variable plurality, commencing dermally with an
apparently homogeneous, fine, isodictyal layer, densely spiculous;
Phloeodictyidae Carter, I 882b: 117. Akaidae Alander, 1942: followed by a coarse, open spiculo-fibrous reticulated one, whose
27. Oceanapiidae Van Soest, 1980: 80. lamination is parallel to the surface, and contrasts strongly in struc-
ture ... with the isodictyal homogeneous one outside; also strongly
Definition with the tissue of sponges generally, whose reticulated structure is
continuous with the deepest portion, and not laminar like the bark of
Haplosclerida with an ectosomal tangential network of free a tree, as in this instance. Internally hollow or more-or-less filled
oxeas or strongyles. Choanosomal network is an isotropic reticula- with a pulpy isodictyal tissue like the outer lamina, i.e. composed of
tion of free spicules, reinforced or not by an irregular network of sarcode (=choanosome) densely charged with the spicule of the
fibres or tracts. species. No evident oscula. Pores in the dermal layer. Spicule of one
kind only, viz. acerate (=oxea), sometimes accompanied by a biha-
Diagnosis (emended from Hooper & Wiedenmayer, mate (=sigma) flesh spicule (fibula).
1994: 332) The history of the family is a relative mess. Carter (1882b)
based his diagnosis on a species described by two different authors
Encrusting, massive, lobate or more frequently spherical or under two different names: Bowerbank (1866), as Isodictya
tubular growth forms, buried in the substrate, with fistules on the robusta and Desmacidon jeffreysii, and Norman (1869b) as
surface, occasionally excavating coralline substrates. The ectoso- Oceanapia jeffreysii, which he synonymised (/. robusta being the
mal skeleton is a multilayered irregular tangential reticulation of most senior name), and assigned to Oceanapia. Carter (1882b) also
oxeas or strongyles, forming a distinct usually detachable crust. recognised Rhizochalina Schmidt, 1870, as a junior subjective syn-
The choanosomal skeleton is an irregular reticulation of multispic- onym of Oceanapia, and in the same work (1882b: 120) erected
ular tracts of diactinal spicules with or without spongin, together Phloeodictyon which he considered different from Oceanapia,
with an irregularly dispersed isotropic reticulation of single despite some obvious similarities between both ('at first sight
spicules scattered between these major tracts. Microscleres may Phloeodictyon looks very much as Isodictya but, on dissection
include microangulate sigmas and toxas. is found to be similar in structure to Desmacidon jeffreysii').
Phloeodictyon Carter was considered by de Laubenfels (1936a) to
Scope be a junior synonym of Oceanapia and Rhizochalina carotta
Schmidt presumably transferred to Oceanapia.
Ten nominal genera have been included in this family (Aka, Ridley & Dendy (1887) conserved the taxon Phloeodictyina at
Biminia, Calyx, Foliolina, Oceanapia, Pachypellina, Pellina, subfamily level and assigned species of Rhizochalina and Oceanapia
Phloeodictyon, Rhizochalina and Siphonodictyon), of which five to it. Dendy (1922b) retained Phloeodictyinae as a subfamily of
are recognised in this work: Aka, Calyx, Oceanapia, Pachypellina Haploscleridae, including in it the genera Phloeodictyon, Oceanapia
and Tabulocalyx. and Rhizochalina. He erected a second subfamily Coelosphaereae
for Coelosphaera, Siderodermella and Amphiastrella, to which
History and biology de Laubenfels (1936a) assigned otherwise typical Phloeodictyidae
genera Oceanapia, Rhizochalina and Phloeodictyon, based exclu-
The group Phloeodictyina, was proposed by Carter (1882b) to sively on their external form. As a consequence of this mess, it
include 'sponges of variable form, chiefly globular, accompanied is not surprising that following Bergquist (1965), there has been
by tubular expansions both above and below, or above only, which a great confusion concerning the synonymy of Oceanapia,
are closed at the extremities, simple, or branched; ... or vasiform, Rhizochalina and Phloeodictyon, and their alleged relationships to

893
894 Porifera· Demospongiae • HaploscIerida • Petrosina • Phloeodictyidae

Coelosphaeridae and Adociidae. Levi (1973) included most of the and Desqueyroux-Faundez (1987). Aka was originally included
genera now considered to belong to Phloeodictyidae in Renieridae, by authors in Clionidae (=Clionaidae) (Hancock, 1849), but
viz., Pellina, Rhizochalina, Oceanapia and Calyx. Riitzler & Stone (1986) recognised in it characters described for
Based on the premise that the name Phloeodictyidae had not Siphonodictyon (Phloeodictyidae), and consequently referred
been in regular use Van Soest (1980: 80) established Oceanapiidae, Aka to this family and as a senior synonym of Siphonodictyon.
which until recently has been widely accepted by contemporary There is still some debate as to the validity of this synonymy,
authors. Unfortunately, however, the name Phloeodictyidae has which is discussed further below.
seniority, irrespective of the synonymies of their respective type
species (Hooper & Wiedenmayer, 1994). Van Soest (1980) also Distribution
included Pellina in his new family Oceanapiidae, but this was later
modified by de Weerdt (1985) and de Weerdt & Van Soest (1986), Nominal genera have the following recorded distributions
based on the absence of oceanapiid characters in the type species, (after Van Soest, pers. comm.): Chile-Peru and Magellanic region
Halichondria semitubulosa Lieberkiihn (1859), which conforms to (Calyx, Oceanapia, Pachypellina), Brasilia (Calyx, Oceanapia),
Halichondria (see chapter on Halichondriidae). In the same work, West Indies (Aka, Calyx, Foliolina, Oceanapia, Rhizochalina,
de Weerdt & Van Soest (1986) proposed a new subgenus, Pellina), Boreal East Atlantic (Aka, Oceanapia), West Africa (Aka,
Rhizochalina Schmidt, 1870, to include species of Oceanapia, like Calyx, Oceanapia), South Africa (Oceanapia), Mediterranean
O. oleracea the type species of Rhizochalina, and three others that Atlantic (Aka, Calyx, Oceanapia), Central Pacific (Aka,
differ from typical pulpy Oceanapia. Siphonodictyon, Oceanapia), Japan, China (Oceanapia, Biminia),
Definitions of taxa belonging to this family are provided by Antarctica (Calyx, Vagocia, Oceanapia), New Zealand
Norman (1869b), Carter (1882b), de Laubenfels (1936a), Bergquist (Oceanapia, Vagocia). Species of Phloeodictyidae are absent in
(1965), Fromont (1991, 1993), Riitzler (1971), Van Soest (1980) Boreal- West Atlantic and Boreal- East Pacific.

KEY TO GENERA

(1) Burrowing into limestone substrate, secreting a layer of mucus, only the fistules are externally visible ...................................... Aka
Non-burrowing sponges ....................................................................................................................................................................... 2
(2) Variable growth form, with fistules ..................................................................................................................................................... 3
Massive globose or tubulose, without fistules ..................................................................................................................................... 4
(3) Hollow body, compact external ectosomal crust, or massive, encrusting from a basal lamina with spreading fistules, oscules only at
the end of fistules, or no oscules. Ectosomal skeleton a single tangential layer of spicules or multilayered compact crust.
Choanosomal skeleton a network of spicule tracts, abundantly ramified irregular-oval meshes, abundant free oxeas. Microscleres may
be present, sigmata and/or toxas .......................................................................................................................................... Oceanapia
Massive to cuplike, coalescent tubes, globose or lamellate, with a short stalk; fistules may be present, oscules in the internal part of
the cup. Ectosomal skeleton a compact network, with a surface-membrane, scattered free spicules, and numerous sphaerulous cells.
Choanosomal skeleton a network of long, entangled spicule tracts, longitudinal in the interior of the body and tangential to the surface
in the peripheral skeleton, with a connecting unispicular reticulation. Spicules oxeas, microscleres if present, toxas ............... Calyx
(4) Massive globose, with a stalk, no fistules, oscules only on the globous body. Ectosomal crust in two separate layers. Choanosomal
skeleton a mass of oxeas in confusion and some irregular multispicular tracts ................................................................ Tabulocalyx
Massive globose or tubulose. Ectosomal skeleton a single tangential crust, no stalk, no fistules, no oscules. Choanosomal skeleton
densely spiculous with large meshes ................................................................................................................................ Pachypellina

AKA DE LAUBENFELS, 1936 Diagnosis (from Johnson, 1899 and Thomas, 1968c)

Synonymy Boring sponges, excavating calcareous substrates (molluscs,

coral rocks); externally visible by their fistulose tubes. Skeleton
[Acca] Johnson, 1899: 461, figs 1-4 (preocc.). Aka de formed only of spicule-bundles in sinuous bands, not in a network,
Laubenfels, 1936a: 155. lining walls of fistules. Spicules short, slender, curved oxeas.
Without microscleres.

Type species
Previous reviews
Aka insidiosa Johnson, 1899 (by subsequent designation;
de Laubenfels, 1936a: 155). Thomas, 1968c, 1979; Riitzler & Stone, 1986; Fromont, 1993.

Definition (from Johnson, 1899)

Description of type species
Excavating sponges boring calcitic substrata. Spicules only
oxeas forming sinuous bands arranged in bundles, producing Aka insidiosa Johnson, 1899 (not figured).
fistulose tubes. Synonymy. Acca insidiosa Johnson, 1899: 461, figs 1-4.
Porifera· Demospongiae • Haplosclerida • Petrosina • Phloeodictyidae 895

Material examined. Holotype: lost (Rtitzler & Smith, Distribution

pers.comm.)
Description (emendedfrom Johnson, 1899; Thomas, 1968c; Species of Aka (as Clionaidae) have been described from
Van So est, unpublished data, pers.comm.). Found on shells of Great Britain, (Hancock, 1849), Madeira (Johnson, 1899), Coast of
Ostrea and Chama, lining holes about 1.5 mm in diameter and issu- Ireland (Stephens, 1915a, as Cliona coralliophaga), West-Central
ing as short tubes from them. The walls of the tubes are composed Pacific (de Laubenfels, 1954), Gulf of Manaar, and Mozambique
of short, small, slender oxeas about 100-200/Lm long, 4-11/Lm Channel (Thomas, 1968c, 1979) and West Indies (Alcolado, 1994).
in diameter, grouped to form sinuous bands of spicules arranged Fossil records of Aka, are Middle and Western Europe, and
in bundles, but not forming a network. The tops of the tubes ArizonaINorthern Mexico, according to Reitner & Keupp (1991b).
are covered over with a sieve-like reticulation also formed
of spicules in bundles. The spicules are short, cylindrical oxeas,
slender, slightly curved. Colour of the sponge when dry, pale CALYX VOSMAER, 1885
brown.
Remarks. De Laubenfels (1936a: 155) replaced Johnson's Synonymy
preoccupied name and nominated its type species A. insidiosa
Johnson, 1899 (burrowing in molluscs). He also included in the [Lieberkuhnial Balsamo-Crivelli, 1863: 293; Schmidt, 1864:
genus Cliona coralliophaga Stephens, 1915a (boring coral) and 43; Gray, 1867: 520 (preocc. for Lieberkuhnia Claparede &
Cliona labyrinthica Hancock, 1849 (boring Tridacna gigas). Lachman, 1859, Rhizopoda (Protozoa». [Lieberkuehnial de
Thomas (1968c) included Aka in Clionaidae, provided a diagnosis Laubenfels, 1936a: 133 (lapsus of [Lieberkuhnial Balsamo-
of the genus, and described a new species from the Gulf of Manaar, Crivelli). Calyx Vosmaer, 1885b: 337. Vagocia de Laubenfels,
Aka diagonoxea, which he considered was related to Cliona 1936a: 133. Taxonomic decision for synonymy: Wiedenmayer, in
labyrinthica. Riitzler & Stone (1986) studied the microscope Hooper & Wiedenmayer (1994).
slide collections of Hancock (1849, 1867), consisting of excavating
sponges from the British coasts and other unspecified localities. Type species
From re-examination of spicules of Cliona labyrinthica they
concluded that Aka was a senior subjective synonym of Reniera ? calyx Schmidt, 1862: 76 (by monotypy) (=Calyx
Siphonodictyon Bergquist, 1965, but since Johnson's original nicaeensis (Risso, 1826: 372), see Topsent, 1925c: 708).
material of the type species, A. insidiosa, is lost, it is impossible
to establish the exact differences between Aka s.s. and Definition
Siphonodictyon from a comparative analysis of the skeleton of the
type species of both genera. In this work, we follow Van Soest Phloeodictyidae with massive to cuplike or lamellate, globose
(pers. comm.), to establish the modem concept of Aka as revived growth forms, supported by a short stalk, or stout coalescing tubes
by Rtitzler & Stone (1986) and by Van Soest's own studies of his growing together. Smooth surface and fibrous choanosome.
collections. Bergquist (1965) erected Siphonodictyon, which she Ectosomal skeleton a multilayered unispicular network with spon-
considered closely related to Phloeodictyon Carter, 1882b, but dif- gin at nodes. Choanosomal skeleton with entangled longitudinal
fered in the great quantities of mucus produced and by the absence tracts in the interior and tangential tracts closer to the periphery.
of a bark-like ectosomal region. Bergquist (1965) remarked that Megascleres oxeas, microscleres if present toxas.
she followed Dendy (1922b), taking into consideration morpholog-
ical details, to include Siphonodictyon in Adociidae. By compari- Diagnosis
son, morphological characters in Aka are reduced to only a few
remarks in Johnson's description of Acca insidiosa. Bergquist Massive, caliculate (Fig. 2A), short pedunculate, lamellar
(pers.comm.) considers that Siphonodictyon is not unequivocally (Fig. 3A, C, Calyx arcuarius as Vagocia) or coalescent tubular
the same as Aka. growth forms, with tubular processes or fistulae may be present.
Van Soest's (pers. comm.) concept of Aka includes boring Surface consists of a multilayered, compact ectosomal network,
sponges with a skeleton of short oxeas, and papillae with oxeas in a under the surface-membrane (Figs 2B, 3B), with a close-knit
reticulate arrangement. Inside the coral, the skeleton becomes unispicular network containing spongin at nodes and scattered free
reduced, consisting of a pulpy organic mass. These characters are spicules with numerous sphaerulous cells. These cells, brown pig-
similar to those of Siphonodictyon mucosum, and differences mented or uncoloured, give to the sponge its dark or clear brown
between species of both genera are reduced to the size of papillae. colour. Small oscules present in the internal part of the calyx
For example, papillae in specimens of Aka infesta collected infest- (Fig. 2C), or absent. Choanosomal skeleton composed of long, dis-
ing Norwegian deep water corals exhibited small papillae (1 mm orientated, entangled spicular tracts, longitudinal in the interior of
or so), whereas in S. mucosum they measured about 5 cm. the body (Fig. 2D), and tangential to the surface in the periphery
Following Rtitzler & Stone's (1986) and Hooper & Wiedenmayer's (Fig. 2B). Between the fibres there is a connecting unispicular retic-
(1994) opinions (the latter based on the former's evidence), in ulation (Figs 2D, 3D). Spongin in variable amount. Megascleres are
this work we consider that it is likely there is insufficient evidence oxeas, of two size categories, bent at the center, with conical points.
to date to differentiate Siphonodictyon from Aka, but we do not Microscleres toxas (Fig. 3D, in Calyx arcuarius).
make a formal decision about this alleged synonymy, which we
leave to future studies, especially on the diverse Indo-west Pacific Previous reviews
fauna (see Siphonodictyon; Fig. 1A-D; based on USNM 23697,
holotype of Siphonodictyon mucosa Bergquist, 1965, Palau, Balsamo-Crivelli, 1863: 293 (Lieberkuhnia calyx); Carter,
Micronesia). 1882b: 116 (Reniera crateriformis); Topsent, 1913b: 638
896 Porifera· Demospongiae • Haplosclerida • Petrosina • Phloeodictyidae

Fig. 1. Siphonodictyon Bergquist. 1965. A-E, type species Siphonodictyon mucosa Bergquist, 1965, USNM 23697, holotype. A , external basal mass con-
taining the gelatinous body of the sponge (scale I em). B, gelatinous body at the interior of the cavity (at left, part of one of the oscular tubes ; scale 1 em).
C, fragment of the ectosomal network of the oscular tube or fistule (scale 50/Lm). D-E, view of the choanosomal network and stout primary longitudinal
fibres (scale 50 /Lm).

(Gellius arcuarius); 1925c: 708; 1928e: 2, figs 1-5 (Calyx nicaeen- Description. Caliculate or vase-shaped, massive, sponge,
sis); de Laubenfels, 1936a: 133; Wiedenmayer, 1977b: 20, 79, Ill, 25 cm high, fixed to the substrate by a short peduncle. Texture hard,
112, 116, 1I8, 119, 121, 156; Hooper & Wiedenmayer, 1994: 333. compact, colour dark brown, greenish. Surface even, rough, or if
missing, the compact stout fibres of the body wall are visible at the
Description of type species surface. Oscules not visible in the external part of the body wall,
some oscules are visible in the cavity of the body. Choanosomal
Calyx nicaeensis (Risso, 1826) (Fig. 2). skeleton: primary multispicular fibres are compact, very variable in
Synonymy. Reniera? calyx Schmidt, 1862: 76 (=Spongia diameter, from 10 to more than 100/Lm, irregularly distributed,
nicaensis Risso, 1826: 372; see Topsent, 1925c: 708). Calyx longitudinally, they form a large irregular mesh, in the interior
lieberkuhni Vosmaer, 1885b: 337 (=Lieberkuhnia calyx (Nardo) of the mesh there is a unispicular network and a few free spicules
Balsamo-Crivelli, 1863). Esperia calyx Nardo, 1863 is a nomen are visible. Spicules are oxeas of two size categories (Fig. 3D,
nudum). [Lieberkuhnia] calyx Balsamo-Crivelli is both preoccu- Vagocia), 99-127-147/Lm long by 5-6mm in diameter and
pied and an unjustified replacement name, so Reniera ? calyx 35-57-80/Lm long by 2-3 /Lm in diameter. Microscleres absent in
Schmidt, 1862 became the oldest available name (from secondary the type species, present in the type species of Vagocia, Calyx
source: Wiedenmayer, 1977b: 118). arcuarius (Topsent).
Material examined. Holotype: BMNH 1867.7.26.71 - Remarks. The type species of Calyx is, in the opinion of
Reniera ? calyx Schmidt, dry, Lesina (specimen of O. Schmidt). Topsent (1925c), very common in the Mediterranean Sea, where it
Other material. BMNH 1926.3.10.4 - Reniera calyx Schmidt, may grow to a big size. This species is the same as described by
1862, Sebenico, dry, Norman Collection. BMNH 1866.7.30.3 - Nardo (1833: 522) as Esperia calix, and later by Schimidt as
Reniera calyx, 'Mer Adriatic', dry, purchased from Schaufuss. Reniera ? calyx, and finally byVosmaer (1885b) as the type species
Additional material. Calyx nicaeensis Risso: BMNH 1958.12.4.5- of his genus Calyx. This sponge was described earlier by Risso
dry, Topsent's specimen purchased from Professor Tortorense, (1826: 372) as Spongia nicaeensis (see Topsent, 1925c), which is the
1958. Holotype of Vagocia, Gellius arcuarius Topsent, 1913b: senior-most available name. Characteristics of this species are:
RSME 1921.143.1419 - Scotia Bay, Scottish Antarctic Expedition, a very variable morphology, stony texture observed in large speci-
1902-1904. mens principally due to the multispicular choanosomal fibres with
Porifera· Demospongiae • HaploscIerida • Petrosina • Phloeodictyidae 897

Fig. 2. Calyx Vosmaer, l885b. A-D, type species Reniera ? calyx Schmidt, l862 (= Calyx nicaeensis Risso) , holotype, BMNH 1867 .7.26.71. A , massive,
caliculate specimen, external view (scale 1 em). B, longitudinal section through the multilayered, compact ectosomal network or crust, covered by the sur-
face-membrane (scale 200 ILm). Note tangential choanosomal tracts under the surface. C,internal surface of the calyx, small oscules present (scale I em).
D, longitudinal section through the choanosomal skeleton, longitudinal tracts and unispicular network (scale 200 ILm).

compact oxeas, distributed longitudinally inside fibres, supported Wiedenmayer, 1977b: 124. Foliolina Schmidt, 1870: 42, pI. 4,
by sparse spongin, and by the multilayered ectosomal skeleton, or fig. 4. (Taxonomic decision for synonymy, Van Soest, 1980: 91;
strong ectosomal crust. In spite of being easily identified, to date concerning Foliolina Schmidt, 1870: Zea, 1987: 136 and Werding &
the only complete description of this species is from Topsent Sanchez, 1991: 204).
(1925c), who illustrated several specimens with different growth
forms, found in the Gulf of Naples. One of Topsent's specimens of Type species
Calyx nicaeensis Risso, BMNH 1958.12.4.5, is illustrated here.
Vagocia de Laubenfels, 1936a, with type species, Gellius arcuarius Isodictya robusta Bowerbank, 1866: 334 (by monotypy).
Topsent, 1913b: 623 (by original designation; Fig. 3), is considered
here to be a junior synonym of Calyx, following Wiedenmayer Definition (from de Weerdt, 1985)
(in Hooper & Wiedenmayer, 1994).
Phloeodictyidae with a compact external ectosomal crust com-
Distribution posed of tangential oxeas, often surrounding a hollow body and
numerous ubiquitous fistulae. Megascleres oxeas. Microscleres, if
Calyx is abundant in the world's oceans, including Magellan present, sigmata or toxas.
Province, Brasilia, West and South Africa, Mediterranean
Atlantic, Antarctica, Subantarctic, New Zealand and West Indies Diagnosis
(Van Soest, 1980, and pers.comm.; Koltun, 1964b; Desqueyroux-
Faundez, 1989). Massive, globular, larnellate growth forms, with long tubular
processes or fistules, opened or closed at their ends. Ectosomal skele-
ton a tangential multilayered compact crust of tangential spicules.
Fistule walls supported by an irregular, network of tracts, with irregu-
OCEANAPIA NORMAN, 1869 lar mesh and abundant spongin. Between the tracts, a fine unispicular,
subisodictyal network is visible and minute sigmata may be present.
Synonymy Choanosomal skeleton (Fig. 4D, F) of the basal part is formed by an
irregular network of strong spicule tracts, parallel to the surface, abun-
Oceanapia Norman, 1869b: 334. Phloeodictyon Carter, dantly ramified to form irregular to oval meshes, with abundant free
1882b: 122. Rhizochalina Schmidt, 1870: 35. Biminia oxeas and less numerous sigmata. Spongin in variable quantities.
898 Porifera· Demospongiae • Haplosc1erida • Petrosina • Phloeodictyidae

Fig. 3. Calyx Vosmaer, 1885b. A-D, Vagocia de Laubenfels, 1936a (type species Gellius arcuarius Topsent, 1913b, holotype RSME 1921.143.1419).
A, external view of the short pedunculate lamellar specimen (scale 1 cm). B, tangential view of the surface membrane over the spicules of the ectosomal
network (scale 50 !-Lm). C, terminal part of the lamellae (scale 1 cm). D, confused connecting unispicular reticulation of the choanosomal skeleton, view of
the oxeas and fine, largely open, toxas (scale 20 !-Lm).

Previous reviews Shetland, 1867, Norman Collection, correlated with 5 slides in the
collection. BMNH 1910.1.1.372-374 - dry, Shetland, 1867, Haaf
Carter, 1882b; Topsent, 1920a: 2 (Rhizochalina); Dendy, Gruney, Off Unst Island, Norman collection. Rhizochalina oler-
1922b; Ridley & Dendy, 1887: 32, 36; Bergquist, 1965: 160; acea: BMNH 1870.5.3.142 - dry and 1 slide, Antillen, purchased
Van Soest, 1980: 85; de Weerdt, 1985: 74; Desqueyroux-Faundez, from O. Schmidt.
1987: 200. Description. Globular, regularly bulbous to irregularly
rounded or turnip-shaped, massive sponge, 5-6 cm in diameter,
Description of type species surrounded by a strong, compact, hard and breakable (in dry state)
external crust (Fig. 4A-C, E). Characteristically one to numerous
Isodictya robusta Bowerbank, 1866: 334 (Fig. 4). long cloacal branched or unbranched tubular appendages or fistu-
Synonymy. Isodictya robusta Bowerbank, 1866: 304. lae, 0.5-3 cm long, opened or closed at their ends. Appendages
Desmacidon jeffreysii Bowerbank, 1866: 347; 1874: 157, pI. 62, issued from the upper part of the sponge are internally branched to
figs 1-5; 1882: 170; Carter, 1882b: 117; Oceanapia jeffreysii; form a strong choanosomal reticulation of cylindrical branching
Norman, 1869b: 334; Esperia jeffreysii; Schmidt, 1870: 77. Not tubes. The basal part of the sponge exhibits long, stout compact
Oceanapia robusta Ridley & Dendy, 1887 (= O. jistulosa, see Van to partly hollow roots (Fig. 4C), also considered fistular
Soest, 1980: 86). appendages by Norman (1869b), terminating in single strong
Material examined. Holotype: BMNH 1877.5.21.2040 - fibres. These appendages are used as basal attachments to the
dry, and 5 slides, Norman collection, Shetland. Other material. substratum. Surface smooth, even, slightly hispid, from ends of
Desmacidonjeffreysii: BMNH 1877.5.21.2041- dry, Norman col- surface spiCUles. Consistency of the sponge body and the fistulae is
lection, Shetland, 1868 (specimen illustrated by Bowerbank, 1874, crumbly, consistency of the roots is resilient. The interior of the
pI. 62). BMNH 1877.5.21.807 - dry, and 2 slides. Specimen with body in the fresh sponge, contains a pulpy, gelatinous, uncoloured
two labels: BMNH 1883.1.2.5.7 - dry, "Desmacidonjeffreysii and mass, which become hard, yellow to brown in the dry sponge.
O. robusta". BMNH 1910.1.1.372 - dry, "Isodictya robusta and Oscules are small, 2-3 mm in diameter, and numerous in the inter-
Desmacidonjeffreysii", small box contains 5 smalliamellate frag- nal part of the body wall. Ectosomal skeleton is a compact, multi-
ments about 0.5 cm. BMNH 1883.1.2.5.7 - dry, Norman layered, tangential, unispicular, unordered network of packed
Collection, presented by H.I. Carter Esq. BMNH 1900.4.4.36-40- oxeas, supported by the subectosomal multi spicular tracts (Fig. 4D,
dry, Shetland, Norman Collection. BMNH 1930.7.3.414-418 - dry, F). The spaces between the tracts are subdivided by thinner
Porifera' Demospongiae • Haplosclerida • Petrosina • Phloeodictyidae 899

Fig. 4. Oceanapia Norman. I 869b. A-B, type species [sodictya robusta Bowerbank, 1866, BMNH 1877.5.21.2040, holotype, internal and external view
of the body wall (scale I cm). C-E, DesmacidonjejJreysii, BMNH 1877.5.21.2041, specimen illustrated by Bowerbank (1874, pI. 57). C, E, lateral view of
the globular, regularly bulbous to irregularly rounded or turnip-shaped massive specimen, with fistules and long root-like structures (scale I cm).
D, choanosomal skeleton, longitudinal section (scale 50 ILm). F-G. Desmacidon jeffreysii, BMNH 1900.4.4.36. F, longitudinal section through the surface
(at top) (scale 500 ILm). G, long choanosomal multi spicular stout fibres and dense unispicular connecting network (scale 500 ILm).

paucispicular tracts forming irregular meshes. Choanosomal skele- The first specimen dredged by Norman in 1861, a portion of
ton is an irregular network of multi spicular longitudinal stout crust, was identified and described by Bowerbank as [sodictya
tracts, parallel to the surface, forming large, irregularly elongate robusta Bowerbank, 1866. A new collection by Norman yielded
meshes, sub-divided by a regular isodictyal network of single fragments of the crust and of detached fistulae of a similar speci-
spicules, with spongin only found at the nodes (Fig. 4G). men. On this occasion Norman remarked that the entire specimen
Megascleres are oxeas, slightly bent, 170-200-260/.Lm long by should be very different from Bowerbank's description. From a
6--8-10/.Lm in diameter. Microscleres are sigmata, thin, irregularly new collection of the same sponge by Norman in 1864, Bowerbank
curved, 10-12-18 /.Lm long. was convinced that he was dealing with a new species that he
Distribution. De Weerdt (1985) remarked that Oceanapia named Desmacidon jeffreysii Bowerbank, 1866. After subsequent
robusta had not been reported from the Mediterranean, nor from collections, Norman retrieved an entire specimen and he described
the Western part of the Atlantic Ocean, and concluded the species this remarkable sponge himself as a new genus Oceanapia,
was geograpically restricted only to the northern part of the North explaining the different interpretations of its type species.
Eastern Atlantic Ocean, viz. from East of Greenland, Iceland and Norman's (1869b) diagnosis of Oceanapia jeffreysii Bowerbank
Norway down to the Azores (37 N). 0
(1869: 334), included its synonymy with Desmacidonjeffreysii and
Remarks. In describing Oceanapia Norman (1869b) under- lsodictya robusta. Bowerbank (1866) only gave indications of the
lined the problem of Bowerbank's identification of this species. sponge's habit, the type of skeleton and the spicules.
900 Porifera· Demospongiae • HaploscIerida • Petrosina • Phloeodictyidae

Fig. 5. Oceanapia Norman. 1869b. A-D, Rhizochalina Schmidt, 1870 (type species Rhizochalina oleracea Schmidt, 1870, holotype BMNH
1870.5.3.142, fragment of Rhizochalina carotta Schmidt, 1870). A, extant fragment of specimen (scale 1 cm). B, ectosomal network, transverse section of
the surface (scale 100 fLm). C, multispicular compact choanosomal tracts with sparse spongin (scale 50 fLm). D, transverse section of choanosomal skele-
ton, below the surface network (scale 50 fLm).

Van Soest (1980) proposed including Rhizochalina Schmidt, (compare with Fig. 6A-D). The method by which water is ingested
1870 (Fig. 5A-D) (type species Rhizochalina oleracea Schmidt, into these Oceanapia species is peculiar. The fistules are inhalant and
1870: 35, by subsequent designation (de Laubenfels, 1936a) the exhalant openings are buried in the sand (Fig. 6A, B). We studied
and Phloeodictyon Carter, 1882b (Fig. 6A-C) (type species the type species and remarked that Foliolina's skeleton is of the same
Phloeodictyon isodictiformis Carter, 1882b: 122, by subsequent type as that of Oceanapia (cf. Figs 4F, G and 6C, D), with an ectoso-
designation (de Laubenfels, 1936a: 72) in synonymy with mal hispid layer of compact spicules, with choanosomal stout multi-
Oceanapia because their respective type species were very similar spicular tracts having only sparse spongin, and thinner paucispicular
in their basic characters. The presence or absence of sigmata tracts forming an irregular mesh. Foliolina peltata is also different in
was judged to be of little value. Wiedenmayer, in Hooper & its peculiar external morphology, having fistules with disc-like or
Wiedenmyaer (1994), also included the genus Biminia platform-like extensions along their length. These characters and the
Wiedenmayer, 1977b into synonymy (Fig. 6D-F) with Oceanapia. absence of sigmas in Foliolina are certainly considered significant at
Actually, we do not know the exact value of microscleres as the species-level, whereas close similarities in their respective skeletal
generic characters of Phloeodictyidae. Characters such as the pres- structures suggest to us that Foliolina is a junior synonym of
ence of an ectosomal crust and the globular habit, which we Oceanapia, as was proposed earlier by Zea (1987: 136) and reiterated
assume are consistently present in O. robusta (e.g., Desqueyroux- by Werding & Sanchez (1991), based on their studies of two species
Faundez, 1987), misled Bowerbank (1874: 158, 161), to propose of Phloeodictyidae (Rhizochalina oleracea and Foliolina peltata,
substituting his first description of fragments of Norman's speci- from Uraba and Cartagena, and Santa Marta, Colombia, respectively).
mens with a new species (Desmacidon jeffreysii). In the same way, From these comparisons, which included living populations of both
the presence or absence of a given type of microsclere in species of species, Foliolina was relegated into synonymy with Oceanapia.
Phloeodictyidae justified the erection of genera which today are Rhizochalina (Fig. 5A-D) and Phloeodictyon (Fig. 7A-C)
considered by most authors to be junior synonyms of Oceanapia differ from Oceanapia only by their lack of sigmas. Biminia
(see above). (Fig. 6D-F) Wiedenmayer, 1977b (type species Oceanapia
Foliolina Schmidt, 1870 (type species Foliolina peltata Schmidt, toxophila Dendy, 1922b: 45, pI. 8, fig. 2, by original designation)
1870: 42, pI. 4, fig. 4, by monotypy; see Fig. 7) is another genus often on the other hand has both sigmas and toxas (Fig. 7D, F), but also
associated with Oceanapia. Van Soest (pers.comm.) considers that exhibiting the same type of skeleton as Oceanapia, and conse-
this genus is very similar to Oceanapia due to its turnip-shaped main quently Wiedenmayer in Hooper & Wiedenmayer (1994) consid-
body, which is also buried in the sand, and held up by a 'root system' ered Biminia also a junior synonym of Oceanapia.
Porifera· Demospongiae • Haplosclerida • Petrosina • Phloeodictyidae 901

Fig. 6. Oceanapia Norman, 1869b. A-C, PhloeodictyDn Carter, 1882b (type species Phloeodictyon isodictyiformis Carter, 1882b, holotype BMNH
1872.5.4.1). A, external view of the massive specimen, erect cylindrical appendages and the basal lamina. B, tangential view of the ectosomal unispicular
network with spongin only at the nodes. C, fistule-wall, unispicular confused dense irregular network. D-F, Bimina Wiedenmayer, 1977b (type species
Oceanapia toxophila Dendy, 1922b, holotype BMNH 1921.11.7.37). D, holotype. E,longitudinal section of the fistulae, with long, multispicular tracts and
connecting unispicular network. F, longitudinal section through the body wall of the tube, etcosomal skeleton at right.

Distribution Type species

Worldwide. Petrosiajistulata Kirkpatrick, 1907a (by original designation).

PACHYPELUNA BURTON, 1934

Definition (from Van Soest, 1980)
Synonymy
Oceanapiidae with a thick ectosomal crust of tangential
Pachypellina Burton, 1934b: 18; de Laubenfels, 1936a: 70; spicules and a densely spiculous choanosome which is riddled with
Van Soest, 1980: 91. holes. Megasc1eres are oxeas, no microsc1eres, no fistules.
902 Porifera· Demospongiae • Haplosclerida • Petrosina • Phloeodictyidae

Fig.7. Oceanapia Norman, 1869b. A-D, Foliolina Schmidt, 1870 (type species Foliolina peltata Schmidt, 1870, specimen BMNH 1948.3.8.37-39, Atlantic
Expedition). A, view of specimen (scale 0.6 cm). B, enlarged view of the same specimen, basal part. C-D, BMNH 1870.5.3.105, slide of Schmidt, Florida.
C, tangential view of the hispid surface. D, longitudinal view of the choanosomal skeleton, through the surface, illustrating identical structure to Oceanapia.

Diagnosis Description. Three fragments of a tubular sponge, with a

finely hispid, smooth internal and external surfaces. Tubes form a
Tubular to massive growth forms. Ectosomal skeleton a dense, thick wall with a terminal oscule, measuring 2-2.5-6 cm high by
tangential network of oxeas. External surface smooth but hispid 3-2.5-2 cm in diameter. A fine ectosomal membrane covers the
due to the intercrossed ends of the ectosomal oxeas. Choanosomal numerous ostia, 0.5-3 mm in diameter. Oscules are only present in
skeleton is a confused spiculous network with numerous aquifer- the internal surface of the tube, they are small about 1 mm in diam-
ous canals. Megascleres are oxeas. Microscleres are absent. eter. Texture is hard and friable. Spongin is not visible, except at
skeletal nodes. Ectosomal skeleton is a multispicular tangential
Previous reviews compact network of closely packed oxeas (Fig. 8A, B), without
visible spongin, fotming a strong, regular crust of spicules of
De Laubenfels, 1936a: 70, 1950: 66; Van Soest, 1980: 91, 900-1520 f-Lm large, supported by terminal brushes of choanoso-
117; de Weerdt & Van Soest, 1986: 44. mal longitudinal principal tracts. Ends of tracts partly visible at the
surface. Choanosomal skeleton is a dense irregular network perme-
Description of type species ated by numerous aquiferous canals (Fig. 8C), with multi spicular
longitudinal tracts, 4--8 spicules wide, running perpendicular to the
Petrosiajistulata Kirkpatrick, 1907a (Fig. 8). surface, connected by short, irregular paucispicular tracts, 3-4
Synonymy. Petrosia jistulata Kirkpatrick, 1907a: 290; spicules wide and 1-2 spicules long. Megascleres are only oxeas,
1908c: 51; [in part] Burton, 1929a: 420, as Chalina dancoi slightly bent with blunt ends: 328-388-439 X 12-16-20 f-Lm,
(Topsent); Pachypellina jistulata; Burton, 1934b: 18, Fig. 2. Not and 238-296-344 X 4--6-8 f-Lm (Burton's specimens; Fig. 8D);
Burton, 1938: 6 (=Haliclona dancoi (Topsent)). 394--430-463 X 12-19-25 f-Lm, and 262-343-385 X 4--6-8 f-Lm
Materiol examined. Holotype: BMNH 1908.2.5.200 - Winter (Kirkpatrick's specimens).
Quarters, 25-30ftm. Holotype (slide): BMNH 1933.3.17.172 - Distribution. Antarctica, Subantarctic (Kirkpatrick, 1907a;
64°20'S 56°38'W, 150m depth. Paratype: BMNH 1908 2.5.202a, b- Burton, 1934b).
McMurdo Bay, 96-120ftm, Antarctica, 'Discovery' Expedition. Remarks. The type species of Pachypellina, Petrosia jistu-
Other material. Holotype of Pachypellina jistulata Burton, 1934b: lata Kirkpatrick, was originally regarded by Burton (1929a: 420)
USNM 20884 - Graham region, SE Seymour Island, 64°20'S as an immature form of Reniera dancoi Topsent, 1901, and conse-
56°38'W, 150m depth. Holotype (slide): BMNH 1933.3.17.172. quently relegated into synonymy with the latter species.
Porifera· Demospongiae • Haplosclerida • Petrosina • Phloeodictyidae 903

Fig. 8. Pachypellina Burton, 1934b. A-B, type species Petrosiajistulata Kirkpatrick, 1907a, BMNH 1908.2.5.202, and BMNH 1908.02.1902. A, frag-
ment of specimen, longitudinal view through the surface, ectosomal crust on top and loose choanosomal network below (scale 200 Il-m). B, the same spec-
imen showing choanosomal network and aquiferous canals (scale 500 Il-m). C-D, holotype of Pachypellina jistulata; Burton, 1934b, RSME 20884.
C, ectosomal network of dense tangential layer of oxeas, or ectosomal crust, tangential view (scale 200 Il-m). D, the same specimen, two types of oxea,
slightly bent, with blunt ends (scale 10 Il-m).

De Laubenfels (1936a: 70) remarked Pachypellina was very jistulata. Kirkpatrick's (1907a) material is clearly tubulose, whereas
much like Adocia except that its spicules were hastate and its ecto- Burton's (1934b) specimen is massive-tubulose, but in both speci-
somal much denser and not typical of Adocia. Van Soest (1980) mens the skeletal structure is identical, and megasclere geometry is
characterised Pachypellina by its distinctive ectosomal crust and its the same (bent oxeas, with blunt or tomote ends). This material indi-
dense choanosomal skeleton. He also included Haliclona podatypa cates to us that the connective tracts in the choanosomal skeleton,
de Laubenfels, 1934, from Puerto Rico, in the genus presumably formed by only few transverse spicules, are very typical and differ
due to its possession of an ectosomal crust and isotropic choano- from the typical Petrosia skeletal structure (Petrosiidae). In addition,
somal reticulation bound together with sparse spongin. We think its tangential surface skeleton makes Pachypellina Burton a well
that this inclusion is a mistake because the spicules of P. podatypa defined genus which we consider to be valid. Undertaking a revision
(158-170.2-180 X 4.5-5.1-6.5 f,Lm) are much smaller than those of species from Antarctica originally described under Reniera and
of the type species of Pachypellina, although in the opinion of Van Petrosia, would certainly assist in recognising new species of
Soest (1980: Table 2), the Atlantic specimens of P. podatypa Pachypellina, and also in providing further biological data as to the
exhibit a variable morphology. De Weerdt & Van Soest (1986) distribution and morphological characteristics of the type species.
recorded only one species of Pachypellina from the North Atlantic
Ocean, Reniera tufa Ridley & Dendy, 1887, from Cape Verde Is.,
referred to the genus by Burton (1934b). Distribution
We examined two species of Pachypellina to assess relation-
ships for the present study: a slide of the holotype of the type species Antarctica, Subantarctic (Burton, 1934b; Kirkpatrick, 1907a;
Petrosiajistulata Kirkpatrick, and Burton's material of Pachypellina Koltun, 1964b).
904 Porifera· Demospongiae • HaploscIerida • Petrosina • Phloeodictyidae

Fig. 9. Tabulocalyx Pulitzer-Finali, 1993. A-F, type species Tabulocalyx pedunculatus Pulitzer-Finali, 1993, MSNG 48327, holotype. A, ectosomal crust
a single tangential layer of free oxeas in confusion, occasionally forming confused tracts (scale 500 j.Lm). B-C, subectosomal two-dimensional network of
multispicular fibres with large and smaller mesh, view taken from below the surface (scales B, 500 j.Lm; C, 200 j.Lm). D, deeper within the interior of the
globous body, the mass of oxeas in confusion (scale 50 j.Lm). E, enlarged view, part of the subectosomal network (scale 50 j.Lm). F, longitudinal view of one
of the subectosomal networks of multi spicular fibres, parallel to the surface (scale bar 200 j.Lm).

TABULOCALYX PULITZER-FINALI, 1992 Definition

Synonymy Phloeodictyidae without fistules. Cortex in two separate lay-

ers with an ectosomal network of a single layer of oxeas, supported
Tabulocalyx Pultizer-Finali, 1992: 322. by subsequent ectosomal layers of multi spicular fibres in the
subectosome, representing earlier growth stages.

Type species Diagnosis (from Pulitzer-Finali, 1992, 1996)

Tabulocalyx pedunculatus Pultizer-Finali, 1992 (by original Irregularly massive, cylindrical-lobate growth fonns with a tan-
designation). gential ectosomal crust of free oxeas in confusion (Fig. 9A), forming
Porifera· Demospongiae • Haplosclerida • Petrosina • Phloeodictyidae 905

occassional thin, vague tracts, supported by a strong two- and smaller meshes, 205-323-533 f.Lm (Fig. 9B, C). Within the inte-
dimensional network of multi spicular fibres (Fig. 9B, C), under rior of the body the mass of oxeas is traversed by two or three
which there is a dense mass of oxeas (Fig. 9D), which are cut networks of multispicular fibres, parallel to the surface, appearing
across by several two-dimensional networks like those mentioned in longitudinal section as different growth stages (Fig. 9F), reminis-
above; each of these two dimensional networks is separate from cent of Callyspongia (Euplacella) (Callyspongiidae) and in
each other and more-or-less parallel to the surface. All the internal Neopetrosia (Petrosiidae), and presumably independently acquired.
part of the body is occupied by a mass of oxeas in confusion and Megascleres oxeas, 185-200 X 10.5 -11.5 f.1m.
some irregular multispicular fibres. Distribution. Known exclusively from the type locality:
East Africa, North Kenya Bank, 110m depth.
Previous reviews Remarks. Pulitzer-Finali (1992) erected this genus princi-
pally due to the characteristic structure of its ectosomal network, and
Pulitzer-Fin ali (1992). included it in Phloeodictydae irrespective that it lacked fistules
(Pulitzer-Finali, 1996: 128, Fig. 24A-B). This skeletal structure con-
Description of type species sisting of the double layered ectosomal network appears to us as a
pivotal character in its diagnosis, differentiating it from other genera
Tabulocalyx pedunculatus Pulitzer-Fin ali, 1993 (Fig. 9). of Phloeodictyidae. Pulitzer-Finali (1996) also included the
Synonymy. Tabulocalyx pedunculatus Pulitzer-Finali, 1993: Petrosiidae species Strongylophora corticata Wilson, 1925, in this
323, figs 61, 62. genus despite the fact that its spicules corresponded to typical
Material examined. Holotype: MSNG 48327 - North Strongylophora, and his action was also probably due to the presence
Kenya Bank, 02°47.5'S, 41°00'E, 110m depth. Other material. of the double layered ectosomal. In our opinion, the spiculation of
Tabulocalyx corticatus (Wilson): MSNG P.60 - Pultizer-Finali S. corticata is typical of Strongylophora, and we maintain here
collections, 1986, Laing island, New Guinea, 17.08.1986. Strongylophora corticata Wilson, 1925, as a valid species of Petrosia
Description. Globose, 5.5 cm in diameter, supported by a (Strongylophora). Pulitzer-Finali (1996) also amended the diagnosis
long stem, 1.5 cm thick, 6 cm long. Oscules, crateriform, numeorus, of Tabulocalyx as follows 'cortex in two separated layers, the sub-
3-5 mm in diameter. Ectosomal crust a single tangential layer of dermal one being a rigid cribriform lamella consisting of a close net-
free oxeas in confusion, occasionally forming confused tracts, sup- work of compact spiculo fibres developed in a tangential plane.
ported by a two dimensional network of multi spicular fibres, Former ectosomal lamellae may be found inside the choanosome,
82-164-287 f.1m in diameter with large meshes, 246-443-820 f.1m, marking earlier growth stages' .
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob WM. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Petrosiidae Van Soest, 1980

R. Desqueyroux-Faundez1 & c. Valentine2

1 Museum d'histoire naturelle, P.O. Box 6434, CH-1211 Geneva 6. Switzerland. (ruth.faundez@mhn.ville-ge.ch).
2 The Natural History Museum. Cromwell Road, London, SW7 5BD, UK. (c.valentine@nhm.ac.uk)

Petrosiidae Van Soest (Demospongiae, Haplosclerida) contains four valid genera and two subgenera. Species are widely distributed in
shallow and deeper warm-temperate to cold waters. Genera share a reticulated tangential ectosomallayer of spicules, or surface crust and
are differentiated by their choanosomal skeletal structure, varying from isotropic-confused to isotropic-lamellate, and their types and size
classes of spicules.
Keywords: Porifera; Demospongiae; Haplosclerida; Petrosiidae; Acanthostrongylophora; Neopetrosia; Petrosia (Petrosia); Petrosia
(Strongylophora); Xestospongia.

DEFINITION, DIAGNOSIS, SCOPE Scope

Synonymy Nine nominal genera and hundreds of species are known

worldwide, of which only four genera and two subgenera are con-
Petrosiidae Van Soest, 1980: 66. sidered valid here: Acanthostrongylophora, Neopetrosia, Petrosia
(Petrosia), Petrosia (Strongylophora) and Xestospongia. Densa
Definition (from Van Soest, 1980) and Quepanetsal are included in synonymy with Xestospongia,
with question, as are Tedaniella and Chalinorhaphis included in
Haplosclerida with an ectosomal skeleton consisting of an Petrosia. Species are distributed in shallow and deeper tropical,
isotropic reticulation of single spicules or spicule tracts and a warm-temperate to cold waters, with depth range 4-280 m.
choanosomal skeleton verging towards an isotropic reticulation of
spicule tracts, in which primary and secondary tracts are indistinct. History and biology

Diagnosis (emended from Van Soest, 1980; Hooper & The family was erected by Van Soest, 1980 to include
Wiedenmayer, 1994) Haplosclerida with an ectosomal skeleton consisting of an
isotropic reticulation of single spicules or spicules tracts and a
Massive, volcano or vase-shaped, bulbous and less com- choanosomal skeleton verging towards an isotropic reticulation of
monly, encrusting-branching sponges. Sponge texture characteris- spicule tracts, in which primary and secondary tracts are indistinct.
tically stony and brittle due the high silica content of the mineral Van Soest (1980) split the genera previously grouped in
skeleton, about 58% relative to spongin content of the skeleton Nepheliospongiidae Clarke, 1900 by Wiedenmayer (1977a) into two
(Desqueyroux-Faundez, 1987). Surface smooth, covered by a new families: Petrosiidae and Oceanapiidae. Van Soest (1980) sug-
smooth, strong crust or variably hispid. Specialised ectosomal gested that it was uncertain whether Recent genera such as Petrosia
skeleton consisting of an isotropic tangential reticulation of free and Xestospongia were closely allied with the fossil genus
spicules or spicule tracts, forming the mineral crust. Choanosomal Nepheliospongia, as implicitly indicated by their inclusion in a
skeleton is a more-or-less regular isotropic network of multispicu- single family Nepheliospongiidae. Wiedenmayer (1977a) defined
lar tracts where primary and secondary fibres are indistinct. Nepheliospongiidae principally by the strong development of
Spongin not visible, spicules bound together with the minimal megascleres in relation to spongin, hence with hard or friable consis-
amount of spongin. Multispicular tracts are regularly distributed to tency, the presence of a skeletal accretive structure with ascending
form, rounded meshes parallel to the surface, that can be ill defined and radially diverging skeletal tracts, and the tendency to have a tan-
or masked by the abundant free spicules. Megascleres: oxeas to gential ectosomal network or peripheral crust. At the same time,
strongylotes. Microscleres if present, smaller oxeas. In one sub- Wiedenmayer (1977a) associated a fossil group HeJiospongiidae
genus (Strongylophora), smaller, sausage-shaped strongyles exist. Finks, 1960 with NepheJiospongiidae, which in his opinion shared
Spicule types and size classes are very variable within different the same characters of spiculation, skeletal architecture and aquifer-
genera, ranging from only one type of spicule in only one size ous system, with living Nepheliospongiidae. Wiedenmayer (1977a)
class, smaller than 200 fLm long (Neopetrosia) or larger than included in this family: Petrosia, Xestospongia, Cribrochalina,
200 fLm long (Xestospongia), up to two types of spicules with five Hemigellius, Vagocia, Calyx, Rhizochalina, Oceanapia, Biminia and
size classes (Strongylophora). Spicule morphologies and sizes are Siphonodictyon.
considered important taxonomic characters among different genera, Bergquist & Warne (1980: 35) accepted, but modified, the
since they are fixed and stable for each genus (Table 1). Oviparous diagnosis of NepheJiospongiidae Clarke given by Wiedenmayer
reproduction is known for only one genus (Xestospongia), which is (1977a). They included only two genera having a mineral skeleton
gonochoric and has synchronous spawning (e.g., Fromont, 1988), predominant over the soft tissues, and a greater development of
but larvae are unknown (Bergquist, 1980a). a spiculose ectosomallayer. In one genus (Petrosia) this is represented

906
Porifera' Demospongiae • Haplosclerida • Petrosina • Petrosiidae 907

Table 1. Spicule forms and dimensions for genera and subgenera of In the opinion of Bergquist et al. (1986) recent improvements
Petrosiidae (from the literature) (measurements in !-Lm, represented as in biochemical technology has provided pivotal information on
minimum-mean- and maximum dimensions). sponge phylogeny, in particular, sterol patterns as markers for
resolving taxonomic problems at the intrageneric level, e.g., within
Genus Type of Spicules size (!-Lm)
spicules Xestospongia, Petrosia and Phakellia. Fromont et al. (1994),
studying structural parameters of sterols, compared sterol percent-
Acanthostrongylophora strongyles 105-154-174 X 5-8-12 age of molecular weights, type of sterol nucleus and position of
Hooper, 1984a thin oxeas 98-139-173 X 1-2--4 alkylation, with previous chemotaxonomic studies in order to
small strongyles contamination determine if these studies showed groups of sponges with similar
Neopetrosia de small oxeas 120X 3 characters. They concluded that grouping species and genera was
Laubenfels, 1949b not possible, although a sterol-set ('fingerprint') was characteristic
Petrosia Vosmaer, 1887 oxeas to oxeotes (1) 182-212-237 X for particular species. Similar results were obtained by Van Soest
10-13-14 & Braekman (1999) for Petrosia species and Cribrochalina
strongylotes (2) 40-80-136 X 7-10 vasculum; both possessing hydroxylated acetylenes as good
strongylotes (3) 30--45-85 X 2-3 markers; Xestospongia testudinaria and X. muta shared bromi-
Strongylophora Dendy, centrangulate 28 X2 nated acetylenes also representing a good marker. In the family
1905 oxeas Niphatidae, Amphimedon compressa, A. viridis and A. erina con-
strongylotes 260 X 20 tained alkylpiperidines which they share with Callyspongia
small strongyles 260X6 species. Nevertheless, one of the major difficulties to be over-
Xestospongia de oxeasto 254-308--400 X come before any confidence can be applied universally to chemo-
Laubenfels, 1930 strongylote-oxeote 16-21-30 taxonomic methods is the confident taxonomic identification of
studied samples and of all the different sterols found in each
sponge.
by a marked crust or non-detachable peripheral crust of spicules,
and in others it is visible in the form of a strongly hispid (e.g., Distribution
Xestospongia) or a shaggy (e.g., Acanthostrongylophora) surface.
Van Soest's (1980) concept of Petrosiidae concerned sponges Species are distributed in shallow and deeper warm-temperate
which often exhibit a tangential surface crust of free spicules - a to cold waters (e.g., Galapagos, Sri Lanka, Falkland Islands, Straits
character considered as an apomorphy for the family. Oceanapia of Magellan, Subantarctic), at depths between 4-280m. More than
and Calyx were excluded from Petrosiidae. The genera a hundred nominal species of Petrosiidae have been recorded
Cribrochalina, Hemigellius, Vagocia, Rhizochalina, Biminia and (Van Soest, pers. comm.) from the West Indies, East Pacific,
Siphonodictyon were considered as Nepheliospongiidae by Galapagos Islands; Chile-Peru, Magellan, Antarctica and the
Wiedenmayer (1977a) but are actually treated in Niphatidae Subantarctic; Australia and New Zealand; Indonesia; Molucca Sea;
or Phloeodictyidae. Van Soest (1980) described only three genera Pacific, Indo-Pacific and Indian Ocean; Mediterranean-Atlantic,
in Petrosiidae, all from the West Indies: Xestospongia, Petrosia and Red Sea, Mediterranean, Boreal, N, and E Atlantic; W and
Strongylophora. S Africa.

KEY TO GENERA AND SUBGENERA

(1) Ectosomal skeleton clearly different from choanosomal skeleton, very dense, tangential unispicular network, finely hispid, with
rounded, regular or isodictyal, or irregular, subrectangular meshes. Choanosomal spicule tracts compact, differentiated or not in
primary and connecting tracts. With or without microscleres ............................................................................................................. 2
Ectosomal skeleton undifferentiated from choanosomal skeleton, consisting of a very dense disordered network of free spicules, with
a single type of spicule of a single size class larger than 200/-Lm long. Choanosomal skeleton consisting of short longitudinal,
undivided, pauci- to multispicular tracts connected by few oxeas, causing irregular meshes ........................................ Xestospongia
(2) Ectosomal skeleton finely hispid, consisting of a tangential network with rounded regular meshes and free spicules. Choanosomal
spicule tracts compact, not differentiated into primary or connecting tracts. Megascleres are oxeas, ranging from oxeote to
strongylote of at least three size classes, large 180-400 X 10-14 /-Lm, medium 40-130 X 5-10 /-Lm, and small curved strongyles
30-100 X 2-3 /-Lm ........................................................................................................................................... Petrosia (Petrosia) ... 3
Ectosomal skeleton a finely hispid, tangential network with isodictyal meshes of single spicules and free spicules at the nodal points.
A second subectosomal tangential network is present. Choanosomal skeleton with irregular paucispicular tracts differentiated into
short connecting and longer longitudinal tracts. Megascleres oxeas in a single size class smaller than 200 /-Lm long ..... Neopetrosia
(3) Ectosomal skeleton finely shaggy, with a tangential network of multispicular tracts of strongyles and free oxeas, with ovoid to sub-
rectangular meshes. Choanosomal spicule skeleton compact, differentiated into multi spicular longitudinal tracts and interconnecting
confused tracts of single strongyles. Spicules: strongyles and thin oxeas of similar length (100-120 /-Lm up 170/-Lm, but of different
diameter: 5-12 and 2-4/-Lm, respectively). Microscleres absent ................................................................... Acanthostrongylophora
Ectosomal skeleton a dense, granular, tangential network of single strongyles forming a subrectangular mesh, with free strongyles
grouped at the nodes of the network. Abundant abruptly curved microxeas. Choanosomal spicule tracts slightly differentiated with
stout longitudinal tracts and connecting short, irregular tracts of strongyles and free strongyles ............. Petrosia (Strongylophora)
908 Porifera· Demospongiae • Haplosclerida • Petrosina • Petrosiidae

Fig. 1. Acanthostrongyla Hooper, 1984a. A-D, type species, A. ashmorica Hooper, 1984a. A, C, holotype NMV F51374 (NTM ZI495), view of the spec-
imen, normal and enlarged view (scale 1 em). B, ectosomallayer (scale 200 fLm). D, choanosomal skeleton forming a network of multispicular, longitudi-
nal tracts with irregular, abundant ovoid meshes (scale 100 fLm).

ACANTHOSTRONGYLOPHORA HOOPER, 1984 Description of type species

Synonymy Acanthostrongylophora ashmorica Hooper, 1984a.

Synonymy. Acanthostrongyla ashmorica Hooper, 1984a:
Acanthostrongylophora Hooper, 1984a: 58; Hooper & 58, figs 4, 5.
Wiedenmayer, 1994: 328. Material examined. Holotype: NMV 51374 (fragment
NTM Z1495) - N of Barracouta Shoals, near Ashmore Reef, Sahul
Type species Shelf, Western Australia. Schizoho1otypes: ZMA POR5294
(fragment), MHNG-INVE 26690 (slides).
Acanthostrongylophora ashmorica Hooper, 1984a (by Description. Massive sponge, erect, firm, friable (Fig. lA, C).
monotypy). Surface uneven, shaggy, with large grooves. Few oscules, large and
flush with the surface, irregularly scattered. Texture firm to stony,
Definition barely compressible, strongly brittle. Surface, light, fragile, covered
by a surface membrane with rounded pores, partly conserved
Petrosiidae with a shaggy surface produced by the ends of pri- between the fine oxea 'tufts' issuing from terminal parts of the pri-
mary fibres and an irregular subrectangular loose reticulation of mary longitudinal fibres (Fig. 1B). Ectosomal skeleton a tangential
strongyles and thin oxeas. irregular subrectangular network of loose strongyles, with free oxeas
and light spongin. Choanosomal skeleton (Fig. lD), partially cav-
Diagnosis (emended from Hooper, 1984a) ernous, forming a network of multispicular, longitudinal tracts with
irregular, abundant ovoid meshes, halichondroid in places.
Petrosiidae with a lamellate-isotropic heavily meshed reticu- Connecting tracts confused, formed by single strongyles and oxeas
lation of spicule tracts cored by thick strongyles and fewer thin cemented by scarce spongin. Megascleres: strongyles, slightly bent,
oxeas. Ectosome with an irregular subrectangular loose reticulation 105-153-174j1m long, 5-8-12j1m wide, and fine, thin oxeas,
of megascleres, interrupted by the terminal part of longitudinal slightly bent, 123-138-173j1m long, 2-4j1m wide.
tracts. Surface membrane thin and fragile with irregular pores. Remarks. Hooper (1984a) described Acanthostrongylophora
Choanosome cavernous, crumb-of-bread texture. Spongin scarce. with spiny microscleres (acanthose microstrongyles), whereas
Spicules strongyles and thin oxeas. subsequently Hooper (in litteris) considered these microscleres
as contaminants, resembling microrhabds of Theonella. Hooper
Distribution (1984a) considered Acanthostrongylophora to be closely related to
Strongylophora, differing only in having acanthose microstrongyles
Northwestern Australia. (which thus potentially negates the genus given that these are now
known to be contaminants, and they were not found subsequently in
Previous reviews our analysis of the type material). However, from our observations
of the holotype we conclude that the two genera differ by the
Hooper & Wiedenmayer, 1994: 328. possession of a shaggy uneven surface in Acanthostrongylophora
Porifera' Demospongiae • Haplosclerida • Petrosina • Petrosiidae 909

Table 2. Comparison of spicule forms and dimensions betwecn hispid, smooth, 'velvet-like' surface (Fig. 2B). Ectosomal skeleton
Acanthostrongylophora and Strongylophora (measurements in fLm taken a tangential isodictyal network with one size of spicule (Fig. 2C),
from type material, represented as minimum-mean- and maximum spongin and spicule brushes only at nodal points, free spicules
dimensions). abundant. Consistency hard, stony, spongin always scarce.
Genus Megasc1eres 'Microscleres' Subdermal region with short spicular tracts, limited by a second
surface skeleton layer, 'peripheral condensation' (Fig. 2D, arrow),
Acanthostrongylophora or a 'paper-thin area about 1 mm below the surface' (see
Hooper, 1984a de Laubenfels, 1953b: 18). Choanosomal skeleton a compact network
Strongyles 66-115-174 X 5-8-12 Absent of irregular paucispicular tracts with rounded meshes, irregular in
Thin oxeas 123-138-173 X 2--4 size. Megascleres: oxeas, less than 200 fLm long.
Small strongyles 16-18-23 X \.5-3--4
contamination
Distribution
Strongylophora Dendy,
1905
Strongyles 156-208-238 X 12-14--20 Central West Atlantic, Indo-West Pacific.
Strongylotes 143-200-230 X 4--7-8
Small strongyles 20--34-53 X 4-7-8 Previous reviews
Microxea-like spicules 21-26-30 X 2-3
Duchassaing, 1850: 26; Green, 1977: 88; Wiedenmayer,
1977b: ll3, pI. 14, figs 1-5, text-fig. 128 (as 'Xestospongia subtri-
(in Strongylophora this is uneven but uniformly granular), by its angularis' (Duchassaing), synonymy from Van Soest, 1980: 72).
breadcrumb texture (hard in Strongylophora), although the texture
of both genera is similar (easily breakable), and the main difference Description of type species
between the two genera being the type of spicules and skeletal
morphology (see Table 2). Haliclona longleyi de Laubenfels, 1932.
Thus, despite the recognition that acanthose microscleres are Synonymy. Haliclona longleyi de Laubenfels, 1932: 54;
contaminants (Hooper, pers. comm.), we suggest that these addi- 1936a: 44, pI. 7, fig. 1, pI. 8, fig. 2. Neopetrosia longleyi de
tional characters detailed above differentiate Acanthostrongylophora Laubenfels, 1949b: 10.
from Strongylophora, with the former a valid genus. Another Material examined. Lectotype (here designated): USNM
species of Acanthostrongylophora is known from Indonesia, where 22475 - near Bird Key, Florida Keys, Dry Tortugas, Gulf of
it is common (Van Soest, pers. comm.), but remains unpublished, Mexico. Schizolectotypes: BMNH 1936.4.8.42 (slide, from
and we await the description of this taxon that may clarify the de Laubenfels' specimen).
taxonomic position of Acanthostrongylophora. Description. Ramose 30-40 cm high, anastomosing
branches 2-5 cm in diameter, erect, with finger-like flattened round
Distribution lobes (Fig. 2A), fixed to the substratum by a narrow basal portion.
Several oscula scattered on one face of the branches, 3-8 mm in
Currently monotypic, known only from the type locality near diameter, sometimes surrounded by a collar. Surface smooth,
Ashmore Reef, Western Australia, 80-91 m depth, with a possible extremely porous, rough, minutely hispid. No special surface
second species from Indonesia (Van Soest, pers. comm.). skeleton. Texture very hard, compact, aquiferous system without
large spaces. Colour alive is yellow green on those surfaces which
NEOPETROSIA DE LAUBENFELS, 1949 are most exposed to light, and dark olive green on shaded surfaces.
Ectosomal skeleton a tangential isodictyal dense surface network
Synonymy composed of one spicule length, with spongin present only at the
nodes. Surface hispidation is produced by fine spicule brushes issu-
Neopetrosia de Laubenfels, 1949b: 10; Bergquist, 1965 ing from the nodes of meshes. Abundant free spicules are present.
(in part): 149, 150, 151. Subectosomal region appears as a compact irregular network of
short spicular tracts. About 1 mm below the surface there is a sec-
Type species ond layer of the same appearance as the surface, described as a
'paper-thin area about 1 mm below the surface' (de Laubenfels,
Haliclona longleyi de Laubenfels, 1932 (by original 1953b: 18). In longitudinal sections this layer is very net-like, eas-
designation). ily observed (Fig. 2D). Choanosomal network confused with irreg-
ular meshes formed by irregular multi- to paucispicular fibres,
Definition 3-20 spicule-rows, ("often more than 20", de Laubenfels, 1932),
coated by a fine spongin sheath. No interconnecting tracts, but 1-3
Petrosiidae with finely hispid surface produced by fine brushes spicules are joined by scarce spongin between tracts. Divergent
of oxeas issued from subectosomal tracts, and a compact choanoso- free ends of longitudinal tracts form the ectosomal skeleton
mal network combining rounded meshes with a superimposed (Fig. 2D). Megascleres: are bent oxeas, 104-120-144 fLm long,
anisotropic reticulation. Megascleres oxeas less than 200 fLm long. 2-3-4 fLm wide, usually of only one size class.

Diagnosis Distribution

Compact sponges (Fig. 2A), with an ectosomal unispicular Dry Tortugas, Florida, near Bird Key Reef, 1 m depth
isodictyal layer and free-brushes of oxeas, producing a finely (de Laubenfels, 1932), Western Bahamas (de Laubenfels, 1949b);
910 Porifera' Demospongiae • Haplosclerida • Petrosina • Petrosiidae

Fig. 2. Neopetrosia de Laubenfels, 1949b. A-D, type species, Neopetrosia longleyi de Laubenfels, I 949b. A, lectotype USNM 22475 (scale 1 cm).
B, ectosomal skeleton, dense and tangential isodictyal surface network of one spicule length (scale 100 j.Lm). C, fine hispid spicule brush issued from nodal
points of mesh (scale 20 j.Lm). D, longitudinal section through the surface. Subectosomal region appears as a compact irregular network of short spicule
tracts, with a fine 'paper-thin area' (arrow) about I mm below the surface (de Laubenfels, 1953b) is clearly visible (scale 200 j.Lm).

Eastern North America (de Laubenfels, 1953b); West of Big Marco (1980 and in litteris) considered Neopetrosia to be a valid genus
Pass, Gulf of Mexico (de Laubenfels, 1953a); West Indian region that could include some species which up until now have been
(de Laubenfels, 1954); La Blanquilla, Veracruz, Gulf of Mexico placed in Xestospongia (e.g., X. subtriangularis, X. exigua and
(Green, 1977). X. pacifica) containing oxeas smaller than 200 f..I..m (atypical for
Remarks. De Laubenfels (1932) observed that his new Xestospongia).
species was for a long time regarded as being 'Haliclona The choanosomal skeleton of Neopetrosia is differently
("Thalysias') subtriangularis Duchassaing & Michelotti", but in his organised from that of Xestospongia, with the latter a reticulate
opinion 'H. longleyi differed from H. ('T.') subtriangularis in hav- coarse arrangement with large meshes lacking any pattern, versus
ing smaller spicules, (the latter being 140 f..I..m in length by 7 f..I..m in a more compact structure in Neopetrosia. Neopetrosia also differs
diameter), and its colour (green instead of yellow in life, and white from Xestospangia in the type of ectosomal network (isodictyal,
instead of drab, when dry, respectively). In H. longleyi we also with one spicule length side-by-side, and spicule brushes at the
observed a different pattern of tracts, with the choanosomal mesh in nodal points), and the possession of a second subectosomal spicu-
H. subtriangularis more variable (from 80-200 f..I..m) than in H. lan- lation (Fig. 2D). When comparing the ectosomal skeletons of
gleyi. De Laubenfels (1932) remarked that it was more certain Neopetrosia and Xestospongia we observe in both genera an ecto-
Schmidt's (1870: 44) Schmidtia aulopara and H. subtriangularis somal layer produced in the same way, but in Neopetrosia this con-
were synonyms, than H. longleyi. sists of a simple tangential unispicular isodictyal network of small
In Bergquist's (1965) opinion Neopetrosia was "incompletely spicules, instead of the dense ectosomal brushes of large spicules
defined" and not different from Xestospongia, as defined by present in Xestospongia. The description of the type species of
de Laubenfels (1950a: 49) and by herself (1965: 151). This is in Neopetrosia by de Laubenfels (1949b) did not mention all the char-
part supported by the fact that both genera, Neopetrosia and acters present in Neopetrosia longleyi, but later (de Laubenfels,
Xestospongia, are defined as having only a single size class of 1954: 83), in describing Neopetrosia pandora, modified its
oxeas. Bergquist (1965) suggested emending "the definition of description and mentioning the presence of a typical 500 f..I..m thick,
Xestospongia to include sponges with a distinct reticulate skeleton special dark subdermal layer that we also observed in N. longleyi
but where the accumulation of spicules may be so great as to (Fig. 2D). We cannot say what the value of this structure is, which
obscure this in parts or in all of the sponge, except the subdermal is produced by the choanosomal skeleton, consisting of longitudi-
region". Wiedenmayer (l977b: 113) considered characters of nal fibres of about 500 f..I..m long. This structure is visible as a
Neopetrosia as integrating with those of Xestospongia, and secondary surface layer overlaying the primary layer, and is similar
suggested the former was a junior synonym of the latter, in accor- to the structure observed in other genera of Haplosclerida (e.g.,
dance with Bergquist's (1965) position. Conversely, Van Soest Callyspongia (Euplacella), or a peripheral condensation at the
Porifera' Demospongiae • Haplosclerida • Petrosina • Petrosiidae 911

Table 3. Comparison of morphology, spicule geometry and spicule sizes rounded meshes. A dense interstitial reticulation of free spicules
between Neopetrosia and Xestospongia (measurements in JLm taken from gives the sponge a stony texture. Megascleres include oxeas and
type material, represented as minimum-mean- and maximum dimensions). strongyles or only strongyles. Microscleres present or absent, when
Genus Morphology Choanosomal Spicules
present are centrangulate microxeas.
Skeleton
Distribution
Neopetrosia Ramose, lobulate, Irregular multi- Oxeas
de Laubenfels, consistency stiff, paucispicular 104-120-l44X Mediterranean; Central Atlantic, North, central and South
1949b surface finely tracts, rounded 2-3-4 Pacific Oceans; Indian Ocean.
hispid, single meshes
isodictyal network
Previous reviews
of small spicules
Xestospongia Lamellar, massive, Dense, short Oxeas Refer to subgenera.
de Laubenfels, surface strongly discontinuous 205-450 X
1932 hispid with tracts( 6-10 sps) 8-30
abundant
Remarks
brushes of larger
spicules Prior to the creation of the family Petrosiidae (Van Soest, 1980)
Petrosia was associated with different genera and often assigned to
Renieridae Schmidt, 1870 (=Chalinidae). Topsent (1928c) consid-
ered differences between micro- and megascleres, observed in some
genera of Haplosclerida, e.g., Strongylophora and Petrosia, as
surface. With regard to the size of its spicules (less than 200 JJ-m invalid, and his concept of P. dura includes specimens exhibiting
long) and the recognizable anisotropic pattern of its skeleton (with microstrongyles and microxeas in the ectosomal skeleton. For the
a relatively small mesh size), Neopetrosia is here considered a same reason, Strongylophora sensu Topsent (1928c) should fall
valid genus. In Van Soest's opinion (in litteris) at least H. longleyi, into synonymy with Petrosia.
H. subtriangularis and X. exigua are members, as well as many of The history of the genus is convoluted. De Laubenfels (1936a)
the similar 'hard' Renieral Haliclona species described in the liter- considered Petrosia as a genus exhibiting the most complicated
ature (e.g., X. pacifica). spiculation of megascleres of the Adociidae (=Chalinidae), in
Neopetrosia has also been compared with Petrosia based on which he also included species originally described as Petrosia that
their similar external morphology and consistency in the dry state. he later transferred to Xestospongia (viz., X. exigua, X. testudinaria
They differ especially in their ectosomal skeleton and the size and X. muta). Levi (1973) remarked on the predominance of silica
categories of their spicules. over spongin as the principal characteristic of Renieridae (includ-
ing Petrosia), and associated genera such as Reniera, Petrosia,
Pachychalina and Gellius, with genera displaying the characters of
PETROSIA VOSMAER, 1885 Phloeodictyidae (e.g., Oceanapia, Calyx, Pellina).
Van Soest (1980) considered that Petrosia and Strongylophora
Synonymy were distinct from these other 'Renieridae' owing to their posses-
sion of small strongyles. More recently, however (Van Soest, pers.
Refer to subgenera. comm.), he suggests that this may not be the case, and furthermore,
the presence or absence of 'microxeas' may have a poor taxonomic
Type species value at the supra-specific level. Supporting evidence to differenti-
ate Strongylophora from Petrosia also came from their different
Rayneria dura Nardo, 1833 (by monotypy). chemical profiles published within the marine natural products
chemistry literature, with Petrosia reported to have hydroxylated
Definition acetylenes and Strongylophora strongylophorines. More recently,
Van Soest (pers. comm.) identified a 'typical' Strongylophora
Petrosiidae with an ectosomal triangular or polygonal reticu- (i.e., with all the spicule types found in the type species), having
lation of spicule tracts or single spicules, usually echinated at the acetylenes very similar to those in many Petrosia. Morphological
nodes or along the tracts by a smaller category of spicules. differences between the type species of both these genera were
Choanosomal skeleton basically a lamellate-isotropic reticulation found mainly in their different spicule morphologies and size
of spicule tracts, and an interstitial unispicular reticulation. classes, but these differences, in the opinion of Van Soest (in
Megascleres with distinct size categories of strongyles or oxeas, litteris), are just two or three size classes which are commonly
often with a special category of ectosomal microxeas. found in Petrosia species: large oxeas (oxeote or strongylote) of
the main skeleton, small oxeas or strongylotes of the surface and
Diagnosis an intermediate class often found, but not always present. When
only strongyles are present in other Petrosiidae there is always
Petrosiidae with a tangential specialised ectosomal unispicu- a category of 'juvenile' thinner oxea spicules, but in typical
lar network (subgenus Petrosia) or a very dense irregular tangential Strongylophora the strongyles and oxeas have similar dimensions
ectosomal reticulation of free strongyles of different sizes, mostly and can be matched easily, and thus the latter clearly do not
large microxeas (subgenus Strongylophora). Choanosomal skele- represent 'juvenile' spicules. In the type species of Strongylophora,
ton a dense network of thickly crowded spicule tracts producing S. durissima, and in several other species the smallest spicules
912 Porifera· Demospongiae • Haplosclerida • Petrosina • Petrosiidae

are sharply pointed oxeas, abruptly curved at the middle. In Van A part of the type specimen, which was deposited in the Bristol
Soest's opinion (in litteris), if a difference between Petrosia and Museum, is kept in the Natural History Museum, as a dry specimen
Strongylophora exists, it is perhaps found in this spicule type. Thus accompanied by two slides (BMNH 1909.8.15.1). The material con-
we relegate Petrosia and Strongylophora to subgenera of a single sists of two pieces of branches, each 2 cm long and 1.5 cm diameter,
genus taxon, with Petrosia being the senior most available name. surface smooth, but undulated, no apparent oscules. The original
Chalinorhaphis Lendenfeld, 1887, with type species specimen is a much larger hand-shaped form with seven branches.
Chalinorhaphis armata Lendenfeld, 1887: 821, pI. XXVIII fig. 1 The skeleton consists of a reticulation of thick spicule bundles, mak-
(by subsequent designation, de Laubenfels, 1936a), is considered ing elongated meshes. At the surface there are ectosomal spicule
here as a probable synonym of Petrosia based on the reported size brushes composed of smaller spicules than those of the subectosomal
of its oxeas (450 X 30 /-lm and the form of its skeletal meshes. It is and choanosomal bundles. Megascleres oxeas in at least two size cat-
described as being irregularly massive, with hard texture, lOX 3 cm, egories, 250-280 X 10 /-lm and 30-40 X 2/-lm. The structure and spic-
with its skeleton a reticulation of spiculofibres (70/-lm diameter) ulation of this sponge conform to the genus Petrosia, and the species
forming large meshes (1000/-lm diameter). However, this syn- may be identical to the species described by Wiedenmayer (1977b:
onymy is tentative as the genus is poorly known only from its type 123) as Cribrochalina dura (Wilson, 1902). Thus, Pharetronema is a
locality (Port Jackson, NSW) (Hooper & Wiedenmayer, 1994). synonym of Petrosia, and because it is a senior name, it has to be
We recognize in the genus Petrosia the possession of a lamel- suppressed under ICZN Article 23.9 as an unused name (not used
late skeleton with regular isodiametric meshes, where the primary after 1899) in favour of Petrosia which has been used more than
and secondary fibres are present but indistinguishable, and the pres- 25 times by 10 different authors in the past 50 years.
ence of spicules typical of the family (oxeas or strongyles), as the
typical diagnostic characters of the genus Petrosia. Aside from these
singular traits we cannot ignore its affinity with genera displaying
similar ectosomal skeletons, of varying degrees of development. SUBGENUS PETROSIA VOSMAER, 1885
For example, the extremely dense tangential ectosomal skeleton
of Petrosia is reminiscent of the crust-like ectosomal skeleton Synonymy
of Cribrochalina (Niphatidae), Sclerochalina (=Siphonochalina)
and Callyspongia (Euplacella) (Callyspongiidae). Wiedenmayer [Schmidtia] Balsamo Crivelli, 1863 (preocc.). [Pharetronema]
(1977b) also associated Cribrochalina [in part] with Petrosia and Sollas, 1879b: 404 (nomen oblitum). Petrosia Vosmaer, 1885b:
Xestospongia, assigning them all to the family Nepheliospongiidae. 338. ? Tedaniella Czemiavsky, 1879: 376 (uncertain affinity). ?
In Van Soest's (1980) opinion the structure of Cribrochalina (with Chalinorhaphis Lendenfeld, 1887: 821.
strong primary fibres, rich in spongin and radially distributed to
form a crust), is very different from the isotropical mesh of Petrosia Type species
and the skeletal structure of Xestospongia.
There is still some debate about the affinities of the type Reniera ? dura (Nardo, 1833) (by monotypy).
species of Petrosia, Reniera dura (Nardo) with other nominal taxa,
and we suggest that this issue cannot be resolved through morpho- Definition
metric data alone. As an example, some authors considered the type
species to be a synonym of P. jiciformis (Poiret, l789a [1980]) Petrosia with spicule complement including two or three size
(e.g., Riitzler, 1965: 39: Wiedenmayer, 1977b: 113: Desqueyroux- categories of oxeas or strongyles, in which the smallest is concen-
Fatindez, 1987: 187), and the micrometries of both species (Table 4) trated at the surface.
illustrate the difficulty in separating these species based on morpho-
metrics alone. Balsamo Crivelli (1863) described four species of Diagnosis (emended from Van Soest, 1980 and
Schmidtia (s. jiciformis, S. fungiformis, S. clavata and S. dura), De Weerdt, 1985)
which according to Schmidt (1864: 42) comprised only two taxa
(S. dura and S. clavata). Vosmaer (1885b) also suggested that Petrosia with a tangential specialised ectosomal unispicular
S. aulopora and S. muta (=Xestospongia), described later by network, and a very dense lamellate-isotropic choanosomal skele-
Schmidt (1870: 44,45), might also belong to S. dura. This potential tal network of thickly crowded spicule tracts producing rounded
synonymy cannot be resolved at present. Topsent (1892a: 68; 1904) meshes, forming layers parallel to the surface. A dense interstitial
described P. clavata from the Azores, emphasising the possession of reticulation of free spiCUles gives the sponge a stony texture.
'kidney shaped' spicules in these populations contrasting to the pos- Megascleres at least two distinct size categories of oxeote or
session of 'short strongyles' in P. dura, with the implication that strongylote spicules. Microscleres absent.
such geometric differences in spicule morphologies may be pivotal
to their specific identities. Similarly, the number, position and size Distribution
of oscules in species of Petrosia vary significantly and might pro-
vide an additional character to differentiate species of Petrosia. Mediterranean; Central Atlantic, North, central and South
Ultimately, however, future revisions of Petrosia must use charac- Pacific Oceans.
ters other than those described above, which are widely open to sub-
jective interpretation (e.g., chemical and molecular data), to more Previous reviews
precisely define species and reconstruct the phylogeny of Petrosia.
We also include in the synonymy of Petrosia, the genus Topsent, 1892a: 68; 1904: 240; Dendy, 1905 (in part): 144;
Pharetronema Sollas, 1879b: 404 (type species Pharetronema Bergquist & Warne, 1980: 35; Van Soest, 1980: 74; Desqueyroux-
zingiberis Sollas, 1879b: 404, pI. XXX, figs 1-15 from Jamaica). Fatindez, 1987: 181; De Weerdt, 1985: 82; Fromont, 1991: 87.
Porifera' Demospongiae • Haplosclerida • Petrosina • Petrosiidae 913

Table 4. Comparisons between various specimens of P. dura (Nardo) and P. ficiformis (Poiret) (measurements in !Lm, taken from the literature or our
*own observations, represented as minimum-maximum, or minimum-mean and maximum dimensions).

Species, Locality Oxeas Oxeotel Affinities

material (Growth Strongylote
form)
----- -------------- - --- -- ----- --- -------- ----- - ----

R. dura
Schmidt, 1862 Sebenico (massive- not recorded not recorded P. ficiformis
lobate) lectotype
*BMNH 1867.7.26. Lesina, Adriatic 30-58 X 2-3 182-237 X 10-14
18 (massive-lobate)
*LMJG 153301169 Lissa, Lesina 38-58-96 X 2-3 107-162-180 X 6-8-12
(massive-lobate) 67-79-112 X 5-6
40-136 X 6-10 150-175-400 X 10
P. ficiformis
Topsent, 1933 Indian Ocean ? 250-280 X 15 microxeas probably
(massive-branched) *280-310 X 20-22 sausage-shaped Strongylophora
Riitzler, 1965 Rovinj (massive 2-160cm 2) 25-260/4-12 not recorded P. ficiformis
Pulitzer-Finali, 1977 Bay of Naples 45-230/6.7 absent P. ficiformis
(habit not
recorded)
Sara, 1972 Mediterranean not recorded not recorded P. ficiformis
(massive,
branched-lobate)
Desqueyroux- Ambon (massive- not recorded 74-140 X 3.2 Petrosial
Faundez, 1987 lobate tubes: 218-286 X 6.4 Strongylophora sp.
21 X 7.5-lOcm)
De Weerdt & SE North Atlantic small oxeas middle-sized P. ficiformis
Van Soest, 1986 (ramose-repent, 45 X 1 oxeas
massive- 50 X 3.5 120-200 X \.5-2.5
flabelliform, 65 X5 140 X 7.5
6 X3.5 X 5cm) large-sized oxeas
240 X 10-15

Description of type species (Table 3) oxeas, strongyloid, oxeote or strongylote in three sizes,
the larger ones oxeote to strongylote, 182-212-400 f.Lm long,
Petrosia dura (Nardo, 1833) (Fig. 3). 10-13-14f.Lm wide; the medium-sized ones oxeas, 40-80-136f.Lm
Synonymy. Rayneria dura Nardo, 1833: col. 519; Reniera ? long, 5-7-1Of.Lm wide; the small ones oxeas, 30-58-96f.Lm long,
dura; Schmidt, 1862: 76, pI. 7 fig. 13, 13 a-e. Schmidtia dura; 2-3 f.Lm wide. Within the last category of oxeas we also observed
Balsamo Crivelli, 1863: 293; pl. 5 figs 1-3. Petrosia dura; some bent strongyles of the same size.
Vosmaer, 1885b: 338. Non Cribrochalina dura sensu Remarks. Refer to genus.
Wiedenmayer, 1977b: 123.
Material examined. Lectotype (here designated): LMJG Distribution
15330 (dry) - Lissa, Lesina. Other material. LMJG 15627 (dry) -
Sebenico. BMNH 1867.7.26.18 (dry) - Lesina, Adriatic. BMNH Naples, Rovinj, Adriatic, SE North Atlantic; Ambon,
1867.3.11.48 (slide), 1877.5.21.321 (dry) - Adriatic (Bowerbank Indonesia; Abrolhos Islands, Western Australia.
collection, 'fragments from Schmidt').
Description. Massive sponge, irregularly globular, with a large
base that produces several fused lobes of 35-40 mm in diameter and
up to 80 mm high. Aquiferous system with a terminal deep aquiferous SUBGENUS STRONGYLOPHORA DENDY, 1905
cavity connected with a unique volcano-shaped osculum at the end of
the lobes. Numerous oscules in the internal wall of the aquiferous Synonymy
cavity. Surface smooth, fine, compact, covered by a fine ectosomal
layer, hispid to the touch. Texture hard, firm. Ectosomal skeleton a Strongylophora Dendy, 1905: 141, pI. 9 fig. 1; Wilson,
fine layer of free spicules, about 1 mm deep, producing a tangential 1925: 391.
compact network with isodiametrical meshes. Choanosomal skeleton
a regular system of multispicular compact tracts or fibres with isodia- Type species
metrical meshes without differences between primary and secondary
fibres. Free spicules abundant. Spongin not visible. Megasc1eres Strongylophora durissima Dendy, 1905: 141 (by monotypy).
914 Porifera· Demospongiae • HapJosclerida • Petrosina • Petrosiidae

Fig. 3. Petrosia (Petrosia) Vosmaer, 1885b. A, lectotype LMJG 15330/169. B, Alcyoniumficiformis Lamarck, Lamarck collection BMNH 1954.2.20.80,
enlarged view of the surface (scale 200 fLm). C, lectotype, surface, tangential view (scale 1 cm). D, Schmidt's specimen LMJG 15330/169, compact
choanosomal fibres (scale 500 fLm).

Definition Description of type species

Petrosia with three to five categories of oxeas or strongyles, Strongylophora durissima Dendy, 1905 (Fig. 4).
including centrangulate microxeas. Synonymy. Dendy, 1905: 141, pI. 9, fig. 1; Wilson, 1925: 393.
Material examined. Lectotype (here designated): BMNH
Diagnosis (emended from Dendy, 1905) 1907.2.1.37 - Ceylon, Indian Ocean, Herdman Collection
(RN 156). Other material. Specimens of Strongylophora duris-
Petrosia with a very dense irregular tangential ectosomal sima: BMNH 1926.9.3.54 (dry)- South Africa, Gilchrist Collection
reticulation of free strongyles and oxeas of different sizes (RN 271). BMNH 1926.9.3.23 (dry) - South Africa, Gilchrist
echinated by small centrangulate microxeas. Large strongyles are Collection (RN 88). BMNH 1926.9.3.84 (dry) - South Africa,
grouped and radiate from the nodes. Small bundles of short Gilchrist Collection (RN 15). BMNH 1926.9.3.85 - South Africa,
strongyles (sometimes sausage-shaped) are also present at the Gilchrist Collection (RN 267). BMNH 1904.12.1.32-44 (several
nodes, giving the surface its granular appearance. Choanosomal dry specimens) - South Africa, Gilchrist Collection.
skeleton a dense network of strongyles which are partly assembled Description. Sponge massive, irregularly depressed or cake-
to form strong tracts with no visible spongin. An interstitial net- like, subcylindrical and slightly ramose (Fig. 4A). Surface very
work of connecting tracts of strongyles and free strongyles gives uneven and wrinkled, with grooves and ridges, minutely and uni-
the sponge a stony texture. formly granular. Sponge relatively cavernous (Fig. 4B) with few
large oscules irregularly scattered, representing the opening of
Distribution wide aquiferous canals. Numerous ostia are scattered in between
the mesh of the ectosomal skeleton. Texture hard, stony, brittle.
Shallow waters of warm-temperate and tropical seas. Indian Ectosomal skeletal network (Fig. 4C) clearly differentiated from
Ocean (Sri Lanka), and Indo-west Pacific, from Western Australia the choanosomal skeleton in the form of a tangential network of
to Vanuatu (Hooper & Wiedenmayer, 1994; Hooper, pers. comm., large and abundant free strongyles and loose tracts of strongyles or
unpublished data). single strongyles forming the sides of irregularly triangular or
quadrangular meshes. At the angular points ('comers') of the
Previous reviews strongyle-tracts are groups of very short strongyles which form
surface nodes that produce the granular-aspect of the surface.
Topsent, 1917; 1928c; de Laubenfels, 1934: 19; 1936a: 70; Abundant smaller spicules, resembling large microxeas, are also
1950a: 64; Van Soest, 1980: 78; Wilson, 1925: 323. present in the meshes. Choanosomal skeleton very dense, compact
Porifera' Demospongiae • HapJosclerida • Petrosina • Petrosiidae 915

Fig. 4. Petrosia (Strongylophora) Dendy, 1905. A-D, type species, Strongylophora durissima Dendy, 1905. A, holotype BMNH 1907.2.1.37 (scale I em).
B, longitudinal section through the surface, on top (scale 500 fLm). C, eetosomal tangential network of large and abundant free strongyles and loose
strongyle-tracts or single strongyles forming the sides of irregularly triangular or quadrangular meshes (scale 200 fLm). D, ehoanosomal tracts and view of
spicules, long and small strongyles, oxeas and mieroxeas (scale 50 fLm).

sub-rectangular network (Fig. 4D) with small meshes produced by Strongylophora and suggested that they did not represent real
stout spicular tracts of closely packed strongyl a and free spicules. microscleres but were reduced and modified megascleres.
Toward the surface are stout multi spicular longitudinal tracts Consequently, Strongylophora sensu Topsent is a synonym of
formed by radiating bundles of single strongyles, closely packed, Petrosia.
without visible spongin. Numerous free strongyles and microxea- De Laubenfels' (1934: 19, 1936a: 70, 1950a: 64) concept of
like spicules are present between the tracts. Megascleres (Fig. 4D) Strongylophora included strongylote megascleres, oxeas,
included larger strongyles, stout, regularly bent or centrangulate, rhaphides and microrhabds or sigmas as microscleres, approaching
with rounded ends, 156-208-238/Lm long, 12-14-20/Lm wide; closely the concept of Sigmadocia (=Haliclona) (e.g., S. amphioxa
slender strongylotes, 143-200-230 fLm long, 4-7-8 fLm wide, and de Laubenfels, 1950a from Bermuda exhibits a typical Chalinidae
oxeotes, 164-193-205 /Lm long, 4 /Lm wide; and a second category skeleton; S. amphioxa and S. rampa de Laubenfels, 1934 with
of strongyles occurs in the ectosomal skeleton (some sausage- large strongyles and oxeas, were transferred to Adocia and
shaped, not considered to be microscleres; Dendy, 1905), Xestospongia, respectively, by Van Soest, 1980). Only a few
20-34-53 fLm long, 4-7-8/Lm wide; and smaller microxea- species of Strongylophora have been described to date although a
like spicules, centrangulate, 28 fLm long, 2/Lm wide (Dendy, thorough revision of Petrosia species is required to distribute
1905), 21-26-30 fLm long, 2-3 fLm wide (re-examination of type species based on possession of strongylote oxeas and smaller
material). These microxea-like spicules are especially abundant sausage-shaped strongyles to subgenus Strongylophora. This is
in the dermal membrane, but less common in the choanosomal the case for Petrosia strongylata Thiele (1900: 938), which was
skeleton. transferred to Strongylophora (Van Soest, 1980: 78) and P. ushit-
Remarks. Strongylophora is a remarkable subgenus, in spite suensis Tanita, 1963 (transferred here to Strongylophora).
of its doubtful taxonomic position due to its ubiquitous possession There are undoubtedly many more putative species of Petrosia
of microxeas (considered as microscleres by Dendy, 1905, but per- that require re-evaluation. Pulitzer-Finali (1996) included
haps representing only 'juvenile' spicules - see remarks for genus Strongylophora corticata Wilson, 1925 in his genus Tabulocalyx
Petrosia). Topsent (1917) compared Strongylophora with his genus Pulitzer-Finali, 1993 (Phloeodictyidae), which is clearly a
Microxina, with resemblance between their respective skeletal Strongylophora since it contains small sausage-shaped strongyles.
structures and the presence of microxeas in both. In addition, The choanosomal network of Strongylophora exhibits a very
both genera shared a 'fibrous skeleton', which in Topsent's (1917) strong, compact skeletal structure, typical of the Petrosiidae type
opinion was reason enough to include them in Axinellidae. Later of skeleton, especially near the surface where roundish surface
(Topsent, 1928c) reconsidered his opinion of the microxeas in meshes are evident and the ectosomal skeleton structure appears as
916 Porifera· Demospongiae • HapJosclerida • Petrosina • Petrosiidae

Fig. 5. Xestospongia de Laubenfels. 1932. A-D, type species, Haliclona diprosopia de Laubenfels, 1930. A, holotype USNM 21509, lamellar subspheri-
cal specimen (scale 1 cm). B, longitudinal section through the surface, showing ectosomal tangential disordered network of abundant spicule brushes
(scale I mm). C, choanosomal skeleton, showing a highly dense network of short longitudinal undivided irregularly parallel tracts, pauci- to multi spicular
(scale 500 !Lm). D, oxeas (scale 100 !Lm).

a single tangential layer, which is clearly different from the Diagnosis

choanosomal skeleton.
Ridley & Dendy (1887: 13) mentioned Reniera craterifonnis Petrosiidae lacking a special ectosomal skeleton, but a tangen-
Carter, 1882b as possessing sausage-shaped spicules although tial disorganised network of free spicules which originate from the
Carter's original description did not mention these. Hooper & terminal ends of choanosomal tracts. Choanosomal skeleton a
Wiedenmayer (1994) included R. craterifonnis in synonymy with dense network of short longitudinal undivided pauci- to multispic-
Xestospongia testudinaria but confirmation of this synonymy is ular tracts of 6-10 spicules, connected by few oxeas. Roundish to
impossible since the BMNH holotype was not found. polygonal large-spaced meshes delimited by unordered spicules.
Free spicules are not present, but spicule brushes formed by the
ends of connecting spicules are visible on the internal part of the
XESTOSPONGIA DE LAUBENFELS, 1932 mesh walls. Megascleres: oxeas with some strongylote and stylote
forms, in a single size class, slightly bent, variable in size but
Synonymy not assigned to more than one class, larger than 200/-Lm long.
Microscleres absent.
Xestospongia de Laubenfels, 1932: 115, fig. 69; 1936a: 46. ?
Densa de Laubenfels, 1934: 14. ? Quepanetsal de Laubenfels, Distribution
1954: 180, fig. 119.
West Indies, Cape Verde Islands, Boreal Pacific,
Type species Mediterranean, Atlantic, Boreal E Atlantic, West Africa, Australia
and Antarctica (Van Soest, pers. comm.).
Haliclona diprosopia de Laubenfels, 1930 (by original
designation). Previous reviews

Definition De Laubenfels, 1936a: 46; 1950a: 49; 1954: 80; Bergquist,

1965: 149; Van Soest, 1980: 66; Bergquist & Warne, 1980: 36;
Petrosiidae with an ectosomal skeleton consisting only of an Desqueyroux-Faundez, 1987: 194; Fromont, 1991: 79; Hooper &
isotropic reticulation of single spicules or spicule tracts. Wiedenmayer, 1994: 329.
Porifera· Demospongiae • HaploscIerida • Petrosina • Petrosiidae 917

Description of type species stages, 254-352-450 and 205-296-387 f.Lm long, 16-23-30 and
8-10-12 f.Lm wide.
Haliclona diprosopia de Laubenfels, 1930 (Fig. 5). Remarks. Slides prepared from the BMNH specimen do not
Synonymy. Haliclona diprosopia de Laubenfels, 1930: 28; correspond to the skeleton described by de Laubenfels (1932).
1932: 115; Wiedenmayer, 1977b: 112. Bergquist (1965) already remarked on this discrepancy, which pri-
Material examined. Holotype: USNM 21509 - California, marily concerns measurements of the choanosomal reticulation
Monterey Bay, colI. Rickets, 1929. Other material. BMNH and well-identified mesh differentiation which were not mentioned
1929.8.22.59, 59a (dry specimen and slide) - Monterey Bay, by de Laubenfels (1932). Bergquist (1965: 150) also remarked on
California (label "Haliclona diprosopata de Laubenfels, MS, 1932"). the practical difficulties in differentiating Xestospongia from
Description. Lamellar subspherical, massive sponge Neopetrosia based only on the generic diagnosis, whereas we con-
(Fig. SA). Aquiferous system represented by numerous oscula, clude, from examination of respective type material that differ-
3 mm in diameter, scattered on one face of the lamellae, connected ences in choanosomal skeletal structure, and especially spicule
with shallow subdermal cavities with numerous ramifications. micrometries, exist between both genera that are sufficient to dis-
The non-oscular side is pierced by numerous ostia, at least 10 f.Lm tinguish them at the generic level, supporting their continued
in diameter (de Laubenfels, 1932). Compact sponge without large recognition as distinct taxa (see remarks for Neopetrosia). Like this
aquiferous spaces. Surface smooth, rough, regularly and strongly latter genus Xestospongia has only a single size class of oxeas,
hispid. Texture, hard, friable. Ectosomal skeleton a tangential although both may be distinguished by the relatively larger size of
disordered network of abundant spicule brushes formed by the oxeas in Xestospongia, larger than 200 f.Lm long.
ectosomal ends of choanosomal longitudinal tracts producing a Zea (1987) included Densa araminta de Laubenfels, 1934
strongly hispid surface (Fig. 5B). Choanosomal skeleton a highly (type species of Densa de Laubenfels, 1934, by original designa-
dense network (Fig. 5C) of short longitudinal undivided, irregu- tion), in the synonymy of Xestospongia proxima (Duchassaing &
larly parallel tracts, pauci- to multispicular (6-10 spicules). Michelotti, 1864). In this work, we tentatively designated the genus
Longitudinal tracts are not continuous and there are a variable Densa as subjective junior synonym of Xestospongia. Van Soest (in
number of short longitudinally interconnecting tracts parallel to the litteris) studied a fragment of Quepanetsal madidus de Laubenfels,
principal tracts, contributing to the high density of the skeleton. 1954, from the west Central Pacific, a species associated with
Primary tracts are interconnected by only a few oxeas or bundles of Halichondriidae by de Laubenfels. In the opinion of Van Soest this
spicules (4-6), with scarce or no visible spongin, forming round to genus is definitely a haplosclerid and probably a Xestospongia.
polygonal meshes, with no free spicules but obscured by ends of The alleged roughened rounded ends of the oxeas are just tele-
connecting oxeas and spicules from the longitudinal tracts of the scoped endings, common to some other Petrosiidae species. The
choanosomal skeleton. Megascleres (Fig. 5D) oxeas with some strongyles are the dominant and mature spicule, the oxeas being
strongylote and stylote forms, slightly bent and with ends of the obvious growth stages (same length and only slightly thinner than
same diameter. Spicules in only one size category showing the oxeas). In this work, we also tentatively consider Quepanetsal
variability only in thickness, and thus they are probably growth as subjective junior synonym of Xestospongia.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Family Calcifibrospongiidae Hartman, 1979

Rob W.M. Van Soest

Zoological Museum, University of Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, Netherlands. (soest@science.uva.nl)

Calcifibrospongiidae Hartman (Demospongiae, Haplosclerida) combines a stromatoporoid grade of coralline skeleton with a uni- or
paucispicular skeleton of thin siliceous strongyles. Since the coralline skeleton of 'sclerosponges' is demonstrated to be polyphyletic, the
family is classified on the basis of its spicule skeleton. The strongyles in a light reticulation reveals the family as a haplosclerid, positioned
in the suborder Petrosina. There is one monotypical Recent genus, Calcijibrospongia Hartman, 1979, possibly related to some fossil
coralline sponges.
Keywords: Porifera; Demospongiae; Haplosclerida; Petrosina; Calcifibrospongiidae; Calcifibrospongia.

DEFINITION, DIAGNOSIS, SCOPE a family of its own. The strongyles in isodictyal reticulation with-
out clear primary and interconnecting tracts are indicative of the
Synonymy suborder Petrosina.

Calcifibrospongiidae Hartman, 1979: 468. Previous reviews

Definition Hartman, 1979; Hartman & Willenz, 1990; Reitner, 1992.

Petrosina with stromatoporoid calcareous basal skeleton and

a reticulation of siliceous strongyles. CALCIFIBROSPONGIA HARTMAN, 1979

Diagnosis Synonymy

Semiglobular or flattened aragonitic masses covered by a thin Calcifibrospongia Hartman, 1979: 468.
veneer of organic tissue. Surface with evenly scattered depressed
oscules. Siliceous skeleton a reticulation of thin strongyles. No ecto- Type species
somal specialization. Aragonitic basal skeleton consists of a mesh-
work of tubes, pillars and lamellae that intergrade peripherally with Calcijibrospongia actinostromarioides Hartman, 1979: 468
the spicular skeleton. The canal system penetrates the aragonitic (by monotypy).
meshwork to a varying degree depending on episodes of growth.
Definition and diagnosis
Scope
See above definition of family.
A single genus and species is recorded.
Description of type species
Taxonomic bistory
Calcifibrospongia actinostromarioides Hartman, 1979
Hartman (1979) referred this 'sclerosponge' to his class (Fig. lA-C).
Sclerospongiae, order Stromatoporoidea, in a separate family Synonymy. Calcifibrospongia actinostromarioides Hartman,
Calcifibrospongiidae. Although some morphological similarities 1979: 468, figs 1-13; Hartman & Willenz, 1990: 231, fig. 4; Reitner,
were noted with mesozoic stromatoporoids, no close relative 1992: 261, pIs 49-51; Willenz, 1993: 279.
appeared to exist among those fossils. Van Soest (1984a: 214) in a Material examined. Holotype (not seen): YPM 9114 -
discussion of the polyphyletic nature of 'sclerosponges' suggested Acklin Island, Bahamas. Paratypes (not seen): BMNH and USNM.
Calcifibrospongia could be a member of Haplosclerida: Chalinidae Other material. ZMA POR. 6157 - Andros Island, Bahamas,
(as Haliclonidae). Hartman & Willenz (1990) conceded that colI. J. Reitner.
Calcifibrospongiidae show convincing similarities to various rep- Description (from Hartman, 1979). Mushroom-shaped
resentatives of Haplosclerida. This was also the conclusion of (smaller specimens), dome-shaped to flattened with a broad base of
Reitner (1992: 261), although he pointed out that both a chalinid attachment. Size up to 30 X 30 X 10 cm. Surface smooth when
and a petrosid affinity were possible. In view of the fact that the alive, provided with many evenly distributed rounded slightly
aragonitic basal skeleton is unique in Haplosclerida, and also depressed oscules, with faint trace of astrorhizae. Color brownish
because affinity with either Chalinidae or Petrosiidae is not firmly tan or tannish orange alive. Soft parts - apart from the pinacoderm -
established, it is proposed here to maintain Calcifibrospongia as entirely incorporated in the basal calcareous skeleton. The inhalant

918
Porifera· Demospongiae • Haplosclerida • Petrosina • Calcifibrospongiidae 919

Fig. 1. A-C, Calcifibrospongia actinostromarioides Hartman, 1979 (all figures reproduced from Hartman, 1979, figs 1,2 and 7). A, ho1otype YPM 9114
(scale 1 em). B, spicular skeleton (scale 100 /-Lm). C, strongy1es (scale 10 /-Lm).

canals lead into the system of cavities of the calcareous skeleton, symbiont prokaryotes. In larger specimens a layer of 1.5-3 cm
and likewise the exhalant canals run through the calcareous mass thick is 'alive', underneath which there are 'dead' layers of similar
over considerable distance to end in vestibulae immediately under- thickness to mark off periods of death and regeneration. Spicules
neath the pinacoderm. The ectosomal region is supported for a are thin strongyles, 130-210 X 3-6 J.Lm. Distribution. Bahamas, in
small distance by free siliceous spicules arranged in a uni- or pau- shaded deep reef habitats.
cispicular isodictyal reticulation. In the subectosomal region the
spicule tracts, which are aligned in a collagenous matrix become Remarks
the focal points for calcification and further down into the sponge
the spicules become entirely enclosed in calcium carbonate. This sponge may be a living representative of a larger now
Siliceous skeletal meshes and calcareous cavities match closely in extinct group of haplosclerid 'sclerosponges' (see review in Reitner,
size, the smaller meshes varying from 55-175 J.Lm in diameter, 1992). However, in most cases the basal skeleton or the microstruc-
many larger represent various canals and tubes. Choanocyte cham- ture of the fossils differ from those of Calcifibrospongia. Some fos-
bers, about 20 J.Lm diameter, and mesohyl tissues are found to a sils have similar basal skeletons but certain traces of spicules are
variable depth within the calcareous basal mass. They are rich in lacking. So far this genus is monotypic.
Systema Porifera: A Guide to the Classification of Sponges, Edited by John N.A. Hooper and Rob W.M. Van Soest
© Kluwer AcademiclPlenum Publishers, New York, 2002

Suborder Spongillina subord. nov.: Freshwater Sponges

Renata Manconi 1 & Roberto Pronzato2

1 Department of Zoology and Anthropological Biology of the University, via Muroni 25, 1-07100, Sassari, Italy. (rmanconi@ssmain.uniss.it)
2 Department for the Study of the Territory and its Resources of the University, Corso Europa 26, 1-16132 Genoa, Italy. (pronzato@
hpe35 .dipteris. unige.it)

Spongillina subord. nov. (Demospongiae, Haplosclerida) consists of seven families of exclusively freshwater sponges together containing
45 genera: Spongillidae (21 valid genera), Lubomirskiidae (three genera), Malawispongiidae fam. nov. (five genera), Metaniidae (five
genera), Metschnikowiidae (monogeneric), Palaeospongillidae (monogeneric), Potamolepidae (six genera), some geographically wide-
spread and others highly endemic. A central body cavity is peculiar to Malawispongiidae. Skeletal network is typically multi spicular
alveolate-reticulate with scanty spongin in Metaniidae, Potamolepidae and Malawispongiidae, paucispicular irregularly reticulate in
Palaeospongillidae, Spongillidae, Metschnikowiidae with a variable amount of spongin, multi- to paucispicular irregularly reticulate with
an abundant amount of spongin in Lubomirskiidae. Smooth or variably ornamented megascleres range from oxeas to strongyles in
Spongillidae, Lubomirskiidae and Metaniidae, but are exclusively oxeas in Malawispongiidae, Metschnikowiidae, Palaeospongillidae,
and exclusively strongy1es in Potamolepidae. Microscleres usually present in Spongillidae, Palaeospongillidae and Metaniidae, are rare in
Potamolepidae, and absent in Lubomirskiidae, Malawispongiidae and Metschnikowiidae. Microscleres if present are oxeas, strongy1es,
aster-like, pseudobirotules. Larvae are always parenchymella. Gemmules are typical of Spongillidae, Metaniidae, Palaeospongillidae, rare
and strictly adhering to the substratum in Potamolepidae, and absent in Lubomirskiidae, Malawispongiidae and Metschnikowiidae.
Gemmular theca is monolayered in Potamolepidae, mono-, bi- or tri-layered in Spongillidae, generally tri-layered in Metaniidae.
Gemrnu1es usually armed by gemrnuloscleres in Metaniidae, Potamo1epidae and Spongillidae, are rarely naked in the latter.
Gemrnuloscleres are boletiform (tubelliform), parmuliform, pseudobirotules in Metaniidae, and oxeas, strongyles, birotules, pseudobiro-
tules, club-like, botryoidal in Spongillidae. Three more genera incertae sedis are included. This work is a relatively critical synthesis of
the literature, however, a critical phylogenetic revision of established taxa is still in progress.
Keywords: Porifera; Demospongiae; Haplosclerida; Spongillina subord. nov.; Spongillidae; Anheteromeyenia; Corvoheteromeyenia;
Corvospongilla; Dosilia; Duosclera; Ephydatia; Eunapius; Heteromeyenia; Heterorotula; Nudospongilla; Pachyrotula; Pectispongilla;
Racekiela; Radiospongilla; Sanidastra; Satumospongilla; Spongilla; Stratospongilla; Trochospongilla; Umborotula; Uruguayella;
Lubomirskiidae; Baikalospongia; Lubomirskia; Swartschewskia; Malawispongiidae fam. nov.; Cortispongilla; Malawispongia;
Ochridaspongia; Pachydictyum; Spinospongilla; Metaniidae; Acalle; Corvomeyenia; Drulia; Houssayella; Metania; Metschnikowiidae;
Metschnikowia; Palaeospongillidae; Palaeospongilla; Potamolepidae; Echinospongilla nom. nov.; Potamolepis; Potamophloios;
Oncosclera; Sterrastrolepis; Uruguaya; Genera incertae sedis; Balliviaspongia; Makedia; Ohridospongilla.

SPONGILLINA SUBORD. NOV. fibres (most families), or with microscleres in the dermal mem-
brane (Potamolepidae). Choanosomal skeleton with pauci- to
Definition multispicular tracts of megascleres forming regular to irregular
reticulate meshes, with or without lacunar or alveolate cavities, or
Exclusively freshwater sponges, with megascleres consisting of occasionally radial ascending paucispicular tracts without anasto-
oxeas or strongyles, smooth or spined, forming pauci- to multispicu- moses (Metschnikowiidae). Spongin usually sparse. Megascleres
lar tracts producing irregular to regular meshes, occasionally with oxeas to strongy1es, smooth, spiny or granular, occasionally with
large alveolate cavities (a central body cavity in one family); spongin larger tuberculate ornamentations, with sharply pointed, rounded
mostly sparse; microscleres present or absent, including smooth or or occasionally inflated points. Microscleres may include oxeas
spined oxeas, aster-like or birotule like spicules. Four families with or strongyles (smooth or microspined), pseudo-asters or pseudo-
gemrnules (resting bodies containing totipotent cells), which may birotules, but frequently absent. Gemmules, resting bodies
contain gemrnuloscleres of diverse morphology that is often diag- containing totipotent cells (thesocytes) are characteristic of
nostic. Three families lack gemmules. Where known reproduction is spongillines but present in only four freshwater sponge families
viviparous, with fully ciliated parenchymella larvae. (Spongillidae, Metaniidae, Palaeospongillidae, Potamolepidae),
and absent in three (Malawispongiidae fam. nov.,
Diagnosis Lubomirskiidae, Metschnikowiidae). If present, gemmules may
be dispersed within the choanosome or restricted to the sponge
Haplosclerida with growth form ranging from encrusting, base in contact with the substratum, occurring singly or forming
bulbose, globular, arborescent to massive. Surface smooth, hispid carpets. Gemmule morphology is variable, with or without a fora-
or conulose. Consistency generally not elastic but ranging to men (occasionally with several), with or without a gemmular
notably elastic in some taxa, and also varying from fragile, to cage connected to the skeletal network or basal plate. Gemmules
firm, to stony hard. Ectosomal skeleton with tangential layer of have a thin or thick gemmular theca formed by one, two or three
megascleres, or tufts of spicules arising from apices of primary layers of spongin. The intermediate one is the pneumatic layer

921
922 Porifera' Demospongiae • HapJoscierida • SpongiIIina

Malawlsponglldsu Melsnildso

-...'-j,....;..;,..-'-i'""....:. - ,- - .... ~{--~. -\r"_~

Melschnlkowlldao Palaeosponglllldae

Poramolupldae

Fig. I. Geographic di tributioll of the p ngillina famil ies.

arranged to fonn chambers, trabecules or fibres. Gemmules History, biology and evolution
can be naked or anned by gemmuloscleres organised radially
or tangentially, partially or totally embedded in the surface. Although sponges may generally provide good paleontological
Gemmulosclere morphology highly diverse, often diagnostic, material, most fossil records of freshwater sponges belong to
including smooth, spiny to granulated oxeas to strongyles, presently widespread genera, and date back to relatively recent
birotules, pseudobirotules, club-like, boletiform (tubelliform), periods of the Miocene, Pliocene and Pleistocene (Rezvoi et al.,
pannulifonn or botryoidal. Larvae hollow parenchymella, where 1971; Harrison & Warner, 1986). It is probable that the coloniza-
known, entirely ciliated, occasionally with larval oxeas. tion of inland waters occurred approximately in the Jurassic Period
Most freshwater sponges have a poor fossil record (not older (210-140 MYA), at the beginning of the disjunction phase between
than Miocene, Pliocene or Pleistocene) although one Africa and South America when the southern Atlantic was proba-
(Palaeospongilla) is recorded from the Cretaceous. bly no more than a narrow fjord characterised by shallow brackish
Porifera' Demospongiae • Haplosclerida • Spongillina 923

waters subject to strong thermal fluctuations (Pearson, 1978). Volkmer-Ribeiro (1990) have debated their mono- or polyphyletic
Among the few fossils known since the Mesozoic Era (Young, nature in detail.
1878; Ott & Volkheimer, 1972), Palaeospongilla chubutensis from
Patagonia, dated 100 MY, is characterised by a notably evolved Scope
gemmular architecture comparable to present taxa of Spongillidae.
The phenomenon of cryptobiosis and the modular bauplan of The present study provides an exhaustive overview of the
freshwater sponges were very precocious in the evolutionary his- taxonomy of freshwater sponges through comparison between
tory of this taxon, and spreading throughout inland waters was original observations and a more modem study of taxa. Our goal
favoured by the twofold specialisation of gemmules as resistant was to perform a detailed description, at the genus level, based on
bodies and propagules (Simpson & Fell, 1974; Frost et al., 1982; holotypes of type species of all existing freshwater sponge fami-
Pronzato & Manconi, 1994b, 1995). Growth rhythms range from lies. This very intriguing but difficult and long-term investigation
cyclic, with periodicity timed by climatic seasonal or annual fluc- resulted in a relative agreement with the taxonomic status of most
tuations, to continuous in perennial forms (Fell & Levasseur, genera, although in many cases our analysis highlighted critical
1991; Manconi & Pronzato, 1991; Fell, 1995). decisions of previous authors. Conversely, we refute the validity
The colonization strategy by gemmule-producing sponges of the recently described genus Clypeatula Peterson & Addis,
seems therefore mainly based on morphogenetic processes by 2000, considered a possible new species belonging to the genus
totipotent cells to regenerate the vegetative mother sponge as Ephydatia. At a higher level we assign to the new suborder
required by the spatial discontinuity of inland waters (Man coni & Spongillina all existing families of Malawispongiidae fam. nov.
Pronzato, 1994a), in the wide variety of unpredictable environmen- (= [Globulospongillidae1 Brien, 1973b), Lubomirskiidae Rezvoi,
tal constraints ranging from the long-term dry-up to permafrost or 1936, Metaniidae Volkmer-Ribeiro, 1986, Metschnikowiidae
anoxic conditions (Poirrier, 1969; Racek, 1969; Holmquist, 1973; Czerniavsky, 1880, Palaeospongillidae Volkmer-Ribeiro & Reitner,
Pronzato et al., 1993). Freshwater sponges are dispersed from the 1991b, Potamolepidae Brien, 1967, and Spongillidae Gray, 1867a.
water surface to a hundred meters depth, from tropical rainforests, A phylogenetic analysis is currently in progress.
cold and hot deserts, from the Arctic Circle to Patagonia (not yet
recorded from Antarctica), and in a wide range of environmental Methodology
conditions: oceanic islands, subterranean caves, thermal fields in
caldera lakes, ancient lakes, springs, ephemeral streams, bill- About five hundred specimens were examined to produce the
abongs, alpine lakes, salt lakes, large rivers, rapids and falls, estu- present review, although these are only partly listed in the materials
aries, brackish seas, man-made basins from tanks and reservoirs to examined, which were obtained from several museum collections
pools in gardens and fonts in archeological sites. throughout the world. This material was tracked down from publi-
During the evolutionary radiation of taxa in inland waters, cations of Gee (1931 et seq.), Potts (1888), Penney & Racek
adaptive traits of resistant bodies were subject to processes of (1968), Volkmer-Ribeiro & Traveset (1987). Holotypes, if avail-
remodelling to attain successful performances of passive dispersal, able, were examined in primis, or where not available we examined
survival and persistence. An extreme morphological plasticity was paratypes, schizotypes and topotypes. Original descriptions and
expressed mainly by the geographically widespread Spongillidae illustrations were repeated only when materials were not available
and Metaniidae (Penney & Racek, 1968). In these taxa resistant (e.g., Corvoheteromeyenia, Metschnikowia and Palaeospongilla).
bodies reached the highest complexity of gemmular architecture, Our observations were also compared to original descriptions of
with the theca arranged in two (outer and inner) compact layers of genera and other literature in order to perform a critical evaluation
spongin and one (middle) alveolar layer of trabeculated spongin, of morphological traits of each taxon, and we also considered
reinforced by spiny spicules in a variable spatial arrangement. eco-ethological characters to achieve a more defined morpho-
On the other hand, the family Potamolepidae, restricted to ancient functional analysis. A large set of diagnostic macro- and micro-
large hydrographic basins of the Afrotropical and Neotropical morphological characters was also considered: (1) body size,
Regions, displayed a limited radiation into few tropical genera external shape, consistency and colour; (2) architecture and fine
characterised by the production of rare gemmules with a simple organisation of ectosomal and choanosomal skeletons; (3) traits of
theca of compact spongin reinforced by non specialised spicules skeletal megascleres and microscleres; (4) gemmular structure and
(Brien, 1970a; Volkmer-Ribeiro, 1990). A perennial life style, gemmulosclere morphology. Methods used to examine skeletal
reproductive modes exclusively sexual, and dispersal by larval and gemmular structure, and spicule geometry using Light (LM) and
stages (Brien & Govaert-Mallebrancke, 1958) characterised Scanning Electron Microscopy (SEM) follow Manconi & Pronzato
Lubomirskiidae, Malawispongiidae and Metschnikowiidae from (2000). Fragments of the sponge surface and choanosome were
ancient lakes scattered in the Afrotropical, Neotropical, Palaearctic critical-point dried, mounted on stubs and gold sputtered to investi-
and Oriental Regions. The simple structure of gemmules in gate skeletal architecture. Entire gemmules and cross sections were
Potamolepidae, and the inability to produce resistant bodies critical-point dried, mounted and sputtered in the same way.
of Malawispongiidae, Lubomirskiidae and Metschnikowiidae Spicule measurements were performed on several hundreds of
could have constrained their dispersal, supporting their restricted megascleres, microscleres, gemmules and gemmuloscleres using
geographic range in ancient hydrographic basins (Fig. 1). The biol- light microscopy and digitizer.
ogy, the life history and the geographic distribution of these fresh-
water sponge families stressed the unsolved problem of their Terminology
taxonomy, systematics and phylogenesis based on the analysis of
classical diagnostic traits such as skeletal organisation and spicular General terminology of freshwater sponge diagnostic traits
morphology. Marshall (1883), Brien (1970a,b), Penney & follows the Thesaurus of sponge morphology (Boury-Esnault &
Racek (1968), Levi (1973), Bergquist (1978), Hartman (1982), Rtitzler, 1997). This Thesaurus, however, contains a misnomer
924 Porifera· Demospongiae • Haplosclerida • Spongillina

regarding the gemmular foramen, erroneously named 'micropyle' outer layer of compact spongin usually present at the gemmule sur-
(which is a pore in the egg membrane of insects, allowing sperm to face is variably developed and varies from armed with tangential
enter and fertilise the egg; Oxford Dictionary of Zoology, 1992). spicules more-or-less embedded, to those pierced by distal apices
Gemmule. A resistant body and asexual propagule produced of radial gemmuloscleres, or to those covering completely or in
by Spongillidae, Metaniidae and Potamolepidae, performing a part the distal gemmulosclere apices. The pneumatic layer, variably
structural and functional role that includes defence against environ- structured and thick, is made of spongin arranged to form cham-
mental injuries from mechanical and chemical stress, and favour- bers, trabecules or fibres. Pneumatic layers may appear to be spa-
ing the adhesion of these bodies to carriers/vectors. Gemmules tially displaced in a few genera (e.g., Saturnospongilla, Eunapius,
may be single, free and scattered in the skeletal network, or restricted Trochospongilla). The inner layer of compact spongin sublayered
to the sponge base in groups, or strictly adherent to the substratum, with a variable number of layers is a consistent trait shared by
singly or in carpets. gemmules of Spongillidae, Metaniidae and Potamolepidae.
Gemmular cage. Several freshwater sponge genera produce Gemmuloscleres. Specialised spicules, peculiar to armed
mono- or multi-gemmular cages of megascleres connected to the gemmules, absent from the naked gemmule of Spongilla lacustris
skeletal network, or to the basal plate. Cages are generally made by and Nudospongilla coggini. Gemmuloscleres encompass various
slightly modified skeletal spicules that assume a regular or irregu- morphotypes, ranging from oxeas to strongyles, birotules, pseudo-
lar spatial arrangement around gemmules. Pseudo-cages may be birotules, parmuliform, boletiform-tubelliform to botryoidal forms.
spatially displaced (e.g., Saturnospongilla, Drulia). The gemmulosclere surface bears ornamentations such as granules,
Gemmular theca. Gemmules contain a dense mass of tubercles, spines and microspines, variably associated and spatially
totipotent cells named thesocytes protected by a theca, generally arranged. (1) Birotule, shared by several genera, bears at the apices
bearing a foramen (an aperture closed by a thin membrane), to of the shaft well-developed plated or curved rotules, sometimes
favour the migration of cells during hatching. Foramen, absent or umbonate with shaft slightly projecting beyond the rotule at both
primitive, and shaped as an inconspicuous concavity in some gen- ends with blunt rounded processes. (2) Pseudo-birotule, shared by
era (e.g., Potamophloios), is usually single but gemmules bearing few genera (e.g., Acalle, Anheteromeyenia, Heteromeyenia), does
several foramina (up to 6 in Spongilla lacustris) are known; foram- not show a developed rotule but a group of curved hooks radiating
inal shape ranges from simple to tubular according to the thickness from the apices of the shaft. (3) Botryoidal gemmulosclere, exclu-
of the theca and it frequently bears a variably-shaped collar. The sive of the genus Pectispongilla, with a curved shaft bearing
theca, made of spongin and specialised spicules, displays a highly towards the convex sides of apices a botryum (group of concavities)
diversified architecture, ranging from naked gemmule to armed that during development appears as spines in rows joined to each
gemmule. Naked gemmules are characterised by a simple layer other by siliceous webs. (4) Parmuliform gemmulosclere, typical of
of laminated compact spongin without spicules (e.g., Spongilla, Drulia, with a single rotule supporting a short acute conical stem.
Nudospongilla). Armed gemmules range from a monolayered (5) Tubelliform-boletiform gemmulosclere, typical of Acalle and
theca of laminated compact spongin with spicules (e.g., Metania, is characterised by a proximal large, irregularly circular,
Potamophloios, Oncosclera), to a notably complex and variably flat rotule with the entire margin supporting a smooth shaft decreas-
arranged tri-layered theca bearing outer, pneumatic, and inner laying in thickness toward the distal apex, and shaped as umbonate
ers with spicules (e.g., most Spongillidae and Metaniidae). The pseudo-rotule with large hooks bearing microspines at their apices.

KEY TO FAMILIES

Keys to families and genera are primarily constructed for specimens that contain all specific characters, and hence care must be taken when
using these for taxa that are 'reduced', or in a non-reproductive 'resting stage'. For example, specimens collected during seasons when gemmules
are not produced might be identified as incertae sedis, or as one of the genera that do not produce gemmules. When gemmules are present their
associated spiculation is often important to the systematics of Spongillina. Among Potamolepidae the genera Oncosclera and Sterrastrolepis
show a trilayered theca, other characters are in agreement with the key; the Spongillidae Nudospongilla show a monolayered theca.
(1) Gemmules present ............................................................................................................................................................................... 2
Gemmules absent .... ... ....... .... ... ......... ................. ........... .......... ... ... ............... ................. .......... ................ .............. ............. .................. 4
(2) Gemmular theca monolayered. Gemmuloscleres, when present, are short irregular strongyles (tangentially arranged); gemmules
are produced in the basal portion (for the few species known to produce them). Skeletal network typically multi spicular alveolate-
reticulate with scanty spongin; megascleres are almost exclusively strongyles, rarely oxeas; microscleres rarely
present .... ...... .... ............ ... .... ... .... ....................... ........... .......... ...... ...................... .......... ...... .... .......................... ............ Potamolepidae
Gemmular theca bi-tri-layered (pneumatic layer generally present) ................................................................................................... 3
(3) Gemmuloscleres are oxeas, strongyles, birotules, pseudobirotules, botryoidal, club-like (radially and tangentially arranged); skeletal
network typically paucispicular irregularly reticulate; smooth or variably ornamented megascleres (oxeas and strongyles); microscle-
res usually present ............. .......... ...... .......... ... ............................. ............................ ..................... ...... .......... .................... Spongillidae
Gemmuloscleres are tubelliform, parmuliform, pseudobirotules (almost exclusively radially arranged); skeletal network typically
multispicular alveolate-reticulate with scanty spongin; smooth or variably ornamented megascleres (oxeas and strongyles);
microscleres usually present .... ... ...... ....... .... ........................ ................... ................ ........................... ................. .......... ...... Metaniidae
Gemmuloscleres are spiny oxeas and strongyles (radially arranged); skeletal network typically paucispicular irregularly reticulate
megascleres exclusively oxeas; microscleres (oxeas) are present; exclusively fossil ............................................ Palaeospongillidae
(4) Skeletal network typically multi spicular alveolate-reticulate, with sparse spongin; megascleres are spiny oxeas and strongyles;
microscleres absent (endemic to Lake Baikal) ........................................................................................................... Lubomirskiidae
Porifera' Demospongiae • Haplosclerida • Spongillina 925

Skeletal network typically paucispicular irregularly reticulate; megascleres exclusively spiny oxeas; microscleres absent (endemic to
Caspian Sea) ........................................................................................................................................................... Metschnikowiidae
Skeletal network typically multi spicular alveolate-reticulate, with sparse spongin; megascleres exclusively spiny or tubercular oxeas;
microscleres absent (endemic to ancient lakes) ..................................................................................... Malawispongiidae fam. nov.
Skeletal network variously structured; megascleres are smooth, spiny or tuberculate oxeas; microscleres absent (endemic to ancient
lakes ............................................................................................................................................................. Spongillina incertae sedis

FAMILY SPONGILLIDAE GRAY, 1867 Foramen present except in Nudospongilla. Gemrnular theca mono-,
bi- or tri-Iayered. Gemrnules, generally anned with gemrnuloscleres,
Synonymy in some cases naked. Gemrnuloscleres, when present, radial or tan-
gential to the surface, partially or totally embedded in the theca.
Spongillidae Gray, 1867a: 550. Spongillidae Carter, 1881e: 77. Gemrnulosclere shape is often species-specific with a large morpho-
Meyeniinae Vejdovsky, 1887-1888: 177. Type genus. Spongilla logical variation from spiny to granulated or smooth, from oxeas to
Lamarck,1816. strongyles, birotules, pseudobirotules, club-like, botryoidal. Larvae,
where described, are always parenchymella.
Diagnosis
Scope
Spongillina with shape ranging from globular to massive,
incrusting, lobate, arborescent. Colour in life from whitish to brown Spongillidae contains 21 valid genera: Anheteromeyenia,
or green, from pale to very dark. Surface smooth, hispid, conulose. Corvoheteromeyenia, Corvospongilla, Dosilia, Duosclera,
Consistency ranging from fragile to notably hard. Ectosomal skeleton Ephydatia, Eunapius, Heteromeyenia, Heterorotula, Nudospongilla,
from spicular brushes at apices of primary fibres to dense tangential Pachyrotula, Pectispongilla, Racekiela, Radiospongilla, Sanidastra,
spicular network. Choanosomal skeleton reticulate with regular or Saturnospongilla, Spongilla, Stratospongilla, Trochospongilla,
irregular meshes. Spongin variable but generally scanty. Megascleres Umborotula, Uruguayella.
are oxeas and strongyles, from smooth to spiny or granulated.
Microscleres when present oxeas, strongyles, aster-like, pseudobiro- Distribution
tules. Gemrnules always present except in very rare perennial speci-
mens. Gemrnular cage of megascleres is present in several species. Cosmopolitan (Fig. 1).

KEY TO GENERA

(1) Birotules and pseudobirotules present ................................................................................................................................................. 5

Birotules and pseudobirotules absent .................................................................................................................................................. 2
(2) Complete absence of microscleres and gemmuloscleres ..................................................................................................................... 3
Presence of microscleres andlor gemrnuloscleres (not pseudobirotules andlor birotules) .................................................................. 4
(3) Nude gemmule lacking pneumatic layer and gemrnuloscleres; microscleres never observed; megascleres are
acanthoxeas ....... ................................................. .......... ................. ........... .......................... ................................. ........... Nudospongilla
Gemrnular theca tri-Iayered, without gemrnuloscleres, covered by a cage of megascleres; microscleres absent; megascleres are spiny
oxeas of two size classes ....................................................................................................................................................... Duosclera
(4) Microsc1eres absent; megascleres are spiny oxeas; gemmuloscleres showing 2-9 spiny axes with rounded or tricuspidate tips
embedded radially in the gemrnular theca ........................................................................................................................... Sanidastra
Microsc1eres absent; megasc1eres are spiny oxeas; gemrnulosc1eres are spiny strongyles radially oriented ............... Radiospongilla
Microsc1eres absent; megascleres are spiny and smooth oxeas; gemmuloscleres are spiny oxeas of two morphologies: strongyles
stout, short and spined, tangentially embedded in the outer layer of gemmular theca; spiny oxeas abruptly pointed, radially arranged
in between the inner and outer layer ............................................................................................................................. Stratospongilla
Mieroscleres absent; megascleres are smooth oxeas; gemmuloscleres are oxeas and strongyles, smooth or with spiny tips, tangentially
arranged on the gemrnular surface ......................................................................................................................................... Eunapius
Microscleres are spiny oxeas; megascleres are smooth oxeas; gemrnuloscleres, when present, are spiny oxeas or strongyles often very
curved ........... ........................................... ...... ......... .... ... ..... .......... ........... ...... .......... ................. ............... ........... ........... ......... Spongilla
Microscleres are spiny oxeas; megascleres are spiny oxeas; gemrnuloscleres show a typical botryoides shape of the two
apices ............................ ...... .......... ............ .... ...... ..................... ........... .................................... ......................................... Pectispongilla
(5) Birotules with rotules showing smooth edges, pseudobirotules absent ............................................................................................... 6
Birotules with rotules showing spiny and indented edges generally associated with pseudobirotules ............................................... 7
(6) Microscleres absent; megascleres are oxeas very spiny; gemmuloscleres are smooth birotules radially arranged .. , Trochospongilla
Microscleres absent; megascleres are microspined short strongyles; gemrnuloscleres are smooth birotules radially
arranged .............................................................................................................................................................................. Uruguayella
Microscleres absent; megascleres are smooth oxeas; gemrnuloscleres are smooth birotules radially arranged in the gemmular
theca, and spiny oxeas tangentially arranged in the typical thoroidal pneumatic layer (endemic to the River Jurua,
Brazil) ......................................................................................................................................................................... Saturnospongilla
926 Porifera· Demospongiae • Haplosclerida • Spongillina

(7) Birotules and/or pseudobirotules are present, exclusively as microscleres ....................................................................................... 8

Birotules and/or pseudobirotules are present, exclusively as gemmuloscleres ................................................................................. 9
Birotules are gemmuloscleres, pseudobirotules are microscleres ................................................................................................... 15
(8) Gemmuloscleres are spiny and/or strongyles tangentially arranged on the gemmular theca; megascleres are smoooth microgranulated;
microscleres are pseudobirotules with a smooth shaft .... ........... ................. ................ .......................... ................ ..... Corvospongilla
(9) Gemmuloscleres birotules and/or pseudobirotules of a single morphology ................................................................................... 11
Gemmuloscleres birotules and/or pseudobirotules belonging to two morphological forms (birotules + pseudobirotules, or
birotules + acanthoxeas) ....... .......... ........... ...... .......... ... ... ............... ................. .......... ................ ............ .... ...................... ... ... ......... 10
(10) Gemmuloscleres are birotules (long shaft) and pseudobirotules (long shaft) radially arranged; microscleres are acanthoxeas;
megascleres are acanthoxeas ...................................................................................................................................... Heteromeyenia
Gemmuloscleres are birotules (short shaft) and pseudobirotules (long and spiny shaft) radially arranged; microscleres are absent;
megascleres are spiny oxeas .... .............................. ...... .... ........... ........... ...... .......... ........... ..................... ................ .............. Racekiela
Gemmuloscleres are birotules (variable rotules) and acanthoxeas tangentially arranged; microscleres are absent; megascleres are
spiny and smooth oxeas (endemic to New Caledonia) .................................................................................................... Pachyrotula
(11) Gemmuloscleres are birotules ......................................................................................................................................................... 12
Gemmuloscleres are pseudobirotules (long and spiny shaft); microscleres are absent; megascleres are spiny
oxeas ..... ........... .......... .... ... .... ...... ...... .... ...................... ...................... .......... ...... .......... ................. ............... ............ Anheteromeyenia
(12) Birotules with a short (usually less than the rotule diameter) shaft ................................................................................................ 13
Birotules with a long spiny shaft (2-3 times the rotule diameter) .................................................................................................. 14
(13) Gemmuloscleres are birotules with a short shaft and rotules of identical diameter; microscleres are absent; megascleres are
microspined and/or smooth oxeas ...................................................................................................................................... Ephydatia
Gemmuloscleres are birotules with a short shaft and rotules of different diameter; microscleres are absent; megascleres are
microspined and/or smooth oxeas .................................................................................................................................. Heterorotula
(14) Gemmuloscleres are birotules with a long spiny shaft; microscleres are absent; megascleres are microspined and rarely smooth
oxeas ...... .......................... ................ ................................. .... ........... ................ ................. ................. .... ...... .......... ...... .... Umborotula
Gemmuloscleres are birotules with a long spiny shaft; microscleres range from acanthoxeas to typical 'euasters' with spiny tips;
megascleres are smooth oxeas .... .......... ........... ...... ..................... .......... ........................... ............. .... .......... ............................. Dosilia
(15) Gemmuloscleres are birotules with a long spiny shaft; microscleres are pseudobirotules showing a long smooth or spiny shaft;
megascleres are smooth oxeas .... ........... .......... ................ ...................... ................ ........................... ........... ...... Corvoheteromeyenia

SPONGILLA LAMARCK, 1816 assigned to other genera since. At present, at least seven other valid
species of Spongilla s.s. have been described: S. alba Carter, 1849
Synonymy (India, SE Asia, Africa, Madagascar, Australia, South America,
Turkey); S. aspinosa Potts, 1880b (E Canada and USA); S. inar-
Spongilla Lamarck, 1816: 98. Euspongilla Vejdovsky, 1883: mata Annandale, 1918 (Japan); S. cenota Penney & Racek, 1968
15. Vejdovsky in Potts, 1888: 172. (Yucatan); S. wagneri Potts, 1889 (SE USA, Florida, S Carolina);
S. shikaribensis Sasaki. 1934 (Japan); S. stankovici Arndt, 1938
Type species (Ochrid Lake, Macedonia).

Spongia lacustris Linnaeus, 1759: 1348 (by subsequent Description of type species
designation; Annandale, 1911b).
Spongilla lacustris (Linnaeus, 1759) (Figs 2-8).
Definition Material examined. Holotype: LSLH 1295.1, LSLH
1295.2 -lake of Smruand, det. Linnaeus. Other material. 'Neotype'
Spongillidae with encrusting, massive, branched to arbores- (now invalid): RMNH 1053 - Vlaardingse Vaart, The Netherlands,
cent body shape. Consistency fragile, soft. Ectosomal skeleton ix.1941. Specimens: MNHN DT2828 - Viene, Limoges. DTRG-FW
consists of spicular brushes. Choanosomal skeleton irregular, from 558 - Sweden, from ZRS. DTRG-FW 10 - Lago d'Orta, ix.1888.
isotropic to anisotropic, pauci- to multi spicular. Abundant spongin DTRG-FW 33 - Coghinas River, Terme di Casteldoria, Sardinia,
sheaths. Megascleres smooth to slightly spined oxeas. Microscleres 31.v.1985. DTRG-FW 68 - Lake Maggiore, Isole Borromee, 1889.
abundant to rare spiny oxeas. Gemmules in dense clusters or scat- DTRG-FW 73 - Scrivia Stream, Ponte della Ferriera, Ronco Scrivia,
tered. Gemmular cage composed of megascleres sometimes present. 4.ix.1985. DTRG-FW431-FW543 -GinevraLake. DTRG-FWI23 -
Gemmules of two types: naked without gemmuloscleres, and Bolle di Magadino, Canton Ticino, 22.iii.1970.
armoured with gemmuloscleres. Gemmular theca from monolayered Description. Body shape encrusting, massive, branched to
to tri-Iayered. Foramen or foramina present. Gemmuloscleres when arborescent. Colour whitish, yellow-orange, greenish, emerald
present spined oxeas or strongyles. green. Consistency fragile, soft. Surface uneven hispid. Oscules not
conspicuous, scattered. Dermal membrane aspicular. Ectosomal
Scope and distribution skeleton occurs as spicular brushes at apices of primary fibres.
Choanosomal skeleton irregular from isotropic in encrusting por-
In addition to the type species, more than 150 nominal species tions, to anisotropic in finger-like projections or branches, pauci- to
have been ascribed to this genus, but most of these have been multispicular primary fibres connected by paucispicular transverse
Porifera' Demospongiae • Haplosclerida • Spongillina 927

/2Jf5". I

Scm

Fig. 2. SpOllgilla laclIslris. syntype (LS LH ). A. paralcctotype. 1295.1. B. lectotype. 1295.2.

fibres. Abundant spongin sheaths. Megascleres (90--350 X 2-18 11m)

smooth fusifonn oxeas with tips from gently to sharply pointed,
slightly spined if associated to gemmules. Microscleres abundant to
rare scattered fusifonn oxeas (25-178 X 2-8 11m) with dense spines
regularly distributed along the entire length and microspinosity on
spines giving them an asterose shape. Gemmules (98-789 11m) from
subspherical to oval in dense clusters, or irregularly scattered in the
skeletal network. Gemmular cage of megascleres sometimes present
in naked gemmules. Gemmules of two types can be found in the
same specimen, naked without gemmuloscleres and armoured with
gemmuloscleres, tangential to partially embedded in the gemmular
theca. Gemmular theca ranging from thick-walled with tri-layers to
thin-walled with a monolayer of compact spongin, with a variable
number of sublayers (3-7), corresponding to the inner layer of the
thick walled gemmule. Outer layer well-developed in some thick-
walled gemmules. Foramen slightly elevated without or with a sim-
ple collar or plate-like collar, generally single in thick-walled
gemmules; multiple foramina (1-6) common in thin-walled gem-
mules, but also present sometimes in thick-walled gemmules.
Gemmuloscleres (21-130 X 1-10 11m), when present, slightly to
strongly bent oxeas or strongyles usually with large spines.
Ecology. Standing and running waters, from pennafrost area
in the Northern Polar Circle, to semi-arid zones in the Mediterranean
islands, from brackish waters to alpine lakes or ancient lakes (Lake
Baikal). Life cycle hibernating. Distribution: Palaearctic.
Remarks. Spongia lacustris was described by Linnaeus
(1759: 1348) as "repens, fragilis, ramis teretibus obtusis" [creep-
ing, fragile, with cylindrical branches showing swellings at their
ends], in the second volume of the Systema Naturae, and its type
material was previously unknown. "Since a type specimen appar-
ently was never designated, and the original collection most cer-
tainly does not exist today", Penney & Racek (1968) subsequently
designated a neotype (RMNH 1053), which they further state Fig. 3. Spongilla lacustris growth forms during an annual life cycle.
" ... correspond[s] in all details with the description of Spongilla A, encrusting. B, arborescent.
928 Porifera· Demospongiae • HapJosclerida • Spongillina

Fig. 4. Spongilla lacustris, skeleton (lectotype, LSLH 1295.2). A, ectosomal skeleton. B, a multi spicular skeletal tract.

o ~
---.. ------ -. -

Fig. 5. Spongi/la lacustris, spicule characteristics (lectotype LSLH Fig. 6. Spong ilia lacustris, gemmules, spicules and skeletal architecture
1295.2). Both Linnean specimens lack gemmules. A, megascleres smooth (previous 'neotype' RMNH 1035). A, naked gemmule. B, foramen.
oxeas. B-C, close-up of megasclere tips. D-E, close-up of megasclere C, gemmular theca with outer, pneumatic and inner layers. D, gemmular
shaft. F-H, microscleres. I, close-up of microsclere shaft. J, close-up of theca without pneumatic layer. E-F, different view of the skeletal network.
microsclere tip. (After Manconi & Pronzato, 2000.) G, ectosome. H, microscleres irregularly arranged in the sponge mesohyl.
Porifera' Demospongiae • Haplosclerida • Spongillina 929

lacustris Auct. in previous literature", from the collection of the

Rijksmuseum van Natuuriijke Historie of Leiden. However, the
rediscovery of Linnaeus's original material (Manconi & Pronzato,
2000) invalidates the neotype designation, although the latter spec-
imen seems to be the best 'representative' material of the species,
being abundantly full of gemmules and easily accessible for scien-
tific study, as recently shown by Manconi & Desqueyroux-Faundez
(1999).

ANHETEROMEYENIA SCHRODER, 1927

Synonymy

Heteromeyenia (Anheteromeyenia) SchrOder, 1927: 108.

Anheteromeyenia de Laubenfels, 1936a: 36.

Type species

Spongilla argyrosperma Potts, 1880b: 357 (by subsequent

designation; Penney & Racek, 1968).

Definition

Spongillidae with encrusting growth form. Ectosomal skeleton

spicular. Choanosomal skeleton irregular network of megascleres
with paucispicular tracts. Sparse spongin. Megascleres acanthoxeas.
Microscleres absent. Gemmular theca tri-layered. Gemmuloscleres
pseudobirotules radially embedded in the theca.

Scope and distribution

Fig. 7. Spongilla lacustris , gemmules and spicules, A-C, G-I, DTRG-FW73 Three other species: A. pictovensis (Potts, 1885a) (from
(ex-IZUG); D-F, DTRG-FWI23 (ex-IZUG). Specimens belonging to
southern populations (Italy) with anned gemmules. A, megascieres.
E-Canada to New York); A. biceps (Lindenschmidt, 1950)
B-C, microscieres. D, armed gemmule. E, foramen. F, trilayered pneumatic (Michigan); A. ornata (Bonetto & Ezcurra de Drago, 1970)
layer showing a thick pneumatic layer. G-I, gemmuloscieres. (S America). Neartic and Neotropical Regions.

Fig. 8. Geographic di slribution of the genus Spollgi/la .

930 Porifera· Demospongiae • Haplosclerida • Spongillina

Description of type species irregular chambers. Inner layer multilayered. Gemmuloscleres

radially embedded in the theca, emerging from the outer layer
Anheteromeyenia argyrosperma (Potts, 1880b) (Figs 9-12). according to their different levels of insertion. Gemmuloscleres
Material examined. Lectotype: ANSP P04538. ParaIectotype. pseudobirotules with a notably variable length (65-160f.Lm) with
ANSP P04587, box 31-A, Leigh River at Leigh Gap, E. Potts leg. umbonate pseudo-rotules formed by 2-4 hooks, and shafts armed
14.xi.l881. Other material. ROM Tl990-015, Lake Temagami, by large spines or more rarely by hooks.
Ko-Ko Bay, 20.viii.l937, leg. J. Oughton, det. N. Gist Gee. Ecology. Both lentic and lotic habitat in waters of about
Description. Sponge is thin crust with inconspicuous neutral pH, low-moderate alkalinity, high conductivity, 9-23°C.
oscu1es. Surface hispid due to emerging spicules. Colour grey or Distribution: Eastern Nearctic region, from Canada (Quebec,
green if in symbiosis with zoochlorellae. Dermal membrane spicu- New Brunswick) to Florida.
lar. Subdermal cavities not observed. Ectosomal skeleton is a spic- Remarks. Penney & Racek (1968) and Volkmer-Ribeiro
ular dermal membrane. Choanosomal skeleton irregular network (1996) report two classes of gemmuloscleres, however Ricciardi &
of megascleres with paucispicular tracts. Sparse spongin. Reiswig (1993) and our observations suggested that the insertion of
Megascleres slightly curved acanthoxeas (250--329 X 10--15 f.Lm) birotules occur at different levels of the gemmular theca.
with sparse small spines except at the tips. Smaller megascleres
(133 X 7 f.Lm) reported by Volkmer (1996) could be those from the
dermal membrane. Microscleres absent. Gemmules subspherical
(400--700 f.Lm). Foramen is a short simple tube without collar. CORVOHETEROMEYENIA EZCURRA DE DRAGO, 1979
Gemmular theca tri-layered. Outer layer thin. Pneumatic layer with
Synonymy

, ~ Corvomeyenia (part) Bonetto & Ezcurra de Drago, 1966: 129.

. '!J
~,

" Corvoheteromeyenia Ezcurra de Drago, 1979: 110.

:./.V,
;;I'"
"b~
~.

~
1 • ~
..
~

~
Type species

Corvomeyenia australis Bonetto & Ezcurra de Drago, 1966:

130 (by monotypy).
.~

;, " "I
::::.>
£'p"
~
--S Diagnosis

S
\,.~
>'
'- \0
"
Spongillidae with encrusting growth form. Choanosomal
"
\> skeleton irregularly alveolate. Megascleres curved oxeas generally
smooth. Microscleres pseudobirotules. Gemmules adherent to the
2cm ~ ~ substratum or free and scattered. Foramen present. Gemmular

I
~ ""
~'
theca tri-layered with gemmuloscleres radially embedded.
r
~. ~
~ , Gemmuloscleres birotules.
~

0
~ .
p
r-. Scope and distribution
Fig. 9. Anheteromeyenia argyrosperma. A. lectotype ANSP P04538.
B, para\ectotype ANSP P04587. One other species: C. heterosclera. Distribution. Neotropical.

Fig. to. Anheteromeyenia argyrosperma, skeletal network (ROM N° B6, 1990-015). A, sponge surface. B, choanosomal skeleton.
Porifera' Demospongiae • Haplosclerida • Spongillina 931

Description of type species

Corvoheteromeyenia australis (Bonetto & Ezcurra de Drago,

1966) (Figs 13-14).
Material examined. None. Description modified after
Bonetto & Ezcurra de Drago (1966). Type material in MABA and
INALI not available. Type locality Laguna de Guadalupe (Setubal),
at the entrance in the Riacho Santa Fe, Santa Fe, Argentina.
Description. Body shape encrusting. Colour brown-greenish.
Hispid surface due to emerging spicules. Oscules inconspicuous.
Dermal membrane spicular. Subdermal cavities not observed.
Ectosomal skeleton not observed. Choanosomal skeleton is irregu-
larly alveolate (?) with large meshes. Megascleres curved oxeas
(300-350 X 13-20/-Lm) generally smooth, sometimes irregularly
microspined. Microscleres pseudobirotules (reported as "isoquelas")
very variable in shape and shaft size (20-35/-Lm up to 90/-Lm) with
slightly curved shaft with pseudo-rotules ranging from small with
three curved hooks (2-8 /-Lm) to large without hooks. Gemmules
(400-500/-Lm) flattened yellowish in dry conditions at the basal por-
tion adherent to the substratum or free and scattered in the sponge
body. Foramen single and circular. Gemmular theca tri-Iayered. Outer
layer with emerging shafts and distal rotules. Pneumatic layer well
developed with gemmuloscleres radially embedded. Inner layer well
developed. Gemmuloscleres birotules (43-90 X 17-23/-Lm) with
spined shafts and circular slightly convex rotules with margins from
crenulated to indented with teeth long and curved.
Ecology. The holotype was found on a stone wall in the
laguna silt bottom in running waters (60 em/sec). The popUlation of
the entire lagoon was notably dense. Distribution. Upper and mid-
dle Parana River, Rio Iguazu, River Uruguay, Guadalupe Lagoon
(Argentina).

CORVOSPONGILLA ANNANDALE, 1911

Fig. 11. Anheteromeyenia argyrosperma, spicules and gemmules, lecto-
type ANSP P04538. A, gemmule. B, gemmular surface. C, foramen. D,
cross section of gemmular theca, pneumatic layer with irregular chambers.
Synonymy
E, multilayered inner layer. F-G, Ha, gemmulosc1eres. Hb, I, spiny oxeas
(megasc1eres). Corvospongilla Annandale, 1911b: 122.

Fig. 12. Geographic distribution of the genus Allileleromeyellia.

932 Porifera' Demospongiae • Haplosclerida • Spongillina

Type species

Spongilla loricata Weltner, 1895: 138 (by subsequent

designation; Annandale, 1911b).

Diagnosis

Spongillidae with encrusting flat to massive, lobate growth

form. Consistency extremely hard to delicate. Megascleres vary from
smooth to granulated strongyles or rarely oxeas. Microscleres often
rare, straight to slightly curved micropseudobirotules with smooth or
spined shaft. Gemmules of two types single or grouped, free or fixed
at the sponge base, with or without a spicular cage of slender
strongyles. Foramen ranging from apical to lateral with a short porus
tube. Gemmular theca from tri-Iayered with a variably developed
pneumatic layer to monolayered of compact spongin without pneu-
matic layer. Gemmuloscleres from elongated slightly curved to oval
stout spined strongyles tangentially arranged in the theca.

Scope and distribution

Eleven other species known: C. burmanica (Kirkpatrick,

1908b) (Burma, India); C. bohmii (Hilgendorf, 1883) (Africa equa-
torial); C. caunteri Annandale, 1911b (India); C. lapidosa
(Annandale, 1908a) (India); C. micramphidiscoides Weltner, 1913
(Congo River); C. novaeterrae (Potts, 1886); C. scabrispiculis
Annandale, 1913a (Tropical Africa, River Nilo); C. ultima
Annandale, 191Oa) (S India); C. zambesiana (Kirkpatrick, 1906)
(River Zambesi); C. sekti Bonetto & Ezcurra de Drago, 1966
(Parana, Paraguay and Uruguay Rivers, Argentina); C. volkmeri
(De Rosa-Barbosa, 1988) (Brasil). Distribution. Tropical regions of
Africa and Asia (from Burma to Iraq) and S America.

Fig. 13. Corvoheteromeyenia, spicules from original papers. A-G,

C. australis, after Bonetto & Ezcurra de Drago, 1966. H-K, C. heterosclera,
Description of type species
after Ezcurra de Drago, 1974. A, E-J, gemmuloscleres. B-C, K, microscle-
res. D , megascleres. Corvospongilla loricata (Weitner, 1895) (Figs 15-18).

Fig. 14. Geographic distribution of the genus Con'oheleromeyenia.

Porifera· Demospongiae • Haplosclerida • Spongillina 933

Material examined. Holotype: 2MB2093 - dry gemmules finely granulated to rarely smooth belong to gemmular cages.
on Aetheria shell, Africa. Other material. BMNH 1882.3.22. Microscleres rare or variably abundant scattered in the
1-3 - C. burmanica, Burma; Corvospongilla sp., River Diyala of choanosome, pseudo-microbirotules (20--40 fLm length) with
R. Tigri Basin, Himmereen Reservoir, Iraq, leg. H.A.A. Saadalla, a smooth shaft bearing pseudo-rotules (4-12 fLm diam.) with few
1997. radial recurved hooks. Gemmules (500-950 fLm) oftwo types, free
Description. Encrusting flat sponge grossly hispid. and fixed at the sponge base, with or without a spicular cage
Consistency hard. Colour light brown when dry. Surface conulose of slender strongyles. Foramen from apical to lateral, with a
due to underlying tufts of megascleres in the convergence of fibres short tube. Gemmular theca from monolayered with compact spon-
toward the surface. Dermal membrane not observed. Ectosomal gin without pneumatic layer, to tri-Iayered with a variably devel-
skeleton dense spicular network. Choanosomal skeleton alveolate oped pneumatic layer. Gemmuloscleres slightly curved stout
from paucispicular to multi spicular tracts, more irregular at the spined strongyles (34-75 X 8-14 fLm) tangentially embedded in
sponge base. Basal plate well developed. Megascleres feebly the theca.
curved smooth stout strongyles (220-226 X 20 fLm) with inflated Ecology. Undefined, in Africa, growing on shells of
tips in the skeletal network; slender strongyles (125-130 X 7 fLm) Aetheria sp.
Remarks. In spite of Penney & Racek (1968) underlining
the existence within the genus of two groups characterised by
oxeas or strongyles as megascleres, we found a gradient from
oxeas to strongyles in the same specimen of Corvospongilla sp.
from Iraq. As stressed by Brien (1973b) with the proposed subfam-
ily Corvospongillinae, the entire set of diagnostic traits presently
studied highlighted a problematic position of this genus between
the families Spongillidae and Potamolepidae.

DOSIUA GRAY, 1867

Synonymy

Dosilia Gray, 1867a: 550. Asteromeyenia Annandale,

Fig. 15. Corvospongilla lorieata, holotype 2MB 2093, is a carpet of 1911a: 593. [Astromeyenial SchrOder, 1927: 102 (lapsus).
gemmules (circle) on a shell of Aetheria. Astroheteromeyenia SchrOder, 1927: 102.

Fig. 16. Corvospongilla, skeletal network, fragment of the holotype of C. burmaniea. BMNH 1882.3.22.1-3 (box 6, dry). A, sponge surface (top view).
B, cross section (top = surface with conules; bottom = basal spongin plate). C, lateral view of a conule with peripheral skeleton. D, choanosome.
E, choanosomal skeleton at the sponge base (cross section).
934 Porifera· Demospongiae • Haplosclerida • Spongillina

Type species

Spongilla plumosa Carter, 1849: 81 (by subsequent designa-

tion; de Laubenfels, 1936a).

Diagnosis

Spongillidae with massive lobate growth form. Choanosomal

skeleton plumose-reticulated anisotropic. Megascleres smooth
oxeas. Microscleres range from acanthoxeas with rays to true
'euasters'. Gemmules scattered. Gemmular theca tri-Iayered with
radially embedded birotules.

Scope and distribution

Three other species: D. palmeri (Potts, 1885b) (Central

America); D. brouni (Kirkpatrick, 1906) (River Nilo E Africa);
D. radiospiculata (Mills, 1888) (USA, Mexico). Tropical and sub-
tropical areas of S America and Africa.

Description of type species

Dosilia plumosa (Carter, 1849) (Figs 19-22).

Material examined. Holotype: Not examined. Other material.
BMNH 1908.2.8.1 - dry, El Obeid, Khartoum, 29.i.1908,
Kirkpatrick. BMNH (DTRG-FW421) - Bowerbank collection, dry
box 6, 3 specimens.
DescriptWn. Sponge shape massive lobate. Consistency
moderately soft and very fragile. Colour from greenish to pale
brown-yellowish. Surface lobate with hispidation due to spicule
tufts at apices of primary fibres. Conspicuous scattered small
oscula. Dermal membrane aspiculous. Subdermal cavities not
observed. Ectosomal skeleton spicule tufts. Choanosomal skeleton
plumose-reticulated, skeleton anisotropic with radial multi spicular
Fig. 17. Corvospongilla, spicular and gemmular traits; A-G, C. lorieata, primary fibres gradually diverging toward the surface and paucispi-
type specimen, 2MB 2093. H, C. burmaniea, BMNH 1882.3.22.1-3. cular secondary tracts. Variable amount of spongin. Microscleres
I, Corvospongilla sp., DTRG-FW574 (ex-IZUG). A, gemmular cage
including two fixed gemmules. B, foramen. C, gemmular surface. D, cross scattered and abundant in the choanosome. Megascleres stout,
section of gemmular theca. E, gemmuloscleres. F-G, megascleres. ranging from straight to slightly curved fusiform smooth oxeas
H, microscleres. I, microscleres in the skeletal network. (400--520 X 15-21 f.Lm). Microscleres range from acanthoxeas,

Fig. 18. Geographic distribution of the genu Con·ospongilla.

Porifera· Demospongiae • Haplosclerida • Spongillina 935

to acanthoxeas with few long central rays, to 'euasters' with 8-12 spines. Rotules from flat to slightly umbonate with margins bearing
spiny rays projecting from a distinct globular centre. Rays gener- numerous small blunt and recurved teeth (spines) (55-85 X 3-4 f,Lm,
ally smooth with recurved spines at their apex. Size highly variable diameter of rotules 23-25 f,Lm).
with rays not exceeding 15-18 f,Lm. Gemmules abundant, scattered Ecology. Unknown. Distribution. India, Philippines.
within the skeletal network, subspherical to ovoid (500-680 f,Lm). Remarks. Asteromeyenia Annandale, 1911a: 593 (for type
Foramen lateral and tube-like in shape. Gemmular theca tri- species Heteromeyenia radiospiculata Mills, 1888 (by original
layered. Outer layer variably developed with emerging free distal designation), now Dosilia radiospiculata, is thus a junior synonym
rotules. Pneumatic layer well developed of minute regular cham- of Dosilia according to Penney & Racek (1968).
bers with radially embedded birotules. Inner layer with 2-
3 sub-layers of compact spongin in contact with proximal rotules.
Gemmuloscleres birotules, straight cylindrical shaft with large
DUOSCLERA REISWIG & RICCIARDI, 1993

Synonymy

Duosclera Reiswig & Ricciardi, 1993: 275.

Type species

Spongilla mackayi Carter, 1885: 19 (by original designation).

Diagnosis

Spongillidae with encrusting growth form. Choanosomal

skeleton moderately developed with fibres of dense spongin. Two
classes of megascleres, one in the skeletal framework and the other
Fig. 19. Dosilia plumosa, specimens of Bowerbank's collection BMNH in the mesohyl, but both also functioning as gemmule spicules.
unreg. (box 6, dry). True microscleres absent. Gemmules typically in hemispherical

Fig. 20. Dosilia plumosa, skeletal network, specimens of Bowerbank's collection, BMNH (box 6, dry). A, a sponge fragment (surface on the right).
B, peripheral skeleton (cross section). C, choanosomal spicular arrangement. D, microscleres in the choanosomal matrix.
936 Porifera· Demospongiae • HaploscIerida • Spongillina

Fig. 21. Dosilia piumosa, spicules and gemmules, Bowerbank's collection, BMNH (box 6, dry). A-B, megascleres smooth oxeas, microscleres and gem-
muloscleres. Ca, megasclere tip. Cb, D, F,gemmuloscleres. Ce, E, G, microscleres. H, foramen. I, surface of the gemmule. J, gemmule. K, trilayered
gemmular theca.

clusters but occasionally occurring as a restricted pavement. Material examined. Holotype: BMNH 1890.1.9.279-4 -
Special gemrnuloscleres absent. Mackay's lake near Pictou, Nova Scotia, 26.viii.1884.
Description. Encrusting (2.5 X 4 X 1 cm). Dermal membrane
Scope and distribution aspicular. Subdermal cavities not observed. Ectosomal skeleton
undifferentiated. Choanosomal skeleton with fibre system moder-
Monotypic, Nearctic (Eastern North America). ately developed with fibres of dense spongin with 3-10 procurved
spicules. Megascleres of two classes: straight slightly curved acan-
Description of type species thoxeas with acute spines directed distally (procurved) in the main
choanosomal skeleton (251 X 1O.2/-Lm); and abundant recurved
Duosclera mackayi (Carter, 1885) (Figs 23-26). shorter acanthoxeas with long robust spines directed proximally
Porifera· Demospongiae • Haplosclerida • Spongillina 937

Fig. 22. Geographic di. tribution of the genus Dosilia.

Fig. 23. Duosclera mackay;, holotype BMNH 1890.1.9.279, with original

labels.

(recurved) scattered in the choanosome, and also functioning Fig. 24. Duosclera mackayi, skeletal network, BMNH 1890.1.9.279.
A, cross section of the dry specimen where the poorly evident sponge skele-
as gemmule spicules (167 X 10,2 f.Lm), Microscleres absent. ton is mixed with vegetal fibres. B, detail of A (rectangle) showing a broken
Gemmules typically in hemispherical clusters but occasionally gemmule enclosed in the choanosomal skeleton. C-D, sponge surface.
occurring as a restricted pavement. Gemmular cage of megascleres
radially oriented within and projecting from the pneumatic layer.
Foramina, simple with collar, directed toward the substrate and Quebec, Massachusetts, New York, Wisconsin, Michigan, New
opening into a restricted cavity, Gemmular theca tri-Iayered, Outer Jersey, Georgia, Florida, Louisiana.
layer conspicuous with polygonal surface, Pneumatic layer with
chambers, Inner layer of multilayered compact spongin, Special
gemmuloscleres absent. EPHYDATIA LAMOUROUX, 1816
Ecology. Restricted to dystrophic lentic waters with a pH
range of 4,7-6.2. Symbiotic with green algae, and commonly asso- Synonymy
ciated to Corvomeyenia everetti and Trochospongilla pennsylvan-
ica. In Quebec it was found active and green in November at 4°C, [Tupha] Oken, 1814 (unavailable). Ephydatia Lamouroux,
whereas T. pennsylvanica under the same conditions was found 1816: 2. Trachyspongilla Dybowsky, 1878: 53, Meyenia Carter,
only as carpets of gemmules (Ricciardi & Reiswig, 1993). 1881c: 90. Pleiomeyenia Mills, 1884: 147. ?Clypeatula Peterson &
Distribution. Newfoundland, Nova Scotia, New Brunswick, Addis, 2000: 269.
938 Porifera· Demospongiae • Haplosclerida • Spongillina

Fig. 25. Duosclera mackayi, spicules and gemmules, holotype BMNH 1890.1.9.279. A, gemmular cage including three gemmules. B, some gemmules
in toto. C, cross section of gemmule. D, foramen. E-F, theca of compact multilayered spongin. G-J, different views of size classes of megascleres.
K, spines of megascleres.

Type species micros pined. Microscleres absent. Gemmules scattered. Gemmular

theca tri-Iayered. Gemmuloscleres birotules.
Spongia fluviatilis Linnaeus, 1759: 1348 (by subsequent
designation; Annandale, 1911b). Scope and distribution

Diagnosis Eight other species: E. meyeni (Carter, 1849) (India, China);

E. muelleri (Lieberkuhn, 1855) (N Hemisphere); E. japonica
Spongillidae with encrusting, bulbous, massive growth form. (Hilgendorf, 1882) (USA?, Manchuria, Japan); E. ramsayi (Haswell,
Choanosomal skeleton anisotropic with paucispicular fibres and 1882) (Australia, New Zealand, New Guinea?); E. robusta (Potts,
tracts. Spongin scarce. Megascleres oxeas from smooth to 1888) (E USA, Mexico, California); E. facunda Wehner, 1895
Porifera· Demospongiae • Haplosclerida • Spongillina 939

Fig. 26. Geographic di Lribution of the genu 0110 c/era.

2cm

NATIOHAA.I. NAnJURJ<lSlOAlSCIl MUSEUM. LEUH

ff"jda.t(.Q. rL . . v,a.l:i{' .. (L )
Neotjpe . chaStI'! by A./J .Ra..cek
lOCQ.Llti t\l\gste{ R Abco...cle
t{oLl ...... cl. 1'2. Sepb!~be.r (Cf~2
Ltg A.W la..colAr l
( trQ.S ........ t "f ("0(0":.1)

Fig. 27. Ephydatiafiuviatilis neotype RMNH 1166.

(central and S America); E. fortis Weltner, 1895 (Indonesia,

Philippines, Japan, Vanuatu); E. millsii (Potts, 1888) (Florida).

Description of type species

Ephydatiafluviatilis (Linnaeus, 1759) (Figs 27-31).

Materio.l examined. Neotype: RMNH 1166 (designated by
A.A. Racek, cf. Penney & Racek, 1968: 84) - A.w. Lacourt,
Angstel River, Abcoude Holland, 12.ix.1942. Other material.
DTRG FW31 - Trasimeno Lake 17.iv.1973). DTRG FW33 -
Coghinas River, Terme di Castel doria, 31.v.1985. DTRG FW63
(from MFSN) - Fosse del Patriarca, Roggia di Udine (1868-70).
DTRG FW73 - Scrivia Stream, Ponte della Ferriera, Ronco
Scrivia (4.9.1985). DTRG FW82 (from MCSN) -Maggiore Lake,
Pallanza Basin 29.iii.l970. DTRG FW133 - Tagliavia SS 118
Sicily, 22.iv.l986. DTRG FW257 - Crete Island, between Fig. 28. Ephydatia fiuviatilis , phases of the life cycle. A, carpet of gem-
Paraskie and Kastelli, dried river, 30.x.l988. DTRG FW263 - mules on a stone. B, a postlarval stage after settlement. C, a mature colony.
Apulia, Lake Capaciotti, dried 5.xi.1988. DTRG FW295 -
Mallorca Island 27.iv.1990. DTRG FW301 - Cernika Lake, Description. Body shapes range from crusts to bulbous or
Bosnia, 27.vii.l990. MNHN DT2820 - Syria, Barara, 1908. MNHN massive, with ridges or rounded lobes. Consistency firm but fragile
DT2834 - Algeria, Seurat-Gautier, 1922. MNHN DT2822 - Yerre, in life, extremely brittle if dry. Colour from whitish to green or
Loire River, 2.iv.1896. brown. Surface hispid due to emerging spicules. Oscules numerous.
940 Porifera· Demospongiae • HaploscIerida • Spongillina

Fig. 29. Ephydatia jluviatiiis, skeletal network, neotype; RMNH 1166. A-B, sponge surface. C, cross section of a sponge fragment (left = surface).
D, superficial his pi dation in a cross section (left). E,choanosomal skeleton.

Dennal membrane aspiculous. Ectosomal skeleton not evident. 1878, with type species Spongia muelleri Lieberktihn, 1855 (by
Choanosomal skeleton anisotropic with paucispicular fibres and subsequent designation in de Laubenfels, 1936a: 37), is also ajun-
tracts. Spongin scarce. Megascleres slightly curved or rarely ior synonym of Ephydatia by inclusion of E. muelleri in this genus.
straight oxeas, from smooth to microspined (210-400 X 6--19 f,Lm).
Microscleres absent. Gemmules abundant and scattered, subspheri-
cal (350-450 f,Lm). Foramen simple without collar. Gemmular EUNAPIUS GRAY, 1867
theca tri-Iayered. Outer layer well developed generally covering
the distal rotule of gemmuloscleres. Pneumatic layer variably Synonymy
developed with irregular chambers and gemmuloscleres radially
embedded in one layer with the proximal rotule in contact with Eunapius Gray, 1867a: 552.
the inner layer. Inner layer of sublayered compact spongin.
Gemmuloscleres birotules (26--30 X 18-21 f,Lm) with a smooth or Type species
spined shaft, flat rotules of equal diameter, microspined, with
incised irregular margins. Spongilla carteri Bowerbank, 1863: 469 (by subsequent
Ecology. Running and standing waters from coastal brack- designation; Annandale, 1911b).
ish waters to inland salt lakes to alpine lakes. Hibernating, aestivat-
ing or perennial life cycle. Sometimes symbiotic with Diagnosis
zoochlorellae. Distribution. Cosmopolitan. Widely distributed
throughout the N Hemisphere with scattered records from the Spongillidae with body shape ranging from encrusting to bul-
S Hemisphere. The wide distribution of E. jluviatilis suggests the bous, from flattish to lobate. Consistency of live sponges from
existence of a species-complex. fragile to moderately soft. Anisotropic choanosomal skeleton with
Remarks. [Tupha] Oken, 1814 has been suggested as a syn- pauci- to multi spicular fibres and tracts. Medium amount of spon-
onym of Ephydatia Lamouroux, but all of Oken's names have been gin. Megascleres smooth oxeas. Microscleres absent. Gemmules,
deemed to be unavailable and therefore the name is unavailable and single and scattered, subspherical, enveloped in cage of gemmular
seniority does not apply (ICZN, 1956: Opinion 417). Meyenia spicules. Gemmuloscleres smooth or spiny oxeas and strongyles.
Carter, 1881c, with type species Spongiajluviatilis L. (by original
designation), is an objective junior synonym of Ephydatia. Scope and distribution
Pleiomeyenia Mills, 1884, with type species ? Pleiomeyenia
calumeticus Mills, 1884: 147 (by page priority), is a suspectedjun- Thirteen other species: E.fragilis (Leidy, 1851) (cosmopolitan);
ior synonym of Ephydatia and possible synonym of the species E. calcuttanus (Annandale, 1911b) (India); E. michaelseni
E. muelleri (Lieberktihn). Similarly, Trachyspongilla Dybowsky, (Annandale, 1914b) (central Africa); E. crassissimus (Annandale,
Porifera· Demospongiae • Haplosclerida • Spongillina 941

Fig.30. Ephydatiaf/uviatilis , spicules and gemmules, neotype RMNH 1166. A, gemmule. B,cross section of gemmule. C,gemmular surface. D,foramen.
E, trilayered theca (right = multilayered inner layer). F , megascieres and a gemmulosciere. G, oxea tip. H, smooth oxea shaft. I, granules on oxea shaft.
J-K, birotule gemmuloscieres with smooth and spiny oxeas.

1907) (India, Australia, tropical SE Asia); E. igloviformis (Potts, Material examined. Holotype: Not seen. Other
1884) (Canada, USA); E. geminus (Annandale, 1911b) (India); material. 2MB6370-4 (from 1M) - India, 8.x.1894. 2MB6365-5
E. coniferus (Annandale, 1916) (China); E. geei (Annandale, 1918) (from 1M) - India, 8.x.1894. BMNH 1914.11.24.13 - Trivandrum,
(China); E. tinei (Gee, 1932) (Philippines); E. nitens (Carter, India. BMNH 1886.2.9.8 - Mauritius. BMNH 1929.4.13.7-
1881c) (Africa); E. sinensis (Annandale, 191Oc) (China, Canton, China. BMNH 1906.5.31.11 - Lake Victoria, Nyanza,
Manchuria, Russia, Australia); E. potamolepis (Annandale, 1918) scraped from shell of Aetheria. 2MB9544 - Balaton Fured,
(Thailand); E. eryuensis (Sasaki, 1970) (Japan). Cosmopolitan. 19.ix.1938. 2MB2774 - Bombay, 1949.
Description. Body shape ranges from encrusting to bulbous-
Description of type species massive to cone-shaped, from flattish to lobate, or bearing finger-like
projections. Colour from yellowish brown to tan to bright green.
Eunapius carteri (Bowerbank, 1863) (Figs 32-35). Consistency oflive sponges ranges from fragile to moderately soft to
942 Porifera' Demospongiae • Haplosclerida • Spongillina

Fig. 31. Geographic diSiributioll of the genus Ephyt/miu.

3cm

Fig. 32. Eunapius carteri, possible type specimen 2MB 2774.

quite compact. Surface hispid due to tufts of emerging spicules. Fig. 33. Eunapius carteri, skeletal network, possible type specimen 2MB
Dermal membrane thin aspiculous. Oscules from several to single, 2774. Aa, sponge fragment (top view). Ab, sponge cross section (right =
from very small to large (2 cm) sometimes at the apex of projections. surface). B, sponge surface (top view). C-D, choanosomal skeleton with
gemmules (detail of Ab).
Subdermal cavities not observed. Ectosomal skeleton undifferenti-
ated. Choanosomal skeleton anisotropic with pauci- to multispicular
fibres and tracts. Spongin relatively abundant. Megascleres stout degeneration after sexual and asexual (gemmulation) reproduction.
fusiform slightly curved smooth oxeas (265-370 X 14-24/-lm) with Fast growth rate. Distribution: India, Ceylon, Malaysia, Indonesia,
tips from sharply pointed to rounded. Microscleres absent. Java, Madura, Burma, Thailand, China, Philippines, Turkestan,
Gemmules from single to grouped, from scattered to restricted as a Afghanistan, Kurdistan, Iran, E Europe (S Russia, Balaton Lake),
carpet at the sponge base. Gemmules subspherical (440-610 /-lm) Africa?, Mauritius Islands.
enveloped in a cage of gemmuloscleres tangentially or partially
embedded in the pneumatic layer. Foramen tubular simple without
collar. Gemmular theca bi-Iayered. Outer layer absent with outlines
of pneumatic chambers evident at the gemmular surface. Pneumatic HETEROMEYENIA POTTS, 1881
layer thick with regular lines of polygonal chambers. Inner layer
sublayered and connected to pneuma by trabecules of compact Synonymy
spongin. Gemmuloscleres smooth oxeas (145-210 X 5-8 /-lm).
Ecology. In standing waters of ponds, lakes and freshwater Heteromeyenia Potts, 1881a: 150. [Carterellal Potts,
tanks on plants, sticks, stones, shells of Unio sp., not in running 1881b: 176 (preocc.). Carterius Petr, 1886: 92. Oxheteromeyenia
waters. Life cycle characterised by vegetative phase and annual SchrOder, 1927: 107. [Oxyheteromeyenial Auct. (Zapsus).
Porifera· Demospongiae • HaploscIerida • Spongillina 943

Fig. 34. Eunapius carteri, spicules and gemmules; A, C, E,G-I, BMNH 1914.11.24.13. B,D,F, 1886.2.9.8. A, two gemmules. B,gemmular cross section.
C, pneumatic layer. D, foramen. E, internal view of the tube-like foramen. F,close-up of the pneumatic layer and sublayered inner layer (bottom) of the
theca. G, close-up of the gemmular external surface with gemmuloscleres. H, smooth oxeas. I ,tip and shaft of oxeas.

Type species tri-Iayered with gemmuloscleres radially embedded. Gemmuloscleres

birotules of two types: birotules and pseudobirotules.
Spongilla baileyi Bowerbank, 1863: 461 (by subsequent
designation; de Laubenfels, 1936a).
Scope and distribution
Diagnosis
Four other species: H. stepanowii (Dybowsky, 1884a)
Spongillidae with encrusting growth form. Choanosomal (Europe, Russia, China, Japan, Australia); H. latitenta (Potts,
skeleton an irregular network of paucispicular parallel fibres and 1881b) (NE USA); H. tentasperma (Potts, 1880a) (NE USA);
undefined secondary tracts. Sparse spongin. Megascleres acanthox- H. tubisperma (Potts, 1881b) (NE America). Palaearctic, Nearctic
eas. Microscleres acanthoxeas. Gemmules scattered in groups. Theca and Australian Regions.
944 Porifera· Demospongiae • HapJosclerida • Spongillina

Fig. 35. Geographic distribution of the genus Eunapius.

eI;

Fig. 37. Heteromeyenia baileyi, skeletal network, lectotype ANSP

Fig. 36. Heteromeyenia baileyi, lectotype from the collection of E. Potts, P04535. A, sponge fragment (side view). B, sponge fragment (top view)
ANSP P04535. with gemmules. C, choanosomal skeleton. D, ectosomal skeleton.

Description of type species fusifonn oxeas with scattered microspines except at the tips.
Microscleres (75-85 X 2-3 /-Lm) sharply pointed oxeas from
Heteromeyenia baileyi (Bowerbank, 1863) (Figs 36-39). curved to straight entirely spined with microspines, increasing in
Material examined. Lectotype: ANSP P04535 - Spongilla length from tips to the central portion, perpendicular and large.
repens type a, colI. E. Potts, Leigh Cape, Pennsylvania. BMNH Gemmules scattered in groups, subspherical (450-480/-Lm).
1890.1.9.216 - Krasetice and Nimency, Boemia, 19.x.1885. Foramen tubular with scarcely developed collar and devoid of cirri.
2MB6275 - Munster, 7.vii.1926, det. Arndt, 1927, det. Schroder. Theca tri-layered with gemmuloscleres radially embedded. Outer
2MB (DTRG-FW538) (H. baileyi var. repens) - Creba, Kr. layer with emerging shafts and distal rotules of both types of gem-
Rotemburg, Schlesia, .x.l929, SchrOder. muloscleres. Pneumatic layer well developed with irregular cham-
Description. Thinly encrusting sponges. Surface hispid due bers. Inner layer composed of compact sublayered spongin.
to spicules emerging through the dennal membrane. Oscules incon- Gemmuloscleres of two types: pseudobirotules (80--85 X 22 /-Lm),
spicuous. Consistency of live sponge soft. Colour bright green in few inserted with their proximal rotules in the outer zone of gem-
life. Dennal membrane spiculose (microscleres). Subdennal cavi- mular theca, cylindrical shaft usually smooth and often irregularly
ties not observed. Ectosomal skeleton as spicules in the dennal bent, pseudo-rotules bear long recurved hooks with microspinosity
membrane. Choanosomal skeleton an irregular network of pau- at the tips; birotules (50--60 X 22 /-Lm) very numerous with straight
cispicular parallel fibres and undefined secondary tracts. Sparse shafts bearing few conspicuous conical spines, variably umbonate
spongin. Megascleres (255-315 X 11-10 /-Lm) sharply pointed rotu1es with incised margins bearing spines.
Porifera' Demospongiae • HapJoscierida • Spongillina 945

Fig. 38. Heteromeyenia baileyi, spicules and gemmules, lectotype ANSP PO 4535. A, gemmule. B, foramen. C, cross section of the foraminal tube.
D, gemmular external surface. E, multi-layered inner layer and pneumatic layer of the trilayered theca. F, megascieres oxeas and two types of gemmu-
loscieres. G, tip of an oxea. H-J, short birotule gemmulosciere. J-K, long pseudo-birotule gemmulosciere.

Ecology. In shallow standing waters on submerged plants, 1884a: 507 as Dosilia) (by original designation) also belongs to
timber and sticks, stems of Nitella, leaves of Potamogeton. this genus according to Penney & Racek (1968). Oxheteromeyenia
Distribution: N America from Canada to Louisiana, Mexico, SchrOder, 1927 and a misspelling [Oxyheteromeyenial Auct. probably
Germany ?, Poland ? also belong here.

Remarks
HETEROROTUIA PENNEY & RACEK, 1968
[Carterella 1Potts, 1881 b, with type species C. latitenta Potts,
1881b: 176 (by monotypy) is both preoccupied and a junior Synonymy
synonym of Heteromeyenia (Penney & Racek, 1968). Likewise,
Carterius Petr, 1886, with type species C. stepanowii (Dybowsky, Heterorotula Penney & Racek, 1968: 96.
946 Porifera· Demospongiae • Haplosclerida • Spongillina

Fig. 39. Geographic di lribulion of lhe genus Heteromeyellia.

Fig. 40. Heterorotula capewelli. type specimen BMNH unregistered.

Type species

Spongilla capewelli Bowerbank, 1863: 447 (by original Fig. 41. Heterorotula capewelli, skeletal network, type specimen BMNH
designation). unregistered. A, cross section (top = sponge surface). B, sponge surface
(top view). C, ectosomal skeleton (top). D, choanosomal skeleton.
Diagnosis
Description of type species
Spongillidae with massive and bulbous growth form and with
lobate surface. Choanosomal skeleton irregularly alveolate with Heterorotula capewelli (Bowerbank, 1863) (Figs 40-43).
paucispicular tracts. Moderate amount of spongin. Megascleres Material examined. Holotype: BMNH (Bowerbank CoIl.) -
spiny oxeas. Microscleres absent. Gemmules scattered with cages Lake Hindmarsh, Victoria, SEAustralia, 1863.
of megascleres. Gemmular theca trilayered with radially embedded Description. Sponge massive and bulbous. Colour light tan
gemmuloscleres. Gemmuloscleres birotules. to brown. Consistency of live sponges moderately hard, firm but
brittle. Surface lobate with high and rounded irregular ridges sepa-
Scope and distribution rated from each other by comparatively deep clefts. Oscules few
and conspicuous. Dermal membrane well developed separated
Six other species: H. nigra (Lendenfeld, 1887) (E Australia); by the underlying symplasm. Subdermal cavities not observed.
H. multidentata (WeHner, 1895) (E Australia, Tasmania); Special ectosomal skeleton absent. Choanosomal skeleton irregu-
H. multiformis (Weltner, 19lO) (W Australia); H. contraversa larly alveolate with paucispicular tracts but also more variably ori-
(Racek, 1969) (E Central Australia); H. kakahuensis ented thick fibres joined by slimmer transverse fibres are present in
(Traxler, 1896) (New Zealand); H. fistula Volkmer & Motta, 1995 some areas of the sponge body at the level of lobes. Moderate
(S America). Australian and Neotropical Regions. amount of spongin. Megascleres (195-330 x 13-18 fLm) stout
Porifera· Demospongiae • Haplosclerida • Spongillina 947

Fig. 42. Heterorotula capewelli. spicules and gemmules. type specimen Bowerbank collection BMNH unregistered. A. megascleres oxeas. B. shaft of
a spiny oxea. C. gemmules in their cage. D. foramen. E. cross section of gemmular theca. F-H, gemmuloscleres.

Fig.43. Geographic distribution ofthe genus Heterorotu/a.

948 Porifera' Demospongiae • Haplosclerida • Spongillina

fusiform oxeas from smooth to microspined. Microscleres absent. Diagnosis

Gemmules (510-600 fl-m) subspherical, scattered in the skeletal
network and more abundant and grouped in basal parts. A loose Spongillidae with encrusting growth form, moderately hard,
cage of irregularly arranged megascleres envelopes gemmules. friable. Dermal membrane supported by spicules. Subdermal
Foramen simple or bearing a shallow peripheral collar. Gemmular cavities notably developed. No well defined central cavity.
theca trilayered. Outer layer distinctly corrugated and of pale yel- Choanosomal skeleton variable, never very stout, ranging from
low colour with tangentially arranged megascleres of the cage. reticulate to alveolate with primary fibres quite distinct, transverse
Pneumatic layer very thick with radially embedded gemmuloscle- fibres not very distinct. Megascleres spined oxeas. Gemmules rare,
res. Inner layer of multilayered compact spongin. Gemmuloscleres when present dome-shaped flattened at the base, adherent to basal
in the gemmular theca with proximal rotule firmly inserted in the plate, devoid of foramen, spicules and pneumatic layer.
inner layer and distal rotule generally not reaching the outer layer.
Gemmuloscleres birotules (34-52 fl-m) with a slender smooth shaft Scope and distribution
bearing sometimes few spines. Rotules flat with small spines and
margins irregularly crenulated with rare teeth. Diameter of rotules Six other species. N. yunnanensis (Annandale, 191Ob) (Yunnan,
unequal (24-28 fl-m; 20-23 fl-m). W China); N. ehraiensis Lizhen, 1998 (yunnan, W China); N.
Ecology. Inland slightly brackish athalassic waters of arid tanganykae (Evans, 1899) (Lake Tanganyika, Africa); N. moorei (Evans,
regions. Distribution. Central Australia, ranging east of the Great 1899) (Lake Tanganyika, Africa); N. cunningtoni (Kirkpatrick, 1906)
Dividing Range and north to Queensland. (Lake Tanganyika, Africa); N. vasta (Weltner, 1901) (Celebes
[Sulawesi], Indonesia). Oriental and Afrotropical Region.

NUDOSPONGlUA ANNANDALE, 1918

Synonymy

NudospongillaAnnandale, 1918: 62.

Type species

Spongilla coggini Annandale, 1910b (by original designation). Fig. 44. Nudospongilla coggini, paralectotype BMNH 1914.11.24.32 (dry).

Fig. 45. Nudospongilla coggini, skeletal network, paralectotype 1914.11.24.32. A, cross section (left = sponge surface). B, ectosomal skeleton (left).
C, choanosomal skeleton. D, sponge surface (top view). E, detail ofD.
Porifera· Demospongiae • Haplosclerida • Spongillina 949

Description of type species network of spicules (according to the original description).

Choanosomal skeleton vaguely alveolate-reticulate, paucispicular in
Nudospongilla coggini (Annandale, 191Ob) (Figs 44--47). the studied syntype (reported in the original descriptions as never very
Material examined. Paralectotype: BMNH 1914.11.24.32 stout), with a close regular network with small meshes, with radiating
(ZEV 3809.7) - dry colI., Coggin, Lake JaJi-Zu (Hai), Yunnan, W (vertical) fibres stout and more coherent of distinguishable transverse
China (without gemmules). Lectotype: 1M not available - Lake fibres. The network is more vague at the base of the sponge. Spongin
Erh Hai, Ta-Li Fu, Yunnan, J. Coggin Brown leg. poor. Megascleres (258-283 X 19-28IJ..m) stout, straight, to feebly
Description. Encrusting rounded sponge to irregularly curved spined oxeas with spines more dense and larger at the tips.
massive, occasionally with short flattened branches. Size from 2 X Few smaller and more sharply pointed oxeas are loose within the
2cm to 5 X 4 X 3.8 cm. Living specimens pale grey to brilliant skeleton. Microscleres absent. Gemrnules when present moderate in
green, light brown when preserved. Consistency moderately hard, size (?), naked (without spicules), ovoid and flattened at the base,
fragile. Surface irregular and minutely hispid due to tufts of spicules dome shaped and adherent to the basal plate. Gemrnular theca of
(3) at the apices of slender fibres (3 spicules thick). Oscules incon- compact spongin thin and brittle, outer layer in continuity with the
spicuous in depressed areas. Dermal membrane closely adherent. basal plate. No foramen, no pneumatic layer, no gemrnuloscleres.
Basal spongin plate delicate. Ectosomal skeleton a horizontal Ecology. In standing waters on the lower surface of stones,
sometimes englobing objects such as fragments of sticks, small
stones and shells. Often on old shells. Larva parenchymula in
October. Distribution. Yunnan Lakes.
Remarks. All species of Nudospongilla were initially
assigned to Spongilla, being spongillid-like with oxeas. To this
genus were assigned species with oxeas and no gemmules col-
lected from W China, Sulawesi, Africa and Tiberiade/Kinneret.
Racek (1974) relegated N. aster, N. mappa, N. reversa of
Annandale (1918) as synonyms of Cortispongilla barroisi. In spite
of this the author declared the genus Nudospongilla a nomen
nudum, which we consider is an invalid action. The gemmular
characters of this genus (theca monolayered without gemmuloscle-
res) are not in agreement with the rules of the family Spongillidae.

PACHYROTULA VOLKMER-RIBEIRO & RUTZLER, 1997

Synonymy
Fig. 46. Nudospongilla coggZn!, skeleton, paralectotype BMNH
1914.11.24.32. A-D, close-up of different characters of tips and shafts of
megascleres (spiny oxeas), with two different kinds of spines. Pachyrotula Volkmer-Ribeiro & Riitzler, 1997: 490.

Fig. 47. Geographic distribution of the genu NlIdospongilla.

950 Porifera· Demospongiae • HapJosclerida • Spongillina

Type species Material examined. Holotype: USNM 238820 - River

Le Diahot, New Caledonia, F. Starmuhler, 18.ix.1965. Paratype:
Spongilla (Stratospongilla) raceki Riitzler, 1968: 60 (by USNM 23883 - same details.
monotypy). Description. Slender, irregular, small soft crusts, ca. 1 mm
thick. Colour greyish-ochre in alcohol. Very soft consistency.
Diagnosis Oscula and pores inconspicuous. Dermal membrane and subder-
mal cavities not observed. Hispid body surface due to tufts of
Spongillidae with encrusting growth form. Alveolate skeleton spicules. Ectosomal skeleton as tufts of spicules. Choanosomal
with sparse spongin. MegascIeres oxeas, ranging from spined to paucispicular skeleton densely and irregularly alveolate with
rarely smooth. MicroscIeres absent. Gemmules at the sponge base sparse spongin. Megascleres (160-245 X 7.5-14 /-Lm) straight to
in cages of oxea megascIeres. Gemmular theca tri-layered. curved oxeas ranging from microspined to rarely smooth, tips
Foramen tube-like. GemmuloscIeres range from birotules with abruptly pointed. MicroscIeres absent. Gemmules (350-650/-Lm)
very variable shape to spiny short oxeas.

Scope and distribution

Monotypic, New Caledonia.

Description of type species

Pachyrotula raceki (RiitzIer, 1968) (Figs 48-51).

lJS~~ ~38g3 PA~tt-\"jff.

5.p~31\\~ ""ac=.e.k.. ~~
New GLledo nl CI-
'1~J~ rrtcd
Fig. 48. Pachyrolllill race/d. paralype M 23883.

Fig. 50. Pachyrotula raceki, spicules and gemmules, A-D, F, holotype

USNM 238820. E, G-I, paratype USNM 23883. A, megasclere spicular
cage. B, external surface of a gemmule showing the spiny tips of gem-
muloscleres (see also F). C, cross section of gemmule and cage. D, trilay-
ered gemmular theca showing a multi-layered inner layer and
Fig. 49. Pachyrotula raceki, skeletal network, paratype USNM 23883. gemmuloscleres embedded in the pneumatic layer. E, rotule of gemmu-
Aa, sponge fragment (top view). Ab, cross section of the sponge fragment losclere. F, modified rotule of a demmulosclere. Ga, microsclere. Gb, gemmu-
(lateral view). Ac, sponge base (top view). B, surface hispidation. losclere. Gc, megasclere smooth oxea. Gd, megasclere spiny oxea.
C, choanosomal network. D, choanosomal skeleton meshes. H, spiny (left) and smooth (right) megascleres. I, tip and shafts of megascleres.
Porifera' Demospongiae • Haplosclerida • Spongillina 951

Fig. 51. Geographic dislribulion of the genu Pocl/yrOllllo.

sub spherical at the base of the sponge, enclosed in a cage of short

acanthoxea megascleres, enveloping one or more gemmules (up to
4). Foramen simple as a short tube (45-50 j.Lm). Gemmular theca
tri-Iayered. Outer layer notably thick with emerging distal rotules.
Pneumatic layer thin with irregular chambers. Inner layer multi-
layered. Gemmuloscleres irregularly tangential to radially embed-
ded in the theca. Gemmuloscleres (20-80 j.Lm length) stout
birotules notably variable in shape with smooth shafts (5-8j.Lm
thick), rotules or knob-like (9-20 j.Lm in diameter) ends orna-
mented by tubercules, rosettes or spines; tips sometimes bear
grouped spines and appear as oxeas (120 X 8j.Lm).
Ecology. On the lower surface or concavities of stones, both
in running and standing waters with low conductivity (28-56) and
pH range 6.6--7.1. Distribution. New Caledonia.
Remarks. Riitzler (1968) originally assigned this species to
Spongilla (Stratospongilla) and considered that it was closely Fig. 52. Pectispongilla aurea, two specimens of the variety subspinosa.
A, schizosyntype from the collection of Penney (90124) with 3 labels
related to Stratospongilla gilsoni (Topsent, 1912) from Fiji. Volkmer- (USNM 34578). B, a specimen from the BMNH 1914.11.24.34 and pre-
Ribeiro & Riitzler (1997) subsequently revised the type material served in the 2MB 7981. Both specimens are represented by the same
and erected this monotypic genus closer to Heterorotula Penney & aquatic plant with scanty sponge remains.
Racek, 1968.

layers. Pneumatic layer of spongin fibres (without chambers).

PECTISPONGILLA ANNANDALE, 1909 Gemmuloscleres radially embedded, botryoides-like.

Synonymy Scope and distribution

Pectispongilla Annandale, 1909a: 103. Three other species: P. subspinosa Annandale, 1912 (India,
Japan); P. stelliferaAnnandale, 1915b (India); P. botryoides Haswell,
Type species 1882 (Australia). Tropical India and Australia, ? Japan.

Pectispongilla aurea Annandale, 1909a: 103 (by original Description of type species
designation).
Pectispongilla aurea Annandale, 1909a (Figs 52-55).
Diagnosis Material examined. Holotype: 1M not available. Other
material. BMNH 1914.11.24.34 - P. subspinosa, Ernakulam,
Spongillidae forming small cushions. Consistency of live Cochin State, India. USNM 34578 - P. subspinosa, Ernakulam,
sponges soft. Choanosomal skeleton consisting of feebly coherent Cochin State, India. DTRG FW568 (from NTM) - Pectispongilla
spicule fibres. Spongin abundant. Megascleres smooth to spined sp., R. Manconi, Kakadu Park, Northern Territory, Australia.
oxeas. Microscleres microspined oxeas with rhomboid outline. Description. Body cushion-shaped. Colour deep golden.
Gemmules scattered in the skeleton. Gemmular theca with three Consistency of live sponges soft. Oscula few and inconspicuous.
952 Porifera· Demospongiae • HapJosclerida • Spongillina

Fig. 53. Pectispongilla aurea var. subspinosa, skeletal network, schizosyntype USNM 34578. A-C, sponge fragment on an aquatic plant at different
magnifications.

Fig. 54. Pectispongilla, spicules and gemmules. A-E, Pectispongilla sp. DTRG-FW568 (ex-IZUG). F-K, P. aurea variety subspinosa, co-type from
BMNH 1914.11.24.34. A, gemmule. B, foramen. C, gemmular outer layer surface. D, cross section of gemmule. E, trilayered gemmular theca.
F, megascleres oxeas. G, smooth oxea. H, spiny oxeas. I, shaft of a spiny oxea. J, tip of a spiny oxea. K, gemmuloscleres.
Porifera' Demospongiae • Haplosclerida • Spongillina 953

Fig. 55. Geographic di. lribulion of the genus Peclispollgil/a.

Dermal membrane well developed. Surface hispid. Ectosomal

skeleton undetected. Choanosomal skeleton consisting of
feebly coherent spicule fibres. Spongin abundant. Megascleres
(270-320 X 13-16 /-Lm) range from fusiform sharply pointed oxeas
from straight to bent, and from smooth to spined. Microscleres
(20-50 X 1.5-3.5 /-Lm) reported in the literature as straight and
sharply pointed oxeas of rhomboid outline usually microspined.
•
Gemmules (190-220/-Lm) subspherical, scattered in the skeleton.
Foramen tubular. Gemmular theca with three layers. Outer layer
surface irregular due to the partial emergence of gemmuloscleres
apices. Pneumatic layer well developed as a network of spongin
fibres. Gemmuloscleres radially embedded in the theca crossing
each other at slanting angles with botryum-like apices variably ori-
ented. Inner layer multilayered. Gemmuloscleres (31-37 X 2.5-4 Q

/-Lm) typical of this genus with smooth distinctly curved shafts Fig. 56. Racekiela ryderi, lectotype ANSP PO 4536.
characterized, in the apex of convex sides, by the disto-lateral
arrangement of spines in rows joined to each other by siliceous
webs to form botryoides-like apices in mature gemmuloscleres, Diagnosis
with well defined polygonal concavities of the botryum. Apices of
young spicules with scarcely developed siliceous webs appear as Spongillidae with body shape encrusting to lobate, ramose or
spines in rows. massive. Ectosomal skeleton with tangential megascleres on the
Ecology. Unknown for the type species. dermal membrane and irregularly scattered erect megascleres.
Choanosomal skeleton consists of irregular paucispicular isotropic
network with sparse spongin. Megascleres are acanthoxeas.
Microscleres absent. Gemmular theca tri-Iayered with gemmuloscle-
RACEKIELA BASS & VOLKMER· RIBEIRO, 1998 res radially embedded. Gemmuloscleres of two types: birotules and
pseudobirotules.
Synonymy
Scope and distribution
[Acanthodiscus] Volkmer-Ribeiro, 1996: 35 [preocc.].
Racekiela Bass & Volkmer-Ribeiro, 1998: 125. Only one other species: R. sheilae (Volkmer-Ribeiro et al.,
1988) (S America). Palaeartic, Nearctic and Neotropical.
Type species
Description of type species
Heteromeyenia ryderi Potts, 1882: 13 (by subsequent designa-
tion; Volkmer-Ribeiro, 1996). Racekiela ryderi (Potts, 1882) (Figs 56-59).
954 Porifera· Demospongiae • Haplosclerida • Spongillina

Fig. 57. Racekiela ryderi, skeletal network, lectotype ANSP P04536. A, section through the entire sponge thickness (left = surface; right = spongin basal
plate). B-C, sponge surface and ectosomal skeleton (top view) at different magnifications. D-E, surface hispidation (lateral and top view). F, spongin basal
plate with gemmules. G, choanosomal skeleton.

Material examined. Lectotype: ANSP P04536 (pott's coll.) - 46-64 X 4-8.5/Lm) with small umbonate pseudo-rotules (17-23/Lm
Cobb's Creek, Delaware River, USA, E. Potts, 1881. Other material. (other material» bearing irregular hooks variably curved; stout
DTRG FW363 - Norway, Oakland. BMNH 191O.1.1.622.C (from spined shafts, spines irregularly distributed and hook-shaped.
INM) - Ireland. BMNH 1866.12.9.3 - dry box 6, Stream and Loch. Birotules (33-49 X 5-8/Lm; 28-41 X 3-5/Lm) with usually
Description. Thinly encrusting to massive ramose or papil- smooth shafts and large disk-like rotules (25-29/Lm; 20-28/Lm)
late sponges. Colour green, brown or white. Small conspicuous bearing serrated margins and irregularly distributed spines or
oscules. Dermal membrane spicular (from protruding megascle- tubercules.
res). Subdermal cavities not observed. Ectosomal skeleton is com- Ecology. Body shape varies significantly according to environ-
posed of irregularly scattered erect megascleres producing a hispid mental conditions, encrusting in running waters and massive in lentic
surface and tangential megascleres lying on the dermal membrane. conditions. Distribution. Amphiatlantic in the northern hemisphere.
Choanosomal skeleton is an irregularly paucispicular isotropic net- From Eastern North America and Canada to W British Islands, Faroes
work with sparse (?) spongin. Megascleres (296-431 X 12-26/Lm and SW Norway. One record (?) from Central America (Belize).
(holotype); 141-279 X 1-21/Lm (other material» stout to slender, Remarks. The type species was moved from the genus
straight to slightly curved, heavily to sparsely spined, abruptly to Anheteromeyenia by Volkmer-Ribeiro (1996). However, the pro-
gradually pointed oxeas; spines range from straight to oriented. posed generic name [Acanthodiscus] was preoccupied and thus
Microscleres absent. Gemmules hemispherical (300-800/Lm; Racekiela was proposed as a replacement name by Bass &
300-400 /Lm), abundant, with no consistent distribution. Foraminal Volkmer-Ribeiro (1998). Moreover in the last paper, the two names
tube inconspicuous, not higher than the shorter gemmuloscleres. Racekiela and Racekiella are used by the authors. We here propose
Gemmular theca trilayered with radially arranged gemmuloscleres. the first name appeared in the text.
Outer layer covering the outer rotules of short gemmuloscleres.
Pneumatic layer thick, with irregular chambers. Inner layer of sub-
layered compact spongin (3-4). Gemmuloscleres of two types: RADIOSPONGILLA. PENNEY & RACEK, 1968
with short birotules completely embedded in the theca, whereas
long pseudobirotules project distal rotules beyond the outer layer. Synonymy
Number and distribution of pseudobirotules notably variable.
Gemmuloscleres of two classes. Pseudobirotules (47-92 X 5-10 /Lm; Radiospongilla Penney & Racek, 1968: 61.
Porifera· Demospongiae • Haplosclerida • Spongillina 955

Fig. 58. Racekiela ryderi, spicules and gemmules, lectotype ANSP P04536. A, gemmule in toto. B, gemmular surface. C, foramen. D, cross section
of foramen. E, trilayered gemmular theca. F, multi-layered inner layer. G, cross section of gemmule. H, spicular complement of spiny oxeas and two types
of gemmulosc\eres. I, spiny megasc\eres. J, microspination of pseudo-birotule gemmulosc\ere. K, pseudo-birotule gemmulosc\ere. L, megasc\ere shaft. M,
megasc\ere tip. N, birotule gemmuiosc\eres.

Type species spined oxeas. Gemmules occur in the basal part of the sponge or
are scattered. Gemmular theca tri-Iayered with gemmuloscleres
Spongilla sceptroides Haswell, 1882: 209 (by original desig- radially embedded. Gemmuloscleres acanthostrongyles with tips
nation; Penney & Racek, 1968). bearing apical spines (oxeas ?).

Diagnosis Scope and distribution

Spongillidae with encrusting growth form with occasional Fourteen other species: R. amazonensis Volkmer-Ribeiro &
delicate cylindrical branches. Choanosomal skeleton paucispicular Maciel 1983 (Neotropical region); R. cantonensis (Gee, 1929)
anisotropic with ill-defined secondary tracts. Megascleres are (China); R. cerebellata (Bowerbank, 1863) (tropical and subtropical
956 Porifera' Demospongiae • Haplosclerida • Spongillina

Fig. 59. Geographic di lributi on of the genu Racekiela.

Fig. 61. Radiospongilla sceptroides, skeletal network, neotype AMS

Africa, India-Pakistan, Indonesia, Philippines, New Guinea, China, Z2837. A, cross section (top = surface). B, choanosomal network. C, ecto-
Russia ?, SE Europe [possibly accidental introduction]); R. cinerea somal skeleton in irregular surface (lateral view). D, ectosomal skeleton in
(Carter, 1849)(Bombay, Himalayas); R. crateriformis (Potts, 1882) smooth surface (top view).
(USA, Canada, Mexico, West Indies; Suriname, China, Japan,
S Asia, Australia); R. hernephydatia (Annandale, 1909) (India,
New Guinea, E Australia); R. hispidula (Racek, 1969) (Australia);
R. indica (Annandale, 1907) (India, Indonesia, Philippines, New South Wales, Australia, on large rock, 0.5 m depth, F. Hersey,
New Guinea ?); R. rnultispinifera (Gee, 1933) (E Australia); 6.iii.1958, det. Penney & Racek, 1968: 66. Other material. BMNH
R. philippinensis (Annandale, 1909c) (Philippines to N Australia); 1886.8.27.665, 13.m.C, Brisbane, Australia. BMNH 1886.8.27.658 -
R. sansibarica (Weltner, 1895) (Zanzibar, N Rhodesia, Belgian dry box 5, Kakalum River, Australia, Lendenfeld.
Congo); R. sinoica (Racek, 1969) (Australia); R. sendai (Sasaki, Description. Body shape consists of flat crusts or with
1936) (Japan); R. hozawai (Sasaki, 1936) (Japan). Nearctic, thin finger-like projections. Consistency of live sponges is firm
Palearctic, Neotropical, Afrotropical, Australian Oriental Regions. and elastic. Colour emerald green in life, grey-light brown in
dry specimens. Oscules numerous with a system of radiating fur-
Description of type species rows in lotic habitats. Surface hispid. Dermal membrane well
developed. Subdermal cavities not reported. Basal spongin plate
Radiospongilla sceptroides (Haswell, 1882) (Figs 6~3). not reported. Ectosome not evident. Choanosomal skeleton
Material examined. Neotype: AMS Z2837 - Merrika River, paucispicular, anisotropic, with ill-defined secondary tracts.
Nadgee Faunal Reserve near Womboyne (37°15'S, 149°55'E), Spongin scarce ? Megascleres fusiform slightly curved oxeas
Porifera· Demospongiae • Haplosclerida • Spongillina 957

(240--310 X 8-11jLm) with scattered minute spines except at their

tips. Microscleres absent. Gemmules in the basal part of the sponge,
subspherical (420--500 jLm). Foramen tubular without collar.
Gemmular theca tri-layered with gemmuloscleres radially embed-
ded. Outer layer with protruding shafts and distal apices of gemmU-
loscleres. Pneumatic layer well developed with irregular chambers.
Inner layer sublayered. Gemmuloscleres slender acanthostrongyles
with tips bearing apical spines (oxeas?). Spines recurved and
grouped at the tips to form a scepter-like shape.
Ecology. On wood or rocks. In lentic or lotic waters. Non-
alkaline environments with a scattered distribution. Colour in life is
due to specific pigments. Distribution: E Australia, New Zealand,
New Guinea, New Caledonia.

SANIDASTRA VOLKMER-RIBEIRO & WATANABE, 1983

Synonymy

Sanidastra Volkmer-Ribeiro & Watanabe, 1983: 151.

Type species

Sanidastra yokotonensis Volkmer-Ribeiro & Watanabe, 1983:

151 (by monotypy).

Diagnosis

Spongillidae with encrusting to massive body shape.

Choanosomal skeleton irregular with vague pauci- to multispicular
tracts. Microscleres absent. Gemmular theca tri-Iayered.
Gemmuloscleres oxeas irregularly radially embedded within the
Fig. 62. Radiospongil/a sceptroides, gemmules and spicules, neotype theca.
AMS Z2837. A, gemmules with skeletal megascleres. B, foramen. C, cross
section of gemmules with radial gemmuloscleres. D, cross section of the
theca with thick pneumatic layer. E, megascleres spiny oxeas and gemmu- Scope and distribution
loscleres. Fa, megascleres. Fb, gemmuloscleres. G, close-up of the megas-
clere shaft. H, close-up of the gemmulosclere spiny tip. Monotypic. Japan, Sardinia and Corsica.

Fig. 63. Geographic di tributi oll of the genus Radiospollgilla.

958 Porifera· Demospongiae • Haplosclerida • Spongillina

Description of type species DTRG FW179 - Rio Su Lernu, Sardinia 18.vii.1987. DTRG FW
217 - Gravona River tributary, near Aiaccio, Corsica, 9. viii. 1987.
Sanidastra yokotonensis Volkmer-Ribeiro & Watanabe, 1983 Description. Body shape (5 X 4-5 cm) encrusting with fla-
(Figs 64-67). bellate surface. Colour greenish-yellow to light brown in life.
Material. Holotype: NSM P04 - Yokotone-gawa Canal, Consistency soft delicate compressible in life. Surface slightly irreg-
Nakajima, Azura-mura, Ibaragi, Japan. Other material. DTRG ular and hispid due to emerging brushes of spicules. Small and scat-
FW156 - Gravona River tributary near Aiaccio, Corsica, 23.iv.1987. tered oscules. Dermal membrane conspicuous and spicular. Dermal
cavities not observed. Ectosomal skeleton as spicules in the dermal
membrane. Choanosomal skeleton irregular, vague, pauci- to multi-
spicular tracts, with long axial fibres in flabellate expansions.
Megascleres oxeas (190-320 X 8.5-15 f.,Lm) ranging from smooth to
3cm
spiny with microspines notably variable in size and density. Tips, in
most cases, sharply pointed, rarely blunt or angulate. Microscleres
absent. Gemmules, scattered singly or grouped, subspherical
(368-514 f.,Lm). Foramen is a short tube (20 f.,Lm in diameter), simple
without collar. Gemmular theca tri-Iayered. Outer layer with irre-
gular surface and emerging apices of gemmuloscleres. Pneumatic
. cl ... TYP -: .Jt.Sc;rnc • .'lu5. n· N5NT-pc";4 layer (20-30 f.,Lm thick) with irregular chambers. Inner layer multi-
_ n:..;..;..;'d::>.J:..::":..::t.:..r·~l I ,.....
",5,;;;." • ot l·"n n,,',:.;,
.. I
n.9cn e l l .Sp.
layered. Gemmuloscleres are irregularly radially embedded within
the theca. Gemmuloscleres oxeas (38.5-65.5 X 5.5-9 f.,Lm), with
Vull..m~.·_ ,bciru : ~"t"n~b~. 198)
2-9 rays, show tips ranging from rounded to pointed to tricuspidate.
Yokuto~ -!Jpwcl Cilnu I ~ 'tJ~t..Ij ima, A:wn ..,-mur/o1 Ecology. In coastal areas of canals and courses in freshwaters
Ibllr"9' Pt'e' ('ctur , Jupun, ·,o,·clhb<'r
"p. Y""o tundbt"
19
I{'~.
,:!. from perennial to ephemeral, under stones, associated to Ephydatia

------------------- jluviatilis. Distribution: highly disjunct, from Japan to Mediterranean

Fig. 64. Sanidastra yokotonensis, holotype NSM P04. (Corsica and Sardinia).

Fig. 65. Sanidastra yokotonensis, skeletal network, holotype NSM PO-4. A, cross section through a small digit (sponge surface = all the perimeter).
B-C, choanosomal skeleton network. D, ectosomal skeleton of an irregular surface (lateral view). E, ectosomal skeleton of a relatively smooth surface.
Porifera· Demospongiae • Haplosclerida • Spongillina 959

Fig. 66. Sanidastra yokotonensis, gemmules and spicules. A-F, holotype NSM PO-4. G-H, DTRG-FW179 (ex-IZUG). A, gemmule with foramen.
B, cross section of gemmule with foramen on the left (middle). C, trilayered theca with sublayered inner layer (bottom). D-E, megascleres spiny
and smooth oxeas. F-H, gemmuloscleres.

Fig. 67. Geographic di tribulion of the genu Sallidaslra.

960 Porifera· Demospongiae • HaploscIerida • Spongillina

Remarks. Volkmer-Ribeiro & Watanabe (1983) included

S. yokotonensis in the family Latrunculiidae, order Poecilosclerida,
on the basis of several similarities with hadromerids, such as skele-
tal subtylostyles and sanidasters as gemmuloscleres. However, E,,· I1~C IJ! 20 ~
re-analysis of skeletal elements showed that spicules were repre- C::J)JC.e.'f-'h> in R4 I1VAJOO ~'O ~lltv4
sented only by smooth and spiny oxeas, rarely modified into "freak C~l J C I)( Ml\.L.o( Ait 41.
oxeas with one blunt or angulate tip", and none of them could be !lATE. ['!IS O
defined as substrongyles or tomotes. Regarding gemmuloscleres, a
comparison between sanidasters of hadromerids and gemmuloscle- f1NR J 003
res of S. yokotonensis indicates that they are clearly not homologous.
The typical radial arrangement of gemmuloscleres in the theca of
S. yokotonensis seems to be allied to the Radiospongilla 'group' and
fossil Palaeospongillidae Volkmer-Ribeiro & Reitner (1991b).
2cm

SATURNOSPONGILLA VOLKMER-RIBEIRO, 1976 Fig. 68. Salllmospollgi//a can'al/wi. ho)otype (FZRG) MN RJ·()()3.

Synonymy

Saturnospongilla Volkmer-Ribeiro, 1976: 271.

Type species

Saturnospongilla carvalhoi Volkmer-Ribeiro, 1976: 271

(by monotypy).

Diagnosis

Spongillidae with bush-like growth form. Consistency

extremely delicate. Ectosomal skeleton occurs as a dense structure
on the apex of primary fibres at the surface. Choanosomal skeleton
is a loose anisotropic paucispicular reticulation with primary fibres
diverging in tufts and becoming more dense towards the sponge
surface and with irregular secondary tracts. Spongin very sparse.
Megascleres oxeas. Microscleres absent. Gemmules scattered at the
sponge base. Satumian-shaped gemmules with a ring-like pneu- Fig. 69. Saturnospongilla carvalhoi, skeletal network, holotype MNRJ·
matic layer. Pseudo-cage of acanthoxeas exclusively at the ring 003. A, skeletal fragment with spiCUlar fibres arising from the basal spon·
surface. Gemmular theca of compact spongin. Gcmmuloscleres gin plate. B, multi spicular fibres. C, choanosomal skeleton trapping
birotules. gemmules. D, spongin basal plate.

Scope and distribution becoming more dense towards the sponge surface, and with irregular
secondary tracts. Spongin very sparse. Megascleres (215-399.5 X
Monotypic. River Jurua. 8.3-25/-lm) of the skeleton range from smooth, slender to almost
stout, straight to slightly curved abruptly pointed oxeas. Acanthoxeas
Description of type species (123-296 X 6.6-15 /-lm) of the gemmular pseudo·cage are curved and
sharply pointed, with central conical large spines curved and more
Saturnospongilla carvalhoi Volkmer-Ribeiro, 1976 dense towards the tips. Microscleres absent. Gemmules abundant at
(Figs 68-71). the sponge base. Gemmules (541-824.6/-lm) with concave·convex
Material examined. Holotype: MNRJ 003 (ex-MRCN 209) - profile with foramen bearing a short porus tube at the middle inferior
Conceicao do Raimundo, middle Rio Jurua, Brazil, J.C. de Mello concavity. Gemmules satumian-shaped with a ring (thoroid) sur-
Carvalho, 25.vi.1950. Other material. FZRG MRCN107 - Rio rounding the central subspherical theca. Tangential acanthoxeas at the
Jurua, lC. de Mello Carvalho. ring surface represent a pseudo·cage. The ring is covered by an outer
Description. Body shape bush-like, from hemispherical to layer chambered spongin, representing a pneumatic layer, shifted
spherical (0.2-2.5 cm). Brownish-yellow colour. Consistency from its usual position. Gemmular theca with gemmuloscleres radi-
extremely delicate, fragile. Oscules inconspicuous. Surface hispid ally embedded in the inner layer of compact sublayered spongin.
due to emerging tufts of spicules from the conspicuous dermal mem- Gemmuloscleres (10-33 X 3.3-5 /-lm) stout birotules with smooth
brane. Subdermal cavities not observed. Basal spongin plate conspic- shaft. Rotules (13.3-16.6/-lm) with recurved circular entire margins
uous and thin. Ectosomal skeleton is formed by the densely arranged and central umbone, proximal rotule flat, distal rotule cup-shaped.
apices of the primary fibres. Choanosomal skeleton is loosely Ecology. This species was reported as dominant in freshwater
anisotropic, paucispicular, with primary fibres diverging in tufts and sponge communities from the middle Rio Jurua, associated with
Porifera· Demospongiae • Haplosclerida • Spongillina 961

Fig. 70. Saturnospongilla carvalhoi, gemmules and spicules, holotype MNRJ-003. A, gemmule from a top view. B, gemmule from a bottom vIew WIln
foramen in the middle (see also D). C, cross section of gemmule (arrows indicate the thoroidal pneumatic layer). D,foraminal area. E, cross section of the
theca lacking outer and pneumatic layer (see C in the middle area not shown by arrows). F, pneumatic layer (left) and surface (right) of the ring with megas-
cleres of the pseudo-cage (see C top and bottom shown by arrows). G-H, gemmulosclere. J, megascleres (smooth oxeas). la, monstrous tip of megasclere.
Ib, K, spiny oxea of the pseudo-cage around the ring .

Acalle recurvata (Carvalho, 1955; Volkmer-Ribeiro & to T. minuta (potts, 1881a) and T. delicata Bonetto & Ezcurra de
De Rosa-Barbosa, 1972). Sponges settled on leaves or small Drago, 1967.
branches periodically reached by flooding waters. Body shape varied
according to the substrate, ranging from hemispherical on flat sur- STRATOSPONGILLA ANNANDALE, 1909
faces to spherical around tiny branches. Distribution. River lurult
Remarks. The peculiar architecture of the gemmules, unique Synonymy
to this genus, seems to be linked to a displacement of the usual
gemmular layers during gemmulogenesis. Volkmer-Ribeiro (1976) Spongilla (Stratospongilla) Annandale, 1909b: 561.
considers the genus closely related to Trochospongilla and particularly Stratospongilla Penney & Racek 1968: 40.
962 Porifera· Demospongiae • HaploscIerida • Spongillina

,.,.
Fig. 71. Geographic di lribulion of lhe genus Salllmospollgilla.

Type species

Spongilla bombayensis Carter, 1882: 369 (by original

designation).

Diagnosis

Spongillidae with hard to moderately hard consistency.

..r;,""fP'/I", 2lomJayn,,,h . C'f./r.
...21on1llny /s.. _h,~," A¥! /,U"/' .If",..A Y)'.
Choanosomal skeleton vaguely reticulate. Megascleres stout oxeas, ". ~ .,~. # .... •'J/:.,H.~ J ... .••• , .. .,IJ ..

smooth or microspined. Microscleres absent. Gemmules strictly

adherent to the substratum. Gemmular theca tri-layered. Outer Fig. 72. StralOspolIgi/la bOllliJayellsis. lype
gemmular coat of compact spongin with strongyles tangentially
arranged. Pneumatic layer not well defined, appearing as an empty
space with thin trabecules of spongin within gemmuloscleres,
which consist of spined oxeas radially arranged.

Scope and distribution

Five other species: S. sumatrana (Weber, 1890) (Indonesia,

India, Africa); S. indica (Annandale, 1908a) (Thailand, India,
Africa); S. gravelyi (Annandale, 1912) (India); S. clementis
(Annandale, 1909c) (Philippines, China, Japan, tropical W Africa);
S. lanei (?) Racek, 1969 (Australia); S. akanensis (Sasaki, 1970)
(Japan). Restricted to the tropical areas of Oriental, Afrotropical
and Australian (?) Regions.

Fig. 73. Stratospongilla bombayensis, type specimen BMNH 1882.11.

29.3. A-C, spicular tracts of skeletal remains of the type.
Description of type species

Stratospongilla bombayensis (Carter, 1882) (Figs 72-75).

Material examined. Holotype: BMNH 1881.11.29.3 - Description. Encrusting with irregular surface. Colour ?
box 2 dry, small fragment. Other material. BMNH 1882.11.29.3 - Consistency moderately hard but brittle. Oscules inconspicuous.
Bombay, on a stick associated with Eunapius carteri and Dermal membrane supported by abundant spicules. Subdermal
Spongilla alba. cavities small. Ectosomal skeleton is a vague reticulation of
Porifera· Demospongiae • Haplosclerida • Spongillina 963

horizontal spicules immediately below the dennal membrane.

Choanosomal skeleton with vertical spicular fibres . Very sparse
spongin. Megascleres (220-300 X 12-17 fLm) slender, ranging
from straight to feebly curved oxeas, from finely and irregularly
spined to smooth. Microscleres absent. Gemmules singular, vari-
able in size, spherical to ovoid, generally flattened at base, finnly
adhered to the basal spongin plate. Foramen (1 or more) tube-like
on lateral side. Gemmular theca tri-layered. Outer layer of compact
spongin with tangential gemmuloscleres. Pneumatic layer not well
defined appearing as an empty space with thin trabecules of spon-
gin between radially arranged oxeote gemmuloscleres. Inner layer
multilayered. Gemmuloscleres of two types: strongyles (34-44 X
4.6-8 fLm) stout, short and spined. tangentially embedded in the
outer layer; oxeas (60-2.5 fLm) spined and abruptly pointed,
radially arranged between the inner and outer layer.
Ecology. Reported as dense populations adhering to the
lower surface of stones. In shallow standing waters subjected
to dessication where sponges remain as carpets of gemmules.
Gemmules are produced between July and October. Growth notably
slow. Distribution: WIndia (Bombay), South Africa (Natal).
Remarks. Penney & Racek (1968) promoted the subgenus
Stratospongilla (Annandale, 1909b) to genus status. Microscleres.
reported in the vicinity of gemmules in the original description of
Carter (1882). and considered up until now to be true rnicroscleres,
are demonstrated here to be true gemmuloscleres through SEM,
being exclusively present within the gemmular theca having
a radial arrangement. Stratospongilla rousseletii (Kirkpatrick,
1906) and S. schubotzi Weltner, 1913 from South Africa, and
S. gilsoni (Topsent, 1912) from Fiji were moved to Oncosclera by
Volkmer-Ribeiro (1970).

Fig. 74. Stratospongilla bombayensis, spicules and gemmules, type speci- TROCHOSPONGILLA VEJDOVSKY, 1883
men BMNH 1882. 11.29.3. A, outer gemmular theca with foramen (C).
B, cross section of gemmular theca (D). Ea, gemmuloscleres from the outer
layer. Eb, Ec, gemmuloscleres of the radial layer. F, surface of the radial Synonymy
gemmular layer. Ga, gemmuloscleres from the outer layer. Gb, Gc, gemmu-
loscleres of the radial gemmular layer. Gd, megascleres spiny oxeas. Trochospongilla Vejdovsky, 1883: 31.

Fig. 75. Geographic distribution of the genu SlralospOl/gil/a.

964 Porifera· Demospongiae • Haplosclerida • Spongillina

Type species Description. Body shape encrusting. Surface flat, hispid.

Consistency fragile to moderately hard. Colour light yellow to dark
Spongilla erinaceus Lieberkiihn, 1856: 496 (by original brown. Dermal membrane spicular. Subdermal cavities undetected.
designation), an unavailable name, replaced by Trochospongilla Ectosomal skeleton consists of spicules in the dermal membrane.
horrida Weltner, 1893: 12. Choanosomal skeleton anisotropic, pauci- or multi spicular fibres and
tracts more dense at the sponge base. Variable amount of spongin.
Diagnosis Megascleres (170-235 X 11-15 /-Lm) straight to feebly curved
fusiform oxeas covered with stout and sharp spines. Microscleres
Spongillidae with encrusting growth form. Consistency fragile absent. Gemmules in the basal part of the sponge grouped in carpets
to moderately hard. Ectosomal skeleton with spicules in the or singly. Gemmules spherical (475-540 /-Lm) enveloped in a single
dermal membrane. Choanosomal skeleton anisotropic, pauci- or or multi-gemmular pneumatic pseudo-cage armoured by acanthox-
mu1tispicular fibres, more dense at the sponge base. Variable eas. Foramen conical with collar. Gemmular theca monolayered.
amount of spongin. Megascleres acanthoxeas. Microscleres absent. Outer layer and pneumatic layer absent. Inner layer well developed
Gemmules grouped in carpets or singly at the sponge base. sublayered with radially embedded gemmuloscleres in one or two
Gemmules spherical, enveloped in single or multi-gemmular pneu- layers with overlapping proximal rotules. Gemmuloscleres are
matic pseudo-cage, armoured by acanthoxea megascleres. Foramen birotules with a short stout smooth shaft and circular rotules.
present. Gemmular theca monolayered consisting of compact spon- Ecology. Substrata from plants to rocks. Sponges of this
gin with radial gemmuloscleres. Gemmuloscleres birotules with a genus display a fast growth rate, overgrowing other associated
short stout smooth shaft and circular rotules. species. Hibernating life cycle. Distribution. Discontinuous in the
Northern Hemisphere, in cold-temperate regions.
Scope and distribution

Eight other species: T. leidii (Bowerbank, 1863) (E USA, one UMBOROTULA PENNEY & RACEK, 1968
locality); T. paulula (Bowerbank, 1863) (River Amazon); T. penn-
sylvanica (Potts, 1882) (N America, Ireland ?, Scotland ?); Synonymy
T. latouchiana Annandale, 1907 (India, China, SE Australia);
T. philottiana Annandale, 1907 (tropical Asia: India, S China, Umborotula Penney & Racek, 1968: 121.
Philippines, Africa ?); T. minuta (Potts, 1881a) (Argentina, Bolivia,
E Brazil); T. delicata Bonetto & De Drago, 1967 (Argentina,
Brazil); T. petrophila Racek, 1969 (E Australia). Palaearctic,
Nearctic, Neotropical, Australian and Afrotropical ? Regions.

Description of type species

Trochospongilla horrida (Weltner, 1893) (Figs 76-79).

Material examined. Probable holotype: 2MB5091
Wellensee Lake, viii.1898, det. Weltner. Other material. 2MB5093 -
Spandau Havel, 24.vii.1906, on wood with bivalves. DTRG
FW4 - Caorso, Po River, 20.vii.1984, Chiappafreddo. DTRG
FW122 - Lago Maggiore, Pallanza, 5.viii.1970, 1. De Drago.

-2cm

Fig. 77. Trochospongilla horrida, skeletal network of 2MB 5093.

A, sponge fragment on a woody substrate. B, sponge surface, probably in
Fig. 76. Trochospongilla horrida, specimen 2MB 5093. Sponge is an area of contact between different specimen. C, surface close-up.
encrusting on a woody substratum shared with some small bivalves. D-G, close-up views of the choanosomal skeleton.
Porifera' Demospongiae • Haplosclerida • Spongillina 965

Fig. 78. Trochospongilla horrida, gemmules and spicules, specimen 2MB 5091. A, group of gemmules trapped in a common cage, B-C, cross section
of gemmule showing that the pneumatic layer is a character of the cage and not of gemmular theca, D, gemmule in toto, E, foramen, F, gemmular theca
(bottom = sublayered inner layer). G, megascleres (gemmuloscleres are also present). H, gemmulosclere. J, spines of megasclere shaft, rounded termina-
tions are considered as monstrosities.

Type species Scope and distribution

Ephydatia bogorensis Weber, 1890: 33 (by original Monotypic, SE Asia to E Australia.

designation).
Description of type species
Diagnosis
Umborotula bogorensis (Weber, 1890) (Figs 80-83).
Spongillidae with small encrusting growth form. Choanosomal Material examined. Holotype: ZMA 6 slides,
skeleton anisotropic, paucispicular. Megascleres oxeas. Microscleres Buitenzorg, 1889, det. M. Weber. Other material. BMNH
absent. Gemmuloscleres birotules. Gemmules scattered. Gemmular 1901.10.22.1-2 - Ephydatia bogorensis var. blembingia Evans,
theca tri-layered. Pneumatic layers of compact spongin trabecules, 1901, Blembing, Blembing River, peninsular Thailand,
with no chambers. 23. vii.1899.
966 Porifera' Demospongiae • HaploscIerida • Spongillina

Fig. 79. Geographic di lribution of the genu Trochospollgilla .

Fig. 80. A, part of the type of Ephydatia blembingia = Umborotula bogorensis, the Natural History Museum of London; catalogue: 1901: 10:22: I. B, six
slides are the remains of the type material of Umborotula bogorensis preserved in ZMA; catalogue: unregistered.
Porifera· Demospongiae • HapJosclerida • Spongillina 967

Description. Shape circular or irregular small crusts (2-3 cm)

on aquatic plants. Colour light brown to dark green. Consistency
soft, texture loose. Oscules inconspicuous. Surface slightly hispid
due to tufts of spicules. Dermal membrane developed. Oscula
inconspicuous. Ectosomal skeleton consists of spicules tufts.
Choanosomal skeleton paucispicular, anisotropic, large reticulation
with slender horizontal fibres branching freely in the sponge and
stout transverse irregular fibres, ill-defined. Spongin sparse.
Megascleres slender sharply pointed oxeas ranging from
microspined to rarely smooth (200-330 X 8-16/Lm). Microscleres
absent. Gemmules spherical (450-600/Lm) not abundant and scat-
tered in the sponge. Foramen tubular without a true collar, in a
depression surrounded by a thickening of the gemmular wall.
Gemmular theca tri-layered. Outer layer not well developed and
distal rotules of gemmuloscleres are evident. Pneumatic layer of

Fig. 82. Umborotula bogorensis (pro Ephydatia blembingia), gemmules

and spicules, BMNH 1901.10.22.1. A, C, spiny megascieres. B, smooth
Fig. 81. Umborotula bogorensis (pro Ephydatia blembingia), skeletal net- megasclere. D, gemmule. E, foraminal area. F, gemmular outer layer sur-
work, BMNH 1901.10.22.1. A, sponge fragments. B-C, skeletal network face with emerging distal rotule of gemmuloscleres. G, trilayered gemmu-
close-up. D, skeleton with gemmules. lar theca with radial birotules gemmuloscieres. H-J, gemmuloscleres.

Fig. 83. Geographic di Iribulion of the genu UmborOlllla.

968 Porifera· Demospongiae • Haplosclerida • Spongillina

anastomosing trabecules of compact spongin without chambers. Inner Description of type species
layer multilayered. Gemmuloscleres radially embedded in the pneu-
matic layer, with umbrella-shaped rotules in contact with the inner Uruguayella repens (Hinde, 1888) (Figs 84-87).
and outer layer. Gemmuloscleres birotules with spiny shaft (62-82 X Material examined. Holotype: BMNH 1881.7.2.1 - dry frag-
5-8/-Lm). Rotules (22-27/-Lm) with distal surface convex and proxi- ment, River Uruguay, A.R. Mackinnon, vii.1888. Other material.
mal concave, incised margins sometimes with recurved spines. BMNH 1919.10.14.1 - dry on a wood fragment, River Uruguay,
Ecology. Creeping over branched vegetable supports, or South America (no locality specified).
on undersides of floating leaves. Distribution: Malaysia, Java, Description. Encrusting body shape. Colour light lead-
Makassar, Sulawesi, Thailand, China (Soochow), E Australia. brown in dry specimens. Consistency rigid and very hard. Surface
smooth, even. Oscules inconspicuous. Dermal membrane and sub-
dermal cavities not observed. Ectosomal skeleton is an alveolate
URUGUAYELLA BONETTO & EZCURRA meshwork with thick spicular tracts much more closely arranged
DE DRAGO, 1969 (130-400/-Lm) than those of the choanosome. Choanosomal skele-
ton alveolate with paucispicular tracts. Megascleres (150-240 X
Synonymy 20-30 /-Lm) slightly curved strongyles covered entirely by granules.
Microscleres absent. Gemmules single or in small groups (2-5)
Uruguayella Bonetto & Ezcurra de Drago, 1969: 356.

Type species

Uruguaya repens Hinde, 1888: 2 (by original designation).

Diagnosis

Spongillidae with encrusting body shape. Choanosomal

skeleton alveolate with paucispicular tracts. Megascleres
strongyles covered entirely by granules. Microscleres absent.
Gemmules single or in small groups strictly adherent to the substra-
tum enclosed in cages of megascleres. Gemmular theca tri-layered.
Gemmuloscleres birotules radially arranged.

Scope and distribution

Four other species: U. macandrewi (Hinde, 1888) (River

Paraguay, River Parana); U. pygmea (Hinde, 1888) (River Paraguay, 3cm
River Uruguay); U. amazonica (Weltner, 1895) (River Amazon);
U. ringueleti (Bonetto & Ezcurra de Drago, 1969) (upper Parana Fig. 84. Uruguayel/a repens, holotype BMNH 1888.7.2.1. Gemmules
River, River Uruguay). adherent to the substratum are indicated by arrows.

Fig. 85. Uruguayella repens, skeletal network, holotype BMNH 1888.7.2.1. Aa, sponge fragment (top view). Ab, a sponge section (left = surface).
B, ectosomal skeleton. C, choanosomal skeleton.
Porifera· Demospongiae • Haplosclerida • Spongillina 969

Fig. 86. Uruguayella repens, spicules and gemmules, holotype BMNH 1888.7.2.1. A, gemmular cage of megasc1ere. B, gemmular surface and cage.
C, gemmule (cross section). D-E, cross section of gemmular theca with multilayered inner layer and alveolate pneumatic layer. F-H, megasc1eres.
I-J, gemmulosc1eres.

strictly adherent to the substratum enclosed by an irregular cage Inner layer sublayered with adhering proximal rotules. Birotules
(100-600 fLm; mean diameter 500 fLm) of finely granulated very regular in size and shape (16 fLm mean length) with smooth
strongyle megascleres (mean 140-180 fLm). Gemmules subspheri- stout cylindrical shafts (14-15 X 6 fLm). Rotules (16 fLm mean)
cal (50-200 fLm), brownish. Foramen not evident, although in a umbonate smooth circular in outline, with both margins invariably
few gemmules there are one or more slightly raised areas in a lat- incurved in the same direction.
eral position, which may represent apertures. Gemmular theca tri- Ecology. On wood debris in running waters. Frequently
layered with a single radial layer of very regularly and closely associated with Uruguaya corallioides. Distribution. River
arranged birotules. Outer layer smooth covering the distal rotules Uruguay, upper Parana River.
of gemmuloscleres. Pneumatic layer with chambers and thickness Remarks. Other species from South America previously
variable in different areas of the same gemmule from thin to thick. referred to Trochospongilla are now assigned to this genus.
970 Porifera· Demospongiae • Hap!osclerida • Spongillina

Fig. 87. Geographic di uibutioll of the genu UrlIf(flaye/la.

FAMILY LUBOMIRSKIIDAE REZVOI, 1936 reticulate. Ectosomal skeleton ranges from spicule tufts, to notably
developed, up to a thick cortical layer. Spongin abundant. Mega-
Synonymy scleres oxeas to strongyles. Microscleres absent. Gemmules
absent. Larvae apparently produced parthenogenetically.
Lubomirskiidae Rezvoi, 1936. Type genus. Lubomirskia
Dybowsky, 1880. Scope

Three genera: Lubomirskia Dybowsky, 1880; Baikalospongia

Diagnosis Annandale, 1914a; Swartschewskia Makuschok, 1927a.

Spongillina with body shape ranging from encrusting to Distribution

cushion-shaped to branched. Subdermal cavities absent.
Choanosomal skeleton from pauci- to multi spicular, from vague to Hydrographic basin of Lake Baikal, Siberia (Fig. 1).

KEY TO GENERA

(1) Megascleres are typically spiny oxeas; microscleres are absent ..................................................................................... Lubomirskia
Megascleres are typically spiny stout curved strongyles with spines arranged in rosettes ......................................... Swartschewskia
Megascleres are typically smooth strongyles with spiny tips, spiny strongyles or/and spiny oxeas may also be
present ......... ........... ..... ............................ ........... .... ............................ ........................... .......... ................ ................. ... Baikalospongia

LUBOMIRSKlA DYBOWSKY, 1880 rigid to flabby. Anisotropic regular choanosomal skeleton.

Abundant spongin envelopes primary fibres and secondary tracts.
Synonymy Megascleres spined oxeas. Microscleres absent. Gemmules absent.

Spongia sensu Pallas, 1776 (in part): 710. Lubomirskia Scope and distribution
Dybowsky, 1880: 12.
Another two species: L. abietina Swartschewsky 1901;
Type species L. fusifera Soukatschoff, 1895. Hydrographic basin of Lake
Baikal, Siberia.
Spongia baikalensis Pallas, 1776 (by subsequent designation,
Rezvoi et al., 1971: 75). Description of type species

Diagnosis Lubomirskia baikaiensis (Pallas, 1776) (Figs 88-92).

Material examined. Type material: not seen. Other mate-
Lubomirskiidae with body shape ranging from encrusting to rial: BMNH 1932.10.3.1 (dry box FWl) - Lake Baikal, colI. R.
cushion-shaped to branched. Consistency elastic, not fragile, from Kirkpatrick.
Porifera· Demospongiae • Haplosclerida • Spongillina 971

Description. Branching shape with an encrusting base (1 cm 193 X 13 /-Lm). Thin, short, smooth oxeas free in the choanosome
thick) bearing erect (30-60 cm up to 1 m high) dichotomous are young megasc1eres. Microsc1eres absent. Gemmules absent.
branches with rounded apices; diameter of branches is not uniform Ecology. Common on rocks, boulders and wood along the
(1-2 to 3-4 cm thick) and ranges from cylindrical to flattened, fan- entire shoreline from 3-4 m to more than 50 m depth. The optimal
shaped (1 cm thick), larger at the apices (up to 5 cm). Colour bril- habitat with flourishing dense populations occurs in the batymetric
liant green in life, light brown in dry specimens. Consistency range of 5-8 m depth (Kozhov, 1931). A peculiar body shape as a
elastic, not fragile. Body surface hispid due to emerging spicules. long branch with both rounded apices and without encrusting base
Oscules (3-4mm diameter) in small pockets, scattered or more is displayed by non-sessile specimens living free on soft bottom.
rarely in lines on both basal portion and branches, with distances of This morphology appears to be due to a fragmentation process
1-3 cm from each other. Dermal membrane aspicular. Subdermal involving detachment oflong branches from sessile specimens, and
cavities absent. Ectosomal skeleton consists of spicule tufts from therefore represents an asexual reproductive mode. A notably
primary fibres. Choanosomal skeleton regularly anisotropic, diversified benthic 'consortium' of endemic invertebrates and
multi spicular primary fibres and paucispicular secondary tracts; fishes is associated with these sponges.
primary fibres diverging in spicule tufts at the surface. Abundant Remarks. Lubomirskia baikalensis was reported by
spongin envelopes primary fibres and secondary tracts. Arndt (1948) in the Behring Sea and around the Kamcatka
Megasc1eres oxeas, cylindrical to fusiform, uniformly spined with Peninsula.
spines of variable size, rarely smooth (145-233 X 9-18 /-Lm, mean

Fig. 88. Lubomirskia baikalensis, BMNH 1932.10.3.1 (dry box FWl),

Lake Baikal, coll. Kirkpatrick. Fig. 89. Lubomirskia baikalensis, specimens in situ in shallow water.

Fig. 90. Lubomirskia baikalensis, skeletal network, BMNH 1932.10.3.1. A, cross section of a sponge digitation (top = sponge surface). B, megasclere tips
extruding from the sponge surface (lateral view). C, sponge surface (top view). D, multi spicular skeleton trait.
972 Porifera· Demospongiae • Haplosclerida • Spongillina

--
Fig. 91. Lubomirskia baikalensis, skeletal spicules, BMNH 1932.10.3. 1. A, spiny megascleres. B, C, E, monstrous spicules. D , F,close-up of spicule
spination.

Fig. 92. Geographic distribution of the genus Lubomirskia.

BAIKALOSPONGIA ANNANDALE, 1914

Synonymy

Lubomirskia, in part, Dybowsky, 1880: 22. Baikalospongia

Annandale, 1914a: 144. Baikalolepis Makushok, 1927a: 99.

Type species

Lubomirskia bacillifera Dybowsky, 1880: 22 (by subsequent

designation, Rezvoi et aI., 1971: 75).

Diagnosis

Body shape encrusting to globose, to rarely branched with

short anastomosing branches (B. dzhegatajensis). Consistency not
Fig. 93. Baikalospongia bacillifera, BMNH 1976.9.1.1, Lake Baikal, SW
elastic, fragile, ranging from rigid to soft or flabby. Ectosomal comer, dredged between Slindyanka and the mouth of River Angara, shal-
skeleton notably developed as a cortical layer of tangential low water, J7.viii.1976, presented by John Massey-Steward (scale = 10 em).
Porifera' Demospongiae • Haplosclerida • Spongillina 973

Fig. 94. Baika/ospongia bacillifera, skeletal network, BMNH 1976.9.1.1. A, cross section throughout the sponge body (top = surface). B, choanosomal
skeleton. C, skeleton organisation at the sponge surface. D, more details of C. E, more details of B. F, top view of the sponge external surface, oscular area
(modified from Rezvoj, 1936). G, basal membrane.

spicules. Choanosomal skeleton irregularly anisotropic. Mega-

scleres oxeas and strongyles. Spongin variably abundant never
forming a continuous layer on fibres and tracts, more developed at
nodes of spicules.

Scope and distribution

Another two species: B. intermedia Dybowsky, 1880; B. dzhe-

gatajensis Rezvoi, 1927 (Djegataj Kul, Urianhajskaja Region).
Hydrographic basin of Lake Baikal, Siberia.

Description of type species

Lubomirskia bacillifera Dybowsky, 1880 (Figs 93-96).

Fig. 95. Baika/ospongia bacillifera, skeletal network, BMNH 1976.9.1.1.
Material examined. Specimen: BMNH 1976.9.1.1 - Lake A, megascleres. B-D, different views of the spiny tips of megascleres.
Baikal, SW comer, dredged between Slindyanka and the mouth
of River Angara, shallow water, 17.viii.1976, presented by John
Massey-Steward.
Description. Body shape encrusting to globose (5-6cm in are rare. Microscleres absent. Gemmules absent. Embryos are
diameter). Colour green. Consistency not elastic, fragile, rigid. often abundant, but a free swimming larval stage is unknown.
Oscules in small pockets, in regularly scattered groups. Subdermal Ecology. Shallow water from 2-lOm of depth. Some mas-
cavities absent. Basal spongin plate, according to Rezvoi's (1936) sive specimens with a blue-green brilliant colour have been
description, is very thick with growth rings evident in sections due collected at 533 m depth.
to the deposition of spongin in successive layers. Ectosomal skele-
ton notably developed as a cortical layer produced by irregular
spicule tufts at the apices of primaries joined to peripheral second- SWARTSCHEWSKIA MAKUSCHOK, 1927
ary tracts in a polygonal network. Choanosomal skeleton variably
developed, irregularly anisotropic with spicules irregularly Synonymy
arranged in primary multispicular fibres and secondary paucispicu-
lar tracts; primary fibres diverge in spicule tufts at the surface. Lubomirskia, in part, Dybowsky, 1880: 13. Baikalospongia,
Spongin variably abundant, but never forming a continuous layer to in part, Annandale, 1914a: 144. Lubomirskia papyracea var. B
cover all spicules of fibres, more developed at nodes of spicules. Swartschewsky, 1901: 10. Swartschewskia Makuschok, 1927a: 79.
Megascleres strongyles, rarely oxeas (262 X 27 f.1m), with spines Type species. Lubomirskia papyracea var. B Dybowsky,
uniformly distributed or more dense at the apices, smooth spicules 1880: 13 (by subsequent designation, Swartschewsky, 1901: 10).
974 Porifera· Demospongiae • Haplosclerida • Spongillina

Fig. 96. Geographic distribulion of the genus Baikalospollgia .

~"...w ~ !MP'::l'A".... (Ll::J"'.)

r'1'1, 1. .if J.- S
L...k.c. B ,I<.>J
W ~I 1(,-.sL.o..., Ie. 1930)

(]), SLD ""..-Q.<;i .,.,... o'-j i>c.c-t:./o:'.l

~.....,~J IL Jt'OJ~ t=' d""""",~
>I' SO .....4~ ~r~e..r1,"tf:;P\/A J VI,S'''''
Rb I'l"l? lei} a>-.r.-......,."

Fig. 98. Swartschewskia papyracea, skeletal network, BMNH

1997.2.14.2-5. A-B , top view of sponge external surface. C-D, spiny oxeas
lying on the internal sponge body layer. E, internal surface of the sponge.

Description of type species

Fig. 97. Swartschewskia papyracea, BMNH 1997.2.14.2-5, Lake Baikal,
de!. M. Kashow (c. 1930), ID confirmed by S. Efremova.
Swartschewskia papyracea (Dybowsky, 1880) (Figs 97-100).
Material examined. Specimen: BMNH 1997.2.14.2-5-
Lake Baikal, det. M. Kashow (c. 1930) (identification confirmed
Diagnosis by S. Efremova).
Description. Body shape globose. Consistency rigid and
Body shape globose to branched. Ectosomal skeleton hard fragile in the ectosome, soft and flabby in the choanosome. Body
and well developed as a polygonal network of thick tangential spic- surface smooth with an evident alveolar network. Oscules from
ular fibres. Choanosomal skeleton sparsely developed with scarce single at the sponge apex, to rounded in shape occurring in small
spicules irregularly arranged in few weak fibres. Abundant spon- pockets with a peripheral ring variably developed, to grouped at the
gin. Megascleres strongyles, from spiny to smooth. apices. Subdermal cavities absent. Ectosomal skeleton a regular
thick cortical layer of dense tangentially arranged spiCUlar tracts,
Scope and distribution shaped as an alveolar network with one inhalant aperture in each
mesh. Choanosomal skeleton sparsely developed with vague fibres
Monotypic. Hydrographic basin of Lake Baikal, Siberia. or scattered spicules; choanosome is therefore soft and flabby.
Porifera • Demospongiae • Haplosclerida • Spongillina 975

Megascleres short, bent acanthostrongy les with spines (112-150 X FAMILY MALAWISPONGIIDAE FAM. NOV.
13-29IJ.m; mean 103 X 21IJ.m), grouped in rosettes, more dense at
the apices, rarely with smooth surfaces; rare spiny oxeas in the Synonymy
choanosome.
Ecology. This rare species is reported prevalently localised in [Globulospongillinae] Brien, 1973b (nomen nudum).
the area B. Kotov of Lake Baikal, from 2-80m depth (Kozhov, 1931). [Globulospongillidae] Racek, 1974. Type genus. Malawispongia
Brien, 1972.

Diagnosis

Spongillina with body shape ranging from encrusting in

young sponges to globular in well-developed sponges. Consistency
firm, not elastic, notably hard, fragile. Central body cavity.
Alveolate-reticulate skeleton with multi spicular tracts also notably
thick. Megascleres spiny or tuberculated oxeas. Microscleres
absent. Gemmulosc1eres absent. Gemmules absent.

Scope

Five genera: Malawispongia Brien, 1972, Cortispongilla

Annandale, 1918, Ochridaspongia Arndt, 1937, Pachydictyum
Weltner, 1901, Spinospongilla Brien, 1974.

Remarks

Brien (1973b) erected the subfamily [Globulospongillinae]

within the family Spongillidae, including all genera listed above.
Racek (1974) subsequently erected the taxon to family level.
However, there is no genus 'Globulospongilla' , and therefore the
taxon is a nomen nudum and requires a new name.

Distribution

Fig. 99. Swartschewskia papyracea, skeletal network, BMNH 1997.2.

Notably disjunct, in ancient lakes: Kinneret (Middle-East),
14.2-5. A, megascleres. B, spiny shaft of a megasclere. C, piny
s tip of a Malawi (African Rift Valley), Ohrida (E-Europe) and Posso
megasclere. (Sulawesi) (Fig. 1).

Fig. 100. Geographic di Lribution of the genu SlI'orlScilell'skia.

976 Porifera· Demospongiae • HaploscIerida • Spongillina

KEY TO GENERA

(1) Megascleres are oxeas with scattered evident buttons (rarely one tip is rounded); (endemic to Lake Malawi Niassa)
.................................................................................... ................ .................................................................................... Malawispongia
Megascleres are stout smooth oxeas occasionally with a microspined surface; (endemic to Kinneret Lake and Jordan River).
............. .................................... ........................................................ ........... ............................................... ....................... Cortispongilla
Megascleres are oxeas evidencing a very spiny surface (endemic to Ochrid Lake) ................................................... Ochridaspongia
Megascleres are microgranulated oxeas (endemic to Lake Posso, Sulawesi (Celebes» ................................................ Pachydictyum
Megascleres are oxeas with scattered microgranules and big spines (endemic to Tanganyika Lake) .......................... Spinospongi/la

MALAWISPONGIA BRIEN, 1972 spicules tufts from apices of primary tracts on conules. Dermal
membrane transparent spicular. Subdermal cavities well developed
Synonymy as open exhalant canals joining to oscules. Ectosomal skeleton con-
sists of tufts of spicules from primary tracts and irregularly arranged
Ma/awispongia Brien, 1972: 85. tangential megascleres converging to form small conules.
Choanosomal skeleton reticulate multispicular with vertical colum-
Type species nar projections connected by an irregular isotropic reticulation to
form notably dense central axis of lobes with spicules diverging in
Ma/awispongia echinoides Brien, 1972: 82 (by monotypy). tufts toward the apical surfaces. Irregularly scattered tracts connect
lobes. Abundant spongin. Megascleres (190-240 X 4-10 fLm) oxeas
Diagnosis ranging from slender to stout, from straight to slightly curved,
from smooth to tuberculated, with acerate to mucronate or blunt
Malawispongiidae with bush-like body, globular to ovate, and tips. Microscleres and gemmules absent. Larva parenchymella
irregular central body cavity. Choanosomal skeleton reticulate. spongillid-like. Spermatic cysts never found, parthenogenetic (?).
Abundant spongin. Megascleres range from smooth to tuberculated Ecology. In standing waters at 20-30 m depth. On inner face
oxeas. Microscleres absent. Gemmules absent. of bivalves shell (Apatharia sp.). Distribution. Malawi Lake.

Scope and distribution

CORTISPONGILLA ANNANDALE, 1918
Monotypic. Lake Malawi-Niasa (12°09'S, 34°25'E), Africa.
Synonymy
Description of type species
Cortispongiila Annandale, 1918: 66. Nudospongilla (part)
Malawispongia echinoides Brien, 1972: 82 (Figs 101-104). Racek, 1974: 146.
Material examined. Paratype: MRAC 001.427 - Monkey
Bay, Lake Malawi-Niasa, 22-23m, D.H. Eccles. 1O.vii.l971. Type species
Description. Globular to ovate shape (bush-like body) with
2-4 lobes (size 2 X 5-12 X 8cm). Colour brown-grey. Consistency Potamolepis barroisi Topsent, I 892d: 85 (by original
hard in dry specimens. A large oscule in each lobe is connected to an designation).
irregular central cavity. Young sponges devoid of central cavity show
a more irregular lobed body with a single apical oscule devoid of Diagnosis
oscular canals. Surface membraneous and hispid. Macro-hispidation
due to the apices of vertical fibres. Micro-hispidation produced by Malawispongiidae with encrusting to globular-massive body
shape. Body central cavity well developed. Reticulate choanoso-
mal skeleton. Megascleres smooth to granulated oxeas.
Microscleres absent. Gemmules absent.

Scope and distribution

Monotypic. Lake Kinneret and River Jordan, Syria? Palestine.

Description of type species

Cortispongilla barroisi (Topsent, 1892d) (Figs 105-108).

Material examined. Holotype: MNHN DT3302 - dry, Lac
de Tiberiade, Syria, Th. Barrois, v.l890, by dredge. Other material.
BMNH 1913.7.31.4C-13.I1I.C - alcohol, Lake Tiberias, del. N.
Annandale. BMNH 1913.7.31.5 - dry, Channel of River Jordan
near Semakh, Lake Tiberias, Palestine, del. N. Annandale. BMNH
Fig. 101. Malawispongia echinoides, paratype, MRAC 1427. 1910.1.1.480 - Norman Collection, Lake Tiberias. BMNH
Porifera· Demospongiae • HapJosclerida • SpongilIina 977

...
Fig. 102. Ma/awispongia echinoides, skeletal network, paratype MRAC 1427. A, a sponge section (top = surface). B, cross section of a columnar projec-
tion (top = surface). C, multispicular connections among columnar projections. D-E, multi spicular tracts cemented by spongin. F, conules. G, spicule tuft
at the sponge surface.

A
W -. 50 ~m - '.
..... .
.

. .

Fig. 103. Ma/awispongia echinoides, spicules, paratype MRAC 1427. A, slim smooth oxea. E , tuberculate oxeas. C-E, tips of skeletal megasc\eres.
978 Porifera· Demospongiae • Haplosclerida • Spongillina

Fig. 104. Geographic dislribu(ion of (he gcnus Ma/all'ispollgia.

-2cm

I
I>J
o
3

Fig. 105. Cortispongilla barroisi, A, type material and originallabcls MNHN DT3302. B, a specimen from Dendy collection BMNH 25.11.1.591.

1914.11.24.24 (ZEV 5213.7) - Channel of River Jordan, Lake upper and lower surfaces, respectively. Oscule single or 2-3 main
Tiberias, Palestine. BMNH 1925.11.1.591 - Dendy Collection, oscules with numerous subsidiary oscules with superficial exha-
SW comer of Lake Tiberias. lant horizontal radiating canals covered by the dermal membrane.
Description. Sponge encrusting to globular-massive Central body cavity well developed, cylindrical, almost symmetri-
shaped (size 3.5 X 2.3 X 2.1-8.5 X 7.5 X 6cm). Compact hard cal, arising from the convergence of canals, leading directly into a
fragile consistency. Colour dull grey to glaucous green. Surface large main oscule or oscules. Ectosomal skeleton consists of tan-
uniformly membraneous, rough but not hispid. Dermal membrane gential spicules in the membrane and peripheral thickening of
spicular. Some large specimens have large and small pores on the fibres. Choanosomal skeleton alveolate (meshes 500-700 j.l.m),
Porifera· Demospongiae • Haplosclerida • Spongillina 979

Fig. 106. Cortispongilla barroisi, skeletal network, type material MNHN DT 3302. Aa, basal spongin plate. Ab, internal view in cross section. Ac, sponge
surface. B, sponge surface. C, choanosomal skeleton.

Fig. 107. Cortispongilla barroisi, spicular traits. B-E, BMNH 19\3.7.31.4C. A, F, BMNH 1914.11.24.24. A, B, F, different oxeas. C, a monster fusion
between spicules. D, the most common oxea tip. E, granulated surface of an oxea.

multi spicular (6-7) primary tracts and paucispicular secondary shallow waters subjected to the action of periodical heavy storms,
tracts more dense and thick toward the surface. Spongin sparse, to deep waters of the lake also subjected to considerable currents
present exclusively between crossing fibres and in the uneven and on substrate devoid of silt in the Jordan channel that traverses
basal plate at the surface of encrusted pebbles, with scattered the lake. Sponges settle on the lower surface of stones, on living
oxeas or groups of parallel oxeas. Megascleres stout oxeas Unio sp., or encrusting on pebbles during growth to become mas-
(180-370 X 30-33 fLm) from straight to slightly curved, from sive specimens. Distribution. Southern Lake Kinneret (Tiberiade),
smooth to granulated with evident axial canal and tips abruptly River Jordan, Israel.
pointed. Slender acerate oxeas are rare. Microscleres absent. Remarks. This species was initially included in the subfam-
Gemmules absent. ily Potamolepidinae. Topsent (l892d) noted its similarity to
Ecology. The original population was reported as being Petrosia and Reniera; Annandale (1918) underlined its similarity
very dense (Annandale, 1918), although it was restricted to a small to Lubomirskia papyracea, Potamolepis and Uruguaya; and
area (two square miles) of the Jordan canal. Until present records de Laubenfels (1936a) remarked on its notable architectural
the species was considered to have disappeared. Sponges are from similarity with Pachydictyum Weltner, 1901.
980 Porifera' Demospongiae • HaploscIerida • Spongillina

Fig. 108. Geographic dislribulion or the genus CortispoJlgilla.

Fig. 110. Ochridaspongia rotunda, was very abundant in the Lake Ochrid
in the 1950s and many specimens were collected by the nets of professional
fishers (figure modified from S. Hadzisce, 1954, Rec. Trav. Stat. Hidrobiol.
Ohrid, W 1--6).

Diagnosis

Fig. 109. Ochridaspongia rotunda, type material and original labels, 2MB Malawispongiidae with body shape globular to massive.
POR9337. Consistency very resilient, very hard. Single elliptical body cavity
and single osculum. Choanosomal skeleton brittle, anisotropic.
Ectosomal skeleton hard (up to 2 mm) due to the branching and
dense primaries. Megascleres oxeas, smooth to spined.
OCHRIDASPONGIA ARNDT, 1937 Microscleres absent. Gemmules absent.

Synonymy Scope and distribution

Ochridaspongia Arndt, 1937: 660. One other species: O. interlithonis Arndt, 1937. Lake Ochrid,
Macedonia.
Type species
Description of type species
Ochridaspongia rotunda Arndt, 1937: 660 (by original
designation). Ochridaspongia rotunda Amdt, 1937 (Figs 109-113).
Porifera' Demospongiae • Haplosclerida • Spongillina 981

Fig. 111. Ochridaspongia rotunda, skeletal network, type material 2MB 9337. A, surface view. B, ectosomal skeleton (top view). C, skeletal arrangement
in cross section. D, choanosomal skeleton. E, spicule tuft at the sponge surface. F, muitispicular tract.

Fig. 112. Ochridaspongia rotunda, spiculation, type material 2MB 9337. A, spiny oxeas. B, variations in spine densities. C, spicule tip.

Material examined. Holotype: 2MB POR9337 - Lake Colour yellowish-brownish, both in life or dry. Consistency very
Ochrid (41°03'N, 20 43'E), by dredge, SOm depth, R. Kenk,
0
resistent, very hard and brittle in dry specimens. Surface velvet-
12.ix.193S. Paratypes: NZMW not seen. like or rough due to the emerging spicule tufts of primary tracts.
Description. Body shape ranges from globular to massive, Dermal membrane supported by few very thin, straight or slightly
with column-, cone- or top-like shape (7.S X 4.S-13 X 7.S em). curved smooth oxeas. Subdermal cavities present on the upper and
982 Porifera· Demospongiae • HapJosclerida • Spongillina

Fig. 113. Geographic distri bution of the genu Och,.ida~pOllg;a .

lateral body sides. Oscules (1-2) in the middle of the upper side
large (8-18mm in diameter) circular due to the fusion of several
oscules (10-100) into a single elliptical body cavity. Basal spongin
plate well developed in specimens living on pebbles. Skeletal
architecture with radial primaries originating from the base up to
the sponge surface. Ectosomal skeleton hard (up to 2 mm) as tan-
gential oxeas and dense branching of primaries (70 f.Lm thick).
Choanosomal skeleton anisotropic with ascending muitispicular
primary tracts gradually becoming more thin and diverging in tufts
toward the surface and uni- to paucispicular (3-5) irregular second-
ary tracts. Spongin very sparse. Megascleres oxeas (175-367/
5-23 /-Lm), ranging from slender to stout, from straight to slightly
curved, from smooth to spined mainly towards the ends; acerate Fig. 114. Pachydictyum globosum, type material and original labels 2MB
tips. Spherical swellings occur in some oxeas. Oxeas of fibre tufts 3003. The single half-specimen is shown from tbe internal (A) and external
(235-279 X 13 f.Lm), ectosomal oxeas (175-220 X 3/-Lm), choano- (B) surfaces.
somal oxeas (264-322 X 13-19 /-Lm). Microscleres absent.
Gemmules absent. Larvae egg-shaped 100-200 f.Lm in diameter Type species
(width anterior 290 f.Lm, posterior 150/-Lm), bearing very thin
smooth oxeas. Pachydictyum globosum Weltner, 1901: 188 (by monotypy).
Ecology. Eastern shore of the lake, 30-70m depth, on
muddy bottoms or covered by bivalve shells or small pebbles, also Diagnosis
in strong currents. Sponges englobe pebbles or shells in the basal
area. Ochridaspongia interlithonis lives near the shore 1.5-2 m Malawispongiidae with globular body. Consistency hard and
between plants (Vallisneria, Miriophyllum), on wooden pillars and brittle. Body cavity elliptical open directly through the single oscu-
pebbles with Dreissena. Distribution. Lake Ochrid. lum. Choanosomal skeleton reticulate with notably thick primary
Remarks. The single preoscular cavity can be considered fibres. Sparse spongin. Megascleres oxeas, styles, tylostyles, ranging
homologous to the central cavity of C. barroisi and P. globosum. The from smooth to granular. Microscleres absent. Gemmules absent.
local trivial name of this species is 'shetsherpare' or 'shekerpare', a
Turkish word that means sweet cake. Scope and distribution

Monotypic. Lake Posso (01°55'S, 1200 37'E) Sulawesi

PACHYDlCTYUM WELTNER, 1901 (Celebes).

Synonymy Description of type species

Pachydictyum Weltner, 1901: 188. Pachydictyum globosum Weltner, 1901 (Figs 114-117).
Porifera· Demospongiae • Haplosclerida • Spongillina 983

Fig. 115. Pachydictyum globosum, skeletal network, type material 2MB 3003. A, cross section from the sponge surface (left) throughout the choanosome
(right). B, sponge surface (top view). C, ectosomal skeleton. D, choanosomal skeleton.

Fig. 116. Pachydictyum globosum, spicules, type material 2MB 3003. A, megascleres oxeas. B, spiny and smooth oxeas. C, megasclere tips. D, granules
at the megasclere surface.

Material examined. Holotype: 2MB 3003A-SE36, 2MB or light brown in alcohol. Consistency hard and brittle. Rough sur-
3003B-SE36 - Lake Posso, Sulawesi (Celebes), P. & F. Sarasin, face with protruding ends of primary tracts. Single osculum con-
ii.1895. nected through a broad barrel-shaped canal with the irregular central
Description. Body shape spherical, oval to club-shaped (size cavity within the sponge body. Thickness of body wall ranges from
3.1-3.3 X 2-2.5 cm). Colour bright light green in life, turned to grey 3.5-10 nun related to the size of inner cavity. Basal spongin plate not
984 Porifera· Demospongiae • Haplosclerida • Spongillina

Fig. 11 7. Geograph i di. tribution of the genus PacilydiclYIIIII .

observed. Dennal membrane not observed. Subdennal cavities evi-

dent. Ectosomal skeleton with spicule tufts at the apices of primary
tracts. Choanosomal skeleton irregularly anisotropic, reticulate in
some areas with primary tracts multi spicular (6--8 to 35--40) diverg-
ing in tufts toward both apical and basal (central cavity) surfaces,
irregular secondary tracts pauci- (1-2) to multispicular (6--10).
Sparse spongin as thin layers around spicules and tracts, with abun-
dant foreign material. Choanocyte chambers range from 20-24 fLm.
Megascleres of various types: stout smooth, straight to slightly curved,
gradually tapering sharp-pointed oxeas with evident axial canal and
very variable in size (slender 374 X 25 fLm, stout 340-410 X
24-32 fLm; mean 360 X25 fLm). Few spicules uniformly thick with
rounded tips (320-28 fLm). Few smooth styles and tylostyles are also
present with common silica spherules. Malfonned tips are frequent Fig. 118. Spinospongilla polli, holotype MRAC 1431.
occurring as bulb-like tips. Microscleres absent. Gemmules absent.
Larvae abundant and scattered in two specimens, on three described
females with numerous cleavage stages and eggs. Oval larvae (750 X Definition
520 fLm) bearing small slender smooth oxeas (230-6 fLm) in their
posterior ends. Malawispongiidae with encrusting body shape, surface with
Ecology. Shallow waters at the eastern shore of Lake Posso. irregularly scattered oscules. Consistency hard. Ectosomal skele-
Sponges settled on shells of Melania kuli Sarasin, M. centaurus ton shaped as conules and irregular uni- or bispicular meshes
Sarasin and Miratesta celebensis var. robusta Sarasin, with the tangential to the surface. Alveolate isotropic multispicular
large osculum directed away from the underside of the living choanosomal skeleton with rare scattered smaller oxeas. Scarce
snails. Shells surfaces showed signs of corrosion in areas of contact spongin. Megascleres oxeas ranging from slender to stout, from
with the sponge. Distribution. Found only in Sulawesi, Indonesia. straight to slightly curved, from smooth to spined and granulated;
tips range from acerate to blunt. Microscleres and gemmules
absent.
SPINOSPONGILLA BRIEN. 1974
Scope and distribution
Synonymy
Monotypic. Lake Tanganyika (06°24'N, 29°38'E) Zaire,
Spinospongilla Brien, 1974: 589. Afrotropical Region.

Type species Description of type species

Spinospongilla polli Brien, 1974: 589 (by monotypy). Spinospongilla polli Brien, 1974 (Figs 118-121).
Porifera' Demospongiae • Haplosclerida • Spongillina 985

Fig. 119. Spinospongilla polli, skeletal network, holotype MRAC 1431, A, cross section of a conule. B, detail of A at the conule base. C, choanosomal
skeleton. D, multispicular axis of a connie. E, choanosomal tract. F, surface hispidation (lateral view). G, sponge surface (top view). H, detail of G (top
view). I, aquiferous openings (modified from Brien, 1974). J, scheme of surface and a cross section (modified from Brien, 1974).

Material examined. Holotype: MRAC 1431, Lake parenchymula (Brien, 1974: 602). Spermatic cysts never found,
Tanganyika, Usumbura, Zaire, v.1973, leg. M. Brichard, by parthenogenetic (?).
diving. Ecology. Unknown.
Description. Body shape encrusting 2-5 mm thick. Colour
buff, green (surface), white-grey (inner part). Consistency hard.
Surface conulose; conules 3-4 mm height. Oscules (2-4 mm) irreg- FAMILY METANllDAE VOLKMER· RIBEIRO, 1986
ularly scattered sometimes crater-like. Dermal membrane spicular.
Basal plate well developed, brown. Ectosomal skeleton irregular Synonymy
network of uni- or bispicular meshes tangential to the surface.
Choanosomal skeleton alveolate isotropic multispicular with rare Metaniidae Volkmer-Ribeiro, 1986: 493. Type genus. Metania
scattered smaller oxeas (128 X 2.3 f1m) in the original description; Gray, 1867a.
at the base of conules fibres converge in vertical thicker tracts.
Sparse spongin. Megascleres oxeas (160-208 X 11-12f1m) rang- Definition
ing from slender to stout, from straight to slightly curved, from
smooth to spined and granulated; tips range from acerate to blunt. Spongillina with body shape encrusting, massive, bulbose,
Microscleres absent. Gemmules absent. Larva spongillid-like globular. Surface smooth, hispid to conulose. Consistency from
986 Porifera· Demospongiae • HapJoscIerida • Spongillina

~ - - ~

----
. 10~m .

---
'.'

Fig. 120. Spinospongilla polli, spicules, holotype MRAC 1431. A, megascleres oxeas. B-D, close-up of megasclere tips with different granulation and
spination. E, monstrosity of a megasclere shaft.

ig. 121. Geographic distribulion of the gen u Spillospollgilla.

fragile to very hard. Ectosomal skeleton consists of tangential Scope

spicules in the dermal membrane or spicule tufts from emerging
fibres. Choanosomal skeleton alveolate-reticulate irregular. Five genera: Metania, Acalle, Drulia, Houssayella,
Megascleres are oxeas to strongyles, from smooth to spiny. Corvomeyenia.
Microscleres, when present, are acanthostrongyles, acanthoxeas,
asters, pseudobirotules. Gemmules subspherical to oval, with or Distribution
without cage. Foramen present. Gemmular theca usually tri-
layered with radially arranged gemmuloscleres. Gemmuloscleres Nearctic, Neotropical, Afrotropical, Oriental and Australian
boletiform (tubelliform), parmuliform, pseudobirotules. Regions (Fig. 1).
Porifera· Demospongiae • Haplosclerida • Spongillina 987

KEY TO GENERA

(i) Pannuliform spicules (gemmuloscleres) are present; squat and smooth oxeas as megascleres; microscleres are acanthoxeas ..... Drulia
Absence of pannuliform spicules (gemmuloscleres) .. ...... .......... ............ .......... ...... ................ ..................... ...................... ................. 2
(2) Tubelliform spicules (gemmuloscleres) are present ............................................................................................................................ 3
Absence of tubelliform spicules as gemmuloscleres ... ................. ............... ............................ .............. ............ ........... ........... ............ 4
(3) Gemmuloscleres show a single tubelliform morphology, radially arranged with the larger rotule embedded in the inner layer of the
gemmular theca; microscleres are strongyles from smooth to spiny; megascleres are oxeas and strongyles, smooth and
spiny ......................................................................................................................................................................................... Metania
Gemmuloscleres of two different morphs: rare tubelliform, arranged radially on the gemmular theca around the foramen; and pseudo-
birotules radially arranged inside the gemmular theca; microscleres are absent; megascleres are few smooth oxeas and abundant
smooth or microspined strongyles .............................................................................................................................................. Acalle
(4) Gemmuloscleres are pseudobirotules with a long slim smooth shaft radially arranged; megascleres are long fusiform oxeas showing
rare spines or tubercles; microscleres are pseudobirotules with a short smooth shaft .................................................. Corvomeyenia
Gemmuloscleres are birotules with a short shaft embedded radially inside the gemmular theca; megascleres are acanthostrongyles
associated with spiny and smooth oxeas; microscleres are acanthostrongyles and acanthoxeas and exclusive 'aster-like' spicules
showing many rays ............................................................................................................................................................ Houssayella

METANIA GRAY, 1867

Synonymy

Metania Gray, i867a: 551. Tubella Carter, i88ic: 96.

Type species

Spongilla retieulata Bowerbank, 1863: 455 (by subsequent

designation; Penney & Racek, 1968: 147).

Diagnosis

Metaniidae with body shape bulbous to massive. Consistency

hard and brittle. Choanosomal skeleton alveolate with multispicular Fig. 122. Metania reticulata, specimen 2MB 3735.

Fig. 123. Metania reticulata, skeletal network, specimen 2MB 3735. A, surface with conules and ectosomal tangential skeleton. B, openings of the aquif-
erous system at the sponge surface. C, ovoidal gemmules within the skeletal meshwork (cross section, left = surface). D, choanosomal skeleton with two
gemmules. E, gemmule with an evident foramen.
988 Porifera· Demospongiae • Haplosclerida • Spongillina

Fig. 124. Metania reticulata, spicules and gemmules, A, E, Bowerbank collection BMNH BK 1435. B-D, F-J, specimen 2MB 3735. A, gemmular cage.
B, gemmule. C, foramen. D, gemmular outer surface. E, gemmuloscieres embedded in the gemmular theca. F, trilayered theca. G, multilayered inner layer.
H, parmuliform gemmulosc1eres. I, microsclere. J, megascieres.

tracts. Spongin sparse. Megasc1eres oxeas to strongyles. Micro- Amazonian Basin); M. ovogel1UJta Stanisic, 1979 (Australia);
sc1eres acanthostrongy1es (acanthoxeas?) to pseudobirotules M. brieni Burton, 1938 (Congo Basin); M. innominata Burton, 1938
variably bent. Gemmules ovoid, scattered or rarely at the sponge (SE Africa); M. rhodesiana Burton, 1938 (SE Africa); M. vanryni
base, enclosed in a variably developed cage of spiny megasc1eres Brien, 1968b (Congo Basin); M. godeauxi Brien, 1968b (Central
ranging from oxeas to strongyles. Theca with three layers and gem- Africa); M. schoutedeni Burton, 1938 (Congo Basin); M. kiliani
mulosc1eres radially embedded. Pneumatic layer as trabecules Volkmer-Ribeiro & Costa, 1992 (Neotropical Region); M. pottsi
(fibres) of spongin. Gemmuloscleres boletiform. (Weltner, 1895) (Congo Basin, Congo, Angola); M. melloleitaoi
(Machado, 1945) (Central Brazil). Distribution. Tropical rain-forests
Scope and distribution of Neotropical, Afrotropical, Oriental and Australian Regions.

Fifteen other species: M. vesparia (von Martens, 1868) Description of type species
(Borneo, Indonesia, Central Africa); M. spinata (Carter, 1881c)
(Amazon Basin); M. vesparioides (Annandale, 1908b) (Burma); Metania reticulata (Bowerbank, 1863) (Figs 122-125).
M. subtilis Volkmer-Ribeiro, 1979 (South America, Amazonian Material examined. Holotype:? BMNH BK1435 - Dark
Basin); M. fittkaui Volkmer-Ribeiro, 1979 (South America, Ygapos in virgin forest, margins of Amazon, Villa Nova,
Porifera· Demospongiae • Haplosclerida • Spongillina 989

Fig. J 25. Geographic distribution of the genus Melal/ia.

20.xi.1854, coli. H. Bates, Bowerbank Collection. Other material. Remarks. The systematics and biogeography of the genus
2MB3735 - Rio Negro, Manaus, colI. Retzloff. was comprehensively studied by Volkmer-Ribeiro (1979, 1984,
Description (most measurements from Volkmer-Ribeiro, 1986), Volkmer-Ribeiro & Costa (1993), da Silva & Volkmer-
1986). Body shape bulbous to massive, often honey-comb shaped. Ribeiro (1998).
Colour from dark grey to dark brown in dry specimens. Consistency The type species of Tubella Carter, 1881c, Spongilla paulata
hard and brittle. Surface reticulate and irregularly conulose due to Bowerbank, 1863: 453 (by subsequent designation, de Laubenfels,
the arrangement of fibres in superficial tracts. Oscules conspicuous 1936a: 37), is possibly a misspelling by Gray (1867a) for Spongilla
and grouped within conules. Dermal membrane supported by paulula Bowerbank, 1863: 453. If the type species designation
conules. Subdermal cavities not observed. Ectosomal skeleton with is correct, then Tubella might become a senior synonym of
spicules grouped in conules. Choanosomal skeleton alveolate with Trochospongilla, because S. paulula is assigned to Trochospongilla
multispicular tracts with meshes increasing from the base to sur- Vejdovsky, 1883 and not to Metania Gray, 1867a, while at the same
face. Spongin sparse. Megascleres cylindrical (106-245 X 11-36 time Tubella is declared a junior synonym of Metania by Penney &
/-Lm) from stout smooth oxeas to strongyles. Microscleres acan- Racek (1968). This proposed synonymy by Penney & Racek is
thostrongyles (acanthoxeas ?) (43-103 X 3-10 /-Lm) with large maintained here in view of the confusion over the type species and
lanceolate spines at the center and granules at the extremities. its subsequent designation.
Gemmules, scattered in the skeletal network or rarely at the sponge
base, enclosed in a variably developed cage of spiny megascleres
ranging from oxeas to strongyles. Gemmular shape ovoidal
(391-588 /-Lm). Foramen with a well-developed tube-shaped collar ACALLE GRAY, 1867
of variable length, distinctly projected above the gemmular sur-
face, ornamented by gemmuloscleres at the basis and with Synonymy
an undulated border. Theca with three layers. Outer layer well
developed. Pneumatic layer trabeculated by spongin fibres. Acalle Gray, 1867a: 551.
Inner layer of compact multilayered spongin. Gemmuloscleres
radially embedded in the theca in one layer with boletiform rotule Type species
in the inner layer and the knoblike pseudo-rotule emerging
from the outer layer. Gemmuloscleres boletiform (22-38 X 2-6 Spongilla recurvata Bowerbank, 1863: 456 (by original
/-lm total length; 16-20/-lm diameter rotule). Shafts with variable designation).
length of spines (2-4 /-lm). Proximal rotule large, stout, polygonal,
with curved, undulated margins. Distal pseudo-rotule knob-like Diagnosis
(7/-lm) or aster-like shaped, with a few recurved, scattered spines
or hooks, or approaching to a true rotule with marginal curved Metaniidae with encrusting growth form. Consistency of dry
spines. sponge fragile. Ectosomal skeleton with a notably dense network
Ecology. On overhanging branches of seasonally submerged of strongyles. Choanosomal skeleton alveolate with paucispicular
trees in periodically flooded habitats. Distribution. Brazilian and (strongyles) tracts. Sparse spongin. Megascleres strongyles to large
Venezuelan Amazonia, in the tropical rain forest realm. oxeas. Microscleres absent. Gemmules subspherical at the base
990 Porifera· Demospongiae • Haplosclerida • Spongillina

of the sponge. Gemmular theca tri-layered. Gemmuloscleres of Choanosomal skeleton alveolate with paucispicular tracts. Sparse
two types: pseudobirotules and tubelliform. spongin. Megascleres stout smooth or rnicrospined abundant
strongyles (120-170 X 14-18/Lm) and few smooth oxeas (350 X
Scope and distribution 20/Lm) of unknown distribution. Microscleres absent. Gemmules
subspherical (350-420/Lm) in one stratum at the base of the sponge,
Monotypic. Neotropical Region. with the foramen of the greater part of them directed downwards.
Foramen is a short tubule with collar surrounded by distal pseudo-
Description of type species rotules of tubelliform gemmuloscleres. Gemmular theca trilayered.
Well-developed outer layer with emerging gemmuloscleres
Acalle recurvata (Bowerbank, 1863) (Figs 126-129). pseudobirotules. Thick pneumatic layer of small chambers with
Material examined. Holotype: BMNH 1931.4.23.3- boletiform (tubelliform) gemmuloscleres radially embedded with
fragment, dry box 6, River Amazon, Villa Nova, colI. H. Bates, large rotules placed at the inner layer. Inner layer with 3--4 sublayers
20.xi.1854. of compact spongin. Gemmuloscleres of two types: pseudobirotules
Description. Sponge consists of thin crusts. Colour light (18-35 X 5-7/Lm) with cylindrical shaft and hooks of umbonate
brown in dry specimens. Consistency of dry sponge fragile, pseudo-rotules stout and notably recurved bearing rnicrospines at
crumbling. Surface even, smooth. Oscules numerous scattered their apices; and tubelliform gemmuloscleres (30-33 X 1-5/Lm) with
inconspicuous simple and of variable size. Dermal membrane thin. the proximal large irregularly circular flat rotule (15-16/Lm) with
Ectosomal skeleton is a notably dense network of strongyles. entire margin supporting a smooth shaft decreasing in thickness
toward the distal end shaped as umbonate knob-like rotule with few
teeth.
2cm
Ecology. In rivers on small stems of trees periodically sub-
merged. Associated with Metania reticulata and Satumospongilla
carvalhoi. In dark Ygapos (dark brown waters) in virgin forest,
margins of Amazon. Distribution. South America Amazon River,
Beni River (Bolivia).

CORVOMEYENIA WELTNER, 1913

Synonymy

Fig. 126. Acalle recurvata, type material and original labels BMNH Corvomeyenia Weltner, 1913: 480. Parameyenia Jewell,
1931.4.23.2. 1952: 455.

Fig. 127. Acalle recurvata, skeletal network, type material BMNH 1931.4.23.2. A-B, sponge surface (top views). C, cross section (top=surface).
D, choanosomal skeleton.
Porifera· Demospongiae • Haplosclerida • Spongillina 991

Fig. 128. Acalle recurvata, spicular and gemmular traits, type material BMNH 1931.4.23.2. A-B, megascieres. C, tubelliform gemmulosciere.
D, gemmule. E, foramen. P, three layered gemmular theca. G, outer layer at the gemmular surface. H-I, gemmuloscieres.

Type species or tuberculate. Microscleres pseudobirotules. Gemmules

subspherical, single or grouped enclosed in cages of mega-
Meyenia everetti Mills, 1884: 146 (by original designation). scleres. Gemmular theca tri-Iayered. Gemmuloscleres
pseudobirotules.
Diagnosis
Scope and distribution
Metaniidae with encrusting growth form. Anisotropic
slightly reticulated choanosomal skeleton with multispicular pri- One other species. C. carolinensis Harrison, 1971 (South
mary fibres and irregular secondary paucispicular tracts. Sparse Carolina), NE USA (Massachusetts), S Canada (Nova Scotia,
spongin. Megascleres oxeas range from smooth to rarely spined Quebec, Newfoundland).
992 Porifera· Demospongiae • Haplosclerida • Spongillina

. ig. 129. Geographic di lribulion of the genus ACldle.

1 2cmr------~
-"<5('l
·-y'dA:,..- t-s
~7~f?e/7/~ ~/B/,C -/-:

Fig. 130. Corvomeyenia everetti, topotype and original labels ANSP P04594.
Fig. 131. Corvomeyenia everetti, skeletal network, topotype ANSP
P04594. A-B, gemmular cages. C, surface hispidation in cross section.
Description of type species D, choanosomal skeleton. E, microsc1eres in the choanosomal matrix.

Corvomeyenia everetti (Mills, 1884) (Figs 130-133).

Material examined. Topotype: ANSP P04594, box shafts and pseudo-rotules dome-shaped (3-7 fLm) ornamented
SO--Gilder Pond, Mt. Everett, Massachusetts, A. Berklin, 23.ii.l986. with 5-8 recurved hooks. Gemmules subspherical (480-530 fLm)
Description. Body shape consists of small thin crusts often single or grouped, each enclosed in a loose cage of mega-
with slender finger-like projections (2 mm). Consistency very soft scleres (710-902 fLm). Shape of foramen not evident. Gemmular
and fragile. Colour emerald green to brown in life. Surface hispid theca tti-layered. Outer layer with emerging distal pseudo-rotules
due to spicule tufts. Oscules not conspicuous. Dermal membrane of gemmuloscleres. Pneumatic layer, not recorded in the present
spicular, closely adherent. Subdermal cavities absent. Ectosomal material, with minute and regular chambers. Inner layer multilay-
skeleton with scattered single spicules. Choanosomal skeleton ered (up to 40 fLm). Gemmulosc1eres radially arranged in a single
anisotropic, slightly reticulate, with multispicular primary fibres series adhering to the inner layer with proximal rotules.
and irregular secondary paucispicular tracts. Sparse spongin. Gemmuloscleres pseudobirotules with straight slender smooth
Megascleres slender feebly curved fusiform oxeas (143-285 X shafts (42-72 X 3-5 fLm), pseudo-rotules (16-19 fLm) with up to 10
3.5-14 fLm) ranging from smooth to sparsely spined or tuberculate, recurved hooks.
with tips gradually or more abruptly pointed. Microscleres (16-26 X Ecology. Moderately acidic (pH 5-6.6) lentic waters, in
1-3.5 fLm) pseudobirotules, with straight to bent slender smooth transparent water, low in calcium (O.4mgll) (Jewell, 1935;
Porifera· Demospongiae • Haplosclerida • Spongillina 993

Fig. 132. Corvomeyenia everetli, spicular and gemmular characters, topotype ANSP P04594. Aa, megasc\ere. Ab, microsclere. Ac, gemmulosc\ere.
B, oxea with scattered spines. C, gemmule with remains of spicular cage. D, gemmule surface. E, cross section of gemmular theca. F, foramen area.
0, gemmulosc\ere. H, microsclere. I, pseudorotule of a gemmulosc\ere.

Harrison, 1974). Very abundant in shallow water on submerged the original material of the type species S. discoides consisted
bushes, water-weeds, grass and wood debris. In symbiosis with of spicules of both Corvomeyenia everetti and Anheteromeyenia
green algae. Often associated with Eunapius mackayi and ryderi. Thus, S. disco ides appears to be a 'chimaera' and conse-
Trochospongilla pennsylvanica. Distribution. NE USA, S Canada. quently the genus Parameyenia is based on faulty characters (but
Remarks. Parameyenia Jewell, 1952, with type species not a nomen nudum as supposed by Penney & Racek). According
Spong ilia discoides Penney, 1933 (by original designation), was to ICZN Article 17.1 the name is available, but needs to be fixed.
considered to be a junior synonym of Heteromeyenia ryderi (Potts, Penney & Racek (1968) say S. disco ides is a synonym of A. ryderi
1882) (see Penney & Racek, 1968: 125), and therefore an objective (and thus a synonym of Racekiela), but it is equally a synonym of
junior synonym of Racekiela. However, Penney & Racek (1968) C. everetti since material of that species is also present in the type
also declared Parameyenia a nomen nudum, and also a junior syn- material, and consequently we propose here this latter course of
onym of Corvomeyenia. This confusing situation arose because action, with Parameyenia a synonym of Corvomeyenia Weltner,
994 Porifera· Demospongiae • Haplosclerida • SpongiUina

Fig. 133. Geographic d i lrihulion of the genu Con'Olll eyenia.

1913, which avoids the problem of semonty of the names

Parameyenia Jewell, 1952 over Racekiela Bass & Volkmer-
Ribeiro, 1998.

DRUUA GRAY, 1867

Synonymy

Drulia Gray, 1867a: 552. Parmula Carter, 1881c: 99.

Type species

Spongilla browni Bowerbank, 1863: 457 (N.B. De

Laubenfels, 1936a cites Bowerbank, 1858: 315) (by subsequent
designation; de Laubenfels, 1936a: 36).
Fig. 134. Drulia browni, BMNH 1931.4.23.8, specimen in the same box of
Diagnosis the type. "This specimen is not the type, but there is a fragment of the actual
type in the collection presented to Bowerbank by R. Brown" (from the orig-
inallabel).
Metaniidae with body shape ranging from encrusting to mas-
sive, from nodular to globular around stems. Consistency of dry
sponges very rigid, hard. Anisotropic choanosomal skeleton with
open large reticulation of diverging multi spicular primary fibres D. batesii (Bowerbank, 1863) (River Amazon, Rio del Plata
and irregularly scattered multispicular secondary tracts. Very Argentina); D. conifera Bonetto & Ezcurra de Drago, 1973 (Rio
sparse spongin. Megasc\eres smooth oxeas. Gemmules scattered Orenoco, Venezuela); D. stenosclera Volkmer-Ribeiro & Mothes
towards the sponge surface, subspherical, enclosed in irregular de Moraes, 1981 (Rio Negro, tributary of the Amazon);
cages of megascleres. Gemmular theca of compact spongin. D. uruguayensis Bonetto & Ezcurra de Drago, 1969 (Uruguay River,
Pneumatic layer atypical as large spaces between few trabecules Parana River, Suriname). Tropical and subtropical South America.
of spongin displaced distally to the outer layer. Gemmuloscleres
parmulifonn regularly embedded in the inner layer of compact Description of type species
spongin, in a very variable number oflayers.
Drulia browni (Bowerbank, 1863) (Figs 134-137).
Scope and distribution Material examined. Holotype: not seen. Other material.
BMNH 1931.4.23.8 - dry box 6, Carter's running N° 527,
Six other species: D. geayi (Gravier, 1899) (River Orinoco, River in British Guyana, R. Shomburgk. DTRG FW551 - Brasil,
Venezuela); D. cristata (Wehner, 1895) (Amazon River, Tapajos); C. Volkmer-Ribeiro.
Porifera· Demospongiae • HaploscIerida • Spongillina 995

Fig. 135. Drulia browni, skeletal network, specimen BMNH 1931.4.23.8. A-B, choanosomal skeleton. C, multispicular tract. D, spicular arrangement at
the junction of primary and secondary tracts.

Description. Body shape from encrusting to massive, from of rotules. The single circular proximal rotule (20--36/-Lm), with
nodular to globular around stems. Bush-like grossly hispid surface lower surface concave, supports a distal short acute conical stem
due to apices of choanosomal fibres on conules. Colour from light oriented toward the outer layer.
green to grey, dark-brown to black in dry specimens. Consistency of Ecology. Sponges settle on periodically submerged (3-5
dry sponges very rigid, hard, often brittle. Dermal membrane pres- months) branches of trees and rocks. Distribution. Rio Amazon,
ent in some areas. Subdermal cavities not observed. Oscules con- Rio Negro (Brazil), Rio Beni.
spicuous. Ectosomal skeleton with apices of primaries forming Remarks. Parmula Carter, 1881c, with type species
conules. Choanosomal skeleton anisotropic with open very large Spongilla batesii Bowerbank, 1858: 315 (by subsequent designa-
(0.5-2 mm) reticulation of stout very firm multispicular ascending tion, de Laubenfels, 1936a: 37), is a synonym of Drulia on account
and diverging primary fibres and irregularly scattered multispicular of the inclusion of this species in this genus (present work).
secondary tracts. Very sparse spongin. Basal spongin plate adherent
to substratum with an open network of flattened fibres. Megascleres
smooth stout bent oxeas (300--550 X 16-43 /-Lm). Small oxeas stout HOUSSAYEUA BONETTO & EZCURRA DE
(128-290 X 22-28 /-Lm) slightly curved acerate are involved in the DRAGO, 1966
gemmular cage structure. Microscleres rare within the skeletal net-
work, slender and fusiform acanthoxeas (34--99 X 2-10 /-Lm) with Synonymy
scattered spines bearing microspines. Gemmules (580--700/-Lm)
subspherical, scattered towards the sponge surface and enclosed, Houssayella Bonetto & Ezcurra de Drago, 1966: 129.
singly or coupled, in a dense variably developed irregular cage of
one or more layers of megascleres. The cage is connected to the Type species
gemmular surface by stout large spongin trabecules (fibres).
Foramen as a short tube with collar. Gemmular theca bi-Iayered of Houssayella iguazuensis Bonetto & Ezcurra de Drago, 1966:
compact spongin with outer layer notably developed and inner layer 129 (by monotypy).
with adhering proximal rotules of gemmuloscleres. Pneumatic layer
structure atypical with large spaces between few trabecules of spon- Diagnosis
gin shifted distally to the outer layer. Gemmuloscleres parmuliform
(25-28/-Lm), regularly embedded in the layer of compact spongin, Metaniidae with encrusting body shape with irregular
in a very variable number oflayers (1-4), with overlapping margins surface. Skeletal network characterised by sparse megascleres
996 Porifera· Demospongiae • HaploscIerida • Spongillina

Fig. 136. Drulia browni, spicules and gemmules, specimen BMNH 1931.4.23.8. A, cage including two gemmules. B-C, gemmules partly free from cage.
D, gemmule. E, foramen. F, spongin projections on outer gemmular layer. G-I, parmuliform gemmuloscleres adhering to the inner layer of the theca.
J, gemmulosclere. K-L, megascleres. M, microscieres. N, spines ofmicroscleres.

and abundant microscleres, with dense choanosome paucispicular Scope and distribution
fibres sometimes arranged perpendicularly. Megascleres range
from oxeas in the skeleton to strongyles in the gemmular Monotypic, Neotropical.
cage. Microscleres range from acanthostrongyles to oxeas and
acanthoxeas with long perpendicular or slanting spines to aster-like Description of type species
shaped spicules. Gemmules in a single layer strictly adhering to the
substrate, enclosed in a multigemmular cage of variably shaped Houssayella iguazuensis Bonetto & Ezcurra de Drago, 1966
acanthostrongyles. Foramen tube-shaped. Gemmuloscleres biro- (Figs 138-140).
tulates with an extremely variable shape radially arranged in a Material examined. Type material. INALI and MABA -
single layer. not available, Saito San Martin, Iguazu faJIs, Parana River,
Porifera· Demospongiae • HapJosclerida • Spongillina 997

Fig. 137. Geographic di tribUli on of the genu Drulia .

·.1

Fig. 138. Houssayella iguazuensis, original plate showing the spiculation of the type specimen (modified from Bonetto & Ezcurra de Drago, 1966).
998 Porifera· Demospongiae • HaploscIerida • Spongillina

Fig. 139. Houssayella iguazuensis, spicular and gemmular characters, (FZRG) MCN 123. A, gemmuloscIeres arrangement on the gemmule surface.
B, gemmuloscIeres. C-D, megascIeres. E, microscIeres. F, acanthostrongyle of the gemmular cage.

Province of Misiones, Argentina, A.A. Bonetto & J.D. Ezcurra ornamented by micro-spines, to an aster-like shaped spicules with
de Drago. Other material. FZRG MCN123 - Rio Itii, 4.vi.1982, up to 7 rays. Gemmules (600-950/Lm) of pale yellow colour,
det. C. Volkmer-Ribeiro. DTRG FW594 - Rio Itil, 28.iv.l970, det. grouped in a single layer at the sponge base strictly adhering to the
C. Volkmer-Ribeiro. substrate. Gemmules enclosed in a multi-gemmular (2-40) brown-
Description. Encrusting sponges with irregular shape up to yellowish cage of acanthostrongyles associated to spined spherules
1.5 cm diameter. Irregular surface from emerging fibres. Dry of silica with spines of variable length up to aster-like shaped
sponges whitish. Oscules not conspicuous. Dermal membrane not spiCUles. Acanthostrongyles (100-210 X 12-18 /Lm) ofthe cage are
observed. Subdermal cavities not observed. Ectosomal skeleton not curved with inflated tips and a highly variable shape, size and distri-
evident. Choanosomal skeleton dense with paucispicular fibres bution of spines. Gemmular theca not observed. Foramen tube-
sometimes arranged perpendicularly up to the sponge surface. shaped (30/Lm) emerging from the gemmular cage. Outer layer
Abundant microscleres irregularly tangentially arranged in a dense not observed. Inner layer not observed. Gemmuloscleres stout
network. Megascleres (210-265 X 7-12 /Lm) from oxeas in the microspined birotulates (17-45/Lm total length, 8-15/Lm shaft
skeleton to strongyles in the gemmular cage from densely spined to length), radially arranged in a single layer with an extremely vari-
smooth particularly at the apices. Microscleres (20-50/Lm) range able shape. Microspined rotules (17-23 /Lm) irregularly indented.
from thin strongyles with spines particularly dense at the tips, to Ecology. Rocky bottoms in rapids on lower side of
oxeas and acanthoxeas, with long perpendicular or slanting spines rocks. Distribution. Argentina, River Parana; Middle River,
Porifera· Demospongiae • Haplosclerida • Spongillina 999

Fig. 140. Geographic distribution of the genu Houssayella.

Uruguay; S Brazil, lower River Uruguay; Brazil, Rio Grande do Type species
SuI, Itil River.
Remarks. Volkmer-Ribeiro & Rtitzler (1997) considered Metschnikowia tuberculata Grimm, 1876: 3 (by original
HoussayeUa to be closely related to Pachyrotula and Heterorotula. designation).

Diagnosis

FAMILY METSCHNIKOWIIDAE CZERNIAVSKY, 1880 Metschnikowiidae with encrusting to globular body shape.
Consistency solid. Ectosomal skeleton as apices of ascending pau-
Synonymy cispicular tracts. Choanosomal skeleton is a paucispicular reticula-
tion of triangular meshes with radial ascending paucispicular tracts
Metschnikowiidae Czemiavsky, 1880. Type genus. without anastomoses. Sparse spongin. Megascleres tuberculate
Metschnikowia Grimm, 1876. spiny oxeas. Microscleres absent. Gemmules absent. Larvae free,
oval, ciliated.
Definition
Scope and distribution
Spongillina with encrusting to globular body shape.
Consistency solid. Ectosomal skeleton occurring at apex of ascend- Monotypic. Caspian Sea.
ing paucispicular tracts. Choanosomal skeleton is a paucispicular
reticulation of triangular meshes with radial ascending paucispicu- Description of type species
lar tracts without anastomoses. Sparse spongin. Megascleres
range from smooth to tuberculate or densely spined oxeas with Metschnikowia tuberculata Grimm, 1876 (Figs 141-142).
acerate tips. Microscleres absent. Gemmules absent. Larvae free, Synonymy. Metschnikowia intermedia Grimm, 1876.
oval, ciliated. Renieraflava Grimm, 1876. Amorphina caspia Grimm, 1877.
Material examined. Type material: not available-
Scope and distribution Karabughas, E Caspian Sea (40 32'N, 2°33'E).
0

Description (from Dybowsky, 1880, Czerniavsky, 1880 and

Monotypic. Caspian Sea (Fig. 1). Koitun, 1962a). Body shape variable, from a thick crust (15 mm)
to globular (5 cm diameter). Consistency solid. Colour from whitish
to red. Surface with radiating exhalant canals. Scattered oscules.
Subdermal cavities as exhalant canal. Ectosomal skeleton with
METSCHNIKOWIA GRIMM, 1876 apices of ascending paucispicular tracts protruding. Choanosomal
skeleton a dense isotropic paucispicular (1-3) reticulation of
Synonymy triangular meshes with ascending paucispicular tracts radially
arranged, without anastomoses. Sparse spongin. Megascleres stout
Metschnikowia Grimm, 1876: 3. acanthoxeas (96-186 X 6-15 /J-m), ranging from straight to slightly
1000 Porifera' Demospongiae • HapJosclerida • Spongillina

, ,
,
A

8 "

.'.'
:./
'::
.. ;
.'.
:: :
.~
i
H
, ',

Fig, 141. Metschnikowia intermedia, spicules and skeleton organization, as represented in the literature. A, D, I, K, skeletal network. B, C, F, G, H, spiny
oxeas. E, drawing of a fresh specimen. L, larva. (A-G, modified from Dybowsky, 1880; H, I, K, L, modified from Koltun, 1962a.)

curved, with dense acute to rounded tubercules for the entire length, t FAMILY PALAEOSPONGILLIDAE
and tips abruptly pointed. Axial canal extremely thin, open at the VOLKMER·RIBEIRO & REITNER, 1991
extremities. Microscleres absent. Gemmules absent. Larvae free,
ciliated, oval (270 X 210 /-Lm) with several tuberculate oxeas (45 /-Lm Synonymy
length), ranging from straight to slightly curved.
Ecology, Common and abundant large specimens in brack- Palaeospongillidae Volkmer-Ribeiro & Reitner, 1991b. Type
ish waters with salinity 10-18%0, 40m depth. On stones or shells. genus. Palaeospongilla Ott & Volkheimer, 1972.
Distribution. Caspian Sea, Karabughas, Baku.
Remarks. Metschnikowia was assigned to the subfamily
Renierinae by Lundbeck (1902), and considered closely related to Definition
Baikalospongia and Reniera by Annandale (1914a). The sponge
fauna of the Black and Caspian Seas were considered in constant Spongillina consisting of a single genus of fossil sponges
contact by Czemiavsky (1880). known from the Cretaceous (100-65 MYA), and sharing similarities
Porifera· Demospongiae • Haplosclerida • Spongillina 1001

Fig. 142. Geographic distribution of the genu Melscllllikovia.

in their skeletal, spicular and gemrnular traits with modern fresh-

water sponges. Encrusting body shape. Reticulate skeleton.
Abundant spongin. Megascleres smooth and spiny oxeas.
Microscleres smooth and spiny oxeas. Gemmules range from
naked, with spicular cage, to armoured by gemrnuloscleres, with a
pneumatic layer variably developed. Foramen well developed.
Gemmuloscleres acanthoxeas to acanthostrongyles.

Scope and distribution

Monogeneric. Chubut Valley, Patagonia (Fig. 1).

t PALAEOSPONGILLA OTT & VOLKHElMER, 1972

Synonymy

Palaeospongilla Ott & Volkheimer, 1972: 53.

- 100~m
Type species

Palaeospongilla chubutensis Ott & Volkheimer 1972: 53

(by original designation). G
b

Diagnosis Fig. 143. Pa/aeospongilla chubutensis, spicules, skeleton architecture and

gemmules, paratype MABA 24748. AI, gemmuloscleres. A2, gemmule.
B, the fossil specimen (arrow) trapped in a sedimentary rock. C, gemmules
Palaeospongillidae consisting of an encrusting fossil freshwa- in formation. D, gemmules and spicular texture of the sponge skeleton.
ter sponge from the Patagonian lower Cretaceous. Choanosomal E, gemmular theca. F, multi spicular main fibre. Ga, Gb, megascleres.
skeleton anisotropic multi spicular of smooth to spiny oxeas with Gc, Gd, microscleres. (A, G, modified from Racek & Harrison, 1975;
B-F, modified from Volkmer-Ribeiro & Reitner, 1991b).
abundant spongin. Microscleres acanthoxeas. Gemmules scattered
in groups or singly. Two gemmular morphs naked without gemrnu-
loscleres, and armoured by acanthostrongyles to acanthoxeas. Description of type species
Gemrnular theca bi- or tri-layered. Gemmular cage of megascleres
present. Palaeospongilla chubutensis Ott & Volkheimer, 1972
(Figs 143-144).
Scope and distribution Material examined. Holotype: Not seen - data based on the
original description (Ott & Volkheimer, 1972), and redescriptions
Monotypic. Distribution. Chubut Valley, Patagonia. (Racek & Harrison, 1975; Volkmer-Ribeiro & Reitner, 1991b).
1002 Porifera • Demospongiae • Haplosclerida • SpongiIlina

Fig. 144. Geographic di tribution of the genu Paiaeospo/igilla.

Description. (measurements from Racek & Harrison (1975) of drought. Sponge settled around stems of aquatic plants
and Volkmer-Ribeiro & Reitner (1991 b), respectively). Body shape (gymnosperm) and plants remains were seasonally submerged.
encrusting (2-6 mm thick) with lobose contour. Surface reported as The sponge surface was covered by a calcareous layer of algae
smooth although not well-discemable due to a calcareous algal (Cyanophyceae). Distribution: Rio Chubut Valley, 16km NNW of
covering. Ectosomal skeleton undetected. Choanosomal skeleton Cerro Condor (type locality), Chubut Province, Argentina.
anisotropic with a regular meshwork and with multispicular (up to Remarks. Palaeospongilla chubutensis from the Lower
40) vertical and horizontal fibres to form a reticulum of square Cretaceous, together the finding of spicular remains in freshwater
meshes (1000 /-Lm). Abundant spongin. Megascleres straight to fee- deposits of the Upper Jura (Young, 1878), represents the oldest
bly curved oxeas, ranging from stout (230--260 X 15-18 /-Lm; confirmed origin of spongillid-like freshwater sponges from fossil
420--540 X 10--22 /-Lm) cylindrical abruptly pointed and smooth, evidence. In spite of "the careful examination of the skeletal
rarely bearing 1-2 spines, to abundant slender fusiform sharply arrangement of P. chubutensis .. . demonstrated beyond doubt that
pointed oxeas finely spined except at their tips (210--260 X the spicular components of the fossil belong to a single species"
7-10 /-Lm; 180--308 X 4-14 /-Lm). Stout oxeas form the main skele- (Racek & Harrison, 1975), from this material a second species was
ton of central and basal sponge body whereas slender oxeas are subsequently described, Spongilla patagonica Volkmer-Ribeiro &
more frequent in peripheral skeletal fibres. Microscleres more Reitner, 1991b, based on the presence of a second gemmular type.
abundant above the layer of gemmules, from straight to feebly The production of two gemmular morphs, however, is also known
curved abruptly pointed oxeas, from incipiently to conspicuously from two genera of Spongillidae (Spongilla and Corvospongilla),
spined (179-190X3-4/-Lm) to smooth (77-147 X 3-5/-Lm; both characterised by naked and armoured gemmules, with or
100--110 X 9-10 /-Lm). Gemmuloscleres (77-168 X 3.5-6 /-Lm; without pneumatic layer, and hence the recognition of a second
70--80 X 8/-Lm) slightly curved slender acanthostrongyles to acan- species may be erroneous.
thoxeas according to the position of spines at the tips. Spines,
scattered along the entire surface, are more grouped and recurved
at the tips, often with a single spine as prolongation of the axis. FAMILY POTAMOLEPIDAE BRIEN, 1967
Gemmules subspherical (483-644 /-Lm; 490--570/-Lm) diffuse in the
skeleton single or grouped. Two types of gemmules: armoured and Synonymy
naked. Naked gemmules without gemmuloscleres, enveloped in a
cage of spined megascleres, bi-Iayered gemmular theca with thin Potamolepidae Brien, 1967. Potamophloiinae Brien, 1969.
pneumatic layer and conspicuous inner layer, foramen simple with- Type genus. Potamolepis Marshall, 1883.
out a porus tube. Armoured gemmules with gemmuloscleres vari-
ably arranged in one layer crossing each other at various angles; Definition
thick tri-Iayered theca, outer layer ill defined, well developed inner
(20/-Lm) and pneumatic (100 /-Lm) layers, nipple-like foramen. Spongillina with body shapes ranging from encrusting,
Ecology. Cretaceous lacustrine sediments of the Chubut massive to arborescent with irregular lobes, ridges or branches.
Valley. The palaeohabitat was a lake subjected to long periods Consistency ranges from rigid to stony hard. Surface smooth
Porifera' Demospongiae • Haplosclerida • Spongillina 1003

or conulated. Ectosomal skeleton, if present, with microscleres in strongyles. Larvae paranchymella,entirely ciliated, with smooth
the dermal membrane. Choanosomal skeleton alveolate-reticulate, oxeas.
notably dense at the surface and more loose and irregular at the
sponge base. Very sparse spongin. Megascleres strongyles, varying Scope and distribution
from smooth to granular or spiny with inflated ends. Microscleres,
if present, are slender oxeas in the ectosome. Gemmules located at Six genera: Potamolepis, Echinospongilla nom. nov.,
the sponge base or strictly adhering to the substr

Common questions