Vol.

2
Manual of Central
National Research Conseil national
Council Canada de recherches Canada

American Diptera
True flies, or Diptera, are an exceedingly diverse and

Manual of Central
American Diptera
tremendously important group of animals, not only
because of their impact on human health, but also
because of their varied roles as decomposers, herbivores,
predators, and parasites of other animals. This two-
volume book, a collaboration of over 70 experts on
Diptera, is the first-ever resource for the identification and
understanding of Central American flies. Together, the two
Volume 2
volumes treat all 106 families of flies found in the region,
including discussions of biology, economic importance,
classification, identification to the genus level, as well as
a synopsis of each genus. In a time of major ecosystem
destruction, this work provides the basis for understanding
the biodiversity of one of the major orders of insects in a edited by:
large tropical region. It will be the foundation for future B.V. Brown (head editor)
taxonomic studies for decades to come.
A. Borkent
While Volume 1 includes several introductory chapters and J.M. Cumming
treats 42 families of flies in the Lower Diptera, Volume 2​ ​
covers the remaining 64 families of flies that make
D.M. Wood
up the Higher Diptera (or Cyclorrhapha). These include N.E. Woodley
families of house flies, fruit flies, bot flies, flower flies and
M.A. Zumbado
many other lesser-known groups. The text is accompanied
by over 1660 line drawings and photographs.
Brown
Borkent
Cumming
Wood
Woodley
Zumbado

ISBN-13 978-0-660-19958-0
ISBN 0 - 660 - 19958 - 0

9 780660 199580
The 113 chapters of this two-volume book represent an extraordinary collaboration of many of the world’s leading Diptera
taxonomists. Their contributions have been edited and organized by an expert committee headed by Dr. Brian V. Brown of
the Natural History Museum of Los Angeles County, Los Angeles, and consisting of Dr. Art Borkent (Royal British
Columbia Museum, Victoria), Dr. Jeffrey M. Cumming, Dr. D. Monty Wood (both of Agriculture and Agri-Food Canada,
Ottawa), Dr. Norman E. Woodley (USDA Systematic Entomology Laboratory, Washington, DC), and Manuel A. Zumbado
(Instituto Nacional de Biodiversidad, Costa Rica).

Financial support for this publication was obtained from the Samuel Wendell Williston Diptera Research Fund, the Natural
History Museum of Los Angeles County, and Agriculture and Agri-Food Canada.

Front cover design by Brian Koehler.

Front cover photo: Glyphidops flavifrons (Neriidae).
Back cover photo: Palpada sp. (Syrphidae).
Insets (top to bottom): Melaloncha elongata (Phoridae); Brachydeutera sp. (Ephydridae); Hystricia amoena (Tachinidae).
 A Publication of the
National Research Council of Canada
Monograph Publishing Program

Manual of Central
American Diptera
Volume 2

Edited by

B.V. Brown
Natural History Museum of Los Angeles County,
Los Angeles, California, USA

A. Borkent
Royal British Columbia Museum,
Victoria, British Columbia, Canada

J.M. Cumming
Agriculture and Agri-Food Canada,
Ottawa, Ontario, Canada

D.M. Wood
Agriculture and Agri-Food Canada,
Ottawa, Ontario, Canada

N.E. Woodley
USDA Systematic Entomology Laboratory,
Washington, DC, USA

M.A. Zumbado
Instituto Nacional de Biodiversidad,
Costa Rica

NRC Research Press

Ottawa 2010
© 2010 National Research Council of Canada
All rights reserved. No part of this publication may be reproduced in a retrieval system, or transmitted by any means,
electronic, mechanical, photocopying, recording or otherwise, without the prior written permission of the National
Research Council of Canada, Ottawa, ON K1A 0R6, Canada. Printed on acid-free paper.

ISBN 978-0-660-19958-0
NRC 49739

1. CIP for print publication:

Library and Archives Canada Cataloguing in Publication

Manual of Central American Diptera. Volume 2 / edited by B.V. Brown ... [et al.].

Issued by: National Research Council Canada.

Includes bibliographical references and index.
ISBN 978-0-660-19958-0 (bound)

1. Diptera--Central America--Classification. 2. Diptera--Central America

--Identification. 3. Flies--Central America--Classification. 4. Flies--Central
America--Identification. I. Brown, Brian Victor II. National Research Council

QL535.2.C35 M36 2010 595.7709728 C2010-980031-1

NRC Monograph Publishing Program

Editor: P.B. Cavers (University of Western Ontario)

Editorial Board: W.G.E. Caldwell, OC, FRSC (University of Western Ontario); M.E. Cannon, FCAE, FRSC
(University of Calgary); K.G. Davey, OC, FRSC (York University); M.M. Ferguson (University of Guelph);
S. Gubins (Annual Reviews); B.K. Hall, FRSC (Dalhousie University); P. Hicklenton Ph.D, P.Ag.; W.H. Lewis
(Washington University); A.W. May, OC (Memorial University of Newfoundland); B.P. Dancik, Editor-in-Chief,
NRC Research Press (University of Alberta)

Inquiries: Monograph Publishing Program, NRC Research Press, National Research Council of Canada, Ottawa,
Ontario K1A 0R6, Canada.
Web site: http://pubs.nrc-cnrc.gc.ca

Correct citation for this publication: Brown, B.V. et al. 2010. Manual of Central American Diptera: Volume 2. NRC
Research Press, Ottawa, Ontario, Canada. 728 pp.
TABLE OF CONTENTS

Table of Contents . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v
Acknowledgements for Volume 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii
Authors of Volume 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ix
Figure Credits for Volume 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xiii
Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xv
50 PLATYPEZIDAE (flat-footed flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 715
51 LONCHOPTERIDAE (spear-winged flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 721
52 PHORIDAE (hump-backed flies, scuttle flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 725
53 SYRPHIDAE (flower flies, hover flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 763
54 PIPUNCULIDAE (big-headed flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 793
55 MICROPEZIDAE (stilt-legged flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 805
56 NERIIDAE (neriid flies, cactus flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 815
57 PSEUDOPOMYZIDAE (pseudopomyzid flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 821
58 TANYPEZIDAE (tanypezid flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 827
59 SOMATIIDAE (somatiid flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 833
60 PSILIDAE (rust flies, psilid flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 837
61 SYRINGOGASTRIDAE (syringogastrid flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 843
62 CONOPIDAE (thick-headed flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 847
63 LONCHAEIDAE (lance flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 857
64 PIOPHILIDAE (skipper flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 865
65 RICHARDIIDAE (richardiid flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 871
66 ULIDIIDAE (picture-winged flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 881
67 PLATYSTOMATIDAE (signal flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 903
68 TEPHRITIDAE (fruit flies, moscas de frutas) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 909
69 PYRGOTIDAE (pyrgotid flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 955
70 CTENOSTYLIDAE (ctenostylid flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 963
71 LAUXANIIDAE (lauxaniid flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 971
72 CHAMAEMYIIDAE (chamaemyiid flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 997
73 PARALEUCOPIDAE (paraleucopid flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1009
74 COELOPIDAE (kelp flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1013
75 SCIOMYZIDAE (snail-killing flies, marsh flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1017
76 ROPALOMERIDAE (ropalomerid flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1025
77 SEPSIDAE (black scavenger flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1031
78 CLUSIIDAE (clusiid flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1041
79 ODINIIDAE (odiniid flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1049
80 AGROMYZIDAE (leaf-mining flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1057
81 ANTHOMYZIDAE (anthomyzid flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1073

v
 82 AULACIGASTRIDAE (aulacigastrid flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1083
83 PERISCELIDIDAE (periscelid flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1087
84 ASTEIIDAE (asteiid flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1093
85 BRAULIDAE (beelice) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1097
86 CARNIDAE (carnid flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1101
87 INBIOMYIIDAE (inbiomyiid flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1105
88 TETHINIDAE (beach flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1109
89 CANACIDAE (surf flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1115
90 CRYPTOCHETIDAE (cryptochetid flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1121
91 MILICHIIDAE (milichiid flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1125
92 CHLOROPIDAE (frit flies, grass flies, eye gnats) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1137
93 CHYROMYIDAE (chyromyid flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1155
94 HELEOMYZIDAE (heleomyzid flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1159
95 SPHAEROCERIDAE (small dung flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1165
96 CURTONOTIDAE (hunchbacked flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1189
97 CAMILLIDAE (camillid flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1193
98 DROSOPHILIDAE (small fruit flies, pomace flies,vinegar flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1197
99 DIASTATIDAE (diastatid flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1207
100 EPHYDRIDAE (shore flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1211
101 NANNODASTIIDAE (nannodastiid beach flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1235
102 HIPPOBOSCIDAE (louse flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1241
103 STREBLIDAE (bat flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1249
104 NYCTERIBIIDAE (bat flies, spider bat flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1261
105 SCATHOPHAGIDAE (dung flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1267
106 ANTHOMYIIDAE (anthomyiid flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1271
107 FANNIIDAE (fanniid flies, latrine flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1277
108 MUSCIDAE (house flies, stable flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1281
109 CALLIPHORIDAE (blow flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1297
110 OESTRIDAE (bot flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1305
111 SARCOPHAGIDAE (flesh flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1313
112 RHINOPHORIDAE (woodlouse flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1337
113 TACHINIDAE (tachinid flies, parasitic flies) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1343
TAXONOMIC INDEX TO VOLUME 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1419

vi
ACKNOWLEDGMENTS FOR VOLUME 2
Brian V. Brown

General acknowledgments arranging the plates, altering digital images, and providing
new images when necessary were all executed with impeccable
The Manual of Central American Diptera project was a precision, talent, and punctuality. We all wish we could have
large and complex endeavor, whose success was determined such illustrators working for us full time. Funding for illus-
by many people. The project was an outgrowth of the Biodi- trations was obtained in part from the Samuel Wendell
versity Resources Development Project, which consisted of Williston Diptera Research Fund (courtesy of Chris Thompson)
two initiatives: the country-wide inventory for Costa Rica, and from Monty Wood.
and the taxon-based special projects. Work in Costa Rica Permission to use illustrations from other sources for both
was funded in part by support to INBio by the Biodiversity volumes was obtained by Giar-Ann Kung.
Resources Development Project (CR-GE-39876), funded by We thank the staff at NRC Press, especially Rob Forrest,
Global Environment Facility (GEF) through its implementing Thad Costigan, and Suzanne Kettley, for working with us to
agency the World Bank. To coordinate this effort, INBio produce the two book volumes. We appreciate all the work
arranged a meeting of interested dipterists at Monteverde, that they put into making this a quality product.
Costa Rica, in May 1998. It was decided that international As scientists, we are all grateful to Costa Rica’s Sistema
collaborators would help train the Diptera parataxonomists Nacional de Areas de Conservación (Ministry of the Envi-
and that the taxon-based special project would be a “Manual ronment) for their conservation efforts that preserve 25% of
of the Costa Rican Diptera.” A second meeting was held in Costa Rica´s territory for Diptera and other wildlife.
June 2000, again at Monteverde, followed by parataxonomist
training at Albergue Heliconias. A third major meeting took
place on August 10–11, 2001 at Rincon on the Osa Peninsula, Further acknowledgments by authors
Costa Rica. At this meeting, it was proposed and accepted
that the scope and title of the manual be changed to “Manual Peter Chandler (Platypezidae) thanks Chris Thompson
of Central American Diptera.” Further organizational meetings and Norm Woodley for enabling him to examine material
were held in Riosparaiso in February 2003 and a smaller one from Central and South America.
at Nacientes Palmichal (editorial committee only), February For support of research associated with this project,
2004. Brian Brown (Phoridae) gratefully acknowledges USA
I am extraordinarily indebted to all the authors of chapters, National Science Foundation (NSF) grants DEB-9407190,
who provided authoritative reviews of their groups. For DEB-0090031, DEB-0315271, DEB-0516420, as well
Volume 2, I am especially thankful for the work of Matthias as NSF support of the Arthropods of La Selva project (to
Buck and Steve Marshall, both of whom tackled large Robert Colwell and Jack Longino). Henry Disney and Sibylle
“orphaned” groups not of their immediate expertise, essentially Noack provided helpful reviews of the phorid chapter. For
becoming experts from the ground up. long-time friendship and camaraderie in Costa Rica, I am
The editorial committee has provided an incredible level grateful to Paul Hanson and Carolina Godoy. For assistance
of expertise and support in making this volume possible. Art
in the laboratory and the field, I thank Vladimir Berezovskiy,
Borkent, Jeff Cumming, Monty Wood, Norm Woodley, and
Emily Corona, Lisa Gonzalez, Giar-Ann Kung, Wendy Porras,
Manuel Zumbado all sacrificed their valuable time to look
Claus Rasmussen, Kelly Walker, and Elvia Zumbado. Illus-
over various versions, make corrections, give opinions, and
trations and cover design were expertly rendered by Brian
help organize this massive undertaking.
The scientific staff at INBio, Manuel Zumbado, Guillermo Koehler. Scanning electron micrographs were produced by
Chaverri, Elvia Zumbado, Annia Picado, as well as Alvaro Giar-Ann Kung.
Herrera, were tremendously helpful and vital to the successful Matthias Buck (Neriidae, Psilidae, and others) was funded by
conclusion of this project. All of the editors and authors of various Natural Sciences and Engineering Research Council
the various chapters are particularly grateful for their mak- (NSERC) of Canada grants to Steve Marshall.
ing the collection and facilities of INBio available to visiting Owen Lonsdale (Somatiidae, Clusiidae) had research
researchers, facilitating collecting permits, and arranging funded by an NSERC grant awarded to him and an NSERC
meetings. Furthermore, the contributions of all the paratax- Discovery Grant awarded to Steve Marshall.
onomists at INBio in sampling Costa Rican biodiversity and For support of research associated with this project,
building the Diptera collection is gratefully acknowledged. Stephen Gaimari (Lauxaniidae, Chamaemyiidae, Odiniidae)
Illustrators Ana Brenes and Ricardo Vargas (both of gratefully acknowledges NSF grant DEB-0075206 to
INBio) were also vital people in this project. Their tasks of S. Gaimari.

vii
viii MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

For support of research associated with this project, Vera John Swann (Milichiidae) thanks Dr. G.G.E. Scudder and
Silva (Lauxaniidae, Sepsidae) gratefully acknowledges Dr. W. Maddison, The University of British Columbia (UBC),
Fundação de Amparo à Pesquisa do Estado de São Paulo for providing him with the space necessary to complete his
(FAPESPS) grants Procs. 94/2031-8, 94/4805-0, 97/13984-5, chapter; Ms. K. Needham and Dr. R. Kenner, UBC, are thanked
and 03/10274‑9. for their comments on earlier drafts of his manuscript.
Stephanie Boucher (Agromyzidae) thanks M. von Tsch- Thomas Pape (Sarcophagidae, Oestridae, Rhinophori-
irnhaus for his comments and suggestions on the key, as dae) thanks Mrs.  E.  Binkiewicz, Mrs.  P.  von  Knorring, and
well as M. Sasakawa, V.V. Zlobin, and O. Lonsdale for re- Mr. M. Meldert, all from Stockholm, Sweden, for producing
viewing the key, and T.A. Wheeler for his comments on the illustrations.
chapter. Any opinions, findings, and conclusions or recommenda-
Jindřich Roháček (Anthomyzidae) acknowledges fi- tions expressed in this book are those of the author(s) and
nancial support from grants No. 206/03/0020 and do not necessarily reflect the views of the National Science
No. 206/06/0287 of the Czech Science Foundation. Foundation.
AUTHORS OF VOLUME 2
Kyle Apigian, Division of Insect Biology, Department of Environmental Science, Policy and Management, University of California, Berkeley,
CA, 94720, USA.

Kevin N. Barber, Natural Resources Canada, Canadian Forest Service, Great Lakes Forestry Centre, 1219 Queen Street East, Sault Ste.
Marie, ON, P6A 2E5, Canada.
E-mail: [email protected].

Stéphanie Boucher, Lyman Entomological Museum, McGill University, Macdonald Campus, Ste-Anne-de-Bellevue, QC, H9X 3V9, Canada.
E-mail: [email protected].

Brian V. Brown, Entomology Section, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles,
CA, 90007, USA.
E-mail: [email protected].

Matthias Buck, Invertebrate Zoology, Royal Alberta Museum, 12845-102nd Avenue, Edmonton, AB, T5N 0M6, Canada.
E-mail: [email protected].

Sidney Camras, Field Museum, Insect Division, Roosevelt Road, at Lake Shore Drive, Chicago, IL, 60605-2496, USA.

Peter J. Chandler, 606B Berryfield Lane, Melksham, Wilts SN12 6EL, United Kingdom.
E-mail: [email protected].

Gregory A. Dahlem, Department of Biological Sciences, Northern Kentucky University, Highland Heights, KY, 41099, USA.
E-mail: [email protected].

Carl W. Dick, Department of Biology, Western Kentucky University, 1906 College Heights Blvd. #11080, Bowling Green, KY, 42101-1080, USA.
E-mail: [email protected].

Amnon Freidberg, Department of Zoology, The George S. Wise Faculty of Life Sciences, Tel Aviv University, Tel Aviv, 69978, Israel.
E-mail: [email protected].

Stephen D. Gaimari, California State Collection of Arthropods, California Department of Food and Agriculture / Plant Pest Diagnostics,
3294 Meadowview Road, Sacramento, CA, 95832-1448, USA.
E-mail: [email protected].

Gustavo Graciolli, Universidade Federal de Mato Grosso do Sul, Centro de Ciências Biológicas e da Saúde, Departamento de Biologia,
Cidade Universitária s/n, Cidade Universitária, 79070-900 Campo Grande, Caixa-Postal 549, MS, Brazil.
E-mail: [email protected].

David Grimaldi, American Museum of Natural History, Central Park West at 79th St., New York, NY, 10024-5192, USA.
E-mail: [email protected].

E. Geoffrey Hancock, Hunterian Museum, Graham Kerr Building, University of Glasgow, G12 8QQ, Scotland, UK.
E-mail: [email protected].

Vicente Hernández-Ortiz, Red de Interacciones Multitróficas, Instituto de Ecología, A. C. Km 2.5 carretera antigua a Coatepec N° 351,
Congregación El Haya, Apartado Postal 63, Xalapa, Veracruz, 91070, México.
E-mail: [email protected].

Sergio Ibáñez-Bernal, Departamento de Biodiversidad y Ecología Animal, Instituto de Ecología, A. C. Km 2.5 carretera antigua a Coatepec
N° 351, Congregación El Haya, Apartado Postal 63, Xalapa, Veracruz, 91070, México.
E-mail: [email protected].

Elena P. Kameneva, General and Applied Entomology Section, I. I. Schmalhausen Institute of Zoology, Bogdan Chmielnicki 15,
01601, Kiev, Ukraine.
E-mail: [email protected].

Ashley H. Kirk-Spriggs, Department of Entomology, National Museum, P.O. Box 266, Bloemfontein 9300, South Africa; and Department of
Zoology and Entomology, University of the Free State, P.O. Box 339, Bloemfontein, 9300, South Africa.
E-mail: [email protected].

ix
x MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

John Klymko, Atlantic Canada Conservation Data Centre, P.O. Box 6416, Sackville, NB, E4L 1G6, Canada.
E-mail: [email protected].

Lloyd Knutson, Salita degli Albito 29, 04024, Gaeta (LT), Italy.
E-mail: [email protected].

Valery A. Korneyev, General and Applied Entomology Section, I. I. Schmalhausen Institute of Zoology, Bogdan Chmielnicki 15,
01601 Kiev, Ukraine.
E-mail: [email protected].

Cheslavo A. Korytkowski, Programa Centroamericano de Entomología, Vice-Rectoria de Investigacion y Post-Grado, Universidad de

Panamá, Estafeta Universitaria, Panamá.
E-mail: [email protected].

Owen Lonsdale, Invertebrate Biodiversity, Agriculture and Agri-Food Canada, K.W. Neatby Bldg., CEF, Ottawa, ON, K1A 0C6, Canada.
E-mail: [email protected].

Luciane Marinoni, Department of Zoology, Universidade Federal do Paraná, Cx. P. 19020, 81.531-980, Curitiba, Paraná, Brazil.
E-mail: [email protected].

Stephen A. Marshall, School of Environmental Sciences, University of Guelph, Guelph, ON, N1G 2W1, Canada.
E-mail: [email protected].

Wayne N. Mathis, Department Entomology, P.O. Box 37012, CE 619, MRC 169, Smithsonian Institution, Washington, DC, 20013-7012, USA.
E-mail: [email protected].

David K. McAlpine, Australian Museum, 6 College Street, Sydney, 2010, Australia.

E-mail: [email protected].

J. Frank McAlpine, Invertebrate Biodiversity, Agriculture and Agri-Food Canada, K.W. Neatby Bldg., CEF, Ottawa, ON K1A 0C6, Canada.

Verner Michelsen, Zoological Museum, University of Copenhagen, Universitetsparken 15, DK-2100 Copenhagen Ø, Denmark.
E-mail: [email protected].

Jeremy A. Miller, Department of Terrestrial Zoology, Nationaal Natuurhistorisch Museum Naturalis, Postbus 9517, 2300 RA Leiden,
The Netherlands.
E-mail: [email protected].

Allen L. Norrbom, Systematic Entomology Laboratory, PSI, Agricultural Research Service, USDA, c/o National Museum of Natural History,
MRC 168, Washington, DC 20013-7012, USA.
E-mail: [email protected].

Andrej L. Ozerov, Zoological Museum, Moscow State University, Bol’shaya Nikitskaya Str. 6, Moscow, 103009, Russia.
E-mail: [email protected].

Thomas Pape, Zoological Museum, University of Copenhagen, Universitetsparken 15, DK-2100 Copenhagen Ø, Denmark.
E-mail: [email protected].

José Albertino Rafael, Instituto Nacional de Pesquisas da Amazônia, P.O. Box 478, 69011-970, Manaus, Amazonas, Brazil.
E-mail: [email protected].

Jindřich Roháček, Slezské zemské muzeum, Tyršova 1, CZ-746 01 Opava, Czech Republic.

E-mail: [email protected].

Graham Rotheray, National Museums of Scotland, Chambers Street, Edinburgh, EH1 1JF, United Kingdom.
E-mail: [email protected].

Alessandra Rung, California State Collection of Arthropods, California Department of Food and Agriculture  / Plant Pest Diagnostics,
3294 Meadowview Road, Sacramento, CA, 95832-1448, USA.
E-mail: [email protected].

Jade Savage, Department of Biological Sciences, Bishop’s University, Sherbrooke, QC, J1M 1Z7, Canada.
E-mail: [email protected].
Authors of Volume 2 xi

Vera C. Silva, Departamento de Morfologia e Fisiologia Animal, Faculdade de Ciências Agrárias e Veterinárias, São Paulo State University,
UNESP, Via de Acesso Prof. Paulo Donato Castellane, s/n, 14884-900 Jaboticabal, São Paulo, Brazil.
E-mail: [email protected].

Jeffrey H. Skevington, Invertebrate Biodiversity, Agriculture and Agri-Food Canada, K.W.  Neatby Bldg., CEF, Ottawa, ON,  K1A  0C6,
Canada.
E-mail: [email protected].

John E. Swann, Department of Biological Sciences, 2500 University Drive N.W., University of Calgary, Calgary, AB, T2N 1N4, Canada.
E-mail: [email protected].

F. Christian Thompson, Systematic Entomology Lab, ARS, USDA, c/o Smithsonian Institution NHB-168, P.O.  Box  37012, Washington,
DC, 20013-7012, USA.
E-mail: [email protected].

John Vargas, Departamento de Ciencias Forenses, Organismo de Investigación Judicial, San Joaquín de Flores, Heredia, Costa Rica; and
Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica.
E-mail: [email protected].

J. Richard Vockeroth, Invertebrate Biodiversity, Agriculture and Agri-Food Canada, K.W.  Neatby Bldg., CEF, Ottawa,
ON, K1A 0C6, Canada.

Terry A. Wheeler, Department of Natural Resource Sciences, McGill University, Macdonald Campus, Ste-Anne-de-Bellevue,
QC, H9X 3V9, Canada.
E-mail: [email protected].

Andrew E. Whittington, FlyEvidence, 2 Newhouse Terrace, Queen’s Road, Dunbar, EH42 1LG, Scotland, UK.
E-mail: [email protected].

D. Monty Wood, Invertebrate Biodiversity, Agriculture and Agri-Food Canada, K.W. Neatby Bldg., CEF, Ottawa, ON, K1A 0C6, Canada.
E-mail: [email protected].

Norman E. Woodley, Systematic Entomology Lab, ARS, USDA, c/o Smithsonian Institution NHB-168, P.O.  Box  37012, Washington,
DC, 20013-7012, USA.
E-mail: [email protected].
FIGURE CREDITS FOR VOLUME 1
Many figures published in this book were originally published elsewhere. We are especially grateful to Agriculture and
Agri-Food, Ottawa, Canada, for permission to re-use many figures from the Manual of Nearctic Diptera, Volumes 1 and 2.
We also thank the following for permission to re-use figures originally published elsewhere (as referenced in figure
captions):
American Entomological Society: Fig. 68.154
American Museum of Natural History: Figs. 53.58; 70.6, 13; 77.33–34
Associated Publishers: Fig. 53.58
California Academy of Sciences: Fig. 66.37
Cambridge University Press: Figs. 103.1–3, 7–8, 21
Cornell University Press: Figs. 68.13, 16–17, 26, 31, 33, 36, 38–40, 51, 56, 58, 60–63
E. Schweizerbart Science Publishers: Figs. 110.3–6, 10
Entomological Society of America: Figs. 65.3; 103.4–6
Entomology Society of Israel: Figs. 66.1, 6–7, 28, 41–42, 44–45
Field Museum: Figs. 103.9, 10–11, 12a–f, 13a–b, 16a–b, 19c, 20, 24; 104.4
Finnish Zoological and Botanical Publishing Board: Figs. 54.18–19, 36–37
Fundação Zoobotânica do Rio Grande do Sul: Figs. 77.4, 11
Hungarian Academy of Sciences/Hungarian Natural History Museum: Fig. 101.1
Institute of Entomology of the Czech Academy of Sciences: Figs. 60.3, 6–15; 81.3; 95.8, 10–11, 13
Institute of Zoology, Slovak Academy of Sciences: Fig. 52.15
Instituto de Ecología A.C.: Figs. 76.2–16
Instituto Nacional de Biodiversidad: Figs. 61.1; 75.1; 96.1; 113.1
Instituto Oswaldo Cruz: Fig. 70.2
John Wiley & Sons, Inc .: Figs. 53.72; 54.2–3, 6–10, 15, 20–24, 26–28, 30–33, 35; 87.1–4, 7; 110.17–21; 112.1–3,
5–6, 9–11, 15–16
Kansas Entomological Society: Figs. 77.14–15
KMK Scientific Press Ltd.: Figs. 77.5, 8, 10, 12, 18–20, 23–26
Magnolia Press: Figs. 52.11, 33, 41, 51–52; 61.2–3; 78.13
Museu de Zoologia da Universidade de São Paulo: Figs. 104.7, 10, 12–14
Muséum d’histoire naturelle de la Ville de Genève: Figs. 68.9, 47, 70, 100, 102, 112
Natural History Museum of Los Angeles County: Figs. 87.5; 95.6, 34, 50
Natural History Museum, London: Figs. 104.5–6, 11
New York Entomological Society: Fig. 102.17
Pontifical Academy of Sciences: Fig. 52.54
Princeps Editions, Sarl: Fig. 110.7
Royal Entomological Society: Figs. 103.15a–b, 17a–b, 18a–b, 19a–b, 22
Schweizerische Entomologische Gesellschaft: Fig. 70.4
Senckenberg Deutsches Entomologisches Institut: Figs. 70.9; 78.14–15; 81.5; 95.24–25, 37–38; 103.23
Sociedade Brasileira de Entomologia: Figs. 54.5, 25, 29, 34; 69.3–4; 70.3, 10, 14; 77.2, 7, 9, 13, 16, 21, 22, 27–29,
30–32; 104.8
Studia dipterologica, AMPYX -Verlag: Figs. 53.66; 56.8; 66.51–53, 55, 57, 59, 69, 73–74, 77, 81–82; 77.3, 6, 17;
108.10; 111.8, 55
Taylor & Francis Ltd.: Figs. 52.19; 68.160; 95.15, 18; 111.56
Western North American Naturalist: Fig. 104.3

xiii
PREFACE
Brian V. Brown

Volume 2 of the Manual of Central American Diptera Each genus found in Central America is discussed briefly in
continues the coverage begun in Volume 1 (Brown et al., the synopsis section of each chapter, giving access to further
2009), treating the remaining 64 families of Diptera that literature and mention of their behavioral or distributional
belong to the Cyclorrhapha, or higher flies. As in Volume 1, peculiarities. This synopsis section varies considerably from
this work comprehensively treats the known (described) Dip- family to family, as our level of knowledge about large, easily
tera fauna of Central America and tropical Mexico (hereafter observed flies is much more complete than for tiny, incon-
“Central America” for simplicity). It is designed to comple- spicuous ones.
ment the coverage of the superb Manual of Nearctic Diptera In many instances, particularly for large, difficult families,
and treats all families occurring south of the Nearctic– this is the first time an identification key for Central American
Neotropical Region dividing line in Mexico and north of the genera has been attempted. Some of these keys are highly
Panama–Colombia border (see fig. 1.1 in Volume 1). The key provisional, representing our best, albeit incomplete knowl-
to families in Volume 1 strictly adheres to this coverage, so edge of the fauna. Numerous specimens that will not key
that all flies that are keyed to this level can be further treated properly are to be expected in some families (such as Phori-
in this book. There are a few families, however, that occur dae and Tachinidae), because much of the diversity of
elsewhere in South America that might eventually be found the Diptera is still unknown or undescribed. Hopefully,
future updates of these keys, either published in book format
in Central America; these are discussed in the introduction
or online at the project website (www.mcadiptera.net), will
to the family key.
improve on this situation.
The entire work is divided into two volumes, with the
introductory chapters, family keys, and treatment of the Conventions
nematocerous families and the lower Brachycera in Volume
1, and the Brachycera–Cyclorrhapha in Volume 2. An index Because it is cited in nearly every chapter throughout this
to taxa is found at the end of each volume. book, the reference for the Manual of Nearctic Diptera has
The area of coverage in this book, restricted as it may been abbreviated to “MND”. The full references for the three
seem, is still vast and encompasses an extraordinary vari- volumes are as follows:
ety of environments. The Central American Diptera fauna is
dominated by diverse tropical groups, with northern (Nearctic McAlpine, J.F., B.V.  Peterson, G.E.  Shewell, H.J.  Teskey,
Region) elements extending their ranges into Central America J.R.  Vockeroth, & D.M.  Wood (coordinators). 1981.
at higher elevations. Thus, each chapter treats the fauna of Manual of Nearctic Diptera. Volume 1. Research Branch,
Agriculture Canada, Monograph No. 27, Ottawa, Canada,
a mixture of tropical and temperate habitats within the geo-
674 pp.
graphical coverage of the book.
McAlpine, J.F., B.V.  Peterson, G.E.  Shewell, H.J.  Teskey,
Many authors of the family treatments, at their own
J.R.  Vockeroth, & D.M.  Wood (coordinators). 1987.
discretion, have extended their coverage to the entire Neo-
Manual of Nearctic Diptera. Volume 2. Research Branch,
tropical Region, the whole of the New World, or in some
Agriculture Canada, Monograph No. 28, Ottawa, Canada,
cases, the entire world. The minimum coverage, however, is pp. 675–1332.
Central America. There is a particular emphasis on the country McAlpine, J.F. & D.M. Wood (coordinators). 1989. Manual
of Costa Rica in many chapters because of the genesis of this of Nearctic Diptera. Volume 3. Research Branch, Agri-
project. The Manual of Central American Diptera was con- culture Canada, Monograph No. 32, Ottawa, Canada, pp.
ceived during a series of meetings in Costa Rica, funded by 1333–1581.
the World Bank (see Acknowledgments). Also, the emphasis
on Costa Rica is because of the extensive inventory activity Many authors have included genera in their keys that do
at the Instituto Nacional de Biodiversidad (INBio) in Santo not occur in Central America, and these are not included in
Domingo, Costa Rica. Led by Manuel Zumbado, this activity the synopsis section following their keys. Such genera are
has generated a large number of specimens and great interest surrounded by square brackets in the key, and their distribu-
in the Costa Rican fauna among the world’s experts. tions given.
For each family we provide a short description and Figure captions include the publication source of the
diagnostic paragraph, a summary of the biology (including illustrations (if not original), and for taxa not from Central
economic importance), classification, and challenges to America, the distribution of the taxon illustrated.
identification. Then an identification key to the genus level The author is given for the genus and species names of
is provided. all flies. The authors of generic and subgeneric names are

xv
xvi MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

given in the key and synopsis, but are not repeated in the nothing of our paucity of information on dipteran phylogenetic
Diagnosis, Biology, Classification, and Identification sections relationships.
(unless they are of extralimital taxa not mentioned in the key When I wrote of the “crisis in Neotropical dipterology”
or synopsis). in 2005, I was concerned about the low rate of species
descriptions of species found in the poorly known New World
The future tropics. The work leading to the Manual of Central American
This project has been a group effort over the last eight Diptera has partly addressed this problem, as many new
years by a large number of people, most of whom are mentioned species and genera (and even one family — Inbiomyiidae)
in the Acknowledgments. Their cumulative impact has made have been described for the purpose of being included in the
the Diptera of Central America more accessible to future books (e. g., Borkent, 2008; Kung & Brown, 2005; Mathis,
taxonomists, ecologists, naturalists, and amateur collectors 2008; Norrbom, 2006; Wheeler & Mlynarek, 2008). It is my
of flies. There is still much more to be done, however. Vast greatest wish that the Manual of Central American Diptera
areas of the region have not been sampled, and nowhere is continues to motivate and stimulate research on the incredibly
the entire fauna well known. In particular, we remain ignorant diverse, but still little-known flies living in the increasingly
of the lifestyles and immature stages of most flies, to say threatened natural habitats of Central America.

Literature cited
Borkent, A. 2008. The frog-biting midges of the world Mathis, W.N. 2008. Two new neotropical genera of the
(Corethrellidae: Diptera). Zootaxa 1804: 1–456. shore-fly tribe Ephydrini Zetterstedt (Diptera: Ephydri-
Brown, B.V. 2005. Malaise trap catches and the crisis in Neo- dae). Zootaxa 1874: 1–15.
tropical dipterology. American Entomologist 51: 180–183. Norrbom, A.L. 2006. A revision of the Neotropical genera
Brown, B.V., A. Borkent, J.M.  Cumming, D.M.  Wood, Molynocoelia Giglio-Tos, Pseudophorellia Lima, and
N.E. Woodley, & M.A. Zumbado. Manual of Central Alujamyia, n. gen. (Diptera: Tephritidae). Israel Journal
American Diptera, Volume 1. NRC Research Press,
of Entomology 35–36: 35–134.
Ottawa, Canada, 714 pp.
Kung, G. & B.V. Brown. 2005. Three unusually divergent Wheeler, T.A. & J.J. Mlynarek. 2008. Systematics of
new genera of phorid flies (Diptera: Phoridae) from Costa Agrophaspidium, a new genus of Neotropical Chloropidae
Rica. Zootaxa 1019: 43–52. (Diptera). Zootaxa 1926: 41–52.
PLATYPEZIDAE (FLAT-FOOTED FLIES) 50
Peter J. Chandler

Fig. 50.1. Female of Polyporivora polypori (Willard), (Nearctic, MND, fig. 50.1, as Plesioclythia agarici).

Diagnosis grey, silver, yellow, or orange markings; males of some

species more brightly colored but females usually still differ
Small to medium-sized flies (body length 1.4–10.0 mm), in coloration.
humpbacked and slender to broad-bodied (Fig. 1). Head Larvae cylindrical or more or less dorsoventrally depressed,
broad, with eyes large and holoptic in male, but separated with small head bearing short antennae and palpi; amphip-
by more or less broad frons in female. Antenna with first neustic with both anterior and posterior spiracles on short
flagellomere rounded or tapered apically, with terminal arista. tubes; usually bearing series of short marginal and dorsal
Wings broad with all long veins reaching margin and cell processes on each body segment, number and form diagnostic
cup well developed. Crossvein dm-cu present or absent and for genera and species.
M1+2 forked or unforked. Posterior tarsi often enlarged and Puparia resemble larvae except head and prothorax fused
more or less flattened, especially in females of Platypezinae. to ventral surface so that mesothorax forms anterior margin;
Secondary sexual characters found in many structures (such may be broader and more depressed than larvae.
as processes on some male hind legs), and sexes often Among cyclorrhaphous flies, the Platypezidae only
differ strikingly in coloration, with males mainly black and resemble Opetiidae, which were formerly included in the
females often lighter in color, variously marked with brown, family, in the holoptic male eyes with larger facets dorsally

715
716
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

and the erect antenna with a terminal arista. They are distin- at least 10 genera, of which eight are known from North
guished from them by the presence of three aristomeres (two America, although it is uncertain how many of these reach
in Opetiidae) and acrostichal setae reduced to a single row or South America. The tropical fauna of Platypezinae is less
absent. In body form they resemble Ironomyiidae (found in diverse generically than that of the Holarctic Region. The
Australia) and Phoridae (including Sciadocerinae), families previous assignment of tropical species to genera has been
that do not have male eyes with this structure, and the arista found to be largely incorrect in revisions of the Oriental and
is terminal in only a few Phoridae, such as Conicera Mei- Australasian species (Chandler, 1994) and Afrotropical species
gen. The wing venation is more complete than in Phoridae, (P. Chandler, unpublished data). The Neotropical fauna has
with vein Sc ending separately in the costa and R1 reach- not yet been critically studied, but preliminary work suggests
ing two-thirds or more of the wing length. The venation of that this is also true there. Most species in these three regions
Ironomyiidae is closer, but they have M1 and M2 arising belong to the large genus Lindneromyia as redefined by
separately from cell dm, while in those Platypezidae with Chandler (1994), who placed in synonymy with it several
cell dm present, M1+2 forks beyond it or does not fork. other genera recognized by Kessel (1987 and earlier works
by him and his co-workers), principally on proportional
differences in wing venation. These other genera (Symme-
Biology tricella Kessel, Penesymmetria Kessel & Maggioncalda,
Plesioclythia Kessel & Maggioncalda, Grossovena Kessel
All known larvae of this family develop in fruiting bodies & Maggioncalda) were found not to be tenable when other
of larger fungi. Chandler (2001) summarized present characters such as male genitalia were studied. Some species
knowledge of host associations, which mainly relates to the placed in Paraplatypeza Kessel & Maggioncalda have also
Holarctic fauna, and provided a key to the larvae or puparia of been re-assigned to Lindneromyia; Paraplatypeza is cur-
13 genera, including those so far recorded in Central America rently maintained for a Holarctic group with more strongly
except Microsania. Most of the fungus hosts grow on wood developed chaetotaxy of the head, but Lindneromyia may be
or have mycorrhizal associations with trees, and most paraphyletic in relation to it.
species of the family are consequently found in forests. There are about 250 described species of the family
Lindneromyia species, however, often develop in Agaricus worldwide. In general, the number of undescribed species in
species and may occur in grassland or other open habitats. collections is similar to the number of names that will be lost
The larval biology of Microsania is unknown, but the to synonymy. Some of this confusion is due to pronounced
adults of this genus are well-known as the “smoke-flies” sexual dimorphism in many species. The Neotropical fauna
because of their strongly fumotropic behavior. The males is the least well known, with only 14 described species. The
form aerial swarms in the smoke of wood fires. Other genera Afrotropical total is more than 40, so at least a comparable
also form aerial male swarms as a prelude to copulation, number of Neotropical species may be expected. An unde-
using gaps in the tree canopy as markers. Both sexes may scribed Lindneromyia species is known to occur in Tierra
be observed rapidly running about on broad leaves of trees, del Fuego, but the only described species from the Chilean
shrubs, or herbaceous plants in a similar fashion to Phoridae. region (Platypeza brunnescens Collin) is actually related to
This activity is considered to be associated with feeding on the Holarctic genus Seri Kessel & Kessel and the Palearc-
honeydew and other surface deposits on the leaves. tic genus Bolopus Enderlein, and probably represents a new
genus.
Classification
Platypezids are a small group of primitive cyclorrhaphous Identification
flies. Their systematic position and composition were dis-
cussed by Chandler (2001), who recognized four subfamilies: Chandler (2001) provided keys to the four subfamilies
Microsaniinae, Melanderomyiinae, Callomyiinae, and Platy- and to the three European genera of Callomyiinae and eight
pezinae. The first two of these are monogeneric. Microsania of Platypezinae. The characters by which other genera are
is worldwide and found in both North and South America, distinguished were also cited. Two other genera (Chydae-
while Melanderomyia Kessel, which develops in Phallaceae, opeza, Metaclythia Kessel) were also included in the key to
is known only from North America. Callomyiinae includes the Palearctic species by Chandler & Shatalkin (1998). The
six genera, of which five are known from North America but Nearctic genera were keyed by Kessel (1987); this key
only two are known to reach South America. These two, Ag- recognized the genera listed above (under Classification)
athomyia and Bertamyia, form a monophyletic group with that are now considered synonymous with Lindneromyia.
the Palearctic genus Chydaeopeza Shatalkin. It is unclear Adult specimens of Platypezidae are usually preserved
if Agathomyia, from which the other two genera have been dry. They can be glued to the side of pins or card pointed, but
split, is monophyletic without them. Platypezinae includes the author prefers to stage them using micro-pins.
PLATYPEZIDAE (flat-footed flies) 50 717

Key to the genera of Platypezidae of Central America and Nearctic Mexico

1. Crossvein dm-cu absent (Fig. 2); vein R1 reaches at most to two-thirds wing length and surrounded
apically by pterostigma; vein M1+2 forked but M1 incomplete basally; hind tarsomere 1 about as
long as tarsomeres 2–4 in both sexes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microsania Zetterstedt
– Crossvein dm-cu present, closing elongate cell dm (Figs. 3–8); vein R1 longer, exceeding
two-thirds wing length and pterostigma absent; vein M1+2 forked or unforked, but complete; hind
tarsomere 1 various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2. Uniserial row of acrostichal setae present; vein M1+2 unforked (Figs. 3–5); hind tarsomere 1
elongate, at least as long as tarsomeres 2–4 combined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
– Acrostichal setae absent; vein M1+2 forked (Figs. 6–8); hind tarsomere 1 much shorter than
tarsomeres 2–4 together. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

3. Vein R1 bearing series of short spines dorsally along most of its length (Fig. 3); midtibia with two
(anteroventral and posteroventral) apical spurlike setae; Durango, Mexico. . [Callomyia Meigen]
– Vein R1 without spines; midtibia with only one (posteroventral) apical spurlike seta. . . . . . . . . . 4

4. Crossvein r-m more basally situated and vein Sc more elongate so that r-m is level with middle
portion of cell c (Fig. 4); postsutural supra-alar seta present above wing base in front of postalar
callus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Agathomyia Verrall
– Crossvein r-m close to level of tip of vein Sc, which is relatively shorter (Fig. 5); postsutural
supra-alar seta absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bertamyia Kessel

5. Larger flies, 5.0–10.0 mm long, sometimes partly yellow in color; males with hind tarsus bearing
elaborate ornamentation; two notopleural setae present. . . . . . . . . . . . . . Calotarsa Townsend

– Smaller flies, 2.0–5.0 mm long, with dark coloration predominating; male hind tarsus flattened
but not ornamented; three or more notopleural setae present . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

6. Head bristling, extending from lower part of occiput onto parafacial, almost or quite reaching
level of antennae; cell cup tapered apically, usually longer than vein beyond (Fig. 7); male
genitalia with hypandrial lobes rounded, large apically bifurcate phallapodeme protruding
between them. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Platypeza Meigen
– Head bristling, at most extending a short distance onto gena from occiput, parafacial bare; cell
cup not so strongly tapered apically, at most as long as vein beyond (usually shorter; Fig. 8);
male genitalia with hypandrial lobes variously produced flanking phallus, phallapodeme absent.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lindneromyia Kessel

Synopsis of the fauna Agathomyia Verrall. Williston (1903) described Callomyia

bella from a single female from Guerrero province, Mexico.
The only previous records of Platypezidae from Central This species was transferred to Agathomyia by Buegler (1974).
America are of the genera Agathomyia and Calotarsa from Other species of this genus develop in polypore fungi.
tropical Mexico and Microsania from Panama, but six spe-
cies in five genera (Microsania, Bertamyia, Calotarsa, Platy- Bertamyia Kessel. A single species, B. notata (Loew), has
peza, and Lindneromyia) have now been examined from been recorded from Canada to Argentina (Kessel, 1970), and a
Costa Rica. Although several Neotropical species have been few females have been examined from Costa Rica. It develops
described in Platypeza, this is the first confirmed occurrence in the soft polypore fungus Tyromyces growing on wood (Kes-
of the genus in this region. A further genus (Callomyia) ex- sel, 1957). Woodley (2005) recorded this species from Costa
tends from the Nearctic Region into northern Mexico. Rica, Panama, Mexico (Chiapas), and Caribbean islands.
718
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Sc
pterostigma R1
Sc R1
R2+3
R2+3
R4+5
R4+5

M1 M1+2
M2

CuA1 CuA1
A1+CuA2 A1+CuA2

2 Microsania 3 Callomyia
c

r-m

dm-cu

4 Agathomyia 5 Bertamyia

M1

M2

6 Calotarsa 7 Platypeza

8 Lindneromyia

Figs. 50.2–8. Wings: dorsal view of (2) Microsania sp., (Nearctic, MND, fig. 50.3); (3) Callomyia calla Kessel, (Nearctic, MND, fig. 50.4);
(4) Agathomyia canadensis Johnson, (Nearctic, MND, fig. 50.6); (5) Bertamyia notata (Loew), (MND, fig. 50.5); (6) Calotarsa insignis
Aldrich, (Nearctic, MND, fig. 50.15); (7) Platypeza consobrina Zetterstedt, (Palearctic, MND, fig. 50.18); and (8) Lindneromyia agarici
(Willard), (Nearctic, MND, fig. 50.12, as Plesioclythia).
PLATYPEZIDAE (flat-footed flies) 50 719

Calotarsa Townsend. Kessel & Young (1974) described Microsania Zetterstedt. There is one published record
Calotarsa mexicana from a single male from Chiapas province, from Panama without further details (Melander, 1922), but
Mexico, confirming that this genus of six species previously some material has now been examined from Panama, Dominica,
known only in the Nearctic extends into the northern Neotropical and Costa Rica. Melander assigned his record to the European
Region. In the same paper C.  durangoensis was described species M.  stigmaticalis Zetterstedt, but he considered New
from Durango province, Mexico, from the female only. The World species of the genus to be conspecific with European
single female of this genus examined from Costa Rica is species. However, Malloch (1935) decided that the two known
5 mm long and probably represents an undescribed species. Nearctic species were distinct and revived the name imper-
It has a grey and black-banded abdomen and most resembles fecta (Loew) for stigmaticalis sensu Melander, but could not
the female of the Nearctic C. pallipes (Loew) in coloration, confirm that the Panama record related to this species. Aczél
but has a strongly darkened wing tip. The male of C. mexicana (1958) also used the European names for South American
has clear wings like the four Nearctic species, while the material of the genus, and the Neotropical fauna of Microsania
female of C. durangoensis has an orange ground color to the still awaits revision. The genus is cosmopolitan, and Micro-
abdomen and a darkened wing tip. A Neotropical origin for sania is the only member of the family to occur in New
this genus is plausible, given its absence from the Old World, Zealand. Larval biology is unknown, but adults are usually
but its closest ally, Protoclythia Kessel, has a Holarctic found around smoke in which the males form aerial swarms.
distribution. One species, C.  insignis Aldrich, develops in
honey fungus (Armillaria species) growing on wood. Platypeza Meigen. Three males and two females of one
species have been examined from Costa Rica. This is probably
Lindneromyia Kessel. Several females, probably belonging an undescribed species, but the several species found in North
to two species, have been seen from Costa Rica. Most America require revision. Although many species from all
described Neotropical Platypezidae belong to this genus, regions were initially described in Platypeza, only two species
which occurs in all regions. It is not yet possible to say if from Burma were previously confirmed to exist outside the
these Costa Rican specimens are of described species. Holarctic Region (Chandler, 1994). Platypeza species develop
Species of this genus mainly develop in terrestrial gill fungi, in a range of gill fungi but especially in honey fungus
especially of the genus Agaricus. (Armillaria species) growing on wood.

Literature cited
Aczél, M.L. 1958. Generos y especies Argentinas de las Clythiidae Kessel, E.L. 1970. Bertamyia, a new genus of Platypezininae (Diptera:
(Platypezidae, Diptera) con catalogo de las especies neotropicales. Platypezidae). Wasmann Journal of Biology 28: 185–190.
Acta Zoologica Lilloana 15: 47–82. Kessel, E.L. 1987. Platypezidae, pp. 681–688. MND, Volume 2.
Buegler, M.E. 1974. A redescription of the Callomyia bella Kessel, E.L. & D.R Young. 1974. The genus Calotarsa. Description of
Williston type and its transfer to the genus Agathomyia (Diptera: three new species and range extension of C. calceata (Snow)
Platypezidae). Wasmann Journal of Biology 32: 95–97. (Diptera: Platypezidae). Wasmann Journal of Biology 32:
Chandler, P.J. 1994. The Oriental and Australasian species of 147–172.
Platypezidae (Diptera). Invertebrate Taxonomy 8: 351–434.
Malloch, J.R. 1935. The North American species of the dipterous
Chandler, P.J. 2001. The flat-footed flies (Diptera: Opetiidae and
genus Microsania. Stylops 4: 65–66.
Platypezidae) of Europe. Fauna Entomologica Scandinavica.
Melander, A.L. 1922. Microsania, a genus of the Platypezidae.
36: 1–276. Brill, Leiden, Boston, Köln.
Chandler, P.J. & A.I.  Shatalkin. 1998. Family Platypezidae, pp. Psyche 29: 43–48.
27–49. In Papp,  L.  & B.  Darvas (editors). Contributions to a Williston, S.W. 1903. Diptera, 2, p. 89. In Godman, F.D. &
manual of Palaearctic Diptera. Volume 3. Higher Brachycera. O. Salvin (editors). Biologia Centrali-Americana. London.
Science Herald, Budapest, 880 pp. Woodley, N.E. 2005. Distributional notes on Bertamyia notata
Kessel, E.L. 1957. Distribution and variation in Agathomyia notata (Loew), including the first report of the family Platypezidae
(Loew) (Diptera: Platypezidae). Wasmann Journal of Biology from the Caribbean (Diptera: Platypezidae). Proceedings of the
15: 69–80. Entomological Society of Washington 107: 731–732
720
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
LONCHOPTERIDAE (SPEAR-WINGED FLIES) 51
Brian V. Brown

Fig. 51.1. Female of Lonchoptera bifurcata (Fallén), (Holarctic, MND, fig. 49.1, as Lonchoptera furcata (Fallén).).

Diagnosis R2+3 and R4+5 converging on wing apex; vein A1+CuA2 ending in
wing margin in male, but ending in CuA1 in female. Femora
Small flies (body length 2–4 mm). Frons broad, with and tibiae with large, bristlelike setae. Male foretarsomeres
three pairs of large setae: lower interfrontal setae, ocellar se- often modified with thickened setae, tarsomeres 4 and 5
tae, and upper orbital setae (Fig. 1). Vertex with two pairs curled basally in dried specimens. Male genitalia with
of convergent inner vertical setae that are continuous with surstyli absent.
row of large ocular, genal, and vibrissal setae, and one Lonchopterids are extremely distinctive and unlikely to
pair of convergent to parallel postocellar setae; some or be confused with any other taxa. The only other families
all of these setae often yellow instead of usual black. First with such pointed wings are psychodids, which have long,
flagellomere short, rounded, arista subapical. Wing narrow, many-segmented antennae, and some phorids whose wings
apically pointed (Fig. 2). Wing venation distinctive, with are reduced and lack all but vestiges of posterior veins.

721
722
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Biology were not accepted by Klymko & Marshall (2008). Grimaldi

& Cumming (1999) described two genera of possible fossil
The little that is known about the natural history and lonchopterids from Cretaceous Lebanese amber, but there is
immature stages of these flies is summarized by Peterson otherwise no fossil record for the group. Recent summaries
(1987), Ferrar (1987), and Klymko & Marshall (2008). on the classification of this family include Barták (1998),
Larvae are considered to be scavengers, found in decay- Peterson (1987), and Klymko & Marshall (2008).
ing vegetation. Adults are found in many habitats, but are
apparently not common in Central America. They feed on Identification
fungi, nectar, pollen, and dead insects (Klymko & Marshall,
2008). At least one species, L. bifurcata (Fallén), is usually There are few recent studies on the taxonomy of New
parthenogenetic, although males are sometimes produced World Lonchopteridae. The North American species were
(e.g., Andersson, 1970). treated by Curran (1934) and recently revised by Klymko &
Marshall (2008). Papavero (1967) listed L. bifurcata (Fallén)
Classification (as L. furcata (Fallén)) from Chile and Argentina, and this
is assumed to be the only species present in the Neotropical
Lonchopteridae are classified at the base of the Cyclor- Region
rhapha, usually placed in a group with some of the families
Opetiidae, Platypezidae, Ironomyiidae, and Phoridae Synopsis of the fauna
(Cumming et al., 1995; Collins & Wiegmann, 2002). Collins
& Wiegmann (2002) found some evidence that they are the Lonchoptera Meigen. I have seen only a single female
sister group of Opetiidae. specimen from our region, from La Rosa, Nicaragua
The family is usually considered to consist of a single (Canadian National Collection). It probably belongs to the
extant genus, Lonchoptera, although Andersson (1984) cosmopolitan, usually parthenogenetic species L. bifurcata,
apparently spoke about, but did not publish, a reclassification but identification of females is problematic. From adjacent
into eight genera. More recently, Vaillant (1989) proposed areas, Klymko & Marshall (2008) list L. uniseta Curran from
recognizing a total of three genera, but the idea has received the southern USA, and I have seen Lonchoptera females
little support (e.g., Andersson, 1991; Barták, 1998; Whit- from Colombia and Peru and a single male from Desierto los
tington, 1991). Yang (1999) described two further genera that Leones, Distrito Federal, Mexico.

R1
Sc R2+3

r-m R4+5
CuA2
CuA1 M1

cup M2

A1+CuA2 A1+CuA2+CuA1

2 Lonchoptera

Fig. 51.2. Wing: dorsal view of (2) Lonchoptera bifurcata (Fallén), (Holarctic, MND, fig. 49.2, as Lonchoptera furcata (Fallén).)

Literature cited
Andersson, H. 1970. Notes on north European Lonchoptera (Dipt., Barták, M. 1998. 3.1. Family Lonchopteridae, pp. 13–16. In
Lonchopteridae) with lectotype designations. Entomologisk Papp, L. & B.  Darvas (editors). Contributions to a manual
Tijdskrift 91: 42–45. of Palaearctic Diptera, Volume 3. Science Herald, Budapest,
Andersson, H. 1984. Evolution within the genus Lonchoptera 880 pp.
(Diptera: Lonchopteridae [abstract], p. 31. In XVII International Collins, K.P. & B.M. Wiegmann. 2002. Phylogenetic relationships
Congress of Entomology, Abstract Volume. Hamburg, Germany. of the lower Cyclorrhapha (Diptera: Brachycera) based on
Andersson, H. 1991. Family Lonchopteridae, pp. 139–142. In Soós, 28S rDNA sequences. Insect Systematics and Evolution 33:
. & L.  Papp. Catalogue of Palaearctic Diptera, Volume  7. 445–456.
Akadeémiai Kiadó, Budapest.
LONCHOPTERIDAE (spear-winged flies) 51 723

Cumming, J.M., B.J. Sinclair, & D.M. Wood. 1995. Homology Papavero, N. 1967. 41. Family Lonchopteridae. In Papavero, N.
and phylogenetic implications of male genitalia in Diptera (editor). A catalogue of the Diptera of the Americas south of
— Eremoneura. Entomologica scandinavica 26: 120–151. the United States. Departamento de Zoologia, Secretaria da
Curran, C.H. 1934. The North American Lonchopteridae (Diptera). Agricultura, São Paulo, 2 pp.
American Museum Novitates 696: 1–7. Peterson, B.V. 1987. 49. Lonchopteridae, pp. 675–680. MND,
Ferrar, P. 1987. A guide to the breeding habits and immature Volume 2.
stages of Diptera Cyclorrhapha. Entomonograph 8, 2 vols. Vaillant, F. 1989. Contribution à l’étude des Diptères Lonchopteridae
E.J. Brill / Scandinavian Science Press, Leiden, Copenhagen, d’Europe et d’Afrique du Nord. Bulletin de la Société Vaudoise
907 pp. des Sciences Naturelles 79: 209–229.
Grimaldi, D. & J. Cumming. 1999. Brachyceran Diptera in Cretaceous Whittington, A.E. 1991. Two new Afrotropical species of Lonchoptera
amber and Mesozoic diversification of the Eremoneura. Bulletin Meigen (Diptera: Lonchopteridae). Annals of the Natal Museum
of the American Museum of Natural History 239: 1–124. 32: 205–214.
Klymko, J. & S.A. Marshall. 2008. Review of the Nearctic Yang, C. 1999. Lonchopteridae of China (Diptera), pp. 49–59. In
Lonchopteridae (Diptera), including the descriptions of three Xue, W. & C. Chao (editors). Flies of China, Vol. 1. Liaoning
new species. The Canadian Entomologist 140: 649–673. Science & Technology Press, Shenyang, 1365 pp.
724
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
PHORIDAE (HUMP-BACKED FLIES, SCUTTLE FLIES) 52
Brian V. Brown

Fig. 52.1. Male of Phora occidentata Malloch, (Holarctic, MND, fig. 51.1).

Diagnosis Eye, wing, and some abdominal tergites reduced or absent

in females of some genera. Abdominal venter usually lack-
Small (body length 0.5–6.0  mm), compact, often con- ing sternites, except for segments 4–6 in some genera; ovis-
spicuously setose flies (Fig. 1). Head usually with 12 frontal cape various, largely membranous to strongly sclerotized,
setae and one or two (rarely more) pairs of supra-antennal and highly modified in parasitic species. Larvae various,
setae, although one or more pairs of setae lacking in many from elongate–cylindrical to oval and flattened, often with
species. Pedicel of antenna usually concealed within first elongate processes. Pupae with well-developed respiratory
flagellomere. Tibiae often with large setae, longitudinal horns.
rows of enlarged setulae (setal palisades), or transverse The wing venation of phorids separates them from most
rows of enlarged setulae (ctenidia). Hind tarsomere 1 with other families, although some species of primitive Diptera
with long antennae, such as Simuliidae, Scatopsidae, and
setal palisades. Wing with distinctive venation composed Cecidomyiidae, are superficially similar. Wingless females
of thick costal and radial veins and thin, linear posterior can be confused with some Sphaeroceridae, but lack a
veins, without crossveins between them. Alula of wing ptilinal suture and have the pedicel of the antenna largely
poorly developed, but alular margin often with strong setae. hidden.

725
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Biology radiation of “bee-killing flies”, genus Melaloncha, is endem-

ic to the Neotropical Region. Some parasitoids of beetles
The natural history of this group has been comprehen- (Cantharidae, Coccinellidae, Lampyridae, Passalidae) and
sively reviewed by Disney (1994). Phorids are among the scale insects (Margarodidae) are also known. Species from
most biologically diverse families of insects, with larvae Europe attack larvae of Diptera (Bibionidae, Sciaridae, Tipu-
that are scavengers, herbivores, predators, kleptoparasites, lidae), and this likely also occurs elsewhere in the world. Old
parasitoids, and true parasites. Adults are less varied in their World termites are attacked by a few phorids, but this has not
feeding habits, apparently subsisting mostly on honeydew yet been found in the Neotropical Region.
on undergrowth leaves, or on nectar, yeasts, carrion fluids, Many phorids have close relationships with other organ-
fungal spores, or pollen. Some are known to be predators isms. Army ant colonies often have large numbers of associ-
on other insects (e.g., Compton & Disney, 1991), to feed on ated phorids, from parasitoids that attack the ant victims of the
dead insects (e.g., Disney, 1985), or to feed on oviposition raids or the raiders themselves (Brown  & Feener, 1998), to
wounds on their hosts (e.g., Brown & Feener, 1991, 1993). those that live within the colony as predators, parasitoids, or
Scavenging phorids, especially Megaselia scalaris (Loew) scavengers (Disney & Kistner, 1989a; Rettenmeyer & Akre,
(reviewed by Disney, 2008a), Dohrniphora cornuta (Bigot) 1968). Many of the latter have unusually modified wingless or
(Barnes, 1990), and Puliciphora borinquenensis Wheeler brachypterous females. A similar, but smaller, fauna is found
(Miller, 1984), are commonly associated with various types in the nests of other social Hymenoptera and termites (e.g.,
of filth, as well as buried carrion, dead invertebrates, and de- Disney  & Kistner, 1989b, 1997, 1998; Salt, 1929; Seevers,
caying plant material, including fruit. Some have been re- 1941). Females of Conicera have been found in the shells of
corded invading wounds on animals, including humans. In living giant snails (Brown, 2000a), but the relationships be-
particular, M. scalaris can be a pest of captive cockroaches, tween the host and flies are unknown. Similarly, adults of a
crickets, tarantulas, hermit crabs, and lizards, attacking in- Johowia species have been reared from puparia found under
jured individuals. the elytra of passalid beetles in Guatemala and Costa Rica, but
A few phorids are known to attack living higher plants, the beetles seem to be undamaged by the flies, and whether the
such as Megaselia species on Aristolochia (Hime & Costa, flies feed on the beetles is unknown (Brown, 2000a).
1985; Disney & Sakai, 2001), violet seeds (Viola; Violaceae) Relatively few phorids have a direct impact on humans.
(Robinson, 1970), and Smilacina leaves (Lilliaceae) (Robin- Some species of Megaselia are pests in mushroom farms,
and some, especially Pseudohypocera kerteszi (Enderlein),
son, 1967). Others, for instance those associated with corn
invade and attack the nests of both honey bees and native
kernels, are found in situations where fungi or bacterial in-
bees kept for apiculture (Chaud-Netto, 1980; Hernández &
vasions have already begun the process of decay. There are
Gutiérrez, 2001; Portugal-Araújo, 1977; Reyes, 1983; Rob-
many records of phorids (mostly Megaselia species) breed- inson, 1981). Species of Melaloncha also attack honey bees,
ing in fungi in the Northern Hemisphere, but similar studies although their impact on colony viability is unknown (Brown,
have not been carried out in the Neotropical Region. 2004a and unpublished observations; Ramírez, 1984; Ronna,
Few predatory species of phorid larvae are known from 1936, 1937). One species of Megaselia has been reared from
the Neotropical Region, but in other parts of the world there corn (Borgmeier, 1962 and personal observation in Califor-
are those that attack Diptera larvae (Deleporte, 1986; Kloter nia, USA), but probably only after invasion of the crop by a
et al., 1977; Robinson & Brown, 1993), aphids (Robinson & fungus. Some phorids are known to attack infected wounds
Brown, 1993; Yarkulov, 1972), and eggs of slugs (molluscs), on humans and other animals, and intestinal myiasis by lar-
spiders, and insects (Disney, 1979, 1994; Robinson & Foote, vae of M. scalaris is commonly reported. Synanthropic spe-
1968). Some phorids found in army ant nests will feed on cies such as Megaselia scalaris and Dohrniphora cornuta
living ant larvae (Rettenmeyer & Akre, 1968). There are also have the potential to mechanically transmit pathogens to
some records of phorids attacking turtle and frog eggs (Au- food and directly to humans. Some South American species
cuña-Mesén & Hanson, 1990; Davis & Disney, 2003; Down- of Pseudacteon are being used for biological control of intro-
ie et al., 1995; Villa & Townsend, 1998), probably some of duced fire ants (Solenopsis) in the southern USA (e.g., Mor-
which were damaged or invaded by fungi prior to attack. rison, 2000), which has spurred a wealth of new research on
Worldwide, parasitic species (including parasitoids) of this phorid genus.
several phorid genera attack a variety of invertebrates, in-
cluding annelid worms, molluscs, arachnids, millipedes,
centipedes, and insects. In Central America, phorids attack Classification
mostly social Hymenoptera (ants, bees, and wasps), with
some hosts attacked by multiple species (Brown, 1999). Phoridae are classified at the base of the Cyclorrhapha, as
Ant-parasitizing species of the genus Apocephalus are called the sister group of the Ironomyiidae, and with various relation-
“ant-decapitating flies” for their habit of developing in the ships to the families Lonchopteridae, Platypezidae, and Opetii-
host head, sometimes causing decapitation before the rest of dae (Brown, 1992; Cumming et al., 1995; Collins & Wiegmann,
the ant body stops moving (this common name is sometimes 2002). The formerly separate family Sciadoceridae is now treated
applied to other ant-parasitizing genera as well). A large as a subfamily (Sciadocerinae) of the Phoridae (Disney, 2001).
PHORIDAE (hump-backed flies, scuttle flies) 52 727

Extant phorids excluding Sciadocerinae are grouped into a higher The Metopina-group of genera has distinctive females
taxon called Euphorida (Brown, 2007). that are usually brachypterous or wingless, with reduced
The details of phorid intrafamilial classification are in dis- eyes and often reduced abdominal tergites. They are almost
pute. The traditional arrangement of New World Euphorida impossible to match with their much more commonly col-
into three subfamilies, Phorinae, Aenigmatiinae, and Meto- lected, but currently unidentifiable, males. Rearing of off-
pininae (Schmitz, 1929), is inadequate because the mono- spring from captured females, collecting males and females
phyly of the Phorinae has not been demonstrated and the in copula, or matching molecular sequences is necessary to
Aenigmatiinae is polyphyletic (three other subfamilies pro- associate the two sexes of these many species and to decide
posed by Schmitz do not occur in the New World, and one on a reasonable classification for them.
was subsequently abolished). Brown (1992) presented a dif- In spite of the above problems, phorid identification re-
ferent classification with five subfamilies  — Hypocerinae, ceived a tremendous boost with the publication of Disney’s
Phorinae, Aenigmatiinae, Conicerinae, and Metopininae — (1994) key to world genera, the first such resource since
but the validity of this arrangement has been contested by earlier in the twentieth century (Schmitz, 1929). Because
Disney (1994, and elsewhere). Furthermore, relationships of the many undescribed species, Disney’s key will not suc-
within the subfamilies are also contentious. More research cessfully work for all specimens, but it will identify most.
is necessary to resolve this situation, including the use of The following key takes into account some of the newly
molecular characters, which has been initiated by both Cook collected, but undescribed, species of various genera, but
et al. (2004) and Brown & Smith (in preparation). it still suffers many of the shortcomings of other published
Within the Metopininae, there are two especially prob- phorid keys.
lematic groups: Megaselia and the Metopina-group of Each method of preservation has advantages and dis-
genera. The genus Megaselia is the largest in the family, ac- advantages in terms of which characters are most eas-
counting for nearly one-half of the described species, and ily observed. Disney advocates slide-mounting of phorid
species within it can be extremely difficult to resolve. There specimens, and his key uses some characters that are difficult
are hundreds, if not thousands, of undescribed species within to see without such preparation. I do not advocate this
Megaselia, making published keys almost worthless. Anoth- type of preparation for all phorids and do not use such
er troubling aspect is that recognition of some other genera characters in my key. In my opinion, phorids should be
probably renders Megaselia paraphyletic (Brown, 1992; Dis- collected into alcohol and dried using a critical-point-drier
ney, 1989). Disney (2003a) classified Megaselia and related or special drying chemicals like HMDS (hexamethyldisila-
genera in tribe Gymnophorini. zane) (Brown, 1993b), or less optimally, ethyl acetate. If
The Metopina-group of genera (sensu Brown, 1992, = such facilities are not available, specimens can be main-
Metopinini of Disney, 2003a) consists of many genera, most tained in alcohol. Slide-mounting of smaller species (of
of which have apterous or brachypterous females. Many are genera such as Puliciphora, Chonocephalus, and others) is
known from one sex only and need to be critically examined often necessary, but limits the possibility of further exami-
to see if they can be justified. Some of the included genera nation of complex, three-dimensional structures like male
have already been synonymized with Puliciphora by Disney terminalia.
(2003b, and elsewhere). Important characters for phorid identification at the ge-
nus level are found mostly in the setation of the frons, wing
venation and form, setation of the legs (mostly tibiae), and
Identification shape of the first antennal flagellomere. The setation of the
frons can be confusing; generally, there are 12 frontal se-
Determining the identity of any phorid specimen is a difficult
tae, roughly arranged in three horizontal rows (referred to as
task because of three factors: the huge abundance of undescribed
“4-4-4” in the phorid literature; refer to Fig. 22). The ventral
taxa, the overwhelming dominance of genus Megaselia, and the
lack of work on the Metopina-group of genera. Worldwide (ex- row consists of the two lower fronto-orbital setae and the
cept for the European fauna), it is likely that 90% of all species two lower interfrontal setae, the middle row is the two upper
are undescribed, and keys published prior to the 1980s, based fronto-orbital and the two upper interfrontal setae, and the
on limited material, are of limited value. More recent keys have dorsal row is the two inner vertical setae and two postocel-
covered a larger percentage of the extant species, but they still lar setae. In many, but not all, phorids there are two or more
represent an incomplete assessment of the entire fauna. Most additional ventral setae near the middle of the frons, termed
of the described species are cataloged by Borgmeier (1968a, supra-antennal setae. Some or all frontal setae can be absent,
1971b), with an update by Brown (1990). leading to confusion when it is not clear if a pair of ventral,
The genus Megaselia comprises about one-half of the de- medial setae on the frons are reclinate supra-antennals or
scribed phorid species. Its monophyly is doubtful (Brown, lower interfrontal setae.
1992; Disney, 1989), but recognition of subordinate groups Further resources for the study of phorids are found in
is complicated by our current ignorance of informative issues of the Phorid Newsletter, available at the author’s
structural characters. website, currently at www.phorid.net.
728
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Key to the genera of Phoridae of Central America

The females of the Costa Rican species of Macrocerides Borgmeier are unknown. The known females of this genus are in
species doubtfully related to the type species of the genus, and thus are not keyed out here.
There are many undescribed genera that will not key out properly in this key. Any identification should be checked against
voucher specimens or illustrations.

1. Midtibia with large isolated (not part of a series) basal setae, usually in basal pair of one anterodorsal
and one dorsal seta (Figs. 2, 5), but in some genera single anterodorsal seta present; supra-antennal
setae, if present, usually reclinate (pointed upwards) (Fig.  13), but one exceptional genus has
proclinate (downward-pointed) setae (as in Fig. 14); anepisternal furrow almost always absent
(Fig. 57; exception is males of Xanionotum). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Midtibia without isolated basal setae, although sometimes with large setae in series (as in Fig. 6);
supra-antennal setae, if present, usually proclinate (Fig. 14; exceptions occur); anepisternal
furrow usually present (Fig. 58). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

2. Midtibia with single basal seta. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

– Midtibia with at least basal pair: one dorsal, one anterodorsal seta (Figs. 2, 5) . . . . . . . . . . . . . . 4

3. Male with greatly elongate pointed first flagellomere; wing vein R2+3 present (as in Fig. 32);
female with well-developed wings and lacking long setae on abdominal tergites . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceratusa Borgmeier, in part
– Male with first flagellomere rounded, only slightly pointed at apex; wing with vein R2+3 absent (as
in Figs. 35, 38); female with wing vestigial and abdominal tergites with transverse row of dense
elongate setae (Fig. 59) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xanionotum Brues, in part

4. Ocelli arranged in nearly straight horizontal line, widely separated, with lateral ocelli closer to eye
margin than anterior ocellus (Fig. 60); ocellar region of frons demarcated by transverse groove,
often outlining rounded lobe for each ocellus. . . . . . . . . . . . . . . . . . . . . . . . . . Stichillus Enderlein
– Ocelli arranged in triangle (one anterior, two posterior), closer together; ocellar region not
outlined into lobes (as in Figs. 11–16). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

5. Hind tibia with one or two dorsal longitudinal setal palisades (Fig. 3). . . . . . . . . . . . . . . . . . . . . 6
– Hind tibia without dorsal longitudinal setal palisades (although transverse ctenidia present in
some genera) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

6. Hind tibia with one dorsal longitudinal setal palisade. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

– Hind tibia with two dorsal longitudinal setal palisades. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

7. Wing with broad area of thickening between apices of veins R1 and R4+5 (Fig. 31); vein R2+3 not
distinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hypocerides Schmitz
– Wing without broad thickening; vein R2+3 distinct (Fig. 32). . . . . . . . . . . . . . . Dohrniphora Dahl

8. Supra-antennal setae present (as in Fig. 13); R2+3 usually present; male cercus on elongate and
narrow stalk, at least as long as length of epandrium (Fig. 61). . . . . . . . . . . . . . . Diplonevra Lioy
– Supra-antennal setae absent; wing vein R2+3 usually absent; male cercus shorter than length of
epandrium. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

9. First flagellomere of male drawn out into long thin apical, highly setose process (Fig. 9); female
with elongate thin mouthparts that extend about 3 times length of head and with frons greatly
extended ventrally between antennae (Fig. 10). . . . . . . . . . . . . . . . . . . . . . Abaristophora Schmitz
– First flagellomere rounded; female mouthparts not so elongate . . . . . . . . . . . . . . . . . . . . . . . . . 10
PHORIDAE (hump-backed flies, scuttle flies) 52 729

s pal
bas pr s
cten
bas pr s

ant s

ant s

2 Triphleba 3 Diplonevra 4 Hypocera 5 Spiniphora

a sp s

notch

proc

7 Apterophora

notch

6 Phalacrotophora 8 Eurycnemis

Figs. 52.2–8. Legs: anterior view of left midleg of (2) Triphleba pachyneura (Loew), (Nearctic, MND, fig. 51.67); anterior view of left hind
tibia and tarsus of (3) Diplonevra nitidula Meigen, (Holarctic, MND, fig. 51.63); anterior view of left hind leg of (4) Hypocera erhmanni
Aldrich, (Nearctic, MND, fig. 51.70); anterior view of left midleg of (5) Spiniphora spinulosa (Malloch), (Nearctic, MND, fig. 51.66); anterior
view of left hind tibia and tarsus of (6)  Phalacrotophora longifrons (Brues), (MND, fig.  51.77); anterior view of left foreleg of
(7) Apterophora sp.; posterodorsal view of left tibia and tarsus of (8) Eurycnemis sp. Figures 7–8 illustrated by B. Koehler.
Abbreviations: ant s, anterior seta; a  sp  s,  anterior spinelike seta; bas  pr  s,  basal pair of setae; cten,  ctenidia; proc,  process; s  pal,  setal
palisade.
730
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

10. M1 strongly curved, such that base of M1 perpendicular to midpoint; vein Rs slightly convex
at origin of M1 (Fig.  76); female with venter of abdomen yellow; costa often greatly thickened
(Fig. 77). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Latiborophaga Brown
– M1 less curved, origin and midpoint forming obtuse angle; vein Rs slightly concave at origin;
female with venter of abdomen black, costa not thickened. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

11. Frons broad, about one-half width of head; midtibia with single anterior seta in basal half. . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Borophaga Enderlein
– Frons narrowed, about one-third (females) to one-quarter (males) head width; midtibia with one
or two anterior setae in basal half . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trineurocephala Schmitz

12. Wing vein R2+3 absent or consisting of small remnant attached to costa but not extending to Rs . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
– Wing vein R2+3 present, extending from costa to Rs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

13. Hind tibia with patches of ctenidia (as in Fig. 4). . . . . . . . . . . . . Chaetocnemistoptera Borgmeier
– Hind tibia without ctenidia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

14. Hind tibia with basal pair of setae: one anterodorsal, one dorsal; male first flagellomere elongate,
pointed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Conicera Meigen
– Hind tibia without basal pair of setae; male first flagellomere various. . . . . . . . . . . . . . . . . . . . 15

15. Midtibia, besides basal pair, with one or more dorsal or near dorsal setae; costa darkened; eye
large and prominent, frons narrow; body color velvety black. . . . . . . . . . . . . . . . . Phora Latreille
– Midtibia, besides basal pair, without dorsal setae (anterior setae may be present); other characters
various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

16. Wing vein Rs with small posterior swelling at origin of M1 (Fig.  78); epandrium of male with
greatly thickened setae on left side (Fig. 40); worldwide, including Caribbean, Brazil. . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Gymnoptera Lioy]
– Wing vein Rs smooth, without posterior process; epandrium of male without greatly thickened
setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

17. Anteroventral seta of apex of midtibia thick and longer than thickness of tibia (as in Fig. 5); male
foretarsomere 1 with thin apical process. . . . . . . . . . . . . . . . . . . . . . . . . Coniceromyia Borgmeier
– Anterior to anteroventral subapical seta of midtibia small, thin, about as long as thickness of tibia
(Fig. 2); foretarsomere 1 of male not modified . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

18. Frons with lower fronto-orbital seta present, in usual position beside eye (as in Fig. 60); male
terminalia with posterodorsal area of epandrium produced posteriorly, ending freely, often in
elongate processes; cerci short, not longer than anterior portion of epandrium; base of cerci
without epandrial ring . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Triphleba Rondani, in part
– Frons apparently lacking lower fronto-orbital seta (Fig. 11); male terminalia with posterodorsal
area of epandrium narrow, continuous below cerci, not produced posteriorly; cerci relatively
elongate; base of cerci with short epandrial ring (Fig. 41) . . . . . . . . . . . Tapantia Brown & Kung

19. Anepisternum with long thick single seta (often missing, but the socket still visible) in addition to
small setulae (Fig. 62) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neopleurophora Brown
– Anepisternum bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

20. Hind tibia without large preapical setae; male with extremely elongate, pointed first flagellomere.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceratusa Borgmeier, in part
– Hind tibia with at least one large preapical seta. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
PHORIDAE (hump-backed flies, scuttle flies) 52 731

21. Wing vein Rs with dorsal row of small setulae along most of length. . . . . . . . . . . . Anevrina Lioy
– Wing vein Rs with at most few basal seta. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

22. Hind tibia with large ventral seta at midlength; hypothetical in Central America, one species
introduced worldwide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Spiniphora Malloch]
– Hind tibia without such seta at midlength. . . . . . . . . . . . . . . . . . . . . . . Triphleba Rondani, in part

23. Hind tibia with one or more dorsal or near dorsal setal palisades (Fig. 3), including genera with
several irregular rows. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
– Hind tibia without dorsal setal palisades. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87

24. Hind tibia with more than one dorsal setal palisade, or numerous incomplete palisades forming
meshwork of setulae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
– Hind tibia with one dorsal setal palisade. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

25. Hind tibia with two straight, parallel, dorsal setal palisades (as in Fig. 3); frons with narrow
ventral process (Fig. 27). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cataclinusa Schmitz, in part
– Hind tibia with two or more dorsal setal palisades that are curved, sinuous, or that often intersect;
frons not as above. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26

26. Forefemur lacking large posterior setae at apex; frons at least as broad as high, with 12 large setae
and one or two pairs of supra-antennal setae; wing vein R2+3 present. . . . . . . . . . . . . . . . . . . . . . . 27
– Forefemur with one or two large posterior setae at apex; frons either extremely broad and with no
large setae, or narrow and lacking only supra-antennal setae (i.e., with 10–12 large setae); wing
frequently lacking vein R2+3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28

27. Supra-antennal setae point dorsally or laterally (as in Figs. 12–13); male with cercus shorter than
dorsal length of epandrium; female with membranous, non-parasitic oviscape (as in Fig. 43) . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Enderleinphora Disney
– With four ventrally directed supra-antennal setae; male with long, thin cercus (Fig. 63); female
with heavily sclerotized, parasitic oviscape. . . . . . . . . . . . . . . . . . Apocephalus Coquillett, in part

28. Frons narrow, less than one-half of head width, with large setae present at least on vertex. . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Melaloncha Brues
– Frons broad, over one-half of head width; frontal setae mostly absent, some barely differentiated.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Melittophora Brues

29. Alula with 10 or more setae (Fig. 79); anal region of wing well developed; R2+3 present, but weaker
than other radial veins, and fork formed by R2+3 and R4+5 small; Rs with small ventral swelling at
origin of M1; body large, robust, about 2.5 mm long; mostly dark brown in color. . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudohypocera Enderlein
– Alula usually with fewer setae; anal region of wing not as well developed; vein R2+3 and fork various;
body color and size various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

30. Supra-antennal setae well developed and laterally or dorsally directed (Figs. 12–13). (Note: in
this lead, supra-antennal setae are said to be present when one or more pairs of medial setae are
present close to lower margin of frons. In some species, lower interfrontal setae occupy this
position; supra-antennal setae can be said to be present in this instance only if the lower interfrontal
setae are first identified. Generally, supra-antennal setae are only found when there are 4-4-4 other
frontal setae present and accounted for; however, one genus has only supra-antennal setae on the
frons) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
– Supra-antennal setae point ventrally or anteroventrally, strongly differing in direction from other
frontal setae (e.g. Fig. 14), or supra-antennal setae absent (in some instances, their place is taken
by divergent lower interfrontal setae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
732
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

9 Abaristophora 10 Abaristophora 11 Tapantia

spant s
spant s
spant s

clyp

12 Beckerina 13 Dohrniphora 14 Trophithauma

spant s

15 Brownphora 16 Chaetaspidia 17 Cootiphora

Figs. 52.9–17. Heads: anterolateral view of male (9) and female (10) of Abaristophora diversipennis Borgmeier, (Nearctic, MND, male,
fig. 51.5); anterior view of (11) Tapantia bicasa Brown & Kung, (Brown & Kung, 2004, fig. 1); anterolateral view of (12) Beckerina sp.,
(Nearctic, MND, fig.  51.13, as B. umbrimargo); (13)  Dohrniphora cornuta (Bigot), (MND, fig.  51.3); (14)  Trophithauma rostratum
(Melander & Brues), (Nearctic, MND, fig. 51.25); and (15) Brownphora sinefurca (Borgmeier), (Brown, 1997c, fig. 1); anterior view of
(16) Chaetaspidia convexifrons (Borgmeier); and (17) Cootiphora angustata Brown. Figures 10, 16–17 illustrated by L. Mui.
Abbreviations: clyp, clypeus; spant s, supra-antennal setae.
PHORIDAE (hump-backed flies, scuttle flies) 52 733

lbl
20 Myrmosicarius
18 Phymatopterella

l infr s

22 Brachycephaloptera

19 Calamiscus
21 Acanthophorides

l frorb s 25 Apocephalus

l infr s 23 Neophora

24 Lecanocerus

26 Neodohrniphora

Figs. 52.18–26. Heads (continued) and frontal setation: anterior view of head of (18) Phymatopterella sp.; anterodorsal view of head of
(19) Calamiscus cryptopalpis Borgmeier, (Brown, 1997b, fig. 1) and (20) Myrmosicarius texanus (Greene), (Nearctic, MND, fig. 51.21);
anterior view of frons of (21)  Acanthophorides  sp.; (22)  Brachycephaloptera trichopleura Borgmeier; and (23) Neophora frontalis
Borgmeier; anterodorsal view of head of (24) Lecanocerus compressiceps Borgmeier, (Nearctic, MND, fig. 51.15); (25) Apocephalus sp.;
and (26) Neodohrniphora curvinervis (Malloch). Figures 18, 26 illustrated by L. Mui; Figures 21–23 illustrated by B. Brown; Figure 25
illustrated by J. Cantley.
Abbreviations: lbl, labellum; l frorb s, lower fronto-orbital seta; l infr s, lower interfrontal seta.
734
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

31. Body small (length 1  mm), limuloid (teardrop-shaped), and extremely flat; frontal setae
consist solely of single pair of supra-antennal setae; costa extends for more than 0.80 wing length
(Fig. 33); only female known. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tabelliphora Kung & Brown
– Body usually larger, not limuloid; frons with more than one pair of setae; costa much less than
0.80 wing length; males and females. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32

32. Male cercus thin, elongate (longer than dorsum of epandrium; Fig. 63); female with rigid, heavily
sclerotized, largely non-retractile, parasitic-type oviscape (as in Fig. 42); female cerci, if present,
extremely small . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apocephalus Coquillett, in part
– Male cercus thick, usually shorter than dorsum of epandrium, but if longer, not as thin (as in
Fig. 63); female with apex of oviscape mostly membranous, often mostly withdrawn into abdomen,
usually with large cerci (as in Fig. 43; female of Chaetaspidia unknown, but assumed not to be
parasitic). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

27 Cataclinusa 28 Chonocephalus

29 Chonocephalus 30 Trophodeinus

Figs. 52.27–30. Heads (concluded): anterodorsal view of (27) Cataclinusa pachycondylae (Brues), (Nearctic, MND, fig. 51.29); anterodorsal
view of male (28) and dorsal view of female (29) of Chonocephalus sp., (MND, figs. 51.31 and 92); lateral view of (30) Trophodeinus
pygmaeus Borgmeier, (Nearctic, MND, fig. 51.93).
PHORIDAE (hump-backed flies, scuttle flies) 52 735

33. Supra-antennal setae located on lower margin of frons; male with elongate, pointed first
flagellomere (Fig. 15); female with foretarsomeres twice as thick as those on mid- and hind legs.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brownphora Disney
– Supra-antennal setae removed dorsally from lower margin of frons (Fig. 16); first flagellomere of
male more rounded; female unknown. . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetaspidia Borgmeier

34. Wing reduced to tiny remnant, or absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35

– Wing of normal size. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36

35. Wing remnant small, triangular (Fig. 34); body limuloid (teardrop-shaped); eyes extremely small
(Fig. 17); only female known. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cootiphora Brown
– Wing rudiment, if present, smaller and not triangular in shape; eyes larger and body less limuoid
(Fig. 64). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Borgmeieriphora Prado, in part

36. Female with greatly enlarged palpus with about 30–40 large, anteriorly projecting setae at apex
(Fig. 51); head extraordinarily flat and broad; foretibia short, about one-half length of forefemur;
hind femur greatly expanded apically; male unknown . . . . . . . . . . . Laciniomyia Kung & Brown
– Female with many fewer, smaller apical setae on palpus; other characters various; all males. . 37

37. Frons short, with 10 large setae, plus pair of ventrally pointing supra-antennal setae (i.e., one pair
of interfrontal setae apparently absent; Fig.  21); costal setae and all frontal setae, including
supra-antennals, elongate; female with sclerotized, parasitic-type oviscape and expanded clypeus;
male with enlarged, elongate flattened first flagellomere. . . . . . . . . . . Acanthophorides Borgmeier
– Frons with either usual 4-4-4 setae or fewer than 10 frontal setae; with or without ventrally
pointing supra-antennal setae; other characters various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38

38. Frons about as broad as high, densely and finely punctate, shiny; wing relatively short and broad,
wing vein R2+3 absent (Fig. 85); hind tibia with enlarged posterodorsal setae, without enlarged
anterodorsal setae in addition to setal palisade; body robust, strongly hump-backed; male cercus
relatively short; female with parasitic-type oviscape. . . . . . . . . . . . . . . . . . . Apodicrania Borgmeier
– Frons various, but usually not densely, finely punctate, shiny, and subequal in height and width, but
if so (Syneura), then wing veins M1 and M2 close together at base (Fig. 36); other characters
various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39

39. Male with notch in epandrium (Fig. 69); female with parasitic-type (rigid, heavily sclerotized,
largely non-retractile) oviscape usually laterally compressed (although sometimes tubular),
narrow, consisting of conspicuously separate dorsal and ventral sclerite separated by narrow to
extensive, often striate, membrane (Fig.  72); female cercus extremely reduced but present;
parasitoids of millipedes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myriophora Brown
– Male without notch in epandrium; oviscape various, but if of parasitic type, not laterally
compressed, but often dorsoventrally flattened; if narrow, parasitic oviscape not separated into
conspicuous dorsal and ventral sclerites separated by striate membrane (e.g., Fig.  65); female
cercus present or absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40

40. Male with broad, rounded labellum (Fig. 18); female with sclerotized spot between wing veins M1
and M2 or M1 and R4+5 (Fig. 35). . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phymatopterella Brues, in part
– Male with narrow, unmodified labellum; female without sclerotized spot between veins . . . . . 41
736
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

thkg R2+3 R4+5

R1

M1

M2

CuA1

31 Hypocerides 32 Dohrniphora

33 Tabelliphora 34 Cootiphora

M1

M2
scl ptch

35 Phymatopterella 36 Syneura

M1

scl ptch M2

CuA1

37 Physoptera 38 Metopina

R4+5

39 Commoptera

Figs.  52.31–39. Wings: dorsal view of (31)  Hypocerides nearcticus Borgmeier, (MND, fig.  51.44); (32)  Dohrniphora cornuta (Bigot),
(MND, fig. 51.42); (33) Tabelliphora teretipenna Kung & Brown, (Kung & Brown, 2005, fig. 5); (34) Cootiphora angustata Brown, (Brown,
1993a, fig.  20); (35)  Phymatopterella luteiclava Borgmeier, (Borgmeier, 1971a, fig.  127); (36)  Syneura cocciphila (Coquillett), (MND,
fig. 51.52); (37) Physoptera vesiculata (Borgmeier), (MND, fig. 51.54); (38) Metopina subarcuata Borgmeier, (Nearctic, MND, fig. 51.56);
and (39) Commoptera solenopsidis Brues, (Nearctic, MND, fig. 51.60). Figure 33 illustrated by L. Mui.
Abbreviations: scl ptch, sclerotized patch; thkg, thickening.
PHORIDAE (hump-backed flies, scuttle flies) 52 737

41. Male terminalia rotated to left, such that right side of terminalia composed almost solely of
hypandrium and left side completely closed by enlarged epandrium (Fig. 45); female oviscape of
parasitic type, setose, highly complex, asymmetrical, slightly laterally flattened (Fig. 48); wing vein
R2+3 extremely thin; supra-antennal setae absent. . . . . . . . . . . . . . . . . . . . . . . . . Cremersia Schmitz
– Male terminalia with both epandrium and hypandrium visible from each side; female oviscape
various, but usually not asymmetrical (if asymmetrical, as in Apocephalus asymmetricus Brown,
then oviscape dorsoventrally flattened, supra-antennal setae and wing vein R2+3 present and well
developed); other characters various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42

42. Wing veins M1 and M2 close together at base (Fig. 36); R1 and Rs close together and parallel; frons
broad (about one-half head width), shiny, deeply punctate; first flagellomere often greatly elongate;
relatively small flies, body length 1–1.8 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Syneura Brues
– Wing veins M1 and M2 not so closely approximated at base; R1 and Rs usually apically divergent;
frons, first flagellomere and body size various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43

43. Wing vein R2+3 present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44

– Wing vein R2+3 absent or vestigial (attached only to costa, not attached to vein Rs) . . . . . . . . . . . 79

44. Wing vein Rs with posterior projection or swelling at base of M1 (Fig. 80); vein R2+3 thin; brown
species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
– Wing veins not as above, or if similar (Neophora Borgmeier), then body yellow . . . . . . . . . . . 46

45. Male cercus deep in lateral view, almost as deep as epandrium; first flagellomere flat (Fig. 20);
female oviscape elongate, rounded, apically narrowed (Fig. 65) . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myrmosicarius Borgmeier, in part
– Male cercus narrower in lateral view; first flagellomere various; female oviscape more
dorsoventrally flattened. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudacteon Coquillett, in part

46. Male with foretarsomere 1 expanded and rounded, strongly differentiated from other tarsomeres
(Fig. 66); female with sclerotized rounded patch at base of wing membrane (Fig. 37). (Note: some
males of this genus do not have expanded tarsomere 1 and will key out to Megaselia Rondani). .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Physoptera Borgmeier, in part
– Male foretarsomere 1 not markedly expanded, subequal in thickness to other tarsomeres; female
without thickened area at base of wing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47

47. Female with long unjointed extension of face encircling extremely elongate proboscis (Fig. 19);
male cannot be separated from Megaselia. . . . . . . . . . . . . . . . . . . . . . . . . . Calamiscus Borgmeier
– Female proboscis, if elongate, jointed near midlength (as in Fig. 10); males of all other genera. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48

48. Frons with six or more ventrally pointing supra-antennal setae . . . . . . . . . Johowia Silva Figuero
– Frons with at most four supra-antennal setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49

49. Lower interfrontal seta and lower fronto-orbital seta close together, directed ventrally and medially,
almost parallel with supra-antennal setae; minute flies, body length approximately 1 mm; costa
short, about 0.3 wing length; female with parasitic-type oviscape with long lateral setae (Fig. 67);
male cercus elongate, about as long as epandrium but thick. . . . . . . . . . . . . Synclinusa Borgmeier
– Lower interfrontal and lower fronto-orbital setae not both inclined medially and ventrally (lower
fronto-orbital seta absent in some species; Neophora have lower fronto-orbital seta medially
inclined, but the lower interfrontal seta is directed laterally, Fig. 23); body size and costal length
various; female oviscape various, but if parasitic type, without long lateral setae; male cercus
various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
738
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

cerc

epand

hypd

cerc
40 Gymnoptera

sg 7

epand

hypd

41 Tapantia 42 Apocephalus
43 Megaselia

cerc

cerc

epand epand
a

hypd

44 Diocophora

b
45 Cremersia

Figs. 52.40–45. Terminalia: lateral view of male terminalia of (40) Gymnoptera simplex (Brues), (modified from Brown, 1992, fig. 33B);
and (41) Tapantia bicasa Brown & Kung, (Brown & Kung, 2004, fig. 3); dorsal view of female terminalia of (42) Apocephalus wheeleri
Brues, (Nearctic, MND, fig. 51.83); and (43) Megaselia iroquoiana (Malloch), (Nearctic, MND, fig. 51.81); lateral view of male terminalia of
(44) Diocophora appretiata (Schmitz), (modified from Brown, 1992, fig. 9C); left lateral view (a) and right lateral view (b) of male terminalia
of (45) Cremersia sp. Figure 45 illustrated by L. Mui.
Abbreviations: cerc, cercus; epand, epandrium; hypd, hypandrium; sg 7, segment 7.
PHORIDAE (hump-backed flies, scuttle flies) 52 739

50. Hind tibia with row of differentiated, enlarged, spinelike setae anterior to dorsal setal palisade that
are similar in size and in addition to usual posterodorsal row of similar setae (Fig. 6). . . . . . . . 51
– Hind tibia with posterodorsal row of setae only; if anterodorsal row present, then setae are much
smaller or not spinelike . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52

51. Posterior portion of anepisternum greatly enlarged posteriorly (Fig. 68); hind tibia with anterodorsal
row of spinelike setae (Fig. 6). . . . . . . . . . . . . . . . . . . . . . . . . . . . Phalacrotophora Brues, in part
– Posterior portion of anepisternum relatively small (Fig. 58); hind tibia with or without anterodorsal
row of spinelike setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Megaselia Rondani, in part

46 Macrocerides
hypd

epand
47 Macrocerides

ovscp ovscp

48 Cremersia 49 Neodohrniphora

epand

hypd

50 Trophodeinus

Figs. 52.46–50. Antenna and terminalia (continued): lateral view of antenna (46) and right lateral view of male terminalia (47) of Macrocerides
curtifrons Borgmeier, (Borgmeier, 1969, figs. 47, 48); lateral view of female terminalia of (48) Cremersia spinicauda Borgmeier, (MND,
fig. 51.77); and (49) Neodohrniphora cognata Prado, (modified from Prado, 1976, fig. 72); lateral view of male terminalia of (50) Trophodeinus
furcatus (Borgmeier), (Nearctic, MND, fig. 51.103, as Bactropalpus).
Abbreviations: epand, epandrium; hypd, hypandrium; ovscp, oviscape.
740
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

52. Frons short, with lower interfrontal setae strongly medially directed (Fig.  22); anepisternum
dorsally setulose; male cercus elongate, narrow (as in Fig. 63); female with heavily sclerotized,
parasitic-type oviscape. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachycephaloptera Borgmeier
– Frons longer; if short then either lower interfrontal setae directed dorsally, anepisternum bare, or
other characters different. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53

53. Frontal setae long, with one pair supra-antennal setae, divergent pair of lower interfrontal setae,
convergent lower frontorbital setae, and other setae unmodified (Fig.  23); wing vein Rs curved
proximal to origin of R2+3 producing slight concavity (Fig. 81); vein R2+3 thinner than R4+5; body yellow
in color; male with thin, elongate cercus; female with thin, sclerotized, parasitic-type oviscape . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neophora Borgmeier
– Lower interfrontal and lower fronto-orbital setae not as long and not curved as above (lower
fronto-orbital seta absent in some species); wing vein Rs usually straight; if somewhat similar to
above, then body color brown; other characters various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54

54. Male. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
– Female . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67

55. First flagellomere platelike (Fig. 24): flat, rounded, greatly enlarged; frons extremely short. . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lecanocerus Borgmeier
– First flagellomere not platelike; if flat and rounded, then not greatly enlarged and frons longer . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56

56. Tergite 6 glossy, shining; wing often marked with spot of pigment posterior to base of vein R2+3
(Fig. 82); if tergite 6 not shining (R. trivittata Malloch), then anepisternum dorsally setulose with
one long seta (as in Fig. 62), dorsum of thorax light brown with median and lateral darker brown
vittae, and cercus elongate, longer than epandrium. . . . . . . . . . . . . . . . . Rhyncophoromyia Malloch
– Tergite 6 matte, not shining; wing without pigment; characters different than those of R. trivittata.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57

57. Cercus elongate (longer than dorsal face of epandrium), narrow (Fig. 63). . . . . . . . . . . . . . . . . 58
– Cercus shorter, or if slightly elongate, then broader (e.g., Fig. 69). . . . . . . . . . . . . . . . . . . . . . . 61

58. Anepisternum with setulae and one long, thick seta (as in Fig. 62). . . . . . . . .Menozziola Schmitz
– Anepisternum at most with small setulae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59

59. First flagellomere greatly elongate, pointed, not flattened; arista absent. . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macrocerides Borgmeier, in part
– First flagellomere not modified in this way; arista present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60

60. Lower interfrontal setae closer to midline of frons than to eye margin, directed upward, parallel or
divergent (Fig. 25). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apocephalus Coquillett, in part,
Dacnophora Borgmeier
Note: Males of these two genera cannot be separated reliably at this time.

– Lower interfrontal setae closer to eye margin, medially directed . . . . Auxanommatidia Borgmeier

61. Anepisternum with setulae and one (in a few species more than one) large seta (as in Fig. 62). . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
– Anepisternum at most with small setulae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
PHORIDAE (hump-backed flies, scuttle flies) 52 741

62. Posterior margin of left side of epandrium with dorsoventral row of thick, dark setae (Fig. 44). .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diocophora Borgmeier
– Epandrium without such distinctive row of setae . . . . . . . . . . . . . . . . . Megaselia Rondani, in part

63. Supra-antennal setae absent; their position occupied by divergent, dorsally pointed lower
interfrontal setae that are closer together than to eye margin (Figs. 25, 26). . . . . . . . . . . . . . . . 64
– Supra-antennal setae usually present; if absent, then lower interfrontal setae not close together and
divergent, instead closer to eye margin (e.g., Fig. 14). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65

64. Either foretibia with dorsal setal palisade, like that of mid- and hind tibia, or cercus no longer than
dorsal face of epandrium; frons slightly narrowed, dark-colored (Fig. 26); abdominal tergites light
brown to yellow with dark brown lateral markings. . . . . . . . . . . . . . . . . Neodohrniphora Malloch
– Foretibia without dorsal setal palisade; cercus longer than dorsal face of epandrium; other
characters various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apocephalus Coquillett, in part

65. Body completely dark brown; first flagellomere elongate, pointed, setulose (Fig. 46); terminalia
as in Fig. 47 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macrocerides Borgmeier, in part
– Body color various, often with lighter markings; first flagellomere not pointed; terminalia
different . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66

66. Fork of wing veins R2+3 and R4+5 extremely broad (Fig. 83); foretarsomeres rectangular, elongate,
broad. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trophithauma Schmitz
– Fork of wing veins narrower; foretarsomeres shorter or narrower . . . . Megaselia Rondani, in part

67. Proboscis elongate (Fig. 70), apical section usually narrowed throughout length, but if basally
broad then tergite 6 glossy; wing often with dark markings (Fig. 82). . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhyncophoromyia Malloch
– Proboscis short; if elongate then apical section slightly broader basally (some Diocophora spp.)
and tergite 6 not glossy; wing unmarked. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68

68. Lower interfrontal setae closer to midline than eye margin, divergent to parallel; if lower
interfrontal setae midway between midline and eye margin, then key females with parasitic-type
oviscape here (as in Figs. 42, 67, 72). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
– Lower interfrontal setae closer to eye margin than midline, parallel to convergent; if lower interfrontal
setae midway between midline and eye margin, then key non-parasitic females this way. . . . . 73

69. Apical tarsomere of foreleg drawn out into long, thin, pointed segment. . . . . . . . . . . . . . . . . . . 70
– Apical tarsomere subequal in thickness to other tarsomeres. . . . . . . . . . . . . . . . . . . . . . . . . . . . 71

70. Apical tarsomeres of all legs drawn out to long, thin, pointed segments; venter of abdominal
segment 6 with only small, inconspicuous setae and without sternite. . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apocephalus Coquillett, in part
– Only apical tarsomere of foreleg modified; venter of segment 6 with horseshoe-shaped sternite
bearing long, thick setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dacnophora Borgmeier
742
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

71. Anepisternum with setulae and large seta (as in Fig. 62); lower interfrontal setae parallel, midway
between midline and eye margin; abdominal segments 5 and 6 greatly narrowed, tergites short. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Menozziola Schmitz
– Anepisternum usually bare (setulae and setae present in few species); lower interfrontal setae
closer to midline than eye margin, divergent; major constriction of abdomen usually after
tergite 5 or 6; tergites not greatly shortened. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72

51 Laciniomyia 52 Laishania

53 Ecitomyia 54 Paurophora 55 Rhabdomyia 56 Puliciphora

Figs. 52.51–56. Adult females (legs omitted in Figs. 53, 54, and 56): lateral view of (51) Laciniomyia dilata Kung & Brown, (Kung & Brown,
2005, fig. 2); and (52) Laishania angustithorax Kung & Brown, (Kung & Brown, 2005, fig. 3); dorsal view of (53) Ecitomyia wheeleri Brues,
(Nearctic, MND, fig. 51.91); (54) Paurophora borgmeieri Silvestri, (Silvestri, 1947, fig. 1); (55) Rhabdomyia setosa (Brues), (Brues, 1925,
fig. 1); and (56) Puliciphora imbecilla Borgmeier, (Borgmeier, 1960a, fig. 91).
PHORIDAE (hump-backed flies, scuttle flies) 52 743

anepst

anepst fur

57 Diplonevra 58 Megaselia

lat oc

oc lb

l frorb s

59 Xanionotum 60 Stichillus

cerc

lg s

61 Diplonevra 62 Neopleurophora

Figs.  52.57–62. Heads, thoraces, and terminalia (continued): lateral view of head and thorax of (57)  Diplonevra setigera (Malloch) and
(58) Megaselia sp.; lateral view of adult female of (59) Xanionotum sp.; anterior view of head of (60) Stichillus sp.; lateral view of male
terminalia of (61) D. setigera; left lateral view of thorax of (62) Neopleurophora sp. Scanning electron micrographs by G. Kung.
Abbreviations: anepst, anepisternum; anepst fur, anepisternal furrow; cerc, cercus; l frorb s, lower fronto-orbital seta; lat oc, lateral ocellus;
lg s, large seta; oc lb, ocellar lobe.
744
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

72. Foretibia with dorsal setal palisade (as on mid- and hind tibia); if not, then oviscape with large,
lateral lobes (subgenus Eibesfeldtphora Disney, Fig.  71), or abdominal segment 6 with lateral
patch of long setae and oviscape laterally compressed, with large median hook (subgenus
Neodohrniphora sensu stricto, Fig. 49). . . . . . . . . . . . . . . . . . . . . . . . . . Neodohrniphora Malloch
– Foretibia without setal palisade; abdomen and oviscape not as above . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apocephalus Coquillett, in part

73. Clypeus enlarged, protruding, shiny; proboscis elongate (Fig. 14). . . . . . . Trophithauma Schmitz
– Clypeus not enlarged and protruding; proboscis not elongate. . . . . . . . . . . . . . . . . . . . . . . . . . . 74

74. Oviscape mostly membranous, without elongate sclerotized portions; cercus well developed . 75
– Oviscape modified for parasitism: elongate, with shiny sclerotized portions; cercus reduced or
absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76

75. Segment 7 of female abdomen enlarged, not strongly narrowed relative to segment 6; venter of
abdominal segment 8 often deeply cleft, with two elongate sclerites protruding; wing as in Fig. 84
with Rs long, straight. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phymatopterella Brues, in part
– Segment 7 of female abdomen small, clearly much narrower than segment 6; venter of abdominal
segment 8 not deeply cleft, with short, relatively non-protruding sclerites only; wing different
than Fig. 84 with Rs more curved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Megaselia Rondani, in part

76. Anepisternum with setulae and large seta (as in Fig. 62); oviscape small, narrow, not extensively
sclerotized . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diocophora Borgmeier
– Anepisternum bare; oviscape conspicuous, mostly sclerotized. . . . . . . . . . . . . . . . . . . . . . . . . . 77

77. Frons short, about twice as wide as long; oviscape dorsally flattened, broad, shiny, apically pointed.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lecanocerus Borgmeier
– Frons longer; if short then oviscape not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78

78. Oviscape long, narrow, strongly curved ventrally (Fig. 65) . . . Myrmosicarius Borgmeier, in part
– Oviscape not so elongate and only slightly curved ventrally. . . . . . . . Auxanommatidia Borgmeier

79. Anepisternum setulose and with two long setae; mid- and hind tibia with anterodorsal row of
spinelike setae (Fig. 6); aberrant specimens of P. halictorum (Melander & Brues) . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phalacrotophora Brues, in part
– Anepisternum usually bare, but if setulose lacking two long setae; mid- and hind tibia without
anterodorsal row of spinelike setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80

80. Costa extremely long, 0.7 or more of wing length; dark brown flies; frontal setae short, bristly
(male only; female unknown) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eremophora Borgmeier
– Costa shorter, 0.6 or less of wing length; body color and frontal setation various . . . . . . . . . . . 81

81. Body shining dark brown to black with widely, evenly spaced, uniform-sized setae (Fig. 73); head
smoothly rounded, body limuloid (teardrop-shaped). . . . . . . . . Platydipteron Borgmeier & Prado
– Body, if shining black, with setae smaller, denser; body not limuloid. . . . . . . . . . . . . . . . . . . . . 82
PHORIDAE (hump-backed flies, scuttle flies) 52 745

ovscp

cerc

63 Apocephalus 64 Borgmeieriphora

ovscp

tsm 1

65 Myrmosicarius 66 Physoptera

anepst

lng lat s

67 Synclinusa 68 Phalacrotophora

Figs. 52.63–68. Terminalia, tarsus, and thorax (continued): lateral view of male terminalia of (63) Apocephalus sp.; lateral view of adult
female of (64)  Borgmeieriphora multisetosa Brown; lateral view of oviscape of (65)  Myrmosicarius  sp.; lateral view of foretarsus of
(66) Physoptera sp.; lateral view of oviscape of (67) Synclinusa sp.; lateral view of thorax of (68) Phalacrotophora sp. Scanning electron
micrographs by G. Kung.
Abbreviations: anepst, anepisternum; cerc, cercus; lng lat s, long lateral seta; ovscp, oviscape; tsm 1, tarsomere 1.
746
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

notch

69 Myriophora 70 Rhyncophoromyia

lat lb

71 Neodohrniphora 72 Myriophora

73 Platydipteron 74 Dinocercus

Figs. 52.69–74. Heads, thoraces, and terminalia (concluded): lateral view of male terminalia of (69) Myriophora sp.; lateral view of head and
thorax of (70) Rhyncophoromyia sp.; lateral view of oviscape of (71) Neodohrniphora mexicanae Disney and (72) Myriophora sp.; dorsal
view of head and thorax of (73) Platydipteron balli Brown; right lateral view of male terminalia of (74) Dinocercus sp. Scanning electron
micrographs by G. Kung.
Abbreviation: lat lb, lateral lobe.
PHORIDAE (hump-backed flies, scuttle flies) 52 747

82. Frons yellow, evenly covered with fine, dense setulae; only lower interfrontal setae, upper
fronto-orbital setae and usual two pairs of vertical setae present (i.e., supra-antennal, lower
fronto-orbital, and upper interfrontal setae absent); small flies, approximately 1.2 mm in length
(measured from head to end of abdominal segment 6); body color mostly yellow; associated with
stingless bees. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Styletta Borgmeier
– Without the above combination of characters; frons usually with more setae; body length, color,
and natural history various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83

83. Male cercus greatly inflated (Fig. 74) and cercal setae near apex short, with length about one-half
depth of cercus; lower interfrontal setae convergent; costa short, approximately 0.36 wing length
(male only; female unknown) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dinocercus Borgmeier
– Male cercus not inflated, cercal setae about as long as depth of cercus; lower interfrontal setae and
costa various; all females. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84

84. Wing vein Rs with posterior projection or swelling at base of M1 (Fig.  80); lower interfrontal
setae parallel or convergent, situated midway between eye margin and median furrow; female with
parasitic-type oviscape; male cercus often pointing posteroventrally. . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudacteon Coquillett, in part
– Wing vein Rs not modified at base of M1, or if slightly modified, then lower interfrontal setae close
to midline and divergent or close to eye margin and convergent; oviscape various; male cercus
pointing more posteriorly. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85

85. Male and most females with lower interfrontal setae near eye margin and strongly inclinate (as in
Fig. 22), in some females only slightly inclinate; all frontal setae small, much shorter than frontal
height; male cercus elongate, slightly inflated; female with parasitic-type oviscape downturned,
held closely against posterior of abdomen, dorsoventrally flattened (Fig. 64), with pair of subapical
dark spots; small species, body length 0.8–1.2 mm. . . . . . . . . . . . Borgmeieriphora Prado, in part
– Lower interfrontal setae usually parallel or divergent, but if strongly inclinate, then frontal setae
usually longer, about as long as frontal height; cercus, oviscape and body length various. . . . . 86

npl clft

anepst

75 Gymnophora

Fig. 52.75. Lateral view of thorax of (75) Gymnophora arcuata (Meigen), (Palearctic). Scanning electron micrograph by G. Kung.
Abbreviations: anepst, anepisternum; npl clft, notopleural cleft.
748
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

76 Latiborophaga 77 Latiborophaga

p swlg

al s

78 Gymnoptera 79 Pseudohypocera

p swlg
cncty

80 Pseudacteon 81 Neophora

82 Rhyncophoromyia 83 Trophithauma

84 Phymatopterella 85 Apodicrania

Figs. 52.76–85. Wings (continued): dorsal view of male (76) and female (77) of Latiborophaga sp.; (78) Gymnoptera sp.; (79) Pseudohypocera
kerteszi (Enderlein); (80)  Pseudacteon  sp; (81)  Neophora frontalis Borgmeier; (82)  Rhyncophoromyia  sp.; (83)  Trophithauma  sp.;
(84) Phymatopterella sp.; and (85) Apodicrania sp. Photographs by B. Koehler.
Abbreviations: al s, alular setae; cncty, concavity; p swlg, posterior swelling.
PHORIDAE (hump-backed flies, scuttle flies) 52 749

86. Lower interfrontal setae close to midline of frons, divergent (Fig. 25); oviscape of female strongly
sclerotized; male cercus long and slender (Fig. 63). . . . . . . . . . . . Apocephalus Coquillett, in part
– Lower interfrontal setae not divergent, often closer to eye margin than midline, parallel to slightly
convergent; female oviscape not modified for parasitism; male cercus usually short, but if longer,
then not slender . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Megaselia Rondani, in part

87. Wing well developed and vein R2+3 present (although sometimes small and inconspicuous) . . . . 88
– Wing absent, reduced, or, if present, vein R2+3 absent; most Metopina-group genera . . . . . . . . . . 91

Note: Either males or females are unknown for many Metopina-group genera, and there is a strong
possibility that some specimens will not end satisfactorily in this part of the key.

88. Wing vein R2+3 and fork of Rs extremely small (Fig. 86); vein M2 strongly curved at base; first
flagellomere elongate, drawn out into long process, lacking arista in male (similar to Fig. 9). . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dahliphora Schmitz
– Wing vein R2+3 and fork large, distinct; vein M2 not strongly curved at base; first flagellomere
rounded; arista present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89

89. Costal setae thick, about as long as wing vein R2+3; male lacking notopleural cleft. . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Allochaeta Borgmeier
– Costal setae thin, short, about one-half length of wing vein R2+3; male (and female of Gymnophora)
with notopleural cleft (Fig. 75). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90

90. Frontal setae small, thin, barely differentiated from frontal setulae; lower interfrontal and lower
fronto-orbital setae not discernable; female with notopleural cleft. . . . . . . Gymnophora Macquart
– Frontal setae large, thick; lower interfrontal and lower fronto-orbital setae well-developed; female
lacking notopleural cleft. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Woodiphora Schmitz

swlg

86 Dahliphora 87 Puliciphora

88 Travassophora

Figs. 53.86–88. Wings (concluded): dorsal view of (86) Dahliphora sp.; (87) Puliciphora sp.; and (88) Travassophora sp. Photographs by
B. Koehler.
Abbreviation: swlg, swelling.
750
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

91. Wing fully developed; vein M2 strongly posteriorly curved at base and vein CuA1 curved anteriorly
at midpoint, such that section between is hourglass-shaped (Fig. 38). . . . . . . Metopina Macquart
– Wing reduced or, if fully developed, without hourglass-shaped space between wing veins. . . . 92

92. Frons produced between antennae, with three to four pairs of strong, ventrally pointing supra-antennal
setae (Fig. 27); female with well-developed wings. . . . . . . . . . . . . . Cataclinusa Schmitz, in part
– Frons, if produced between antennae, with only two pairs of supra-antennal setae or
supra-antennal setae smaller than other frontal setae (as in Fig. 28); female usually brachypterous
or apterous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93

93. Male (fully winged). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94

– Female (fully winged, apterous, or brachypterous). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105

94. Frons short, narrowly produced between antennae (Fig. 28). . . . . . . . . . . . . . . . . . . . . . . . . . . . 95

– Frons longer, not narrowly produced. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96

95. First flagellomere greatly elongate, pointed apically. . . . . . . . . . . . . . . Cyphocephalus Borgmeier

– First flagellomere rounded to slightly elongate (Fig. 28). . . . . . . . . . . Chonocephalus Wandolleck

96. Costa with abrupt swelling between humeral crossvein and R1 (Fig. 87). . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Puliciphora Dahl, in part
– Costa without such swelling. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97

97. Foretibia with ventral patch of long setae . . . . . . . . . . . . . . . . . . . . . . . . Homalophora Borgmeier

– Foretibia without such patch of setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98

98. Foretibia apically flattened, with apicoventral process and usually with ventral notch (Fig. 7). . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apterophora Brues
– Foretibia not modified in this way. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99

99. Tarsomere 1 of foreleg with ventrally excavated notch (Fig. 8) . . . . . . . . . Eurycnemis Borgmeier
– Tarsomere 1 not modified in this way . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100

100. First flagellomere drawn out in short process; frontal setae extremely long and strong. . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adelopteromyia Schmitz
– First flagellomere round; frontal setae small. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101

101. Wing broad, with greatly enlarged anal region (Fig. 88); vein R1 incomplete, not reaching costa. . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Travassophora Borgmeier
– Wing less developed in anal region; vein R1 complete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102

102. Wing vein Sc strongly developed and reaching R1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103

– Wing vein Sc weak and ending free of R1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104

103. Terminalia highly modified; epandrium arched, ventrally excavated; hypandrium with narrow,
often with darkly sclerotized and shiny pointed processes (Fig. 50); palpus broad with extremely
small setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trophodeinus Borgmeier
– Terminalia normal, epandrium not arched, hypandrium consists of mixture of membrane and light
colored sclerites; palpus narrower, usually with large setae . . . . . . . . . . . . . . . . Puliciphora Dahl,
Commoptera Brues

Note: Separation of males of Commoptera and Puliciphora is questionable.

PHORIDAE (hump-backed flies, scuttle flies) 52 751

104. Costa slightly less than one-half wing length; wing vein Sc evident, sometimes with one to two
small setae at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acontistoptera Brues
– Costa extending slightly more than one-half wing length; wing vein Sc without setae and often
absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ecituncula Schmitz,
Ecitophora Schmitz,
Ecitoptera Borgmeier & Schmitz,
Ecitomyia Brues

Note: This couplet is from Disney (1994). Probably many other described and undescribed genera key
here as well. Much work is needed on generic limits in the Metopina group, as well as the association of
Metopina-group males with their respective females.

105. Oviscape of parasitoid type, heavily sclerotized, pointed, downturned (Fig. 64). . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Borgmeieriphora Prado, in part
– Oviscape not of parasitoid type; not heavily sclerotized, or pointed and downturned . . . . . . . 106

106. Wing and usually halter completely absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107

– At least rudimentary wing or wing stump present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120

107. Body without usual dense setae, instead with long, sparse setae (Fig. 52); head elongate, rounded;
foretarsomere 1 with extremely dense comb of elongate yellow setae; forefemur with ventral
expansion at midlength . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Laishania Kung & Brown
– Body with smaller, denser setae; head usually flattened; foretarsomere 1 without comb of elongate
yellow setae; forefemur not expanded ventrally at midlength. . . . . . . . . . . . . . . . . . . . . . . . . . 108

108. Frons broadly produced in a rounded lobe between antennae (Figs. 29, 54). . . . . . . . . . . . . . . 109
– Frons not produced in this way . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 110

109. Posterior segments of abdomen with long setae (Fig. 54). . . . . . . . . . . . . . . Paurophora Silvestri
– Posterior abdominal segments with short, inconspicuous setae only . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chonocephalus Wandolleck

110. Proboscis large, triangular in lateral view (Fig.  30); eyes greatly reduced; body laterally
flattened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trophodeinus Borgmeier
– Proboscis, if enlarged, not triangular in lateral view; other characters various. . . . . . . . . . . . . 111

111. Tergites absent or greatly reduced from abdominal segments 3–6 (Figs. 53, 55). . . . . . . . . . . 112
– Most abdominal segments with well-developed tergites. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115

112. Abdominal membrane dorsally with sparse, long, thick setae (Fig. 55). . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhabdomyia Borgmeier, in part
– Abdominal membrane dorsally without prominent setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113

113. Except for supra-antennal setae, prominent frontal setae absent; head and thorax dark colored,
abdomen contrastingly whitish-gray . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apterella Borgmeier
– At least postocellar setae present, in addition to supra-antennal setae; body color different . . 114

114. Body relatively uniform yellowish-brown in color. . . . . . . . . . . . . . . . . . . . . . Ecituncula Schmitz

– Head and thorax dark, reddish-brown; abdominal segments 1–2 white, in strong contrast with
blackish-gray segments 3–6. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Zikania Borgmeier
752
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

115. Head and thorax orangish-brown, in strong contrast with abdominal tergites that are almost black;
ocelli absent; inner face of hind tibia with bifid posterior setae at apex (visible with compound
microscope only). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adenophora Borgmeier
– Body color usually different; if similar then ocelli present; posterior setae of hind face of hind
tibia simple. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116

116. Proboscis elongate, much longer than head, narrow, geniculate (as in Fig. 10) . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apterophora Brues
– Proboscis shorter, thicker. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117

117. No large frontal setae present between supra-antennal setae and those at vertex. . . . . . . . . . . 118
– At least one pair of large setae present between supra-antennal setae and those at vertex. . . . 119

118. Abdominal segment 5 without dorsal gland opening or associated flaplike lid. . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Homalophora Borgmeier
– Abdominal segment 5 with dorsal gland opening and usually flaplike lid (Fig. 56). . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Puliciphora Dahl, in part

119. Posterior margin of scutum without setae. . . . . . . . . . . . . . . . . . . . . . . . Placophorina Borgmeier

– Posterior margin of scutum with at least one pair of large setae (Fig. 56). . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Puliciphora Dahl, in part

120. Wing with membrane and well-developed post-radial veins present . . . . . . . . . . . . . . . . . . . . 121
– Wing rudimentary, consisting of a linear stump, without wing membrane or membrane extremely
reduced and lacking more than rudimentary post-radial veins . . . . . . . . . . . . . . . . . . . . . . . . . 123

121. Costal setae extremely long, about as long as depth of wing. . . . . . . . . . Adelopteromyia Schmitz
– Costal setae much shorter. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122

122. Some abdominal tergites posterior to tergite 2 greatly reduced in size, much shorter than tergite 2
such that most of abdomen is membranous (Fig. 53) . . . . . . . . . Ecitoptera Borgmeier & Schmitz
– Abdominal tergites posterior to tergite 2 only gradually reduced in size . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Holopterina Borgmeier

123. Wing rudiment with extremely long setae that are more than twice length of rudiment itself. . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124
– Wing rudiment with only short setae that are much shorter than rudiment itself . . . . . . . . . . . 125

124. Dorsum of abdomen with rows of long, often thick setae along posterior margin of each tergite
(Fig. 59). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xanionotum Brues, in part
– Dorsum of abdomen with short, inconspicuous setae. . . . . . . . . . . . . . . . . . Acontistoptera Brues

125. Abdomen with sparse, long, thick setae (Fig. 55). . . . . . . . . . . . . Rhabdomyia Borgmeier, in part
– Abdomen with short, inconspicuous setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 126

126. Scutellum and scutellar setae absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127

– Scutellum or scutellar setae present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 128

127. Tergites 3–5 rudimentary, much smaller than the large, trapezoidal tergite 2 (Fig. 53) . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ecitomyia Brues
– Tergites 3–5 at most gradually reduced in size posteriorly (Fig. 56). . . . Puliciphora Dahl, in part
PHORIDAE (hump-backed flies, scuttle flies) 52 753

128. Tergite 2 broader than thorax. . . . . . . . . . . . . . . . . . . . . . . . . . . Thalloptera Borgmeier & Schmitz

– Tergite 2 (the largest tergite of the abdomen) narrower than thorax. . . . . . . . . . . . . . . . . . . . . 129

129. Wing rudiment consists of simple, linear remnant of costa, or wing halterlike. . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ecitophora Schmitz
– Wing rudiment with at least some membrane and usually with post-costal veins evident
(Fig. 39). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Commoptera Brues

Synopsis of the fauna from Mexico (Disney  & Bragança, 2000), is probably of
limited value in our region.
The number of species of Phoridae that inhabit Central
America is not known with any degree of accuracy, but Anevrina Lioy. Specimens of an undescribed species
Brown (1998b) estimated a total of about 1100–2500 species similar to the European Anevrina unispinosa (Zetterstedt)
for Costa Rica alone. These numbers were obtained based are known from Monteverde, Costa Rica. This is the only
on extrapolations of “known” faunas (for instance, certain Neotropical Region record for this otherwise Holarctic and
genera) with the number described for the Neotropical Oriental Region genus. Larvae feed on carrion.
Region as a whole. The currently described World total is
Apocephalus Coquillett. This is currently the second
about 3700 species, a number that probably represents less
largest genus of phorids (after Megaselia), and species are
than 10% of the actual total.
common throughout Central America. Single sites may
Abaristophora Schmitz. A few undescribed species of have over 100 species of this genus (Brown  & Feener,
this genus have been collected at high elevations in Costa 1995; Brown, 2004b), many of which are undescribed. All
Rica and South America. Nothing is known of their life are parasitoids, mostly attacking ants. Some that develop
history. Other species occur in New Zealand, Southeast Asia, in the head of ants have earned the name “ant-decapitating
and the Holarctic Region. fly”, a term that is also applied to some other genera. Other
species attack stingless bees, vespid wasps, and cantharoid
Acanthophorides Borgmeier. Both sexes of these flies beetles (Brown, 1994a, 1996). The last comprehensive key
are found at the front of army ant raids (Brown & Feener, to species (Borgmeier, 1971a) is of marginal use because
1998), where the females presumably oviposit in army ants. of the vast number of undescribed species and needs to be
A key to species is given by Disney & Berghoff (2007). supplemented by more recent revisions of Apocephalus
Acontistoptera Brues. The distinctive, brachypterous subgroups (Brown, 1993c, 1994b, 1996, 1997a, 2000b,
females of this genus are associated with army ants of the 2002).
genera Labidus and Neivamyrmex, but are occasionally also
Apodicrania Borgmeier. Of the at least two Central
collected by Malaise traps. The species were keyed by
Borgmeier (1963c). American species of this genus, A. molinai Borgmeier is
attracted to and oviposits in injured Camponotus carpenter
Adelopteromyia Schmitz. These flies are found in ants. Another, A. termitophila (Borgmeier), has been
association with army ants. Females have short wings with collected with termites and carpenter ants. The three described
extremely long costal setae, but at least some are still able to species are keyed by Borgmeier (1963b).
fly. A key to the species, as well as a critical discussion of the
genus, is given by Disney & Kistner (1990). Apterella Borgmeier. The single species, A. molliventris
Borgmeier from Costa Rica, is known only from females
Adenophora Borgmeier. Disney & Kistner (1997) described that are similar to those of Puliciphora species.
a species of this genus from Guatemala and discussed its
differences from the only other species, known from Brazil. Apterophora Brues. The females of this genus are
The genus is found throughout Central America, and females wingless, associated with ant and termite colonies, and
can be collected by searching in Nasutitermes termite mostly undescribed. There is a key to males (Prado,
nests. 1976).

Allochaeta Borgmeier. Specimens of this genus are Auxanommatidia Borgmeier. At least some of the several
occasionally collected in Malaise traps, although some Neotropical Region species of this genus probably should be
have also been collected associated with leaf-cutter ants. placed in Veruanus Schmitz (Brown, 1998a), but there has
The key to the described species, four from Brazil and one been no modern revision of the group. Borgmeier (1971a)
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

gave up trying to classify some of the species placed in Commoptera Brues. The brachypterous females of this
Auxanommatidia, noting that the principle characters New World genus are associated with ants of the genus
also occurred in other genera. There is a key to females Eciton, Pheidole (Seid  & Brown, 2009), and Solenopsis.
(Borgmeier & Prado, 1975), but it is probably of little use No specimens are known from Central America, but it
until the group is revised. The problem of the diagnosis of doubtlessly occurs there.
the genus is discussed by Disney (2004b).
Conicera Meigen. The Neotropical fauna of this genus
Borophaga Enderlein. A single female specimen of is virtually unstudied; probably several species are found
this otherwise mostly Holarctic Region genus is known in Central America. The larvae of Holarctic Region species
from Costa Rica, and several specimens of B. verticalis are scavengers on buried carrion, and one species associ-
Borgmeier are known from Chiapas, Mexico. ated with human cadavers, C. tibialis Schmitz, is known
as the “coffin fly.” There are some unusual associations
Brachycephaloptera Borgmeier. This genus is rep- for this genus in the Neotropical Region, such as one spe-
resented by a single species found in Central and South cies associated with living land snails (Brown, 2000a) and
America, B. trichopleura Borgmeier, that parasitizes several specimens associated with an army ant raid of a
ants of the genus Gnamptogenys (B.  Brown, personal small vespid wasp nest in Ecuador (B.  Brown, personal
observation). observation).

Brownphora Disney. Recently, Disney (2004a) separated Coniceromyia Borgmeier. There are many species of
this genus from Beckerina for the single species B. sinefurca this genus in Central America, but most are undescribed.
(Borgmeier). Specimens are known from mid-elevation sites Kung  & Brown (2000) keyed the patterned-wing species,
in Costa Rica (Brown, 1997c) and Bolivia. but other resources are scattered. The last key was that of
Borgmeier (1963b), but other species within our region
Calamiscus Borgmeier. One of the two species of this were subsequently described by Borgmeier (1969), Peter-
genus (both found in Central America) is attracted to, son & Arntfield (1971), and Peterson (1982).
and parasitizes, injured or fighting stingless bees (Brown,
Cootiphora Brown. Specimens of the single known spe-
1997b).
cies, C. angustata Brown, are from Ecuador, Costa Rica, and
Cataclinusa Schmitz. One of the two described species Texas, USA.
of this rarely collected genus, C. pachycondylae (Brues), Cremersia Schmitz. The key to females of the described
was reared in nests of the ant Pachycondyla harpax in Neotropical members of this New World genus (Borgmeier,
Texas, USA (this ant also occurs in Central America). The 1961) is of limited value, as there are many undescribed spe-
two described species are somewhat different in structure, and cies. They are parasitoids of non-Eciton army ants (Labidus,
it is questionable whether they represent a single genus. Neivamyrmex, Nomamyrmex).

Ceratusa Borgmeier. One species of this genus, Cyphocephalus Borgmeier. The validity of this genus is
C. crinicornis Borgmeier, is known from several Central questionable, as it might only represent some Chonocephalus
American sites. males with long, pointed antennae. There is a single described
species, C. caviceps Borgmeier, from Costa Rica.
Chaetaspidia Borgmeier. Although only the Costa Rican
C. convexifrons (Borgmeier) has been described, there is at Dacnophora Borgmeier. The species of this genus are par-
least one other undescribed species in this genus. asitoids of Labidus and Neivamyrmex army ants. There are six
described species, but there are at least seven species, mostly
Chaetocnemistoptera Borgmeier. Several species of this undescribed, in Costa Rica alone. A key to females is given by
genus are known from Central America (Brown, 1992), but Borgmeier (1961). Males cannot be reliably separated from
they need revision. Formerly, they were considered part of those of Apocephalus.
the genus Chaetopleurophora Schmitz, and are still treated
Dahliphora Schmitz. This genus needs to be critically
this way by Disney (1994).
studied, as it possibly consists only of Metopina species with
elongate antennae. A key to species is given by Borgmeier &
Chonocephalus Wandolleck. A full revision of the Neo-
Prado (1975).
tropical species of this genus is desperately needed. The
females are flattened and apterous and found associated Dinocercus Borgmeier. It is not clear if the specimens
with decaying organic material. Disney’s (2008b) revision found in Central America are the same as the single described
of the Neotropical fauna recognized 28 species, many from species, D. caudatus Borgmeier, from Brazil. Only males are
Central America. known.
PHORIDAE (hump-backed flies, scuttle flies) 52 755

Diocophora Borgmeier. This is a problematic genus that (Borgmeier, 1971a; Disney  & Berghoff, 2005; Silvestri,
grades into Rhyncophoromyia. Most species of both genera 1947) and one reinstated (Disney & Rettenmeyer, 2007). The
are parasitoids of Camponotus carpenter ants, with both number of species found in Central America is unknown, as
also attacking Dolichoderus species. Diocophora species the genus needs revision relative to the similar Ecitophora.
also attack Cephalotes atratus. One species, D. appretiata
Schmitz, sits beside army ant trails, possibly to lay eggs in dead Enderleinphora Disney. Disney (2004a) created this
Camponotus workers carried by the army ants (Brown  & genus to house E.  fuscohalterata (Borgmeier), which was
Feener, 1998). A new key to species is given by Disney & formerly a species of Beckerina, and E.  hansoni Disney.
Berghoff (2007), but further undescribed species exist. Specimens of both species have been collected in Costa Rica
at mid-elevation sites.
Diplonevra Lioy. Found much more frequently at mid- to
high elevations, this genus includes six Central American Eremophora Borgmeier. The male of a single species,
species (Corona & Brown, 2005), two of which are the larg- E. hyalina Borgmeier, was described from Panama, but also
est phorids in Central America. Holarctic Region species are occurs in Costa Rica.
known to be scavengers and parasites of earthworms.
Eurycnemis Borgmeier. A single species, E. alvarengai
Dohrniphora Dahl. After Megaselia, this is the most Borgmeier, is described from Brazil, but there are also
commonly collected genus of phorids, probably because they unidentified specimens from Costa Rica. The character for
are relatively large and conspicuous. Preliminary sorting has recognition of this genus, an excavated first foretarsomere, is
uncovered about 50 Central American species. The larvae probably a modification to carry wingless females in flight.
Only males are known; the females were probably described
are scavengers, predators, kleptoparasites, and parasitoids,
in another, female-only genus.
and some are associated with ant, termite, and bee nests.
The identification literature is scattered (Borgmeier, 1960a, Gymnophora Macquart. Two species of this scavenging
1961, 1969; Borgmeier & Prado, 1975; Disney, 1983; Prado, genus are found in Central America (Brown, 1987). One,
1976), but the New World species of the genus are being G. spiracularis Borgmeier, is readily attracted to crushed
revised (e.g., Kung & Brown, 2005a, 2006; Brown & Kung, millipedes.
2007).
Holopterina Borgmeier. The single species, H. longi-
Ecitomyia Brues. This is the first of a series of genera palpis Borgmeier (and a subspecies, H.  longipalpis sedula
(i.e., Ecitomyia, Ecitophora, Ecitoptera, Ecituncula) with Borgmeier) should probably be placed in Adelopteromyia. It
brachypterous females that have strongly reduced abdominal is found in colonies of the army ant Labidus praedator.
tergites and that are found associated with army ants. Nearly
all records of this genus have been in association with non- Homalophora Borgmeier. The six species of this genus
Eciton army ants (Labidus, Neivamyrmex, Nomamyrmex). A have wingless females that are extremely similar to the
key to species is given by Borgmeier (1960a), and some new genus Puliciphora, and one species has been transferred to
information on the genus is given by Disney & Rettenmeyer the latter genus (Disney, 2005). They are sometimes found
(2007). in nests of ants and termites. The important literature about
this group is that of Borgmeier (1958, 1960a, 1969).
Ecitophora Schmitz. The brachypterous females of this
genus are commonly found in the raids and emigrations Hypocerides Schmitz. Unidentified specimens of this
of their host army ants (mostly Eciton species). Disney  & cosmopolitan genus are known from Costa Rica (in the
Rettenmeyer (2007) give a new key to species. collection of Utah State University) and Honduras (in the
collection of the Natural History Museum of Los Angeles
Ecitoptera Borgmeier  & Schmitz. Although less County – LACM).
commonly collected than Ecitophora, the brachypterous
females of this genus are more reliably found in Malaise trap Johowia Silva Figuero. I am skeptical that all the
samples. The latest key is that of Borgmeier (1960b), and species placed in this genus belong together; it appears that
some new information on the genus is given by Disney  & Johowia has been a place to put unusual Megaselia species
Rettenmeyer (2007). with extra supra-antennal setae. One such species might be
a parasite of passalid beetles (Brown, 2000a).
Ecituncula Schmitz. This is another genus of army ant-
associated phorids with greatly reduced wings and abdominal Laciniomyia Kung  & Brown. There are two female
tergites; one species has also been found with termites. The specimens of these bizarre flies known from Estación
key of Borgmeier (1960a) is the most recent identification Cacao, Guanacaste, Costa Rica (Kung  & Brown, 2005b).
aid, although three further species have been described They are easily recognized by their densely setose palpus.
756
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Laishania Kung  & Brown. One wingless female more. The females of Holarctic Region species are parasi-
specimen of this highly unusual genus was collected in toids of queens of Camponotus and Crematogaster ants. One
the Peñas Blancas Valley in Costa Rica (Kung  & Brown, species in Costa Rica has been found parasitizing queens of
2005b). Its long legs and general form indicate that it Azteca ants (P. Hanson, personal communication).
probably lives with army ants.
Metopina Macquart. An unknown but probably small
Latiborophaga Brown. One or possibly two species of number of species of these tiny flies are found in Central
this genus (formerly considered part of Borophaga) are America. Their larvae are scavengers. Neotropical species
found in mid- to low elevation rain forests. are keyed by Borgmeier (1969), who also redescribed one
species under the now synonymized generic name Typhlo-
Lecanocerus Borgmeier. There is a single species of phorina Silvestri. Further species are added by Borgmeier &
this genus, L. compressiceps Borgmeier, described from the Prado (1975), and amplified descriptions of some species are
Nearctic Region. Specimens are commonly collected from given by Borgmeier (1969) and Disney & Kistner (1989b).
mid-elevation sites in Central America south to Colombia,
possibly representing further species. Myriophora Brown. Although Disney (1994) prefers
to recognize this group as a subgenus of Megaselia, it is
Macrocerides Borgmeier. This is one of the most prob- as recognizable as many other genera found in our region.
lematic genera in the Neotropical Region, and the species One difficulty, however, is that a few Megaselia species
desperately need critical review. Even without detailed have males with a notched epandrium (a character of male
study, however, it is clear that Macrocerides is an artificial Myriphora) but females without a sclerotized oviscape.
assemblage of species with little in common other than an There are approximately 50 species of Myriophora found
enlarged first flagellomere. The type species, M. curtifrons in Central America, all of which apparently parasitize milli-
Borgmeier (or an extremely similar species), is present in pedes (although centipedes and harvestmen are known hosts
Costa Rica.
elsewhere). There are no useful keys to this genus, as most
Megaselia Rondani. This is the largest genus of phorids species are undescribed, but a few have been named under
and accounts for the majority of specimens gathered by Plastophora Brues (a synonym of Megaselia).
general collecting methods (sweeping, Malaise trapping, pan
Myrmosicarius Borgmeier. There are a few species of
trapping). Although there are some older keys to species
Myrmosicarius in Central America, and the known species
(especially Borgmeier, 1958, 1962, 1971a), they are of limited
(mostly South American) are keyed by Disney et al. (2006).
use because of the tremendous number of undescribed spe-
Most species are parasitoids of Atta and Acromyrmex leaf-
cies. The phorid most commonly submitted for identification
cutting ants.
is Megaselia scalaris (Loew), which is a general, polypha-
gous scavenger. Photographs of this species can be seen at Neodohrniphora Malloch. The two subgenera of Neodohr-
www.phorid.net/phoridae/mscalaris.html. Worldwide, other niphora, Neodohrniphora sensu stricto and Eibesfeldtphora
species are scavengers, predators, parasitoids, and herbi- Disney, are parasitoids of Acromyrmex and Atta leaf-cutting
vores, but the life history of most species is unknown. Two ants, respectively. The five Central American species of N.
distinctive new species from Costa Rica were described by Eibesfeldtphora were recently revised (Brown, 2001b), but
Kung & Brown (2004). the species of N. Neodohrniphora are in disarray and require
revision.
Melaloncha Brues. This exclusively Neotropical genus is
represented by approximately 75 species in Central America. Neophora Borgmeier. The sole species of this genus,
All are parasitoids of adult bees, including stingless bees, N.  frontalis Borgmeier, is found at middle elevations in
honey bees, bumble bees (all Apidae), and one recent record Costa Rica.
from a halictid bee (Wcislo et al., 2004). The entire genus of
over 150 species has recently been revised (Brown, 2004a, Neopleurophora Brown. There are about 15 species,
2004c, 2005, 2006; Gonzalez  & Brown, 2004; Brown  & mostly undescribed, of this genus in Central America.
Kung, 2006; Kung, 2008). Formerly, they were considered part of Chaetopleuro-
phora Schmitz, and are still considered to be so by Disney
Melittophora Brues. The single known species, M. salti (1994).
Brues, lives as a kleptoparasite in the nests of stingless bees
(Salt, 1929). The adults are rarely collected, but a few Costa Paurophora Silvestri. A single specimen of this genus,
Rican specimens are known. otherwise known from termite nests in Brazil, was collected
near Guapiles in Costa Rica (LACM collection).
Menozziola Schmitz. There are three described Neotro-
pical species of this genus (Borgmeier, 1961; Borgmeier & Phalacrotophora Brues. There are about 20–30, mostly
Prado, 1975, as genus Stylusa Borgmeier  & Prado), but undescribed, species of this questionably monophyletic
a review of the available material probably would uncover genus in Central America. Although the life history of most
PHORIDAE (hump-backed flies, scuttle flies) 52 757

Neotropical species is unknown, the larvae of some are nest carried in flight by winged males. Disney (2003b, 2005) is
parasitoids of solitary Hymenoptera, such as halictid bees revising this group, and there are probably about 20 species
and sphecid wasps (Coville & Griswold, 1984; Coville et al., in Central America.
2000; Wcislo, 1990). One South American species has been
reared from a coccinellid beetle, a lifestyle more typical of Rhabdomyia Borgmeier. Two species are known from
European species. The most recent key to the Neotropical Mexico (Brues, 1925 as Ecitomyia and Ecituncula), but
species (Borgmeier, 1971a) is hopelessly outdated and of Disney & Kistner (1989c) have rightly questioned the valid-
little value. ity of the genus relative to others in the Metopina-group of
genera. Only females are known.
Phora Latreille. In tropical regions, such as Central
America, these distinctive, black-colored phorids are found Rhyncophoromyia Malloch. This genus is problematic, as
at mid- to high elevations. The three Costa Rican species its species grade into the genus Diocophora (see discussion
have been revised (Brown, 2001a) and are the same as those of Diocophora). Species with known lifestyles are para-
occurring in other parts of Central America. sitoids of ants in the genus Camponotus and Dolichoderus.
The life history of one species, R. maculineura Borgmeier,
Phymatopterella Brues. Borgmeier (1971a) provided a that occurs in Central America has been investigated in detail
key to the five Neotropical species of this genus, but there are (Brown & Feener, 1993). Females of R. trivittata are attract-
further undescribed species. Previously, the genus was based ed to injured ants of Camponotus sericeiventris (B. Brown,
on females with a prominent sclerotized spot on the wing, personal observation). The most recent published key
and males with enlarged labella, but some undescribed (Prado, 1976) is of limited use in identifying specimens, as
species have females without wing spots. Furthermore, by there are many undescribed species.
Borgmeier’s (1971a) redefinition of the genus, at least one
species described in the genus Megaselia, M.  miristigma Stichillus Enderlein. About a dozen species of this genus
Borgmeier (which occurs in Central America), should be are found in Central America. They occur at all elevations,
transferred to Phymatopterella. Much further study is needed but are common at high elevations and are relatively rare in
to resolve the relationships of this questionable genus, which the lowlands. The last key to species found in our region is
might have to be integrated into Megaselia. that of Borgmeier (1959).
Physoptera Borgmeier. There are at least 22 species of Styletta Borgmeier. The single species, S.  crocea
this genus from Costa Rica. The key (Borgmeier & Prado, Borgmeier, is a parasitoid of small stingless bees (Brown,
1975) to the 13 described New World species is of limited 2000a).
value for identifying specimens from Central America.
Synclinusa Borgmeier. There are two described Brazil-
Placophorina Borgmeier. The single described species, ian species of this genus (Borgmeier, 1971a; Borgmeier  &
P. obtecta Borgmeier from Costa Rica, is known only from Prado, 1975), but it also ranges throughout South America
females that are similar to those of Puliciphora species. and into at least Panama. Although the female has a parasitic-
type oviscape, the hosts are unknown.
Platydipteron Borgmeier  & Prado. One of the two
described species, P. balli Brown, is found at mid-elevations Syneura Brues. The type species, S. cocciphila (Brues),
in Costa Rica, as well as in Bolivia and Colombia. is a well-known parasitoid of scale insects, but is only ques-
tionably related to others described in this genus. The key by
Pseudacteon Coquillett. There are many, mostly unde- Prado (1976) is of limited use, and the genus needs revision.
scribed, species of this genus in Central America, but there is Some species have the first flagellomere greatly enlarged and
no modern, useful revision or key for this region (Borgmeier, are known under the name Tubiceroides Borgmeier
1969 gives the last comprehensive key). An exception is the
key of Plowes et al. (2009) to the species that are parasitoids Tabelliphora Kung & Brown. Several specimens of this
of fire ants of the Solenopsis geminata complex. All Pseu- tiny, limuloid (teardrop-shaped) fly are known from the
dacteon are parasitoids, known to attack species of Azteca, Osa Peninsula in Costa Rica. The female has a parasitic-
Crematogaster, Solenopsis, and other small ants. type oviscape, but the hosts are unknown (Kung & Brown,
2005b).
Pseudohypocera Malloch. The single species, P. kerteszi
(Enderlein) is a well-known pest in both honey bee and Tapantia Brown  & Kung. The single species of this
stingless bee nests. Adult females visit flowers of Theobroma genus, T. bicasa Kung  & Brown, is known from Costa
(Sterculieaceae) and possibly are pollinators of some species Rica (Brown & Kung, 2004).
of this genus (A. Young, personal communication). Disney &
Bartareau (1995) found pollen in the gut of adult females. Thalloptera Borgmeier  & Schmitz. This is another
army ant-associated genus whose females are brachypterous.
Puliciphora Dahl. The unusual, apterous females of this The species were keyed most recently by Disney & Kist-
genus are commonly found on decomposing organic material, ner (1998), and some new information on the genus is
especially dead or injured insects. They often arrive by being given by Disney & Rettenmeyer (2007).
758
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Travassophora Borgmeier. The only described species The females are apterous and laterally flattened, whereas the
in this genus is the Brazilian T. plaumanni Borgmeier. A males are winged and more normal in appearance. Borg-
single unidentified species is known from Costa Rica. meier (1963a) keyed the known males (as Bactropalpus
Borgmeier) and separately, the females (Borgmeier, 1963c),
Trineurocephala Schmitz. These Neotropical flies are but there are other undescribed species.
relatively rare, but are known from a number of low to mid-
elevation sites, especially San Antonio de Escazú in Costa Woodiphora Schmitz. Although they are relatively
Rica. commonly collected in pan and Malaise traps, there prob-
ably are only a few species of this genus found in Central
Triphleba Rondani. Several species, including those
formerly classified as Citrago Schmitz, are found in mid- America. The latest key to species is that of Borgmeier
to high elevation sites in Central America. Larvae are (1967).
presumably scavengers in buried, decomposing plant or
animal material, as they are elsewhere in the world. Xanionotum Brues. These flies, with their brachypter-
ous, setose females, are associated with non-Eciton army
Trophithauma Schmitz. Disney (1994) is correct in ants (Labidus, Neivamyrmex, Nomamyrmex). A key to
stating that recognition of this genus is questionable. females is given by Borgmeier (1968b), and some new
There is a single described Neotropical species, but a few information on the genus is given by Disney  & Retten-
undescribed species also exist. meyer (2007).

Trophodeinus Borgmeier. A few species of these flies Zikania Borgmeier. A single, rarely collected species,
are found at mid- to high elevations in Central America. Z. schmidti Borgmeier, is known from Costa Rica.

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Cook, C.E., J.J. Austin, & R.H.L. Disney. 2004. A mitochondrial Disney, R.H.L. & S.M. Berghoff. 2005. New species and new
12S and 16S rRNA phylogeny of critical genera of Phoridae records of scuttle flies (Diptera: Phoridae) associated with
(Diptera) and related families of Aschiza. Zootaxa 593: 1–11. army ants (Hymenoptera: Formicidae) in Trinidad and Venezuela.
Corona, E.M. & B.V. Brown. 2005. The Central American species Sociobiology 45: 887–898.
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Coville, R.E., C. Griswold, & P.L. Coville. 2000. Observations on of Phoridae (Diptera) associated with leaf-cutter ants (Hy-
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Disney, R.H.L. 1979. ��������������������������������������������
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Disney, R.H.L. 1994. Scuttle flies: the Phoridae. Chapman and lochiaceae) in Panama. European Journal of Entomology 98:
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Entomological Society of Washington 106: 751–756. predatory on gall-inhabiting insects. Proceedings of the Ento-
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lated to D. longirostrata (Enderlein) (Diptera: Phoridae). Annals Robinson, W.H.  & B.A. Foote. 1968. Biology and immature
of the Entomological Society of America 98: 55–62. stages of Megaselia aequalis, a phorid predator of slug
Kung, G.  & B.V. Brown. 2005b. Three unusually divergent new eggs. Annals of the Entomological Society of America 61:
genera of phorid flies (Diptera: Phoridae) from Costa Rica. 1587–1594.
Zootaxa 1019: 43–52. Ronna, A. 1936. Observaçôes biologicas sobre dois dipteros par-
Kung, G. & B.V. Brown. 2006. The Caribbean species of Dohrni- asitas de Apis mellifica L. (Dipt. Phoridae, Sarcophagidae).
phora Dahl (Diptera: Phoridae). Journal of Natural History 40: Revista de Entomologia 6: 1–9.
1931–1945. Ronna, A. 1937. Novos dados sobre os habitos de Melaloncha
Miller, P.L. 1984. Alternative reproductive routines in a small fly, ronnai Borgmeier (Dipt. Phoridae), endoparasita de Apis
Puliciphora borinquenensis (Diptera: Phoridae). Ecological mellifica L. Revista de Entomologia 7: 409–413.
Entomology 9: 293–302. Salt, G. 1929. A contribution to the ethology of the Meliponinae.
Morrison, L.W. 2000. Biology of Pseudacteon (Diptera: Phoridae) Transactions of the Royal Entomological Society of London
ant parasitoids and their potential to control imported Solenop- 77: 431–470.
sis fire ants. Recent Research Developments in Entomology 3: Schmitz, H. 1929. Revision der Phoriden. F. Dümmler, Bonn &
1–13. Berlin, 211 pp.
Peterson, B.V. 1982. A new species of Coniceromyia (Diptera: Seid, M.  & B.V. Brown. 2009. Behavioral observations and a
Phoridae) from Panama. Memoirs of the Entomological Society new host association of Commoptera solenopsidis (Phori-
of Washington 10: 136–138. dae) with the ant Pheidole dentata. Florida Entomologist 92:
Peterson, B.V. & P.W. Arntfield. 1971. A new species of Conicero- 309–313.
myia from Chiapas, Mexico (Dipt., Phoridae). Studia Entomo- Seevers, C.H. 1941. New termitophilous Diptera from the Neo-
logica 14: 395–398. tropics. Zoological Series of Field Museum of Natural History
Plowes, R.M., E.G. LeBrun, B.V. Brown, & L.E. Gilbert. 2009. A 24: 175–193.
review of Pseudacteon (Diptera: Phoridae) that parasitize ants of Silvestri, F. 1947. Contributo alla conoscenza dei Phoridae ter-
the Solenopsis geminata complex (Hymenoptera: Formicidae). mitofili del Brasile. Acta Pontificiae Academiae Scientarum
Annals of the Entomological Society of America 102: 937–958. 10: 281–296.
Portugal-Araújo, V. de. 1977. Contribuiçâo para o conhecimento Villa, J. & D.S. Townsend. 1983. Viable frog eggs eaten by phor-
da biologia de Pseudohypocera kerteszi (Enderlein, 1912), seu id fly larvae. Journal of Herpetology 17: 278–281.
acasalamento e captura (Diptera, Phoridae). Acta Amazonica 7: Wcislo, W.T. 1990. Parasitic and courtship behavior of Phalac-
153–166. rotophora halictorum (Diptera: Phoridae) at a nesting site of
Prado, A.P. do. 1976. Records and descriptions of phorid flies, Lasioglossum figuersi (Hymenoptera: Halictidae). Revista de
mainly of the Neotropical Region (Diptera; Phoridae). Studia Biologia Tropical 38: 205–209.
Entomologica 19: 561–609. Wcislo, W.T., L. Arneson, K. Roesch, V. Gonzalez, A. Smith, &
Ramírez, W. 1984. Biología del género Melaloncha (Phoridae), H.  Fernández. 2004. The evolution of nocturnal behaviour
moscas parasitoides de la abeja doméstica (Apis mellifera L.) en in sweat bees, Megalopta genalis and M. ecuadoria (Hy-
Costa Rica. Revista de Biologia Tropical 32: 25–28. menoptera: Halictidae): an escape from competitors and
Rettenmeyer, C.W.  & R.D. Akre. 1968. Ectosymbiosis between enemies? Biological Journal of the Linnean Society 83:
phorid flies and army ants. Annals of the Entomological Society 377–387.
of America 61: 1317–1326. Yarkulov, F. 1972. Phora holosericea (Diptera, Phoridae), a
Reyes, F. 1983. A new record of Pseudohypocera kerteszi, a pest of predator of root aphids. Zoologischeskii Zhurnal 51: 1415–
honey bees in Mexico. American Bee Journal 1983: 119–120. 1418. [In Russian.]
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SYRPHIDAE (FLOWER FLIES) 53
F. Christian Thompson, Graham E. Rotheray, and Manuel A. Zumbado

Fig. 53.1. Female of Meromacrus loewi Williston, (illustrated by F. Zeledón).

Diagnosis well developed. Abdomen slender or petiolate (Fig. 59, 60)

to elongate (Fig. 55, 57) or oval (Fig. 58).
Small to large flies (body length 4–25 mm). Fully winged; Larvae (third instar) (Figs. 62–72) and puparia (Fig. 73)
usually with holoptic males (Figs.  2–3, 23–25, 27–28); recognized by combination of characters: anal segment
females (Figs.  1, 26) and some males (Fig.  29) dichoptic; bearing single, sclerotized breathing tube (except Syrphus
shorae Fluke, in which the two tubes are separate); anus on
three ocelli present; antenna short to elongate, with distinct
anteroventral margin of anal segment, not separated from it
scape, pedicel, first flagellomere, and apical style (Figs.  2,
by transverse integumental fold; dorsum of prothorax with
5–6) or dorsal to subbasal arista (Figs. 1, 3–4, 7–29); with- longitudinal folds (inconspicuous in microdontines and
out head bristles, rarely with bristles on thorax (Cheilosia, some syrphines).
Copestylum, and Ornidia); plumula (fringed posteroventral Four basic forms of larvae exist: (1) Microdontinae:
extension of subalar sclerite) short to long, rarely absent. mandibles at apex of head skeleton, bladelike with serrated
Wing with large basal cells r, bm, and cup; with closed api- ventral margins. Larva hemispheric, with distinct lateral
cal cell (cell r4+5) (Figs. 43–53); spurious vein often present fringe (Fig.  64). Mesothorax, prothorax, and mouthparts
between radial and medial fields (Fig. 51); calypter usually concealed in ventral pocket of metathorax, which consequently

763
764
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

forms anterior margin of larva. Anal segment and poste- With specialized potlike structure, formed from mandibles
rior respiratory process short. (2) Syrphinae: apex of head and their lobes. Anal segment usually elongate, extended, and
skeleton consists of elongated, thin, and tapered labrum partially retractile (Fig. 67). Prolegs with crochets (Figs. 67,
and labium forming black sclerotized upper and lower rods; 69), except in most eumerines and cheilosines and lacking in
mandibles thin and styletlike and appear on each side of syrphines and microdontines.
these rods. Anal segment and posterior respiratory pro- Syrphid flies are easily recognized by a combination of
cess short. (3)  Rhingiini and Merodontini: mandibles also large basal cells (cells r, bm, and cup) with a closed apical
at apex of head skeleton but hooklike in form and project- cell (cell r4+5) (Fig. 51). A long spurious vein between the
ing from mouth. Anal segment and posterior respiratory radial and medial sectors is a useful diagnostic character, but
process usually short. (4) Eristalinae, exclusive of Rhingiini is not found in all species, and shorter spurious veins are
and Merodontini: mandibles reduced and inconspicuous. found in some Conopidae.

2 Ceriana 3 Ocyptamus

4 Cepa

Figs. 53.2–4. Habitus: dorsal view of (2) Ceriana cacica Walker and (3) Ocyptamus (Mimocalla) tristani Zumbado; lateral view of (4) Cepa
alex Thompson (Thompson, 1999, fig. 12, as Xela alex). Figure 2 illustrated by F. Zeledón; Figure 3 illustrated by T. Litwak.
SYRPHIDAE (flower flies) 53 765

Biology There are 202 genera and 96 non-typic subgenera of

Syrphidae currently recognized in the world, 60 of which
Most syrphid flies visit flowers, and many are pollinators, occur in the Neotropical Region. The traditional view of the
although the microdontines are only found in association higher classification of flower flies based on adult data is
with ant colonies. Males hover or rest near flowers and/or recognition of three subfamilies, 13 tribes, and 12 subtribes
breeding sites awaiting females. Syrphid larvae have a wide (Thompson, 1972; Vockeroth, 1969). However, character
range of niches. Saprophages occur in all types of wet envi- evidence from the immature stages support a different view
ronments, from tree sap to bromeliads, decaying plant parts, (Rotheray & Gilbert, 1999, 2008; Katzourakis et al., 2001),
and specialized niches such as the refuse dumps of Atta leaf- and new data from molecular sequences when used in a total
cutter ants. Predatory species also vary in the niches occupied. evidence analysis strongly suggest that neither immature nor
Some live concealed in ant nests, attacking the early stages adult characters provide a complete picture of the relation-
of ants or other myrmecophiles. Many syrphines live on ships among the genera of flower flies (Ståhls et al., 2003;
plants, attacking colonial insects such as sternorrhynchous Mengual et al., 2008). Two clades (subfamilies Microdonti-
Hemiptera����������������������������������������������������
. Given this diversity, we have included in the syn- nae and Syrphinae) are supported as monophyletic based on
opsis below more specific information on the biology of each all evidence, but the Eristalinae is not. Hence, for pragmatic
group. reasons, we follow the traditional classification here.
Because of the diverse life histories of flower flies, they
are of great importance. Adult flies are beneficial pollinators Identification
(Ssymank et al., 2009), some being used for greenhouse
pollination of flowers and seed-producing plants. A number Thompson (1999) provided a key to genera for the Neo-
of the predaceous species are valuable biological control tropical Region as a whole. The West Indian species were
agents of plant pests on agricultural crops (Tenhumberg & revised by Thompson (1981). Work on the Costa Rican fauna
Poehling, 1995), and phytophagous species have been used has stimulated revisions of many groups (see below) and will
for weed control (Cheilosia). Some of the saprophagous lead to a monograph of the Mesoamerican fauna. For other
species (Palpada, Ornidia) have been used to recycle wastes areas of the Neotropical Region, keys date back to the 1930s
from coffee and orange juice production. In Europe, syrphids and 1940s; these works are noted in Thompson et al. (1976).
are used as ecological indicators to assess environmental The following adult key includes all genera known from
quality (Sommaggio, 1999). A few species (of Eumerus and Neotropical Mexico, the West Indies, and Central America.
Merodon) are pests of ornamental flowers, and occasionally Most Central American syrphid larvae are unknown and
some species cause accidental myiasis. have yet to be reared. Data on some genera are included in
the larval key on the basis of material from other regions.
Classification This potential limitation on the utility of the key should be
borne in mind when attempting to identify larvae. General
Syrphidae are members of the cyclorrhaphan grade information on syrphid larvae can be found in Rotheray
Aschiza and have long been considered to be the sister group (1993) and references therein. Figure 62 is a morphological
to the Pipunculidae (e.g., Brauer, 1883; Hennig, 1948; Grif- map of a syrphid larva and gives the names and locations of
fiths, 1972; McAlpine, 1989; Cumming et al., 1995). While the important features.
some authors (Thompson, 1972) have split off the basal Note that the genus Nothomicrodon Wheeler, based on
clades of Syrphidae, recognizing two separate families (Micro- a single Panamanian species, N. aztecarum Wheeler (1924)
dontidae, Syrphidae), the monophyletic status and contents (Fig. 75), is only known from larvae found in the carton nest
of the Syrphidae sensu lato have remained unchanged since of the ant Azteca trigona Emery. Although it was placed in
the group was first recognized by Latreille (1802). family Syrphidae, we suspect it belongs elsewhere.

Key to the genera of Syrphidae of the northern Neotropical Region

1. Postpronotum pilose (Fig.  30); male abdomen with four unmodified pregenital segments;
tergum 5 not visible in dorsal view . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Postpronotum bare, male abdomen with five unmodified pregenital segments; tergum 5 visible in
dorsal view. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63

2. Antenna with terminal style (Figs. 2, 5, 6). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

– Antenna with subbasal dorsal arista (Figs. 4, 7–29) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

5 Callicera 7 Allograpta
6 Polybiomyia

8 Toxomerus 9 Rhopalosyrphus 10 Rhingia

11 Quichuana 12 Pseudodoros 13 Paramicrodon

14 Orthonevra 15 Ornidia 16 Microdon

Figs.  53.5–16. Heads: lateral view of (5)  Callicera erratica (Walker), (Nearctic, MND, fig.  52.10), (6)  Polybiomyia townsendi (Snow),
(MND, fig. 52.12), (7) Allograpta (Rhinoprosopa) sp., (8) Toxomerus geminatus (Say), (Nearctic, MND, fig. 52.5), (9) Rhopalosyrphus
guentherii (Lynch Arribalzaga), (10) Rhingia nigra Macquart, (11) Quichuana angustiventris Macquart, (12) Pseudodoros clavatus (Fabricius),
(MND, fig. 52.6, mouthparts removed), (13) Paramicrodon sp., (14) Orthonevra sp., (15) Ornidia obesa (Fabricius), and (16) Microdon sp.
Figures 7, 9–11, and 13–16 illustrated by D. Marques.
SYRPHIDAE (flower flies) 53 767

17 Lycopale 18 Lycastrirhyncha 19 Lepidomyia

20 Chrysotoxum 21 Chalcosyrphus 22 Argentinomyia

frgn sut

a tnt pit

23 Argentinomyia 24 Trichopsomyia 25 Cheilosia

27 Paragus 28 Melanostoma

26 Allograpta 29 Lejops

Figs.  53.17–29. Heads (concluded): lateral view of (17)  Lycopale chrysotaenia Fluke, (18)  Lycastrirhyncha nitens Bigot, (19)  Lepidomyia
micheneri (Fluke), (Nearctic, MND, fig.  52.14, mouthparts removed), (20)  Chrysotoxum derivatum Walker, (Nearctic, MND, fig.  52.4,
mouthparts removed), (21)  Chalcosyrphus (Neplas) schildi Shannon, and (22)  Argentinomyia  sp.; frontal view of (23)  Ar. rugosonasus
Williston, (24) Trichopsomyia sp., and (25) Cheilosia capillata (Loew), (Nearctic, MND, fig. 52.35, mouthparts removed); dorsal view of
(26) Al. neotropica Curran, (27) Paragus (Pandasyophthalmus) haemorrhous Meigen, (28) Melanostoma sp., and (29) Lejops (Asemosyrphus)
polygrammus Loew, (MND, fig. 52.39). Figures 17, 18, 21, 23, 24, and 27 illustrated by D. Marques; Figure 22 illustrated by R. Vargas;
Figures 26 and 28 illustrated by F. Zagonel.
Abbreviations: a tnt pit, anterior tentorial pit; frgn sut, frontogenal suture.
768
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

3. Eye pilose (Fig. 5); scutellum with ventral fringe; crossvein r-m basal to middle of cell dm. . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callicera Panzer
– Eye bare; scutellum without ventral fringe; crossvein r-m apical to middle of cell dm . . . . . . . . 4

4. Frontal prominence absent or short, much shorter than scape (Fig.  6); postmetacoxal bridge
complete or incomplete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
– Frontal prominence elongate, at least as long as scape (Fig. 2); postmetacoxal bridge incomplete.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

5. Postmetacoxal bridge complete (Fig. 38). . . . . . . . . . . . . . . . . . . . . . . . . . . Polybiomyia Shannon

– Postmetacoxal bridge incomplete, with membranous area posterodorsally to bases of hind coxae.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sphiximorpha Rondani

6. Abdomen petiolate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Monoceromyia Shannon

– Abdomen elongate, only weakly constricted (Fig. 2) . . . . . . . . . . . . . . . . . . . . . Ceriana Rondani

7. Vein R4+5 strongly sinuate (Figs. 52–53); hind femur usually with basoventral patch of black
setulae (Fig. 40). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
– Vein R4+5 straight, not or only slightly sinuate (Figs. 45, 49); hind femur without basoventral patch
of black setulae (Figs. 41–42) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

8. Cell r1 open (as in Fig. 53). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

– Cell r1 closed and petiolate (as in Fig. 52). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

9. Arista plumose, with pile many times longer than basal diameter of arista (as in Fig. 15); vein R4+5
only slightly sinuate; hind femur without basoventral patch of setulae. . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sericomyia (Arctophila) Schiner, in part
– Arista bare; vein R4+5 strongly sinuate; hind femur with basoventral patch of black setulae
(Fig. 40). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

10. Eye pilose (as in Fig. 11). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Quichuana Knab

– Eye bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

11. Ocellar triangle extremely large; frons broad in both sexes (Fig. 29); scutum with pruinose vittae;
male dichoptic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lejops (Asemosyrphus) Bigot
– Ocellar triangle normal size, small; frons not broad; scutum usually not vittate; male dichoptic or
holoptic. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

12. Dark elongate flies; thorax and abdomen without distinct pale maculae or tomentose pile, with
only short sparse pile; wing with anterior one-third dark; male broadly dichoptic. . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Habromyia Williston
– More brightly colored flies; thorax and abdomen with pale maculae and/or yellow tomentose pile;
wing with or without dark anterior margin; male narrowly dichoptic or holoptic. . . . . . . . . . . . 13

13. Frontoantennal region greatly produced anteriorly, subconical to conical (Fig. 17); scutum and
frequently abdomen with distinctive pruinose patterns; hind femur not greatly swollen . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lycopale Hull
– Frontoantennal region not produced anteriorly; scutum and abdomen without distinctive pruinose
patterns; hind femur greatly swollen. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mallota Meigen

14. Face drawn out into long slender porrect snout (Fig. 18) . . . . . . . . . . . . . . Lycastrirhyncha Bigot
– Face not produced into snout. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
SYRPHIDAE (flower flies) 53 769

15. Katepimeron bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

– Katepimeron pilose (as in Fig. 30). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

16. Eye bare; thorax usually with maculae of opaque tomentose pile. . . . . . . . . Meromacrus Rondani
– Eye pilose; thorax without tomentose pile. . . . . . . . . . . . . . . . . . . Eristalis (Eoseristalis) Kanervo

17. Meron and metepisternum with pile anterior to and/or ventral to metathoracic spiracle; eye usually
without contrasting vittae or pile; wing with or without microtrichia. . . . . . . . Palpada Macquart
– Meron and metepisternum without any pile near spiracle (as in Fig. 30); eye with contrasting
vittae of light and dark colored pile; wing bare. . . . . . . . . . . . . . . . . . Eristalis (Eristalis) Latreille

18. Arista plumose, with pile at least 3 times as long as basal diameter of arista (Fig. 15) . . . . . . . 19
– Arista bare or pubescent, with pile not more than twice as long as basal diameter of arista
(Figs. 16–22) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

19. Eye bare; meron without pile anterior to metathoracic spiracle . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sericomyia (Arctophila) Schiner, in part
– Eye pilose; meron with patch of long pile anterior to metathoracic spiracle. . . . . . . . . . . . . . . . 20

20. Face with medial and two lateral tubercles (Fig. 15); posterior anepimeron pilose; notopleuron
enlarged and produced posteriorly. . . . . . . . . . . . . . . . . . . . . . . . . . . Ornidia Lepeletier & Serville
– Face with only medial tubercle; posterior anepimeron bare; notopleuron normal, not produced. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Copestylum Macquart

21. All femora with strong short ventral spinelike setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

– Femoral spinelike setae, if present, restricted to hind femur. . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

22. Face with tubercle in both sexes (Fig. 19); antenna greatly elongate, with first flagellomere more
than twice as long as broad, frequently with scape and pedicel elongate . . . . . . Lepidomyia Loew
– Face with tubercle only in male; female with face concave; antenna short, with first flagellomere
oval or subquadrate, less than twice as long as broad, with pedicel and scape not elongate. . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myolepta Newman

23. Postmetacoxal bridge complete (Fig. 39). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

– Postmetacoxal bridge incomplete, with membranous area ventroposterior to bases of hind coxae.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43

24. Face bare, carinate; oral margin notched anteriorly; anterior tentorial pit elongate, not forming
small round pit; vein R4+5 without appendix; hind femur swollen; scutum with two pairs of
transverse yellow pruinose fasciae. . . . . . . . . . . . . . . . . . . . . . . . Sterphus (Ceriogaster) Shannon
– Face pilose, straight or convex, not carinate; oral margin evenly rounded, usually not notched
anteriorly; anterior tentorial pit small, round; vein R4+5 frequently with appendix into cell r4+5 (as
in Fig. 46); hind femur and scutum various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

25. Vein M1 processive (strongly curved towards apex), directed outwardly and cell r4+5 with acute
apex (Fig. 50); metasternum underdeveloped, bare. . . . . . . . . . . . . . . . . . . . Aristosyrphus Curran
– Vein M1 not processive, either straight or slightly recurrent and cell r4+5 with broader apex (Figs. 44,
46); metasternum various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26

26. Abdomen petiolate; metasternum underdeveloped, reduced to thin line, medially bare (Fig. 37) .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mixogaster Macquart
– Abdomen usually not petiolate, oval to elongate; if petiolate, then metasternum well developed,
not reduced, and usually pilose (as in Fig. 36). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
770
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

27. Anepimeron bare; antenna short, only about one-half as long as face; abdomen oval. . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paragodon Thompson
– Anepimeron pilose; antenna usually long, usually longer than one-half as long as face; if shorter,
then abdomen elongate, not oval. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28

28. Antenna short, less than one-half as long as face; scape not more than twice as long as broad
(Fig. 13); abdomen elongate, with parallel sides; vein R4+5 without appendix into cell r4+5 . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paramicrodon de Meijere
– Antenna long, more than one-half as long as face; scape much more than twice as long as broad
(Figs. 9, 16); abdomen frequently oval or petiolate; vein R4+5 frequently with appendix into cell
r4+5. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

29. Katepimeron pilose; abdomen petiolate. . . . . . . . . . . . . . . . . . . . . . . . Rhopalosyrphus Giglio-Tos

– Katepimeron bare; abdomen various; Microdon Meigen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

30. Vein R4+5 without appendix; occiput uniform, as wide dorsally as ventrally; hind tibia narrow, with
long hairs; abdomen elongate; small, pale orange to yellow flies. . . . . Microdon (Rhoga) Walker
– Vein R4+5 with appendix extending posteriorly into cell R4+5; other characters various. . . . . . . . 31

31. Abdomen petiolate; second segment flattened, sometimes constricted or with large basolateral
pale macula; third and fourth (males) and or fifth (females) forming club. . . . . . . . . . . . . . . . . 32
– Abdomen oval or parallel sided. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

32. Metasternum pilose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microdon (abnormis group)

– Metasternum bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

33. Vertex swollen, shiny, bare at least anteriorly; face convex, not produced ventrally; gena reduced,
narrow, usually not visible in lateral view. . . . . . . . . . . . . . . . . Microdon (Pseudomicrodon) Hull
– Vertex not swollen nor shiny, pilose; face straight, produced ventrally; gena broad. . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microdon (diaphanus group)

34. Hind tibia without hair brush along dorsal edge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35

– Hind tibia with long hair along dorsal edge, forming distinct brush of pile . . . . . . . . . . . . . . . . 40

35. Face with distinct mystax; scutellum deeply sulcate; large flies (26–28 mm). . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microdon (Syrphipogon) Hull
– Face without mystax; scutellum not deeply sulcate; smaller flies (under 20 mm) . . . . . . . . . . . 36

36. Antenna inserted under large shelflike extension of frons; face bulging and prominent ventrally;
head and thorax strongly punctate. . . . . . . . . . . . . . . . . . . . . . . . . . Microdon (Chrysidimyia) Hull
– Antenna not so inserted; face not bulging nor prominent ventrally; head and thorax not strongly
punctate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37

37. Face carinate, medially swollen and laterally sunken; hind metatarsus enlarged, quadrate; abdomen
oval, with terga and sterna overlapping; with fourth segment bent perpendicular to second . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microdon (Ceratophya) Wiedemann
– Face not carinate, evenly convex from eye to eye; abdomen oval or triangular; terga and sterna not
overlapping; abdominal apex not perpendicular to base. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38

38. Vein M1 (apical crossvein) strongly recurrent on anterior one-third, usually with appendix; second
abdominal segment with anterior margin rectangular; abdomen broadly triangular, broadest at
base. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microdon (Chymophila) Macquart
– Vein M1 rounded, not angulate, without appendix; second abdominal segment usually without
rectangular anterior margin; abdomen various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
SYRPHIDAE (flower flies) 53 771

pprn

a anepst

anepm

p anepst
kepm
mtepst
mtepm
npl wg
shield

cx 3
kepst mr

30 Eristalinus
anepst

anepm
sctl

cx 1 kepst sctl

v sctl
31 Nausigaster fringe 33 Syritta
mtst

mtst 32 Allograpta

mtst cx 3
cx 3 mtst

34 Melanostoma 37 Mixogaster
cx 3
cx 3

35 Platycheirus 36 Microdon

cx 3

mtepm cx 3
pmtcx brg mtepm
pmtcx brg

st 1 st 1
tg 1 tg 1

38 Polybiomyia 39 Microdon

Figs. 53.30–39. Thorax and thoracic structures: thoraces: lateral view of (30) Eristalinus aenus (Scopoli), (Holarctic, MND, fig. 52.64);
anterior part of (31) Nausigaster punctulata Williston, (MND, fig. 52.74); lateral view of scutellum of (32) Allograpta obliqua (Say), (Nearctic,
MND, fig. 52.63), and (33) Syritta flaviventris Macquart; ventral view of metasternum of (34) Melanostoma mellinum (Linnaeus), (Nearctic,
MND, fig. 52.70), (35) Platycheirus quadratus (Say), (Nearctic, MND, fig. 52.71), (36) Microdon piperi Knab, (Nearctic, MND, fig. 52.73),
and (37) Mixogaster delongi Hull, (Nearctic, MND, fig. 52.72); ventral view of postmetacoxal bridge of (38) Polybiomyia townsendi (Snow),
(MND, fig. 52.81) and (39) Mic. cothurnatus Bigot, (Nearctic, MND, fig. 52.79). Figure 33 illustrated by F. Zagonel.
Abbreviations: a anepst, anterior anepisternum; anepm, anepimeron; anepst, anepisternum; cx, coxa; kepm, katepimeron; kepst, katepisternum;
mr, meron; mtepm, metepimeron; mtepst, metepisternum; mtst, metasternum; npl wg shield, notopleural wing shield; p anepst, posterior
anepisternum; pmtcx brg, postmetacoxal bridge; pprn, postpronotum; sctl, scutellum; st, sternite; tg, tergite; v sctl fringe, ventral scutellar
fringe.
772
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

39. First two abdominal segments with three distinct depressions, two lateral and one anteromedial;
abdomen more or less parallel sided; scutellum with apical calcar. . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microdon (Omegasyrphus) Loew
– First two abdominal segments without such depressions; abdomen oval or triangular; scutellum
with or without calcar. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microdon (Microdon) Meigen

40. Abdomen oval or rectangular, not triangular. . . . . . . . . . . . . . . . . . . . Microdon (Ubristes) Walker

– Abdomen triangular, broad basally, strongly narrowed apically. . . . . . . . . . . . . . . . . . . . . . . . . 41

41. Male first flagellomere furcate; female first flagellomere greatly elongate, about 2.5 times as long
as scape; first tarsomere of midleg enlarged, quadrate; vertex greatly produced dorsally. . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microdon (Carreriamyia) Doesburg
– First flagellomere not furcate, usually not greatly elongate, usually about twice as long as scape;
first tarsomere of midleg not enlarged, elongate; vertex not greatly produced. . . . . . . . . . . . . . 42

42. Abdomen short, almost equilateral in shape . . . . . . . . . . . . . . . . Microdon (Hypselosyrphus) Hull

– Abdomen much longer, not equilateral . . . . . . . . . . . . . . . . . . . . . . Microdon (Stipomorpha) Hull

43. Vein M1 recurrent or perpendicular; cell r4+5 with obtuse or rectangular apex (Figs. 47, 49) . . . 44
– Vein M1 processive, directed apically; cell r4+5 with acute apex (Figs. 45, 50). . . . . . . . . . . . . . 46

44. Eye bare, usually with metallic pattern (Fig. 14); first flagellomere elongate, at least twice as long
as broad; antenna usually elongate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Orthonevra Macquart
– Eye sparsely or densely pilose, without pattern; first flagellomere orbicular or oval, less than twice
as long as broad; antenna short . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45

45. Crossvein r-m basal to middle of cell dm; vein M1 once angulate (Fig. 49); propleuron bare; head
fitting close to thorax, thus occiput reduced laterally to thin line, anterior thoracic spiracle hidden
from lateral view and postpronotum greatly reduced . . . . . . . . . . . . . . . . . . . Alipumilio Shannon
– Crossvein r-m apical to middle of cell dm; vein M1 twice angulate (Fig. 47); propleuron pilose;
head not as above, occiput broad on dorsal one-third, anterior thoracic spiracle visible laterally
and postpronotum normal; Old World, introduced elsewhere. . . . . . . . . . . . . . [Eumerus Meigen]

46. Anepisternum uniformly raised, not differentiated into flattened anterior and convex posterior part
(Fig. 31); antennal cavities broadly separated; scutum with large flaplike extension above wing
base (notal wing shield); body densely punctate, with punctures large. . . . . Nausigaster Williston
– Anepisternum clearly differentiated into flat anterior part and convex posterior part; antennal
cavities usually confluent, rarely narrowly separated; notal wing shield absent; body usually not
punctate; if punctate, then antennal cavities confluent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47

47. Eye densely pilose (as in Fig. 24) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48

– Eye bare or nearly so, rarely with sparse short pile (as in Fig. 21). . . . . . . . . . . . . . . . . . . . . . . 50

48. Oral margin notched anteriorly; anterior tentorial pit elongate, not forming small pit (Fig. 25). . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cheilosia Meigen, in part
– Oral margin evenly rounded, not notched anteriorly; anterior tentorial pit small, round (Fig. 24).
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49

49. Anterior anepisternum pilose, with row of long erect pile posterior to mesothoracic spiracle;
katepimeron pilose. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trichopsomyia Williston
– Anterior anepisternum bare; katepimeron bare; Nearctic Region, Chile. . . . . . . . [Pipiza Meigen]
SYRPHIDAE (flower flies) 53 773

50. Face drawn out into long porrect snout (Fig. 10); costa and vein R4+5 ending well posterior to apex
of wing. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhingia Scopoli
– Face not produced into snout; costa and vein R4+5 ending at or anterior to apex of wing. . . . . . 51

51. Antenna elongate, much longer than face; scape more than 3 times as long as broad . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cacoceria Hull
– Antenna short, only as long as or shorter than face; scape not more than twice as long as broad. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52

52. Metasternum reduced, difficult to see as mid- and hind legs are close together, pilose; face straight
or slightly concave in profile; crossvein  r-m basal to middle of cell  dm and at level of end of
vein  sc; scutellum without bristles; abdomen oval, strongly recurved; small to medium-sized,
compact, metallic flies (Fig. 4). . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cepa Thompson & Vockeroth
– Metasternum not reduced; other characters various, but not occurring all together in combination .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53

40 Palpada 41 Allograpta

42 Syritta

Figs.  53.40–42. Legs: lateral view of male hind leg of (40)  Palpada scutellaris (Fabricius), (41)  Allograpta exotica (Wiedemann), and
(42) Syritta flaviventris Macquart. Figures 40–42 illustrated by D. Marques.
774
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

53. Metasternum bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54

– Metasternum pilose, with pile as long as that on hind coxa. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57

54. Face straight in profile, with distinct carinae, rarely with oral margin slightly produced. . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sterphus (Crepidomyia) Shannon, in part
– Face either concave or tuberculate in profile, not straight. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55

55. Face concave; wing not bicolored. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xylota Meigen

– Face tuberculate, usually with distinct tubercle, or carinate; wing bicolored or not. . . . . . . . . . 56

56. Abdomen oval, with second tergum wider than third tergum; male narrowly dichoptic, with eyes
separated by aristal width only; crossvein r-m at midlength of cell dm; wing not bicolored, hyaline
or uniformly gray. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sterphus (Sterphus) Philippi
– Abdomen petiolate, with third tergum about twice as wide as base of second tergum; male
holoptic; crossvein r-m beyond middle of cell dm; wing bicolored, with anterior margin dark and
posterior part hyaline. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mutillimyia Hull

57. Cell r1 closed and petiolate (Fig. 45). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Milesia Latreille

– Cell r1 open. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58

58. Anterior anepisternum pilose; hind femur with single small ventroapical spine. . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Spilomyia Meigen
– Anterior anepisternum bare; hind femur without single small ventroapical spine (Fig. 42). . . . 59

59. Scutellum without ventral fringe (Fig.  33); wing extensively bare, with microtrichia almost
completely absent on basal two-thirds or more, sparse and scattered on apical one-third or less;
metathoracic pleuron pilose, with some fine pile ventral to spiracle; Old World, introduced into
North and South America. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Syritta Lepeletier & Serville]
– Scutellum with ventral pile fringe (as in Fig. 32); wing extensively microtrichose, apical one-half
or more densely microtrichose, with only limited bare areas on basal one-third or less;
metathoracic pleuron bare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60

60. Crossvein r-m basal, basal to middle of cell dm and at level of end vein sc; scutellum with
marginal bristles; face distinctly tuberculate (Fig. 25); abdomen oval; medium-sized, black flies.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cheilosia Meigen (in part)
– Crossvein r-m medial to apical, at or beyond middle of cell dm and beyond level of end of vein sc;
scutellum without bristles; other characters various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61

61. Face indistinctly tuberculate, not carinate nor concave; abdomen oval; large, robust bumble bee
mimic, with long pilosity on body. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Criorhina Meigen
– Face concave and subcarinate; abdomen elongate; small narrow flies, not bumble bee mimics,
without long pilosity on body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62

62. First flagellomere oval, only as long as broad; face straight; posterior thoracic spiracle large, about
as large as first flagellomere. . . . . . . . . . . . . . . . . . . . . . Sterphus (Crepidomyia) Shannon, in part
– First flagellomere elongate, longer than broad; face slightly but distinctly concave in profile
(Fig. 21); posterior thoracic spiracle small, much smaller than first flagellomere. . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chalcosyrphus (Neplas) Porter

63. Anterior anepisternum usually with some distinct short pile posterodorsally; metathoracic pleuron
usually with some long erect or subappressed pile ventral to spiracle; with pile on one of these two
places (see Fig. 30). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
– Anterior anepisternum bare; metathoracic pleuron bare (see Fig. 30 for reference). . . . . . . . . . 66
SYRPHIDAE (flower flies) 53 775

R4+5 R4+5

M1
M1
43 Salpingogaster 44 Mixogaster
r1 R4+5 R4+5

r4+5
r4+5
M1

45 Milesia 46 Microdon
h Sc sprs vn R1
R2+3

R4+5
r-m R4+5
M1
CuP

CuA2
M1
M2
A2 A1 CuA1 dm-cu

47 Eumerus 48 Dideomima

R4+5
r-m R4+5

M1 M1
r4+5
r4+5
49 Alipumillo 50 Aristosyrphus
sprs vn r1 r1
R4+5

bm
cup r4+5

51 Allograpta 52 Palpada
r1
R4+5

u calyp

l calyp

hlt

53 Quichuana 54 Syrphus

Figs. 53.43–54. Wings and portion of thorax: dorsal view of wing of (43) Salpingogaster nigra Schiner, (44) Mixogaster conopsoides
Macquart, (45) Milesia pulchra Williston, (46) Microdon fulgens Wiedemann, (47) Eumerus strigatus (Fallén), (Holarctic, MND, fig. 52.48),
(48)  Dideomima  sp., (49)  Alipumilio femoratus Shannon, (50)  Aristosyrphus  sp., (51)  Allograpta exotica (Wiedemann), (52)  Palpada
mexicana (Macquart), (53) Quichuana sp., and lower calypter of (54) Syrphus ribesii Linnaeus, (Palearctic, MND, fig. 52.62). Figures 43, 44,
46, 48, and 49 illustrated by T. Litwak; Figures 45 and 50–53 illustrated by L. Donzo.
Abbreviations: hlt, halter; l calyp, lower calypter; sprs vn, spurious vein; u calyp, upper calypter.
776
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

64. Metasternum and/or eye pilose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ocyptamus Macquart, in part

– Metasternum and eye bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65

65. Eye on posterior margin with distinct triangular emargination at or dorsal to level of insertion
of antenna; facial tubercle well developed, beginning immediately ventral to antennal bases and
sometimes laterally compressed; face often produced anteriorly, sometimes strongly so (Fig. 8);
abdomen usually oval, with at least weak premarginal sulcus, not strongly petiolate or long and
thin; wing lacking dark markings; male genitalia with sclerotized, short to long triangular process
arising from fused bacilliform sclerites and projecting caudad between bases of surstyli; phallus
simple, unsegmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Toxomerus Macquart
– Eye on posterior margin with emargination usually indistinct or shallow and rounded, if distinct
and subtriangular, then situated ventral to level of antennal insertion; facial tubercle usually weak,
not as described above, if distinct, then with strong concavity between it and antennal bases;
face not produced anteriorly; abdomen usually petiolate, frequently long and thin, rarely parallel
sided, not oval; wing frequently with dark markings; male genitalia without sclerotized process
projecting between bases of surstyli, with at most weak semimembranous process in this position;
phallus complex, segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ocyptamus Macquart, in part

66. Face and scutellum black in ground color. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67

– Face and/or scutellum partially pale in color, usually yellow or yellowish-brown in ground color.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71

67. Abdomen petiolate, petiole much narrower than thorax (as in Figs. 59–60); face without tubercle,
flat. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leucopodella Hull
– Abdomen oval (Fig. 58) or with parallel sides (Fig. 55), not narrower than thorax; face tuberculate.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68

68. Antennal cavities confluent (as in Fig. 24); metathoracic pleuron with fine subappressed pile
ventrad of spiracle; katepisternum with pile patches continuous anteriorly; face straight; hind
coxa with pile tuft at posteromedial apical angle. . . . . . . . . . . . . . . . . . . . . . . Xanthandrus Verrall
– Antennal cavities separated (Fig. 23); metathoracic pleuron bare; katepisternum with pile patches
usually broadly separated throughout; face and legs various. . . . . . . . . . . . . . . . . . . . . . . . . . . . 69

69. Metasternum greatly reduced, with deep posterior incision on each side so that median portion
narrowly joined to lateral arms (Fig. 34); face not produced below, with small tubercle; facial
pruinescence not punctate nor rippled; legs of male slender, without bristles, hair tufts, or
modified hairs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Melanostoma Schiner
– Metasternum not greatly reduced, with median portion broadly joined to lateral arms (Fig. 35);
face various, almost straight in profile with weak tubercle or moderately or strongly produced
forward below, sometimes with pruinescence forming punctate or rippled pattern; legs of male
sometimes slender and unornamented, sometimes with foretibia and tarsus broadened, or
sometimes with femora or tibiae with bristles, hair tufts, or modified hairs. . . . . . . . . . . . . . . . 70

70. Face frequently produced anteriorly, densely pruinose and with puncturelike bare maculae or
ripples; antenna short, with scape not more than twice as long as broad; male legs at least with
strong black setae on foretibia; abdomen without pale-colored maculae, with silvery-gray maculae.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Platycheirus (Carposcalis) Enderlein
– Face straight in profile, not produced anteriorly, usually uniformly pruinose, rarely with shiny
(bare) punctate maculae; antenna frequently long, with scape much longer than broad (Fig. 22);
male legs simple; abdomen frequently with pale-colored maculae. . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Argentinomyia Lynch Arribálzaga, in part

71. Metasternum pilose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72

– Metasternum bare (Fig. 35). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
SYRPHIDAE (flower flies) 53 777

55 Allograpta 56 Syrphus premg sulc

tg 1
tg 1

tg 5
tg 4

st 8

57 Paragus 58 Syrphus

tg 1
tg 1

tg 1

tg 5
tg 5
tg 5

59 Salpingogaster 60 Ocyptamus 61 Chrysotoxum

Figs. 53.55–61. Abdomens: dorsolateral view of (55) Allograpta exotica (Wiedemann) and (56) Syrphus phaeostigma Wiedemann; dorsal
view of (57) Paragus (Pandasyophthalmus) haemorrhous Meigen, (MND, fig. 52.97), (58) Sy. phaeostigma, (redrawn by A.G. Brenes from
Fluke 1942, fig. 3), (59) Salpingogaster (Salpingogaster) punctifrons Curran, (MND, fig. 52.95), (60) Ocyptamus (Mimocalla) bonariensis
(Brèthes), and (61) Chrysotoxum derivatum Walker, (Nearctic, MND, fig. 52.94, as Chrysotoxum integre Williston). Figures 55, 56, and 60
illustrated by D. Marques.
Abbreviations: premg sul, premarginal sulcus; st, sternite; tg, tergite.
778
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

72. Abdomen with distinct strong premarginal sulcus, extending from middle of tergum 2 to end of
tergum 5 (as in Fig. 56); scutum entirely dark or at most obscurely yellowish pruinose laterally;
face not strongly produced anteriorly; oral opening not more than 2.5 times as long as broad . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eupeodes (Metasyrphus) Matsumura
– Abdomen without premarginal sulcus (Fig.  55); scutum often with well-defined bright yellow
lateral or sublateral vitta extending at least from postpronotum to suture; face often produced
anteriorly so that oral opening may be more than 3 times as long as broad . . . . . . . . . . . . . . . . 73

73. Scutellum with ventral fringe absent or nearly so on at least median third; male terminalia
extremely large and globose. . . . . . . . . . . . . . . . . . . . . . . . . . Sphaerophoria Lepeletier & Serville
– Subscutellar fringe complete, well developed, moderately dense; male terminalia small, incon-
spicuous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74

74. Face produced anteriorly; oral apex distinctly more prominent than antennal bases; oral opening
3 or more times longer than broad; facial tubercle high, distinctly differentiated. . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Allograpta (Fazia) Shannon
– Face straight, not produced anteriorly; oral apex distinctly less prominent that antennal bases; oral
opening only about 2 times as long as broad; facial tubercle usually low (except in A. armillata
Fluke), not differentiated dorsally. . . . . . . . . . . . . . . . . . . . Allograpta (Allograpta) Osten Sacken

75. Eye pilose (as in Fig. 20). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76

– Eye bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79

76. Tergum 1 well developed, especially medially on disc where it is frequently one-half as long as
tergum 2 and extends well beyond scutellum (as in Fig. 57), sublaterally about three-quarters as
long as tergum 2; terga minutely punctate; length 7.5 mm or less. . . . . . . . . . . . Paragus Latreille
– Tergum 1 greatly reduced, on disc frequently almost linear and practically covered by scutellum,
sublaterally at most one-half as long as tergum 2; terga not punctate; length 7.5 mm or more,
usually about 10 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77

77. Antenna elongate, as long as or longer than face; first flagellomere 4 times as long as broad, longer
than arista (Fig. 20); abdomen strongly convex dorsally, strongly marginated, with posterolateral
angles of terga projecting (Fig. 61) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chrysotoxum Meigen
– Antenna short, shorter than face; first flagellomere short, at most about 2.5 times as long as broad,
shorter than arista; abdomen not strongly convex dorsally; terga without posterolateral angles
projecting. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78

78. Calypter with ventral lobe pilose, with long, coarse, erect yellow pile dorsally, especially on
posteromedian portion (Fig. 54). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Syrphus Fabricius, in part
– Calypter with ventral lobe bare, without pile. . . . . . . . . . . . . . . . . . . . . . . . Dasysyrphus Enderlein

79. Hind femur with distinct anteroventral and posteroventral rows of spinose setae on apical one-half;
vein M1 abruptly and strongly sinuate; vein R4+5 slightly to strongly sinuate (Fig. 43); postmetacoxal
bridge complete; abdomen strongly petiolate (Fig. 59); Salpingogaster Schiner. . . . . . . . . . . . 80
– Hind femur without spinose setae; vein M1 at most weakly sinuate; vein R4+5 usually straight or
nearly so, if strongly sinuate, then postmetacoxal bridge incomplete and/or abdomen oval. . . . 81

80. Vein R4+5 strongly sinuate (Fig. 43); first tergum produced laterally into strong spur (Fig. 59);
upper occipital cilia reduced to single row . . . . . . . . . . . Salpingogaster (Salpingogaster) Schiner
– Vein R4+5 only weakly sinuate; first tergum not produced into spur (as in Fig. 60); upper occipital
cilia in three to four rows. . . . . . . . . . . . . . . . . . . . . . . . . Salpingogaster (Eosalpingogaster) Hull
SYRPHIDAE (flower flies) 53 779

81. Calypter with ventral lobe pilose, with long, coarse, erect yellow pile dorsally, especially on
posteromedian portion (Fig. 54). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Syrphus Fabricius, in part
– Calypter with ventral lobe bare, without long pile. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82

82. Thorax without yellow maculae except on scutellum. . . . . . . . . . . . . . . . . . . Pseudodoros Becker

– Postpronotum yellow; scutum with lateral yellow vitta at least in front of suture; anepisternum
and katepisternum partially yellow; frequently pleuron more extensively yellow . . . . . . . . . . . 83

83. Vein R4+5 strongly sinuate (Fig.  48); face not produced anteriorly, with antennal bases slightly
anterior to oral margin; oral opening less than twice as long as broad; abdomen oval, with strong
premarginal sulcus (as in Figs. 56, 58) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dideomima Vockeroth
– Vein R4+5 straight or nearly so (Fig. 51); face produced or not; oral opening various; abdomen
petiolate or parallel-sided, without premarginal sulcus (Fig. 55) . . . . . . . . . . . . . . . . . . . . . . . . 84

84. Face vertical, not produced anteriorly, with antennal bases slightly anterior to oral margin; oral
opening less than twice as long as broad; antenna elongate, with scape and first flagellomere twice
as long as broad (Fig. 22). . . . . . . . . . . . . . . . . . . . . . . . Argentinomyia Lynch Arribálzaga, in part
– Face strongly produced anteriorly, with oral margin greatly anterior to antennal bases; oral
opening 3 or more times as long as broad (Fig. 7); antenna short, with scape and first flagellomere
only slightly longer than broad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85

85. Face with abrupt tubercle; abdomen parallel-sided. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Allograpta (Costarica) Mengual & Thompson
– Face with indistinct or no tubercle; abdomen petiolate. . . . . . . . . Allograpta (Rhinoprosopa) Hull

Key to the subfamilies and genera of larvae of Syrphidae of Central America

1. Larva with metathorax at front of body, prothorax and mesothorax narrow and telescoped into
pocket underneath metathorax (Fig. 64); locomotory organs consisting of slight projections on
ventral surface without crochets or suckers; band of setae around lateral margins of body; mandibles
bladelike with serrated ventral margin; living in and around ant nests. . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . MICRODONTINAE
– Larva with prothorax at front of body, not telescoped into ventral pocket of metathorax (Figs. 62–63,
65–72); usually lacking marginal band of setae (except some Copestylum living in bromeliads
(Fig. 70), but these have crochets and/or suckers (Fig. 71)); mandibles without serrated ventral
margin; various habitats. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2. Anal segment without lappets (fleshy projections on lateral margins of anal segment) (Fig. 63);
thorax narrower than abdomen; apex of head skeleton elongate and pointed (serrated apically in
leaf-mining Allograpta); ventral surface smooth, without setae, prolegs, or crochets; living in and
on plant surfaces; often with color pattern; SYRPHINAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
– Anal segment with up to three pairs of lappets (Fig. 66) (reduced and inconspicuous in long-tailed
larvae (Fig. 67)); thorax as broad or broader than abdomen (Fig. 68); ventral surface usually with
setae, prolegs, and crochets (Fig. 69) often present; living in decaying organic material, e.g., mud,
but also in decay on live and dead plants; ERISTALINAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

3. Larva translucent, white or cream, lacking bright color . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

– Larva with bright color pattern formed from hemolymph pigments, fat bodies, or markings on
body surface. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
780
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

4. Ventral surface with at least one sucker; tip of posterior respiratory process elongate (Fig. 65),
with one pair of spiracles above other two pairs; predatory in bromeliads. . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ocyptamus Macquart, in part
– Ventral surface without suckers; tip of posterior respiratory process flat with spiracles all on
one plane; predatory in various habitats. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

5. Tip of anal segment with pair of rounded projections; posterior respiratory process with spiracles
short, not extending over sides of dorsal plate; one European species predatory on gall-inducing
psyllids and one Australasian species in ant nests. . . . . . . . . . . . . . . . . . . Trichopsomyia Williston
– Tip of anal segment without rounded projections; spiracles elongate, extending over sides of
dorsal plate; in Europe, found in or around ant nests feeding on ant-attended Sternorrhyncha on
plant roots. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chrysotoxum Meigen

6. Larva with dorsal projections on abdominal segments 1–7 (viewed from side). . . . . . . . . . . . . . 7

– Larva without dorsal projections on abdominal segments 1–7. . . . . . . . . . . . . . . . . . . . . . . . . . . 8

7. Tip of anal segment with pair of posteriorly directed tapering projections. . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dasysyrphus Enderlein
– Four bristlelike projections between posterior respiratory process and tip of anal segment. . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paragus Latreille

8. Dorsal surface of larva with aggregated groups of dark, sclerotized spicules; stem of posterior
respiratory process short with spiracles almost reaching integument. . . . Eupeodes Osten Sacken
– Dorsal surface of larva without dark sclerotized spicules, vestiture absent or fleshy; stem of
posterior respiratory process longer, spiracles not reaching integument. . . . . . . . . . . . . . . . . . . . 9

isg intg fold sg bnd

a spr
sens

long fold p res proc

a fold
lap

antmx
cmplx

d lip
mth an sg
l lip prth an
msth mtth 1st abd sg 2nd abd sg

62 Third stage syrphid larva

Fig. 53.62. Morphological map of a third stage syrphid larva, lateral view, dorsal side uppermost, segments 3–6 not shown. Illustrated by
G.E. Rotheray.
Abbreviations: a fold, anterior fold; a spr, anterior spiracle; abd sg, abdominal segment; an, anus; an sg, anal segment; antmx cmplx,
antennomaxillary complex; d lip, dorsal lip; isg intg fold, intra-segmental integumental fold; l lip, lateral lip; lap, lappets; long fold,
longitudinal fold on dorsum of prothorax; msth, mesothorax; mth, mouth; mtth, metathorax; p res proc, posterior respiratory process;
prth, prothorax; sen, sensilla; sg bnd, segmental boundary.
SYRPHIDAE (flower flies) 53 781

p res proc

p res proc

64 Microdon

sucker

63 Syrphus

p res proc

65 Ocyptamus

a spr pup spr

lap

p res proc

a fold
antmx
cmplx
l lip
msth
prlg
an
66 Ornidia

Figs. 53.63–66. Third stage larvae: lateral view of (63) Syrphus sp. feeding on an aphid; dorsolateral view of (64) Microdon ruficrus
Williston, (Nearctic, MND, fig. 52.107); ventral view of (65) Ocyptamus sp.; and lateral view of (66) Ornidia major Curran, (Rotheray et al.,
2005, fig. 1). Figure 63 illustrated by G.E. Rotheray; Figure 65 illustrated by A.G. Brenes.
Abbreviations: a fold, anterior fold; a spr, anterior spiracle; an, anus; antmx cmplx, antennomaxillary complex; lap, lappets; l lip, lateral lip;
msth prlg, mesothoracic proleg; p res proc, posterior respiratory process; pup spr, pupal spiracle.
782
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

9. Posterior respiratory process with dorsal spurs; spiracles dark lined . . . . . . . . . Syrphus Fabricius
– Posterior respiratory process without dorsal spurs; spiracles pale in outline but may become dark
after pupariation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

10. Apicoposterior margin of posterior respiratory process with posterior triangular-shaped projection;
spiracles straight and parallel to each other; predatory on caterpillars . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xanthandrus Verrall
– Apicoposterior margin of posterior respiratory process smooth, without posterior triangular-shaped
projection; spiracles not all parallel to each other; lifestyle various. . . . . . . . . . . . . . . . . . . . . . 11

11. Larva subrectangular in cross-section; mid-dorsal chevron-shaped fat bodies on abdominal

segments; dorsal lateral margins of abdomen marked by thin, linear fat body. . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Platycheirus Lepeletier & Serville
– Larva flattened, subcyclindrical or irregularly shaped in cross-section; fat bodies not arranged as
above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

12. Posterior respiratory process apex longer than broad . . . . . . . . . . . . Ocyptamus Macquart, in part
– Posterior respiratory process apex shorter than broad. . . . . . . . . . . . . . . Allograpta Osten Sacken

Note: The poorly-known larvae of Toxomerus Macquart and Pseudodoros Becker may also key out here.

13. Anterior fold of prothorax with fleshy vestiture; without posteriorly directed sclerotized spicules.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
– Anterior fold of prothorax with posteriorly directed sclerotized spicules. . . . . . . . . . . . . . . . . . 17

14. Larva with mouthhooks usually protruding from mouth; in exuding tree sap. . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Alipumilio Shannon
– Larva without protruding mouthhooks; various habitats. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

15. Larva with fleshy ridged mandibular lobes external, surrounding lateral margins of mouth;
posterior end of abdomen coated in dark, sticklike projections. . . . . . . . . . . . . . . Rhingia Scopoli
– Larva with mandibular lobes internal, not surrounding lateral margins of mouth; mouth edge
smooth; posterior end without dark, sticklike projections. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

16. Anal segment with setae not borne on fleshy papillae; anterior spiracles absent; crochets arranged
in linear rows, little developed on abdominal segments  1–4; living in decaying vegetation in
ponds and streams . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Orthonevra Macquart
– Anal segment with tufts of setae borne on fleshy papillae; anterior spiracles present; crochets
arranged in curved rows, well developed on abdominal segments 1–7; living in tree holes. . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myolepta Newman

17. Thorax with dorsal or lateral brown or black sclerotized hooks (as in Fig.  74) (sclerotized
structures larger than spicules on anterior fold). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
– Thorax without hooks; dorsal and lateral margins of thorax may have isolated groups of spicules
of mixed size . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

18. Anal segment narrow, elongate and tail-like, about as long as rest of body; anterior fold of prothorax
with up to four pairs of laterally directed sclerotized pads bearing hooks (Fig. 74); living in
bromeliad tanks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Quichuana Knab, in part
– Anal segment narrowing towards apex, not tail-like, shorter than rest of body; anterior fold of
prothorax with spicules; sclerotized pads bearing hooks on dorsal and lateral margins of thorax;
various habitats. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
SYRPHIDAE (flower flies) 53 783

p res proc

an sg

67 Meromacrus
prlg

p res proc

68 Copestylum crochet

p res proc

69 Copestylum

p res proc

70 Copestylum sucker

p res proc

71 Copestylum

Figs. 53.67–71. Third stage larvae (continued): lateral view of (67) Meromacrus sp.; dorsal (68) and ventral (69) views of Copestylum sp.;
dorsal (70) and ventral (71) views of Copestylum sp. Figures 67–71 illustrated by A.G. Brenes.
Abbreviations: an sg, anal segment; p res proc, posterior respiratory process; prlg, proleg.
784
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

p res proc

72 Copestylum lap

pup spr

p res proc

73 Copestylum

74 Quichuana
75 Nothomicrodon

Figs. 53.72–75. Third stage larvae (concluded) and puparium: dorsal view of larva of (72) Copestylum sp., (Rotheray et al., 2007, fig. 87);
lateral view of puparium of (73)  Copestylum  sp.; frontal view of head of larva of (74)  Quichuana  sp.; and lateral view of larva of
(75) Nothomicrodon aztecarum Wheeler, (Wheeler, 1924, fig. 2a). Figures 73 and 74 illustrated by A.G. Brenes.
Abbreviations: lap, lappets; p res proc, posterior respiratory process; pup spr, pupal spiracle.
SYRPHIDAE (flower flies) 53 785

19. Larva with one or two pairs of hooks on lateral margin of thorax; anal segment slightly extended,
up to 1.5 times length of abdominal segment 7; living under bark in wet conditions. . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chalcosyrphus (Neplas) Porter
– Larva with hooks on dorsal surface of thorax; anal segment more than 1.5 times length of abdominal
segment 7; various habitats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

20. Mid-dorsum of thorax with “Y” or triangular-shaped sclerotized pad bearing one to two pairs of
small and large posteriorly directed hooks; lateral to this pad with pair of “cow-horn” laterally
directed hooks; living in tree holes and decaying tree roots . . . . . . . . . . . . . . . . Criorhina Meigen
– Mid-dorsum of thorax without “Y” or triangular-shaped sclerotized pad bearing one to two pairs
of small and large posteriorly-directed hooks; various habitats. . . . . . . . . . . . . . . . . . . . . . . . . . 21

21. Thorax with one pair of sclerotized pads bearing one to three laterally directed hooks. . . . . . . 22
– Thorax with groups of hooks mid-dorsally and/or laterally in addition to main pair of sclerotized
pads bearing three or more hooks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

22. Larva with blue-colored hemolymph; abdominal segments 1–7 each with pair of separate prolegs
oval shaped in outline; living in decaying cacti and terrestrial bromeliads. . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nausigaster Williston
– Larva white to pale gray, not blue; prolegs on abdominal segments 1–7 fused medially forming
figure-8 shape in outline; living in tree holes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callicera Panzer

23. Dorsolateral pads bearing more than three hooks; mid-dorsum of prothorax lacking single pair of
hooks; living in tree holes and decaying tree roots . . . . . . . . . . . . . . . . . . . . . . . . Milesia Latreille
– Dorsolateral pads bearing one to three hooks; mid-dorsum of prothorax with single pair of hooks;
living in tree holes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Spilomyia Meigen

24. Base of first pair of lappets with short fleshy projection. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

– Base of first pair of lappets, if clearly present, without short fleshy projection. . . . . . . . . . . . . . 26

25. Three pairs of approximated lappets about equal in length and covered with setae (Fig. 66). . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ornidia Lepeletier & Serville
– One of first two pairs of lappets longer than other (Fig. 72), or lappets short and inconspicuous
(Figs. 68–70), or, if lappets about equal, then bare, not covered with setae . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Copestylum Macquart

26. Anterior fold of prothorax with dense coating of spicules discontinuous, divided into three or
four groups with clear gaps; living in exuded tree sap and decaying cacti. . . . . Ceriana Rondani,
Polybiomyia Shannon
– Anterior fold of prothorax with continuous coating of spicules, not divided; various habitats. . 27

27. Anal segment not as long as body length (mouth to anus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28

– Anal segment extended to body length or more; long-tailed or rat-tailed larvae (Fig. 67). . . . . 29

28. Anal segment with three pairs of about equally long and equidistant lappets. . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sterphus (Ceriogaster) Shannon,
Xylota Meigen
– Anal segment with first or second or both lappets reduced and inconspicuous. . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepidomyia Loew,
Myolepta Newman
786
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

29. Anterior spiracles about as tall as broad, flat tipped and not retractile . . . . . . . Sericomyia Meigen
– Anterior spiracles longer than broad and retractile into pockets on thorax (often partially or
completely retracted in preserved and live specimens). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

30. Abdominal prolegs with crochets arranged in two main transverse rows. . . . . . . . . . . . . . . . . . 31
– Abdominal prolegs with crochets in three or more distinct semicircular rows (arranged in curved,
not transverse row). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

31. Abdominal segments 2–6 with sensilla 4–6 in the same horizontal plane. . . . . . . Mallota Meigen
– Abdominal segments 2–6 with sensilla 4 above 5 and 6. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32

32. Thorax with hooks in addition to spicules (Fig. 74) . . . . . . . . . . . . . . . . Quichuana Knab, in part
– Thorax lacking hooks, only spicules present. . . . . . . . . . . . . . . . . . . . . . Eristalis Latreille, in part

33. Spicules on upper margin of anterior fold interrupted and not forming continuous line and not
extending to dorsum of prothorax (Fig. 74). . . . . . . . . . . . . . . . . . . . . . . Quichuana Knab, in part
– Spicules on upper margin of anterior fold continuous, either forming complete transverse line or
extending from anterior fold on to dorsum of prothorax. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

34. Lateral margins of abdomen without rows of setae or, if present, only on segments 5 to 8 . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eristalis Latrielle, in part,
Palpada Macquart
– Lateral margins of abdominal segments 1–8 with rows of conspicuous setae, becoming shorter
towards head (Fig. 67) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35

35. Spicules of anterior fold long, longer than wide at base; abdominal sensilla on short and
inconspicuous fleshy projections not as tall as wide. . . . . . . . . . . . . . . . . . . Habromyia Williston
– Spicules of anterior fold short, shorter than wide at base; abdominal sensilla on conspicuous
fleshy projections, as tall as wide or taller. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36

36. Anteroventral corners of metathorax with groups of spicules (Fig. 67) . . . . Meromacrus Rondani
– Anteroventral corners of metathorax without spicules . . . . . . . . . . . . . . . . . . . . . . . Lycopale Hull

Synopsis of the fauna Region. The genus is currently organized into eight groups
(Mengual et al., 2009), of which four  subgenera occur in
About 5825 species of Syrphidae are currently recognized. Central America. The typical subgenus (Allograpta) is cos-
The greatest species richness is in the Neotropical Region, mopolitan and has nine species in Central America; the larvae
where some 1560 have been described, but many more await are predators of aphids and relatives (Hemiptera: ��������
Sternor-
description and discovery. In the last few years in Costa Rica rhyncha). The subgenus Costarica Mengual & Thompson is
alone, some 265 new species have been discovered, but have endemic to Costa Rica and contains only two species; the
not yet been described. The estimated total number of flower larvae are stem-borers. The subgenus Fazia Shannon is a
fly species for Central America is nearly 700. Neotropical endemic, with 22 species known from Central
America; the larvae are leaf-miners (Nishida et al., 2003).
Alipumilio Shannon. Found from Mexico to Brazil, this The subgenus Rhinoprosopa Hull is also a Neotropical
genus includes four species, of which only A. nigrocoeruleus endemic, with three species known from Central America;
Vockeroth is found in Costa Rica (Vockeroth, 1964; Thompson the immatures are unknown. The last key was that of Fluke
& Wynne, in press). Larvae are saprophagous in exuding (1942) as Epistrophe and Allograpta.
plant sap.
Argentinomyia Lynch Arribálzaga. Found in the Neo-
Allograpta Osten Sacken. This genus is cosmopolitan, tropical Region only, except absent from the Chilean subregion,
with 104 species, 63 of which occur in the Neotropical this genus includes 32 species. There are four named and 15
SYRPHIDAE (flower flies) 53 787

undescribed species from Costa Rica. The last key was that including 299 in the Neotropical Region, is an important part
of Fluke (1945) as Rhysops and Melanostoma. Larvae are of the Central American fauna. There are 30 named and 51
unknown. undescribed species in Costa Rica. The last key was that of
Curran (1939a); see also Fluke (1951) and Curran (1953).
Aristosyrphus Curran. The six species of this genus are Larvae are saprophagous in a wide variety of decaying ma-
found from Mexico to Brazil, and two of them occur in Costa terials, including exuding tree sap, decaying cacti and aloes,
Rica. A revision is forthcoming (Thompson & Marioni, in decaying forest flowers, fruits, stems of non-woody plants
press). Larvae are unknown. including ferns and terrestrial bromeliads, and watertanks of
bromeliads and heliconias (Rotheray et al., 2007).
Cacoceria Hull. There are two species in this genus, found
from Mexico to Argentina. Only C. cressoni Hull is found in Criorhina Meigen. This is a north temperate genus whose
Costa Rica. No key is available, and larvae are unknown. distribution extends into the Oriental (India) and Neotropical
Regions (south to Costa Rica). There are 50 species, with
Callicera Panzer. This is a north temperate genus with two species in Mexico and three undescribed species in Costa
extensions into the Oriental (India, Taiwan) and Neotropical Rica. No key to species is available. Larvae are sapropha-
Regions (south to Costa Rica). Of the 16 species, only gous in wet decaying wood.
C. poultoni Verrall is found in Costa Rica and the Neotropical
Region. A revision of the New World species exists (Thomp- Dasysyrphus Enderlein. Another north temperate group
son, 1980). Larvae of Palearctic species are saprophagous in with extensions into the Oriental (Indonesia) and Neotro-
tree holes. pical Regions (south to Brazil), Dasysyrphus includes two
or three species in the Neotropical Region. Only one unde-
Cepa Thompson & Vockeroth. Three species of this ge- scribed species occurs in Costa Rica. Keys to species are in
nus are known: two from Brazil and one from Costa Rica. The Fluke (1942) and Vockeroth (1986b). Larvae are predatory
last key was that of Thompson (2007). Larvae are unknown. on Sternorrhyncha.

Ceriana Rondani. This is a north temperate genus whose Dideomima Vockeroth. There are four species in this
distribution extends into the Neotropical Region, south to genus, found from the southwestern USA (Arizona) to
Costa Rica. There are 66 described species, with one named Costa Rica. Only D.  vockerothi Thompson is known from
and one undescribed species in Costa Rica and the Neotropi- Costa Rica. A revision by Thompson will be available soon
cal Region. The last key was that of Curran (1941) as Ceri- (Thompson, in press b). Larvae are unknown, but probably
oides. Larval stages are saprophagous, mostly in decaying are predatory.
plant sap.
Eristalis Latreille. Eristalis is organized into two sub-
Chalcosyrphus Curran. This is a cosmopolitan group, genera. Subgenus Eristalis is restricted to the Palearctic,
with its greatest richness in the north temperate region. It is Afrotropical, and Oriental Regions (two species), but
divided into 10 subgenera, of which subgenus Neplas Porter E. tenax Linnaeus has been introduced into the cooler areas
is restricted to the New World (Arizona southward to Bra- of the Neotropical and Australian Regions. The subgenus
zil, absent from the Chilean subregion). Of the 29 species in Eoseristalis Kanervo is northern temperate in distribution,
subgenus Neplas, there are four named and five undescribed with extensions into the Oriental (Indonesia) and Neotropical
species in Costa Rica. The last key was that of Curran (1941) Regions (along cordillera). There are 96 species of Eoseri-
as Planes. Larvae are saprophagous under tree bark. stalis, of which nine are found in the Neotropical Region, but
only E. alleni Thompson and E. gatesi Thompson occur in
Cheilosia Meigen. This is a north temperate genus whose Costa Rica. A revision was published by Thompson (1997a).
distribution extends into the Oriental (Indonesia) and Neo- Larvae are saprophagous in decaying plant material.
tropical Regions. There are 439 species, with one species in
Chiapas, Mexico, and another in Guatemala. The last key Eupeodes Osten Sacken. Cosmopolitan except absent
was that of Hull & Fluke (1950). Larvae are phytophagous. from New Zealand, the main diversity of this genus is in
north temperate areas. There are 90 species, with two in the
Chrysotoxum Meigen. Chrysotoxum is another north Neotropical Region, and only E. americanus Wiedemann in
temperate group with extensions into the Oriental (India, Costa Rica. Larvae are predatory on Sternorrhyncha.
Indonesia), Afrotropical (one species), and Neotropical Re-
gions (to Guatemala). There are 97 species, including one Habromyia Williston. The four species of this genus are
undetermined species in Guatemala. Shannon (1926) was the found from Costa Rica to Brazil. Only H.  coeruleithorax
last to key the species. Larvae are predatory on root ��������
Sternor- Williston is present in Costa Rica (Thompson, in press a).
rhyncha in ant nests. Larvae are saprophagous in wet decaying wood.

Copestylum Macquart. Found from Canada south to Lejops Rondani. Within Lejops, subgenus Asemosyrphus
Chile and Argentina, this large genus, with 309 species, Bigot is found from Canada to Guatemala (also Colombia),
788
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

with three described species. The last key was that of Curran typic group (Microdon), which is cosmopolitan, and Ome-
(1939b). Larvae are saprophagous in wetlands. gasyrphus Loew, which also occurs in the Nearctic Region.
Overall, there are 352 species worldwide, including 141 in
Lepidomyia Loew. This genus is distributed from the the Neotropical Region and 23 named and 31 unnamed spe-
southern USA (Texas) to Argentina, although it is absent cies in Costa Rica. The latest key is that of Curran (1941).
from the Chilean subregion. Sixteen species are present, Larvae are found in ant colonies.
including 15 in the Neotropical Region. Two named and five
undescribed species occur in Costa Rica. The last key was Milesia Latreille. Found in all regions except the Austra-
that of Hull (1946b). Larvae are saprophagous in exuding lian, this genus is distributed in the Neotropical Region south
tree sap and in tree holes. only to Panama. There are 82 species, with only M.  nigra
Fluke and M. pulchra Williston in Costa Rica and Panama.
Leucopodella Hull. This genus is found in the Neotro- The last monograph was that of Hippa (1990). Larvae are
pical Region only, except not from the Chilean subregion; saprophagous in tree holes.
of the 14 species, six are named and two undescribed spe-
cies are from Costa Rica. The latest key to species is that Mixogaster Macquart. Found in the New World only,
of Thompson (1981). Larvae are unknown, but are probably this genus is distributed from the USA (Massachusetts)
predatory on Sternorrhyncha. south to Argentina, but is absent from the Chilean subregion.
There are 17 species, with only M.  mexicana Macquart
Lycastirhyncha Bigot. Five described species are found and M. orpheus Hull occurring in Costa Rica. A revision was
from Mexico to Brazil, but only L.  willistoni Coquillett is given by Hull (1954). Larvae live in ant nests.
found in Costa Rica. In the last key, Doesburg (1966) rec-
ognized five species, but we suspect there is only one wide- Monoceromyia Shannon. There are 45 species in this
spread species, L. nitens Bigot. Larvae are unknown. genus, of which nine species are found in the Neotropical
Region. Two named and two undescribed species are found
Lycopale Hull. One species, L. wygodzinskyi Thompson, in Costa Rica. The last key was that of Curran (1941), as
occurs in Costa Rica and has been reared from a tree hole. Cerioides. Larvae live in decaying plant sap.
Otherwise, there are nine species, found from Mexico to Bra-
zil. A revision has been prepared (Thompson, in press a). Mutillimyia Hull. The single species of this genus,
M.  auricaudata Williston, is known only from type series
Mallota Meigen. The distribution of this genus is north collected in Neotropical Mexico (Guerrero, Morelos). Larvae
temperate, with extensions into the Oriental (Indonesia) and are unknown.
Neotropical Regions (south to Brazil). There are 64 described
species, with 12 in the Neotropical Region and six in Costa Myolepta Newman. Found in all regions except the Aus-
Rica. A revision of Central American species was published tralasian, there are 37 species of this genus, including 11 in
by Thompson & Zumbado (2002). Larvae live in tree holes the Neotropical Region. One named and four undescribed
and decaying trunks of tree ferns. species occur in Costa Rica. The genus is usually split into
subgenera (Myolepta, Eumyiolepta Shannon, and Protolepi-
Melanostoma Schiner. Found in all biogeographical re- dostola Hull), but the status of these divisions, especially
gions, this genus is only found in the Neotropical Region in with respect to Lepidomyia, remains unclear. The last key to
Chiapas, but may also occur in the adjacent highlands of Gua- species is that of Fluke & Weems (1956); also see Thompson
temala. There are 57 species, with only M. bellum Giglio-Tos (1968). Larvae of known species (mainly Palearctic) are sap-
in Mexico. Larvae are predatory on Sternorrhyncha. rophagous in tree holes.

Meromacrus Rondani. Found from the southern USA to Nausigaster Williston. This genus is found from the
Argentina and northern Chile, the 43 species of this genus southern USA to Brazil and is absent from the Chilean sub-
include 40 in the Neotropical Region and seven from Costa region. There are 15 species, with nine in the Neotropical
Rica. The last key to the entire genus was that of Hull (1942), Region, and only N. meridionalis Townsend in Costa Rica.
whereas a revision of the Central American species was pre- The latest key to species is Curran (1941). Larvae are sap-
sented by Blatch et al. (2003). Larvae are saprophagous in rophagous in wet decaying cacti, bromeliads, and probably
tree holes and decaying plant stems. other plants in arid or open habitats.

Microdon Meigen. This genus is cosmopolitan, except Ocyptamus Macquart. This is a New World genus, found
absent in New Zealand and the smaller oceanic islands. There from southern Canada southward. There are 282 species,
are many distinctive groups within the clade here treated as including 273 in the Neotropical Region, and 26 named and
the genus Microdon. Unfortunately, their status and rank are 64 undescribed species in Costa Rica. The latest key is that
not well understood (Cheng & Thompson, 2008), but they of Hull (1949), but it includes less than half the described
are included here as the adults are easily recognized. All the species. There is a revision of the subgenus Mimocalla Hull
groups are restricted to the Neotropical Region except the (Thompson & Zumbado, 2000). Larvae are predators of
SYRPHIDAE (flower flies) 53 789

gregarious soft-bodied insects (such as aphids and psyllids) Zealand. A number of subgenera are recognized, but only
on plants, but some species have been reared as predators in the subgenus Carposcalis Enderlein is found in the Neotro-
epiphytic bromeliads (Rotheray et al., 2000). pical Region, where it ranges along the Andean Cordillera
and is widespread in the south temperate areas. One hundred
Ornidia Lepeletier & Serville. This is a New World and sixty-six species are known world-wide, with 13 in the
endemic genus, found from the USA south to Chile and Neotropical Region and a couple of species in Costa Rica.
Argentina, with one species introduced into the Old World The last key was that of Fluke (1945). Larvae are predatory
tropics. Four species are found in the Neotropical Region on Sternorrhyncha.
and Costa Rica (Thompson, 1990). Larvae are saprophagous
in a wide variety of decaying plant materials from exuding Polybiomyia Shannon. This genus is found from the
tree sap to decaying flowers, fruits, and stems of non-woody southern USA to Argentina, but is absent from the Chilean
plants. They are also associated with animals, apparently subregion. Nineteen species are described, including 14
breeding in wounds, dung, and other waste materials including Neotropical species. There are four named species and five
plant and animal compost (Rotheray et al., 2005). undescribed species in Costa Rica. The last key was that of
Curran (1941), as Cerioides. Larvae are predatory, living in
Orthonevra Macquart. This is a predominantly north
decaying plant sap.
temperate genus, with extensions into the Oriental and Neo-
tropical Regions (south to Brazil). There are 59 species, Pseudodoros Becker. This genus is found in Africa and
eight in the Neotropical Region, including two named and New World, from the USA south. There are three species,
one undescribed species in Costa Rica. Larvae of Palearctic
including two in the Neotropical Region and only P. clavatus
species are saprophagous in accumulations of wet, decaying
Fabricius in Costa Rica. The latest key was that of Kassebeer
vegetation in slow-moving water bodies such as pools and
(2000). Larvae are predatory on Sternorrhyncha.
lakes. No keys are available.

Palpada Macquart. Palpada species are found from the Quichuana Knab. Found from Mexico to Brazil and
southern USA, south to Chile and Argentina. There are 85 Argentina, but absent from the Chilean subregion, there are
species, 82 in the Neotropical Region, 21 named and three 23 species of this genus. Seven named and three undescribed
undescribed species in Costa Rica. The last key was that of species are known from Costa Rica. The latest key is that
Curran (1934). Larvae are saprophagous in decaying plant of Hull (1946a). Larvae are saprophagous in pockets of wet
material, including dung. decay on woody and non-woody plants.

Paragodon Thompson. In Costa Rica, this genus is Rhingia Scopoli. All regions except the Australian have
represented only by P.  paragoides Thompson. Otherwise, species of Rhingia. Of the 37 species, two are found in the
there are four species, found from Mexico to Brazil. A revi- Neotropical Region, but only R.  nigra Macquart in Costa
sion of the genus is given by Thompson (1969). Larvae are Rica. Fluke (1943) provided diagnostic notes to separate the
unknown. two Neotropical species. Larvae of Palearctic species are
saprophagous in animal dung.
Paragus Latreille. This genus is cosmopolitan, except
that it is absent from New Zealand and the Neotropical Region Rhopalosyrphus Giglio-Tos. This genus is distributed
south of Costa Rica. There are 90 species organized into from the southern USA (Arizona, Texas, and Florida) to Ar-
three major clades: subgenus Pandasyophthalmus Stucken- gentina. There are four species, two of which are found in
berg with only P. haemorrhous Meigen in Costa Rica; sub- Costa Rica. The latest revision is by Weems et al. (2003).
genus Paragus restricted to the north temperate region; and Larvae live in arboreal ant nests.
the P. serratus complex, restricted to the Old World tropics.
Some authors want to recognize a number of new genera and Salpingogaster Schiner. The 35 species of this genus are
species, but according to Vockeroth (1986a), there is only found in the southern USA (Florida, Texas), south to Chile
one variable species in this group in the New World, and only and Argentina. There are five named and 13 undescribed spe-
one genus is necessary for all the species. Larvae are preda- cies from Costa Rica. Two subgenera are recognized, with
tory on Sternorrhyncha. most species in the subgenus Salpingogaster, and only one
(S.  cochenillivorus Guérin-Méneville) in subgenus Eosal-
Paramicrodon de Meijere. One undescribed species of
pingogaster Hull in Central America. The latest key is that
this genus is found in Costa Rica. Otherwise, there are 10
species found in the Australian, Oriental, and Neotropical of Curran (1941). Salpingogaster nigra Schiner is a predator
Regions. Larvae live in ant nests. of spittlebugs on sugar cane. The species of the subge-
nus Eosalpingogaster are all predators of scales (Hemiptera:
Platycheirus Lepeletier & Serville. This is largely a north Coccoidea), and the Central American species S. cochenil-
temperate group with extensions into Neotropical Region, livorus is a predator of the famous red (cochineal) scale,
Oriental Region (Philippines and Taiwan only), and New Dactylopius coccus Costa (Dactylopiidae).
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Sericomyia Meigen. This genus is found in north tem- S. stimulans Thompson and an undescribed species in Costa
perate areas with extensions into the Oriental (Taiwan) and Rica (last key: Thompson, 1973). Larvae are unknown.
Neotropical Regions (south to Costa Rica). There are a total
of 30 species, with only S. fairmanorum Fairman in Costa Syrphus Fabricius. Largely north temperate with exten-
Rica. A second species, S. meyersi Fluke, is found in central sions into the Oriental and Neotropical Regions (Brazil,
Mexico. The latest key is that of Thompson et al. (2000). The Argentina, and Chile), this genus has 70 species, including
genus is usually organized into subgenera, with the two Me- 17 in the Neotropical Region and three in Costa Rica. A key
soamerican species placed in subgenus Arctophila Schiner. to the Neotropical species is in Thompson et al. (2000).
Larvae of Palearctic species are saprophagous in pools and Larvae are predatory on Sternorrhyncha.
wet boggy ground, including seepages containing decaying
vegetation. Toxomerus Macquart. Found in the New World (south-
ern Canada and southward), Toxomerus has 141 species,
Sphaerophoria Meigen. This is a predominantly north including 132 in the Neotropical Region. There are 22 named
temperate genus with limited extensions into the south tem- species and nine undescribed species in Costa Rica. The last
perate areas (South Africa and Australia). In the New World, key to species is that of Hull (1943b), but Metz & Thompson
one species, S. contigua Macquart, extends south to Chiapas, (2001) revised the larger species. Some larvae are predatory
Mexico. The New World species were revised by Knutson on Sternorrhyncha, but some feed on pollen. The most
(1973). famous pollen-feeder is T. politus (Say), which feeds on corn
(Zea mays) pollen and was known to the Aztecs as an indica-
Sphiximorpha Rondani. Cosmopolitan, except absent tor of good corn harvests.
from New Zealand, this genus has 70 described species.
There are 23 species in Neotropical Region, with two named Trichopsomyia Williston. This is a north temperate genus
species and five undescribed species in Costa Rica. The latest whose distribution extends into Australia and the Neotropi-
key is that of Curran (1941), as Cerioides. Larvae are sap- cal Region (south to Brazil). There are 28 species, including
rophagous in wet decaying wood and decaying plant sap. 12 in the Neotropical Region. In Costa Rica, there are one
named and five undescribed species. The latest key is that
Spilomyia Meigen. This is a predominantly north temper- of Fluke (1937). Larvae are predatory on psyllids and other
ate genus with extension into the Neotropical Region, southward Sternorrhyncha.
to Argentina. There are 36 species, with eight in the Neotro-
pical Region, and three named and two undescribed species Xanthandrus Verrall. This genus is cosmopolitan,
in Costa Rica. The last key is that of Thompson (1997b). except absent from New Zealand. There are 26 species, in-
Larvae are saprophagous in tree holes. cluding 14 in the Neotropical Region. Two species are pres-
ent in Costa Rica. The last keys are those of Fluke (1937)
Sterphus Philippi. The 41 species of Sterphus are distrib- and Borges & Pamplona (2003). Some species are predators
uted from Mexico southward to Chile and Argentina. There of caterpillars.
are four subgenera, three of which occur in Costa Rica: sub-
genus Ceriogaster Shannon, with 25 species and six unnamed Xylota Meigen. The 137 Xylota species are found in all
species in Costa Rica (last key: Hull, 1943a); subgenus regions except the Australian. Seven are found in the Neo-
Crepidomyia Shannon, with 30 species and six unnamed tropical Region, while eight undescribed species are in Costa
species in Costa Rica (key to Costa Rican species: Zum- Rica. A revision has been prepared (Thompson et al., in
bado & Thompson, 1997; also Hippa & Thompson, 1994); press). Larvae are saprophagous in wet decaying tree sap
and subgenus Sterphus Philippi, with three species, but only and wood.

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Syrphidae) including the description of a new species. Studia of two new genera from Africa and China. Zootaxa 1879:
dipterologica 10: 13–36. 21–48.
Borges, Z.M. & D.M. Pamplona. 2003. Revision of the Neotropical Cumming, J.M., B.J. Sinclair, & D.M.  Wood. 1995. Homology
Xanthandrus Verrall (Diptera, Syrphidae). Revista Brasileira de and phylogenetic implications of male genitalia in Diptera —
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Brauer, F. 1883. Die Zweiflügler des Kaiserlichen Museums zu Curran, C.H. 1934. Diptera of Kartabo, Bartica District, British
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Curran, C.H. 1939b. Two new American Diptera with notes on Knutson, L.V. 1973. Taxonomic revision of the aphid-killing flies
Asemosyrphus Bigot. American Museum Novitates 1031: of the genus Sphaerophoria in the Western Hemisphere (Syr-
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Fluke, C.L. 1942. Revision of the Neotropical Syrphini related mitochondrial COI and nuclear 28S rRNA genes: Conflict and
to Syrphus (Diptera, Syrphidae). American Museum Novi- congruence with the current tribal classification. Cladistics 24:
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Fluke, C.L. 1943. A new genus and new species of Syrphidae Mengual, X., C. Ruiz, S. Rojo, G. Ståhls, & F.C. Thompson. 2009.
(Diptera) from Ecuador. Annals of the Entomological Society A conspectus of the flower fly genus Allograpta (Diptera: Syr-
of America 36: 425–431. phidae) with description of a new subgenus and species.
Fluke, C.L. 1945. The Melanostomini of the Neotropical Region Zootaxa 2214: 1–28.
(Diptera, Syrphidae). American Museum Novitates 1272: 1–29. Metz, M.A. & F.C. Thompson 2001. A revison of the larger spe-
Fluke, C.L. 1951. Syrphid flies related to Volucella scutellata Mac- cies of Toxomerus (Diptera: Syrphidae) with the description of
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Fluke, C.L. & H.V. Weems, Jr. 1956. The Myoleptini of the Ameri- Nishida, K., G.E. Rotheray, & F.C. Thompson. 2003. First non-
cas (Diptera, Syrphidae). American Museum Novitates 1758: predaceous syrphine flower fly (Diptera: Syrphidae): a new
1–23. leaf-mining Allograpta from Costa Rica. Studia dipterologica
Griffiths, G.C.D. 1972. The phylogenetic classification of Diptera 9: 421–436.
Cyclorrhapha with special reference to the structure of the male Rotheray, G. 1993. Colour guide to hoverfly larvae (Diptera, Syr-
postabdomen. Dr. W. Junk N. V., The Hague, 340 pp. phidae) in Britain and Europe. Dipterists Digest 9: 1–155.
Hennig. W. 1948. Die Larvenformen der Dipteren. 1. Teil. Rotheray, G. & F. Gilbert. 1999. Phylogeny of Palaearctic Syrphi-
Akademie–Verlag, Berlin, 185 pp. dae (Diptera): Evidence from larval stages. Zoological Journal
Hippa, H. 1990. The genus Milesia Latreille (Diptera, Syrphidae). of the Linnean Society 127: 1–112.
Acta Zoologica Fennica 187: 1–226. Rotheray, G. & F. Gilbert. 2008. Phylogenetic relationships and
Hippa, H. & F.C. Thompson. 1994. Revision of the Sterphus cybele the larval head of the lower Cyclorrhapha (Diptera). Zoologi-
species group (Diptera: Syrphidae). Proceedings of the Entomo- cal Journal of the Linnean Society 153: 287–323.
logical Society of Washington 96: 483–495. Rotheray, G.E., M.A. Zumbado, E.G.  Hancock,  & F.C. Thomp-
Hull, F.M. 1942. The flies of the genus Meromacrus. American Mu- son. 2000. Remarkable aquatic predators in the genus Ocypta-
seum Novitates 1200: 1–19. mus (Diptera, Syrphidae). Studia dipterologica 7: 385–398.
Hull, F.M. 1943a. The genus Ceriogaster Williston (Syrphidae). Rotheray, G.E., E.G. Hancock, M.A. Marcos-García, & M. Zum-
Revista de la Sociedad Entomologica Argentina 12: 138–140. bado. 2005. Early stages and breeding sites of three species
Hull, F.M. 1943b. The genus Mesogramma. Entomologica Ameri- of Neotropical Ornidia (Diptera, Syrphidae). Studia diptero-
cana 23: 1–41. logica 12: 419–427.
Hull, F.M. 1946a. The genus Quichuana Knab. American Museum Rotheray, G.E., E.G. Hancock,  & M.A.  Marcos-García. 2007.
Novitates 1317: 1–17. Neotropical Copestylum (Diptera, Syrphidae) breeding in bro-
Hull, F.M. 1946b. The genus Lepidostola Mik. American Museum meliads (Bromeliaceae) including 22 new species. Zoological
Novitates 1326: 1–15. Journal of the Linnean Society 150: 267–317.
Hull, F.M. 1949. The genus Baccha from the New World. Entomo- Shannon, R.C. 1926. The chrysotoxine syrphid-flies. Proceedings
logica Americana 27: 89–291. of the United States National Museum 70: 1–33.
Hull, F.M. 1954. The genus Mixogaster Macquart (Diptera: Syrphi- Sommaggio, D. 1999. Syrphidae: can they be used as biologi-
dae). American Museum Novitates 1652: 1–32. cal indicators? Agriculture, ecosystems and environment 74:
Hull, F.M. & C.L. Fluke. 1950. The genus Cheilosia Meigen (Dip- 343–356.
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letin of the American Museum of Natural History 94: 299–402. Caring for pollinators: Safeguarding agro-biodiversity and
Kassebeer, C.K. 2000. Zur Gattung Pseudodoros Becker, 1903 (Di- wild plant diversity. Federal Agency for Nature Conservation
ptera, Syrphidae). Dipteron 3: 73–92. (BfN). BfN-Skripten 250, Bonn-Bad Godesberg, Germany,
Katzourakis, A., A. Purvis, S. Azmeh, G. Rotheray, & F. Gilbert. [7] + 191 pp.
2001. Macroevolution of hoverflies (Diptera: Syrphidae): The Ståhls, G., H. Hippa, G.E. Rotheray, J. Muona, & F. Gilbert. 2003.
effect of using higher-level taxa in studies of biodiversity, and Phylogeny of Syrphidae (Diptera) inferred from combined
correlates of species richness. Journal of Evolutionary Biology analysis of molecular and morphological characters. System-
14: 219–227. atic Entomology 28: 433–450.
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Tenhumberg, B. & H. Poehling. 1995. Syrphids as natural enemies Thompson, F.C. & L. Marinoni. in press. Flower flies of the genus
of cereal aphids in Germany: Aspects of their biology and effi- Aristosyrphus Curran (Diptera: Syrphidae). Studia dipterologi-
cacy in different years and regions. Agriculture, Ecosystems and ca Supplement.
Environment 52: 39–43. Thompson, F.C. & I.R. Wynne. in press. Flower flies of the genus
Thompson, F.C. 1968. The placement of the subgenus Protolepi- Alipumilio Shannon (Diptera: Syrphidae). Studia dipterologica
dostola Hull (Diptera: Syrphidae) with description of two Supplement.
new species. Journal of the Kansas Entomological Society 41: Thompson, F.C. & M. Zumbado. 2000. Flower flies of the subgenus
270–277. Ocyptamus (Mimocalla Hull) (Diptera: Syrphidae). Proceedings
Thompson, F.C. 1969. A new genus of microdontine flies (Dip- of the Entomological Society of Washington 102: 773–793.
tera: Syrphidae) with notes on the placement of the subfamily. Thompson, F.C. & M. Zumbado. 2002. Mesoamerican Mallota
Psyche 76: 74–85.
flower flies (Diptera: Syrphidae) with the description of four
Thompson, F.C. 1972. A contribution to a generic revision of the
new species. Studia dipterologica 9: 89–107.
Neotropical Milesinae (Diptera: Syrphidae). Arquivos de Zoo-
Thompson, F.C., J.R. Vockeroth, & Y.S. Sedman. 1976. 46. Family
logia 23: 73–215.
Thompson, F.C. 1973. Review of the genus Sterphus Philippi (Dip- Syrphidae. In Papavero, N. (editor). A Catalogue of the Diptera
tera: Syrphidae). Part I. Entomologica Americana 46: 185–240. of the Americas South of the United States. Departamento de
Thompson, F.C. 1980. The North American species of Callicera Zoologia, Secretaria da Agricultura, São Paulo, 195 pp.
Panzer (Diptera: Syrphidae). Proceedings of the Entomological Thompson, F.C., B.J. Thompson,  & J.E.  Fairman. 2000. Only in
Society of Washington 82: 195–211. Costa Rica: new Neotropical flower flies (Diptera: Syrphidae).
Thompson, F.C. 1981. The flower flies of the West Indies (Diptera: Studia dipterologica 7: 33–43.
Syrphidae). Memoirs of the Entomological Society of Washing- Thompson, F.C., M. Zumbado, & H. Hippa. in press. Neotropical
ton 9: 1–200. Xylota flower flies (Diptera: Syrphidae). Studia dipterologica
Thompson, F.C. 1990. The flower fly genus Ornidia (Diptera: Syr- Supplement.
phidae). Proceedings of the Entomological Society of Washing- Vockeroth, J.R. 1964. The genus Alipumilio Shannon (Diptera: Syr-
ton 93: 248–261. phidae). The Canadian Entomologist 96: 922–924.
Thompson, F.C. 1997a. Revision of the Eristalis flower flies of Vockeroth, J.R. 1969. A revision of the genera of the Syrphini
the Americas south of the United States (Diptera: Syrphidae). (Diptera: Syrphidae). Memoirs of the Entomological Society of
Proceedings of the Entomological Society of Washington 99: Canada 62: 1–176.
209–237. Vockeroth, J.R. 1986a. Revision of the New World species of Para-
Thompson, F.C. 1997b. Spilomyia flower flies of the New World gus Latreille (Diptera: Syrphidae). The Canadian Entomologist
(Diptera: Syrphidae). Memoirs of the Entomological Society of 118: 183–198.
Washington 18: 261–272. Vockeroth, J.R. 1986b. Nomenclatural notes on Nearctic Eupeodes
Thompson, F.C. 1999. A key to the genera of the flower flies of the
(including Metasyrphus) and Dasysyrphus (Diptera: Syrphidae).
Neotropical Region with the description of two new genera and
The Canadian Entomologist 118: 199–204.
eight new species. Contributions on Entomology, International
Weems, H.V., Jr., F.C. Thompson, G.E.  Rotheray,  & M.  Deyrup.
3: 319–378.
Thompson, F.C. 2007. A new Costa Rican flower fly (Diptera: Syr- 2003. The genus Rhopalosyrphus (Diptera: Syrphidae). Florida
phidae) and a replacement name for a Neotropical flower fly Entomologist 86: 186–193.
genus. Studia dipterologica 14: 167–170. Wheeler, W.M. 1924. Two extraordinary larval myrmecophiles from
Thompson, F.C. in press a. The genera Habromyia and Lycopale Panama. Proceedings of the National Academy of Sciences 10:
and related Neotropical species (Diptera: Syrphidae). Studia 237–244.
dipterologica Supplement. Zumbado, M. & F.C. Thompson. 1997. Nuevas especies de Sterphus
Thompson, F.C. in press b. Flower flies of the genus Dideomima (Diptera: Syrphidae) de Costa Rica con notas sobre otras especies
(Diptera: Syrphidae). Studia dipterologica Supplement. presentes en Costa Rica. Southwestern Entomologist 22: 79–90.
PIPUNCULIDAE (BIG-HEADED FLIES) 54
José Albertino Rafael and Jeffrey H. Skevington

Fig. 54.1. Male of Pipunculus luteicornis Cresson, (Nearctic, MND, fig. 53.1).

Diagnosis Abdomen subcylindrical, coloration and tomentum similar

to thorax, occasionally with yellow markings. Tergite 1 usu-
Adults (Fig.  1) small (body length 2.0–11.5  mm), dark. ally with row or cluster of bristles laterally. Male terminalia
Head semiglobose with narrow occiput in Chalarinae or globose twisted left (Fig. 17). Male syntergosternite 8 large, forming
with broadened occiput in Nephrocerinae and Pipunculinae. tip of abdomen (Fig.  17). Female terminalia modified into
Compound eyes occupying most of head (Figs. 1–4). Female piercing ovipositor (Fig. 16).
with enlarged anterior ommatidia (not distinct in Nephrocerus). Pipunculids can be differentiated from syrphids by the
Antenna with first flagellomere appearing as single segment, large compound eyes that occupy most of their globose or
obtuse to filiform in shape, with dorsal arista. Thorax with semiglobose head, lack of spurious vein, and open r4+5 cell.
darkish ground color. Proepisternum sometimes bearing fan- A few bombyliids such as Metacosmus Coquillett resemble
like tuft of bristles (Fig.  3). Claws and pulvilli larger in pipunculids when encountered in the field, but these round-
females. Wings long and narrow, usually hyaline, iridescent headed bombyliids can be easily differentiated by their
in direct light, pterostigma present or absent, cell r4+5 open. antennae that lack the dorsal arista found in pipunculids.

793
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Biology Region (Skevington  & Marshall, 1997) and the Australian

Region (Skevington, 2001). Most New World pipunculid
Most pipunculids that have been studied are endoparasitoids species have no biological data available. In the Neotropical
of several Hemiptera (Auchenorrhyncha) families, including Region and Mexico, the following papers treat pipunculid–
Cicadellidae, Cercopidae, Delphacidae, Membracidae, Issidae, host associations: Hardy (1963) for southern Colombia and
Cixiidae, and Flatidae. The only known exception occurs in northern Ecuador, Vega et al. (1991) for Mexico, Virla & Ra-
the genus Nephrocerus. These unusual flies attack adult crane fael (1996) for Argentina, and recently Virla et al. (2009) for
flies (Tipulidae) (Koenig  & Young, 2007). The potential val- all Neotropical species.
ue of Pipunculidae for biological control has stimulated some Most pipunculid species attack more than one species of
work on the bionomics of this family. For example, research host, but show a preference for a particular set of host spe-
into the control of the potato leafhopper, Empoasca fabae (Har- cies. Only two species have been reared from more than one
ris), a major pest of alfalfa in midwestern and eastern USA and subfamily of host (Skevington, 2001). Superparasitism and
Canada, involved exploration in Europe for natural enemies to multiple parasitism are rare, and survival of more than one
be introduced into the USA (Jervis, 1992) and included rear- pipunculid in these cases is exceptional (Jervis, 1980; Mora-
ing of Chalarus for release (Skevington  & Marshall, 1997). kote & Yano, 1988; Waloff & Jervis, 1987).
Despite the economic importance of the group, few data on
host relationships exist, and some genera have no known hosts Classification
(for example Allomethus, Basileunculus, Elmohardyia, and
Protonephrocerus). Pipunculidae are considered to be a monophyletic lineage
Adults are frequently seen hovering among vegetation, and and sister group of the flower flies (Syrphidae). Together,
their flight is similar to that of Syrphidae. Adults utilize all ter- pipunculids and syrphids are grouped in the monophyletic
restrial habitats, but diversity and numbers are greatest in forest clade Syrphoidea, sister group of Schizophora (Cumming
openings and along forest edges. In semi-arid regions, adults are et al., 1995). Worldwide, 1401 valid species of pipunculids
commonly found at seeps or along small creeks. Adult flies feed are recognized, and it is likely that over 2700 species exist
on honeydew and can occasionally be found in large, mixed (De Meyer, 1996; De Meyer & Skevington, 2000; J.H. Skev-
feeding groups on Auchenorrhyncha secretions (Skevington & ington, unpublished data). Twenty-one extant genera are
Marshall, 1998). Because of this behavior, they are often at- currently recognized in three subfamilies: Chalarinae, Neph-
tracted to artificial honeydew sprayed on vegetation (a mixture rocerinae, and Pipunculinae (De Meyer, 1996; De Meyer &
of honey, Coca-Cola®, and water). Many pipunculids employ Skevington, 2000; Rafael & De Meyer, 1992; Skevington &
the landmark mating system known as hilltopping (Skevington, Yeates, 2001). The latter are divided into five tribes: Pipun-
2000, 2001). Hilltop collecting is an excellent way to sample culini, Cephalopsini, Microcephalopsini, Eudorylini, and
much of the pipunculid diversity in an area and often provides Tomosvaryellini. Molecular analyses made by Skevington &
large numbers of specimens. Other effective methods frequent- Yeates (2000) corroborated most of Rafael  & DeMeyer’s
ly employed to capture pipunculids include sweeping, Malaise (1992) phylogenetic results, except for the position of the
trapping, and less successfully, using yellow pan traps. Occa- Nephrocerinae genera (Nephrocerus and Protonephrocerus).
sionally, mating pairs are discovered hovering in copula or tem- The combined molecular data suggested that Nephrocerus
porarily settled on vegetation. Pronounced sexual dimorphism is a basal lineage of subfamily Chalarinae, and Protoneph-
in pipunculids makes association of the sexes difficult, so these rocerus is a basal lineage of subfamily Pipunculinae. When
occasional discoveries are useful. these data were combined with the morphological data set
Females of most pipunculid species deposit their eggs in ear- of Rafael & De Meyer (1992), Nephrocerus and Protoneph-
ly instar nymphs or adults of their hosts. After visually locating rocerus appeared between the basal Chalarinae and the ter-
a host, some species remove the host from the food plant and minal Pipunculinae, reflecting the hypothesis of Rafael  &
oviposit one egg inside each host while in flight (Huq, 1982; De Meyer with the difference that these two genera do not
May, 1979; Williams, 1919), whereas others leave the host form a monophyletic group. These results suggest that the
in situ (Benton, 1975). Larvae develop to maturity within the subfamilial taxonomy of the pipunculids may need modifica-
nymph or adult hosts. When mature, the larva leaves the host tion and further study. Generic concepts are relatively well
through a break in one of the dorsal intersegmental membranes resolved but more work is still needed to clarify Eudorylini
of the abdomen in order to pupate in the soil, in leaf litter, at genera. Despite recent work on Eudorylini, problems remain
the base of plants, or rarely, attached to leaves (Ferrar, 1987; within this large lineage (Skevington & Yeates, 2001).
Perkins, 1905). More detailed information on immature stages
together with information on rearing techniques can be found in
Benton (1975), Coe (1966), Ferrar (1987), Freytag (1988), Huq Identification
(1984), Jervis (1980), Morakote  & Yano (1987), and Perkins
(1906). Because of their small size, black coloration, and the pau-
Pipunculid–host associations are best known for the Euro- city of external characters, pipunculid species are difficult
pean fauna (Waloff & Jervis, 1987), followed by the Nearctic to identify. It is particularly important to have male and
PIPUNCULIDAE (big-headed flies) 54 795

female specimens from the same sample in order to associate In order to secure the ground color and tomentum of pipun-
the sexes. For secure identification, it is necessary to dissect culids, specimens should ultimately be dried and pinned. Pin-
the male terminalia. To accomplish this, remove the abdo- ning fresh material killed with cyanide or ethyl acetate is the
men and heat in 85% lactic acid for at least 30 minutes. Re- easiest method, and a number of techniques can be used. The
move the macerated abdomen and place it directly into clean best method for obtaining views of the insect’s genitalia in-
glycerin. Dissection involves separating syntergosternite  8 volves double mounting. Pinning through the right anepis-
from the remainder of the abdomen. Store the abdomen and ternum with a minuten pin, then pinning to pith is more
dissected terminalia in glycerin in plastic microvials on the expensive and time consuming than pointing, but produces
same pin as the source specimen. Surstyli shape, presence or excellent specimens that are less likely to require dissection
absence of a membranous area on syntergosternite 8, phal- than other pinning techniques. Material collected into alcohol
lic guide shape, phallus shape and ramification, sperm pump is preferably dried with a critical point drier and then pointed
shape and sclerotization, ejaculatory apodeme shape, hypan- or glued directly to the side of a pin. Chemical drying using
drium shape, and modifications of the sternites are often in- HMDS (hexamethyldisilazane) or ethyl acetate is also accept-
formative. In female specimens, the ovipositor shape is often able. Although some researchers prefer to study pipunculids
useful, but few other diagnostic characters are available. preserved in 70% alcohol, specimens preserved in this way
Description of new species on the basis of females often com- discolor over time and become brittle and difficult to dissect.
plicates the taxonomy of the family and should be avoided Terminology used to describe pipunculid morphology
unless new character systems are discovered for females. We has varied considerably over time and between authors, but
encourage DNA barcoding (sequencing cytochrome oxidase recent efforts have been made to standardize this language.
I) of new species of pipunculids to faciliate association of Most of the current morphological terminology is illustrated
sexes and larvae and to stabilize nomenclature across wider here (Figs. 1–37), but see Skevington & Yeates (2001) for a
geographical areas. more thorough treatment of this topic.

Key to the genera of Pipunculidae of Central America and Nearctic Mexico

(modified from Albrecht, 1990; Rafael, 1996; Skevington & Yeates, 2001)

1. Hind margin of eye deeply excised medially (Fig.  4); first flagellomere reniform (Fig.  4);
NEPHROCERINAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nephrocerus Zetterstedt
– Hind margin of eye straight or nearly so (Figs. 2, 3, 8); first flagellomere acute (Fig. 5), filiform
(Fig. 6) to aristiform. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2. Ocellar bristles distinct; occiput narrow, scarcely projecting behind eyes (Fig. 8); head hemispherical
(Fig. 8); margin of scutum and scutellum with strong bristles (Fig. 8); CHALARINAE. . . . . . . 3
– Ocellar bristles reduced or absent (Figs. 2, 3, 7); occiput swollen and plainly visible in lateral
view (Figs. 2, 3, 7); head spherical (Figs. 2, 3, 7); margin of scutum and scutellum without strong
bristles (Fig. 7); PIPUNCULINAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

3. Wing venation incomplete, cross vein dm-cu absent, vein M reduced (Fig. 9). . . Chalarus Walker
– Wing venation complete, M and dm-cu present (as in Fig. 12). . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4. M2 present (Fig. 11). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Verrallia Mik

– M2 absent (as in Fig. 13) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Jassidophaga Aczél

5. Proepisternum with fanlike tuft of bristles (Fig. 3). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

– Proepisternum bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

6. At least anterior portion of scutum evenly setose (Fig. 1). . . . . . . . . . . . . . . . Pipunculus Latreille

– Scutum bare except for differentiated rows of dorsocentral bristles (Fig. 7). . . . . . . . . . . . . . . . . 7
796
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

ocp

prepst s

2 Collinias 3 Cephalosphaera
4 Nephrocerus

5 Elmohardyia
ocp

7 Dasydorylas

6 Eudorylas 8 Jassidophaga
C Sc R1 R2+3

M R4+5

A1 CuA2

CuA1

9 Chalarus 10 Collinias
ptstg

br
bm
cup dm
M1

A1+CuA2 dm-cu M2
11 Verrallia 12 Pipunculus

r-m

dm
dm-cu

13 Dorylomorpha 14 Tomosvaryella

15 Cephalops

Figs. 54.2–15. Heads, wings, and associated structures: lateral view of head of (2) Collinias sp., (Australasian, Skevington & Yeates, 2001,
fig. 2B); dorsolateral view of head and portion of thorax of (3)  Cephalosphaera parthenopipis Skevington, (Australasian, Skevington &
Yeates, 2001, fig. 2I); lateral view of head of (4) Nephrocerus slossonae Johnson, (Nearctic, MND, fig. 53.7); lateral view of antenna of
(5) Elmohardyia doelloi (Shannon), (South America, Rafael, 1987b, fig. 9); (6) Eudorylas mutillatus (Loew), (Australasian, Skevington &
Yeates, 2001, fig.  2A); dorsal view of head, thorax, and base of abdomen of (7)  Dasydorylas eucalypti (Perkins), (Australasian,
Skevington & Yeates, 2001, fig. 3A); same, with head rotated almost 90º to right, of (8) Jassidophaga sp., (Australasian, Skevington & Yeates,
2001, fig. 3C); dorsal view of wing of (9) Chalarus sp., (Australasian, Skevington & Yeates, 2001, fig. 4D); (10) Collinias sp., (Australasian,
Skevington & Yeates, 2001, fig. 4C); (11) Verrallia virginica Banks, (Nearctic, MND, fig. 53.3); (12) Pipunculus fuscus Loew, (Nearctic,
MND, fig. 53.6); (13) Dorylomorpha exilis (Malloch), (Nearctic, MND, fig. 53.4); (14) Tomosvaryella sylvatica (Meigen), (Holarctic, MND,
fig. 53.5); dorsal view of hind leg of (15) Cephalops cochleatus De Meyer, (Australasian, Skevington & Yeates, 2001, fig. 3F).
Abbreviations: ocp, occiput; prepst s, proepisternal setae; ptstg, pterostigma.
PIPUNCULIDAE (big-headed flies) 54 797

7. Vein M2 present (as in Fig. 11); Cephalosphaera Enderlein. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

– Vein M2 absent (as in Fig. 13) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

8. Hind tibia with comb of longer setae apically; male abdominal syntergosternite 8 with membranous
area reaching epandrium (Fig. 23). . . . . . . . . . . Cephalosphaera (Neocephalosphaera) De Meyer
– Hind tibia without comb of longer setae apically; male abdominal syntergosternite 8 with mem-
branous area not reaching epandrium (Fig. 22). . . . Cephalosphaera (Cephalosphaera) Enderlein

9. Face narrower than lower portion of frons; frons inflated in lateral view (as in Fig.  2); first
flagellomere small, slightly larger than pedicel (as in Fig. 2); cell dm not expanded medially (as
in Fig. 10). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microcephalops De Meyer
– Face equal or subequal in width to lower portion of frons; frons not inflated in lateral view;
first flagellomere distinctly larger than pedicel (as in Fig. 3); cell dm expanded medially (as in
Fig. 13); Cephalops Fallén. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

10. Tibiae lacking strong median and apical spinelike setae; abdomen broad and shortened; male
abdominal syntergosternite 8 with membranous area reaching epandrium (as in Fig. 23). . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cephalops (Semicephalops) De Meyer
– Tibiae with strong median and apical spinelike setae (Fig. 15); abdomen long and narrow; male
abdominal syntergosternite 8 with membranous area not reaching epandrium (as in Fig. 22). . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cephalops (Cephalops) Fallén

11. Pterostigma absent (Figs. 13, 14). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

– Pterostigma present (Figs. 9–12); Eudorylini (only males identifiable to genus). . . . . . . . . . . . 16

12. Crossvein r-m meeting cell dm near middle (Fig.  14); abdominal tergites usually with at least
some grey or brown tomentum laterally; abdomen shorter and not distinctly clavate, usually
widest at middle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tomosvaryella Aczél
– Crossvein r-m meeting cell dm near basal third (Fig. 13); abdominal tergites (except tergite 1)
shining, entirely without tomentum; abdomen elongate and clavate, widening distally; Dorylomorpha
Aczél . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

13. Males . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
– Females . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

14. Frons at narrowest point at least as wide as anterior ocellus; inner lobe of surstylus with ventral
projection (Figs. 36b, c); apices of surstyli usually dilated (normally visible without dissection)
(Fig. 36a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dorylomorpha (Dorylomyia) Albrecht
– Frons at narrowest point less than half as wide as anterior ocellus or eyes contiguous; inner lobe
of surstyli without ventral projection (Figs. 37b, c); apices of surstyli not dilated (Fig. 37a);
Neotropical (Colombia, Brazil) and Holarctic Regions (USA: south to Colorado and Utah in
Nearctic Region) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Dorylomorpha (Pipunculina) Albrecht]

15. Piercer without lateral flanges (Fig. 18) or if distinct flanges present then abdomen short and
robust in form. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dorylomorpha (Dorylomyia) Albrecht
– Piercer of ovipositor with distinct lateral flanges (Fig. 19); abdomen long and slender; Neotropical
(Colombia, Brazil) and Holarctic Regions (USA: south to Colorado and Utah in Nearctic
Region). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Dorylomorpha (Pipunculina) Albrecht]

16. Tergite 6 visible dorsally in males (Fig. 20) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

– Tergite 6 not visible dorsally in males (Fig. 21) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
798
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

tg 5

spr 6
st 4 tg 6
st 5
st 6 tg 7
ph
sur
spr 7 hypd
st 7
st 6 tg 7 cerc syntgst 8
st 7

st 8
tg 8 epand spm ej apod
pmp
ovp tg 9 17 Dorylomorpha
cerc

16 Pipunculus epand

mem

syntgst 8
tg 6

st 7
18 Dorylomorpha (Dorylomyia) 19 Dorylomorpha (Pipunculina)
20 Elmohardyia

epand

syntgst 8

tg 1 tg 2 tg 3 tg 4 tg 5 22 Cephalosphaera (Cephalosphaera)
21 Allomethus syntgst 8 epand
sur

mem

23 Cephalosphaera (Neocephalosphaera)

Figs. 54.16–23. Abdomens and terminalia: lateral view of female abdomen of (16) Pipunculus torus Skevington, (Nearctic, MND, fig. 53.9);
lateral of male abdomen of (17) Dorylomorpha canadensis Hardy, (Nearctic, MND, fig. 53.8); ventral view of ovipositor of (18) D. (Dorylomyia)
beckeri Aczél, (Holarctic, Albrecht, 1990, fig. 44); (19) D. (Pipunculina) borealis Wahlgren, (Holarctic, Albrecht, 1990, fig. 62); dorsal view
of male abdomen of (20) Elmohardyia atlantica (Hough), (Nearctic, Skevington & Yeates, 2001, fig. 4J); (21) Allomethus rotundicornis
(Hardy), (Neotropical, Skevington & Yeates, 2001, fig. 4M); ventral view of male terminalia of (22) Cephalosphaera (Cephalosphaera)
prionotaina Skevington, (Australasian, Skevington, 1999, fig.  5D); (23)  Ce. (Neocephalosphaera) petila Skevington, (Australasian,
Skevington, 1999, fig. 4e).
Abbreviations: cerc, cercus; ej apod, ejaculatory apodeme; epand, epandrium; hypd, hypandrium; mem, membranous area; ovp, ovipositor;
ph, phallus; spr, spiracle; spm pmp, sperm pump; st, sternite; sur, surstylus; syntgst, syntergosternite; tg, tergite.
PIPUNCULIDAE (big-headed flies) 54 799

st 7
ph

syntgst 8

hypd
ph gd a b

ph gd sur
24 Eudorylas 25 Eudorylas
sur

ph gd
26 Eudorylas
ej apod

hypd
27 Basileunculus
hypd

ej apod

ph

28 Basileunculus

spm pmp

29 Allomethus
30 Allomethus
31 Allomethus

ph gd sbepand scl
a

sur

b epand

32 Elmohardyia 33 Elmohardyia 34 Elmohardyia

35 Elmohardyia

Figs. 54.24–35. Male terminalia: dorsal view of male terminalia, phallus removed, of (24) Eudorylas moffattensis Skevington, (Australasian,
Skevington & Yeates, 2001, fig. 6I); lateral view of phallus and phallic guide of (25) Eu. schreiteri (Shannon), (South America, Rafael, 1987b,
fig. 6); lateral (a) and dorsolateral (b) views of ejaculatory apodeme and sperm pump of (26) Eu. mutillatus (Loew), (Australasian, Skevington &
Yeates, 2001, figs. 8E–F); dorsal view of surstyli, phallic guide, and hypandrium of (27) Basileunculus rex (Curran), (Skevington & Yeates,
2001, fig. 6E); dorsal view of phallus, sperm pump, and ejaculatory apodeme of (28) B. rex, (Skevington & Yeates, 2001, fig. 7J); lateral
view of male sperm pump and ejaculatory apodeme of (29) A. brimleyi Hardy, (Nearctic, Rafael, 1990b, fig. 14); dorsal view of male surstyli,
phallic guide, and hypandrium of (30) A. rotundicornis Hardy, (Neotropical, Skevington & Yeates, 2001, fig. 6F); dorsal view of phallus of
(31) A. brimleyi Hardy, (Nearctic, Skevington & Yeates, 2001, fig. 8C); lateral (a) and dorsal (b) views of male ejaculatory apodeme and
sperm pump (32) and dorsal view of male terminalia, phallus removed (33) of Elmohardyia atlantica (Hough), (Nearctic, Skevington &
Yeates, 2001, figs. 8P–Q and fig. 7C); right lateral view of phallus of (34) El. doelloi (Shannon), (South America, Rafael, 1987b, fig. 13);
right lateral view of phallus of (35) El. atlantica (Hough), (Nearctic, Skevington & Yeates, 2001, fig. 7I).
Abbreviations: ej apod, ejaculatory apodeme; epand, epandrium; hypd, hypandrium; ph, phallus; ph gd, phallic guide; sbepand scl, subepan-
drial sclerite; spm pmp, sperm pump; st, sternite; sur, surstylus; syntgst, syntergosternite.
800
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

17. Apex of first flagellomere obtuse below (Fig. 5); ventral spinelike setae (ctenidium) present on
midfemur only; at least tergite 5 with oblique gray tomentose spots posterolaterally; phallus
simple, apically swollen and membranous (Figs.  34, 35); sperm pump usually elongate with
square distal process (Fig. 32) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Elmohardyia Rafael
– Apex of first flagellomere acuminate (Fig.  6) to subfiliform or aristiform in some species;
ventral spinelike setae present on forefemur and midfemur; tergites without oblique gray tomentose
spots; phallus trifid (Fig. 25); sperm pump usually elongate with two distal processes (Fig. 26). .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eudorylas Aczél, in part

18. Abdomen with small, inconspicuous bristles (as in Fig. 20) . . . . . . . . . . Eudorylas Aczél, in part
– Abdomen with conspicuous bristles, notably laterally (Fig. 21). . . . . . . . . . . . . . . . . . . . . . . . . 19

19. Distal section of vein M1 relatively straight (as in Fig. 12); phallus trifid, with hooks at apices and
carpet of setae laterally (Fig. 31); ejaculatory apodeme linear (Fig. 29). . . . . . . Allomethus Hardy
– Distal section of vein M1 curved (as in Fig. 13); phallus simple and heavily sclerotized, swollen
distally (Fig. 28); ejaculatory apodeme shaped like ball on stem (Fig. 28). . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Basileunculus Rafael

b b

c c

36 Dorylomorpha (Dorylomyia) 37 Dorylomorpha (Pipunculina)

Figs. 54.36–37. Male surstyli: (a) ventral view of both surstyli, (b) left lateral view of left surstylus, and (c) right lateral view of right surstylus
of (36)  Dorylomorpha (Dorylomyia) beckeri Aczél, (Holarctic, Albrecht, 1990, fig.  47) and of (37)  D. (Pipunculina) borealis Walhgren,
(Holarctic, Albrecht, 1990, fig. 64).

Synopsis of the fauna studied, and only 14 genera and approximately 90 species are
known. Pipunculids are so poorly sampled that many new
The following synopses are based on De Meyer (1996), species can be expected, and it is likely that there are well
Rafael & Menezes (1999), De Meyer & Skevington (2000), over 200 species. As stated above, pipunculids are found in
and unpublished material from the Canadian National almost all terrestrial habitats, including canopy, lowland rain
Collection of Insects (CNC), the National Museum of forest, dry forests, open areas, riparian regions, and woodland
Natural History (USNM), and the Instituto Nacional de edges, being more diverse in wet environments and along
Biodiversidad (INBio). ecotones.
Of the 1401 valid pipunculid species, 270 occur in the
Neotropical Region (De Meyer, 1996; De Meyer & Skeving- Allomethus Hardy. This genus contains four described
ton, 2000). The Central American Pipunculidae are poorly species. A species formerly known only from Brazil
PIPUNCULIDAE (big-headed flies) 54 801

(A. catharinensis Rafael) and three undescribed species have the relevant species can be formally transferred into these
recently been collected in Costa Rica (based on material from genera. Of nine species of Eudorylas recorded from Costa
INBio). Rica, six were described by Rafael & Menezes (1999) (one
as Metadorylas). Mexican and Central American species
Basileunculus Rafael. All three recognized species in this were revised by Rafael (1993) but not keyed. The Neotropi-
Neotropical genus (Rafael, 1987a) have been recorded in cal species of Eudorylas (=Metadorylas sensu Rafael) were
Central America and the West Indies. revised and keyed by Rafael (1990a).
Cephalops Fallén. Eight species have been identified (six Jassidophaga Aczél. A male and a female of this ge-
described) plus at least two undescribed species from Costa nus from the Durango region of Mexico are in the CNC.
Rica that are known only from females. Neotropical species One specimen has associated elevational data (collected at
can be keyed using Rafael (1990c). 2743  m). Based on these Mexican specimens, this genus
Cephalosphaera Enderlein. Nine described species could reasonably be expected to occur in the mountains else-
occur in Central America, Mexico, and the West Indies. where in Central America.
Keys to the Neotropical (Rafael, 1992), the West Indian
Microcephalops De  Meyer. Microcephalops latifrons
(Rafael, 1996), and the Costa Rican species (Rafael  &
(Hardy) from the West Indies and M.  williamsi (Hardy),
Menezes, 1999) are available.
which is widespread across the Neotropical Region, are the
Chalarus Walker. Five species have been identified (four only described species. Two unidentified species (based on
are described) plus at least two more from Costa Rica that females) have also been recorded in Costa Rica. The Neotro-
are known only from females. Neotropical species can be pical Region species were revised by Rafael (1990c) under
keyed using Rafael (1990d). Cephalops, group latifrons.

Dorylomorpha Aczél. This widespread genus has not pre- Nephrocerus Zetterstedt. Two female specimens in
viously been recorded in Central America. However, a male the CNC were collected in San Cristobal  de Las Casas
of a new species of Dorylomorpha (Dorylomyia) is in the (2160  m, Chiapas, Mexico), and an additional female was
USNM, and a female of a new species (possibly the same) collected near El Salto (2499 m, Durango, Mexico). As with
is in INBio. Albrecht (1990) provides keys to the world Jassidophaga, this genus should be searched for elsewhere
species. in the mountains of Central America.

Elmohardyia Rafael. All seven Central American species Pipunculus Latreille (= Parapipunculus Rafael). Ten spe-
occur in Costa Rica and can be identified using the key in cies have been identified from Central America and Mexico
Rafael & Menezes (1999). Eleven undescribed species were (nine described). Skevington & Marshall (1998) provided a
discovered in recent collections made by INBio. key to all New World species.

Eudorylas Aczél (= Metadorylas Rafael as synonymized Tomosvaryella Aczél. This is the second largest genus
recently by Skevington  & Yeates, 2001). This is the larg- in this region with 15 recorded species. The species from
est genus in the family and certainly the largest in Central Mexico, Central America, and the West Indies were revised
America. Many undescribed Costa Rican species exist in and keyed by Ale-Rocha & Rafael (1995). The West Indian
collections. Recent work on the phylogeny of the Eudorylini species were keyed by Rafael (1996).
has created a firm footing for the classification of the tribe in
all but the Neotropical Region (Skevington & Yeates, 2001). Verrallia Mik. One Central American specimen is known
There appears to have been a rapid radiation of members of from Chichicastenango, Guatemala. Previously, Verrallia
this tribe within the Neotropical Region, leaving an appar- was not recorded south of 37°N in the New World. We are
ently heterogeneous mix of species that are currently all con- now aware of specimens from the southwestern USA (New
fined to the genus Eudorylas. Efforts to place this fauna into Mexico), Mexico, and Guatemala. Only two Nearctic species
a world context will continue. It is apparent that the genera are known, but the genus is in need of revision in the New
Claraeola Aczél and Dasydorylas Skevington occur in the World. The Guatemalan specimen likely belongs to an un-
New World tropics, but considerable work remains before described species.
802
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Literature cited

Albrecht, A. 1990. Revision, phylogeny and classification of the genus Rafael, J.A. 1987a. Basileunculus, um gênero novo de Pipunculi-
Dorylomorpha (Diptera, Pipunculidae). Acta Zoologica Fennica dae (Diptera) da região Neotropical, com chave para as espé-
188: 1–240. cies. Acta Amazonica 16–17: 627–634.
Ale-Rocha, R. & J.A. Rafael. 1995. Tomosvaryella Aczél do México, Rafael, J.A. 1987b. Two new genera of Pipunculidae (Diptera)
América Central e Antilhas (Diptera, Pipunculidae). Revista Brasi- from the New World: Metadorylas,  gen.  n.  & Elmohardy-
leira de Zoologia 12: 407–427. ia gen. n., with new synonyms, designation of lectotypes and
Benton, F.P. 1975. Larval taxonomy and bionomics of some British revalidation of a species. Revista Brasileira de Entomologia
Pipunculidae. Ph.D. thesis, University of London, 208 pp. 31: 35–39.
Coe, R.L. 1966. Diptera Pipunculidae. Handbooks for the Identification Rafael, J.A. 1990a. As espécies Neotropicais de Metadorylas
of British Insects 10: 1–83. Rafael (Diptera: Pipunculidae). Boletim do Museu Paraense
Cumming, J.M., B.J. Sinclair, & D.M. Wood. 1995. Homology and phy- Emilio Goeldi, Séries Zoologia 6: 127–164.
logenetic implications of male genitalia in Diptera — Eremoneura. Rafael, J.A. 1990b. O Gênero Allomethus Hardy (Diptera, Pipun-
Entomologica scandinavica 26: 121–151. culidae). Revista Brasileira de Entomologia 34: 811–815.
De Meyer, M. 1996. World catalogue of Pipunculidae (Diptera). Institut Rafael, J.A. 1990c. Revisão das espécies Neotropicais do gênero
Royal des Sciences Naturelles de Belgique, Documents de Travail Cephalops Fallén (Diptera: Pipunculidae). Acta Amazonica
86: 1–127. 20: 353–390.
De Meyer, M. & J.H. Skevington. ���������������������������������������
2000. First addition to the world cata- Rafael, J.A. 1990d. Revisão das espécies Neotropicais do gênero
logue of Pipunculidae. Bulletin de l’Institut Royal des Sciences Natu- Chalarus Walker, 1834 (Diptera: Pipunculidae). Iheringia, sé-
relle de Belgique, Entomologie 70: 5–11. rie Zoologia 70: 45–53.
Ferrar, P. 1987. A guide to the breeding habits and immature stages of Rafael, J.A. 1992. Chapter 42. A review of the Neotropical species
Diptera Cyclorrhapha. Entomonograph 8(1), E.J. Brill / Scandinav- of big-headed flies, genus Cephalosphaera (Diptera: Pipuncu-
ian Science Press, Leiden, Copenhagen, 478 pp. lidae), pp. 633–646. In Quintero, D. & A. Aiello (editors). In-
Freytag, P.H. 1988. Rearing techniques for Dryinidae and their hyperpar- sects of Panama and Mesoamerica. Selected studies. Oxford
asites (parasitic Hymenoptera), pp. 519–524. In Gupta, V.K. (editor). University Press, Oxford, New York, Tokyo, xxii + 692 pp.
Advances in Parasitic Hymenoptera Research. E.J.  Brill, Leiden/ Rafael, J.A. 1993. Espécies de Eudorylas Aczél (Diptera, Pi-
New York, 546 pp. punculidae) do México e América Central. Revista Brasileira
Hardy, D.E. 1963. Studies in Pipunculidae (Diptera) of Colombia. Pro- de Entomologia 37: 751–762.
ceedings of the Hawaiian Entomological Society 18: 259–266. Rafael, J.A. 1996. Pipunculidae (Insecta, Diptera) of the Domini-
Huq, S.B. 1982. A contribution to the biology of pipunculid-flies (Pipun- can Republic: new records and description of new species. An-
culidae: Diptera). Ph.D. thesis, Free University, Berlin, 128 pp. nals of the Carnegie Museum 65: 363–381.
Huq, S. 1984. Breeding methods for Pipunculidae (Diptera), endop- Rafael, J.A. & M. De Meyer. 1992. Generic classification of the
arasites of leafhoppers. International Rice Research Newsletter 9: family Pipunculidae (Diptera): a cladistic analysis. Journal of
14–15. Natural History 26: 637–658.
Jervis, M.A. 1980. Studies on oviposition behaviour and larval devel- Rafael, J.A. & M.D. da S. Menezes. 1999. Taxonomic review of
opment in species of Chalarus (Diptera, Pipunculidae), parasites Costa Rican Pipunculidae (Insecta: Diptera). Revista de Biolo-
of typhlocybine leafhoppers (Homoptera, Cicadellidae). Journal of gia Tropical 47: 513–534.
Natural History 14: 759–768. Skevington, J.H. 1999. Cephalosphaera Enderlein, a genus of
Jervis, M.A. 1992. A taxonomic revision of the pipunculid fly genus Pipunculidae (Diptera) new for Australia, with descriptions
Chalarus Walker, with particular reference to the European fauna. of four new species. Australian Journal of Entomology 38:
Zoological Journal of the Linnean Society 105: 243–352. 247–256.
Koenig, D.P. & C.W. Young. 2007. First observation of parasitic rela- Skevington, J.H. 2000. Pipunculidae (Diptera) systematics: Spot-
tions between big-headed flies, Nephrocerus Zetterstedt (Diptera: light on the diverse tribe Eudorylini in Australia. Ph.D. thesis,
Pipunculidae) and crane flies, Tipula Linnaeus (Diptera: Tupulidae: University of Queensland, Brisbane, 446 pp.
Tipulinae), with larval and puparial descriptions for the genus Neph- Skevington, J.H. 2001. Revision of Australian Clistoabdominalis
rocerus. Proceedings of the Entomological Society of Washington (Diptera: Pipunculidae). Invertebrate Taxonomy 15: 695–761.
109: 52–65. Skevington, J. & S.A. Marshall. 1997. First record of a big-headed
May, Y.Y. 1979. The biology of Cephalops curtifrons (Diptera: Pipuncu- fly, Eudorylas alternatus (Cresson) (Diptera: Pipunculidae),
lidae), an endoparasite of Stenocranus minutus (Hemiptera: Delpha- reared from the subfamily Cicadellinae (Homoptera: Cicadel-
cidae). Zoological Journal of the Linnean Society 66: 15–29. lidae), with an overview of pipunculid–host associations in the
Morakote, R. & K.J. Yano. 1987. Morphology of immature stages of Nearctic Region. The Canadian Entomologist 129: 387–398.
some Japanese Pipunculidae (Diptera) parasitizing Nephotettix cinc- Skevington, J. & S.A. Marshall. 1998. Systematics of New World
ticeps (Hemiptera, Deltocephalidae). Kontyû 55: 176–186. Pipunculus (Diptera: Pipunculidae). Thomas Say Publications
Morakote, R. & K.J. Yano. 1988. ���������������������������������
Biology of some Japanese Pipuncu- in Entomology: Monographs. Entomological Society of Amer-
lidae (Diptera) parasitizing Nephotettix cincticeps (Hemiptera, Del- ica, Lanham, Maryland, 201 pp.
tocephalidae). Bulletin of the Faculty of Agriculture, Yamaguchi Skevington, J.H.  & D.K. Yeates. 2000. Phylogeny of the Syr-
University 35: 9–22. phoidea (Diptera) inferred from mtDNA sequences and
Perkins, R.C.L. 1905. Leaf-hoppers and their natural enemies morphology with particular reference to classification of the
(pt. IV. Pipunculidae). Bulletin. Division of Entomology. Hawai- Pipunculidae (Diptera). Molecular Phylogenetics and Evolu-
ian Sugar Planters Association Experiment Station 1: 123–157. tion 16: 212–224.
Perkins, R.C.L. 1906. Leaf-hoppers and their natural enemies. Skevington, J.H. & D.K. Yeates. 2001. Phylogenetic classification
(Introduction). Bulletin. Division of Entomology. Hawaiian of Eudorylini (Diptera, Pipunculidae). Systematic Entomology
Sugar Planters Association Experiment Station 1: i–xxxii. 26: 421–452.
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Vega, F.E., P. Barbosa,  & A.P.  Panduro. 1991. Eudorylas (Meta- parasitoid of the corn leafhopper (Hemiptera: Cicadellidae)
dorylas)  sp. (Diptera: Pipunculidae): a previously unreported in Argentina, with a table of pipunculid–host associations
parasitoid of Dalbulus maidis (Delong and Wolcott) and Dal- in the Neotropical Region. Neotropical Entomology 38:
bulus elimatus (Ball) (Homoptera: Cicadellidae). The Canadian 152–154.
Entomologist 123: 241–242. Waloff, N.  & M.A. Jervis. 1987. Communities of parasitoids
Virla, E.G. & J.A. Rafael. 1996. Datos bionómicos preliminares y associated with leafhoppers and planthoppers in Europe,
descripcion de la hembra de Cephalops penepauculus (Hardy) pp. 281–402. In Macfayden, A. & E.D. Ford (editors). Ad-
(Diptera: Pipunculidae) um parasitoide de Delphacidae (Homop- vances in Ecological Research. Vol. 17. Academic Press,
tera: Auchenorryncha) em Argentina. Revista de Investigación London, 382 pp.
Centro de Investigaciones para la Regulacion de Poblaciones Williams, F.X. 1919. Some observations of Pipunculus flies
de Organismos Nocivos, Tucumán 10: 33–36. which parasitize the cane leafhopper, at Pahala, Hawaii,
Virla, E.G., G. Moya-Raygoza, & J.A. Rafael. 2009. First Record Feb. 11–April 25, 1918. Proceedings of the Hawaiian Ento-
of Eudorylas schreiteri (Shannon) (Diptera: Pipunculidae) as a mological Society 4: 68–71.
804
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
MICROPEZIDAE (STILT-LEGGED FLIES) 55
Stephen A. Marshall

Fig. 55.1. Female of Poecilotylus similis Hennig, (illustrated by J. Knopp).

Diagnosis one Nearctic species, and spiracles vestigial in one Australian

species). Mandibles of known Neotropical species lack teeth
Medium to large (body length 5–17  mm), distinctively (the mandible is ventrally toothed in at least one Australian
long-legged flies (Fig. 1). Wing veins R4+5 and M1+2 conver- species). Anterior spiracles usually fan-shaped, but sometimes
gent towards wing tip. Ocellar bristles and vibrissae absent. reduced or divided into two lobes. Larvae of genera that occur
Most Neotropical species with distinctive vertical, fanlike in Central America were described by Albuquerque (1972),
row of bristles on katepisternum. Females with tergite and Steyskal (1964), Müller (1957), and Sabrosky (1942).
sternite  7 fused to form rigid oviscape housing ovipositor. Neriidae, the long-legged flies most similar to Micropezi-
Males of Micropezinae and Eurybatinae with surstyli; Tae- dae, have an apical arista in contrast with the dorsobasal arista
niapterinae without. Males of most species with prominent of Micropezidae. Neriids can also usually be distinguished
forked process on sternite 5. from micropezids by the presence of vibrissae and spine-
Larvae of relatively few species known, but known spe- like femoral bristles. Tanypezidae are superficially similar to
cies of genera that occur in the Neotropical Region with two micropezids, but are easily distinguished by the presence of
dorsally pointed posterior respiratory processes bearing spi- strong anepisternal bristles and the absence of katepisternal
racular discs, each with three or four slits (numerous pores in bristles.

805
806
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Biology Adults of many micropezid species are attracted to dung,

and they can often be seen in great numbers on leaves that have
Micropezids are common flies, often seen standing prom- been spattered by monkeys from the trees above. Many of the
inently on leaves or similar surfaces where they closely re- specimens on which this chapter is based were collected using
semble sympatric Hymenoptera, such as ants, spider wasps, small dung baits placed on logs, leaves, or the ground. Some
and ichneumonoid wasps. Most Neotropical micropezids micropezids are attracted to artificial honeydew (honey and water
seem to mimic ichneumonoids in both morphology and sprayed on leaves), and a few species are attracted to decompos-
movement. The forelegs, which are much shorter than the ing fruit. Some groups (Tenthes, Metasphen) are not attracted to
mid- and hind legs, are usually pigmented like the anten- dung or sugar baits, but occur in aggregations on broad leaves,
nae of sympatric wasps and are waved in front of the body where they appear to feed on insect feces and other debris on the
in an antenna-like fashion. A few species, especially in the leaf surfaces. Some micropezid adults feed on living or dead
genus Plocoscelus, are striking ant mimics in both shape and insects, and Taeniaptera lasciva has been recorded as a predator
behavior. of adult sugarcane moth borers (Diatraea saccharalis) in
Micropezids often engage in elaborate courtship behaviors Barbados (Bennett & Alam, 1985).
involving conspicuous male displays, transfer of regurgi- Some Micropezidae are associated with edge habitats
tated fluids from the male to the female, and other complex (Plocoscelus harenosus Cresson, for example) or even highly
behaviors. Wheeler (1924) described the remarkable court- disturbed areas, especially a few Taeniaptera species such as
ship behavior of Plocoscelus arthriticus (Wiedemann) (as T. albibasis (Enderlein) and T. lasciva. Other taxa are usually as-
Cardiacephala myrmex (Schiner)), recording a sequence sociated with native forests, and a few taxa (such as Mesoconius)
of behaviors including posturing, stroking, and exchange are exceptionally habitat-restricted. Mesoconius species rarely
of oral fluids. Similar behaviors occur in Ptilosphen (Ortiz, occur outside pristine or almost pristine, high elevation forests.
2001), Taeniaptera, and Scipopus. Ortiz (2001) described
the behavior of Ptilosphen viriolatus Enderlein in detail. This Classification
species forms “sleeping aggregations”, in which individual
flies spend the night on leaves, facing the petiole, while The Micropezidae belong to the superfamily Nerioidea, along
continuously and slowly waving their forelegs. with the Cypselosomatidae, Pseudopomyzidae, and Neriidae.
Little is known about micropezid larval habitats, but The family is currently divided into five subfamilies, of which
some Taeniapterinae have been reared from dung and three (Micropezinae, Eurybatinae, and Taeniapterinae) occur in
decaying vegetable matter. Several genera are associated the Neotropical Region (Steyskal, 1968; McAlpine, 1975). The
with decaying wood, and Scipopus limbativertex Enderlein primarily Pacific–Oriental subfamily Eurybatinae has only re-
and Ptilosphen species have been observed ovipositing in cently been recorded from the Neotropical Region, based on a
stumps and the ends of logs in Costa Rica. The Nearctic species described from Costa Rica (Marshall, 2002). The over-
species Taeniaptera trivittata Macquart has been observed whelming majority of Neotropical micropezids belong to the
ovipositing in, and emerging from, the below-ground parts diverse subfamily Taeniapterinae. The generic classification of
of rotting Typha stems. The Nearctic species Rainieria the Taeniapterinae is currently under revision and is likely to
antennaepes (Say) and Calobatina geometroides (Cresson) undergo significant change in the coming years.
have been reared from rotting material in dead or damaged
trees in the United States (Sabrosky, 1942; Wallace, 1969).
Scipopus belzebul (Schiner) can develop, under laboratory Identification
conditions, on rotting bananas and human feces (Albuquerque,
1972). Several Neotropical micropezids, for example, Despite their relatively large size and conspicuous be-
Plocoscelus conifer (Hendel), Taeniaptera lasciva (Fabr- havior, most micropezids are not easily identified. Hennig’s (1934,
icius), Taeniaptera annulata (Fabricius), have been reared 1935a, 1935b, 1936) keys to Neotropical species remain the most
from decaying banana stumps or decaying sugar canes. Else- important references for identification of Central America
where in the world, taeniapterine larvae have been recorded species, even though they do not include all of the species now
from live ginger roots (Steyskal, 1964), in banana roots, and known to occur in this region. Other works of value in identifying
under the bark of dead trees. The only Neotropical member Central American Micropezidae include Aczél’s (1951) review
of the subfamily Eurybatinae is unknown as a larva, but one of the Argentine species of Micropezidae, and Albuquerque’s
wingless Australian species lives in pitcher plants (Yeates, (1989) key to species of Plocoscelus. Steyskal (1968) prefaced
1992). Neotropical Micropezinae also remain unknown as his catalogue of Neotropical Micropezidae with the statement
larvae, but larvae of the secondarily widespread Micropeza that “most of the described species are still known only from their
corrigiolata (Linnaeus) develop in fresh, healthy root nodules type localities”. This remains true today; however, a significant
of several leguminous plants (Müller, 1957). Other Micro- number of species previously known only from type locali-
peza species, which are commonly collected on legumes, are ties in South America or elsewhere in Central America are
likely to have similar habits. now represented in the Costa Rican INBio collections.
MICROPEZIDAE (stilt-legged flies) 55 807

Key to the genera of Micropezidae of the Neotropical Region

1. Fronto-orbital bristles absent; crossvein CuA2 transverse; cells  bm and dm confluent

(crossvein  bm-cu absent)  (Fig.  5); upper calypter bare or with small, inconspicuous setulae
only; one to four katepisternal bristles, not forming a dense row (Fig.  2); surstyli present;
MICROPEZINAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– One to three fronto-orbital bristles present (often small) (Figs. 11–24); crossvein CuA2 slightly (Fig. 6)
to extremely (Fig. 7) sloped distally, posteroapical angle of cell cup slightly to extremely acute;
cells bm and dm separate (crossvein bm-cu present); upper calypter with tightly packed comb
of short bristles; katepisternal setae almost always forming dense, vertical, fanlike row (Fig. 1);
surstyli normally absent, but one rare species of Eurybatinae lacking katepisternal fan has surstyli
present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

2. Dorsocentral bristles present in one or two pairs; Argentina and Chile. . . . . [Cryogonus Cresson]
– Dorsocentral bristles absent; Micropeza Meigen. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3. One notopleural bristle present; cell r4+5 open (Fig. 5) or closed at margin. . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Micropeza (Micropeza) Meigen
– Two notopleural bristles present; cell r4+5 closed at or before margin, often petiolate. . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Micropeza (Neriocephalus) Enderlein

4. Katepisternum with only three large posterior bristles, not forming fanlike row; subscutellum
strongly developed; surstyli of male present; EURYBATINAE. . . . . . . . . . . . Notenthes Marshall
– Katepisternum with dense, fanlike, vertical row of posterior bristles; subscutellum not developed;
surstyli of male absent (Figs. 1, 4); TAENIAPTERINAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

5. Thoracic bristles conspicuously long, four to six dorsocentral bristles present including at least
three long presutural bristles; subcosta fused with R1 distally; Bolivia, Ecuador. . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Chaetotylus Hendel]
– Dorsocentral bristles usually in zero to two pairs, rarely (one Brazilian Taeniaptera) in four pairs;
subcosta free from R1 distally (Figs. 6, 7), usually divergent from R1 but parallel with and close to
R1 in some small species with one dorsocentral bristle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

6. Arista pectinate (long-haired on upper side only) (Fig. 19); scutellum upturned; mid- and hind
femur with abrupt, knotlike swelling. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
– Arista bare to plumose or long-pubescent (Fig. 21), not haired on one side only; scutellum not
upturned; femora rarely with abrupt knotlike swellings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

7. Wing clear with brown crossbands; vertex flattened, if medially sunken then dull tomentose. . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plocoscelus Enderlein
– Wing brownish, with round hyaline spot(s); vertex cordate (medially sunken to give the posterior
margin a heart-shape), smooth and shining; northern South America, Trinidad. . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Cardiacephala Macquart]

8. Anterior margin of clypeus with few short, erect setulae; Mexico, Caribbean, Florida, Brazil . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Hoplocheiloma Cresson]
– Anterior margin of clypeus bare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

9. Cell cup long, CuA2 as long as or longer than A1 + CuA2 (Fig. 7) . . . . . . . . . . . . . . . . . . . . . . . 10

– Cell cup short, CuA2 shorter than A1 + CuA2 (Fig. 6) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
808
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

2 Micropeza 3 Taeniaptera

4 Pseudeurybata

9 Cliobata
A1+CuA2
5 Micropeza
br

CuA2

bm cup A1+CuA2

6 Rainiera

7 Taeniaptera 8 Cliobata 10 Tenthes

Figs. 55.2–10. Adult males, male terminalia, wings, and legs: male of (2) Micropeza lineata Van Duzee, (MND, fig. 56.1); (3) Taeniaptera
vulgata Hennig, (Caribbean); lateral view of male terminalia of (4)  Pseudeurybata  sp.; dorsal view of wing of (5)  M. lineata, (MND,
fig. 56.11); (6) Rainieria antennaepes (Say), (MND, fig. 56.13); (7) Ta. trivittata Macquart, (MND, fig. 56.12); anterior view of hind tibia of
(8) Cliobata sp., (South America); cross section of hind tibia of (9) Cliobata sp.; anterior view of hind leg of (10) Tenthes citatus Cresson,
(South America). Figure 4 illustrated by M. Musial. Figures 3 and 8–10 illustrated by K. Bethune-Leamen and A. Cormier.
MICROPEZIDAE (stilt-legged flies) 55 809

10. Arista plumose (hairs much longer than width of arista at base); postocellar bristle absent (Figs. 21,
22) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
– Arista bare or nearly so (hairs no longer than width of arista at base) (Figs. 11–18); postocellar
bristle usually present (Figs. 19, 23) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

11. Third costal section (costa between end of R2+3 and R4+5) short (less than 0.3 times as long as M
beyond crossvein  dm-cu); frons convex in profile; frontal vitta not clearly differentiated from
fronto-orbital plates (Fig. 21); Brazil, Ecuador, Guyana. . . . . . . . . . . . . . . [Parasphen Enderlein]
– Third costal section long (more than 0.3 times as long as M beyond crossvein dm-cu); frons
almost flat in profile; frontal vitta differentiated from fronto-orbital plates (Fig. 22). . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ptilosphen Enderlein

12. Mid- and hind femora with knotlike swelling (Fig. 10); subcosta and R1 approximated distally;
hind tibia usually with only small setulae dorsally, sometimes with few short distal dorsal bristles;
Peru, Ecuador. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Tenthes Cresson]
– Mid- and hind femora of uniform width or gradually swollen; subcosta and R1 separate distally;
hind tibia with one to two rows of dorsal bristles (Fig. 8). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

13. Scutellum densely setose, entire disc covered by fine setulae about half as long as marginal
bristles; dorsocentral bristles absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Grallomyia Rondani
– Scutellar disc mostly bare; setulae, if present, small and mostly restricted to margin or near
margin; dorsocentral bristles usually present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

14. Palpus parallel sided, apex broad and rounded; wing pigmentation usually with discal band
straight at least on distal edge (often convex proximally, giving the band a triangular appearance);
frontal vitta usually broad and raised anterior to ocellar plate, at least half as wide as frons. . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Poecilotylus Hennig
– Palpus apically pointed with ventrolateral edge strongly convex, usually broadest basally and
gradually tapering to apex (Fig. 13); discal wing band usually convex both proximally and
distally, giving band rounded appearance; frontal vitta flat, usually long and narrow, less than half
as wide as frons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Taeniaptera Macquart

15. Abdomen somewhat petiolate, basal two segments elongate and narrow; males without copulatory
fork (sternite 5 unmodified); frontal vitta broad, not reaching lower margin of frons (Fig. 20); hind
tibia cylindrical, not flattened or grooved; katatergite usually swollen, sometimes prominently so
(Fig. 1); large species, usually over 10 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .16
– Abdomen not petiolate; if weakly petiolate (some small species, under 9 mm) then frontal vitta
narrow; frontal vitta reaching lower margin of frons; copulatory fork present on male sternite 5
(Fig. 4) (males unknown for Planipeza); katatergite not swollen; size and hind tibia various. . 17

16. Katatergite strongly tuberculate, usually tapered to nipplelike tip (Fig. 1); robust species, often
brown or black and yellow. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mesoconius Enderlein
– Katatergite flat to slightly swollen, not tuberculate; slender species with elongate legs, often black
or black and red. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Zelatractodes Enderlein

17. Antennal bases contiguous; face narrow and recessed (Fig. 18); palpus thin, cylindrical (Fig. 17);
hind tibia strongly swollen in basal third; small (5–7 mm), dark brown flies with lower frons
orange and upper frons black; Venezuela, Ecuador. . . . . . . . . . . . . . . . . . . . [Globopeza Marshall]
– Antennae separated by more than half width of scape; face prominent at least on upper half;
palpus flat, not cylindrical; hind tibia not swollen, or with gradual swelling extending onto distal
half; size and color various.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
810
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

18. Subcosta distally approximated to R1, apices of Sc and R1 separated by no more than apical width
of R1; antennae separated by less than width of scape (Figs. 23, 24); head globose, conspicuously
large, length equal to greatest thoracic width; small (6–9 mm) flies with body and legs usually
orange and apex of epandrium yellow; if body darker then postocellar bristles absent (Fig. 23);
hind tibia cylindrical, neither flattened nor grooved . . . . . . . . . . . . . . . . . . . . . . . Metasphen Frey
– Subcosta distally diverging from R1; antennae separated by more than width of scape (Fig. 15);
head not disproportionately large, length conspicuously less than thoracic width; body length
usually over 10 mm, color, head chaetotaxy, and hind tibia various. . . . . . . . . . . . . . . . . . . . . . 19

19. Frons evenly reddish dull pruinose and evenly swollen or with frontal vitta slightly depressed;
no clear division between frontal vitta and fronto-orbital plates; frontal vitta of uniform width
and more than three-quarters as wide as frons; wing usually evenly infuscated, sometimes with
hyaline marks; large flies (body length over 15 mm) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
– Fronto-orbital plates shining relative to pruinose frontal vitta; frontal vitta either less than half as
wide as frons or strongly elevated anterior to ocelli; wing various, often banded; size various but
usually less than 14 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

20. Body color black or dark brown; postocellar bristles present; thoracic length subequal to height.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scipopus Enderlein
– Body color usually brown or reddish-brown, if dark brown then postocellar bristles absent; thorax
much longer than high. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

21. Postocellar bristles absent; body color brown to black with golden sheen; R4+5 almost straight
distally; South America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Phaeopterina Frey]
– Postocellar bristles present; body color orange to reddish-brown; R4+5 convex distally. . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudeurybata Hennig

22. Hind tibia cylindrical, without sulcus on outer face or with weak sulcus on basal quarter only;
parafacial expanded dorsally almost to antennal base. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
– Hind tibia flattened, with sulcus on outer face at least on basal half (Figs. 8, 9); parafacial widely
separated from antennal base dorsally.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

23. Arista pubescent on at least basal two-thirds (Fig. 16), although microtrichia sometimes fine and
hard to see; frontal vitta broad at each end, not strongly elevated in profile. . . . . . . . . . . . . . . . 24
– Arista bare or with fine pubescence on basal one-third only (Fig. 11); frontal vitta anterior to
ocellar triangle elevated in profile, usually tear-drop shaped and tapered anteriorly . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calosphen Hennig

24 Wing mostly clear, usually with basal, discal and distal bands; thorax compact, anterior half not
shallow and elongate; palpus narrow; most species orange and relatively delicate.. . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Grallipeza Rondani
– Wing heavily infuscated on anterior half with clear patches over crossveins; anterior half of thorax
shallow and strikingly elongate, extending well anterior of prolegs; palpus extremely broad; body
dark brown, robust; male unknown; Brazil. . . . . . . . . . . . . . . . . . . . . . . . . . [Planipeza Marshall]

25. Wing membrane infuscated, usually with hyaline marks or bands; palpus spatulate, less than
one-quarter as wide as long, ventral margin straight; frontal vitta at least half as wide as frons;
third costal section (costa between end of R2+3 and R4+5) usually less than half as long as distal part
of M; distal part of Cu usually more than half as long as crossvein dm-cu; stem of vein CuP visible
in cell cup; body length usually over 13 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . Cliobata Enderlein
– Wing membrane clear with brown marks or bands; palpus usually conspicuously broad, with
convex and pale ventral margin; frontal vitta usually less than half as wide as frons; third costal
section various, but usually more than half as long as distal part of M; distal part of Cu at most half
as long as crossvein dm-cu; stem of CuP not visible in cell cup; body length 7–13 mm; Rainieria
Rondani . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .26
MICROPEZIDAE (stilt-legged flies) 55 811

11 Calosphen 12 Cliobata 13 Taeniaptera 14 Rainieria

18 Globopeza
15 Rainieria (Rainieriella) 16 Grallipeza 17 Globopeza
poc s
o vt s

i vt s

orb s
fr s

21 Parasphen
20 Mesoconius
19 Plocoscelus

22 Ptilosphen 23 Metasphen 24 Metasphen

Figs.  55.11–24. Heads of Taeniapterinae: anterolateral view of head of (11)  Calosphen  sp., (South America); (12)  Cliobata  sp.;
(13)  Taeniaptera vulgata Hennig, (Caribbean); (14)  Rainieria alternata Cresson; (15)  R. (Rainieriella)  sp.; (16)  Grallipeza unifasciata
(Fabricius); anterior and lateral views (17,18) of head of Globopeza ecuadorensis Marshall (South America); lateral view of head of
Gl. equadorensis; anterolateral view of head of (19) Plocoscelus harenosus Cresson; anterior view of head of (20) Mesoconius sp., (South
America); (21) Parasphen frontalis Cresson, (South America); (22) Ptilosphen sp., (South America); (23) Metasphen gilvimana (Cresson);
(24) Me. obscura (Hennig), (South America). Figures 11–24 illustrated by K. Bethune-Leamen and A. Cormier.
Abbreviations: fr s, frontal setae; i vt s, inner vertical seta; orb s, orbital seta; o vt s, outer vertical seta; poc s, postocellar seta.
812
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

26. One frontal bristle present (Fig. 15). . . . . . . . . . . . . . . . . . . . . . . . Rainieria (Rainieriella) Hennig

– Two frontal bristles present (Fig. 14). . . . . . . . . . . . . . . . . . . . . . . . Rainieria (Rainieria) Rondani

Synopsis of the fauna here following Merritt & James (1973), but the internal phy-
logeny of this poorly known genus remains unresolved.
There are about 50 species of Micropezidae known from
Central America, but the actual fauna is probably closer to Notenthes Marshall. This is the only New World ge-
200 species. nus in the Eurybatinae. One species, Notenthes mcalpinei
Marshall, is known only from a few specimens from cloud
Calosphen Hennig. This genus is currently recorded only forest sites in Costa Rica. The other species, N.  rubella
from Bolivia, Colombia, and Venezuela, but occurs from (Wulp), is known only from the type specimens collected
Costa Rica to Bolivia. Calosphen and Grallipeza are closely in northern Mexico.
related and together form a group that needs to be revised. Plocoscelus Enderlein. This is a Neotropical genus
Cliobata Enderlein. Although currently recorded only common from Mexico to Brazil. Five of the 17 New World
species are recorded from Central America; four are record-
from South America, this genus also occurs in Central Amer-
ed from Costa Rica. Some species are common along forest
ica, including Costa Rica. It needs to be revised and rede-
edges and similar disturbed sites. They strongly resemble
fined with respect to the closely related genus Rainieria. ants and often occur in large numbers on open foliage.
Grallipeza Rondani. Species of this genus occur from the Poecilotylus Hennig. Four of the 15 species in this genus,
USA south to Argentina. Although only four of the 38 Neo- which occurs from Mexico south to Argentina, are recorded
tropical species are currently recorded from Central America from Central America, but at least 12 species of Poeciloty-
(only one from Costa Rica), at least eight species occur in lus actually occur in Costa Rica. The division of the genus
Costa Rica. The genus needs to be revised and redefined with into two subgenera based on the number of frontal bristles
respect to the closely related genus Calosphen. is of dubious value, and the genus as currently defined is
probably paraphyletic with respect to Taeniaptera.
Grallomyia Rondani. There is one species in this genus,
G. tarsata Wiedemann, distributed from Mexico to Brazil. It Pseudeurybata Hennig. This genus includes one de-
is treated as part of Taeniaptera by recent authors. scribed species from Panama and at least one undescribed
species from Mexico and Costa Rica. Pseudeurybata is
Mesoconius Enderlein. This group includes some of the closely related to, and possibly belongs within, the genus
largest and most striking of all Micropezidae, many of which Scipopus.
are brightly colored mimics of ichneumonoid wasps. Although
the only species in this genus currently recorded from outside Ptilosphen Enderlein. Occurring from Mexico to Peru
the Andean countries is M.  hemithorax Frey, described from and Brazil, these are usually relatively large flies, often com-
Panama, M. hemithorax and a number of other Mesoconius spe- mon in forests. Three of the 20 (15 described) species in
cies are reasonably common in undisturbed high elevation for- this genus are recorded from Costa Rica, and an additional
ests in Costa Rica. All records of Mesoconius seem to be from five undescribed or unrecorded species occur in the country.
deep in the interior of undisturbed cloud forest. Probably five One additional Central American species is recorded from
described and two undescribed species occur in Costa Rica. Panama. Some common Ptilosphen exhibit striking sexual
dimorphism in color, and males and females of some species
Metasphen Frey. Described as a monotypic genus for were described under different names.
M. vigilans Frey, from Brazil, this genus has recently been re-
Rainieria Rondani. This genus includes 17 described
defined to include Rainieria (Globomyia) Hennig (Marshall, Neotropical species distributed from Mexico to Bolivia.
2004). Although not formerly recorded from Costa Rica, one Three species have Central American ranges including Costa
species, Metasphen gilvimana (Hennig), is common in low el- Rica, although only one species has been formally record-
evation forests there. Metasphen includes four species, distrib- ed from Costa Rica. A total of seven Rainieria species are
uted from Mexico to Peru. known from Mexico and Central America. Although the ge-
nus is usually divided into three subgenera, one (Globomyia)
Micropeza Meigen. This is a large and widespread ge- belongs in Metasphen. The other two subgenera (Rainieria
nus well represented in both temperate and tropical regions. and Rainieriella), currently separated by the number of fron-
Micropeza is the only genus in the Micropezinae other than tal bristles, are of dubious value.
the southern South American (Chile and Argentina) genus
Cryogonus. Ten of the 46 Neotropical species of Micropeza Scipopus Enderlein. This genus occurs from Mexico south
are recorded from Costa Rica; 15 species are recorded from to Argentina, with five of the 22 Neotropical species recorded
Central America. The genus is divided into two subgenera from Central America, all probably occurring in Costa Rica.
MICROPEZIDAE (stilt-legged flies) 55 813

Taeniaptera Macquart. Probably the most common a presumably synapomorphic character. Species formerly
genus of Micropezidae, Taeniaptera occurs throughout the placed in Taeniaptera that lack this character will key here
New World tropics. Seventeen of the 43 Neotropical species to the paraphyletic genus Poecilotylus.
of this genus are known from Central America, and most of
these species should occur in Costa Rica. Some Taeniaptera Zelatractodes Enderlein. This is a distinctive genus
species are synanthropic and are common flies in disturbed with all 11 species currently recorded only from the An-
areas where they are attracted to feces and other decompos- dean countries but also present in the Costa Rican fauna.
ing material. Taeniaptera as diagnosed here includes only Zelatractodes and Mesoconius are closely related and ex-
species with a strongly modified, wide and tapered palpus, tremely similar.

Literature cited

Aczél, M.L. 1951. Morfologia externa y división sistemática de Merritt, R.W. & M.T. James. 1973. The Micropezidae of Califor-
las «Tanypezidiformes» con sinópsis de las espécies Argentinas de nia (Diptera). Bulletin of the California Insect Survey 14: 1–29.
«Tylidae» («Micropezidae») y «Neriidae» (Dipt.). Acta Zoologica McAlpine, D.K. 1975. The subfamily classification of the Micro-
Lilloana 11: 483–589, pls.1–4. pezidae and the genera of Eurybatinae (Diptera: Schizophora).
Albuquerque, L. 1972. Sobre uma espécie do genero Scipopus En- Journal of Entomology B 43: 231–245.
derlein, 1922. Acta Amazonica 2: 89–94. Müller, H. 1957. Leguminosenknöllchen als Nahrungsquelle hei-
Albuquerque, L. 1989. Cardiacephalina subtr. n. de Taeniapterini mischer Micropezidae Larven. Beiträge zur Entomologie 7:
(Diptera, Micropezidae, Taeniapterinae) Memorias do Instituto 247–262.
Oswaldo Cruz 84: 19–26. Ortiz, P. 2001. Mimicry, Sleep and Sex: The natural history of a
Bennett, F.D. & M.M. Alam. 1985. An annotated check-list of the micropezid fly. Film presented to the Animal Behavior Society
insects and allied terrestrial arthropods of Barbados. Caribbean Annual Meeting, Corvallis, Oregon, 2001.
Agricultural Research and Development Institute, 81 pp. Sabrosky, C.W. 1942. An unusual rearing of Rainieria brunneipes
Hennig, W. 1934. Revision der Tyliden (Dipt., Acalypt.). I. Teil: Die Tae- (Cresson) (Diptera: Micropezidae). Entomological News 53:
niapterinae Amerikas. Stettiner Entomologische Zeitung 95: 65–108,
283–285.
294–330.
Steyskal, G.C. 1964. Larvae of Micropezidae (Diptera) including
Hennig, W. 1935a. Revision der Tyliden (Dipt. Acalypt.). I. Teil: Die Tae-
two species that bore in ginger roots. Annals of the Entomo-
niapterinae Amerikas. Stettiner Entomologische Zeitung 96: 27–67.
Hennig, W. 1935b. Revision der Tyliden (Dipt. Acalypt.). II. Teil: logical Society of America 57: 292–296.
Die ausseramerikanishcen Taeniapterinae die Trepidariinae und Steyskal, G.C. 1968. 49. Micropezidae. In Papavero, N. (editor).
Tylinae. Allgemeines über die Tyliden. Konowia 14: 68–92, A Catalogue of the Diptera of the Americas South of the United
192–216, 289–310. States. Departamento de Zoologia, Secretaria da Agricultura,
Hennig, W. 1936. Revision der Tyliden (Dipt. Acalypt.). II. Teil: Die São Paulo, 33 pp.
ausseramerikanishcen Taeniapterinae, die Trepidariinae und Tylinae. Wallace, J.B. 1969. The mature larvae and pupae of Calobatina
Allgemeines über die Tyliden. Konowia 15: 129–144, 201–239. geometroides (Cresson) (Diptera: Micropezidae). Entomologi-
Marshall, S.A. 2002. Notenthes a new genus of Eurybatinae and a cal News 80: 317–321.
first record of Eurybatinae from the New World (Diptera Micro- Wheeler, W.M. 1924. Courtship of the Calobatas. Journal of He-
pezidae). Studia dipterologica 9: 83–88. redity 15: 485–495.
Marshall, S.A. 2004. A review of the genus Metasphen Frey includ- Yeates, D.K. 1992. Immature stages of the apterous fly Badisis
ing Rainieria (Globomyia) Hennig new synonym. Studia dip- ambulans McAlpine (Dipera: Micropezidae). Journal of Natu-
terologica 11: 529–536. ral History 26: 417–424.
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
NERIIDAE (NERIID FLIES, CACTUS FLIES) 56
Matthias Buck

Fig. 56.1. Male of Odontoloxozus longicornis (Coquillett), (MND,  fig. 57.1).

Diagnosis with stout, spinelike ventral bristles, mid- and hind tibia
with ventroapical bristle, foretibia with rows of spine- or
Medium-sized to large flies (body length 5–15  mm), tubercle-like bristles in males of some species. Wing elon-
with long, slender legs (Fig. 1). Head usually longer than gate, with R4+5 and M strongly convergent; subcostal break
high. Frons with one to five fronto-orbital bristles, ocellar usually absent, additional crossveins present in one Central
bristles absent, vertical bristles sometimes reduced, vibrissa American species. Male terminalia elongate, flexed below
present in some species. Upper face deeply divided medi- abdomen. Female abdominal segment 7 developed as con-
ally and forming conspicuous antennal bases in Neriinae spicuous oviscape.
(Fig.  2: ant  bas). Antennae porrect; pedicel with elongate Neriidae can be distinguished from other long-legged
to triangular projection on inner side; scape and/or pedi- flies with apically narrowed wing cell  r4+5 (Micropezidae
cel elongated in some species; arista apical to dorsoapical, and Tanypezidae) by the porrect antennae with dorsoapical
bare or pubescent. Thorax with relatively short and stout or apical arista, the apicomedial projection of the pedicel,
bristles in Central American species; katatergite protuber- the peculiar development of the upper face, and the presence
ant. Foreleg with elongated coxa, at least forefemur usually of ventral femoral spines (at least on forefemur).

815
816
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

hum car
sct
ant bas

ppl

aprn rg

2 Cerantichir

3 Nerius

prepst sta

4 Odontoloxozus

6 Glyphidops (Oncopsia) 7 Glyphidops (Oncopsia)

5 Nerius
Sc R1
R2+3
bm R4+5
br r-m
cup
bm-cu M1
CuA2
dm-cu

8 Longina A1+CuA2 CuA1

9 Odontoloxozus

Figs. 56.2–9. Heads, thoraces, antennae, coxae, and wings: lateral view of head (antenna omitted) and anterior part of thorax of
(2) Cerantichir enderleini (Hennig); anterior view of right forecoxa of (3) Nerius plurivittatus Bigot; lateral view of head (antenna omitted)
and anterior part of thorax of (4) Odontoloxozus longicornis (Coquillett); medial view of right antenna (apex of arista omitted) of
(5) N. plurivittatus; and (6) Glyphidops (Oncopsia) flavifrons (Bigot); anterior view of right forecoxa of (7) G. (Oncopsia) flavifrons; dorsal
view of wing of (8) Longina anguliceps Buck & Marshall, (South America, Buck & Marshall, 2004, fig. 4); and (9) O. longicornis, (MND,
figs. 4.54, 57.4). Figures 2–4 and 6–7 illustrated by A. Cormier. Figure 5 illustrated by A. Brenes.
Abbreviations: ant bas, antennal base; aprn  rg, antepronotal ridge; hum  car, humeral carina; ppl,  postpronotal lobe; prepst  sta,  patch of
proepisternal setulae; sct, scutum.
NERIIDAE (neriid flies, cactus flies) 56 817

Biology Nerioidea. The Neriidae consist of two subfamilies: the

less derived and possibly paraphyletic Telostylinae and the
All Neriidae for which biological information is avail- more derived Neriinae (subfamily limits after Aczél, 1954,
able breed in decaying plant tissue. Species of the genus not Hennig, 1937). Only the Neriinae occur in the New
Odontoloxozus that inhabit arid areas of southwestern World. This subfamily is defined by the presence of promin-
North America and Central America develop in necrotic ent antennal bases, which are formed by the expanded and
cactus tissues (Ryckman & Olsen, 1963). Other neriid gen- medially divided upper face and lunule. Among the nine
era have been reported to use fresh decaying wood (large Neotropical genera, four occur in Central America and two
logs to small branches) (Eberhard, 1998), beetle borings reach the southern extreme of the Nearctic Region. The
in decaying wood (Preston-Mafham, 2001), injured trees other genera are restricted to South America or the Carib-
(including banana stems), or diseased trees as breeding bean (Antillonerius). The Neotropical genera of Neriidae
sites. Adults of the most common Central American genera, have not yet been analyzed from a phylogenetic point of
Glyphidops and Nerius, are often observed in these micro- view, and the validity of some genera and subgenera needs
habitats. The males of several species guard territories that critical evaluation.
represent favorable oviposition sites and/or guard females
after copulation and during oviposition (Mangan, 1979; Identification
Preston-Mafham, 2001). Adult Neriidae feed on sappy
excretions of trees and to a lesser degree on rotten fruit, The New World Neriidae were revised by Aczél
other decaying plant matter, small dung, and even carrion. (1961), who provided keys to genera and species. Since
Odontoloxozus longicornis (Coquillett) is the only
then, few taxonomic papers on New World Neriidae have
Neotropical species of the family whose immature stages
been published (Steyskal, 1965b; Mangan  & Baldwin,
have been described (Olsen  & Ryckman, 1963; Steyskal,
1986; Buck  & Marshall, 2004). Chaetotaxy has been
1965a). The polytene chromosomes of the salivary glands
used extensively in neriid taxonomy, but its usefulness
of third instar larvae of the sibling species O. longicornis
is limited because of high variability. At the generic
and O. pachycericola Mangan & Baldwin were described
level, thorax morphology, structure of the antenna and
by Mangan & Baldwin (1986).
antennal bases, as well as wing venation provide good
characters. Thoracic tomentum patterns appear to be
Classification quite constant and are very useful at the species level. Un-
fortunately, the male genitalia are highly uniform even
The Neriidae, together with the Micropezidae, Cypselo- between genera and provide little taxonomic or phyloge-
somatidae, and Pseudopomyzidae, belong to the superfamily netic information.

Key to the genera of Neriidae of the New World

1. Wing with supernumerary crossveins originating from both sides of R2+3 and M. . . . . . . . . . . . . 2
– Wing without supernumerary crossveins. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

2. Vibrissa absent; only one pair of dorsocentral bristles; forecoxa with one anterior apical to
preapical bristle, mid- and hind coxa each with one lateral bristle; mid- and hind femur with
strong, spinelike bristles ventrally. . . . . . . . . . . . . . . . . Glyphidops (Oncopsia) Enderlein, in part
– Vibrissa present; five to six pairs of dorsocentral bristles; forecoxa with two anteroapical to preapical
bristles (similar to Fig. 3), mid- and hind coxa each with two lateral bristles; mid- and hind femur
without strong, spinelike bristles; ventral bristles only slightly stronger than surrounding hairs;
South America. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eoneria Aczél, in part

3. Antennal bases at least in part shining black or brown (inner margin often dull and paler); one pair
of dorsocentral bristles present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
– Antennal bases completely tomentose, brown or pale, at most subshining; one to six pairs
of dorsocentral bristles present; Caribbean, South America. . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

4. Scape at most slightly longer than broad (as seen from above). . . . . . . . . . . . . . . . . . . . . . . . . . . 5
– Scape at least twice as long as broad; South America. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
818
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

5. Length of pedicel (excluding apicomedial process) subequal to or greater than length of

first flagellomere; arista whitish pubescent (except sometimes basal third). . . . . . . . . . . . 6
– Pedicel short, about half as long as first flagellomere, if subequal then arista brown and
bare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

6. Vein dm-cu not oblique (similar to Fig. 8), sometimes bent near middle, strongly diverging
from wing hind margin; anterior margin of frons at most weakly convex, not produced between
antennal bases; proepisternum bare anterodorsally; scutum anteriorly projecting distinctly
beyond level of humeral carina (Fig. 2: hum car); antepronotal ridge (Fig. 2: aprn rg) prominent,
distinctly anterior to humeral carina. . . . . . . . . . . . . . . . . . . . . . . . . Cerantichir Enderlein

– Vein dm-cu strongly oblique, almost parallel to wing hind margin (Fig. 9); anterior margin
of frons triangularly produced between antennal bases; proepisternum with patch of black
setulae (at least two) anterodorsally (Fig. 4: prepst sta); scutum anteriorly ending at level
of humeral carina (Fig.  4); antepronotal ridge inconspicuous, almost at level of (below)
humeral carina (Fig. 4). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Odontoloxozus Enderlein

7. Apicomedian process of pedicel short, triangular or rounded (Fig.  5), at most as long as
wide at base; prosternum broad, fused to proepisternum or scarcely separated from latter by
narrow membranous area; forecoxa with two anterior apical to preapical bristles (Fig. 3);
midcoxa with two lateral bristles; vibrissa present or absent. . . . . . . . . . . Nerius Fabricius
– Apicomedian process of pedicel long, narrow and pointed (Fig. 6), distinctly longer than its
width at base; prosternum narrow, broadly separated from proepisternum by membranous
area; forecoxa with one anterior apical to preapical bristle (Fig. 7); midcoxa with one to
two lateral bristles; vibrissa absent; Glyphidops Enderlein, in part . . . . . . . . . . . . . . . . . . 8

8. Arista whitish pubescent, at least in basal third. . . . . . Glyphidops (Glyphidops) Enderlein

– Arista dark, bare or with scattered dark pubescence; short, thickened base sometimes whitish
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Glypidops (Oncopsia) Enderlein, in part

9. Antennae separated by more than twice diameter of scape; pedicel distinctly longer than scape;
apex of first flagellomere broadly rounded; vein dm-cu strongly oblique, almost parallel to wing
hind margin (similar to Fig. 9); wing extensively microtrichose towards base (including cells c,
dm, and br beyond level of humeral crossvein); South America. . . . . . . . . [Loxozus Enderlein]
– Antennae more approximated, separated by less than diameter of scape; pedicel (excluding
apicomedial process) usually shorter than scape, if longer, then first flagellomere tapered towards
apex; vein dm-cu at most slightly oblique (Fig. 8); wing membrane with large bare areas towards
base (including cells c, br, and basal part of dm); South America. . . . . . . . [Longina Wiedemann]

10. One pair of dorsocentral bristles present; vibrissa absent; Caribbean. . . . . [Antillonerius Hennig]
– Five to six pairs of dorsocentral bristles present; vibrissa present; South America. . . . . . 11

11. Arista white pubescent; first flagellomere slightly longer than pedicel; vein dm-cu strongly
oblique; midcoxa with three lateral bristles; South America . . . . . . . . . . [Eoloxozus Aczél]
– Arista brown, bare to pubescent; first flagellomere distinctly longer than pedicel; vein  dm-cu
slightly oblique; midcoxa with two lateral bristles; South America . . . . . [Eoneria Aczél, in part]
NERIIDAE (neriid flies, cactus flies) 56 819

Synopsis of the fauna Central American species G. (O.) flavifrons (Bigot) appear to

develop under the bark of fresh, decaying tree branches or in
The Central American fauna of this family appears to be injured banana stems (M. Buck, personal observation).
relatively small, consisting of about 20 species.
Nerius Fabricius. Five described species of this Neotro-
Cerantichir Enderlein. Previously, this genus included pical genus occur in Central America. Females of the common
only the endemic Central American species C.  enderleini Central American species N. plurivittatus Bigot oviposit in
(Hennig). Based on the shared apomorphic condition of cracks of bark of decaying tree trunks (Eberhard, 1998). The
the anterior part of the thorax (see couplet 6 of key), Odon- larvae probably develop in the decaying cambium layer
toloxozus peruanus Hennig is transferred here to Cerantichir below the bark.
(new combination). Cerantichir peruana ranges from Costa
Rica south to Bolivia. A female of this species was observed Odontoloxozus Enderlein. After the removal of
ovipositing under the bark of a decaying log (S.A. Marshall, O.  peruanus Hennig (see above under Cerantichir), the
personal communication). genus Odontoloxozus now includes only two closely relat-
ed cryptic species that can be identified using Mangan  &
Glyphidops Enderlein. This primarily Neotropical genus Baldwin (1986). Odontoloxozus longicornis (Coquillett) is
ranges into the southern Nearctic Region. Two species of the distributed from the southern USA to northern Costa Rica,
nominate subgenus and four species of the subgenus Onco- whereas O. pachycericola Mangan & Baldwin appears to be
psia Enderlein have been reported from Central America. restricted to Baja California Sur, Mexico. Larvae of both spe-
At least two Central American and many South American cies develop in decaying tissues of various cacti (Ryckman &
species remain to be described. Larvae of the most common Olsen, 1963; Mangan & Baldwin, 1986).

Literature cited

Aczél, M.L. 1954. Results of the Archbold Expeditions. Neriidae California Sur (Mexico). Proceedings of the Entomological
von Neuguinea (Diptera). Treubia 22: 505–531. Society of Washington 88: 110–121.
Aczél, M.L. 1961. A revision of American Neriidae (Diptera, Olsen, L.E.  & R.E. Ryckman. 1963. Studies on Odontoloxozus
Acalyptratae). Studia Entomologica (nova série) 4: 257–446. longicornis (Diptera: Neriidae). Part I. Life history and descrip-
Buck, M. & S.A. Marshall 2004. A review of the genus Longina tions of immature stages. Annals of the Entomological Society
Wiedemann, with descriptions of two new species (Diptera, of America 56: 454–469.
Neriidae). Studia dipterologica 11: 23–32. Preston-Mafham, K. 2001. Resource defence mating system in two
Eberhard, W.G. 1998. Reproductive behavior of Glyphidops flies from Sulawesi: Gymnonerius fuscus Wiedemann and
flavifrons and Nerius plurivittatus (Diptera, Neriidae). Journal Telostylinus sp. near duplicatus Wiedemann (Diptera: Neriidae).
of the Kansas Entomological Society 71: 89–107. Journal of Natural History 35: 149–156.
Hennig, W. 1937. Übersicht über die Arten der Neriiden und über Ryckman, R.E. & L.E. Olsen. 1963. Studies on Odontoloxozus lon-
die Zoogeographie dieser Acalyptraten-Gruppe (Diptera). gicornis (Diptera: Neriidae). Part II. Distribution and ecology.
Stettiner Entomologische Zeitung 98: 240–280. Annals of the Entomological Society of America 56: 470–472.
Mangan, R.L. 1979. Reproductive behavior of the cactus fly, Odon- Steyskal, G.C. 1965a. The third larval instar and puparium of
toloxozus longicornis, male territoriality and female guarding Odontoloxozus longicornis (Coquillett) (Diptera, Neriidae).
as adaptive strategies. Behavioral Ecology and Sociobiology 4: Annals of the Entomological Society of America 58: 936–937.
265–278. Steyskal, G.C. 1965b. Synonymy of the genera Antillonerius and
Mangan, R.L.  & D. Baldwin. 1986. A new cryptic species of Imrenerius. Proceedings of the Entomological Society of
Odontoloxozus (Neriidae: Diptera) from the cape region of Baja Washington 67: 60.
820
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
PSEUDOPOMYZIDAE (PSEUDOPOMYZID FLIES) 57
Matthias Buck and David K. McAlpine

Fig. 57.1. Female of Latheticomyia tricolor Wheeler, (MND, fig. 55.1, as Cypselosomatidae, crossvein bm-cu removed).

Diagnosis two katepisternal bristles; anepisternum usually bare, some-

times setulose with outstanding bristle; scutellum with two
Acalyptrate flies of small to medium size (body length to four (rarely five) pairs of marginal bristles, decreasing in
1.5–5.0 mm), legs relatively long in Latheticomyia (Fig. 1). size anteriorly, often with additional setulae on disc or margin;
Head with three to four pairs of reclinate or latero-reclinate prosternum anteriorly narrowed, precoxal bridge absent.
orbital bristles (first pair sometimes small), distinct vibrissae, Tibiae without preapical dorsal bristles, sometimes (Helo-
cruciate postocellar bristles, and usually with differentiated clusia) with scattered bristles. Wing (Fig. 4) clear to moder-
paraverticals (Figs. 2, 3). Face ventromedially desclerotized ately infuscated, without conspicuous markings; costa with
to various degree, facial carina setulose (setulae minute and subcostal break and humeral break or weakening; subcosta
often hard to discern in Pseudopomyza, seemingly absent incomplete, apically desclerotized and merging with R1;
in Heloclusia Malloch). Antenna porrect; first flagellomere cells bm and dm confluent in New World taxa; CuA2 (clos-
rounded or with dorsoapical angle, arista short pubescent. ing vein of anal cell) often reduced, A1+CuA2 sclerotized.
Thorax with at least four pairs of dorsocentral bristles, one to Abdomen with tergite 6 large in both sexes (Figs. 5, 6); female

821
822
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

o vt s i vt s
i vt s
orb s
poc s

pavt s pavt s

o vt s

vb

2 Latheticomyia
R1
R2+3

R4+5
3 Pseudopomyza (Rhinopomyzella)
M1

A1+CuA2
CuA1 tg 6

4 Latheticomyia

tg 5
sg 7

5 Latheticomyia
tg 6

ph

st 6 cerc
ej apod

sur syntgst
7+8

cerc phapod

epand

6 Latheticomyia

Figs. 57.2–6. Heads, wing, and terminalia: anterolateral view of head of (2) Latheticomyia tricolor Wheeler, (MND, figs. 4.108, 55.2) and
(3) Pseudopomyza (Rhinopomyzella) nigrimana (Hennig); dorsal view of wing (4) and lateral view of male terminalia (5) of L. tricolor,
(MND, figs. 55.3 (crossvein bm-cu removed) and 55.5); lateral view of female terminalia of (6) L. longiterebra Hennig, (South America,
MND, fig. 55.4). Figure 3 illustrated by A. Brenes.
Abbreviations: cerc, cercus; ej apod, ejaculatory apodeme; epand, epandrium; i vt s, inner vertical seta; orb s, orbital seta; o vt s, outer
vertical seta; pavt s, paravertical seta; ph, phallus; phapod, phallapodeme; poc s, postocellar seta; sg, segment; st, sternite; sur, surstylus;
syntgst, syntergosternite; tg, tergite; vb, vibrissa.
PSEUDOPOMYZIDAE (pseudopomyzid flies) 57 823

segment 7 developed as a sclerotized oviscape (Fig. 5); two deme basal of phallus (not on each side of phallus); female
spermathecae, sometimes of different sizes. segment 7 modified as sclerotized oviscape. The Pseudopo-
Pseudopomyzidae are easily distinguished from other myzidae were first recognized as a family and placed within
Neotropical acalyptrate families by the presence of three the Nerioidea by D.K.  McAlpine (1966). It retains many
to four pairs of reclinate orbital bristles (some Australasian groundplan characters of the Nerioidea and is in many
Pseudopomyza have only two pairs) in conjunction with respects the least derived clade within the superfamily. The
well-developed vibrissae and porrect antennae. Additional Pseudopomyzidae has been considered monophyletic by
diagnostic characters include an incomplete wing venation most authors (except Griffiths, 1972), but none of the auta-
with reduced subcosta and missing bm-cu (present in two Old pomorphies suggested in support of this hypothesis appear
World genera), as well as two costal breaks, and in the female very convincing (see J.F. McAlpine, 1989; Shatalkin, 1995;
a sclerotized oviscape. The genera Pseudopomyza and D.K. McAlpine, 1996; D.K. McAlpine & Shatalkin, 1998).
Pseudopomyzella superficially resemble certain Drosophili- Several supposedly autapomorphic characters are shared
dae or Ephydridae, from which they can readily be separated with other Nerioidea families (e.g., with Neriidae: porrect
by the characters mentioned. Latheticomyia is reminiscent of antennae with distally shifted arista, medially desclerotized
a miniature neriid, though Neriidae are usually significantly face, scapular bristles that are present in Longina Wiedemann
larger (5–15  mm), lack ocellar bristles, and usually lack and Cerantichir Enderlein; with Cypselosomatidae: reduced
vibrissae (if present they are short and hair- or spinelike), subcosta) or are of uncertain polarity (e.g., convergent
and have cell r4+5 strongly narrowed apically. postocellar bristles) depending on whether Diopsoidea or
Sphaeroceroidea are considered the sister group of the
Biology Nerioidea (D.K. McAlpine, 1996).
Given the difficulties of characterizing the Pseudopomyz-
The biology and immature stages of New World Pseudo- idae, it is not surprising that its sister group (if Pseudopo-
pomyzidae are unknown. Krivosheina (1979) found larvae myzidae are indeed monophyletic) also remains uncertain.
of the eastern Palearctic species Polypathomyia stackelbergi Various authors have tentatively suggested the Neriidae
Krivosheina under bark of rotten logs, and Frey (1952) (D.K.  McAlpine, 1996), the remainder of the Nerioidea
reported adults of the trans-Palearctic Pseudopomyza (sensu (D.K. McAlpine, 1966), or the Cypselosomatidae (Griffiths,
stricto) atrimana Meigen “swarming” over recently felled 1972; J.F. McAlpine, 1989) as sister groups, sometimes unit-
logs. In the Neotropical Region, adults of several Pseu- ing Pseudopomyzidae and Cypselosomatidae in a single
dopomyza (Rhinopomyzella) species are attracted to de- family (Griffiths, 1972; Prado, 1984; J.F. McAlpine, 1987,
caying vegetation, especially fallen trees, where they can 1989). D.K. McAlpine (1996) rejected the latter hypothesis
be collected in pan traps or by sweep net. A female of and argued that the similarities between both families are
the rare genus Pseudopomyzella was obtained from pan mostly due to symplesiomorphy.
traps in kitchen compost at the edge of primary rain forest Currently the family is composed of six extant genera
in Ecuador. Adults of Latheticomyia tricolor Wheeler and and one fossil genus, which were divided into three probably
L.  lineata Wheeler are attracted to banana baits (Wheeler, monophyletic groups by D.K. McAlpine (1996): (1) Latheti-
1956). Several Neotropical Latheticomyia species visit small comyia (New World) and Tenuia Malloch (Palearctic,
dung baits, especially when those are placed on logs or moss Oriental), (2)  Pseudopomyzella (Neotropical) and Eopseu-
near small streams (S.A.  Marshall, personal communica- dopomyza Hennig (Baltic amber), and (3)  Pseudopomyza
tion). Dung baits are more attractive to males than to females, (Neotropical, Palearctic, Oriental, Australasian), Heloclusia
and pan traps baited with dung sometimes exclusively yield (Chile), and Polypathomyia Krivosheina (eastern Palearc-
males of Latheticomyia. tic). The genus Pseudopomyza is here defined to include
five subgenera (D.K. McAlpine, 1994; Papp, 2005), one of
which, Rhinopomyzella Hennig, occurs in the New World.
Classification
The monotypic Oriental taxon Macalpinella Papp was de-
The Pseudopomyzidae belong in the well-defined super- scribed recently as a separate genus (Papp, 2005), but clear-
family Nerioidea, which is characterized by several striking ly represents a subgenus of Pseudopomyza. It shares with
synapomorphies of the male and female genitalia: epandrium Pseudopomyza (sensu lato) a similarly modified sternite  6
elongate with apical surstyli that are widely separated from (erroneously considered part of syntergosternite 7+8 by Papp,
the articulation point of the hypandrium near the epandrial 2005) and forms a monophyletic group with Rhinopomyzella
base; phallus borne on an elongate cone that is formed by and Pseudopomyza (sensu stricto) defined by a completely
a posterior extension of the hypandrial bridge dorsally and symmetrical, ringlike syntergosternite 7+8. More primitive
the elongate, rodlike pregonites (ventral arms of parameres members of Pseudopomyza (sensu lato) including the Aus-
of Shatalkin, 1995; gonopods of J.F. McAlpine, 1987, 1989; tralasian subgenera Apops McAlpine and Dete McAlpine as
sclerite “X” of Griffiths, 1972); postgonites (pregonites of well as some Australasian species of “Pseudopomyza (sensu
Hennig, 1969) shifted anteriorly, articulated to phallapo- stricto)” (e.g., P.  aristata (Harrison) from New Zealand)
824
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

possess an asymmetrical, laterally more or less open syn- Identification

tergosternite 7+8. The subgeneric placement of Pseu-
dopomyza (sensu stricto) species with asymmetrical A key to extant World genera was provided by Krivoshei-
syntergosternite is highly questionable and needs to be na (1979). The subgenera of Pseudopomyza (excluding
revisited within the context of a phylogenetic revision of Macalpinella) were keyed by D.K.  McAlpine (1994). The
the family. The monophyly of the Neotropical subgenus nine described New World species were treated by Wheeler
Rhinopomyzella, however, is clearly established by the (1956), Hennig (1969), and Malloch (1933). Shatalkin (1995)
autapomorphic fusion and strong dorsal arching of the keyed the Palearctic genera and species. The single fossil
pregonites. genus Eopseudopomyza was described by Hennig (1971).

Key to the genera of Pseudopomyzidae of the New World

1. Anepisternum with few setulae and posterior bristle; katepisternal bristle strongly curved forward;
acrostichal setulae in three rows on anterior half of scutum; four pairs of postsutural dorsocentral
bristles present; male pregenital sclerite (sternite 8) with two pairs of long bristles. . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudopomyzella Hennig
– Anepisternum bare; katepisternal bristle (posterior one if two are present) curved posteriorly;
acrostichal setulae in more than three rows on anterior half of scutum and/or only three pairs of
postsutural dorsocentral bristles present; male pregenital sclerite setulose, without outstanding
bristles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2. Four pairs of postsutural dorsocentral bristles present (Fig. 1); genal setulae arranged more or less
in single row (Fig. 2). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Latheticomyia Wheeler
– Three pairs of postsutural dorsocentral bristles present; genal setulae in more than one row or
scattered (Fig. 3). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3. Subvibrissal bristle greatly enlarged, subequal to vibrissa; costa with series of large, spaced anterior
bristles in second section; two dorsal katepisternal bristles present; tibiae with long dorsal bristles;
Chile. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Heloclusia Malloch]
– Subvibrissal bristle at most half as long as vibrissa (Fig. 3); costa without differentiated bristles in
second section; one dorsal katepisternal bristle present; tibiae without dorsal bristles. . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudopomyza (Rhinopomyzella) Hennig

Synopsis of the fauna demic subgenus Rhinopomyzella Hennig, which is

distributed throughout Central and South America from
At least 15 species are found in Central America, although Mexico to Bolivia and northern Argentina, including
knowledge of the fauna is likely to be incomplete. the Caribbean (Cuba, Puerto Rico, Dominican Repub-
lic, United States Virgin Islands, British Virgin Islands,
Latheticomyia Wheeler. This genus ranges from the Jamaica, Grenada, Trinidad and Tobago). The Central
southwestern United States through Central America to American fauna includes at least 10 species, most of
Bolivia and northern Argentina, including the Caribbe- which are undescribed. The two described species, P. ni-
an. The genus includes five described species (Wheeler, grimana (Hennig) and P.  albimana (Hennig) (Hennig,
1956; Hennig, 1969), two of which, L. infumata Wheeler 1969), both occur in Central America, including Costa
and L. tricolor, occur in Central America. The latter was Rica, but Central American specimens show more ex-
described from Arizona and Utah but ranges as far south tensive dark leg coloration than specimens from the type
as Honduras. A third species, L.  rotundicornis Hennig, localities in Brazil.
was described from Nearctic Mexico (Durango, 3000 m),
close to the boundary of the Neotropical region. At least Pseudopomyzella Hennig. The genus includes only
two undescribed species occur in Central America. one described species, P.  flava Hennig, from Ecuador
and Peru. Pseudopomyzella is also found in Costa Rica
Pseudopomyza Strobl. The widespread genus Pseu- and Venezuela, but the identity of the Central American
dopomyza is represented in the New World by the en- specimens has not been studied.
PSEUDOPOMYZIDAE (pseudopomyzid flies) 57 825

Literature cited

Frey, R. 1952. Über Chiropteromyza n. gen. und Pseudopomyza za. Proceedings of the Linnean Society of New South Wales
Strobl (Diptera Haplostomata). Notulae Entomologicae 32: 114: 181–187.
5–8. McAlpine, D.K. 1996. Relationships and classification of the
Griffiths, G.C.D. 1972. The phylogenetic classification of Diptera Pseudopomyzidae (Diptera: Nerioidea). Proceedings of the
Cyclorrhapha with special reference to the structure of the male Linnean Society of New South Wales 116: 223–232.
postabdomen. Series entomologica 8. Dr. W. Junk, The Hague, McAlpine, D.K. & A.I. Shatalkin. 1998. 3.8. Family Pseudo-
340 pp. pomyzidae, pp. 155–163. In Papp, L. & B. Darvas (editors).
Hennig, W. 1969. Neue Gattungen and Arten der Acalypteratae. Contributions to a manual of Palaearctic Diptera (with spe-
The Canadian Entomologist 101: 589–633. cial reference to flies of economic importance). Vol. 3, Higher
Hennig, W. 1971. Die Familien Pseudopomyzidae und Milichiidae Brachycera. Science Herald, Budapest, 880 pp.
im Baltischen Bernstein (Diptera: Cyclorrhapha). Stuttgarter McAlpine, J.F. 1987. 55. Cypselosomatidae, pp. 757–760. MND
Beiträge zur Naturkunde 233: 1–16. Volume 2.
Krivosheina, N.P. 1979. A new member of the family Pseudopo- McAlpine, J.F. 1989. 116. Phylogeny and classification of the
myzidae (Diptera) in the Palaearctic and the position of the fam- Muscomorpha, pp. 1397–1518. MND Volume 3.
ily in the system of the Diptera. Entomologiceskoye Obozrenie Papp, L. 2005. Some acalyptrate flies (Diptera) from Taiwan.
58: 179–189. (In Russian; English translation in Entomological Acta Zoologica Academiae Scientiarum Hungaricae 51:
Review 58: 106–113.) 187–213.
Malloch, J. R. 1933. Acalyptrata. (Helomyzidae, Trypetidae, Scio- Prado, A.P. do. 1984. Family Cypselosomatidae. In Papavero, N.
myzidae, Sapromyzidae, etc.), pp. 177–391. Diptera of Patago- (editor). A Catalogue of the Diptera of the Americas South of
nia and South Chile, Part VI – Fascicle 4. British Museum of the United States. Departamento de Zoologia, Secretaria da
Natural History, London. Agricultura, São Paulo, 2 pp.
McAlpine, D.K. 1966. Description and biology of an Australian Shatalkin, A.I. 1995. Palaearctic species of Pseudopomyzidae
species of Cypselosomatidae (Diptera), with a discussion (Diptera). Russian Entomological Journal 3(3–4): 129–145.
of family relationships. Australian Journal of Zoology 14: (1994)
673–685. Wheeler, M.R. 1956. Latheticomyia, a new genus of acalyptrate
McAlpine, D.K. 1994. A new Australian species of pseudopomyz- flies of uncertain family relationship. Proceedings of the
id fly (Diptera: Nerioidea) and the subgenera of Pseudopomy- United States National Museum 106: 305–314.
826
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
TANYPEZIDAE (TANYPEZID FLIES) 58
Kyle Apigian and F. Christian Thompson

Fig. 58.1. Male of Neotanypeza elegans (Wiedemann), (illustrated by T. Litwak).

Diagnosis Tanypezids are occasionally misidentified as Dolichopo-

didae; however, close inspection will reveal the typical
Small to medium-sized (body length 5.0–11.5  mm), schizophoran ptilinum, lunule, and other characteristics that
slender flies, with large head and long legs (Fig.  1). Head readily separate Tanypezidae from Dolichopodidae.
large, hemispherical; compound eye covering most of lateral
aspect; ptilinal suture and lunule distinct; antennal pedi- Biology
cel without dorsal suture. Thorax without greater ampulla;
anepisternum with prominent bristle near posterodorsal mar- Little is known about the ecology and behavior of
gin; katepisternum setose, without bristles; legs long, slender, adults except that they commonly sit on leaves. The lar-
yellow; wing hyaline, vein R1 setose dorsally; veins R4+5 and vae of one species are apparently saprophagous, as dem-
M converging apically. With silvery tomentose maculae on onstrated by Foote (1970), who reared Tanypeza from
head and thorax. eggs to third instar maggots on decaying watermelon rind

827
828
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

and pulp. Chandler (1975) suggested that Tanypeza larvae lein’s classification to three genera, Tanypeza, Neotanypeza,
develop in rotten wood. and Scipopeza, with the genus Neotanypeza being divided
into four subgenera, Neotanypeza, Polphopeza Enderlein,
Tritanypeza Enderlein, and Tripolphopeza Enderlein. These
Classification subgenera appear to be only phenetic concepts based on vari-
ous permutations of the number of orbital and dorsocentral
Rondani (1856) first recognized Tanypeza as being a dis- bristles.
tinct group or clade. In the past, the Tanypezidae have been
considered part of the Ulidiidae (Hendel, 1903, as Ortalidae)
or within the superfamily Tephritoidea (Enderlein, 1913, Identification
1936, as Otitoidea). Most recent authors place the Tanypezi-
dae as a distinct family within the superfamily Diopsoidea The only keys available are those of Enderlein (1913,
(=Nothyboidea of Hennig, 1973). However, McAlpine 1936). Care must be used with the older literature, however,
(1997) proposed placing the Tanypezidae within the Neri- as different terminology was employed for the bristles of the
oidea. The sister group to Tanypezidae is Strongylophthal- head. Enderlein counted only the “fronto-orbital” bristles,
myiidae (Steyskal, 1987); the only remaining question is that is, the orbital bristles between the antenna and the ocel-
whether they should be considered a single family (Griffiths, lar triangle, whereas modern authors also count as an orbital
1972; McAlpine, 1997) or separate families (most authors). bristle the large bristle posterior to the ocellar triangle and
Roháček (1998) provides the best current treatment of the at the posterior dorsomedial corner of the eye. A revision
family. based on re-examination of all types is now being prepared
The family currently consists of three genera: Tanypeza (O. Lonsdale, in preparation).
(Holarctic) and Neotanypeza and Scipopeza (Neotropical). Species determination in Tanypezidae is made using a va-
Hendel proposed Neotanypeza as a subgenus for the South riety of characters, including the number of orbital and tho-
American species, as they lack ocellar and postvertical racic bristles, the color of the palpus and first flagellomere,
bristles. Enderlein (1913, 1936) elevated Neotanypeza to the shape of the first flagellomere, the color of the legs, and
generic status and described a series of new genera based the extent and shape of the silvery tomentum posterior to the
on the number of orbital and dorsocentral bristles. Hennig ocelli. Wing markings and patterns are also important char-
(1936), followed by Steyskal (1967, 1987), reduced Ender- acters in determining species.

Key to the genera of Tanypezidae of the world

1. Tarsomere 1 of hind leg not produced ventrally as distinct tubercle at base; three orbital bristles
present; one dorsocentral bristle present; one postpronotal bristle present, pre-alar bristle absent;
Holarctic Region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Tanypeza Fallén]
– Tarsomere 1 of hind leg produced into distinct basoventral tubercle (Fig. 1); other characters
various; Neotropical Region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2. Frons with single orbital bristle (Fig. 8); postpronotal bristle absent; pre-alar bristle absent; one
dorsocentral bristle present (Fig. 5); Ecuador, Peru. . . . . . . . . . . . . . . . . . . . [Scipopeza Enderlein]
– Frons with two or more orbital bristles (Figs.  6–7, 9–10); postpronotal bristle present; one to
two pre-alar (supra-alar) bristles present; usually two or more dorsocentral bristles present
(Figs. 2–4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neotanypeza Hendel
TANYPEZIDAE (tanypezid flies) 58 829

2 Neotanypeza 3 Neotanypeza

pprn s

dc s

4 Neotanypeza 5 Scipopeza

Figs. 58.2–5. Thoraces: dorsal view of (2) Neotanypeza elegans (Wiedemann); (3) N. flavicalx (Enderlein); (4) N. ornatipes (Bigot); and
(5) Scipopeza grandis Enderlein. Figures 2–5 illustrated by T. Litwak.
Abbreviations: dc s, dorsocentral seta; pprn s, postpronotal seta.
830
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

6 Neotanypeza 7 Neotanypeza

8 Scipopeza

9 Neotanypeza 10 Neotanypeza

Figs. 58.6–10. Heads, anterodorsal view of (6) Neotanypeza apicalis (Wiedemann); (7) Neotanypeza sp.; (8) Scipopeza grandis Enderlein;
(9) N. elegans (Wiedemann); and (10) N. ornatipes (Bigot). Figures 6–10 illustrated by T. Litwak.
TANYPEZIDAE (tanypezid flies) 58 831

Synopsis of the fauna are absent from the Chilean subregion. There are two
named species and a couple of undescribed species in
Neotanypeza Hendel. Sixteen species of this genus Costa Rica. The most recent key is that of Enderlein
are found from southern Mexico to Argentina, but they (1936).

Literature cited

Chandler, P.J. 1975. Notes on the British status of three unusual aca- der Zoologie. Eine Naturgeschichte der Stämme des Tierreiches.
lypterate flies (Diptera). Proceedings of the British Entomologi- IV. Band: Arthropoda — 2. Hälfte: Insecta. 2. Teil: Spezielles.
cal and Natural History Society 8: 66–72. W. De Gryter, Berlin.
Enderlein, G. 1913. Dipterologische Studien II. Zur Kenntnis der McAlpine, D.K. 1997. Gobryidae, a new family of Acalyptrate flies
Tanypezinen. Zoologischer Anzeiger 42: 224–229. (Diptera: Diopsoidea), and a discussion of relationships of the Di-
Enderlein, G. 1936. Klassifikation der Tanypeziden. Deutsche Ento- opsoid families. Notes of the Australian Museum 49: 167–194.
mologische Zeitschrift 1936: 39–47. Roháček, J. 1998. Family Tanypezidae, pp. 165–171. In Papp, L. &
Foote, B.A. 1970. The larvae of Tanypeza longimana (Diptera: B. Darvas (editors). Contributions to a manual of Palaearctic
Tanypezidae). Annals of the Entomological Society of America Diptera (with special reference to flies of economic impor-
63: 235–238. tance). Vol. 3, Higher Brachycera. Science Herald, Budapest,
Griffiths, G.C.D. 1972. The phylogenetic classification of Diptera 880 pp.
Cyclorrhapha, with special reference to the structure of the male Rondani, C. 1856. Dipterologiae Italicae prodromus. Vol: 1. Genera
postabdomen. Series entomologica 8. Dr. W. Junk, The Hague, italica ordinis dipterorum ordinatim disposita et distincta et in
340 pp. familias et stirpes aggregata. A.  Stoschi, Parmae [= Parma],
Hendel, F. 1903. Ueber die systematische Stellung von Tanypeza 226 + [2] pp.
Fall. (Dipt.). Wiener Entomologische Zeitung 22: 201–205. Steyskal, G.C. 1967. Family Tanypezidae. In Papavero, N. (editor).
Hennig, W. 1936. Beiträge zur Kenntnis des Kopulationsapparates A Catalogue of the Diptera of the Americas South of the United
und der Systematik der Tanypeziden (Dipt., Acalypt.). Deutsche States. Departamento de Zoologia, Secretaria da Agricultura,
Entomologische Zeitschrift 1936: 27–38. São Paulo, 4 pp.
Hennig, W. 1973. Diptera (Zweiflügler). Vol. 4(2), pp. 1–337. In Steyskal, G.C. 1987. Family Tanypezidae, pp. 773–776. MND, Vol-
Helmcke, J.-G., D.  Starck  & H. Wermuth (editors). Handbuch ume 2.
832
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
SOMATIIDAE (SOMATIID FLIES) 59
Owen Lonsdale and David K. McAlpine

Fig. 59.1. Female of Somatia aestiva (Fabricius), (illustrated by O. Lonsdale).

Diagnosis central, one prescutellar acrostichal, two or three scutellars,

one anepisternal. Midtibia with three ventroapical bristles.
Moderately small (body length 3.5–5.0 mm), robust, shin- Wing without distinct alula or anal lobe; cells bm and cup
ing, usually black and yellow flies (Fig.  1). Ptilinal suture long and closed; subcostal break distinct. Abdomen broad
extending onto gena (Fig. 2); frons produced anteromedially. with tergites  1–6 wide and strongly convex, covering re-
Pedicel slit apically; first flagellomere elongate and setulose; maining segments (Fig.  3); sternites highly reduced. Male
arista long bipectinate. Fronto-orbital plate setulose; postocellar with epandrium thin, semicircular and setose; fused with
bristles small and convergent; outer vertical bristle absent; surstyli to form ring (Figs.  4–5). Internal genitalia highly
vibrissa short and slender. Pronotum well developed and reduced: only distiphallus, basiphallus, phallapodeme, and
“necklike”. Scutum convex with central transverse suture. large ejaculatory apodeme (Fig.  6) remaining. Distiphallus
Scutellum convex and with short pile. Coxae remote from long, exposed below abdomen. Female with annular tergos-
abdomen, with intervening postmetacoxal bridge. Thoracic ternite 7 (Fig. 7). Cerci small and truncated. Two papillose
chaetotaxy reduced: two posterior notopleurals, one dorso- spermathecae present (Fig. 8).

833
834
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Syringogastridae are most similar to Somatiidae, also similar venation and head morphology, but the abdomen is
having an enlarged pronotum, postmetacoxal bridge, bipec- slender, the anal region of the wing is unreduced, there are
tinate arista, narrowed wing, and reduced cephalic chaeto- no vibrissae, the pronotum is not enlarged, and the arista is
taxy, but the abdomen is petiolate, the hind femur is enlarged pubescent. Other taxa with a widened abdomen can be con-
and spinose, the vibrissae and postverticals are absent, and fused for somatiids, particularly Richardiidae, which can be
the head is relatively spherical in outline. Psilids also have a similarly patterned in some cases, but none of these have the
dorsal seam on the pedicel, reduced thoracic chaetotaxy, and characteristic venation or chaetotaxy unique to the Somatiidae.

poc s i vt s
tg 1

tg 2

st 1
st 2
tg 3
st 3

st 4 tg 4

st 5
tg 5
ph st 6

tg 6
cerc
tg 7

2 Somatia 3 Somatia

epand

cerc
cerc epand

tg 7
bph

phapod
st 6

sur

sur pgt distph

4 Somatia 5 Somatia

6 Somatia

Figs. 59.2–6. Head, abdomen, and male terminalia: anterolateral view of head (2), ventral view of abdomen of male (3), lateral view of male
terminalia (4), posterior view of male terminalia (5), and lateral view of ejaculatory apodeme (6) of Somatia aestiva (Fabricius). Figures 2–6
illustrated by O. Lonsdale.
Abbreviations: bph, basiphallus; cerc, cercus; distph, distiphallus; epand, epandrium; i vt s, inner vertical seta; pgt, postgonite; ph, phallus;
phapod, phallapodeme; poc s, postocellar seta; st, sternite; sur, surstylus; tg, tergite.
SOMATIIDAE (somatiid flies) 59 835

The pigmentation of Somatia in combination with a shiny ous hypotheses of relationship, but was unable to infer a sis-
frontal plate is also reminiscent of some Chloropidae. ter group or superfamily placement.

Biology Identification
Almost nothing is known of somatiid biology, although Somatia, the only genus, includes seven described spe-
adults have been observed on the undersides of leaves of ri- cies. These were keyed by Steyskal (1968) and catalogued
parian Leguminosae in Bolivia, and one adult has been ob- by Steyskal (1970).
served feeding on a dead caterpillar (S.A. Marshall, personal
communication). Immature stages are unknown. Synopsis of the fauna
Classification
Somatia Schiner. This genus is endemic to the Neotropi-
Somatiidae have been associated with Periscelididiae, cal Region. Two species are known from Costa Rica, S. aes-
Richardiidae, or Heleomyzidae at different times (Griffiths, tiva (Fabricius) and S. schildi Steyskal, although S. sophiston
1972). More recently, the family was placed in the Diop- Steyskal is also likely found there because it is known from
soidea as the sister group to the Psilidae by J.F. McAlpine Guatemala, Panama, and Venezuela (Steyskal, 1970). Only
(1989). D.K. McAlpine (1997) considered evidence for vari- S. aestiva is encountered frequently.

spmth
cerc
hyprct
tg 8
st 8

tgst 7

7 Somatia 8 Somatia

Figs. 59.7–8. Female abdomen and genitalia: ventral view of terminal segments of abdomen, positions of spermathecae shown by dotted
lines (7), and spermathecae (8) of Somatia aestiva (Fabricius). Figures 7–8 illustrated by O. Lonsdale.
Abbreviations: cerc, cercus; hyprct, hypoproct; spmth, spermatheca; st, sternite; tg, tergite; tgst 7, tergosternite 7.

Literature cited

Griffiths, G.C.D. 1972. The phylogenetic classification of Diptera Steyskal, G.C. 1968. The genus Somatia Schiner, II. (Diptera: So-
Cyclorrhapha, with special reference to the structure of the male matiidae). Proceedings of the Entomological Society of Wash-
postabdomen. Series entomologica 8. Dr. W. Junk, The Hague, ington 70: 255–257.
340 pp. Steyskal, G.C. 1970. 94. Family Somatiidae. In Papavero, N. (edi-
McAlpine, D.K. 1997. Gobryidae, a new family of acalyptrate flies
tor). A Catalogue of the Diptera of the Americas South of the
(Diptera: Diospsoidea), and a discussion of relationships of
the diopsoid families. Records of the Australian Museum 49: United States. Departamento de Zoologia, Secretaria da Agri-
167–194. cultura, São Paulo, 2 pp.
McAlpine, J.F. 1989. Phylogeny and classification of the Musco-
morpha, pp. 1397–1518. MND, Volume 3.
836
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
PSILIDAE (RUST FLIES, PSILID FLIES) 60
Matthias Buck

Fig. 60.1. Female of Psila (Xenopsila) collaris Loew, (Nearctic, MND, fig. 60.1).

Diagnosis subcosta vanishes before curving anteriorly to C. Male abdo-

men with well-developed symmetrical tergite and sternite 6,
Small to medium-sized flies (body length 3–12 mm), with dorsal pregenital sclerite (possibly tergite  7) present (Psili-
more or less elongated body (Fig.  1). Head chaetotaxy nae) or absent (Chylizinae). Surstyli present (Chylizinae) or
including zero to two fronto-orbital bristles, ocellar bristles, absent (Psilinae). Phallapodeme connected to hypandrium,
inner and outer vertical bristles (the latter duplicated in in Psilinae forming large phallapodemic plate (Buck & Mar-
many Chyliza and some Psila), divergent postocellar bristles shall, 2006a). Phallus usually short, but long with bifid apex
(sometimes reduced), and no vibrissa. Antenna with elon- in Psila (Xenopsila). Ejaculatory apodeme small or absent.
gated first flagellomere in Loxocera; arista short- to medi- Female with elongate ovipositor; in Loxocera laterally com-
um- pubescent; pedicel with distinct dorsal seam. Thorax pressed and coriaceous, tergite 10 and cerci fused in subgenus
relatively sparsely bristled, with one to two dorsocentral, Imantimyia (Fig.  6). Spermathecae desclerotized, ventral
one notopleural, one supra-alar, one postalar, and no pleural receptacle present.
bristles. Laterotergite with prominent callus in Chyliza. Legs Psilidae can be distinguished from all other Neotropical
normal to elongated, without bristles. Wing with subcostal break; acalyptrate families by the presence of a subcostal break,

837
838
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

incomplete subcosta, complete cell cup, and the absence of Loew (Buck & Marshall, 2006a). Larvae of several Nearctic
vibrissae and pleural bristles. The two families that come and Palearctic Chyliza are phytophagous on trees or herba-
closest to this diagnosis, Somatiidae (with weak vibrissae and ceous plants (Ferrar, 1987), live in the pitch mass of pine
an outstanding but fairly inconspicuous anepisternal bristle) trees (Teskey, 1976), under the bark of trees infested by
and Syringogastridae (without subcostal break), can also be buprestid beetles (Gates et al., 2006), and in galls induced
separated based on the following characters: a plumose by Cecidomyiidae (Sugiura  & Yamazaki, 2006). In some
arista (pubescent in Psilidae) and enlarged pronotum (small cases, the larvae apparently induce galls in the host plant.
in Psilidae). Syringogastridae is furthermore distinct because Infestation of trees usually starts from wounds, with larvae
of the enlarged, spinose hind femora and the lack of inner tunnelling into healthy tissue between the bark and sap wood
vertical bristles. (Lyneborg, 1987). Larvae of the North American Loxocera
cylindrica Say attack shoots of Carex (Valley et al., 1969),
while the European L. albiseta (Schrank) apparently devel-
Biology ops in Juncus stems (Ferrar, 1987). Loxocera females have
a specialized, laterally compressed ovipositor that serves to
There are no biological observations on Neotropical insert eggs into narrow crevices (i.e., probably between the
Psilidae. Eggs have been described and illustrated for three leaf sheaths and stems of their host plants). Larvae of the
Neotropical species: Chyliza consanguinea Schiner (Albu- genus Psila live in herbaceous plants, with some Holarctic
querque, 1957), Loxocera ignyodactyla Buck, and L. collaris species being pests of cultivated plants (Ferrar, 1987).

bm

2 Chyliza

ltg

4 Chyliza

3 Loxocera 5 Chyliza

6 Loxocera

Figs. 60.2–6. Wings, thorax, hind femur and female terminalia: dorsal view of wing of (2) Chyliza apicalis Loew, (Nearctic, Mexico, MND,
fig. 60.5) and (3) Loxocera (Imantimyia) ignyodactyla Buck, (Buck & Marshall, 2006a, fig. 55); dorsal view of thorax of (4) Chyliza sp.;
ventral view of hind femur of (5) Chyliza sp.; lateral view of female terminalia from segment 8 onward of (6) L. (Imantimyia) ignyodactyla,
(Buck & Marshall, 2006a, fig. 64). Figures 4 and 5 illustrated by A. Cormier.
Abbreviation: ltg, laterotergite.
PSILIDAE (rust flies, psilid flies) 60 839

Classification cap with regularly placed, large aeropyles and wide, longi-
tudinal, chorionic air canals. Six subgenera are recognized
The family Psilidae is a relatively small group of flies that in Psila (sensu lato) (Buck  & Marshall, 2006b) but only
is most diverse in the Old World and the Nearctic Region. Psila (sensu stricto) and Xenopsila Buck (=Pseudopsila
Only a few species occur in the Neotropical Region. The auct., nec  Johnson, in part) occur in the New World. The
family is currently divided into three subfamilies: Psilinae, subgenera of Psila appear to be reasonably well charac-
Chylizinae, and Belobackenbardiinae. The last was described terized by male genitalic characters, but the monophyly of
only recently and appears to be restricted to the Afrotropical some groups, especially Psila (sensu stricto), remains to be
Region (Shatalkin, 2002). The monogeneric Chylizinae is established, and the phylogenetic relationships within the
well defined by a protuberant laterotergite, the presence of a genus have never been studied. The second Psilinae genus,
postmetacoxal bridge, a shortened anal cell, and the absence Loxocera (sensu lato) is defined by a coriaceous and com-
of abdominal spiracle 7 in both sexes. The peculiar, shallow, pressed female ovipositor. The subgenera and species groups
microtrichose, ventrodistal, oval pit of the hind femur of of Loxocera were recently redefined and their phylogenetic
Chyliza (Fig. 5), which has apparently not been mentioned relationships clarified (Buck  & Marshall, 2006a). Three
in the previous literature, might be another synapomorphy subgenera are currently included in the genus: Loxocera
for the genus, even though the pit is absent (possibly second- (sensu stricto), Tropeopsila Shatalkin from the Palearctic
arily lost) in at least one South American species. Charac- and Oriental, and Imantimyia Frey from the Holarctic and
ters considered autapomorphic for the Psilinae include a northern parts of the Neotropical Region; the status of the
receding face and the absence of surstyli (Steyskal, 1987; Afrotropical taxon Loxocerosoma Verbeke remains unclear.
Shatalkin, 2002). The two subordinate clades of Psilinae,
Psila (sensu lato) and Loxocera (sensu lato), have recently Identification
been redefined (Buck  & Marshall, 2006a,  b), resulting in
the removal of the Oriental subgenus Asiopsila Shatalkin The Psilidae are represented by mostly the same genera
from Loxocera to Psila. Psila (sensu lato) is defined by a and subgenera in both Neotropical and Nearctic Regions.
peculiar egg morphology, including a disklike micropylar The Nearctic genera were keyed by Steyskal (1987).

Key to the genera and subgenera of Psilidae of the New World

Adults
1. Laterotergite strongly convex, protruding (Fig.  4); postmetacoxal bridge developed; cell
cup distinctly shorter than bm (Fig.  2); ventral surface of hind femur usually with shallow,
microtrichose, oval pit near apex (Fig. 5); face more or less vertical, not receding towards margin
of subcranial cavity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chyliza Fallén
– Laterotergite slightly convex, not protruding; postmetacoxal bridge absent; cell cup almost as
long as bm (Fig. 3); ventral surface of hind femur without oval pit; face slightly to strongly
receding towards margin of subcranial cavity. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2. First flagellomere greatly elongated, over 7 times as long as wide in lateral view; posteroventral
corner of anepisternum with patch of dense, ventrally directed hairlike setae in New World
species; ventral surface of hind femur with elongate patch of dense, feltlike microtrichia near
apex; ovipositor laterally compressed and coriaceous (Fig. 6) . . . . . . Loxocera (Imantimyia) Frey
– First flagellomere at most slightly elongated, 1.5–4.0 times as long as wide; posteroventral corner
of anepisternum with sparse setae; ventral surface of hind femur without feltlike microtrichia near
apex; ovipositor simple; Psila Meigen. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3. �����������������������������������������������������������������������������������������������������
First flagellomere�����������������������������������������������������������������������������������
shorter, 1.5–2.1 times as long as wide, if longer then tergites 2–4 ���������������������������
setose to lat-
eral margin; male sternite 6 rectangular; asetose pregenital sclerite (possibly tergite 7) developed
as small median sclerite, closely associated with epandrium but usually separate from it; phallus
short, with simple apex; median portion of hypandrium present, forming distinct plate below
phallapodemic sclerite.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Psila (Psila) Meigen
– First flagellomere longer, 2.5–4.0 times as long as wide; tergites 2–4 conspicuously bare laterally;
male sternite 6 deeply emarginate posteriorly, (almost) bipartite; asetose pregenital sclerite (possibly
tergite  7) divided into two lateral portions, fused to epandrium; phallus long, with bifid apex;
median portion of hypandrium absent; Nearctic Region. . . . . . . . . . . . . [Psila (Xenopsila) Buck]
840
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

aero

7 Psila 10 Chyliza
mic

14 Loxocera
mic pole

long rg

p pole

8 Psila

11 Chyliza
long rg

9 Psila 12 Chyliza 13 Loxocera 15 Loxocera

Figs. 60.7–15. Eggs: lateral view of micropylar cap (7), lateral view of egg (8), and fine structure of chorion (9), showing reticulate pattern of
inner surface of air canals within longitudinal ridges, of Psila (Xenopsila) lateralis Loew, (Nearctic, Buck & Marshall, 2006b, figs. 16–18);
lateral view of micropylar pole (10), lateral view of egg (11), and surface structure of chorion (12) of Chyliza notata Loew, (Nearctic, Buck &
Marshall, 2006b, figs. 20, 21, 19); surface structure of chorion (13), frontal view of micropylar area (14), and lateral view of egg (15) of
Loxocera (Imantimyia) ignyodactyla Buck, (Buck & Marshall, 2006a, figs. 83, 81, 82).
Abbreviations: aero, aeropyle; long rg, longitudinal ridge; mic, micropyle; mic pole, micropylar pole; p pole, posterior pole.
PSILIDAE (rust flies, psilid flies) 60 841

Eggs
1. Egg with disklike micropylar cap, bearing 6–10 regularly placed, large aeropyles around
its circumference (Fig.  7:  aero); egg surface with coarse, broadly rounded longitudinal ridges
(Fig. 8), sometimes connected by short transverse ridges, each ridge with wide internal air
canal; air canals with polygonal reticulation on inner surface (visible in slide-mounted specimens)
(Fig. 9); egg relatively stout (Fig. 8) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Psila Meigen
– Egg without clearly defined micropylar cap; micropylar pole usually without large aeropyles
(Figs. 10, 14), but if so then aeropyles irregularly scattered around micropyle; egg surface either
even, usually with polygonal reticulation (Fig. 13), or with fine polygonal or longitudinal ridging;
in this case ridges narrow, often thin and lamellar (Fig. 12), sometimes low and ripplelike; chorion
without wide, microscopically visible, internal air canals; egg shape various (Figs. 11, 15). . . . 2

2. Micropylar pole simple, truncate or obtusely rounded (Figs.  14, 15); chorion surface either
unridged (Fig.  15), with or without polygonal reticulation (Fig.  13), or with fine polygonal or
longitudinal ridging; ridges either low and ripplelike or higher but never lamellate; egg long and
slender (Fig. 15). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Loxocera Meigen
– Micropylar pole with corona of small, club-shaped, solid processes (Fig. 10); chorion with fine
longitudinal ridging, connected by short transverse ridges (Fig. 11); longitudinal ridges sharply
defined and lamellate (Fig. 12); egg stout, noticeably arched (Fig. 11). . . . . . . . . . . Chyliza Fallén

Synopsis of the fauna the Neotropical Region (Buck & Marshall, 2006a). All New
World species belong to the L.  (Imantimyia) achaeta
Eleven or twelve species of this family are found in species group, and two are found in the Neotropical Region.
Central America. The transcontinental Nearctic L. collaris Loew (=L. califor-
nica Capelle) ranges as far south as Durango, Mexico, at
Chyliza Fallén. This is the most diverse psilid genus in the boundary between the Nearctic and Neotropical Regions
the Neotropical Region and the only one that also occurs (Buck & Marshall, 2006a). The only species endemic to the
in South America. Chyliza is distributed through all bio- Neotropical Region, L.  ignyodactyla Buck, is known only
geographical regions, but has not yet been formally recorded from elevations around 2000 m in Puntarenas Province, Costa
from Central America south of Mexico. The four Mexican Rica (Buck  & Marshall, 2006a). It is characterized by a
species were keyed by Melander (1920). Three of these conspicuous ventral process on the male hind femur, a
species were described or recorded from the Neotropical feature otherwise unknown in the genus.
part of Mexico (C. enthea Giglio-Tos, Veracruz; C. varipes
Wulp, Guerrero and Tabasco). Giglio-Tos’s (1895) record of Psila Meigen. Also known from all biogeographical regions
the eastern Nearctic Chyliza apicalis Loew from Veracruz (introduced in New Zealand), this genus only reaches the
needs verification. Chyliza bigoti Melander was described northernmost parts of the Neotropical Region in Mexico.
from “Mexico” without reference to any region or locality. It has not yet been recorded from Central America south
Three Brazilian species were (re-)described by Albuquerque of Mexico, but its occurrence there is to be expected. Psila
(1957). The Central American fauna south of Mexico consists (sensu stricto) exigua Wulp was described from Guerrero,
of at least four species, all of which are undescribed. and one undescribed species occurs in Chiapas. The former
is included in Melander’s (1920) key to Nearctic species. The
Loxocera Meigen. This genus is distributed through all subgenus Xenopsila has not yet been recorded from south of
biogeographical regions but was only recently recorded from the United States.
842
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Literature cited

Albuquerque, D. de O. 1957. Fauna do Distrito Federal. XXXVII — Accademia delle Scienze di Torino, Classe di Scienze Fisiche,
Redescrição de Chyliza consanguinea Schiner, 1858 e descrição Matematiche e Naturali 45: 1–74.)
de duas espécies novas (Diptera — Psilidae). Boletim do Museu Lyneborg, L. 1987. On the life history of Chyliza annulipes Mac-
Nacional, N.S., Zoologia 162: 1–11. quart, 1835 (Diptera: Psilidae). Entomologiske Meddelelser 55:
Buck, M. & S.A. Marshall. 2006a. Revision of New World Loxo- 27–29.
cera Meigen (Diptera: Psilidae), with phylogenetic redefinition Melander, A.L. 1920. Synopsis of the dipterous family Psilidae.
of Holarctic subgenera and species groups. European Journal of
Psyche 27: 91–101.
Entomology 103: 193–219.
Buck, M.  & S.A. Marshall. 2006b. The identity of Pseudopsila, Shatalkin, A.I. 2002. Afrotropical Psilidae (Diptera). 1. Genera
description of a new subgenus of Psila, and redefinition of Psila Belobackenbardia gen.  n. and Psila Meigen, 1803. Russian
sensu lato (Diptera: Psilidae). European Journal of Entomology Entomological Journal 10: 417–424. (2001)
103: 183–192. Steyskal, G.C. 1987. 60. Psilidae, pp. 781–784. MND, Volume 2.
Ferrar, P. 1987. A guide to the breeding habits and immature stages Sugiura, S.  & K. Yamazaki. 2006. First record of a psilid fly
of Diptera Cyclorrhapha. Entomonograph 8(1), E.J.  Brill  / feeding on cecidomyiid galls. The Canadian Entomologist 138:
Scandinavian Science Press, Leiden, Copenhagen, 478 pp. 235–237.
Gates, M.W., H. Liu, L.S.  Bauer,  & M.E.  Schauff. 2006. A new Teskey, H.J. 1976. Diptera larvae associated with trees in North
species of Pediobius (Hymenoptera: Eulophidae) parasitizing America. Memoirs of the Entomological Society of Canada 100:
Chyliza apicalis (Diptera: Psilidae) in ash trees attacked by
53 pp.
Agrilus planipennis (Coleoptera: Buprestidae). The Great Lakes
Entomologist 38: 76–82. (2005) Valley, K., T. Wearsch, & B.A. Foote. 1969. Larval feeding habits
Giglio-Tos, E. 1895. Ditteri del Messico, Parte 4. Carlo Clausen, of certain Chloropidae (Diptera). Proceedings of the Entomo-
Torino, 74  pp. (Also published 1896 in Memorie della Reale logical Society of Washington 71: 29–34.
SYRINGOGASTRIDAE (syringogastrid flies) 61
Stephen A. Marshall and Matthias Buck

Fig. 61.1. Male of Syringogaster brunnea Cresson, (Zumbado, 2006, p. 139).

Diagnosis anteroventral and posteroventral row of stout, short spinelike

bristles. Scutum with distinct supra-alar carina, metathorax
Small (body length 4–6 mm), antlike flies with relatively with large, cylindrical posterior extension, postmetacoxal
large, spherical head separated from mesothorax by long, bridge high. Abdomen distinctly petiolate, with first three
collarlike prothorax (Fig. 1). Chaetotaxy reduced, head bris- or four segments fused (Fig. 3). Male with fully developed
tles limited to outer verticals and (usually) ocellar bristles; and symmetrical tergite 6. Synsternite 7+8 forming complete
thorax only with postalar and one pair of scutellar bristles. ring separate from epandrium, which bears surstyli and cer-
Pedicel with dorsal seam, arista bipectinate. Wing partially ci. Hypandrium with prominent, setose, ventrolateral lobes;
infuscated or banded, constricted basally through reduction hypandrial bridge usually developed; distiphallus large and
of anal part of wing, including loss of alula. Costa without complex, usually curled up the left side of the postabdomen
breaks; subcosta incomplete, more or less coalescing with R1 (Fig.  2). Female lacking oviscape; apical segments form
before costa; cell cup complete. Forelegs short and widely short telescoping ovipositor.
separated from midlegs; hind femur swollen and with

843
844
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

ej apod

phapod
cerc

sur
hypd

anterior part of
pregt
hypd arm
posterior part of
hypd arm

distph
pgt

2 Syringogaster

syntg 1-3

3 Syringogaster

Figs. 61.2–3. Abdomen and male terminalia of Syringogaster: lateral view of male terminalia of (2) Syringogaster brunneina Marshall &
Buck, (Marshall et al., 2009, fig. 21); lateral view of male abdomen of (3) S. brunneina, (Marshall et al., 2009, fig. 25).
Abbreviations: cerc, cercus; distph, distiphallus; ej apod, ejaculatory apodeme; hypd, hypandrium; hypd arm, hypandrial arm; phapod,
phallapodeme; pgt, postgonite; pregt, pregonite; sur, surstylus; syntg, syntergite.
SYRINGOGASTRIDAE (syringogastrid flies) 61 845

Syringogaster, the only genus in the family Syringo- Syringogastridae (Prado, 1969), most authors (Curran, 1934;
gastridae, is superficially similar in both habitus and habits Papavero, 1964; Hennig, 1958) considered Syringogaster to
to some other ant mimics, especially small Micropezidae in be similar to, or even part of, the family Megamerinidae.
the genera Metasphen Frey and Globopeza Marshall. Syrin- Feijen (1989) made a strong case for a sister group relation-
gogastrids and micropezids, however, are not closely related ship between the Syringogastridae and the Centrioncidae,
as indicated by details of wing venation, chaetotaxy, and ab- a group considered by all other workers (McAlpine, 1997;
dominal structure. Syringogastrids are easily distinguished Meier & Hilger, 2000; Baker et al., 2001; Meier  & Baker,
from Psilidae (the family in which they were originally 2002; Kotrba, 2004; Marshall et al., 2009) to be part of the
described by Cresson, 1912), and most other Neotropical Diopsidae. Neither Diopsidae nor Megamerinidae occur in
flies, by the long collarlike pronotum, the swollen ventrally the Neotropical Region.
spinose hind femur, the greatly reduced head chaetotaxy, the
basally narrowed wing, and the antlike body shape. Identification

Biology The species of Syringogaster have recently been revised

(Marshall et al., 2009), more than doubling the number of
Syringogastridae occur in lowland tropical forests, usu- known species from 9 to 20. Previously, Prado (1969) pre-
ally below 800  m (in Costa Rica rarely up to 1400  m). sented a key to the six species of Syringogaster occurring
Adults are uncommon and most often seen on large leaves in Brazil and provided notes on three other putative Syrin-
or similar surfaces. They closely resemble ants, and observa- gogastridae recorded from countries other than Brazil. Of
tions on Brazilian species suggest that they are mimics of these, two (S. rufa and S. brunnea Cresson) occur in Costa
Pseudomyrmex muticus (Mayr) (Papavero, 1964). Syringo- Rica, and the other (the Mexican Megamerina fulvida Bigot,
gaster rufa Cresson in Costa Rica and a related undescribed in fact a species of Odontomera Macquart) belongs in the
species in Ecuador have been observed on the same large Richardiidae (Marshall et al., 2009).
leaf surfaces frequented by the remarkably similar micro-
pezids Metasphen and Globopeza (Marshall, 2005). All Synopsis of the fauna
three genera walk much like their ant models and seem to
feed on honeydew or other materials on large leaves, such Syringogaster Cresson. This genus is organized into four
as those of Heliconia. Syringogaster adults are occasionally species groups, three of which are extant. The Central Amer-
found in large numbers on single plants, presumably forming ican fauna is composed of three species in the S.  brunnea
mating aggregations. Several mating events were observed group and one species each in the S.  rufa and the S.  figu-
in one such aggregation of S. rufa in Costa Rica. No elabo- rata groups. While one of the three species of the S. brunnea
rate courtship behavior was observed, with males mounting group, S. subnearctica Feijen, is endemic to Central America
females and then using their foretarsi to grip the female’s
(range: Mexico to Costa Rica), the other two, S. brunnea and
notopleura.
S.  brunneina Marshall  & Buck, range from Guatemala to
Larvae are unknown, but Feijen (1989) and Meier  &
northern South America. Syringogaster subnearctica is eas-
Hilger (2000) provide information on egg morphology.
ily distinguished from all other Central American species by
its black (as opposed to largely reddish) thorax. The other
Classification two species differ by the absence (S. brunnea) versus pres-
ence (S.  brunneina) of ocellar bristles. The only represen-
Syringogastrids are exclusively Neotropical, and all nine tative of the S.  rufa group and type species of the genus,
described species are included in the single genus Syringo- S. rufa, ranges from Costa Rica south to Ecuador on the west
gaster. The Diopsidae is currently considered to be the fam- side of the Andes. The single Central American species of
ily most closely related to the Syringogastridae (Griffiths, the S. figurata group (S. figurata Marshall & Buck) is known
1972; McAlpine, 1997; Meier  & Hilger, 2000; Marshall from Costa Rica only. The South American fauna includes at
et al., 2009), although prior to the erection of the family least an additional 15 species (Marshall et al., 2009).
846
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Literature cited

Baker, R., G.S. Wilkinson, & R. DeSalle. 2001. Phylogenetic anal- Marshall, S.A., M. Buck, J.H. Skevington, & D. Grimaldi. 2009. A
ysis of sexual dimorphism and eye-span allometry in stalk-eyed revision of the family Syringogastridae (Diptera: Diopsoidea).
flies (Diopsidae). Evolution 55: 1373–1385. Zootaxa 1996: 1–80.
Cresson, E.T. 1912. Descriptions of several new Neotropical aca- McAlpine, D.K. 1997. Gobryidae, a new family of acalyptrate flies
lyptrate Diptera. Entomological News 23: 389–396. (Diptera: Diopsoidea), and a discussion of relationships of the diop-
Curran, C.H. 1934. The families and genera of North American soid families. Records of the Australian Museum 49: 167–194.
Diptera. Published by the author, New York, 512 pp. Meier, R. & R. Baker. 2002. A cladistic analysis of Diopsidae (Dip-
Feijen, H.R. 1989. Diopsidae. In Griffiths, G.C.D. (editor). Flies of tera) based on morphological and DNA sequence data. Insect
the Nearctic Region 9(12): 1–122. E. Schweizerbart’sche Ver- Systematics and Evolution 33: 325–336.
lagsbuchhandlung, Stuttgart. Meier, R. & S. Hilger. 2000. On the egg morphology and phyloge-
Griffiths, G.C.D. 1972. The phylogenetic classification of the Dip- netic relationships of Diopsidae (Diptera: Schizophora). Journal
tera Schizophora with special reference to the structure of the of Zoological Systematics and Evolutionary Research 38: 1–36
male postabdomen. W. Junk, The Hague, 340 pp. Papavero, N. 1964. Notes on the myrmecomimicry of Syringogaster
Hennig, W. 1958. Die Familien der Diptera Schizophora und ihre rufa Cresson, 1912 (Diptera, Acalyptratae, Megamerinidae). Papé-
phylogenetischen Verwandtschaftbeziehungen. Beiträge zur is avulsos do Departamento de Zoologia, São Paulo 16: 110–113.
Entomologie 8: 505–688. Prado, A.P. do. 1969. Syringogastridae, uma nova familia de dipteros
Kotrba, M. 2004. Baltic amber fossils reveal early evolution of Acalyptratae, com a descrição de seis espécies novas do gênero Sy-
sexual dimorphism in stalk-eyed flies (Diptera: Diopsidae). Or- ringogaster Cresson. Studia Entomologica (nova série) 12: 1–32.
ganisms, Diversity and Evolution 4: 265–275. Zumbado, M. A. 2006. Dípteros de Costa Rica y la América tropi-
Marshall, S.A. 2005. Globopeza, a new genus of micropezid fly cal  — Diptera of Costa Rica and the New World tropics. In-
from Ecuador and Venezuela (Diptera, Micropezidae, Taeniap- stituto Nacional de Biodiversidad, Santo Domingo de Heredia,
terinae). Studia dipterologica 11: 537–541. (2004) Costa Rica, 272 pp.
CONOPIDAE (thick-headed flies) 62
Jeffrey H. Skevington, F. Christian Thompson, and Sidney Camras

1 Stylogaster

2 Physoconops

Figs. 62.1–2. Females of (1) Stylogaster stylosus Townsend, (Skevington & Dang, 2002, fig. 6) and (2) Physoconops brachyrhynchus
(Macquart), (Nearctic, MND, fig. 54.1).

847
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Diagnosis of orthopteroids, specifically of cockroaches and crickets

(Smith  & Cunningham-van Someren, 1985; Woodley  &
Small to large (body length 2.5–30.0 mm) flies (Figs. 1–2). Judd, 1998).
Relatively bare and elongate, usually tomentose and colored Most conopid flies are found at flowers, where mating
black and yellow, reddish-brown or blackish, often with takes place and, in many cases, females find suitable hosts in
striking resemblance to their common hosts, wasps and bees. which to lay their eggs. Females attack their hosts by insert-
Head large, broader than thorax, with bare eyes, dichoptic in ing their eggs into the dorsum of the abdomen. The larvae
both sexes; proboscis usually long and slender, frequently develop first by feeding on haemolymph, and in their last
two or more times longer than head, singly or doubly genicu- instar attack the tissue of the thorax, weakening and then
late; palpus one-segmented or absent. Antenna usually long, killing the host. Pupation takes place in the abdomen. Most
with pedicel generally longer than scape, flagellum with conopids are not host specific and will strike a variety of
short, thick apical stylus (Conopinae) or dorsal arista. Ocelli aculeate hymenopterans (Smith, 1966). With the exception
usually absent in Conopinae, present in other subfamilies. of Stylogaster, where more than one egg is laid per host, only
Thorax subquadrate, relatively bare, though postpronotal, one adult is known to emerge (Smith, 1969a; Schmid-Hempel &
notopleural, supra-alar, postalar, and scutellar bristles often Schmid-Hempel, 1989). Host records are summarized by
present. Wing frequently dark along costal margin or macu- Meijere (1912), Freeman (1966), and Smith (1966).
late or unmarked; veins Sc, R1, and R2+3 closely aligned to Stylogastrine conopids are mainly found in association
costa, cell r4+5 closed or nearly closed, spurious vein usually with army ants, where they are concentrated at the front of the
present, cell bm much shorter than cell br, discal cell (dm) swarm and up to 2 m in advance of the swarm (Rettenmeyer,
elongate, cell cup normally elongate, always closed and 1961). They have not been recorded at flowers. Females seek
petiolate. Legs moderately long and usually nearly bare, al- out and dive at their hosts, which have been disturbed by the
though sometimes spinose; tarsi with two distinct pulvilli. ants, using their impact to secure recurrently barbed eggs in
Abdomen cylindrical or constricted at base. Males usually the host cuticle (Kotrba, 1997). The larvae develop in the
with shorter abdomen with six unmodified segments, except host the same way as other conopids. Not all stylogastrine
for Dalmanniinae, which have five; 7th and 8th segments of conopids are associated with army ants, as they occur in
males modified into double cushion-shaped process beneath areas where there are none, so much remains to be learned
5th and 6th segments. Male genitalia (excluding Stylogaster) about their behavior. Also, because of the chaos associated
symmetrical, simple, most complicated in Zodion. Female with army ant swarms, accidents do happen and other non-
with seven visible abdominal segments, 8th appearing to orthopteroid species appear to be attacked, such as tachinids,
form tip of abdomen; 5th prominently produced ventrally in other Stylogaster, and the ants themselves. No successful
form of theca in myopines, except in Myopa where this may rearing, however, has been noted from these other victims,
be replaced by tuft of hairs or bristles; 7th and 8th segments but see Stuckenberg (1963) and Smith (1969b, 1979) for cir-
prolonged and lined with palpate pads to form clasping or- cumstantial evidence of parasitization of Diptera.
gan and theca, these modifications absent in Dalmanniinae While many conopid flies are found at flowers, thus con-
and Stylogastrinae; ovipositor developed in Stylogastrinae tributing to the economy as pollinators, they are also para-
and Dalmanniinae. Sclerotized spermathecae present, two sites of bees. Hence, their overall importance is perhaps
in Conopinae and Myopinae, one in Dalmanniinae and balanced. They have been noted as an important pest of hon-
Stylogastrinae. ey bees (VanDuzee, 1934; Severin, 1937; Jamieson, 1941;
Conopid flies are easily recognized among higher flies by Smith, 1966; Zimina, 1973; Huttinger, 1974; Mei, 1999), so
a combination of characters: ptilinal suture present, lack of in some commercial situations they are harmful. Stylogas-
distinct bristles (except Stylogaster), cell r4+5 frequently peti- trine conopids apparently are of little economic interest to
olate; cell cup usually large (small in Stylogaster), often with humans, as they appear to be minor pollinators and are only
short spurious vein between radial and medial fields, antenna endoparasites of wild orthopteroid species.
and proboscis elongate, usually greatly so. The only group Much remains to be discovered about conopid biology.
that conopids may be confused with are syrphids, but the Rearing conopids is an effective way to generate specimens
well-developed ptilinal suture will separate these two fami- for taxonomy and adds much-needed biological informa-
lies. Many conopids and syrphids are sorted as hymenopter- tion. Parasitized hosts can be easily obtained as the conopid
ans, but obviously differ in the number of wings and other larva always weakens its host before it completes its own
characters. development. Parasitized bees may be found at the entrance
to their hives or nest or may remain in the field overnight.
Biology These parasitized host bees may be simply collected and
kept, and with a little luck adult conopids will emerge. In
Two different biologies are found among the conopids. addition to rearing, a variety of collecting methods are useful
Most conopids are endoparasites of social Hymenoptera, for capturing adult conopids. Malaise traps are the best type
bees and wasps. Stylogastrine conopids are endoparasites of trap to use, while hand collecting at flowers, on hilltops
CONOPIDAE (thick-headed flies) 62 849

(Mei  et  al.,  2010), at army ant swarms (Stylogaster only), Nearly 800 extant species of Conopidae have been de-
and by sweeping are also effective. scribed. They are grouped into four extant subfamilies
Accounts of the immature stages of conopids are widely (Conopinae, Dalmanniinae, Myopinae, Stylogastrinae), plus
scattered in the literature, but Ferrar (1987) gives a summary the fossil subfamily Palaeomyopinae, eight tribes, and 47
and Woodley & Judd (1998) provide additional information genera. All subfamilies, five tribes, 14 genera, and 212 spe-
on Stylogaster. cies are found south of the United States. A catalog to the
Neotropical species was provided by Papavero (1971), but it
is now obsolete. A new conspectus is provided by Thompson
Classification et al (2010). Stuke & Skevington (2007) have provided the
first part of a review of the Costa Rican fauna.
Conopids are placed in Schizophora (Cyclorrhapha), but
their exact relationships and monophyly have not been ful-
ly resolved. Hennig (1966) has reviewed the relationships
Identification
among the included clades, with Camras (1994) providing
further details. The stylogastrines differ from other conopids Most conopids are easily identified to genus, with the
in biology and have a number of distinct characters that have only likely confusion being between males of Parazodion
led some (Rohdendorf, 1964; Smith  & Cunningham-van and Zodion. Species identification is largely based on color
Someren, 1985) to recognize them as a separate family. In the characters, antennal shape, and relative size of antennal seg-
past, most authors placed conopids next to Syrphidae, but the ments. Subgenera are recognized in the large genera Conops
close resemblances are merely due to primitive similarities and Physoconops, but these are not used in identification keys
(symplesiomorphy). Presence of a fully developed ptilinum since they are currently poorly circumscribed and not clearly
indicates that conopids clearly belong to Schizophora. Some monophyletic. Male genitalic characters have been neglected
authors have, however, continued to consider conopids as except among the stylogastrines, but these will certainly yield
the sister to all other schizophorans in the group Archischiza valuable species characters when adequately studied.
(Chvála, 1988; Colless & McAlpine 1991). Hennig (1958) As color and pollinosity are critical for species identifica-
and others rejected the group because once again it is based tion, conopids should ultimately be pinned and dried. Pin-
on primitive characters, and the sister group, Muscaria, re- ning fresh material killed in cyanide is the best. Material
mains undefined. Most current authors view the conopids to collected in alcohol should be degreased and then dried with
be most closely related to the Tephritoidea (Griffiths, 1972; a critical point drier or chemically dried using ethyl acetate
McAlpine, 1989; Korneyev, 2000). or HMDS.

Key to the genera of Conopidae of the northern Neotropical Region

1. First flagellomere with dorsal (Figs.  17–19) or subapical (Fig.  16) arista, with two or three
aristomeres; ocelli present; abdomen not noticeably constricted basally, with segment two broader
than scutellum (Figs. 1, 4). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– First flagellomere with short, thick apical arista, with three aristomeres (Figs. 3, 5, 6, 12–15, 20);
ocelli usually absent (except in Physoconops  & Tropidomyia); abdomen petiolate, constricted
basally, with segment 2 narrower than scutellum or rarely as broad as scutellum (Fig. 3). . . . . . 6

2. Anepimeron with long strong bristle; face with prominent medial carina (Fig.  16); thorax and
scutellum with long distinct black bristles; cell cup shorter than cell bm (Fig. 24); alula narrow,
only about as wide as costal cell (Fig. 24); female with long ovipositor, usually about as long as
rest of abdomen (Fig. 1). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stylogaster Macquart
– Anepimeron without bristles; face without distinct carina; thorax and scutellum without or with
only short bristles; cell cup longer than cell bm (Fig. 25); alula broad, much broader than costal
cell; female ovipositor short, not more than half as long as abdomen (Fig. 4). . . . . . . . . . . . . . . 3

3. Proboscis geniculate, with one bend (Fig. 18). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

– Proboscis doubly geniculate, with two bends (Fig. 17). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

4. Ovipositor of piercing type (Fig. 10); male with five unmodified pre-genital segments (Figs. 11).
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parazodion Kröber
– Ovipositor of clasping type (Fig. 26); male with six unmodified pre-genital segments . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Zodion Latreille
850
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

3 Physoconops 4 Zodion

prepst
prepst

5 Physocephala 6 Conops

h cx

pmtcx brg
p spr

abd sg 1

7 Physocephala 8 Physoconops 9 Physoconops

10 Parazodion 11 Parazodion

Figs. 62.3–11. Adults, heads, thoraces, and abdomens; lateral view of male of (3) Physoconops bulbirostris (Loew); lateral view of female
of (4) Zodion americanum (Wiedemann); lateral view of head and anterior portion of thorax of (5) Physocephala inhabilis (Walker); and
(6) Conops nobilis Williston; posteroventral view of metathorax, hind coxae (below), and base of abdomen (above) of (7) Physocephala
inhabilis (Walker); (8) Physoconops sylvosus (Williston); and (9) Physoconops ornatifrons (Kröber); lateral view of apex of abdomen of
female (10) and male (11) of Parazodion schmidti Kröber (Kröber, 1927, fig. 11). Figures 3–9 illustrated by T. Litwak.
Abbreviations: abd sg 1, abdominal segment 1; h cx, hind coxa; pmtcx brg, postmetacoxal bridge; prepst, proepisternum; p spr, posterior
(thoracic) spiracle.
CONOPIDAE (thick-headed flies) 62 851

black mac

12 Physoconops 13 Tropidomyia

14 Physocephala
15 Physoconops

fc car

16 Stylogaster

17 Myopa 18 Zodion

Figs. 62.12–18. Heads: anterolateral view of (12) Physoconops sylvosus (Williston); anterior view of (13) Tropidomyia bimaculata Williston;
anterolateral view of (14) Physocephala inhabilis (Walker); (15) Physoconops ornatifrons (Kröber); (16) Stylogaster stylosus Townsend;
(17) Myopa fasciata Coquillett; and (18) Zodion americanum (Wiedemann). Figures 12–18 illustrated by T. Litwak.
Abbreviations: fc car, facial carina; black mac, black macula.
852
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

21 Physocephala
19 Thecophora

20 Physocephala

r-m
dm

al
r1
bm c br
r1

22 Physocephala r2+3

r4+5
cup sprs vn
dm

bm

23 Physoconops

cup
C R1
Sc R2+3

24 Stylogaster r-m
R4+5

M1
dm-cu
CuA2
A2 A1
CuA1
A1+CuA2

25 Zodion

st 5
st 6 tg 5
cerc
syntg 8+9 tg 6
st 7

26 Zodion

Figs.  62.19–26. Antennae, leg, wings, and female abdomen: lateral view of antenna of (19)  Thecophora occidensis (Walker), (Nearctic,
MND, fig.  54.6); and (20)  Physocephala furcillata (Williston), (Nearctic, MND, fig.  54.7); posterior view of right hind leg of
(21) Physocephala texana (Williston), (Nearctic, MND, fig. 54.5); dorsal view of wing of (22) Physocephala texana (Williston), (Nearctic,
MND, fig. 54.8); (23) Physoconops fronto (Williston), (Nearctic, MND, fig. 54.9); (24) Stylogaster neglecta Williston, (Nearctic, MND,
fig. 54.10); and (25) Zodion obliquefasciatum (Macquart), (MND, fig. 54.13); lateral view of abdomen of female of (26) Z. perlongum
Coquillett, (Nearctic, MND, fig. 54.16).
Abbreviations: al, alula; cerc, cercus; sprs vn, spurious vein; st, sternite; syntg, syntergite; tg, tergite.
CONOPIDAE (thick-headed flies) 62 853

cerc

hyprct

epand
sur

phapod

ph
hypd

27 Parazodion

spm pmp

epand

ej apod

phapod
cerc

pgt

sur

ph

28 Stylogaster

Figs. 54.27–28. Male terminalia: lateral view of (27) Parazodion schmidti Kröber and (28) Stylogaster neglecta Williston, (Nearctic, MND,
fig. 54.18). Figure 27 illustrated by T. Litwak.
Abbreviations: cerc, cercus; ej apod, ejaculatory apodeme; epand, epandrium; hypd,  hypandrium; hyprct, hypoproct; pgt,  postgonite;
ph, phallus; phapod, phallapodeme; spm pmp, sperm pump; sur, surstylus.
854
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

5. Propleuron bare dorsal to proepisternal bristles; gena usually at least half as high as vertical
eye diameter (Fig. 17); midcoxa bare on posteromedial surface; species usually reddish brown;
Mexico, Chile. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Myopa Fabricius]
– Propleuron pilose dorsomedially; gena usually less than half as high as vertical eye diameter (as
in Fig. 18); midcoxa pilose posteromedially; species usually black. . . . . . . Thecophora Rondani

6. Wing with crossvein r-m well apical of middle of cell dm, apical of crossvein sc-r (Fig. 22); ocelli
absent (Fig. 14); proepisternum bare (Fig. 5); hind femur irregularly thickened on basal one-third
(Fig. 21); postmetacoxal bridge connected to fused metepimera with strong uniformly sclerotized
portion running between coxae (Fig. 7). . . . . . . . . . . . . . . . . . . . . . . . . . . . Physocephala Schiner
– Wing with crossvein r-m at or only slightly beyond middle of cell dm, in line with or basal to
crossvein  sc-r (Figs.  3, 23); ocelli present or absent; proepisternum usually with at least one
bristle, rarely bristle small or absent (Fig. 6); hind femur uniformly thickened basally, thickest
medially; postmetacoxal bridge not connected to fused metepimera, area ventrad of bridge not
sclerotized and area between coxae only weakly sclerotized (Figs. 8–9). . . . . . . . . . . . . . . . . . . 7

7. Face with medial carina, without submedial grooves, with black macula on lateral edge of frons
adjacent to eye (Fig. 13); with two proepisternal bristles; ocellar triangle distinct, with two ocelli
(Fig. 13); cell cup elongate, extending to level of crossvein r-m. . . . . . . . . Tropidomyia Williston
– Face with indistinct medial carina, with distinct, broad submedial grooves, frons without black
macula (Fig. 15); other characters various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

8. Vertex without ocellar tubercle; proepisternum usually with more than one bristle (Fig. 6). . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Conops Linneaus
– Vertex with distinct ocellar tubercle (Fig. 2, 15); proepisternum usually with only one bristle. . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Physoconops Szilady

Synopsis of the fauna America (15 in Costa Rica). The most recent keys are by
Camras (1955) and Stuke & Skevington (2007).
The fauna of this region contains about 60 species, most
of which are already described. There are some unresolved Stylogaster Macquart. This genus is found in all regions,
species complexes, however, that will result in further new except the Palearctic. There are 94 described species, 70 of
species. which are known from the Neotropical Region and 24 from
Central America (16 in Costa Rica). The last key is that of
Conops Linnaeus. This genus is cosmopolitan, absent Camras & Parrillo (1985).
only from New Zealand and most oceanic islands. There
are 168 described species, eight in the Neotropical Region, Thecophora Rondani. This is another genus found in all
regions, but absent from New Zealand and most oceanic is-
and in Costa Rica only C. nobilis (Walker) and C. geminatus
lands. There are 37 species, with six in the Neotropical Re-
Camras. The last key is by Camras (1955).
gion, and while the genus ranges south to Argentina, only
Parazodion Kröber. Endemic to the Neotropical Region, T. nigripes Camras has been found in Central America. The
this genus has two known species, including the type spe- last key (Camras, 1945) only covered North America and is
of little use. No key is available for the Neotropical Region.
cies, P. schmidti Kröber, described from Costa Rica.
Tropidomyia Williston. This genus is found in most re-
Physocephala Schiner. This genus is cosmopolitan, ex-
gions, except the Nearctic and Australian Regions. There are
cept absent from New Zealand and most oceanic islands. Of seven species, two of which are found in the Neotropical Re-
133 species, 31 are found in the Neotropical Region, with gion, of which T. bimaculata Williston occurs in Costa Rica.
eight in Central America (seven in Costa Rica). The latest The last key was that of Camras (1955).
keys are by Camras (1996, see also 1957) and Stuke & Skev-
ington (2007). Zodion Latreille. Zodion species are found in all regions,
but are absent from New Zealand and most oceanic islands.
Physoconops Szilady. Found in most regions, except absent Of 57 species, 32 are found in the Neotropical Region and
from the Palearctic and Australian Regions, Physoconops has about eight in Central America (seven in Costa Rica). The
62 species, 49 found in Neotropical Region and 16 in Central last key was in Pearson & Camras (1978).
CONOPIDAE (thick-headed flies) 62 855

Literature cited
Camras, S. 1945. A study of the genus Occemyia in North Amer- Mei, M., J. Gibson & J.H. Skevington. 2010. Observations on hill-
ica. Annals of the Entomological Society of America 38: topping in the Conopidae (Diptera). Journal of Insect Science 10:
216–222. 27, available online at insectscience.org/10.27.
Camras, S. 1955. A review of the New World flies of the genus Meijere, J.C.H. de. 1912. Neue Beiträge zur Kenntnis der Conopiden.
Conops and allies (Diptera: Conopidae). Proceedings of the Tijdschrift voor Entomologie 55: 184–207.
United States National Museum 105: 155–187. Papavero, N. 1971. 47. Conopidae. In Papavero, N. (editor). A Cata-
Camras, S. 1957. A review of the New World Physocephala (Dip- logue of the Diptera of the Americas South of the United States.
tera: Conopidae). Annals of the Entomological Society of Amer- Departamento de Zoologia, Secretaria da Agricultura, São Paulo,
ica 50: 213–218. 28 pp.
Camras, S. 1994. A new subfamily for the fossil conopid fly, Pa- Pearson, D.L. & S. Camras. 1978. Notes and key to Neotropical Zo-
laeomyopa tertiaria (Diptera: Conopidae). Entomological News dion (Diptera: Conopidae). Journal of the Kansas Entomological
105: 175–177. Society 51: 198–206.
Camras, S. 1996. New information on the New World Physocephala Rettenmeyer, C.W. 1961. Observations on the biology and taxon-
(Diptera: Conopidae). Entomological News 107: 104–112. omy of flies found over swarm raids of army ants (Diptera: Ta-
Camras, S. & P.P. Parrillo. 1985. Review of New World Stylogaster chinidae, Conopidae). The University of Kansas Science Bulletin
(Diptera: Conopidae). Annals of the Entomological Society of 42: 993–1066.
America 78: 111–126. Rohdendorf, B.B. 1964. [Historical development of dipterous in-
Chvála, M. 1988. Family Conopidae, pp. 245–272. In Soos, A. & sects.] Trudy Paleontologischeskogo Instituta, Moscow, 100,
L. Papp (editors). Catalogue of Palaearctic Diptera. Volume 8, 311  pp. [In Russian, translation 1974, University of Alberta
Syrphidae-Conopidae. Elsevier Science Publishers, Amsterdam, Press, Edmonton, 360 pp.]
Oxford, 363 pp. Schmid-Hempel, R.  & P. Schmid-Hempel. 1989. Superparasitism
Colless, D.H.  & D.K. McAlpine. 1991. Diptera, pp.  717–786. In and larval competition in conopid flies (Dipt., Conopidae), para-
Division of Entomology, CSIRO. The Insects of Australia, Sec- sitizing bumblebees (Hym., Apidae). Mitteilungen der Schweiz-
ond Edition. Melbourne University Press and Cornell University erischen Entomologischen Gesellschaft 62: 279–289.
Press, Ithaca, 1137 pp. Severin, H.C. 1937. Zodion fulvifrons Say, a parasite of the honey
Ferrar, P. 1987. A guide to the breeding habits and immature stages bee. Entomological News 48: 243–244.
of Diptera Cyclorrhapha. Entomonograph 8(1). E.J. Brill / Scan- Skevington, J.H. & P.T. Dang (editors). 2002. Exploring the diver-
dinavian Science Press, Leiden, Copenhagen, 478 pp. sity of life: Diptera. Biodiversity 3: 3–27.
Freeman, B.A. 1966. Notes on conopid flies, including insect host, Smith, K.G.V. 1966. The larva of Thecophora occidensis, with com-
plant and phoretic relationships (Diptera: Conopidae). Journal of ments upon the biology of Conopidae (Diptera). Journal of Zool-
the Kansas Entomological Society 39: 123–131. ogy 149: 263–276.
Griffiths, G.C.D. 1972. The phylogenetic classification of Diptera Smith, K.G.V. 1969a. Diptera : Conopidae. Handbooks for the iden-
Cyclorrhapha, with special reference to the structure of the male tification of British Insects 10(3(a)): 1–19.
postabdomen. Series entomologica 8. Dr. W. Junk, The Hague, Smith, K.G.V. 1969b. Further data on the oviposition by the genus
340 pp. Stylogaster Macquart (Diptera: Conopidae, Stylogasterinae)
Hennig, W. 1958. Die familien der Diptera Schizophora und ihre upon adult calyptrate Diptera associated with ants and animal
phylogenetischen verwandtschaftsbeziehungen. Beiträge zur dung. Proceedings of the Royal Entomological Society of Lon-
Entomologie 8: 505–688. don. (A) General Entomology 44: 35–37.
Hennig, W. 1966. Conopidae im Baltischen Bernstein (Diptera: Smith, K.G.V. 1979. The genus Stylogaster (Diptera: Conopidae:
Cyclorrhapha). Stuttgarter Beiträge zur Naturkunde 154: Stylogasterinae) in the Australian region. Australian Journal of
1–24. Zoology 27: 303–310.
Huttinger, E. 1974. Little attention paid to Conopidae (Diptera, Smith, K.G.V. & G.R. Cunningham-van Someren. 1985. The larva
Insecta) as parasites of honey bees. Bienenvater 95: 102–103. of Stylogaster varifrons Malloch (Dipt., Stylogastridae). Ento-
Jamieson, C.A. 1941. A dipterous parasite (Myopa sp.) of the honey- mologist’s Monthly Magazine 121: 81–85.
bee. American Bee Journal 81: 324. Stuckenberg, B.R. 1963. A study on the biology of the genus Sty-
Korneyev, V.A. 2000. Phylogenetic relationships among the fami- logaster, with a description of a new species from Madagascar
lies of the superfamily Tephritoidea, pp.  3–22. In Aluja, M.  & (Diptera: Conopidae). Revue de Zoologie et de Botanique Afri-
A.L.  Norrbom (editors). Fruit Flies (Tephritidae): Phylogeny caines 68: 251–276.
and evolution of behavior. CRC Press, Boca Raton, London, Stuke, J.-H. & J.H. Skevington. 2007. The Conopidae of Costa Rica
New York, Washington, D.C., [xvi] + 944 pp. (Diptera) (Part  1: Conopinae  — Conopini  & Tropidomyiini).
Kotrba, M. 1997. Shoot or stab? Morphological evidence on the Zootaxa 1528: 1–40.
unresolved oviposition technique in Stylogaster Macquart (Dip- Thompson, F.C., J.H. Skevington, & S. Camras. 2010. Conspectus
tera: Conopidae), including discussion of behavioral observa- to the Neotropical Conopidae (Diptera). Neotropical Diptera.
tions. Proceedings of the Entomological Society of Washington VanDuzee, M.C. 1934. Conopidae from North Dakota and the
99: 614–622. Rocky Mountain region. Annals of the Entomological Society of
Kröber, O. 1927. Beiträge zur kenntnis der Conopidae. Konowia 6: America 27: 315–323.
122–143. Woodley, N.E. & D.D. Judd. 1998. Notes on the host, egg, and pu-
McAlpine, J.F. 1989. Phylogeny and classification of the Musco- parium of Stylogaster biannulata (Say) (Diptera: Conopidae).
morpha, pp. 1357–1518. MND, Volume 3. Proceedings of the Entomological Society of Washington 100:
Mei, M. 1999. Pseudophysocephala cfr. vitripennis (Curran) (Dipt., 658–664.
Conopidae), parasitizing the honeybee (Hym., Apildae) in Guin- Zimina, L.V. 1973. Physocephala truncata (Diptera, Conopidae),
ea, West Africa. Entomologist’s Monthly Magazine 135(1620-3): a parasite of honey bees in Tuva. Zoologicheskii Zhurnal 52:
141–142. 1732–1733.
856
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
LONCHAEIDAE (lance flies) 63
Allen L. Norrbom and Cheslavo A. Korytkowski

Fig. 63.1. Female of Lonchaea polita Say, (Nearctic, MND, fig. 62.1).

Diagnosis sometimes with one to several setulae medially. Meron bare.

Legs mostly dark, tarsi sometimes partially or entirely yel-
Moderately small (body length 3.0–6.0 mm), stout, rela- low. Wing (Figs. 22–23) somewhat triangular, tapered, with
tively setose, shiny black to metallic blue or green flies, with anal lobe and alula well developed. Venation with crossveins
wings usually hyaline, sometimes partly yellowish or fumose bm-cu, r-m, and dm-cu and vein  CuA2 present. Costa ex-
(Fig. 1). Body stout, abdomen broad and flat. Frons narrower tended to vein  M, with costagial and humeral weakenings
in male than in female; setulose, with one reclinate orbital and subcostal break. Subcosta complete, apex free from R1.
seta. Ocellar and inner and outer vertical setae strong (Fig. 2). Vein R1 nonsetulose dorsally. Crossvein r-m near or proximal
Postocellar setae divergent, weak. Without vibrissa, but one to midlength of cell dm. Vein A1+CuA2 sometimes reaching
or more subvibrissal setae sometimes vibrissa-like. Lunule wing margin as fold. Dorsal calypter well developed, with
large, exposed, bare or setulose. Face broad, depressed, usu- whitish to brownish fringe of hairs, those at calyptral fold
ally without strong carina. Eye large, pilose or bare. First sometimes longer and darker than others (Neosilba, some
flagellomere short and porrect to long and decumbent; arista Dasiops). Halter dark brown. Abdomen broad and flat. Male
bare, pubescent or plumose. Thorax with one postpronotal, with epandrium and hypandrium symmetrical. Phallus short
two notopleural, one presutural, and one postsutural supra- and unsegmented (Dasiops; Figs. 14–15); or elongate, biseg-
alar, two  postsutural dorsocentral, two  prescutellar acros- mented, and ornamented (other genera; Figs.  16–19). Sur-
tichal, often one  weak intra-alar, and two  postalar setae. stylus usually with short, stout prensisetae (Figs.  14–15).
Scutum strongly arched, setulose. Scutellum with two large Pregonite (=gonopod of McAlpine, 1987) with base fused
setae and sometimes additional setulae on margin or disk. to hypandrium, with tiny setula on medial surface near apex;
Proepisternum with one seta, proepimeron with one to many postgonite reduced and semimembranous to strongly devel-
setulae. Anepisternum with numerous setae, at least posteri- oped and toothed. Female with sclerotized telescopic ovi-
orly, often smaller anteriorly. Katepisternum with one to three positor (Figs.  20–21). Seventh tergite and sternite fused to
dorsal setae and numerous setulae. Anepimeron usually bare, form oviscape. Eighth tergite and sternite comprising four

857
858
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

o vt s
orb s i vt s
poc s

sbvb s

2 Dasiops 3 Protearomyia 4 Chaetolonchaea

5 Earomyia 6 Lamprolonchaea 7 Neosilba

8 Protearomyia 9 Chaetolonchaea 10 Earomyia

anepst

pstg s kepst

11 Dasiops 12 Chaetolonchaea 13 Neosilba

Figs. 63.2–13. Heads, scutella, and thoraces: anterolateral view of head of (2) Dasiops alveofrons McAlpine, (Nearctic, MND, fig. 62.2);
(3) Protearomyia trichopleura McAlpine, (Nearctic, MND, fig. 62.3); (4) Chaetolonchaea americana McAlpine, (Nearctic, MND, fig. 62.4);
(5) Earomyia aberrans (Malloch), (Nearctic, MND, fig. 62.5); (6) Lamprolonchaea smaragdi (Walker), (Old World, MND, fig. 62.6);
(7) Neosilba batesi (Curran), (Nearctic, MND, fig. 62.7); dorsal view of scutellum of (8) P. trichopleura, (Nearctic, MND, fig. 62.8);
(9) C. americana, (Nearctic, MND, fig. 62.9); (10) E. aberrans, (Nearctic, MND, fig. 62.10); lateral view of thorax of (11) D. alveofrons,
(Nearctic, MND, fig. 62.11); (12) C. americana, (Nearctic, MND, fig. 62.12); (13) N. batesi, (Nearctic, MND, fig. 62.13).
Abbreviations: anepst, anepisternum; i  vt  s, inner vertical seta; kepst, katepisternum; orb  s,  orbital seta; o  vt  s,  outer vertical seta;
poc s, postocellar seta; pstg s, poststigmatal seta; sbvb s, subvibrissal seta.
LONCHAEIDAE (lance flies) 63 859

elongate rods, closely associated with or occasionally fused Neosilba and Lonchaea spp. in the Neotropical Region)
to usually oval or pointed apical sclerite formed by fused breed in damaged fruit (particularly those previously at-
epiproct, hypoproct, and cerci. Three spermathecae present tacked by Tephritidae), vegetables, or cacti. Many species
(McAlpine, 1987). of Dasiops and Lonchaea develop under the bark of dead
Larvae (Fig. 24) typical muscomorph maggots, relative- or dying trees, often in association with bark beetles and
ly long and slender, cylindrical, tapered anteriorly, and with weevils; at least some species of Lonchaea feed on dead
spiculose intersegmental rings and creeping welts. Mandi- beetles or are facultative predators (perhaps only on sick
ble without secondary teeth, dental sclerite quadrangular or injured beetles), but their exact role and importance in
to triangular, parastomal bar slender, hypopharyngeal sc- controlling beetle populations are unclear. Adult Lonchaei-
lerite elongate, tentoropharyngeal sclerite with large dorsal dae are unusual for Schizophora in that they form mating
and ventral wings. Anterior spiracle (Fig. 25) usually fan- swarms (McAlpine & Munroe, 1968).
shaped, with 5–10 digits. Posterior spiracle (Fig. 26) raised
above surface of caudal segment, with distinct peritreme,
sometimes strongly sclerotized, often with one to several
Classification
dorsal or lateral spines, and with three openings, the dorsal
and ventral openings aligned at nearly 180°, the medial one The Lonchaeidae belong to the superfamily Tephri-
more or less perpendicular to them. Sometimes (in Dasi- toidea and may be the most basal clade of the superfamily
ops) with sclerotized area lateral to posterior spiracle. (McAlpine, 1989; Korneyev, 1999). There are nine genera,
Lonchaeidae are sometimes confused with Agromyzi- divided into two subfamilies: Dasiopinae, which includes
dae, but differ in having a longer cell  dm (shorter than only Dasiops; and Lonchaeinae, which includes the other
distal section of vein  M in Agromyzidae) and no vibris- genera, divided into two tribes, Earomyiini and Lonchaei-
sa or frontal setae (at least one present in Agromyzidae). ni, each with four genera. McAlpine (1962) provided an
The brown halter is also a useful diagnostic character for analysis of the phylogenetic relationships among and with-
Lonchaeidae. in the genera. No fossil species are known. Those fossils
previously described in the family have been transferred
elsewhere (McAlpine, 1987).
Biology The family is cosmopolitan, with more than 500 species
currently recognized. There are four genera known from
Ferrar (1987) reviewed the biology of the family; Fer- the Neotropical Region, and 93 species reported from Mex-
rar (1987) and McAlpine (1987) compiled what little is ico to South America, but the majority of the Neotropical
known of the immature stages. Most lonchaeids are scav- species are undescribed.
engers as larvae, breeding mainly in damaged plant tis-
sues or decaying vegetation, but a few species are primary
invaders of living plants. Some species of Dasiops are Identification
pests of Passiflora (Norrbom & McAlpine, 1997) or vari-
ous cacti (e.g., D. saltans Townsend), and Neosilba perezi McAlpine & Steyskal (1982) and McAlpine (1987) pro-
(Romero & Ruppel) is a stem miner on cassava (Manihot vided keys to the world genera, the latter copied below.
esculenta) (McAlpine & Steyskal, 1982). Some species of Korytkowski & Ojeda (1971) provided keys to the species
Earomyia breed in conifer cones where they feed on the of Peru, and Norrbom  & McAlpine (1997) keyed those
seeds, sometimes in association with Cecidomyiidae and species of Dasiops associated with Passiflora. However,
other primary invaders. Other species live in flower heads because of the large number of undescribed species,
and root crowns. Many lonchaeid species (particularly Neotropical lonchaeids are extremely difficult to identify.

Key to genera of Lonchaeidae of the world

1. Poststigmatal setae present (Fig. 11); DASIOPINAE. . . . . . . . . . . . . . . . . . . . . Dasiops Rondani

– Poststigmatal setae absent (Figs. 12, 13); LONCHAEINAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2. Lunule bare (Figs. 3–6); EAROMYIINI. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

– Lunule setulose (Fig. 7); LONCHAEINI. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

3. Scutellum with four large setae only, without additional setulae (Fig. 8); Holarctic. . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Protearomyia McAlpine]
– Scutellum with two or more setulae in addition to usual four large setae (Figs. 9, 10). . . . . . . . . 4
860
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

phapod

pregt
hypd pgt

ej apod
pregt

ph ph

epand
sur

sur
epand
prens

cerc

14 Dasiops 15 Dasiops
16 Neosilba

pregt

ph gd

pgt

17 Lonchaea 18 Lonchaea 19 Neosilba

sg 7 sg 8 cerc

20 Lonchaea
sg 7 sg 8 cerc

21 Lonchaea

Figs. 63.14–21. Male terminalia and female abdomen: ventral view (14) and left lateral view (15) of male terminalia of Dasiops alveofrons
McAlpine, (Nearctic, MND, figs. 62.14, 15); ventral view of male terminalia of (16) Neosilba batesi (Curran), (Nearctic, MND, fig. 62.16)
and (17) Lonchaea polita Say, (Nearctic, MND, fig. 62.17); left lateral view of male terminalia of (18) L. polita, (Nearctic, MND, fig. 62.18)
and (19) N. batesi, (Nearctic, MND, fig. 62.19); dorsal view (20) and lateral view (21) of female abdomen of L. polita, (Nearctic, MND,
figs. 62.20, 21).
Abbreviations: cerc, cercus; ej apod, ejaculatory apodeme; epand, epandrium; hypd, hypandrium; pgt, postgonite; ph, phallus; phapod,
phallapodeme; ph gd, phallic guide; pregt, pregonite; prens, prensiseta; sg, segment; sur, surstylus.
LONCHAEIDAE (lance flies) 63 861

4. Body brilliant golden green or bronzy blue; eye bare; setulae of frons sparse and short (Fig. 6);
anepisternum with one to several clearly distinguishable anterodorsal setae and two well-differentiated
posterodorsal setae (as in Fig. 13); katepisternum with one clearly distinguishable seta; mainly
Oriental and Australasian, one  species (L.  smaragdi (Walker)) widely introduced in southern
Palearctic Region, Afrotropical Region and Bermuda. . . . . . . . . . . . . . . [Lamprolonchaea Bezzi]
– Body dull to subshiny brownish- to bluish-black; eye bare or pilose; setulae of frons fairly dense
and long (Figs. 4, 5); anepisternum with (as in Fig. 13) or without (Fig. 12) anterodorsal setae,
with more numerous posterior setae, sometimes poorly differentiated from setulae; katepisternum
usually with several setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

Sc R1

R2+3

br
bm R4+5
cup dm

M1

CuA1
A1+CuA2

22 Lonchaea 23 Dasiops

24 Lonchaea

pastm b a spr

d corn

spr plt

v corn

ant md hyphar scl

25 Lonchaea 26 Lonchaea

Figs. 63.22–26. Wings and larval structures; dorsal view of wing of (22) Lonchaea polita Say, (Nearctic, MND, fig. 62.22); (23) Dasiops
alveofrons McAlpine, (Nearctic, MND, fig. 62.3); lateral view of mature larva of (24) L. corticis Taylor, (Nearctic, MND, fig. 62.24); head,
anterior thoracic segments, and internal cephalopharyngeal skeleton of (25) L. corticis, (MND, fig. 62.25); posterolateral view of posterior
spiracles and last abdominal segment of (26) L. corticis, (MND, fig. 62.26).
Abbreviations: ant, antenna; a spr, anterior spiracle; d corn, dorsal cornu of tentoropharyngeal sclerite; hyphar scl, hypopharyngeal sclerite;
md, mandible; pastm b, parastomal bar; spr plt, spiracular plate; v corn, ventral cornu of tentoropharyngeal sclerite.
862
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

5. Scutellum with one or more marginal setulae anterior to basal scutellar seta (Fig. 9); anepisternum
with or without weak anterodorsal setae (Fig. 12); gena and parafacial wide (Fig. 4); calypteres
with margins and fringes whitish; Holarctic . . . . . . . . . . . . . . . . . . . . . . [Chaetolonchaea Czerny]
– Scutellum bare anterior to basal scutellar seta (Fig. 10); anepisternum with long anterodorsal setae
(as in Fig. 13); gena and parafacial usually narrow (Fig. 5); calypteres with margins and fringes
brown; Holarctic. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .[Earomyia Zetterstedt]

6. Katepisternum with one strong seta (as in Fig. 11); arista usually bare or short pubescent . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lonchaea Fallén, in part
– Katepisternum with two strong setae (Fig. 13); arista bare to long plumose. . . . . . . . . . . . . . . . . 7

7. Arista usually bare or short pubescent, or if long pubescent, then fringes of calypteres uniform in
length; prosternum bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
– Arista plumose (Fig. 7), or if somewhat short plumose, then fringes of calypteres with cluster of
longer black hairs at fold (Fig. 13); prosternum bare or setulose . . . . . . . . . . . . . . . . . . . . . . . . . 9

8. Abdominal sternite one setulose; Holarctic. . . . . . . . . . . . . . . . . . . . [Setisquamalonchaea Morge]

– Abdominal sternite one bare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lonchaea Fallén, in part

9. Fringes of calypteres with cluster of longer black hairs at fold (Fig.  13); anepisternum with
posterior row of five to seven evenly spaced setae (Fig. 13); prosternum bare or setulose. . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neosilba McAlpine
– Fringes of calypteres uniform throughout (as in Figs. 11, 12); anepisternum with three to four
strong posterior setae, with gap in middle; prosternum setulose; Old World tropics, Peru. . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Silba Macquart]

Synopsis of the fauna Lonchaea Fallén. Lonchaea is a large, cosmopolitan ge-

nus with 29 described species reported from the Neotropical
Four genera and 93 species of Lonchaeidae are reported Region or Nearctic Mexico. Six species are recorded from
from the Neotropical Region (or Mexico), of which three Mexico or Central America, but others probably occur in the
genera and 29 species are known from Mexico and Central region. Luna (1987, 1988) revised and keyed the Neotropical
America. The genus Silba is restricted mainly to the Old fauna, but additional species remain undescribed.
World tropics and subtropics, but one species, S.  devians Neosilba McAlpine. McAlpine  & Steyskal (1982) pro-
Hennig, was described from Peru (McAlpine  & Steyskal, vided a revision of most of the 17 known species of this
1982). Many species now in Neosilba were previously in- Neotropical genus, but there are numerous undescribed spe-
cluded in Silba. cies (C.A. Korytkowski, unpublished data). Ten species are
reported from Mexico or Central America. Some Neosilba
Dasiops Rondani. This is a large, cosmopolitan genus species have been reared from a wide variety of fruits, but
with 53 described species reported from the Neotropical they are probably secondary invaders in damaged tissues
Region or Nearctic Mexico. Twelve species are known (McAlpine & Steyskal, 1982). Neosilba perezi (Romero &
from Mexico or Central America. Numerous species remain Ruppel) is a pest of cassava (Manihot esculenta) in which it
undescribed. mines the stems or terminal shoots or buds.

Literature cited
Ferrar, P. 1987. A guide to the breeding habits and immature stages Luna, I.G. 1987. Revisión taxonómica del genero Lonchaea Fallén
of Diptera Cyclorrhapha. Entomonograph 8(1). E.J.  Brill  / en la Región Neotropical (Diptera: Lonchaeidae). Tesis. Pro-
Scandinavian Science Press, Leiden, Copenhagen, 478 pp. grama de Maestria en Entomología, Universidad de Panamá,
Korneyev, V.A. 1999. Phylogenetic relationships among the fami- 329 pp.
lies of the superfamily Tephritoidea, pp. 3–22. In Aluja, M. & Luna, I.G. 1988. Clave taxonómica para la identificación del genero
A.L. Norrbom (editors). Fruit flies (Tephritidae): Phylogeny and Lonchaea Fallén en la Región Neotropical. Scientia 3: 27–42.
evolution of behavior. CRC Press, Boca Raton, [16] + 944 pp. McAlpine, J.F. 1962. The evolution of the Lonchaeidae (Diptera).
Korytkowski, C.A. & D. Ojeda. 1971. Revision de las especies de Ph. D. dissertation, University of Illinois, Urbana, 233 pp.
la familia Lonchaeidae en el Peru (Diptera: Acalyptratae). Re- McAlpine, J.F. 1987. Family Loncheidae, pp. 791–797. MND,
vista Peruana de Entomología 14: 87–116. Volume 2.
LONCHAEIDAE (lance flies) 63 863

McAlpine, J.F. 1989. Phylogeny and classification of the Musco- McAlpine, J.F.  & G.C. Steyskal. 1982. A revision of Neosilba
morpha, pp. 1397–1518. MND, Volume 3. McAlpine with a key to the world genera of Lonchaeidae. The
McAlpine, J.F.  & D.D. Munroe. 1968. Swarming of lonchaeid Canadian Entomologist 114: 105–137.
Norrbom, A.L. & J.F. McAlpine. 1997. A revision of the Neotropi-
flies and other insects, with descriptions of four new species cal species of Dasiops Rondani (Diptera: Lonchaeidae) attack-
of Lonchaeidae (Diptera). The Canadian Entomologist 100: ing Passiflora (Passifloraceae). Memoirs of the Entomological
1154–1178. Society of Washington 18: 189–211.
864
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
PIOPHILIDAE (skipper flies) 64
Andrej L. Ozerov and Allen L. Norrbom

Fig. 64.1. Male of Piophila casei (Linnaeus), (MND, fig. 69.1).

Diagnosis abdomen, with well-developed anal lobe and alula; en-

tirely hyaline in Neotropical species. Costa without
Moderately small (body length 3.0–8.0 mm) shiny black humeral break; with subcostal break. Crossvein bm-
to dull brown or yellow flies, with wings frequently glassy cu sometimes absent (present in Neotropical species),
(Fig.  1). Frons setulose, sometimes indistinctly; with zero crossveins r-m and dm-cu and vein  CuA 2 present. Sub-
to two lateroclinate orbital setae, occasionally with one to costa complete. Vein R1 setulose dorsally (Neottiophili-
two inclinate frontal setae. Postocellar setae divergent. one nae) or bare (Piophilinae). Veins R4+5 and M parallel or
to two vibrissae present. Antenna usually short (except in divergent apically. Crossvein dm-cu at or distal to mid-
male of Nearctic Prochyliza xanthostoma Walker), porrect; length of cell dm. Vein A 1+CuA 2 usually reaching wing
flagellomere 1 oval; arista dorsobasal, bare or short pubes- margin at least as fold. Male with phallus short to long,
cent. Thorax with zero to two postpronotal, one to two no- bare to pubescent. Surstylus usually present and not
topleural (two in Neotropical spp.), zero to one presutural fused to epandrium. Pregonite and postgonite present.
and one postsutural supra-alar, zero to one presutural and Female with telescopic ovipositor. Seventh and eighth
one to three postsutural dorsocentral, zero to one prescutellar tergites and sternites elongate. Epiproct, hypoproct, and
acrostichal, usually one intra-alar, one postalar, two scutel- cerci fused. Two spermathecae present (J.F.  McAlpine,
lar, one proepisternal, usually zero to one anepisternal (two 1977; Ozerov, 2000).
in Protopiophila latipes (Meigen), up to four in one row in Larvae (Fig. 6) typical muscomorph maggots, relatively
Neottiophilum and Actenoptera, which are not Neotropical), long and slender, cylindrical, with spiculose intersegmental
zero anepimeral, and one (often small) to several katepis- rings and creeping welts. Mandible without secondary teeth,
ternal setae. Anepisternum and/or scutellum often finely discrete dental sclerite lacking, parastomal bar slender, hypo-
setulose, anepimeron rarely so. Scutellum sometimes pharyngeal sclerite and labial sclerites well developed, ten-
flattened and elongate, especially in male Thyreophorini. toropharyngeal sclerite with more or less equally large dorsal
Tibiae without dorsal preapical setae. Wing longer than and ventral flanges. Anterior spiracle (Fig.  7) fan-shaped,

865
866
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

with 4–12 digits. Terminal segment with three to four pairs Psilidae lack a vibrissa and have an incomplete subcosta; and
of tubercles surrounding spiracular plates (Fig. 8). Posterior Sepsidae do not have a subcostal break in the costa.
spiracles not on raised tubercles, each with distinct peritreme
and with three parallel or slightly radiating slits.
Piophilidae may be confused with other Tephritoidea, Biology
particularly species with dark bodies and unpatterned wings,
but differ by having a vibrissa. No Neotropical piophilids The natural history of these flies is poorly known for the
have patterned wings. Some species of several other families Neotropical fauna, except for the introduced species. Ferrar
are superficially similar, but differ as follows: Agromyzi- (1987), J.F. McAlpine (1977, 1989), and Ozerov (2000) have
dae have well-developed, inclinate frontal setae (absent reviewed the biology and immature stages of the family, and
in Neotropical Piophilidae) and a much shorter (or open) Freidberg (1981) described the behavior and other aspects
cell dm; Clusiidae have well-developed, inclinate frontal or of the biology of the Palearctic Centrophlebomyia furcata
interfrontal setae and usually a lobe on the pedicel projecting (Fabricius). Most piophilids, and probably all of the Neotro-
into the first flagellomere on the lateral side; Milichiidae have pical species, are scavengers as larvae, breeding mainly in
an incomplete subcosta and/or a humeral break in the costa protein-rich plant and animal matter. Larvae of the Palearc-
and usually have well-developed, inclinate frontal setae; tic Neottiophilum praeustum (Meigen), the only species of

vb

2 Stearibia 3 Bocainamyia

pprn s

npl s
presut dc s
psut dc s presut
acr s spal s

psut
spal s
ial s
pal s

4 Bocainamyia 5 Protopiophila

Figs.  64.2–5. Heads, thorax, and scutellum: anterodorsal view of head of (2)  Stearibia nigriceps (Meigen), (MND, fig.  69.2); and
(3) Bocainamyia necrophila Albuquerque, (MND, fig. 69.10); dorsal view of scutellum of (4) B. necrophila, (J.F. McAlpine, 1977, fig. 21);
dorsal view of thorax of (5) Protopiophila latipes (Meigen), (Holarctic, MND, fig. 69.12).
Abbreviations: acr s, acrostichal seta; ial s, intra-alar seta; npl s, notopleural seta; pal s, postalar seta; pprn s, postpronotal seta; presut dc s,
presutural dorsocentral seta; presut spal s, presutural supra-alar seta; psut dc s, postsutural dorsocentral seta; psut spal s, postsutural supra-alar
seta; vb, vibrissa.
PIOPHILIDAE (skipper flies) 64 867

Neottiophilinae whose biology is known, live in bird nests behavior may occur in most or all Piophilidae, but it is not
where they are intermittent blood-sucking ectoparasites on exclusive to the family, as it is also known in various other
the nestlings. Species of Mycetaulini, so far as known, breed Tephritoidea and some other acalyptrate flies (some Clusii-
in fungi or rotting wood. Some European species are attracted dae and Agromyzidae, according to Oldroyd, 1964).
to fermenting liquids (e.g., banana–yeast baits, beer traps).
Other Piophilinae have been reared from materials such as
carrion (e.g., of mammals, birds, fish, snails), especially in an Classification
advanced state of decay or when relatively dry, bone marrow,
The Piophilidae belong to the superfamily Tephritoidea
discarded cervid antlers, dung, and decaying leaves. Several
and appear to be most closely related to the Pallopteridae
species are synanthropic and have become widespread, es-
(J.F. McAlpine, 1977, 1989; Korneyev, 1999).
pecially Piophila casei (L.), which is a pest of stored foods The higher classification of the Piophilidae has been un-
such as meat, cheese, and hides. If ingested, the larvae can settled. Some authors recognize the separate families Neot-
cause serious intestinal lesions. tiophilidae and Thyreophoridae (e.g., Soós & Papp, 1984),
Neotropical piophilids are rare in collections, in part be- but they were included within the Piophilidae in the treat-
cause of their low diversity in the region, but also because of ments in the Nearctic and Palearctic Manuals (J.F. McAlpine,
their specialized habits. Because of the late stage at which 1987; Ozerov, 2000), and that classification is followed here.
adults appear on carrion, they are rarely taken by general J.F. McAlpine (1977) revised the generic classification and
collectors. provided a phylogenetic analysis, although D.K. McAlpine
The larvae of at least P. casei, commonly known as the (1989) questioned whether some genera and lower clades
cheese skipper, Centrophlebomyia furcata, the bone skip- were supported by synapomorphies. The classification of
per, and Protothyreophora grunini Ozerov are able to jump J.F. McAlpine (1977), as slightly modified by Ozerov (2000),
or skip (Freidberg, 1981; Ozerov, 2000). This remarkable recognizes 23 genera in two subfamilies: Neottiophilinae, with

6 Lasiopiophila

7 Lasiopiophila 8 Lasiopiophila

Figs. 64.6–8. Larva: lateral view of larva (6), lateral view of anterior spiracle (7), and posterolateral view of terminal segment (8) of
Lasiopiophila pilosa (Staeger), (Holarctic, MND, figs. 29–31).
868
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

two genera and four species; and Piophilinae, with the three The Piophilidae is a small, but cosmopolitan family.
tribes Mycetaulini (6 genera; 15 species), Piophilini (8 gen- There are only 72 described species worldwide, although
era; 41 species), and Thyreophorini (six genera; nine species), J.F. McAlpine (1989) indicated that he had seen 15 addi-
and one unplaced genus (one  species). The Thyreophorini tional undescribed Nearctic species. The greatest species
was considered a subtribe of Piophilini by J.F.  McAlpine diversity is in the temperate and boreal parts of the Holartic
(1977). Only the Thyreophorini (one genus, two species) and Region.
Piophilini (four genera, eight species) are represented in the
Neotropical Region. Based on a study of the four Australian Identification
species of Piophilini, D.K. McAlpine (1989) considered Pro-
topiophila a synonym of Piophila, which if justified, would J.F. McAlpine (1977) provided keys to most of the known
probably require the other genera of Piophilini to also be genera and species. J.F. McAlpine (1987) and Ozerov (2000)
considered synonyms of Piophila. Pending a comprehensive, provided keys to the Nearctic and Palearctic genera, respec-
phylogenetic analysis of the group, we continue to follow the tively. D.K. McAlpine (1989) keyed the Australian species,
J.F. McAlpine / Ozerov classification. and Merz (1996) keyed the Swiss species.

Key to the genera of Piophilidae of the Neotropical Region

1. Head (Fig. 3) with two vibrissae; frons with setulae near anterior margin larger, sometimes forming
transverse row; scutellum at least 0.8 times as long as wide, strongly flattened, with numerous
fine setulae on margins and disk in addition to marginal setae (Fig. 4); anepisternum with strong
posterior seta in addition to setulae; Brazil, Guyana, Peru . . . . . . . . . [Bocainamyia Albuquerque]
– Head (Figs. 1, 2) with one vibrissa; frons bare anteriorly or with setulae along anterior margin no
larger than other setulae; scutellum at most 0.75 times and usually less than 0.66 times as long
as wide, flattened to convex, without setulae (with only normal two pairs of marginal setae);
anepisternum without seta posteriorly.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2. Scutum with presutural supra-alar seta well developed, and with one presutural and three postsutural
dorsocentral setae (sometimes only posteriormost well developed) (Fig.  5); postpronotal lobe
with two well-developed setae, the medial one inclinate. . . . . . . . . . . . . . . . . Protopiophila Duda
– Scutum with presutural supra-alar seta weak or absent, and with only one postsutural dorsocentral
seta; postpronotal lobe without outstanding setae, not more than twice as long as setulae.. . . . . 3

3. Anepisternum bare; frons entirely dark brown.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stearibia Lioy

– Anepisternum finely setulose (Fig. 1); frons with anterior margin yellow or orange.. . . . . . . . . . 4

4. Scutum uniformly setulose, setulae not in isolated rows. . . . . . . . . . . . . . . . . . . Prochyliza Walker

– Scutum largely bare, setulae restricted to margins, one medial acrostichal row, and two dorsocentral
rows (Fig. 1). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Piophila Fallén

Synopsis of the fauna Prochyliza Walker. Three of the eight species of this Ho-
larctic and Neotropical genus are Neotropical. Of these, only
Only five genera and 10 species of Piophilidae are known P. azteca McAlpine is known from Central America. It is re-
from the Neotropical Region, and two of the genera and corded from southern Mexico (Chiapas, San Cristobal), Costa
three of the species are not native. Four genera and four spe- Rica (Cartago; Estación Cuerici; Estación La Casona; Monte-
cies are known from Central America, only two of which are verde; Zurquí), and Panama (Cerro Punta) and appears to be
native. The species in Central America have been found at restricted to mid- to high elevations (1500–3000 m). Prochyliza
various elevations, from nearly at sea level to 3000 m. inca McAlpine is known only from Ecuador, whereas P. nigri-
mana (Meigen) is a Holarctic species that has been introduced
Piophila Fallén. The only species occurring in the Neotropi- to South America (Argentina, Chile). J.F. McAlpine (1977) pro-
cal Region is the nearly cosmopolitan P. casei, a pest of stored vided a key to the known species. The biology of the native spe-
meat, cheese, and hides, which was presumably spread by man. cies is unknown, but P. nigrimana has been bred from bones
In the Neotropical Region, P. casei has been reported from Ar- and decaying leaves in Europe.
gentina, Bolivia, Brazil, Chile, Costa Rica, Cuba, Ecuador, Ja-
maica, Mexico, Paraguay, and Venezuela (Steyskal, 1967), and Protopiophila Duda. Three of the 11 known species of
we have seen specimens from Guatemala, Panama, and Peru. this cosmopolitan genus are Neotropical (J.F.  McAlpine,
PIOPHILIDAE (skipper flies) 64 869

1977), of which one is known from Central America. Pro- discarded cervid antlers and the Holarctic P. latipes from car-
topiophila atrichosa McAlpine was described from Peru rion and the corpse of a cat (Ozerov, 1989; Bonduriansky,
and Belize (St. Hermans Cave, 120 m elevation) and is also 1995).
known from Costa Rica (Parque Nacional Corcovado, San
Pedrillo). Protopiophila nigriventris (Curran) is known Stearibia Lioy. The single known species, S.  nigriceps
from Guyana and P. pallida McAlpine from Brazil, Guyana, (Meigen), is known from Costa Rica (Estación Magsasay;
and Peru. All three species appear to be restricted to natu- Estación Sirena; San José). It is otherwise widespread in
ral areas at low elevations. J.F.  McAlpine (1977) provided the Holarctic Region and South America (Argentina, Chile,
a key to most of the described species, which was emended Colombia, Ecuador, Peru) and has probably been spread by
by Bonduriansky (1995). Little is known of the biology of human activity. The Costa Rican specimens have entirely
the Neotropical species of Protopiophila. The paratype of yellow mid- and hind legs, but appear to be conspecific with
P.  atrichosa from Belize was collected in a bat cave. The those from other areas, which have at least the femora par-
Nearctic species P. litigata Bonduriansky has been bred from tially brown.

Literature cited

Bonduriansky, R. 1995. A new Nearctic species of Protopiophila McAlpine, J.F. 1989. Phylogeny and classification of the Musco-
Duda (Diptera: Piophilidae), with notes on its behaviour and morpha, pp. 1397–1518. MND, Volume 3.
comparison with P. latipes (Meigen). The Canadian Entomolo- Merz, B. 1996. Die Piophilidae (Diptera) der Schweiz mit Bes-
gist 127: 859–863. chreibung einer neuen Art. Mitteilungen der Schweizerischen
Ferrar, P. 1987. A guide to the breeding habits and immature stages Entomologischen Gesellschaft 69: 345–360.
of Diptera Cyclorrhapha. Entomonograph 8(1). E.J.  Brill  / Oldroyd, H. 1964. The Natural History of Flies. Weidenfeld & Ni-
Scandinavian Science Press, Leiden, Copenhagen, 478 pp. colson, London, 324 pp.
Freidberg, A. 1981. Taxonomy, natural history and immature stag- Ozerov, A.L. 1989. Kizucheniyu nekrobiontnykh dvukrylykh
es of the bone-skipper, Centrophlebomyia furcata (Fabricius) Dal’nego Vostoka SSSR [Study of carrion flies of the Far East
(Diptera: Piophilidae, Thyreophorina). Entomologica scandi- of the USSR], pp. 114–142. In Shatalkin, A.I. (editor). Naseko-
navica 12: 320–326. mye v ekosistemakh Sibiri i Dal’nego Vostoka. Trudy Zoo-
Korneyev, V.A. 1999. Phylogenetic relationships among the fami- logicheskogo muzeya MGU [Archives of Zoological Museum
lies of the superfamily Tephritoidea, pp. 3–22. In Aluja, M. & Moscow State University] 27: 1–192.
A.L. Norrbom (editors). Fruit flies (Tephritidae): Phylogeny and Ozerov, A.L. 2000. Family Piophilidae, pp. 355–365. In Papp, L. &
evolution of behavior. CRC Press, Boca Raton, [16] + 944 pp. B. Darvas (editors). Contributions to a manual of Palaearctic
McAlpine, D.K. 1989. A synopsis of the Australian Piophilidae Diptera (with special reference to flies of economic importance).
(Diptera: Schizophora). General and Applied Entomology 21: Appendix. Science Herald, Budapest, 604 pp.
17–24. Soós, Á.  & L. Papp. 1984. Catalogue of Palaearctic Diptera,
McAlpine, J.F. 1977. A revised classification of the Piophilidae, Vol. 9. Micopezidae — Agromyzidae. Akadémiai Kiadó, Buda-
including ‘Neottiophilidae’ and ‘Thyreophoridae’ (Diptera: pest, 460 pp.
Schizophora). Memoirs of the Entomological Society of Canada Steyskal, G.C. 1967. 68. Family Piophilidae. In Papavero, N. (edi-
103: 1–66. tor). A Catalogue of the Diptera of the Americas South of the
McAlpine, J.F. 1987. Family Piophilidae, pp.  845–852. MND, United States. Departamento de Zoologia, Secretaria da Agri-
Volume 2. cultura, São Paulo, 2 pp.
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
RICHARDIIDAE (richardiid flies) 65
E. Geoffrey Hancock

Fig. 65.1. Male of Richardia eburneosignata Hennig, (MND, fig. 67.1).

Diagnosis long coiled phallus. Females with piercing-style ovipositor

typical of tephritoid group; two spermathecae present.
Small to medium-sized flies (body length 3–15  mm) Some Ulidiidae that do not have the usual acute apical ex-
(Fig. 1). Head with single orbital seta. Stalk-eyes or genal ex- tension to cup (Figs. 66.83–86) are likely to be confused with
tensions present in some Richardia. Frons may be consider- richardiids. Two orbital setae distinguish ulidiids. Platysto-
ably extended or swollen; occiput depressed above neck, but matids may be difficult to separate on the basis of orbital
in Sepsisoma, for example, enlarged behind eyes. Body color setae but do not have a break at subcosta. Sepsidae can be
yellow to brown or black, sometimes with darker or pruinose confused with some darker, antlike richardiids in unsorted
bands or stripes on scutum. Dorsum of thorax often setulose; collections, but sepsids usually have clear wings with a dis-
thoracic chaetotaxy not extensive, presutural dorsocentrals tinct spot at tip of R2+3 and male legs often heavily adorned
absent. Scutellum with two or four marginal setae; if four with bristles, tubercles, or emarginations. Male sepsids do
then one pair basolateral. Postmetacoxal bridge complete in not have a long coiled phallus, and females do not have a
many species. Wings with one subcostal break, sometimes piercing tephritoid-type ovipositor.
with humeral break also (in Automola); cup closed apically
by short sinuous or slightly convex vein. Wings frequently Biology
with apical spot, median band, basal and other markings.
Legs often banded and usually bearing ventral femoral bris- The majority of known larvae of Richardiidae are
tles on all or some legs except in Epiplateinae and some saprophagous, found in decaying vegetable matter. Published
Setellia spp. Two basal abdominal tergites fused and may be data associate them with the bracts or flowers of Heliconia
narrowed (pedunculate) or tapering basally more gradually. (Musaceae) (Seifert & Seifert, 1976), Xanthosoma (Arace-
Usually strong preapical bristles present laterally on tergite 2 ae) (Kitching, 2000), and diseased coconut palms (Steyskal,
(see Fig. 7). Some species with metallic blue or green abdo- 1958). Adults of Melanoloma decrepita Hendel have been
men contrasting with thorax. Male genitalia complex, with found in dung-baited traps, and Richardia  spp. are greatly

871
872
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

cerc

tg 8

2 Automola 3 Epiplatea

hum brk sc brk

R1
R2+3

R4+5
dm
cup
M1

5 Coniceps
A1+CuA2 CuA1

4 Automola

6 Coniceps

8 Setellia 7 Setellia

9 Odontomera

Figs. 65.2–9. Female terminalia, wings, head, and abdomen: dorsal view of apex of female abdomen of (2) Automola rufa Cresson, (Nearctic,
MND, fig. 67.17); dorsal view of wing of (3) Epiplatea sp., (Steyskal, 1958, pl.2, fig. 14); and (4) A. rufa, (Nearctic, MND, fig. 67.2); lateral
view of head of (5) Coniceps niger Loew, (MND, fig. 67.13); dorsal view of wing of (6) C. niger, (MND, fig. 67.4); dorsal view of abdomen
of (7) Setellia apex Hendel, (Hendel, 1911a, fig. 22); dorsal view of wing of (8) S. costalis Schiner, (Hendel, 1911a, fig. 28); and
(9) Odontomera ferruginea (Macquart), (MND, fig. 67.6).
Abbreviations: cerc, cercus; hum brk, humeral break; sc brk, subcostal break; tg, tergite.
RICHARDIIDAE (richardiid flies) 65 873

attracted to fruit baits. Inouye (1999) found Richardia sp. as are regarded as basal within the Tephritoidea, a position
one of four dominant flies in fallen fruit at La Selva, Costa shared with Pallopteridae and Piophilidae. The Epiplateinae
Rica. These data probably reflect the relative attractiveness and Richardiinae are the only two subfamilies, with no fur-
of the rotting material for oviposition. The larvae of a species ther subdivisions. The former includes three genera and the
of Sepsisoma feed on the extrafloral nectaries of some shrubs latter 26 genera as listed by Steyskal (1968), who updated an
in Brazil (Pie & Del-Claro, 2002). earlier catalog (Aczél, 1950).
There have been a number of instances in which richar-
diids occur as pests. Larvae of Sepsisoma erythrocephalum
(Schiner) that burrow into stems are responsible for ‘dead Identification
heart’ damage to growing points of grasses (Deeming, 1985).
In association with fruit growing, particularly pineapple, There has been no published key to all Neotropical genera
Melanoloma viatrix Hendel can have a significant effect on of Richardiidae since Hendel (1911a, 1911b). Draft, unpub-
crop yields (Peñaranda  & Ospina, 1995). With its sibling, lished keys were drawn up and used by Steyskal (manu-
M. canospila Hendel, adults have been identified after rear- script., undated), which were adapted from Hendel’s keys,
ing from damaged fruit in various parts of South America, al- adding genera subsequently described. There are extralimital
though they have not yet been recorded in Central America. treatments that work only for North America (Curran, 1934;
Few immature stages are known, but larvae or puparia Steyskal, 1987). The key presented here utilizes a number
have been figured for Epiplatea hondurana Steyskal (Stey- of standard morphological features. Important characters in-
skal, 1958) and Melanoloma viatrix (Peñaranda & Ospina, clude ventral spines on the femora, relative position of ocel-
1995). lar triangle and the orbital seta, position of crossveins and
Adults of some species such as Automola atomaria other venational features. Examination of the profile of the
(Wiedemann) have been observed lekking on the surface head in lateral view is necessary for couplets dealing with a
of tree trunks, and such behavior is described also in Setell- protruding occiput behind the vertex as opposed to a reduc-
ia spp. (Pie, 1998). It is likely that other species indulge in tion in that area, leaving the lower part relatively wider. The
similar mating rituals if they have prominent wing markings presence of a postmetacoxal bridge has some value.
or sexual dimorphism in head structures, as found in species Careful card pointing or micro-pinning and then staging
such as Richardia telescopica Gerstaecker. Adult behavior are suitable for preserving dry adults. Direct pinning with
in an ant-mimicking Sepsisoma sp. has been studied (Pie & fine long pins should not damage characters except with
Del-Claro, 2002). The flies successfully use their behavior the smaller species. Reared specimens should be kept alive
and appearance, closely resembling the locally common long enough for the integument to harden and develop full
Camponotus ants, to protect extrafloral nectaries that are pigmentation. Puparia complete with the caps forced off dur-
used as a resource by their larvae. ing eclosion should be carefully preserved with the adults.
Adults can be found on various plant surfaces, displaying, The best option for this is to place each puparium in a gela-
mating, or ovipositing. They occur in baited and Malaise trap tin capsule and put them on the same pin as the adult that
samples in most habitats but are not found so frequently at emerged from it. Some of the mature larvae from a reared
higher elevations. batch could be preserved in 80% alcohol after killing in hot
water to render them relaxed and show features of value for
Classification study. Fluid preservation is an option for adults until pinning
can be achieved at a later date after careful drying to avoid
Of the tephritoid Diptera families, the Richardiidae are shrinkage, such as the use of critical point drying techniques,
unique in being restricted to the New World. Richardiidae if available.

Key to the genera of Richardiidae of the New World

1. Femora lacking spiny ventral setae (although forefemur may have one strong setae in Setellia
unispinosa Bigot, or a small group of fine setae in Automola rufa and Schnusimyia); Sc complete
to costal margin; male phallus tip simple, except in Setellia (see couplet 2 below); female cerci
separate (Fig. 2) or fused.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Femora with spiny ventral setae on at least one pair (Fig. 1); Sc sometimes interrupted adjacent to
costal break; male with phallus tip complex; female cerci fused, adapted for piercing; RICHAR-
DIINAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

2. Postmetacoxal bridge complete; male with phallus tip complex; female cerci fused; RICHARDIINAE.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Setellia Robineau-Desvoidy
– Postmetacoxal bridge incomplete (i.e., non-chitinized membrane visible above hind coxa); male
with phallus simple at tip (at most slightly specialized); female ovipositor with separate cerci;
EPIPLATEINAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3. One pair of scutellar setae present; wing either broad with large pigmented area bearing clear
spots (Fig. 17) or with four transverse bands. . . . . . . . . . . . . . . . . . . . . . . . . . Schnusimyia Hendel
– Two pairs of scutellar setae present; wing pattern not as above . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4. Proepisternal seta absent; vein R1 not setose (Fig. 3) . . . . . . . . . . . . . . . . . . . . . . Epiplatea Loew

– Proepisternal seta well developed; R1 setose or not. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

5. R1 setose; costa finely and uniformly clothed with proclinate hairs; wings with conspicuous
pattern, including complete brown fascia (Fig. 4). . . . . . . . . . . . . . . . . . . . . . . . . Automola Loew
– R1 not setose; costa with fine proclinate hairs and several more widely spaced stronger setae; wing
pattern of infuscated costal cells and brown spot at tip of R3; USA . . . . . . . [Omomyia Coquillet]

6. Head twice as long as high or projecting beyond eye for at least three-quarters eye diameter
(Fig. 5). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
– Head little or no longer than high (may project up to three-quarters eye length in Macrostenomyia,
but in which case ocellar triangle equilateral). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

7. Ocellar triangle and fronto-orbital setae behind middle of eye; vein A1+CuA2 reaches wing margin
(Fig. 6); USA. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Coniceps Loew]
– Ocellar triangle anterior to middle of eye; vein A1+CuA2 shortened. . . . . . . . . . . . . . . . . . . . . . . 8

8. Ocellar triangle equilateral; frons inflated anteriorly. . . . . . . . . . . . . . . . . . . . Oedematella Hendel

– Ocellar triangle elongate; frons not inflated. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

9. Fronto-orbitals nearly level with hind ocelli or actually behind them (i.e., nearer vertex); South
America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Ozaenina Enderlein]
– Fronto-orbitals level with anterior ocellus; Brazil. . . . . . . . . . . . . . . . . . . . [Oceanicia Enderlein]

10. Hind femur without ventral spines; abdomen petiolate (Fig. 7); wing apically or marginally dark-
ened (Fig. 8). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Setellia Robineau-Desvoidy
– Hind femur with one or more ventral spines; abdomen various; wing clear or patterned. . . . . . 11

11. Hind femur more swollen than others (Fig. 1). . . . . . . . . . . . . . . . . Richardia Robineau-Desvoidy
– Hind femur not more swollen than fore- and midfemora . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

12. Vein A1+CuA2 shortened, not reaching wing margin even as fold (Fig. 9). . . . . . . . . . . . . . . . . 13
– Vein A1+CuA2 reaching margin at least as fold (as in Fig. 13) . . . . . . . . . . . . . . . . . . . . . . . . . . 18

13. Head without orbital setae or lateral vertical setae; size large, body length 15 mm, wing length
11 mm; Dominican Republic. . . . . . . . . . . . . . . . . . [Johnrichardia Perez-Gelabert & Thompson]
– Head with orbital and lateral vertical setae; size smaller, body length less than 10 mm, wing length
less than 6 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
RICHARDIIDAE (richardiid flies) 65 875

10 Odontomera 11 Sepsisoma 12 Macrostenomyia

13 Hemixantha 14 Setellida

16 Neoidiotypa

17 Schnusimyia

15 Euolena

Figs. 65.10–17. Heads, wings, and adult: lateral view of head of (10) Odontomera ferruginea (Macquart), (MND, fig. 67.10); (11) Sepsisoma
flavescens Johnson, (MND, fig. 67.11); and (12) Macrostenomyia guerinii (Bigot), (Curran, 1934, fig. 1); dorsal view of wing of
(13) Hemixantha flavicornis Wiedemann, (South America, Hendel, 1911a, fig. 50); and (14) Setellida caerulescens Hendel, (Hendel, 1911a,
fig. 77); lateral view of adult of (15) Euolena egregia (Gerstaecker), (Hendel, 1911a, fig. 64); dorsal view of wing of (16) Neoidiotypa
appendiculata (Loew), (Hendel, 1911a, fig. 63); and (17) Schnusimyia parvula (Hendel), (South America, illustrated by G. Hancock).
876
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

14. Crossveins r-m and dm-cu closer together than length of dm-cu; twp  pairs of scutellar setae
present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Antineuromyia Hendel
– Crossveins r-m and dm-cu further apart than length of dm-cu; with one apical pair of scutellar
bristles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

15. Occiput in lateral view as wide dorsally as ventrally (i.e., equally wide above and below insertion
into thorax, Fig. 11). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sepsisoma Johnson
– Occiput much narrower dorsally (i.e., wider below neck, Fig. 10). . . . . . . . . . . . . . . . . . . . . . . 16

16. Ocellar triangle nearly equilateral, well anterior to posterior eye margin; vertex produced anteriorly
up to three-quarters eye width (Fig. 12). . . . . . . . . . . . . . . . . . . . . . . . . . . Macrostenomyia Hendel
– Ocellar triangle elongate, more or less level with posterior eye margin. . . . . . . . . . . . . . . . . . . 17

17. Abdomen petiolate (as in Figs. 7 and 15) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Odontomera Hendel

– Abdomen oval; Brazil . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Odontomerella Hendel]

18. Crossveins r-m and dm-cu not further apart than length of dm-cu (Fig. 13), usually much closer,
often with stump vein(s) on R2+3 (as in Fig. 16). . . . . . . . . . . . . . . . . . . . . . . . . Hemixantha Loew
– Crossveins r-m and dm-cu further apart than length of dm-cu (sometimes by nearly the same
length as in Euolena, Fig. 15); stump veins present or absent. . . . . . . . . . . . . . . . . . . . . . . . . . . 19

19. Abdomen slender, petiolate, two basal segments fused and much narrower than following (as in
Figs. 7 and 15). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
– Abdomen not petiolate, gradually narrowed towards base, relatively short and broad. . . . . . . . 25

20. Crossvein r-m basal to middle of cell dm; cell  r4+5 noticeably widened in middle of lower
side, being significantly wider at that point than cell above but narrowing again towards apex
(Fig. 14). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Setellida Hendel
– Crossvein r-m at or apical to middle of cell dm; cell r4+5 not significantly wider at midpoint and
parallel-sided in distal half. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

18 Poecilomyia 19 Coilometopia

20 Melanoloma

Figs. 65.18–20. Heads and wing: lateral view of head of (18) Poecilomyia longicornis Hendel, (Hendel, 1911a, fig. 43); anterior view of
head of (19) Coilometopia trimaculata (Fabricius), (Hendel, 1911a, fig. 40); dorsal view of wing of (20) Melanoloma affinis Loew, (South
America, Hendel, 1911a, fig. 53).
RICHARDIIDAE (richardiid flies) 65 877

21. Calypteres and alula lacking, wing cuneate at base; Brazil, Peru. . . . . . . [Spheneuolena Hendel]
– Calypteres and alula well developed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

22. R1 ending before midwing. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

– R1 ending at midwing. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

23. Occiput swollen more below neck than above; not antlike flies; legs, especially in males, long and
slender (Fig. 15). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Euolena Loew
– Occiput equally or more swollen above neck than below; antlike flies with legs of normal length;
Bolivia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Acompha Hendel]

24. All femora spinose ventrally; R2+3 usually with stump vein(s) (Fig. 16); wasplike flies with costa
broadly darkened; wing lacking medial spot. . . . . . . . . . . . . . . . . . . . . Neoidiotypa Osten Sacken
– Only hind femur with spines; R2+3 without stump vein; body not wasplike; costa not darkened;
wing with large medial spot; Mexico. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Panermya Wulp]

25. R2+3 doubly sinuate at tip; thorax relatively long and high compared with head and abdomen; male
costa expanded beyond midpoint; Ecuador, Peru . . . . . . . . . . . . . . . . . . [Megalothoraca Hendel]
– R2+3 not undulate; thorax smaller; male costa not expanded. . . . . . . . . . . . . . . . . . . . . . . . . . . . 26

26. Eyes protruding; head wider than anteriorly tapering thorax; r-m distal to middle of cell  dm;
Peru . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Batrachophthalmum Hendel]
– Eyes not protruding; head not wider than rounded or quadrate thorax, or r-m near middle of
cell dm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27

27. Basal antennal segment lengthened, as long as second; M straight basal to r-m. . . . . . . . . . . . . 28
– Basal antennal segment short, more or less hidden; M basal to r-m usually curved posteriorly
(Fig. 20). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

28. Four scutellar bristles present; frons with anterior ocellus on broad elevation (Fig.  18); R2+3
without stump vein. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Poecilomyia Hendel
– Two scutellar bristles present; frons flat; R2+3 with stump veins; Brazil . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Cladiscophleps Enderlein]

29. Frons more or less concave between eyes, broader than width of eye (Fig. 19); r-m at or before
middle of cell dm; tip of R1 at same level . . . . . . . . . . . . . . . . . . . . . . . . . Coilometopia Macquart
– Frons flat, level with eyes; r-m before or after middle of cell dm; tip of R1 basal to r-m . . . . . . 30

30. Crossvein r-m at or basal to middle of cell dm . . . . . . . . . . . . . . . . . . . . . . . . Beebeomyia Curran

– Crossvein r-m apical to middle of cell dm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

31. Mid- and hind tibiae thickened; midfrontal stripe black, tomentose; Bolivia. . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Maerorichardia Hennig]
– Tibiae not thickened; midfrontal stripe bare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32

32. Abdomen slender; M not curved posteriorly into cell dm before r-m; Brazil, Peru . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Richardiodes Hendel]
– Abdomen oval, broad basally; M curved posteriorly into cell dm (Fig. 20). . Melanoloma Curran
878
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Synopsis of the fauna Neoidiotypa Osten Sacken. The only described species of
this genus, N. appendiculata (Loew), is from Cuba. The ge-
I estimate that there are over 50 species of Richardiidae in nus appears to be rare, but three or four possible undescribed
Central America placed in 18 genera. Of these, only 15 spe- species are represented by specimens from Costa Rica.
cies in eight genera have been mentioned in publications; the
increase is based on examination of museum collections. Odontomera Hendel. Eighteen species are listed in the
The most recent checklist (Steyskal, 1968) gives infor- checklist, of which one is Nearctic and three are from Cen-
mation on the Richardiidae, and no new species have been tral America, including Costa Rica. One of these, O. nitens
described from Central America since then. (Schiner) is represented in INBio collections, and other pos-
sible undescribed species are present. It is unknown if O. ba-
Antineuromyia Hendel. In addition to the two known
from South America, one undescribed species of this genus sistriga (Walker) and O.  ferruginea Macquart have been
is known from Costa Rica. collected in recent years.

Automola Loew. Two of the three known species occur in Oedematella Hendel. The single species, O. czernyi Hen-
the region: A. atomaria, which is found also in the Nearctic del, formerly recorded from Peru is now known from Central
Region, and A.  caloptera (Bigot). A key to the species of America.
Automola is provided by Steyskal (1958).
Poecilomyia Hendel. The single species, P.  longicornis
Beebeomyia Curran. None of the four species in the Hendel, previously known from Panama, is represented by
checklist (Steyskal, 1968) are listed from Central America. one specimen in INBio. Two other taxa mentioned in the
Several undescribed species are now recognized in Costa checklist are probably synonyms of this widespread and dis-
Rica from specimens reared from flowers and bracts of An- tinctive species.
thurium and Heliconia spp.
Richardia Robineau-Desvoidy. The largest and most
Coilometopia Macquart. Of the six known species,
C. trimaculata (Fabricius), C. costalis Hendel, and C. longi- characteristic genus, defined by their enlarged hind femora,
cornis Hendel are represented in the collections of INBio. contains some species with dramatic genal extensions or
No representatives of this genus were known previously stalk-eyes. Of the 29 known species, at least six or seven
from Central America. occur in Central America.

Epiplatea Loew. Two of the five described species are Schnusimyia Hendel. Two undescribed species of this
known from Central America, and a key is available to them genus are known from Costa Rica. The strongly patterned
(Steyskal, 1958). wings are superficially reminiscent of some other tephritoid
family representatives with which they have been confused
Euolena Loew. One species, E. egregia (Gerstaecker), is
in the past.
widely distributed in South America and now has been found
as far north as Costa Rica. Other probably undescribed spe- Sepsisoma Johnson. Prior to the checklist (Steyskal,
cies also occur in Central America.
1968), no species of Sepsisoma were recorded from Cen-
Hemixantha Loew. The checklist (Steyskal, 1968) has tral America even though representatives of the genus were
one out of 19 species recorded from Costa Rica. There are at known from both South and North America. Three spe-
least five or six species represented in museum collections, cies have been seen in museum collections from Central
including INBio. A key to Hemixantha spp. is provided by America.
Hennig (1937).
Setellia Robineau-Desvoidy. There are 19 species listed
Macrostenomyia Hendel. The single species M. guerinii in the checklist, of which one is recorded previously from
(Bigot) is known from Central America and the West Indies. Costa Rica and three others from elsewhere in Central Amer-
Melanoloma Curran. None of the 11 species in the ica. At INBio, there are examples of five described species
checklist (Steyskal, 1968) were listed from Central America, and a possible new species close to S. apex Hendel, all from
but at least two species are well represented in the INBio Costa Rican localities.
collections. Specimens appeared in numbers in dung-baited
traps in Osa, Costa Rica, in 2001. Two species, M.viatrix Setellida Hendel. The only species, S. caerulescens Hen-
and M. canospila, are known as pests of pineapple crops in del, is known from Panama, but there are also numerous
South America. specimens in the INBio collections from Costa Rica.
RICHARDIIDAE (richardiid flies) 65 879

Literature cited

Aczél, M.L. 1950. Catalogo de la subfamilia Americana «Rich- Peñaranda, E.A.  & M.A.O. Ospina. 1995. Consideraciones gen-
ardiinae» (Dipt. Acalyp. «Otitidae»). Acta Zoologica Lil- erales sobre Melanoloma viatrix Hendel, una nueva plaga de la
loana 9: 5–47. piña. Revista Colombiana de Entomología 21: 1–8.
Curran, C.H. 1934. Family Otitidae — the pictured-wing flies, Pie, M.R. 1998. Lek behaviour as the mating strategy of Setell-
pp.  271–284. In Curran, C.H. The families and genera of ia sp. (Diptera: Richardiidae). Journal of Insect Behaviour 11:
North American Diptera. Published by the author, New 823–832.
York, 512 pp. Pie, M.R. & K. Del-Claro. 2002. Male–male agonistic behaviour and
ant-mimicry in a Neotropical richardiid (Diptera: Richardiidae).
Deeming, J.C. 1985. A description of the adult and imma-
Studies on Neotropical Fauna and Environment 37: 19–22.
ture stages of Ectecephala capillata (Coquillet) (Diptera:
Seifert, R.P.  & F.H. Seifert. 1976. Natural history of insects liv-
Chloropidae). Proceedings of the Entomological Society of ing in inflorescences of two species of Heliconia. Journal of the
Washington 87: 402–407. New York Entomological Society 84: 233–242.
Hendel, F. 1911a. Diptera, Fam. Muscaridae, Subfam. Rich- Steyskal, G.C. 1958. Notes on the Richardiidae, with a review of
ardiinae. Fascicle 113, pp  1–56. In Wytsman,  P. (editor). the species known to occur in the United States (Diptera, Aca-
Genera Insectorum. Bruxelles. lyptratae). Annals of the Entomological Society of America 51:
Hendel, F. 1911b. Die Arten der Dipteren-Subfamilie Richardi- 302–310.
inae. Deutsche Entomologische Zeitschrift 1911: 181–212; Steyskal, G.C. 1968. 53. Family Richardiidae. In Papavero, N. (edi-
239–270; 367–396. tor). A Catalogue of the Diptera of the Americas South of the
Hennig, W. 1937. Beiträge zur Systematik der Richardiiden. United States. Departamento de Zoologia, Secretaria da Agri-
Revista de Entomología 7: 306–312. cultura, São Paulo, 25 pp.
Inouye, B.D. 1999. Integrating nested spatial scales: impli- Steyskal, G.C. 1987. Richardiidae, pp. 833–837. MND, Volume 2.
cations for the coexistence of competitors on a patchy re- Steyskal, G.C. (manuscript undated) [Manuscripts on Richardiidae
source. Journal of Animal Ecology 68: 150–162. by Steyskal preserved at the Smithsonian Institution include
Kitching, R.L. 2000. Food webs and container habitats, the draft keys to both genera and species. To some extent he also
natural history and ecology of phytotelmata. Cambridge examined types, and these notes provide much information in
University Press, Cambridge & New York, xiii + 431 pp. addition to published data.]
880
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
ULIDIIDAE (picture-winged flies) 66
Elena P. Kameneva and Valery A. Korneyev

Fig. 66.1. Female of Euxesta stigmatias Loew, (illustrated by V. Korneyev).

Diagnosis with a pattern of dark spots or bands, rarely completely hya-

line; Sc vein complete, C with distinct humeral break, sub-
Medium-sized (body length 2–14  mm, most commonly costal break indistinct. Legs moderately developed, usually
4–7 mm long) flies with moderately elongate body (Fig. 1); without rows of spines or strong setae; tibiae without dorsal
some species that mimic ants with elongate or somewhat peti- setulae or setae. Females, like in all other Tephritoidea, with
olate abdomen, while some species associated with cacti with telescopic ovipositor, whereas males with long phallus, hid-
body short and stout, resembling bugs (Heteroptera). Color- den in membranous pocket on ventral side of abdomen and
ation yellow to brown or black with pattern of microtrichose bearing no glans at apex.
spots or bands, often with metallic blue or green sheen. Head Central American picture-winged flies are usually recog-
variously shaped; inner and outer vertical, ocellar, divergent nized by the distinctive wing patterns, which are often similar
postocellar and orbital (one or two, anterior one shortest) to those of fruit flies and other tephritoid families, lauxani-
setae present; frontal setae absent (except in Chaetopsis ids, snail-killing flies, and some drosophilids. Ulidiidae dif-
and most Aciuroides). Frons sometimes pitted or wrinkled. fer from tephritids by their complete vein  Sc, and lack of
Clypeus moderately large. Proboscis with moderately devel- presutural supra-alar (except Dyscrasis) and anepimeral seta
oped prementum. Thorax usually longer than broad. Wings (except Diacrita). Richardiidae and Platystomatidae have

881
882
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

cell cup closed by an arcuate or straight crossvein, whereas Classification

in Ulidiidae this crossvein is sinuate, forming an acute pos-
teroapical lobe (except in Eumecosomyia and Aciuroides); Ulidiidae are the third largest family, after Tephritidae
however, richardiids usually possess well-developed pre- and Platystomatidae, of the superfamily Tephritoidea. Together
sutural supra-alar seta, and Platystomatidae have distinc- with these two families and Pyrgotidae, they form a mono-
tive anepimeral setae; Schnusimyia Hendel (Richardiidae), phyletic group within Tephritoidea, the so-called “higher
which sometimes was assigned to Ulidiidae, can be sepa- Tephritoidea”, which differs from the “lower Tephritoidea”
rated with certainty by its genital characters, and abdominal by completely reduced spiracles  6 and 7 and setae on ab-
spiracles 6 and 7 in the male are well developed. Like other dominal sternite  6 of males. Monophyly of the family is
tephritoids, Ulidiidae never have rows of strong setae on the supported by the absence of the presutural supra-alar seta
dorsum of tibiae, distinguishing them from Lauxaniidae and (synapomorphy) in combination with the absence of anepi-
Sciomyzidae. Drosophilids with wing patterns usually have meral setulae and setae (symplesiomorphy differentiating
cell bm open. Ulidiidae from other higher Tephritoidea families). Nomen-
clature of the family has been controversial throughout the
history of its study. The family name Ortalidae was often
Biology used for this family or for part of it, including Platystoma-
tidae and Richardiidae. Since Ortalis Meigen was found to
Larvae of most Ulidiidae are saprophagous. Larvae of be a preoccupied name, the name Otitidae was proposed
Holarctic Pseudotephritis Johnson, Myennis Robineau- instead (Aldrich, 1932). Both names Ulidiidae and Otitidae
Desvoidy (Otitinae: Myennidini), Pseudoseioptera Stackel- remained in use before the priority of the first was noted
berg, and Homalocephala Zetterstedt (Ulidiinae: Seiopterini) (Kameneva  & Korneyev, 1994). The most comprehensive
live under bark of fallen poplars, aspens, larches, and pines and detailed classification based upon phylogenetic analysis
(Krivosheina & Kovalev, 1972; Krivosheina & Krivosheina, was proposed by Kameneva & Korneyev (2006). The family
1995b, 1997a, 1997b; Lukasheva, 1987). Flies of the tribe is divided into two subfamilies: Ulidiinae Macquart and Oti-
Lipsanini (Chaetopsis, Eumetopiella Hendel, Eumecosomyia, tinae Aldrich. The subfamily Ulidiinae includes four tribes:
Euxesta) are associated mostly with monocotyledons (cere- predominantly Neotropical Lipsanini Enderlein (= Euxesti-
als and palms), living in tissues damaged by other insects, nae Hennig, preoccupied name) and Pterocallini Loew,
young inflorescences, or new growth. Euxesta stigmatias Holarctic Seiopterini Kameneva & Korneyev, and the mostly
Loew, E.  mazorca Steyskal, and E.  eluta Loew infest ear Old World Ulidiini Macquart. The subfamily Otitinae con-
corn in Puerto Rico, Colombia, and Chile (App, 1938; Stey- sists of three tribes: Cephaliini Schiner, Otitini Aldrich, and
skal, 1974; Huepe et al., 1986). In Guatemala, maggots of Myennidini Kameneva & Korneyev. The majority of genera
Chaetopsis major (Wulp) were reported to be one of the most belonging to this subfamily occur in the Holarctic Region. A
destructive pests of seedling corn (Melhus  & Harris, 1949), few genera of Myennidini occur in South Africa and South
often killing buds, whereas E. stigmatias Loew, E.  sororcula America (Venezuela south to central Chile), whereas South
(Wiedemann), and Eumecosomyia nubila (Wiedemann) American genera of Cephaliini are predominantly Patago-
were reared from decaying corn silk (Barber, 1939; Painter, nian and Andean in distribution. The Otitini also contains
1955). North American Otitinae Cephaliini (Delphinia, Tritoxa a few representatives in the Oriental, Australasian, and
Loew) and the possibly related Idana Loew (? Otitini) Neotropical Regions.
either infest bulbs of liliaceous plants or live in rotting plant
matter (Allen & Foote, 1967, 1975). Tetanops (Eurycepha-
lomyia) myopaeformis (Röder) (Otitini) are pests of sugar
beet (Mahrt & Blickenstaff, 1979), and Pterotaenia fasciata Identification
(Wiedemann) (Otitinae: Cephaliini) cause economic damage
to cherries (Porter, 1922; Riesle, 1922; Malloch, 1933). Flies Hendel’s (1909a, 1909b, 1910) Pterocallinae and Ulidii-
of the genera Euacaina Steyskal, Notogramma, Diacrita, nae chapters in the Genera Insectorum series, together with
and Stictomyia Bigot are usually associated with rotting Hennig’s (1939, 1940) monographs on Palearctic Otitinae
cacti, although some Notogramma have also been reared and Ulidiinae are still the most important literature for iden-
from rotting tomatoes, immature coconuts, bananas, liver, tification of genera. Steyskal (1961, 1987) has provided the
wild tubers, watermelon, Solanum fruits, and fruits of Attalea most comprehensive treatment and a key to North Ameri-
palms (Steyskal, 1963, 1991). Larvae of Pseudeuxesta prima can genera, which includes some genitalic characters. He
Osten Sacken were found in a human corpse under natural also compiled a key to the genera of “Otitinae” (including
conditions (Bohart  & Gressitt, 1951). The most detailed Pterocallini and some Lipsanini) south of the United States
synopsis of existing biological data was given by Allen  & (Steyskal, 1982), the most diverse regional fauna of picture-
Foote (1967) and Ferrar (1987). However, larval habits of winged flies.
the majority of species remain unknown. Adults are often The following key is based upon a broad review of Neo-
attracted to various decaying matter, like feces, carrion, and tropical, mainly Central American, material involving
hydrolyzed proteins and compost. genitalic characters of both sexes for most species (Kameneva,
ULIDIIDAE (picture-winged flies) 66 883

2004a; E.P. Kameneva & V.A. Korneyev, in preparation). It is this key complete by discussing, where possible, unpublished
evident now that concepts of many genera need to be com- data on morphology, taxonomy, and distribution.
pletely revised, which cannot be done in the current chapter. The ratio of the second (between bm and r-m crossveins)
This is especially evident concerning Euxesta, the largest to the third (between r-m and dm crossveins) sections of the
genus in the Neotropical fauna. We tried, therefore, to make medial vein is abbreviated below as M2:M3.

Key to the genera of Ulidiidae of Central America

1. Vertex, scutum, or other body parts with greenish or bluish sheen, or body entirely yellow to
reddish-yellow without dark pattern . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Body without greenish or bluish sheen, brownish to black or yellow with darker pattern; if
abdomen with metallic blue sheen (rare), then additional crossvein present between pterostigma
and vein R2+3 or abdomen deeply pitted. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

2. Frons bare or with fine setulae only (Fig. 2); cell r4+5 almost or completely closed (Fig. 10); male
with phallus bare, with sclerotized subapical hooks; female with three  spermathecae;
ULIDIINAE: ULIDIINI. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Physiphora Fallén
– Frontal setae or setulae well developed (Figs. 3–7); cell r4+5 not apically closed (Figs. 8–9); male
with phallus setulose or bare, but without subapical sclerotized hooks (Fig.  24); female with
two spermathecae (Fig. 20); ULIDIINAE: LIPSANINI. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

fr s infr s
fr s

2 Physiphora 3 Euxesta
4 Chaetopsis

5 Euphara 6 Siopa
7 Acrosticta

Figs. 66.2–7. Heads of Ulidiinae: anterior view of (2) Physiphora clausa (Macquart) (antennae omitted), (Oriental); (3) Euxesta stigmatias
Loew; (4) Chaetopsis aenea (Wiedemann) (antennae omitted); (5) Euphara caerulea (Macquart) (antennae omitted); (6) Siopa longicornis
Hendel (right antenna omitted); and (7) Acrosticta profunda Hendel (right antenna omitted). Figures 2, 4 redrawn from Hendel (1910) by
V. Korneyev; Figures 3 and 5–7 illustrated by V. Korneyev.
Abbreviations: fr s, frontal seta; infr s, interfrontal setula.
884
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

3. Vein A1+CuA1 strongly reduced or absent (Fig. 8), if reaching halfway to wing margin, then cell
cup closed with straight or arcuate crossvein without triangular lobe along vein A1+CuA1; vein R1
bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
– Vein A1+CuA1 reaching posterior margin of wing; if short, then cell cup with triangular lobe along
vein A1+CuA1 or also vein R1 setulose (Figs. 31–39) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

4. Eye higher than long; clypeus low, hidden (Fig. 8). . . . . . . . . . . . . . . . . . . Eumecosomyia Hendel
– Eye longer than high; clypeus high, protruding (Fig. 27) . . . . . . . . . . . . . . . Undescribed Genus A

5. Frons with rows of strong setae along its sides, but completely bare medially (Fig. 4); first
flagellomere usually slightly pointed; lower margin of face somewhat produced; clypeus moderately
developed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetopsis Loew, in part
– Frons with setae in medial portion, at least above lunule (Figs. 3, 5); first flagellomere usually
rounded, if pointed, then prescutellar acrostichal seta absent; lower margin of face and clypeus
various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

6. Scutellum strongly flattened, its tip acute in profile; vertex meeting occiput at acute angle (Fig. 29);
scutum with numerous irregular stripes of coalescent dots. . . . . . . . . . . . . . . . Notogramma Loew
– Scutellum rounded; if vertex rarely acute, then scutum without stripes . . . . . . . . . . . . . . . . . . . . 7

7. Gena more than half eye height (Fig. 30); occiput concave in upper portion; scutum silver
microtrichose, with pattern of numerous round spots; Peru . . . . . . . . . . . . . . [Euacaina Steyskal]
– Gena less than half eye height (Figs. 26, 28); if occiput concave, then scutum uniformly shining
or microtrichose. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

8. Prescutellar acrostichal setae rudimentary or lacking; thorax and abdomen usually elongate,
scutum more than 1.4 times as long as wide; aculeus apex often wide and flattened (Fig. 19), if
narrow and round, then eye conspicuously longer than high. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
– Prescutellar acrostichal setae well developed; if absent, then thorax and abdomen robust; scutum
usually less than 1.4 times as long as wide; aculeus apex narrow and round (Fig. 21); eye as high
as, or higher than long (Fig. 28). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

9. Ventral portion of face strongly protruding anteriorly, in front of anterior margin of frons (Fig. 26);
aculeus various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
– Ventral portion of face receding or slightly protruding anteriorly (Fig. 25); aculeus apex flattened
truncate (Fig. 19). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

10. Eye higher than long (Fig. 26); scutellum flattened; aculeus apex wide, flat and pointed; anal lobe
narrower than or as wide as cell cup . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Zacompsia Coquillett
– Eye longer than high; scutellum strongly convex, protruding dorsally; aculeus apex rounded and
narrow (as in Fig. 21); anal lobe wider than cell cup. . . . . . . . . . . . . . . . . . . Vladolinia Kameneva

11. Head twice as long as high (Fig. 9); North America; Cuba; South America. . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Eumetopiella Hendel]
– Head almost as long as high or higher than long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

12. Ventral portion of face slightly protruding anteriorly . . . . . . . . . . . . . . . . . . . . . Chaetopsis Loew

– Ventral portion of face receding (Fig. 25); North and South America. . . . . . . [Stenomyia Hendel]
ULIDIIDAE (picture-winged flies) 66 885

9 Eumetopiella
8 Eumecosomyia

r4+5

10 Physiphora

A1+CuA2

11 Goniaeola 12 Megalaemyia

Figs.  66.8–12. Lateral views of adults: (8)  Eumecosomyia nubila (Wiedemann); (9)  Eumetopiella rufipes (Macquart), (Nearctic);
(10) Physiphora clausa (Macquart); (11) Goniaeola foveolata Hendel, (South America); (12) Megalaemyia elsae Hendel, (South America).
Figures 8–12 redrawn from Hendel (1909b, 1910) by V. Korneyev.
886
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

13. Vein R1 setulose in distal portion and A1 completely discontinued about halfway to wing margin
(Fig. 37). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pareuxesta Coquillett
– Vein R1 bare or, if rarely with setulae, then vein A1 in apical half turning apically and becoming
foldlike, but continuing to wing margin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

14. Frons smooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

– Frons pitted or wrinkled. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

15. Prescutellar acrostichal setae absent; vein R2+3 sinuate; crossvein r-m proximal to Sc vein apex
(Fig. 31); scutum as long as wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Axiologina Hendel
– Prescutellar acrostichal setae present; vein R2+3 straight, and if sinuate, then crossvein r-m distal
to Sc vein apex (Fig. 33); scutum usually slightly longer than wide. . . . . . . . . . . . . Euxesta Loew

13 Paragorgopis 14 Pterocalla

r4+5

15 Pterocerina 16 Cyrtomostoma

Figs. 66.13–16. Lateral views of adults (concluded): (13) Paragorgopis mallea Hendel, (South America); (14) Pterocalla plumitarsis Hendel,
(South America); (15) Pterocerina fenestrata Hendel, (South America); (16) Cyrtomostoma gigas Hendel, (South America). Figures 13–16
redrawn from Hendel (1909b, 1910) by V. Korneyev.
ULIDIIDAE (picture-winged flies) 66 887

spmth

tg 4
ej apod plr memb
tg 5
tg 6
st 6 st 6
st 7
oviscape
st 8
tg 6
epand aculeus

ph

17 Delphinia
18 Chaetopsis 19 Chaetopsis
20 Acrosticta
st 5

spmth
st 6

st 7

st 8

plr memb
21 Acrosticta ph tg 6
am apod
ej apod
23 Dyscrasis
epand
prct
sur

prens prct

22 Pterocerina

pgt
pregt
sur

ph

24 Acrosticta

Figs. 66.17–24. Abdomens and terminalia: ventral view of male abdomen of (17) Delphinia picta (Fabricius), (Nearctic, MND, fig. 63.26);
ventral view of abdomen of female (aculeus not shown) (18), and enlarged ventral view of apex of aculeus (19), of Chaetopsis major (Wulp);
dorsal view of abdomen of female (20), and enlarged ventral view of apex of aculeus (21), of Acrosticta profunda Hendel; ventral view of
male terminalia of (22) Pterocerina fenestrata Hendel; ventral view of abdomen of female of (23) Dyscrasis hendeli Aldrich; posteroventral
view of male terminalia of (24) A. profunda. Figures 18–24 illustrated by V. Korneyev.
Abbreviations: am apod, anteromedial apodeme of sternite; ej apod, ejaculatory apodeme; epand, epandrium; pgt, postgonite; ph, phallus;
plr memb, pleural membrane of abdomen; prct, proctiger; pregt, pregonite; prens, prensiseta; spmth, spermatheca; st, sternite; sur, surstylus;
tg, tergite.
888
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

16. Frontal vitta with four to six shallow pits or smooth in middle, laterally with few well-separated
shining pits at bases of major setae (Fig. 5) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Euphara Loew
– Frontal vitta with numerous pits, sometimes broadly coalescent (Figs. 6–7). . . . . . . . . . . . . . . 17

17. Frons anterior of ocellar triangle with bare triangular callus and pit at middle (Fig.  6); first
flagellomere elongate oval, more than twice as long as wide . . . . . . . . . . . . . . . . . . Siopa Hendel
– Frons uniformly pitted or wrinkled (Fig. 7); first flagellomere  oval, less than 1.8 times as long as
wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

18. Veins R2+3 and R4+5 strongly convergent, attaining costa anterior of wing apex (Fig. 35). . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cenchrometopa Hendel
– Veins R2+3 and R4+5 not deflected anteriorly (Fig. 32) . . . . . . . . . . . . . . . . . . . . . . Acrosticta Loew

19. Scutum with three or more dorsocentral setae; male genitalia with unpaired metaphallic plate
between surstyli well developed (Fig. 43, 46) and ejaculatory apodeme with small apical portion;
female abdominal sternites  4–6 each with thin anteromedial apodemes (Fig.  23); OTITINAE:
MYENNIDINI. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
– Scutum with two or one dorsocentral setae; male genitalia with no metaphallic plate between
surstyli; ejaculatory apodeme with apical portion at least as long as fanlike portion; female
abdominal sternites either with or without apodemes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

20. Thorax with five acrostichal, five dorsocentral, three supra-alar (one of each presutural),
two  intra-alar, and one  postalar seta, but no minor setulae on scutum (Fig.  40); Sc  sinuate;
extension of cell cup conspicuously longer than anal vein apical to it (Fig. 40); southern USA;
Mexico. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Dyscrasis Aldrich]
– Scutum with both setae and setulae; before suture, at most one supra-alar and one intra-alar larger
than surrounding setulae (Figs 42, 45); Sc almost straight, bowed before end; extension of cell cup
shorter or, at most, as long as anal vein apical to it (Figs. 41, 44). . . . . . . . . . . . . . . . . . . . . . . . 21

21. Scutum with large shining supra-alar areas; two supra-alar setae present (Fig. 45); basal cells of
wing with isolated black dots and stripes having hyaline borders (Fig. 44) . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neodyscrasis Kameneva & Korneyev
– Scutum without large shining areas; one supra-alar seta present (Fig. 42); wing with uniformly
darkened basal cells (Fig. 41) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudodyscrasis Hernández

22. Palpus broad, subtriangular or crescentlike (Fig.  56); proepisternal seta rudimentary; cell cup
closed by straight or arcuate vein with short posteroventral lobe, or posteroventral lobe lacking;
male genitalia with phallus spinulose (Fig. 17); female abdominal sternites 4–6 each with thin
anteromedial apodemes; OTITINAE: CEPHALIINI. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
– Palpus not broad, subtriangular, or crescentlike; proepisternal seta usually well developed, but if
rudimentary, then cell cup with long posteroventral lobe; male genitalia and female sternites
various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

23. Costal vein conspicuously bowed between apices of crossvein h and Sc vein, costal cell convex
anteriorly; wing dark yellowish-brown, with four hyaline incisions (Fig. 48); thorax reddish-
yellow, white microtrichose. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Delphinia Robineau-Desvoidy
– Costal vein straight in basal half, costal cell not convex; wing hyaline with two elongate brown
spots along costal margin (Fig. 49); thorax black, pleuron non-tomentose, with bluish metallic
sheen. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Proteseia Korneyev & Hernández
ULIDIIDAE (picture-winged flies) 66 889

24. Cell cup with posteroventral lobe 1.5 times longer than cell itself (Fig.  47); proepisternal seta
rudimentary; face with deep antennal grooves separated by rounded keel; male genitalia with
phallus spinulose; female abdominal sternites  4–6 each with thin anteromedial apodemes;
OTITINAE: OTININI. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diacrita Gerstaecker
– Cell cup usually with shorter posteroventral lobe; if lobe long and proepisternal seta rudimentary,
then face without antennal grooves or keel; male genitalia with phallus bare, ejaculatory apodeme
with apical portion at least as long as fanlike portion (Fig. 22); female abdominal sternites 4–6
without apodemes; ULIDIINAE: PTEROCALLINI. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

25. Cell cup with distinct posteroapical lobe extended along A1 vein; if inconspicuous (in males of
Plagiocephalus and Chondrometopum), then head at least 2 times wider than long. . . . . . . . . . 26
– Cell cup closed by arcuate or straight vein without posteroapical extension . . . . . . . . . . . . . . . 43

26. Vein R1 bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27

– Vein R1 setulose. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36

27. Cell r4+5 greatly broadened, cells posterior to it narrowed (Fig. 11); scutum with broad longitudinal
depression; female unknown; Bolivia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Goniaeola Hendel]
– Wing venation not as above; scutum without depression in both sexes . . . . . . . . . . . . . . . . . . . 28

28. Cell cup with long posteroapical lobe as long as apical portion of A1 vein; R2+3 sinuate; scutum
with pattern of darker microtrichial spots at bases of setae and setulae (Fig. 14). . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocalla Rondani, in part
– Cell cup with short posteroapical lobe as long as apical portion of A1 vein; R2+3 almost straight in
both sexes; scutum uniformly gray microtrichose or shining . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

25 Stenomyia 26 Zacompsia
27 Undescribed
Genus A

30 Euacaina
28 Axiologina 29 Notogramma

Figs.  66.25–30. Heads: lateral view of (25)  Stenomyia nasoni Cresson; (26)  Zacompsia colorata Steyskal; (27)  Undescribed Genus A;
(28) Axiologina ferrumequinum Hendel; (29) Notogramma cimiciforme Loew; and (30) Euacaina cactipeodes (Steyskal). Figures 25–30
illustrated by V. Korneyev.
890
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

R1

R2+3

r-m
R4+5
M1

lb cup Cu A1
A1+Cu A2

31 Axiologina 32 Acrosticta

33 Euxesta 34 Euphara

35 Cenchrometopa 36 Siopa

37 Pareuxesta 38 Chaetopsis

39 Notogramma

Figs. 66.31–39. Wings: dorsal view of (31) Axiologina ferrumequinum Hendel; (32) Acrosticta scrobiculata Loew; (33) Euxesta eluta Loew;
(34) Euphara caerulea (Macquart); (35) Cenchrometopa curvinervis Hendel; (36) Siopa longicornis Hendel; (37) Pareuxesta latifasciata
Coquillett; (38) Chaetopsis major (Wulp); and (39) Notogramma cimiciforme Loew, (Nearctic, MND, fig. 63.18). Figures 31, 32, 34, and
35 redrawn from Hendel (1910); and Figure 37 redrawn from Steyskal (1966c) by V.  Korneyev. Figures 33, 36, and 38 illustrated by
V. Korneyev.
Abbreviation: lb cup, lobe of cell cup.
ULIDIIDAE (picture-winged flies) 66 891

41 Pseudodyscrasis

40 Dyscrasis

dc s dc s 44 Neodyscrasis

spal s

47 Diacrita

Sc
h

42 Pseudodyscrasis 48 Delphinia
45 Neodyscrasis
R1

mtph plt mtph plt

43 Pseudodyscrasis 46 Neodyscrasis 49 Proteseia

Figs. 66.40–49. Adult thoraces and abdomens, wings, and male terminalia of Otitinae: dorsal view of adult of (40) Dyscrasis hendeli Aldrich,
(Mexico); dorsal view of wing (41), dorsal view of thorax and abdomen of female (42), and of epandrium and metaphallic plate of male (43),
of Pseudodyscrasis scutellaris (Wiedemann), (Mexico, Kameneva & Korneyev, 2006, figs. 107, 108; Hernández-Ortiz, 1988, fig. 2A); dorsal
view of wing (44), dorsal view of thorax and abdomen of female (45), and epandrium and metaphallic plate of male (46), of Neodyscrasis
steyskali (Hernández-Ortiz), (Kameneva & Korneyev, 2006, figs. 112, 113; Hernández-Ortiz, 1988, figs. 2B); dorsal view of wing (47), of
Diacrita costalis Gerstaecker, (Mexico, Hendel, 1914b, fig. 41); (48) Delphinia picta (Fabricius), (Nearctic, Hendel, 1914b, fig. 43); and
(49) Proteseia steyskali Hernández-Ortiz & Kameneva, (Mexico). Figures 40 and 49 illustrated by V. Korneyev. Figures 43 and 46 redrawn
from Hernández-Ortiz (1988) by V. Korneyev.
Abbreviations: dc s, dorsocentral setae or setulae; spal s, supra-alar setae; mtph plt, metaphallic plate.
892
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

50 Bothrometopa 51 Chondrometopum 52 Chondrometopum

53 Paragorgopis 54 Paragorgopis 55 Dasymetopa

vb
plp

58 Rhyparella

56 Proteseia 57 Ophthalmoptera

fr s
ar

61 Apterocerina
59 Terpnomyennis 60 Aciuroides

Figs. 66.50–61. Heads of Cephaliini and Pterocallini: dorsal view of (50) Bothrometopa determinata (Walker), (South America); anterior view
of male (51), and female (52), of Chondrometopum bifenestratum Kertész, (Kameneva, 2004a, figs. 5, 6); anterior view of (53) Paragorgopis
euryale Kameneva, (Kameneva, 2004a, fig. 53); (54) Pa. mallea Hendel, (South America, Hendel, 1909b, fig. 58); and (55) Dasymetopa
lutulenta Loew, (Kameneva, 2004a, fig.  31); lateral view of (56)  Proteseia steyskali Hernández-Ortiz  & Kameneva, (Mexico);
(57)  Ophthalmoptera undulata Hendel, (Kameneva, 2004a, fig.  39); (58)  Rhyparella decempunctata Hendel; (59)  Terpnomyennis nigra
(Hendel), (Kameneva, 2004a, fig. 59); (60) Aciuroides sp.; and (61) Apterocerina argentea (Hendel). Figures 50, 56, 60, and 61 illustrated by
V. Korneyev. Figure 58 redrawn from Hendel (1909b) by V. Korneyev.
Abbreviations: ar, arista; fr s, frontal seta; plp, palpus; vb, vibrissa.
ULIDIIDAE (picture-winged flies) 66 893

29. Crossvein r-m basal to middle of discal medial cell (M2:M3  = 0.5–1.2); if (in males of
Chondrometopum) apical of middle (M2:M3 = 2.0–3.0), then head more than twice as wide as
high. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
– Crossvein r-m at apical two-fifths of discal medial cell or more apically (M2:M3 > 5) (Figs. 77,
80); if (in Neomyennis zebra Hendel) M2:M3 = 2.2–2.5, then head less than twice as wide as
high. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

30. Pterostigma short, less than 1.5 times as long as wide; r-m basal of middle of cell dm (M2:M3 =
0.5–0.7); apex of wing with hyaline crescent border (Fig. 71) . . . . . . . . . . . Cymatosus Enderlein
– Pterostigma elongate, at least 1.5 times as long as wide; r-m at middle of dm (M2:M3 > 0.9); wing
apex dark, without hyaline border (Fig. 74). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

31. Pterostigma in males large, dipping posteriorly (Fig. 73); scutum densely gray microtrichose, if
subshining (in Chondrometopum leve Hendel), then acrostichal seta lacking. . . . . . . . . . . . . . . 32
– Pterostigma neither longer than costal cell, nor dipping posteriorly, but if slightly dipping, then
not sexually dimorphic (Fig. 12) and scutum shining or gray tomentose with shining black spots
(in Megalaemyia radiata Hendel, M.  punctulata Hendel) or silvery microtrichose vitta
(M. albostriata Hendel), and acrostichal seta present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

32. Acrostichal seta usually lacking, but if present (in some Chondrometopum arcuatum Hendel specimens),
then close to level of anterior dorsocentral seta; costal vein between humeral and subcostal veins
bowed anteriorly; pterostigma in males strongly enlarged, in females dipping posteriorly (Figs. 73,
74); vein R2+3 conspicuously bowed anteriorly . . . . . . . . . . . . . . . . . . . . Chondrometopum Hendel
– Acrostichal seta closer to level of posterior dorsocentral seta; costal vein between humeral and
subcostal veins straight (Fig. 81); pterostigma in females not dipping posteriorly; vein R2+3 almost
straight, not bowed anteriorly (Fig. 82). . . . . . . . . . . . . . . . . . . . . . . . Dasymetopa Hendel, in part

33. Wing with three to four spots in apical portion; pterostigma gradually narrowing apically, not
dipping posteriorly (Fig. 69) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ophthalmoptera Hendel
– Wing without spots; if subapical hyaline area 8-shaped, medially constricted or forming two spots,
then pterostigma dipping posteriorly at least slightly (Fig. 12). . . . . . . . . . . Megalaemyia Hendel

34. Scutellum swollen and shining or with additional swellings (Figs. 65–68); body bright yellow to
orange with darker markings (Fig. 64); without additional crossveins. . . . . . . Xanthacrona Wulp
– Scutellum microtrichose, unmodified, if somewhat swollen (in Neomyennis), then wing with
additional crossvein between subcostal cell and vein R2+3 (Fig. 80); body brownish-black. . . . 35

35. Wing with additional crossvein between subcostal cell and vein  R2+3 (Fig.  80); scutum with
distinct pattern of gray and brown microtrichiae; one or two additional setae at upper margin of
katepisternum and anterior of supra-alar seta; South America . . . . . . . . . . . [Neomyennis Hendel]
– Without additional katepisternal and supra-alar setae, or additional crossveins (Fig. 77); scutum
uniformly grayish microtrichose or at most with pale brown pattern. . Terpnomyennis Kameneva
36. Cell cup with long posteroapical lobe twice as long as maximal cell width; R1 in male often
dipping posteriorly. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
– Cell cup with posteroapical lobe shorter than cell width; R1 in male not dipping posteriorly. . . 38

37. Vein M ending anterior to or at wing apex (Fig. 75); cell r2+3, in female, with supernumerary
crossvein in apical third (Fig. 76); in male, frons with deep pit (Fig. 50); South America . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Bothrometopa Hendel]
– Vein M ending posterior to wing apex; cell r2+3 in females without supernumerary crossvein; frons
without pit in both sexes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocalla Rondani, in part
894
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

38. Crossvein r-m proximal to or at the middle of discal medial cell (M2:M3 = 0.7–1.1). . . . . . . . 39
– Crossvein r-m at apical two-fifths of discal medial cell or more apical (M2:M3 > 5) . . . . . . . . 41

39. Scutum shining; wing with yellow–brown pattern, with extensive yellow areas and three to four
hyaline spots (Fig. 70). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parophthalmoptera Hendel
– Scutum uniformly tomentose; wing with reticulate or banded pattern . . . . . . . . . . . . . . . . . . . . 40

40. Anteriormost subvibrissal setula strong, vibrissa-like (Fig. 58); wing pattern reticulate (Fig. 72),
with quadrate spots. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhyparella Hendel
– All subvibrissal setulae subequal, or anteriormost shorter than posterior (Fig. 55); wing pattern
reticulate or banded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dasymetopa Hendel, in part

41. R2+3 sinuate; wing predominantly dark brown, with few dark and hyaline spots; with two
supra-alar and one dorsocentral setae . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudopterocalla Hendel
– R2+3 almost straight; wing hyaline with dark banded pattern (Figs. 62, 78–79); with one supra-alar
and two dorsocentral setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42

42. Head more than twice as wide as long in females (Fig. 63); males with stalked eyes (Fig. 62);
frons much wider than long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plagiocephalus Wiedemann
– Head less than twice as wide as long, frons narrower than wide in both sexes. . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Terpnomyia Hendel

43. Prescutellar acrostichal seta absent; strong frontal setae present (Fig.  60); vein  R1 setulose;
pterostigma narrow and elongate (Fig. 84) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aciuroides Hendel
– Prescutellar acrostichal seta present; other characters various . . . . . . . . . . . . . . . . . . . . . . . . . . 44

44. Arista long setulose (Fig. 15). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocerina Hendel

– Arista bare or short setulose (Fig. 61). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45

45. Vein R2+3 and cell r4+5 sinuate, turning posteriorly, then gently anteriorly toward wing tip; wing
large, 12–15 mm long (Fig. 16); arista short setulose; R4+5 bare; Bolivia. . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Cyrtomostoma Hendel]
– Vein R2+3 almost straight and cell r4+5 at most narrowed towards apex, but not sinuate; smaller flies,
wing 2.5–10 mm long; arista bare; R4+5 setulose or bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46

46. Pterostigma narrow and elongate, vein R1 close to costa (Fig. 85); South America. . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Tetrapleura Schiner]
– Pterostigma gradually tapered towards apex or widened in male (as in Fig. 84) . . . . . . . . . . . . 47

47. Wing pattern banded (Fig. 86). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apterocerina Hendel

– Wing pattern reticulate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48

48. Forefemur of both sexes not swollen (Fig. 13); vein R2+3 strongly sinuate; head in male often
widened (Fig. 54); if head not widened (Fig. 53), then pterostigma in male widened . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paragorgopis Giglio-Tos
– Forefemur of male strongly swollen; vein R2+3 almost straight; neither head nor pterostigma
widened; South America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Coscinum Hendel]
ULIDIIDAE (picture-winged flies) 66 895

63 Plagiocephalus

62 Plagiocephalus

65 X. phyllochaeta 66 X. tripustulata

64 Xanthacrona bipustulata

67 X. bipustulata 68 X. tuberosa

Figs. 66.62–68. Adults, head and scutella: dorsal view of adult male (62), and anterior view of head of female (63), of Plagiocephalus
latifrons (Hendel), (Kameneva, 2004b, figs. 1, 4); lateral view of adult of (64) Xanthacrona bipustulata (Wulp); dorsolateral view of scutellum
of (65) X. phyllochaeta Hendel; (66) X. tripustulata Enderlein; (67) X. bipustulata; and (68) X. tuberosa Cresson. Figure 64 redrawn from
Hendel (1909b) by V. Korneyev. Figures 65–68 illustrated by V. Korneyev.
896
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

ptstg

69 Ophthalmoptera
70 Parophthalmoptera
ptstg R1
R2+3

71 Cymatosus 72 Rhyparella
ptstg ptstg
R1

R2+3

73 Chondrometopum 74 Chondrometopum

75 Bothrometopa 76 Bothrometopa

77 Terpnomyennis

Figs. 66.69–77. Wings: dorsal view of (69) Ophthalmoptera undulata Hendel, (Kameneva, 2004a, fig. 40); (70) Parophthalmoptera bipunctata
(Hendel); (71) Cymatosus bestifer (Hendel); (72) Rhyparella decempunctata Hendel; male (73) and female (74) of Chondrometopum leve
Hendel, (Kameneva, 2004a, figs. 10, 11); male (75) and female (76) of Bothrometopa determinata (Walker); and (77) Terpnomyennis nigra
(Hendel), (Kameneva, 2004a, fig. 62). Figures 70–72 and 75–76 redrawn from Hendel (1909b) by V. Korneyev.
Abbreviation: ptstg, pterostigma.
ULIDIIDAE (picture-winged flies) 66 897

78 Terpnomyia 79 Terpnomyia
ptstg
R1
Sc
R2+3

R4+5

80 Neomyennis 81 Dasymetopa
ptstg R1
c

R2+3

R4+5

82 Dasymetopa 83 Pterocerina

ptstg R1

84 Aciuroides 85 Tetrapleura

86 Apterocerina

Figs. 66.78–86. Wings (concluded): dorsal view of male (78), and female (79), of Terpnomyia angustifrons Hendel; (80) Neomyennis cyaneiventris
(Hendel); male (81) and female (82) of Dasymetopa lutulenta Loew, (Kameneva, 2004a, figs. 16, 17); (83) Pterocerina ruficauda Hendel;
(84) Aciuroides sp.; (85) Tetrapleura picta Schiner; (86) Apterocerina argentea Hendel. Figures 78 and 79 redrawn from Hendel (1909b) by
V. Korneyev. Figures 80, 83–86 illustrated by V. Korneyev.
Abbreviation: ptstg, pterostigma.
898
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Synopsis of the fauna species, is obviously heterogeneous. Chaetopsis major and

C.  praeceps Hendel have interfrontal setae (as shown in
Based upon available collection material, approximately Fig.  3) as in Stenomyia Hendel, but the ventral margin of
135 species in 31 genera of the family Ulidiidae are found in the face is slightly produced, as in the type species of this
Central America, of which at least 63 species and one genus genus. Chaetopsis, Eumetopiella, Stenomyia, Zacompsia,
are undescribed. The full Mesoamerican fauna is estimated and a few other Neotropical genera share narrow abdominal
to include about 40 genera and 160 species. pleura (Fig. 18) and a flattened aculeus (Fig. 19) and form a
monophyletic group, which needs a revision to refine generic
Aciuroides Hendel. Three nominal species described limits. There is no comprehensive key to species.
from Paraguay and Brazil (Nova Teutônia) (Steyskal, 1968)
and three nominal species described from Brazil and Peru Chondrometopum Hendel. Three species are known
under other generic names fit the concept of this genus. They from Central America, Colombia, Venezuela, Peru, and Bo-
are readily recognized by distinct frontal setae (Fig. 60) com- livia. Two of them differ by having a strongly dilated head
bined with the lack of acrostichal seta, and relatively small in both sexes (Figs.  51–52), whereas C.  leve Hendel from
body size (wing length 2.5–6  mm). Fourteen undescribed Bolivia, Peru, Colombia, and Venezuela has an unmodified
species from Central America, Mexico, South America, head. Chondrometopum bifenestratum Kertész was recorded
Dominican Republic, and Trinidad are known in collections. from Costa Rica and Panama (Kameneva, 2004a).
Some of them show sexual dimorphism in wing venation and
pattern. Names of some species described from one sex may Cymatosus Enderlein. Three species occur in Peru, Ecuador,
be synonyms (E.P. Kameneva, in preparation). and Brazil, and one in Mexico. Cymatosus icarus Kameneva
was described from Costa Rica (Kameneva, 2009).
Acrosticta Loew. This genus includes 14 species (Steyskal,
1965, 1968). The Central American fauna has five species, Dasymetopa Hendel (= Euxestina Curran). Twelve spe-
all recently recorded from Costa Rica (Kameneva, 2004a), cies of this genus are described so far, predominantly from
with two also known from Honduras and El Salvador; two or Peru and Bolivia. Further specimens are known from Mexico,
three undescribed Central American species are known from Nicaragua, Costa Rica, Panama, Colombia, Venezuela, and
collections. Hendel (1909c) keyed only seven species, and Ecuador. Five species are found in Central America (Kame-
there is no comprehensive key. neva, 2004a). Sexual dimorphism in head shape and wing
pattern is characteristic for many species; however, some
Apterocerina Hendel. Five species from Ecuador, Peru, were described from one sex only. Species can be identified
and Bolivia are included in Apterocerina (Steyskal, 1968; using Kameneva (2004a).
E.P.  Kameneva  & V.A.  Korneyev, unpublished data). They
can be readily distinguished by the combination of setulose Delphinia Robineau-Desvoidy. One common North
veins R1 and R4+5 and short pubescent arista. The only species American species, D. picta (Fabricius), was recorded from
Mexico and El Salvador (Steyskal, 1971b).
of the South American genus Coscinum Hendel (C.  clavi-
pes Hendel) has the forefemur swollen and microspinulose, Diacrita Gerstaecker. Diacrita costalis Gerstaecker, one
but otherwise is closely related to Apterocerina amoena of the two known species occurring in the USA and Mexico,
(Enderlein), so concepts of these genera must be refined. was recently recorded from Nicaragua (Kameneva, 2004a).
One undescribed species of Apterocerina from Costa Rica is See also Steyskal (1947).
known from collections.
Eumecosomyia Hendel. Three species, including two de-
Axiologina Hendel. Axiologina ferrumequinum Hendel scribed — E. nubila (associated with corn) and E. hambletoni
is widespread from Mexico (Chiapas) and USA (Florida) to Steyskal (larvae in sugar cane stems) — and one undescribed
Ecuador, Bolivia, Peru, and Brazil. In Central America, it is species are known from Central America (Steyskal, 1968;
recorded from Belize, Guatemala (Kameneva, 2004a), Costa Kameneva, 2004a). A key to species is given by Steyskal
Rica, and Panama. The two species of this genus can be (1966a).
identified using Steyskal (1973).
Euphara Loew. In addition to the type species, E. caerulea
Chaetopsis Loew. There are 13 described species, eight (Macquart), from Guyana, Venezuela, Brazil, Peru (Stey-
of which occur in North America, southward to Mexico, skal, 1968), Costa Rica, and Panama (Kameneva, 2004a),
Cuba, and Puerto Rico; five species are known from Brazil two undescribed species, both occurring in Mesoamerica,
and Argentina (Steyskal, 1965, 1968). The only Central are known in collections (G.C. Steyskal, unpublished data).
American species, C.  major, recorded from Mexico, USA,
El Salvador (Steyskal, 1968), and Nicaragua (Painter, 1955), Euxesta Loew. There are 90 species of this genus in the
is represented also by material from Honduras, Colombia, New World and Oceania; of them, a few were accidentally in-
and Costa Rica (Kameneva, 2004a). The genus, as delim- troduced into Europe and Asia. About 70 species are known
ited by Steyskal (1968), with the inclusion of Neotropical from America south of the USA; 18 described species have
ULIDIIDAE (picture-winged flies) 66 899

been recorded from Central America (Steyskal, 1965, 1968, (E.P. Kameneva, unpublished data). A key to species is given
1974; Steyskal & Ahlmark, 1995; Kameneva, 2004a). In ad- by Kameneva (2004a).
dition, 15 undescribed Central American species were seen
in collections during this study. Larvae of Euxesta stigmatias, Pareuxesta Coquillett. This genus, in the strict sense, is
E. mazorca, E. sororcula, and E. eluta infest ear corn (App, found only in the Eastern Pacific. Six species are described
1938; Barber, 1939; Painter, 1955; Steyskal, 1974; Huepe from the Galápagos Islands (Steyskal, 1966c). Two unde-
et al., 1986). Some other species live in rotting trees and in scribed species occur on Cocos Island (Costa Rica). Some
young shoots of palms (Krivosheina & Krivosheina, 1995a; undescribed mainland species that key to this genus possibly
Yoon et al., 1983). The most recent key to species by Curran are non-congeneric with the island species.
(1935) needs to be critically revised. Species of Acrosticta
Parophthalmoptera Hendel. One described species,
and some Euxesta have similarly reduced female abdominal
P. picea Hendel from Bolivia and two undescribed species
tergites 4–6 (Fig. 20); Euxesta in its current concept is not
from Costa Rica belong here. “Ophthalmoptera” bipunctata
monophyletic (Kameneva & Korneyev, 2006).
from Peru and Bolivia (Steyskal, 1968; as Ophthalmoptera
Megalaemyia Hendel. There are about 18 species (of (Parophthalmoptera)), runs to Megalaemyia in the key
them, only five described) occurring from Nicaragua and and apparently is closely related or belonging to the latter
the Antilles (Dominica) to Peru, Ecuador, and Brazil (São genus.
Paulo). From Central America, M.  punctulata Hendel and
at least seven undescribed species are known in collections Physiphora Fallén. There are two subcosmopolitan
(Kameneva, 2004a, in preparation). Hendel’s (1909a) key to synanthropic species known from the Americas. One of
species is out of date and includes species currently assigned them, P. clausa (Macquart), is introduced in the USA, Mexico,
to Cymatosus. Some species currently assigned to Ophthal- Cuba, and Brazil, and was recently found in urban localities
moptera Hendel may belong here. Megalaemyia differs from in Costa Rica.
the latter genus by its pterostigma dipping posteriorly and
Plagiocephalus Wiedemann. There are three species of
having one elongate area in the apical portion of the r2+3 cell.
this genus, of which one is known only from Brazil, Para-
“Ophthalmoptera” bipunctata Hendel and “O.”  elegans
guay, and Argentina. The second, P.  latifrons (Hendel) is
Hendel have intermediate states of these characters and are
widespread from Mexico (Chiapas) and Trinidad to Peru,
often keyed as Megalaemyia.
Bolivia, Brazil, and Paraguay; in Central America, it is
Neodyscrasis Kameneva  & Korneyev. Neodyscrasis known from Nicaragua, Costa Rica, and Panama. The third
steyskali (Hernández-Ortiz) is known from material collect- species, P.  intermedius Kameneva, is known from Costa
ed in Mexico, Honduras, Guatemala, Venezuela, and Peru Rica (Kameneva, 2004b). Males in all three species have the
(Hernández-Ortiz, 1988; Kameneva & Korneyev, 2006). head strongly widened, extremely so in males of P. latifrons
(Fig.  62), whereas females have only moderately widened
Notogramma Loew. These are bluish- or greenish-black, heads and differ mostly in minute details of frons coloration
swiftly running flies with flattened body and appressed wings, and wing pattern. A key to species is given by Kameneva
resembling small bugs or alticine leaf beetles. They are as- (2004b).
sociated predominantly with rotting cacti and other plant
matter (e.g., watermelon fruits). Of the four species occur- Proteseia Hernández-Ortiz  & Korneyev. One species,
ring from California and Texas to Chile, two species occur in P.  steyskali Hernández-Ortiz  & Kameneva was described
Central America: N. cimiciforme Loew and N. purpuratum from Neotropical Mexico (Hernández-Ortiz et al., 1999), but
Cole (Steyskal, 1968; Kameneva, 2004a). Keys to species has not yet been found elsewhere in Central America.
are given by Steyskal (1963, 1991).
Pseudodyscrasis Hernández-Ortiz. One species,
Ophthalmoptera Hendel. There are five described spe- P.  scutellaris (Wiedemann), occurs only in Neotropical
cies of this genus; O.  undulata Hendel (Figs.  57, 69) and Mexico.
at least one undescribed species are known from collections
Pseudopterocalla Hendel. Two species, Pseudoptero-
from Costa Rica (Steyskal, 1968; Kameneva, 2004a). Oph-
thalmoptera elegans from Venezuela and Bolivia is keyed to calla obscura (Wiedemann) and P.  scutellata (Schiner),
Megalaemyia and apparently is closely related to or belongs both occur from Central America to Guyana, Peru, Brazil,
to that genus. and Bolivia. The third species was recently moved to Ptero-
calla (Kameneva, 2004a). Hendel (1909b) recognized Pseu-
Paragorgopis Giglio-Tos. Eleven described species as- dopterocalla as a subgenus of Pterocalla. Later, it was raised
signed here are widespread from southern Mexico and Trini- to full generic rank (Curran, 1934; Aczél, 1951; Steyskal,
dad to Bolivia and Brazil, and six occur in Central America. 1968). Although Steyskal (1982) again reduced its rank to
Paragorgopis maculata Giglio-Tos is a nomen dubium; another subgenus, we maintain it as a genus. Hendel (1909a) gives a
species from Bolivia possibly belongs in the genus Apterocerina key to species.
900
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Pterocalla Rondani. One of the most common and wide- added from Argentina and Uruguay actually belong in the
spread in the Neotropical Region, this genus in its strict sense subfamily Otitinae (E.P.  Kameneva  & V.A.  Korneyev, in
comprises 19 described species (we consider Pseudoptero- preparation) and must be excluded from Terpnomyia.
calla a separate genus). The subgenus Parapterocalla Aczél
was proposed to include three species with an enlarged male Vladolinia Kameneva. One species of this genus, V. dolini
subcostal cell. Eight species occur in Central America (Steyskal, Kameneva, is known from Costa Rica and other localities in
1968; Kameneva, 2004a), and four undescribed species from the Neotropical Region. It superficially resembles Zacomp-
this region are known from collections. Hendel’s (1909a) sia in the elongate head and body and similar wing pattern,
key to species is out of date and includes species currently but differs by the wide anal lobe, acute fronto-facial angle,
assigned to Pseudopterocalla and Callopistromyia. and strongly produced clypeus, like in Eumecosomyia,
Aspistomella Hendel, and Paraphyola Hendel, but having
Pterocerina Hendel (= Sympaectria Hendel). This is the the eye longer than high (Kameneva, 2005). It shares with
largest and most diverse genus of the Pterocallini, with 29 all these genera the long abdomen with narrow pleura, dif-
described and a number of undescribed species. Some names fering from any other genera related to Chaetopsis and
of species, described from a single sex only, are obviously Zacompsia by the aculeus neither flattened nor widened.
synonyms. This genus was originally established to include The scutellum is strongly produced dorsally.
species with a long pubescent arista, setulose R4+5, and cell
cup closed with arcuate vein (Fig. 83). Later, Hendel (1914a) Xanthacrona Wulp. This genus includes five species
established additional genera and subgenera for species with and is widespread from southern USA (Texas) to northern
a bare or short pubescent arista, including Apterocerina and Argentina. All are easily recognizable by the yellow-and-
Micropterocerus Hendel (= Tetrapleura Schiner). Pteroceri- black shining scutellum and oblique-banded wing pattern
na ruficauda Hendel, P. psidii Capoor, and seven undescribed (Fig. 64). Three species occur in Central America (key in
species are known from collections from Central America. Steyskal, 1966b).
There are no comprehensive keys.
Zacompsia Coquillett (=  Metopocampta Enderlein).
Rhyparella Hendel. One species, R.  decempunctata The range of the three described species is from southern
Hendel, is known from Costa Rica, Bolivia, and Brazil. All USA to southern Brazil, and all are found in Central America
other species assigned to Rhyparella belong to Dasymetopa (Kameneva, 2004a). The genus is related to Chaetopsis,
(Kameneva, 2004a). Eumecosomyia, and Stenomyia, sharing with them the
narrow abdominal pleura and flattened aculeus, but are
Siopa Hendel. Siopa longicornis Hendel, which occurs readily differentiated by the narrow anal lobe. Steneretma
in Costa Rica, Panama, Peru (Steyskal, 1968), El Salvador, Loew (see Steyskal, 1987: fig. 63.17) from the USA is an
and Nicaragua (Kameneva, 2004a) is apparently an aberrant aberrant Zacompsia with not only the anal lobe reduced,
species of Acrosticta. but also the cup and cua1 cells lacking (E.P. Kameneva &
V.A.  Korneyev, in preparation). Steyskal (1971a) gives a
Terpnomyennis Kameneva. One species, T. nigra (Hendel) key to species.
from Peru, Venezuela, and Costa Rica, fits near Terpnomyia
in head shape (Fig. 59), body coloration, and wing pattern Undescribed Genus A. There is a single species dis-
and venation. It differs by having vein R1 bare, and veins r-m tantly resembling Euxesta in its shortened and narrowed
and dm-cu approximated (Kameneva, 2004a). tergites  4–6 of the female and wing pattern. Otherwise, it
differs from any known Ulidiidae species by its wide and low
Terpnomyia Hendel. One species occurs in Costa Rica, head, similar to that of Holarctic Homalocephala Zetterstedt,
Peru, and Bolivia (Steyskal, 1968; Kameneva, 2004a). This but with a high, produced, and sagittally flattened clypeus
genus was originally established to include five species from (Fig.  27), well-developed anepisternal setae, and a single
Peru and Bolivia, of which one was later transferred to katepisternal seta. It differs from any mainland Lipsanini
Plagiocephalus. The other three species must be transferred by its non-metallic blackish-brown body. The only known
to Pterocerina. Steyskal (1968) erroneously synonymized female specimen was not dissected, and its tribal position
Proterpnomyia Blanchard with Terpnomyia. Two species he cannot be determined with certainty.

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Acalyptratae). Proceedings of the California Academy of Sci- Steyskal, G.C. 1982. A key to the genera of the subfamily Otitinae
ences, Fourth series 34: 483–498. of the Americas south of the United States (Diptera: Otiti-
Steyskal, G.C. 1968. 54. Family Otitidae (Ortalidae; including dae). Memoirs of the Entomological Society of Washington 10:
Pterocallidae, Ulidiidae) In Vanzolini, P.E. & N. Papavero (edi- 139–144.
tors). A Catalogue of the Diptera of the Americas South of the Steyskal, G.C. 1987. Otitidae, pp. 799–808. MND, Volume 2.
United States. Departamento de Zoologia, Secretaria da Agri- Steyskal, G.C. 1991. Description of new species of Otitidae from
cultura, São Paulo, 31 pp.
Chile (Diptera). Acta Entomológica Chilena 16: 25–28.
Steyskal, G.C. 1971a. A new Central American species of Zacomp-
sia Coquillett, with a key to the described species (Diptera: Steyskal, G.C. & K.M. Ahlmark. 1995. Two new species of Euxesta
Otitidae). Proceedings of the Entomological Society of Wash- Loew (Diptera; Otitidae). Insecta Mundi 9: 189–192.
ington 73: 247–248. Yoon, S.J., M.T. Mathew, & R.E. Holman. 1983. Biology of Eux-
Steyskal, G.C. 1971b. Delphinia picta (Fabricius) in Central America. esta quaternaria Loew (Diptera: Otitidae). Entomological News
Proceedings of the Entomological Society of Washington 73: 445. 94: 122–126.
PLATYSTOMATIDAE (signal flies) 67
Andrew E. Whittington

Fig. 67.1. Female of Rivellia flavimana Loew, (Nearctic, MND, fig. 64.1).

Diagnosis ent (one pair of each, unless stated): postpronotal, two pairs

notopleural; anepisternal; postsutural supra-alar; postalar;
Body length 3.3–12.0  mm (world fauna: 2.5–20.0  mm) intra-alar; prescutellar acrostichal; prescutellar dorsocentral;
and form ranging from small, slender species (Fig.  1) to basal (sometimes), lateral and apical scutellar. Wing usu-
large, robust individuals. Body colors often with metallic ally with pigmented pattern; humeral break in costa present;
luster and wings usually patterned with dark spots or bands. R1 and R4+5 dorsally setulose for almost entire length; cup
Head shape various, with either frons and face bulging subrectangular at apex — CuA2 perpendicular to CuA at its
forwards or gena swollen and extending backwards. Head base and either straight or convexly curved toward middle
higher than long (Fig. 2); ocelli present; lower frontal setae (Fig. 3). Legs moderately developed, without armature. Ab-
absent, one or two reclinate orbital setae present; inner and domen subcylindrical to narrow-ovate. Male postabdomen
outer verticals present; genal seta present; vibrissa absent (Fig.  4) with sternite  6 reduced or absent; strongly sclero-
to rudimentary; postocellar setae weak, divergent. Pedicel tized bilobed cap on ejaculatory apodeme; phallapodeme
with dorsal seam, first flagellomere elongate, sometimes present; inner surstyli well developed. Female postabdomen
with apical point; arista finely setose to pubescent. Proboscis with segment 6 reduced or absent; segment 7 present as coni-
stout, palpus flattened. Thorax longer than broad; setae pres- cal oviscape, which forms rigid base to flattened ovipositor

903
904
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

r-m r4+5
cup
CuA2 dm

3 Senopterina

2 Senopterina
tg 5 ej apod
phapod
ph gd gls

syntgst 7+8

epand

hypd
cerc
i sur
ph

o sur

st 5 st 6 st 7

4 Rivellia

cu-bm
cubm

5 Amphicnephes

6 Senopterina

7 Rivellia

Figs. 67.2–7. Heads, wings, and abdomen: lateral view of head of (2) Senopterina mexicana (Macquart), (Shewell, 1962, fig. 8); dorsal view
of wing of (3) S. caerulescens Loew, (Nearctic, MND, fig. 64.3); ventrolateral view of abdomen of male of (4) Rivellia inaequata Namba,
(Nearctic, MND, fig. 64.9); dorsal view of wing of (5) Amphicnephes stellatus Wulp; lateral view of head of (6) S. brevipes (Fabricius),
(Shewell, 1962, fig. 1); dorsal view of wing of (7) R. occulta Wulp. Figures 5 and 7 illustrated by A.E. Whittington.
Abbreviations: cerc, cercus; ej apod, ejaculatory apodeme; epand, epandrium; gls, glans; hypd, hypandrium; i sur, inner surstylus; o sur, outer
surstylus; ph, phallus; phapod, phallapodeme; ph gd, phallic guide; st, sternite; syntgst, syntergosternite; tg, tergite.
PLATYSTOMATIDAE (signal flies) 67 905

sheath protecting retractable (by invagination) segment 8 or Classification

aculeus, with apical sensory setae on either side. Larvae with
generalized, subcylindrical schizophoran morphology, typi- Like many flies, the first described species of Platystom-
cal of many Tephritoidea. atidae were placed in the large family Muscidae. Hendel’s
The species of this family are frequently mixed with un- significant contribution and reorganization of the genera of
sorted specimens of other higher Tephritoidea. The absence the world (Hendel, 1914) firmly placed the Platystomatidae
of frontal setae, the lack of any extension of the posterior as a subfamily of Muscaridae, divided into several tribes.
apex of cup, and the presence of setulae along the whole The oldest family-group name for this family is in fact not
length of R1 (and frequently all of R4+5) distinguishes Platys- Platystomatidae, but Achiasidae Fleming, based on the genus
tomatidae from most Tephritidae, Ulidiidae, and Pyrgotidae. Achias Fabricius. There has, however, been an overwhelm-
All Central American Platystomatidae lack a katepisternal ing predominance of the usage of names based on the genus
seta. Furthermore, Tephritidae have between one and five Platystoma Meigen. After petition by Steyskal & McAlpine
frontal setae and a subcostal break. In Ulidiidae, postocellar (1974), the International Commission on Zoological Nomen-
setae are better developed, katepisternal setae present, and clature (1979) ruled under plenary power that family-group
the coiled aedeagus lacks a distinct apical glans. Pyrgotidae names based on Achias should be suppressed, giving those
often have a proepisternal ridge bearing strong setae (absent based on Platystoma precedence.
in all Platystomatidae) and lack ocelli. Frey (1930, 1932) used the family status of Platystomati-
dae and included subfamily divisions. Later, Hennig (1958),
Biology followed by Steyskal (1961), clearly separated the Otitidae
(=Ulidiidae) and Platystomatidae with substantial character
The biology of Central American Platystomatidae is gen- states, suggesting that these families were not only interrelat-
erally unknown, and our knowledge of the biology and im- ed, but formed distinct lineages. From this springboard, the
mature stages must be drawn from examples farther afield. modern approach to the Tephritoidea (Korneyev, 2000) and
Of the four genera, Rivellia is the only one for which any the definitions of the families comprising it have developed.
detailed information is known. Based on species from other The most significant modern publication dealing with
biogeographical regions, Rivellia has a notable association the family status and characterization of the Platystomati-
with leguminous plants, and many other genera of the fam- dae was McAlpine’s (1973) revision of the Australasian
ily prefer shaded, densely vegetated locations, resting on the members of this family. He provided good characters and
underside of foliage. The wings are often held up at an acute reasoning to give family status to the Platystomatidae, sepa-
angle and waved around in signaling motions for sexual dis- rate from the Tephritidae and other tephritoid families and
play. Adults may be attracted to malodorous substances and clearly defined and delimited five subfamilies. More recent-
decay. ly, McAlpine (2001) revised the number of subfamilies to
Larvae of Platystomatidae are poorly known and im- four, incorporating Angitulinae into Platystomatinae. The
mature stages of only a few species have been described. other three subfamilies are Scholastinae, Trapherinae, and
Contrary to Steyskal’s (1987) statement that no larvae of Plastotephritinae
any genus in the Americas had been described, there are lar- The Central American fauna belongs entirely to the cos-
val descriptions or biological notes for half a dozen species mopolitan subfamily Platystomatinae. The other three sub-
of Rivellia (Bibro  & Foote, 1986; Ferrar, 1987; Seeger  & families have an Old World distribution, predominating
Maldague, 1960) and one Nearctic species of Amphicnephes across the Afrotropical, Oriental, and Australasian Regions,
(Blanton, 1938). but are absent from the Palearctic and with Plastotephritinae
Pest species of Rivellia are known from root nodules probably not extending into Australia.
of the following range of leguminous plants: hog peanut
(Amphicarpa bracteata), groundnut (Apios americana), pea- Identification
nut (Arachis hypogaeai), southern pea (Vigna unguiculata),
and black locust tree (Robinia psuediacacia) in USA; soy- Although there are no published keys with which to iden-
bean (Glycine max) in Japan and USA; pigeon pea (Cajanus tify Central American specimens, the Central American fau-
cajan) in India and USA; pasture legume (Glycine javanica) na is clearly similar to that of the Nearctic in both paucity
in Australia; and mung bean (Phaseolus aureus), cow pea and structure. Thus, Steyskal (1961, 1987), dealing with the
(Vigna sinensis), and Bambarra groundnut (Voandzeia sub- Nearctic fauna, provided a fitting key requiring little revision
terraneana) in Congo. They have also been recorded from for the four known Central American genera. The only ex-
the fruit of eggplant (Solanum melongena) in China, dried ceptions were the genus Pogonortalis, introduced from Aus-
flowers of black locust tree (Robinia pseudoacacia), false tralia to California, and Himeroessa (a possible synonym of
indigo (Amorpha fruticosa), and panicled tick trefoil (Des- Senopterina), endemic to Cuba. As far as is presently known,
modium paniculatum, Fabaceae) in USA and from piles of Pogonortalis has not spread to the Central American region;
rotting Narcissus bulbs in USA (Ferrar, 1987). nevertheless, Pogonortalis and Plagiostenopterina are in-
906
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

cluded in the key below as genera introduced from the Old postabdomen. It is sometimes possible to extract the male
World, in anticipation that they might be found in Central genitalia from the genital pouch (Whittington, 2003) at the
America. time of mounting — an action that can considerably assist
No keys are available for determination of Central later examination.
American species. Thus, identification requires reference The basal medial cell (bm) is extended in some genera of
to the original descriptions, which are, for the most part, Platystomatidae, such that there is an unusually long por-
inadequate. Namba (1956) revised the Nearctic species of tion of CuA1 arching between the posterior apex of cross-
Rivellia, including just three species listed for the Central vein bm-cu and the anterior apex of CuA2. This portion of
American fauna (and one of those is dubiously delimited), CuA1 is short in most Acalyptratae and unnoticed; hence, it
and Shewell (1962) treated some Senopterina species. Revi- generally goes unnamed. For ease of interpretation, Whit-
sions of the Central American genera are needed to clarify tington (2003) used cu-bm for this part of CuA1 (Fig. 5). It
the known fauna and should include thorough morphologi- is used in the following key, as it is an important diagnostic
cal examination, including dissection and description of the feature.

Key to the genera of Platystomatidae of the Neotropical Region

1. Veins R4+5 and Ml divergent or subparallel at apex (beyond cell dm), such that cell r4+5 either
widened or remains even to apex (endemic genera) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Veins R4+5 and M1 distinctly convergent at apex (beyond cell dm), such that cell r4+5 narrowed at
apex (genera introduced from Old World). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

2. Angle between frons and median occipital sclerite approximately 55–60°, meeting in sharp ridge
along top of head with frons steeply inclined; cell bm longer than cell cup, cu-bm long and gently
arching into bm, approximately equal to length of bm-cu or CuA2 (Fig. 5); scutellum flat on top
and square to subsquare posteriorly. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amphicnephes Loew
– Angle between frons and median occipital sclerite approximately 70–80°, usually curved over
apex (Fig. 2), but if meeting as sharp ridge (Fig. 6), then frons not noticeably steeply inclined;
cell  bm approximately equal in length to cell cup, cu-bm short, shorter than bm-cu or CuA2
(Fig. 3); scutellum rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3. Crossveins r-m and bm-cu almost perpendicular; Ml curved just before crossvein r-m (Fig. 7),
dipping gently into dm (occasionally imperceptibly so); arista pubescent along full length. . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rivellia Robineau-Desvoidy
– Crossveins r-m and bm-cu oblique (angled basally); Ml straight along dm before crossvein r-m;
arista bare, or pubescent only at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4. Antenna long, reaching lower facial margin; cell  r4+5 without short remnant crossvein on M
beyond dm-cu (Fig. 3). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Senopterina Macquart
– Antenna not reaching lower facial margin; cell r4+5 with short remnant crossvein on M beyond
dm-cu; Cuba. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Himeroessa Loew]

5. Hind femur with low anteroventral keel in distal third; crossvein r-m situated basal to midway
along cell dm, hence basal section of vein M on dm much shorter than distal section of M on dm;
two pairs of reclinate orbital bristles present; face without erect hairs; Australia, introduced to
USA. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Pogonortalis Hendel]
– Hind femur plain, lacking any elaboration; crossvein r-m situated midway (or slightly distal)
along cell dm, hence basal section of vein M on dm much longer than distal section of M on dm;
reclinate orbital bristles absent; face with fine erect hairs; Old World tropics, introduced to
Trinidad, Peru, Bolivia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Plagiostenopterina Hendel]
PLATYSTOMATIDAE (signal flies) 67 907

Synopsis of the fauna this genus (Namba, 1956). Five species were described from
Guerrero in the Yucatán by Wulp (1898), although with poor
Within four genera, Steyskal (1968) listed 26 Neotropical descriptions and in all probability requiring some revision.
species, a figure almost certainly requiring revision. Com- Species of this genus are known to have associations with
pared with a world fauna of a little over 1000 species, this Leguminosae.
fauna is indeed sparse.
Senopterina Macquart. The name is sometimes incor-
Amphicnephes Loew. A single species, A. stellatus Wulp, rectly written Stenopterina. The generic names Bricinnia
is known from Mexico to Panama; extralimital specimens Walker and Bricinniella Giglio-Tos are junior synonyms.
are also known from Colombia. No identification key exists for the Central American spe-
cies. This is the most widespread of the four endemic gen-
Rivellia Robineau-Desvoidy. No identification key ex- era; 12 species are listed for Senopterina by Steyskal (1968)
ists for all eight species of Rivellia in Central America, but for the Neotropical Region, with an additional unrecognized
three species from Mexico are also found in the Nearctic species from Brazil mentioned, which is probably a new spe-
Region and appear in the key to 34 Nearctic members of cies. Seven of these occur in Central America.

Literature cited

Bibro, C.M. & B.A. Foote. 1986. Larval description of Rivellia pal- McAlpine, D.K. 2001. Review of the Australian genera of signal
lida (Diptera: Platystomatidae), a consumer of the nitrogen-fixing flies (Diptera: Platystomatidae). Records of the Australian Mu-
root nodules of hog-peanut, Amphicarpa bracteata (Legumino- seum 53: 113–199.
sae). Proceedings of the Entomological Society of Washington Namba, R. 1956. A revision of the flies of the genus Rivellia (Otiti-
88: 578–584. dae, Diptera) of America north of Mexico. Proceedings of the
Blanton, F.S. 1938. Some dipterous insects reared from narcissus United States National Museum 106: 21–84.
bulbs. Journal of Economic Entomology 31: 113–116. Seeger, J.R. & M.E. Maldague. 1960. Infestation de nodules de lé-
Ferrar, P. 1987. A guide to the breeding habits and immature stag- gumineuses en région équatoriale par des larves de Rivellia sp.
es of Diptera Cyclorrhapha. Entomonograph 8(1). E.J.  Brill  / (Dipt.). Parasitica 16: 75–84.
Scandinavian Science Press, Leiden, Copenhagen, 478 pp. Shewell, G.E. 1962. A new Canadian species of Stenopterina Macq.
Frey, R. 1930. Philippinische Dipteren. VII. Fam. Platystomidae. with notes on the species allied to brevipes (Fab.) (Diptera:
Notulae Entomologicae 10: 46–64. Otitidae). The Canadian Entomologist 94: 194–200.
Frey, R. 1932. On African Platystomidae (Diptera). Annals and Steyskal, G.C. 1961. The genera of Platystomatidae and Otitidae
Magazine of Natural History Series 10, 9: 242–264. known to occur in America north of Mexico (Diptera, Acalyp-
Hendel, F. 1914. Diptera Fam. Muscaridae Subfam. Platystominae, tratae). Annals of the Entomological Society of America 54:
Fascicle 157, pp. 1–179. In Wytsman, P. (editor). Genera Insec- 401–410.
torum. Bruxelles. Steyskal, G.C. 1968. 55. Family Platystomatidae. In Papavero, N.
Hennig, W. 1958. Die Familien der Diptera Schizophora und ihre (editor). A Catalogue of the Diptera of the Americas South of
phylogenetischen Verwandtschaftsbeziehungen. Beiträge zur the United States. Departamento de Zoologia, Secretaria da Ag-
Entomologie 8: 505–688. ricultura, São Paulo, 4 pp.
International Commission on Zoological Nomenclature. 1979. Steyskal, G.C. 1987. Platystomatidae, pp.  809–812. MND,
Opinion 1142. Family-group names based on Platystoma Mei- Volume 2.
gen, 1803, given precedence over those based on Achias Fabri- Steyskal, G.C. & D.K. McAlpine. 1974. Platystomatidae Schiner,
cius, 1805 (Diptera). Bulletin of Zoological Nomenclature 36: 1862: Proposed conservation as a family-group name over
125–129. Achiidae Fleming, 1821 (Insecta, Diptera). Z.N.(S.) 2053. Bul-
Korneyev, V.A. 2000. Phylogenetic relationships among the fami- letin of Zoological Nomenclature 31: 59–61.
lies of the superfamily Tephritoidea: higher groups of Tephriti- Whittington, A.E. 2003. Taxonomic revision of the Afrotropical
dae, pp.  3–22. In Aluja,  M.  & A.L.  Norrbom (editors). Fruit Plastotephritinae (Diptera; Platystomatidae). Studia diptero-
Flies (Tephritidae): Phylogeny and evolution of behavior. CRC logica Supplement 12: 1–300.
Press, Boca Raton, London, New York, Washington, D.C., Wulp, F.M. van der. 1898. Family Muscidae, pp. 380–384. In God-
[xvi] + 944 pp. man, F.D. & O. Salvin (editors). Biologia Centrali-Americana, or,
McAlpine, D.K. 1973. The Australian Platystomatidae (Diptera, contributions to the knowledge of the fauna and flora of Mexico
Schizophora) with a revision of five genera. The Australian Mu- and Central America. Zoologia. Class Insecta. Order Diptera.
seum, Sydney, Memoir 15: 1–256. Vol. 2. London, 489 pp.
908
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
TEPHRITIDAE (fruit flies, moscas de frutas) 68
Allen L. Norrbom

Fig. 1. Dorsal habitus of Alujamyia farri Norrbom, (Norrbom, 2006b, fig. 29).

Diagnosis extension due to bend in vein  CuA2 (except in Myopitini

and a few other taxa). Female with ovipositor of piercing type
Small to large (body length, including female ovipositor, (Figs. 163–165), including strongly sclerotized aculeus (fused
2–35 mm), generally colorful flies. Body variously colored segment  8  + cerci) telescoping into fused oviscape (synter-
but not metallic in American taxa (except abdomen of Ta- gosternite 7). Male with correspondingly long, coiled phallus
chinisca). Frons with both inclinate frontal setae (sometimes (Fig. 156), stored between postabdomen and tergite 5 at rest
reduced or absent in wasp mimics such as Toxotrypana and and bearing stout apical glans (absent in some Anastrepha);
some Pseudophorellia or rarely in other species) and one to lateral surstylus fused to epandrium; and medial surstylus,
two (rarely three to four) orbital setae, at least the anterior closely associated with lateral surstylus and bearing two or
one reclinate (Figs. 21, 27) or rarely (male of Ceratitis capi- rarely zero to one pairs of prensisetae (Figs. 155–156).
tata (Wiedemann)) anterior seta proclinate and expanded Eggs 0.35–1.65 mm long, white to pale brown, elongate-
apically (Fig. 17). Three ocelli present (anterior one absent oval to pedicillate, often slightly curved, and smooth to
in Ceratodacus longicornis Hendel). Anepisternum with rugose-textured, often with raised areas in hexagonal or sim-
vertical phragma on posterior third (Fig. 43) (difficult to see ilar shapes. Micropyle apical to subapical and usually notice-
in densely microtrichose species). Anepimeron with at least able. Some species with slender lobe, sometimes extremely
one seta medially and with greater ampulla usually well de- elongate, on anterior end. Chorionic respiratory system well
veloped. Wing almost always patterned. Costa with humeral developed, with aeropyles confined to anterior end, plastron
and subcostal breaks. Subcosta bent sharply forward subapi- and hatching lines absent (Hinton, 1981; Ferrar, 1987).
cally and usually weaker or foldlike beyond the bend. Vein R1 Larvae white to yellow, occasionally with darker caudal or
dorsally densely setulose, setulae extending basally to fork ventral areas; in most species elongate cylindrical, but in gall
with Rs. Cell cup usually with at least small posteroapical and flower-breeding taxa sometimes stout and subspherical.

909
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

First instar metapneustic, second and third instars amphip- in Cucurbitaceae (Condon  & Norrbom, 1999), and except
neustic. Anterior spiracle sessile or nearly so, and usually for a few species of Eutreta and several Old World taxa, spe-
fan-shaped or bimodal, with 2–53 lobes. Cephalopharyngeal cies of Tephritinae breed in Asteraceae (Prado et al., 2002).
skeleton typically muscomorphan, with right and left man- Genera of Trypetinae breed in various plant families. Most
dibles equally developed, with one to two subapical teeth in American species are fruit or seed-feeders, but some Trype-
first and second instars, zero to two in third instar. Posterior tini are leaf or stem miners. Adult tephritids generally feed
spiracular plate weak, unpigmented, without peritreme, spi- only on liquid materials or substances soluable in saliva,
racular slits straight, in third instars the three slits oriented such as honeydew, bird feces, plant exudates (including those
radially from the ecdysial scar, subparallel to perpendicular from oviposition wounds), and microbes or debris on plant
to one another (in frugivorous species angle between dor- surfaces, but adults of Blepharoneura have labellar teeth and
rasp plant tissues (Condon & Norrbom, 1999). The behavior
sal and ventral slits usually no more than 90°, but greater in
of many fruit fly species has been extensively studied (see
some species of Tephritinae).
Aluja  & Norrbom, 1999). Among the observed behaviors
Puparia usually barrel-shaped, but rarely bean-shaped or
with flattened caudal segment; smooth or wrinkled, with dis- are elaborate courtship (e.g., various wing and body move-
tinct or indistinct segmentation. Color from white to yellow, ments, production of nuptial gifts), lekking, male combat for
brown or black, and color may vary from the anterior to the mating sites or guarding oviposition sites, and female phero-
posterior end. mone marking of oviposition sites. Mature third instar lar-
Adult tephritids are most commonly confused with other vae of some Tephritidae (e.g., many Dacinae, Adramini, and
picture-winged flies, in the Neotropical Region particularly Blepharoneura) can jump. Some fruit fly species are mimics
with Ulidiidae. The shape of the subcosta (bent sharply for- of wasps (e.g., Toxotrypana, some Anastrepha and Pseudo-
ward subapically and usually weaker or foldlike beyond the phorellia spp.) or jumping spiders (Zonosemata, Fig. 12; at
bend) will diagnose most New World species from Ulidii- least some Rhagoletis, and probably Cecidochares, Fig.  3;
dae, although in some taxa, especially species with elongate Tomoplagia, Fig. 10; and other genera with similarly banded
wings (e.g., Toxotrypana, some species of Tachiniscinae) the wing patterns).
bend is less than the typical 80–90° found in most tephrit- Tephritidae are the most agriculturally important fam-
ids and/or the apical part is not as distinctly weaker. Some ily of flies, including major pest species of mango, citrus,
Plastotephritinae (Platystomatidae; Old World), various Pyr- and many other important fruit and vegetable crops. In
gotidae, and a few other species of Tephritoidea also have a the Neotropical Region, the most important pests are vari-
similarly shaped, strongly bent subcosta, although these flies ous species of Anastrepha, Rhagoletis, and Toxotrypana,
lack distinct frontal setae, which within the Tephritoidea and the introduced Ceratitis capitata, which is widespread
are present only in Tephritidae (including Tachiniscinae)
in the New World tropics, and two species of Bactrocera:
and some Ulidiidae. The non-metallic body color of New
World tephritids (except the abdomen of Tachinisca) is a B.  carambolae Drew  & Hancock, which is invasive in the
useful field character, as many Ulidiidae, Platystomatidae, Guianas, and B. oleae (Rossi), which was introduced to the
and Richardiidae are metallic blue or green. Larvae and pu- USA (California) and Mexico (detected in 2000). Some Neo-
paria are typically muscomorphan, recognized by the lack tropical species of Tephritinae (e.g., Cecidochares connexa
of a sclerotized peritreme surrounding the posterior spiracle, (Macquart), Eutreta xanthochaeta Aldrich) have been used
which is not elevated off the surface of the caudal segment as biological control agents of their Asteraceae hosts that are
(Fig. 169) (B.A. Foote, 1991). Ferrar (1987) compiled much invasive weeds in other regions.
of the published information on these stages.
Classification
Biology
Higher classification within the Tephritidae has been un-
Neotropical Tephritidae, as far as known, are all phytopha- stable. Six subfamilies are currently recognized (Korneyev,
gous, except for the Tachiniscinae, which are presumably 1999a): Tachiniscinae, Blepharoneurinae, Phytalmyiinae,
parasitoids. The only tachiniscine species that has been Trypetinae, Dacinae, and Tephritinae. The bulk of the Neo-
reared was reportedly bred from larvae or pupae of Afrotro- tropical genera and species belong to the subfamilies Try-
pical saturniid moths (Korneyev & Norrbom, 2006), but all petinae and Tephritinae. The Phytalmyiinae do not occur in
tachiniscine species have similar female terminalia, suggest- the New World, nor do the Dacinae, except for three intro-
ing they are also parasitoids. A number of Old World gen- duced species of Ceratitis and Bactrocera. The Tachinisci-
era of Phytalmiinae are saprophages, often with specialized nae and Blepharoneurinae each include only a few genera
habits, such as breeding in damaged bamboo or under bark and are thought to be among the basal clades of the family.
of newly fallen trees. The larvae of the phytophagous spe- Of the former, only Tachinisca, Neortalotrypeta, Agnitrena,
cies feed within various plant tissues, including fruits, seeds, Ischyropteron, and the fossil genus Protortalotrypeta are
flowers, leaves, stems, or roots. Many Tephritinae feed with- known from the Neotropical Region, but none have been re-
in stem, root, bud, or flower galls that they induce on the host ported from Mexico or Central America. Of the three known
plant. Blepharoneura (Blepharoneurinae) breed exclusively Neotropical genera of Blepharoneurinae (two extant), only
TEPHRITIDAE (fruit flies, moscas de frutas) 68 911

Blepharoneura occurs in the Central American region (in- in relation to the levels of the postsutural supra-alar and
cluding Neotropical Mexico). intra-alar setae, which are relatively consistent in posi-
tion (Figs.  42, 52). Several other characters that are sig-
Identification nificant for tephritid identification and may not be familiar
to the general dipterist are as follows. The anepisternum
Various information on fruit fly taxonomy and natural of Tephritoidea has an internal, slightly oblique, vertical
history, such as the generic classification, world names and phragma (sometimes referred to as a suture) that is often
New World specimen databases, a host plant database for visible externally (Fig. 43) and is a useful landmark. Par-
some New World genera, and pages on some individual ticularly in many genera of Tephritinae, some setae (e.g.,
genera and species, is available on the fruit fly pages on the posterior orbital seta, posterior notopleural seta, some or
Internet at the Diptera site (http://www.sel.barc.usda.gov/ all postocular setae, and some or all scutal setulae) tend
Diptera/tephriti/tephriti.htm). Interactive keys to the adults to be paler in color and often more lanceolate (i.e., stouter
and larvae of fruit fly pest species (Carroll et al., 2002, 2004) medially) than the other typically slender, or acuminate, se-
are available on the Delta site (http://delta-intkey.com/ffl/ tae (e.g., Figs. 25, 49). The wings of some tephritid genera
www/_wintro.htm). Adult tephritid specimens preserve well have a bulla, a darker, usually concave, spotlike area, or
either pinned or in ethanol. argents, which are usually spotlike, silver or white areas
No attempt has been made to key all tephritid genera due to microtrichia that change in color or appearance (or
worldwide since that of Hendel (1914a), which is badly disappear) when viewed at different angles. Common gen-
outdated. However, generic keys for most of the major bio- eral types of tephritid wing patterns include the following:
geographic regions are available. Foote et al. (1993) keyed banded (e.g., Figs. 1–8, 10); radiate, with the center of the
the genera of the continental United States and Canada, and wing mostly patterned, and raylike marks extending to the
Foote (1980) provided a key to the genera of Mexico and the margins (e.g., Figs.  74, 119, 146); reticulate, with netlike
Neotropical Region. The latter key, especially the first cou- or mostly dark pattern with numerous hyaline spots (e.g.,
plets, can be difficult to use, as it followed the higher classi- Figs.  78, 86–87, 94); and stellate, with a somewhat star-
fication then in use. The key provided here is more practical shaped anterior subapical mark with rays radiating to the
and is not phylogenetic for the most part. One of the most wing margin (e.g., Figs. 125, 126, 144, 151). Some species
significant characters it uses is the position of the dorsoce- have patterns that are intermediate or combine these ele-
ntral seta, which varies in location anteriorly to posteriorly, ments (e.g., Figs. 67–68).

Key to the genera of Tephritidae of Mexico and the Neotropical Region

1. Thorax with most major setae short or absent, acrostichal setae less than 1.2 times as long as
distance between them (Fig. 41); eye relatively small, genal height more than half long diameter
of eye (Fig. 34); lunule large and setulose (Fig. 14); antenna short, first flagellomere no longer
than pedicel, sometimes setulose; wing elongate (more than 2.5 times as long as wide) and
predominantly brown, at most with one large hyaline area on anterior margin and two on posterior
margin or sometimes narrow hyaline streaks or spots medially (Figs. 69, 136); scutellum usually
high and convex or produced dorsoapically, entirely microtrichose, and without distinct yellow
markings; large (wing length greater than 7 mm), predominantly brown. . . . . . . . . . . . . . . . . . . 2
– At least some major thoracic setae well developed, or if short, genal height less than half long
diameter of eye or wing shorter or with different pattern; lunule rarely setulose; first flagellomere
nonsetulose; other characters various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

2. Wing with only narrow irregular hyaline streaks in addition to hyaline triangle on anterior margin
distal to apex of vein R1 (Fig. 69); male forefemur stout, with row of ventral setae stout and spinelike
(Fig. 55); tibiae slender to moderately stout . . . . . . . . . . . . . . . . . . . . . . . . . Agallamyia Norrbom
– Wing with two large hyaline areas on posterior margin and with or without hyaline triangle on
anterior margin (Fig. 136); male forefemur sometimes moderately stout, but if so, row of ventral
setae more slender and not spinelike; tibiae, especially hind tibia, sometimes stout (Fig. 57). . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pyrgotoides Curran

3. Scutellum with three or more pairs of marginal setae (Fig. 43) and/or at least basal segment of
vein CuA setulose dorsally (Fig. 59). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
– Scutellum with one to two pairs of marginal setae (Figs. 42, 48, 49) and vein CuA nonsetulose. . 9
912
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

2 Anastrepha 3 Cecidochares

4 Cryptodacus 5 Haywardina

6 Molynocoelia 7 Pseudophorellia

Figs.  68.2–7. Adults: dorsal view of (2)  Anastrepha obliqua (Macquart); (3)  Cecidochares fluminensis (Lima); (4)  Cryptodacus lopezi
Norrbom; (5) Haywardina cuculi (Hendel); (6) Molynocoelia lutea Giglio-Tos, (Norrbom, 2006b, fig. 30); (7) Pseudophorellia anypsilon
Norrbom, (Norrbom, 2006b, fig. 55). Figures 2–5 illustrated by T.R. Litwak.
TEPHRITIDAE (fruit flies, moscas de frutas) 68 913

8 Rhagoletotrypeta 9 Tachinisca

10 Tomoplagia 11 Xanthaciura

12 Zonosemata

Figs.  68.8–12. Adults (concluded): dorsal view of (8)  Rhagoletotrypeta rohweri Foote, (Nearctic); (9)  Tachinisca cyaneiventris
Kertész, (Korneyev & Norrbom, 2006, fig. 4); (10) Tomoplagia stonei Aczél; (11) Xanthaciura sp.; and (12) Zonosemata guybushi Norrbom,
(Norrbom, 2002b, fig. 2C). Figure 8 illustrated by A. Cushman; Figures 10–11 by T.R. Litwak.
914
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

flgm 1

13 Acidogona 14 Agallamyia

15 Blepharoneura
orb s

17 Ceratitis

16 Campiglossa 18 Ceratodacus
i vt s
orb s
oc s orb s
oc s
fr s o vt s
fr s pocl s

ar

flgm 1
gn s

19 Chetostoma 20 Cryptodacus
lbl

21 Dioxyna

Figs. 68.13–21. Heads: lateral view of (13) Acidogona melanura (Loew), (Foote et al., 1993, fig.  70); anterior view of (14)  Agallamyia
pendula, (Norrbom et al., 2010, fig. 12); lateral view of (15) Blepharoneura sp., (redrawn from R.H. Foote, 1980, fig. 17); (16) Campiglossa
genalis (Thomson), (Nearctic, redrawn from Foote et al., 1993, fig. 102); (17) Ceratitis capitata (Wiedemann), (Foote et al., 1993, fig. 35);
(18)  Ceratodacus longicornis Hendel, (R.H.  Foote, 1980, fig.  19); (19)  Chetostoma  sp., (redrawn from R.H.  Foote, 1980, fig.  20);
(20) Cryptodacus tau (Foote), (R.H. Foote, 1980, fig. 26); (21) Dioxyna picciola (Bigot), (MND, fig. 66.3).
Abbreviations: ar, arista; flgm 1, first flagellomere; fr s, frontal setae; gn s, genal seta; i vt s, inner vertical seta; lbl, labella; oc s, ocellar seta;
orb s, orbital setae; o vt s, outer vertical seta; pocl s, postocular setae.
TEPHRITIDAE (fruit flies, moscas de frutas) 68 915

4. Thorax and abdomen stout (Fig. 9); thorax yellow, abdomen metalic blue to purple; with numerous
large, stout setae including: vibrissa; three or more postpronotal setae; two to three postsutural
supra-alar setae; four to five scutellar setae; eight or more anepisternal setae, including four anterior
to phragma; three anepimeral setae on protuberance; and rows of stout erect setae posteriorly
on abdominal tergites and sternites; veins  R4+5 and M strongly and evenly divergent apically
(Fig. 142); vein Sc subapically anteriorly turned at angle of approximately 45°; Ecuador, Peru,
Bolivia, Brazil . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Tachinisca Kertész]
– Thorax and abdomen shape and color various, but not metallic; vibrissa absent; at most with two
postpronotal, three postsutural supra-alar, three scutellar, four anepisternal (at most one anterior
to phragma), and two anepimeral setae, the latter not on protuberance; abdominal setae not erect;
veins R4+5 and M at most slightly divergent apically; vein Sc subapically anteriorly turned at angle
of 80–90°. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

5. Anepisternum with large seta just anterior to vertical phragma (Fig. 43); one to two postpronotal
setae; usually two frontal setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
– Anepisternum without outstanding setae anterior to vertical phragma; zero to one postpronotal
setae; usually three to four frontal setae (zero in Agnitrena, zero to two in some Pseudophorellia
spp.). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

6. Wing with distinct markings, usually predominantly brown (Figs.  75–76); vein  CuA1 setulose
dorsally (Fig. 59); crossvein r-m near or distal to midlength of cell dm (measured along vein M);
prementum elongate and strongly convex (Fig. 15); labella with rows of spinules (sometimes not
exposed in dry specimens); face without numerous black spots; antenna moderately long, but
usually not reaching ventral margin of face. . . . . . . . . . . . . . . . . . . . . . . . . . Blepharoneura Loew
– Wing with only diffuse markings (at least in extant species) (Fig. 81); vein CuA1 nonsetulose dorsally;
crossvein r-m at or proximal to basal two-fifths of cell dm; prementum small and not strongly
convex; labella without rows of spinules; face yellow with numerous black spots (Fig.  18);
antenna elongate, reaching at least ventral margin of face; Brazil, Guyana, Peru. . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Ceratodacus Hendel]

7. Head with anterior margin strongly receding (as in Fig. 29); eye small, genal height more than half
long diameter of eye; frontal setae absent; Argentina . . . . . . . . . . . . [Agnitrena Korneyev, in part]
Head with anterior margin not receding; eye large, genal height less than one-third long diameter
of eye; usually two or more frontal setae present (zero to two in some Pseudophorellia spp.) . . 8

8. Abdominal syntergite 1+2 with one to several lateral setae distinctly larger than others (Fig. 7);
ocellar, postocellar, acrostichal, posterior orbital, and/or presutural supra-alar setae often small
or absent; scutum without microtrichia; vein CuA1 with at least basal third between bm-cu and
dm-cu not bordered by brown anteriorly (Figs. 7, 134); base of vein CuA always and vein CuA1
usually setulose dorsally; phallus with large bilobed membranous basal lobe on glans. . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudophorellia Lima
– Abdominal syntergite 1+2 with lateral setae similar in size; ocellar seta various, but other
setae mentioned above usually well developed; scutum microtrichose; vein CuA1 with basal half
between bm-cu and dm-cu usually bordered by brown (Fig. 104); base of vein CuA sometimes
setulose, CuA1 usually not setulose; phallus without membranous basal lobe on glans. . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hexachaeta Loew

9. Scutum with two dorsocentral setae, one presutural, one postsutural (Fig. 3). . . . . . . . . . . . . . . 10
– Scutum with zero to one postsutural dorsocentral setae, sometimes only slightly posterior to
transverse suture (Figs. 42, 49, 52) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
916
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

23 Epochrinopsis

22 Ensina

24 Euaresta

fc car

26 Goedenia
27 Gymnocarena

25 Eutreta 28 Molynocoelia

29 Neortalotrypeta

30 Neotephritis 31 Oedicarena

Figs. 68.22–31. Heads (continued): lateral view of (22) Ensina sp.; (23) Epochrinopsis sp.; (24) Euaresta sp., (R.H. Foote, 1980, fig. 22);
anterior view of (25) Eutreta distincta (Schiner); lateral view of (26) Goedenia grineliae (Coquillett), (Nearctic, Foote et al., 1993, fig. 20);
dorsolateral view of (27)  Gymnocarena diffusa (Snow), (Nearctic, MND, fig.  66.7); lateral view of (28)  Molynocoelia lutea Giglio-Tos,
(Norrbom, 2006b, fig. 2); (29) Neortalotrypeta bicolor Norrbom, (redrawn from Norrbom, 1994a, fig. 1C); (30) Neotephritis finalis (Loew),
(R.H. Foote, 1980, fig. 27); and (31) Oedicarena persuasa (Osten Sacken), (Nearctic, Foote et al., 1993, fig. 294). Figures 22 and 23
illustrated by J.M. Metz; Figure 25 by S. Stewart.
Abbreviation: fc car, facial carina.
TEPHRITIDAE (fruit flies, moscas de frutas) 68 917

32 Paracantha 33 Parastenopa 34 Pyrgotoides

35 Rachiptera
36 Rhagoletis

37 Tephritis

38 Terellia 39 Xanthaciura

40 Zonosemata

Figs. 68.32–40. Heads (concluded): anterodorsal view of (32) Paracantha gentilis Hering; lateral view of (33) Parastenopa limata (Coquillett),
(Nearctic, Foote et al., 1993, fig. 263); (34) Pyrgotoides sp., (R.H. Foote, 1980, fig. 24); anterior view of (35) Rachiptera sp.; lateral view
of (36) Rhagoletis pomonella (Walsh), (Nearctic, Foote et al., 1993, fig. 61); and (37) Tephritis sp.; anterior view of (38) Terellia palposa
(Loew), (Nearctic, Foote et al., 1993, fig. 44); dorsal view of (39) Xanthaciura insecta (Loew), (Foote et al., 1993, fig. 77); lateral view of
(40) Zonosemata vittigera (Coquillett), (Nearctic, Foote et al., 1993, fig. 34). Figure 32 illustrated by T.B. Griswald; Figures 35 and 37 by
J.M. Metz.
918
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

10. Wing banded, with basal or subbasal band covering crossvein  h, and three to four additional
bands, including band from pterostigma to posterior wing margin covering bm-cu and CuA2, band
covering r-m and dm-cu, apical band, and sometimes band across cell  cu1 (Figs.  3, 130–131);
cells  r1 and r2+3 sometimes with marginal hyaline marks or band, but if extended to vein  R4+5,
reaching it distal to level of crossvein r-m; vein R4+5 dorsal setation various, sometimes bare or
setulose only proximal to crossvein dm-cu. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
– Wing radiate or reticulate (Figs. 77, 97), or if somewhat banded, bands distal to crossvein dm-cu
or cells r1 and r2+3 with triangular marginal hyaline mark reaching vein R4+5 at or proximal to level
of crossvein r-m (Figs. 114, 128); vein R4+5 densely setulose dorsally beyond crossvein dm-cu. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

11. Posterior orbital seta absent; scutum with presutural and postsutural lateral shiny nonmicrotrichose
areas; anepisternum sometimes with shiny nonmicrotrichose area; wing without band across middle
of cell cu1, and with hyaline area in distal half of cell br (Fig. 130) . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Procecidochares Hendel, in part
– Posterior orbital seta present; scutum with or without presutural and postsutural shiny nonmicrotrichose
areas; anepisternum entirely microtrichose; wing often with band across middle of cell cu1 or with
distal half of cell br entirely brown (Fig. 3). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

12. Thoracic pleuron usually entirely brown (sometimes bluish-gray if densely microtrichose), postpronotal
lobe and anepisternum rarely yellow; distal half of cell br without hyaline area (Fig. 3) or notopleural
microtrichia denser than microtrichia of scutum, appearing velvety; posterior orbital seta
reclinate; some or all postocular setae often brown. . . . . . . . . . . . . . . Cecidochares Bezzi, in part
– Postpronotal lobe, at least dorsal half of anepisternum, and often additional parts of thoracic
pleuron yellow; distal half of cell br with large hyaline area (Fig. 131); notopleural microtrichia
similar to microtrichose parts of scutum; posterior orbital seta often inclinate; all postocular setae
yellow. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Procecidocharoides Foote

13. Wing with apical dark rays radiating from middle of cell r4+5 separated by large marginal hyaline
marks, cell r2+3 with at least two marginal hyaline marks and cell r4+5 with one marginal or submarginal
hyaline mark (Figs. 77, 97) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
– Wing apically without dark rays radiating from middle of cell  r4+5, usually with two to three
oblique or transverse dark bands separated by hyaline bands or wedge-shaped marks extending
from posterior margin, margins of cells r2+3 and/or r4+5 often entirely brown (Figs. 114, 128). . 15

14. Middle of wing brown, with paler areas but without hyaline marks, including all of cells br and
dm, and except for apices, cells r1, r2+3, and r4+5 (Fig. 77); pterostigma also entirely brown. . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caenoriata Foote
– Middle of wing and pterostigma with hyaline spots or marks (Fig. 97) . . . . . Euarestopsis Hering

15. Cell r2+3 with large triangular marginal hyaline mark at apex of vein R2+3 (Fig. 114); cell r1 without
large triangular marginal hyaline mark . . . . . . . . . . . . . . . . . . . . . . . . . . Neotaracia Foote, in part
– Cell r2+3 without large triangular marginal hyaline mark at apex of vein R2+3 (Fig. 128); cell r1 with
large triangular marginal hyaline mark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polionota Wulp

16. Cell cup without posteroapical lobe (Fig. 61), CuA2 meeting A1 at angle of at least 90°; frons with
one orbital seta (Fig.  26); postocular setae all acuminate and dark brown; two scutellar setae;
vein R4+5 nonsetulose; wing mostly (rarely entirely) hyaline with narrow brown bands or spots
(Fig. 110). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
– Cell cup usually with at least small posteroapical lobe (Figs. 60, 62, 63) or CuA2 meeting A1 at
acute angle (Fig. 140), if doubtful, at least one of following present: frons with two or more orbital
setae; postocular setae mixed acuminate, brown and pale, lanceolate; one scutellar seta; vein R4+5
setulose dorsally; and/or wing mostly infuscated, reticulate, or stellate (Figs. 107, 151). . . . . . 19
TEPHRITIDAE (fruit flies, moscas de frutas) 68 919

17. Anepisternum, anepimeron, and katepisternum each at least partially nonmicrotrichose and shiny;
face moderately to deeply concave; western United States, northern Mexico . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Goedenia Freidberg & Norrbom]
– Anepisternum, anepimeron, and katepisternum entirely microtrichose, matte; face at most slightly
concave. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

18. Proboscis short to long geniculate, labella length more than half head height (at least as long as
in Fig. 26); abdominal tergites at least partially yellow; scutellum medially yellow; hind femur
without outstanding anteroventral setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhynencina Johnson
– Proboscis capitate to short geniculate, labella length usually less than half head height (no longer
than in Fig. 26); abdominal tergites entirely brown; scutellum medially yellow or entirely brown;
hind femur often with one to several outstanding anteroventral setae. . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neomyopites Freidberg & Norrbom

19. Dorsocentral seta absent or closer to level of intra-alar seta than to level of postsutural supra-alar
seta (Figs. 42, 47). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
– Dorsocentral seta at or anterior to level of postsutural supra-alar seta or closer to it than to level of
intra-alar seta (Figs. 44, 49–50, 52). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

20. Presutural supra-alar seta absent or if weakly present (some Toxotrypana), postpronotal,
acrostichal, dorsocentral, anepisternal, and basal scutellar setae also weak or absent; ocellar and
posterior orbital setae minute or absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
– Presutural supra-alar seta well developed or if absent (Oedicarena nigra Hernández-Ortiz),
ocellar seta and posterior orbital seta well developed (as in Fig. 31) . . . . . . . . . . . . . . . . . . . . . 23

21. Anatergite with long, fine setulae (Fig. 46); lobe of cell cup short, about as long as width of cell
(Fig. 98); wing with transverse bands; postpronotal, acrostichal, dorsocentral, and basal scutellar
setae well developed; mostly Old World tropical, in America south to central Mexico. . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Euphranta Loew]
– Anatergite without long, fine setulae; lobe of cell cup elongate, much longer than width of cell;
wing without transverse bands (in New World species), with only long marginal band and infuscation
along cell cup and vein CuA1 (Figs. 72, 148); postpronotal, acrostichal, dorsocentral, and/or basal
scutellar setae often small or absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

22. Abdominal syntergite 1+2 elongate and petiolate, basal half more or less parallel-sided; vein R2+3
with three sharp bends, often with spur veins (Fig. 148); cell bm less than 1.5 times as wide as cell
cup; scutum with weak medial longitudinal depression; first flagellomere of antenna no more than
2.5 times as long as wide, and not reaching ventral margin of face; oviscape of female at least as
long as rest of abdomen, often strongly curved. . . . . . . . . . . . . . . . . . . . Toxotrypana Gerstaecker
– Syntergite 1+2 short and not petiolate; vein R2+3 without sharp bends or spur veins (Fig.  72);
cell bm broad, approximately 2 times as wide as cell cup; scutum without medial depression; first
flagellomere more than 3 times as long as wide, usually reaching ventral margin of face; oviscape
usually no longer than rest of abdomen and not strongly curved . . . . . . . . . Bactrocera Macquart

23. Vein M curved forward apically, meeting costa in gradual curve or at slight angle (Figs. 2, 71);
wing pattern various but usually similar to Fig. 2, typically including the following: costal band
extended to apex of vein R1; “S-band” extending obliquely from cell cup, across r-m, to wing
margin in cell r1, then bordering costa to wing apex; and subapical and posterior apical bands often
forming inverted “V-band”; or with only uninterrupted costal band, mark along cubital veins,
and sometimes another along dm-cu (Fig. 71); ocellar seta weak (except in A. tripunctata Wulp);
eversible membrane of female with dorsobasal cluster of usually long, hooklike teeth (Fig. 159).
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anastrepha Schiner
– Vein M not curved forward apically, meeting costa at distinct angle (Figs. 12, 122); wing pattern
various, different from above; ocellar seta often strong; eversible membrane without dorsobasal
cluster of hooklike teeth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
920
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

vt phgm

presut area
sct pprn s

trn sut
41 Agallamyia npl s 43 Blepharoneura
presut spal s
psut area
sct psut spal s

dc s

pal s

acr s

sctl ial s

a sctl s

p sctl s

42 Anastrepha

44 Carpomya 45 Ceratitis

Figs. 68.41–45. Thoraces: dorsal view of (41) Agallamyia pendula, (Norrbom et al., 2010, fig. 13); and (42) Anastrepha ludens (Loew),
(MND, fig. 68.8); lateral view of (43) Blepharoneura femoralis Wulp, (Norrbom & Condon, 1999, fig. 6.2D); dorsal view of (44) Carpomya
tica Norrbom, (Norrbom 1997a, fig. 1); and (45) Ceratitis capitata (Wiedemann). Figures 41 and 45 illustrated by T.R. Litwak.
Abbreviations: acr s, acrostical seta; a sctl s, anterior scutellar seta; dc s, dorsocentral seta; ial s, intra-alar seta; npl s, notopleural setae;
pal s, postalar seta; pprn s, postpronotal seta; presut area sct, presutural area of scutum; presut spal s, presutural supra-alar seta; p sctl s,
posterior scutellar seta; psut area sct, postsutural area of scutum; psut spal s, postsutural supra-alar seta; sctl, scutellum; trn sut, transverse
suture; vt phgm, vertical phragma.
TEPHRITIDAE (fruit flies, moscas de frutas) 68 921

sctl

anatg

mtg

46 Euphranta

48 Lamproxyna

47 Ischyropteron

51 Procecidochares

49 Neotephritis

50 Paracantha

Figs. 68.46–51. Thoraces (continued): posterior view of scutellum, mediotergite, and setose anatergite of (46) Euphranta canadensis (Loew),
(Nearctic); dorsal view of (47) Ischyropteron nigricaudatum Bigot, (Korneyev & Norrbom, 2006, fig. 45); (48) Lamproxyna nitidula Hendel;
(49) Neotephritis finalis (Loew), (MND, fig. 66.11); (50) Paracantha gentilis Hering; lateral view of scutellum and adjacent thorax and
abdomen of (51) Procecidochares gibba (Loew), (Nearctic, Foote et al., 1993, fig. 36). Figure 46 illustrated by A.L. Norrbom & L.H. Rodriguez;
Figure 48 by T.R. Litwak; and Figure 50 by T.B. Griswald.
Abbreviations: anatg, anatergite; mtg, mediotergite; sctl, scutellum.
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

24. Crossveins r-m and dm-cu both within subapical band (band reaching costa in cell r1) (Fig. 1);
acrostichal seta absent; scutum without microtrichia; arista almost bare, with only few minute
basal hairs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Alujamyia Norrbom
– Crossvein r-m not in band or it and dm-cu in separate bands (Figs. 6, 12, 106, 117) or sometimes
(some Cryptodacus) both within discal band (band crossing pterostigma) or (some Parastenopa)
within large area covering most or all of cell dm (Fig. 121); acrostichal seta present (except in
Oedicarena nigra, in which scutum is mostly microtrichose); arista usually with at least some
minute hairs on apical half (rarely bare in Parastenopa; unknown for Ischyropteron). . . . . . . . 25

25. Wing hyaline except for pterostigma and entire apex distal to dm-cu (Fig.  106); scutum nonmi-
crotrichose dorsally, yellow with two pairs of broad brown stripes on scutum (Fig. 47); basal scutellar
seta relatively close to apical seta, at midlength of scutellum; Brazil. . . . . . [Ischyropteron Bigot]
– Wing with more extensive markings proximal to dm-cu; scutal color and microtrichia various;
basal scutellar seta more separated from apical seta, proximal to midlength of scutellum (Figs. 6,
12) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26

26. Frontal setae absent; head with anterior margin strongly receding (as in Fig. 29) and eye small,
genal height more than half long diameter of eye; wing with large medial brown area as broad as
long covering pterostigma and crossvein r-m and extending to vein CuA1 (Fig. 70); Argentina . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Agnitrena Korneyev, in part]
– At least two well-developed frontal setae present; head with anterior margin not receding; eye
usually large and genal height less than two-fifths its long diameter (Figs. 20, 33), or if smaller
(Fig. 31), wing with band or spots crossing pterostigma and r-m narrow (Fig. 117) . . . . . . . . . 27

27. Middle of wing with large, broad, usually ovoid brown area covering pterostigma, crossveins
bm-cu and r-m, cell br distal to bm-cu, and basal half or more of cell dm (Figs. 121–122); distal to
dm-cu, two apical bands present, anterior band bordering costa, and posterior band more or less
parallel to it crossing cell r4+5; ocellar seta usually small and weak (Fig. 33). . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parastenopa Hendel, in part
– Crossvein bm-cu not covered by brown, or it and r-m in different bands, sometimes connected
posteriorly or anteriorly, but separated by hyaline area at least in cell br and usually in base of
cell dm (Figs. 4, 6, 12, 117); apical bands various; ocellar seta well developed except in most
Cryptodacus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28

28. Arista short to long plumose, hairs at least half width of first flagellomere (Fig. 28); wing brown
on distal half or with six bands, including subcostal band (from pterostigma to apex of cell cup),
small band originating in middle of cell r1 sometimes covering r-m and connected posteriorly to
discal band to form Y-shaped mark, and posterior apical band (crossing distal half of cells r4+5
and m) (Fig. 6); scutum without microtrichia, yellow, with two to four pairs of brown mark-
ings including presutural ovoid dark spots and postsutural stripes or spots; frons with two frontal
setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Molynocoelia Giglio-Tos, in part
– Arista pubescent, hairs less than one-fourth width of first flagellomere; wing pattern various, but
lacking at least posterior apical band; crossvein r-m covered by discal band (Figs. 4, 12, 117);
anterior apical band often interrupted or reduced to apical spot; scutal microtrichia and color
various, but usually not mostly yellow with pairs of brown markings; frons usually with three or
more frontal setae (two in some Cryptodacus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

29. Crossvein r-m distal to middle of cell dm, more than 0.60 distance from bm-cu to dm-cu (Figs. 4,
83); vein CuA1 usually bordered with brown between bm-cu and discal band; accessory costal
band absent; antenna elongate, extended beyond ventral facial margin; face usually with brown
spot(s) or inverted U-shaped marking (Fig. 20); aculeus tip sagittate and finely serrate (Fig. 161).
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cryptodacus Hendel, in part
– Crossvein r-m near middle of cell dm, less than 0.60 distance from bm-cu to dm-cu (Figs. 12,
117); vein CuA1 without brown border between bm-cu and discal band; accessory costal band
usually present; antenna not extended beyond ventral facial margin (Figs. 31, 40); face without
markings; aculeus tip various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
TEPHRITIDAE (fruit flies, moscas de frutas) 68 923

pprn s

npl s
presut spal s

psut spal s

dc s

pal s
acr s

ial s

a sctl s

p sctl s

52 Rhagoletis
53 Terellia

54 Aciurina 55 Agallamyia 56 Campiglossa

57 Pyrgotoides 58 Tephritis

Figs. 68.52–58. Thoraces (concluded) and legs: dorsal view of thorax of (52) Rhagoletis pomonella (Walsh), (Nearctic, MND, fig. 66.9); and
(53) Terellia palposa (Loew), (Nearctic); anterior view of left foreleg of (54) Aciurina thoracica Curran, (Nearctic); posterior view of right
foreleg of (55) Agallamyia pendula, (Norrbom et al., 2010, fig. 14); anterior view of left hind femur of (56) Campiglossa genalis (Thomson),
(Nearctic, Foote et al., 1993, fig. 96); anterior view of left hind leg of (57) Pyrgotoides crassipes Curran, (R.H. Foote, 1980, fig. 10); anterior
view of left hind femur of (58) Tephritis subpura (Johnson), (Nearctic, Foote et al., 1993, Fig. 95). Figure 53 illustrated by J.M. Metz; Figures
54–55 by T.R. Litwak.
Abbreviations: acr s, acrostical seta; a sctl s, anterior scutellar seta; dc s, dorsocentral seta; ial s, intra-alar seta; npl s, notopleural setae;
pal s, postalar seta; pprn s, postpronotal seta; presut spal s, presutural supra-alar seta; p sctl s, posterior scutellar seta; psut spal s, postsutural
supra-alar seta.
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

30. Antennal first flagellomere rounded apically (Fig. 31); usually three frontal setae; scutum without
white or yellow stripes, and microtrichose except small paired bare spots at medial end of
transverse suture and at base of dorsocentral seta and sometimes three narrow stripes; postocular
setae slender, sometimes poorly differentiated from postgenal setulae; western United States to
central Mexico. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Oedicarena Loew, in part]
– Antennal first flagellomere with small, acute dorsoapical lobe (Fig.  40); usually four or more
frontal setae; scutum with single medial and paired sublateral white or yellow stripes (Fig. 12),
without microtrichia; postocular setae strong . . . . . . . . . . . . . . . . . . . . . . . Zonosemata Benjamin

31. Posterior orbital seta distinctly inclinate (Fig. 38); vein R4+5 bare dorsally or with at most five setulae
on or near basal node (Fig. 62). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
– Posterior orbital seta reclinate (Fig.  27) or absent; vein  R4+5 often more extensively setulose
dorsally. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35

32. Wing pattern reticulate (Fig. 78); posterior orbital seta white, lanceolate, paler than anterior seta;
proboscis short geniculate (Fig. 16); phallus with small spines proximal to glans. . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Campiglossa Rondani, in part
– Wing pattern banded or highly reduced, not reticulate; posterior orbital seta similar to anterior seta
in color and shape; proboscis capitate; phallus without small spines proximal to glans. . . . . . . 33

Cu A1

cup

59 Blepharoneura

60 Euaresta 61 Goedenia 62 Terellia

63 Tetreuaresta

Figs. 68.59–63. Wings and wing bases (minus pattern): dorsal view of wing of (59) Blepharoneura sp.; dorsal view of wing base, showing
shape of cell cup, of (60) Euaresta aequalis (Loew), (Nearctic, Foote et al., 1993, fig. 18); (61) Goedenia caurina (Doane), (Nearctic, Foote
et al., 1993, fig. 17); wing base, with enlarged inset of setulae on node at fork of radial sector, of (62) Terellia occidentalis (Snow), (Nearctic,
Foote et al., 1993, fig. 47); dorsal view of wing of (63) Tetreuaresta sp., (redrawn from Foote et al., 1993, fig. 107). Figure 59 illustrated by
T.R. Litwak.
TEPHRITIDAE (fruit flies, moscas de frutas) 68 925

33. Thorax dark brown except dorsal margin of anepisternum and most of scutellum; first
flagellomere with dorsoapical point (Fig. 36) . . . . . . . . . . . . . . . . . . . . . . Rhagoletis Loew, in part
– Thorax predominantly yellow, scutum usually with lyre-shaped dark pattern (Fig.  53); first
flagellomere rounded dorsoapically. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

34. Scutum entirely microtrichose; acrostichal and usually dorsocentral setae on yellow area, the
former lacking brown spot at base; wing mostly hyaline, or if banded, apex of cell r4+5 hyaline;
vein R4+5 usually bare, at most with one to two setulae on node; abdomen without spots or if spots
present, usually broader than long; male foretarsus and claws usually asymmetric; mainly Nearctic,
south to northern Mexico and the Bahamas. . . . . . . . . . . . . . . . . . . . . . [Neaspilota Osten Sacken]
– Scutum microtrichose except small shiny bare spots at bases of dorsocentral and acrostichal setae
(Fig. 53); dorsocentral seta on large brown area, acrostichal seta with small brown spot at base;
wing banded, apex of cell r4+5 brown; vein R4+5 with two to five setulae on or near node (Fig. 62);
abdomen with submedial and usually sublateral rows of brown spots; male foretarsus and claws
symmetric; mainly Palearctic, in Nearctic Region south to northern Mexico. . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Terellia Robineau-Desvoidy]

35. Proboscis capitate (Fig. 31); scutellum with two pairs of marginal setae (Fig. 52); scutum with dors-
ocentral seta at or posterior to level of postsutural supra-alar seta or closer to it than to transverse
suture (Figs. 44, 52); wing usually banded, dark areas of pattern without numerous small paler
or hyaline spots; scutal and anepisternal setulae and postocular, posterior orbital, and posterior
notopleural setae various in color, but all acuminate (Figs. 19, 33, 40, 44, 45, 52) . . . . . . . . . . 36
– Any of five following characters present: proboscis geniculate (Figs. 16, 21); scutellum with only
one (basal) pair of marginal setae (Fig. 48); scutum with dorsocentral seta closer to transverse
suture than to level of postsutural supra-alar seta (Fig. 49); wing pattern reticulate, i.e., dark areas
with numerous small paler or hyaline spots (Figs. 65, 67, 86–87), or radiate (Figs. 74, 90); or some
or all of following setae or setulae lanceolate, white, and distinctly paler than other setulae or setae
(multiple setae, like postocular row or scutal setulae, may be mixed acuminate and lanceolate):
scutal or anepisternal setulae, postocular setae (Figs. 24, 25, 39), posterior orbital seta (Fig. 21),
or posterior notopleural seta (Fig. 49). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52

36. Scutellum high and strongly convex in posterior or lateral view (similar to Fig. 51), shiny nonmi-
crotrichose medially or on margin, mostly brown with distinct narrow yellow markings (Figs. 44,
45); notopleuron distinctly bicolored, yellow posteriorly, brown anterior part and side of scutum
posterior to notopleuron with brown velvetlike microtrichia more dense than elsewhere on
scutum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
– Scutellum flattened or if convex, entirely microtrichose or not distinctly bicolored; notopleuron
and scutum laterally without velvetlike, more densely microtrichose areas. . . . . . . . . . . . . . . . 38

37. Wing with small dark spots in cells bc, c, br, bm, and cup (Fig. 80); crossvein r-m near midlength
of cell dm, covered by band crossing pterostigma; band covering crossvein dm-cu not extending
across cell r4+5; posteroapical lobe of cell cup narrower near base than near middle; scutellum with
irregular yellow mark across basal half (Fig. 45), medially with erect setulae; male with anterior
orbital seta proclinate and expanded apically (Fig. 17). . . . . . . . . . . . . . . . . . . Ceratitis MacLeay
– Wing with small band across cells bc, c, br, bm, and cup (Fig. 79); crossvein r-m distinctly distal
to midlength of cell dm, almost covered by band covering crossvein dm-cu; band covering crossvein
dm-cu extending to costal margin; posteroapical lobe of cell cup evenly tapered; scutellum with
inverted, U-shaped yellow mark extending to apex (Fig. 44), setulose only laterally; both sexes
with anterior orbital seta reclinate and not expanded apically. . . . . . . . . . . . . . . . Carpomya Costa

38. Frons with one orbital and one to two weak frontal setae; gena large, its height approximately half
long diameter of eye (Fig. 29); body brown, except for entirely bright yellow to orange head; wing
predominantly brown, with 4 narrow transverse hyaline bands or spots medially, cell r4+5 brown
except basally (Fig. 112); Brazil, Argentina. . . . . . . . . . . . . . . . . . . . . [Neortalotrypeta Norrbom]
– Frons usually with two orbital setae (one in Haywardina cuculi (Hendel)), which has three frontal
setae and is predominantly yellow); genal height and body color various; wing pattern various,
but usually more predominantly hyaline, cell r4+5 usually with subapical hyaline area or mostly
hyaline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
926
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

64 Acidogona 65 Acinia

66 Aciurina 67 Acrotaenia

68 Acrotaeniacantha 69 Agallamyia

70 Agnitrena 71 Anastrepha

72 Bactrocera 73 Baryplegma

Figs. 68.64–73. Wings (continued): dorsal view of (64) Acidogona sp.; (65) Acinia picturata (Snow); (66) Aciurina mixteca Hernández-Ortiz;
(67) Acrotaenia testudinea (Loew); (68) Acrotaeniacantha radiosa Hering; (69) Agallamyia pendula Norrbom, (Norrbom et al., 2010, fig. 33);
(70)  Agnitrena igniceps Korneyev, (Korneyev  & Norrbom, 2006, fig.  34); (71)  Anastrepha avispa Norrbom, (Norrbom, 1998, fig.  5B);
(72) Bactrocera carambolae Drew & Hancock; and (73) Baryplegma apiata (Wulp). Figures 64–68, 72, and 73 illustrated by L.H. Rodriguez
and A.L. Norrbom.
TEPHRITIDAE (fruit flies, moscas de frutas) 68 927

74 Baryplegma 75 Blepharoneura

76 Blepharoneura 77 Caenoriata

78 Campiglossa 79 Carpomya

80 Ceratitis 81 Ceratodacus

82 Chetostoma 83 Cryptodacus

Figs.  68.74–83. Wings (continued): dorsal view of (74)  Baryplegma pertusa (Bates); (75)  Blepharoneura atomaria (Fabricius);
(76) Blepharoneura sp.; (77) Caenoriata pertinax (Bates); (78) Campiglossa sp.; (79) Carpomya tica Norrbom, (Norrbom 1997a, fig. 2);
(80) Ceratitis capitata (Wiedemann); (81) Ceratodacus longicornis Hendel, (Norrbom & Condon 1999, fig. 6.3A); (82) Chetostoma sp., and
(83) Cryptodacus obliquus Hendel. Figures 74–78, 80, 82, and 83 illustrated by L.H. Rodriguez and A.L. Norrbom.
928
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

39. Arista short to long plumose, rays at least half width of first flagellomere (Fig. 28); wing brown
on distal half or with six brown bands, including subcostal band (from pterostigma to apex of
cell cup), small band originating in middle of cell  r1 sometimes covering r-m and connected
posteriorly to discal band to form Y-shaped mark, and posterior apical band (crossing distal half
of cells r4+5 and m) (Fig. 6); scutum without microtrichia, yellow, with two to four pairs of brown
markings including presutural ovoid dark spots and postsutural stripes or spots; frons with two
frontal setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Molynocoelia Giglio-Tos, in part
– Arista pubescent or bare (Figs. 20, 33); wing pattern various, but lacking at least one of above
bands; scutal microtrichia and color various, but if mostly yellow usually without at least
presutural brown spots; frons usually with three or more frontal setae (two in some Cryptodacus spp.
and one Parastenopa sp., zero in Tomoplagia quadriseriata Hendel). . . . . . . . . . . . . . . . . . . . . 40

40. Scutum with medial white spot or stripe (Figs. 4, 5, 8). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41

– Scutum without medial white spot or stripe. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43

41. Vein CuA1 bordered with brown between bm-cu and discal band (Figs. 4, 83) and/or antenna
elongate, extended beyond ventral facial margin (Fig. 20); crossvein r-m more than 0.60 distance
from bm-cu to dm-cu; subapical band and posterior orbital seta present; face usually with brown
spot(s) or inverted U-shaped marking; aculeus tip sagittate and finely serrate (Fig. 161). . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cryptodacus Hendel, in part
– Vein CuA1 without brown border between bm-cu and discal band (rarely with short extension
from discal band in Rhagoletotrypeta) (Figs. 5, 8); antenna not extended beyond ventral facial
margin; crossvein r-m less than 0.60 distance from bm-cu to dm-cu, or if greater (Haywardina
cuculi), subapical band and posterior orbital seta absent (Fig. 5); face without brown markings;
aculeus tip neither sagittate nor serrate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42

42. First flagellomere with small dorsoapical point (as in Figs. 36, 40); scutum with white postsutural
sublateral stripe (Fig. 5); aculeus tip less than one-third aculeus length, entire or with minute
subapical notches (Fig. 162); medial surstylus long, prensisetae well beyond level of posterior
margin of epandrium (similar to Fig. 156); Ecuador to Argentina. . . . . . . . . . [Haywardina Aczél]
– First flagellomere usually rounded dorsoapically; scutum with or without white postsutural
sublateral stripe (Fig. 8); aculeus tip either more than one-half aculeus length or deeply trilobed
(Figs. 166, 167); medial surstylus short, prensisetae only slightly beyond level of posterior margin
of epandrium (Fig. 157). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhagoletotrypeta Aczél

43. Wing medially or basomedially with large, usually broad and ovoid brown area covering pterostigma,
crossvein bm-cu, basal half or more of cell dm, and usually crossvein r-m and cell br distal to
bm-cu (Figs. 121–123); distal to dm-cu, two apical bands present, anterior band bordering costa,
and posterior band more or less parallel to it crossing cell r4+5 . . . . . . . . . . . . . . . . . . . . . . . . . . 44
– Wing usually lacking one or both apical bands (Figs. 100–102, 117) and/or with either crossvein
bm-cu or r-m in hyaline area (Fig. 118, 139) or in different bands (Figs. 82, 152–153), sometimes
connected posteriorly or anteriorly, but separated by hyaline area at least in cell br and usually in
base of cell dm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45

44. Wing with crossvein dm-cu covered by same large medial brown area as crossvein r-m (Fig. 121),
or if these crossveins in separate brown bands, cell r1 with two basal marginal hyaline incisions
(Fig. 122) or r-m and dm-cu separated by hyaline area that does not extend anteriorly to costa
(Fig. 123); ocellar seta usually absent or small and weak, less than twice as long as ocellar tubercle,
much weaker than posterior orbital and postocellar setae (Fig. 33), if larger, scutum without obvious
vittae due to microtrichia; facial carina weak or absent. . . . . . . . . . . . Parastenopa Hendel, in part
– Wing with crossveins r-m and dm-cu in separate brown bands, separated by hyaline band extending
anteriorly to costa; cell r1 without other marginal hyaline marks except sometimes small apical
spot or extension from cell r2+3; ocellar seta and facial carina well developed (Fig. 36); scutum
with four vittae of denser microtrichia. . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhagoletis Loew, in part
TEPHRITIDAE (fruit flies, moscas de frutas) 68 929

45. Crossvein r-m at least two-thirds distance from bm-cu to dm-cu along vein M (Figs. 10, 153);
pterostigma, r-m, and dm-cu covered by same band, isolated from other markings except anteriorly,
or if such band incomplete or connected elsewhere (rarely in Tomoplagia), scutum yellow with
pair of oval brown sublateral spots near posterior margin (Fig. 10); body mostly yellow, usually
with sublateral brown spots on one or more abdominal tergites. . . . . . . . . . . . . . . . . . . . . . . . . 46
– Crossvein r-m usually less than two-thirds distance from bm-cu to dm-cu along vein  M, if
two-thirds or more, then pterostigma, r-m, and dm-cu not covered by same isolated band and
scutum not mostly yellow with brown oval sublateral spots near posterior margin; body color and
abdominal pattern various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47

46. Scutum with pair of oval sublateral brown spots near posterior margin (Fig. 10) (absent only in
Antillean species T. propleuralis Aczél), occasionally with additional brown markings; wing usually
with band between that covering dm-cu and apical band (band along costa); aculeus tip shape
various, but not broad and finely serrate; surstyli short. . . . . . . . . . Tomoplagia Coquillett, in part
– Scutum entirely yellow to orange; wing without band between that covering dm-cu and apical
band (Fig. 153); aculeus tip broad and finely serrate (Fig. 168); surstyli elongate. . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trypeta Meigen, in part

47. Gena with row of large setae on ventral margin (Fig. 19); face nearly flat, carina weak or absent;
wing with weak humeral and four strong narrow brown bands: from pterostigma to apex of cell
cup; from costa in middle of cell r1 crossing r-m; bordering dm-cu; and along costa on distal third
of wing (Fig. 82); aculeus tip flattened in sagittal plane, broader in lateral view than in ventral
view, dorsal and ventral margins serrate (Fig. 160). . . . . . . . . . . . . . . . . . . . Chetostoma Rondani
– Gena without row of large setae on ventral margin; facial carina various; wing without band from
costa in middle of cell r1 crossing r-m and otherwise differing from above; aculeus tip flattened
dorsoventrally, broader in ventral view than in lateral view, lateral margin serrate or entire. . . 48

48. Wing with crossvein r-m in hyaline band extending from costa to cell cu1 (Fig. 118); cell bc,
base of br, and most or all of pterostigma covered by brown area also extending from pterostigma
to posterior wing margin crossing bm-cu; distal half of wing with somewhat F-shaped pattern,
including band crossing cell  m; scutellum entirely brown, convex, shiny dorsally, without
microtrichia; male forefemur stout, with short, spinelike ventral setae (similar to Fig. 55). . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ostracocoelia Giglio-Tos
– Wing with crossvein r-m infuscated, or with totally different pattern; scutellum various, but not
convex and shiny, nonmicrotrichose; male forefemur not exceptionally stout, ventral setae sometimes
strong but usually not spinelike. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49

49. Scutum microtrichose except small paired bare spots at medial end of transverse suture and at
base of dorsocentral seta and sometimes three narrow stripes; facial carina strong, dorsal two-thirds
at most slightly concave in profile (Fig.  31); postocular setae slender, sometimes poorly
differentiated from occipital setulae; wing with narrow bands or series of spots, but without
complete bands from either crossvein h or pterostigma to apex of cell cup (Fig. 117); western
United States to central Mexico. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Oedicarena Loew, in part]
– Scutum entirely microtrichose or with more extensive bare areas; facial carina various, often
weaker; postocular setae strong; wing pattern various, if banded often with complete band from
either crossvein h or pterostigma to apex of cell cup (Figs. 138, 152) . . . . . . . . . . . . . . . . . . . . 50

50. Wing with three to six bands, including one from pterostigma to at least vein  CuA1, crossing
r-m, another covering dm-cu and extending to anterior margin, and usually subbasal band from
crossvein h to apex of cell cup (Figs. 138, 139), if latter absent or incomplete, postocular and pos-
tocellar setae yellow, contrasting with blackish frontal, orbital, and ocellar setae; first flagellomere
usually with small dorsoapical point (Fig. 36); facial carina strong; male surstyli elongate, lateral
surstylus usually much longer than medial surstylus (Fig. 156). . . . . . . . Rhagoletis Loew, in part
– Wing, if banded, without band from crossvein h to apex of cell cup (Fig. 152); postocular and
postocellar setae same color as or at most slightly paler than frontal, orbital, and ocellar setae
(Fig. 27); first flagellomere rounded dorsoapically; facial carina various; male surstyli short, or if
elongate, lateral surstylus only slightly longer than medial surstylus. . . . . . . . . . . . . . . . . . . . . 51
930
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

84 Cryptotreta 85 Dictyotrypeta

86 Dictyotrypeta 87 Dioxyna

88 Dracontomyia 89 Dracontomyia

90 Dyseuaresta 91 Dyseuaresta

92 Ensina 93 Epochrinopsis

Figs.  68.84–93. Wings (continued): dorsal view of (84)  Cryptotreta cislimitensis Steyskal, (Nearctic); (85)  Dictyotrypeta incisa (Wulp);
(86) Dictyotrypeta sp.; (87) Dioxyna peregrina (Loew); (88) Dracontomyia borrichia (Bush & Huettel), (Nearctic); (89) Dr. riveti Becker;
(90) Dyseuaresta sobrinata (Wulp); (91) Dy. trinotata Bates; (92) Ensina longiceps (Hendel); and (93) Epochrinopsis sp. Figures  84–93
illustrated by L.H. Rodriguez and A.L. Norrbom.
TEPHRITIDAE (fruit flies, moscas de frutas) 68 931

51. Frons bare medially (Fig. 27); setae yellow to moderate brown; wing not banded, with brown
or yellow areas usually broad and connected (Figs. 100–102); crossvein r-m 0.60–0.75 distance
from bm-cu to dm-cu along vein M; aculeus varied in width, tip finely serrate or nonserrate;
surstyli short; glans with spines on distal membranous part . . . . . Gymnocarena Hering, in part
– Frons with fine setulae medially; setae dark brown to black; wing with bands or spots, usually
isolated, and narrow except sometimes for apical spot (Fig. 152); crossvein r-m often less than
0.60 distance from bm-cu to dm-cu along vein M; aculeus broad, tip extensively finely serrate
(Fig. 168); surstyli elongate; glans without spines on membranous part. . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trypeta Meigen, in part

52. Anterior notopleural seta absent (Fig. 48); wing pattern reticulate or mostly dark with hyaline
spots (Fig. 107) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
– Anterior notopleural seta present, or if absent (one Cecidochares species), wing banded (similar
to Figs. 3, 130). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54

53. Proboscis capitate (Fig. 24); most setae yellow to pale brown; frons with one to two frontal
setae; scutum densely covered with white lanceolate setulae, especially anterior to transverse
suture; anepisternum with numerous white, lanceolate setulae on posterior half, more ventral
ones only slightly smaller and less dense than row on dorsal margin; all of scutellum and sub-
scutellum, most of mediotergite, and at least most of abdominal syntergite 1+2 microtrichose;
abdominal setulae often white and lanceolate. . . . . . . . Euaresta Loew, in part (E. toba group)
– Proboscis geniculate (similar to Figs. 21, 26); frons with three frontal setae; most setae black;
scutum with white lanceolate setulae small and sparse, anterior to transverse suture at most in
10 irregular rows; anepisternum with only setulae in dense row on dorsal margin white, lanceo-
late, more ventral setulae smaller, acuminate and much less conspicuous; margin of scutellum,
subscutellum, most of mediotergite, and abdominal tergites except base of syntergite 1+2 nonmi-
crotrichose and shining; abdominal setulae acuminate; Peru, Bolivia. . . . . . [Lamproxyna Hendel]

54. Frons narrow, width at level of anterior ocellus approximately equal to width of eye (Fig. 39);
postocular setae all white, lanceolate; wing usually mostly brown (mostly yellow in X. bipuncta
(Aczél)), with two large, similar sized, usually triangular, marginal hyaline marks in cell r1, usually
two (up to four) hyaline marks in cell m, and without marginal hyaline marks in cell r4+5 and usually
cell r2+3 (Fig. 11). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xanthaciura Hendel
– Frons usually broader than eye, if not broader, cell r1 with at most one large, triangular, marginal
hyaline mark (Figs. 3, 67); postocular setae and wing pattern various. . . . . . . . . . . . . . . . . . . . 55

55. Wing predominantly dark, including alula and usually most or all of basal cells (at least most of
basal half of br), often with numerous small white or silvery dots medially, and with apical or
anteroapical hyaline band on margin or separated from margin by no more than width of band
(Figs. 99, 129, 132); frons at least anteromedially with white or yellow, often lanceolate, setulae
(Fig. 25). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
– Wing pattern various, but if predominantly dark, except for small dots, and with apical hyaline
band, then alula and basal fourth of cell br mostly yellowish or hyaline and frons bare anteromedi-
ally. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58

56. Posterior distal corner of cell dm acute, usually strongly so (Fig. 129); crossvein r-m at least 0.60
distance from bm-cu to dm-cu; anepisternal setulae all white, lanceolate, more or less evenly
distributed on posterior half; cell r1 often with one to two large marginal hyaline triangles; cells cu1
and m, exclusive of apical hyaline band, with one to three large marginal hyaline triangles or elon-
gate marks; middle of wing with at most few hyaline or white spots; face and parafacial without
brown markings, male face often extended ventrally . . . . . . . . . . . . . . . . Polymorphomyia Snow
– Posterior distal corner of cell dm not acute (Figs. 99, 132); position of crossvein r-m various, but
often less than 0.60 distance from bm-cu to dm-cu; anepisternal setulae acuminate or lanceolate,
often in denser row dorsally; cell r1 without large marginal hyaline triangles, or rarely (one species
of Eutreta) with three; cells cu1 and m, exclusive of apical hyaline band, usually with no more
than one large, marginal hyaline triangle or elongate mark; middle of wing usually with extensive
pattern of dots (sometimes faint, due only to microtrichial color); parafacial often with brown
spot at level of antennal base (Fig. 25); face often with brown markings, male face not extended
ventrally. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
932
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

57. Scutellum with only one (basal) pair of marginal setae; hyaline apical or anteroapical band of
wing sometimes partially narrowly separated from margin (Fig. 132); middle of cell cu1 usually with
large, oblique hyaline mark (if lacking, apical hyaline band submarginal); anepisternal setulae
all acuminate, more or less evenly distributed; face without brown markings; Bolivia, Paraguay,
Argentina, southern Brazil. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Pseudeutreta Hendel]
– Scutellum with two pairs of marginal setae; hyaline apical or anteroapical band of wing on margin
(Fig. 99); middle of cell cu1 without large, oblique hyaline mark; some or all anepisternal setulae
lanceolate, dorsally often in denser row; face often with brown markings (Fig. 25) . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eutreta Loew

58. Scutellum with only one (basal) marginal seta (as in Fig. 48). . . . . . . . . . . . . . . . . . . . . . . . . . . 59
– Scutellum with two marginal setae (Figs. 49, 50). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69

59. Proboscis long geniculate (Fig. 21), labella length at least 0.6 times head height.. . . . . . . . . . . 60
– Proboscis capitate or short geniculate; labella length less than half head height.. . . . . . . . . . . . 61

60. Scutellum largely nonmicrotrichose and shiny dorsally; frons with three to four frontal and
one orbital seta; head 1.1 times as high as long; wing with large brown and hyaline areas, including
solid brown marking extending from pterostigma almost to apex of vein CuA1 and covering most of
cell dm, and uninterrupted hyaline area from cell r1 to cell m (Fig. 103); Peru . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Hetschkomyia Hendel]
– Scutellum entirely densely microtrichose and matte dorsally; frons with two  frontal and
two orbital setae (Fig. 21); head at least as long as high; wing pattern reticulate, with smaller,
more discontinuous brown and hyaline areas (Fig. 87). . . . . . . . . . . . . . . . . Dioxyna Frey, in part

61. Vein M with distal section strongly arched anteriorly, cell m 2.5 times as wide as cells r2+3 and r4+5
together; wing mostly dark brown with four small hyaline triangles on anterior margin (two in cell c
and two in r1) and four large, elongate marks on posterior margin (two in cu1 and two in m); body
blackish, scutum and abdomen subshining, sparsely microtrichose; Argentina. . [Lilloaciura Aczél]
– Vein M with distal section at most slightly arched anteriorly, cell m less than 1.5 times as wide as
cells r2+3 and r4+5 together; other characters various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62

62. Wing banded (Fig.  140), with oblique hyaline band between r-m and dm-cu from costa to
vein  CuA1, and another from apex of CuA1 across base of cell  m and apical half of cell  r4+5;
scutum dark brown with pair of broad, gray dorsocentral vittae; abdomen largely shining, nonmi-
crotrichose; Peru, Bolivia, Argentina. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Rhithrum Hendel]
– Wing pattern various, if banded (Fig. 133), hyaline band between r-m and dm-cu interrupted or
perpendicular, and without oblique band from apex of CuA1 to apex of cell r4+5; scutum color and
abdomen microtrichia various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63

63. Male forefemur ventrally with dense irregular rows of short, often stout setulae anterior to row
of ventral setae (Fig. 54), female often with fewer such setulae; cell r4+5 usually with medial bulla
in species south of United States; abdominal tergites with only acuminate setulae, often partially
shiny, nonmicrotrichose; male medial and lateral surstyli elongate (Fig. 154). . . . . . . . . . . . . . 64
– Male and female forefemur ventrally at most with few setulae anterior to row of ventral setae;
cell r4+5 without medial bulla except in some Dyseuaresta and Lamproxynella; abdominal tergites
with at least some setulae lanceolate, except in some Pseudoedaspis, and usually entirely
microtrichose; male medial and lateral surstyli elongate or short. . . . . . . . . . . . . . . . . . . . . . . . 65

64. Wing pattern radiate, apex with rays extending from central dark part of pattern to apices of
veins R4+5 and M with large hyaline apical spot between them (similar to Fig. 97); cell r4+5 with
strong bulla; western North America, south to northern Mexico. . . . [Valentibulla Foote & Blanc]
– Wing banded, spotted or more extensively dark, but not radiate, apex mostly or entirely dark or
with marginal spots or band (Fig. 66); cell r4+5 with or without bulla. . . . . . . . . . Aciurina Curran
TEPHRITIDAE (fruit flies, moscas de frutas) 68 933

94 Euaresta 95 Euaresta

96 Euarestoides 97 Euarestopsis

98 Euphranta 99 Eutreta

100 Gymnocarena 101 Gymnocarena

102 Gymnocarena 103 Hetschkomyia

Figs.  68.94–103. Wings (continued): dorsal view of (94)  Euaresta reticulata (Hendel); (95)  Euaresta stigmatica Coquillett, (Nearctic);
(96)  Euarestoides acutangulus (Thomson); (97)  Euarestopsis  sp; (98)  Euphranta mexicana Norrbom, (Nearctic); (99)  Eutreta  sp;
(100) Gymnocarena brunnea Norrbom, (Norrbom, 2006a, fig. 2); (101) G. mexicana (Aczél), (Nearctic); (102) G. macalpinei Norrbom,
(Nearctic), (Norrbom, 2006a, fig.  4); and (103)  Hetschkomyia maculipennis Hendel. Figures 94–99, 101, and 103 illustrated by
L.H. Rodriguez and A.L. Norrbom.
934
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

65. Frons with three frontal setae, or if (rarely) with two, wing pattern as in Fig.  149; male glans
small and mostly membranous except for large, hooklike basolateral sclerite (Fig. 158); wing pat-
tern usually stellate (Fig. 151) but sometimes more extensively dark or rarely banded (Figs. 149,
150) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trupanea Schrank
– Frons with two frontal setae; male glans without large, hooklike basolateral sclerite; wing pattern
not stellate except in some Plaumannimyia and Dyseuaresta bilineata (Foote). . . . . . . . . . . . . 66

66. Hind femur with at least one small anterodorsal preapical seta and usually with one postero-
dorsal preapical seta (Figs. 56, 58); wing pattern radiate, reticulate or banded, with or without rays
extending to apices of veins R4+5 and M; male surstyli long and posteriorly curved, prensisetae
subequal. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
– Hind femur without anterodorsal or posterodorsal preapical setae; wing pattern radiate or stellate,
with narrow rays extending to apices of veins R4+5 and M (Fig. 125); surstyli usually short, if long
and curved, medial prensisetae greatly enlarged. . . . . . . . . . . . . . Plaumannimyia Hering, in part

67. Wing banded (Fig. 133), apical margin of cells r2+3 and r4+5 entirely infuscated; dorsum of abdomen
usually partially nonmicrotrichose and shiny laterally and apically; Chile, Argentina. . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Pseudoedaspis Hendel]
– Wing pattern radiate, stellate or reticulate (Figs. 90, 91, 108), cell r2+3 with at least one marginal
hyaline mark, and cell  r4+5 usually with marginal or submarginal hyaline spot (varied in Lam-
proxynella); abdomen usually entirely microtrichose, male tergite 5 and female tergite 6 some-
times partly nonmicrotrichose and shiny. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68

68. Wing pattern radiate (Fig. 90) or stellate (similar to Fig. 125), extensively dark with apical rays,
or rarely reticulate with large dark marks on anterior margin (Fig. 91); cell r4+5 with apical hyaline
spot broadly touching margin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dyseuaresta Hendel
– Wing pattern reticulate or almost banded (Fig. 108); cell r4+5 sometimes without apical hyaline
spot or with spot narrowly or not touching margin; Venezuela to Chile and Argentina, southern
Brazil . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Lamproxynella Hering]

69. First flagellomere with distinct dorsoapical point, in female long and broad in lateral view, almost
reaching ventral facial margin (Fig. 13); scutellum convex, with distinct brown and yellow areas,
and with white, lanceolate setulae only on margins; frons broad, approximately 2 times width of
eye in dorsal view; wing pattern reticulate or with irregularly margined bands (Fig. 64); aculeus
tip sagittate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acidogona Loew
– First flagellomere shorter or narrower, not reaching ventral facial margin, and usually without
distinct dorsoapical point; scutellum various, but if convex, without distinct brown and yellow
areas; frons and wing pattern various; aculeus tip not sagittate.. . . . . . . . . . . . . . . . . . . . . . . . . 70

70. Scutellum convex in posterior or lateral view (Fig. 51), often shiny nonmicrotrichose medially
or entirely, without setulae except sometimes on margins (usually only basal to basal seta); whitish,
lanceolate scutal setulae often in clusters posteriorly on scutum and/or on scutellum (Fig.  3);
scutum often with lateral or sublateral shiny nonmicrotrichose area; notopleuron and/or scutum
bordering notopleuron or scutellum often with velvetlike microtrichia more dense than elsewhere
on scutum; wing with brown bands (Figs. 89, 111, 130, 141). . . . . . . . . . . . . . . . . . . . . . . . . . . 71
– Scutellum flattened or only slightly convex, microtrichose and often setulose medially; scutal
setulae not in clusters; scutal microtrichosity various, but notopleuron and scutum usually not
with lateral or sublateral nomicrotrichose areas nor velvety, more densely microtrichose areas;
wing pattern various, sometimes banded, but often reticulate or mostly dark with hyaline spots or
incisions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
TEPHRITIDAE (fruit flies, moscas de frutas) 68 935

104 Hexachaeta 105 Homoeothrix

106 Ischyropteron 107 Lamproxyna

108 Lamproxynella 109 Lewinsohnia

110 Neomyopites 111 Neorhagoletis

112 Neortalotrypeta 113 Neosphaeniscus

Figs. 68.104–113. Wings (continued): dorsal view of (104)  Hexachaeta valida Lima; (105)  Homoeothrix lindigi (Hendel); (106)  Ischy-
ropteron nigricaudatum Bigot, (redrawn from R.H. Foote, 1979b, fig. 1); (107) Lamproxyna nitidula Hendel; (108) Lamproxynella impluviata
(Blanchard); (109) Lewinsohnia magna Norrbom & Prado; (110) Neomyopites tresmilius (Steyskal); (111) Neorhagoletis latifrons Hendel;
(112) Neortalotrypeta bicolor Norrbom, (Korneyev & Norrbom, 2006, fig. 52); (113) Neosphaeniscus m-nigrum (Hendel), (Norrbom et al.,
2010, fig. 48). Figures 104–105 and 107–111 illustrated by L.H. Rodriguez and A.L. Norrbom.
936
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

114 Neotaracia 115 Neotephritis

116 Neotephritis 117 Oedicarena

118 Ostracocoelia 119 Paracantha

120 Paracantha 121 Parastenopa

122 Parastenopa 123 Parastenopa

Figs. 68.114–123. Wings (continued): dorsal view of (114) Neotaracia imox (Bates); (115) Neotephritis cinerea (Blanchard); (116) Neotephritis
rava Foote (Nearctic); (117)  Oedicarena nigra Hernández-Ortiz (Nearctic); (118)  Ostracocoelia mirabilis Giglio-Tos; (119)  Paracantha
ruficallosa Hering; (120) Paracantha trinotata (Foote); (121) Parastenopa elegans (Blanchard); (122) Parastenopa sp; and (123) Paras-
tenopa sp. Figures 114–123 illustrated by L.H. Rodriguez and A.L. Norrbom.
TEPHRITIDAE (fruit flies, moscas de frutas) 68 937

71. Crossvein r-m distinctly distal to midlength of cell dm, in middle of same oblique band covering
pterostigma and crossvein dm-cu (Figs. 3, 124, 130) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72
– Crossvein r-m usually proximal to or near midlength of cell dm, in hyaline area or covered by
different band than that covering crossvein dm-cu (Figs. 89, 111, 141), or if more distal, at proximal
margin of transverse band covering dm-cu but not pterostigma (Fig. 88). . . . . . . . . . . . . . . . . . 74

72. Posterior orbital seta small or absent, if present less than half as long as anterior orbital seta; wing
without band across middle of cell cu1 (Fig. 130); scutum with presutural and postsutural lateral
shiny nonmicrotrichose areas; anepisternum often with shiny nonmicrotrichose area. . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Procecidochares Hendel
– Posterior orbital seta present and usually more than half as long as anterior orbital seta; wing
usually with band across middle of cell cu1 (Figs. 3, 124); scutum with or without presutural and
postsutural shiny nonmicrotrichose areas; anepisternum usually entirely microtrichose (partially
bare in a Cecidochares species from Argentina). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73

73. Scutellum with five marginal clusters of setulae, including one between basal and apical setae and
unpaired cluster between apical setae; lateral vertical seta white, lanceolate, less than half as long
as blackish medial vertical seta; dark basal area of wing broadly connected in cells c, bm, and cup
to band across crossvein bm-cu and vein CuA2 (Fig. 124); Ecuador, Peru, Bolivia, over 2000 m.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phacelochaeta Norrbom
– Scutellum without setulae or with setulae present only proximal to basal seta; lateral vertical seta
more than half as long as medial vertical seta and similar to it in shape and color; dark basal area
of wing usually distinctly separated except sometimes posteriorly from band across crossvein
bm-cu and vein CuA2 (Fig. 3), rarely with narrow connections in cells br or cup . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cecidochares Bezzi, in part

74. Cells r1 and r2+3 crossed by two hyaline bands (Fig. 141); costa with several enlarged setulae at
humeral break; scutellum entirely microtrichose, although less densely so than scutum; Nearctic,
south to central Mexico. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Stenopa Loew]
– Cells r1 and r2+3 crossed by only one hyaline band (Figs. 88, 89, 111); costa with setulae at humeral
break not enlarged; scutellum shiny, nonmicrotrichose medially . . . . . . . . . . . . . . . . . . . . . . . . 75

75. Lateral vertical seta dark brown, same color as and two-thirds or more length of medial vertical
seta; scutum with four clusters of white lanceolate setulae near posterior margin (two anterior to
corners of scutellum and two more medially), and with postsutural lateral shiny nonmicrotrichose
area, microtrichia uniformly colored; subscutellum with ventral half shiny, nonmicrotrichose; ab-
dominal tergal setulae fine, acuminate and brown except on lateral margins white and strongly
lanceolate; crossvein dm-cu less than two-thirds width of radial cells measured directly anterior
to it (Fig. 111); Bolivia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Neorhagoletis Hendel]
– Lateral vertical seta at most half length of medial vertical seta, often paler brown or yellow;
scutum with two clusters of white lanceolate setulae near posterior margin (anterior to corners of
scutellum), entirely microtrichose, or if nonmicrotrichose posterolaterally, with band of distinctly
darker microtrichia along posterior margin; subscutellum entirely densely microtrichose; abdomi-
nal tergal setulae various, but lateral setulae not more lanceolate than others; crossvein dm-cu at
least 0.85 times as long as width of radial cells measured directly anterior to it (Fig. 89), except
in D. borrichia (Fig. 88); southern United States, South America . . . . . [Dracontomyia Becker]

76. Parafacial with brown spot near level of antennal base (Figs. 32, 35); cell r4+5 often with bulla. 77
– Parafacial without brown spot near level of antennal base; cell r4+5 usually without bulla. . . . . 81
938
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

77. Face microtrichose, matte, with or without pair of brown spots; lunule without brown marks; eye
often with one or three narrow stripes (often faded or absent after death) . . . . . . . . . . . . . . . . . 78
– Face nonmicrotrichose, shiny yellow with one to four dark brown spots (Figs.  32, 35); lunule
often with dark brown mark or spots; eye green with two rows of large irregular red spots (often
faded or absent after death) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80

78. Frons nonsetulose medially; wing with isolated brown spots on apices of veins  R4+5 and M
(Fig.  84); at least apical abdominal tergite (male fifth, female sixth) largely shiny, nonmi-
crotrichose; Mexico (Baja California). . . . . . . . . . . . . . . . . . . . . . . . [Cryptotreta Blanc & Foote]
– Frons setulose medially (Fig. 25); wing without isolated brown spots on apices of veins R4+5 and
M (Figs. 85, 86); abdominal tergites entirely matte, microtrichose. . . . . . . . . . . . . . . . . . . . . . . 79

79. Face with pair of dark brown spots (Fig. 25); cell r4+5 without bulla; body largely tan or brown
microtrichose. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eutreta Loew, in part
– Face usually without pair of dark brown spots, if present, cell r4+5 with bulla and/or body predomi-
nantly grayish microtrichose, spotted with tan or brown. . . . . . . . . . . . . . . . Dictyotrypeta Hendel

80. Scutellum without erect white lanceolate setae; frons with two orbital setae, anterior seta darker
than posterior seta; face with two or four dark brown spots, but without medial spot near mid-
height (Fig. 35); wing reticulate, sometimes with broad paler or hyaline area on posterior margin,
without bullae (Fig. 137); Colombia to Chile, southern Brazil, Argentina. . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Rachiptera Bigot]
– Scutellum with three pairs of erect white lanceolate setae on disc (Fig.  50); frons with two to
three orbital setae, all white; face with one to four dark brown spots, including medial spot near
midheight (Fig.  32); wing usually radiate, cells  r2+3, r4+5, and dm with orange or brown bullae
(sometimes faint) (Figs. 119, 120). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paracantha Coquillett

81. Wing entirely brown, completely without hyaline marks; Brazil . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Cipomyia Norrbom & Prado]
– Wing not entirely brown.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82

82. Wing broad, oval, and medially with numerous small hyaline or pale spots or streaks, not banded
except sometimes distally (Figs. 67, 68, 73); vein R4+5 densely setulose dorsally to level of apex
of R2+3 (as in Fig. 63).. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83
– Wing usually without numerous small hyaline or pale spots medially, if present (some Epochrinopsis)
wing more or less banded (Fig. 93); vein R4+5 setulose or bare dorsally.. . . . . . . . . . . . . . . . . . . 86

83. Wing with uninterrupted hyaline apical band from middle of cell  r2+3 into cell  m (similar to
Fig. 99); Brazil. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Eutretopsis Norrbom & Prado]
– Wing without uninterrupted hyaline apical band (Figs. 67, 68, 73). . . . . . . . . . . . . . . . . . . . . . . 84

84. Wing with subapical transverse bands, and anteriorly with large dark brown area in cell  c,
pterostigma, and cells r1 and r2+3, often narrowly divided by yellow into three to eight large spots
(Fig. 67). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acrotaenia Loew
– Wing without subapical transverse bands (Figs.  68, 73); infuscated area in cells  c and r1 and
pterostigma sometimes with minute hyaline or pale brown spots, but not divided by narrow yel-
low areas. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85

85. Wing distally with marginal hyaline marks in cells r1, r2+3, r4+5, and m large and wedge-shaped,
usually longer than wide (Fig. 68); medial wing spots mostly minute; junction of crossvein dm-cu
and vein M surrounded by bulla; Venezuela . . . . . . . . . . . . . . . . . . . . . [Acrotaeniacantha Hering]
– Wing distally with marginal hyaline marks in cells r1, r2+3, r4+5, and m shallow, broader than long
(Fig. 73); medial wing spots larger, orange; without bulla . . . . . . . . . . . Baryplegma Wulp, in part
TEPHRITIDAE (fruit flies, moscas de frutas) 68 939

124 Phacelochaeta 125 Plaumannimyia

126 Plaumannimyia 127 Plaumannimyia

128 Polionota 129 Polymorphomyia

130 Procecidochares 131 Procecidocharoides

132 Pseudeutreta 133 Pseudoedaspis

Figs.  68.124–133. Wings (continued): dorsal view of (124)  Phacelochaeta quinquefasciata (Hendel), (Norrbom et al., 2010, fig. 55);
(125) Plaumannimyia eugenia (Wulp); (126) Pl. flava (Adams); (127) Pl. pallens Hering; (128) Polionota kohnae Norrbom; (129) Polymor-
phomyia pilosula (Wulp); (130) Procecidochares alani Steyskal; (131) Procecidocharoides pullatus Foote, (Nearctic); (132) Pseudeutreta
anteapicalis Hendel; and (133) Pseudoedaspis decorata (Blanchard). Figures 125–133 illustrated by L.H. Rodriguez and A.L. Norrbom.
940
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

134 Pseudophorellia 135 Pseudopolionota

136 Pyrgotoides 137 Rachiptera

138 Rhagoletis 139 Rhagoletis

140 Rhithrum 141 Stenopa

142 Tachinisca 143 Tephritis

Figs.  68.134–143. Wings (continued): dorsal view of (134)  Pseudophorellia hansoni Norrbom, (Norrbom, 2006b, fig.  82);
(135) Pseudopolionota radians Lima; (136) Pyrgotoides peruvianus (Korytkowski); (137) Rachiptera luctulenta (Hendel); (138) Rhagoletis
nova (Schiner); (139)  Rhagoletis striatella Wulp, (Nearctic); (140)  Rhithrum vittatum Aczél; (141)  Stenopa mexicana Norrbom;
(142) Tachinisca cyaneiventris Kertész; and (143) Tephritis sp. Figures 134–143 illustrated by L.H. Rodriguez and A.L. Norrbom.
TEPHRITIDAE (fruit flies, moscas de frutas) 68 941

144 Tephritis 145 Tetreuaresta

146 Tetreuaresta 147 Tetreuaresta

148 Toxotrypana 149 Trupanea

150 Trupanea 151 Trupanea

152 Trypeta 153 Trypeta

Figs. 68.144–153. Wings (concluded): dorsal view of (144) Tephritis sp. (145) Tetreuaresta bartica Bates; (146) Te. obscuriventris (Loew);
(147)  Te. heringi Norrbom; (148)  Toxotrypana curvicauda Gerstaecker; (149)  Trupanea bullocki Malloch; (150)  Tru. simulata Malloch;
(151) Tru. vicina (Wulp); (152) Trypeta costaricana Han & Norrbom; and (153) Try. inclinata Han & Norrbom, (Han & Norrbom, 2005,
fig. 6T). Figures 144–152 illustrated by L.H. Rodriguez and A.L. Norrbom.
942
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

st 5 st 6 st 7 syntgst 7+8
epand

cerc

epand

154 Aciurina tg 5

155 Rhagoletis

epand
ej apod
cerc

spm sac
phapod
hypd
spm dt
ph gd

epand
ph

157 Rhagoletotrypeta
cerc

prens

sur

gls

156 Rhagoletis

158 Trupanea

Figs. 68.154–158. Male terminalia: left lateral view of epandrium and surstyli of (154) Aciurina semilucida (Bates), (Nearctic, Norrbom,
1989, fig. 1, as A. bigeloviae (Cockerell)); ventral view of abdomen of (155) Rhagoletis pomonella (Walsh), (Nearctic, MND, fig. 66.38);
lateral view of terminalia of (156) Rhagoletis pomonella (Walsh), (Nearctic, MND, fig. 66.41); left lateral view of epandrium and surstyli of
(157) Rhagoletotrypeta morgantei Norrbom, (Norrbom, 1994b, fig. 9B); glans of (158) Trupanea nigricornis (Coquillett), (Nearctic). Figure
158, illustrated by A.L. Norrbom.
Abbreviations: cerc, cercus; ej apod, ejaculatry apodeme; epand, epandrium; gls,  glans; hypd, hypandrium; ph,  phallus; phapod, phallapodeme;
ph  gd,  phallic guide; prens, prensiseta; spm  dt, sperm duct; spm  sac, sperm sac; st,  sternite; sur,  surstylus; syntgst, syntergosternite;
tg, tergite.
TEPHRITIDAE (fruit flies, moscas de frutas) 68 943

86. Vein R4+5 evenly and usually densely setulose dorsally to level of apex of vein  R2+3 (Fig.  63);
lateral vertical seta shorter than medial vertical seta but similar in shape and color (Fig. 27), or
if slightly paler (some Baryplegma), wing broad, less than 2 times as long as wide; cell cup with
apical lobe at least one-third (usually more than half) as long as width of cell (Fig. 63); frons non-
setulose medially (except Epochrinopsis) and with three or more frontal setae (Fig. 27); proboscis
not geniculate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
– Vein R4+5 dorsally bare, with few scattered setulae, or if setulose beyond level of dm-cu (some
species of Acinia, Neotephritis, and Lewinsohnia), lateral vertical seta distinctly paler and more
lanceolate than medial vertical seta (Fig. 30); cell cup with apical lobe at most half as long as
width of cell (Fig. 60), often less than one-third as long; frons often setulose medially and with
two to four frontal setae (Figs. 21, 30); proboscis sometimes geniculate. . . . . . . . . . . . . . . . . . 93

87. Wing usually with three to five oblique, parallel, usually mostly yellow bands (Fig. 10); crossvein
r-m close to dm-cu and covered by same band; body mostly yellow, scutum usually with pair of
oval sublateral brown spots near posterior margin (Fig. 10), occasionally with larger or additional
brown markings; abdomen yellow, usually one or more tergites with pair of small, discrete
sublateral brown spots. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tomoplagia Coquillett, in part
– Wing not banded (Figs. 74, 101, 145) or if so bands nearly perpendicular to long axis of wing and/
or present only distal to crossvein dm-cu (Figs. 93, 114, 135); body color various, but if scutum
mostly yellow, without brown posterolateral spots; abdomen without sublateral spots . . . . . . . 88

88. Frons usually with fine setulae medially; wing usually more or less banded, with numerous small
hyaline or white spots within bands and/or paler areas (Fig. 93), occasionally with reduced pat-
tern of small brown markings; crossvein r-m near or proximal to midlength of cell dm, 0.33–0.60
distance from bm-cu to dm-cu along vein M; gena and postgena well developed, ventral margin
of head usually strongly slanted upward towards face (Fig. 23); major setae usually dark brown to
black; Bolivia, Ecuador, Venezuela (high elevations). . . . . . . . . . . . . . . . [Epochrinopsis Hering]
– Frons bare medially (Fig. 27); wing pattern various, but not banded with numerous small hyaline
spots; position of crossvein r-m and head shape various; setae yellow to moderate brown . . . . 89

89. Wing distally with two to three oblique or transverse hyaline bands arising on posterior margin of
cells m or r4+5, at least most distal band reaching or almost reaching R4+5, but none extending to
anterior wing margin, most distal band proximal to or including apex of vein M (Figs. 114, 135);
cell r2+3 with zero to one hyaline marginal marks, if present, at apex of vein R2+3 (Fig. 114). . . 90
– Wing distally usually without oblique or transverse hyaline bands arising on posterior margin of
cells m or r4+5 (Figs. 74, 100, 101), if bands present, not or only slightly extended into cell r4+5
(Figs. 102, 145), or most distal band reaching margin distal to apex of vein M (Tetreuaresta
heringi Norrbom, Fig. 147); cell r2+3 with zero to two hyaline marginal marks . . . . . . . . . . . . . 91

90. Wing with wedge-shaped marginal hyaline mark on apex of vein R2+3 and with at most one wedge-
shaped marginal hyaline mark basally in cell r1 (Fig. 114); cell r1 with small spotlike to elongate
nonmarginal hyaline mark; cell c with two hyaline spots. . . . . . . . . . . . Neotaracia Foote, in part
– Wing without wedge-shaped marginal hyaline mark on apex of vein  R2+3 and with two broad
wedge-shaped marginal hyaline marks in cell r1 (Fig. 135); cell r1 without nonmarginal hyaline
mark; cell c with single elongate hyaline mark posteriorly; Brazil. . . . . . [Pseudopolionota Lima]

91. Wing distal to crossvein dm-cu without dark rays, or if present in cells r2+3 or m, small and
oriented perpendicular to long axis of wing (Figs. 100–102); either cell r1 without marginal
hyaline markings or posterobasal fourth or more of wing hyaline or much paler than other infus-
cated areas; glans with spines on distal membranous part. . . . . . . . . Gymnocarena Hering, in part
– Wing pattern usually radiate distally, with dark markings radiating from middle of apical part of
cell r4+5, two marginal or submarginal hyaline wedge-shaped marks or spots in cell r2+3 and one
medial mark or spot in cell r4+5 (Figs. 74, 146), if not radiate (some Tetreuaresta) cell r1 with two or
more wedge-shaped marginal hyaline marks and anal lobe mostly brown (Figs. 145, 147); glans
without spines on distal membranous part. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92
944
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

161 Cryptodacus

160 Chetostoma

162 Haywardina
159 Anastrepha
ac gl
spmth
ev mem
acul

sg 7
v rep ovscp st 8 cloac op

164 Rhagoletis

ovscp
165 Rhagoletis
163 Rhagoletis

166 Rhagoletotrypeta 167 Rhagoletotrypeta

168 Trypeta

169 Rhagoletis

Figs.  68.159–169. Female terminalia and larva: dorsal view of base of eversible membrane of (159)  Anastrepha grandicula Norrbom,
(Norrbom, 1991, fig.  10D); lateral view of apex of aculeus of (160)  Chetostoma californicum Blanc, (Nearctic, Han, 1999, fig.  11.10J);
ventral view of apex of aculeus of (161) Cryptodacus lopezi Norrbom, (Norrbom, 1994b, fig. 5D); (162) Haywardina bimaculata Norrbom,
(Norrbom, 1994b, fig. 8H); lateral view of abdomen of (163) Rhagoletis pomonella (Walsh), (Nearctic, MND, fig. 66.42); lateral view
of extended ovipositor of (164) Rhagoletis pomonella (Walsh), (Nearctic, MND, fig. 66.43); ventral view of aculeus of (165) Rhagoletis
pomonella (Walsh), (Nearctic, MND, fig. 66.44); (166) Rhagoletotrypeta intermedia Norrbom, (Norrbom, 1994b, fig. 10B); (167) Rhagoletotrypeta
parallela Norrbom, (Norrbom, 1994b, fig. 10C); and (168) Trypeta rufata (Wulp); posterodorsal view of posterior end of third instar larva of
(169) Rhagoletis sp. (MND, fig. 5.105). Figure 168 illustrated by A.L. Norrbom.
Abbreviations: ac gl, accessory gland; acul, aculeus; cloac op, cloacal opening; ev mem, eversible membrane; ovscp, oviscape; sg, segment;
spmth, spermathecae; st, sternite; v rep, ventral receptacle.
TEPHRITIDAE (fruit flies, moscas de frutas) 68 945

92. Wing at least distally with narrow darker brown margins on infuscated areas (Fig. 74) and with
broad distal rays; cell  r 4+5 often with argent (ovoid area of silvery microtrichia on dark
background) proximal to level of crossvein dm-cu. . . . . . . . . . . . . . . . . Baryplegma Wulp, in part
– Wing with infuscated areas more or less evenly brown (Figs. 145–147) or at most with yellowish
areas along vein M medially, occasionally not radiate distally; cell r4+5 without argent. . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetreuaresta Hendel

93. Proboscis geniculate; head often longer than high in lateral view (Figs. 16, 21) . . . . . . . . . . . . 94
– Proboscis capitate; head higher than long in lateral view (Figs. 24, 30). . . . . . . . . . . . . . . . . . . 97

94. Frons with one acuminate orbital seta, posterior seta absent (Fig. 22); lateral vertical seta brown,
acuminate, more than half as long as medial vertical seta; wing mostly hyaline or diffusely infus-
cated with hyaline posterior margin (Fig. 92); aculeus usually with indentation or ridge delimiting
tip; Old World, South America (Venezuela to Bolivia and Chile). . . [Ensina Robineau-Desvoidy]
– Frons with two orbital setae, posterior seta white, lanceolate (Fig. 21); lateral vertical seta white,
lanceolate; wing pattern reticulate (Figs. 78, 87); aculeus without indentation delimiting tip.. . 95

95. Wing with most dark markings restricted to two to three bands with hyaline spots within them
and few or no dark marks between them; frons usually with only one of three to four frontal
setae brown; posterior notopleural seta white, lanceolate (as in Fig. 49); abdomen evenly gray
microtrichose; Palearctic and Nearctic (south to northern Mexico) Regions . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Oxyna Robineau-Desvoidy]
– Wing more evenly reticulate (Figs. 78, 87); frons with two or occasionally three brown frontal setae
(Figs. 16, 21); posterior notopleural seta usually brown, acuminate; abdomen gray microtrichose,
usually with two submedial brown stripes or rows of spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96

96. Head usually no longer than high (Fig. 16); labella length less than 0.75 times head height; apical
scutellar seta usually longer than scutellum; phallus with small spines proximal to glans. . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Campiglossa Rondani, in part
Head longer than high (Fig. 21); labella length at least 0.75 times head height; apical scutellar seta
usually shorter than scutellum; phallus without small spines proximal to glans. . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dioxyna Frey, in part

97. Hind femur without anterodorsal or posterodorsal preapical setae; wing pattern often stellate or
paler on basal half, but sometimes reticulate (Figs. 125–127). . . . Plaumannimyia Hering, in part
– Hind femur with at least one small anterodorsal preapical seta and usually with one posterodorsal
preapical seta (Figs. 56, 58); wing pattern various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98

98. Frons with two frontal setae (Figs. 24, 37) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99

– Frons with three to four frontal setae (Fig. 30) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100

99. Frons and face, in lateral view, meeting in gradual curve or at slight angle (Fig. 24); wing pattern
reticulate to radiate (Figs. 94–95); setae yellow to orange–brown; male forefemur much stouter
than that of female, three or more times as broad as tibia; epandrium and surstyli, in posterior
view, broader than high, posterior surface flattened or concave, with striations. . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Euaresta Loew, in part
– Frons and face meeting at distinct angle (Fig. 37); wing pattern reticulate to stellate (Figs. 143–144);
setae usually dark brown; male forefemur similar to that of female, two or less times as broad as
tibia; epandrium and surstyli, in posterior view, higher than broad, convex, without striations .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tephritis Latreille, in part
946
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

100. Wing with subapical dark brown stellate mark broadly reaching costa in cell r1 and with narrow rays
to apices of veins R4+5 and M and in cell m, basal half of wing predominantly hyaline with yellow
to pale brown or gray reticulate markings usually distinctly paler than stellate mark (Fig. 96 or
similar to Fig. 126). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Euarestoides Benjamin
– Wing with margin of cell r4+5 brown (Figs. 65, 105, 113) or without distinct subapical stellate mark
(Figs. 109, 115, 143), if narrow rays present to apices of veins R4+5 and M and in cell m, originating
in middle of cell r4+5 from much more extensive brown area (Fig. 109); infuscated areas usually
more or less uniform in color or at least not paler on basal half of wing. . . . . . . . . . . . . . . . . . 101

101. Wing pattern reticulate, infuscated areas orange and brown, orange areas usually at least partially
with narrow brown margins (Fig. 65); scutum without brown markings surrounding acrostichal
setae; setae yellow to orange–brown. . . . . . . . . . . . . . . . . . . . . . . . . . . Acinia Robineau‑Desvoidy
– Wing pattern various, but infuscated areas usually more or less evenly brown (Figs. 105, 109, 113,
115), if margins of some areas narrowly darker brown, scutum with brown markings surrounding
acrostichal setae; setal color various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102

102. Wing with hyaline apical band extending from apex of vein R2+3 into cell m, without brown spots
on apices of veins  R4+5 and M, and with three  hyaline markings arising on anterior margin in
cell r1, proximal 2 connected to form V-shaped mark. . . . . . . . . . . . . . . . . . Eutreta Loew, in part
– Wing pattern various, hyaline apical band, if present, usually not extending to apex of vein R2+3
or not extending into cell m, and usually with brown spot on apex of vein R4+5 and/or M; cell r1
without all of above hyaline markings. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103

103. Wing banded (Fig. 113); abdomen dark brown, with at least male tergite 5 or female tergite 6 with
large shiny nonmicrotrichose area; Argentina, Chile. . . . . . . . . . . . . . . . Neosphaeniscus Norrbom
– Wing not banded (Figs. 105, 109, 116); abdominal tergites entirely microtrichose, matte. . . . 104

104. Wing radiate, with narrow brown rays extending to apices of veins R4+5 and M surrounding large
wedge-shaped apical hyaline area in cell  r4+5 (Fig.  109), and with middle half predominantly
brown; R4+5 dorsally with numerous setulae, extending well beyond level of dm-cu; cell r4+5 with
strong bulla anterior to dm-cu; southern Brazil. . . . . . . . . . . . . . [Lewinsohnia Norrbom & Prado]
– Wing usually reticulate, cell r4+5 with apical margin brown (Figs. 105, 115–116) or with broad
apical rays surrounding small ovoid hyaline spot, or if narrow apical rays present, wing with
middle half more extensively hyaline; R4+5 dorsally usually without setulae beyond level of r-m,
often bare; cell r4+5 at most with small weak bulla anterior to dm-cu . . . . . . . . . . . . . . . . . . . . 105

105. Lateral vertical seta dark brown to black, acuminate, similar in color and shape and more than half
as long as medial vertical seta; both orbital setae dark brown to black; vein R4+5 with numerous
long setulae ventrally to well beyond level of crossvein dm-cu; Colombia, Venezuela (high
elevations) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Homoeothrix Hering]
– Lateral vertical seta distinctly paler and more lanceolate than, and no more than half as long as,
medial vertical seta (Fig. 30); posterior orbital seta whitish, distinctly paler than anterior seta
unless anterior seta yellow; vein R4+5 bare ventrally or with few short setulae proximal to level of
dm-cu (most or all setulae usually proximal to r-m). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106

106. Hind femur without posterodorsal preapical seta, only anterodorsal preapical seta present (Fig. 58);
pterostigma entirely brown; scutum more or less evenly gray or brownish . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tephritis Latreille, in part
– Hind femur usually with posterodorsal and anterodorsal preapical setae (as in Fig. 56); pterostigma
usually with one to two hyaline or distinctly paler spots or mostly hyaline or paler than bordering
area in cell r1 (Figs. 115–116), if entirely brown (some males of N. semifusca (Wulp)), scutum
darker brown posterior to acrostichal setae and basolaterally on scutellum. . Neotephritis Hendel
TEPHRITIDAE (fruit flies, moscas de frutas) 68 947

Synopsis of the fauna from the Central American region (including Neotropical
Mexico). Their biology is poorly known, but some species
The world fruit fly species were cataloged by Norrbom have been bred from flowers of Asteraceae, including species
et al. (1999a). The updated version of their database is of Calea (Neurolaeneae) and Otopappus, Perymenium, and
available online at http://www.sel.barc.usda.gov:591/dip- Wedelia (Heliantheae). The latest key by Hendel (1914b) is
tera/Tephritidae/TephIntro.html. As of February 2008, outdated; only four of the included species are currently clas-
there were 4541 valid, extant species or subspecies of fruit sified in Acrotaenia.
flies worldwide, grouped in 493 genera, of which 71 genera
and 817 species (two genera and two species introduced) oc- Agallamyia Norrbom. This genus includes a single spe-
cur in the Neotropical Region. Two fossil genera and three cies, A. pendula Norrbom, from highland areas of Guate-
fossil species also have been described from the region, mala. It probably is most closely related to Ostracocoelia. It
in amber from the Dominican Republic (Norrbom, 1994a; was reared from large spherical stem galls on two species of
Norrbom & Condon, 1999). At least 62 genera and 506 spe- Perymenium (Asteraceae: Heliantheae).
cies, many of which are undescribed, are known from Mex-
ico (including Nearctic areas) and Central America. The Alujamyia Norrbom. This Neotropical genus and the
number of species known for Mexico (including Nearctic four known species were described and keyed by Norrbom
part) and the individual countries of Central America are as (2006b). Two species occur in Neotropical Mexico and Gua-
follows: Mexico, 289; Belize, 21; Guatemala, 136; Hondu- temala; the other two are in the Greater Antilles. Their biol-
ras, 25; El  Salvador, 24; Nicaragua, 29; Costa Rica, 203; ogy is unknown.
and Panama, 116.
Anastrepha Schiner. This is the largest genus in the New
Acidogona Loew (=Xenochaeta Snow). This genus in- World, with more than 200 currently recognized species. It
cludes three recognized species, A. melanura (Loew), from may be paraphyletic with respect to Toxotrypana (Norrbom
eastern North America, and A.  dichromata (Snow), from et al., 1999b; McPheron et al., 1999). It is mainly Neotropi-
western North America, and A. stecki Norrbom from Chia- cal, although 18 species have been recorded as far north as
pas and Guatemala. A single teneral male from Chihuahua, the extreme southern United States. The latest comprehen-
the only specimen known from northern Mexico, may be sive keys, by Stone (1942) and Steyskal (1977), are useful
A. melanura. An undescribed species occurs in Chiapas and but out of date. Hernández-Ortiz (1992) keyed the Mexi-
Guatemala. All three species breed in flowers of Hieracium can species, and Norrbom (1991, 1997b, 1998, 2002a) re-
species (Asteraceae: Cichorieae) (Foote et al., 1993; Goeden vised various species groups. At least 90 described and 11
and Teerink, 1997; Norrbom et al., 2010). undescribed species occur in the Central American region
Acinia Robineau-Desvoidy. This genus includes 13 spe- (including Neotropical Mexico). Except for A.  manihoti
cies from the Palearctic Region and New World, although Lima, which breeds in buds and stems of cassava, the larvae
Korneyev (1999b) suggested that the nine American species of Anastrepha species feed in fruits or seeds. They attack
should be removed and placed in a new genus. Only A. pic- a wide variety of plants (Norrbom, 2004), although many
turata (Snow) is known from the Central American region species breed in plants with milky latex, especially Sapota-
(Neotropical Mexico, Belize, Guatemala). It also occurs ceae, Moraceae, and Apocynaceae. One large species group
northward to the eastern coastal and southern United States breeds in Passiflora, and many others breed in various Myrt-
and in the West Indies. The other American species are known aceae. A number of species are generalist pests, including
from Ecuador to Chile, Argentina, and southern Brazil. They the A.  fraterculus (Wiedemann) complex (South American
breed in flowers of Pluchea and Tessaria (Asteraceae: Inu- fruit fly), A.  ludens (Loew) (Mexican fruit fly), A.  obliqua
leae) (Prado et al., 2002). Most of the American species can (Macquart) (West Indian fruit fly), and A. suspensa (Loew)
be identified using the key of Aczél (1958). (Caribbean fruit fly), and various others are minor or poten-
tial pests.
Aciurina Curran. This endemic North American genus
includes 15 currently recognized species. Only A.  mixteca Bactrocera Macquart. This is the largest genus of Te-
Hernández-Ortiz (1994), from Oaxaca, Mexico, is known phritidae, with more than 550 recognized species, many of
from the Neotropical Region. At least three other species,
which are major pests, together attacking a wide variety of
treated by Foote et al. (1993), occur in northern Mexico.
Aciurina species are gall-formers on Asteraceae, including fruits. Bactrocera is restricted to the Old World, except for
species of Baccharis, Chrysothamnus, and Ericameria (As- two invasive pest species: B. carambolae in the Guianas and
tereae) and Artemisia (Anthemideae). Brazil (Amapá) and B. oleae (Rossi) in California and north-
western Mexico. This genus is not yet known from Central
Acrotaenia Loew. This American, mainly Neotropical, America, although it is likely to be introduced as specimens
genus includes seven described species. Two of the described of various species are commonly intercepted at ports and
species and three to four undescribed species are known other points of entry worldwide.
948
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Baryplegma Wulp. This endemic Neotropical genus in- United States (New Mexico, Texas), and numerous un-
cludes 11 described species. At least four described and six described species. At least three described and three probably
undescribed species occur in the Central American region undescribed species occur in the Central American region
(including Neotropical Mexico). Little is known of their biol- (including Neotropical Mexico). Aczél (1953a) provided
ogy, but the four species that have been reared attack species the most recent key (to 12 species). Cecidochares species
of Heliantheae: Ecliptinae (Asteraceae). One species breeds have been reared mainly from plants of the tribe Eupatorieae
in flowers of a species of Melanthera, and two undescribed (Asteraceae). The majority form stem, or less commonly,
species have been reared from a Lasianthaea species, one flower galls, but several species breed in flowerheads without
from flowerheads, and one from stem galls (A.L. Norrbom, forming galls (Prado et al., 2002). Cecidochares connexa has
unpublished data). The latest key, by Hendel (1914b, as been introduced to various Old World countries for biologi-
Pseudacrotaenia), included only five species that are cur- cal control of the weed Chromolaena odorata.
rently classified in the genus.
Ceratitis MacLeay. This genus of nearly 100 described
Blepharoneura Loew. This endemic Neotropical genus species is restricted to the Afrotropical Region, except for
includes only 22 described species, but its diversity has been C. capitata, the Mediterranean fruit fly, which has been in-
estimated to be nearly 10 times as great (Condon & Norrbom, troduced to many other areas, including much of the Neotro-
1999). It is the only genus of Blepharoneurinae known from pical Region. Ceratitis capitata is one of the most generalist
the Central American region (including Neotropical Mexi- species of Tephritidae and is a major pest of many types of
co), where 10 described and at least 30 undescribed species fruit. In Central America, it occurs from Guatemala to Pan-
occur. No modern species keys have been published. Species ama, and it has been eradicated from southern Mexico and
of Blepharoneura are highly host-specific and breed only in Belize.
native species of Cucurbitaceae, but in a variety of host tis-
sues (fruits, seeds, male or female flowers, or stems). Adults Chetostoma Rondani. This genus includes 15 described
are unusual in having labellar spinules used to rasp and feed species, mainly from the Palearctic and Oriental Regions.
upon plant tissues (Condon & Norrbom, 1999). Two species occur in the Nearctic Region and were treat-
ed by Foote et al. (1993). Within the Neotropical Region,
Caenoriata Foote. This genus is known from only the type Chetostoma is so far known from only three specimens
species, C. pertinax (Bates), from southern Brazil (R.H. Foote, from Chiapas, Mexico, which are similar to C. californicum
1980), and an undescribed species from Mexico (Jalisco). It is Blanc. Chetostoma species breed in fruit of Lonicera species
likely to occur in elsewhere in central and southern Mexico (Caprifoliaceae) (Foote et al., 1993; Han, 1999).
and Central America. Caenoriata pertinax breeds in flower-
heads of a Piptocarpha species (Asteraceae: Vernonieae). Cryptodacus Hendel. This Neotropical genus of 10
described species was revised by Norrbom (1994b) and
Campiglossa Rondani (=Paroxyna Hendel). This is one Norrbom & Korytkowski (2008). Seven species are known
of the few cosmopolitan genera of Tephritidae, and it is also from the Central American region (Neotropical Mexico,
among the largest with nearly 180 valid species. At least 12 Guatemala, Costa Rica, Panama). A host plant is known only
species occur in Mexico (including Nearctic part) and Cen- for C. silvai (Lima) from Brazil, which was reared from fruit
tral America, mainly at higher elevations, but their taxonomy of a Loranthus sp. (Loranthaceae).
and distributions are poorly understood. The only species
key for the region, by Wulp (1899, as Ensina), is badly out- Dictyotrypeta Hendel. This mainly Neotropical genus is
dated. Campiglossa species breed in Asteraceae. The Neo- in need of revision. It currently includes six described spe-
tropical species probably have host plants in various tribes cies, but numerous undescribed species will also key here.
of the family like the Nearctic species (Goeden, 1994), but Whether they all form a monophyletic group is uncertain.
many seem to be associated with species of Senecioneae. Several species, such as D. incisa (Wulp) from central and
Most Campiglossa species breed in flowerheads, but at least southern Mexico and Guatemala and D.  crenulata (Wulp)
one central Mexican species mines stems (A.L.  Norrbom, from Mexico (Sinaloa, Guerrero, Veracruz) have been in-
unpublished data). cluded only tentatively. In addition to the above two species,
the Central American fauna also includes two undescribed
Carpomya Costa. This genus includes five species from species and the type species, D.  syssema Hendel, which is
the Palearctic, Oriental, and Afrotropical Regions and one known from Costa Rica to Peru. The known host plants of
species, C. tica Norrbom (1997a), known only from Costa Dictyotrypeta are species of Asteraceae, mostly in the tribes
Rica (Zurquí). The Old World species, several of which are Heliantheae and Vernonieae (Prado et al., 2002; Norrbom et
considered pests, breed in fruit of Cucurbitaceae, Rhamna- al., 2010).
ceae, or Rosaceae, but the host plant of C. tica is unknown.
Dioxyna Frey. This genus of 11 described species is
Cecidochares Bezzi. This mainly Neotropical genus in- widespread, occurring in all faunal regions except the Ant-
cludes 13 described species, one of which ranges north to the arctic. Six species occur in the Americas, although there are
TEPHRITIDAE (fruit flies, moscas de frutas) 68 949

at least several more undescribed Neotropical species. Two mined specimens from Colombia, Ecuador, and Brazil may
described and one undescribed species are known from the be congeneric (R.H. Foote, 1980). The biology of the type
Central American region (including Neotropical Mexico). species, E. paupera Hering, is unknown, but its host plant is
Dioxyna species breed in flowers of various Asteraceae, but presumably an Asteraceae species.
the American species have been reared mainly from species
Eutreta Loew. This genus includes 35 described and nu-
of Bidens, Coreopsis, and other genera of Coreopsideae; the
merous undescribed species ranging from southern Canada
larvae feed within individual achenes within the flowerhead
to South America. At least six described and six undescribed
(Headrick et al., 1996; Prado et al., 2002). No key is avail-
species occur in the Central American region (including
able for the Neotropical species. Neotropical Mexico), and 10 additional species are known
Dracontomyia Becker. This genus includes five described from Nearctic Mexico. The described species were treated in
and at least three undescribed species, mostly from South the latest revision of the genus by Stoltzfus (1977). Eutreta
America (Ecuador, Peru, Bolivia, southern Brazil, Argentina) species are gall-formers, mainly on species of Asteraceae in
(Norrbom et al., 2010). Dracontomyia borrichia (Bush  & a variety of tribes, but a few species attack Lamiaceae, Lo-
Huettel) is recorded only from southern Texas (USA) but un- ganiaceae, Scrophulariaceae, Solanaceae, or Verbenaceae.
doubtedly occurs in at least northern Mexico, and the genus Eutreta xanthochaeta, the lantana gall fly, has been used as
is likely to occur in the Central American region. Draconto- a biological control agent against the invasive weed Lantana
myia species breed in Asteraceae. At least two of the South camara in Hawaii.
American species are gall-formers (A.L.  Norrbom, unpub- Gymnocarena Hering. This genus includes 16 described
lished data), but D. borrichia breeds in flowers of Borrichia species, revised by Norrbom (1992, 2006a), and at least three
frutescens (Bush  & Huettel, 1970). The described species undescribed species (B.D.  Sutton & G.J.  Steck, personal
were keyed by Aczél (1953a) and Bush & Huettel (1970, as communication). It ranges from Canada to the mountains of
Cecidocharella). central and southern Mexico and Guatemala, with one spe-
Dyseuaresta Hendel. This genus includes 10 described cies known from Costa Rica. The Neotropical species oc-
Neotropical species, two of which, D. mexicana (Wiedemann) cur at high elevations, such as Cerro de la Muerte in Costa
and D. sobrinata (Wulp), occur in the Central American re- Rica. Gymnocarena species breed in flowers of Asteraceae,
gion and extend northward to the United States. Dyseuaresta including species of Dahlia (Coreopsideae) and sunflowers
and the closely related genera Lamproxynella and Pseudoe- and related plants of the genera Helianthus, Heliomeris, and
daspis are poorly delimited and all are in need of revision. Verbesina (Heliantheae).
No key to identify the species is available. Dyseuaresta spe- Hexachaeta Loew. This genus, which includes 31 de-
cies breed in flowerheads of Asteraceae (mainly Heliantheae, scribed species, ranges from the USA (southern Texas) to
such as Aspilia and Melanthera) (Prado et al., 2002). northern Argentina and the Greater Antilles (Jamaica). At
Euaresta Loew. This genus includes 15 valid species least 11 described and two undescribed species occur in the
occurring from Canada to Argentina. Five are known from Central American region (including Neotropical Mexico).
Nearctic Mexico, and a sixth species is from El Salvador. There are keys to the subgenera and some species groups
The Nearctic species were treated by Foote et al. (1993), and (Hernández-Ortiz, 1999, 2002, 2006; Lima, 1953; Lima  &
the Neotropical species were keyed by Norrbom (1993), who Leite, 1952), and the rest of the genus is being revised by
also provided a phylogeny of the species. Euaresta species V. Hernández-Ortiz. The biology of Hexachaeta is not well
breed in flowers of ragweeds, bursages, and cockleburs of known, but nine species have been reared; the reported host
the closely related genera Ambrosia, Xanthium, and Dico- plants, except for one species of Bignoniaceae and one spe-
ria (Asteraceae: Heliantheae: Ambrosiinae), some of which cies of Verbenaceae, belong to the Moraceae (Norrbom et
are serious allergens and agricultural weeds. Some Euaresta al., 1999b). In at least one species, the larvae feed on the
species have therefore been studied or used as biological developing seeds within the fruit.
control agents.
Molynocoelia Giglio-Tos. This Neotropical genus of
Euarestoides Benjamin. This genus includes three de- four known species was revised by Norrbom (2006b). Three
scribed North American species, two of which are known species occur in the Central American region (Neotropical
from Mexico (south to Oaxaca and Chiapas) and Guatemala, Mexico, Guatemala, Costa Rica), and the fourth is known
and two undescribed species from South America. The three only from Brazil. Their biology is unknown.
described species were treated by R.H. Foote (1958). Euares-
toides species breed in flowers of various genera of Asteraceae Neomyopites Freidberg & Norrbom. This genus, which
(Wasbauer, 1972; Foote et al., 1993; Prado et al., 2002). includes 28 described species, occurs in the West Indies
and from the USA (southern Texas) to Bolivia and southern
Euarestopsis Hering. The single described species of this Brazil. At least eight species occur in the Central American
genus is recorded only from Costa Rica, although undeter- region (including Neotropical Mexico). Most of the species
950
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

were keyed by Steyskal (1979, as Urophora). Neomyopites undescribed species, two of which occur in the Central Amer-
species breed in flowerheads or form seed or stem galls on a ican region (Neotropical Mexico, Guatemala, Costa Rica). A
variety of genera of Asteraceae, mainly in the Heliantheae, few species have been keyed (Malloch, 1933, five species in
Eupatorieae, and Astereae (Freidberg  & Norrbom, 1999; a key to Tephritis; Hering, 1940, 1941, seven species), but no
Prado et al., 2002). comprehensive key is available. Most of the South American
species previously placed in Tephritis have been transferred
Neotaracia Foote. This Neotropical genus includes three here. Plaumannimyia species breed in flower heads of Bac-
described and one undescribed species, three of which occur charis and its close relatives (Asteraceae: Astereae) (Prado et
in the Central American region (including Neotropical Mex- al., 2002; Norrbom et al., 2010).
ico). R.H. Foote (1979a) revised the three described species.
Neotaracia species occur at low to mid-elevations; Neotara- Polionota Wulp. This Neotropical genus was revised by
cia imox (Bates) has been reared from flowers of Synedrella Norrbom (1988). Seven of the eight known species occur in
nodiflora (Asteraceae: Heliantheae). the Central American region (including Neotropical Mexi-
co). The only reared species was from a gall on an unidenti-
Neotephritis Hendel. Ten described species are current- fied plant, presumably a species of Asteraceae.
ly included in this genus, although it may not be a natural
group. Five described and seven undescribed species occur Polymorphomyia Snow. This Neotropical genus includes
in the Central American region (including Neotropical Mex- five described and at least one undescribed species, of which
ico). Steyskal (1972a) keyed the described species, but revi- two occur in the Central American region (Neotropical Mex-
sion of this genus is needed. Neotephritis species have been ico, Costa Rica). Korytkowski (1971) provided a key to four
reared from flowers of a variety of genera of Heliantheae species. Only two species have been reared, both from Aster-
(Asteraceae). aceae: P. footei Korytkowski from Peru forms leaf galls on a
species of Tessaria (Inuleae) (Korytkowski, 1971), and P. ba-
Ostracocoelia Giglio-Tos. This genus includes a single silica Snow from the Antilles was reared from a stem gall on
species, O. mirabilis Giglio-Tos, known from Mexico, Gua- Chromolaena odorata (Eupatorieae) (Wolcott, 1936).
temala, and Costa Rica. It has been reared from stem galls on
a species of Ageratina (Asteraceae: Eupatorieae) (Norrbom Procecidochares Hendel. This genus includes 18 de-
et al., 2010). scribed species, most of which are restricted to North Amer-
ica. At least three described and three undescribed species
Paracantha Coquillett. This genus, which includes 11 occur in the Central American region (Neotropical Mexico,
described species, ranges from Canada to Argentina and Guatemala, Costa Rica), and an additional eight are known
Chile. Four described and at least two undescribed species from Nearctic Mexico. Foote et al. (1993) and Goeden  &
occur in the Central American region (including Neotropical Norrbom (2001) keyed the species occurring in the United
Mexico). Keys to most of the species are available in Aczél States and Canada, but no keys are available for the Neo-
(1952a) and Foote et al. (1993, Nearctic species only). Para- tropical species. Procecidochares species breed in Asterace-
cantha species breed in flowerheads of various Asteraceae, ae, mostly as gall formers, although some are flower feeders
including thistles, Cirsium spp. (Cardueae), Dahlia (Coreop- and at least one species has alternate flower-feeding and
sideae), Calea (Neurolaeneae), and sunflowers and related gall-forming generations. Many species breed in goldenrods
genera, such as Borrichia, Helianthus, and Viguiera (He- (Solidago) and related genera (Astereae), but others breed in
liantheae) (Foote et al., 1993; Prado et al., 2002; Norrbom genera of Heliantheae, such as Ambrosia and Viguiera, and
et al., 2010). the few known hosts of the Mexican and Central American
P. utilis group are species of Ageratina and Brickellia (Eu-
Parastenopa Hendel. This genus includes one described patorieae) (Foote et al., 1993; Goeden  & Norrbom, 2001).
Nearctic species and nine described and numerous un- Procecidochares alani Steyskal and P. utilis Stone have been
described Neotropical species (Norrbom  & Han, in prepa- introduced to Hawaii, Australia, and New Zealand, and the
ration). At least 18 species, only one of them described, latter species also to additional countries, as biological con-
occur in Costa Rica and/or Panama. Some of the described trol agents of weedy Ageratina species.
species were revised by Aczél (1956). Several species have
been reared from species of Ilex (Aquifoliaceae), includ- Procecidocharoides Foote. This genus, which was re-
ing I. paraguariensis (yerba-mate), one from the fruit, two vised by R.H. Foote (1960), includes three described species
mining shoots, and one from leaf galls of a psyllid. Another from Canada and the United States, one of which occurs in
species was reared from bud galls caused by a species of Mexico south to Puebla. An undescribed species is known
Cecidomyiidae on a Marcetia sp. (Melastomataceae) (Aczél, from central Mexico (Distrito Federal), and specimens simi-
1956). lar to P. flavissima Foote were collected in Panama. Proceci-
docharoides penelope (Osten Sacken), the only species that
Plaumannimyia Hering (=Trypanaresta Hering). This has been reared, breeds in flowerheads of Ageratina altissi-
New World genus includes 20 described and numerous ma (Asteraceae: Eupatorieae).
TEPHRITIDAE (fruit flies, moscas de frutas) 68 951

Pseudophorellia Lima. This genus was revised by Tetreuaresta Hendel. This Neotropical genus includes
Norrbom (2006b). It includes 25 species from southern Mex- 19 described species, six of which occur in the Central
ico (Chiapas) to Bolivia and from Hispaniola, 13 of which American region (including Neotropical Mexico), and nu-
are known from the Central American region. Their biology merous undescribed species. Steyskal (1972b) provided a
is unknown. key to 15 species, but the genus is in need of a modern
revision. The biology of most species is unknown, but at
Pyrgotoides Curran. This Neotropical genus includes least five species breed in flowerheads of genera of Ver-
three described and at least four undescribed species, mainly nonieae (Asteraceae) (Prado et al., 2002). Tetreuaresta ob-
from Ecuador, Peru, and Bolivia. The type species, P. cras- scuriventris (Loew) has been used as a biological control
sipes Curran, was described from Panama, and single male agent against Elephantopus mollis in Hawaii and other Pa-
specimens examined from Costa Rica (Estación Pitilla) and cific Islands.
Ecuador may be this species. Two species were keyed by
Korytkowski (1976, as Gerrhoceras). Only P.  peruvianus Tomoplagia Coquillett. This New World genus includes
(Korytkowski) has been reared; it forms large stem galls on a 61 described and numerous undescribed species, of which at
species of Ophryosporus (Asteraceae: Eupatorieae) (Koryt- least 15 (six described) occur in the Central American region
kowski, 1976). (including Neotropical Mexico). Aczél (1955a, b) and Prado
et al. (2004) treated the known species. Tomoplagia species
Rhagoletis Loew. This mainly Holarctic and Neotropical breed in Asteraceae, mainly in genera of Vernonieae and to
genus includes 70 described species, of which 21 occur in the a lesser extent Mutiseae, Heliantheae, and other tribes. Most
Neotropical Region. At least seven described and two unde- species breed in flowerheads, but one South American spe-
scribed species are known from Neotropical Mexico or else- cies forms a large stem gall (Prado et al., 2002).
where in the Central American region. Bush (1966) revised
the Nearctic species, and R.H. Foote (1981) and Hernández- Toxotrypana Gerstaecker. This Neotropical genus in-
Ortiz  & Frías (2000) treated most of the Neotropical spe- cludes seven described and at least a similar number of un-
cies. Species of Rhagoletis breed in fruits of various plant described species, all of which are wasp mimics. At least
families, with the majority of the Neotropical species attack- four species occur in Mexico and Central America, including
ing Solanaceae or Juglans species (Juglandaceae) (Smith & T.  curvicauda Gerstaecker, the papaya fruit fly. Blanchard
Bush, 1999). Several species are pests in the region, includ- (1960) published the only key to species. Toxotrypana species
ing R.  lycopersella Smyth, R.  nova (Schiner), R.  striatella breed in fruits of species of Asclepiadaceae and Caricaceae
Wulp, and R. tomatis Foote. that have thick casks and produce milky latex (Norrbom et
al., 1999b). The larvae feed on the developing seeds and as-
Rhagoletotrypeta Aczél. This genus, which includes 10 sociated tissues.
described species, ranges from the northeastern United Sates
to Argentina and Cuba. One species is reported from Mexico Trupanea Schrank. With 226 valid species, this is the
and Costa Rica and another from central Mexico (Morelos). largest genus of Tephritinae. It occurs in all biogeographic
Norrbom (1994b) keyed most of the known species, which regions and includes 88 described and numerous undescribed
breed in fruit of Celtis spp. (Ulmaceae). Neotropical species, of which at least 17 occur in the Central
American region. An additional 12 species are known from
Rhynencina Johnson. This genus includes five spe- Nearctic Mexico. Some Neotropical species were revised or
cies: one from the eastern United States; one known from included in keys by Hendel (1914b), Malloch (1933), Her-
Mexico (Veracruz, Chiapas), Guatemala, and Honduras; and ing (1941), Aczél (1953b, c), Frías (1985), and Foote et al.
three from the Andes (Colombia, Ecuador, Venezuela). The (1993), but the Neotropical fauna is badly in need of com-
four Neotropical species were keyed by Steyskal (1979, as prehensive revision. Trupanea species have been reared from
Urophora). Rhynencina species have been reared from flow- a variety of genera of Asteraceae; most species breed in the
erheads of Espeletia and Smallanthus species (Asteraceae: flowerheads (Prado et al., 2002).
Heliantheae) (Freidberg  & Norrbom, 1999; Steck et al.,
2003). Trypeta Meigen. This genus includes nearly 60 valid
species and occurs in the Oriental, Palearctic, and Nearc-
Tephritis Latreille. This mainly Holarctic genus includes tic Regions, as well as the Neotropical Region at least as
150 species, of which at least seven occur in Mexico and far south as Costa Rica. Twelve species are known from
Guatemala (three endemic). The South American species pre- highland areas of Mexico, including several from Oaxaca
viously classified in Tephritis have been transferred, mostly and Chiapas and another also known from Guatemala, and
to Plaumannimyia. The key of Foote et al. (1993) includes three species are known from Costa Rica (Han & Norrbom,
four of the species known from Mexico. Most New World 2005). Trypeta species are leaf-miners in species of Aster-
Tephritis species breed in flowerheads or in a few cases form aceae; the host plants of the New World species that have
galls or feed in stem mines on species of Asteraceae (Foote been reared are mainly species of Senecio and related gen-
et al., 1993; Goeden, 1993). era (Senecioneae).
952
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Xanthaciura Hendel. This New World genus includes 17 Zonosemata Benjamin. This genus, which includes eight
described species and at least several more undescribed spe- described species, ranges from Canada to Colombia and
cies. At least 10 occur in the Central American region (in- northern Brazil and the Greater Antilles (Jamaica). Three
species occur in the Central American region (including
cluding Neotropical Mexico). Most of the valid species were
Neotropical Mexico). Norrbom (2002b) keyed the known
revised by Aczél (1950, 1952b). Xanthaciura species breed species, which breed in fruits of Solanaceae species. The
in flowers of various genera of Asteraceae (mainly Eupato- eastern North American Z. electa (Say) is a pest of peppers
rieae, Coreopsideae, and Heliantheae) (Prado et al., 2002). and eggplant.

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954
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Norrbom, A.L. 1993. New species and phylogenetic analysis of nae: Toxotrypanini) based on morphology, pp.  299–342. In
Euaresta Loew (Diptera: Tephritidae) with a key to the spe- Aluja, M. & A.L. Norrbom (editors). Fruit Flies (Tephritidae):
cies from the Americas south of Mexico. Proceedings of the Phylogeny and evolution of behavior. CRC Press, Boca Raton,
Entomological Society of Washington 95: 195–209. London, New York, Washington, D.C., [16] + 944 pp.
Norrbom, A.L. 1994a. New genera of Tephritidae (Diptera) from Norrbom, A.L., B.A. Sutton, G.J. Steck, & J. Monzón. 2010. New
Brazil and Dominican amber, with phylogenetic analysis of the genera, species and host plant records of Nearctic and Neotropi-
tribe Ortalotrypetini. Insecta Mundi 8: 1–15. cal Tephritidae (Diptera). Zootaxa 2398: 1–65.
Norrbom, A.L. 1994b. New species and phylogenetic analysis of Prado, P.I., T.M.  Lewinsohn, A.M.  Almeida, A.L.  Norrbom,
Cryptodacus, Haywardina, and Rhagoletotrypeta (Diptera: Te- B.D.  Buys, A.C.  Macedo,  & M.B.  Lopes. 2002. The fauna of
phritidae). Insecta Mundi 8: 37–65. Tephritidae (Diptera) from capitula of Asteraceae in Brazil.
Norrbom, A.L. 1997a. The genus Carpomya Costa (Diptera: Te- Proceedings of the Entomological Society of Washington 104:
phritidae): new synonymy, description of first American species, 1007–1028.
and phylogenetic analysis. Proceedings of the Entomological Prado, P.I., A.L. Norrbom, & T.M. Lewinsohn. 2004. New species
Society of Washington 99: 338–347. of Tomoplagia Coquillett (Diptera: Tephritidae) from capitula of
Norrbom, A.L. 1997b. Revision of the Anastrepha benjamini spe- Asteraceae in Brazil. Neotropical Entomology 33: 189–211.
cies group and the A. pallidipennis complex (Diptera: Tephrit- Smith, J.J. & G.L. Bush. 1999. Phylogeny of the subtribe Carpo-
idae). Insecta Mundi 11: 141–157. myina (Trypetinae), emphasizing relationships of the genus
Norrbom, A.L. 1998. A revision of the Anastrepha daciformis spe- Rhagoletis, pp.  187–217. In Aluja,  M.  & A.L.  Norrbom (edi-
cies group (Diptera: Tephritidae). Proceedings of the Entomo- tors). Fruit Flies (Tephritidae): Phylogeny and evolution of
logical Society of Washington 100: 160–192. behavior. CRC Press, Boca Raton, London, New York, Wash-
Norrbom, A.L. 2002a. A revision of the Anastrepha serpentina spe- ington, D.C., [xvi] + 944 pp.
cies group (Diptera: Tephritidae). Proceedings of the Entomo- Steck, G.J., B.D. Sutton, & D. De Foe. 2003. Biology of Rhynenci-
logical Society of Washington 104: 390–436. na longirostris Johnson (Diptera: Tephritidae). Proceedings of
Norrbom, A.L. 2002b. A new species and key for the genus Zonose- the Entomological Society of Washington 105: 542–547.
mata Benjamin (Diptera: Tephritidae). Proceedings of the Ento- Steyskal, G.C. 1972a. A preliminary key to the species of Neote-
mological Society of Washington 104: 614–623. phritis Hendel (Diptera: Tephritidae). Proceedings of the Ento-
Norrbom, A.L. 2004. Host plant database for Anastrepha and Tox- mological Society of Washington 74: 414–416.
otrypana (Diptera: Tephritidae: Toxotrypanini). Diptera Data Steyskal, G.C. 1972b. A preliminary key to the species of the
Dissemination Disk 2 (CD-ROM). Neotropical genus Tetreuaresta Hendel (Diptera: Tephritidae).
Norrbom, A.L. 2006a. New species and host records for Gym- Proceedings of the Entomological Society of Washington 74:
nocarena Hering (Diptera, Tephritidae). In Merz,  B. (editor). 403–405.
Phylogeny, taxonomy, and biology of tephritoid flies (Diptera, Steyskal, G.C. 1977. Pictorial key to species of the genus Anas-
Tephritoidea). Proceedings of the 3rd Tephritoid Taxonomist’s trepha (Diptera: Tephritidae). Entomological Society of Wash-
Meeting, Geneva, 19–24 July 2004. Instrumenta Biodiversitatis ington, Washington, D.C, 35 pp.
7: 217–226. Steyskal, G.C. 1979. Taxonomic studies on fruit flies of the genus
Norrbom, A.L. 2006b. A revision of the Neotropical genera Mo- Urophora (Diptera: Tephritidae). Entomological Society of
lynocoelia Giglio-Tos, Pseudophorellia Lima, and Alujamyia, Washington, Washington, D.C., 61 pp.
n.  gen. (Diptera: Tephritidae). Israel Journal of Entomology Stoltzfus, W.B. 1977. The taxonomy and biology of Eutreta (Diptera:
35–36: 35–134. Tephritidae). Iowa State Journal of Research 51: 369–438.
Norrbom, A.L. & M.A. Condon. 1999. Phylogeny of the subfamily Stone, A. 1942. The fruitflies of the genus Anastrepha. United
Blepharoneurinae, pp. 135–155. In Aluja, M. & A.L. Norrbom States Department of Agriculture Miscellaneous Publication
(editors). Fruit Flies (Tephritidae): Phylogeny and evolution of 439. pp. 1–112.
behavior. CRC Press, Boca Raton, London, New York, Wash- Wasbauer, M.S. 1972. An annotated host catalog of the fruit flies
ington, D.C., [16] + 944 pp. of America north of Mexico (Diptera: Tephritidae). Occasional
Norrbom, A.L.  & C.A. Korytkowski. 2008. New Cryptodacus Papers, California Department of Agriculture, Bureau of Ento-
(Diptera: Tephritidae) from Panama, with a key to the known mology 19: 1–172.
species. Zootaxa 1773: 31–43.
Wolcott, G.N. 1936. “Insectae borinquenses” a revised annotated
Norrbom, A.L., L.E. Carroll, F.C.  Thompson, I.M.  White,  &
checklist of the insects of Puerto Rico. Journal of Agriculture of
A. Freidberg. 1999a. Systematic database of names, pp. 65–251.
the University of Puerto Rico 20: 1–600.
In Thompson, F.C. (editor). Fruit Fly Expert Identification Sys-
tem and Systematic Information Database. Myia 9, vii + 524 pp. Wulp, F.M. van der. 1899. Fam. Muscidae, pp. 393–416. In God-
(1998), and Diptera Data Dissemination Disk 1 (CD-ROM) man, F.D. & O. Salvin (editors). Biologia Centrali-Americana,
(1998). or, contributions to the knowledge of the fauna and flora of
Norrbom, A.L., R.A. Zucchi, & V. Hernández-Ortiz. 1999b. Phy- Mexico and Central America. Zoologia. Class Insecta. Order
logeny of the genera Anastrepha and Toxotrypana (Trypeti- Diptera. Vol. 2. London, 489 pp.
PYRGOTIDAE (pyrgotid flies) 69
Vicente Hernández-Ortiz

Fig. 69.1. Female of Pyrgota undata Wiedemann, (Nearctic, MND, fig. 65.1).

Diagnosis 1+2 elongate, usually with several long hairs on lateral sides
near base. Female terminalia (Figs. 8–12) with syntergoster-
Medium to large-sized flies (body length 5–30  mm), nite 7 robust and strong basally forming conical tube turned
body usually robust with long legs, wing pattern often spot- ventrally, sometimes longer than rest of abdominal length.
ted, banded, or reticulated, rarely hyaline (Fig.  1). Head Apical margin of syntergosternite 7 sometimes provided
(Figs. 2–7) spherical with frons large, antennal bases project- with ventral hooklike sclerite, or two apicolateral clawlike
ed anteriorly, pedicel elongate; arista setiform; ocelli usually sclerites (according to Steyskal, 1972, probably derived from
lacking in American species (except in Descoleia Aczél); tergum and sternum 8); aculeus usually shorter than length
frons weakly depressed, frontal bristles and vibrissa absent, of syntergosternite 7. Male terminalia similar to Tephritidae
face broad, with deep antennal grooves; mouthparts of mod- and Platystomatidae; epandrium globose, curved ventrally;
erate size. Scutum mostly bare, setulose, or distinctly haired, lateral surstyli usually short and robust basally, medial sur-
sometimes black spotted, major bristles usually shorter than styli without prensisetae developed; aedeagus coiled and
in Tephritidae; scutellum short and broad with one to four elongated; glans more or less developed and partially sclero-
pairs of marginal bristles. Legs (Figs.  13–15) long in both tized in some species.
sexes, frequently without strong bristles, but covered by short Pyrgotidae are most similar to certain members of the
setulae; hind tibia tapering to base or with strong constriction family Conopidae by the similar shape of the body, and par-
in basal third or near midlength. Wing (Figs.  16–23) with ticularly the head shape, in addition to the absence of ocelli
or without subcostal break, C weakening after vein R4+5; Sc (subfamily Conopinae). However, the Pyrgotidae possess
complete to C or disappearing close to C; R1 dorsally setu- a normally developed, sponging proboscis, and veins  R4+5
lose; R2+3 sometimes with stump vein at apical third; cell cup and M1+2 are not convergent, ending well separated at the
closed by CuA2 usually forming acute angle in lower api- wing margin. In contrast, Conopidae possess a long (usually
cal corner, sometimes short. Preabdomen elongate, robust, geniculate) proboscis, and veins R4+5 and M1+2 are strongly
more or less petiolate; lateral membrane broad; syntergite convergent apically, and sometimes fused distally.

955
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Biology their crepuscular or nocturnal habits that are similar to those

of most of their hosts. Some other species, such as the Austra-
Little information is available on the biology of the Pyr- lian beetle Anoplognathus olivieri (Dalman) and its pyrgotid
parasite Maenomenus ensifer Bezzi (with ocelli developed),
gotidae, although it is well known that larvae are endopara-
have diurnal habits (Aczél, 1956a). Probably nocturnal or
sitic on adult scarabaeid beetles (Coleoptera: Scarabaeidae). diurnal activity explains the presence or absence of ocelli
The highly specialized morphology of the female terminalia throughout the family.
suggests that all pyrgotid species are parasites. For Central and South America, there is no biological in-
Forbes (1908) described the oviposition behavior of formation on this family. Most life history data are for some
Pyrgota undata Wiedemann females. He observed the flies North American species, such as Pyrgota undata.
waiting on the body of Phyllophaga adults until the beetles Some work has be done on the value of pyrgotids as con-
opened their elytra before flying. The female deposited an trol agents of pest scarabaeids in the New World. Pyrgota
egg (or larva?) into the abdomen of the host while both were undata was introduced in Puerto Rico as a potential control
of some injurious Phyllophaga species (Wolcott, 1923);
in flight. The life cycle of P. undata was described by Davis
Peltodasia flaviseta (Aldrich), a pyrgotid native to the Ori-
(1919), who found that female beetles are more frequently
ental Region, was introduced into the USA for the control
used as hosts. Parasitized beetles live 10–14 days, with only of the Japanese beetle, Popillia japonica (Gardner & Park-
one larva developing in each host. The fly pupates within the er, 1942). In both cases results were not satisfactory. More
host body, and developmental time from oviposition to pupa- recently, Crocker et al. (1996) conducted an evaluation of
tion is about three weeks. the flight period and egg production of P. undata, with re-
Although both males and females are attracted to light, spect to seasonal occurrence of pest Phyllophaga species
adult Pyrgotidae are uncommon in collections, because of in Texas.

2 Leptopyrgota 3 Leptopyrgota 4 Idiopyrgota

5 Pyrgota 7 Stenopyrgota

6 Anapyrgota

Figs. 69.2–7. Heads: lateral view of (2) Leptopyrgota undulata Becker; anterior view (antennae omitted) of (3) L. albitarsis Aczél, (South
America, Aczél, 1956b, fig. 29); and (4) Idiopyrgota setiventris Aczél, (South America, Aczél, 1956c, fig. 78); lateral view of (5) Pyrgota
lugens Wulp; (6) Anapyrgota sp.; and (7) Stenopyrgota sp. Figures 2 and 5–7 illustrated by V. Hernández-Ortiz.
PYRGOTIDAE (pyrgotid flies) 69 957

Classification sternum and tergum  6 in males, synsternite 7 + 8 absent,

phallus extremely short, female sternum 1 largely desclero-
Formerly this family was classified in three subfami- tized, aculeus short and blunt, and two rudimentary cerci
lies: Lochmostyliinae, Pyrgotinae, and Toxurinae (Aczél, bearing setae.
1956a; 1958; McAlpine, 1978; Steyskal, 1987), all of I distinguish two subfamilies in Pyrgotidae: Teretrurinae
them represented in the New World. In a recent treatment, (formerly Teretrurini), characterized by sternites  1 and 2
however, McAlpine (1989) proposed a different subfamily fully developed and free, and Pyrgotinae (including Tox-
arrangement. The Ctenostylinae (=Lochmostyliinae) was urini) with sternites 1 and 2 fused into one synsternite. The
removed from the Pyrgotidae as the family Ctenostylidae, subfamily Teretrurinae has an exclusively southern Neo-
suggesting that further studies could support its exclusion tropical distribution with the genera Pyrgotosoma Malloch
from the Tephritoidea. This was based on the well-developed and Teretrura Bigot recorded from Argentina and Chile.

9 Sphecomyiella

tg 6
ovpung

sg 7

8 Sphecomyiella 10 Idiopyrgota 11 Undescribed Genus A

12 Undescribed Genus A 13 Pyrgota 14 Stenopyrgota 15 Anapyrgota

Figs.  69.8–15. Female abdomen, terminalia, and legs: lateral view of abdomen, ovipositor extended, with offset of hooked apex of
sternite 7 in ventral view (8), and dorsal view of aculeus, or sclerotized apical portion of ovipositor (9) of Sphecomyiella valida (Harris),
(MND, figs. 65.8–9); lateral view of apex of abdomen and syntergosternite 7 of (10) Idiopyrgota sp.; ventral (11) and lateral (12) view of
apex of syntergosternite 7 of Undescribed Genus A; posterior view of left hind tibia of (13) Pyrgota lugens Wulp; (14) Stenopyrgota sp.; and
(15) Anapyrgota sp. Figures 10–15 illustrated by V. Hernández-Ortiz.
Abbreviations: ovpung, ovipositunguis; sg, segment; tg, tergite.
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

The subfamily Pyrgotinae includes two tribes: Toxurini Differences between Stirothrinax Enderlein and Spheco-
with a mainly Australian distribution, but also represented myiella could not be established based on currently known
in the Americas by the genus Descoleia, with one species, species, as well as additional studied specimens that repre-
D. teretrura Aczél from Argentina; and Pyrgotini, which sent undescribed species. For this reason, Stirothrinax will
is well represented in the Neotropical Region. In my eventually be considered a synonym of Sphecomyiella, as
study of the fauna of Central America, I found the genera also noted by Kondratieff & Fitzgerald (1993). Much work
Anapyrgota, Boreothrinax, Idiopyrgota, Leptopyrgota, is necessary to gain a better understanding of the relation-
Neopyrgota, Pyrgota, Sphecomyiella, and Stenopyrgota ships among pyrgotid genera.
to be present, in addition to three undescribed genera.
Recently, Bernardi (1990) proposed the synonymy of
Anapyrgota with Carrerapyrgota Aczél mainly based on the Identification
absence of the subscutellum. However, his proposal does not
discuss characters supporting the segregation of Carrerapy- The most comprehensive works for the recognition of
rgota by Aczél (1956), which are missing in Anapyrgota.
Neotropical taxa were published by Aczél (1956a, 1956b,
Such characters include the costal vein ending almost at the
level of vein R4+5 and the last section of vein M being barely 1956c). He described, illustrated, and keyed most of the spe-
evident after vein dm-cu, disappearing before the wing mar- cies, except those in the genera Sphecomyiella and Boreo-
gin. In addition, the discal cell is shorter and its lower thrinax, which are treated by Steyskal (1978, 1987).
apical corner ends distant from the wing margin, and there Some genera cannot be presently recognized in the male
are other outstanding differences in wing pattern. Therefore, sex (see couplet 7). Further study of male terminalia is nec-
in the present study this synonymy is not accepted. essary to remedy this situation.

Key to the genera of Pyrgotidae of Central America

1. Face without median keel, or antennal grooves completely joined (Fig. 3); if present, median keel
weak or poorly differentiated; alula indistinct to well developed. . . . . . . . . . . . . . . . . . . . . . . . . 2
– Face with antennal grooves separated by strongly developed median keel (Fig. 4); alula well
developed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

2. Alula well developed (Figs. 16, 18–21); several pairs of scutellar bristles usually present . . . . . 3
– Alula greatly reduced or absent (Figs. 17, 22); scutellar bristles usually absent, or sometimes with
only short pair of slender apical bristles present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

3. Subscutellum indistinct or absent; wings with brownish bands well differentiated by hyaline
areas; legs normally developed; female ovipositor sheath with neither apicolateral ovipositungues
nor apicoventral unguis; stump vein in R2+3 present (Fig. 21). . . . . . . . . . . . . Anapyrgota Steyskal
– Subscutellum well developed; wings mostly hyaline, with diffuse pattern, or hyaline spotted; legs
various; female ovipositor sheath with two apicolateral ungues or one apicoventral unguis; stump
vein in R2+3 usually absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4. Wing broadly hyaline or with slightly infuscate pattern (Fig. 18); legs elongated and slender with
hind tibia constricted near basal half; female ovipositor sheath with two claw-shaped apicolateral
ungues (Figs. 11–12) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Undescribed Genus A
– Wing brownish, hyaline spotted; legs normally developed with hind tibia tapering gradually to
base; female usually with hook-shaped apicoventral unguis (as in Fig. 8). . . . Neopyrgota Hendel

5. Vein R2+3 straight throughout length, becoming parallel with respect to R4+5; alula completely
absent; wing pattern irrorate, with small hyaline spots throughout surface (Fig. 22); pedicel
usually shorter than flagellum; head strongly depressed anteriorly, triangular shaped in lateral
view (Fig. 7). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stenopyrgota Malloch
– Vein R2+3 strongly sinuate on apical one-third, becoming convergent with R4+5; alula greatly
reduced; wing mostly hyaline usually with large anteroapical dark spot (Fig.  17); pedicel
elongate, usually longer than flagellum; head not as above (Fig. 2). . . . . . . Leptopyrgota Hendel
PYRGOTIDAE (pyrgotid flies) 69 959

6. Hind tibia tapering to base (as in Fig. 13).. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

– Hind tibia strongly constricted near basal one-third (as in Fig. 15).. . . . . . . . . . . . . . . . . . . . . . 10

7. Abdominal sternites 3 and 4 in females each with comb-shaped row of strong long bristles;
syntergosternite 7 lacking apicoventral unguis (Fig. 10); ocellar and postocellar bristles usually
well differentiated (Fig. 4). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Idiopyrgota Aczél
– Abdominal sternites in females without any distinct strong bristles; other characters various . . 8

8. Syntergosternite 7 without apicoventral hook-shaped unguis; body dark brown to black; wing
pattern brown with extensive irregular hyaline spots . . . . . . . . . . . . . . . . . . Undescribed Genus B
– Syntergosternite 7 with strong apicoventral hook-shaped unguis (as in Fig.  8); body yellow,
reddish or brownish, rarely black; wing pattern uniformly darkened, or pale brown and hyaline
spotted on most of surface. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

Sc R1
R2+3

cup R4+5

M1

CuA1
CuA2 A1+CuA2

16 Sphecomyiella 17 Leptopyrgota

18 Undescribed Genus A 19 Pyrgota

20 Idiopyrgota 21 Anapyrgota

22 Stenopyrgota 23 Undescribed Genus C

Figs.  69.16–23. Wings: dorsal view of (16)  Sphecomyiella valida (Harris), (MND, fig.  65.2); (17)  Leptopyrgota undulata Becker;
(18)  Undescribed Genus  A; (19)  Pyrgota lugens Wulp; (20)  Idiopyrgota  sp.; (21)  Anapyrgota  sp.; (22)  Stenopyrgota  sp.; and
(23) Undescribed Genus C. Figures 17–23 illustrated by V. Hernández-Ortiz.
960
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

9. Robust flies; wing pattern with extensive large brownish dark areas, not mottled; vein R2+3
usually with stump vein in apical one-third (Fig. 19); parafacial and genal groove broad and
usually wrinkled (Fig. 5) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pyrgota Wiedemann
– Slender flies; wing pattern pale brown and hyaline spotted throughout surface; vein  R 2+3
without stump veins (Fig. 16); parafacial and genal groove short and smooth (including Stirothrinax
Enderlein). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sphecomyiella Hendel

10. Female ovipositor sheath without apicoventral unguis; wing pattern with numerous hyaline spots;
legs of normal length. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Boreothrinax Steyskal
– Female ovipositor sheath with apicoventral hook-shaped unguis well developed; wing pattern
mostly pale brown (Fig. 23); legs long and slender. . . . . . . . . . . . . . . . . . . . Undescribed Genus C

Synopsis of the fauna but the present study added three undescribed species from
Costa Rica and Mexico.
The pyrgotid fauna south of the United States includes
around 59 known species (Aczél 1956a, 1956b, 1956c; Stey- Neopyrgota Hendel. Two species of this genus are known
skal 1967, 1978; Kondratieff  & Fitzgerald 1993; Bernardi to occur in Costa Rica, N. major Hennig and N. picea Hen-
1991), but only 11 species have been recorded for Central nig. An additional undescribed species from that country was
America. The present study revealed that approximately 22 also encountered.
other undescribed species and three undescribed genera are
Pyrgota Wiedemann. Five species are known in this ge-
present in the region.
nus, including two found in Central America. Pyrgota lugens
Most of the genera occurring in the Americas were found
Wulp was originally described from Guatemala, but there is
in Central America. Only the North American genus Pyr-
gotella Curran, and the tropical genera Lopadops Enderlein additional material from southern Mexico, and P.  longipes
(Argentina), Tropidothrinax Enderlein (Bolivia), and Car- Hendel is known from Guatemala and Brazil.
rerapyrgota Aczél (Brazil, Paraguay, and Argentina) were
Sphecomyiella Hendel. This genus includes two known
not recorded. Most Central American genera were found in
species that occur in the USA, S. valida Harris and S. nelsoni
Mexico (10) and Costa Rica (8). Despite the great number of
Kondratieff  & Fitzgerald. The specimens examined in this
undescribed species so far uncovered in the family, I expect
that further collecting will show that the pyrgotids are even study revealed the presence of S. valida in central Mexico.
more diverse in the region. Within Sphecomyiella, the following species of Stirothrinax
Enderlein should also be included: S.  cribratus Enderlein,
Anapyrgota Steyskal. Only one species, A.  personta recorded from Mexico and Nicaragua, and S. knudseni May-
(Lutz & Lima), from Brazil has been recognized in this ge- er from Costa Rica. Additional specimens examined suggest
nus. Several specimens examined in this study represent a the presence of three undescribed species: one from Mexico
second undescribed species from Panama. and two others from Costa Rica.
Boreothrinax Steyskal. Three species have been re- Stenopyrgota Malloch. The genus currently comprises
corded from the USA, of which B. dichaetus Steyskal is also two species, S. mexicana Malloch from Mexico and S. cras-
present in Mexico (Durango). In addition, two undescribed sitibia Aczél known from Argentina. Studied specimens in-
Mexican species were examined from Querétaro, Guerrero, dicate that three other unnamed species occur in Costa Rica
and Oaxaca. and Mexico.
Idiopyrgota Aczél. This is a monotypic genus represented Undescribed Genus A. This genus comprises at least two
by I.  setiventris Aczél from Brazil, but at least three other new species, one represented by specimens from Mexico and
species occur in Mexico, Guatemala, Costa Rica, Panama, Guatemala, and the other from Costa Rica.
the Bahamas, and Ecuador.
Undescribed Genus B. This genus is represented by
Leptopyrgota Hendel. In the catalog of the Pyrgotidae three undescribed species all occurring in Costa Rica, but
south of the USA, Steyskal (1967) recognized 12 species, two of which are also present in Mexico.
and more recently Bernardi (1991) described 21 Brazilian
species, most of them on the basis of only one specimen. In Undescribed Genus C. One undescribed species is in-
Central America, two species were previously recorded from cluded in this genus, represented by a large series of speci-
Costa Rica, L. amplipennis Hendel and L. undulata Becker, mens from Mexico, Costa Rica, and Panama.
PYRGOTIDAE (pyrgotid flies) 69 961

Literature cited

Aczél, M. 1956a. Revisión parcial de las Pyrgotidae neotropicales Bulletin of the Illinois Agricultural Experimental Station 116:
y antárticas, con sinopsis de los géneros y especies (Diptera, 447–480.
Acalyptratae), Parte  I. Revista Brasileira de Entomologia 4: Gardner, T.R. & L.B. Parker. 1942. Investigations of the parasites
161–184. of Popillia japonica and related Scarabaeidae in the Far East
Aczél, M. 1956b. Revisión parcial de las Pyrgotidae neotropicales y from 1929 to 1933 inclusive. Review of Applied Entomology
antárticas, con sinopsis de los géneros y especies (Diptera, Aca- (series A) 30: 51–52.
lyptratae), Parte II. Revista Brasileira de Entomologia 5: 1–70. Kondratieff, B.C.  & S.C. Fitzgerald. 1993. A new species of
Aczél, M. 1956c. Revisión parcial de las Pyrgotidae neotropicales Sphecomyiella (Diptera: Pyrgotidae) from North America.
y antárticas, con sinopsis de los géneros y especies (Diptera, Proceedings of the Entomological Society of Washington 95:
Acalyptratae), Parte  III. Revista Brasileira de Entomologia 6: 554–557.
1–38. McAlpine, D.K. 1978. Notes on nomenclature and type-speci-
Aczél, M. 1958. Pyrgotidae (Diptera, Acalyptrata). Exploration du mens of Australian Pyrgotidae (Diptera, Schizophora). Austra-
Parc National de l’Upemba Mission G.F. de Witte (1946–1949), lian Entomological Magazine 5: 27–33.
Brussels 50: 3–53. McAlpine, D.K. 1989. The taxonomic position of the Ctenostyli-
Bernardi, N. 1990. Caracterização do gênero Carrerapyrgota Ac- dae (=Lochmostyliinae; Diptera: Schizophora). Memorias do
zél, 1956 e sinonimização de Anapyrgota Steyskal, 1967 (Dip- Instituto Oswaldo Cruz 84: 365–371.
tera, Pyrgotidae). Biotemas 3: 101–104. Steyskal, G.C. 1967. 56. Family Pyrgotidae. In Papavero, N. (edi-
Bernardi, N. 1991. Notas sobre o gênero Leptopyrgota Hendel, com tor). A Catalogue of the Diptera of the Americas South of the
descrição de vinte e uma novas espécies do Brasil (Diptera, Pyr- United States. Departamento de Zoologia, Secretaria da Agri-
gotidae). Revista Brasileira de Zoologia 7: 327–350. cultura, São Paulo, 8 pp.
Crocker, R.L., L.A. Rodríguez-del-Bosque, W.T.  Nailon, Jr.,  & Steyskal, G.C. 1972. African Pyrgotidae (Diptera) collected by
X. Wei. 1996. Flight periods in Texas of three parasites (Dip- Willi Richter, Erwin Lindner, and others. Stuttgarter Beiträge
tera: Pyrgotidae) of adult Phyllophaga spp. (Coleoptera: Scara- zur Naturkunde 238: 1–11.
baeidae), and egg production by Pyrgota undata. Southwestern Steyskal, G.C. 1978. Synopsis of the North American Pyrgotidae
Entomologist 21: 317–324. (Diptera). Proceedings of the Entomological Society of Wash-
Davis, J.J. 1919. Contributions to a knowledge of the natural en- ington 80: 149–155.
emies of Phyllophaga. Bulletin of the Illinois Natural History Steyskal, G.C. 1987. Pyrgotidae, pp. 813–816. MND, Volume 2.
Survey 13: 53–138. Wolcott, G.N. 1923. Insect parasite introduction in Porto Rico.
Forbes, S.A. 1908. On the life history, habits, and economic re- Journal of the Department of Agriculture of Porto Rico. Review
lations of the white-grubs and may-beetles (Lachnosterna). of Applied Entomology (series A) 11: 60–61.
962
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
CTENOSTYLIDAE (ctenostylid flies) 70
Valery A. Korneyev

Fig. 70.1. Female of Lochmostylia borgmeieri Hendel, (illustrated by V. Korneyev).

Diagnosis ternum, setulose anepimeron, and large swollen anatergite.

Thoracic spiracles circular, almost devoid of hairs on mar-
Medium-sized to large (body length 3.5–13.0 mm) noc- gins. Legs with short setulae only, without setae; midtibia
turnal, long-legged flies with short yellow (often red or without apicoventral spur. Abdominal syntergite 1+2 more
black spotted) body and patterned wings (Fig. 1). Ocelli and or less petiolate. Sternites  1 and 2 at least partially fused.
proboscis, and often palpus lacking. Head large, almost as Sternites 5–6 indistinguishable, and pleural membrane setu-
bulky as thorax, sexually dimorphic: with arista dendritical- lose in abdomen of Ctenostylum (Fig.  25). Male with five
ly branched and frons wide in females; arista simple, short abdominal spiracles and sternite 8 large; in Ctenostylum and
trichose, and frons often narrowed in males. Wing with only Lochmostylia postabdomen symmetrical, with additional
subcostal break of costa; vein Sc obsolete in apical one-third, setulose tergite and sternite (possibly the sixth). Male geni-
vein R1 setulose, cell bm open anterobasally, cell bcu without talia simple, with strongly reduced hypandrium and vanes
posteroapical lobe, vein A2 usually distinct distad of alula, at of phallapodeme; phallus simple, in Ramuliseta Keiser, Fur-
least as a fold. Thorax short, wide, with large notopleuron ciseta, and Nepaliseta Barraclough, long and coiled in pocket on
bearing single seta (in Lochmostylia lopesi Keiser with two right dorsal side of abdomen, in Furciseta bearing weak mem-
unequal setae) and two to three setulae, with short anepis- branous glans (Figs. 19–23), in Ctenostylum and Lochmostylia

963
964
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

short and pipelike (Figs. 28–29). Females, as in other Tephri- Ctenostylids are usually recognized by the subequal size
toidea, with telescopic ovipositor with sclerotized oviscape, of the head and thorax, their long slender legs, and patterned
or tergosternum  7, consisting of narrow tergite and large, wings. They are readily distinguished from all other fami-
setulose, and convex sternite (Fig. 25); eversible membrane lies by the lack of both a proboscis and ocelli. Long-legged
finely microtrichose, without taeniae; aculeus reduced to Pyrgotidae (Leptopyrgota Hendel, Toxopyrgota Hendel), in
non-sclerotized tube (tergosternite  8), with short and blunt which ocelli are also lacking, have a functional proboscis,
cerci dorsoapically (Figs. 30–32). No spermathecae evident.
though in Toxopyrgota it is small. Agromyzidae, which have
Viviparous or ovoviviparous, vagina usually full of first in-
star larvae or eggs with larvae (Fig. 32). a similarly reduced apical section of Sc and basal section of

2 Ctenostylum 3 Tauroscypson 4 Lochmostylia

5 Ctenostylum 6 Tauroscypson 7 Lochmostylia

8 Furciseta 9 Furciseta 10 Tauroscypson

Figs. 70.2–10. Heads and antennae: anterior view of head of (2) Ctenostylum sp. near rufum Macquart, (redrawn from McAlpine, 1990,
fig. 7); (3) Tauroscypson andina Aczél, (South America, Aczél, 1956, fig. 94); and (4) Lochmostylia borgmeieri Hendel, (Keiser, 1951, fig. 1);
lateral view of head of (5) C. rufum, (South America); (6) T. guiana Curran, (South America, redrawn from Curran, 1934, fig. 52); (7) L. lopesi
Keiser, (South America); and (8) Furciseta plaumanni (Hennig), (South America); antenna of (9) F. plaumanni, (redrawn from Hennig, 1952,
fig. 6); and (10) T. andina Aczél, (redrawn from Aczél, 1956, fig. 96). Figures 5, 7, and 8 illustrated by V. Korneyev.
CTENOSTYLIDAE (ctenostylid flies) 70 965

M, differ in having hyaline wings, dorsally setulose tibiae, species. Korneyev (2001) suggested that if Ctenostylidae
and a normally developed proboscis and ocelli. belong in the Tephritoidea, they are related to the Platys-
tomatidae  + Pyrgotidae  + Tephritidae group of families.
Further studies of the male terminalia in Furciseta have
Biology confirmed that Ctenostylidae, indeed, belong in this group
of families, but further relationships remain vague, though
Unknown. Adults of Ramuliseta ashleyi Barraclough the most probable related group is Pyrgotidae (V.A.  Kor-
were collected shortly after dusk at a mercury vapor lamp by neyev, in preparation). At present, the family includes 12
A. Kirk-Spriggs (Barraclough, 1998). described species and seven nominal genera: Ctenostylum
(Neotropical; monotypic), Lochmostylia (Neotropical;
Classification two species), Tauroscypson (Neotropical; two species), Ra-
muliseta (Paleotropical; three species), Furciseta (Neotro-
Ctenostylidae were usually treated as the subfam- pical; monotypic), Nepaliseta (Oriental; two species), and
ily Lochmostyliinae of Pyrgotidae (Hendel, 1934; Lopes, the monotypic Oriental genus Sinolochmostylia Yang.
1935; Aczél, 1956; Steyskal, 1968). McAlpine (1990)
excluded them from the latter family and advanced argu- Identification
ments against inclusion of Ctenostylidae in the superfam-
ily Tephritoidea. Barraclough (1994, 1995, 1998), Yang Aczél (1956) provided keys to the species of the New
(1995), and Korneyev (2001) reviewed the Old World World genera Lochmostylia and Tauroscypson.

Key to the genera of Ctenostylidae of the New World

1. Cell r1 with additional crossveins between C and R1 veins (Figs.  11, 12, 15, 16); lunule deep,
setulose (Figs. 2, 3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Cell r1 without crossveins (Figs. 13, 14, 17, 18); lunule narrow, transverse, bare (Fig. 4). . . . . . 3

2. Palpus, alula and anal lobe absent; male head holoptic; frons narrow; arista pectinate (Figs. 2, 5).
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ctenostylum Macquart
– Palpus present; alula and anal lobe developed; male head dichoptic, eyes not approximated; frons
not narrowed (Figs. 3, 6); arista bushlike, branching (Fig. 10); Guyana, Peru, & Bolivia. . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Tauroscypson Curran]

3. Palpus absent (Figs. 4, 7); veins R2+3 and R4+5 at least slightly convergent; vein M distinct in distal
half of cell bm (Figs. 13, 14); wing 8–12 mm long; male head dichoptic, eyes not approximated;
frons not narrowed; female arista with five to seven primary branches, bushlike, male arista sim-
ple. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lochmostylia Hendel
– Palpus present (Fig.  8); veins R2+3 and R4+5 strongly divergent; vein  M absent in distal half of
cell bm (Figs. 17, 18); wing 3.5–4.5 mm long; male head holoptic, eyes approximated; frons
narrow; female arista basally bifurcated (Fig. 8), male arista simple (Fig. 9). . . . . Furciseta Aczél

Synopsis of the fauna Furciseta Aczél. Originally known only from southeast-
ern Brazil, this genus has also been collected in Venezuela
Six species in four genera of Ctenostylidae occur in the and southern Mexico (San Luis Potosi) (D. Yanega, personal
Neotropical Region; three genera and three species are rec- communication).
ognized in collections from Central America.
Lochmostylia Hendel. There are two species of this ge-
Ctenostylum Macquart. This genus includes the type nus, differing in wing pattern and venation; the type species
species C.  rufum Macquart from Brazil and Venezuela L. borgmeieri (Figs. 1, 4, 7, 13, 24, 26–28) is represented by
(Fig. 12); specimens from Costa Rica (Figs. 11, 29) apparently specimens from Costa Rica, and L. lopesi (Fig. 14) is known
belong to an undescribed species, showing minor differences
from Brazil.
in wing venation.
966
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

R1 r1 Sc R1 r1

R2+3

bm

cup R4+5

A1+CuA2 M
CuA1

11 Ctenostylum 12 Ctenostylum

Sc R1 r1
R2+3

R4+5

A2

13 Lochmostylia 14 Lochmostylia

15 Tauroscypson 16 Tauroscypson

R2+3
Sc R1

R4+5

17 Furciseta 18 Furciseta

Figs. 70.11–18. Wings: dorsal view of (11) Ctenostylum sp. near rufum Macquart; (12) C. rufum, (South America); (13) Lochmostylia
borgmeieri Hendel; (14) L. lopesi Keiser, (South America); (15) Tauroscypson guiana Curran, (South America, redrawn from Curran, 1934,
fig. 51); (16) T. andina Aczél, (South America, redrawn from Aczél, 1956, plate 5, fig. T); and male (17) and female (18) of Furciseta plau-
manni (Hennig), (South America). Figures 11–14, 17, and 18 illustrated by V. Korneyev.
CTENOSTYLIDAE (ctenostylid flies) 70 967

ej apod

phapod

hypd
cerc sur epand

ph

19 Furciseta 20 Furciseta

ph

cerc sur epand

hypd

phapod

21 Furciseta 22 Furciseta

Figs. 70.19–22. Male terminalia: right lateral view, apex of phallus removed (19), apex of phallus (20); ventral view, apex of phallus
removed (21), and ejaculatory apodeme (22) of Furciseta plaumanni (Hennig), (South America). Figures 19–22 illustrated by V. Korneyev.
Abbreviations: cerc, cercus; ej apod, ejaculatory apodeme; epand, epandrium; hypd, hypandrium; ph,  phallus; phapod, phallapodeme;
sur, surstylus.
968
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

st 1
st 1

st 2 st 2

tg 3
st 5
tg 3

st 8 st 5
ph
st 6
tg 6
23 Furciseta

24 Lochmostylia
25 Ctenostylum
cerc

tg 6
epand
phapod

tgst 7
sur
phapod

hypd
ph st 6

ph

26 Lochmostylia 27 Lochmostylia 28 Lochmostylia

ev memb

epand

cerc tgst 8
cerc
sur
31 Furciseta

29 Ctenostylum
cerc
cerc

30 Ctenostylum 32 Furciseta

Figs.  70.23–32. Abdomens and terminalia: ventral view of male abdomen of (23)  Furciseta plaumanni (Hennig), (South America); and
(24) Lochmostylia borgmeieri Hendel; dorsal view of female abdomen of (25) Ctenostylum sp., near rufum Macquart; ventral view (26) and
lateral view (27) of male terminalia, and lateral view of hypandrium and phallus (28) of L. borgmeieri; lateral view of male terminalia of
(29) Ctenostylum sp., near rufum Macquart; dorsal view of female terminalia of (30) Ctenostylum sp., near rufum Macquart; dorsal view of
cerci (31) and lateral view of female terminalia (32) of F. plaumanni. Figures 23–32 illustrated by V. Korneyev.
Abbreviations: cerc, cercus; epand, epandrium; ev  memb, eversible membrane; hypd, hypandrium; phapod, phallapodeme; ph,  phallus;
sur, surstylus; st, sternite; tg, tergite; tgst, tergosternite.
CTENOSTYLIDAE (ctenostylid flies) 70 969

Literature cited

Aczél, M.L. 1956. Revisión parcial de las Pyrgotidae neotropicales Keiser, F. 1951. Die Unterfamilie der Lochmostyliinae (Dipt.,
y antárticas, con sinopsis de los géneros y especies (Diptera, Pyrgotidae) nebst beschreibung einer neuen Gattung und
Acalyptratae). Revista Brasileira de Entomologia 6: 1–38. Art aus dem indo-australischen Faunengebiet. Mitteilun-
Barraclough, D.A. 1994. A review of Afrotropical Ctenostylidae gen der Schweizerischen Entomologischen Gesellschaft 24:
(Diptera: Schizophora: ?Tephritoidea), with redescription of 113–124.
Ramuliseta lindneri Keiser, 1952. Annals of the Natal Museum Korneyev, V.A. 2001. New records of Oriental Ctenostylidae
35: 5–14. (Diptera Acalyptrata), with discussion of the position of the
Barraclough, D.A. 1995. Nepaliseta mirabilis, a remarkable new family. Vestnik zoologii 35: 47–60.
genus and species of Ctenostylidae (Diptera: Schizophora) from Lopes, H.S. 1935. Um novo gênero de Diptera acalyptratae e
Nepal. Annals of the Natal Museum 36: 135–139. considerações sobre a família Ctenostylidae. Archivos do In-
Barraclough, D.A. 1998. The missing males of Ramuliseta Keiser stituto de biologia vegetal 2: 247–253.
(Diptera: Schizophora: Ctenostylidae). Annals of the Natal Mu- McAlpine, D.K. 1990. The taxonomic position of the Ctenostyli-
seum 39: 115–126. dae (=Lochmostyliinae: Diptera: Schizophora). Memorias do
Curran, C.H. 1934. The Diptera of Kartabo, Bartica District, British Instituto Oswaldo Cruz 84, Suplemento IV: 365–371. (1989)
Guiana. Bulletin of the American Museum of Natural History Steyskal, G.C. 1968. 56. Family Pyrgotidae. In Papavero,  N.
66: 287–533. (editor). A Catalogue of the Diptera of the Americas South of
Hendel, F. 1934. Übersicht über die Gattungen der Pyrgotiden, the United States. Departamento de Zoologia, Secretaria da
nebst Beschreibung neuer Gattungen und Arten. Encyclopédie Agricultura, São Paulo, 8 pp.
Entomologique. (B) II. Diptera 7: 141–156. Yang, J. 1995. Diptera: Pyrgotidae, pp.  247–249. Insects and
Hennig, W. 1952. Bemerkenswerte neue Acalyptraten in der Sam- macrofungi of Gutianshan, Zhejiang. Zhejiang Science
mlung des Deutschen Entomologischen Institutes (Diptera: and Technology Publishing House, Hangzhou, 318  pp. [In
Acalyptrata). Beiträge zur Entomologie 2: 604–618. Chinese.]
970
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
LAUXANIIDAE (lauxaniid flies) 71
Stephen D. Gaimari and Vera C. Silva

Fig. 71.1. Male of Minettia lupulina (Fabricius), (Nearctic, MND, fig. 87.1).

Diagnosis (Fig.  44), or continued to R4+5 in Homoneurinae (Fig.  43),

although fine hairs often continue around wing beyond
Small to moderately large (body length 2–11 mm) flies; spinules; Sc complete, separated from R1 (Figs.  43–52);
compact bodied (Fig. 1) or elongate; varied in color patterns, cell cup small with rounded apex; A1 usually short (long in
often with marks, spots, bands, or reticulating patterns. Head Eurychoromyiinae), not reaching wing margin; membrane
with postocellar setae convergent to cruciate, rarely absent; various, from hyaline to darkened crossveins to intricately
ocellar setae proclinate, divergent to parallel, small and patterned, irrorated, or banded. Legs each with preapical
hairlike to strong; orbital plate with two fronto-orbital se- dorsal tibial seta (except in some Eurychoromyiinae). Abdo-
tae (rarely zero to one), usually reclinate, but anterior seta men short and stout to elongate and tapering; segment 7 in
sometimes slightly (Fig. 23) to strongly (Fig. 27) inclinate; female usually normal (Fig.  55), but occasionally forming
lunule small, narrowly exposed and bare; face various, from complete sclerotized ring; sternite 8 often variously modified
slightly concave or flat to convex or bulging; subgena usually (e.g., Figs. 53, 54). Some genera display sharp reductions or
thin and inconspicuous, but occasionally enlarged and bulg- losses of setae.
ing; vibrissa absent. Scutum setulose, not hairy, occasionally Eggs white, ovoid in dorsal view, sometimes banana-
nearly bare; with two to four dorsocentral setae (rarely only shaped in lateral view or flattened on dorsal surface; an-
teriorly with distinct tubercle shielding apical micropyle;
one), either posteriorly located (0+2, 0+3) (Figs. 33, 34) or
posteriorly with knoblike swelling or constricted tubercle;
evenly distributed with one presutural (1+3, 1+2) (Fig. 35); chorion with distinct longitudinal ridges (often connected by
prescutellar acrostichal seta usually present, sometimes with transverse septa, occasionally with spicules along ridges),
additional acrostichal setae in row (Fig. 35); scutellum with and often with pitted or reticulated patterns at each pole.
two pairs of setae; anepisternum setulose, with one strong Mature larvae (Fig. 56) musciform, tapering slightly an-
seta near posterior margin; katepisternum setulose, with teriorly and less so posteriorly, often compressed dorsoven-
one to two setae dorsally. Wing with C unbroken, weakened trally; oral area with multiple rows of spicules (Fig. 59); all
beyond R4+5 and ending at M1, with stout black spinules on segments with bands of spinules anterodorsally, and seg-
anterodorsal surface ending slightly beyond apex of R2+3 ments 3–12 with similar bands of spinules anteroventrally, in

971
972
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

addition to ventral creeping welts; often with other spinules phytophagous existence. Adults are fungal grazers on leaves
on body surface and/or near anal plate; segment 12 with up (Broadhead, 1984, 1989; Mello & Silva, 2006) and display
to three pairs of fleshy lobes or tubercles dorsolaterally, later- related mouthpart modifications. Attesting to their biologi-
ally, and ventrolaterally (Fig. 60); anterior spiracles arranged cal importance, lauxaniids are probably among the more
fanlike with 4–14 lobes at end of spiracular stalk (Fig. 57), important dipterans responsible for turnover of enormous
or with 11–19 fingerlike lobes in cluster at end of stalk; pos- quantities of organic litter in the tropics, likely playing a sig-
terior spiracles on short, separate, usually darkened and pos- nificant role in the healthy functioning of tropical ecosys-
teriorly directed tubular mountings on segment 12 (Fig. 60), tems (Freidberg & Yarom, 1990). In addition, despite their
with each spiracular plate circular and bearing three ovoid generalization as saprophages, there are many examples
to elongate oval slits in radiating pattern from slightly diver- of specialization to different media and feeding strategies.
gent (Fig. 58) to right angled, and with four 6–20 branched Miller (1977a) summarized the known larval biologies for
spiracular hairs (Fig.  58). Cephalopharyngeal skeleton 70  species in 21  genera  & subgenera, reporting that more
(Fig.  59) with mandible slender to stout, with or without than half bred in fallen leaves of deciduous trees (=leaf litter).
accessory tooth, but usually with small basal window; dental Of these, only Homoneura (three subgenera) (Miller, 1977a;
sclerite and ventral pharyngeal ridge present; labial sclerite Hardy & Delfinado, 1980; Sasakawa & Ikeuchi, 1982) and
arched, with slender posteriorly directed rodlike rami medial Tricholauxania Hendel (Meijere, 1906) appear restricted to
to parastomal bars; parastomal bars present, small, not fused this medium. Interestingly, among leaf litter feeders, there
apically or to epipharyngeal sclerite; tentoropharyngeal and is a subspecialization to feed in decaying mesophyll tissues
hypopharyngeal sclerites separate, with left and right hypo- between the upper and lower epidermis, as leaf miners of
pharyngeal sclerites connected by dorsal bridge; dorsal cornu fallen leaves (Hering, 1951; Miller, 1977a; Ferrar 1987). Of
slender, usually shorter and narrower than ventral cornu; the relative minority of non-leaf litter feeders, several gen-
ventral cornu often with window along upper margin. era and subgenera have species that are nidicolous, feeding
Puparia ovoid, tapering slightly more posteriorly than on decaying matter in bird nests, including Pseudocalliope
anteriorly; convex dorsally, flattened ventrally; anterior end and Poecilominettia (Hicks, 1953; Miller  & Foote, 1975),
dorsoventrally flattened to produce lateral ridge; surface the subgenus Minettia (Frendelia) (Armstrong, 1953), the
often covered with white calcareous deposit secreted from subgenus Meiosimyza (Lyciella) (Thienemann, 1926; Miller,
anus prior to pupariation. 1977a), “Sapromyza” (Colless  & McAlpine, 1970), and
Families superficially similar to lauxaniids (e.g., some Calliopum (Collin, 1933; Miller 1977a), or within mam-
Drosophilidae) can be readily separated from them by the malian nests, such as Minettia (Séguy, 1946), Sapromyza
following lauxaniid characteristics: two  fronto-orbital setae (Lyayster, 1967), and Pseudogriphoneura (Miller, 1977a).
(rarely zero to one), both usually reclinate, but anterior seta Other known feeding sites include rotting fungus (Chan-
inclinate in some genera; vibrissa absent (although subvibris- dler, 1978), under bark of rotting tree stumps (Meijere,
sal setae can be enlarged near oral margin in some genera, 1909; Collin, 1948; Laurence, 1952), and within rotting
such as Hypagoga, Trigonometopus, and Oncodometopus); gymnosperm cones (Keen, 1958; Miller, 1977a) and club
ocellar setae proclinate; postocellar setae convergent; anepis- tops (Miller, 1977a). Just before pupariation, larvae of
ternum with at least one strong seta on posterior margin; some species first excrete their gut contents and then cover
C unbroken; Sc complete, separated from R1; presence of themselves with a calcareous anal secretion originating
costal spinules ending slightly beyond R2+3 (ending at R4+5 from large extensions of the Malpighian tubules that hard-
in Homoneurinae, with only one genus in the Neotropical ens after being smeared over the body surface (Miller  &
Region, but common in the Nearctic). From the most simi- Foote, 1975). At least one species of Pseudocalliope in-
lar family, the closely related Chamaemyiidae, they can be stead forms a mud-and-fiber cocoon around its puparium
separated on the basis of the prescutellum being absent, the (Miller & Foote, 1975).
presence of preapical dorsal tibial setae, and the presence of Morphology of the immature stages is known for few
distinct black costal spinules; some genera of chamaemyiids species, summarized in part by Ferrar (1987). Foote (1991)
share the presence of reclinate fronto-orbital setae and an suggested that many larval lauxaniids are difficult if not im-
anepisternal seta. possible to separate from other saprophagous acalyptrates.
Details and extensive figures are provided for all stages
Biology and instars for species (none Neotropical) in the following
genera: Camptoprosopella, Homoneura (Tarsohomoneura),
Larval lauxaniids are widely documented as saprophages Poecilolycia, Minettia (Minettia), Minettia (Frendelia), Po-
(Miller, 1977a), feeding in a diverse array of decaying plant ecilominettia, Pseudocalliope, and Melanomyza (Melano-
matter, likely consuming associated microorganisms, fungi, myza) (Miller & Foote, 1976). The eggs are described and
bacteria, or yeasts (Miller  & Foote, 1976). Silva  & Mello illustrated for two species of Eurychoromyiinae: Tauridion
(2008) also record one species, Physoclypeus farinosus shewelli Papp  & Silva and Physegeniopsis albeto Gaim-
Hendel, feeding in flower heads of Asteraceae in Brazil; al- ari  & Silva (Gaimari & Silva, 2010). Few other studies
though saprophagy could not be ruled out, the data suggest a have dealt with morphology of lauxaniid immatures, so
LAUXANIIDAE (lauxaniid flies) 71 973

scp

flgm 1

2 Melanomyza 3 Neodeceia 4 Trivialia

i vt s
p frorb s
poc s
a frorb s o vt s

7 Trigonometopus

5 Xenochaetina 6 Physoclypeus

8 Pseudocalliope 9 Xeniconeura 10 Oncodometopus

Figs. 71.2–10. Heads: lateral view of (2) Melanomyza (Melanomyza) gracilipes (Loew), (Nearctic, MND, fig. 87.2); (3) Neodeceia
cineracea (Coquillett), (Nearctic, MND, fig.  72.3); (4)  Trivialia fuscocapitata (Malloch), (Nearctic, MND, fig.  87.4); (5)  Xenochaetina
muscaria (Loew), (Nearctic, MND, fig. 87.5); (6) Physoclypeus coquilletti (Hendel), (Nearctic, MND, fig. 87.6); (7) Trigonometopus sp.,
(Nearctic, MND, fig. 87.7); (8) Pseudocalliope flaviceps (Loew), (Nearctic, MND, fig. 87.8); (9) Xeniconeura costalis (Curran), (Nearctic,
MND, fig. 87.10); and (10) Oncodometopus umbrosus (Loew), (Nearctic, MND, fig. 87.35).
Abbreviations: a frorb s, anterior fronto-orbital seta; flgm 1, first flagellomere; i vt s, inner vertical seta; o vt s, outer vertical seta; poc s,
postocellar seta; p frorb s, posterior fronto-orbital seta; scp, scape.
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

there is still much to learn, as this represents a small subset the presence of costal setulae extending to the apex of wing
of the known genera in the family. vein R4+5. Stuckenberg (1971) also noted that most species in
this subfamily possess an anteroventral ctenidium (=comb)
on the forefemur, while few non-homoneurines display this
Classification state. Shewell (1977) later added the presence of one or two
extra midtibial spurs (actually not true spurs but stout apical
Frey (1941) was the first to recognize subfamilial divi- setae) as further support for the subfamily, also proposing the
sions of the family, adding Homoneurinae and Minettiinae, paraphyletic subfamily Lauxaniinae for the remainder of the
but the subfamilial names were without descriptions. Stuck- family, based on costal setulae ending just beyond the apex
enberg (1971) validly proposed the subfamily Homoneurinae of vein R2+3 and a single midtibial spur. The conditions cited
for a (potentially) monophyletic group of genera based on for Lauxaniinae are clearly plesiomorphic and are likely part

i vt s
p frorb s poc s oc s poc s
a frorb s
o vt s

flgm 1

11 Neogriphoneura 12 Trisapromyza 13 Lauxania

14 Allominettia 15 Mallochomyza 16 Poecilolycia

Figs. 71.11–16. Heads (continued): lateral view of (11) Neogriphoneura sordida (Wiedemann), (Nearctic, MND, fig. 87.11); (12) Trisapromyza
vittigera (Coquillett), (Nearctic, MND, fig. 87.12); (13) Lauxania sp., (Nearctic, MND, fig. 87.13); (14) Allominettia geniseta (Malloch),
(MND, fig. 87.39); (15) Mallochomyza citreifrons (Malloch), (Nearctic, MND, fig. 87.40); and (16) Poecilolycia serrata (Malloch), (Nearctic,
MND, fig. 87.37).
Abbreviations: a frorb s, anterior fronto-orbital seta; flgm 1, first flagellomere; i vt s, inner vertical seta; oc s, ocellar seta; o vt s, outer vertical
seta; poc s, postocellar seta; p frorb s, posterior fronto-orbital seta.
LAUXANIIDAE (lauxaniid flies) 71 975

of the groundplan for the entire Muscomorpha (McAlpine, showed that many of the setae thought to be “lost” are pres-
1989), and both Stuckenberg (1971) and Shewell (1977) ent but greatly reduced, and the costal spinules are hairlike,
acknowledged the likely paraphyly for the group. Within but are distinguishable from the standard wing vestiture.
Homoneurinae, Kim (1994) presented phenetically based Although lauxaniids are uncommon in the fossil record,
dendrograms for the Australian members, resulting in sev- several species have been described from Eocene/Oligocene
eral new genera, but it must be noted that monophyly was Baltic and Chinese amber (Hennig, 1965; Hong, 1981). In
apparently not a criterion in their definitions and in their addition, no less than 15 new species from 24 specimens in
separation from Homoneura or other genera. No additions or Dominican amber are being described by the authors, all in
changes to the suprageneric classification were proposed. extant New World genera.
Although Lauxaniidae is among the largest families of
Acalyptratae, the internal classification of the family is in
its infancy. The monophyly of the family is supported by
Identification
several autapomorphies (McAlpine, 1989), including the
male accessory glands forming a dense tangle due to repeated The literature for identifying the described species of
branching (Sturtevant, 1925, 1926), the phallus being reduced Lauxaniidae is sparse, and many species remain unde-
to a rigid tubular process, and the lunule being unexposed. scribed worldwide, particularly in the Old and New World
Within the Lauxanioidea, Lauxaniidae is clearly separated tropics. There are few modern synthetic works at the spe-
from Chamaemyiidae, but the monophyly of the family rela- cies level. For genera, the most significant work was the
tive to Celyphidae and Eurychoromyiidae has been conten- monumental review of all Old World genera by Stuckenberg
tious. The latter was revised to subfamily status (Gaimari & (1971), including a key to the genera. Regionally, keys to
Silva, 2010), with a thorough discussion of the history of genera and general reviews were provided for the Nearctic
its classification. The relationships of Eurychoromyia had Region by Shewell (1987) and for the Palearctic Region by
been previously obscured by the apparent loss of setae (and Papp & Shatalkin (1998). Shatalkin (2000) further provid-
costal spinules of the wing), presence of four spermathecae, ed an overview of all Palearctic genera, with keys to the de-
and reductions in features, which are of great importance to scribed species of the region; Schacht et al. (2004) provided
understanding acalyptrate relationships. The evidence sug- updates and translation into English for these keys. For the
gesting that Eurychoromyiidae is subordinate within the Nearctic Region, Miller (1977b, c) revised the fauna of the
Lauxaniidae is based upon Gaimari & Silva’s (2010) assess- large genus Homoneura, and Pérusse & Wheeler (2000) re-
ment of the clear relationship of Eurychoromyia with sev- vised the fauna of Lauxania. For the Oriental Region, the
eral unequivocal lauxaniid genera, including Tauridion and many works of Sasakawa (e.g., 1992, 2001) and colleagues
Physegeniopsis, based on male and female genitalic features, (e.g., Sasakawa & Ikeuchi, 1982) have provided faunistic
many characteristics of the head and wing, and presence of and revisionary studies, particularly for the Homoneuri-
two pairs of spermathecae (most other known lauxaniids nae, in this highly species-rich region. In Australia, Kim
have spermathecae in 1+2 configuration). The genus Chory- (1994) revised the fauna of Homoneurinae, providing keys
euromyia, although known only from one female specimen, to the genera and species of the region. For the Neotropi-
is so remarkably similar to Eurychoromyia that its sister- cal Region, the current work is the first key to all genera
group relationship appears likely. Other evidence includes of the New World, including several recently described.
the modifications of the adult labellum for fungal grazing A conspectus of Neotropical lauxaniids is in progress by
(see Broadhead, 1984), such as the nonbifurcating labellar the authors, including a catalog to all species in the region
canals that direct fungal spores and hyphae directly into the and descriptions for all genera keying out here as “Unde-
oral opening. In this regard, the gut of dissected specimens scribed Genus A–S”. Gaimari & Silva (2010) revised and
of several of the included genera of Eurychoromyiinae have provided a key for all species of Eurychoromyiinae, with
been found full of entire fungal spores and hyphae, and with one additional genus with three species yet to be described
the similar mouthpart modifications, indicate the same adult (Undescribed Genus S). In the following key, all genera are
feeding habits as other lauxaniids. In addition, examina- currently classified as Lauxaniinae except where indicated
tion of specimens of Eurychoromyia at high magnification otherwise.

Key to the genera of Lauxaniidae of the New World

1. Frons with two pairs fronto-orbital setae, posterior seta strong and reclinate, anterior seta strong
or weak and reclinate to inclinate; prescutellar acrostichal seta usually present, strong. . . . . . . . 2
– Frons with fronto-orbital setae absent, or with one pair only; prescutellar acrostichal seta absent;
EURYCHOROMYIINAE, in part. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

2. Subgena usually inconspicuous, narrow strip below gena; combined height of gena and subgena
less than half height of eye (usually closer to 0.2 times); wing length not exceeding 2.5 times
height; anal vein short . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
– Subgena broad, 0.6 times height of gena or greater, and clearly differentiated from gena by suture;
combined height of gena and subgena 0.7 times height of eye or greater; wing elongate, length
3 times height or greater; anal vein long, nearly reaching wing margin; EURYCHOROMYIINAE,
in part. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103

3. Antenna long and slender (Figs. 19, 20), with first flagellomere at least 7 times longer than basal
height. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
– Antenna relatively short (Fig. 1), with first flagellomere no longer than 6 times basal height (up to
length in Fig. 22) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

4. Wing short; R2+3 distinctly curved and parallel to anterior margin, such that cell r2+3 wide, more
than twice width of cell r1 (Fig. 46); Argentina, Brazil, Paraguay. . . . . . . [Steganolauxania Frey]
– Wing normal; R2+3 not curved and parallel to anterior margin (e.g., Figs. 44, 45) . . . . . . . . . . . . 5

5. Face with two or three conspicuous protuberances. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

– Face lacking protruding bumps, although face may be protuberant . . . . . . . . . . . . . . . . . . . . . . . 7

6. Face yellow with two contrasting shiny black protuberances; fronto-orbital setae reclinate . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cephalella Malloch
– Face yellowish-brown with three pointed protuberances darkened at tips; anterior fronto-orbital
seta inclinate; Argentina, Brazil, Paraguay, Uruguay. . . . . . . . . . . . . . . . . . . [Triconopsis Hendel]

7. Face strongly convex, with facial margin narrow; arista equal to or extending beyond apex of first
flagellomere. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Asilostoma Hendel
– Face slightly prominent, facial margin wide; arista ending before apex of first flagellomere
(Fig. 19). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

8. Prescutellar acrostichal seta present; dorsocentral setae 0+3; anterior fronto-orbital setae strongly
converging; face protuberant on lower two-thirds, usually with polished black spot (Fig. 19);
thorax usually with contrasting colors: yellow with blackish spots or black with yellow spots. . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pachyopella Shewell
– Prescutellar acrostichal seta reduced or absent; dorsocentral setae 0+2; anterior fronto-orbital
setae inclinate, but not strongly; face lacking contrasting spots and not tuberculate; thorax slender,
without contrasting colors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stenolauxania Malloch

9. Wing vein C with black costal spinules extending to apex of R4+5 (Fig. 43); HOMONEURINAE,
in part. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
– Wing vein C with black costal spinules ending well before apex of R4+5 (Fig. 44). . . . . . . . . . . 12

10. Body grayish pruinose with brown maculations; anepisternum with extra seta near or slightly
below middle in addition to posterior seta; wing mostly brownish, strongly patterned, irrorated
with whitish spots; forefemur lacking ctenidium (as in Fig. 37); midtibia with one apical spur (as
in Fig. 41). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trypetisoma Malloch, in part
– Body light yellow to brownish-yellow; anepisternum lacking extra seta, only with posterior seta;
wing hyaline, sometimes patterned with brown but never irrorated; forefemur usually with ctenidium
(Fig. 36); midtibia with two apical spurs (Fig. 40); Homoneura Wulp. . . . . . . . . . . . . . . . . . . . 11
LAUXANIIDAE (lauxaniid flies) 71 977

11. Postocellar setae usually shorter than ocellar setae, strongly convergent, their bases much farther
apart than posterior ocelli (Fig. 26); hind tarsus pale, unornamented; Nearctic Region, widespread
Old World. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Homoneura (Homoneura) Wulp]
– Postocellar setae equal to or longer than ocellar setae, gradually convergent, their bases as close
together as posterior ocelli (Fig. 25); hind tarsus with at least one tarsomere blackened, often
enlarged and ornamented in males (Figs. 38, 39); Nearctic Region . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Homoneura (Tarsohomoneura) Hendel]

12. Anepimeron with one to several short fine setulae dorsally . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

– Anepimeron bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

i vt s
p frorb s
oc s poc s
o vt s
a frorb s

inc frorb s

flgm 1

19 Pachyopella

17 Calliopum 18 Camptoprosopella

flgm 1

frgn sut
scp

22 Melanomyza

20 Steganolauxania 21 Neogriphoneura

Figs.  71.17–22. Heads (continued) and antenna: lateral view of head of (17)  Calliopum quadrisetosum (Thomson), (Nearctic, MND,
fig. 87.32); (18) Camptoprosopella borealis Shewell, (Nearctic, MND, fig. 87.33); (19) Pachyopella sp., (Nearctic, MND, fig. 87.34); anterior
view of head, left antenna omitted, of (20) Steganolauxania latipennis (Coquillett), (Nearctic, MND, fig. 87.44); and (21) Neogriphoneura
sordida (Wiedemann), (Nearctic, MND, fig. 87.45); lateral view of antenna of (22) Melanomyza (Lauxaniella) manuleata (Loew), (Nearctic,
MND, fig. 87.16).
Abbreviations: a frorb s, anterior fronto-orbital seta; flgm 1, first flagellomere; frgn sut, frontogenal suture; inc frorb s, inclinate fronto-orbital
seta; i vt s, inner vertical seta; oc s, ocellar seta; o vt s, outer vertical seta; poc s, postocellar seta; p frorb s, posterior fronto-orbital seta;
scp, scape.
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

13. Presutural dorsocentral seta absent; one katepisternal seta present; anterior fronto-orbital seta long
(as long as posterior) (Fig. 11), inclinate, but not strongly (Fig. 21), close to anterior margin of
frons (Fig. 23); arista plumose (Fig. 11) . . . . . . . . . . . . . . . . . . . . . . . . . Neogriphoneura Malloch
– Presutural dorsocentral seta present; two katepisternal setae present; anterior fronto-orbital seta normal,
reclinate, not close to anterior margin of frons (Fig. 17); arista pubescent (as in Fig. 16). . . . . . . 14

14. Face and body dull yellow to yellowish-brown; Holarctic Region. . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Meiosimyza (Lyciella) Collin]
– Face mostly polished black, sometimes with grayish pruinescence dorsally; body shining black,
sometimes with yellowish legs; Holarctic Region. . . . . . . . . . . . . . . . . . . . . . . [Calliopum Strand]

15. Wing with supplementary crossvein joining R4+5 and M1 beyond crossvein dm-cu (Figs. 47, 52).
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
– Wing lacking supplementary crossvein beyond crossvein dm-cu (e.g., Fig. 44). . . . . . . . . . . . . 17

16. Frons with protruding, conical profile, with transverse depression (Fig. 9); arista thickened and
thickly pubescent (Fig. 9); postsutural intra-alar seta absent. . . . . . . . . . . . . Xeniconeura Shewell
– Frons at most slightly convex in profile, without transverse depression; arista normal, pubescent,
neither thickened nor thickly pubescent; postsutural intra-alar seta present. . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xenopterella Malloch

17. With 0+3 dorsocentral setae present, with anterior pair much weaker than others; length of
crossvein r-m two-thirds that of dm-cu; abdomen black. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
– Dorsocentral setae various; length of crossvein r-m less than two-thirds that of dm-cu (Fig. 45);
abdomen various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

18. Ocellar setae absent; first flagellomere tapering distally, arista plumose; wing membrane dark
with white spots and usually white along distal margin . . . . . . . . . . Baliopteridion Papp & Silva
– Ocellar setae present; first flagellomere rounded, arista pubescent; wing membrane banded, not
spotted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Undescribed Genus A

19. Frons protruding, conical or with bulging profile (Fig. 15). . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

– Frons various, at most slightly convex in profile. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26

20. Upper lunule and anterior margin of frons depressed (slightly concave in profile). . . . . . . . . . . 21
– Upper lunule and anterior margin of frons not depressed (various, but not concave). . . . . . . . . 23

21. Ocellar seta long; intra-alar seta present; anterior katepisternal seta present but weak, half length
of posterior; wing spotted; Ecuador, Brazil. . . . . . . . . . . . . . . . . . . . . . . . [Undescribed Genus B]
– Ocellar seta short; intra-alar seta absent; anterior katepisternal seta present and strong or absent;
wing hyaline. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

22. Arista pubescent; ocellar seta short and strong; 1+2 dorsocentral setae present; anterior katepisternal
seta present; prescutellar acrostichal seta weak; wing brownish hyaline . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Undescribed Genus C
– Arista short plumose; ocellar seta short and weak; 0+3 dorsocentral setae present; anterior
katepisternal seta absent; prescutellar acrostichal seta strong; wing tinged yellow; Brazil. . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Hirtodeceia Shewell]
LAUXANIIDAE (lauxaniid flies) 71 979

23. Frons with anterior margin projecting over antennal bases, flat and horizontal in profile (fronto-facial
angle slightly less than 90º), conspicuously setulose (Fig.  10); sometimes with distinct orbito-
antennal spot; dorsalmost subvibrissal seta elongate (Fig. 10); arista pubescent; anterior katepis-
ternal seta short; wing usually spotted or darkened. . . . . . . . . . . Oncodometopus Shewell, in part
– Frons with anterior margin swollen, either rounded (Fig.  15) or upturned distally (Fig.  9),
sometimes with longitudinal groove; no subvibrissal setae conspicuously enlarged; other
characters various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

24. Entirely yellow, unmarked; most setae pale; head lacking dark pruinose patch; Nearctic Region.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Mallochomyza Hendel, in part]
– Dark with various dark and silvery-gray pruinose markings; setae black; head with velvety black
orbito-antennal spot. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

25. Head length less than two-thirds that of height; frons with anterior margin strongly bulging
forward, bearing shiny protuberance above pruinose orbito-antennal spot; wing brown with
yellowish translucent areas; Brazil, Peru. . . . . . . . . . . . . . . . . . . . . . . . . . . [Paracestrotus Hendel]
– Head length more than two-thirds that of height; frons with anterior margin slightly bulging,
sometimes with shiny area above pruinose orbito-antennal spot, but lacking protuberances, sometimes
with shallow medial groove; wing often marked, but not as above. . . . . . . . Siphonophysa Hendel

26. Postsutural intra-alar seta present (Fig. 33). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27

– Postsutural intra-alar seta absent (Fig. 34). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53

27. Scutellum with many setulae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28

– Scutellum bare (Fig. 33). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

28. R4+5 bare; ocellar seta as strong as postocellar seta. . . . . . . . . . . . . . . . . . . . Deutominettia Hendel
– R4+5 setulose dorsally; ocellar seta weaker than postocellar seta. . . . . . . . . . Scutominettia Hendel

29. Wing vein M with pronounced forwardly rounded curvature apically, closely approaching the tip
of R4+5 and almost closing cell r4+5 (Fig. 48) . . . . . . . . . . . . . . . . . . . . . . . . . Griphoneura Schiner
– Wing vein M nearly straight, not approaching R4+5 near apex (Fig. 44) . . . . . . . . . . . . . . . . . . . 30

30. At least one radial vein with setulae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

– Radial veins bare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

31. R1 bare; R4+5 with setulae dorsally; midtibia with distinct short posterior setae . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetominettia Malloch
– R1 with setulae dorsally; R4+5 with or without setulae; midtibia lacking posterior setae. . . . . . . 32

32. R4+5 with setulae; basal part of Rs bare . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dryomyzothea Hendel

– R4+5 bare; basal part of Rs with setulae (as in Fig. 49). . . . . . . . . . . . . . . . . . . . . . Setulina Malloch

33. Frons with fronto-orbital setae on small raised bases; wing deeply fuscous brown on costal half or
more, hind margin often with irregular hyaline patches . . . . . . . . . . . . . . Chaetocoelia Giglio-Tos
– Frons lacking raised bases to fronto-orbital setae; wing various. . . . . . . . . . . . . . . . . . . . . . . . . 34

34. Scape and pedicel subequal in length. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35

– Scape shorter than pedicel (Fig. 14). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
980
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

a frorb s poc s

p frorb s
25 Homoneura

o vt s

i vt s

23 Neogriphoneura 24 Trisapromyza

poc s

26 Homoneura

inc frorb s
rc frorb s
oc s

oc s
poc s

27 Camptoprosopella 28 Minettia
poc s

29 Poecilominettia

30 Melanomyza 31 Oncodometopus 32 Poecilolycia

Figs.  71.23–32. Heads (concluded): dorsal view of (23) Neogriphoneura sordida (Wiedemann), (Nearctic, MND, fig.  87.14):
(24) Trisapromyza vittigera (Coquillett), (Nearctic, MND, fig. 87.15); (25) Homoneura (Tarsohomoneura) americana (Wiedemann), (Nearctic,
MND, fig. 87.50); (26) H. (Homoneura) philadelphica (Macquart), (Nearctic, MND, fig. 87.51); (27) Camptoprosopella borealis Shewell,
(Nearctic, MND, fig. 87.17); (28) Minettia flaveola (Coquillett), (Nearctic, MND, fig. 87.18); (29) Poecilominettia macula (Loew), (Nearctic,
MND, fig. 87.19); (30) Melanomyza (Melanomyza) gracilipes (Loew), (Nearctic, MND, fig. 87.41); (31) Oncodometopus umbrosus (Loew),
(Nearctic, MND, fig. 87.42); and (32) Poecilolycia serrata (Malloch), (Nearctic, MND, fig. 87.43).
Abbreviations: a frorb s, anterior fronto-orbital seta; inc frorb s, inclinate fronto-orbital seta; i vt s, inner vertical seta; oc s, ocellar seta; o vt s,
outer vertical seta; poc s, postocellar seta; p frorb s, posterior fronto-orbital seta; rc frorb s, reclinate fronto-orbital seta.
LAUXANIIDAE (lauxaniid flies) 71 981

35. Two katepisternal setae present; proepisternal seta long and strong; Venezuela. . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Dryosapromyza Hendel]
– One katepisternal seta present (if two, anterior seta weak, hairlike); proepisternal seta absent or
weak. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sciosapromyza Hendel, in part

36. Anterior katepisternal seta present (Fig. 1), sometimes shorter than posterior, but strong. . . . . 37
– Anterior katepisternal seta absent or greatly reduced, hair- or setula-like. . . . . . . . . . . . . . . . . . 50

37. Arista plumose; dorsocentral setae 0+3; fore- and hind tibiae and tarsi laterally compressed; wing
yellowish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudominettia Papp & Silva
– Arista bare to plumose; dorsocentral setae 1+2, 0+2, or 0+3; legs not modified; wing various. . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38

38. Midtibia with short, strong posterior setae (Fig. 42); ocellar seta weak; bases of anterior fronto-
orbital setae separated by smaller distance than that between posterior fronto-orbital setae . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Allominettia Hendel, in part
– Midtibia without short, strong posterior setae (Fig.  41); ocellar seta various; bases of anterior
fronto-orbital setae often separated by distance subequal to that of posterior fronto-orbital setae
(Fig. 28). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39

39. Arista bare, pubescent or short plumose (all rays shorter than half height of first flagellomere) . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
– Arista long plumose (some rays as long or longer than half height of first flagellomere). . . . . . 46

40. Ocellar seta absent or setula-like; scutellum with shallow longitudinal groove apically . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Agriphoneura Hendel
– Ocellar seta present, various; scutellum without longitudinal groove apically. . . . . . . . . . . . . . 41

41. Ocellar seta equal to or longer than postocellar seta; face glossy, slightly convex (Fig. 8); arista
bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudocalliope Malloch
– Ocellar seta various; face dull, sometimes slightly convex; arista pubescent to short plumose. . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42

42. Ocellar setae long and parallel, with bases separated by at least diameter of one ocellus (Fig. 28);
anterior fronto-orbital seta closer to posterior fronto-orbital seta than to anterior margin of frons
(Fig. 1); Minettia Robineau-Desvoidy, in part. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
– Ocellar setae short and divergent, with bases closer to each other than diameter of one ocellus
(Fig.  29); anterior fronto-orbital seta closer to anterior margin of frons than to posterior
fronto-orbital setae (Fig. 29) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44

43. Prosternum bare; black species; Holarctic, Oriental Regions. . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Minettia (Frendelia) Collin, in part]
– Prosternum haired; color various, but not black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Minettia (Minettia) Robineau-Desvoidy, in part

44. Apical pair of scutellar setae convergent. . . . . . . . . . . . . . . . . . . . Poecilominettia Hendel, in part

– Apical pair of scutellar setae divergent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45

45. Ocellar seta tiny, weak and setula-like; parafacial row with dorsalmost setula longest and strongest;
1+2 dorsocentral setae present; intra-alar seta short; slender, pale yellow species; thorax
sometimes with setigerous spots; Brazil . . . . . . . . . . . . . . . . . . . . . . . . . . [Undescribed Genus D]
– Ocellar seta short, but strong and not setula-like; parafacial row of setulae uniform; 0+3
dorsocentral setae present; intra-alar seta strong; stout species; thorax lacking setigerous spots;
Brazil, Argentina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Undescribed Genus E]
982
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

presut
spal s

presut
spal s

a dc s
psut
ial s

p dc s

33 Minettia 34 Xenochaetina 35 Homoneura

cten

36 Homoneura

37 Minettia
38 Homoneura 39 Homoneura

tb spur

tb spur

40 Homoneura 41 Minettia 42 Allominettia

Figs. 71.33–42. Thoraces and legs: dorsal view of thorax of (33) Minettia flaveola (Coquillett), (Nearctic, MND, fig. 87.31); (34) Xenochaetina
muscaria (Loew), (Nearctic, MND, fig. 87.49); and (35) Homoneura (Homoneura) philadelphica (Macquart), (Nearctic, MND, fig. 87.47);
anterior view of left forefemur of (36) H. bispina (Loew), (Nearctic, MND, fig. 87.29); and (37) M. flaveola, (Nearctic, MND, fig. 87.30);
anterior view of left hind tarsus of male and female of (38) H. (Tarsohomoneura) ornatipes (Johnson), (Nearctic, MND, fig. 87.52); and
(39) H. (Tarsohomoneura) americana (Wiedemann), (Nearctic, MND, fig. 87.53); ventral view of left midtibia of (40) H. bispina, (Nearctic,
MND, fig. 87.27); (41) M. flaveola, (Nearctic, MND, fig. 87.28); and (42) Allominettia geniseta (Malloch), (MND, fig. 87.54).
Abbreviations: a dc s, anterior dorsocentral seta; cten, ctenidium; p dc s, posterior dorsocentral seta; presut spal s, presutural supra-alar seta;
psut ial s, postsutural intra-alar seta; tb spur, tibial spurs.
LAUXANIIDAE (lauxaniid flies) 71 983

46. Color entirely black; Holarctic, Oriental Regions. . . . . . . . . [Minettia (Frendelia) Collin, in part]
– Color various, yellowish to dark grayish pruinose, not entirely black. . . . . . . . . . . . . . . . . . . . . 47

47. Anterior fronto-orbital seta closer to anterior margin of frons than to posterior fronto-orbital seta
(Fig. 29); ocellar seta short (Fig. 29); if arista with long rays, then with distinct spots on face,
frons, scutum, and scutellum, and thorax with dark stripes; yellowish species. . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Poecilominettia Hendel, in part
– Anterior fronto-orbital seta equally placed between anterior margin of frons and posterior fronto-
orbital seta; ocellar seta short or long; if arista with long rays, then face and frons sometimes with
spots, but thorax without dark stripes; body color various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48

48. Fronto-facial angle approximately 90º; frons drawn out anteromedially to form small process
between antennal bases; face shining, with silvery dusting on parafacial. . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sciosapromyza Hendel, in part
– Fronto-facial angle obtuse; frons lacking process between antennal bases; face dull or pruinose .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49

49. Extra seta between intra-alar and supra-alar setae present, distinctly larger than surrounding
setulae; body yellow to dark yellow; wing with brown spots. . . . . . . . . . . . . Neominettia Hendel
– Lacking extra seta between intra-alar and supra-alar setae; body color various; wing usually
hyaline, rarely with brown spots . . . . . . . . . . . . . Minettia (Minettia) Robineau-Desvoidy, in part

50. Midtibia with strong posterior setae (Fig. 42). . . . . . . . . . . . . . . . . . . Allominettia Hendel, in part
– Midtibia lacking posterior setae (as in Fig. 41). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51

51. First flagellomere rounded; arista plumose with long rays both dorsally and ventrally; femora and
tibiae long haired anteriorly, without setae; Peru. . . . . . . . . . . . . . . . . . . . . . . . . [Eriurgus Hendel]
– First flagellomere elongated oval; arista plumose, with rays shorter ventrally; femora and tibiae
without long setulae or hairs anteriorly, with setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52

52. With 0+3 dorsocentral setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ocellominettia Hendel

– With 0+2 dorsocentral setae; Brazil. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Undescribed Genus F]

53. Scutellum more than half length of scutum. . . . . . . . . . . . . . . . . . . . . . . Celypholauxania Hendel

– Scutellum of normal length, less than one-third length of scutum . . . . . . . . . . . . . . . . . . . . . . . 54

54. Fronto-facial angle less than 90º . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55

– Fronto-facial angle equal to or greater than 90º. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56

55. Wing marked, solidly dark in anterior half with posteriorly pointed extensions; crossvein dm-cu
before middle of wing; cell dm with pointed apex; head not sharply pointed anteriorly, eye higher
than long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Procrita Hendel
– Wing hyaline or variously marked, never solidly dark in anterior half with posteriorly projecting
points; crossvein dm-cu beyond middle of wing, cell dm with rounded apex; head sharply pointed
anteriorly (Fig. 7); eye at least slightly longer than high. . . . . . . . . . . . Trigonometopus Macquart

56. First flagellomere with elongated dorsal pubescence, length at least half height of first
flagellomere. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
– First flagellomere without long pubescence. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
984
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

57. R4+5 setulose; first flagellomere with dorsal pubescence as long or longer than its height . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Blepharolauxania Hendel
– R4+5 bare; first flagellomere with dorsal pubescence half as long as its height . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Platygraphium Hendel

58. Anterior fronto-orbital seta distinctly inclinate (Fig. 27). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59

– Anterior fronto-orbital seta reclinate (Fig. 24) (or slightly inclinate only at apex). . . . . . . . . . . 63

59. Body dark to black overall, often with some brownish or grayish pruinescence (especially on
scutum), and always with extensive areas of shiny black (especially abdomen, face, frons,
pleuron, and lateral scutum); face not yellow to orange . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
– Body color various, but not black overall with pruinescence on scutum; if abdomen and/or thorax
black (e.g., some Camptoprosopella), then head mostly yellow to orange, with frons not convex.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61

60. Face with deep transverse depression through middle; frons with transverse depression anteriorly;
with 0+1 dorsocentral setae; prescutellar acrostichal seta absent or minute; anal vein reduced. . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Freyia Malloch
– Face convex with slight transverse depression in lower part; frons flat or rounded, never with
transverse depression; with 0+2 dorsocentral setae; prescutellar acrostichal seta present; anal vein
not reduced; Nearctic Region. . . . . . . . . . . . . . . . . . [Melanomyza (Lauxaniella) Malloch, in part]

61. Ocellar seta strong and long; anterior and posterior fronto-orbital setae arise close together
(Fig. 27), normal sized; facial grooves convergent on upper half of face; face usually flat; parafacial
broad (Fig. 18). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Camptoprosopella Hendel
– Ocellar seta minute or absent; anterior and posterior fronto-orbital setae not arising close together,
anterior fronto-orbital seta short; face convex; parafacial usually narrow . . . . . . . . . . . . . . . . . 62

62. Face strongly convex, protruding beyond eye in profile; frons broad, anteriorly concave; anterior
fronto-orbital seta strongly inclinate; acrostichal setulae biseriate; scutellum almost as long as
wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Physegenua Macquart
– Face almost vertical in profile; frons slightly wider than long, not concave anteriorly; anterior
fronto-orbital seta inclinate, with apex slightly reclinate; acrostichal setulae multiseriate; scutellum
wider than long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neoxangelina Hendel

63. Face strongly convex or with protuberances; in profile clearly convex (Fig. 6), sometimes more
distinct in lower part. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
– Face in profile not distinctly convex, sometimes flat, concave, or slightly rounded. . . . . . . . . . 70

64. Arista pubescent or short plumose (rays not longer than half height of first flagellomere). . . . . 65
– Arista long plumose (rays longer than half height of first flagellomere). . . . . . . . . . . . . . . . . . . 66

65. First flagellomere more than 3 times longer than wide; arista white pubescent (Fig. 13); ocellar
seta slightly shorter than postocellar seta; scutellum lacking apical longitudinal groove; Holarctic
Region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Lauxania Meigen, in part]
– First flagellomere less than 2.5 times longer than wide; arista dark pubescent or short plumose;
ocellar seta minute (Fig. 6); scutellum often with more or less distinct apical longitudinal groove.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Physoclypeus Hendel

66. Rs with one to several setulae (Fig. 49); face with transverse groove along oral margin (Fig. 5). .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xenochaetina Malloch, in part
– Rs bare; face without transverse groove along oral margin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
LAUXANIIDAE (lauxaniid flies) 71 985

C sta C sta

R4+5
R4+5

M1

M1

43 Homoneura 44 Minettia
Sc R1
R2+3

R4+5

M1

CuA1

45 Mallochomyza 46 Steganolauxania

R4+5

M1

47 Xenopterella 48 Griphoneura

v hairs

50 Neogriphoneura
49 Xenochaetina

R4+5

dm-cu

51 Trypetisoma 52 Xeniconeura

Figs. 71.43–52. Wings: dorsal view, with offset of enlargement of apex, of (43) Homoneura bispina (Loew), (Nearctic, MND, fig. 87.20); and
(44) Minettia flaveola (Coquillett), (Nearctic, MND, fig. 87.21); dorsal view of (45) Mallochomyza citreifrons (Malloch), (Nearctic, MND,
fig. 87.55); (46) Steganolauxania latipennis (Coquillett), (Nearctic, MND, fig. 87.22); (47) Xenopterella beameri Steyskal, (Nearctic, MND,
fig. 87.24); and (48) Griphoneura sp.; wing base, with setulae at base of Rs encircled, of (49) Xenochaetina muscaria (Loew), (Nearctic,
MND, fig. 87.23); ventral view of costal area of wing at apices of Sc and R1 of (50) Neogriphoneura sordida (Wiedemann), (Nearctic, MND,
fig. 87.56); dorsal view of wing of (51) Trypetisoma sticticum (Loew), (Nearctic, MND, fig. 87.25); and (52) Xeniconeura costalis (Curran),
(Nearctic, MND, fig. 87.26). Figure 48 illustrated by R. Vargas.
Abbreviations: C sta, costal setulae; v hairs, ventral hairs.
986
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

67. Small (2–3 mm) species; face convex with deep grooves in upper part; outer vertical seta absent;
prescutellar acrostichal seta absent . . . . . . . . . . . . . . . . . . . . . . . . . . . Minilauxania Papp & Silva
– Size greater than 4 mm; face convex, but lacking deep grooves in upper part; outer vertical seta
present; prescutellar acrostichal seta present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68

68. Large, robust species; facial margin wide; first flagellomere oval; Brazil. . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Undescribed Genus G]
– Size various, more elongate; facial margin narrow; first flagellomere long, rodlike; Melanomyza
Malloch, in part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69

69. Antennal scape not longer than wide, bare below; first flagellomere less than 3 times longer than
wide; face moderately swollen (Fig. 2), without well-defined V-shaped depression below. . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Melanomyza (Melanomyza) Malloch, in part
– Antennal scape longer than wide, usually with apical hairs below (Fig. 22); first flagellomere
usually more than 3 times longer than wide; face distinctly swollen, with V-shaped depression
below; Nearctic Region. . . . . . . . . . . . . . . . . . . . . . [Melanomyza (Lauxaniella) Malloch, in part]

70. Lower facial margin with three to seven vibrissa-like setae oriented anteriorly. . . . Hypagoga Hendel
– Lower facial margin without strong setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71

spmth tg 8 cerc

st 8

54 Sapromyza

epiprct

53 Melanomyza

spr 7 hyprct

55 Minettia

Figs. 71.53–55. Female terminalia, each with offset of ventral view of sternite 8, of (53) Melanomyza (Melanomyza) gracilipes (Loew),
(Nearctic, MND, fig.  87.61); (54)  Sapromyza nigrans (Melander), (Nearctic, MND, fig.  87.65); and (55)  Minettia lupulina (Fabricius),
(Nearctic, MND, fig. 87.63).
Abbreviations: cerc, cercus; epiprct, epiproct; hyprct, hypoproct; spmth, spermathecae; spr, spiracle; st, sternite; tg, tergite.
LAUXANIIDAE (lauxaniid flies) 71 987

71. Anterior fronto-orbital seta slightly inclinate only at apex (Fig. 30); face smooth, shining; arista
plumose (Fig. 2); tarsi usually paler in contrast with tibiae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Melanomyza (Melanomyza) Malloch, in part
– Anterior fronto-orbital seta reclinate; face smooth or dull; arista pubescent or plumose; tarsi
various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72

72. Crossvein r-m close to crossvein dm-cu, forming zigzag line. . . . . . . . . . . . . . . . . Ritaemyia Frey
– Crossvein r-m not close to crossvein dm-cu . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73

73. Presutural dorsocentral seta present; arista short pubescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74

– Presutural dorsocentral seta absent; arista pubescent or plumose. . . . . . . . . . . . . . . . . . . . . . . . 81

74. Wing strongly patterned, irrorated (Fig.  51); vein  R2+3 usually wavy; HOMONEURINAE, in
part. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trypetisoma Malloch, in part
– Wing never strongly patterned nor irrorated, occasionally with markings on crossveins; vein R2+3
straight. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75

75. First flagellomere more than 3 times longer than basal height (Fig. 13); frons entirely brown and
shining, with anterior margin elevated; face and parafacial mostly shining brown; Holarctic
Region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Lauxania Meigen, in part]
– First flagellomere less than 2.5 times longer than basal height; frons, face and parafacial dull, with
anterior margin not elevated. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76

76. Entirely pale yellow, unmarked, delicate species; setae mostly pale; wing sometimes with faint spots
at crossveins; forefemur lacking ctenidium; Nearctic Region. . . . . [Mallochomyza Hendel, in part]
– Yellowish to gray, often with face, frons, scutum, legs, and abdomen variously marked; setae
black; wing hyaline; forefemur sometimes with ctenidium. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77

77. With 1+2 dorsocentral setae (anterior seta sometimes close to suture). . . . . . . . . . . . . . . . . . . . 78
– With 1+3 dorsocentral setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79

78. Frons and fronto-orbital plates grayish pruinose, sometimes with brown markings; face with
spots; scutum with median vittae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trisapromyza Shewell
– Frons yellow with fronto-orbital plates contrasting grayish pruinose; face lacking spots; scutum
not vittate; Argentina, Chile. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Undescribed Genus H]

79. Katepisternum with one seta; face yellow with two brown longitudinal stripes; Brazil. . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Undescribed Genus I]
– Katepisternum with two setae; face various, but without two brown longitudinal stripes . . . . . 80

80. More or less uniformly colored, usually dark yellowish to orange, with few, if any, distinctive
markings. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sapromyza Fallén
– Yellowish to gray, variously marked or mottled; gena with dark brown mark below eye (Fig. 16);
frons, face, scutum, legs, and abdomen variously marked, strongly contrasting (e.g., Fig. 32). . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Poecilolycia Shewell

81. Wing with some R veins with setulae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82

– Wing with R veins bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83

82. Scutellum setulose dorsally on disc, in addition to four marginal setae; Peru. . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Scutolauxania Hendel]
– Scutellum bare dorsally, with only four marginal setae. . . . . . . . . . Xenochaetina Malloch, in part
988
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

83. Arista appearing thick, due to short, dense black setulae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84

– Arista without short, dense black setulae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85

84. Eye small; first flagellomere slender and tapering apically; Uruguay. . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Neopachycerina Malloch]
– Eye normal; first flagellomere ovoid and rounded apically; Brazil. . . . . . . [Undescribed Genus J]

85. Frons projecting (Fig. 10) and thickly setulose anteriorly (Fig. 31); dorsalmost subvibrissal seta
usually longest and strongest (Fig. 10); wing cloudy or spotted. . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oncodometopus Shewell, in part
– Frons not projecting anteriorly (Figs. 4, 12), sometimes setulose; subvibrissal setulae more or less
uniform; wing hyaline or patterned. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86

86. Body dark brown in upper part and contrastingly paler below; ocellar setae short, divergent; arista
short plumose, with basal-most dorsal ray usually longest. . . . . . . . . . . . . . . . . Trivialia Malloch
– Body color various, but not distinctly contrasting between upper and lower parts; ocellar seta
short or long, various in orientation; arista plumose or pubescent . . . . . . . . . . . . . . . . . . . . . . . 87

87. Katepisternum with two setae, both well developed; face without marks. . . Undescribed Genus K
– Katepisternum with one seta (if two, then anterior seta reduced and hairlike); face various, often
marked. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88

88. With 0+3 dorsocentral setae, all well developed and strong; ocellar seta long, well developed. . . 89
– With 0+2 dorsocentral setae (if 0+3, anterior seta weak and setula-like or ocellar seta weak);
ocellar seta various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90

89. Arista long plumose; acrostichal setulae uniformly small, in eight rows. . . . . Undescribed Genus L
– Arista pubescent; acrostichal setulae in four to six rows, with median rows usually slightly
enlarged; Argentina, Chile. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Undescribed Genus M]

90. Eye with posteroventral margin strongly concave (as in Fig. 2, 14) to straight (Fig. 3); body often
with grayish pruinosity. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
– Eye with posteroventral margin convexly rounded; body usually lacking grayish pruinosity . . 95

91. Frons flat in profile (Fig. 3); thorax grayish pruinose with black marks. . . . . . Neodeceia Malloch
– Frons convex in profile; thorax without black marks, but sometimes vittate . . . . . . . . . . . . . . . 92

92. Face narrow; frons smooth, with orbito-antennal spot; prescutellar acrostichal seta absent or
reduced; legs slender . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gibbolauxania Papp & Silva
– Face not narrow; frons dull or shining, without orbito-antennal spot; prescutellar acrostichal seta
present, well developed; legs normal. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93

93. Ocellar seta minute; Brazil. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Undescribed Genus N]

– Ocellar seta various, but not minute. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94

94. Eye with posteroventral margin straight (as in Fig.  3); ocellar setae long, parallel; Nearctic
Region, Caribbean. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Deceia Malloch]
– Eye with posteroventral margin strongly concave; ocellar setae various . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudogriphoneura Hendel
LAUXANIIDAE (lauxaniid flies) 71 989

95. Wing with dark transverse bands; veins R4+5 and M distinctly divergent at apex; Brazil. . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Undescribed Genus O]
– Wing without transverse bands; veins R4+5 and M subparallel . . . . . . . . . . . . . . . . . . . . . . . . . . 96

96. Eye wider than high; face short, with brown transverse band; scutum brown vittate; abdomen with
brown setigerous spots; male with basal tarsomere of foreleg swollen. . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Marmarodeceia Shewell
– Eye higher than wide; face long, sometimes with longitudinal stripes; scutum vittate or not;
abdomen without setigerous spots; if basal tarsomere of foreleg swollen, then also laterally
compressed along with foretibia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97

97. Ocellar setae short and divergent; gena with brown mark below eye; foretibia and basal tarsomere
of foreleg thickened and laterally compressed; hind tibia thickened; Brazil. . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Undescribed Genus P]
– Ocellar seta long; gena without mark below eye; legs not modified. . . . . . . . . . . . . . . . . . . . . . 98

56 Homoneura
lab scl

md hyphar d brg

ant

58 Homoneura
57 Homoneura den scl b
59 Homoneura

60 Homoneura

Figs. 71.56–60. Larva and associated structures of Homoneura sp., (Nearctic, MND, figs. 87.66–70); lateral view of third instar larva (56);
anterior spiracle (57); posterior spiracular plate (58); dorsal (59a) and lateral (59b) views of cephalopharyngeal skeleton; and posterior
spiracular disc (60).
Abbreviations: ant, antenna; d brg, dorsal bridge; den scl, dental sclerite; hyphar, hypopharynx; lab scl, labial sclerite; md, mandible.
990
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

98. Arista short plumose; ocellar setae divergent; scutum and pleuron without contrasting stripes;
Brazil . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Undescribed Genus Q]
– Arista long plumose; ocellar setae parallel; scutum and pleuron grayish pruinose with brown
stripes; Brazil. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Undescribed Genus R]

99. Body and legs elongate, with mid- and hind femora each longer than scutum; hind tarsus either
longer than hind femur or with basal tarsomere expanded into flattened pad; Colombia. . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Undescribed Genus S]
– Body and legs more compact, with femora shorter than scutum; hind tarsus unmodified, shorter
than hind femur . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100

100. Antenna greatly elongate, altogether 1.5  times longer than head height; all antennal segments
elongated, with scape and pedicel subequal in length, and first flagellomere only slightly longer
than either; scutum with silvery gray dorsocentral vittae on brown background; tibiae unmodified;
Ecuador . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Eurystratiomyia Gaimari & Silva]
– Antennal length shorter than head height; scape and pedicel much shorter than first flagellomere;
scutum not vittate; at least hind tibia greatly expanded and laterally flattened. . . . . . . . . . . . . 101

101. Antennal first flagellomere elongated, 3 times longer than combined length of scape and pedicel;
postocellar seta present but small; scutellum with shallow longitudinal depression, appearing
weakly bilobed in dorsal view. . . . . . . . . . . . . . . . . . . . . . . . . . . Choryeuromyia Gaimari & Silva
– Antennal first flagellomere short, only slightly longer than combined length of scape and pedicel;
postocellar seta absent; scutellum unmodified. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102

102. Body predominantly yellow pruinose with brown markings; setae of head (e.g., fronto-orbital
seta, inner and outer vertical setae) and thorax (e.g., two pairs of dorsocentral setae, two pairs
of scutellar setae, anepisternal seta) small, but distinct; gena slightly higher than subgena, and
combined height of gena and subgena less than eye height; frons yellow and brown pruinose;
Ecuador . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Roryeuchomyia Gaimari & Silva]
– Body dark brown to black, with some whitish pruinosity; setae of head and thorax at most tiny
and inconspicuous; gena at least 1.4 times higher than eye and 5 times higher than subgena; frons
shiny black, with only sparse pruinosity; Bolivia . . . . . . . . . . . . . . . . . . [Eurychoromyia Hendel]

103. Face with paired velvety black pruinose spots; gena with velvety black pruinose spot below eye;
foretarsus at least twice length of hind tarsus . . . . . . . . . . . . . . . Physegeniopsis Gaimari & Silva
– Face lacking spots, instead with reticulated pattern of brown and yellowish pruinosity; gena
lacking spot below eye; foretarsus less than 1.5 times longer than hind tarsus. . . . . . . . . . . . . 104

104. Ocellar seta as strong as postocellar seta; anterior fronto-orbital seta strong, more than half length
of posterior seta; anterior part of fronto-orbital plate without small brown spots; antenno-orbital
spot small, elongated black mark; Colombia. . . . . . . . . . . . . . [Euryhendelimyia Gaimari & Silva]
– Ocellar seta weak, hairlike, much smaller than postocellar seta; anterior fronto-orbital seta tiny,
hairlike; anterior part of fronto-orbital plate with small brown spots at bases of all setulae;
antenno-orbital spot large, rounded, velvety black pruinose. . . . . . . . . . . Tauridion Papp & Silva

Synopsis of the fauna contains no species of Homoneura, and in fact no species of

Homoneurinae except for two species of Trypetisoma, a genus
Lauxaniidae are extremely diverse worldwide, with their that may or may not be a homoneurine. A major radiation at
greatest diversity in both the Old and New World tropics both the generic and species levels occurred in the Neotropi-
(Stuckenberg, 1971). Four genera are nearly cosmopolitan, cal Region. Of the 93 total New World genera and subgenera,
including Homoneura, Minettia, Sapromyza, and Trigonome- only nine are not found in the Neotropical Region, which has
topus, each of which, after substantial revisionary work, will a described fauna of nearly 400 species. Of the Neotropical
likely be found to represent several distinct lineages. Of these genera and subgenera, 27 are strictly South American. The
genera, it is interesting to note that the Neotropical fauna Central American fauna, including Neotropical Mexico and
LAUXANIIDAE (lauxaniid flies) 71 991

the islands of the Caribbean, consists of nearly 200 described South America and have both been recorded from Costa Rica.
species in 57 genera. For the region, it is readily apparent af- Chaetocoelia caloptera Hendel is known only from Mexico.
ter study that only a small fraction of the fauna is described, The other two species are known from South America and the
with hundreds of new species recognized. West Indies. More than 20 new species are known from Costa
Rica, and seven from Guatemala and Mexico.
Agriphoneura Hendel. The one described species is
known only from Bolivia, but new species are known as Chaetominettia Malloch. This strictly Neotropical ge-
far north as Neotropical Mexico and Puerto Rico in the nus has four valid species, two of which occur in Costa
Caribbean. Rica. The first, C. corollae (Fabricius) is widespread from
Mexico through southern Brazil and Paraguay. The second,
Allominettia Hendel. One species, A. geniseta (Malloch), C. spinitibia Malloch, is only known from Costa Rica and
occurs throughout Central America, from the southern USA Guyana. The other two species are known only from South
(Texas) through Panama. The other three species are South America.
American, with A. approximata (Malloch) and A. assimilis
(Malloch) also occurring in Panama and Trinidad. Choryeuromyia Gaimari & Silva. The single species of this
eurychoromyiine genus, C. xenisma Gaimari & Silva, is known
Asilostoma Hendel. This strictly Neotropical genus has only from Costa Rica. Details of this species are given in Gaim-
five described species, four of which are Central American. ari & Silva (2010).
Two occur in Costa Rica, A. atriceps Malloch and A. flavi-
facies Malloch, the former being widespread from Mexico Deutominettia Hendel. Both of the two described species
through Ecuador. The other two species, A. pallipes Malloch of this genus occur south of Costa Rica, D. bimaculata Mal-
and A. palpalis Malloch, are only known from Panama. For loch in Panama, and the other in South America. Malloch
the genus, six new species are known from Costa Rica, one (1928) provided a key to the described species.
from Guatemala, and two from Panama. Malloch (1928)
provided a key to the few described species, although more Dryomyzothea Hendel. Although the two described species
than 20 new species are known throughout the Neotropical are known only from South America, at least one new species is
Region. known from Panama.

Blepharolauxania Hendel. The single described species Freyia Malloch. The single species of this genus, F. nigrita
of this genus occurs in Brazil and Peru. One new species is Malloch, is known only from Costa Rica.
known from Costa Rica.
Gibbolauxania Papp & Silva. Two of the three described
Camptoprosopella Hendel. This large (32 described spe- species of this genus, G. aliena (Malloch) and G. lateralis
cies) New World genus has 22 species in the Neotropical (Malloch), are known only from Costa Rica, whereas the
Region. Of these, 16 species occur in Central America; six other is known only from Peru. At least one new species is
of these extend north into the USA, three extend south into known from Costa Rica.
South America, one also occurs in the Greater Antilles, and
Griphoneura Schiner. This strictly Neotropical genus
six are strictly Central American. Two additional species are
has eight described species, four of which occur in Central
found in the Greater Antilles, one of which also occurs in the
America. The other three species are known only from South
southeastern USA. The remaining four species are strictly
America. More than 15 new species are known from Costa
South American. One new species is known from Guatema-
Rica. Curran (1938) provided a key to the known species,
la. This genus was last revised by Shewell (1939).
although many new species are recognized.
Celypholauxania Hendel. Although the single described
species is known only from northern South America, there Hypagoga Hendel. The single described species of this ge-
is at least one undescribed species in Costa Rica, and eight nus is from Venezuela, but five new species are known to oc-
others are known in South America. cur in Costa Rica, several are known from elsewhere in Central
America, four from the Caribbean Islands, and 21 from South
Cephalella Malloch. The single species of this genus, America.
C. fumipennis Malloch, was described from Costa Rica and
occurs throughout Central America from Mexico through Marmarodeceia Shewell. Only one of the three valid species
Panama. Silva (1999) provides details on this genus. is known from Costa Rica, M. marmorata (Malloch), which is
widespread from Costa Rica south to Brazil and Paraguay, and
Chaetocoelia Giglio-Tos. This strictly Neotropical genus has into the West Indies. The other two species are from South
five valid species, two of which, C. distinctissima (Schiner) and America and the West Indies, but absent from Central America.
C. palans Giglio-Tos, are known from Mexico through northern One new species is known from Panama.
992
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Melanomyza Malloch. This New World genus has one Oncodometopus Shewell. The single described species
described Neotropical species, M.  (Melanomyza) proana in this genus, O. umbrosus (Loew), is widespread from the
(Curran), known only from Panama. Several new species are eastern USA south through Colombia and Trinidad.
known from all over Central America, including Costa Rica,
South America, and the islands of the Caribbean. The other Pachyopella Shewell. All four described species in this
subgenus, M. (Lauxaniella), is strictly Nearctic. genus are Neotropical, with only one species, P. ornata (Me-
lander), occurring in Central America from southern Mexico
Minettia Robineau-Desvoidy. This large, nearly cos-
mopolitan genus has 21 valid species known from the through Panama. One of the remaining species is only known
Neotropical Region, all of which are thought to be in the from Jamaica, and the other two occur in South America.
subgenus Minettia. In the New World, an additional 20 valid One of these latter species, P.  flavida (Wiedemann), is re-
species are known from the Nearctic Region, 15 of which corded from Florida and Argentina, suggesting it is either
are in this subgenus, and five from the subgenus Frendelia, introduced or one of these is an erroneous record. Several
which is widespread but does not occur in the Neotropical new species are known from Central America and the West
Region. None of the described Neotropical species of Mi- Indies.
nettia (sensu stricto) are known from Central America, but
M. hubbardii (Coquillett) occurs in northern Mexico into the Physegeniopsis Gaimari & Silva. This eurychoromyiine
southwestern USA, and M. aibonito Curran occurs in Puerto genus has three described species; P. albeto and P. ankhoidea
Rico. Many new species are known from the New World, Gaimari & Silva are known only from Costa Rica, and P. had-
including Central America. rocara Gaimari & Silva is known from Costa Rica through
Minilauxania Papp & Silva. The single species in this Ecuador, Bolivia, and Brazil. Additional undescribed species
genus, M. bulbifacies Papp & Silva, is known only from Cos- are known from Brazil and Ecuador. A key to the described
ta Rica. There is at least one new species in Costa Rica. species is provided in Gaimari & Silva (2010).

Neodeceia Malloch. As far as is known, the species of Physegenua Macquart. The 11 species in this genus are
this genus occur primarily on the islands of the Caribbean, all Neotropical, with six species known from Costa Rica.
although some species occur in southern Florida, and there Although the key provided by Curran (1942) is incomplete
are doubtful records in South America. One species, N. exul for the genus, it does include all of the described Central
(Williston), is known from Honduras, and one new species is American species.
known from Costa Rica and Panama.
Physoclypeus Hendel. The 14 described species in this
Neogriphoneura Malloch. All 11 valid species of this ge- genus are all Neotropical, with one species, P.  coquilletti
nus are Neotropical. Three species, N. sordida (Wiedemann),
Hendel, being known from the southeastern USA, the Great-
N. striatifrons Hendel, and N. striga Curran, are known from
the eastern or southeastern USA, also occurring in South er Antilles, and Guatemala. Two other species are known
and Central America. Two species, including N.  sordida, from the Greater Antilles, one of which, P. hendeli Mello &
are known from the islands of the Caribbean. Two species, Silva, is also known from southern Mexico to Costa Rica.
N. laevifrons (Hendel) and N. tertia Curran, are known from Two species, P. zebrinus Mello & Silva and P. unimaculatus
Central and South America, and N. timida Curran is known Mello  & Silva, are known from Central America, with the
throughout Central America. The remaining four species are former species extending through northern South America.
strictly South American. Mello & Silva (2008a) recently re- The remaining nine species are known only from South
vised this genus and provided a key to all species. America. Mello & Silva (2008b) recently revised this genus
and provided a key to all species.
Neominettia Hendel. This strictly Neotropical genus has
six described species, only one of which, N. contigua (Fab- Platygraphium Hendel. The single species of this genus,
ricius), occurs in Central America and is widespread from
P. penicillatum Hendel, is known from Panama and Bolivia.
southern Mexico south to Brazil and Bolivia. The remaining
species are all from South America, although several new Poecilolycia Shewell. This mainly Nearctic genus has
species are known.
only three species described from the Neotropical Region,
Neoxangelina Hendel. Only one of the four described one from Jamaica and two from Chile, with none known
species, N.  ornata (Schiner), occurs in Central America, from Central America. At least one new species is known
where it is known from Costa Rica and Panama, south into from Guatemala and Costa Rica and another from Neo-
Venezuela and Guyana. One new species is known from tropical Mexico.
Guatemala.
Poecilominettia Hendel. This large (74 described spe-
Ocellominettia Hendel. The two species in this genus are cies) genus was regionally revised, for Panama, by Broad-
known from South America, but two new species are known head (1989), including keys to species. More than half of
from Neotropical Mexico and Venezuela. the species are known or described from Barro Colorado
LAUXANIIDAE (lauxaniid flies) 71 993

Island, Panama, and 12 additional species are known from from Costa Rica and is also known from Panama and Vene-
other parts of Panama; others are known from throughout the zuela; S. coronata (Broadhead) and S. ficulnea (Broadhead)
Neotropical Region. Several new species are known from the are both known only from Panama. Broadhead (1989) pro-
West Indies, although the genus was treated relatively com- vided a key to four of the five known species of the genus
pletely by Broadhead (1989). (but as Floriminettia Broadhead).

Procrita Hendel. The single valid species in this genus, Scutominettia Hendel. Both described species in this
P. pectinata Hendel, is known from Mexico and Costa Rica. genus are found in Central America. Scutominettia innuba
One possibly new species is known from Panama. (Giglio-Tos) is known only from Mexico and Costa Rica,
and S.  mallochi Hendel is known from Mexico through
Pseudocalliope Malloch. Several described species occur Venezuela, Brazil, and Paraguay. One new species is
in the New World, with none recorded from Central Amer- known from Panama, and several new species are known
ica, although P. eucephala (Loew) is known from northern from Venezuela and Peru.
Mexico, and one described species is known from Argen-
tina. Additionally, several new species are known from Costa Setulina Malloch. The four known species in this genus
Rica and Neotropical Mexico, and at least 10 new species are are all found in Central America. Setulina geminata (Fabr-
known from the southwestern USA. Species of this genus icius) and S. interjecta (Walker) are widespread from Mex-
are most commonly collected in flowers of Yucca. ico and throughout Central America into northern South
America. Setulina prima Malloch and S. stata (Giglio-Tos)
Pseudogriphoneura Hendel. This large (28 described are both known from Mexico, with the former species also
species) genus is known only from the Neotropical Region. occurring in Guatemala.
Seven species are known from Costa Rica, one is known only
from Mexico, and 11 species are known only from Panama. Siphonophysa Hendel. This genus has two described
The remaining 12 species are known only from South Amer- species, although many new species are recognized in Cen-
ica. One new species is known from Costa Rica. tral and South America. Of the two described species, only
S.  albipes (Coquillett) is known from Central America,
Pseudominettia Papp & Silva. One of the two species, from Mexico through Panama. The other species is from
P. platypeza Papp & Silva, was described from Brazil, but is South America. Numerous new species are known from the
also known from Neotropical Mexico. The other species is Neotropical Region, including many from Costa Rica and
found on the islands of the Caribbean. the rest of Central America.
Ritaemyia Frey. The described species are known from Stenolauxania Malloch. Two of the seven described
Brazil and Colombia. Several new species are also known species, S. striata Malloch and S. nubecula (Malloch), are
from Costa Rica and from the collections of Broadhead in known from Costa Rica, with the latter species also known
Panama. from Panama. In addition, several undescribed species oc-
cur in Central America, five of which occur in Costa Rica.
Sapromyza Fallén. The Neotropical species currently
recognized as members of this genus are likely not “true” Tauridion Papp  & Silva. The single species in this
Sapromyza, as this is the largest “dumping-ground” genus genus, T.  shewelli, was described from a single specimen
in the family worldwide, with nearly 300 described species. from Peru. Gaimari & Silva (2010) redescribed the species,
It is likely that the Neotropical fauna does not have any true recognizing it as a member of the subfamily Eurychoro-
members of this genus, yet many specimens are sorted to this myiinae and recording it from Costa Rica, Bolivia, and
name. The numerous currently valid species in this genus Ecuador.
from the Neotropical Region are a reflection of the need to
work out their true affinities. The conspectus of Neotropical Trigonometopus Macquart. Like Sapromyza, although
lauxaniids in progress by the authors addresses nearly all of not nearly as species-rich, this genus is considered to be
these species. That said, all except three of these species are cosmopolitan, but more likely represents numerous inde-
from South America. One species, S. picticornis (Curran), is pendent lineages. Six described species are recognized in
known from Panama, and another, S. vinnula Giglio-Tos, is the Neotropical fauna, including three only known from
known from Neotropical Mexico. The remaining non-South islands of the Caribbean, and one known only from South
American species, S. slossonae Coquillett, is from southern America. Many new species are known from Central and
Florida and Puerto Rico. South America and the West Indies, including at least four
from Costa Rica.
Sciosapromyza Hendel. The five described species in this
genus are all Neotropical, three of which are Central Ameri- Trisapromyza Shewell. Of the two species of this genus,
can. Sciosapromyza tinctinervis (Malloch) was described one is only known from South America, and the other,
994
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

T. vittigera (Coquillett), is distributed mainly in the USA and are only known from South America and one from South
Canada, but extends south through Costa Rica. America and the West Indies. The remaining two species,
X. flavipennis (Fabricius) and X. pallida Malloch, are known
Trivialia Malloch. Of the seven described species in this from Central and South America, the former also occurring
genus, six are Neotropical and one is found only in the east- in the Nearctic. Many new species are known from Central
ern USA. Of the Neotropical species, only three are known and South America and the islands of the Caribbean.
from Central America. Trivialia celeste (Curran) and T. ni-
groapicata (Malloch) are known from Panama, and T. puella Xenopterella Malloch. This genus has two valid spe-
(Williston) is known from Costa Rica, Panama, and the West cies, one of which is Nearctic. The other species, X. obliqua
Indies (St. Vincent). One of the remaining species is known Malloch, is known only from Neotropical Mexico. Steyskal
only from Ecuador, and the other two are only known from (1965) is the most current resource for identifying the spe-
the West Indies (St. Vincent). At least five new species are cies of this genus.
known from Central America and the West Indies.
Undescribed Genus A. The single undescribed species in
Trypetisoma Malloch. Most of the 24 currently valid this undescribed genus is known only from southern Mexico
species of Trypetisoma (sensu stricto) are from Australia, al- and Guatemala.
though the type species is from the New World. Of the four
New World species, two are Neotropical, only one of which Undescribed Genus C. Two new species in this unde-
is known from Central America, T. zacatecasense Arnaud & scribed genus are known from Costa Rica, one of which also
Gelhaus, from central Mexico. Arnaud (1968) provided a occurs in Brazil.
key to the three American species known at the time, which
is supplemented by the addition of the Central American Undescribed Genus K. Only one of the four species
species by Arnaud & Gelhaus (1980). in this undescribed Neotropical genus, Sapromyza sororia
Williston, occurs in Central America, also occurring on the
Xeniconeura Shewell. The single species in this genus, islands of the Caribbean. The remaining species are all in
X.  costalis (Curran), is distributed in Neotropical Mexico, South America.
extending into the southwestern USA.
Undescribed Genus L. The single species of this unde-
Xenochaetina Malloch. This genus is represented by 15 scribed genus is Sapromyza frontalis Macquart, described
described species in the Neotropical Region, 12 of which from Brazil, but also known from Costa Rica.

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Broadhead, E.C. 1984. Adaptations for fungal grazing in lauxaniid tor). Immature Insects, Volume 2. Kendall/Hunt Publishing,
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Chandler, P. 1978. Associations with plants. Fungi, pp.  199–211. mann A.B., Helsinki, 63 pp.
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ix + 255 pp. 2342: 1–64.
Colless, D.H. & D.K. McAlpine. 1970. Diptera (flies), pp. 656–740. Hardy, D.E.  & M.D. Delfinado. 1980. Family Lauxaniidae,
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Press, Melbourne, 1029 pp. Volume 13. Diptera: Cyclorrhapha III. University Press of Ha-
Collin, J.E. 1933. Five new species of Diptera. Entomologist’s waii, Honolulu, 451 pp.
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Collin, J.E. 1948. A short synopsis of British Sapromyzidae (Dip- tgarter Beiträge fur Naturkunde 145: 1–215.
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Hicks, E.A. 1953. Observations on the insect fauna of birds’ nests. Papp, L.  & A.I. Shatalkin. 1998. Family Lauxaniidae, pp.  383–
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
CHAMAEMYIIDAE (chamaemyiid flies) 72
Stephen D. Gaimari

Fig. 72.1. Female of Chamaemyia polystigma (Meigen), (Holarctic, MND, fig. 88.1).

Diagnosis posteriorly (Figs.  4, 6); lunule small, narrowly exposed,

and bare (Figs. 1–4) to large, broadly exposed, and setulose
Small, compact-bodied or elongate (Fig.  1) flies (body (Figs.  5–7); vibrissa absent; antenna short (rarely twice as
length 1–6 mm); usually densely silvery-gray pruinose, occa- long as high, Fig. 2), porrect; first flagellomere rounded or
sionally brown or polished black; often with brown to gold- pointed and upturned apically; subbasal arista bare or short
ish vittae on scutum and/or brown to black pruinose spots or pubescent, not plumose. Scutum setulose, not hairy; with one
bands on abdominal tergites. Head either compact, shorter to four (rarely five) dorsocentral setae, either all postsutural
than high and rounded anteriorly (Figs. 5–7) to subequal or (Fig. 8) or with one presutural (Fig. 1); one postpronotal, one
longer than high and rounded (Fig. 4) or pointed (Figs. 1–3) presutural (absent in Cremifania Czerny), and one postsutural
anteriorly; postocellar setae convergent or absent; ocellar supra-alar, one postalar, and two notopleural setae present;
triangle occasionally enlarged, prominent (Fig.  7); ocellar prescutellar acrostichal seta present or absent; prescutellum
setae normal-sized and proclinate (Figs.  1–2, 4), reduced usually present (Fig. 8) (absent in Anchioleucopis Tanasijt-
and reclinate (Figs.  5–6), or absent; fronto-orbital setae shuk and the Palearctic genus Lipoleucopis Meijere); scutellum
absent or with two (rarely three, Fig. 6) reclinate setae, either with two pairs of setae, otherwise bare; anepisternum with
distributed evenly on frontal orbits (Figs.  1–3) or placed or without seta(e) or setulae; anepimeron bare (except some

997
998
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Cremifania); katepisternum with one strong seta in addition Eggs ovoid, either uniformly rounded at each end or
to setulae; proepisternal and proepimeral setae absent; pros- slightly tapering anteriorly; chorion sculptured, either with
ternum bare. Wing (Figs. 10–12) with C unbroken, extending distinct longitudinal ridges, some anastomosing, or with
to M (R4+5 in Lipoleucopis, Fig. 12); Sc complete, separated more reticulated pattern of ridges; with small depressed
from R1; cells  bm and dm usually separated by crossvein micropylar opening anteriorly; posteriorly with small pad
bm-cu (except Melaleucopis); cell cup present; A1 short, not with several large pores of unknown function.
reaching wing margin; membrane sometimes patterned (e.g., Larvae musciform, tapering anteriorly and rather blunt
Fig.  11), hyaline or milky white. Legs lacking preapical posteriorly (Fig.  13), usually somewhat dorsoventrally
dorsal tibial seta. Abdominal tergites with (Fig. 1) or without compressed, with conspicuous secondary annulations dorsally,
(Fig. 9) paired brown to black pruinose spots or bands. each appearing as three pseudosegments covered with small

poc s
p frorb s i vt s
oc s

o vt s
lun

sbvb s

2 Ortalidina
3 Plunomia 4 Pseudodinia

5 Neoleucopis 6 Leucopis (Xenoleucopis) 7 Leucopina

pprn s

a npl s

presut
spal s

spal s
presctl acr s
presctl
pal s
dc s

8 Leucopomyia 9 Anchioleucopis

Figs. 72.2–9. Heads, thorax, and abdomen: dorsolateral view of head of (2) Ortalidina punctata (Coquillett), (Nearctic, MND, fig. 88.2);
(3) Plunomia elegans Curran, (Nearctic, MND, fig. 88.3); (4) Pseudodinia varipes Coquillett, (Nearctic, MND, fig. 88.4); (5) Neoleucopis
pinicola Malloch, (Holarctic, MND, fig. 88.6); (6) Leucopis (Xenoleucopis) sp., (Nearctic, MND, fig. 88.7); and (7) Leucopina ocellaris
(Malloch), (Nearctic, MND, fig. 88.8); dorsal view of thorax of (8) Leucopomyia pulvinaria (Malloch), (Nearctic, MND, fig. 88.13); dorsal
view of abdomen of female of (9) Anchioleucopis macalpinei Tanasijtshuk, (Nearctic, MND, fig. 88.14, as Leucopis geniculata).
Abbreviations: a npl s, anterior notopleural seta; dc s, dorsocentral seta; i vt s, inner vertical seta; lun, lunule; oc s, ocellar seta; o vt s, outer
vertical seta; pal s, postalar seta; p frorb s, posterior fronto-orbital seta; poc s, postocellar seta; pprn s, postpronotal seta; presctl, prescutellum;
presctl acr s, prescutellar acrostichal seta; presut spal s, presutural supra-alar seta; sbvb s, subvibrissal setae; spal s, supra-alar seta.
CHAMAEMYIIDAE (chamaemyiid flies) 72 999

Sc R1 Sc R1

R2+3
bm R4+5
CuA2 dm
cup
A1+CuA2
M1

CuA1

10 Cremifania 11 Ortalidina

12 Lipoleucopis

m psdsg

p spr sk

13 Leucopis

14 Leucopis 15 Undescribed Genus C

Figs. 72.10–15. Wings and larval structures: dorsal view of wing of (10) Cremifania nigrocellulata Czerny, (Palearctic, introduced Nearctic,
MND, fig. 4.61); (11) Ortalidina punctata (Coquillett), (Nearctic, MND, fig. 88.9, as Toropamecia); and (12) Lipoleucopis praecox
de  Meijere, (Palearctic, MND, fig. 88.11); lateral view of third instar larva of (13) Leucopis (Leucopis) simplex Loew, (Nearctic, MND,
fig. 88.21); lateral view of cephalopharyngeal skeleton of (14) Leucopis (Leucopis) simplex Loew, (Nearctic, MND, fig. 88.18); lateral view
of posterior spiracle of (15) Undescribed Genus C, (Nearctic, MND, fig. 88.20, as Leucopis (Leucopis) sp.).
Abbreviations: m psdsg, middle pseudosegment; p spr sk, posterior spiracular stalk.
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

chaetoids (rarely smooth); middle pseudosegment usually leucopis Malloch (Clark & Brown, 1957; McAlpine, 1971;
with transverse row of enlarged chaetoids or fleshy spines; Sluss & Foote, 1973), Anchioleucopis (Tanasijtshuk, 1997),
usually with lateral (not shown in generalized Fig. 13) and Undescribed Genus C (Kohler et al., 2008), and Cremifania
ventral welts. Anterior spiracles protruding laterally, arranged (Delucchi & Pschorn-Walcher, 1954; Clark & Brown, 1962);
fanwise with 3–12 lobes. Posterior spiracles usually on Cinara aphids on gymnosperms are attacked by species of
distinctly separated divergent stalks (Fig.  13) (flattened in Undescribed Genus B and possibly Undescribed Genus C;
Echinoleucopis Gaimari  & Tanasijtshuk and Leucopomyia scales and mealybugs on dicots are attacked by species of
Malloch), each with three lobes bearing one opening each Melaleucopis (Beingolea, 1957) and Leucopis (Leucopella)
(Fig.  15); first instar with only two lobes on short stalk (Malloch, 1927; James, 1934; Gaimari  & Raspi, 2002);
bearing one opening each. Cephalopharyngeal skeleton eggs within ovisacs of coccoids are preyed upon by species
(Fig.  14) simple; dental sclerite usually absent (present in
of Echinoleucopis (Griot, 1954; Gaimari  & Tanasijtshuk,
Cremifania), hypopharyngeal and tentoropharyngeal sclerites
2001), Leucopomyia (Malloch, 1922; Tanasijtshuk, 1959;
fused, parastomal bar absent; gular region sometimes with
Babaev & Tanasijtshuk, 1971; Kaydan et al., 2006), and Un-
cephalic ring.
Puparia subovate, anterior end abruptly tapered; usually described Genus A. Some genera have broader feeding strat-
slightly dorsoventrally compressed, more or less convex egies, such as Leucopina attacking various coccoids, from
dorsally (strongly convex in Echinoleucopis) and flattened mealybugs to scales, from free-living to feeding on eggs
ventrally; anterior end usually dorsoventrally flattened to within an ovisac, and species of Leucopis sensu stricto feed-
produce lateral ridge; surface sculpturing and welts reflect ing on almost the entire range of available prey. Genera with
third instar conditions, but more flattened; anterior and pos- unknown feeding habits are often found to be specifically asso-
terior spiracles as in third instar; usually with hardened shiny ciated with particular habitats frequented by particular poten-
black droplet present posteriorly, excreted at pupariation. tial hosts. For example, species of Ortalidina and Plunomia
Families superficially similar to chamaemyiids (i.e., small Malloch are always swept from grasses, but have never been
gray acalyptrates, such as some heleomyzids, chloropids, directly associated with the mealybugs in the leaf sheaths
odiniids, ephydrids, tethinids) can be readily separated from that they are likely feeding upon. The immatures likely remain
them by the following chamaemyiid characteristics: zero to undiscovered simply because of a lack of collecting effort.
two reclinate fronto-orbital setae; vibrissa absent; postocellar Species in many genera feed in tight enclosures where other
setae convergent or absent; C unbroken; Sc complete, separated predators cannot typically enter, including the above, and ad-
from R1; presence of a prescutellum (absent only rarely), and ditionally within galls of Phylloxeridae (Stevenson, 1967) and
lack of preapical dorsal tibial setae. Chamaemyiids can be Aphididae (Raspi, 1990; Valenti & Gaimari, 1992; Valenti et
separated from the most similar family, the closely related al., 1996), within leaf sheaths of grasses housing colonies of
Lauxaniidae, on the basis of the developed prescutellum, the aphids (Gaimari & Turner, 1996a, 1998) or mealybugs, and
lack of distinct dense black costal spinules, the absence of within root systems infested with aphids or mealybugs. With
preapical dorsal tibial setae, and in some genera the lack of their predaceous habits, exotic species have been introduced
fronto-orbital and/or anepisternal setae.
into North America for biological control efforts against Rus-
sian wheat aphid, Diuraphis noxia (Mordvilko) (Gaimari &
Biology Turner, 1996b; Noma et al., 2005) and the balsam woolly
adelgid, Adelges picea (Ratzeburg) (McAlpine, 1978), and
All known members of the family are predators on against the Eurasian pine adelgid, Pineus pini (Macquart) in
sternorrhynchous aphidoid or coccoid Hemiptera, although Hawaii (Culliney et al., 1988; Greathead, 1995).
one species has been reported to attack psylloid Hemiptera Beyond host records, larval and adult behavior is poorly
as well (McLean, 1998a). Specific biological data are known known. McLean (1992) made mostly anecdotal observations
for few Central American species, and specific host informa- on larval feeding on aphids and their being ignored by the
tion is spotty for known chamaemyiid species outside the ants tending the aphids, on the adults approaching an aphid
Palearctic Region, although predatory habits appear to be rel- colony to oviposit and their avoidance of ants. Additionally,
atively uniform at the genus-group level (Ferrar, 1987). As a the adults are known to stroke the aphids with their foretarsi
group, they display a spectrum of feeding strategies from ac- to elicit honeydew secretion (presumably mimicking the ants
tive predation within free-living prey colonies to completely antennal actions for the same purpose), which is fed upon by
sessile predation on eggs within a single coccoid egg sac. the adult flies. Gaimari & Turner (1998) made observations of
The following are several more or less genus-specific feeding larval searching and feeding behavior by instar, molting and
strategies: mealybugs in leaf sheaths of grasses are attacked pupariation, adult courtship behavior (including wing flutter-
by species of Chamaemyia (Tanasijtshuk, 1970; Sluss  & ing, wing extension, body postures, and touching), and ovi-
Foote, 1973; Raspi, 1983), Parochthiphila (Tanasijtshuk, position. Gaimari & Turner (1996b) studied the differential
1963, 1968; Raspi, 1983), and Pseudodinia (Barber, 1985); feeding and developmental rates of two species of Leucopis
adelgids on gymnosperms are attacked by species of Neo- at varying temperatures and aphid densities, demonstrating
CHAMAEMYIIDAE (chamaemyiid flies) 72 1001

their life cycle and effectiveness in destroying aphid popula- apical bend of vein Sc, absence of a preapical dorsal tibial
tions. Interestingly, Smith & McLean (1998) discussed aerial seta, presence of heavily gray pruinose body vestiture, and
distribution of the puparium of a species via airborn thistle the length of tergite 6 in males being less than half that of
pappus elements (on which plant the prey aphids were feed- tergite 5. Griffiths (1972) disagreed with McAlpine’s initial
ing), including several pappus feather elements being firmly (1963) assessment and treated Cremifaniinae at the family
glued to a single puparium, ensuring a longer airborn pe- level as an unplaced member of the Sciomyzoidea, separated
riod and farther dispersal. Also of great interest, Eisner et al. from the Lauxanioidea by the asymmetry of male abdominal
(1994) reported on a species of Leucopina feeding on Dacty- segments 6–8 and the articulated phallic sclerites that allow
lopius spp. scales and using dietarily acquired carminic acid swinging through a wide arc against the phallapodeme to
(the red dye from cochineal bugs) for defense. The liquid reach a more anteriorly directed position. Griffiths’ (1972)
was stored in a special rectal pouch and in the Malphigian opinion was later followed by Tanasijtshuk (1984, 1992),
tubules, and larvae secreted a droplet from the rectum in re- although McAlpine (1989) pointed to several misinterpreta-
sponse to being walked over or attacked by ants. In some tions by Griffiths, including, for example, the complex phal-
species, just prior to pupariation, a sticky liquid is secreted lic structure found in many lauxaniids functioning similarly
from the anus, likely from the Malpighian tubules. This liq- to Cremifania, allowing for the possibility that this sort of
uid is spread over the entire ventral surface of the larva and arcing phallus could be in the lauxanioid ground plan, which
hardens after pupariation, adhering the puparium to the sub- was secondarily changed (an autapomorphy) in his Chamae-
strate (Gaimari & Turner, 1998). After this secretion, the gut myiinae and some lauxaniids. The same can be said for the
purges a thin, sticky black fluid that hardens to form a char- presence of three spermathecae in Cremifania, a character-
acteristic shiny black droplet attached to the posterior end of istic seemingly in opposition to the four spermathecae typi-
the puparium (Gaimari & Turner, 1998). cally cited in other chamaemyiids (Papp, 1998). Not only is
Morphology of the known immature stages is summarized this condition consistent with the ground plan of Lauxan-
in part by Ferrar (1987). Details and extensive figures are pro- ioidea, but other chamaemyiid genera have been found with
vided for all stages and instars for species (none Neotropical) two, three, or four spermathecae (e.g., Echinoleucopis), so
in the following genera: Pseudodinia (Barber, 1985), Leu- this characteristic provides no definitive data to contradict
copis (Raspi, 1983, 1990; Gaimari & Turner, 1996b), Leu- the placement of Cremifania within the family.
copina (Sluss & Foote, 1971), Neoleucopis (Sluss & Foote, In McAlpine’s (1963, 1989) scheme, which is followed
1973; Gaimari et al., 2007), Chamaemyia (Sluss  & Foote, herein, Chamaemyiinae are supported by four autapomor-
1973; Raspi, 1983), and Parochthiphila (Raspi, 1983). Eggs phies, including the lack of discrete surstyli on the epan-
are discussed generally by Tanasijtshuk (1974) for several drium, fusion of the phallapodeme to the hypandrium, and
genera, including Leucopis, Chamaemyia, Parochthiphila, uniform sclerotization of the phallus into a rigid and pos-
and Acrometopia Schiner, noting some fundamental differ- teroventrally directed structure. Chamaemyiinae are divided
ences in surface sculpturing among the genera. Mazzini & into two tribes, Chamaemyiini and Leucopini, with Chamae-
Raspi (1983) provide extensive figures and descriptions of myiini weakly supported by the reduction of the gonopods,
eggs for species of Parochthiphila and Chamaemyia. Larvae and Leucopini more strongly supported by the lunule being
and puparia have also been figured and discussed briefly in broadly exposed, the absence of postocellar setae, reduction
various revisionary studies. or absence of fronto-orbital setae, the atrophied wing vein A1
beyond CuA2, and tergite 6 in males being lost or fused with
syntergosternite 7+8. Relationships among chamaemyiid
Classification genera have been considered by Tanasijtshuk (1986, 1992,
2003). In considering relationships among 16 (Tanasijtshuk,
The family group was proposed by Hendel (1910), but was 1992) and 22 (Tanasijtshuk, 2003) chamaemyiid genera, a
initially treated by Malloch (1922) as a subfamily of Agro- hierarchical classification was proposed with two subfami-
myzidae, a position he later abandoned (Malloch, 1930) to lies, Chamaemyiinae and Leucopinae, which were eleva-
follow Hendel (1916) in allying the family with Lauxaniidae. tions in rank for the tribes recognized by McAlpine (1963)
McAlpine (1963) divided the family into two subfamilies, after exclusion of Cremifaniinae. Chamaemyiinae sensu
Cremifaniinae and Chamaemyiinae, which was strongly en- Tanasijtshuk (2003) were subdivided into three tribes, two
dorsed by Hennig (1965), and McAlpine (1989) cited at least of which occur in the New World, Acrometopiini (including
10 autapomorphic character states supporting this position, Ortalidina — as Toropamecia Cogan, Parapamecia, and Pa-
including larval predation on sternorrhynchous hemipterans, rochthiphila in the Neotropical Region) and Chamaemyiini
fusion of the larval hypopharyngeal and tentoropharyngeal (including Chamaemyia and Pseudodinia found in the Neo-
sclerites and the parastomal bar with the hypopharyngeal tropical Region). A dendrogram was presented with characters
sclerites, presence of unbranched interspiracular hairs on mapped, although the conclusions were largely intuitive,
the posterior spiracles, a bare adult prosternum, absence of based on relatively few characters (many of which represent
a proanepimeral seta, flexion of wing vein  R1 towards the symplesiomorphies) without quantitative analysis, leaving
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

the monophyly of “Chamaemyiinae” in question. The char- to the genera and species, but many undescribed species
acteristics supporting Leucopinae (i.e., Leucopini) are more remain. Resources to identify Neotropical chamaemyi-
convincing as synapomorphies, although quantitative analysis ids are sparse. Few generic revisions exist that include
is still lacking. Neotropical species, such as Barber’s (1985) revision of the
mainly Nearctic genus Pseudodinia, and McAlpine’s (1971)
revision of the more widespread (but rare in the Neotropi-
Identification cal Region) genus Neoleucopis. Several Neotropical genera
have also been described or revised, such as Echinoleuco-
Keys to genera are available for the Nearctic Region pis (Gaimari & Tanasijtshuk, 2001) in the southern portion
(McAlpine, 1987) and the Palearctic Region (McLean, of South America, Ortalidina (as Toropamecia) throughout
1998b), although Cremifaniinae was treated as a separate the Neotropical Region (Cogan, 1978), and Melaleucopis
family in the latter manual by Papp (1998), with a key to in Peru and the West Indies (Sabrosky, 1957); the latter two
the three species of Cremifania provided. The resources for genera include recognized new species. Otherwise, the lit-
the Palearctic Region are most complete, with much of the erature consists mainly of isolated descriptions of species
fauna revised by Tanasijtshuk (1986). For the Australian and smaller faunal treatments.
fauna, Tanasijtshuk (1996) provides descriptions and keys

Key to the genera of Chamaemyiidae of the New World

1. Pterostigmal space darkly clouded, subequal in length to cell  c (Fig.  10); prescutellum and
presutural supra-alar seta absent; male with movable bilobate surstylus and flexible phallus with
spinelike subapical armature (predators of Adelgidae); CREMIFANIINAE; Holarctic Region. . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Cremifania Czerny]
– Pterostigmal space usually lightly clouded, seldom even half as long as cell c (Figs. 11–12);
prescutellum usually and supra-alar seta always present (Fig.  8); male with fixed surstylus,
phallus rigid and without spinelike armature; CHAMAEMYIINAE . . . . . . . . . . . . . . . . . . . . . . 2

2. Body compact, stout posteriorly; head (Figs. 5–7) with lunule high and often with setulae; fronto-
orbital setae present or absent; postocellar setae absent; ocellar setae absent (Fig.  7) or small;
Leucopini. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
– Body elongate, tapering posteriorly (Fig. 1); head (Figs. 1–4) with lunule low, bare; one to two
strong fronto-orbital setae usually present; postocellar setae present; ocellar setae strong,
proclinate (absent or minute in some species of Plunomia, Fig. 3); “Chamaemyiini”. . . . . 15

3. Ocellar setae present, proclinate and diverging; two fronto-orbital setae present (posterior one
subequal to inner vertical seta, anterior one slightly weaker) with anterior seta equidistant
between posterior seta and lunule; prescutellar seta present (predators of Ceroplastes scales);
Brazil . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Undescribed Genus A]
– Ocellar setae absent, or present and reclinate (Figs. 5-6), but if proclinate and divergent then
one fronto-orbital seta present (Melaleucopis); otherwise, fronto-orbital setae usually absent, ot
weak and posteriorly located (Fig. 6) or rarely with more than four along eye margin; prescutellar
setae present or absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4. Body without pruinosity, displaying shiny black ground color; ocellar setae present, proclinate and
diverging; with one strong fronto-orbital seta; crossvein bm-cu stublike, incomplete, not fully
separating cells dm and bm (predators of Ortheziidae). . . . . . . . . . . . . . . . Melaleucopis Sabrosky
– Body with at least some pruinosity, usually silvery-gray, occasionally goldish-brown, dull gray or
brown; rarely, abdomen partially shiny; crossvein bm-cu complete, separating cells dm and bm
(Figs. 11–12) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

5. Anepisternum with one to several setae, setulae, or both, along posterior edge (predators of
Coccoidea); Afrotropical Region, South America. . . . . . . . . . . . [Leucopis (Leucopella) Malloch]
– Anepisternum lacking setae or setulae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
CHAMAEMYIIDAE (chamaemyiid flies) 72 1003

6. Ocellar setae lacking in both sexes; ocellar triangle neither densely pilose nor fuzzy in males,
but if moderately fuzzy (i.e., not completely covering area), then posterior ocelli equidistant in
comparison with distance from one posterior ocellus to anterior ocellus. . . . . . . . . . . . . . . . . . . 7
– Ocellar setae well developed and reclinate (Fig. 6), or ocellar triangle (males only) densely pilose
and fuzzy with posterior ocelli widely separated (Fig. 7). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

7. Prescutellar acrostichal seta present (Fig. 8). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

– Prescutellar acrostichal seta absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

8. Prescutellum present; head higher than long; scutum with dorsoventral vittae (predators of eggs
within coccoid ovisac); Holarctic Region . . . . . . . . . . . . . . . . . . . . . . . . . [Leucopomyia Malloch]
– Prescutellum absent; head with length and height subequal; scutum unicolorous gray, lacking
vittae (predators on Adelgidae); Holarctic Region. . . . . . . . . . . . . . [Anchioleucopis Tanasijtshuk]

9. Abdomen shiny black with only sparse pruinosity; frons with one to two rows of stiff black
lateroclinate setulae along eye margin; in males, frons protruding rooflike over lunule, and with
enlarged laterally directed setulae in anterolateral corners (sometimes clumped into hornlike pencils);
in female, maxillary palpus greatly enlarged (subequal to first flagellomere) and flattened
(predators of Cinara aphids on Pinus); western North America. . . . . . . . [Undescribed Genus B]
– Abdomen pruinose, often with spots but not shiny black; frons not protruding over lunule, lacking
stiff lateroclinate setulae, and lacking enlarged laterally directed setulae in anterolateral corners;
maxillary palpus normal, much smaller than antennal first flagellomere . . . . . . . . . . . . . . . . . . 10

10. Scutum unicolorous gray, lacking vittae; frons flat, sometimes slightly concave medially, ending
at 90° angle with upper edge of lunule; plane of face and parafacial at an acute angle with frons
(predators of Adelgidae); Holarctic Region. . . . . . . . . . . . . . . . . . . . . . . . [Undescribed Genus C]
– Scutum with dorsocentral vittae (rarely absent); frons gently curved and sloping through lunule;
plane of face and parafacial at obtuse angle with frons. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

11. Head high, height to length ratio nearly 2:1; eye height to length ratio 2:1; lunule high, subequal
in length to frons, with proclinate black setulae; upper part of frontal orbit with one or two slightly
enlarged setulae; anterior ocellus surrounded by ring of short black setulae (predators of eggs
within coccoid ovisac); Argentina, Brazil, Chile. . . . . . [Echinoleucopis Gaimari & Tanasijtshuk]
– Neither head, eye, nor lunule so high; lunule setulose or not, or with white hairs; upper part of
frontal orbit lacking enlarged setulae; ocellar area sometimes setulose, but not in ring around
anterior ocellus (predators on various Sternorrhyncha). . . . . . . . . . . Leucopis (Leucopis) Meigen

12. Prescutellar acrostichal seta present (as in Fig.  8); body dull, dark brown to grayish pruinose,
lacking dorsocentral vittae on scutum (predators on Adelgidae and rarely Coccoidea); Holarctic
Region, Chile, Ecuador . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Neoleucopis Malloch]
– Prescutellar acrostichal seta absent; body silvery to silver–gray pruinose, usually with goldish to
brownish dorsocentral vittae on scutum. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

13. Fronto-orbital area lacking enlarged setulae; ocellar plate greatly expanded, posterior ocelli widely
spread, much farther apart than distance between anterior and one posterior ocellus; ocellar setae
often absent in males (present in females) that instead have ocellar plate densely pilose and fuzzy
(Fig. 7) (predators of various coccoids). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leucopina Malloch
– Fronto-orbital area usually with enlarged setulae (e.g., Fig. 6) or setae; ocellar plate not expanded,
pilose nor densely fuzzy, but posterior ocelli sometimes slightly farther apart than distance to
anterior ocellus; ocellar setae present in both sexes (Fig. 6). . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

14. Upper part of fronto-orbital area with one to two (rarely zero or three) enlarged setulae (Fig. 6)
(predators of various coccoids) . . . . . . . . . . . . . . . . . . . . . . . . . Leucopis (Xenoleucopis) Malloch
– Fronto-orbital area with series of five to six setae along eye margin from level of anterior ocellus
to lunule edge (possibly predators of Cinara aphids on Pinus); southwestern USA. . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Undescribed Genus D]

15. Anepisternum with one to several setae or setulae along posterior edge. . . . . . . . . . . . . . . . . . . 16
– Anepisternum bare (Fig. 1) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

16. Anepisternum with one strong seta; frons, thorax and/or abdomen often shiny black, or variously
dark grayish pruinose; setulae of frons, if present, not in transverse line; abdomen lacking black
pruinose tergal spots (predators of mealybugs in grasses). . . . . . . . . . . . . . Pseudodinia Coquillett
– Anepisternum with one to several setulae, with no outstanding seta; body entirely silvery-gray
pruinose, lacking areas of polished black; frons with transverse line of small black setulae between
anterior fronto-orbital setae, and often with accompanying transverse black band; abdomen
usually with paired black pruinose spots on some tergites (predators of mealybugs in grasses) . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parochthiphila Czerny, in part

17. Wing with various brown markings on membrane (Fig. 11), but not restricted to costal cell and
along veins (likely predators of mealybugs in grasses). . . . . . . . . . . . . . . . . Ortalidina Blanchard
– Wing hyaline (except one species, Chamaemyia fumicosta Malloch, with only costal cell and
some crossveins darkened). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

18. Scutum with four (occasionally five) dorsocentral setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

– Scutum with two or three dorsocentral setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

19. Scutum with four evenly spaced and evenly sized dorsocentral setae; lacking prescutellar
acrostichal seta; anterior fronto-orbital seta at lunule edge (Fig. 3), lacking transverse line of small
black setulae between (likely predators of mealybugs in grasses); Nearctic Region . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Plunomia Malloch]
– Scutum with four (occasionally five) dorsocentral setae, often with anterior 2 (or 3) reduced in
size; prescutellar acrostichal seta present; anterior fronto-orbital seta at least slightly removed
from lunule border, with transverse line of small black setulae between . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parochthiphila Czerny, in part

20. Scutum with three dorsocentral setae, including one presutural (Fig.  1); anterior fronto-orbital
seta varied in size and position (predators of mealybugs in grasses). . . . . . . Chamaemyia Meigen
– Scutum with two postsutural dorsocentral setae only; anterior fronto-orbital seta smaller than
posterior seta, and closer to posterior seta than to lunule (likely predators of mealybugs in grasses);
Caribbean, Nearctic Mexico. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Parapamecia Cogan]

Synopsis of the fauna fauna, including the islands of the Caribbean, consists of 38
described species in nine genera and subgenera, including
Although a small family, Chamaemyiidae have several only nine species in six genera from Central America. For
widespread genera and subgenera. Most, however, are rela- the entire region, dozens of undescribed species are known.
tively restricted in their distribution. Only Leucopis sensu Chamaemyia Meigen. This genus is nearly cosmopolitan,
stricto is truly cosmopolitan, being found on all continents but only a single described species, C. fumicosta extends into
except Antarctica; the genera Neoleucopis, Parochthiphila, Neotropical Mexico. Few specimens of this genus are known
and Chamaemyia are also widely distributed but are notably ab- from the Neotropical Region, but they represent several new
sent from some regions. Most genera and subgenera, though, species, including one from southern Mexico (Veracruz).
are regionally restricted, including several strictly in the All known members of this genus are predators as larvae on
New World and several even more restricted. The Neotropical mealybugs in the leaf sheaths of grasses.
CHAMAEMYIIDAE (chamaemyiid flies) 72 1005

Leucopina Malloch. Species in this genus attack scales species are known from southern Mexico and Brazil. All
and mealybugs on a variety of plants, some feeding on eggs are predators of species of Orthezia scales, feeding on eggs
within an ovisac; typically, each species attacks only one host within an ovisac (Beingolea, 1957).
type. Only two described species are known from the Neo-
tropical Region, both in Mexico. Leucopina bellula (Willi- Ortalidina Blanchard. This genus is nearly all Neo-
ston) is widespread in California and the southern United tropical, but extends into the Nearctic Region (southeastern
States and is widespread throughout Mexico. The larvae of United States) through the West Indies and Bahamas. Most
this species are egg predators within the ovisacs of scales in of the described species are from Chile, Argentina, and Bra-
the genus Dactylopius on cacti in the genera Opuntia and zil, with three known strictly from the Caribbean. One spe-
Platyopuntia. Mann (1969) and Gilreath  & Smith (1988) cies, O.  longipennis (Cogan), is known from Ecuador, and
discussed L.  bellula as predators of Dactylopius confusus another, O.  nigripalpis (Cogan), is known from southern
(Cockerell) on Opuntia species. Leucopina bivittata (Mal- Mexico, Guatemala, and El  Salvador. Several new species
loch) is widespread in California and the southern United are also known, including some from Neotropical Mexico
States, and some specimens are known from Neotropical (one from Tamaulipas, two from Chiapas, one from Micho-
Mexico. This species has been reared from numerous mealy- acán, and one from Veracruz), one from Costa Rica, and six
bug hosts, including species in the genera Anisococcus, Dac- from the Caribbean. Many additional undescribed species
tylopius, Lecanium, Phenacoccus, and Pseudococcus on a are known throughout South America. Cogan (1978) is the
variety of plants. At least five undescribed species occur in best resource for identifying species (as Toropamecia), but
the Neotropical Region, including three in Central America; many new species are known that may fit concepts in the
at least 12 new species are known from the Nearctic Region.
key. Also, the type species, O.  cellularis Blanchard, is not
The recognized new species include egg predators within
included.
ovisacs of mealybugs in the genera Planococcus, Phenacoc-
cus, Pseudococcus, and Ferrisia on a variety of plants, and Parochthiphila Czerny. This genus has not yet been re-
free-living predators of other coccoids, including species of corded from the New World, and no described species are
Orthezia, Lecanium, Dactylopius, and Olliffiella on a variety known from the Neotropical Region. However, several new
of plants.
species are recognized, including at least two from Costa
Leucopis Meigen. Subgenus Leucopis Meigen. This cosmo- Rica, of which one also occurs in Ecuador and the other also
politan subgenus is the largest in the family, including nearly in Mexico, and additional new species are known from Bra-
half of all described chamaemyiid species. Greatest diversity zil, the Dominican Republic, and the Nearctic Region. All
occurs in the northern temperate regions, where the species known members of this genus are predators of mealybugs in
attack mainly aphidoid Sternorrhyncha (Aphididae, Adel- leaf sheaths of grasses. They are often collected by sweep-
gidae, Phylloxeridae), although many also attack various ing grasses, although the biology of Parachthiphila species
coccoids. Despite its great diversity, few species are known in the Neotropical Region has not been observed. Some of the
from the Neotropical Region, probably because of the lack new species from the Neotropical Region differ from Parochthi-
of aphidoid diversity. Leucopis charactophalla Tanasijtshuk phila species from other parts of the world in having only
is known mainly from Texas and California (USA), but also two strong dorsocentral setae located posteriorly on the
occurs in Neotropical Mexico (southern Tamaulipas). Leuco- scutum (the anterior two are reduced in some, while reduc-
pis flavicornis Aldrich is known only from the southern tip tion or loss of only one of these is known to occur in other
of Texas (Brownsville) and likely occurs in adjacent areas of species).
northern and eastern Mexico. Many new species are known
throughout the New World, including Central America. Pseudodinia Coquillett. This mainly Nearctic genus has
Subgenus Xenoleucopis Malloch. The few described three described Neotropical species, P.  meridionalis Hen-
Neotropical members of this genus are from southern South nig from southern Mexico (Chiapas) through Panama and
America, but undescribed species are known from the P.  tuberculata Barber and P.  obscura Barber from south-
Nearctic Region and throughout South America, with a few ern Mexico (Chiapas). The remaining five species found
specimens known from Central America. The known species
in Mexico are found farther north, in Nearctic Mexico, es-
of this subgenus are predators of scales and mealybugs on
pecially Durango. All members of this genus with known
various plants.
biology are associated with various grasses, with larvae
Melaleucopis Sabrosky. Two described species of this feeding on mealybugs in the leafsheaths (Barber, 1985),
Neotropical genus are known, one from Peru and the oth- and many can be collected by sweeping roadside grasses.
er from the West Indies. The species from the Caribbean, Barber (1985) is the best available resource for identifying
M. simmondsi Sabrosky, is also known from Honduras. New species.
1006
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

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Culliney, T.W., J.W. Beardsley  Jr.,  & J.J.  Drea. 1988. Population ciden (Dipt.). Entomologische Mitteilungen 5: 294–299.
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Ferrar, P. 1987. A guide to the breeding habits and immature stages ogy 37: 494–504.
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311–328. United States National Museum 256: 1–158.
Gaimari, S.D. & W.J. Turner. 1996a. Larval feeding and develop- Mazzini, M.  & A. Raspi. 1983. Sulla fine struttura del micropilo
ment of Leucopis ninae Tanasijtshuk and two populations of negli insetti. XVIII. L’uovo di Chamaemyia flavipalpis (Ha-
Leucopis gaimarii Tanasijtshuk (Diptera: Chamaemyiidae) on liday) e di Parochthiphila coronata (Loew) (Diptera, Chamae-
Russian wheat aphid, Diuraphis noxia (Mordvilko) (Homoptera: myiidae). Frustula Entomologica, Nuova Serie 6: 1–12.
Aphididae), in Washington. Proceedings of the Entomological McAlpine, J.F. 1963. Relationships of Cremifania Czerny (Diptera:
Society of Washington 98: 667–676. Chamaemyiidae) and description of a new species. The Cana-
Gaimari, S.D.  & W.J. Turner. 1996b. Immature stages of Leuco- dian Entomologist 95: 239–253.
pis ninae Tanasijtshuk and two variant populations of Leucopis McAlpine, J.F. 1971. A revision of the subgenus Neoleucopis
gaimarii Tanasijtshuk (Diptera: Chamaemyiidae) feeding on (Diptera: Chamaemyiidae). The Canadian Entomologist 103:
Russian wheat aphid, Diuraphis noxia (Mordvilko) (Homoptera: 1851–1874.
Aphididae). Proceedings of the Entomological Society of Wash- McAlpine, J.F. 1978. A new dipterous predator of balsam woolly
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Gaimari, S.D. & W.J. Turner. 1998. Behavioral observations on the tomologica Germanica 4: 349–355.
adults and larvae of Leucopis ninae and L.  gaimarii (Diptera: McAlpine, J.F. 1987. Chamaemyiidae, pp.  965–971. MND,
Chamaemyiidae), predators of Russian wheat aphid, Diuraphis Volume 2.
noxia (Homoptera: Aphididae). Journal of the Kansas Entomo- McAlpine, J.F. 1989. Phylogeny and classification of the Musco-
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McLean, I.F.G. 1992. Behaviour of larval and adult Leucopis (Dip- Tanasijtshuk, V.N. 1963. On the biology of the genus Parochthiphila
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ural History 5: 35–36. 1876–1880. [In Russian.]
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Digest 5: 49–54. reniye 47: 633–651. [In Russian; English translation in 1968,
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records and geographic range. BioControl 50: 97–111. myiid flies (Diptera). Entomologicheskoye obozreniye 53:
Papp, L. 1998. Family Cremifaniidae, pp. 409–414. In Papp, L. & 304–308 [In Russian; English translation, 1970, Entomological
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Raspi, A. 1983. Contributi alla conoscenza dei Ditteri Camemiidi. Volume 9. Akadémiai Kiadó, Budapest, 460 pp.
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Raspi, A. 1990. Contributi alla conoscenze dei Ditteri Camemiidi. Tanasijtshuk, V.N. 1992. Morphological differences and phyletic re-
V. Su alcune specie del genere Leucopis viventi a spese di afidi lations between genera of chamaemyiid flies (Diptera, Chamae-
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(1988) Russian; English translation in 1993, Entomological Review 72:
Sabrosky, C.W. 1957. A new genus and two new species of Chamae- 66–100.]
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the Brooklyn Entomological Society 52: 114–117. Australia. International Journal of Dipterological Research 7:
Sluss, T.P.  & B.A. Foote. 1971. Biology and immature stages of 1–62.
Leucopis verticalis (Diptera: Chamaemyiidae). The Canadian Tanasijtshuk, V.N. 1997. A new Holarctic genus of Chamaemyiidae
Entomologist 103: 1427–1434. (Diptera). International Journal of Dipterological Research 8:
Sluss, T.P. & B.A. Foote. 1973. Biology and immature stages of Leu- 113–116.
copis pinicola and Chamaemyia polystigma (Diptera: Chamae- Tanasijtshuk, V.N. 2003. Geographical distribution and system
myiidae). The Canadian Entomologist 105: 1443–1452.
of silver-flies (Diptera, Chamaemyiidae). Proceedings of the
Smith, K.G.V. & I.F.G. McLean. 1998. Leucopis glyphinivora Ta-
nasijtshuk (Dipt., Chamaemyiidae) new to Britain and the aerial Zoological Institute of the Russian Academy of Sciences 299:
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Monthly Magazine 134: 85–87. Valenti, M.A. & S.D. Gaimari 1992. The manzanita leafgall aphid,
Stevenson, A.B. 1967. Leucopis simplex (Diptera: Chamaemyiidae) Tamalia coweni (Cockerell) (Homoptera: Aphididae), and its
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
PARALEUCOPIDAE (paraleucopid flies) 73
Terry A. Wheeler

Fig. 73.1. Male of Paraleucopis mexicana Steyskal, (illustrated by R. Vargas).

Diagnosis Legs without outstanding setae except for posterior and pos-
teroventral setae on forefemur and apicoventral bristle on
Small (body length 2.0–3.0 mm), shining black or metallic midtibia. Wing (Fig. 4) unpatterned; costa extending to M1,
flies (Fig. 1). Head (Figs. 1, 2) with vertical bristles strong, subcostal break present but distinct only at high magnifica-
other setae short and weak or absent, interfrontal setulae tion; Sc incomplete, not fused with R1 in Central American
numerous and evenly distributed; frons broad, microstriate; species, cell cup complete, anal vein usually long. Abdomen
lunule broad; face broad, sclerotized, with vertical median short, broad. Male epandrium small, offset to right side of
sulcus; antenna small, first flagellomere round with dorsal midline at rest; surstylus long, narrow; phallus short, mem-
arista; eye bare or short-haired; gena broad in Central Ameri- branous; cerci distinct, weakly sclerotized, setulose. Female
can species, approximately 35% eye height, curved inward below postabdomen telescopic posterior to segment  5; tergite  6
eye; palpus and proboscis short. Scutum (Fig. 3) usually with one well sclerotized, varied in shape; cerci long, narrow, setose;
postpronotal, two notopleural, one dorsocentral, two postalar spermathecae two or three in number, varied in shape.
bristles; scutal setulae dense, evenly distributed over surface. The combination of shining metallic body, broad incurved
Scutellum short, convex, with two pairs of marginal bristles. gena, and wing venation distinguishes paraleucopids from
Thoracic pleurites mostly bare, katepisternal bristle present. most other acalyptrate flies in the region. Chloropidae lack

1009
1010
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

a closed cell cup in the wing and have a vertical propleural around the eyes of colonial nesting seabirds and lizards in
carina. Most Sepsidae have a more slender, antlike body Mexico. Adult flies are also attracted to humans, sometimes
form. Most small shining flies in other families (e.g., Carnidae, in large numbers, where they congregate around the eyes,
Milichiidae, Drosophilidae, Ephydridae) have more extensive mouth, and superficial wounds, in much the same way as the
cephalic chaetotaxy and different wing venation. eye gnats of the family Chloropidae (Steyskal, 1971; Smith,
1981). These adult swarms are only active during warm
Biology days; at night the flies form large roosting aggregations on
the underside of building balconies, cave overhangs, or large
The following account applies mainly to Paraleucopis leaves (Smith, 1981).
because few specimens of other genera have been collected
and their habits are virtually unknown. Although larvae of Classification
Paraleucopis have not been described, they are apparently
saprophagous, possibly in association with bird nests. Adults The relationships of the Paraleucopidae have been the
have been reared from nests (Malloch, 1913; M.R. Wheeler, subject of confusion and controversy for almost 100 years.
1959) and can be abundant near seabird colonies (Smith, Malloch (1913) originally assigned Paraleucopis to the
1981). Adults have been collected on flowers and other veg- Chamaemyiidae (as Ochthiphilinae: Agromyzidae), and it
etation, especially in saline areas, including ocean beaches. was treated as a chamaemyiid by most later authors, includ-
Smith (1981) reported large numbers of adults feeding ing McAlpine (1987), although he noted that this placement

2 Paraleucopis 3 Paraleucopis

4 Paraleucopis

Figs. 73.2–4. Head, thorax, and wing: anterodorsal view of head (2), dorsal view of thorax (3), and dorsal view of wing (4), of Paraleucopis
corvina Malloch, (Nearctic, MND, figs. 88.5, 88.12, and 88.10, as Chamaemyiidae).
PARALEUCOPIDAE (paraleucopid flies) 73 1011

was for convenience only. Subsequently, McAlpine (1989) United States and Mexico and a second species group
left Paraleucopis and the related genera Mallochianamyia in Patagonia. Mallochianamyia and Schizostomyia are re-
Santos-Neto (as Gayomyia Malloch) and Schizostomyia stricted to Patagonia.
Malloch as unplaced acalyptrates more closely related to
Asteiidae than to Chamaemyiidae. T.A. Wheeler (in press) Synopsis of the fauna
erected a separate family for Paraleucopis.

Identification Paraleucopis Malloch. This genus contains at least

six species in the southwestern United States and western
Described species of Paraleucopis may be identified us- Mexico (T.A. Wheeler, in press) that are generally found in
ing the key in T.A. Wheeler (in press). Identification of the arid regions. The habits of the genus are discussed above
genus and related genera relies primarily on external charac- under the family overview. Although most Mexican re-
ters (wing venation, halter color, head shape) and geographic cords of Paraleucopis are from areas adjacent to the Gulf
distribution, so identification to that level should not pres- of California (Baja California, Baja California Sur, Sonora),
ent a problem with specimens in good condition. Because of specimens of Paraleucopis may eventually be found in arid
their small size and the importance of genitalic characters in
regions, especially in association with nesting seabirds, far-
species identification, adult specimens should be preserved
in ethanol and critical-point-dried or chemically dried soon ther south along the Pacific coast. Smith (1981) found that
afterwards. P. mexicana Steyskal was particularly abundant from mid-
Of the three described genera, Paraleucopis is the only March to mid-June in the northern Gulf of California, often
one found in Mexico or Central America; its distribution causing sunbathers to abandon the beaches and seek shelter
is disjunct, with one group of species in the southwestern from the flies.

Literature cited

Malloch, J.R. 1913. A synopsis of the genera of Agromyzidae, Proceedings of the Entomological Society of Washington 83:
with descriptions of new genera and species. Proceedings of the 406–412.
United States National Museum 46: 127–154. Steyskal, G.C. 1971. The genus Paraleucopis Malloch (Diptera:
McAlpine, J.F. 1987. Family Chamaemyiidae, pp. 965–971. MND, Chamaemyiidae), with one new species. Entomological News
Volume 2. 82: 1–4.
Wheeler, M.R. 1959. Notes on some flies reared from bird’s nests.
McAlpine, J.F. 1989. Phylogeny and classification of the Muscomorpha, The Southwestern Naturalist 4: 154.
pp. 1397–1518. MND, Volume 3. Wheeler, T.A. In press. Systematics of Paraleucopis Malloch, with
Smith, R.L. 1981. The trouble with “bobos,” Paraleucopis mexicana proposal of Paraleucopidae, a new family of acalyptrate Dip-
Steyskal, at Kino Bay, Sonora, Mexico (Diptera: Chamaemyiidae). tera. Zootaxa.
1012
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
COELOPIDAE (kelp flies) 74
Wayne N. Mathis

Fig. 74.1. Female habitus of Coelopa (Fucomyia) frigida (Fabricius), (Holarctic, illustrated by G. Venable).

Diagnosis apex; CuA2+A1 reaching posterior margin. Apical tarsomere

broadly subtriangular. Abdominal sternite  1 short or vesti-
Adults (Fig. 1) small to large flies (body length 3–16 mm); gial. Male terminalia as in Fig. 7. Female with 3 (1+2) small
robust, squat, strongly setose; brown to blackish; occurring spermathacae.
on seashores. Vertex flat to shallowly convex; postocellar Eggs undescribed.
setae convergent. First flagellomere subcircular, decumbent. Third-instar larvae typically maggotlike, cylindrical, an-
Face slightly convex to concave (Figs.  2–3), sclerotized, terior end tapered, posterior end generally broadly rounded;
parafacial prominence near lower margin of face. Thorax
anterior spiracle on short process, fan-shaped, digitate, with
distinctly flattened; metepisternum bearing several short
numerous terminal openings; posterior spiracles separate,
setulae; katepisternal seta anteroclinate to anterodorso-
clinate (Fig.  4); katepisternal suture shallowly projected on short or elongate respiratory tubes, each spiracle with
dorsally, forming arc along posterior portion above katepis- three slits, surrounded by circlet of plumose hydrofuge hairs
ternal seta. Wing (Fig. 6) usually macropterous, hyaline to (not in fascicles); anal opening on protuberance, this some-
faintly infuscate; C unbroken; Sc complete; apical section of times subdivided into lobes that bear spines or starlike
M not bent strongly forward, terminating posterior of wing sclerites around anus.

1013
1014
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

2 Coelopa 3 Coelopa

4 Coelopina
5 Coelopa
Sc R1
R2+3
br
cup r-m R4+5

dm M1
st 6
CuA1
A1+CuA2 st 7
st 5 epand
sur
6 Coelopa cerc

tg 6 pgt
tg 7 tg 8

ph hypd
tg 5 ph gd
pregt

7 Coelopa

Figs. 74.2–7. Heads, thoraces, wing, and male terminalia: lateral view of head of (2) Coelopa (Neocoelopa) vanduzeei Cresson, (Nearctic and
Neotropical); and (3) Coelopa (Fucomyia) frigida (Fabricius), (Holarctic); lateral view of thorax of (4) Coelopina anomala Cole; dorsal view
of thorax of (5) Coelopa (Fucomyia) frigida; dorsal view of wing of (6) Coelopa (Neocoelopa) vanduzeei, (MND, fig. 82.2); ventral view of
male terminalia of (7) Coelopa (Fucomyia) nebularum Aldrich, (Nearctic, MND, fig. 82.5). Figures 2–5 illustrated by Y. Sohn.
Abbeviations: cerc, cercus; epand, epandrium; hypd, hypandrium; pgt,  postgonite; ph,  phallus; ph  gd, phallic guide; pregt, pregonite,
st, sternite; sur, surstylus; tg, tergite.
COELOPIDAE (kelp flies) 74 1015

Puparia dark brown, ovoid with moderately rugose Meier & Wiegmann (2002) have proposed the most recent
surface. and complete phylogenies for the family, using morpho-
Adult kelp flies are similar to Helcomyzidae and Dryo- logical data, and in the latter paper, also DNA sequences as
myzidae (neither of which are found in Central America) and evidence. Although the morphological evidence confirms the
are distinguished from these and other families of the Scio- monophyly of the family (McAlpine identified nine synapo-
myzoidea by the generally compact, flattened, and setose morphies), the DNA data were equivocal, especially regarding
appearance, the convergent postocellar setae, the flattened the position of the Australian genus Lopa McAlpine. Meier &
scutum (Fig.  5), the anteroclinate to anterodorsoclinate Wiegmann (2002: p. 401) suggested that genes other than the
katepisternal seta, and the setose metepisternum. Adults can ones used might better resolve this ambiguity.
vary considerably in their size and vestiture, which has led McAlpine’s classification not only divided the Coelopidae
to misinterpretation of species, and several synonyms have into two subfamilies (Lopinae and Coelopinae), but within
resulted. the Coelopinae also recognized four tribes and 12 genera.
McAlpine’s classification is relatively finely divided (seven
Biology of the 13 genera are monotypic) and did not always accord
with the “preferred” cladogram in Meier  & Wiegmann’s
All stages of kelp flies live in stranded seaweeds, espe- total-evidence analysis. Thus, the subfamily Coelopinae now
cially where extensive wrack beds of kelp, sea grasses, and comprises three tribes, 12 genera, and 28 species (Mathis &
other jetsam have accumulated. Adults are often abundant McAlpine, In press). Only the subfamily Coelopinae occurs
and sometimes are a nuisance to people at the beach. Adults in the New World, and of the four tribes assigned to this
are also attracted to certain organic solvents and can be a subfamily, only two, Glumini (Coelopina) and Coelopini
pest to facilities where these solvents are produced or used. (Coelopa), are found in the New World. The Neotropical
Coelopa frigida (Fabricius) has been reared in the labora- Region is notable in lacking any species, with the exception
tory and used for genetic studies. Egglishaw (1960) and of Coelopina anomala Cole and Coelopa vanduzeei Cresson
McAlpine (1991) reviewed the ecology of kelp flies. Spe- in Baja California, and elsewhere only a few species are
cies of Coelopidae largely have temperate distributions and found in the tropics and subtropics.
reveal greatest diversity in the Southern Hemisphere, espe-
cially in the Australasian Region where most genera (10) and
species (18) are found. Identification
McAlpine (1991) published the only recent key to genera
Classification of the world, and his paper is essential to understanding the
family even though its focus is on the Australian fauna. Al-
Coelopidae are in the superfamily Sciomyzoidea and are drich’s (1929) revision of North American species of Coelo-
similar and closely related to the Helcomyzidae, perhaps pa and Malloch’s (1933) summary of Coelopa are still useful
as the sister group (McAlpine 1991). McAlpine (1991) and to identify species from the New World.

Key to the genera of Coelopidae of the New World

1. Anepisternum bare on posterior half; ventrally fused metakatepisternal area (metasternum)
setulose; widespread, mostly in temperate regions . . . . . . . . . . . . . . . . . . . . . . . . Coelopa Meigen
– Anepisternum bearing many setulae and setae on posterior half (Fig.  4); ventrally fused
metakatepisternal area (metasternum) bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . Coelopina Malloch

Synopsis of the fauna Coelopa Meigen. Three subgenera have been recognized
within this genus, but I am not convinced of their validity.
The Coelopidae include 29 species in 13  genera Only the monotypic subgenus Neocoelopa Malloch occurs
(McAlpine, 1991). The family is largely temperate in distri- in Central America, with C. vanduzeei as its only included
bution with greatest diversity occurring in the Australasian species. This species is often found abundantly in association
Region, where most genera and species are found. The New with stranded beds of kelp (Poinar, 1977; Arnaud, 1983). Ap-
World fauna comprises two genera and five species (Steyskal, parently based on genetics and nutrition, there is consider-
1967; Vockeroth, 1965, 1987), with four species in Coelopa able variation in the size of specimens and also in the number
and a single species in Coelopina. Two species occur in and size of setae. In the region, the species is known thus far
Central America. No fossils are known. from Baja California and Baja California Sur.
1016
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Coelopina Malloch. This is a monotypic genus with investigation, as there are no large brown kelps, the typical
C.  anomala as its only included species. The type locality food of coelopids, in the Gulf of California where this spe-
is Las Animas Bay (Baja California), Mexico. The biology cies occurs.
and immature stages of this species are unknown but invite

Literature cited
Aldrich, J.M. 1929. Revision of the two-winged flies of the genus Meier, R.  & B.M. Wiegmann. 2002. A phylogenetic analysis of
Coelopa Meigen in North America. Proceedings of the United Coelopidae (Diptera) based on morphological and DNA se-
States National Museum 76: 1–6. quence data. Molecular Phylogenetics and Evolution 25:
Arnaud, P.H., Jr. 1983. Aggregation of Coelopa (Neocoelopa) van- 393–407.
duzeei Cresson on the Monterey Peninsula coast, California, Poinar, G. 1977. Observations on the kelp fly, Coelopa vanduzeei
and notes on the family (Diptera: Coelopidae). Pan-Pacific En- Cresson in Southern California. Pan-Pacific Entomologist 53:
tomologist 58: 245–249. 81–86.
Egglishaw, H.J. 1960. Studies on the family Coelopidae (Diptera). Steyskal, G.C. 1967. 61. Family Coelopidae. In Papavero, N. (edi-
Transactions of the Royal Entomological Society of London tor). A Catalogue of the Diptera of the Americas South of the
112: 109–140. United States. Departamento de Zoologia, Secretaria da Agri-
Malloch, J.R. 1933. The genus Coelopa Meigen (Diptera, Coelo- cultura, São Paulo, 1 p.
pidae). Annals and Magazine of Natural History, Series 10, 11: Vockeroth, J.R. 1965. Family Coelopidae, pp. 679–680. In Stone, A.,
339–350. C.W. Sabrosky, W.W. Wirth, R.H. Foote, & J.R. Coulson (edi-
Mathis, W.N. & D.K. McAlpine. In Press. A catalog and conspectus tors). A catalog of the Diptera of America north of Mexico.
on the family Coelopidae (Diptera: Schizophora). Myia. United States Department of Agriculture, Agriculture Handbook
McAlpine, D.K. 1991. Review of the Australian kelp flies (Diptera: 276, Washington, D.C., iv + 1696 pp.
Coelopidae). Systematic Entomology 16: 29–84. Vockeroth, J.R. 1987. Coelopidae, pp. 919–922. MND, Volume 2.
SCIOMYZIDAE (snail-killing flies, marsh flies) 75
Luciane Marinoni and Lloyd Knutson

Fig. 75.1. Male of Thecomyia chrysacra Marinoni & Steyskal, (Zumbado, 2006, p. 157).

Diagnosis reaching margin. Legs slender; some or all tibiae with preapi-
cal dorsal seta. Male terminalia symmetrical to asymmetrical;
Adults (Fig.  1) small to large flies (body length 2.0– tergite 6 usually absent; right spiracles 6 and 7 mostly in terg-
13.0 mm), mostly yellow to brown, sometimes black. Head ite; epandrium well developed, symmetrical; usually with an-
with antenna porrect, pedicel usually elongate; arista bare to terior and posterior surstylus, sometimes anterior one vestigial
plumose; ocellar setae present in most species; inner and outer or absent; cercus well developed, setose. Female sternites 6, 7,
vertical setae developed; postocellar setae parallel or slightly and 8 separate or fused; two spermathecae present.
divergent. Face slightly to strongly concave, uniformly scle- Egg length 0.6–2.0 mm; white or yellowish, often becom-
rotized; vibrissa absent; clypeus small, withdrawn. Thorax ing brown or gray as larva develops; usually unicolorous.
with zero or one proepisternal seta; zero or one presutural su- Mostly oval, with dorsal surface often less convex than
pra-alar; zero, one, or two postalar; zero or one postpronotal; ventral surface. Each end usually with low or prominent
most with two notopleural; usually two pairs of scutellar se- tubercle, anterior tubercle often reduced. Micropyle terminal
tae. Wing elongate, completely hyaline (Fig. 7) to patterned or subterminal. Chorion slightly reticulate or with ridges and
(Figs.  6, 8); C and Sc complete; C unbroken; A1 + CuA2 grooves.

1017
1018
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Larva white, integument transparent, diaphanous (semi- arch connected posterolaterally on each side of mandibles;
terrestrial and terrestrial species); or brownish or yellowish mandibles with or without row of accessory teeth.
(aquatic species). Third instar with elongate and cylindrical Puparium with integument opaque, yellowish, reddish-
body, tapered anteriorly; posterior spiracular disc surrounded by brown to dark brown or almost black. Ovoid, barrel-shaped,
two to five pairs of short to elongate lobes. Surface smooth or tapered at both ends, posterior spiracles in most aquatic
with areas of microscopic spinules; Sciomyzini species with species elevated above dorsum.
ventral spinules; terrestrial Tetanocerini with cuticle com- Among acalyptrate cyclorrhaphans, the sciomyzids can
pletely naked; aquatic and semi-aquatic Tetanocerini with be distinguished by the following set of morphological char-
rings of tubercles and welts. Aquatic species with elongate acters: costal vein  (C) without breaks; subcostal vein  (Sc)
float hairs between posterior spiracular slits; float hairs much complete, free from vein R1; vein A1 complete; oral vibrissae
reduced in terrestrial species. Cephalopharyngeal skeleton in absent; postvertical setae divergent to parallel and at least
the first three segments, long, with paired mandibles; ventral one tibia with a preapical dorsal seta.

2 Neuzina

3 Sepedonea

4 Euthycera 5 Protodictya

Figs. 75.2–5. Heads: lateral view of (2) Neuzina diminuta Marinoni & Zumbado; (3) Sepedonea lagoa Steyskal; and (4) Euthycera arcuata
(Loew); ventrolateral view of head of (5) Protodictya nubilipennis (Wulp). Figures 2–5 illustrated by F. Zagonel.
SCIOMYZIDAE (snail-killing flies, marsh flies) 75 1019

Biology and/or in the Neotropical Region: Atrichomelina (Foote et

al. 1960); Anticheta Haliday (Fisher & Orth, 1964; Knutson,
Life cycles and immature stages of approximately one- 1966; Knutson  & Abercrombie, 1977); Dictya (Valley  &
third of the 534 described species and half of the 61 described Berg, 1977); Dictyodes Malloch (Abercrombie  & Berg,
genera are known, with the best-studied species occurring 1978); Ditaeniella (Bratt et al., 1969); Euthycera Latreille
in the Nearctic and Palearctic Regions. Larval feeding hab- (Vala et al., 1983); Hoplodictya (Neff & Berg, 1962); Per-
its are complex and varied within the family; most are pri- ilimnia Becker (Kaczynski et al., 1969); Pherbellia (Bratt
marily parasitoids or predators on freshwater or terrestrial et al., 1969); Protodictya (Neff & Berg, 1961); Sepedomer-
mollusks. This behavior has been interpreted as one of the us (Neff  & Berg, 1966); Sepedon Latreille (Neff  & Berg,
characters that define the monophyly of the family (Grif- 1966); Sepedonea (Freidberg et al., 1991; Knutson & Valley,
fiths, 1972; McAlpine, 1989; Marinoni  & Mathis, 2000). 1978; Neff & Berg, 1966); Shannonia Malloch (Kaczynski
The only known trophic deviation from eating mollusks et al., 1969); Tetanocera (Foote, 1996); and Thecomyia (Ab-
are the African species Sepedonella nana Verbeke (Vala ercrombie  & Berg, 1975). Only Atrichomelina, Ditaniella,
et al., 2000) and Sepedon knutsoni Vala et al. (Vala et al., and Pherbellia from this list belong to the Sciomyzini. Most
2002), whose larvae were reared in the laboratory feeding on species of this tribe are parasitoids on exposed aquatic or
aquatic Oligochaeta (Annelida). Behavioral studies and/or terrestrial snails, but Atrichomelina pubera (Loew), depend-
descriptions of immature stages are available for at least one ing on circumstances, has predatory, parasitoid, or even sap-
species in the following genera occurring in Central America rophagous behavior. Species of other genera, all in the tribe

7 Sepedonea
6 Neuzina

8 Euthycera 9 Sepedonea

10 Sepedomerus 11 Hoplodictya

Figs. 75.6–11. Wings, legs, and antenna: dorsal view of wing of (6) Neuzina diminuta Marinoni & Zumbado; (7) Sepedonea lagoa Steyskal;
and (8) Euthycera arcuata (Loew); anterior view of hind leg of (9) Sepedonea lagoa; and (10) Sepedomerus macropus (Walker); lateral view
of antenna of (11) Hoplodictya spinicornis (Loew). Figures 6–11 illustrated by F. Zagonel.
1020
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Tetanocerini, are predaceous on aquatic or terrestrial snails, Griffiths (1972) characterized the family as comprising
slugs, and a few on fingernail clams. Species of aquatic Tet- Salticellinae and Sciomyzinae. He removed Pelidnoptera
anocerini have been best studied experimentally because of Rondani to a separate family, Phaeomyiidae. Griffiths
their potential to help control diseases of man and domestic defined Sciomyzidae and established its monophyly based
animals, such as fascioliasis and schistosomiasis, that have on the malacophagous habits of the larvae, on the presence
of a ventral arch in the cephalopharyngeal skeleton, and on
aquatic snails as intermediate hosts. Most of the aquatic
four characters of the adults. The tribes Sciomyzini and Tet-
predators attack freshwater pulmonate snails, killing up to anocerini of Steyskal (1965) were not recognized within the
30 snails during the three larval instars. Recently, attempts to Sciomyzinae.
understand the evolution of feeding behavior within the Sci- Barnes (1979, 1981), studying morphological characters
omyzidae have been made. Knutson & Vala (2002), based on of adults, recognized the Sciomyzidae with the subfamilies
the phylogeny of the family provided by Marinoni & Mathis Phaeomyiinae, Huttonininae, Sciomyzinae, and Salticellinae.
(2000), observed that there is general agreement between the Similar to Hennig (1965) and Griffiths (1972), he regarded
cladogram and some attributes of the behavioral groups at the the small clypeus as an indication of monophyly for Scio-
family, subfamily, and tribe levels, and some concordance myzidae without, however, a cladistic interpretation of his
between the relative degree of specialization of behavior and data. His subfamily Sciomyzinae, as in Steyskal (1965), was
position of certain genera. They observed also that the pattern composed of two tribes, Sciomyzini and Tetanocerini.
of dispersion of behavioral groups is not highly concordant McAlpine (1989) did not recognize the family Phaeomyi-
idae and in a comprehensive discussion of the relationships
with the cladogram for many genera, and they recognized 15
among the Sciomyzoidea treated Sciomyzidae with four
behavioral groups within Sciomyzidae. Barker et al. (2004), subfamilies: Huttonininae, Phaeomyiinae, Sciomyzinae, and
in an overview of the biology of the family with special ref- Salticellinae.
erence to predators and parasitoids of terrestrial gastropods, Marinoni  & Mathis (2000), in a cladistic analysis of
corroborated the higher level classification for the family, 50 genera belonging to the family, recognized two subfami-
subfamilies, and tribes of Marinoni & Mathis (2000). They lies, Salticellinae and Sciomyzinae. They concluded that both
concluded that some of the genera should be placed in dif- subfamilies were monophyletic, and within the Sciomyzinae
ferent positions in the cladogram. Based on their phylogeny, concluded that the tribes Sciomyzini and Tetanocerini were
they examined the evolution of prey preferences within the monophyletic. Most genera in the analysis remained
Sciomyzidae and identified nine behavioral groups that they in unresolved polytomies. The most corroborated lineage
named “eco-groups”. Berg et al. (1982) defined five distinct within their analysis comprised genera of the Sepedon group
(Steyskal, 1965): Thecomyia, Sepedoninus Verbeke, Sepedo-
patterns of seasonality for Sciomyzidae living in cool, tem-
nella Verbeke, Sepedon, Sepedomerus, and Sepedonea.
perate latitudes. In the Neotropical Region, seasonality has Barker et al. (2004), in an overview of the Sciomyzidae
been studied for Thecomyia longicornis Perty (Knutson  & (mainly predators and parasitoids of terrestrial gastropods),
Carvalho, 1989), five species of Sepedonea, and Sepedomerus revised the cladistic analysis of Marinoni & Mathis (2000).
bipuncticeps Malloch (Mello & Bredt, 1978). They added seven morphological characters of adults in the
analysis and removed the two related to biological character-
istics. The subfamilies and tribes remained the same, and the
Classification relationships of the genera of Sciomyzini were congruent.
However, differences in placement of some genera within
The Sciomyzidae belong to the superfamily Scio- the Tetanocerini were observed. Seven genera identified
myzoidea, a group that also includes the Coelopidae, Dryo- as higher Tetanocerini by Marinoni  & Mathis (2000) were
myzidae, Helosciomyzidae, Ropalomeridae, and Sepsidae placed at the base of the tribe. The genera Thecomyia, Sepe-
(McAlpine, 1989). doninus, Sepedonella, Sepedon, Sepedomerus, and Sepe-
Verbeke (1950) was the first modern author to define a donea remained at the higher position in the cladogram.
classification for the Sciomyzidae. He divided the family
into the Phaeomyiinae, Sciomyzinae, Renocerinae, Tetano-
cerinae, Sepedoninae, and Salticellinae. Identification
Steyskal (1965) recognized five subfamilies: Hutton-
ininae, Salticellinae, Helosciomyzinae, Phaeomyiinae, and Wulp (1897) treated 13 species in the genera Sciomyza
Sciomyzinae, the last consisting of two tribes, Sciomyzini Fallén, Tetanocera, and Sepedon, all from Mexico, with 11
and Tetanocerini. of these being new species. According to modern taxonomy
Hennig (1965) organized the family into eight subfami- and nomenclature, four of his new species are synonyms,
lies: Phaeomyiinae, Helosciomyzinae, Huttonininae, Scio- one has been transferred to the Lauxaniidae, and the species
myzinae, Tetanocerinae, Neoliminiinae, Renocerinae, and of Sciomyzidae represent 10 genera.
Sepedoninae. According to him, the monophyly of the fam- The most recent key to Neotropical genera is Steyskal &
ily is supported by the clypeus being small, withdrawn, and Knutson (1975) and to the Nearctic genera is Knutson (1987).
separated from the anterior margin of the face by a large The names of the Central American species are covered in
membranous area. Knutson et al. (1976). Some revisionary papers have been
SCIOMYZIDAE (snail-killing flies, marsh flies) 75 1021

published, including keys to species of Central American Most Neotropical sciomyzids are not difficult to identify
and Neotropical genera: Protodictya (Marinoni & Knutson, at the genus level, since they have conspicuous taxonomic
1992); Sepedonea (Freidberg et al., 1991); and Thecomyia characters and their size is relatively large compared with
(Marinoni et al., 2003). other acalyptrate flies.

Key to the genera of Sciomyzidae of Central America

1. Pedicel approximately half length of first flagellomere (Fig.  2); head in profile as in Fig.  2;
proepisternal seta present; SCIOMYZINI. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Pedicel subequal to or twice length of first flagellomere (Figs. 3–5); head in profile as in Figs. 3–4;
proepisternal seta absent; TETANOCERINI. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

2. Proepisternal seta short, fine, hairlike; thoracic pleuron with fine hairs, without setae; wing
without pattern . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Atrichomelina Cresson
Proepisternal seta well developed; thoracic pleuron with setae; wing with or without pattern . . 3

3. One fronto-orbital seta present; midfrontal vitta subshiny, nearly reaching anterior margin of
frons; wing without pattern . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ditaeniella Sack
– One or two fronto-orbital setae present; midfrontal vitta absent or short, triangular, pruinose; wing
patterned. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4. Face with median rounded black spot; one fronto-orbital seta present; body with spots on bases of
hairs and setae; wing strongly patterned (Figs. 2, 6). . . . . . . . . . . . Neuzina Marinoni & Knutson
– Face without such spot; two fronto-orbital setae present; wing weakly to strongly patterned . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pherbellia Robineau-Desvoidy

5. Face with median rounded black spot (Fig. 5). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

– Face without median rounded black spot. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

6. Head longer than high, as in Fig. 4; lunule exposed; two fronto-orbital setae present; arista white;
wing patterned (Fig. 8); Mexico (Durango). . . . . . . . . . . . . . . . . . . . . . . . . . [Euthycera Latreille]
– Head higher than long, as in Fig. 3; lunule exposed or covered; one fronto-orbital seta present;
arista white or black; wing patterned or not. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

7. Lunule exposed; wing translucent, not patterned or with pattern; hind femur slender, longer than
tibia; fifth abdominal sternite of male with process. . . . . . . . . . . . . . . . . . . . . Protodictya Malloch
– Lunule not exposed; wing strongly patterned; hind femur short, almost same length as tibia; fifth
abdominal sternite of male without process. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dictya Meigen

8. Ocellar seta absent; one pair of scutellar setae present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

– Ocellar seta present; two pairs of scutellar setae present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

9. Hind tibia not arch-shaped, without such projection; head conically developed downwards
(Fig. 1); palpus absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thecomyia Perty
– Hind tibia arch-shaped, with spinelike projection ventroapically (Figs. 9–10); head not conically
developed; palpus present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

10. Face with dark brown spot on anteroventral corner (Fig. 3); postocellar setae well developed; hind
femur without markings (Fig. 9) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sepedonea Steyskal
– Face without dark brown spot on anteroventral corner; postocellar setae absent; hind femur well
developed, with lateral dark marks (Fig. 10). . . . . . . . . . . . . . . . . . . . . . . . Sepedomerus Steyskal
1022
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

11. Hind tibia arch-shaped; small flies; head dorsoventrally flattened; pedicel wide and flattened; all
basitarsi white. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Teutoniomyia Hennig
– Hind tibia not arch-shaped; small to medium-sized flies; head, pedicel, and basitarsi not as
above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

12. Anepisternum and anepimeron without setae or hairs; two fronto-orbital setae present; wing not
patterned. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetanocera Duméril
– Anepisternum and sometimes anepimeron with setae or hairs; one or two fronto-orbital setae pres-
ent; wing patterned. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

13. Face not strongly concave, with dark brown vitta on ventral corner; lunule not exposed; two fron-
to-orbital setae present; frons without spots; arista with sparse black hairs, not densely plumose
(Fig. 11). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplodictya Cresson
– Face strongly concave, without dark brown vitta on ventral corner; lunule exposed; one
fronto-orbital seta present; frons with black spots laterad of antenna and ocellar triangle; arista
black plumose. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Guatemalia Steyskal

Synopsis of the fauna analysis, the genus is in the apex of Tetanocerini. In the
second, it appears as the sister group of Tetanoceroides
The Sciomyzidae of Central America are poorly known, Malloch in a more basal position in the cladogram.
with 11 genera and 24 species recorded for the region.
Hoplodictya Cresson. This is a mainly Nearctic genus
Atrichomelina Cresson. This monotypic genus, with the (Fisher & Orth, 1972). Of the five species, only the primarily
species A.  pubera, ranges throughout North America from Nearctic H. spinicornis (Loew) is found in the Neotropical
southernmost Canada southward to Mexico (Distrito Federal, Region, occurring in Mexico and Guatemala.
Chiapas, Tamaulipas).
Neuzina Marinoni & Knutson. This genus has been re-
Dictya Meigen. This genus, the third largest in the fam- cently described for the species Neuzina diminuta Marinoni &
ily, includes 33 described species (Orth, 1984, 1991), and Zumbado (Marinoni et al., 2004), which occurs in Costa
only D.  umbrarum  (L.) (Palearctic) is not a New World Rica, Venezuela, and Brazil. The specimens from Costa Rica
species. Six species have been recorded for Central America, were collected in Malaise traps during the months of June
with D. guatemalana Steyskal in Guatemala and Costa Rica and July in the province of Guanacaste as part of the surveys
and D. bergi Valley in Costa Rica and Colombia (Knutson et that have been conducted by the Instituto Nacional de
al., 1976; Orth, 1984, 1991). Dictya is close to Hoplodictya Biodiversidad de Costa Rica.
(Marinoni & Mathis, 2000; Barker et al., 2004) and can be
separated from it by the presence of a midfacial black spot Pherbellia Robineau-Desvoidy. The genus Pherbellia
in Dictya (absent in Hoplodictya); by a row of two or three occurs worldwide except in the Subantarctic Region, and
spines dorsally at the distal end of the pedicel, which are with 95 species, it is the most species-rich genus in the family.
strong in Hoplodictya and less so in Dictya; and by the male In Central America only P. trabeculata (Loew) (Guatemala)
terminalia (Fisher & Orth, 1972). is known. In the cladograms of Marinoni & Mathis (2000)
and Barker et al. (2004), Pherbellia is in the Sciomyzini in
Ditaeniella Sack. Ditaeniella includes one Palearctic spe- a polytomy with Pteromicra Lioy and Calliscia Steyskal +
cies, one Neotropical species (D. patagonensis (Macquart), Parectinocera Becker. The last two genera are exclusively
which is found throughout South America), and two Nearctic Neotropical.
species, one of which, D. parallela (Walker), ranges as far
south as Mexico (Distrito Federal, Veracruz, Michoacán) and Protodictya Malloch. Protodictya is a Neotropical
Costa Rica (Cartago). genus that occurs mainly in Brazil and ranges from southern
Mexico to southern Argentina. Marinoni & Knutson (1992)
Guatemalia Steyskal. Guatemalia is a Neotropical genus reviewed Protodictya, which now includes eight species, and
with two species described, both of which occur in Central presented a key to them. Protodictya nubilipennis (Wulp) is
America: G. straminata (Wulp) is known from Mexico and the only species occurring in Central America (Mexico,
Guatemala and G.  nigritarsis Marinoni from Mexico Guatemala, Honduras, Nicaragua, Costa Rica, Panama, and
(Veracruz), Guatemala, and Honduras (Marinoni, 1992). The Venezuela). Marinoni & Carvalho (1993) did a cladistic anal-
phylogenetic position of Guatemalia in Marinoni & Mathis ysis of the eight species. In their analysis, P. nubilipennis is
(2000) differs from that of Barker et al. (2004). In the first at the apex of the cladogram along with P. iguassu Steyskal
SCIOMYZIDAE (snail-killing flies, marsh flies) 75 1023

(Argentina, Brazil) and P. brasiliensis (Schiner) (Brazil). As Tetanocera Duméril. This genus includes 40 spe-
synapomorphies for the genus, they found a median black cies, with 20 Nearctic, nine Palearctic, 10 Holarctic, and
facial spot and a medioapical process in the fifth sternite of one Oriental species. Tetanocera spreta Wulp is known
the male. from Mexico (Distrito Federal, Jalisco and Toluca), and
T. plumosa Loew, widespread in the Nearctic Region, is
Sepedomerus Steyskal. This genus has three Neotropi-
known from high elevations in Mexico (Distrito Federal)
cal species. The species occurring in Central America is
S.  macropus (Walker), which ranges from Peru to Texas and Venezuela (Caracas).
(USA), also including Mexico, Guatemala, Nicaragua, Costa
Teutoniomyia Hennig. This genus has two named
Rica, and Panama. Sepedomerus macropus was introduced
into Hawaii for biological control of the liver-fluke snail species, T.  plaumanni Hennig found in Brazil and
Lymnaea ollula Gould and has become established there T.  costaricensis Steyskal in Costa Rica, Mexico, and
(Berg, 1964). Sepedomerus + Sepedonea is the sister group the southern USA (Brownsville, Texas). Teutoniomyia is
of Sepedon (Marinoni & Mathis, 2000; Barker et al., 2004). basal to the Sepedon group in the analyses of Marinoni &
Mathis (2000) and Barker et al. (2004).
Sepedonea Steyskal. Like Protodictya, Sepedonea is
a Neotropical genus that is found mainly in Brazil. Three Thecomyia Perty. This genus is known only from
species have been reported from Central America: S. isthmi Central and South America. It is highly distinctive and
(Steyskal) (Canal Zone, Panama, Trinidad to Brazil, and can be readily distinguished from other genera by the
Bolivia), S.  lagoa (Steyskal) (Costa Rica to Brazil), and following characters: (1)  the narrow, conical, and ventral
S.  guatemalana (Steyskal) (Chiapas, Mexico, and Costa development of the head; (2)  the loss of the palpus;
Rica). Freidberg et al. (1991) reviewed the genus, described
(3) the strong reduction of most body setae; and (4) the
and illustrated the immature stages of six species, and pre-
sented a key to the 12  species. The biology and immature striking yellow and brown body color. Marinoni et al.
stages of S. guatemalana were treated by Neff & Berg (1966) (2003) reviewed the genus, which comprises 12 Neotro-
and those of S. isthmi by Knutson & Valley (1978). This is pical species, and presented a key and a cladistic analysis
the sister group of Sepedomerus (Marinoni & Mathis, 2000; for the species. From Central America, four species have
Barker et al., 2004). been recorded.

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Entomological Society of Washington 77: 355 –368. dia 36: 1–29.
Abercrombie, J.  & C.O. Berg. 1978. Malacophagous Diptera of Fisher, T.W. & R.E. Orth. 1972. Synopsis of Hoplodictya Cresson
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Barnes, J.K. 1979. The taxonomic position of the New Zealand Freidberg, A., L.V. Knutson,  & J. Abercrombie. 1991. A revision
genus Prosochaeta Malloch (Diptera: Sciomyzidae). Proceed- of Sepedonea, a Neotropical genus of snail-killing flies (Dip-
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Barnes, J.K. 1981. Revision of the Helosciomyzidae (Diptera). 1–48.
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Berg, C.O. 1964. Snail control in trematode diseases: the possible Cyclorrhapha, with special reference to the structure of the male
value of sciomyzid larvae snail-killing Diptera, pp. 259–309. In postabdomen. Series entomologica 8. Dr. W. Junk, The Hague,
Dawes, B. (editor). Advances in Parasitology, Vol. 2. Academic 340 pp.
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Berg, C.O., B.A. Foote, L.V. Knutson, & J.K. Barnes. 1982. Adaptive garter Beiträge zur Naturkunde 145: 1–212.
differences in phenology in sciomyzid flies. Memoirs of the Kaczynski, V.W., J. Zuska, & C.O. Berg. 1969. Taxonomy, imma-
Entomological Society of Washington 10: 15–36. ture stages, and bionomics of the South American genera Per-
Bratt, A.D., L.V. Knutson, & C.O. Berg. 1969. Biology of Pherbellia ilimnia and Shannonia (Diptera: Sciomyzidae). Annals of the
(Diptera: Sciomyzidae). Cornell University Agricultural Exper- Entomological Society of America 62: 572–592.
iment Station, New York State College of Agriculture, Ithaca, Knutson, L.V. 1966. Biology and immature stages of malacopha-
246 pp. gous flies: Antichaeta analis, A.  atriseta, A.  brevipennis and
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A. obliviosa (Diptera: Sciomizydae). Transactions of the Ameri- dae). Transactions of the American Entomological Society 88:
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Knutson, L.V.  & J. Abercrombie. 1977 Biology of Antichaeta malacophagous Diptera of the genus Sepedon (Sciomyzidae).
melanosoma (Diptera: Sciomyzidae), with notes on parasitoid Bulletin of the Agriculture Experiment Station, Virginia Poly-
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Knutson, L.V. & C.J.B. de Carvalho. 1989. Seasonal distribution of Sciomyzidae). Proceedings of Entomological Society of Wash-
a relatively rare and a relatively common species of Thecomyia ington 86: 893–897.
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Knutson, L. & J-C. Vala. 2002. An evolutionary scenario of Scio- dae (Diptera) from the Americas south of the United States, with
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Knutson, L. & K. Valley. 1978. Biology of a Neotropical snail-killing Vala, J.-C., J.M. Reidenbach, & C. Gasc. 1983. Biologie des sta-
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the Entomological Society of Washington 80: 197–209. de gastéropodes terrestres. Premiére découverte du cycle ches
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Sciomyzidae). Revista Brasileira de Zoologia 9: 247–249. myzidae). Bulletin de la Société Entomologique de France 88:
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Marinoni, L. & L.V. Knutson. 1992. Revisão do gênero Neotropi- killing flies. Comptes Rendus de l’Académie des Sciences. Série
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McAlpine, J.F. 1989. Phylogeny and classification of the Musco- sels, 97 pp.
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ROPALOMERIDAE (ropalomerid flies) 76
Sergio Ibáñez-Bernal and Vicente Hernández-Ortiz

Fig. 76.1. Male of Ropalomera femorata (Fabricius), (MND, fig. 85.1).

Diagnosis and broadened (Figs. 8–12). Female terminalia with tergite

and sternite  7 partially fused, ending conically; ovipositor
Medium to large robust flies (body length 6–12  mm) soft and short with small setulae; two globose sclerotized
(Fig.  1) with body generally gray or dark brown. Vertex spermathecae. Male terminalia (Figs. 13–16) with epandri-
commonly sunken (Fig.  3); gena high (Figs.  2, 4–7); face um curved ventrally; cerci basally fused as in Ropalomera
with median ridge (Fig. 4) or rounded protuberance (Figs. 2, (Fig.  13), Kroeberia (Fig.  15), and Lenkokroeberia, but
3, 5–7); antenna with first flagellomere ovoid (Fig.  4) or sometimes completely fused as in Willistoniella (Fig. 14);
elongate (Fig.  2), arista subbasal, bare (Figs.  5–6), with surstyli spatulate; postgonite with one to three lobes; phal-
short flattened hairs (Fig. 7) or plumose (Figs. 2–4); vibris- lus short, membranous with spurlike epiphallus at posterior
sae absent. Scutum with tomentum pattern of longitudinal base.
stripes; scutellum frequently grooved; greater ampulla pres- Ropalomerids grossly resemble Sarcophagidae in terms
ent. Wings mostly hyaline to brownish or spotted; R1 bare; of their body coloration and presence of a greater ampulla,
R4+5 and M strongly convergent towards wing margin; dm-cu but the usually sunken vertex, wing venation, broad femo-
curved sharply forward in line with M. Femora swollen, ra, and flattened tibia are sufficient characteristics for easy
particularly on hind leg; hind tibia often laterally flattened recognition.

1025
1026
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Biology and two spermathecae present. The species of Ropalol-

meridae can be distinguished from those of Sepsidae by the
The biology of the species of this family is poorly known. following autapomorphies: vertex concave (sunken), face
Lopes (1932) provided descriptions of the egg and larva of with median protuberance, precoxal bridge present, greater
Ropalomera stictica Wiedemann and figured the puparium of ampulla developed, vein M bent forward, hind tibia laterally
Ropalomera clavipes (Fabricius). Fischer (1932) described compressed and strongly expanded, sternite 6 of male great-
the puparium of Willistoniella pleuropunctata (Wiedemann), ly reduced or absent, and basiphallus broadly expanded cau-
and later Prado (1966) observed the eggs of this species. dally forming a pseudohypandrial bridge (McAlpine, 1989).
The latter author presented the only known bionomic data A study of sepsid phylogeny using larval characters found
on Ropalomeridae, including rearings of Ropalomera  sp. only one character (the presence of spines on the creeping
from a gummy secretion of Enterolobium timbouva, of welts) supporting the relationship with ropalomerids (Meier,
W. pleuropunctata apparently from wood of Cocos nucifera 1995). On the other hand, there were more characters from
and a stump of a species of Musa, and of R.  stictica from immatures that suggested a sister-group relationship be-
resin of Spondias lutea. Aldrich (1932) mentioned Rhytidops tween the Sepsidae and the Coelopidae. We still find the
floridensis (Aldrich) breeding in decayed wood of Sabal relationship based on adult morphology convincing, how-
ever, because it is based on characters found throughout both
palmetto and S. minor (Steyskal, 1987).
families, whereas the larval synapomorphies supporting the
alternative point of view are restricted to basal sepsids.
Classification The Ropalomeridae consist of eight genera and 26 valid
species restricted to tropical and subtropical America. Only
Ropalomeridae belong to the superfamily Sciomyzoidea, one species has been found north of Mexico, Rhytidops
a group including Coelopidae, Dryomyzidae, Helosciomyzi- floridensis, and three species occur as far south as northern
dae, Sciomyzidae, and Sepsidae. The morphological evidence Argentina. Most species occur in the central portion of South
from adults suggest that Ropalomeridae are the sister group America (Steyskal, 1967, 1987). Other genera that do not
of Sepsidae, based on the following synapomorphic charac- occur in Central America are Apophorhynchus Williston and
ters: presence of spiracular setulae, vein R1 bare, A2 absent, Dactylissa Fischer.

2 Ropalomera 3 Ropalomera 4 Willistoniella

5 Kroeberia 6 Lenkokroeberia 7 Mexicoa

Figs. 76.2–7. Heads: lateral view of (2) Ropalomera femorata (Fabricius); anterior view of (3) R. femorata; lateral view of (4) Willistoniella
pleuropunctata (Wiedemann); (5) Kroeberia fuliginosa Lindner; (6) Lenkokroeberia chryserea Prado; and (7) Mexicoa mexicana Steyskal.
Figures 2–7 redrawn from Ramírez-García & Hernández-Ortiz (1994) figs. 1A, 1B, 1F, 2B, 2D, and 2F, respectively.
ROPALOMERIDAE (ropalomerid flies) 76 1027

Identification Specimens can be obtained by the use of fruit fly traps,

such as the Van Someren Rydon trap baited with fermented
The Ropalomeridae (formerly spelled Rhopalomeridae by fruit, with the use of Malaise traps, or by catching them
many authors) were first revised by Lindner (1930) and later with entomological nets especially when they rest on tree
by Prado (1963, 1966). The catalog of the Diptera of Ameri- trunks. The specimens should be pinned and stored dry. For
ca north of Mexico (Foote, 1965) includes only one species, species recognition, it is sometimes necessary to examine
Rhytidops floridensis. In the last catalog of this family for the the male genitalia. The best method is to separate the ter-
Neotropical Region, Steyskal (1967) recorded eight genera minalia from the rest of the body, macerate the soft tissues
and 29 species, of which three species are unrecognizable, with 10% NaOH or KOH at room temperature for about
and two are now synonymized.
24–48 h, wash with acidified water, and dehydrate with a
Recent contributions to knowledge of the taxonomy of the
family were published by Baez (1985), who recorded five species gradual ethanol series (50%, 70%, 96%, 100%), 10 min in
for Venezuela; Ibáñez-Bernal et al. (1992), who published on the each. Transfer the terminalia to Haya’s creosote or clove
species found in the Biosphere Reserve of Sian Ka’an (Quintana oil for clearing. The terminalia can be stored on the same
Roo, Mexico); and Ramírez-García & Hernández-Ortiz (1994), pin with the rest of the specimen inside a microvial filled
who presented a revision of Mexican ropalomerids, including with pure glycerol and a drop of 70% ethanol. The termina-
five genera and six species, proposing one new synonymy and lia can be observed, variously oriented, under a compound
describing one new species. A taxonomic revision of two genera microscope using a depression microscope slide without
was presented by Marques and Ale-Rocha (2004, 2005). deforming the structures.

8 Ropalomera 10 Kroeberia

9 Willistoniella

11 Lenkokroeberia 12 Mexicoa

Figs.  76.8–12. Hind legs: posterior view of (8)  Ropalomera femorata (Fabricius); (9)  Willistoniella pleuropunctata (Wiedemann);
(10) Kroeberia fuliginosa Lindner; (11) Lenkokroeberia chryserea Prado; and (12) Mexicoa mexicana Steyskal. Figures 8–12 redrawn from
Ramírez-García and Hernández-Ortiz (1994) figs. 4A, 4C, 4D, 4E, and 4F, respectively.
1028
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Key to the genera of Ropalomeridae of the world

1. Postpronotal bristles present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

– Postpronotal bristles absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

2. Arista bare or microscopically pilose (Figs. 5–6) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

– Arista with long rays or bipectinate (Figs. 2–4). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

3. Face with median vertical carina (similar to Fig. 4); genal surface striated; scutellum bare on disc
with only one pair of marginal and one pair of apical scutellar bristles; hind tibia weakly flattened;
Florida (USA), Argentina. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Rhytidops Lindner]
– Face with small median rounded tubercle (Figs. 3, 5–7); genal surface smooth; scutellum setulose on
disc and with two to five pairs (including apical pair) of scutellar bristles; hind tibia strongly
flattened (Figs. 10–11). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

cerc

pgt
cerc

sur

pgt
13 Ropalomera
sur

14 Willistoniella

sur

pgt

cerc
cerc

sur

15 Kroeberia
pgt

16 Mexicoa

Figs. 76.13–16. Male terminalia: posterior view of (13) Ropalomera femorata (Fabricius); (14) Willistoniella pleuropunctata (Wiedemann);
(15) Kroeberia fuliginosa Lindner; and (16) Mexicoa mexicana Steyskal. Figures 13–16 redrawn from Ramírez-García and Hernández-Ortiz
(1994) figs. 5C, 7C, 8C, and 9C, respectively.
Abbreviations: cerc, cercus; pgt, postgonite; sur, surstylus.
ROPALOMERIDAE (ropalomerid flies) 76 1029

4. Two postpronotal bristles on each side; ocellar bristles well developed; posterior thoracic spiracle
with more than three bristles; scutellum convex dorsally; scutellum with two to three pairs of
scutellar bristles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kroeberia Lindner
– One postpronotal bristle on each side; ocellar bristles absent; posterior thoracic spiracle with one
to three bristles; scutellum weakly concave or grooved dorsally with four to five pairs (including
apical pair) of scutellar bristles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lenkokroeberia Prado

5. Face with median vertical carina that ends as hump between antennal bases (Fig. 4); posterior
thoracic spiracle with four to seven bristles; scutellum flattened on dorsal surface . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Willistoniella Mik
– Face with rounded protuberance distant from antennal bases (Figs.  2–3); posterior thoracic
spiracle with one to three bristles; dorsal surface of scutellum weakly concave. . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ropalomera Wiedemann

6. Arista furnished on all sides with flattened hairs (Fig. 7); scutellum with two pairs (including apical
pair) of scutellar bristles; posterior thoracic spiracle with five to six bristles. . . Mexicoa Steyskal
– Arista microscopically pubescent or bipectinate, but not densely plumose; only apical pair of
scutellar bristles present; posterior thoracic spiracle with one to four bristles . . . . . . . . . . . . . . . 7

7. Arista bipectinate or long plumose; ocellar bristles absent; first flagellomere short and rounded,
less than 2 times as long as broad; Bolivia, Brazil. . . . . . . . . . . . . . . . [Apophorhynchus Williston]
– Arista microscopically pubescent; ocellar and postocellar, inner and outer vertical bristles
present; first flagellomere digitiform and elongated, more than 2 times longer than broad;
Brazil, Paraguay. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Dactylissa Fischer]

Synopsis of the fauna (type locality), Mexico (Oaxaca), Venezuela, and Costa Rica
(unpublished data); and L. ciliata (Williston) only recorded
Published records for Central America and Mexico con- from Brazil (type locality), and Bolivia.
firm the presence of five genera and six species in this region.
In Mexico all five genera and six species are present (Ramírez- Mexicoa Steyskal. This is a monotypic genus with a dis-
García & Hernández-Ortiz, 1994), but in the other countries tinctive species, M.  mexicana Steyskal, known only from
of Central America only three genera and four species have the states of Sonora (type locality), Guerrero, Jalisco, and
been recorded, demonstrating the scarcity of studies in the Morelos in Mexico.
area (there are only records from Costa Rica, Guatemala,
Nicaragua, and Panama). Ropalomera Wiedemann. This genus contains more than
half of all known ropalomerid species. Fifteen valid species
Kroeberia Lindner. This genus includes two species, can be recognized using the key presented by Prado (1966),
K. fuliginosa Lindner and K. minor Marques & Ale-Rocha. except for Ropalomera latiforceps Ramírez-García  &
Kroeberia fuliginosa was previously recorded for Mexico Hernández-Ortiz, described from Mexico and Panama in
(Campeche, Chiapas, Guerrero, Jalisco, Oaxaca, Quintana 1994. There are records of two species in Central America
Roo, Veracruz, Yucatán), Panama, Venezuela, and Brazil, but and Mexico. Newly examined material from Costa Rica
we have seen specimens from Costa Rica. Kroeberia minor included the well-known R. femorata (Fabricius), at least.
is known only from Brazil (Pará and Amazonas). Marques &
Ale-Rocha (2004) revised this genus. Willistoniella Mik. This genus is represented by W. pleuro-
punctata, a species widely distributed from central Mexico
Lenkokroeberia Prado. Two described species are includ- to northern Argentina. Marques & Ale-Rocha (2005) revised
ed in this genus: L. chryserea Prado known to occur in Brazil the genus.

Literature cited

Aldrich, J.M. 1932. New Diptera, or two-winged flies, from Amer- Baez, M. 1985. Los Ropaloméridos de Venezuela (Diptera:
ica, Asia, and Java, with additional notes. Proceedings of the Ropalomeridae). Boletín Entomológico Venezolano, N.S. 4:
United States Natural Museum 81: 1–28. 77–81.
1030
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Fischer, C.R. 1932. Um gênero e duas espécies novas de Rho- Marques, A.P.C.  & R. Ale-Rocha. 2005. Revisão do gênero Wil-
palomeridae do Brasil, e o pupário de Willistoniella pleuro- listoniella Mik, 1895 (Diptera, Ropalomeridae) da Região Neo-
punctata Wied. (Dipt.). Revista de Entomologia 2: 441–450. tropical. Revista Brasileira de Entomologia 49: 210–227.
Foote, R.H. 1965. Ropalomeridae, p. 679. In Stone, A., C.W. Sa- McAlpine, J.F. 1989. Phylogeny and Classification of the Musco-
brosky, W.W. Wirth, R.H. Foote, & J.R. Coulson (editors). A morpha, pp. 1397–1518. MND, Volume 3.
Meier, R. 1995. Cladistic analysis of the Sepsidae (Cyclorrhapha:
catalog of the Diptera of America north of Mexico. United
Diptera) based on a comparative scanning electron microscopic
States Department of Agriculture, Agriculture Handbook study of larvae. Systematic Entomology 20: 99–128.
276, Washington, D.C., iv + 1696 pp. Prado, A.P. do. 1963. Primeira contribuição ao conhecimento da
Ibáñez-Bernal, S., O. Canul-González, & J.F. Camal-Mex. 1992. familia Rhopalomeridae (Diptera). Memórias do Instituto Os-
Ropalomeridae (Diptera) de la Reserva de la Biosfera de Sian waldo Cruz 61: 459–470.
Ka’an, Quintana Roo, México. Folia Entomologica Mexi- Prado, A.P. do. 1966. Segunda contribuição ao conhecimento da
cana 84: 85–103. familia Rhopalomeridae (Diptera, Acalyptratae). Studia Ento-
Lindner, E. 1930. Revision der amerikanischen Dipteren-Familie mologica 8: 209–268. (1965)
der Rhopalomeridae. Deutsche Entomologische Zeitschrift Ramírez-García, E.  & V. Hernández-Ortiz, 1994. Revisión de la
1930: 122–137. familia Ropalomeridae (Diptera) en México. Acta Zoológica
Mexicana (N.S.) 61: 57–85.
Lopes, H.S. 1932. Sôbre a Rhopalomera stictica Wied., 1828
Steyskal, G.C. 1967. 60. Family Ropalomeridae. In Papavero, N.
(Diptera: Rhopalomeridae). Anais da Academia Brasileira de (editor). A Catalogue of the Diptera of the Americas South of
Ciências 4: 127–129. the United States. Departamento de Zoologia, Secretaria da Ag-
Marques, A.P.C. & R. Ale-Rocha. 2004. Revisão do gênero Kroe- ricultura, São Paulo, 7 pp.
beria Lindner (Diptera, Ropalomeridae) da Região Neotropical. Steyskal, G.C. 1987. Ropalomeridae, pp.  941–944. MND,
Revista Brasileira de Entomologia 48: 315–322. Volume 2.
SEPSIDAE (black scavenger flies) 77
Vera C. Silva

Fig. 77.1. Male of Sepsis punctum (Fabricius), (Nearctic, MND, fig. 86.1).

Diagnosis with basal constriction, responsible for antlike appearance

of the members of this family.
Small to medium-sized (body length 2–7  mm) slender Sepsids are recognized by the unbroken costal vein of the
flies (Fig.  1). Body usually brown to blackish, sometimes wing, the presence of one or more distinct setae on the
shining or with metallic purplish color. Wing hyaline, but posteroventral margin of the posterior spiracle, and the
dark spot near tip sometimes present (Figs.  6–7). Head antlike habitus.
more or less globular (Figs.  2–4); eye large; vibrissa lack-
ing, but vibrissal angle bearing several strong setae; antenna Biology
short with arista bare; palpus usually minute. Thorax usu-
ally shining, but sometimes dull dorsally; anepisternum with The larvae feed on a range of decaying organic materials,
some whitish pruinose areas. Male forefemur and tibia often but most frequently on mammalian feces. Nothing has been
modified, with bristles, spines, tubercles, or emarginations published about the biology of the family in the Neotropi-
used to grasp base of wing of females during copulation cal Region, except a few studies focusing on reproductive
(Figs.  9–13). Abdomen (Figs.  14–17) narrow, sometimes behavior of sepsids in Costa Rica (see below). Pont (1979)

1031
1032
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

and Pont & Meier (2002) summarized the knowledge of the in detail aspects of sexual selection of sepsids in Costa Rica
Palearctic Region fauna. with particularly interesting results (see Eberhard 2000a, b,
Neotropical sepsids have been collected around cowpats 2001a,  b,  c, 2002; Eberhard  & Pereira, 1997; Eberhard  &
and sometimes around feces of wild mammals (V.C. Silva, Huber, 1998). Males wait near oviposition sites such as dung
unpublished data). They are easily found resting on low pats or carrion and mount females when they arrive to lay
herbage, even in places away from cattle pats, or away from eggs. The male typically grasps the female wings with two
water (Eberhard, 2000a). different sets of structures. The modified front leg is used to
The family has fascinating courtship patterns and repro- clasp the base of each of the female’s wings (Hennig, 1949;
ductive behavior. Parker (1972a, b) studied the reproductive Parker, 1972a, b), and, when initiating copulation, the male
behavior of Sepsis cynipsea (Linneaus). Eberhard has studied grasps the female near the tip of her abdomen with a pair

2 Meropliosepsis 4 Lateosepsis

3 Sepsis

5 Microsepsis 6 Sepsis

7 Meropliosepsis 8 Pseudopalaeosepsis

Figs. 77.2–8. Heads and wings: lateral view of head of (2) Meropliosepsis sexsetosa Duda, (Silva, 1992, fig. 1); (3) Sepsis secunda
(Melander & Spuler), (Silva, 1995, fig. 4); and (4) Lateosepsis laticornis (Duda), (South America, Silva, 1993, fig. 45); dorsal view of wing
of (5) Microsepsis furcata (Melander & Spuler), (Ozerov, 1994, fig. 19); (6) S. secunda, (Silva, 1995, fig. 1); (7) Me. sexsetosa, (Silva, 1992,
fig. 2); and (8) Pseudopalaeosepsis nigricoxa Ozerov, (Ozerov, 1994, fig. 173).
SEPSIDAE (black scavenger flies) 77 1033

of clasping organs, the genitalic surstyli (Parker, 1972a, b; Classification

Eberhard & Pereira, 1997). Males only mate with females
after the oviposition from a prior copulation is finished (Ward Sepsidae have been assigned to the superfamily Scio-
et al., 1992; Eberhard, 2000a). myzoidea by Hennig (1958, 1973), Griffiths (1972), and
The egg and larval morphology of sepsids have been McAlpine (1989), although the concept of the superfamily
extensively studied by Meier (1995a, b; 1996) and Ozerov & varies among these authors. McAlpine (1989) considered
Meier (1995). Meier (1996) described the third larval stage the Sciomyzoidea to be composed of Coelopidae, Dryo-
of three Neotropical species (Figs. 30, 31). Previous studies myzidae, Sciomyzidae, Ropalomeridae, Sepsidae, and
are summarized in Ferrar (1987). Helosciomyzidae.

9 Meroplius 10 Meropliosepsis

11 Microsepsis 12 Palaeosepsis

13 Archisepsis

Figs.77.9–13. Forefemora and tibiae: posterior view of (9)  Meroplius albuquerquei Silva, (South America, Silva, 1990, fig.  2);
(10) Meropliosepsis sexsetosa Ozerov, (Ozerov, 1994, fig. 56); (11) Microsepsis furcata (Melander & Spuler), (Silva, 1993, fig. 140);
(12) Palaeosepsis dentata (Becker), (Ozerov, 1994, fig. 70); (13) Archisepsis discolor (Bigot), (Silva, 1993, fig. 115).
1034
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

The Ropalomeridae is usually regarded as the sister group The Neotropical sepsid fauna includes the genera
of the Sepsidae (McAlpine, 1989; Meier, 1996). Meier’s Archisepsis, Lateosepsis, Meropliosepsis, Meroplius,
(1995a) analysis of larval features included some characters Microsepsis, Palaeosepsioides, Palaeosepsis, Pseudo-
suggesting a sister-group relationship between Sepsidae and palaeosepsis, Sepsis, and Themira. The species among the
Coelopidae. He favored the traditional view, however, based genera are morphologically quite similar, making it difficult
on characters that are found throughout the Sepsidae, where- to characterize the genera. Species are usually based on male
as the larval characters that supported a connection with the features, and in some genera, females cannot be identified at
Coelopidae are restricted to basal sepsids (mainly Orygma the species level.
Meigen). It is not easy to estimate the undescribed sepsid diver-
The monophyly of the family is widely accepted sity in the Neotropical Region. Ozerov (1995) described
five new genera and five new species from Africa. Possibly,
(Griffiths,1972; Hennig, 1973; McAlpine, 1989). Two sub-
there are many more in other poorly sampled areas, such as
families are recognized (Zuska, 1977, 1980; Steyskal, 1987):
Papua New Guinea and the Neotropical Region. Recently,
Orygmatinae, with one Holarctic monotypic genus, and Sep- two authors have published the results of their studies of
sinae. Traditionally, the Sepsinae have been broken into three the Neotropical sepsid fauna. The first two papers (Silva,
tribes: Saltellini, Sepsini, and Toxopodini (Steyskal, 1987). 1990, 1991) covered small areas of Brazil, but resulted in
Ozerov (1994) proposed a tribe, Palaeosepsini, to include the description of two new species. In a study that included
most Neotropical genera. Both Sepsini and Palaeosepsini have the examination of more than 7600 specimens from many
representatives in the Neotropical Region. Meier (1995a, areas of the Neotropical Region, Silva (1993) found eight
1996) discussed the classification schemes proposed for the new species and described two new genera. The Nicaraguan
Sepsidae, but did not consider the tribe Palaeosepsini. His fauna was revised by Maes & Silva (1999). Ozerov, another
cladistic analyses pointed out that Sepsini may not be mono- specialist on Neotropical sepsids, has described 19 new spe-
phyletic and that there is evidence for the monophyly of Tox- cies and five new genera from the region (Ozerov, 1992a,
opodini and Saltellini. 1993b, 1994, 1998, 2000, 2004). The most studied areas are

tg 1
st 1

tg
st 2
tg
st 2

tg
st 3
tg
st 3

st 4
tg
st 4
tg
16 Palaeosepsis
st
tg 5
st
tg 5

st 6 tg
st 6
st 7
tg 6 st 7
tg

14 Archisepsis 15 Archisepsis

17 Sepsis

Figs. 77.14–17. Abdomens: ventral view (14) and lateral view (15) of Archisepsis ecalcarata (Thomson), (Eberhard and Pereira, 1997, fig. 5);
lateral view of first three segments of (16) Palaeosepsis dentatiformis (Duda), (Silva, 1993, fig. 3); and (17) Sepsis secunda (Melander &
Spuler), (Silva, 1995, fig. 3).
Abbreviations: st, sternite; tg, tergite.
SEPSIDAE (black scavenger flies) 77 1035

St. Vincent (West Indies), Nicaragua, Costa Rica, Panama, The chaetotaxy of the head and thorax, as well as the
Ecuador, Peru, Brazil, Paraguay, and Argentina. shape and texture of the abdomen, are used for identifying
genera. When identifying specimens to the species level, it
Identification is necessary to observe the male forefemur and tibia for their
emarginations and vestiture and also the wing venation. Dry
Duda (1926a,  b) wrote a monograph of the family, and specimens are best observed when double mounted. There
Steyskal (1968) published the Neotropical catalog, now are no keys to larvae and pupa for the Neotropical Sepsidae,
superseded by the World catalog published by Ozerov but the keys provided by Teskey (1981) and Ozerov & Meier
(2005). Since the recent efforts to describe new Neotropical (1995) are useful. Meier (1996) provided a key to some gen-
species (see above), Ozerov (2004) published a key to the era and a key for the four species of Palaeosepsis he had
genera, but it does not include all the genera known to occur studied.
in the Neotropical Region.

19 Archisepsis

18 Archisepsis 20 Pseudopalaeosepsis

21 Pseudopalaeosepsis 22 Archisepsis

23 Archisepsis

Figs. 77.18–23. Male terminalia: posterior view (18) and lateral view (19) of epandrium and surstyli of Archisepsis pusio (Schiner), (Silva,
1993, figs. 61, 62); posterior view (20) and lateral view of epandrium and surstyli (21) of Pseudopalaeosepsis nigricoxa Ozerov, (Ozerov,
1994, figs. 175, 176); posterior view (22) and lateral view of epandrium and surstyli (23) of A. discolor (Bigot), (Ozerov, 1994, figs. 100, 101).
1036
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

cerc

cerc

24 Microsepsis 25 Microsepsis 26 Microsepsis

27 Microsepsis 28 Microsepsis 29 Microsepsis

fc mask a bd spn

ant l tub
d tub

mx plp spr sk

v tub

l lip

vl sens

30 Archisepsis p bd spn

31 Archisepsis

Figs. 77.24–31. Male terminalia (concluded): posterior view (24), lateral view of left side (25), and lateral view of right side (26) of
Microsepsis armillata (Melander & Spuler), (Silva, 1993, figs. 125–127); posterior view (27), lateral view of left side (28), and lateral view
of right side (29) of M. simplicula (Steyskal), (Silva, 1993, figs. 174, 176, 177); lateral view of cephalic region (30) and terminal segment (31)
of third instar larva of Archisepsis polychaeta (Ozerov), (Meier, 1996, figs. 115, 119).
Abbreviations: a bd spn, anterior band of spines; ant, antenna; cerc, cercus; d tub, dorsal tubercle; fc mask, facial mask; l tub, lateral
tubercle; l lip, lower lip; mx plp, maxillary palpus; p bd spn, posterior band of spines; spr sk, spiracular stalk; vl sens, ventrolateral sensilla;
v tub, ventral tubercle.
SEPSIDAE (black scavenger flies) 77 1037

Key to the genera of Sepsidae of the Neotropical Region

1. One pair of strong, well-developed fronto-orbital setae present (Fig. 2); abdomen of both sexes
without strong setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Fronto-orbital setae lacking, or when present, only vestigial (Figs. 3–4); abdomen of some species
with setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

2. Three pairs (1+2) of strong dorsocentral setae and three pairs of vibrissae; base of wing, subcostal
cell and apex of R2+3 with dark spot (Fig. 7) . . . . . . . . . . . . . . . . . . . . . . . . . Meropliosepsis Duda
– Only one pair of strong dorsocentral setae present; R2+3 without apical dark spot . . . . . . . . . . . . 3

3. Outer vertical seta present; supra-alar seta present; basal scutellar seta absent; worldwide, one
species in southern Brazil. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Meroplius Rondani]
– Outer vertical seta absent or short, hairlike; supra-alar seta absent; basal scutellar seta present . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Themira Robineau-Desvoidy

4. Male terminalia asymmetrical (Figs.  24–29); vein CuA2+A1 extremely reduced, shorter than
length of cell cup (Fig. 5); male forefemur with (Fig. 11) or without emarginations. . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microsepsis Silva
– Male terminalia symmetrical (if asymmetrical, other characters do not correspond) (Figs. 18–23);
vein CuA2+A1 of normal length (as in Figs. 6, 8); male forefemur with elaborate emarginations
(Figs. 12–13) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

5. Syntergite 1+2 without posterior, transverse, wide elevation; abdominal tergites usually rugose,
with hairs and setae usually fully developed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
– Syntergite 1+2 with elevated posterior margin (Figs.  16, 17); abdominal tergites polished and
shining, with hairs and setae undeveloped. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

6. Postmetacoxal bridge present; forecoxa black. . . . . . . . . . . . . . . . . . . Pseudopalaeosepsis Ozerov

– Postmetacoxal bridge absent; forecoxa yellowish brown to yellow . . . . . . . . . . . Archisepsis Silva

7. Postalar seta weak; abdominal sternite 4 absent in males; tergite  4 modified with constriction
before posterior margin; tergite 5 reduced to two small lateral plates (Fig. 17) . . . . . Sepsis Fallén
– Postalar seta present, well developed; sternite 4 present in males; tergites 4 and 5 not modified. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

8. Postmetacoxal bridge absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Palaeosepsioides Ozerov

– Postmetacoxal bridge present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

9. Male abdominal sternite 4 simple; anal lobe of wing well developed; alula entirely covered with
microtrichia, width of alula approximately 2 times as long as wide cell bm; basal costal cell clear;
South America. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Lateosepsis Ozerov]
– Male abdominal sternite 4 with tuft of setae laterally; anal lobe of wing narrow; alula without
microtrichia, width of alula approximately equal to width of cell bm; basal costal cell blackish. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Palaeosepsis Duda
1038
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Synopsis of the fauna tropicanus Ozerov, is known from Costa Rica. References
for this genus are Silva (1991) and Ozerov (1992a, 2004).
Sepsidae are well represented in all zoogeographic re-
gions, with 37 recent genera and 318 species in the world Palaeosepsis Duda. Nine species of Palaeosepsis are
(Ozerov, 2005). The family is noticeably absent in Chile. known to be present in the Neotropical Region, of which
Central America contains 25 described species in eight gen- three occur in Central America. It used to be the largest
era. They are found in all terrestrial habitats, but are more Neotropical genus until it was revised independently by
diverse in rain and cloud forests. Shannon traps using dung Silva (1991, 1993) and Ozerov (1992a, b, 1994, 1997).
as bait can collect huge numbers of specimens.
Pseudopalaeosepsis Ozerov. There are three species
Archisepsis Silva. Thirteen valid species have been de- in this genus that are endemic to the Neotropical Region,
scribed for this mainly Neotropical genus, of which eight are one of which, P. nigricoxa Ozerov, is known to occur in
found in Central America. One species, A. discolor (Bigot), Central America. References for this genus are Ozerov
was introduced in the Azores Islands. (Ozerov, 2005). Ref- (1992a, 1993a, b, 1994) and Silva (1993).
erences to this genus can be found in Melander  & Spuler
(1917, in Sepsis), Silva (1993), and Ozerov (1993a, b, 1994, Sepsis Fallén. This worldwide genus has 78 valid
2004). species, but just three are known to occur in North and
Central America, and only one, S. secunda (Melander &
Meropliosepsis Duda. This genus is known from a single Spuler), is found in the Neotropical Region (Costa Rica,
species, M. sexsetosa Duda, distributed from Costa Rica to Cuba, and Nicaragua). References can be found in Silva
southern Brazil (Duda 1926b; Silva, 1992). (1995, as a new genus Nicarao) and Ozerov (2000,
Microsepsis Silva. Thirteen species have been described 2005).
for this exclusively Neotropical genus, of which nine are
Themira Robineau-Desvoidy. This genus has 32
found in Central America. Ozerov (1994) has a key to nine
valid species worldwide, mostly Holarctic, impinging on
of the Neotropical species; other references dealing with
Microsepsis are Silva (1993) and Ozerov (1992b, 1993b). the Neotropical Region in Mexico and Belize (Pont  &
Meier, 2002; Ozerov, 2005). Only one species is known
Palaeosepsioides Ozerov. Three species were described to occur in Central America. Ozerov (1999) presented a
in this exclusively Neotropical genus, of which one, P. neo- review of the world species with keys.

Literature cited
Duda, O. 1926a. Monographie der Sepsiden (Dipt.). I. Annalen des Eberhard, W.G. & B.A. Huber. 1998. Copulation and sperm trans-
Naturhistorischen Museums Wien 39: 1–153. fer in Archisepsis flies (Diptera, Sepsidae) and the evolution of
Duda, O. 1926b. Monographie der Sepsiden (Dipt.). II. Annalen des their intromittent genitalia. Studia dipterologica 5: 217–248.
Naturhistorischen Museums Wien 40: 1–110. Eberhard, W.G. & F. Pereira. 1997. Functional morphology of male
Eberhard, W.G. 2000a. Mating systems of sepsid flies and sexual genitalic surstyli in the dungflies Achisepsis [sic] diversiformis
behavior away from oviposition sites by Sepsis neocynipsea and A.  ecalcarata (Diptera: Sepsidae). Journal of the Kansas
(Diptera: Sepsidae). Journal of the Kansas Entomological So- Entomological Society 69 (Suppl.): 43–60.
ciety 72: 129–130. Ferrar, P. 1987. A guide to the breeding habits and immature stages
Eberhard, W.G. 2000b. Behavior and reproductive status of Microsepsis of Diptera Cyclorrhapha. Entomonograph 8(1). E.J.  Brill  /
armillata (Diptera: Sepsidae) flies away from oviposition sites. Scandinavian Science Press, Leiden, Copenhagen, 478 pp.
Annals of Entomological Society of America 93: 966–971. Griffiths, G.C.D. 1972. The phylogenetic classification of Diptera
Eberhard, W.G. 2001a. The functional morphology of species- Cyclorrhapha, with special reference to the structure of the male
specific clasping structures on the front legs of male Archisepsis postabdomen. Series entomologica 8. Dr. W. Junk, The Hague,
and Palaeosepsis flies (Diptera: Sepsidae). Zoological Journal 340 pp.
of the Linnean Society 133: 335–368. Hennig, W. 1949. 39a. Sepsidae. In Lindner,  E. (editor). Die
Eberhard, W.G. 2001b. Species-specific genitalic copulatory court- Fliegen der palaearktischen Region. 157, 159: 1–91.
ship in sepsid flies (Diptera, Sepsidae, Microsepsis) and theories E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart.
of genitalic evolution. Evolution 55: 93–102. Hennig, W. 1958. Die Familien der Diptera Schizophora und ihre
Eberhard, W.G. 2001c. Multiple origins of a major novelty: move- phylogenetischen Verwandtschaftsbeziehungen. Beiträge zur
able abdominal lobes in male sepsid flies (Diptera: Sepsidae), Entomologie 8: 505–688.
and the question of developmental constraints. Evolution and Hennig. W. 1973. Ordnung Diptera (Zweiflügler), pp.  1–337. In
Development 3: 206–222. Helmcke, J.-G., D. Starck, and H. Wermuth (editors). Handbuch
Eberhard, W.G. 2002. The function of female resistance behavior: der Zoologie. 4(2) 2/31 (Lfg. 20). Walter De Gruyter, Berlin &
intromission by male coercion vs. female cooperation in sep- New York.
sid flies (Diptera: Sepsidae). Revista de Biología Tropical 50: McAlpine, J.F. 1989. Phylogeny and classification of the Musco-
485–505. morpha, pp. 1397–1518. MND, Volume 3.
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Maes, J.M. & V.C. Silva. 1999. Familia Sepsidae, pp. 1569–1570. Ozerov, A.L. 2005. World Catalogue of the family Sepsidae (In-
In Maes, J.M. (editor). Insectos de Nicaragua, Volume 3. SET- secta: Diptera). Zoologicheskie issledovania 8: 1–74.
AB, MARENA, Managua, Nicaragua, 700 pp. Ozerov, A.L. & R. Meier. 1995. A key to the larvae of the Palaearctic
Meier, R. 1995a. Cladistic analysis of the Sepsidae (Cyclorrhapha: genera of Sepsidae (Diptera: Cyclorrhapha). Annales de la So-
Diptera) based on a comparative scanning electron microscopic ciété Entomologique de France, Nouvelle Série 31: 259–283.
study of larvae. Systematic Entomology 20: 99–128. Parker, G.A. 1972a. Reproductive behaviour of Sepsis cynipsea (L.).
Meier, R. 1995b. A comparative SEM study of the eggs of the Sep- I. A preliminary analysis of the reproductive strategy and its
sidae (Diptera) with a cladistic analysis based on egg, larval and associated behavioural patterns. Behaviour 41: 172–206.
adult characters. Entomologica scandinavica 26: 425–438. Parker, G.A. 1972b. Reproductive behaviour of Sepsis cynip-
Meier, R. 1996. Larval morphology of the Sepsidae (Diptera: Scio- sea (L.). II. The significance of the precopulatory passive phase
myzoidea), with cladistic analysis using adult and larval charac- and emigration. Behaviour 41: 241–250.
ters. Bulletin of the American Museum of Natural History 228: Pont, A.C. 1979. Sepsidae. Diptera. Cyclorrapha Acalyptrata. Hand-
1–147. books for the Identification of British Insects 10(5c): 1–35.
Melander, A.L. & A. Spuler. 1917. The dipterous family Sepsidae Pont, A.C.  & R. Meier. 2002. The Sepsidae (Diptera) of Europe.
and Piophilidae. Bulletin of the Washington Agricultural Exper- Fauna Entomologica Scandinavica. 37: 1–198. Brill, Leiden,
imental Station 143: 1–103. Boston, Köln.
Ozerov, A.L. 1992a. New data on the neotropical sepsids (Diptera, Silva, V.C. 1990. Revision of the family Sepsidae of the Neotropi-
Sepsidae). Russian Entomological Journal 1: 81–86. cal Region. I. The genus Meroplius Rondani (Diptera, Schizo-
Ozerov, A.L. 1992b. The results of the examination of Sepsis in- phora). Revista brasileira de Entomologia 34: 709–711.
sularis Williston, 1896 and Sepsis furcata Melander et Spuler, Silva, V.C. 1991. Levantamento preliminar de Sepsidae (Diptera,
1917 (Diptera, Sepsidae). Russian Entomological Journal 1: Schizophora) em Roraima, Projeto Maracá, com descrição de
75. uma espécie nova. Acta amazônica 21: 369–374.
Ozerov, A.L. 1993a. Four new synonyms in the family Sepsidae Silva, V.C. 1992. Revision of the family Sepsidae of the Neotropi-
(Diptera). Entomologicheskoye obozreniye 72: 248–249 cal Region. II. The genus Meropliosepsis Duda, 1926 (Diptera,
Ozerov, A.L. 1993b. Six new species of the genus Palaeosepsis Schizophora). Revista brasileira de Entomologia 36: 549–552.
Duda (Diptera, Sepsidae). Russian Entomological Journal 2: Silva, V.C. 1993. Revisão da família Sepsidae na região neotropical.
63–71. III. Os gêneros Palaeosepsis Duda, 1926, Archisepsis, gen. n. e
Ozerov, A.L. 1994. A review of the sepsid flies of the Palaeosepsini, Microsepsis, gen. n.: Chave para os gêneros neotropicais (Dip-
trib. n. (Diptera, Sepsidae). Russian Entomological Journal 2: tera, Schizophora). Iheringia (Zoologia) 75: 117–170.
55–103. (1993) Silva, V.C. 1995. A new genus of Sepsidae (Diptera, Schizophora)
Ozerov, A.L. 1995. To the fauna and taxonomy of African Sepsidae. from Nicaragua. Studia dipterologica 2: 203–206.
Russian Entomological Journal 4: 127–144. Steyskal, G.C. 1968. 63. Family Sepsidae. In Papavero, N. (editor).
Ozerov, A.L. 1997. Some types of Sepsidae in the Berlin and A Catalogue of the Diptera of the Americas south of the United
Eberswalde Museums (Diptera). Beiträge zur Entomologie 47: States. Departamento de Zoologia, Secretaria da Agricultura,
477–487. São Paulo, 6 pp.
Ozerov, A.L. 1998. Two new species of Sepsidae (Diptera). Russian Steyskal, G.C. 1987. Sepsidae, pp. 945–950. MND, Volume 2.
Entomological Journal 6: 83–85. Teskey, H.J. 1981. Key to families: Larvae, pp.  125–149. MND,
Ozerov, A.L. 1999. A review of the genus Themira Robineau- Volume 1.
Desvoidy, 1830 (Diptera, Sepsidae) of the World, with a revi- Ward, P.I., J. Hemmi, & T. Roösli. 1992. Sexual conflict in the dung
sion of the North American species. Russian Entomological
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Journal 7: 169–208. (1998)
Ozerov, A.L. 2000. A revision of the Nearctic species of the genus Zuska, J. 1977. A revision of the genus Parapalaeosepsis (Diptera,
Sepsis Fallén, 1810 (Diptera, Sepsidae). Russian Entomological Sepsidae). Acta entomologica Bohemoslovaca 74: 50–61.
Journal 9: 161–176. Zuska, J. 1980. Family Sepsidae, pp. 591–596. In Crosskey, R.W.
Ozerov, A.L. 2004. New genera and species of Sepsidae (Diptera). (editor). Catalogue of the Diptera of the Afrotropical Region.
Russian Entomological Journal 12: 399–420. (2003) British Museum (Natural History), London, 1437 pp.
1040
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
CLUSIIDAE (clusiid flies) 78
Owen Lonsdale and Stephen A. Marshall

78.1. Female of Sobarocephala picta Hennig, (illustrated by O. Lonsdale).

Diagnosis (Clusiodes and Hendelia; Fig. 2). Ocellar and/or postocellar

bristles usually small to absent, divergent when present. All
Central American species with one or two pairs of postsutur-
Relatively elongate flies, body length 2.5–6.0 mm in the
al dorsocentral bristles; sometimes with one well-developed
New World. Color various, but usually all or partly yellow
setula in front of anterior dorsocentral. Subcosta complete
(Craspedochaeta and Heteromeringia are often dark brown)
and subcostal break present (sometimes indistinct in Clusio-
with wing frequently infuscated (Fig.  1). Arista pubescent
dinae). Abdomen slender in all Central American species and
to densely plumose; vibrissa well developed; face usually
particularly tapered in females.
soft, receding. Outer and (sometimes) inner distal margins of
pedicel with triangular extensions, outer extension distinct Eggs (known for temperate Clusiodes, Australian Heter-
and acute in Clusiinae and Sobarocephalinae, blunt and ob- omeringia, and Neotropical Procerosoma, Sobarocephala,
tuse in Clusiodinae. Three to five (occasionally two) fronto- and Craspedochaeta) translucent, usually three to four times
orbital bristles in New World taxa, with the first, second, or longer than wide (2.5  times in species of Sobarocephala)
third bristle usually inclinate (Fig.  3); if all fronto-orbital and approximately as long as female sternite  6. Both ends
bristles reclinate, then one pair of interfrontal bristles present tapered, most pronounced anteriorly, with micropyle small

1041
1042
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

and terminal (nearly indistinct in Clusiodes). Egg surface rounded with ridge broken and shallow (Clusiodes). Anal
minutely tuberculate; usually with no more than 12 longitu- hooks bifid (Heteromeringia) or unbranched, curving dor-
dinal wrinkles that sometimes fade basally. sally (Clusiodes) or dorsomedially, with ovate (Clusia
Larvae (of European Clusiodes) with minute cephalic and Clusiodes) or semi-circular (Sobarocephala) cross
papillae, small well-sclerotized mandibles, and vestigial un- section.
pigmented cephalopharyngeal skeleton (Soós, 1987). Anterior Clusiids differ from similar slender acalyptrate flies in
spiracles with six openings, each on small raised tubercle having a porrect antenna with a dorsoapical arista (as op-
(Soós, 1987); posterior spiracles elevated on chitinized plates posed to dorsobasal) and strong triangular extensions on the
produced as dorsally curving hooks with three elongate-oval outer and sometimes inner distal margins of the pedicel. The
openings on ventral half (Malloch, 1918). Anal plate large, Neriidae have a strong extension on the inner margin, and
heavily pigmented, wider than long and tapered laterally the Ropalomeridae have shallow extensions on both sides,
(Malloch, 1918). but these are characteristically large-bodied flies. The Aul-
Puparia (known from Australian Heteromeringia and acigastridae have well-developed extensions on both sides
north temperate Clusia Haliday, Sobarocephala, and Clusiodes) of the pedicel, but the extension on the inner margin is more
approximately 5%–10% longer than adult and covered prominent and the subcosta is incomplete. Some species of
with numerous minute transverse wrinkles (see Lons- Chloropidae and Richardiidae superficially resemble clusi-
dale & Marshall, 2007b). Posterior face smooth with vari- ids, but chloropids have an incomplete subcosta, and the anal
ous sculpturing, and either sharply curved and bordered (cup) and basal medial cells are open; Richardiidae have no
by high ridge (Clusia and Sobarocephala) or broadly vibrissae and only a single pair of fronto-orbital bristles.

i vt s

frorb s poc s

o vt s

infr s

gn s
vb

sbvb s

2 Clusiodes 3 Sobarocephala

preap d s

4 Sobarocephala 5 Heteromeringia

Figs. 78.2–5. Heads and legs: anterodorsal view of head of (2) Clusiodes albimanus (Meigen), (Palaearctic, MND, fig. 70.2); (3) Sobarocephala
flaviseta (Johnson), (Nearctic, MND, fig. 70.3); anterior view of midtibia of (4) S. flaviseta, (MND, fig. 70.4); and (5) Heteromeringia nitida
Johnson, (Nearctic, MND, fig. 70.5).
Abbreviations: frorb s, fronto-orbital seta; gn s, genal seta; infr s, interfrontal seta; i vt s; inner vertical seta; o vt s; outer vertical seta; poc s,
postocellar seta; preap d s, preapical dorsal seta; sbvb s, subvibrissal seta; vb, vibrissa.
CLUSIIDAE (clusiid flies) 78 1043

Some Agromyzidae have similar venation and chaetotaxy, Classification

but they are smaller and stouter with a large, dark ovipositor
and usually with two or more inclinate fronto-orbital bristles, Frey (1960) divided the Clusiidae into two subfamilies,
three or more dorsocentral bristles, and/or several mediolat- Clusiodinae, characterized by reclinate fronto-orbital bris-
eral tibial bristles. The Anthomyzidae share a similar tles, and Clusiinae, with at least one pair of inclinate fron-
habitus, but the first flagellomere is distinctly downturned, to-orbital bristles. Subsequent authors, including Steyskal
the postocellar bristles are convergent or absent, and the sub- (1965), Soós (1987), Sasakawa (1977), and Pitkin & Even-
costa is incomplete. Some species of Lauxaniidae are also huis (1989), restricted the Clusiinae to those genera with only
similar in their habitus to clusiids, but the postocellar bristles the anterior fronto-orbital bristle inclinate, moving Craspe-
are convergent and the vibrissae are absent. dochaeta to the Clusiodinae. Neither of these subfamily
concepts, based on head chaetotaxy, correspond with clades
Biology defined by numerous synapomorphies of the male and female
genitalia and previously unrecognized external morphologi-
Although adult clusiids are rarely noticed in the field, cal characters. A new subfamilial classification proposed by
they are regularly collected in Malaise and pan traps and can Lonsdale & Marshall (2006a) divided the Clusiidae into the
be common on some substrates and in certain microhabitats. Clusiodinae, Clusiinae, and Sobarocephalinae (formerly part
Neotropical clusiids often occur in mossy, humid habitats, of the Clusiinae).
and many species are attracted to bird droppings and small Reclinate fronto-orbitals are probably synapomorphic for
deposits of mammal dung. Elsewhere, clusiids are known the redefined Clusiodinae, but this character is reversed in
Heteromeringia, which was redefined by Lonsdale & Mar-
to feed on nectar, rotting vegetation, and sap (Soós, 1987).
shall (2008b). The New World Clusiodinae includes four
Many Neotropical species, especially of Hendelia and Soba-
genera, all of which also occur throughout the Old World.
rocephala, have been collected along waterways. Conversely,
The New World Hendelia is restricted to the tropics, Craspe-
species of Heteromeringia prefer relatively open areas such dochaeta and Heteromeringia are tropical with several spe-
as tree falls, forest borders, pastures, and landslides. Some cies extending into the temperate regions to the north and
species frequent tree trunks or logs, but this habit seems less south, and Clusiodes is a predominantly north temperate
widespread in Neotropical species than in temperate species, group that has a single species extending into Neotropical
possibly because of the greater abundance of ants in the trop- Mexico (Lonsdale & Marshall, 2006b, 2007a, 2007b). The
ics. Males sometimes use bare logs or tree trunks as lek sites remaining extant clusiodine genus, Allometopon Kertész, is
(Roháček, 1995; Caloren & Marshall, 1998; Marshall, 2000; Indoaustralian in distribution.
Lonsdale  & Marshall, 2006b, 2007a), defending territories The Clusiidae with inclinate lower fronto-orbital bristles,
from other males and mating with females that later leave excluding the clusiodine Heteromeringia, are divided into
the lek sites to oviposit elsewhere. Although no information two clades, the subfamilies Clusiinae and Sobarocephalinae.
is available on the oviposition behavior of Neotropical spe- The Sobarocephalinae includes the large, primarily Neotro-
cies, some north temperate and Australian species oviposit pical Sobarocephala and two small, closely related Neotro-
in soft, rotting wood. Selection of oviposition site appears to pical genera (Chaetoclusia and Procerosoma) (Lonsdale &
be limited by humidity, amount of shade, and the presence Marshall, 2006a). The Clusiinae includes the Chilean and
of certain fungi (Roháček, 1995). Larvae and puparia have Argentinean genus Apiochaeta Czerny, the small Old World
been recovered from rotting deciduous wood and termite genera Phylloclusia Hendel, Tetrameringia McAlpine, and
galleries (Soós, 1987). Larvae are known to be slow mov- Melanoclusia Lonsdale  & Marshall, and the Holarctic and
ing, although observations of “leaping” behavior have been Oriental genus Clusia (Lonsdale & Marshall, 2006a, 2008a,
reported (Malloch, 1918). 2008c).
Fossil Clusiidae are known from Baltic amber, includ-
Males of some clusiids, particularly Hendelia, have en-
ing Electroclusiodes Hennig and several tentatively placed
larged heads or other conspicuous modifications used in
species (Hennig, 1965, 1969), but no Clusiidae have been
mutual assessment on lek sites (McAlpine, 1976; Marshall, described from Dominican amber.
2000). Hendelia kinetrolicros (Caloren  & Marshall), Hen-
delia mirabilis (Frey), and Procerosoma alini (Shatalkin)
have spectacular long genal processes that are probably used
Identification
in male–male agonistic interactions, although these species
The New World species of most clusiid genera can be
have not been observed while engaged in such behavior. identified using keys from recently published revisions that
Neotropical Craspedochaeta males have been observed to are listed below in the genus synopses. The majority of
exhibit head-to-head contact and associated agonistic inter- the neotropical fauna belongs to the genus Sobarocephala,
action (Lonsdale  & Marshall, 2006b), but intrasexual in- however, which contains dozens of undescribed species.
teractions otherwise remain unknown amongst Neotropical Neotropical and Old World revisions of this large genus are
Clusiidae. currently being developed.
1044
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Key to the genera of Clusiidae of the New World

1. Anterior fronto-orbital bristle reclinate (Fig. 2); mid- and hind tibiae with dorsal preapical bristles
(Fig. 4); CLUSIODINAE, excluding Heteromeringia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Anterior fronto-orbital bristle inclinate (Fig. 3); hind tibia lacking dorsal preapical bristle (Fig. 5);
midtibia various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

2. Interfrontal bristle absent; four or five pairs of fronto-orbital bristles present with third fronto-
orbital from posterior margin of head inclinate; genal bristles small and hairlike; mid- and hind
tibiae with one pair of dorsal preapical bristles; phallapodeme rodlike; hypandrial arm weakly
attached to remainder of hypandrium; ejaculatory apodeme mushroom-shaped; spermatheca clear
and annulated, but not telescoped; ventral receptacle rarely with minute subapical disc (Fig. 13).
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Craspedochaeta Czerny
– Interfrontal bristle present (Fig. 2); two or three well-developed fronto-orbital bristles present, all
reclinate, with anterior and posterior bristles reduced in length; several medial genal bristles more
strongly developed; mid- and hind tibiae with two pairs of dorsal preapical bristles; phallapodeme
keel-like (Figs. 9–10); hypandrial arm strongly fused to remainder of hypandrium + pregonite;
ejaculatory apodeme thin along length, sometimes with apex slightly widened; spermatheca
pigmented and strongly telescoped; ventral receptacle with large subapical disc (Fig. 12) . . . . . 3

3. Arista variably plumose, but not pubescent; if three fronto-orbital bristles present, anterior
two pairs usually closely spaced at anterior margin of frons; scutum variously patterned but not
with white lateral stripes; male: fore- and midfemora with at most one anterior and one posterior
row of ventral ctenidial bristles; perianal area longer than wide; surstylus usually small, lobate
and perpendicular to the long axis of the epandrium; ejaculatory apodeme widest apically . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hendelia Czerny
– Arista pubescent; if three pairs of fronto-orbital bristles present, middle pair midway between
anterior and posterior pairs; scutum usually with one pair of white lateral stripes; male: fore- and
midfemora with one anterior and two posterior rows of ventral ctenidial bristles; perianal area
wider than long; surstyli parallel to long axis of epandrium and usually large; ejaculatory
apodeme of uniform width or widest medially; Nearctic Region . . . . . . . . . [Clusiodes Coquillett]

4. Triangular extension on outer margin of pedicel obtuse and blunt; scutellum flat and wrinkled;
only one pair of hairlike lateral scutellar bristles present; midtibia without dorsal preapical tibial
bristle; middle fronto-orbital bristle not reduced, cell bm closed, and head without wide truncate
notch; ratio of ultimate to penultimate section of vein M various, but rarely less than 3.0–4.0;
epandrium small and usually much narrower than tergite 5; distiphallus extremely long, coiled,
dark, with one pair of well-sclerotized lateral ribs, not flanked by one pair of laterobasal lobes;
spermatheca strongly telescoped and dark (Fig. 14), elongate (Fig. 15) or more irregular in outline,
but not spherical; ventral receptacle with subterminal flagellum. . . . . . . . Heteromeringia Czerny
– Triangular extension on outer margin of pedicel acute and projecting; scutellum smooth and
distinctly (if only slightly) convex; usually two pairs of lateral scutellar bristles present, but if one
pair present, then well developed and subequal in size to notopleural bristles; if dorsal preapical tibial
bristle absent from midtibia, then middle fronto-orbital bristle reduced in length and cell bm open
(Procerosoma), or posterodorsal margin of head with wide truncate notch and ratio of ultimate
to penultimate section of vein M less than 2.7 (Clusia); epandrium well developed and usually
as wide as tip of abdomen; distiphallus rodlike (sometimes bent medially), often flanked by one
pair of laterobasal lobes (Figs. 7–8); spermatheca spherical and usually clear; ventral receptacle
lacking subterminal flagellum (Fig. 16). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
CLUSIIDAE (clusiid flies) 78 1045

5. Ratio of ultimate to penultimate section of vein  M greater than 3.0 (often less in species of
Sobarocephala with elaborately patterned wings); forefemur setulose on posterodorsal surface;
presutural intra-alar bristle almost always absent; frons shiny or covered with pale, inconspicuous
setulae; posterior margin of frons flat to gradually concave; segment 7 of female cylindrical and
inconspicuous; surstylus rarely with basal process; distiphallus straight, not longer than phallapodeme
and not articulated medially; SOBAROCEPHALINAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
– Ratio of ultimate to penultimate section of vein  M less than 2.7 (3.0 in Apiochaeta bicolor);
forefemur with well-developed bristles on posterodorsal surface; presutural intra-alar bristle
present; frons usually covered with minute black setulae; posterior margin of frons with shallow
truncate notch behind ocelli; segment 7 of female conical, well sclerotized, and usually more thin,
elongate, and heavily sclerotized than preceding segments; surstylus usually with basal process
on inner face; distiphallus elongate and either bent or articulated medially; CLUSIINAE. . . . . . 8

ann
epand
sbepand scl hypd
phapod

phapod

hypd arm

cerc ej apod

distph
v lb hypd
l lb distph

pregt
sur
bph
b shield distph
6 Sobarocephala pgt
epiph

7 Sobarocephala 8 Sobarocephala

hypd

bph
phapod

hypd

pregt

distph

9 Clusiodes 10 Clusiodes

Figs. 78.6–10. Male terminalia: lateral view (6) of Sobarocephala latifrons (Loew), (Nearctic), hypandrium and subepandrial sclerite shaded;
enlarged lateral view (7) and anterior view (8) of hypandrial complex of S. flava Melander & Argo, (Nearctic); lateral view (9) and anterior
view (10) of hypandrium and associated sclerites of Clusiodes johnsoni Malloch (Nearctic). Figures 6–10 illustrated by O. Lonsdale.
Abbreviations: ann, annulus; bph, basiphallus; b shield distph, basal shield of distiphallus; cerc, cercus; distph, distiphallus; ej apod, ejaculatory
apodeme; epand, epandrium; epiph, epiphallus; hypd, hypandrium; hypd arm, hypandrial arm; l lb distph, lateral lobe of distiphallus; phapod,
phallapodeme; pgt, postgonite; pregt, pregonite; sbepand scl, subepandrial sclerite; sur, surstylus; v lb hypd, ventral lobe of hypandrium.
1046
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

12 Hendelia

13 Craspedochaeta
11 Apiochaeta

spmth
14 Heteromeringia spmth

sbterm flg

v rep v rep

spmth dt spmth dt

15 Heteromeringia 16 Sobarocephala

Figs. 78.11–16. Female terminalia: ventral view of spermathecae, spermathecal ducts, and associated structures of (11) Apiochaeta
connexa Malloch, (South America); (12) Hendelia sp.; (13) Craspedochaeta concinna (Williston), (Lonsdale & Marshall, 2006b, fig. 106);
(14)  Heteromeringia fumipennis Melander  & Argo, (Lonsdale  & Marshall, 2007a, fig.  28); (15)  Heteromeringia czernyi Kertesz,
(Londsdale & Marshall, 2007a, fig. 48); and (16) Sobarocephala sp. Figures 11, 12, and 16 illustrated by O. Lonsdale.
Abbreviations: sbterm flg, subterminal flagellum; spmth, spermatheca; spmth dt, spermathecal duct; v rep, ventral receptacle.
CLUSIIDAE (clusiid flies) 78 1047

6. Vein R1 setulose dorsally; gena usually shiny (rarely pilose); anterior subapical scutellar bristle
minute to absent; cell bm closed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetoclusia Coquillett
– Vein R1 bare dorsally; gena pilose, silvery tomentose or shiny; subapical scutellar bristles various;
cell bm usually open. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

7. Midtibia with dorsal preapical bristle; ocellar and/or postocellar bristles present (usually small);
middle fronto-orbital bristle usually well developed; surstylus sometimes comparable in height to
epandrium (Fig. 6); common. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sobarocephala Czerny
– Midtibia without dorsal preapical bristle; ocellar and postocellar bristles absent; middle fronto-
orbital bristle small or absent; surstylus small and rounded; uncommon . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Procerosoma Lonsdale & Marshall

8. Interfrontal bristle present; postocellar bristle minute; two pairs of dorsocentral bristles; midtibia
without dorsal preapical bristle; surstylus small and triangular; hypandrial bristles absent; ventral
receptacle smooth; North America. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Clusia Haliday]
– Interfrontal bristle absent; postocellar bristle long; three pairs of dorsocentral bristles; midtibia
with dorsal preapical bristle; surstylus longer than wide and usually strikingly elaborated; hypandrial
bristles present; ventral receptacle with numerous transverse wrinkles and two longitudinal
furrows (Fig. 11); Argentina and Chile.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Apiochaeta Czerny]

Synopsis of the fauna Rica, H. kinetrolicros and H. mirabilis, have strikingly modi-
fied heads with long, distinctive genal processes. Hendelia
There are over 200 species of Clusiidae in Central America, species sometimes exhibit male agonistic behavior at lek
the vast majority of which belong to Sobarocephala, with sites, usually on exposed tree trunks, but the behavior of
more than half of these undescribed. Neotropical species is unknown. The species of Hendelia
were revised by Caloren & Marshall (1998) as the Clusiodes
Chaetoclusia Coquillett. This genus is made up of mirabilis group.
13 species in the Neotropical Region that were revised and
keyed by Lonsdale & Marshall (2006a). Five of these species Heteromeringia Czerny. There are 80 described species
are known from Central America. Most species of Chaeto- of this genus worldwide, most of which occur in the Ori-
clusia are uncommon, with restricted Central American or ental and Australian Regions. Eighteen species are found in
Caribbean distributions, but C. bakeri Coquillett (character- the New World, two of which are predominantly or entirely
ized by a densely plumose arista) is regularly collected from Nearctic in distribution. Most of the 11 Central American
southern Mexico to southern Brazil. These small, brightly Heteromeringia are predominantly black, although sever-
colored flies superficially resemble species of Sobarocephala, al species have a yellow thorax with brown markings.
but they are readily characterized by setulae along most of Heteromeringia species are often associated with relatively
R1 dorsally. open areas such as tree falls and pastures, and many spe-
cies have been collected using dung, mushrooms, and car-
Craspedochaeta Czerny. This genus of 49 described spe- rion. In Australia, one species has been observed moving its
cies is most diverse throughout the Oriental and Australian wings in a fashion similar to that seen in the Sepsidae and
Regions. Thirty-one species occur in the Neotropical Region, Platystomatidae (McAlpine, 1960), and in the Nearctic Re-
12 of which are found in Central America. These species gion, males of H. nitida Johnson have been observed while
were revised and keyed by Lonsdale  & Marshall (2006b), engaged in lekking behavior (Lonsdale & Marshall, 2007a).
who also included a discussion of lekking in one species. The New World fauna was revised and keyed by Lonsdale &
Most New World Craspedochaeta are South American in dis- Marshall (2007a), who also redefined the genus (Lonsdale &
tribution, but several extend northward into Central America, Marshall, 2008b), subsequently synonymizing the Oriental
and a few species are restricted to Central America and the Tranomeringia Sasakawa.
Caribbean. These generally small, dark flies are often found
on small dung baits, especially on fallen trees or branches. Procerosoma Lonsdale  & Marshall. Two species of
Procerosoma are known from Mexico (Chiapas) and south-
Hendelia Czerny. The 44 species in this genus (as ern Brazil. These species closely resemble Sobarocephala,
redefined by Lonsdale  & Marshall, 2007b) are primar- but they can be distinguished by a reduction or absence of
ily pantropical and Australian in distribution, although the the cephalic, thoracic, and dorsal preapical tibial bristles,
type species, H. beckeri Czerny, is found throughout the in addition to being comparatively large, slender, and elon-
Palearctic Region. Males of the two species found in Costa gate. Males of P.  alini have elongate genal processes that
1048
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

are presumably used in male agonistic behavior. The two tree trunks, but also around damp, mossy areas near water,
known species were differentiated by Lonsdale & Marshall usually on the underside of leaves or on small dung baits.
(2006a). Most species of Sobarocephala are small and yellow with a
sparsely plumose or pubescent arista, but many species have
Sobarocephala Czerny. This is a predominantly Central a densely plumose arista and/or are large and elaborately pat-
and South American genus, with a number of species found terned, particularly on the head, thorax, and forelegs. The
throughout North America and the Old World excluding species of Sobarocephala with densely plumose aristae were
Europe. Twenty-eight of the 76 described world species are keyed by Soós (1963, 1964) as the “plumata group”, but this
currently known from Central America (16 recorded from key included only 18 of the 59 species now recognized as
Costa Rica), although dozens of species and locality records part of this species group and includes one species (S. lanei
remain undescribed. It is likely that the total size of this ge- Curran) that belongs to another group. Furthermore, there
nus will ultimately exceed 200  species. In North America, are dozens of undescribed Sobarocephala in several other
adults are often found on fallen tree trunks, but in the Neo- species groups that cannot be identified with currently pub-
tropical Region individuals have been observed not only on lished keys.

Literature cited

Caloren, D.C. & S.A. Marshall. 1998. A revision of the New World Lonsdale, O.  & S.A. Marshall. 2008c. Revision of the temperate
species of Clusiodes Coquillett (Diptera: Clusiidae). Studia dip- South American genus Apiochaeta Czerny, 1903, including Al-
terologica 5: 261–321. loclusia Hendel, 1917 new synonym (Diptera: Clusiidae). Zoot-
Frey, R. 1960. Studien über indoaustralische Clusiiden (Dipt.) nebst axa 1944: 1–33.
Katalog der Clusiiden. Commentationes Biologicae 22: 1–31. Malloch, J.R. 1918. A revision of the dipterous family Clusiodidae
Hendel, F. 1936. Ergebnisse einer zoologischen Sammelreise nach (Heteroneuridae). Proceedings of the Entomological Society of
Brasilien, insbesondere in das Amazonasgebiet, ausgeführt von Washington 20: 2–8.
Dr. H. Zerny. Annalen des Naturhistorischen Museums in Wien Marshall, S.A. 2000. Agonistic behaviour and generic synonymy in
47: 61–106. Australian Clusiidae (Diptera). Studia dipterologica 7: 3–9.
Hennig, W. 1938. Beiträge zur Kenntnis der Clusiiden und ihres McAlpine, D.K. 1960. A review of the Australian species of Clusii-
Kopulationsapparates (Dipt. Acalypt.). Encyclopédie Ento- dae (Diptera: Acalyptrata). Records of the Australian Museum 25:
mologique 9: 121–138. 63–94.
Hennig, W. 1965. Die Acalyptratae des baltischen Bernsteins und McAlpine, D.K. 1976. Spiral vibrissae in some clusiid flies (Diptera:
ihre Bedeutung für die Erforschung der phylogenetischen En- Schizophora). Australian Entomological Magazine 3: 75–78.
twicklung dieser Dipteren-Gruppe. Stuttgarter Beiträge zur Melander, A.L.  & N.G. Argo. 1924. Revision of the two-winged
Naturkunde 145: 1–215. flies of the family Clusiidae. Proceedings of the United States
Hennig, W. 1969. Neue Übersicht über die aus dem Baltischen National Museum 64: 1–54.
Bernstein bekannten Acalyptratae (Diptera: Cyclorrhapha). Pitkin, B.R.  & N.L. Evenhuis. 1989. Family Clusiidae, pp.  534–
Stuttgarter Beiträge zur Naturkunde 209: 1–42. 536. In Evenhuis, N.L. (editor). Catalog of the Diptera of the
Lonsdale, O. & S.A. Marshall. 2006a. Redefinition of the Clusii- Australasian and Oceanian Regions. Bishop Museum Press &
nae and Clusiodinae, description of the new subfamily Sobaro- E.J. Brill, Honolulu & Leiden, 1155 pp.
cephalinae, revision of the genus Chaetoclusia and a description Roháček, J. 1995. Clusiidae (Diptera) of the Czech and Slovak Re-
of Procerosoma gen. n. (Diptera: Clusiidae). European Journal publics: Faunistics and notes on biology and behaviour. Casopis
of Entomology 103: 163–182. Slezského Muzea Opava (A) 44: 123–140.
Lonsdale, O. & S.A. Marshall. 2006b. Revision of the New World Sasakawa, M. 1977. Family Clusiidae, pp. 234–239. In Delfina-
Craspedochaeta Czerny. Zootaxa 1291: 1–101. do, M.D. & D.E. Hardy (editors). A catalog of the Diptera of
Lonsdale, O. & S.A. Marshall. 2007a. Revision of the New World the Oriental Region, Volume III. University Press of Hawaii,
Heteromeringia (Diptera: Clusiidae: Clusiodinae). Beiträge zur Honolulu, x + 854 pp.
Entomologie 57: 37–80. Soós, Á. 1962. A review of the species of Czernyola Bezzi, 1907
Lonsdale, O. & S.A. Marshall. 2007b. Redefinition of the gen- (Diptera: Clusiidae). Acta Zoologica 8: 449–457.
era Clusiodes and Hendelia (Diptera: Clusiidae: Clusiodi- Soós, Á. 1963. Identification key to the species of the “plumata-
nae), with a review of Clusiodes. Studia dipterologica 14: group” of the genus Sobarocephala Czerny (Diptera: Clusii-
117–159. dae). Acta Zoolica Hungarica 9: 391–396.
Lonsdale, O. & S.A. Marshall. 2008a. Synonymy within Clusia and Soós, Á. 1964. New Sobarocephala — species from the “plumata-
description of the new genus Melanoclusia (Diptera: Clusiidae: group” (Diptera: Clusiidae). Annales Historico-Naturales Mu-
Clusiinae). Annals of the Entomological Society of America sei Nationalis Hungarici 56: 449–455.
101: 327–330. Soós, Á. 1987. Clusiidae, pp. 853–857. MND, Volume 2.
Lonsdale, O.  & S.A. Marshall. 2008b. The Clusiidae (Diptera: Steyskal, G.C. 1965. Family Clusiidae, pp. 805–807. In Stone, A.,
Schizophora) of Fiji, with redefinition of Heteromeringia Cz- C.W. Sabrosky, W.W. Wirth, R.H. Foote, & J.R. Coulson (edi-
erny and synonymy of Tranomeringia Sasakawa, pp.  3–14. tors). A catalog of the Diptera of America north of Mexico.
In Evenhuis, N.L. & D.J. Bickel (editors). Fiji Arthropods XI. United States Department of Agriculture, Agriculture Handbook
Bishop Museum Occasional Papers 98. 276, Washington, D.C., iv + 1696 pp.
ODINIIDAE (odiniid flies) 79
Stephen D. Gaimari

Fig. 79.1. Male of Odinia betulae Sabrosky, (Nearctic, MND, fig. 72.1).

Diagnosis with posterior marginal setae. Legs (Fig.  1) stout; femora

often enlarged (especially hind femur of males); often with
Small to medium-sized (body length 2.5–6.0  mm), tibial bands; preapical dorsal tibial setae present on one or
compact-bodied, strongly bristled flies (Fig.  1). Body pru- more legs. Wing often with patterns, from simple darkened
inose silvery to dull gray and variously mottled or marked crossveins (e.g., Fig.  1) to complexly pictured or mottled
with brownish or yellowish, or rarely solid brown pruinose. membrane (e.g., Fig. 3); C extending to R4+5 or to M1, with
Head higher than long; postocellar setae divergent (absent in subcostal break only; Sc incomplete. Abdomen relatively
Helgreelia and Shewellia), usually well developed; ocellar short and broad, usually patterned with gray and/or brown
plate normal (Fig. 1) to unusually prominent (Fig. 2); ocellar pruinescence.
setae well developed, proclinate; with three or more pairs of Eggs and first two larval instars unknown. Mature larvae
fronto-orbital setae, posterior two reclinate (rarely one, with (Fig. 4) typical cyclorrhaphans; body essentially bare except
second missing), anterior one to five (usually one) inclinate; for small fringe of hairs anterior to oral opening (Fig.  6),
frons and face in line or sloping (Traginopinae) to distinctly and paired creeping welts anteroventrally on abdominal
angulate (Odiniinae); lunule bare, broadly exposed, some- segments 2–7; each welt consists of pad of large spines di-
times with dorsomedial black pruinose spot; gena broad, rected anteriorly and posteriorly, surrounded by similarly
often with upturned genal seta (Fig.  2); vibrissa strong; directed smaller spines. Caudal segment bears paired fleshy
antennal first flagellomere about as high as long, rounded lobes (expansion likely aids locomotion; Lewis, 1979) near
at apex; arista bare to plumose. Scutum (Fig.  1) with one bilobed perianal pad. Anterior spiracles placed dorsolater-
presutural and three to four postsutural dorsocentral setae; ally (not so dorsal as in the related Agromyzidae), each with
one prescutellar acrostichal seta; prescutellum usually pres- 4–12 papillae arranged in marginal rosette around expanded
ent; scutellar disc setulose or bare; anepisternum bare or apex on short stalk (Fig. 5). Posterior spiracles small, each

1049
1050
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

with three diverging oval openings distally on tubular exten- Of the families with three or more fronto-orbital setae,
sion set on dome-shaped protuberance; paired protuberanc- only a few are even superficially similar to odiniids, and few
es separated distally but fused basally. Cephalopharyngeal specifically have two posterior reclinate fronto-orbital setae
skeleton (Fig.  6) similar to many other acalyptrate larvae, and one (to several) anterior inclinate fronto-orbital setae.
with separate hypopharyngeal and tentopharyngeal sclerites, Most agromyzids lack preapical dorsal tibial setae, have
long parastomal bar, separate labial sclerite, and dental scler- slender legs, only rarely have patterned wings, and some-
ite; mandible simple, curved; ventral cornu larger than dorsal times have a more elongated, tapered abdomen (in females
cornu. Puparium barrel-shaped, tapered both anteriorly and distinctly specialized for piercing plant tissue with a sclero-
posteriorly. tized oviscape). Of particular note is the genus Paraneossos

i vt s
rc frorb s poc s

o vt s
R1

R2+3

cup R4+5
gn s dm
vb M1
sbvb s

A1+CuA2 CuA1

3 Traginops
2 Traginops

4 Neoalticomerus

d corn

pastm b

v corn

den scl
md hyphar scl
5 Neoalticomerus 6 Neoalticomerus

Figs. 79.2–6. Head, wing, and larval structures: anterodorsal view of head (2) and dorsal view of wing (3) of Traginops irroratus Coquillett,
(Nearctic, MND, figs. 72.2, 72.3, as T. irrorata); lateral view of third instar larva (4), anterior spiracle (5), and head and cephalopharyngeal
skeleton of third instar larva (6) of Neoalticomerus seamansi Shewell, (Nearctic, MND, figs. 72.7–9).
Abbreviations: d corn, dorsal cornu; den scl, dental sclerite; gn s, genal seta; hyphar scl, hypopharyngeal sclerite; i vt s, inner vertical seta;
md, mandible; o vt s, outer vertical seta; pastm b, parastomal bar; poc s, postocellar seta; rc frorb s, reclinate fronto-orbital setae; sbvb s,
subvibrissal seta; vb, vibrissa; v corn, ventral cornu.
ODINIIDAE (odiniid flies) 79 1051

Wheeler (Heleomyzidae), which is superficially similar to then ate the beetle frass, crawling towards the beetle’s pu-
Odinia, with three fronto-orbital setae, identical head shape, pal chamber, where they then attacked the pupa externally
overall body form and coloration; however, the species of and bored internally, also displaying cannibalism. Also noted
this genus are small (1–2 mm), the anterior fronto-orbital seta was the fact that the flies apparently did not need to devour
is reclinate, the postocellar setae are convergent to parallel, the beetle pupa to fully develop and occasionally did not
and the costal vein is weakly spinose. Other families, such even attack the pupa.
as Clusiidae, Anthomyzidae, and Acartophthalmidae are also As interesting exceptions, several species are known as
distinctly more slender-bodied. Clusiids sometimes have the associates, and possibly predators, of lepidopteran caterpil-
same typical fronto-orbital setal arrangement as odiniids lars in enclosed spaces such as tunnels and galls. Traginops
(two reclinate posterior, one inclinate anterior), but the setae purpurops Steyskal was reared from a species of Robinia
are much thinner and more delicate, as is the case for all infested with a tortricid moth species in the genus Ecdytol-
setae on the head and thorax; the gena is narrow; the an- opha (Steyskal, 1963); Odinia meijerei Collin was reared
tennal pedicel often has a characteristic triangular extension from larvae of the cossid moth species Zeuzera pyrina Lin-
laterally; first flagellomere often bears the arista terminally naeus (Campadelli, 1995) and from galls of the aegerid moth
to subterminally; Sc is complete; and the abdomen is slender. species Paranthrene tabaniformis Rottemburg (Bescho-
In acartophthalmids, the fronto-orbital setae are all laterorec- vski  & Georgiev, 1993); and Odinia boletina (Zetterstedt)
linate, and the scutum lacks a presutural dorsocentral seta. In was reared from tunnels of gelechiid moth larvae in Juglans
anthomyzids, the fronto-orbital setae are all reclinate with twigs (Kato, 1952), and adults of this species are apparently
the anterior seta weak, the postocellar setae are convergent if attracted to bracket fungi (Chandler, 1978), as are the adults
present, and the scutum lacks a presutural dorsocentral seta. of Odinia rossi MacGowan  & Rotheray (MacGowan  &
Other families that can superficially resemble odiniids in Rotheray, 2002). Krivosheina (1981) reported that sev-
coloration and habitus, such as lauxaniids and chamaemyiids eral Palearctic species attack larvae of other wood-infest-
(some odiniids even share the characteristic prescutellum, ing Diptera. One Asian species, Turanodinia coccidarum
which is an unusual state), are easily differentiated by their Stackelberg, was reared from egg masses of the mealybug
lack of vibrissae, unbroken C, convergent postocellar setae, Pseudococcus comstocki Kuwana. Although appearing odd
and having at most two fronto-orbital setae. as a life history for an odiniid, this is perfectly reasonable
for a variety of reasons. First, the females of this mealybug
typically move under the tree bark to become sessile egg
Biology masses, thickly gathered together into waxy clusters. The
microcosm associated with the egg masses of this mealy-
Few studies describe the biology of species of Odiniidae, bug is quite diverse, with many different organisms involved
and none discuss their biology in the Neotropical Region. (from mites to predaceous fly larvae, to saprophagous moth
Adults are closely associated with trees, especially those in- larvae, many beetles, etc.), and the dense honeydew secreted
fested with other insects, rotting, with fungi, or exuding sap. by the mealybugs has intensive fungal growth. Ultimately,
Some species of Odinia have been collected in McPhail traps the odiniids could be associated with this sub-bark micro-
baited with vinegar (Carles-Tolrá, 1996) or various yeast habitat in a variety of ways, from consuming fungi, to in-
mixtures (Sabrosky, 1959; Steyskal, 1963). sect frass, to predation. Krivosheina & Krivosheina (1996)
Although few species have known biologies, those with also reported this species, and other species of Turanodinia
known larvae are associated with trees such as those that at- Stackelberg, living within congestions of sap and in galler-
tract the adults. Usually, they are associated with galleries ies of wood-destroying insects. One Afrotropical species,
of wood-boring beetles, with several life history possibili- Afrodinia (nr. medleri Cogan) was reared from “gum in tun-
ties from saprophagy to predation. Krivosheina (1981), Fer- nels” of Acacia karroo, without a known predacious habit
rar (1987), and Kirk-Spriggs & Barraclough (2009) discuss (Kirk-Spriggs & Barraclough, 2009).
the known biologies (e.g., logs of Betula trees infested with
Mature larvae or puparia have been described for 13 spe-
platypodine weevils and a species of the buprestid genus
cies (none of which are Neotropical) in five genera (only one
Agrilus (Sabrosky, 1959); rotten Ulmus trees infested with
of which has Neotropical species). These include the Holarc-
species of the oedemerid beetle genus Ischnomera (Col-
lin, 1952) or of the scolytine weevil genus Scolytus (Lewis, tic species Odinia xanthocera (Yang, 1984), the Palearctic
1979); under bark of dead Abies in tunnels of the scolytine species Odinia czernyi Collin and Odinia foliata Krivosheina
weevil genus Pitiogenes (Zubkov  & Kovalev, 1975); and (Krivosheina, 1981), Odinia meijerei (Lewis, 1979), Odinia
Populus and Salix trees infested with species of the weevil boletina (Kato, 1952), Odinia ornata (Zetterstedt) (Zub-
genus Cryptorhynchus (Shewell, 1960) or the tenebrionid kov & Kovalev, 1975), Turanodinia stackelbergi Krivoshei-
genus Scolobates (Sabrosky, 1959)). Yang (1984) and Pulk- na & Krivosheina (Krivosheina & Krivosheina, 1996), and
kinen & Yang (1984) provided details on the biology of the Turanodinia tisciae Papp (Papp, 1995), an Afrotropical spe-
Holarctic species Odinia xanthocera Collin, finding that ovi- cies of Afrodinia (nr. medleri) (Kirk-Spriggs & Barraclough,
position of 15–20 eggs occurred at the entrance to a tunnel of 2009), and the Nearctic species Neoalticomerus seamansi
the cerambycid species Saperda populnea Linnaeus. Larvae Shewell and Traginops irroratus Coquillett (Shewell, 1960).
1052
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

McAlpine (1987), Ferrar (1987), Smith (1989), and Foote the frons more or less in line or sloping with the face; (ii) the
(1991) also provide useful figures and discussion of imma- position of the ocellar triangle, with the odiniines having
ture stages. A detailed scanning electron microscope study of the posterior ocelli in line with the inner vertical setae and
the puparium of an Afrotropical species of Afrodinia Cogan the traginopines having the ocellar triangle more anteriorly
was done by Kirk-Spriggs & Barraclough (2009), along with placed.
a review of the species of the family with known, or partially
known, immature stages.
Identification

Classification Only a few key resources are necessary for identifying

the described species of Odiniidae, although many species
Current evidence (McAlpine, 1989) suggests that Odinii- remain undescribed. Hennig (1969) provided a key to the
dae forms a monophyletic group within the suprafamily genera of the world, with descriptions or re-diagnoses of
Agromyzoinea along with Agromyzidae and Fergusonini- several genera and their species, although without keys to
dae, within the superfamily Opomyzoidea. This suprafamily species. Cogan (1975) provided keys and some descrip-
is supported by several synapomorphies relative to the per- tions for the known Afrotropical odiniid fauna. McAlpine
ceived acalyptrate ground plan (McAlpine, 1989), including (1987) provided a key to the genera present in the Nearctic
the more dorsally located anterior spiracles in the larvae. Region, as did Papp (1998) for the Palearctic Region. The
There are two monophyletic lineages, Agromyzidae + Fer- Palearctic Region has received some attention to the spe-
gusoninidae and Odiniidae. Odiniids appear to be the most cies level, particularly for the genera Odinia (Collin, 1952;
generalized member of the Agromyzoinea, with larval Krivosheina, 1981) and Turanodinia (Krivosheina & Kri-
mouthparts simpler, the anterior spiracles more dorsolater-
vosheina, 1996). Sabrosky (1959) revised the genus Odinia
ally located, and the larval habitats within galleries of wood-
for the New World, including descriptions of taxa and keys
boring insects rather than as primary tissue invaders of living
to species (although numerous species remain undescribed);
plants. Odiniids were originally considered as a subfamily of
Agromyzidae, until Hendel (1922–1923, 1928) recognized only two additional Neotropical species of Odinia have
them as a separate family. Hennig (1958) recognized odini- been described since. The most important contribution to
ids and agromyzids as sister taxa, but it was not until Spencer Neotropical odiniid studies is that of Prado (1973), which
(1969) that specific character evidence was used to support includes reassessments of described taxa, descriptions of
this relationship, citing the strong aedeagal apodeme (phal- new taxa, keys to species, and adequate drawings of diag-
lapodeme), modified hypandrium, the form of the surstyli, nostic characteristics. Gaimari (2007) provides a key to the
the whole general genitalic arrangement, and the presence currently described New World genera, and Gaimari & Ma-
of scutellar hairs in several odiniid genera and the more this (in press) provide a key to all described genera and a
primitive agromyzid genera such as Selachops Wahlberg. world catalog for the family. Filho et al. (2009) concatenated
The monophyly of the family is supported by the apparent several published keys to present a unified key to the
absence of an ejaculatory apodeme, the presence of several described Neotropical species.
setae on the katepisternum, and the presence of a preapical Odiniids are most commonly found in forested ecosys-
dorsal seta on one or more tibiae (McAlpine, 1989). tems, where their biological activities occur. Specimens
The family Odiniidae is divided into two subfamilies, are most commonly collected in Malaise traps or associ-
Odiniinae and Traginopinae. Several characters support this ated with tree trunks or beetle galleries, and certain species
division: (i) the shape of the head, with the odiniines having a are collected in McPhail traps baited with vinegar or yeast
distinctly angulate frons to face, while the traginopines have mixtures (e.g., Torula yeast).

Key to the genera of Odiniidae of the New World

1. Posterior ocelli in line with inner vertical setae (Fig.  1); ocelli arranged in equilateral triangle
or with distance between posterior pair greater than distance between either posterior ocellus
and anterior ocellus; frons flat to slightly concave; scutellar disc bare; in profile, face distinctly
angulate with frons at antennal insertion; ODINIINAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Posterior ocelli in front of line between inner vertical setae (Fig. 2); ocelli usually arranged in
isosceles triangle with distance between posterior pair conspicuously less than distance between
either posterior ocellus and anterior ocellus; frons usually relatively arched (sometimes
conspicuously tuberculate); scutellar disc sometimes setulose; in profile, face and frons more or
less in line or sloping; TRAGINOPINAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
ODINIIDAE (odiniid flies) 79 1053

2. Postocellar setae reduced, short and thin; with four dorsocentral setae (1+3); anepisternum with
one to two strong setae plus several setulae; dorsal preapical tibial setae well developed; Holarctic
Region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Neoalticomerus Hendel]
– Postocellar setae well developed, subequal to ocellar setae; with five dorsocentral setae (1+4);
anepisternum bare; dorsal preapical tibial setae thin, hairlike. . . . . . . Odinia Robineau-Desvoidy

3. Ocelli on moderately large to large tubercle, well in front of vertex (as in Fig.  2, or more
prominent) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
– Frons lacking enlarged tubercle, but occasionally with ocelli on raised bump little larger than
diameter of anterior ocellus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

4. Arista long plumose; vein M1 strongly curved anteriorly at apex. . . . . . . . Paratraginops Hendel
– Arista pubescent; vein M1 straight (Fig. 3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

5. Propleuron bearing two setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neotraginops Prado

– Propleuron lacking setae; Holarctic, Afrotropical, Oriental Regions . . . . . [Traginops Coquillett]

6. Costa extended to vein M1; genal seta present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

– Costa extended to vein R4+5 or slightly beyond; genal seta absent. . . . . . . . . . . . . . . . . . . . . . . . . 9

7. Scutellum at most bearing two to five setulae along margin. . . . . . . . . . . . . . . Lopesiodinia Prado
– Scutellar disc and margins bearing numerous setulae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

8. Only anterior fronto-orbital seta inclinate (as in Fig. 2) . . . . . . . . . . . . . . . . Schildomyia Malloch

– Several anterior fronto-orbital setae inclinate . . . . . . . . . . . . . . . . . . . . . Neoschildomyia Gaimari

9. Postocellar setae present; posterior part of head straight from dorsal view. . . Pradomyia Gaimari
– Postocellar setae absent; posterior part of head concave from dorsal view. . . . . . . . . . . . . . . . . 10

10. Several anterior fronto-orbital setae inclinate; scutellar disc setulose. . . . . . . . . Shewellia Hennig
– Only anterior fronto-orbital seta inclinate; scutellar disc bare. . . . . . . . . . . . . . Helgreelia Gaimari

Synopsis of the fauna Filho, Esposito & Santos, is known only from the Brazilian
Amazon.
No concise resources are available to readily identify the
Central American members of this family, largely because Lopesiodinia Prado. The two described species, L. alva-
rengai Prado and L.  diversa Prado, are only known from
of the high number of undescribed taxa in the Neotropical
Brazil. At least two new species of this genus are known
Region as a whole, although Prado (1973), Sabrosky (1959),
from Costa Rica.
Gaimari (2007), and Filho et al. (2009) are the most use-
ful publications for the New World fauna. Of the 62 species Neoschildomyia Gaimari. The single described spe-
and three subspecies described worldwide, 26 are Neotropi- cies, N.  fusca Gaimari, is known only from Costa Rica
cal, of which 14 are from Central America plus the islands (Alajuela).
of the Caribbean (nine of the 11 species known from main-
land Central America are in Costa Rica). Despite this, many Neotraginops Prado. The single described species,
new species are recognized, and the Central American fauna N.  clathratus (Hendel), is known from Paraguay through
includes an additional 30 or more new species. Extending Colombia. At least three new species of this genus are known
south into South America and north into the western Nearc- from Costa Rica, one of which is also known from Belize
tic Region are dozens more undescribed species. and one from Panama.

Helgreelia Gaimari. Two of the three described species, Odinia Robineau-Desvoidy. This cosmopolitan genus of
H. albeto Gaimari and H. parkeri Gaimari, are only known 26 described species is represented by 12 species in the New
from Costa Rica (Guanacaste). The other species, H. gaimarii World, of which only one, O. coronata Sabrosky, is known
1054
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

from Central America, recorded from Belize, El Salvador Schildomyia Malloch. Four species of this genus are
(type locality), Guatemala, Panama, and southern Mexico known from Central America. Schildomyia punctifrons Mal-
(Morelos, Chiapas), as well as the southwestern USA, north- loch and S.  vittithorax Malloch were both described from
ern South America, and the West Indies. The remaining New Costa Rica (Alajuela) and are also known from Panama
World species are from the islands of the Caribbean, South (Canal Zone); the former species also occurs in Brazil. The
America, and the Nearctic Region. At least 20 new species latter species was also recorded from Peru (Hennig, 1969),
are known to occur in Costa Rica alone, and many additional but specimens identified as such from Peru and Brazil are
new species are known from throughout Central and South subtly different and may not be conspecific. Schildomyia
America and the western Nearctic Region. reticulata Prado was previously known only from Peru, but
also occurs in Brazil, Panama, Costa Rica (Guanacaste),
Paratraginops Hendel. The only described species Belize, and southern Mexico (Veracruz). The other Central
known from Central America is P.  plaumanni Shewell, American species, S. brevihirta Malloch, is known only from
which occurs in Panama (Canal Zone), but is otherwise Panama and Peru, and S. trinidadensis Hennig is known only
known only from Brazil. The other New World species, from Trinidad. Four other described species are known from
P.  pilicornis Cresson, is known only from Brazil and Brazil, one from Peru, and two from the far eastern Palearctic
Region. At least two new species are known from Costa Rica
Guyana. The remaining species is from Africa and may or
and one from Panama.
may not be congeneric.
Shewellia Hennig. The single described species of this
Pradomyia Gaimari. This genus is represented by the genus, S.  agromyzina Hennig, was previously known only
single species P. hadromera Gaimari, described from Costa from Peru, but also occurs in Panama and Costa Rica. Several
Rica (Guanacaste). new species are known from northern South America.

Literature cited

Beschovski, V.L. & G.T. Georgiev. 1993. Three species of Diptera – Hendel, F. 1922–1923. Die paläarktischen Muscidae Acalyptratae
Acalyptrata (Diptera) dwelling galls of Parathrene tabaniformis Girsch.  = Haplostomata Frey nach ihren Familien und Gat-
Rott. (Lepidoptera, Aegeridae). Acta Zoologica Bulgarica 46: tungen. I. Die Familien. Konowia 1: 145–160 (1922), 253–265
44–49. (1923).
Campadelli, G. 1995. Su alcuni parassitoidi di Zeuzera pyrina L. Hendel, F. 1928. Zweiflügler oder Diptera. II. Allgemeiner Teil. In
(Lep. Cossidae) in Emilia-Romagna. Bollettino dell’Istituto di Dahl, F. (editor). Die Tierwelt Deutschlands. G. Fischer, Jena,
Entomologia “Guido Grandi” del’Università di Bologna 50: 135 pp.
127–131. Hennig, W. 1958. Die Familien der Diptera Schizophora und ihre
Carles-Tolrá, M. 1996. Odinia trifida sp.n., a new odiniid spe- phylogenetischen Verwandtschaftsbeziehungen. Beiträge zur
cies from Spain (Diptera: Odiniidae). Stuttgarter Beiträge zur Entomologie 8: 505–688.
Naturkunde, Serie A (Biologie) 545: 1–3. Hennig, W. 1969. Neue Gattungen und Arten der Acalypteratae.
Chandler, P.J. 1978. Associations with plants. Fungi, pp.  199–211. The Canadian Entomologist 101: 589–633.
In Stubbs, A. & P.J. Chandler (editors). A Dipterist’s Handbook. Kato, S. 1952. Odiniidae of Japan, with descriptions of a new spe-
The Amateur Entomological Society, Hanworth, Middlesex, cies and a new subspecies. Insecta Matsumurana 18: 1–8.
ix + 255 pp. Kirk-Spriggs, A.H. & D.A. Barraclough. 2009. The puparium of an
Cogan, B.H. 1975. New taxa in two families previously unrecorded Afrodinia species (Diptera: Odiniidae) from South Africa, with
from the Ethiopian Region (Diptera: Odiniidae and Diastatidae). a review of the known immature stages of Odiniidae and notes
Annals of the Natal Museum 22: 471–488. on biology and functional morphology. African Entomology 17:
Collin, J.E. 1952. On the European species of the genus Odinia 161–169.
R.-D. (Diptera, Odiniidae). Proceedings of the Royal Entomo- Krivosheina, N.P. 1981. [Systematics and biology of Palaearctic
logical Society of London (Series B) 21: 110–116. species of the family Odiniidae (Diptera)-entomophages and
Ferrar, P. 1987. A guide to the breeding habits and immature stag- xylophilous insects], pp. 130–157. In [Insects of Wood and their
es of Diptera Cyclorrhapha. Entomonograph 8(1). E.J.  Brill  / Entomophages]. Academy of Sciences of the USSR, A.N. Sev-
Scandinavian Science Press, Leiden, Copenhagen, 478 pp. ertsov Institute of Evolutionary Morphology and Ecology of
Filho, F.S.C., M.C. Esposito, & R.C.O. Santos. 2009. A new species Animals, Nauka Publishing, Moscow, 254  pp. [In Russian.]
of Helgreelia Gaimari (Diptera: Odiniidae) from Brazil, with (1979)
a key to the Neotropical species of Odiniidae. Zootaxa 2219: Krivosheina, N.P.  & M. G. Krivosheina. 1996. New data on the
61–68. taxonomy and biology of the flies from the genus Turanodinia
Foote, B.A. 1991. Odiniidae (Opomyzoidea), pp. 818–819. In Stackelberg (Diptera, Odiniidae). International Journal of Dip-
Stehr, F.W. (editor). Immature Insects, Volume 2. Kendall/Hunt terological Research 7: 141–146.
Publishing, Dubuque, 975 pp. Lewis, D.C. 1979. The larva and puparium of Odinia meijerei Collin
Gaimari, S.D. 2007. Three new Neotropical genera of Odiniidae (Dipt., Odiniidae). Entomologist’s Monthly Magazine 114: 233–235.
(Diptera: Acalyptratae). Zootaxa 1143: 1–16. MacGowan, I.  & G.E. Rotheray. 2002. A new species of Odinia
Gaimari, S.D. & W.N. Mathis. In press. World catalog and conspec- (Diptera, Odiniidae) from Scotland. Dipterists Digest (second
tus on the family Odiniidae (Diptera: Schizophora). Myia 12. series) 9: 67–69.
ODINIIDAE (odiniid flies) 79 1055

McAlpine, J.F. 1987. Odiniidae, pp. 863–867. MND, Volume 2. Shewell, G.E. 1960. Notes on the family Odiniidae with a key to the
McAlpine, J.F. 1989. Phylogeny and classification of the Musco- genera and descriptions of new species (Diptera). The Canadian
morpha, pp. 1397–1518. MND, Volume 3. Entomologist 42: 625–633.
Papp, L. 1995. Morphology of Periscelis annulata third instar larva Smith, K.G.V. 1989. An introduction to the immature stages of Brit-
and Turanodinia tisciae larva and puparium (Diptera: Perisce- ish flies. Diptera larvae, with notes on eggs, puparia and pu-
lididae and Odiniidae). Acta Zoologica Academiae Scientiarum pae. Handbooks for the identification of British insects 10(14):
Hungaricae 41: 15–24. 1–280.
Papp, L. 1998. Family Odiniidae, pp. 233–242. In Papp,  L.  & Spencer, K.A. 1969. The Agromyzidae of Canada and Alaska.
B. Darvas (editors). Contributions to a Manual of Palaearctic Memoirs of the Entomological Society of Canada 64: 1–311.
Diptera. Volume 3, Higher Brachycera. Science Herald Publish- Steyskal, G.C. 1963. A second North American species of Tragin-
ers, Budapest, 880 pp. ops Coquillett (Diptera: Odiniidae). Proceedings of the Entomo-
Prado, A.P. do. 1973. Contribuição ao conhecimento da família logical Society of Washington 65: 51–54.
Odiniidae (Diptera, Acalyptratae). Studia Entomologica (N.S.) Yang, Z.-Q. 1984. Notes on the larva and puparium of Odinia xan-
16: 481–510. thocera Collin (Diptera, Odiniidae). Annales Entomologici Fen-
Pulkkinen, M. & Z.-Q. Yang. 1984. The parasitoids and predators
nici 50: 93–94.
of Saperda populnea (Linnaeus) (Coleoptera, Cerambycidae) in
Finland. Annales Entomologici Fennici 50: 7–12. Zubkov, G.A.  & V.G. Kovalev. 1975. [New data on the develop-
Sabrosky, C.W. 1959. Flies of the genus Odinia in the Western ment of flies of the family Odiniidae (Diptera)]. Nauchnye
Hemisphere (Diptera: Odiniidae). Proceedings of the United doklady vysshei shkoly. Biologicheskie nauki 1975: 14–19. [In
States National Museum 109: 223–236. Russian.]
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
AGROMYZIDAE (leaf-mining flies) 80
Stéphanie Boucher

Fig. 80.1. Female of Agromyza albipennis Meigen, (Holarctic, MND, fig. 73.1).

Diagnosis present, forming small cell cup (Fig.  11); A1 not reaching
wing margin. Abdomen tapered, composed of six visible
Minute to medium-sized flies (wing length 0.9–6.5 mm, segments. Females with abdominal segment 7 modified into
usually 2–3 mm). Often yellow and/or black, brown, or gray, large conical nonretractile oviscape (Fig. 12).
few have some metallic greenish, bluish, or coppery color- Some female Tephritidae have an oviscape similar to that
ation (genus Japanagromyza and Melanagromyza). Head in Agromyzidae, but their subcostal vein is abruptly bent for-
(Figs. 1–6) with first flagellomere small, round, or sometimes ward at an angle of 90˚ (not so in Agromyzidae), and they
enlarged, elongate, or with anterodorsal projection or spine frequently have patterned wings (rare in Agromyzidae).
and with well-developed dorsobasal arista. Fronto-orbital Chloropidae usually have much reduced chaetotaxy on the
and ocellar bristles strong; postocellar bristles divergent; head and thorax, and wing vein A1 and cell cup are absent. In
fronto-orbital setulae in one or more rows, usually reclinate Clusiidae, the subcostal vein is more pronounced and com-
(Fig. 2), but sometimes proclinate (Fig. 3), upright or absent; plete, the subcostal break is sometimes absent, and many
vibrissa well developed, some males (genus Ophiomyia) species have a triangular extension on the outer side of the
with vibrissal fasciculus (Fig.  4). Scutum with two to five pedicel. Members of the family Odiniidae have the anepis-
dorsocentral bristles present, but many genera in Phyto- ternum bare (always with some setae in Agromyzidae).
myzinae with three postsutural and one presutural dorsocentral
bristles; acrostichals present (in up to 10  rows) or absent.
Costal break present near the end of R1, humeral break Biology
absent; Sc incomplete distally (Fig. 9), or reaching C as lin-
ear fold, independently of R1 (Fig. 10), or Sc complete and Larvae of all species of Agromyzidae feed on living
fusing with R1 before reaching costa (Fig. 11); CuA2 and A1 plant tissues. Although many species are leafminers, many

1057
1058
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

other parts of plants such as roots, stems, seeds, flower head, Central America. For example, Liriomyza sativae Blanchard
trunk, and twigs are also attacked. Most known agromyzids is a major pest of cultivated plants in South America and
are host-specific and will feed on plants from a single ge- southern United States that has recently been recorded in
nus (monophagous) or from different genera included in the Costa Rica (Spencer, 1983). This species attacks a wide
same family (oligophagous). Only a few species are truly variety of plants but prefers the families Cucurbitaceae
polyphagous and feed on a number of unrelated families, and (cantaloupe, cucumber, squash), Fabaceae (various beans),
these include some of the most serious agromyzid pests in and Solanaceae (pepper, tomato, eggplant, and potato)

i vt s
poc s
orb s

o vt s

fr s
frorb sta

vb

2 Liriomyza 3 Phytomyza 4 Ophiomyia

a b

6 Cerodontha 7 Cerodontha

5 Cerodontha

8 Cerodontha

Figs. 80.2–8. Heads, lunules, and antenna: dorsolateral view of head of (2) Liriomyza septentrionalis Sehgal, (Nearctic, MND, fig. 73.2);
(3) Phytomyza pedicularidis Spencer, (Nearctic, MND, fig. 73.3); (4) Ophiomyia maura (Meigen), (Holarctic, MND, fig. 73.4); anterior view
of head and/or lunule of (5) Cerodontha (Dizygomyza) luctuosa (Meigen), (Holarctic and Neotropical, MND, fig. 73.5); (6a) C. (Poemyza)
pygmaea (Meigen), (Holarctic, MND, fig. 73.8); (6b) C. (Poemyza) muscina (Meigen), (Holarctic); (7) C. (Butomomyza) sp.; lateral view of
antenna of (8) C. (Cerodontha) dorsalis (Loew), (Nearctic and Neotropical, MND, fig. 73.6). Figures 6b and 7 illustrated by S. Boucher.
Abbreviations: fr s, frontal bristle; frorb sta, fronto-orbital setula; i vt s, inner vertical bristle; orb s, orbital bristle; o vt s, outer vertical bristle;
poc s, postocellar bristle; vb, vibrissa.
AGROMYZIDAE (leaf-mining flies) 80 1059

(Spencer, 1973a). Most other species that are potential pests of the most important agromyzid pests in Venezuela (Spen-
in Central America are also in the genus Liriomyza. These cer, 1973b), has also been recorded in Costa Rica. This is a
include the polyphagous and widespread species Liromyza highly polyphagous species, known to feed on nine different
trifolii (Burgess) that has been recorded from Costa Rica plant families, but more commonly on beets, peas, beans,
(Spencer, 1983) and that attacks many different crops, such and some ornamental plants.
as onions, beans, and tomatoes, as well as ornamentals like Most of the damage to plants is caused by the larvae, but
chrysanthemums. Liriomyza huidobrensis (Blanchard), one the female can also cause damage during oviposition. Females

bm-cu

bm-cu r-m
bm-cu

9 Pseudonapomyza 10 Aulagromyza
Sc R1

R2+3

bm br
cup
dm R4+5

M1
A1+CuA2
CuA1

11 Melanagromyza 12 Cerodontha

13 Agromyza 14 Phytomyza

15 Melanagromyza

Figs. 80.9–15. Wings, oviscape, and larval features: dorsal view of wings of (9) Pseudonapomyza lacteipennis (Malloch), (Holarctic, MND,
fig.  73.12); (10)  Aulagromyza nitida (Malloch), (Nearctic, MND, fig.  73.11); (11)  Melanagromyza laetifica Spencer, (Nearctic, MND,
fig. 73.9); lateral view of oviscape of (12) Cerodontha dorsalis (Loew), (Nearctic and Neotropical); lateral view of larval cephalopharyngeal
skeleton of (13) Agromyza albipennis Meigen, (Holarctic, MND, fig. 73.27); (14) Phytomyza chelonei Spencer, (Nearctic, MND, fig. 73.23);
dorsal view of larval posterior spiracles of (15) Melanagromyza sp., (Nearctic, MND, fig. 73.30). Figure 12 illustrated by S. Boucher.
1060
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

insert their well-sclerotized ovipositor into the plant tissues The Phytomyzinae are characterized by having the subcostal
to deposit their eggs. Some females also use these punctures vein incomplete (Fig. 9) or running straight to the costa as a
as feeding sites before laying eggs, sucking plant sap through linear fold (Fig. 10) and the larval cephalopharyngeal skel-
the punctures. After a few days, the larva emerges and starts eton with only one upper arm (Fig. 14). However, there are
feeding on the plant tissues, causing a mine or sometimes a few exceptions. For example, in the genus Phytobia (Phy-
a gall. The mines can be of different forms: linear, serpen- tomyzinae), the course of Sc is often most similar to that of
tine (Fig.  27), blotchlike (Fig.  28), or irregular (Fig.  29). Agromyzinae, but this does not contradict the monophyly of
In some cases the first instar larva forms a linear mine and the subfamilies (Spencer, 1987; Dempewolf, 2001).
later instars form a blotch mine. These feeding patterns are Worldwide there are over 2800 species of Agromyzidae
species-specific and often helpful in the identification of described, with the majority occurring in temperate areas of
Agromyzidae. Most species feed in the upper surface of the the Northern Hemisphere, but the family is also well repre-
leaf, in the palisade parenchyma, or less often immediately sented in the tropics. There are still many undescribed spe-
beneath the leaf epidermis. Others feed on the lower surface cies, and Spencer & Steyskal (1986) estimated that only half
of the leaf, in the spongy parenchyma. The part of the leaf the actual number of species are known to date.
where the mine is formed is also important. Some species
feed along the margin of the leaf, while others stay along the
midrib or along a strong lateral vein as in the pest species
Identification
Liriomyza huidobrensis (Spencer & Steyskal, 1986). Other
than the mine itself, the arrangement of the frass inside the Fourteen agromyzid genera are known from Central Amer-
mine is also characteristic. The frass may be in the form of ica. In two of these (Amauromyza, Haplopeodes), species are
small pellets or short strips, and these may be widely sepa- present but as yet undescribed. Although all 14 genera are
rated, interconnected, deposited alternately on each side of also present in the Nearctic Region, keys to Nearctic Agro-
the mine, or in some cases the frass is excreted only at the myzidae (e.g., Spencer, 1987) will only work for some of the
very end of the mine. The majority of agromyzid species pu- Central American species. Existing keys to Neotropical agro-
pate on the ground, but for most seed feeders, stem borers, myzids (e.g., Spencer, 1963; Spencer & Stegmaier, 1973) are
and a few leafminers, pupation takes place within the plant, out of date, as many new species have been described since
at the feeding site. these keys were published. Most of these species were de-
Agromyzid species are usually naturally controlled by a scribed from Argentina (Sasakawa, 1992a; Valladares, 1981,
large complex of wasp parasites. The use of insecticides has 1992, 1998a), Bolivia (Sasakawa, 1992d), Brazil (Sasakawa,
removed many of these natural enemies, however, and has 1992d; Esposito & Prado, 1993a, b; Esposito, 1994), Chile
caused a great increase in the damage caused by agromyzid (Spencer, 1982; Sasakawa, 1994), Colombia (Spencer, 1984;
larvae to a wide range of cultivated plants (Spencer, 1973a, Sasakawa, 1992b; Sanabria de Arévalo, 1993a,  b, 1994),
1983). This phenomenon has been observed in L.  sativae, Costa Rica (Spencer, 1983; Woodley & Janzen, 1995), Ec-
which has caused increased damage since the widespread uador (Sasakawa, 1992c), El Salvador and Guatemala (Sa-
use of chemical insecticides. sakawa, 2005), Peru (Sasakawa, 1992d), Venezuela (Spencer
1973b; Sasakawa, 1992d), Guadeloupe (Martinez, 1994; Eti-
enne & Martinez, 2003; Spencer et al., 1992), and other Ca-
Classification ribbean islands (Etienne & Martinez, 1996; Zlobin, 1996a).
Martinez & Etienne (2002) published a checklist of Neotro-
The relationship of Agromyzidae to other Diptera fami- pical Agromyzidae. Few publications deal specifically with
lies is unclear, and there is still no widespread agreement the Central American fauna. There is no doubt that many of
on the sister group of the family. Hennig (1958) suggested the species described from South America or the Caribbean
that Agromyzidae were most closely related to Odiniidae (or are more widespread than currently known, and many spe-
possibly Clusiidae), and Spencer (1969) pointed out geni- cies (described and undescribed) await discovery in Central
talic similarities between Odiniidae and Agromyzidae. The America.
Odiniidae was the sister group of Agromyzidae in the phy- Many agromyzid genera can be identified based on ex-
logenetic analysis of Dempewolf (2001). However, Griffiths ternal characters, such as body coloration and size, wing
(1972) concluded that Agromyzidae and Clusiidae were sis- venation, head, and body chaetotaxy. However, other more
ter groups, based on a detailed study of the male postabdo- specific characters are sometimes necessary to differenti-
men. More recently, McAlpine (1989) placed Agromyzidae, ate certain genera. For example, the presence of a stridu-
Odiniidae, and the Australian family Fergusoninidae togeth- latory mechanism is a specific character used to separate
er in his suprafamily Agromyzoinea. male Liriomyza species from similar species in other genera
Agromyzidae are generally divided into two subfami- (Tschirnhaus, 1971). Although this character is considered
lies: Agromyzinae and Phytomyzinae. The Agromyzi- a synapomorphy for Liriomyza, it is sometimes secondarily
nae are characterized by having the subcostal vein joining lost, in which case the correct identification must be based
with R1 before reaching the costa (Fig.  11) and the larval on genitalic structures (Zlobin, 1996b, 1999). When present,
cephalopharyngeal skeleton with two upper arms (Fig. 13). the stridulatory mechanism consists of a line of vertical scales
AGROMYZIDAE (leaf-mining flies) 80 1061

syntg 1+2

hlt
syntg 1+2
hlt

synst 1+2

strd mch
strd mch 16 Liriomyza
st 1 st 2
17 Agromyza

pgt
strd mch

epand

18 Agromyza sbepand
scl

cerc

19 Cerodontha

sur

ph
epand

cerc

sur

cerc
20 Liriomyza
21 Japanagromyza

Figs. 80.16–21. Stridulatory mechanism and male genitalic features: lateral view of stridulatory mechanism on abdominal pleura of a male
(16) Liriomyza sp.; below lower margin of syntergite 1+2 in (17) Agromyza albipennis Meigen, (Holarctic, MND, fig. 73.13); on inner surface
of hind femur in (18) A. albipennis, (Holarctic, MND, fig. 73.14); ventral view of epandrium with subepandrial sclerites of (19) Cerodon-
tha dorsalis (Loew), (Nearctic and Neotropical); ventral view of epandrium and surstylus of (20) Liriomyza sp.; lateral view of male genitalia
of (21) Japanagromyza phaseoli Spencer, (Neotropical). Figures 16, 19–21 illustrated by S. Boucher.
Abbreviations: cerc, cercus; epand, epandrium; hlt,  halter; pgt,  postgonite; ph,  phallus; sbepand  scl, subepandrial sclerite; sur,  surstylus;
st, sternite; strd mch, stridulatory mechanism; synst, synsternite; syntg, syntergite.
1062
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

sur pgt
epand

epand

hypd
cerc

cerc sur

22 Phytobia 23 Nemorimyza

sur

epand

24 Calycomyza 25 Phytoliriomyza 26 Phytoliriomyza

27 Liriomyza 28 Aulagromyza 29 Phytomyza

Figs. 80.22–29. Male genitalic features and leaf mines: ventral view of epandrium and surstylus of (22) Phytobia sp.; lateral view of male
genitalia of (23) Nemorimyza posticata (Meigen), (Nearctic and Neotropical); ventral view of epandrium and surstylus of (24) Calycomyza
meridiana (Hendel), (Neotropical); left side of epandrium and surstylus of (25) Phytoliriomyza leechi Spencer, (Nearctic); left side of
epandrium and surstylus of (26) Phytoliriomyza sp.; leaf mines of (27) Liriomyza arctii Spencer, (Nearctic, MND, fig. 73.33); (28) Aulagromyza
orbitalis (Melander), (Nearctic, MND, fig. 73.34); and (29) Phytomyza gregaria (Frick), (Nearctic, MND, fig. 73.35). Figures 22–26
illustrated by S. Boucher.
Abbreviations: cerc, cercus; epand, epandrium; hypd, hypandrium; pgt, postgonite; sur, surstylus.
AGROMYZIDAE (leaf-mining flies) 80 1063

along the center of the abdominal pleura (Fig. 16) and a ridge importance in the identification of these genera. Larval
on the inner surface of the hind femur. In some cases the line morphology can also be useful in identifying Agromyzi-
of scales can only be seen at higher magnification (400×). dae at the generic level. For example, larvae of the genus
A similar stridulatory mechanism is also present in males Melanagromyza are easily distinguished by their posterior
of a few species of Cerodontha (Tschirnhaus, 1971). In the spiracles having numerous bulbs surrounding a central horn
genus Agromyza (Figs.  17, 18), a stridulatory mechanism (Fig. 15) or at least the scar of it. Frick (1952) published
is also present, but it is present in both males and females, a generic key for North American species based on larval
and the line of scales is just below the lower margin of syn- morphology, but more information on the larvae of Neo-
tergite 1+2. Members of the genus Phytobia are sometimes tropical species will be needed before a key can be made.
externally similar to Agromyza, but lack the stridulatory Other information such as the larval biology; the shape,
mechanism. form, and color variation of the puparium; characteristics
Male genitalic characters are often important in the de- of leaf mines; and identity of the host plants can be used in
termination of agromyzid genera. For example, the sym- the identification of Agromyzidae.
metrical phallus of Melanagromyza (asymmetrical in Like most other groups of small Diptera, Agromyzidae
Ophiomyia), the enlarged cerci of many species of Japa- should be collected in 70% alcohol and subsequently dried
nagromyza (Fig.  21), the L-shaped subepandrial sclerites using a critical-point dryer or chemicals like hexamethyl-
(also known as “bacilliform sclerites”, “processus longi”, disilazane (HMDS). Spencer & Steyskal (1986) suggested
and “Langfortsatz” in the literature) of Cerodontha species collecting agromyzids dry, but this often results in shriv-
(Fig. 19), and the patch of spines at the hind corner of the eled specimens, with structures such as the stridulatory
epandrium in Calycomyza (Fig. 24) are all of outstanding mechanism difficult to see.

Key to the genera of Agromyzidae of Central America

1. Subcostal vein developed throughout its length and joined with vein  R1 before reaching costa
(Fig. 11); distal margin of R1 often clearly expanded and sinuous near junction (Fig. 11); vein R4+5
ending closer to wing tip than does M1, or both veins equidistant from wing tip; AGROMYZINAE
(and some Phytobia species in PHYTOMYZINAE). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Subcostal vein either incomplete distally (Fig. 9) or extending to costa as thin vein or fold
independently from R1 (Fig. 10); distal margin of R1 straight at junction (Figs. 9, 10), at most
slightly expanded; usually vein M1 (very rarely R4+5) ending closer to wing tip than does R4+5, or
both veins equidistant from wing tip; PHYTOMYZINAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

2. Stridulatory mechanism present on lateral margin of fused first and second abdominal tergites
and on inner surface of hind femur in male and female (Figs. 17, 18); halter usually white (black
in A. fusca also with infuscated wings); usually three or more dorsocentral bristles (anterior pair
sometimes weak or absent, e.g. A.  megaepistoma Sasakawa with only two  pairs); prescutellar
bristles well developed; gena usually angular, deepest at rear; body without metallic sheen;
surstylus with numerous spines and fused with epandrium. . . . . . . . . . . . . . . . . . Agromyza Fallén
– Stridulatory mechanism absent; other characters various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3. Normally at least four dorsocentral bristles (usually 3+1 or 4+1), but anterior pairs sometimes
smaller; halter white; calyptral fringe dark; body brightly colored with yellow or completely dark
without metallic sheen; lunule often conspicuously broad and silvery; wings sometimes pictured
or slightly infuscated; distal margin of R1 usually not expanded and sinuous near junction; wing
length sometimes over 3.0  mm; surstylus broad, lobate, separated from epandrium by distinct
suture (Fig. 22); hypandrium usually broad and rounded, sometimes V-shaped; postgonite well
developed (Phytomyzinae). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phytobia Lioy, in part
– Usually only two postsutural dorsocentral bristles; if three, anterior pair usually short and close
to second pair; if three or more well-developed pairs (few South American Melanagromyza), then
halter brown. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4. Knob of halter partially or completely brown or black. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

– Knob of halter completely white or yellow. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
1064
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

5. Prescutellar bristles present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

– Prescutellar bristles absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

6. Wing tinged with brown or foretibia with one posterolateral bristle; abdomen shining black or
with metallic reflection; eye bare. . . . . . . . . . . . . . . . . . . . . . . . Japanagromyza Sasakawa, in part
Note: J. jamaicensis Spencer and J. nebulifera Sasakawa; males unknown.

– Wing clear and foretibial bristle absent; abdomen with metallic reflection, eye with at least short
scattered hairs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Melanagromyza Hendel, in part

7. Antennae often divided by facial keel; vibrissal angle often acute (Fig. 4); male often with distinct
vibrissal fasciculus consisting of partly fused bristles (Fig. 4); uniformly black species, without
prominent metallic sheen; upper margin of clypeus truncate, sometimes pointed, not uniformly
rounded; phallus usually asymmetrical in ventral view; basiphallus usually not U-shaped or circular,
mostly elongate with one side often reduced in length . . . . . . . . . Ophiomyia Braschnikov, in part
– Facial keel absent, at most antennal bases separated by flat ridge; male with vibrissa normal;
scutum and/or abdomen mostly with some metallic bluish, greenish, or coppery color; upper
margin of clypeus usually more or less uniformly rounded; eye sometimes with scattered or dense
pilosity in upper third; phallus usually symmetrical in ventral view; basiphallus mostly U-shaped
(the U is open towards the distiphallus) or circular in ventral view . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Melanagromyza Hendel, in part
Note: Japanagromyza orbitalis (Frost) also keys here; see discussion under Japanagromyza.

8. Body color usually dark, often with metallic sheen; calyptral fringe almost always white (except
J. polygoni Spencer); only two postsutural dorsocentral bristles, rarely with short third pair also
present; prescutellar bristles usually present, sometimes absent; phallus in form of elongated
tubule (simple or divided); epandrium usually with narrow, elongate prolongation (the surstylus)
at anteroventral corner or lower (Fig. 21); cercus usually large and often with tubercles or spinules
on inner side (Fig. 21); ejaculatory apodeme usually with narrow blade . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Japanagromyza Sasakawa, in part
– Body without metallic sheen; usually brightly colored with yellow especially on legs and
abdomen (except Phytobia unica Spencer with brown body); calyptral fringe dark or white; two
well-developed postsutural dorsocentral bristles present, sometimes with small third pair close
to second pair; prescutellar bristles well developed (except P. unica); phallus usually ending in
enlarged process or paired tubules; surstylus covered with numerous strong spines and almost
completely separated from epandrium except for narrow connection on posteroventral margin;
cercus without tubercles or spinules on inner side; ejaculatory apodeme with large blade and large
base (usually bowl-shaped) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

9. Calyptral fringe dark or white; phallus usually in form of simple tubule with enlarged process at
apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Japanagromyza Sasakawa, in part
– Calyptral fringe dark; phallus ending in paired tubules of various length and width . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phytobia Lioy, in part

10. Fronto-orbital setulae erect, reclinate or absent; if proclinate, halter completely white and costa
extending to vein M1, or first flagellomere with distinct anterodorsal spine or projection . . . . . 11
– Fronto-orbital setulae distinctly proclinate; halter usually darkened or costa extends to vein R4+5;
first flagellomere without distinct anterodorsal spine or projection. . . . . . . . . . . . . . . . . . . . . . . 30

11. Crossvein dm-cu absent and only one upper orbital bristle present, or costa extending only to
vein R4+5. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
– Crossvein dm-cu usually present; if absent, then two upper orbital bristles present and costa
extending to vein M1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
AGROMYZIDAE (leaf-mining flies) 80 1065

12. Costa extending only to vein R4+5; crossvein r-m close to or in line with crossvein bm-cu (Fig. 9);
only three postsutural dorsocentral bristles present, with anterior pair weaker; first flagellomere
with acute angle at upper tip in New World species; head and body black; halter yellow; wing
length over 1.5 mm; phallus well sclerotized. . . . . . . . . . . . . . . . . . . . . .Pseudonapomyza Hendel
– Costa extending to vein M1; crossvein r-m well separated from crossvein bm-cu; three postsutural
dorsocentral bristles and one presutural present; first flagellomere rounded, normal; body usually
partly yellow, if completely black, then halter also at least partially darkened; some small species,
with wing length sometimes less than 1.5 mm; phallus greatly reduced and little differentiated. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Haplopeodes Steyskal

13. Knob of halter completely white and R4+5 ending closer to wing tip than M1, or ends of both veins
equidistant from wing tip; foretibia sometimes with one posterolateral bristle and midtibia often
with two or three strong posterolateral bristles near midpoint; lunule sometimes silvery (espe-
cially in dorsal view); frons dark, sometimes reddish anteriorly; male genitalia without large L-
shaped subepandrial sclerites. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
– Knob of halter partially or completely black or if completely white, M1 usually ending closer to
wing tip than R4+5, if ends of both veins equidistant from wing tip then lunule large, not silvery
and male genitalia with large L-shaped subepandrial sclerites (Fig. 19); other characters various.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

14. Body completely brown or with contrasting yellow coloration (usually on scutum, scutellum,
pleura, and abdomen); calyptral fringe dark; lunule often conspicuously broad, less than
semicircular and silvery; normally at least four dorsocentral bristles (usually 3+1 or 4+1), with
anterior ones sometimes smaller (if only two pairs, sometimes with an extra short one: discussion
under Phytobia); prescutellar bristles present or absent; foretibia rarely with posterolateral bristle
near midpoint; wing sometimes pictured or partially infuscated; surstylus usually broad, lobate,
separated from epandrium by distinct suture (Fig. 22) . . . . . . . . . . . . . . . . . Phytobia Lioy, in part
– Body mostly brown except male abdomen (rarely females) with anterior and/or posterior tergites
yellow (sometimes only the epandrium); calyptral fringe white; lunule not especially large,
higher than semicircular and usually silvery; three well-developed postsutural dorsocentral
bristles (sometimes with an additional smaller pair close to suture); prescutellar bristles well
developed; foretibia usually with posterolateral bristle near midpoint; wings clear; surstylus com-
pletely separated from epandrium with dorsal part projecting deeply inside epandrium (Fig. 23).
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nemorimyza Frey, in part

15. Stridulatory mechanism on the membrane connecting the abdominal tergites and sternites usually
present in males (Fig. 16); knob of halter completely white or yellow; frons and scutellum usually
bright yellow (frons and/or scutellum may be dark in a few species); three postsutural and one
presutural dorsocentral bristles (if 2+1, body partly yellow); lunule not especially high or large
(Fig. 2); surstylus usually present and with one or two (rarely three) spines and sometimes few
weak hairs (Fig. 20); epandrium often with one or two spines (sometimes more) at posteroventral
margin (Fig. 20), or conspicuously sclerotized on inner margin. . . . . . . . . . . . . . . Liriomyza Mik
– Stridulatory mechanism absent in males; other characters various. . . . . . . . . . . . . . . . . . . . . . . 16

16. Knob of halter at least partially darkened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

– Knob of halter completely white or yellow. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

17. Frons strongly projecting above and in front of eye in profile, forming broad ring below eye;
gena deep, one-third to one-quarter eye height, extended at rear; body, including knob of halter,
completely brownish-black; three postsutural and one presutural dorsocentral bristles; three lower
orbital and two upper orbital bristles; calyptral fringe black; foretibial bristle absent; wing clear;
distiphallus surrounded by spinular membrane; sperm pump with enlarged bowl-shaped base;
Nearctic Region species A. abnormalis Hendel. . . . . . . . . . . . . . . . . Amauromyza Hendel, in part
– Not exactly fitting above description. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
1066
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

18. Usually one upper and three lower orbital bristles present; wing with cells r1 and r2+3 conspicuously
infuscated; lunule large, with silvery pubescence (especially in dorsal view); base of antennae
widely separated; body mostly brown with foreknee (femur–tibia joint) and tarsi pale yellow;
knob of halter completely brown; calyptral fringe dark; male genitalia with large L-shaped
subepandrial sclerites (Fig. 19) . . . . . . . . . . . . . . . . . . . Cerodontha (Dizygomyza) Hendel, in part
– Not exactly fitting above description. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

19. Lunule often with silvery or grayish pubescence; R4+5 usually ending closer to wing tip than M1, or
ends of both veins equidistant from wing tip; body usually completely black; usually with three
postsutural and one presutural dorsocentral bristles; posterolateral bristle often present on foretibia
near midpoint; epandrium and surstylus without black spines; surstylus completely separated
from epandrium with its dorsal part projecting deeply inside epandrium (Fig. 23) . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nemorimyza Frey, in part
– Lunule brown without silvery or grayish pubescence; M1 usually ending closer to wing tip than
R4+5; body often partly yellow; if body entirely dark then either presutural dorsocentral bristles
absent or two postsutural and one presutural bristles present; posterolateral bristle on foretibia
absent; epandrium and surstylus with comb of black spines (Figs. 24–26). . . . . . . . . . . . . . . . . 20

20. Presutural dorsocentral bristle absent or if present, only two postsutural dorsocentral bristles
present and body completely brown; gena narrow, usually one-seventh or less eye height . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calycomyza Hendel, in part
– Usually three postsutural and one presutural dorsocentral bristles present; if 2+1 dorsocentral
bristles, then anterior bristle close to suture and body partly yellow; gena deep, usually approximately
one-quarter eye height. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phytoliriomyza Hendel, in part

21. Presutural dorsocentral bristle absent (or present but weak in few South American species; well
developed in C. illustris Spencer, Venezuela, but identity unconfirmed without males); scutellum
dark brown or black; first flagellomere brown or black (yellow in C. meridiana (Hendel)); frons
usually yellow or pale yellowish-brown, but may be darker (e.g., C. devia Spencer and C. obscura
Spencer); lunule not especially high or large; epandrium with conspicuous patch of spines at hind
corner (Fig. 24); distiphallus well sclerotized. . . . . . . . . . . . . . . . . . . Calycomyza Hendel, in part
– Presutural dorsocentral bristle usually present; other characters various . . . . . . . . . . . . . . . . . . 22
22. Distiphallus surrounded by spinular membrane; sperm pump with enlarged bowl-shaped base;
lunule not especially high or large; head and body black in Neotropical species, various in North
American species; usually three or four lower and one (sometimes two) upper orbital bristles; gena
often deep and extended posteriorly; first flagellomere sometimes slightly angulate, two pairs of
scutellar bristles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amauromyza Hendel, in part
– Distiphallus not surrounded by spinular membrane; sperm pump without enlarged bowl-shaped
base; other characters various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

23. First flagellomere with distinct anterodorsal projection or spine (Fig. 8); scutellum with only one
pair of bristles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
– First flagellomere without spine, at most slightly angulate; scutellum with two pairs of bristles,
basal scutellar bristles sometimes reduced. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

24. Body at least partially bright yellow . . . . . . . . . . . . . . . . . . . . Cerodontha (Cerodontha) Rondani

– Body completely black or brown (except halter); North America, Jamaica . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Cerodontha (Xenophytomyza) Frey)]
AGROMYZIDAE (leaf-mining flies) 80 1067

25. Lunule conspicuously higher than a semicircle (Figs.  6a,  b, 7) or lunule conspicuously broad
(Fig. 5); scutellum dark; male subepandrial sclerites dark, L-shaped (Fig. 19); Cerodontha
Rondani (in part) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
– Lunule usually normal, not especially high or large (Figs. 2–4), but if conspicuously high or large,
scutellum partially yellow and epandrium and/or surstylus with conspicuous comb of bristles
(some Phytoliriomyza). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

26. Lunule conspicuously higher than a semicircle (Figs. 6a, b, 7). . . . . . . . . . . . . . . . . . . . . . . . . . 27

– Lunule conspicuously broad, nearly semicircular (Fig. 5). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28

27. Lunule narrow, sometimes triangular (Figs.  6a,  b); surstylus usually without patches of strong
bristles or spines (but short spines sometimes present on lower margin of epandrium). . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cerodontha (Poemyza) Hendel
– Lunule higher than semicircle but still broad (Fig. 7); surstylus and sometimes lower margin of
epandrium with patches of strong bristles or spines . . . . Cerodontha (Butomomyza) Nowakowski

28. Head usually with frons, lunule, face, or gena bright yellow; ocellar triangle extended anteriorly
beyond level of upper orbital bristle; first flagellomere not enlarged in males; surstylus usually
without patches of strong bristles (but short stout spines sometimes present on lower margin of
epandrium). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cerodontha (Icteromyza) Hendel
– Head usually mostly brown or partially yellowish-brown (usually not bright yellow); ocellar triangle
shorter, not extended beyond level of upper orbital bristle; first flagellomere often enlarged in
males; surstylus and sometimes lower margin of epandrium with patches of strong bristles or
spines. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cerodontha (Dizygomyza) Hendel, in part

29. Frons and scutellum dark blackish-brown; first flagellomere often bright yellow (sometimes
completely black); phallus usually ending in short oval or rounded section, sometimes divided
medially; surstylus and epandrium without comb of fingerlike projections and rarely with long
bristles; epandrium with or without spines; North and South America. . . . . [Galiomyza Spencer]
– Scutellum usually grayish, sometimes with faint yellow undertone, or mostly yellow; color of
first flagellomere various; phallus ending in elongate paired tubules; surstylus and/or epandrium
with characteristic comb ovf long bristles or fingerlike projections; hind corner of epandrium
sometimes with short stout spines (Figs. 25, 26). . . . . . . . . . . . . . . Phytoliriomyza Hendel, in part

30. Costa extending to vein M1; crossvein dm-cu usually present (absent in P. conjunctimontis (Frick));
knob of halter usually darkened, yellowish-brown . . . . . . . . . . . . . Phytoliriomyza Hendel, in part
– Costa extending only to vein R4+5; crossvein dm-cu usually absent; halter white. . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phytomyza Fallén

Synopsis of the fauna (A.  animata Spencer, A.  fusca Spencer, A.  megaepistoma
Sasakawa, and A. venezolana Spencer) occurring in Central
Approximately 85 species of Agromyzidae have been America. Agromyza fusca was previously known from Brazil
recorded from Central America, mostly from Costa Rica. and Dominica (Spencer & Stegmaier, 1973), but this species,
Many more species are known from neighboring areas (e.g., distinctive in having brown halteres and infuscated wings, is
over 50 species known from Colombia but not yet recorded also present in Argentina and in Costa Rica. Most species whose
in Central America), and the diversity of Central American feeding habits are known are leafminers, and a wide range
Agromyzidae is expected to be much higher. In terms of of host plants have been recorded for this genus. The male
known diversity, Liriomyza, Melanagromyza, and Calyco- genitalia of the three Costa Rican species suggest that they
myza are the dominant genera in Central America. belong to a group of species that feed on grasses (Poaceae).

Agromyza Fallén. There are approximately 195 species Amauromyza Hendel. There are close to 60 species of
of Agromyza, with the highest diversity in temperate areas of this genus worldwide, and most of them are found in temper-
the Holarctic Region. Only 11 described species have been ate areas of the Holarctic Region. A few Neotropical species
recorded in the Neotropical Region, including four species previously included in Amauromyza (Annimyzella) have been
1068
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

transferred to the genus Nemorimyza (Zlobin 1996b). The this genus resemble those in the genus Liriomyza or Phyto-
genus Amauromyza now includes only two species known to liriomyza, but the absence of crossvein dm-cu, the presence
occur in the Neotropical Region, including one species from of only one orbital seta, and the absence of the stridulatory
Cuba not formally described (Spencer & Stegmaier 1973). mechanism (present in males of Liriomyza) confirms the
The other Neotropical species, A. boliviensis Sasakawa, is so identity of Haplopeodes. Most species feed on host plants
far known only from Bolivia. The Nearctic species A. abnor- in the family Solanaceae, but other hosts are in the families
malis (Malloch) has been recorded from Tlaxcala, Mexico Amaranthaceae, Chenopodiaceae, and Portulacaceae. Stey-
(Bautista-Martinez et al. 1997). If this species continues to skal (1980) provided a key to the 11 Haplopeodes species
expand its distribution southwards, it is likely to occur in known at that time, and an extension to this key was pro-
Central America in the near future. The genus is apparently vided by Valladares (1998b). One species known from Chile
present in Central America, but the only specimen known to is not included in these keys, but its description can be found
date is a female whose identity cannot be confirmed. The lar- in Sasakawa (1994).
vae of many Amauromyza species form large blotch mines,
while others, such as A. abnormalis, are stem borers. Japanagromyza Sasakawa. This is a small genus with
approximately 60  species worldwide. The genus is well
Calycomyza Hendel. There are over 80 species of Caly- represented in the Neotropical Region with approximately
comyza known, and they are essentially restricted to the New 25 species now recorded. Eleven species are known to occur
World. In the Neotropical Region, approximately 60  spe- in Central America, including J. orbitalis (Frost) (Panama)
cies have been recorded. Surprisingly only 11  species are with uncertain generic status (Spencer & Stegmaier, 1973).
known in Central America (eight in Costa Rica), compared This species was previously included in Melanagromyza
with Venezuela where over 20 species have been recorded. (Frick, 1952), but based on the shape of the phallus it was
Many new species of Neotropical Calycomyza have been later transferred to Japanagromyza by Spencer (1963), al-
described since the early 1980s (Esposito, 1994; Esposi- though the black halteres, absence of prescutellar bristles,
to  & Prado, 1993b; Etienne  & Martinez, 1996; Sasakawa, and the shape of the surstyli are not typical of the genus.
1992b, d, 1994; Spencer et al., 1992; Valladares, 1981, 1992; Other Neotropical Japanagromyza species (J.  jamaicensis
Zlobin, 1996a), and many additional species will probably Spencer and J. nebulifera Sasakawa) with black halteres, but
be discovered in Central America in the future. Calycomyza well-developed prescutellar bristles, are known from female
larvae are leafminers (usually forming large blotch mines) specimens only. Other atypical Japanagromyza, namely
and most are host-specific. J.  maculata (Spencer) and J.  spadix (Spencer) (couplet 9
Cerodontha Rondani. There are over 260 species of in the key), were previously included in the genus Gerato-
Cerodontha known, now classified into seven subgenera (Bu- myza Spencer (in Spencer & Stegmaier, 1973: p.140). These
tomomyza Nowakowski, Cerodontha, Dizygomyza Hendel, brightly colored Japanagromyza differ from typical Japana-
Icteromyza Hendel, Phytagromyza Hendel, Poemyza Hendel, gromyza not only by the partly yellow color but also by the
and Xenophytomyza Frey). To date approximately 30 species presence of a very small third anterior dorsocentral bristle
have been recorded in the Neotropical Region, in all sub- and by a few male genitalic characters, including the surstyli,
genera except Phytagromyza, which is exclusively Holarc- the cercus, and the ejaculatory apodeme that are all atypical
tic. The previously Holarctic subgenus Xenophytomyza has of the genus. The genus Geratomyza was synonymized with Ja-
recently been recorded from Jamaica (Boucher, 2003). Only panagromyza by Spencer (1984), but further studies will be
one species, C. (Cerodontha) dorsalis (Loew), was previ- necessary to confirm the identity of the species included in
ously recorded in Central America. The presence of C. (Po- the Japanagromyza maculata group. An undescribed Gua-
emyza) muscina (Meigen) in Costa Rica (Spencer, 1983) was temalan species of this previously Caribbean species group
presumably a mistake (Martinez & Etienne, 2002). At least has been identified, and additional Central American species
four additional species are now known in Central America, are to be expected. Species of Japanagromyza are well repre-
with most of them in the subgenus Dizygomyza (Boucher, sented on host plants in the family Fabaceae (Leguminosae)
2005). Cerodontha species are leafminers exclusively on (Spencer, 1990). Among Central American species, J. phase-
Monocotyledoneae. Host plants are known in the families oli Spencer is noteworthy as it is a pest of cultivated beans
Cyperaceae, Iridaceae, Juncaceae, and Poaceae. The larvae (Phaseolus  spp.). This species has been recorded in Costa
of C.  dorsalis feed on many genera of Poaceae, including Rica, Venezuela, Guadeloupe, and Peru (where it was also
many cereal crops. This species is considered to be of some reported on Brassica oleracea, but this might be an isolated
economic importance (Spencer, 1983). occurrence; see Spencer, 1983) and is probably widespread
in the Neotropical Region. Like most other members of the
Haplopeodes Steyskal. This small genus contains 15 genus, the larvae of J.  phaseoli cause conspicuous blotch
species that are restricted to the New World. In the Neotro- mines on the leaves of the host plant and pupate externally.
pical Region, 13  species have been recorded, mostly from A few species are known to cause galls, including one Costa
Argentina and Brazil. The genus is also known from some Rican species that has been reared from galls on Lonchocar-
undescribed female specimens from Costa Rica. Members of pus (Fabaceae) (Boucher & Hanson, 2006).
AGROMYZIDAE (leaf-mining flies) 80 1069

Liriomyza Mik. This is one of the largest genera with Nemorimyza maculosa was originally described from New
over 360 species described worldwide. Although Liriomyza York, USA, but is now widespread in the Neotropical Re-
is primarily a north temperate genus, it is still well repre- gion. The larvae of this economically important species cause
sented in the Neotropical Region with approximately 85 large blotch mines on leaves of many genera of Asteraceae,
species recorded. A total of 16 species of Liriomyza have and this species has caused a total loss of a small lettuce
been recorded from Central America. In addition, L. cortesi field in Venezuela (Spencer, 1973a). The Holarctic species
Spencer previously known from Chile (Spencer, 1982) has Nemorimyza posticata is widespread in North America as far
now been recorded in Costa Rica. Many other Liriomyza south as Florida and is now also common in Costa Rica. This
species are known from neighboring areas (e.g., Spencer, species forms large blotch mines on Asteraceae, which is the
1973b, 1984; Sanabria de Arevalo, 1993a), and it is ex- only host plant family recorded for the genus.
pected that the number of species from Central America
will be much higher. Within the family Agromyzidae, the Ophiomyia Braschnikov. There are over 175 species
genus Liriomyza has the widest host range with 76 plant in this genus, including approximately 35 species recorded
families recorded (Spencer, 1990). Most Liriomyza spe- from the Neotropical Region. Thirteen species have been re-
cies are leafminers, and the majority are monophagous or corded in Central America. Larvae are mainly external stem
oligophagous. A few Liriomyza species are polyphagous, miners (as opposed to similar Melanagromyza species, which
and these include some of the major Liriomyza pest species are internal stem borers), but many species are also leafmin-
recorded in Costa Rica, such as L. huidobrensis, L. sativae, ers. For the stem-mining species, pupation takes place in the
and L. trifolii. stem with anterior spiracles projecting through the epidermis
(as opposed to Melanagromyza species, which pupate deep
Melanagromyza Hendel. This primarily tropical genus is in the stem). One of the major agromyzid pests worldwide,
among the largest of the genera of Agromyzidae, with over O. phaseoli (Tryon) occurs throughout the Old World tropics
300 described species. Most of the species are internal stem where it attacks a wide variety of beans. Ophiomyia phaseoli
borers, pupating inside the stem, but a few are internal feed- has been recorded in Hawaii but has not so far been recorded
ers in the roots, seeds, or flower heads. In the Neotropical in the Neotropical Region. There are so far no Ophiomyia
Region, approximately 90 species have been recorded, and species of major economic importance known in Central
there are probably many more. Among the Neotropical mate- America, and it is important to monitor O. phaseoli in order
rial studied by Spencer & Stegmaier (1973), at least 10 new to keep it out of Central and South America, where it could
species from Costa Rica were discovered but not described cause serious damage (Spencer, 1973a). Other species are
because of the poor condition of the specimens (Spencer, considered beneficial, such as O. lantanae (Froggatt). This
1983). There are 11 described species recorded in Central species is native to Mexico and Central America, but was de-
America, including M. rosales Woodley, a leafminer of Bro- liberately introduced in Hawaii, Australia, India, South Af-
melia pinguin (Bromeliaceae). This is the only agromyzid rica, and elsewhere for the biological control of Lantana, a
known to feed on Bromeliaceae and only the second known major weed pest (Spencer 1973a). Unlike most species in the
species of Melanagromyza (along with M.  cyrtorchid- genus, O. lantanae feeds in the seed heads of its food plant.
is Spencer, East Africa) that feeds exclusively on leaves
(Woodley  & Janzen, 1995). Many Melanagromyza species Phytobia Lioy. This genus contains approximately 65 de-
are pests of cultivated plants. In Central America, there is scribed species, distributed throughout the world. In the Neo-
only one potential pest species, M. caerula (Malloch), which tropical Region, close to 20 species have been recorded, but
feeds on the seeds of various Ipomoea species, including the diversity is apparently much higher. Eight species were
sweet potato (Spencer, 1973a). Other pest species such as previously known in Central America, including two Costa
M. tomaterae Steyskal, M. caucensis Steyskal (both feeding Rican species known only from females and not formally de-
in tomato stems), and M. phaseolivora Spencer (in pods of scribed (Spencer & Stegmaier, 1973). An additional species,
beans) occur in neighboring areas (Spencer, 1984; Bautista- P.  unica Spencer, previously known only from Jamaica, is
Martinez  & Morales-Galvan, 2000) and could possibly be also present in Costa Rica. This brown species has only two
present in Central America. dorsocentral bristles and no prescutellar bristles, which is a
unique combination in the genus; P. unica also has distinc-
Nemorimyza Frey. This genus contains five described tive male genitalia with a long projection on the anteroventral
species (four of which were previously in the Amauromyza margin of the epandrium, narrow and elongated hypandrium
subgenus Annimyzella; see Zlobin, 1996b), all present in the (usually broader in most Phytobia spp.), surstyli covered
Neotropical Region. Only two species, N.  maculosa (Mal- with short spines and narrowly connected to the posteroven-
loch) and N. posticata (Meigen), were previously recorded in tral margin of the epandrium. The presence of only two well-
Central America, but N. ranchograndensis (Spencer) known developed dorsocentral bristles, the shape of the hypandrium,
from Venezuela, Colombia, and Chile (Spencer, 1973b; Sa- and surstyli are also found in some colorful Phytobia species
sakawa, 1992b, 1994) has now been recorded from Costa (P.  mentula Sasakawa, P.  pipinna Sasakawa, and P.  guate-
Rica. Two additional undescribed species occur in Central malensis Sasakawa), which together with P. unica form what
America, and at least five more occur in South America. I have called the Phytobia unica group (see couplet 9 in the
1070
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

key). The wing venation of the P. unica group is very similar including five species previously included in the genus Chro-
to that of Agromyzinae because the subcostal vein joins R1 matomyia Hardy, which was recently synonymyzed with
before reaching the costa (also occurs in other Phytobia), and Phytomyza (Winkler et al., 2009). Only three species are
the distal margin of R1 is usually clearly expanded near the known from Central America (Costa Rica). Among these,
junction. These characters make the P.  unica group hardly one has been recorded from an empty leaf mine on Myr-
differentiated from the Japanagromyza maculata group (see rhidendron donnell-smithii (Apiaceae), and another (previ-
discussion under Japanagromyza). The larvae of all known ously included in Chromatomyia) is known from puparia in
species of Phytobia are cambium borers in twigs or trunks leaf mines on Brachiaria mutica (Poaceae), but no adults of
of trees. Some of the largest species in the family belong these species were found (Spencer, 1983). The other species,
to this genus and have a wingspan reaching 6.5 mm, while Phytomyza loewii Hendel, is a Nearctic species originally
other species such as P. rabelloi Spencer (Brazil, Ecuador, described from Washington, D.C., that may have expanded
Costa Rica) and P. kallima (Frost) (Panama) are conspicuous its distribution southward during the Pleistocene (Spencer &
because of their pictured wings. Stegmaier, 1973). Other Neotropical species may also be
present in Central America, including the widespread Hol-
Phytoliriomyza Hendel. There are approximately 70 arctic species Phytomyza syngenesiae (Hardy) that has re-
described species worldwide in this genus. In the Neotropical cently been discovered in Colombia. This species is a pest of
Region, approximately 30 species are known. One of these spe- chrysanthemums in greenhouses, and it is a potential threat
cies, not yet formally described, was previously known from to the flower industry in Colombia (Spencer, 1984). Phy-
Argentina (Spencer, 1990) and possibly from Guadeloupe tomyza rufipes Meigen, a pest species on Brassica  spp., is
(Spencer et al., 1992). In Argentina, adults were reared from an another species not yet recorded in Central America but is
unidentified fern, while in Guadeloupe mines containing dead present in Colombia, where it has probably been introduced
larvae were found on the tree fern Cyathea arborea (L.). Only with cabbages coming from Europe (Spencer, 1984). Most
six species have been recorded from Central America, but the Phytomyza species are leafminers, but some bore inside
diversity is apparently much higher. Phytoliriomyza costaricen- seeds or stems. A total of 75 families have been recorded as
sis Spencer is so far known only from Costa Rica but is prob- host plants (Spencer, 1990).
ably of Nearctic origin, based on the shape of the male genitalia
(Spencer & Stegmaier, 1973). There are no pest species known Pseudonapomyza Hendel. This genus contains approxi-
in the genus, but host plants are known for only a few leaf-min- mately 60 species worldwide. In the New World, this group
ing and stem-mining species. Phytoliriomyza is the dominant is not well represented, with only four species (Boucher,
genus on ferns and the only one feeding on liverworts (class 2004), including one, P.  asiatica, recently recorded from
Hepaticae) (Spencer, 1990). Guadeloupe and from Costa Rica (Etienne  & Martinez,
2003; Boucher, 2004). Members of this genus are known to
Phytomyza Fallén. This is the largest agromyzid genus feed on Poaceae and Acanthaceae. In the New World, only
with over 560 species, reaching its highest diversity in the the Poaceae feeders have been recorded and can easily be
Holarctic Region. Few species are known from the tropics. distinguished from the Acanthaceae feeders by the angulate
In the Neotropical Region, there are only 17 species recorded, first flagellomere.

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AGROMYZIDAE (leaf-mining flies) 80 1071

Etienne, J. & M. Martinez. 2003. Les Agromyzidae de Guadeloupe: Spencer, K.A. 1982. Agromyzidae (Diptera) in Chile. Stuttgarter
espèces nouvelles et notes additionnelles (Diptera). Nouvelle Beiträge zur Naturkunde, Serie A (Biologie) 357: 1–55.
Revue d’Entomologie 19: 249–272. Spencer, K.A. 1983. Leaf mining Agromyzidae (Diptera) in Costa
Frick, K.E. 1952. A generic revision of the family Agromyzidae Rica. Revista de Biología Tropical 31: 41–67.
(Diptera) with a catalogue of New World species. University of Spencer, K.A. 1984. The Agromyzidae (Diptera) of Colombia, in-
California Publications in Entomology 8: 339–452. cluding a new species attacking potato in Bolivia. Revista Co-
Griffiths, G.C.D. 1972. The phylogenetic classification of Diptera Cyclo- lombiana de Entomología 10: 3–33.
rrhapha, with special reference to the structure of the male postabdo- Spencer, K.A. 1987. Agromyzidae, pp. 869–879. MND, Volume 2.
men. Series Entomologica 8. Dr. W. Junk, The Hague, 340 pp. Spencer, K.A. 1990. Host specialization in the World Agromyzidae
Hennig, W. 1958. Die Familien der Diptera Schizophora und ihre (Diptera). Series Entomologica 45. Kluwer Academic Publish-
phylogenetischen Verwandtschaftbeziehungen. Beiträge zur ers, Dordrecht, 440 pp.
Entomologie 8: 505–688. Spencer K.A.  & C.E. Stegmaier. 1973. Agromyzidae of Florida
Martinez, M. 1994. Japanagromyza etiennei n. sp. (Diptera: Agro- with a supplement on species from the Caribbean. Arthropods
myzidae) ravageur potential des Phaseolus spp. [Leguminosae] of Florida and Neighboring Land Areas 7: 1–205.
dans les caraïbes. Revue Française d’Entomologie (Nouvelle Spencer, K.A. & G.C. Steyskal. 1986. Manual of the Agromyzidae
Serie) 16: 81–85. (Diptera) of the United States. United States Department of Ag-
Martinez, M. & J. Etienne. 2002. Liste systématique et biogéographi- riculture, Agriculture Handbook 638: vi + 478 pp.
que des Agromyzidae (Diptera) de la région néotropicale. Bolle- Spencer, K.A., M. Martinez, & J. Etienne. 1992. Les Agromyzidae
tino di Zoologia agraria e di Bachicoltura, Serie II 34: 25–52. (Diptera) de Guadeloupe. Annals de la Société Entomologique
McAlpine, J.F. 1989. Phylogeny and classification of the Musco- de France (N.S.) 28: 251–302.
morpha, pp. 1397–1518. MND, Volume 3. Steyskal, G.C. 1980. Haplopeodes, a new genus for Haplomyza of
Sanabria de Arévalo, I. 1993a. Nuevas especies del género Lirio- authors (Diptera, Agromyzidae). Proceedings of the Entomo-
myza Mik (Diptera: Agromyzidae) en el altiplano de Bogotá logical Society of Washington 82: 140–151.
(Cundinamarca, Colombia). Caldasia 17: 265–281. Tschirnhaus, M. von. 1971. Unbekannte Stridulationsorgane bei
Sanabria de Arévalo, I. 1993b. Dos especies nuevas del género Dipteren und ihre Bedeutung für Taxonomie und Phylogenetik
Ophiomyia Braschnikov (Diptera: Agromyzidae) de San- der Agromyziden (Diptera: Agromyzidae et Chamaemyiidae).
tafé de Bogotá (Cundinamarca, Colombia). Caldasia 17: Beiträge zur Entomologie 21: 551–579.
283–289. Valladares, G.R. 1981. Contribución al conocimiento de las espe-
Sanabria de Arévalo, I. 1994. Insectos minadores (Diptera: Agro- cies de Calycomyza Hendel (Diptera, Agormyzidae), minadoras
myzidae) de la Sabana de Bogotá (Cundinamarca, Colombia). de hojas en la República Argentina. Revista de la Sociedad En-
Revista Colombiana de Entomología 20: 61–100. tomológica Argentina 40: 221–229.
Sasakawa, M. 1992a. The Neotropical Agromyzidae (Diptera). Valladares, G.R. 1992. Contribucion al conocimiento de las espe-
Part 1. New or little-known species from Argentina. Japanese cies de Calycomyza Hendel (Diptera, Agormyzidae), minadoras
Journal of Entomology 60: 346–357. de hojas en la republica Argentina. II. Revista de la Sociedad
Sasakawa, M. 1992b. The Neotropical Agromyzidae (Diptera). Entomologica Argentina 50: 179–200.
Part 2. New or little-known species from Colombia. Akitu, New Valladares, G.R. 1998a. A new species of Amauromyza Hendel
Series 132: 1–20. (Dipt., Agromyzidae) from Argentina. Entomologist’s Monthly
Sasakawa, M. 1992c. The Neotropical Agromyzidae (Diptera). Magazine 134: 235–237.
Part  3. New or little-known species from Ecuador. Japanese Valladares, G.R. 1998b. Three new species of Haplopeodes Stey-
Journal of Entomology 60: 815–825. skal (Dipt., Agromyzidae) from Argentina. Entomologist’s
Sasakawa, M. 1992d. The Neotropical Agromyzidae (Diptera). Monthly Magazine 134: 31–38.
Part 4. New or little-known species from Peru, Venezuela, Bra- Winkler, I.S., S.J. Scheffer & C. Mitter. 2009. Molecular phylogeny
zil and Bolivia. Scientific Reports of the Kyoto Prefectural Uni- and systematics of leaf-mining flies (Diptera: Agromyzidae):
versity, Agriculture 44: 1–33. delimitation of Phytomyza Fallén sensu lato and included spe-
Sasakawa, M. 1994. The Neotropical Agromyzidae (Diptera). Part 5. cies groups, with new insights on morphological and host-use
New or little-known species from Chile. Scientific Reports of evolution. Systematic Entomology 34: 260–292.
the Kyoto Prefectural University, Agriculture 46: 15–41. Woodley, N.E. & D.H. Janzen. 1995. A new species of Melanagro-
Sasakawa, M. 2005. The Neotropical Agromyzidae (Insecta: Dip- myza (Diptera: Agromyzidae) mining leaves of Bromelia pin-
tera). Part 6. New or little-known species from El Salvador and guin (Bromeliaceae) in a dry forest in Costa Rica. Journal of
Guatemala. Species Diversity 10: 151–169. Natural History 29: 1329–1337.
Spencer, K.A. 1963. A synopsis of the Neotropical Agromyzidae Zlobin, V.V. 1996a. Seven new species of mining flies (Diptera:
(Diptera). Transactions of the Royal Entomological Society of Agromyzidae) from Antilles. International Journal of Diptero-
London 115: 291–389. logical Research 7: 147–155.
Spencer, K.A. 1969. The Agromyzidae of Canada and Alaska. Zlobin, V.V. 1996b. The genus Amauromyza Hendel (Diptera, Agromyz-
Memoirs of the Entomological Society of Canada 64: 1–311. idae): a clarification of species of the subgenus Annimyzella Spencer.
Spencer, K.A. 1973a. Agromyzidae (Diptera) of economic impor- International Journal of Dipterological Research 7: 271–280.
tance. Series Entomologica 9. Dr. W. Junk, The Hague, 436 pp. Zlobin, V.V. 1999. The correction of species composition in the
Spencer, K.A. 1973b. The Agromyzidae of Venezuela. Revista de genera Liriomyza Mik and Phytoliriomyza Hendel (Diptera:
la Facultad de Agronomía, Universidad Central, Caracas 7: Agromyzidae). International Journal of Dipterological Re-
5–107. search 10: 129–131.
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
ANTHOMYZIDAE (anthomyzid flies) 81
Kevin N. Barber and Jindřich Roháček

Fig. 81.1. Habitus, male of Mumetopia nigrimana group, (illustrated by A. Brenes).

Diagnosis and sometimes marked, or brachypterous. Head sometimes

distinctly elongate; antenna short with pedicel not swollen
Small (body length 1.1–3.4  mm) usually slender flies or caplike, first flagellomere turned downward, arista
(Fig.  1), ranging in color from yellow to black, shiny to dis- pubescent to pectinate. One to three reclinate fronto-orbital
tinctly microtomentose, with wings usually long, narrow, setae with short setulae anteriorly on fronto-orbital plates

1073
1074
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

and usually on frons. Ocellar and vertical setae always pres- Sweep netting is probably the most productive means
ent; postocellar setae short, convergent, absent only in one of collection. Some species can also be aspirated from the
species. Forefemur usually with pronounced posteroventral leaves of broad-leaved plants. Malaise, light, pan, and pitfall
ctenidial spine (actually a spinelike seta; Fig. 3); sometimes traps are only occasionally a productive means of captur-
foretibia and some tarsomeres black, contrasting with pale ing anthomyzids, but the latter two sometimes yield large
femur. Midtibia with distinct ventroapical seta. Hind femur numbers of those species usually found close to the ground.
in male usually with posteroventral row of shortened, thick- Aspiration of adults from the disturbed thatch below thick
ened setae. Vein R1 of wing with distinctive preapical kink graminoids is especially productive for certain smaller spe-
(best seen in Figs.  2, 11). Male genitalia characterized by cies, especially those with short wings. Some members of
phallapodeme with robust ventral modified phallic guide the Mumetopia nigrimana group have been aspirated near
the bases of grasses as well as swept from the same habi-
(fulcrum) connected with hypandrium and by distiphallus
tats. These same flies can be found on nearby broad-leaved
bifid apically (composed of membranous saccus and slender
plants, although they may be here after flushing from neigh-
sclerotized filum). Female tergite 7 entire or medially split
boring graminoids.
(rarely desclerotized or missing), sometimes fusing with Anthomyzidae appear to be of little economic importance
sternite 7 in various ways. despite being locally abundant in certain habitats.
Most anthomyzids can be recognized by their general
habitus and the presence of the forefemoral ctenidial spine.
Some Neotropical drosophilids have a similar spine (often Classification
part of a series or ctenidium) but are differentiated by their
loosely pectinate arista that is ventrally bare in the basal half. The Anthomyzidae consistently have been considered
Periscelididae, particularly Stenomicra Coquillett, can be most closely related to the Opomyzidae for some time (Hennig,
distinguished by the presence of facial setae. In those few an- 1958; McAlpine, 1989; Roháček, 1998b). McAlpine (1989)
thomyzid species with the femoral spine missing or greatly treated these two families as the suprafamily Opomyzoin-
reduced, the presence of a preapical kink in the Vein R1 is an ea and associated them with 11 others in the superfamily
extremely useful diagnostic character. Opomyzoidea (including Clusioinea, Agromyzoinea, and
Asteioinea).
Roháček (1998b, 2006) has most recently dealt with the
Biology Anthomyzidae in a synthetic manner. He refined the lim-
its of the family and provided a revised subfamily classi-
Biological information for Neotropical species is mostly fication wherein the subfamily Protanthomyzinae contains
limited to label data. Ferrar (1987) and Roháček (1998a) only the fossil genus Protanthomyza Hennig (one  species;
provide the most comprehensive discussions of the biology Baltic amber). The remainder of the world fauna is held in
of Anthomyzidae as summarized here, supplemented with the subfamily Anthomyzinae and is composed of the other
known fossil genus (Grimalantha Roháček  — one  species;
personal observations.
Dominican Republic amber) and 19 extant genera includ-
The adults are usually associated with damp habitats ing Fungomyza, Margdalops Roháček  & Barraclough,
including graminoids in both open and wooded areas. One Epischnomyia Roháček, Receptrixa Roháček, and Zealantha
Nearctic species is associated with grasses on well-drained, Roháček (Roháček, 1999b, 2006, 2007; Roháček & Barra-
sandy soils. Adults have also been taken at fungi and under clough, 2003).
fallen plant stems in varying degrees of decay. Larvae have The Palearctic fauna is well documented (cf. Roháček,
been found feeding in, and adults have been reared from, a 1996, 1999a,  b, 2006), while other regions require much
range of microhabitats, suggesting that many are phytosap- more work. Generic concepts are in need of full integration
rophagous. These include the sheathing leaves of the tillers on a world basis, particularly when considering the New
or terminal shoots of various grasses, sedges, and rushes World fauna. The Nearctic fauna (about 40 species with
(Anthomyza Fallén, Stiphrosoma, Typhamyza Roháček); the many undescribed) is currently organized into five genera
galls of Lipara  spp. (Diptera: Chloropidae) and Steneotar- but may require another two or three, whereas the Afrotropi-
sonemus (Acari: Tarsonemidae) (Anthomyza); leafmining cal Region is expected to eventually yield about 90 species
in dicotyledons of Caryophyllaceae and Lamiaceae (Paran- in more than 12 genera (most currently unnamed).
thomyza Czerny, Anagnota Becker); small mammal nests — The described Neotropical fauna currently consists of
only seven species in three genera (Mumetopia, Chamae-
possibly brought in with nesting material (Stiphrosoma,
bosca Speiser, and Stiphrosoma), and the available generic
Anagnota); and fungi (Fungomyza Roháček). All specimens concepts cannot at this time be applied unambiguously to the
of an undescribed species from Ecuador (Undescribed variety of forms that are known to exist in this region.
Genus C) were aspirated from the inflorescences of two spe- Mumetopia, Chamaebosca, and Stiphrosoma together rep-
cies of Heliconia L. (Heliconiaceae) in early May (M. Buck resent a distinctive group in the Neotropical fauna (female
collector). The female cerci may be modified for inserting internal abdominal characters of wide ring sclerite and pyri-
eggs into crevices or flesh of these flowers. form spermathecae with tubercles), but the limits of each are
ANTHOMYZIDAE (anthomyzid flies) 81 1075

not well defined. Ischnomyia Loew and Amygdalops Lamb 2006). A key to 11 Palearctic or Holarctic genera is found in
share affinities with many of the Neotropical forms treated Roháček (2006), and four Afrotropical genera are described
here. There are about 80 Neotropical species known, but the or revised by Roháček (1993, 2004) and Roháček & Barra-
fauna is expected to rival that of the Afrotropical Region. clough (2003). A revision of the Oriental, Australasian, and
The extant world fauna should thus be expected to climb Oceanian Amygdalops is found in Roháček (2008), while
easily from 101 species (Roháček, 1998b, 1999a,  b, 2004, treatment of two other genera (Mumetopia and Ischnomyia)
2006, 2007, 2008; Sueyoshi & Roháček, 2003; Roháček & appears in Vockeroth (1987) along with the Holarctic An-
Barraclough, 2003; Roháček  & Barber, 2004, 2005) to at
thomyza. Two genera, Apterosepsis Richards and Chamae-
least four times that figure with increased sampling and
bosca, are redescribed and discussed but not keyed by
revisionary work.
Roháček (1998b), while the monotypic Zealantha has been
described most recently by Roháček (2007). Sueyoshi  &
Identification Roháček (2003) provide a key to the anthomyzid species of
Japan, while Roháček & Barber (2005) provide a key to the
Roháček (1998b) provides a checklist of world genera world species of Stiphrosoma, including three Neotropical
and species and a comprehensive bibliography (Roháček, species.

ct

2 Undescribed Genus A 3 Chamaebosca

i vt s
oc s
o vt s
fr s

vb

5 Stiphrosoma

6 Undescribed Genus C
4 Mumetopia

7 Undescribed Genus H
Figs. 81.2–7. Wing, forefemur and tibia, heads: dorsal view of wing of (2) Undescribed Genus A (Caribbean, female); posterior view of
forefemur and tibia of (3) Chamaebosca microptera Speiser (South America, male holotype; Roháček, 1998b, fig. 13); lateral view of heads
of (4) Mumetopia occipitalis Melander (Nearctic, male; MND, fig. 75.2); (5) Stiphrosoma lucipetum Roháček & Barber (Bahamas, holotype
male; adapted from Roháček & Barber, 2005, fig. 42); (6) Undescribed Genus C (South America, male); and (7) Undescribed Genus H (South
America, male). Figure 2 illustrated by A. Brenes; Figures 6 and 7 illustrated by K. Barber.
Abbreviations: ct, ctenidial spine; fr s, frontal-orbital seta; i vt s, inner vertical seta; o vt s, outer vertical seta; oc s, ocellar seta; vb, vibrissa.
1076
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Most anthomyzids bear a characteristic posteroventral offer a clear view and interpretation of tomentosity by
ctenidial spine on the forefemur, which is usually read- reducing shriveling and greasing, and more importantly
ily visible. However, many of the important, diagnostic they more often exhibit maximum distention of terminal
characters for generic and specific identification are as- abdominal segments and greater exposure of male geni-
sociated with the cephalic and thoracic chaetotaxy and the talia while facilitating removal of the abdomen. Direct
terminalia. Specimens that are killed in 70% ethanol and gluing to the side of a pin or mounting on card points are
then critical-point-dried have several advantages. They equally acceptable.

8 Undescribed Genus H 9 Undescribed Genus J

10 Undescribed Genus K 11 Undescribed Genus K

Figs. 81.8–11. Wings: dorsal view of (8) Undescribed Genus H (South America, male); (9) Undescribed Genus J (Caribbean, male); and
(10, 11) Undescribed Genus K (Caribbean, female, male of additional species, respectively). Figures 8–11 illustrated by A. Brenes.

Key to the genera of Anthomyzidae of the Neotropical Region

The key here has been developed with reference to the entire Neotropical Region, including South America and the Caribbean.
Attempts to overlay existing generic concepts on this fauna have met with limited success, necessitating an interim step of
providing a key to groups of species tentatively offered as generic concepts. A much more exhaustive study is required, which
will be particularly aided with more extensive collections of representatives outside the Mumetopia nigrimana group. For
clarification, Chamaebosca cursor sensu Malloch (1933) is treated here as Undescribed Genus M, whereas C. cursor sensu
Hennig (1955) is likely the same as Undescribed Genus B.

1. Wing blade reduced, even straplike; Colombia, Ecuador, Chile. . . . . . . . [Chamaebosca Speiser]
– Wing blade not noticeably reduced. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2. Vein dm-cu missing (Fig.  2); cell cup absent or incomplete; surstylus elongate; spermathecae
reduced; Caribbean, Brazil. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Undescribed Genus A]
– Vein dm-cu present (as in Figs.  8–11) (rarely missing on one wing only); cell cup usually
complete; surstylus various; spermathecae of usual size. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
ANTHOMYZIDAE (anthomyzid flies) 81 1077

3. Forefemoral ctenidial spine absent or length reduced to no more than 3 times width of basal
diameter. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
– Forefemoral ctenidial spine present and strong (Fig. 3), length approaching or exceeding opposite
diameter of tibia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

4. Cephalic and thoracic chaetotaxy unusually strong including presutural intra-alar seta as long as
scutellum; ctenidial spine strongly reduced but discernible in front of long posteroventral setae on
forefemur; legs entirely yellow and halter knob white; arista long-ciliate; Chile (Juan Fernandez
Islands). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Undescribed Genus B]
– Chaetotaxy not unusually strong, presutural intra-alar seta shorter than scutellum and usually
indistinguishable from acrostichal setulae; forefemoral ctenidial spine strongly reduced or absent;
color of legs and halter various; arista long-pectinate or if short-pectinate then thorax, femoral
apex, tibial base, and halter knob darkened. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

5. Arista no more than short-pectinate (as in Figs. 4, 7) . . . . . . . . . . . . . . . . . . . . . Unplaced Species

– Arista long-pectinate (Fig. 6). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

6. Only one pair of dorsocentral setae; notum with glabrous stripe immediately laterad of dorsocentral
lines; notum predominantly black with scutellum yellow apically; male sternite 5 with single
medial patch of short setulae near posterior margin; female with cerci apically pointed; Ecuador.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Undescribed Genus C]
– Two pairs of dorsocentral setae, anterior seta short but at least twice length of acrostichal setulae;
notum without glabrous stripe; notal coloration including wide yellow vitta filling notal area
between dorsocentral lines, extending over scutellum and onto tergites 1 and 2 medially; male
sternite 5 with longer setulae in one patch or short in two distinct patches near posterior margin;
female cerci apically blunt. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Undescribed Genus D

7. Only one fronto-orbital seta well developed (Fig. 4), anterior seta, if present, less than one-third
length of posterior; fronto-orbital plates bare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
– Usually two (rarely three) fronto-orbital setae (Figs. 5, 7) well developed, anterior (middle) seta
at least one-third length of posterior; if only one seta, then fronto-orbital plates tomentose (bare
immediately surrounding insertion of seta in one species) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

8. Inner vertical and ocellar setae reduced to one-half length or less, respectively, of outer vertical
seta; postocellar setae absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Undescribed Genus E
– Inner vertical and ocellar setae stronger, at least two-thirds length of outer vertical setae (Fig. 4);
postocellar setae present though often weak. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

9. Foretibia and at least basal tarsomere usually entirely black with some apical tarsomeres usually
white contrasting with yellow femur, if not so colored, then wing with cell cup present; midtibial
ventroapical seta shorter than tarsomere  2; female tergite  7 complete or narrowly divided
medially. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mumetopia Melander, in part
– Foreleg entirely concolorous yellow; wing with cell cup absent; midtibial ventroapical seta
elongated, subequal to length of tarsomere 2 or longer; female tergite 7 complete or missing or
completely desclerotized dorsally (middle one-third of tergite) and represented only by lateral
extremities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Undescribed Genus F

10. Foretibia and tarsus entirely black, contrasting with yellow femur and other legs; frons and fronto-
orbital plates densely tomentose, appearing black or silver; arista long-pubescent especially dense
in basal one-third. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mumetopia Melander, in part
– Foretibia and tarsus not contrasting in color with femur or other legs; frons, fronto-orbital plates,
and arista various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
1078
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

11. Presutural dorsocentral seta present along with three postsutural dorsocentrals; anterior of three
fronto-orbital setae about one-quarter length of posterior fronto-orbital seta; female sternite 7 and
tergite 7 fused into ring although sternal area pale, contrasting with dark tergal elements (male
unknown). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Undescribed Genus G
– Presutural dorsocentral seta absent, with two or three postsutural dorsocentrals; only two pairs of
fronto-orbital setae well developed; female sternite 7 and tergite 7 elements various . . . . . . . . 12

12. Wing usually with extensive dark markings (sometimes reduced to diffuse infumation of cell r1 or
diffuse clouding at apex of vein R2+3) or if clear, then at least scutellum contrastingly marked with
yellow at least medially; head often elongate; subvibrissa usually shorter than first flagellomere or
absent (Fig. 7). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
– Wing clear or evenly lightly infumated; scutellum concolorous with notum, not with contrasting
medial or apical yellow markings; head not elongate; subvibrissa well developed, subequal to or
longer than first flagellomere (Fig. 5) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

13. Anterior margin of frons projecting apically and dorsally (Fig. 7) (in questionable cases cell r1 and
at least anterior half of cell r2+3 darkened in at least distal half (Fig. 8), wing without discrete
hyaline spots); wing cell  r4+5 usually slightly narrowing apically, vein  R2+3 running in close
proximity to C . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Undescribed Genus H
– Frons not projecting; wing cell r4+5 not narrowing apically, vein R2+3, cells r1 and r2+3 various, wing
pattern sometimes with hyaline spots (Figs. 9,11). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

14. Ocellar triangle entirely tomentose; scutellum medially or apically yellow and abdominal tergites
usually with contrasting pattern of medial and/or anterolateral yellow markings; if scutellum and
abdomen entirely dark then wing markings reduced to small subapical cloud surrounding tip of
vein  R2+3 (as in Fig. 10); head with one or two well-developed fronto-orbital setae but if with
two setae, then halter knob completely or partially darkened. . . . . . . . . . . . Undescribed Genus I
– Ocellar triangle glabrous at least medially anterior to anterior ocellus; scutellum and abdomen
usually completely black and head with two pairs of well-developed fronto-orbital setae; if with
yellow markings on notum, scutellum, or abdomen (two species) then halter entirely white; wing
markings various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

15. Wing pattern extensive and including three hyaline spots (Fig. 9); vein R4+5 straight; thorax and
abdomen without contrasting yellow markings; fronto-orbital plates widening anteriorly; female
without asymmetrical internal abdominal sclerites. . . . . . . . . . . . . . . . . . . . Undescribed Genus J
– Wing with various markings, with only apical clouding and vague longitudinal band (Fig. 10) or
more extensive and sometimes including only two hyaline spots (Fig. 11); vein R4+5 with slight
bend forward or with yellow markings on thoracic notum, scutellum, and abdomen; fronto-orbital
plates parallel-sided, not widening anteriorly; female with heavily sclerotized asymmetrical
internal abdominal sclerites; Caribbean. . . . . . . . . . . . . . . . . . . . . . . . . . . [Undescribed Genus K]

16. Occiput without white tomentum above foramen; notum tomentose gray or if glabrous brown,
with two secondary, short, spinelike setae anterad of forefemoral ctenidial spine; Chile. . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Undescribed Genus L]
– Occiput with white tomentose supracervical patch; notum yellow or glabrous brown; only usual
single forefemoral ctenidial spine present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

17. White, tomentose supracervical patch medially divided; notum yellow with at least medial brown
vitta usually continuing onto scutellum or concolorous brown. . . . . . . . . . . . Stiphrosoma Czerny
– White, tomentose supracervical patch not medially divided; notum concolorous brown, without
medial vitta; Ecuador, Peru, Bolivia, Chile, Argentina . . . . . . . . . . . . . . . [Undescribed Genus M]
ANTHOMYZIDAE (anthomyzid flies) 81 1079

Synopsis of the fauna Specimens have been taken at elevations that range from
0–100 m (Dominica) to 2500–3000 m (Ecuador).
Approximately 25 species of Anthomyzidae are known
to occur in Central America, but the fauna will likely be Stiphrosoma Czerny. This Holarctic genus is represented
found to include about 35 species. Six of these species are in Central America by three species. Stiphrosoma sororium
represented by single specimens, while 12 others are not yet Roháček  & Barber is known only from Chiapas, Mexico
known outside the region. Several species of the Mumetopia (2400  m elevation). A second more widespread species
nigrimana group are widely distributed and together pre- (including Bahamas and Florida), S. lucipetum Roháček &
dominate the examined collections of Anthomyzidae from Barber, is known from a single female from Costa Rica
Central America and the Neotropical Region in general. The (Manglar, Rio Corcovado) and a series of males from Be-
overall distribution of the family appears to be quite wide, lize (Twin Cays). Stiphrosoma pullum Roháček & Barber is
including high and low altitudes, although most collections much darker, has silvery tomentose fronto-orbital plates, and
are concentrated at mid-elevations. is known from a single male from Costa Rica (Punta Coy-
Five of the seven described species of Neotropical ote). These three species are heavily bristled flies that are
Anthomyzidae are known to occur in Central America (see related to a group of Nearctic species (Roháček  & Barber,
Mumetopia and Stiphrosoma). The other two are short- 2005). There are also four described Palearctic species and
winged Chilean species (Chamaebosca microptera Speiser two Holarctic species (Roháček, 1996, 2006; Roháček  &
and C. cursor (Kieffer); see discussion in Roháček, 1998b). Barber, 2005).
However, two primary factors have limited this preliminary Habitats include both lowland areas (mangrove) and
breakdown of the fauna at the generic level: world generic higher altitudes (2200 m). Stiphrosoma lucipetum has been
concepts are just beginning to be developed, and many Neo- taken at lights in Florida. Nearctic species, especially those
tropical species are represented by single specimens or with short wings, are most readily collected with an aspirator
extremely short series (unique specimens of either sex were at the bases of graminoids.
not dissected for this study). Except for the three species of
Stiphrosoma (Roháček & Barber, 2005), there are no keys to
Undescribed Genus D. This undescribed genus is repre-
species of any of the Neotropical fauna.
sented by about six similar species known from Costa Rica,
Mumetopia Melander. This New World genus is known Panama, Colombia, Ecuador, Trinidad, Brazil, and Peru.
from the primarily Nearctic M. occipitalis Melander (M. ter- Outwardly, these specimens resemble Amygdalops, but sev-
minalis (Loew) is referable elsewhere) and the mostly Neo- eral characteristics suggest that they may represent a distinct
but closely related grouping. It is preferred to note this close
tropical M. nigrimana Coquillett. The usually darkened tibia
relationship rather than to begin using Amygdalops in refer-
and basal tarsomeres of the foreleg help to define the M. nigri-
ence to New World fauna at this time.
mana group, which contains most of the Mumetopia treated
here, including the widespread M. nigrimana. A few species Undescribed Genus E. A unique male from Costa Rica
with pale forelegs and usually wing infuscation belong (La Suiza, Cartago) that is slender and shiny black is tenta-
here as supported by male and female genitalic characters. tively treated separately, based primarily on the loss of the
There is a single specimen from Panama and two others from postocellar setae and the strong reduction of the inner ver-
northeastern Mexico that appear to be disjunct records for tical and ocellar setae. The head is flattened and elongate,
M. occipitalis. More significantly, there are specimens from but the frons does not project as in Undescribed Genus H.
the Galápagos Islands that appear to represent three species The weakly patterned wings have the R2+3 running in close
that are similar to M. occipitalis. At least 18 Neotropical spe- approximation to C, a pattern that is reminiscent of Un-
cies are included here of which about half are likely to be described Genus K, but the exposed genitalia are much more
found in Central America. Two species (from Costa Rica and compact.
Mexico) are remarkable in having two strong fronto-orbital
Undescribed Genus F. Only one of the four species of
setae and heavy tomentosity of the frons and fronto-orbital
this group occurs in Central America, ranging from Mexi-
plates (reminiscent of the Palearctic Anagnota).
co to Colombia. It is the smallest species of Anthomyzidae
Common species of the M. nigrimana group are frequent- presently known from this region. Another species is known
ly encountered in a variety of habitats, even in urban areas from Jamaica, while two others are known from Trinidad
where they are associated with long grasses on disturbed (unique male) and Brazil (two females). Two of the three
sites. Specimens referred here usually dominate collections smaller species are known to have the female tergite 7 broadly
of Neotropical Anthomyzidae. They are most commonly col- desclerotized dorsally, which appears to be a unique feature
lected by sweeping, but Malaise, flight intercept, pan, and within the Anthomyzidae. The larger species (Brazil) has the
light traps have yielded specimens. Mumetopia nigrimana female tergite 7 entire, which may indicate that this species is
alone has been taken at lights (Florida, Mexico), black lights improperly placed, but it is not a Mumetopia (spermathecae
(Mexico), and in light traps (Mexico, Panama, Dominica). neither pyriform nor tuberculate). Collection records include
1080
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

reference to Malaise traps and a stream bed in Jamaica. One a striking similarity to members of Undescribed Genus I in
species has been taken in the Mojinga Swamp (Fort Sher- body shape and coloration, but the female has the character-
man, Canal Zone, Panama), at 10 m altitude (Osa Peninsula, istic asymmetrical internal abdominal sclerites). Five species
Costa Rica), 212 m (Palo Verde National Park, Costa Rica), of Undescribed Genus  I are known only from Costa Rica,
but also at 2400 m (Medellín, Colombia). and a sixth species is known only from Costa Rica and Ni-
caragua. Another four species are known from St. Lucia (1),
Undescribed Genus G. Represented by only two females Venezuela (2), and Brazil (1). Some records are from a Mal-
(questionably conspecific) from Panama and Mexico, this aise trap and a flight intercept trap.
genus is unique in the Neotropical fauna for its possession
of 1+3 dorsocentral and three fronto-orbital setae. It does not Undescribed Genus J. Only three species are included
appear to be related to Santhomyza Roháček (1984), as it dif- here, of which one is represented by a unique male (from
fers in having the 7th tergite and sternite forming a complete Cuba). A second species is known from Jamaica and a third
uninterrupted ring, although the ventral area is not as heav- from Costa Rica and Mexico (two males). The wing pattern
ily sclerotized. It is preferred not to begin using a European is unique with three hyaline spots arranged in a triangle,
name for a poorly associated species. the medial spot (cell  r4+5) shifted apicad of the other two.
The male from Costa Rica was swept from vegetation at
Undescribed Genus H. Four species of this genus are the Monteverde Ecolodge San Luis, and the male from
known only from Costa Rica, while a fifth species ranges Mexico was taken in a pan trap among coastal shrubs (Xpu-
from Panama to Ecuador and Venezuela. The members of Há, Quintana Roo). Most of the specimens of the Jamaican
this genus are usually recognized by the combination of the species were taken in a “stream bed” with another single
prominent extension of the frons and the darkened pattern of Malaise capture represented.
the wing. However, internal genitalic characteristics of most
species have not been examined, and more clear distinction Unplaced Species. These three species are represented
from Undescribed Genus I is not yet possible. Another nine by a total of four females and bear attention for the lack,
species are known from South America. At least a superfi- or strong reduction, of the forefemoral ctenidial spine. Two
cial resemblance to Margdalops is notable. Specimens have species were taken in Costa Rica and are relatively large
been taken in a Malaise trap, at lights, and swept from várzea (over 3  mm). The combination of ctenidial spine loss and
habitat. large size suggest that these two species are closely related.
The unique female was taken on foliage outside the entrance
Undescribed Genus I. All but two of the species included
to Tapanti National Park (1150 m), while one female of the
here have a medial yellow marking evident on the scutellum.
There is also usually some indication of yellow markings on other species was taken in a Malaise trap (24 km west of Pie-
the abdomen and the thorax. One species (3 km southeast of dras Blancas, 200 m). The third species has a greatly reduced
Río Naranjo, Costa Rica) has the frons slightly projecting, forefemoral ctenidial spine and is known from Paucartambo,
but the wing pattern is not as extensive as in Undescribed Peru (50 km northwest of Pilcopata, 1600 m). It is smaller
Genus H (apical pattern similar to that in Undescribed Ge- than the other two unplaced species and has the acrostichal
nus K, Fig. 10; this species of Undescribed Genus K bears setulae arranged in two rows.

Literature cited

Ferrar, P. 1987. A guide to the breeding habits and immature stages Roháček, J. 1984. Santhomyza gen. n., a new genus of Anthomyzi-
of Diptera Cyclorrhapha. Entomonograph 8(1). E.J.  Brill  / dae (Diptera) from the Mediterranean area. Bolletino del Museo
Scandinavian Science Press, Leiden, Copenhagen, 478 pp. Regionale di Scienze Naturali, Torino 2: 531–543.
Hennig, W. 1955. Los insectos de las Islas Juan Fernandez. Roháček, J. 1993. Two new Afrotropical genera of Anthomyzidae
16.  Phryneidae, Helomyzidae, Lonchaeidae, Piophilidae, (Diptera), with descriptions of seven new species. Annals of the
Natal Museum 34: 157–190.
Anthomyzidae und Muscidae (Diptera). Revista Chilena de Roháček, J. 1996. Revision of Palaearctic Stiphrosoma, including
Entomología 4: 21–34. the Anthomyza laeta-group (Diptera: Anthomyzidae). European
Hennig, W. 1958. Die Familien der Diptera Schizophora und ihre Journal of Entomology 93: 89–120.
phylogenetischen Verwandtschaftsbeziehungen. Beiträge zur Roháček, J. 1998a. 3.22. Family Anthomyzidae, pp.  267–278. In
Entomologie 8: 505–688. Papp, L. & B. Darvas (editors). Contributions to a Manual of
Malloch, J.R. 1933. Acalyptrata. (Helomyzidae, Trypetidae, Scio- Palaearctic Diptera. Volume  3, Higher Brachycera. Science
myzidae, Sapromyzidae, etc.), pp. 177–392. Diptera of Patago- Herald Publishers, Budapest, 880 pp.
Roháček, J. 1998b. Taxonomic limits, phylogeny and higher classi-
nia and South Chile, Part VI – Fascicle 1. British Museum of
fication of Anthomyzidae (Diptera), with special regard to fossil
Natural History, London. record. European Journal of Entomology 95: 141–177.
McAlpine, J.F. 1989. Phylogeny and classification of the Musco- Roháček, J. 1999a. Taxonomy and distribution of West Palaearctic
morpha, pp. 1397–1518. MND, Volume 3. Anthomyzidae (Diptera) with special regard to the Mediterra-
ANTHOMYZIDAE (anthomyzid flies) 81 1081

nean and Macaronesian faunas. Bolletino del Museo Regionale and Oceanian Regions. Acta Zoologica Academiae Scientia-
di Scienze Naturali, Torino 16: 189–224. rum Hungaricae 54: 325–400.
Roháček, J. 1999b. A revision and re-classification of the genus Roháček, J.  & K.N. Barber. 2004. A new species of the genus
Paranthomyza Czerny, with description of a new genus of Fungomyza from the Nearctic Region (Diptera: Anthomyzi-
Anthomyzidae (Diptera). Studia diperologica 6: 373–404. dae). Časopis Slezského zemského Muzea, Opava (A) 53:
Roháček, J. 2004. Revision of the genus Amygdalops Lamb, 1914 131–141.
(Diptera: Anthomyzidae) of the Afrotropical Region. African Roháček, J.  & K.N. Barber. 2005. Revision of the New World
Invertebrates 45: 157–221. species of Stiphrosoma Czerny (Diptera: Anthomyzidae).
Roháček, J. 2006. A monograph of Palaearctic Anthomyzidae (Dip- Beiträge zur Entomologie 55: 1–107.
tera), Part  1. Časopis Slezského zemského Muzea, Opava (A) Roháček, J. & D. Barraclough. 2003. Margdalops, a new Afri-
55(Suppl. 1): 1–328. can genus of Anthomyzidae (Diptera), comprising six new
Roháček, J. 2007. Zealantha thorpei gen. et sp.  nov. (Diptera: species. African Invertebrates 44: 157–190.
Anthomyzidae), first family representative from New Zealand. Sueyoshi, M. & J. Roháček. 2003. Anthomyzidae (Diptera: Aca-
Zootaxa 1576: 1–13. lyptrata) from Japan and adjacent areas. Entomological Sci-
Roháček, J. 2008. Revision of the genus Amygdalops Lamb, ence 6: 17–36.
1914 (Diptera, Anthomyzidae) of the Oriental, Australasian Vockeroth, J.R. 1987. Anthomyzidae. MND, Volume 2.
1082
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
AULACIGASTRIDAE (aulacigastrid flies) 82
Alessandra Rung and Wayne N. Mathis

Fig. 82.1. Male of Aulacigaster sp., (Nearctic, MND, fig. 76.1).

Diagnosis above phallapodeme (Fig.  6). Segments  6 and 7 of female

abdomen each with separate tergite and sternite.
Small to medium-sized flies (body length 1.5–4.0  mm), Third instar larvae (Fig. 7) bearing long, partly retractile
predominantly dark brown to black (Fig. 1). Face completely terminal respiratory tube; anterior spiracle small, retracted
sclerotized, often protruding. Pedicel not caplike, sinuous into deep, narrow pocket; posterior spiracles on apex of re-
dorsally, but without slit; first flagellomere porrect or decum- spiratory tube, joined at their base, bifurcating posteriorly.
bent; arista unsegmented, often with microtrichia on apical Puparia light brown, oval in shape, with prothoracic spir-
two-thirds, microtrichia short to elongate, zigzagged, dorsal acle completely everted; posterior spiracular tube strongly
and ventral microtrichia alternating; ocellar seta, when pres- retracted.
Adults of Aulacigaster are readily distinguished from
ent, minute to moderately long (up to three-quarters length
genera of Periscelididae and Anthomyzidae by the shape of
of lateral vertical seta); with two fronto-orbital setae; intra-
the pedicel, which is not caplike, by the confluent cells bm
frontal seta absent; peristomal chaetotaxy typically consist-
and dm (also in some Cyamops Melander and Stenomicra
ing of weak setulae posterior to pseudovibrissal seta. Two
Coquillett) and by the position of crossvein r-m at the basal
scutellar setae present, apical pair nearly parallel, oriented
one-third of cell dm.
posteriorly; anepisternum bearing one–two posterior setae.
Wing (Figs. 2–3) hyaline or infuscate, brown; cell dm flat,
lacking longitudinal crease or fold; crossvein r-m basal to Biology
middle of confluent cells dm + bm; Sc partially fused with
R1. Male preabdomen consisting of five  segments; sursty- Adults and immatures of the A.  leucopeza group (Ma-
lus fused with epandrium (Fig. 4); hypandrium bearing lat- this & Freidberg, 1994) are attracted to weeping wounds and
eral, anterior apodeme arising from base of each gonopod sap fluxes of deciduous trees (Robinson, 1953; Teskey, 1976).
(Fig. 5), halves of hypandrium connected by narrow bridge The larvae of a group of species have been found to inhabit

1083
1084
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

bromeliads in the Neotropical Region (A. Rung & W.N. Ma- sister group of the family, according to J.F. McAlpine’s phy-
this, in preparation), where they are fully aquatic and prob- logeny (1989), is the family Periscelididae. Characterization
ably feed on decaying organic matter. The adults of some of Aulacigastridae differs widely among authors (see below),
species of Aulacigaster have also been collected on freshly and up to five genera have been assigned to it, besides the
cut logs and a few species on leaves of Heliconiaceae. nominate genus, Aulacigaster. The Aulacigastridae sensu
stricto, defined by the characters listed in the diagnosis, in-
cludes only one other genus, Curiosimusca Rung et al. (2005),
Classification which is from the Oriental Region and which represents an
addition to what recent authors (Mathis & Freidberg, 1994;
The family Aulacigastridae belongs to the superfamily Papp, 1997; Baptista, 1998b; Hilger & Kassebeer, 2001) have
Opomyzoidea (Teskey, 1987; J.F. McAlpine, 1989), and the adopted in their characterizations of the family. A broader

R1 R2+3

R4+5

M1

A1+CuA2
CuA2

2 Aulacigaster 3 Aulacigaster

4 Aulacigaster 5 Aulacigaster 6 Aulacigaster

7 Aulacigaster

Figs. 82.2–7. Wings, male terminalia, and larva: dorsal view of wing of two different undescribed species (2) and of Aulacigaster spp. (3);
lateral view of epandrium and fused surstylus (4), lateral view of hypandrium and gonopod (5), and ventral view of subepandrial plate (6) of
Aulacigaster spp., (Mathis & Friedburg, 1994, figs. 7–9); lateral view of third instar larva of (7) Aulacigaster sp., (Nearctic, MND, fig. 76.6).
Figures 2–3 photographed by A. Rung.
AULACIGASTRIDAE (aulacigastrid flies) 82 1085

characterization for the family includes the genera Cyamops, Identification

Stenomicra, Planinasus Cresson, and a fossil species from
Baltic amber, Protoaulacigaster electra Hennig. Hennig Mathis & Freidberg (1994) revised the Nearctic species
(1958, 1965, 1969, 1971) first advocated this definition, fol- of Aulacigaster and provided a key to the species of that re-
lowed by Griffiths (1972, with the exclusion of Stenomicra), gion. A key to the Palearctic species was published by Papp
Teskey (1987), and J.F.  McAlpine (1989). D.K.  McAlpine (1997). Barraclough (1993) and Hilger & Kassebeer (2001)
(1978, 1983) proposed a rather different characterization that treated the Afrotropical species of Aulacigaster.
(1) transferred the genera Stenomicra, Cyamops, and Plani-
nasus from Aulacigastridae to Periscelididae, (2)  excluded
Protoaulacigaster electra from Aulacigastridae, and (3) de- Synopsis of the fauna
scribed two new genera in the family: Ningulus McAlpine
(Afrotropical Region) and Nemo McAlpine (Australian Re- Aulacigaster Macquart. There are 14 valid species of
gion). Freidberg (1994) created the family Neminidae for Aulacigaster in the Neotropical (Papavero, 1967; Hennig,
Ningulus, Nemo, and his new Afrotropical genus, Nemula 1969), Nearctic (Mathis  & Freidberg, 1994), Palearctic
Freidberg. Freidberg’s work resulted again in a monotypic (Papp, 1984; Teskey 1987), and Afrotropical (Barraclough,
Aulacigastridae. Protaulacigaster electra Hennig was most 1993) Regions. Two species, A.  melanoleuca Hennig and
recently listed in Aulacigaster by Evenhuis (1984, 1994), A.  minuta Hennig, are recorded from Central America.
and Roháček (1998) tentatively transferred the genus Echid- However, better sampling has revealed Aulacigaster to be
nocephalodes Sabrosky (Seychelles) from the Anthomyzi- far richer in species (Mathis  & Freidberg, 1994), and ap-
dae to the Aulacigastridae. In spite of these varied concepts proximately 37 undescribed Neotropical species are known,
of the family, we consider Aulacigastridae to consist of only of which 12 occur in Central America (Baptista, 1998a, b;
Aulacigaster and Curiosimusca A. Rung & W.N. Mathis, in preparation).

Literature cited

Baptista, A.R. 1998a. Systematics of the family Aulacigastridae. Mathis, W.N. & A. Freidberg. 1994. A review of North American
Abstracts of the fourth International Congress of Dipterology, Aulacigaster Macquart (Diptera: Aulacigastridae). Proceedings
6–13 September, Oxford, UK. of the Entomological Society of Washington 96: 583–598.
Baptista, A.R. 1998b. Preliminary cladistic analysis of the family McAlpine, D.K. 1978. Description and biology of a new genus
Aulacigastridae sensu lato (Diptera, Cyclorrhapha). Abstracts of of flies related to Anthoclusia and representing a new family
the 17th meeting of the Willi Hennig Society, 21–25 September, (Diptera, Schizophora, Neurochaetidae). Annals of the Natal
São Paulo, Brazil. Museum 23: 273–295.
Barraclough, D.A. 1993. The Afrotropical species of Aulacigaster McAlpine, D.K. 1983. A new subfamily of Aulacigastridae (Dip-
Macquart (Diptera, Aulacigastridae: Aulacigastrinae). Annals of tera: Schizophora), with a discussion of aulacigastrid classifica-
the Natal Museum 34: 31–42. tion. Australian Journal of Zoology 31: 55–78.
McAlpine, J.F. 1989. Phylogeny and classification of the Musco-
Evenhuis, N.L. 1984. Catalog of the Diptera of the Australasian and
morpha, pp. 1397–1518. MND, Volume 3.
Oceanian Regions. Bishop Museum Special Publication 86, Bish- Papavero, N. 1967. 91. Family Aulacigastridae. In Papavero,  N.
op Museum Press & E.J. Brill, Honolulu & Leiden, 1155 pp. (editor). A Catalogue of the Diptera of the Americas South of the
Evenhuis, N.L. 1994. Catalogue of the Fossil Flies of the World (In- United States. Departamento de Zoologia, Secretaria da Agri-
secta: Diptera). Backhuys Publishers, Leiden, 600 pp. cultura, São Paulo, 2 pp.
Freidberg, A. 1994. Nemula, a new genus of Neminidae (Diptera) Papp, L. 1984. Family Aulacigastridae, pp. 60–61. In Sóos, Á. and
from Madagascar. Proceedings of the Entomological Society of L. Papp (editors). Catalogue of Palaearctic Diptera. Volume 10.
Washington 96: 471–482. Elsevier Science Publishers and Akadémiai Kiadóm, Budapest,
Griffiths, G.C.D. 1972. The phylogenetic classification of Diptera 402 pp.
Cyclorrhapha with special reference to the structure of the male Papp, L. 1997. The Palaearctic species of Aulacigaster Macquart
postabdomen. Dr. W. Junk, The Hague, 340 pp. (Diptera, Aulacigastridae). Acta Zoologica Academiae Scientia-
Hennig, W. 1958. Die Familien der Diptera Schizophora und ihre rum Hungaricae 43: 225–234.
phylogenetischen Verwandtschaftsbeziehungen. Beiträge zur En- Robinson, I. 1953. The postembryonic stages in the life cycle of
tomologie 8: 505–688. Aulacigaster leucopeza (Meigen) (Diptera, Cyclorrhapha: Aul-
Hennig, W. 1965. Die Acalyptraten der Baltischer Bernsteins. Stutt- acigasteridae). Proceedings of the Royal Entomological Society
garter Beiträge zur Naturkunde 145: 1–215. of London (A) 28: 77–84.
Hennig, W. 1969. Neue Gattungen und Arten der Acalyptratae. The Roháček, J. 1998. Taxonomic limits, phylogeny and higher classifi-
Canadian Entomologist 101: 589–633. cation of Anthomyzidae (Diptera), with special regard to fossil
Hennig, W. 1971. Neue Untersuchungen uber die Familien der Di- record. European Journal of Entomology 95: 141–177.
Rung, A., W.N. Mathis, & L. Papp. 2005. Curiosimusca, gen. nov.,
ptera Schizophora (Diptera: Cyclorrhapha). Stuttgarter Beiträge
and three new species in the family Aulacigastridae from the Ori-
zur Naturkunde 226: 1–76. ental Region (Diptera: Opomyzoidea). Zootaxa 1009: 21–36.
Hilger, S. & C.F. Kassebeer. 2001. A new species of Aulacigaster Teskey, H.J. 1976. Diptera larvae associated with trees in North
Macquart, 1835 (Diptera, Aulacigastridae) from Réunion. Dip- America. Memoirs of the Entomological Society of Canada 100:
teron 3: 167–172. 1–53.
Teskey, H.J. 1987. Family Aulacigastridae, pp.  887–891. MND,
Volume 2.
1086
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
PERISCELIDIDAE (periscelid flies) 83
Wayne N. Mathis and Alessandra Rung

Fig. 83.1. Male of Periscelis annulata (Fallén), (Nearctic, MND, fig. 77.1).

Diagnosis Male preabdomen consisting of six segments, with postab-

domen (Figs.  6–7) either symmetrical or asymmetrical;
Moderately small to medium-sized flies (body length 2.5– surstylus discrete, articulated or fused with epandrium;
5.0 mm), broad to slender (Fig. 1). Face uniformly sclerotized, cercus weakly sclerotized. Female abdomen (Fig.  8) with
protruding, ventral portion of face receding. Pedicel swollen seventh tergite and sternite separate or fused to each other
and caplike, with dorsal cleft, bearing one to two dorsoapical forming complete ring.
setae; first flagellomere moderately to sharply deflexed, aris- Eggs (based on three European species of Periscelis) sky
ing from ventral surface of pedicel; arista with dorsal and blue to violet; boat-shaped, long ovoid in dorsal view, with-
ventral rays. Ocellar seta, when present, inserted laterally to out filaments; ventral portion convex, lighter in color than
margin of ocellar tubercle; inner vertical seta absent or pres- dorsum, with fine, irregular, hexagonal network; dorsal portion
ent; postocellar seta, when present, reduced and divergent; darker, less convex but not flat, with two fine ribs that are
one to two fronto-orbital setae present with one seta reclinate confluent at apices, lacking network but microsculptured;
and other proclinate; interfrontal seta absent or present, micropyle inconspicuous.
upright and slightly reclinate. Dorsocentral setae 0+1 or 0+2; Papp (1995) summarized the larval features of
posterior intra-alar seta reduced; scutellar setae one to two Periscelis (subgenera Periscelis and Myodris) as follows
marginal pairs, disc of scutellum bare. Wing (Fig. 3) hyaline (Fig.  9): body strongly flattened dorsoventrally; segments
or with infuscate markings; true costal breaks absent; Sc with fleshy digitiform processes laterally and with several
incomplete, not fused with R1 apically; C extended to R4+5 other bulbous processes, all processes with large, partly
or M; cell dm with flat longitudinal fold; cell cup typically curved spinules; subanal pads indistinct (there is a bifid
present, although CuA2 either well developed or reduced. inflated protuberance that can be extroverted from anus in

1087
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

P.  annulata (Fallén)); anterior spiracles short, bulbiform Adults of Periscelididae are readily distinguished from
with four to five small bulbous papillae; posterior spiracles genera of Anthomyzidae and Aulacigastridae by the shape
(with lateroclinate intraspiracular hairs) on pair of long of the pedicel, which is caplike; by the arista, which bears
divergent spiracular processes; cephalopharyngeal skeleton dorsal and ventral rays; by the longitudinal fold in cell dm;
with robust body (pharynx); mandibles strong, composed and cell cup typically present, although CuA2 either well
of curved apical part and robust basal (posterior) part and developed or reduced.
with one pair of ventrobasal, midventral, and dorsal processes
each; extremely long and thin parastomal bar; well-sclerotized Biology
dental sclerites and weakly sclerotized but complex labial sc-
lerite; dorsal cornu much shorter than ventral; ventral cornu The immature stages of Periscelis, and to an extent the
robust but simple. Larvae of Myodris plump, those of P. (Peri adults, are associated with sap from bleeding deciduous trees
scelis) slim; lateral process of the posterior spiracle distal (oak, elm, cottonwood, etc.). Papp (1998) recently described
in Myodris or proximal in Periscelis (extended to middle of the habits and habitats of three European species. Williams
posterior spiracle); aerial sack (Papp, 1995) behind posterior (1939) reported larvae of Stenomicra orientalis Malloch
spiracle conical (Myodris) or sausage-shaped (Periscelis). at the water-holding leaf bases of several monocot species

i vt s
poc s o vt s

oc s
R1
Sc
orb s

R2+3

R4+5

fc

gn M1

sbvb s clyp CuA1

A1+CuA2

3 Periscelis

2 Periscelis

o vt s

i vt s

orb s

4 Cyamops 5 Stenomicra

Figs. 83.2–5. Heads and wing: anterior view of head of (2) Periscelis annulata (Fallén), (Nearctic, MND, fig. 77.2); dorsal view of wing of
(3) P. annulata, (Nearctic, MND, fig. 77.3); anterior view of head of (4) Cyamops nebulosus Melander, (Nearctic, MND, fig. 76.2, as
Aulacigastridae); and (5) Stenomicra angustata Coquillett, (MND, fig. 76.3, as Aulacigastridae).
Abbreviations: clyp, clypeus; fc, face; gn, gena; i vt s, inner vertical seta; oc s, ocellar seta; orb s, orbital seta; o vt s, outer vertical seta; poc s,
postocellar seta; sbvb s, subvibrissal seta.
PERISCELIDIDAE (periscelid flies) 83 1089

st 6 st 6

tg 6
tg 6
hypd ph

syntgst
7+8 sur ej
apod
ph sur
pgt syntgst
cerc pgt 7+8
epand cerc epand

6 Periscelis 7 Periscelis
spmth

tg 7 tg 8

cerc

st 7 st 8
8 Periscelis

9 Periscelis

Figs. 83.6–9. Male terminalia, female abdomen, and larva: ventral view (6) and lateral view (7) of male terminalia of Periscelis annulata
(Fallén), (Nearctic, MND, figs. 77.4–5); lateral view of abdomen of female of (8) P. annulata, (Nearctic, MND, fig. 77.6); dorsal view, with
enlarged lateral view of anterior spiracle, of third instar larva of (9) P. annulata, (Nearctic, MND, fig. 77.7).
Abbreviations: cerc, cercus; ej apod, ejaculatory apodeme; epand, epandrium; hypd, hypandrium; pgt,  postgonite; ph,  phallus; spmth,
spermatheca; st, sternite; sur, surstylus; syntgst, syntergosternite; tg, tergite.
1090
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

in Hawaii, including pineapple (Ananas sativus Schultes, two to three posteroventral setae on apical half; scutellum
Bromeliaceae) and some Liliaceae and Poaceae. bearing a single pair of marginal setae, these apical; reclinate
fronto-orbital seta inserted behind proclinate fronto-orbital
seta; each tibia with a dorsoapical seta; hind femur with a
Classification subapical dorsal seta; anepisternum with one to two setae
along posterior margin.
The concept of Periscelididae, as adopted here, follows Papp (1984) proposed Stenomicridae for the genus
McAlpine (1978, 1983) and includes a few genera previously Stenomicra after McAlpine (1978, 1983) had transferred
assigned to Aulacigastridae (Cyamops, Planinasus, and that genus, together with Planinasus and Cyamops, from
Stenomicra). McAlpine characterized Periscelididae primarily the Aulacigastridae to the Periscelididae. Baptista  &
by the swollen, caplike pedicel, which has a dorsal cleft, and Mathis (1994) and Mathis & Papp (1998) recognized two
its caplike relationship to a sharply and ventrally deflexed subfamilies (Periscelidinae and Stenomicrinae) in the
first flagellomere. Although these characters are common Periscelididae, although the monophyly of only Periscelidinae
to all Periscelididae, they also occur in Neurochaetidae is well corroborated as follows: occiput with a silvery
(McAlpine, 1978; Woodley, 1982) and other acalyptrate white, microtomentose area immediately adjacent to the
genera. posterior margin of the compound eye; only one fronto-
The family to which Planinasus belongs has been contro- orbital seta, reclinate; costa short, extended only to R4+5;
versial. Although originally described in the family Ephydridae, CuA2 reduced or absent; several characters of the male
the genus was subsequently assigned to Drosophilidae terminalia (see Griffiths, 1972). The genera comprising
(Curran, 1934), Periscelididae (Malloch, 1934), Aulaci- Periscelidinae are those that Hennig (1969) included in
gastridae (Hennig, 1969), and more recently back to his more restricted concept of the family: Periscelis, Mar-
Periscelididae (McAlpine, 1983). McAlpine (1983: p.  56) benia, Neoscutops, Scutops, and Diopsosoma Malloch.
presented some evidence of a close phylogenetic relationship Baptista  & Mathis (1994) questioned the monophyly of
between Planinasus and Cyamops, pointing out that Plani- Stenomicrinae and presented evidence that Planinasus
nasus and the Australian species of Cyamops are the only might be more closely related to Periscelidinae. Griffiths
acalyptrate flies he has seen with branched hairs on the arista. (1972) considered the genera Diopsosoma and Somatia
Planinasus is distinct from other genera of Periscelididae, Schiner as Periscelididae. Mathis  & Papp (1992) and
and its monophyly is established by the following putative Grimaldi  & Mathis (1993), however, questioned this
synapomorphies: frons bearing a pair of interfrontal setae classification, and Mathis (1993) considered Somatia to
that are usually upright to slightly reclinate; forefemur with be related to the Psilidae (Diopsoinea).

Key to the genera of Periscelididae of the Neotropical Region

1. Two fronto-orbital setae present (Figs. 4–5); ocellar setae absent (Figs. 4–5); costa long, extended
to vein M; vein CuA2 usually well developed, usually with distinct cell cup (weak or lacking in
some Stenomicra); postpronotum lacking well-developed seta; STENOMICRINAE . . . . . . . . . 2
– One fronto-orbital seta present (Figs. 1–2); ocellar setae present; costa short (Fig. 3), extended to
vein R4+5; vein CuA2 weak or lacking, cell cup absent; postpronotum bearing well-developed seta;
PERISCELIDINAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

2. Frons with one pair of interfrontal setae; eye bare; katepisternum with two subequal setae; hind
femur bearing anterodorsal, preapical seta. . . . . . . . . . . . . . . . . . . . . . . . . . . . Planinasus Cresson
– Frons lacking interfrontal setae; eye microsetulose, sometimes sparsely; katepisternum with one
prominent seta; hind femur lacking anterodorsal, preapical seta. . . . . . . . . . . . . . . . . . . . . . . . . . 3

3. Fronto-orbital setae reclinate or occasionally mesoclinate, lacking proclinate seta (Fig. 5); inner
vertical seta present but with proclinate orientation (Fig. 5); face in profile angulate, dorsal surface
flattened; supra-alar seta lacking; with one pair of apicolateral scutellar setae; crossvein bm-cu
absent, making cells bm and dm confluent; vein CuA2 weak or lacking; cell cup lacking. . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stenomicra Coquillett
– Fronto-orbitals comprising one proclinate and one reclinate setae (Fig. 4); inner vertical seta
absent; face in profile shallowly and vertically arched, lacking flattened, dorsal area; supra-alar
seta present, well developed; lateral scutellar setae various but usually with two; crossvein bm-cu
well developed, cell bm distinct from dm; vein CuA2 present, well developed; cell cup present. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyamops Melander
PERISCELIDIDAE (periscelid flies) 83 1091

4. Eye borne on conspicuous stalk; antenna semiporrect; presutural supra-alar seta well developed;
anepisternum setose; scutellum triangular, apex pointed and bearing apical patch of setae; Peru .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Diopsosoma Malloch]
– Eye not borne on conspicuous stalk; antenna not porrect; lacking presutural seta; anepisternum
bare of setae; scutellar apex broadly rounded to truncate, lacking apical patch of setae. . . . . . . . 5

5. Vein R1 bearing numerous setulae along dorsum. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

– Vein R1 lacking setulae along dorsum. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

6. Vein R2+3 curved anteriorly toward costa to just beyond R1, thereafter reverse curved more or less
evenly, shallowly, and parallel to costa until merger with latter; apical section of vein M usually
conspicuously arched; with one posterior dorsocentral seta . . . . . . . . . . . . . . Neoscutops Malloch
– Vein R2+3 more or less evenly arched throughout length except just before apex; apical section of
vein M straight or shallowly arched; with two posterior dorsocentral setae. . . Marbenia Malloch

7. Dorsal area of face not distinctly flattened and shieldlike; wing various but usually mostly
hyaline; [Periscelis Loew]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
– Dorsal area of face distinctly flattened, shieldlike; wing with at least apical one-third
conspicuously infuscate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

8. Prescutellar acrostichal setulae moderately well developed, distinct from other acrostichal setulae;
crossvein dm-cu straight and well developed throughout length; male genitalia lacking digitiform
process at base of epandrium between surstylus and cercus; Holarctic, with amber fossils from the
Dominican Republic and Chiapas, Mexico. . . . . . . . . . . . . . . . . . . . . [Periscelis (Myodris) Lioy]
– Prescutellar acrostichal setulae undifferentiated; crossvein dm-cu weakened to completely attenuate
anteriorly, usually angulate or curved toward base, sometimes nearly straight; male genitalia with
ventrally oriented, narrow process at ventral margin of epandrium between surstylus and cercus;
Holarctic and Neotropical (Argentina). . . . . . . . . . . . . . . . . . . . . . . [Periscelis (Periscelis) Loew]

9. Wing irrorate (speckled); scutum dull, densely microtomentose, mostly gray, with several brown
spots; pair of distinct prescutellar acrostichal setae present; arista with three to four dorsal branches;
Mexico (DF, Mexico state). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Parascutops Mathis & Papp]
– Wing darkly infumate on apical one-half to one-third and with subapical, transverse, white area
or band; scutum thinly to moderately microtomentose, subshiny to shiny; prescutellar acrostichal
setae lacking; arista with six or more dorsal branches. . . . . . . . . . . . . . . . . . . . . Scutops Coquillett

Synopsis of the fauna Marbenia Malloch. Although Marbenia is presently

monotypic (M.  peculiaris Malloch) and has only been
Periscelididae includes 96 species that are placed in nine recorded from Peru (Iquitos), there is an undescribed species
genera. The New World fauna comprises nine genera and from Costa Rica.
54 species, including seven fossils in the genera Periscelis
(Myodris), Planinasus, and Stenomicra (Grimaldi & Mathis, Neoscutops Malloch. This genus includes three described
1993). There are approximately 35 species found in Central species from the New World tropics, and one of these,
America. N. rotundipennis Malloch, occurs in Costa Rica. There is a
second species from Costa Rica and Panama that is apparently
Cyamops Melander. Baptista  & Mathis (1996, 2000) undescribed.
last revised the New World species of Cyamops, recognizing
12  species (two species remain undescribed). The Central Planinasus Cresson. In a study that is still in progress
American fauna includes four species found primarily in (W.N. Mathis & A. Rung, in preparation), numerous additional
montane habitats, usually near aquatic environments (pools, species of Planinasus have been discovered, and the genus
streams, waterfalls). Adults, although rarely collected, are now includes one fossil and at least 16 extant species that
usually found on broad-leafed plants adjacent to aquatic occur in the New World tropics. Three species are found in
habitats. Central America. Specimens of Planinasus are generally
1092
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

rare in collections, and nothing is known about their immature species, P. electrus Grimaldi & Mathis (1993), from Dominican
stages or life cycle. Adults are collected by sweeping dense, amber.
understory vegetation, some bearing flowers, that is associated
with shaded, damp, riparian habitats. We found specimens Scutops Coquillett. Seven species, all from the New
to be relatively common on exposed sand to mud substrates World tropics, have been recorded (Prado, 1975), and two,
of shaded, riparian habitats, and numerous specimens were S.  chapmani (Curran) and S.  fascipennis Coquillett, are
captured alive by carefully lowering a vial over each one. We known from Central America, where the latter species is
widespread and relatively common.
also observed up to four species occurring sympatrically. Usu-
ally one species at these sites would predominate in number Stenomicra Coquillett. Although four species are known
of individuals. Several species exhibit considerable sexual from the New World, including three from Central America,
dimorphism, especially in the width and coloration of the there are over 100 that await description. Worldwide, species
face. Males in these species tend to have wider faces, i.e., of Stenomicra occur in most zoogeographic regions, but
larger facial ratios, and frequently there is a distinct color there is greater diversity in the tropics. Grimaldi & Mathis
pattern. The facial pattern usually involves microtomentum (1993) described two fossil species, S.  sabroskyi and
or its absence in additional to color. There is one fossil S. anacrostichalis, from Dominican amber.

Literature cited

Baptista, A.R. & W.N. Mathis. 1994. A revision of the New World of Palaearctic Diptera. Volume 3, Higher Brachycera. Science
species of the genus Cyamops Melander (Diptera: Periscelidi- Herald Publishers, Budapest, 880 pp.
dae). Smithsonian Contributions to Zoology 563: iv + 1–28. McAlpine, D.K. 1978. Description and biology of a new genus
Baptista, A.R.P. & W.N. Mathis. 1996. A new species of Cyamops of flies related to Anthoclusia and representing a new family
Melander (Diptera: Periscelididae) from Brazil, with distribu- (Diptera, Schizophora, Neurochaetidae). Annals of the Natal
tional notes on another species. Proceedings of the Entomologi- Museum 23: 273–295.
cal Society of Washington 98: 245–248. McAlpine, D.K. 1983. A new subfamily of Aulacigastridae (Dip-
Baptista, A.R.P. & W.N. Mathis. 2000. Notes on the genus Cyam- tera: Schizophora), with a discussion of aulacigastrid classifica-
ops Melander (Diptera: Periscelidididae), including description tion. Australian Journal of Zoology 31: 55–78.
of ten new species. Proceeding of the Entomological Society of Papp, L. 1984. Family Stenomicridae, pp.  61–62. In Soós, A.  &
Washington 102: 481–506. L. Papp (editors). Catalogue of Palaearctic Diptera, Volume 10.
Curran, C.H. 1934. The families and genera of North American Akadémiai Kiadó, Budapest, 402 pp.
Diptera. Published by the author, New York, 512 pp. Papp, L. 1995. Morphology of Periscelis annulata third instar larva
Griffiths, G.C.D. 1972. The phylogenetic classification of Diptera Cy- and Turanodinia tisciae larva and puparium (Diptera: Perisce-
clorrhapha, with special reference to the structure of the male post-
lididae and Odiniidae). Acta Zoologica Hungarica 41: 15–24.
abdomen. Series entomologica 8. Dr. W. Junk, The Hague, 340 pp.
Papp, L. 1998. Life-habits of the Central European species of
Grimaldi, D.A. & W.N. Mathis. 1993. Fossil Periscelididae (Dip-
Periscelididae (Diptera). Folia Entomologica Hungarica 59:
tera). Proceedings of the Entomological Society of Washington
119–123.
95: 383–403.
Prado, A.P. do. 1975. 67. Family Periscelididae. In Papavero,  N.
Hennig, W. 1969. Neue Gattungen und Arten der Acalyptratae. The
Canadian Entomologist 101: 589–633. (editor). A Catalogue of the Diptera of the Americas South of
Malloch, J.R. 1934. A remarkable new genus of the family Peris- the United States. Departamento de Zoologia, Secretaria da Ag-
celidae. Stylops 3: 52–53. ricultura, São Paulo, 3 pp.
Mathis, W.N. 1993. A new species and subgenus of Periscelis Loew Williams, F.X. 1939. Biological studies in Hawaiian water-loving
from Australia (Diptera: Periscelididae). Journal of the Austra- insects. Part  iii. Diptera or flies. B. Asteiidae, Syrphidae and
lian Entomological Society 32: 13–19. Dolichopodidae. Proceedings of the Hawaiian Entomological
Mathis, W.N. & L. Papp. 1992. A new genus of Periscelididae (Dip- Society 10: 281–315.
tera) from the Neotropics. Proceedings of the Biological Society Woodley, N.E. 1982. Two new species of Neurochaeta McAlpine
of Washington 105: 366–372. (Diptera: Neurochaetidae), with notes on cladistic relationships
Mathis, W.N. & L. Papp. 1998. Family Periscelididae, pp. 285–294. within the genus. Memoirs of the Entomological Society of
In Papp, L. & B. Darvas (editors). Contributions to a Manual Washington 10: 211–218.
ASTEIIDAE (asteiid flies) 84
Amnon Freidberg

Fig. 84.1. Female of Asteia beata Aldrich, (Nearctic, MND, fig. 78.1).

Diagnosis tergites; spiracle seven lacking. Male terminalia symmetric

in Leiomyza, asymmetric in most other taxa. Surstylus fused
Small (body length 1–5 mm), winged, generally weakly with epandrium. Phallus usually large and heavily sclero-
sclerotized flies (Fig. 1). Coloration varies from predominant- tized. Female terminalia simple, although segment 7 some-
ly yellow to predominantly black, but more or less elaborate times developed as an incomplete oviscape. Two sclerotized
and contrasting pattern usually present on some or all major spermathecae present.
body parts. Head rectangular or oval in lateral view, with Superficially, Asteiidae resemble some Anthomyzidae,
zero to two short to long fronto-orbital setae. Antenna short, Periscelididae Stenomicra Coquillett, and Drosophilidae,
with first flagellomere usually rounded. Arista usually well but these other flies do not have the combination of a con-
developed, with short to long alternating rays (microtrichia), cave M1, alternating rays on the arista, and male interfrontal
sometimes reduced or lacking. Proboscis short, with fleshy stripes.
labella. Males with interfrontal stripes. Thorax with one to
four dorsocentral setae; when four then anterior dorsocentral
seta presutural. Scutellum short, convex, usually with two
Biology
pairs of setae, basal seta much shorter than apical seta or
lacking. Acrostichal setulae zero to two rows, without setae. Little is known about the biology of asteiids and almost
Pleura without setae; anepisternum sometimes with setulae. nothing of the biology of Central American species. Phy-
Legs without overt features. Wing usually elongate, cover- tophagous (Asteia), fungivorus (Leiomyza), and sapropha-
ing abdomen and extending beyond it when at rest. Vein R2+3 gous Phlebosotera Duda species are known (Papp, 1998;
short, ending at tip of vein R1 or slightly more distally ex- Freidberg, 2002), and immatures belonging to these three
cept in Leiomyza Macquart, in which it reaches distal 0.75 of guilds have been described (Papp, 1998) or are being de-
wing. Vein M1 concave anteriorly, distally convergent with scribed (A. Freidberg, in preparation). Adult asteiids are gen-
vein  R4+5. Crossvein dm-cu present or lacking. Alula bare erally uncommon in collections, despite the fact that they
or with marginal cilia. Abdomen with weakly sclerotized may be abundant under certain natural circumstances, such
tergites, except in Leiomyza. Spiracles two to six situated in as on certain trees (e.g., Pinus, Tamarix, Quercus), herbaceous

1093
1094
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

vegetation, and in entrances to caves. Courtship and mat- Within the family, Sabrosky (1956) proposed the subfam-
ing are sometimes elaborate, including mating trophallaxis ilies Asteiinae (as Asteinae) and Sigaloessinae, and in the
(Freidberg, 1984). No economically important species of same paper gave the most comprehensive key to the world
Asteiidae are known. genera. Anarista Papp was subsequently added to the family
(Papp, 1972). Freidberg (2002) transferred Leiomyza from
Classification Sigaloessinae to the new subfamily Leiomyzinae and made
it the sole representative of this subfamily, whereas all other
This family is one of a number of groups whose relation- genera were included in Asteiinae. The latter subfamily will
ships have historically been obscure. Hennig (1973) treated be divided into two tribes, mostly corresponding to Sabro-
Asteiidae as part of his superfamily Anthomyzoidea, with sky’s subfamilies, although with the one exception noted
the family as part of the Periscelididea family group (with above (A.  Friedberg, in preparation). Based on cladis-
Periscelididae, Aulacigastridae, and Teratomyzidae). Griffiths tic analysis, Anarista, which shares characters of both
(1972) tentatively proposed a closer relationship of Asteiidae, tribes, should be assigned to Sigaloessini (Freidberg,
Opomyzidae, and Trixoscelididae (this last family considered 2002).
part of Heleomyzidae in this manual) based on the loss of the
seventh abdominal spiracles in both sexes. McAlpine (1989)
considered asteiids to be part of his superfamily Opomyzoidea, Identification
including some of the families in Hennig’s and Griffiths’
groups, in particular agreeing with Hennig in recognizing the The most recent identification resource for New World
relationship of Asteiidae, Periscelididae, Aulacigastridae, and species is the synopsis by Sabrosky (1957). This work is
Teratomyzidae in his suprafamily Asteioinea. The classifica- outdated, however, and much further work is needed on this
tion of the acalyptrate Diptera is still developing, however, family. Forrest  & Wheeler (2002) described some further
and future changes to these groups are likely. new species from the Galápagos Islands.

Key to the genera of Asteiidae of the Neotropical Region

1. Vein R2+3 relatively long, reaching costa at about distal 0.75 of wing (Fig.  2); epandrium
symmetrical; LEIOMYZINAE; Holarctic Region. . . . . . . . . . . . . . . . . . . . . [Leiomyza Macquart]
– Vein R2+3 relatively short, reaching costa at about middle of wing (Fig. 3); epandrium asymmetrical;
ASTEIINAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2. Crossvein dm-cu present; alula present, margin with cilia; SIGALOESSINI. . . . . . . . . . . . . . . . 3

– Crossvein dm-cu absent; alula absent, margin of alular region without cilia; ASTEIINI. . . . . . . 5

3. Anepisternum setulose; North America, Caribbean. . . . . . . . . . . . . . . . . . . . [Phlebosotera Duda]

– Anepisternum bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4. Face strongly narrowed between vibrissae, ventral margin projecting almost snoutlike between
vibrissal angles (Fig. 4); scutellum yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sigaloessa Loew
– Face not narrowed between vibrissae, ventral margin not projecting snoutlike between vibrissal
angles; scutellum black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tucumyia Sabrosky

5. Arista present; one pair of strong, erect, reclinate, fronto-orbital setae present; acrostichal setulae
usually absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Asteia Meigen
– Arista absent (Fig. 5); strong erect reclinate fronto-orbital setae absent; acrostichal setulae
present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Loewimyia Sabrosky
ASTEIIDAE (asteiid flies) 84 1095

Synopsis of the fauna from Panama. At least two species were recently found
in Costa Rica by the author, one by sweeping grass and
Approximately 12 species of this small family are found other herbaceous vegetation, the other by sweeping the
in Central America. underside of leaves of large banana plants, both on a Pacific
beach.
Asteia Meigen. This is the largest genus in the family,
containing over half of the family’s species. Only four species Sigaloessa Loew. This is essentially a New World genus,
are known from Central America, and two of these, A. anten- with one further species known from French Polynesia. Sev-
nata (Sabrosky) and A. spinosa (Sabrosky), were originally eral species are known from Central America, and the adults
described in Asteimyia Sabrosky, which was later synony- have been collected on various herbs and flowers, including
mized with Asteia (Sabrosky, 1956). Based on cladistic coastal vegetation.
analysis, it appears that Asteimyia should be resurrected
from synonymy. Tucumyia Sabrosky. This is a southern New World genus
of four species, of which T. liniaris Sabrosky was described
Loewimyia Sabrosky. This genus contains only four from Panama. An unidentified species was collected recently
described species, of which L. bifurcata Sabrosky is known in Costa Rica by the author.

R2+3 Sc R1 R2+3

R4+5
r-m
M1
cup CuA2 dm
dm-cu
A1+CuA2

CuA1

2 Leiomyza 3 Phlebosotera

4 Sigaloessa 5 Loewimyia

Figs.  84.2–5. Wings, head, and antenna: dorsal view of wing of (2)  Leiomyza laevigata (Meigen), (Holarctic, MND, fig.  78.7); and
(3) Phlebosotera setipalpis Sabrosky, (Nearctic, MND, fig. 78.6); anterolateral view of head of (4) Sigaloessa bicolor Loew, (Nearctic, MND,
fig. 78.3); lateral view of antenna of (5) Loewimyia sp., (Nearctic, MND, fig. 78.5).
1096
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Literature cited

Forrest, J. & T.A. Wheeler. 2002. Asteiidae (Diptera) of the Galápa- McAlpine, J.F. 1989. Phylogeny and classification of the Musco-
gos Islands, Ecuador. Studia dipterologica 9: 307–317. morpha, pp. 1397–1518. MND, Volume 3.
Freidberg, A. 1984. The mating behavior of Asteia elegantula with Papp, L. 1972. Entomological explorations in Ghana by Dr. S. En-
biological notes on some other Asteiidae. Entomologia Genera- drody-Yunga. 18. A new genus and species of Asteiidae from
lis 9: 217–224. Ghana (Diptera). Annales Historico-Naturales Musei Nationalis
Freidberg, A. 2002. Recent advances in the study of Asteiidae,
Hungarici 64: 319–321.
p. 73. In Yeates, D. (editor). 5th International Congress of Dip-
terology. Abstract Volume. The University of Queensland, Bris- Papp, L. 1998. Family Asteiidae, pp. 295–303. In Papp,  L.  &
bane, 283 pp. B. Darvas (editors). Contributions to a Manual of Palaearctic
Griffiths, G.C.D. 1972. The phylogenetic classification of Diptera Cy- Diptera. Volume 3. Science Herald, Budapest, 880 pp.
clorrhapha, with special reference to the structure of the male post- Sabrosky, C.W. 1956. Additions to the knowledge of Old World
abdomen. Series entomologica 8. Dr. W. Junk, The Hague, 340 pp. Asteiidae [Diptera]. Revue Française d’Entomologie 23:
Hennig, W. 1973. Ordnung Diptera (Zweiflügler), pp.  1–337. In 216–243.
Helmcke, J.-G., D. Starck, & H. Wermuth (editors). Handbuch Sabrosky, C.W. 1957. Synopsis of the New World species of the
der Zoologie. 4(2) 2/31 (Lfg. 20). Walter De Gruyter, Berlin & dipterous family Asteiidae. Annals of the Entomological Society
New York. of America 50: 43–61.
BRAULIDAE (beelice) 85
Brian V. Brown and F. Christian Thompson

Fig. 85.1. Female of Braula coeca Nitzsch, (MND, fig. 81.1).

Diagnosis suture and the visible pedicel of the antenna (in phorids the
pedicel is inserted into the first flagellomere). They differ
Minute flies, body length approximately 1.5 mm (about from sphaerocerids by having pectinate tarsal claws (Fig. 2)
double this size in an Oriental Region genus). Eye greatly re- and by the general structure of the legs and setation of the
duced to clear, depressed area without facets. Ocelli absent. body. Beelice are also confused with adult Varroa mites, be-
Ptilinal suture extremely prominent. Antennae widely sepa- ing similar in size and shape, but are separated by the num-
rated, placed in deep frontal pits. Arista unsegmented. Entire ber of legs.
body covered with stout, bristlelike setae (Fig. 1). Scutellum
absent. Mid- and hind legs, including coxae, widely sepa- Biology
rate. Tarsomeres extremely short, broad, with long lateral
setae. Claws modified, pectinate; pulvilli reduced, clavate. The natural history of this group was reviewed by
Wing and halter absent. Abdomen highly sclerotized, with Grimaldi & Underwood (1986) and Ferrar (1987); most in-
ventral sclerotization probably derived from tergites. formation exists for the cosmopolitan Braula coeca Nitzsch.
These flies resemble some wingless Phoridae and Sphaero- All species of Braula are associated with the honey bee,
ceridae. They differ from phorids by the prominent ptilinal Apis mellifera  (L.), which is an introduced species in the

1097
1098
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

New World. Adult flies cling to bees in the nest, occasion- Identification
ally climb to the oral region, and claw at the bee’s clypeus
to solicit regurgitation of food. Flies rarely leave the nest on The only reference for species identification of Braula is
foragers and prefer to cling to queens. Larvae (Fig. 3) form Örösi-Pal (1966); since this work, the only additional descrip-
mines in the comb and feed on the pollen, honey, and other
tion is that of B. hansruttneri Huttinger (1980). Unfortunately,
debris in the wax and perhaps the wax itself.
Although many individual beelice can be on a single Örösi-Pal’s types are apparently lost (Papp, 1998). The Cen-
queen bee, they apparently cause no economic damage. The tral American fauna so far consists of two species.
only impact is that the wax-lined larval tunnels spoil the ap-
pearance of honeycombs intended for show or sale (Dobson,
1999). Control measures for Varroa mites probably elimi- Synopsis of the fauna
nate Braula effectively.
Braula Nitzsch. Of the seven taxa (including two sub-
Classification species of B.  coeca Nitzsch) described, only B.  coeca and
B.  schmitzi Örösi-Pal are known from Central America
The classification of this family has changed frequently (Örösi-Pal, 1980). Örösi-Pal (1980) states that B.  schmitzi
through history. Most recently, McAlpine (1989) placed predominates in the Neotropical Region and gives charac-
Braulidae in the Carnoidea, but as he notes they have been ters to separate the two species. While some authors (e.g.,
placed differently by nearly every author who has studied McAlpine, 1989) believe that only one species, B.  coeca,
them. Use of molecular analysis would be highly interesting
exists worldwide, there is clear morphological and biologi-
in this situation.
There are two genera recognized in Braulidae: Braula, cal evidence to document that B. coeca and B. schmitzi are
which is found nearly worldwide with honey bee culture, distinct species. Dobson (1999) provides a comprehensive
and the two species of Megabraula Grimaldi & Underwood, overview of the literature and an excellent key with figures
found in Nepal associated with Apis laboriosa Smith. to distinguish these species.

clw

2 Braula
ant
mx plp
md

3 Braula

Figs. 85.2–3. Tarsus and larva: ventral view of midtarsus (2) of Braula coeca Nitzsch, (MND, fig. 81.2); lateral view of third instar larva (3)
of B. coeca, (MND, fig. 81.5).
Abbreviations: ant, antenna; clw, claw; md, mandible; mx plp, maxillary palpus.
BRAULIDAE (beelice) 85 1099

Literature cited

Dobson, J.R. 1999. A ‘bee-louse’ Braula schmitzi Örösi-Pal (Dip- Huttinger, E. 1980. Braula hansruttneri sp. n., eine neue Bienenlaus
tera: Braulidae) new to the British Isles, and the status of Brau- aus Österreich (Braulidae, Diptera). Entomofauna 1: 298–301.
la  spp. in England and Wales. British Journal of Entomology McAlpine, J.F. 1989. Phylogeny and classification of the Musco-
and Natural History 11: 139–148. (1998) morpha, pp. 1397–1518. MND, Volume 3.
Ferrar, P. 1987. A guide to the breeding habits and immature stages Örösi-Pal, Z. 1966. Die Bienenlaus-Arten. Angewante Parasitolo-
gie 7: 138–171.
of Diptera Cyclorrhapha. Entomonograph 8(1). E.J.  Brill  /
Örösi-Pal, Z. 1980. Bee lice in the Americas. American Bee Journal
Scandinavian Science Press, Leiden, Copenhagen, 478 pp. 1980: 438–440.
Grimaldi, D. & B.A. Underwood. 1986. Megabraula, a new genus Papp, L. 1998. 3.29. Family Braulidae, pp. 325–330. In Papp, L.
for two species of Braulidae (Diptera), and a discussion of brau- and B. Darvas (editors). Contributions to a Manual of Palaearc-
lid evolution. Systematic Entomology 11: 427–438. tic Diptera. Volume 3. Science Herald, Budapest, 880 pp.
1100
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
CARNIDAE (carnid flies) 86
Terry A. Wheeler

Fig. 85.1. Dealated female of Carnus hemapterus Nitzsch, (Holarctic, MND, fig. 80.1).

Diagnosis setulae various; two pairs of scutellar bristles; pleurites vari-

ous (Fig. 1). Wing (Figs. 3–4) usually well developed (bro-
Small (body length 1.0–3.0  mm) black or brown flies ken off at base in most specimens of Carnus), C with humeral
(Fig.  1). Frons with two lateroclinate upper fronto-orbital and subcostal breaks (subcostal break only in Neomeoneu-
bristles and two inclinate lower fronto-orbitals (one inclinate, rites), Sc complete but weak, joining C separately from R1,
three reclinate fronto-orbitals in Neomeoneurites Hennig); other venational characters (extent of C, length of cell dm,
postocellar bristles usually parallel or divergent; antennae presence of cell cup) various. Legs with bristles various but
in deep foveae separated by distinct facial carina (Fig.  2); usually weak, dorsal preapical tibial bristles absent. Abdomi-
gena usually broad and curved inward from ventral margin nal tergites well developed but small, sternites often reduced
of eye to oral margin, with one or two strong vibrissae and (absent in female Carnus), abdominal pleura with setae aris-
a row of long genal bristles; proboscis usually short, with ing from small sclerotized spots. Male genitalia often with
broad, swollen prementum (Fig.  2). Thoracic chaetotaxy: complex modifications of surstylus and subepandrial sclerite
one strong postpronotal (weaker postpronotal setae also or hypoproct (Fig. 5); phallus membranous, usually long and
present); two notopleurals; one presutural and one post- spinulose. Female postabdomen telescopic, cerci fused into
sutural supra-alar; one to four dorsocentrals, acrostichal single sclerotized projection (Fig. 6).

1101
1102
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

orb s
i vt s
r-m

fr s o vt s

3 Carnus
hum sc R1
vb brk brk
sbvb s R2+3

premnt r-m R4+5

lbl
dm
cup M1
dm-cu
2 Carnus A1+CuA2
CuA1
4 Hemeromyia
st 5 tg 5

hypd

syntgst 6-8
ph

ej apod sg 6

hypd arm

pgt
sg 7
epand

proc of hyprct

cerc

sur
sg 8

hyprct
cerc

5 Meoneura 6 Meoneura

Figs. 85.2–6. Head, wings, and terminalia: anterodorsal view of head (2) and dorsal view of wing (3) of Carnus hemapterus Nitzsch,
(Holarctic, MND, figs. 80.4, 80.3); dorsal view of wing of (4) Hemeromyia sp., (Nearctic, MND, fig. 80.2); lateral view of male terminalia (5)
and ventral view of female terminalia (6) of Meoneura obscurella (Fallén), (Holarctic, MND, figs. 80.8, 80.5).
Abbreviations: cerc, cercus; ej apod, ejaculatory apodeme; epand, epandrium; fr s, frontal seta; hum brk, humeral break; hypd, hypandrium;
hyprct, hypoproct; i vt s, inner vertical seta; lbl, labellum; orb s, orbital seta; o vt s, outer vertical seta; pgt, postgonite; ph, phallus; premnt,
prementum; proc, process; sbvb s, subvibrissal seta; sc brk, subcostal break; sg, segment; st, sternite; sur, surstylus; syntgst, syntergosternite;
tg, tergite; vb, vibrissa.
CARNIDAE (carnid flies) 86 1103

The combination of deep antennal foveae, arrangement of distinct family by most authors since then. The Australasian
the fronto-orbital bristles, parallel or divergent postocellars, genus Australimyza Harrison has been placed in Carnidae by
broad incurved gena, wing venation, and fused female cerci some authors (e.g., Colless & McAlpine, 1991), but morpho-
distinguishes carnids from other acalyptrate families. They logical characters do not support a close relationship, and the
are most easily confused with small, black Milichiidae or two groups are best treated as separate families (Buck, 2006;
Chloropidae, but may be distinguished from both families by Brake  & Mathis, 2007). Carnidae is apparently the sister
the above characters. group to Inbiomyiidae plus Australimyzidae (Buck, 2006).
Generic relationships within Carnidae are well resolved.
Neomeoneurites and the Baltic amber fossil genus Meoneu-
Biology rites Hennig are basal lineages (Wheeler, 1994; Grimaldi
1997; Buck  & Marshall, 2007). Of the remaining genera,
The only carnid genus for which biological information is Meoneura and Carnus are sister groups, with Enigmocar-
available in Central America is Carnus (see Synopsis of the nus Buck as the sister group to these two and Hemeromyia
fauna). Habits of species in other genera are based on records Coquillett as the sister group to these three (Buck, 2006;
from other geographic regions. Larvae of Carnidae are sap- Buck & Marshall, 2007).
rophagous in a range of substrates rich in decaying organic Of the five extant genera, Meoneura is by far the largest,
material. Adults are usually collected in bird or mammal
with approximately 70 described species, mostly in the
nests and burrows, on decaying vegetation, dung, carrion,
Holarctic Region (Sabrosky, 1987; Barraclough, 1994; Papp,
mud, or shoreline debris (Ferrar, 1987). The most specialized
1998). There are 10 described species of Hemeromyia
habits are found in species of the genus Carnus, in which
(Holarctic and southern Africa) (Sabrosky, 1987; Barra-
larvae and adults are almost entirely restricted to bird nests.
clough, 1994; Papp, 1998), five described species of Carnus
The larvae are apparently saprophagous, feeding on the nest
(Holarctic, Oriental, Afrotropical) (Grimaldi, 1997), Neome-
material, but the adults attach themselves to nestling birds
and feed on them, although there is still some disagreement oneurites has two described species restricted to Patagonia
as to whether they feed preferentially on blood or dermal (Wheeler, 1994), and Enigmocarnus is known from a single
secretions (Grimaldi, 1997). None of the species have a southern Nearctic species (Buck & Marshall, 2007).
significant economic impact. Carnus species are primarily
restricted to cavity-nesting wild birds and appear to have a Identification
negligible or minor effect on nestling growth. On rare occa-
sions species of Meoneura build up high numbers in dung Central American genera of Carnidae were previously
or, less commonly, in stored grain (Ferrar, 1987), but this is keyed by Sabrosky (1987). Identification of the genera relies
a sporadic and minor nuisance. on external characters, especially the wing venation and
externally visible abdominal characters, so identification to
Classification that level does not usually present a problem with specimens
in good condition. Because of their small size, adult specimens
The Carnidae were previously considered a subfamily of should be preserved in 70% ethanol and critical-point dried
Milichiidae (e.g., Sabrosky, 1959) but have been treated as a or chemically dried soon afterwards.

Key to the genera of Carnidae of Central America

1. Wing dehiscent, usually broken off at base, when present (rarely), crossvein dm-cu absent (Fig. 3);
abdominal sternites small or absent; abdomen (especially of female) often greatly swollen (Fig. 1);
adults usually restricted to bird nests. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carnus Nitzsch
– Wing complete, crossvein dm-cu present (Fig.  4); abdominal sternites present, usually well
developed; abdomen not greatly swollen; adults found in a variety of habitats . . . . . . . . . . . . . . 2

2. Wing with C extending to R4+5, crossveins r-m and dm-cu close together in basal third of wing;
ultimate section of CuA1 longer than penultimate section; M1 beyond dm-cu much weaker than
other veins (as in Fig. 3). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Meoneura Rondani
– Wing with C extending to M1, crossveins r-m and dm-cu more widely separated; dm-cu in distal
half of wing; ultimate section of CuA1 less then half as long as penultimate section; M1 well
developed beyond dm-cu (Fig. 4); northern Mexico. . . . . . . . . . . . . . . . [Hemeromyia Coquillett]
1104
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Synopsis of the fauna Because of their close association with bird nests, specimens
of Carnus species are rarely collected by entomologists, and
There are no published records of described species of more species and host records may await discovery in
Carnidae from Central America, and there are few specimens Central America.
in museum collections. Although collecting in appropriate
habitats such as bird nests or arid coastal or saline areas Meoneura Rondani. This is the only genus of Carnidae
might be expected to document additional species, the rarity that tends to be found in high numbers and in a wide range
in collections of even the diverse, generalist genus Meoneura of habitats. Nevertheless, specimens are rarely collected in
from Central America suggests that overall diversity of Carnidae Mexico and Central America. The only Neotropical records
in the region is lower than that in the Nearctic Region, are one described species (M.  sinclairi Wheeler) in the
probably in the range of 15–20 species. Galápagos Islands, Ecuador, and a few specimens of uniden-
tified species from Mexico (Morelos), Venezuela, and Peru
Carnus Nitzsch. Adults and larvae of Carnus are almost (Wheeler, 2000). The presence of a widespread native spe-
always collected in bird nests, usually cavity nests. Newly cies of Meoneura throughout the Galápagos Islands almost
emerged adult flies are fully winged and capable of dispersing certainly means that additional species are (or were) present
and colonizing a new host nest, after which the wings are on the Pacific coast of Central America or northern South
broken off. There are four described Nearctic species, including America (Wheeler, 2000). In addition to the records of ex-
one in Mexico (Durango) (Grimaldi, 1997); the only records tant species, Grimaldi (1997) described a fossil species of
from southern Mexico or Central America are females of Meoneura in Miocene Dominican amber. Sabrosky (1959)
two undescribed species from Chiapas, Mexico, collected provided a key to the Nearctic species of Meoneura, includ-
from the nests of an American Kestrel (Falco sparverius L., ing Mexican records, but there are many undescribed species,
Falconidae) and an unidentified owl (Grimaldi, 1997). and his key should be used with caution.

Literature cited

Barraclough, D.A. 1994. Hemeromyia australis sp. n. and Meoneura Grimaldi, D.A. 1997. The bird flies, genus Carnus: species revision,
prima (Becker, 1903) from Namibia and South Africa, the first generic relationships, and a fossil Meoneura in amber (Diptera:
Carnidae (Diptera: Schizophora) recorded from subequatorial Carnidae). American Museum Novitates 3190: 1–30.
Africa. Annals of the Natal Museum 35: 15–24. Papp, L. 1998. Family Carnidae, pp. 211–217. In Papp,  L.  &
Brake, I. & W.N. Mathis. 2007. Revision of the genus Australimyza B. Darvas (editors). Contributions to a Manual of Palaearctic
Harrison (Diptera: Australimyzidae). Systematic Entomology Diptera. Volume 3, Higher Brachycera. Science Herald Publishers,
32: 252–275. Budapest, 880 pp.
Buck, M. 2006. A new family and genus of acalypterate flies from
Sabrosky, C.W. 1959. The Nearctic species of the filth fly genus
the Neotropical region, with a phylogenetic analysis of Carnoidea
family relationships (Diptera, Schizophora). Systematic Entomology Meoneura (Diptera, Milichiidae). Annals of the Entomological
31: 377–404. Society of America 52: 17–26.
Buck, M.  & S.A. Marshall. 2007. Enigmocarnus chloropiformis Sabrosky, C.W. 1987. Carnidae, pp. 909–912. MND,
gen. et sp. nov., and parallel evolution of protandrial symmetry Volume 2.
in Carnidae (Diptera). Annals of the Entomological Society of Wheeler, T.A. 1994. A new species of Neomeoneurites Hennig
America 100: 9–18. (Diptera: Carnidae) from Argentina, with a discussion of
Colless, D.H.  & D.K. McAlpine. 1991. Diptera, pp.  717–786. In male postabdominal structure in the genus. The Canadian
Division of Entomology, CSIRO. The Insects of Australia, Entomologist 126: 435–441.
Volume  II. Melbourne University Press, Melbourne, pp.  i–vi, Wheeler, T.A. 2000. Carnidae of the Galapagos Islands, Ecuador:
543–1137.
description and phylogenetic relationships of a new species
Ferrar, P. 1987. A guide to the breeding habits and immature stages
of Diptera Cyclorrhapha. Entomonograph 8(1). E.J.  Brill  / of Neotropical Meoneura Rondani, 1856 (Diptera: Carnidae).
Scandinavian Science Press, Leiden, Copenhagenm, 478pp. Studia dipterologica 7: 115–120.
INBIOMYIIDAE (inbiomyiid flies) 87
Matthias Buck

Fig. 87.1. Female of Inbiomyia mcalpineorum Buck, (Buck, 2006, fig. 1).

Diagnosis bristles absent. Face bare, almost flat, with weakly developed,
rounded longitudinal median ridge. Vibrissal angle evenly
Small (body length 1.3–1.6  mm), mostly dark flies of rounded, gena narrow. Antennae porrect, of moderate size,
stocky, humpbacked build (Fig.  1). Head (Fig.  2) broad, broadly separated. First flagellomere spherical, small. Arista
short; eyes protruding. Frons approximately half as wide as dorsoapical, extremely long, with long pubescence. Palpus
head, ocellar plate large, occupying most of frons. Head bris- with conspicuous preapical ventral bristle. Labella short, tri-
tles include one pair of long inclinate fronto-orbitals, inner angular, with tapered apex, separate from each other. Thorax
and outer verticals, and vibrissae. Ocellar and postocellar with one presutural and two postsutural dorsocentrals; one

1105
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

i vt s Sc R1
frorb s
o vt s R2+3

R4+5

M1

A1+CuA2
CuA1

3 Inbiomyia

2 Inbiomyia

4 Inbiomyia

mp phapod

hypd

epand

5 Inbiomyia

pgt

tg 6 epand clft
distph

epiph
tg 8
spmth
sur

v proc epand

6 Inbiomyia

7 Inbiomyia

Figs. 87.2–7. Head, wing, forefemur, egg, and terminalia: anterior view of head (2), dorsal view of wing (3), and posterior view of
forefemur (4) of Inbiomyia mcalpineorum Buck, (Buck, 2006, figs. 3A, E, D); dorsal view of egg of (5) I. zeugodonta Buck, (South America,
Buck  & Marshall, 2006, fig.  12); lateral view of male terminalia  (6) and dorsal view of female terminalia and spermathecae  (7) of
I. mcalpineorum, (Buck, 2006, figs. 6B, 7A).
Abbreviations: distph, distiphallus; epand, epandrium; epand clft, epandrial cleft; epiph, epiphallus; frorb s, fronto-orbital seta; hypd,
hypandrium; i vt s, inner vertical seta; mp, micropyle; o vt s, outer vertical seta; pgt, postgonite; phapod, phallapodeme; spmth, spermathecae;
sur, surstylus; tg, tergite; v proc epand, ventral process of epandrium.
INBIOMYIIDAE (inbiomyiid flies) 87 1107

postpronotal, one presutural, and one postsutural supra-alar; raised above the surface. The micropyle is slightly projecting
and one postalar and two notopleural bristles. Scutellum bare, at the apex of the egg in dorsoventral view. The eggs of the
with two or four marginal bristles. Katepisternum setulose, three examined species show little interspecific variation.
with dorsal bristle, pleuron otherwise bare. Legs slender. Inbiomyia eggs are strikingly similar to macrotype
Forefemur (Fig. 4) with antero- and posteroventral bristles. Tachinidae eggs. In Tachinidae, this egg morphology is as-
Tibiae lacking preapical or ventroapical bristles. Wing sociated with a parasitoid lifestyle, where eggs are attached
unmarked (Fig. 3). Costa extending to apex of M, with sub- to the host’s cuticle. Future studies will have to investigate
costal and humeral break. Subcosta incomplete. Epandrium the possibility that Inbiomyia larvae parasitize arthropods in-
with slender, ventral, surstylus-like process (Fig. 6). Phallus habiting decaying plant matter.
with long phallapodeme; distiphallus short, membranous, Adult feeding has not been observed, but gut contents of
and tubular; epiphallus well developed. Female terminalia numerous specimens showed fungal matter, algae (including
extremely truncate (Fig. 7); sternite 8, segments 9, 10, and
diatoms), and probably bacteria. The proboscis shows no
cerci absent. Two sclerotized spermathecae.
obvious modifications for specialized feeding habits.
Inbiomyiidae can easily be distinguished from other Aca-
lyptratae by their extremely shortened frons, head chaeto-
taxy (only one pair of inclinate fronto-orbital bristles, ocellar Classification
bristles absent, vibrissa present, palpus with strong ventral
preapical bristle), and porrect antenna with extremely long, The family Inbiomyiidae has only recently been recog-
dorsoapical arista. nized (Buck, 2006; Buck & Marshall, 2006) and was erected
for the single Neotropical genus Inbiomyia. Based on the
presence of an inclinate fronto-orbital bristle, simple pedicel
Biology (without dorsal seam or apical notch), incomplete subcosta,
two costal breaks, simple and microtrichose distiphallus, two
Little is known about the biology of Inbiomyia. Adults spermathecae, and the absence of preapical tibial bristles, the
are most frequently collected in Malaise traps in tropical for- family is best placed in the superfamily Carnoidea (sensu
ests from sea level to about 2000 m altitude. Two species, McAlpine, 1989). An analysis of Carnoidea relationships
I. mcalpineorum Buck and I. exul Buck, have been collected (Buck, 2006) places the Inbiomyiidae as the sister group
in tree falls by sweeping and pan trapping. Recently fallen of Australimyzidae, a family restricted to the Australasian
trees are important breeding microhabitats for various fam- Region. The highly autapomorphic Inbiomyiidae have di-
ilies of saprophagous Acalyptratae (e.g., Sphaeroceridae, verged considerably from Australimyzidae, which preserve
Drosophilidae). However, the peculiar egg morphology and many character states plesiomorphic for Carnoidea. Austra-
the low number of mature eggs in gravid females points to- limyzidae have been considered part of Carnidae by some
wards a non-saprophagous lifestyle. authors (Colless & McAlpine, 1991; Grimaldi, 1997). How-
The immature stages of Inbiomyia are unknown except ever, there are no synapomorphies linking Australimyzidae
for eggs obtained by dissection from gravid females (Buck, to Carnidae with the exclusion of Inbiomyiidae. Carnidae is
2006; Buck & Marshall, 2006). The number of mature eggs the putative sister group of Inbiomyiidae + Australimyzidae
(Buck, 2006).
per female varies from one to four. Eggs (Fig.  5) are rela-
tively large (length 0.39–0.48  mm, width 0.21–0.27  mm),
elliptical in outline, and extremely flattened dorsoventrally Identification
(thickness ~0.05 mm). The chorion is reticulated and unpig-
mented on the dorsal surface of the egg and is membranous The eleven described species of Inbiomyia can be identi-
and transparent on the ventral surface. The reticulation of the fied with the key provided by Buck & Marshall (2006). The
upper side is polygonal (predominantly hexagonal) and not species groups are distinguished as follows.

Key to the species groups of Inbiomyiidae (males)

1. Hypandrium with long, basomedial, ventrally directed, Y-shaped process; postgonites absent; surstylus
slightly tapered apically, with one stout, short, toothlike bristle at apex. . . . . . . . . . . I. exul group
– Hypandrium simple; postgonites well developed; surstylus slightly to greatly expanded apically,
with two toothlike bristles or none. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1108
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

2. Expanded apical portion of surstylus with several scattered discal bristles, apex usually with two
stout, short, toothlike bristles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. scoliostylus group
– Expanded apical portion of surstylus with at most one discal bristle, other bristles arranged in
more or less regular, marginal, or submarginal row (row sometimes duplicated near apex), apex
without toothlike bristles (Fig. 6). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3. Surstylus moderately expanded at apex (Fig. 6); ventral process of epandrium curved medially,
articulated to epandrium; apex of postgonite with marginal row of at least eight toothlike, short
bristles and no long hairs (Fig. 6). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. mcalpineorum group
– Surstylus greatly expanded at apex; ventral process of epandrium more or less straight, fused to
epandrium; postgonite with fewer (about four) toothlike marginal bristles (apical one sometimes
enlarged), and usually with some longer hairs; South America. . . . . . . . . . . [I. anemosyris group]

Synopsis of the fauna Ecuador and one undescribed species (known from the
female only) from Guatemala. The I.  scoliostylus group
Five species of this family are found in Central America. is represented in Costa Rica by I.  scoliostylus Buck;
two other species occur in Colombia. The I.  mcalpine-
Inbiomyia Buck. The genus is distributed throughout the orum group includes two closely related species, one of
New World tropics (but apparently absent from the Carib- which, I.  mcalpineorum, is known from Costa Rica, the
bean), with records from Guatemala, Honduras, Costa Rica, other one from Ecuador. One undescribed species from
Colombia, Venezuela, French Guiana, Ecuador, Peru, and Honduras (known from a single female) might also be-
Bolivia. A total of eleven species is described (Buck, 2006; long in this group. Unlike species of the other three groups,
Buck and Marshall, 2006), but undoubtedly more species which are restricted to tropical lowland forests, the I. mcalpi-
will be discovered in the future. The I. exul group includes neorum group is mainly distributed in the cloud forest
one described species, I.  exul Buck, from Costa Rica and zone (560–2000 m).

Literature cited

Buck, M. 2006. A new family and genus of acalypterate flies from second edition. Melbourne University Press & Cornell Univer-
the Neotropical Region, with a phylogenetic analysis of Car- sity Press, Ithaca, 1137 pp.
noidea family relationships (Diptera, Schizophora). Systematic Grimaldi, D. 1997. The bird flies, genus Carnus: species revision,
Entomology 31: 377–404. generic relationships, and a fossil Meoneura in amber (Diptera:
Buck, M. & S.A. Marshall. 2006. Revision of the Neotropical fam- Carnidae). American Museum Novitates 3190: 1–30.
ily Inbiomyiidae (Diptera, Schizophora). Contributions in Sci- McAlpine, J.F. 1989. Phylogeny and classification of the Musco-
ence 512: 1–30.
morpha, pp. 1397–1518. MND, Volume 3.
Colless, D.H.  & D.K. McAlpine. 1991. Diptera, pp.  717–786. In
Division of Entomology, CSIRO. The insects of Australia,
TETHINIDAE (beach flies) 88
Wayne N. Mathis

Fig. 88.1. Male of Tethina parvula Loew, (Nearctic, MND, fig. 101.1).

Diagnosis spermathecae; cercus subcylindrical or compressed, some-

times bearing strong, spinelike setulae (pseudacanthophorites);
Small to medium-sized flies (body length 1.6–4.0 mm); fre- tergites 7 and 8 mostly with characteristic pigmented areas;
quently invested with pale yellowish to brown microtomen- epiproct generally small, bearing pair of setulae dorsally on
tum. With one to four orbital and zero to four fronto-orbital apical third; hypoproct large.
setae (Figs. 1–7); postocellar seta convergent or absent. Face Adult Tethinidae are similar to some Heleomyzidae
sometimes characterized by two shiny protuberances laterad (Trixoscelidinae), Canacidae, and Chloropidae but are dis-
of facial cavity above vibrissal pore (Tethina); vibrissal seta tinguished by lacking well-developed dorsally curved genal
various, if present usually weak. With one proepisternal seta; setae, by having a single costal break (Sc), by having a com-
one proepimeral seta; anepisternum with one or more setae plete, although small, cell cup, and by lacking a preapical
and some setulae posteriorly. Precoxal bridge present. Wing dorsal seta on the tibiae.
(Figs. 8–11) hyaline to infuscate or pale yellow; C with Sc
break only; cell cup present but small; A1 weakened apically, Biology
not reaching wing margin. Tibiae lacking preapical dorsal
seta. Male terminalia (Figs.  12–13) with epandrium bear- Tethinidae are mostly halophiles/thalassophiles, occur-
ing two surstyli ventrally (partially fused with epandrium in ring in coastal marine habitats, although some species are
Tethina); anterior surstylus distinctive; posterior surstylus found inland, usually associated with saline biotopes, such
generally strongly setulose and lobelike; phallapodeme long, as salt lakes and alkaline hot springs. A few species, such as
slender; ejaculatory apodeme usually large; phallus usually Pelomyia coronata (Loew), are exceptions, being associated
elongate, ribbonlike, sinuous, subcylindrical, with a more or with meadows that occur in mountain passes, forests, and
less dense ventral pubescence, often with several microscopic oases (Melander, 1952: p.  194). Adults of thalassophilous
papillae. Female terminalia (Fig.  14) with two sclerotized species are commonly found in coastal marine habitats

1109
1110
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

2 Tethina 3 Neopelomyia 4 Pelomyiella

i vt s
orb s
oc s o vt s

5 Dasyrhicnoessa 6 Pelomyia 7 Masoniella

Sc R1
R2+3

R4+5

M1

CuA1

8 Masoniella 9 Tethina

10 Pelomyia 11 Pelomyiella

Figs. 88.2–11. Heads and wings: lateral view of head of (2) Tethina horripilans (Melander), (Nearctic, MND, fig. 101.3); (3) Neopelomyia
rostrata (Hendel), (Nearctic, MND, fig. 101.4); (4) Pelomyiella mallochi (Sturtevant), (Holarctic, MND, fig. 101.7); (5) Dasyrhicnoessa
insularis (Aldrich), (cosmopolitan, Foster and Mathis, 1998, fig. 3); (6) Pelomyia coronata (Loew), (MND, fig. 101.5); and (7) Masoniella
richardsi Vockeroth, (Nearctic, MND, fig. 101.2); dorsal view of wing of (8) M. richardsi, (Nearctic, MND, fig. 101.9); (9) T. horripilans,
(Nearctic, MND, fig. 101.10); (10) Pelomyia. coronata, (MND, fig. 101.11); and (11) Pelomyiella mallochi, (Holarctic, MND, fig. 101.12).
Abbreviations: i vt s, inner vertical seta; oc s, ocellar seta; orb s, orbital seta; o vt s, outer vertical seta.
TETHINIDAE (beach flies) 88 1111

phapod

tg 6
ph

ej apod
tg 6

syntgst 7+8

hypd
syntgst
7+8
hypd
pregt
ph

pregt
pgt

i lb bac scl
pgt of epand
epand
bac scl vl lb
sur of epand

sur
cerc

cerc

12 Masoniella 13 Masoniella

tg 8 epiprct

tg 7

hyprct cerc

st 7 st 8

14 Masoniella

Figs. 88.12–14. Male and female terminalia: ventral view (12) and lateral view (13) of male terminalia of Masoniella richardsi Vockeroth,
(Nearctic, MND, figs. 101.13–14); lateral view of female terminalia of (14) M. richardsi, (MND, fig. 101.8).
Abbreviations: bac scl, bacilliform sclerite; cerc, cercus; ej  apod, ejaculatory apodeme; epand, epandrium; epiprct, epiproct; hypd,
hypandrium; hyprct, hypoproct; i lb of epand, inner lobe of epandrium; pgt, postgonite; ph, phallus; phapod, phallapodeme; pregt, pregonite;
st, sternite; sur, surstylus; syntgst, syntergosternite; tg, tergite; vl lb of epand, ventrolateral lobe of epandrium.
1112
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

(Karl, 1930), including the intertidal zone, wrack heaps be considered tentative (1976: p. 337), “...as much data are
(usually brown algae that are most abundant along temperate still to be obtained on certain character complexes, such as
seashores bathed by cold currents), salt marshes, dune veg- the male genitalia and the immature stages.” Foster proposed
etation, lagoon–littoriparian zones, mangroves (particularly two monophyletic subfamilies, Tethininae and Pelomyiinae,
species of Dasyrhicnoessa), and on salty soils or bare sand. with Pelomyia, Pelomyiella, and Neopelomyia in the latter
Two species of Apetaenus, A. littoreus (Hutton) and A. wat- and all other Nearctic species in the genus Tethina as part of
soni Hardy, occur on seabird dung and are associated with Tethininae.
colonies of penguins and other seabirds. Some species occur Foster & Mathis (2002) revised Masoniella and Pelomyia
in habitats that man has dramatically and usually adversely and included a cladistic analysis of the subfamily Pelomyii-
modified, such as meadows polluted by industrial emissions nae. Synapomorphies that demonstrate the monophyly of
Pelomyiinae are as follows: (1)  crossvein bm-cu gener-
(Bährmann, 1982) or slaughterhouses and poultry farms
ally absent; (2)  surstylus shifted dorsally; (3)  gena setose;
(Zuska & Laštovka, 1969). These synanthropic habitats are
(4)  acrostichal setae reduced or absent; and (5)  internal
usually sites of salt accumulation and enrichment.
copulatory apparatus complex (well-developed pregonites;
The biology and immature stages of the family are incom-
postgonites may be bilobed, dentate, or fused). Genera of
pletely known. Hardy  & Delfinado (1980) reared Dasyrh-
Pelomyiinae are divided into two clades: the first comprises
icnoessa vockerothi Hardy  & Delfinado from deposits of
Pelomyiella and Neopelomyia, and the second, which is the
seaweed on beaches in Hawaii, and Ferrar (1987) provided
sister group to the first clade, includes Masoniella and Pelo-
some observations on the puparia of Tethina grisea (Fallén).
myia. The latter two genera are closely related and similar
Séguy (1940) and Hardy (1962) described the larva of two
to each other, and Foster & Mathis (2002) considered com-
species of the subantarctic genus Apetaenus Eaton. Gorczyt-
bining them. With evidence that the two clades are mono-
za (1988) reported on the spatial and seasonal distribution of
phyletic, however, they advocated keeping them separate,
some European species, Pelomyiella mallochi (Sturtevant),
especially as they are usually easily distinguished and rec-
Tethina albosetulosa (Strobl), T.  illota Haliday, T.  flavi-
ognizable in the field. With the exception of Pelomyiella,
genis (Hendel), and T.  grisea (Fallén), from a study using
which is Holarctic in distribution, the genera of Pelomyiinae
color traps on the Frisian Islands of Mellum and Memmert.
are found primarily in the New World. The genera Masoniella
Tréhen & Vernon (1982) and Tréhen et al. (1985) conducted and Neopelomyia occur exclusively in the New World, as does
extensive ecological investigations on the genera Apetaenus
Pelomyia with the exception of P. occidentalis Williston, which
and Listriomastax Enderlein on the Crozet Islands. In nature, is apparently adventive in central Europe (Roháček, 1992).
an abundance of individuals and a paucity of species usually A comprehensive phylogenetic analysis of Tethinidae is
characterize sites where tethinids occur. now being investigated (G.A.  Foster, personal communica-
Rossi (1988) described a species of Laboulbeniales (As- tion), especially at the tribal and subfamilial levels. Placement
comycetes) that is parasitic on Pseudorhicnoessa rattii Munari of Masoniella (Vockeroth, 1987, 1995) within the existent
from the Seychelles. classification, for example, has cast uncertainty on the rela-
tionships between the subfamilies Tethininae and Pelomyii-
Classification nae. Moreover, the relationships between the subfamilies
with single or few included genera, such as Horaismopterinae,
The phylogenetic relationships of the Tethinidae, either Apetaeninae, and Zaleinae, are attractive for phylogenetic
with other families (outgroups, sister groups) or among the analysis and invite further consideration.
included taxa, are not entirely resolved, and further clarifi-
cation will require cladistic analysis at all taxonomic levels Identification
(see “Classification” section under Canacidae, Chapter 89).
At the subfamilial and generic levels, Foster (1976) pro- Foster  & Mathis (1998) should be adequate to identify
posed a hypothesis of phylogenetic relationships that remains most species of Tethinidae from Central America. If speci-
the only analysis available. Foster’s hypothesis primarily mens of Pelomyia and/or Masoniella are found in Central
concerned Nearctic taxa (Tethina illota, from the Palearctic America, the keys, illustrations, and descriptions in Foster &
Region, was also included), and he suggested that his analysis Mathis (2002) should be consulted.

Key to the genera of Tethinidae of the New World

1. Gena bare (Fig. 2); peristoma bearing row of strong, dorsally curved setae; frons with two rows of
interfrontal setulae, often setalike; cell bm and discal cell distinct; crossvein bm present (Fig. 9);
prescutellar acrostichal setae present; scutum bearing numerous rows of coarse setae arising from
punctures; postpronotum with three or more setae, with ventral seta dorsoclinate. . . . . . . . . . . . 2
– Gena with scattered setulae (Figs. 3–7); peristoma bearing row of weak setulae; frons bare or with few
weak and scattered setulae; cell bm and discal cell confluent, crossvein bm absent (Figs. 8, 10–11);
prescutellar acrostichal setae absent; scutum smooth, at most with few scattered setulae; postpronotum
with one or two setae, lacking dorsoclinate ventral seta; PELOMYIINAE, in part . . . . . . . . . . . . . . 4
TETHINIDAE (beach flies) 88 1113

2. Gena about one-eighth eye height (Fig.7) (except M.  advena Foster  & Mathis: 0.25); two
well-developed fronto-ortibal setae present; cells bm and dm confluent; thorax sometimes shiny;
abdomen shiny (except M. argentinaensis Foster & Mathis); PELOMYIINAE, in part; California,
USA. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Masoniella Vockeroth]
– Gena about one-third eye height; three to five fronto-orbital setae present; cells bm and dm separate;
thorax and abdomen usually microtomentose, mostly appearing dull; TETHININAE. . . . . . . . . 3

3. Eye generally densely covered with small, pale, interfacetal setulae; true vibrissal seta present
(Fig. 5); shiny tubercle above foremost strong subvibrissal seta lacking; scutellar disc bare except
for marginal setae; ctenidium of forefemur present in most species; epandrium with two pairs of
surstyli; anterior surstylus varied in shape but not curved, hooklike. . . . . Dasyrhicnoessa Hendel
– Eye appearing bare (Figs. 1–2), setulae sparse or lacking; vibrissal seta absent on apex of vibrissal
angle; vibrissal apex occupied by shiny tubercle (sometimes as a scarcely visible fold); scutellar
disc bare, shiny; ctenidium of forefemur lacking; epandrium with one pair of surstyli, partially
fused with epandrium. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tethina Haliday

4. Paravertical setae proclinate; lacking forefemoral comb of setae along anteroventral surface;
subcranial and parafacial margins with at most remnant of shiny, chitinous stripe. . . . . . . . . . . . 5
– Paravertical setae convergent; forefemur usually bearing anteroventral ctenidial comb; subcranial
and parafacial margins with distinct, shiny, chitinous stripe; Holarctic Region . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Pelomyia Williston]

5. Head elongate, longer than high (Fig. 3); eye oblique; ventral portion of face protruding; three
fronto-orbital setae present; oral vibrissae stronger; distinct subvibrissal setae absent; labellum
elongate; palpus with many fine setae; west coast of North America. . . . . [Neopelomyia Hendel]
– Head at least as high as long (Fig. 4); eye round; face vertical; one to two fronto-orbital setae present;
oral vibrissae weak to absent; subvibrissal setae distinct; labellum short; palpus with only one
weak seta; nearly worldwide, including southern Nearctic Region. . . . . . . . [Pelomyiella Hendel]

Synopsis of the fauna this species from the New World (as D. ferruginea (Lamb)),
and Foster & Mathis (1998) provided detailed locality and
The family now comprises 166 valid species that are descriptive data for the species. We first discovered D. insu-
placed in 17 genera (Mathis & Munari, 1996; Munari, 2004, laris in Central America on barrier islands off the coast of
plus updates). No species of fossil Tethinidae have been de- Belize and more recently at the western margin of the Carib-
scribed (Evenhuis, 1994), although undetermined specimens bean in Costa Rica (Puerto Limón). We have found this spe-
have been recorded but not verified from the Oligocene/ cies in the USA (Florida), Mexico (Tabasco), on the Lesser
Miocene amber of Chiapas, Mexico (Hurd et al., 1962; Poi- Antilles (Dominica, St. Lucia, St. Vincent), and Bermuda in
nar, 1992). I estimate about 12 species of this family occur the western North Atlantic. Elsewhere, the genus occurs pri-
in Central America. marily within the Pacific and Indian Ocean basins, where 14
species have been described (Mathis & Munari, 1996).
Dasyrhicnoessa Hendel. This genus is distinctive, espe-
cially from other genera of the subfamily Tethininae, and is Tethina Haliday. Among genera of Tethinidae, Tethina is
easily distinguished by the densely setulose eyes, prominent by far the most species rich worldwide, with almost half (72
oral vibrissal seta (Fig. 5), and an anterior surstylar-like lobe species) of the 166 described species currently recognized
and a posterior (true) surstylus in males. In the New World, in the family (Mathis & Munari, 1996, plus updates). Nine
a single species, D.  insularis (Aldrich), is known and was species of Tethina are known from Central America (Fos-
probably introduced through human commerce. Woodley & ter & Mathis, 1998), and they occur on both the Atlantic and
Hilburn (1994) and Mathis & Munari (1996) first recorded Pacific maritime coasts.
1114
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Literature cited

Bährmann, R. 1982. Zur Vorkommen sogenannter halophile Munari, L. 2004. Beach flies (Diptera: Tethinidae: Tethininae) from
Dipteren-Arten in einer industriell belasteten Immissiongebiet. Australia and Papua New Guinea, with descriptions of two new
Entomologische Nachrichten Berichte 26: 75–78. genera and ten new species. Records of the Australian Museum
Evenhuis, N.L. 1994. Catalogue of fossil flies of the world (Insecta: 56: 29–56.
Diptera). Backhuys Publishers, Leiden, 600 pp. Poinar, G.O., Jr. 1992. Life in Amber. Stanford University Press,
Ferrar, P. 1987. A guide to the breeding habits and immature stages Stanford, California, xiii + 350 pp.
of Diptera Cyclorrhapha. Entomonograph 8(1). E.J.  Brill  / Roháček, J. 1992. Tethinidae (Diptera) of Czechoslovakia: A fau-
Scandinavian Science Press, Leiden, Copenhagen, 478 pp. nistic survey. Časopis Slezského zemského muzea, Opava (A)
Foster, G.A. 1976. Notes on the phylogeny of the Nearctic Tethini- 41: 127–131.
dae and a review of the genus Neopelomyia Hendel, and the Rossi, W. 1988. New or interesting Laboulbeniales (Ascomycetes)
Tethina milichioides group (Diptera). Proceedings of the Ento- parasitic on Diptera. Webbia 42: 171–178.
mological Society of Washington 78: 336–352. Séguy, E. 1940. IV. Diptères, pp. 203–267. In Jeannel, R. (editor)
Foster, G.A. & W.N. Mathis. 1998. A revision of the family Tethin- Croisière du Bougainville aux îles australes françaises. Mémoi-
idae (Diptera) from the Caribbean, Gulf of Mexico, and Ber- res du Muséum national d’Histoire naturelle, new series 14:
muda. Proceedings of the Entomological Society of Washington 203–267.
100: 601–632. Tréhen, P. & P. Vernon. 1982. Peuplement diptérologique d’une île
Foster, G.A. & W.N. Mathis. 2002. A revision of the genera Pelo- subantarctique: la Possession (46°S, 51°E; Îles Crozet). Revue
myia Williston and Masoniella Vockeroth (Diptera: Tethinidae). d’Écologie et de Biologie du Sol 19: 105–120.
Smithsonian Contributions to Zoology 619: 1–63. Tréhen, P., M. Bouché, P. Vernon, & Y. Frenot. 1985. Organization
Gorczytza, H. 1988. Die Tethiniden der Nordseeinseln Mellum und and dynamics of Oligochaeta and Diptera on Possession Island,
Memmert (Diptera: Tethinidae). Drosera 1988: 303–310. pp.  606–613. In Siegfried,  W.R., P.R.  Condy,  & R.M.  Laws
Hardy, D.E. 1962. Insects of Macquarie Island. Diptera: Coelopi- (editors). Antarctic Nutrient Cycles and Food Webs. Springer-
dae. Pacific Insects 4: 963–971. Verlag, Berlin, 700 pp.
Hardy, D.E. & M.D. Delfinado. 1980. Tethinidae, pp. 369–379. In Vockeroth, J.R. 1987. Tethinidae, pp.  1073–1078. MND,
Hardy, D.E. & M.D. Delfinado (editors). Insects of Hawaii 13: Volume 2.
369–379. Diptera: Cyclorrhapha III. University Press of Hawaii, Vockeroth, J.R. 1995. Validation of nomina nuda of Nearctic
Honolulu, 451 pp. Tethinidae, Scathophagidae, and Muscidae proposed in Manual
Hurd, P.D., Jr., R.F. Smith, & J.W. Durham. 1962. The fossiliferous
of Nearctic Diptera. Proceedings of the Entomological Society
amber of Chiapas, México. Ciencia 21: 107–118.
Karl, O. 1930. XI. Thalassobionte und thalassophile Diptera of Washington 97: 732–734.
Brachycera. Die Tierwelt der Nord- und Ostsee 19: 33–84. Woodley, N.E.  & D.J. Hilburn. 1994. The Diptera of Bermuda.
Mathis, W.N.  & L. Munari. 1996. World catalog of the family Contributions of the American Entomological Institute 28: ii +
Tethinidae (Diptera). Smithsonian Contributions to Zoology 1–64.
584: iv + 1–27. Zuska, J.  & P. Laštovka. 1969. Species-composition of the
Melander, A.L. 1952. The North American species of Tethinidae dipterous fauna in various types of food-processing plants
(Diptera). Journal of the New York Entomological Society 59: in Czechoslovakia. Acta entomologica bohemoslovaca 66:
187–212. 201–221.
CANACIDAE (surf flies) 89
Wayne N. Mathis

Fig. 89.1. Female of Canacea macateei (Malloch), (MND, fig. 102.1).

Diagnosis Eggs simple, ovoid; with microscopic reticulations.

Third-instar larvae 5–6  mm (Fig.  7); tapered anteriorly
Adults minute to small flies (body length 1.6–5.0  mm); and posteriorly from about 4th abdominal segment and ter-
blackish, brownish, yellowish, or gray, often invested with minated posteriorly in slender retractable respiratory tube.
whitish to grayish microtomentum. Head (Figs. 1–2, 4) in- Abdominal segments 2–7 with creeping welts. Prothoracic
cluding subcranial cavity large; three to five lateroclinate spiracle a slender retractable filament. Posterior spiracles
(Fig.  9) with three oval spiracular openings arranged with
fronto-orbital setae present. Face slightly convex to con-
longitudinal axis at slightly less than right angles to adja-
cave; setae usually sparse except for inclinate vibrissal seta;
cent opening; each spiracular plate with four tufts of inter-
clypeus prominent, wide. Gena high, bearing one to four spiracular setae. Cephalopharyngeal skeleton (Fig.  8) with
dorsoclinate genal setae. Scutum with four or more dorsoce- ventral cornu truncate, appearing broken at apical margin;
ntral setae. Wing (Fig. 6) usually hyaline; C extended to M mandibles approximate anteriorly, separated posteriorly by
and with subcostal break only; Sc complete, separate from small V-shaped accessory oral sclerite; anterior ventrolateral
R1 almost to its apex; cells br, bm, dm, and cup complete; A1 extensions of tentoropharyngeal sclerite narrowly fused with
short. Precoxal bridge present. Male tergites 1–6 exposed; ventral bridge of hypopharynx; parastomal bars prominent,
spiracles 1–6 in posteroventral portion of tergite, spiracle 7 united by thin fenestrated epipharyngeal sclerite.
also in tergite 6; terminalia (Fig. 3) symmetrical; surstylus Puparia (Fig. 10) brown, similar in size and form to third-
fused with epandrium; hypandrium usually with lateral arms instar larva, spindle-shaped, curved at each end; integu-
extended above aedeagus, fused into posteriorly directed mental spinules more prominent than on larva and anterior
process; cercus usually weak. Female cercus (Fig. 5) long, respiratory processes fully extended.
approximate, bearing strong apical seta, sometimes preceded Adult canacids are similar to, and sometimes confused
by similar but smaller setae; ventral wall of genital chamber with, shore flies (Ephydridae). Canacids are distinguished by
with V- or ring-shaped sclerite; with two spermathacae. the wing venation (cells bm and cup complete) and by the

1115
1116
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

tg 6 syntgst 7+8

epand
infr s

cerc

phapod
hypd spn

gn s
pgt
vb
sbvb s
pregt

ph
hypd
sur
2 Canaceoides
3 Canacea

tg 7 tg 8

epiprct

cerc

4 Nocticanace st 7 st 8 hyprct

5 Canacea
Sc R1
R2+3

R4+5

M1

CuA1
6 Canacea

Figs. 89.2–6. Heads, terminalia, and wing: anterodorsal view of head of (2) Canaceoides nudatus (Cresson), (MND, fig. 102.2); lateral view
of male terminalia of (3) Canacea macateei (Malloch), (MND, fig. 102.3); anterodorsal view of head of (4) Nocticanace arnaudi Wirth,
(Nearctic, MND, fig.  102.4); lateral view of female terminalia of (5)  Canacea  macateei, (MND, fig.  102.5); dorsal view of wing of
(6) Canacea macateei, (MND, fig. 102.6).
Abbreviations: cerc, cercus; epand, epandrium; epiprct, epiproct; gn s, genal seta; hypd, hypandrium; hypd spn, hypandrial spine; hyprct,
hypoproct; infr s, interfrontal seta; pgt, postgonite; ph, phallus; phapod, phallapodeme; pregt, pregonite; sbvb s, subvibrissal seta; st, sternite;
sur, surstylus; syntgst, syntergosternite; tg, tergite; vb, vibrissa.
CANACIDAE (surf flies) 89 1117

number of abdominal segments (five in ephydrid males, six (Oriental and Afrotropical Regions) is found in freshwater
in canacids), which in females terminate as an elongate and environments (usually mountain streams), and on Hawaii,
fused epiproct + cercus that bears enlarged, apical setae. members of the Procanace nigroviridis group also occur in
freshwater habitats. Although poorly known, the larvae and
adults are probably grazers on algae or are saprophytic.
Biology
In the New World, only the larva and puparium of Cana- Classification
cea macateei Malloch are known. Teskey & Valiela (1977)
successfully reared this species from larvae and puparia that The phylogenetic relationships of the Canacidae, either
were collected on mats of bluegreen algae at Cape Cod, with other families (outgroups, sister groups) or among the
Massachusetts. Larvae of C. macateei, along with those of included taxa, are not entirely resolved, and further clarifica-
Dolichopodidae, were the principal contributors to the bio- tion will require cladistic analysis at all taxonomic levels. At
mass of invertebrates in the algal mats. Puparia were quite the familial level, J.F. McAlpine (1989: p. 1472) identified
common, and the puparial stage is brief, perhaps no more five synapomorphies that link Canacidae with Tethinidae and
than 2–3  days. Adults were observed to aggregate, espe- noted that “There are clear indications of a sister-group rela-
cially at night, on the tops of grass-covered dunes. tionship between them ... and may even indicate that they are
All canacids from the New World occur in intertidal subgroups of a single family.” Other authors (Hennig, 1958;
habitats. In the Old World, Procanace grisescens Hendel Griffiths, 1972; D.K. McAlpine, 1982; Freidberg, 1995) have

prn pd

7 Canacea

d corn

ant
pastm b

mx plp

md

den scl lab scl hyphar scl v corn

8 Canacea 9 Canacea

10 Canacea
Figs. 89.7–10. Larval and pupal structures of Canacea macateei (Malloch), (MND, figs. 102.7–10): lateral view of third instar larva (7);
lateral view of cephalopharyngeal skeleton of third instar larva (8); posterior view of posterior spiracles of third instar larva (9) and lateral
view of puparium (10).
Abbreviations: ant, antenna; d corn, dorsal cornu; den  scl, dental sclerite; hyphar scl, hypopharyngeal sclerite; lab  scl, labial sclerite;
md, mandible; mx plp, maxillary palpus; pastm b, parastomal bar; prn pd, perianal pad; v corn, ventral cornu.
1118
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

also suggested a relationship with the family Tethinidae, and Paracanace. Procanace, the fifth New World genus, was
and Griffiths (1972) further noted some affinities with the initially placed in Nocticanacinae, but it is now evident that
Chloropidae and Milichiidae. According to J.F. McAlpine’s it is a more basal lineage to both subfamilies (i.e., the sister
(1989) cladogram, which included an analysis of 25 charac- group of the remainder of the family), but no formal family-
ters for the families Canacidae and Tethinidae, the Canaci- group category has been proposed, pending phylogenetic
dae, together with Australimyzidae, Braulidae, Carnidae, studies of the Canacidae–Tethinidae complex (W.  Mathis,
Chloropidae (including Minidae and Siphonellopsidae), unpublished data).
Cryptochetidae, Milichiidae, Risidae, and Tethinidae, com-
prise the superfamily Carnoidea (= Chloropoidea). Identification
Freidberg (1995) reviewed the phylogenetic status of Zalei-
nae as part of his description of Suffomyia Freidberg. Although Although Wirth (1951) reviewed the entire family, his
clearly associating Suffomyia with Zaleinae within the Canaci- paper is of limited value now, not including even half of
dae–Tethinidae complex, Freidberg concluded that Zaleinae is
the described fauna. Mathis (1982) revised the subfamily
an intermediate taxon between Canacidae and Tethinidae and
Canacinae, which is primarily Old World, and provided keys
further suggested that all three taxa be recognized as a single
family, with Canacidae as the oldest family-group name. to all included genera and species. Mathis’ paper for some
It is evident that the Canacidae and Tethinidae are closely regions has been supplanted (Afrotropical Region, Mathis &
related and perhaps should be combined into a single family, Freidberg, 1991; Australian Region, Mathis, 1996, 1999),
and there are ongoing studies to resolve the relationships and and these papers also treat species of the subfamily Nocti-
classification. For now, however, the concept of Canacidae is canacinae. Mathis (1992) also published a catalog of all
as proposed in regional catalogs and manuals during the last species and included references to keys and illustrations for
50 years. all species for which these are available. In the New World,
Mathis (1982) proposed a classification for the Canacidae Mathis (1989, 1997) recently reviewed the surf fly fauna for
that includes two subfamilies, Canacinae and Nocticanacinae, the Caribbean and Gulf of Mexico, and the included keys
and within the subfamily Canacinae, two tribes, Dynomiel- and illustrations should be adequate to identify most spe-
lini and Canacini. The Canacinae are represented in the New cies occurring in Mesoamerica. The Neotropical fauna has
World fauna by a single genus, Canacea, which belongs to numerous undescribed species, especially in the subfamily
the tribe Dynomiellini. The Nocticanacinae are represented by Nocticanacinae, and undoubtedly there are many others still
three genera in the New World, Canaceoides, Nocticanace, to be collected.

Key to the genera of Canacidae of the New World

1. With four or more lateroclinate fronto-orbital (Fig. 1); forefemur bearing row of usually four to
five spinelike setae along apical one-half of anteroventral surface; apical one-third of arista bare
(Fig. 1); 2 supra-alar setae present (Fig. 1) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Canacea Cresson
– With three lateroclinate fronto-orbital setae (Figs. 2, 4); forefemur lacking row of spinelike setae;
arista evenly haired throughout length (Figs. 2, 4); one supra-alar seta present. . . . . . . . . . . . . . 2

2. Intrafrontal setae absent, although anterior one-third of frons occasionally with scattered setulae;
eastern North America, West Indies, Brazil. . . . . . . . . . . . . . . . . . . . . . . . . . . [Procanace Hendel]
– One or more intrafrontal setae present in addition to any setulae. . . . . . . . . . . . . . . . . . . . . . . . . 3

3. Two intrafrontal setae present; postocellar setae well developed, proclinate, and slightly
divergent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paracanace Mathis & Wirth
– One intrafrontal seta present (Figs. 2, 4); postocellar setae either much reduced or absent. . . . . 4

4. Three large, dorsoclinate genal setae present (Fig.  2); disc of scutellum usually bearing 2–15
setulae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Canaceoides Cresson
– Two large, dorsoclinate genal setae present (Fig. 4); disc of scutellum lacking setulae. . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nocticanace Malloch
CANACIDAE (surf flies) 89 1119

Synopsis of the fauna species, and four occur in Central America. Three of the four
Central American species belong to the N.  texensis group,
Worldwide, the Canacidae now include 118 valid species which Mathis (1989) revised. The fourth species, N. chilen-
that are placed in 12  genera (Wirth, 1951; Mathis, 1992). sis Cresson, has been recorded from Panama and belongs
The New World fauna comprises five genera and 35 spe- to a species group that has numerous species in temperate
cies (Wirth, 1965, 1975, 1987; Mathis, 1992). No fossils are South America, including many that are undescribed. On the
known. Galápagos Islands, Wirth (1969b) recorded eight apparently
endemic species.
Canacea Cresson. This is a New World genus with four
species that are primarily tropical in distribution, although Canaceoides Cresson. This genus is closely related and
C. macateei occurs northward along the east coast of North similar to Nocticanace and Paracanace. Wirth (1969a) re-
America as far as the maritime provinces of Canada. Ma- vised the genus, which now includes nine species. Seven
this (1982, 1992) last reviewed the genus. Three species are species are known from Central America (Mathis, 1992),
found in Central America, but only C. macateei is relatively and the fauna of Baja California is especially diverse with
common. In Belize, adults of C. macateei were collected by six species, including four that are apparently endemic.
sweeping just above mats of bluegreen algae that occurred
along the mud banks of brackish pools (Mathis, 1989). The Paracanace Mathis  & Wirth. Like Canacea, all de-
surface of the mud banks, which was covered by water only scribed species of Paracanace occur in the New World
at higher tides, was broken by deep cracks that had divided and have primarily tropical or subtropical distributions
the bank into irregularly shaped mud plates that were 15– (Mathis & Wirth, 1978; Mathis, 1992). There are two spe-
25 cm in diameter. The algal mats were comparatively thick, cies groups in Paracanace (Mathis & Wirth, 1978), and all
and as a protective covering, they helped keep the underlying known Central American species belong to the P.  hoguei
mud moist. The species of Canacea are similar externally group, which includes three species (Mathis, 1997). Speci-
but can be distinguished by structures of the male terminalia mens of Paracanace from Florida were collected on a
and the position, number, and color of certain setae. causeway between the Gulf Coast of Florida and Sanibel
Island. The sides of the causeway were partially lined with
Nocticanace Malloch. This is the most species-rich ge- broken pieces of concrete and large rocks, and much of the
nus of surf flies (34 species) and has greatest species diver- surface of the concrete and rocks was covered with algae
sity in the Old World (Mathis, 1992). The New World has 13 where the flies were found.

Literature cited

Freidberg. A. 1995. A study of Zaleinae, a taxon transitional between Mathis, W.N. & W.W. Wirth. 1978. A new genus near Canaceoides
Canacidae and Tethinidae (Diptera), with the description of a new Cresson, three new species and notes on their classification
genus and species. Entomologica scandinavica 26: 447–457. (Diptera: Canacidae). Proceedings of the Entomological Society
Griffiths, G.C.D. 1972. The phylogenetic classification of Diptera Cy- of Washington 80: 524–537.
clorrhapha, with special reference to the structure of the male post- McAlpine, D.K. 1982. A new genus of Australian littoral flies (Dip-
abdomen. Series entomologica 8. Dr. W. Junk, The Hague, 340 pp. tera: ?  Canacidae). Memoirs of the Entomological Society of
Hennig, W. 1958. Die Familien der Diptera Schizophora und ihre Washington 10: 108–117.
phylogenetischen Verwandtschaftsbeziehungen. Beiträge zur McAlpine, J.F. 1989. 116. Phylogeny and classification of the
Entomologie 8: 505–688. Muscomorpha, pp. 1397–1518. MND, Volume 3.
Mathis, W.N. 1982. Studies of Canacidae (Diptera), I: suprageneric Teskey, H.J. & I. Valiela. 1977. The mature larva and puparium of
revision of the family, with revisions of new tribe Dynomiellini Canace macateei (Diptera: Canaceidae). The Canadian Ento-
and new genus Isocanace. Smithsonian Contributions to Zool- mologist 109: 345–347.
ogy 347: 1–29. Wirth, W.W. 1951. A revision of the dipterous family Canaceidae. Oc-
Mathis, W.N. 1989. A review of the beach flies of the Caribbean casional Papers of the Bernice P. Bishop Museum 20: 245–275.
and Gulf of Mexico (Diptera: Canacidae). Proceedings of the Wirth, W.W. 1965. Family Canaceidae, pp. 733–734. In Stone, A.,
Biological Society of Washington 102: 590–608. C.W. Sabrosky, W.W. Wirth, R.H. Foote, & J.R. Coulson (edi-
Mathis, W.N. 1992. World catalog of the beach-fly family Canacidae tors). A catalog of the Diptera of America north of Mexico.
(Diptera). Smithsonian Contributions to Zoology 536: iv + 1–18. United States Department of Agriculture, Agriculture Handbook
Mathis, W.N. 1996. Australian beach flies (Diptera: Canacidae). Pro- 276, Washington, D.C., iv + 1696 pp.
ceedings of the Biological Society of Washington 102: 326–348. Wirth, W.W. 1969a. The shore flies of the genus Canaceioides
Mathis, W.N. 1997. A review of the hoguei group of Paracanace Cresson (Diptera: Canaceidae). Proceedings of the California
Mathis and Wirth (Diptera: Canacidae). Memoirs of the Ento- Academy of Sciences (Fourth Series) 36: 551–570.
mological Society of Washington 18: 140–148. Wirth, W.W. 1969b. New species and records of Galápagos Diptera.
Mathis, W.N. 1999. A review of the beach-fly genus Isocanace Proceedings of the California Academy of Sciences (Fourth
Mathis (Diptera: Canacidae). Proceedings of the Entomological Series) 36: 571–594.
Society of Washington 101: 347–358. Wirth, W.W. 1975. 76. Family Canaceidae. In Papavero, N. (editor).
Mathis, W.N.  & A. Freidberg. 1991. Review of the Afrotropical A Catalogue of the Diptera of the Americas South of the United
beach flies of the tribe Canacini and subfamily Nocticanacinae States. Departamento de Zoologia, Secretaria da Agricultura,
(Diptera: Canacidae). Proceedings of the Entomological Society São Paulo, 5 pp.
of Washington 93: 70–85. Wirth, W.W. 1987. 102. Canacidae, pp. 1079–1083. MND, Volume 2.
1120
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
CRYPTOCHETIDAE (cryptochetid flies) 90
J.F. McAlpine and F. Christian Thompson

Fig. 90.1. Female of Cryptochetum iceryae (Williston), (Australian, introduced to Nearctic, MND, fig. 100.1).

Diagnosis spinelike ovipositor (Figs. 7–9), with one weakly sclerotized

spermatheca.
Small (body length 2–4 mm) stout-bodied flies (Fig. 1), Eggs about 0.2  mm long and less than 0.1  mm wide,
metallic bluish-black with all bristles absent or scarely kidney shaped.
distinguishable; head as broad as thorax and twice as high as Mature larvae (Fig.  10) 1.8–3.2  mm long, ovoid,
wide; frons nearly square in both sexes; eyes large, occupy- yellowish-white, tapered to anterior end, with 10 clearly
ing most of side of head; first flagellomere large, without marked segments behind head, with each segment bearing
arista, only with small apical sensory spine; wing short, wide, belt of minute spines; terminal segment with pair of long
about twice as long as broad, with strong anal angle, with tails (6–9  mm); anterior spiracle retractile into body, with
costagial, humeral, and subcostal breaks, with cells bm and 5–12 elongate digits. Posterior spiracles forming pair of
dm confluent, and cell cup small, incomplete; legs stocky,
posteroventrally curved hooks situated on heavily pigmented
setulose, lacking outstanding bristles; abdomen short, broad
(Figs.  6, 7), with six well-developed terga in both sexes, plates on posterodorsal surface of segment 10.
with last tergum considerably longer than preceding ones; Cryptochetids are easily recognized by their short, stout
male with elongate phallus (Figs. 2–6); female with curved, bodies, metallic color, and lack of arista.

1121
1122
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

hypd
phapod ej apod hypd
ej apod ph ph gd phapod

epand
pregt
pgt

sur
cerc pgt pregt

ph
ph

2 Cryptochetum 3 Cryptochetum 4 Cryptochetum

syntg 1+2

hypd

tg 3

phapod

tg 4

pregt
tg 5

pgt
tg 6

ph
tg 6 st 7
epand
cerc

5 Cryptochetum 6 Cryptochetum st 8
tg 8
prct

st 7

7 Cryptochetum

st 7

st 8 tg 8

cerc

hyprct

8 Cryptochetum 9 Cryptochetum 10 Cryptochetum

Figs. 90.2–10. Terminalia, abdomens, and mature larva: ventral view (2) and lateral view (3) of male terminalia, lateral view (4) and ven-
tral view (5) of hypandrium and phallus, lateral view of male abdomen (6), ventral view of female abdomen (7), with offset of enlarged
ventral view of sternite  8, and lateral view, withdrawn  (8) and extended  (9), of ovipositor of Cryptochetum iceryae (Williston),
(Australian, introduced to Nearctic, MND, figs. 100.2–9); lateral view of third instar larva of (10) C. yokohama Kuwana, (Palearctic, MND,
fig. 100.10).
Abbreviations: cerc, cercus; ej apod, ejaculatory apodeme; epand, epandrium; hypd, hypandrium; hyprct, hypoproct; pgt,  postgonite;
ph,  phallus; phapod, phallapodeme; ph  gd, phallic guide; prct, proctiger; pregt, pregonite; st,  sternite; sur,  surstylus; syntg, syntergite;
tg, tergite.
CRYPTOCHETIDAE (cryptochetid flies) 90 1123

Biology separate family for it. Various authors (Hennig, 1958, 1969,
1973; Griffiths, 1972; D.K. McAlpine, 1976; J.F. McAlpine,
Larvae are endoparasitic on monophlebine scale insects. 1989) have discussed the relationships of the family, with
Their biology and economic importance is described in current consensus being that cryptochetids are most closely
detail by Thorpe (1931). Our single New World species, related to Milichiidae and Chloropidae.
Cryptochetum iceryae (Williston), was introduced from
Australia for the control of cottony cushion scale (Icerya Identification
purchasi Maskell, Homoptera: Margaroididae), a serious
pest of citrus. The last comprehensive key to the species of Cryp-
tochetum includes only 16 species (Thorpe, 1941). Ghesquières
Classification (1943) treats the Afrotropical species, to which Bruggen
(1960) added, with revised subgeneric classification. Stey-
The family consists of Cryptochetum, with four poorly skal (1971) provides a key to the species of subgenus Trito-
differentiated subgenera (Bruggen, 1960). Two other lestes Ghesquiere.
genera, Phanerochaetum Hennig (Hennig, 1965) from
Baltic amber (one  species, P.  tuxeni Hennig) and Librella Synopsis of the fauna
McAlpine (D.K.  McAlpine, 1976) from Australia (one
species, L.  demetrius McAlpine) were originally referred
to Cryptochetidae by their authors, but both genera have Cryptochetum. This genus is native to the southern
typical elongate aristae. For this and many other reasons, Palearctic, Afrotropical, Oriental, and Australian Regions.
J.F. McAlpine (1989) excluded them from the family. In the past, Of the 29 described species, one species, Cryptochetum
Cryptochetum has been referred to many families, including iceryae, has been introduced widely for the biological
Agromyzidae, Carnidae, Chamaemyiidae, Drosophilidae, control of cottony cushion scale. This species is known from
Milichiidae, and Chloropidae. Brues  & Melander (1932) USA, Chile, and Peru. Its status in Central America is
resolved the uncertain placement of the genus by making a unknown, although it is expected to occur there.

Literature cited

Brues, C.T. & A.L. Melander. 1932. Key to the families of North Hennig. W. 1973. Ordnung Diptera (Zweiflügler), pp. 1–337. In
American insects. Brues & Melander, Boston & Pullman, vii + Helmcke, J.-G., D. Starck, & H. Wermuth (editors). Handbuch
140 pp. der Zoologie. 4(2) 2/31 (Lfg. 20). Walter De Gruyter, Berlin &
Bruggen, A.C. van. 1960. A new subgenus and four new species of
New York.
Cryptochaetum Rondani (Diptera: Cryptochaetidae) from South
Africa. Journal of the Entomological Society of southern Africa McAlpine, D.K. 1976. A new genus of flies possibly referable to
23: 286–295. Cryptochetidae (Diptera, Schizophora). Australian Entomological
Ghesquières, J. 1943. Recherches sur les Dipteres d’Afrique. II. Magazine 3: 45–56.
Notice monographique sur les Muscoïdes Cryptochaetidae, McAlpine, J.F. 1989. Phylogeny and classification of the
parasites de Coccides Monophlebinae. Revue de Zoologie et de Muscomorpha, pp. 1397–1518. MND, Volume 3.
Botanique Africaines 36: 390–410. (1942) Steyskal, G.C. 1971. The subgenus Tritolestes Ghesquiere of the
Griffiths, G.C.D. 1972. The phylogenetic classification of Diptera
genus Cryptochetum Rondani with a new species from Pakistan
Cyclorrhapha, with special reference to the structure of the male
postabdomen. Series entomologica 8. Dr. W. Junk, The Hague, (Diptera: Cryptochetidae). Proceedings of the Entomological
340 pp. Society of Washington 73: 48–51.
Hennig, W. 1958. Die Familien der Diptera Schizophora und ihre Thorpe, W.H. 1931. The biology, post-embryonic development,
phylogenetischen Verwandtschaftsbeziehungen. Beiträge zur and economic importance of Cryptochaetum iceryae (Dip-
Entomologie 8: 505–688. tera, Agromyzidae) parasitic on Icerya purchasi (Coccidae,
Hennig, W. 1965. Die Acalyptratae des Baltischen Bernsteins und Monophlebini). Proceedings of the Zoological Society of
ihre Bedeutung für die Erforschung der phylogenetischen London 1930: 929–971.
Entwicklung dieser Dipteren-Gruppe. Stuttgarter Beiträge zur Thorpe, W.H. 1941. A description of six new species of the genus
Naturkunde 145: 1–215.
Cryptochaetum (Diptera—Agromyzidae) from East Africa and
Hennig. W. 1969. Neue Ubersicht über die aus dem Baltischen
Bernstein bekannten Acalyptratae (Diptera: Brachycera). East Indies; together with a key to the adults of larvae of all
Stuttgarter Beiträge zur Naturkunde 209: 1–42. known species. Parasitology 33: 131–148.
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MILICHIIDAE (milichiid flies) 91
John E. Swann

Fig. 91.1. Female of Desmometopum m-nigrum (Zetterstedt), (cosmopolitan, MND, fig. 79.1).

Diagnosis Biology
Small (body length 1–7  mm) acalyptrate flies (Fig.  1), Although the biology of the majority of species is
often largely brown or black, occasionally orange or yellow. unknown, many milichiids have general saprophagous
Males of some genera with silvery microtomentum on or coprophagous larvae (Sabrosky, 1977, 1987a; Papp  &
abdomen. Usually with two or three pairs of medioclinate Wheeler, 1998; Brake, 2000; Swann, 2000). Ferrar (1987)
frontal bristles and two or three pairs of lateroclinate to provides a good overview of the larval habits, noting they are
proclinate orbital bristles; rarely with uniform row of 8–10 in decaying matter ranging from rotting fish and animal dung
pairs of reclinate fronto-orbital bristles (Eusiphona) or fron- to rotting plant material. Ferrar (1987) notes the widespread and
tal and orbital setae greatly reduced to absent (Undescribed species-rich genus Desmometopa has many species reared
Genus D). Frons usually with two rows of interfrontal setae, from decaying fruit and vegetation.
sometimes on distinctly shining stripes; lunule often with In addition to these more mundane lifestyles, larvae of
one or two pairs of setae; proboscis often elongate, geniculate. many milichiids have a close association with Hymenoptera.
Wing with humeral and subcostal break; region of latter often Melo (1996) found a species of an undescribed milichiid
modified into costal lappet; cell cup closed, small. Male genus breeding as inquilines in the nests of a species of
terminalia with pregonites fused with hypandrium and Melipona (stingless bees) in Brazil. In the Nearctic Region,
postgonites absent. Krombein (1967) found larvae of the genus Eusiphona feeding
Milichiids are generally recognizable based on the pattern on the pollen in megachilid bee nests; this association may
of head bristling, humeral and subcostal breaks, and the also hold true for Neotropical species of this genus. Of the
closure of cell cup. Some carnids with similar head bristling six species of milichiids in four genera that are known to
lack closed cell cup and lack lunule setae, and all carnids have larvae associated with ant nests (Ferrar, 1987), Costalima
have postgonites in males. Chloropids with a similar general myrmicola Sabrosky is the only species known from the
appearance lack a humeral break and cell cup. Neotropical Region. Some Phyllomyza species have been

1125
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

found in association with formicine and ponerine ants in the feeding behavior with Argiope spiders. Therefore, it is pos-
Palearctic and Oriental Regions (Ferrar, 1987; J.E. Swann, sible that these Phyllomyza may also be egg parasitoids of
unpublished data); these groups of ants may also be hosts Nephila spiders.
of Phyllomyza species in the Neotropical Region. One Robinson & Robinson (1977) also discovered species of
species of Undescribed Genus  B was found in association Neophyllomyza feeding on a stingless bee being digested by
with Azteca sp. ant nests in Trinidad; Undescribed Genus C a reduviid in Panama. Sivinski et al. (1999) note that Neo-
was found in Azteca constructor nests in Cecropia obtusifo- phyllomyza wulpi (Hendel) exhibited a phoretic relationship
lia trees in Costa Rica (J.E. Swann, unpublished data), and with New Guinean asilid flies, presumably also as kleptopara-
Undescribed Genus D was found in Azteca trigona nests in sites. Eisner et al. (1991) found that species of Neophyllomyza,
Guyana (J.E. Swann, unpublished data). A close association Milichiella, Desmometopa, and Paramyia nitens (Loew)
with Azteca ants, which are arboreal, might account for the could be attracted to defensive compounds from coreids and
fact that these three undescribed genera and Costalima (also pentatomids that were commonly captured in Nephila spider
associated with Azteca ants) have been rarely collected. webs. Although Leptometopa latipes is documented as a
Deeming (1998) notes an unidentified species of Milichia spider kleptoparasite (Sivinski et al., 1999), the larvae are also
in Nigeria pursuing ants that were trying to avoid them. This commonly associated with bird nests (Ferrar, 1987).
is the one recorded instance of milichiid behavior that Eisner et al. (1991) and Sivinski  & Stowe (1980) both
indicates they might be true parasitoids of ants rather than report that the vast majority of specimens of all milichiid
just inquilines. Other species of Milichia have larvae that genera collected in association with spiders are female. The
have been reared from various types of dung. extra protein from kleptoparasitic meals may be necessary for
Two Nearctic species of Pholeomyia have been recorded egg maturation (Brake, 2000), and as suggested by Robin-
from the refuse heaps in Atta texana nests (Waller, 1980). son & Robinson (1977), associated flies might possibly be
Neither of these species of Pholeomyia occur in Central egg parasitoids of spiders. Numerous species of Paramyia and
America, although species of Atta are widespread throughout one species of Pholeomyia have been collected in Central and
the Neotropical Region. The only species of Pholeomyia with South America in association with wounded Hymenoptera,
a known life history from Central America is Pholeomyia especially ants (B.V. Brown, personal communication); without
dampfi Sabrosky, recorded as adults following bats deep into exception all specimens examined have been females. Brake
caves in Guatemala and breeding in the bat guano (Sabrosky, (2000) asserts that in addition to nectar feeding, kleptoparasitic
1959). Breeding in bat guano has also been documented in feeding behavior of females is part of the ancestral behavior
the Neotropical Region by Ferreira  & Martins (1999) of milichiids.
(Milichiella spp.) and inferred by Stringer & Meyer-Rochow Many species of several genera can be collected at flowers
(1996, 1997) (Pholeomyia sp.). Papp (1978) has recorded an (Sabrosky, 1987a, b; Papp & Wheeler, 1998). Brantjes (1980)
adult of Pholeomyia leucozona Bilimek from a cave in and Wolda  & Sabrosky (1986) have documented milchiids
Guatemala, and it seems reasonable to assume it is also as important pollinators of Neotropical species of the genus
associated with bat guano. Other genera recorded as adults Aristolochia, finding seven genera (two undescribed) and a
from caves from other regions include Leptometopa and total of 18 species of milichiids pollinating Aristolochia pilosa
Phyllomyza (Papp, 1978, 1982, 1984; Sabrosky, 1964) in Panama. As with the kleptoparasitic records, the flies were
Many adult milichiid flies have fascinating kleptoparasitic all female. Wolda & Sabrosky (1986) noted that there was no
habits. Sivinski et al. (1999) list the genera Phyllomyza, oviposition occuring in association with the plants and that it
Desmometopa, Neophyllomyza, Paramyia, Milichiella, and was possible that the flies were receiving nutrition essential
Leptometopa (listed as Desmometopa latipes (Meigen)) as for egg maturation.
having species with adults that are kleptoparasitic on a wide Adult male milichiids have equally well-documented,
range of predaceous arthropods. Large orb-weaving spiders interesting behaviors. Males of some species of Milichiella,
(genus Nephila) have been observed with several associated Pholeomyia, Milichia, and Pseudomilichia have silvery
milichiid genera feeding on partially digested prey items abdomens and form mating swarms that can be seen from long
(Robinson  & Robinson, 1977; Sivinski  & Stowe, 1980; distances (Sabrosky, 1973; Monteith, 1982; Deeming, 1998).
Eisner et al., 1991). Of particular interest are the observations Although milichiids do not have any known appreciable
of Robinson & Robinson (1977) on Nephila clavipes on Barro economic impact, Ferrar (1987) indicates that the ‘filth-fly’
Colorado Island, Panama, where they found numerous biology of several species makes them prone to dispersal by
examples of female Phyllomyza sp. riding on the spider’s ce- human activities and to be potential vectors of fecal-borne
phalothorax and feeding on prey items as they were partially diseases. Okaeme (1983) found numerous acalyptrate flies,
digested. They hypothesized from their observations that the including species of Leptometopa and Desmometopa, associated
flies might be associated with the same spider for a prolonged with cattle wounds in Nigeria. He indicated that the irritation
period of time, only leaving their perch on the cephalothorax by these flies could reduce cattle productivity and that the flies
to feed. Furthermore, they noted that all observed Phyllomyza could serve as mechanical vectors for pathogens. The only
were females and that the spider egg parasitoid Conioscinella documented case of a milichiid having a significant economic
Duda (Diptera: Chloropidae) exhibits the same sort of adult impact was Desmometopa varipalpis Malloch infesting the
MILICHIIDAE (milichiid flies) 91 1127

biofiltration system of a corn processing plant in Indiana (Wil- including Phyllomyza and Pholeomyia sensu stricto, that
liams & Walker, 1981); pesticides were required to eradicate have from one to four pairs of dorsocentral bristles. There-
the flies. Desmometopa varipalpis is considered to be a fore, for this treatment Eccoptomma is considered a synonym
cosmopolitan species and occurs in Central America. of Milichiella.
Brake (2000) was probably correct in noting that several
genera may be paraphyletic. What is needed is thorough
Classification revisions of the genera in virtually every zoogeographic
region and a higher phylogeny of the genera utilizing
Most current treatments of the family list two subfamilies, numerous species from each genus in the data matrix. From
Madizinae and Milichiinae (Sabrosky, 1980, 1987a; Papp & such a phylogeny, a more sound classification system can
Wheeler, 1998). Sabrosky (1973) included subfamily be developed. Based on current knowledge, with many
Carninae; however, this is now generally recognized as undescribed taxa having suites of characters that blur all
family Carnidae and is thought to be relatively phylogenetically subfamily boundaries, all of the subfamilies that have been
distant from the Milichiidae (Papp & Wheeler, 1998; Brake, used in the past are of doubtful monophyly.
2000). Brake (2000) recognized three subfamilies: besides
Milichiinae, she separated Madizinae sensu lato into two
subfamilies, Phyllomyzinae and Madizinae. This scheme is Identification
not utilized here for several reasons, including that character state
distributions of the six undescribed genera do not match well Because of their apparent lack of economic importance
with Brake’s phylogeny. One could potentially put two of and small size, milichiids have traditionally been overlooked
these undescribed genera in two different subfamilies each, in mass collections, such as those from Malaise traps and
with two others altering the phylogenies within two of the pan traps.
subfamilies. Also, several of the taxa in Phyllomyzinae had At ground level, Malaise trapping usually yields the
missing data for some of the characters used as synapomorphies greatest number of species and genera. Pan traps tend to
for the subfamily. Hennig (1958) argued that Milichiinae are largely collect the genera Neophyllomyza, Paramyia, and
clearly monophyletic, but he noted that Madizinae do not Pholeomyia. Hand collecting around blossoms, UV light
have any clear synapomorphy and may be paraphyletic, as traps, and flight intercept traps, all at ground level, may also
suggested by Brake (2000). collect specimens but usually with much less success than
Brake (2000) noted that Milichia and Leptometopa may Malaise traps. Based on collections from Panama, it would
not be monophyletic and she synonymized Pseudomilichia appear that UV light trapping is the most effective method
with Pholeomyia. According to Brake, the latter association for collecting milichiids in the canopy and is much more
is because the three synapomorphies defining Pholeomyia effective than UV light trapping at ground level. Malaise
sensu stricto also define Pseudomilichia, and the emarginate trapping and canopy fogging are also effective in the canopy,
eye of Pseudomilichia is an insufficient character to define but as with ground level collecting, pan trapping collects a
the genus. Of the three synapomorphies given for Pholeomyia limited number of genera, and flight intercept trapping is the
sensu lato, the presence of more than one postpronotal setae least effective method. While they can be collected in
is acknowledged to also occur in Eusiphona and Microsimus virtually every habitat, milichiids appear to be the most
and thus is homoplasious both within and outside the species-rich in xeric habitats and are relatively rare in aquatic/
subfamily. The presence of three frontal setae is actually a semi-aquatic environments such as stream sides, swamps,
range of three to six frontal setae in Pholeomyia sensu stricto or bogs.
as compared with only three in Pseudomilichia. The presence Air drying tends to result in shriveled specimens, so
of three or four strong anepisternal setae also occurs in some critical-point-dried specimens are preferred. Milichiids are
Eusiphona and in Undescribed Genera A and B. Sabrosky best glued to the side of pins or pointed, as opposed to double
(1955) suggested that the presence of anepisternal bristles mounting, which may destroy important thoracic chaetotaxic
may indicate a close relationship between Eusiphona and characters necessary for generic and specific identification.
Pholeomyia, but in actuality it may unite Eusiphona, With the exceptions of Brake (1999, 2000) and Brake &
Pholeomyia, Pseudomilichia, and Undescribed Genera A and Freidberg (2003), there have been no revisions that have
B. Within this group, Pseudomilichia is the only genus with closely examined genitalic characters. The only recently
a pronounced emargination of the eye; thus for this treatment published worldwide generic revisions are Papp (2001) and
it is recognized as a separate genus. This conforms with both Sabrosky (1983); however, even these revisions have species
Hennig (1939) and Sabrosky (1973). that were not described because of insufficient material.
Sabrosky (1973) listed the genus Eccoptomma Becker, Sabrosky (1959) is useful along with Becker (1907) for keys
but Sabrosky (1980) and Brake (2000) considered it to be a to identify some of the species of Pholeomyia and Milichiella.
synonym of Milichiella. Papp & Wheeler (1998) separated The only modern worldwide generic key is Brake (2000),
Eccoptomma from Milichiella based on the shape of the exci- which will be problematic for some of the poorly known
sion of the posterior margin of the eye, while Becker (1907) Neotropical fauna. Many species descriptions of Neotropical
stated that Eccoptomma have four dorsocentral bristles taxa are more than 50 years old and are sufficiently vague
versus one or two pairs in Milichiella. This difference in the that species-level identifications are extremely difficult and
eye excision is not significant, and there are several genera, will probably necessitate examination of types.
1128
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Key to genera of Milichiidae of the New World

1. Vibrissal angle scarcely or not at all evident; vibrissa well above lower margin of eye (Figs. 2–8) .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Vibrissal angle 90o or less, with vibrissa below level of lower margin of eye (Figs. 9–20) . . . . . 9

2. Frons with 8–10 pairs of uniformly arranged fronto-orbital bristles, all erect and reclinate, showing
no distinction between orbital and frontal series (Fig. 2). . . . . . . . . . . . . . . . Eusiphona Coquillett
– Frons with five or fewer pairs of medioclinate frontal bristles and two or three pairs of orbital
bristles (Figs. 3–8). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3. Hind margin of eye not excised (Figs. 3–5). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

– Hind margin of eye excised at level of antennae; excision narrow or may continue to lower corner
of eye (Figs. 6–8). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

4. Thorax with two rows of greatly enlarged presutural and postsutural acrostichal setae (Fig. 35);
medial supra-antennal setae greatly enlarged, approximately equal in size to orbital setae (Fig. 3);
Argentina. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Undescribed Genus A]
– Thorax with no enlarged presutural and at most two enlarged pairs of postsutural acrostichal setae;
medial supra-antennal setae small, much smaller than orbital setae (Figs. 4–5). . . . . . . . . . . . . . 5

5. Anepisternum bare posteriorly. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Milichia Meigen

– Anepisternum posteriorly with row of strong bristles (Fig. 30). . . . . . . . . . . . . . . . . . . . . . . . . . . 6

6. Ptilinal suture extending well posterior of the antennae, to vicinity of orbital bristles and ocellar
triangle (Fig. 4). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Undescribed Genus B
– Ptilinal suture near antennae, anterior to medioclinate frontal bristles (Fig. 5). . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pholeomyia Bilimek

7. Anepisternum posteriorly with row of strong bristles (Fig. 31). . . . . . . . . Pseudomilichia Becker

– Anepisternum bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

8. Head and thorax rough, with fine pubescence; posterior margin of eye with excision from level
of antennal insertion extending to lower margin of eye; proboscis long and strongly geniculate
(Fig. 6). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulia Becker
– Head and thorax smooth, hairs and bristles normal; excision in posterior margin of eye usually not
extending to lower margin of eye; proboscis long or short, strongly to weakly geniculate (Fig. 7).
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Milichiella Giglio-Tos

9. Wing with subcostal break developed into prominent costal notch (Fig. 21). . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Undescribed Genus C
– Wing without subcostal break developed into prominent notch (Figs. 22–29). . . . . . . . . . . . . . 10
10. C extending only to R4+5 (Figs. 22–23) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
– C extending to M1 (Figs. 24–29). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

11. Crossvein dm-cu absent (Fig. 22); arista slender, pubescent (Fig. 9) . . . . . . . . Paramyia Williston
– Crossvein dm-cu present (Fig.  23); arista thickened at base, densely long-haired (Fig.  10);
Nearctic Region, south to South Carolina, Palearctic, Afrotropical, and Oriental Regions . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Aldrichiomyza Hendel]
MILICHIIDAE (milichiid flies) 91 1129

frorb s

2 Eusiphona 3 Undescribed Genus A 4 Undescribed Genus B

5 Pholeomyia 6 Ulia 7 Milichiella

o vt s poc s
orb s i vt s

infr s
fr s

vb sbvb s

8 Pseudomilichia 9 Paramyia 10 Aldrichiomyza

Figs.  91.2–10. Heads: anterodorsal view of (2)  Eusiphona mira Coquillett, (Nearctic, MND, fig.  79.6), (3)  Undescribed Genus A,
(4) Undescribed Genus B, (5) Pholeomyia sp., (6) Ulia sp., (7) Milichiella lacteipennis (Loew), (Nearctic, MND, fig. 79.7), (8) Pseudomilichia sp.,
(9) Paramyia sp., and (10) Aldrichiomyza agromyzina (Hendel), (Nearctic, MND, fig. 79.2). Figures 3–6 and 8–9 illustrated by A. Brenes.
Abbreviations: frorb s, fronto-orbital seta; fr s, frontal seta; infr s, interfrontal seta; i vt s, inner vertical seta; orb s, orbital seta; o vt s, outer
vertical seta; poc s, postocellar seta; sbvb s, subvibrissal seta; vb, vibrissa.
1130
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

11 Undescribed Genus D 12 Stomosis 13 Microsimus

14 Costalima 15 Undescribed Genus E 16 Phyllomyza

vb

18 Leptometopa 19 Madiza

17 Desmometopa 20 Undescribed Genus F

Figs.  91.11–20. Heads (continued): anterodorsal view of (11)  Undescribed Genus  D, (12)  Stomosis  sp., (13)  Microsimus  sp.,
(14)  Costalima  sp., (15)  Undescribed Genus  E, (16)  Phyllomyza securicornis Fallén, (Holarctic, MND, fig.  79.3), (17)  Desmometopa
m-nigrum (Zetterstedt), (MND, head from fig. 79.1), (18) Leptometopa latipes (Meigen), (Holarctic, MND, fig. 79.4), (19) Madiza glabra
Fallèn, (Holarctic, MND, fig. 79.5), and (20) Undescribed Genus F. Figures 11–15 and 20 illustrated by A. Brenes.
Abbreviation: vb, vibrissa.
MILICHIIDAE (milichiid flies) 91 1131

12. Frons appearing bare, all frontal and orbital bristles absent or reduced to microsetae (Fig.  11)
(known only from females collected from an Azteca trigona nest in Guyana). . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Undescribed Genus D]
– Frons with two or three pairs of enlarged lateroclinate orbital bristles (Figs. 12–20) . . . . . . . . 13

13. Frons with three pairs of lateroclinate orbital bristles (Figs. 12–16). . . . . . . . . . . . . . . . . . . . . . 14

– Frons with two pairs of lateroclinate orbital bristles (Figs. 17–20). . . . . . . . . . . . . . . . . . . . . . . 18

14. Two enlarged postpronotal setae present (Fig. 33); eye small (Fig. 13); male with long fringe of
hairs on hind margin of wing (Fig. 24) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microsimus Aldrich
– One enlarged postpronotal seta present (Fig. 34); eye normal (Figs. 12, 14–16); male lacking long
fringe of hairs on hind margin of wing (Fig. 28). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

15. Lunule with at least three prominent setae, dorsal pair approximately twice size of ventral
(Fig. 15). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Undescribed Genus E
– Lunule with only one pair of setae (Figs. 12, 14, 16) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

16. Lunule setae extremely large, approximately equal in length to antenna; middle lateroclinate
orbital seta, if present, much weaker than anterior and posterior lateroclinates (Fig. 14). . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Costalima Sabrosky, in part
– Lunule setal length at most approximately one-half antennal length; middle lateroclinate seta
strong, approximately equal in length to anterior and posterior lateroclinates (Figs. 12, 16). . . 17

17. Proepimeral setae present (Fig. 32); lower margin of gena obviously shorter than greatest length
of head (Fig. 16); body color various but if orange or yellow, dorsal margin of katepisternum also
orange or yellow. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phyllomyza Fallén
– Proepimeral setae absent; lower margin of gena equal to slightly subequal to greatest length of
head (Fig. 12); body orange or yellow with dorsal margin of katepisternum brown. . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stomosis Melander

18. Frons with two rows of interfrontal setae borne on gray sclerotized interfrontal strips; strips and
similarly-colored fronto-orbital plates and ocellar triangle demarcating dull to velvet black
M-shaped area (Fig. 17). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Desmometopa Loew
– Interfrontal setae, if present on frons, not on sclerotized strips that are the same color as
fronto-orbital plates (Figs. 14, 18–20). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
19. Lunule extended ventrally on midline of face as flat plate between antennae, reaching nearly to
lower margin of face; lower margin of face usually extended dorsally, subtriangular (Fig. 18);
anepimeron with short stout bristle; hind tibia of male greatly broadened (Fig. 36). . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leptometopa Becker
– Lunule not extended ventrally on midline of face as flat plate dividing antennae; lower margin
of the face not extending dorsally (Figs. 14, 19–20); anepimeron bare; hind tibia not enlarged in
male . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

20. Body polished black. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

– Body dull or subshining black, brown, or orange . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

21. Crossvein dm-cu almost equidistant from wing margin (along CuA1) and from crossvein r-m
(Fig. 25); lunule small and often slightly concave (Fig. 19); body and coxae uniformly black, wing
hyaline; Holarctic and Oriental Regions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Madiza Fallén]
– Crossvein dm-cu much closer to wing margin (along CuA1) than to crossvein r-m (Fig. 29); lunule
large, forming shelflike protrusion from frontal region (Fig. 20); body black with forecoxa often
bright orange; wing often fumose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Undescribed Genus F
1132
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

21 Undescribed Genus C 22 Paramyia

23 Aldrichiomyza 24 Microsimus

25 Madiza 26 Undescribed Genus E

27 Neophyllomyza 28 Costalima

29 Undescribed Genus F

Figs.  91.21–29. Wings: dorsal view of (21)  Undescribed Genus  C, (22)  Paramyia nitens (Loew), (Nearctic, MND, fig.  79.12),
(23)  Aldrichiomyza agromyzina (Hendel), (Nearctic), (24)  Microsimus  sp., (25)  Madiza glabra Fallèn, (Holarctic, MND, fig.  79.13),
(26)  Undescribed Genus  E, (27)  Neophyllomyza quadricornis Melander, (Nearctic, MND, fig.  79.14), (28)  Costalima  sp., and
(29) Undescribed Genus F. Figures 21, 23–24, 26, and 28–29 illustrated by A. Brenes.
MILICHIIDAE (milichiid flies) 91 1133

30 Undescribed Genus B 31 Pseudomilichia 32 Phyllomyza

33 Microsimus 34 Phyllomyza

36 Leptometopa

35 Undescribed Genus A

Figs.  91.30–36. Thorax and leg: lateral view of anterior portion of thorax showing anepisternum of (30)  Undescribed Genus  B and
(31) Pseudomilichia sp.; ventrolateral view of anterior portion of thorax showing proepimeron of (32) Phyllomyza sp.; dorsolateral view of
anterior portion of thorax showing postpronotal setae of (33) Microsimus sp.; and dorsal view of the thorax of (34) Phyllomyza sp. and
(35) Undescribed Genus A; hind tibia of male of (36) Leptometopa latipes (Meigen), (Holarctic, MND, fig. 79.18). Figures 30–35 illustrated
by A. Brenes.
1134
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

22. Cell dm short; crossvein dm-cu 2 or 3 times as far from wing margin as from crossvein r-m
(Fig. 27); body black or dark brown; lunule setae small, less than one-half length of antenna. . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neophyllomyza Melander
– Cell dm long; crossvein dm-cu almost equidistant from wing margin (along CuA1) and from
crossvein r-m (Fig. 28); body orange or light brown; lunule setae extremely large, at least equal in
length to entire antenna (Fig. 14). . . . . . . . . . . . . . . . . . . . . . . . . . . . . Costalima Sabrosky, in part

Synopsis of the fauna match Nearctic specimens of E. mira. I suspect Eusiphona

is widespread but rare throughout Central America and that
The number of described species in most areas of the there are several undescribed species in the region.
world, including Central America, is a serious underestima-
tion of the total number present. Stork (1991) collected 169 Leptometopa Becker. Leptometopa halteralis (Coquillett)
milichiid specimens comprising 25 species through canopy is recorded from the Caribbean and Neotropical portions of
fogging of Bornean rain forest trees. Swann (2000) examined Mexico. Leptometopa latipes is also recorded from the
107 specimens comprising 29 species, of which only 2 could Caribbean, is considered to be widespread, and probably occurs
be confirmed as described species, in a preliminary faunal in Central America. The two species may be separated using
survey of Namibia. There are 30 species in 10 genera known Cole (1969). I have seen some apparently undescribed
to occur in Central America and 14 species in seven genera species from Nearctic portions of Mexico and suspect that
known from Costa Rica. I have seen 73 species in 12 genera there are several more in Central America.
from 489 specimens from Costa Rica and expect, based on Microsimus Aldrich. This is a monotypic genus known
the ratio of known genera and species relative to the material from adults in the nests of Crematogaster stolli Forel from
I have seen from the country, at least 156 species in 17 gen- Bolivia (Aldrich, 1926). I have seen additional specimens of
era in Central America, based on the number of currently Microsimus from Panama, Guyana, and Peru. The distribution
known genera and species from this region. of the host ant Crematogaster stolli is Guatemala to Colombia,
Costalima Sabrosky. Costalima myrmicola, the only Venezuela, and Trinidad, with a similar species replacing it
described species in this genus, is found in Azteca ant nests further to the south. It is likely that as an inquiline, Microsimus
in Cecropia trees in Minas Gerais, Brazil. There is at least matches the distribution of the host.
one, and possibly two, undescribed species in Panama. The Milichia Meigen. I have seen two specimens of what
genus may be widespread in the Neotropical Region, but its appears to be Milichia aethiops Malloch from Costa Rica.
distribution is poorly known. This may in part be because of This species is known in the literature from Baja California,
the requirement for highly specialized collecting techniques Nuevo León, and Zacatecas in the Nearctic portion of Mexico
to sample from nests of arboreal Azteca ants. and may also occur in Neotropical Mexico.
Desmometopa Loew. There are several inaccuracies in Milichiella Giglio-Tos. There are 17 species of Milichiella
the interpretation of Nearctic and Neotropical ranges for recorded from the Neotropical Region, of which only two,
Mexican species of Desmometopa in both Sabrosky (1973) the almost cosmopolitan Michiella lacteipennis (Loew)
and Brake (2000). Based on distributional data in Sabrosky (Costa Rica) and Milichiella bisignata Melander (Costa
(1983), there are 13 species of Desmometopa known from Rica, Mexico), are known from Central America. I have seen
Central America. There are several other cosmopolitan and 13 species from Costa Rica, of which many appear to be
Neotropical species that might also occur, but there are no undescribed, and there are probably at least 20 species from
records to date. Of the eight species I have examined from Central America as a whole. There is no satisfactory
Costa Rica, five could not be identified with certainty using Neotropical key for this genus.
Sabrosky’s (1983) revision, and they might be undescribed.
I expect several more undescribed species from the rest of Neophyllomyza Melander. Neophyllomyza magnipalpus
Central America. (Williston) from St. Vincent is the only described Neotropical
species, although the genus is widespread throughout the
Eusiphona Coquillett. There are two species of Eusiphona region. There are at least seven species in Costa Rica, and I
described from the Neotropical Region, of which Eusiphona would expect at least twice that number of species in Central
mira Coquillett occurs in both the Nearctic Region and in America.
Neotropical Mexico. The only described, exclusively
Neotropical species is Eusiphona vittata Sabrosky from Paramyia Williston. There are seven species of Paramyia
Argentina (Sabrosky, 1982). Sabrosky (1955, 1982) provides known from the Neotropical Region, of which the only three
adequate means to sort the described species. In addition, I Central American species are known from Costa Rica. I have
have seen single specimens, each a different species, from El seen 16 species from Costa Rica and would expect there to
Salvador, Trinidad, Jamaica, and Peru. None of these specimens be many more from Central America.
MILICHIIDAE (milichiid flies) 91 1135

Pholeomyia Bilimek. Pholeomyia is widespread and a species complex. There are probably at least three or four
species-rich throughout the Neotropical Region. There are species of Stomosis in Central America.
28 described Neotropical species, of which seven are known
to occur in Central America. Sabrosky’s (1959) key includes Ulia Becker. Brake (2000) lists Ulia as occurring in Peru,
eight Neotropical species, including all those recorded from Bolivia, and Costa Rica. Besides these countries, I have
Central America. I have seen 13 species from Costa Rica, also seen specimens of the single species, U. poecilogastra
of which 10 could not be identified with certainty using Becker, from El Salvador and suspect that it is widespread in
Sabrosky (1959) and Becker (1907). It is difficult to say what Central America and the northern half of South America.
proportion of these are undescribed species and which are
simply range extensions of described South American species, Undescribed Genus B. This genus contains at least three
but I suspect that there are many undescribed Pholeomyia in undescribed Neotropical species. One is known from a col-
Central America. lection in association with Azteca sp. ant nests and a black
light collection in Trinidad; one is known from collections
Phyllomyza Fallén. Sabrosky (1973) noted that Phyllomyza of pollinators of Aristolochia pilosa and canopy fogging in
occurs in Costa Rica but that there are no described Neo- Panama; and one species is known from a light trap collection
tropical species. I have seen 10 undescribed species from from Colombia. Undescribed Genus B may occur throughout
Costa Rica and would expect twice that number from all of Central and South America.
Central America. Phyllomyza appears to be relatively uncommon
in collections from the Neotropical Region compared with Undescribed Genus C. This genus, with its single
collections from other tropical regions; however, it seems to species, is only known from four collections from nests of
be equally species-rich in all tropical regions. the ant Azteca constructor in Cecropia obtusifolia trees in
Costa Rica. If the species is host-specific then one would
Pseudomilichia Becker. This may or may not be a valid
predict its range to be the same as A.  constructor, from
genus. For the purposes of this chapter, I have followed the
generic definition of Sabrosky (1973) and the species definition Guatemala to northern South America (Longino, 1991).
of Brake (2000). As such, Pseudomilichia is valid and has Undescribed Genus E. This genus contains two identified,
three described species from Bolivia and Peru. I have also
but undescribed, species, one Ecuadorian and one Costa
seen an additional species from Panama.
Rican.
Stomosis Melander. There are two uncommon described
species of Neotropical Stomosis: S.  innominata (Williston) Undescribed Genus F. This genus appears to be wide-
(widespread in Central America and in the Nearctic Region) spread and relatively species-rich in the Neotropical region.
and S. rufula (Frey) (Brazil). Sabrosky (1973) lists, in part, There are 12 identified, but undescribed, species occurring
S.  innominata from Costa Rica, El  Salvador, Trinidad, in Argentina, Ecuador, Bolivia, Peru, Costa Rica, El Salvador,
Neotropical Mexico, and Arizona. I have seen specimens from and Trinidad. There are six species known to occur in
these countries, and there appear to be marked differences Costa Rica and El Salvador. It is likely widespread in Central
between them that may indicate that S. innominata is actually America.

Literature cited

Aldrich, J.M. 1926. Descriptions of new and little-known Diptera or Cole, F.R. 1969. The Flies of Western North America. University of
two-winged flies. Proceedings of the United States National California Press, Berkeley & Los Angeles, California, 694 pp.
Museum 69: 1–26. Deeming, J.C. 1998. Milichiidae and Carniidae (Diptera: Cyclorrhapha)
Becker, T. 1907. Die Dipteren-Gruppe Milichiinae. Annales Musei from the Arabian peninsula. Fauna of Arabia 17: 147–157.
Nationalis Hungarici 5: 507–550. Eisner, T., M. Eisner,  & M.  Deyrup. 1991. Coattraction of
Brake, I. 1999. Prosaetomilichia DeMeijere: a junior subjective kleptoparasitic flies to heteropteran insects caught by orb-
synonym of Milichia Meigen, with a phylogenetic review of the weaving spiders. Proceedings National Academy of Sciences
myrmecophila species-group (Diptera, Milichiidae). Tijdschrift 88: 8194–8197.
voor Entomologie 142: 31–36. Ferrar, P. 1987. A guide to the breeding habits and immature
Brake, I. 2000. Phylogenetic systematics of the Milichiidae (Diptera,
stages of Diptera Cyclorrhapha. Entomonograph 8(1).
Schizophora). Entomologica scandinavica Supplement 57: 1–120.
E.J. Brill / Scandinavian Science Press, Leiden, Copenhagen,
Brake, I. & A. Freidberg. 2003. Revision of Desmometopa Loew
(Litometopa Sabrosky) (Diptera: Milichiidae), with descriptions 478 pp.
of six new species. Proceedings of the Entomological Society of Ferreira, R.L & R.P. Martins. 1999. Trophic structure and natural
Washington 105: 265–282. history of bat guano invertebrate communities, with special
Brantjes, N.B.M. 1980. Flower morphology of Aristolochia reference to Brazilian caves. Tropical Zoology 12: 231–251.
species and the consequences for pollination. Acta Botanica Hennig, W. 1939. Beiträge zur Kenntnis des Kopulationsapparates
Neerlandica 29: 212–213. und der Systematik der Acalyptraten. II. Tethinidae, Milichiidae,
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Anthomyzidae und Opomyzidae (Diptera). Arbeiten über Sabrosky, C.W. 1977. Family Milichiidae, pp.  270–274. In Del-
morphologische und taxonomische Entomologie 6: 81–94. finado, M.D. & D.E. Hardy (editors). A catalog of the Diptera
Hennig, W. 1958. Die familien der Diptera Schizophora und ihre of the Oriental Region, Volume III. University Press of Hawaii,
phylogenetischen verwandtschaftbeziehungen. Beiträge zur Honolulu, x + 854 pp.
Entomologie 8: 505–688. Sabrosky, C.W. 1980. Family Milichiidae, pp. 686–689. In Cross-
Krombein, K.V. 1967. Trap-nesting wasps and bees. Life histories, key, R.W. (editor). Catalogue of the Diptera of the Afrotropical
nests and associates. Smithsonian Press, Washington, D.C., Region. British Museum (Natural History), London, 1437 pp.
570 pp. Sabrosky, C.W. 1982. Una nueva especie de Eusiphona de la
Longino, J.T. 1991. Taxonomy of the Cecropia-inhabiting Azteca Republica Argentina. Revista del Museo de La Plata (Nueva
ants. Journal of Natural History 25: 1571–1602. Serie) 13: 7–9.
Melo, G.A.R. 1996. Notes on the nesting biology of Melipona
Sabrosky, C.W. 1983. A synopsis of the world species of Desmometopa
capixaba (Hymenoptera, Apidae). Journal of the Kansas
Loew (Diptera, Milichiidae). Contributions of the American
Entomological Society 69: 207–210.
Monteith, G.B. 1982. Dry season aggregations of insects in Entomological Institute 19: 1–69.
Australian monsoon forests. Memoirs of the Queensland Sabrosky, C.W. 1987a. Milichiidae, pp. 903–908. MND,
Museum 20: 533–543. Volume 2.
Okaeme, A.N. 1983. Notes on acalyptrates (Diptera) as pest of Sabrosky, C.W. 1987b. A new species of Leptometopa (Diptera,
cattle. Bulletin of Animal Health and Production in Africa Milichiidae) from Madagascar pollinating Ceropegia (Ascle-
31: 255–256. piadaceae). Proceedings of the Entomological Society of
Papp, L. 1978. Some cavernicolous Diptera of the Geneva Museum. Washington 89: 242–243.
Revue Suisse de Zoologie 85: 99–106. Sivinski, J. & M. Stowe. 1980. A kleptoparasitic cecidomyiid and
Papp, L. 1982. Cavernicolous Diptera of the Geneva Museum. other flies associated with spiders. Psyche 87: 337–348.
Revue Suisse de Zoologie 89: 7–22. Sivinski, J., S. Marshall,  & E.  Petersson. 1999. Kleptoparasitism
Papp, L. 1984. Cavernicolous Milichiidae (Diptera): three new species and phoresy in the Diptera. Florida Entomologist 82: 179–197.
from Fiji and Sri Lanka. Revue Suisse de Zoologie 91: 241–247. Stork, N.E. 1991. The composition of the arthropod fauna of
Papp, L. 2001. A revision of the species of Paramyia Williston Bornean lowland rain forest trees. Journal of Tropical Ecology
(Diptera, Milichiidae) with the description of a new genus. Acta 7: 161–180.
Zoologica Academiae Scientiarum Hungaricae 47: 321–347. Stringer, I.A.N.  & V.B. Meyer-Rochow. 1996. Distribution of
Papp, L.  & T.A. Wheeler. 1998. 3.28. Family Milichiidae, flying insects in relation to predacious web-spinning larvae
pp. 315–324. In Papp, L. & B. Darvas (editors). Contributions of Neoditomyia farri (Diptera: Mycetophilidae) in a Jamaican
to a Manual of Palaearctic Diptera. Volume 3, Higher Brachycera. cave. Annals of the Entomological Society of America 89:
Science Herald Publishers, Budapest, 880 pp. 849–857.
Robinson, M.H. & B. Robinson. 1977. Associations between flies Stringer, I.A.N.  & V.B. Meyer-Rochow. 1997. Flight activity of
and spiders: bibiocommensalism and dipsoparasitism? Psyche insects within a Jamaican cave: in search of the zeitgeber.
84: 150–157. Invertebrate Biology 116: 348–354.
Sabrosky, C.W. 1955. A third species of Eusiphona, with remarks
Swann, J.E. 2000. Milichiidae (Diptera: Carnoidea), pp. 265–268.
on the systematic position of the genus (Diptera, Milichiidae).
In Kirk-Spriggs, A.H. & E. Marais (editors). Dâures — biodiversity
Entomological News 66: 169–173.
Sabrosky, C.W. 1959. A revision of the genus Pholeomyia in North of the Brandberg Massif, Namibia. Cimbebasia Memoir  9,
America (Diptera, Milichiidae). Annals of the Entomological National Museum of Namibia, Windhoek, 389 pp.
Society of America 52: 316–331. Waller, D.A. 1980. Leaf-cutting ants and leaf-riding flies. Ecological
Sabrosky, C.W. 1964. Milichiidae and Chloropidae (Diptera) from Entomology. 5: 305–306.
the Batu Caves, Malaya. Pacific Insects 6: 308–311. Williams, R.E. & C.L. Walker. 1981. A unique fly problem in an
Sabrosky, C.W. 1973. 75. Family Milichiidae. In Papavero,  N. industrial bio-filter system. Pest Control 49: 14–16.
(editor). A catalogue of the Diptera of the Americas south of Wolda, H. & C.W. Sabrosky. 1986. Insect visitors to two forms of
the United States. Departamento de Zoologia, Secretaria da Aristolochia pilosa in Las Cumbres, Panama. Biotropica 18:
Agricultura, São Paulo, 12 pp. 295–299.
CHLOROPIDAE (frit flies, grass flies, eye gnats) 92
Terry A. Wheeler

Fig. 92.1. Female of Chlorops certimus Adams, (Nearctic, MND, fig. 99.1).

Diagnosis symmetrical, genitalia simple. Female abdomen telescoping,

cerci long, spermathecae unsclerotized.
Minute to small (body length 1.0–5.0 mm, rarely larger) The combination of wing venation (missing cup and CuA2,
flies (Fig. 1). Shining black to dull gray or yellow, often with kink in CuA1), vertical propleural carina, reduced fronto-
patterns of black and yellow. Antenna with first flagellomere orbital setae (in most species), and prominent frontal triangle
usually round, arista dorsal, bare or pubescent. Cephalic (in most species) distinguishes chloropids from other acalyp-
chaetotaxy reduced: ocellar, postocellar, inner, and outer trate flies. Some shiny black species of Milichiidae and some
vertical setae usually present, fronto-orbital setae usually yellow and black Agromyzidae and Clusiidae are externally
reduced, only slightly longer than surrounding setulae, occa- similar but can be distinguished by the number and orien-
sionally with one to two longer setae present, frontal setulae tation of fronto-orbital bristles and by differences in wing
usually uniformly distributed, except on surface of frontal venation. Ephydridae with similar wing venation generally
triangle; distinct frontal triangle usually present, occupy- have a projecting face and longer fronto-orbital bristles.
ing half or more of frons; thoracic chaetotaxy usually reduced
(less so in Apotropina), one to two postpronotal, one anterior,
one to two posterior notopleural, one to two postsutural
Biology
dorsocentral setae present, acrostichal setulae in several rows,
two to three pairs of scutellar setae present; propleuron with Species of Chloropidae have a wider range of lifestyles
complete, sharp vertical carina, prothoracic spiracle round, than almost any other family of flies. Larvae of most species
wing with subcostal break (secondarily lost in some species are apparently saprophagous, but many others are phytophagous
of Pseudogaurax), sc incomplete, cell cup and CuA2 absent, in stems or flower heads, particularly of grasses (Poaceae) or
kink in CuA1 usually present basal to dm-cu. Legs usually sedges (Cyperaceae). Smaller numbers of species are gall
without outstanding bristles, but apicoventral hind tibial spur inducers or inquilines; predators of insects and other inver-
(actually a spurlike seta) present in some genera; hind tibia tebrates; predators in egg masses of spiders, amblypygids,
often with elongate or oval tibial organ. Male postabdomen and Lepidoptera; and species of one Australian genus are

1137
1138
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

subcutaneous parasites of amphibians. Some adults are placement of many species should be expected in the future
kleptoparasites of prey captured by larger arthropods (Ferrar, as additional species are described and phylogenetic studies
1987; Sivinski et al., 1999). Chloropidae are species-rich and are undertaken.
abundant in most habitats, including marine beaches, fresh-
water wetlands, riparian habitats, fields and roadsides, rain
forest, and cloud forest, including the canopy. They are Identification
particularly dominant in disturbed habitats.
Several species are of economic importance. The pest spe- Many genera of Chloropidae are restricted to one or a
cies fit into two general ecological groups: some phytophagous few zoogeographic realms, so keys to other regions will
species in the genera Chlorops, Meromyza, and Oscinella be less effective for identification of the Central American
are pests of cereal crops; the eye gnats and yaws flies in the fauna. Sabrosky’s (1987) Nearctic key will be of some use in
genera Hippelates, Liohippelates, and, rarely, Siphunculina Mexico, although some widespread Neotropical genera are
are attracted to eyes, nostrils, mouths, perspiration, or open not included. Most of the Neotropical genera are endemic,
wounds where they feed on fluids. Normally this represents
and existing keys to the Neotropical fauna are outdated or
only a nuisance, but some species are also mechanical vectors
incomplete. Enderlein (1911) described many of the cur-
of human and livestock diseases, including yaws, conjunctivitis,
and Brazilian Purpuric Fever (Sabrosky, 1987; Paganelli & rently recognized Neotropical genera, but his descriptions
Sabrosky, 1993). and generic key were based primarily on material from
southern Brazil. Becker’s (1912) monograph of the Neotro-
pical Chloropidae provided keys to the genera and species
Classification described at that time. Unfortunately, Becker’s concepts of
some of Enderlein’s genera were in error and his keys are
The sister group of the Chloropidae is the Milichiidae out of date because of the subsequent description of many
(Brake, 2000); Acartophthalmidae may be the sister group genera and species. Duda’s (1929, 1930, 1931, 1933) mono-
to these two families (Brake, 2000), although that relationship graph of the Neotropical Chloropidae included descriptions
is less clearly resolved. The classification of Buck (2006) of 26 new genera and more than 100 new species, but omit-
placed the Cryptochetidae as the sister group to Chloropidae ted 12 genera and 75 species described by previous authors,
plus Milichiidae. which created unnecessary synonyms and errors in his keys.
The family is generally divided into three subfamilies: Keys to the fauna of other parts of the Neotropical Region,
Siphonellopsinae, Chloropinae, and Oscinellinae. Some such as Curran (1928) for Puerto Rico or Malloch (1934)
authors have divided the Oscinellinae into additional for Patagonia, omit many genera present in Central America.
subfamilies (e.g., Hippelatinae, Rhodesiellinae), but this Paganelli (1985) revised the Neotropical Chloropinae at the
division is not well supported, and some of the resulting taxa generic level, but that work is not available except in thesis
are undoubtedly paraphyletic. There is currently no accept-
form, so names proposed for new taxa are not valid.
able phylogenetic hypothesis on higher level relationships
Identification of chloropid genera relies mostly on external
within the Chloropidae, including the monophyly of the
characters, often in combinations of presence–absence
three recognized subfamilies.
Over 200 genera of Chloropidae are currently recognized, character states. Color characters, which form the basis of
although many are probably not monophyletic and some (e.g., many older species descriptions and generic limits, are some-
Conioscinella, Oscinella, Chlorops) are large repositories of times varied, especially in species with a wide geographic or
species that cannot be accommodated in smaller genera as elevational range. Furthermore, several undescribed species
currently defined. The Siphonellopsinae is the smallest have combinations of characters intermediate between those
subfamily with three genera worldwide, one of which occurs of recognized genera.
in Central America. There are 10 genera of Chloropinae in the Specimens of Chloropidae are well sclerotized (for their
region. All remaining genera belong to Oscinellinae, which size) and have reduced chaetotaxy, so they are less prone
is the largest and most morphologically diverse subfamily. than many flies to damage during collecting and processing.
Thirty-five genera of Oscinellinae have been recorded in However, given their small size, adult specimens should be
Central America. Given the lack of phylogenetic information preserved in ethanol and critical-point dried or chemically
on the family, I am not in a position to make major changes dried as soon as possible thereafter. Air-dried specimens may
to generic classification and, consequently, the classification shrivel badly and are often difficult to identify to the generic
followed here is conservative, using current limits of most level. Specimens that have been stored for some time in eth-
genera despite their possible non-monophyly. Exceptions anol (especially at room temperature) lose their color, which
are noted in the text and significant changes in the generic makes comparison to well-prepared specimens difficult.
CHLOROPIDAE (frit flies, grass flies, eye gnats) 92 1139

Key to the genera of Chloropidae of Central America

1. Costa ending at apex of M1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Costa ending at apex of R4+5, rarely slightly beyond R4+5; Chloropinae. . . . . . . . . . . . . . . 44

2. Fronto-orbital bristles long, proclinate or lateroclinate, ocellar bristles long, proclinate

and divergent (Fig. 2), postgenal margin with long ventral bristle; postpronotum with two strong
bristles; SIPHONELLOPSINAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apotropina Hendel
– Fronto-orbital bristles usually short; if long, then reclinate or erect, ocellar bristles usually erect or
reclinate, rarely proclinate and divergent; postpronotum usually with one long bristle, rarely with
two or more; OSCINELLINAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3. Hind tibia with dark, ventral apical or pre-apical spur, as long or longer than tibial diameter
(Figs. 29–30) (specimens with dark spur slightly shorter than tibial diameter will key to both sides
of this couplet). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
– Hind tibial spur absent or (rarely) shorter than tibial diameter (Fig. 31). . . . . . . . . . . . . . . . . . . . 8

4. Eye with dense, long hairs; first flagellomere reniform, expanded dorsally basal to arista, arista
long and pubescent (Fig. 9); scutellum long, triangular, flattened dorsally, scutellar bristles pale;
rarely collected in Central America, usually on marine beaches. . . . . . . . . . . . . . Cadrema Walker
– Eye bare or sparsely short-haired; first flagellomere rounded, arista bare or pubescent; scutellum
rounded apically or trapezoidal, usually not flattened dorsally, scutellar bristles usually dark. . . 5

5. Scutellum trapezoidal with apical bristles on small tubercles at posterolateral corner; notopleuron
with one anterior, one posterior bristle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Goniaspis Duda
– Scutellum rounded apically, tubercles present or absent, apical scutellar bristles not as widely
separated; notopleuron usually with one anterior, two posterior bristles. . . . . . . . . . . . . . . . . . . . 6

6. Frontal triangle shining, rest of frons and gena bare but microscopically striate so that surface
appears subshining; thorax mostly shining, scutal setae usually arising from small punctures or
tubercles, such that surface of scutum appears punctate. . . . . . . . . . . Liohippelates Duda, in part
– Frontal triangle shining or dull, rest of frons and gena densely pruinose; gena dull, pruinose;
thorax with at least some pruinosity in posterior part of scutum and on pleurites, usually more
extensively pruinose, scutal setae not arising from punctures or tubercles, surface of scutum
usually appearing smooth, although sometimes heavily pruinose. . . . . . . . . . . . . . . . . . . . . . . . . 7

7. Frontal triangle usually entirely shining, at least partly shining anteriorly; gena usually as broad
as first flagellomere, densely pruinose and paler than rest of head; thorax and legs usually black.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apallates Sabrosky, in part
– Frontal triangle dull (rarely with three shining spots adjacent to ocelli or single median shining
stripe); gena usually narrower than first flagellomere and not strongly contrasting with color and
pruinosity of rest of head; thorax and legs usually yellow or gray . . . . . . Hippelates Loew, in part

8. Eye reniform, concave posteroventrally; vibrissa placed well above ventral margin of eye, with
row of at least four subvibrissal setae below this (Fig. 6); scutellum with long, curved, marginal
projections, densely covered with setulae (Fig. 15); anepisternum with long dense setae dorsally;
body shining black. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acanthopeltastes Enderlein
– Not with the above combination of characters. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

9. Wing with R2+3 short, second costal sector less than half as long as third costal sector (Fig. 22);
frons and thorax with distinctive pattern of shining black and dull gray irregular lines; rarely
collected in Central America. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Siphunculina Rondani
– Wing with R2+3 longer; second costal sector as long or longer than third costal sector; frons and
thorax not patterned as above. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
1140
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

2 Apotropina 3 Olcella 4 Fiebrigella

5 Monochaetoscinella 6 Acanthopeltastes 7 Medeventor

8 Elachiptera 9 Cadrema 10 Parectecephala

Figs. 92.2–10. Heads and antennae: anterodorsal view of head of (2) Apotropina sp.; anterolateral view of head of (3) Olcella cinerea (Loew),
(MND, fig. 99.16); (4) Fiebrigella sp.; (5) Monochaetoscinella nigricornis (Loew), (Nearctic, MND, fig. 99.12); (6) Acanthopeltastes sp.;
and (7) Medeventor nubosus Wheeler; lateral view of left antenna of (8) Elachiptera costata (Loew), (Nearctic, MND, fig. 99.20);
(9) Cadrema pallida (Loew), (MND, fig. 99.21); and (10) Parectecephala eucera (Loew), (Nearctic, MND, fig. 99.29). Figures 2, 4, and 7
illustrated by R. Vargas; Figure 6 illustrated by B. Koehler.
CHLOROPIDAE (frit flies, grass flies, eye gnats) 92 1141

pprn s

npl s

pal s
dc s
ial s
acr s

sbap sctl s

ap sctl s

11 Chaetochlorops 12 Tricimba 13 Olcella

14 Pseudogaurax 15 Acanthopeltastes 16 Onychaspidium

17 Pseudogaurax 18 Thaumatomyia 19 Loxobathmis

Figs.  92.11–19. Thoraces: dorsal view of scutum and scutellum of (11)  Chaetochlorops inquilinus (Coquillett), (MND, fig.  99.54);
(12) Tricimba trisulcata (Adams), (Nearctic, MND, fig. 99.53); (13) Olcella cinerea (Loew), (MND, fig. 99.55); and (14) Pseudogaurax sp.;
dorsal view of scutellum of (15)  Acanthopeltastes  sp.; (16)  Onychaspidium  sp.; (17)  P. signatus (Loew), (Nearctic, MND, fig.  99.47);
(18) Thaumatomyia glabra (Meigen), (Nearctic, MND, fig. 99.51); and (19) Loxobathmis sp. Figures 14–16, 19 illustrated by B. Koehler.
Abbreviations: acr s, acrostichal seta; ap sctl s, apical scutellar setae; dc s, dorsocentral seta; ial s, intra-alar seta; npl s, notopleural setae; pal s,
postalar seta; pprn s, postpronotal seta; sbap sctl s, subapical scutellar setae.
1142
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

10. Postocellar bristles parallel; gena sharply divided into pale, densely pruinose upper portion and
shining dark lower portion (Fig. 4); head and thorax shining dark brown or black; scutum with
dense setulae over dorsal surface. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
– Postocellar bristles convergent; gena not divided as above; head and thorax color and texture
various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

11. Mouthparts long, geniculate (as in Fig. 3); scutum with numerous extra bristles; at least two
postpronotals, two anterior notopleurals, three posterior notopleurals; prescutellar area with at
least one row of strong bristles between posterior dorsocentrals (Fig. 11); rarely collected. . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetochlorops Malloch
– Mouthparts not long and geniculate; scutum usually with at most two postpronotals, one anterior
notopleural, two posterior notopleurals; prescutellar area usually with at most only one pair of
prescutellar acrostichals between posterior dorsocentrals. . . . . . . . . . . . . . . . . . Fiebrigella Duda

12. Scutum with at least two postpronotals, two anterior and three posterior notopleurals, at least one
transverse row of strong bristles between posterior dorsocentrals; body usually mostly yellow
(two undescribed species of Olcella Enderlein have extra notopleural bristles and transverse
prescutellar row, but are distinguished by complete, narrow facial carina, projecting vibrissal
angle, and geniculate mouthparts, as in Fig. 3). . . . . . . . . . . . . . . . . . . . . . . . . Eugaurax Malloch
– Scutum with at most two postpronotals, one anterior and two posterior notopleurals, usually at
most one  pair of prescutellar acrostichals between posterior dorsocentrals (except in some
undescribed species of Olcella; see first half of couplet); body color various. . . . . . . . . . . . . . . 13

13. Arista broad, swordlike, with dense pubescence (Fig.  8); frontal triangle shining; scutellum
trapezoidal with apical scutellar bristles on small tubercles, or longer, with marginal bristles on
longer tubercles; notopleuron with one anterior, one posterior bristle . . . . . Elachiptera Macquart
– Arista narrow, sometimes slightly thickened but not swordlike, arista bare to long pubescent;
frontal triangle, scutellum, notopleural bristles various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

14. Frons with one to two fronto-orbital bristles clearly longer and stronger than the rest (as in Fig. 5);
frontal triangle usually shining; notopleuron with one anterior, one posterior bristle (upper
posterior bristle sometimes weakly developed). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
– Frons with all fronto-orbital bristles short and subequal (as in Fig. 3); frontal triangle shining or
pruinose; notopleuron usually with one anterior, two (occasionally one) posterior bristles. . . . 16

15. Frons with one long, strong fronto-orbital bristle (Fig. 5); frontal triangle shining, extending
almost to anterior margin of frons; gena usually with single row of setulae. . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Monochaetoscinella Duda
– Frons with two, rarely three long, strong fronto-orbital bristles; frontal triangle usually shining,
rarely partly or entirely pruinose, one-half to two-thirds as long as frons; gena usually with two
rows of distinct setulae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Biorbitella Sabrosky

16. Facial carina broad; gena broad and rugose; cephalic bristles reduced, only outer verticals well
developed, head with dense setulae (Fig. 7); 4–5 mm long; rarely collected . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Medeventor Wheeler
– Facial carina narrow and knifelike or absent; gena usually narrow and not rugose; head with at
least ocellar and outer vertical bristles well developed, usually other bristles present and dark,
head not densely setulose; less than 3 mm long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

17. Head mostly yellow, frontal triangle shining yellow with black median stripe; vibrissal angle
projecting and gena broad; mouthparts long, geniculate; scutum yellow with five black stripes,
median stripe continued onto scutellum; rarely collected. . . . . . . . . . . . . . . Metasiphonella Duda
– Not with the above combination of characters. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
CHLOROPIDAE (frit flies, grass flies, eye gnats) 92 1143

18. Ocellar bristles long, proclinate, divergent; rarely collected in Central America (specimens with
missing or misdirected ocellar bristles should be keyed first to couplet 19 to eliminate those easily
distinguished genera). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
– Ocellar bristles erect or reclinate, usually short. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

19. Anepisternum extensively setulose; scutellum long, triangular, punctate dorsally; wing with M1
bent anteriorly at crossvein dm-cu (Fig. 26) . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhodesiella Adams
– Anepisternum bare, or with few setulae near dorsal margin; scutellum short, rounded, smooth
dorsally; wing with M1 straight or concavely curved at or beyond dm-cu (as in Fig. 24). . . . . . 20

20. Wing with crossvein r-m in basal one-quarter or one-third of cell dm, cell dm narrow (Fig. 25);
gena usually narrow, often silvery-gray; tibial organ varied in size . . . . . . Notaulacella Enderlein
– Wing with crossvein r-m near middle of cell dm (as in Fig. 21), cell dm broader; gena usually
broader, same color as most of head; tibial organ large, oval . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

21. Wing with alula absent and anal angle greatly reduced, R2+3 not greatly elongate, second costal
sector approximately equal to third (Fig. 21); head and thorax shining black; hind tarsal segments
not swollen, all segments the same color, usually black. . . . . . . . . . . . . . . . . Stenoscinis Malloch
– Wing with alula and anal angle well-developed, R2+3 greatly elongate, second costal sector at least
2.5–3.0 times as long as third; head and thorax with at least some yellow; hind tarsal segments
swollen, basal segments yellow, distal segments black. . . . . . . . . . . . . . . . . . . . Psilacrum Becker

22. Hind tibia with dark, ventral apical spur, shorter than tibial diameter, sometimes extremely short
and indistinct (Fig. 31). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
– Hind tibial spur absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26

23. Body mostly yellow, scutum yellow with four black stripes; gena twice as wide as first flagellomere
and almost one-half eye height; eye oval; frons projecting in front of eye in lateral view; female
cerci sclerotized, black, knifelike; rarely collected in Central America . . . . . . Opetiophora Loew
– Body usually primarily black or gray, rarely yellow, scutum often black or gray, rarely yellow
with four black stripes; gena usually narrower than first flagellomere; eye usually round; frons not
projecting; female cerci not modified as above. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

24. Frontal triangle shining, rest of frons and gena bare but microscopically striate so that surface
appears subshining; thorax mostly shining; scutal setae and setulae often arising from small
punctures or tubercles (species assigned by previous authors to Lioscinella Duda key here). . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Liohippelates Duda, in part
– Frontal triangle pruinose or shining, rest of frons densely pruinose; gena dull, pruinose; thorax
with at least some pruinosity in posterior part of scutum, usually more extensively pruinose;
scutal setae not arising from punctures or tubercles, surface of scutum usually appearing smooth,
although sometimes heavily pruinose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

25. Thorax and legs usually mostly black; frontal triangle usually entirely shining, at least with round
shining spot at anterior apex; gena usually as broad as first flagellomere, densely pruinose and
much paler than rest of head . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apallates Sabrosky, in part
– Thorax and legs usually yellow or gray; frontal triangle dull (sometimes with three shining spots
adjacent to ocelli or single median shining stripe); gena usually narrower than first flagellomere
and same color as rest of head (some undescribed, small, yellow species tentatively assigned to
Conioscinella key here). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hippelates Loew, in part
1144
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

20 Trigonomma 21 Stenoscinis

22 Siphunculina 23 Gaurax

24 Loxobathmis 25 Notaulacella

26 Rhodesiella

Figs. 92.20–26. Wings: dorsal view of (20) Trigonomma fossulatum (Loew), (MND, fig. 99.43); (21) Stenoscinis longipes (Loew), (Nearctic,
MND, fig. 99.33); (22) Siphunculina striolata (Wiedemann), (MND, fig. 99.32); (23) Gaurax festivus Loew, (Nearctic, MND, fig. 99.34);
(24) Loxobathmis sp.; (25) Notaulacella sp.; and (26) Rhodesiella brimleyi Sabrosky, (Nearctic, MND, fig. 99.37). Figure 24 illustrated by
R. Vargas; Figure 25 illustrated by B. Koehler.
CHLOROPIDAE (frit flies, grass flies, eye gnats) 92 1145

fem
tb org
tb

tb spur

ts

27 Meromyza 28 Thaumatomyia 29 Hippelates

30 Liohippelates 31 Apallates

Figs. 92.27–31. Hind legs: anterior view of (27) Meromyza americana (Fitch), (Nearctic, MND, fig. 99.62); dorsal view of (28) Thaumatomyia
glabra (Meigen), (Nearctic, MND, fig. 99.60); anterior view of (29) Hippelates plebejus Loew, (Nearctic, MND, fig. 99.57); (30) Liohippelates
bishoppi (Sabrosky), (MND, fig. 99.58); and (31) Apallates coxendix (Fitch), (MND, fig. 99.59).
Abbreviations: fem, femur; tb, tibia; tb org, tibial organ; tb spur, tibial spur; ts, tarsus.
1146
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

26. Scutum with distinct incised lines of medial and dorsocentral punctures, sometimes with
additional intra-alar lines (Fig. 12); if lines only shallowly incised (some Olcella, Fig. 13), then
frontal triangle and scutum dull. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
– Scutum smooth dorsally, median and dorsocentral rows of setulae usually present but not
distinctly incised (as in Fig. 1) (some Coryphisoptron have shallow incised rows of setulae, but a
shining frontal triangle and scutum) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

27. Head usually elongate; scutellum long rectangular or trapezoidal with apical bristles at
posterolateral corners; anal angle of wing reduced; abdomen long and cylindrical; male genitalia
usually enlarged. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Enderleiniella Becker
– Head usually short, taller than long; scutellum rounded or short trapezoidal, with apical bristles
usually closely approximated; anal angle of wing usually broad; abdomen broad; male genitalia
small. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28

28. Vibrissal angle projecting; facial carina complete and narrow; mouthparts usually long, sclerotized,
geniculate (Fig. 3); notopleuron with one anterior, two posterior bristles (one undescribed species
has 0+1 notopleurals). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Olcella Enderlein, in part
– Vibrissal angle not projecting, usually rounded; facial carina small or absent; mouthparts small,
not geniculate; notopleuron with one anterior, one to two posterior bristles. . . . . . . Tricimba Lioy

29. Scutellum distinctly flattened dorsally, with four, usually long, marginal projections bearing stout
bristles; male genitalia enlarged in most species, surstylus and cercus usually branched. . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Agrophaspidium Wheeler & Mlynarek
– Scutellum not usually flattened dorsally, marginal projections or tubercles either absent, much
shorter than scutellum, or more than four in number (Figs. 16, 19); male genitalia usually not
enlarged, surstylus and cercus usually simple. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

30. Frontal triangle, and usually scutum, dull, pruinose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

– Frontal triangle, and usually scutum, shining . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

31. Arista thickened and pubescent; scutum flattened in posterior half; northern Mexico. . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Eribolus Becker]
– Arista thin, bare or pubescent; scutum convex posteriorly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32

32. Notopleuron with one anterior, one posterior bristle; rarely collected in Central America. . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Incertella Sabrosky
– Notopleuron with one anterior, two posterior bristles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

33. Vibrissal angle projecting; facial carina complete, narrow; mouthparts usually elongate, geniculate
(Fig. 3); usually gray or brown flies, with some yellow in some species . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Olcella Enderlein, in part
– Vibrissal angle usually rounded; facial carina usually absent or incomplete (species with a
complete facial carina and projecting vibrissal angle are usually entirely yellow); mouthparts
usually short, not geniculate; body color various (several morphologically dissimilar species,
probably representing multiple genera, key here) . . . . . . . . . . . . . . . . . . . . . . Conioscinella Duda

34. Scutellum longer than broad, triangular, with pale apical bristles closely approximated at base and
divergent (Figs. 14, 17); eye hairy; thorax usually with yellow and black pattern dorsally; arista
usually apical. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudogaurax Malloch
– Scutellum usually broader than long, rounded (if longer than broad, then marginal tubercles
present), apical bristles not closely approximated at base; eye hairy or bare; thorax usually dark,
if with black and yellow pattern (Gaurax), then cell R1 broad (Fig. 23); arista usually dorsal. . 35
CHLOROPIDAE (frit flies, grass flies, eye gnats) 92 1147

35. Scutellum longer than broad, triangular, with four to eight marginal tubercles, each bearing
bristles (Fig. 16); scutellum black at base, yellow at apex. . . . . . . . . . . Onychaspidium Enderlein
– Scutellum broader than long; rounded, marginal tubercles usually absent (present in Loxobathmis,
Fig. 19); scutellum uniformly dark or yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36

36. Wing short and broad, veins R4+5 and M1 curved anteriorly, cell br noticeably broadened at middle,
crossvein dm-cu strongly reflexed, almost parallel to hind margin of wing (Fig. 24); scutellum
with bristles arising from distinct tubercles (Fig. 19) . . . . . . . . . . . . . . . . . Loxobathmis Enderlein
– Wing narrower, veins R4+5 and M1 almost straight distally, cell br not usually broadened in middle,
crossvein dm-cu not reflexed, almost parallel with crossvein r-m; scutellum usually without
tubercles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37

37. Gena curved inwards in anterior half; frons strongly sloping downwards in anterior half; crossvein
dm-cu absent; minute flies 1.0–1.2 mm long. . . . . . . . . . . . . . . . . . . . . . . . . . . Oscinicita Wheeler
– Gena vertical below eye; frons relatively horizontal in lateral view; crossvein dm-cu present; size
various, usually more than 1.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38

38. Clypeus pointed and projecting anteriorly; forecoxa with stout spinelike anterior bristle . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Malloewia Sabrosky, in part
– Clypeus rounded anteriorly; forecoxa without spinelike anterior bristle. . . . . . . . . . . . . . . . . . . 39

39. Mouthparts long, geniculate; gena linear, much narrower then first flagellomere; vibrissal angle
acute. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Malloewia Sabrosky, in part
– Mouthparts short, not geniculate; gena broader, usually at least half as high as first flagellomere;
vibrissal angle rounded or only slightly projecting. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40

40. Gena at least as broad as first flagellomere, densely pruinose and pale, contrasting with dark face
and frons. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apallates Sabrosky, in part
– Gena usually narrower, at most half as broad as first flagellomere, bare or sparsely pruinose, usually
same color as face and frons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41

41. Scutum densely and evenly covered dorsally with long setulae; eye hairy; gena narrow, linear . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Coryphisoptron Enderlein
– Scutum with setulae either in rows, or sparse, or short; eye usually bare; gena various. . . . . . . 42

42. Frontal triangle usually extending just over halfway to anterior margin of frons; cell  R1 broad
(Fig. 23); eye hairy; thorax usually with yellow and black pattern; male genitalia sometimes
enlarged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gaurax Loew
– Frontal triangle usually extending almost to anterior margin of frons; cell R1 narrow; eye bare;
thorax entirely black; male genitalia small . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43

43. Notopleuron with one anterior, one posterior bristle. . . . . . . . . . . . . . . . . . . Rhopalopterum Duda
– Notopleuron with one anterior, two posterior bristles. . . . . . . . . . . . . . . . . . . . . Oscinella Becker

44. Hind femur inflated, much broader than midfemur, with two rows of ventral tubercles; hind tibia
curved to fit against enlarged femur (Fig. 27). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
– Hind femur not inflated, not significantly broader than midfemur, ventral rows of tubercles absent,
rarely with one or more spines; hind tibia straight or only slightly curved. . . . . . . . . . . . . . . . . 46
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45. Antenna with first flagellomere short, round or slightly oval; body color mostly pale, green or
yellow, with black stripes; male genitalia with large, sclerotized postgonite, often visible in dried
specimens; rarely collected in region. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Meromyza Meigen
– Antenna with first flagellomere elongate, usually at least twice as long as high; body color usually
darker, extensively black or brown, with yellow on head, thorax, and abdomen; male genitalia
without large sclerotized postgonites. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neodiplotoxa Malloch

46. Crossvein dm-cu absent or separated from crossvein r-m by less than length of dm-cu. . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diplotoxa Loew
– Crossvein dm-cu present, separated by more than its own length from crossvein r-m (Fig. 20). . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47

47. Frontal triangle with at least two rows of setulae on surface; tibial organ, if present, narrow,
linear. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
– Frontal triangle bare or with single row of setulae near lateral margin; if two rows present (some
Thaumatomyia), then tibial organ large, oval (Fig. 28). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49

48. First flagellomere elongate, arista pubescent (as in Fig. 10); frons projecting anteriorly; palpus
elongate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ectecephala Macquart
– First flagellomere short, arista bare; frons broad, not projecting; palpus usually short; rarely
collected in Central America. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Homaluroides Sabrosky

49. Arista broad, sword-shaped (as in Fig. 8), first flagellomere kidney-shaped. . . Steleocerellus Frey
– Arista sometimes long-haired or densely pubescent, but not sword-shaped, first flagellomere
round or oval . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50

50. Frontal triangle usually dark, metallic, with longitudinal median sulcus; gena usually linear, much
narrower than height of first flagellomere; scutum uniformly dark, with dense granular texture;
body entirely or mostly black. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trigonomma Enderlein
– Frontal triangle not metallic, usually without median sulcus; gena usually broader, at least half
height of first flagellomere; scutum usually yellow with dark stripes, rarely completely black;
smooth or slightly punctate; body almost always with more extensive yellow color, especially on
head and sides of thorax. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51

51. Scutellum flat on dorsal surface, with sharp dorsolateral margin, apical scutellar bristles closely
approximated at their bases (Fig. 18); tibial organ broad, oval (Fig. 28). . .Thaumatomyia Zenker
– Scutellum usually convex dorsally without sharp dorsolateral margin, apical scutellar bristles not
as closely approximated; tibial organ narrow and linear, or absent. . . . . . . . . . . . . . . . . . . . . . . 52

52. Arista thick, densely setulose, white with yellow base; first flagellomere usually at least 1.5 times
longer than broad (Fig. 10). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parectecephala Becker
– Arista not noticeably thickened, not densely setulose, usually black; first flagellomere round or
slightly oval; rarely collected in Central America. . . . . . . . . . . . . . . . . . . . . . . . Chlorops Meigen

Synopsis of the fauna 2008). However, recent collections and studies of museum
specimens indicate that this is only a fraction of the total
The number of species of Chloropidae in Central America diversity of Central American Chloropidae, which apparently
is unknown, but it is undoubtedly one of the most species-rich exceeds several hundred species.
families of Brachycera in the region. Sabrosky & Paganelli
(1984) recorded 125 named species from Central America Acanthopeltastes Enderlein. The long, fingerlike projec-
and Neotropical Mexico, and several species have been tions on the scutellum (Fig.  15) and the shape of the head
described since that time (e.g., Sabrosky, 1994, 1996; Wheeler, (kidney-shaped eyes, high ventral facial margin) are distinctive
2007; Mlynarek & Wheeler, 2008, 2009; Wheeler & Mlynarek, in this genus. The five described species recorded from
CHLOROPIDAE (frit flies, grass flies, eye gnats) 92 1149

Central America can be identified using the key in Enderlein Conioscinella Duda. This genus is species-rich, abundant,
(1911), although there are additional undescribed species. and morphologically diverse in Central America (as well as
Acanthopeltastes is endemic to the Neotropical Region. the rest of the world). However, this is partly due to the fact
that the genus is a repository for species that cannot be accom-
Agrophaspidium Wheeler & Mlynarek. This genus, so modated in other genera that are defined on small sets of
far known only from six species in Central America, is distinguishing characters. Phylogenetic analyses will probably
recognized by the structure of the scutellum and the distinctive result in the division of Conioscinella into multiple genera
male genitalia (Wheeler & Mlynarek, 2008). Although it may or the synonymy of smaller genera with it. Six described
be the sister group to the Neotropical genus Enderleiniella, species have been recorded from the region, but this is only
the relationship of these two genera to other Oscinellinae is a small fraction of the actual species richness. Specimens are
unknown (Wheeler & Mlynarek, 2008). abundant in habitats associated with decaying organic material,
especially in disturbed areas.
Apallates Sabrosky. This New World genus is diverse
and abundant throughout the region. Although 13 described Coryphisoptron Enderlein. This poorly-known genus is
species have been recorded in Central America, there are endemic to the Neotropical Region. Four described species
apparently many undescribed species. Species with a minute have been recorded from Central America, but others are
hind tibial spur (and at least one species in which the spur apparently present. The limits of this genus are problematic,
is absent) are difficult to key out, but can be recognized as and the species assigned to Coryphisoptron by earlier authors
Apallates by the color pattern and texture of the head. Specimens are a diverse assemblage without clear distinguishing
of Apallates are especially abundant in disturbed areas such characters (see also comments under Onychaspidium).
as roadsides and margins of agricultural fields.
Diplotoxa Loew. The limits of Diplotoxa are confus-
Apotropina Hendel. This cosmopolitan genus is the only ing especially with regard to the genera Elliponeura Loew
representative of the subfamily Siphonellopsinae in the region, and Pseudopachychaeta Strobl, which are included here in
so identification to the generic level is easy. Although five Diplotoxa sensu lato (Spencer, 1986; Wheeler  & Forrest,
2003). The loss of dm-cu (Elliponeura) or the development
species have been recorded from Central America, at least 4
of longer fronto-orbital setae (Pseudopachychaeta) is probably
times that many are represented in museum collections.
not sufficient for generic recognition, especially given that
Specimens of Apotropina are occasionally collected in
other characters in these genera vary widely. The male genitalia
large swarms, especially over dark vertical surfaces like are varied and insufficiently studied to provide reliable
cave openings or overhanging banks. defining characters. There are few Central American records
Biorbitella Sabrosky. This New World genus is distinguished of Diplotoxa sensu lato, and all are of unidentified species.
by the two long fronto-orbital bristles, although other external Larvae of known New World species are phytophagous in
characters (texture of frontal triangle, presence of small hind sedges (Cyperaceae).
tibial spur) are various. Four species of Biorbitella have Ectecephala Macquart. This is one of the most species-
been recorded from Central America but additional species rich genera of Chloropinae in the Neotropical Region,
are present. Specimens in this genus are often abundant in although only five species have been recorded from Central
disturbed habitats. America. Ectecephala is divided into two externally different
species groups: the E.  albistylum group comprises species
Cadrema Walker. Only one species of this genus has been with elongated antennae and a narrow frontal triangle; the
recorded in the region, the pantropical species C. pallida E.  capillata group includes species with shorter antennae
(Loew). It is most commonly collected near seashores. and a broad frontal triangle. Predictably, there are species
intermediate between these two groups.
Chaetochlorops Malloch. Two described species of this
New World genus have been recorded from Central America Elachiptera Macquart. Species of this genus are easily
(Hibbert  & Wheeler, 2007); the other described species is distinguished from other Central American Oscinellinae on
Nearctic. Chaetochlorops is apparently closely related to the basis of the broad arista, although this character occurs
Fiebrigella. Members of this genus are probably inquilines in additional genera in other regions. Steleocerellus, present
in galls and feeding galleries made by other insects (Sabrosky, in Costa Rica, also has a broad arista but is in the sub-
1950; Hibbert & Wheeler, 2007). family Chloropinae. Eight species of Elachiptera have been
recorded in Central America (Sabrosky & Paganelli, 1984;
Chlorops Meigen. This primarily temperate genus is Mlynarek  & Wheeler, 2008), and the Costa Rican species
much more diverse in the Nearctic Region; only one described were revised by Mlynarek  & Wheeler (2008). Although
species has been recorded in Central America or southern there is little biological information on the Central American
Mexico, and there are few specimens in museum collections species, most known species in other geographic regions are
from the region. Species whose biology is known are saprophagous secondary invaders of damaged plant tissues,
phytophagous in grasses or sedges. especially in moist habitats.
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Enderleiniella Becker. This genus of two described within this group remain confusing. Paganelli  & Sabrosky
species (and at least 10 undescribed species) is endemic to (1993) described three new species of Hippelates from
the Neotropical Region. See comments under Agrophaspidium Brazil and provided a partial key to species. Becker (1912),
on the relationships of these two genera. Some museum Paganelli & Sabrosky (1993), and Wheeler & Forrest (2003)
specimens of Enderleiniella examined during this study were treated Olcanabates Enderlein as a synonym of Hippelates,
reared from flowers of Araceae, although it is not known and that decision is followed here. Six described species of
whether the larvae were phytophagous or saprophagous. Hippelates have been recorded from Central America, but
the group is considerably more diverse in the region.
Eugaurax Malloch. Members of this genus are easily
recognized on the basis of the extra thoracic bristles and Homaluroides Sabrosky. Although no described species
primarily yellow body (species of Chaetochlorops, Hippelates, have been recorded from Central America, I have seen
Olcella, and Fiebrigella with extra thoracic bristles are specimens from southern Mexico (Chiapas), Colombia, and
primarily black or gray). One described species of Eugaurax Jamaica, and Homaluroides may be recorded elsewhere in
has been recorded from Central America, but at least six to the region with additional collecting in appropriate habi-
seven species are present in Costa Rica alone. The north- tats. Homaluroides is usually associated with arid regions
ern South American species E. setigena Sabrosky is a stem or ocean beaches and is rarely collected throughout its New
miner in water hyacinth (Eichhornia  spp., Pontederiaceae) World range.
(Sabrosky 1974).
Incertella Sabrosky. Although this genus is abundant and
Fiebrigella Duda. This genus is distinctive and easily diverse in North America, only a single described species has
recognized on the basis of the head structure, but there is been recorded in Central America and specimens are rarely
great morphological diversity of species within the group. collected. Additional species from the region are represented
Although only one described species has been recorded in museum collections, but the limits of Incertella are not
from Central America, the true species richness is apparently clearly defined and some of the Central American species
much higher. The biology of most species is unknown, but may eventually have to be assigned to other genera. Nearctic
some are predaceous on grasshopper eggs (Sabrosky, 1967), species whose biology is known are saprophagous.
and others may be inquilines or predators of egg masses of
other arthropods. Liohippelates Duda. This genus, along with Hippelates,
contains most of the species referred to as “eye gnats”.
Gaurax Loew. Although usually defined on the basis of There has been confusion as to the limits of Liohippelates
wing venation, the generic limits of Gaurax are tenuous and Lioscinella, as well as the correct names and status of
because of considerable variation in most other characters. many included species. The two genera are combined in the
The enlarged male epandrium applies to the Nearctic fauna key because the only character distinguishing them is the
but not necessarily to the Neotropical species (Wheeler  & length of the hind tibial spur (long in Liohippelates, short
Forrest, 2003), although some of the Central American species in Lioscinella), a highly variable character in many genera.
have highly autapomorphic male genitalia. The biology of Ten species of Liohippelates and three of Lioscinella have
the three recorded Central American species is unknown, been recorded in Central America, and they are abundant in
but Nearctic species of Gaurax are more frequently found a range of habitats. Additional species are undoubtedly
in forest areas, where they have been reared in association present, but the group is in serious need of systematic
with feeding tunnels and frass of other insects (Sabrosky, revision to resolve species limits and distributions. Known
1951). Specimens of Gaurax are rarely collected in Central larvae are saprophagous.
America.
Loxobathmis Enderlein. This endemic Neotropical
Goniaspis Duda. This genus includes species closely genus is known from a single species, L. obliquum Enderlein,
related to Elachiptera. The southern Nearctic species from Brazil and Costa Rica. There is at least one additional
Elachiptera eulophus (Coquillett) is intermediate in appearance species in the region. The wing venation is characteristic for
between Goniaspis and Elachiptera, having the long hind this genus, but external characters do not give any indication
tibial spur of Goniaspis, the broad arista of Elachiptera, and as to its relationships to other genera.
the trapezoidal scutellum found in species of both genera
(Wheeler & Forrest, 2002). Five species of Goniaspis have Malloewia Sabrosky. Two described species have been
been recorded from Central America (Mlynarek & Wheeler, recorded from Central America, although the generic limits
2009). as defined by Sabrosky (1980) do not hold for either one.
Although I am not convinced that the Neotropical species
Hippelates Loew. This genus includes some of the species are congeneric, I have left them in Malloewia for the present.
commonly referred to as eye gnats. Many species of Hippelates An additional problem with recognition of Malloewia is that
cited in older papers are now assigned to other genera some of the Nearctic species overlap in some characters with
(primarily Liohippelates and Apallates), and the generic limits species previously assigned to Lioscinella (= Liohippelates).
CHLOROPIDAE (frit flies, grass flies, eye gnats) 92 1151

Little is known of the biology of Malloewia, although Neo- (Sivinski et al., 1999), and some species appear to be attracted
tropical specimens have been collected in a range of habitats, even by damaged insects in sweep nets.
often in large numbers. They are probably saprophagous.
Onychaspidium Enderlein. This is one of the more easily
Medeventor Wheeler. This distinctive genus is represented recognized genera in the region because of the long, flattened
by a single described species from a number of localities in scutellum with distinct marginal tubercles (Fig.  16). Some
Costa Rica. Because of its large size and setulose body, it is species of Coryphisoptron have a similar scutellar structure,
probably the most atypical Central American chloropid genus although the tubercles are not as well developed. The limits
(Wheeler, 2007). Nothing is known of its biology, although of these two genera should be assessed. Onychaspidium is
the apparently closely related Palearctic genera Lipara Meigen endemic to the Neotropical Region.
and Pseudeurina de Meijere are both phytophagous.
Opetiophora Loew. This genus is known from a single
Meromyza Meigen. This is primarily a temperate genus, species, O. straminea Loew, from the southern Nearctic Region
with most species restricted to the Holarctic Region. One and Neotropical Mexico. Although the female ovipositor is
species, M. opacula Fedoseeva, was described from southern
clearly modified for piercing, the biology of this species is
Mexico (Chiapas). I have seen no additional specimens from
poorly known.
Central America or southern Mexico. Larvae of Meromyza
species are phytophagous in stems of grasses, including Oscinella Becker. Although diverse elsewhere (especially
cereal crops. the Holarctic Region), there are few Neotropical species as-
Metasiphonella Duda. This monotypic genus is endemic signed to Oscinella except for the aberrant O. blanda group.
to the Neotropical Region, and the single described species, Placement of the O. blanda group in Oscinella is question-
M. magnifica Duda, is easily recognized by the characters in able, but the species are retained here for consistency with
the key. It may be related to the Tricimba group of genera. published catalogs (and the absence of any phylogenetic
evidence to the contrary).
Monochaetoscinella Duda. This small genus is dis-
tinguished by the single, long fronto-orbital bristle. Only Oscinicita Wheeler. This monotypic genus was described
one described species, the widespread New World species by Wheeler (2007) for a single species of tiny flies from a
M. anonyma (Williston), has been recorded in Central America cacao plantation in Costa Rica. The characteristic wing
and is found throughout the region. venation (dm-cu absent, CuA1 straight and incomplete) is
probably related to the minute size of individuals of this
Neodiplotoxa Malloch. Although there are no published species.
records of this genus from Central America, several undescribed
species are present in the region. Four species have been Parectecephala Becker. There are three described species of
described in western North America and temperate South Parectecephala in Costa Rica, but the genus is more diverse
America, but the actual number is probably at least 4–5 in the Nearctic Region and in South America. It is closely
times higher. Although Neodiplotoxa appears to be related related to Chlorops, but the limits of these genera will not
to Diplotoxa sensu lato based on genitalic characters, exter- be clearly established without a generic revision of the
nal characters are not so clear-cut. Chloropinae.
Notaulacella Enderlein. This endemic Neotropical genus Pseudogaurax Malloch. Species of Pseudogaurax are
provides an excellent example of the potential diversity of not frequently collected in the field. They are usually reared
Chloropidae in Central America. Previously known from
from the egg masses of spiders, with scattered records from
four described species in the region, Sabrosky (1994, 1996)
mantid oothecae and Lepidoptera cocoons. One species of
described 17 new species, mostly from a single canopy
Pseudogaurax has been reared from amblypygids (Arachnida:
fogging project in Panama, an indication of how species-rich
the Chloropidae may be in unexplored habitats such as the Amblypygi) in Costa Rica. Sabrosky (1966, 1990) published
forest canopy. There are also several new species represented keys to the described New World species of Pseudogaurax.
by relatively few specimens in museum collections examined Five species have been recorded in Central America;
during this study. however, based on examination of museum specimens, there
are at least 4 times that many species in the region.
Olcella Enderlein. This genus is especially diverse in the
Neotropical Region. Ten species have been recorded from Psilacrum Becker. This genus was recorded by
Central America and Mexico, but there are several undescribed Sabrosky  & Paganelli (1984) as Oscinoides. There is a
species. Olcella is one of the dominant chloropid genera in single described species in Central America, although at
arid and saline areas of Central America and the southwestern least another five to six species are represented in museum
Nearctic Region. Adults of some species are apparently collections. Psilacrum is more diverse in the Old World
attracted to the prey of spiders and predaceous insects tropics (Ismay, 1986).
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Rhodesiella Adams. No described species of this genus Stenoscinis Malloch. Two species of this small genus
have been recorded in Central America, but undescribed have been described in Central America, but the generic
species are present, although rarely collected, throughout the limits are confused with respect to Rhopalopterum (see notes
region (Sabrosky & Paganelli, 1984). Rhodesiella is most under that genus).
diverse in the Old World tropics, and there is a single recorded
Nearctic species. Early Neotropical records (e.g., Becker, Thaumatomyia Zenker. This is one of the more morpho-
1912) of Meroscinis de Meijere, a junior synonym of logically distinctive genera of Chloropinae. There are few
Rhodesiella, refer to species now assigned to other genera. described species in the region, although there are apparently
Rhopalopterum Duda. Although the type species, undescribed species in Central America. The larvae of
R. limitatum Duda, is Neotropical, there are few described species whose habits are known are predators on soil-
species of this genus in the region, and the status of some of dwelling Homoptera.
these is questionable. There is also some confusion as to the
limits of Rhopalopterum and Stenoscinis, especially based Tricimba Lioy. This is a large and morphologically diverse
on the Neotropical species (Sabrosky, 1961). The two genera genus (Ismay, 1993) whose limits worldwide are poorly
are currently distinguished on the orientation of the ocellar resolved. In the Neotropical Region, Tricimba is divided into
bristles, but there is considerable variation and overlap in two subgenera: Tricimba sensu stricto includes mostly small,
some other morphological characters. dark species, whereas Pentanotaulax Enderlein includes
larger, often yellow and black species. The monotypic genus
Siphunculina Rondani. One described species of Siphun- Discogastrella Enderlein, known from Costa Rica, keys to
culina has been recorded from Central America: the widespread
Tricimba; its status is unresolved.
pantropical species S. striolata (Wiedemann). Specimens of this
genus are occasionally encountered as eye gnats, although much Trigonomma Enderlein. This genus, along with Ectecepha-
less so than species of Hippelates or Liohippelates.
la, is one of the few species-rich genera of the subfamily
Steleocerellus Frey. There are no published records of Chloropinae in Central America, with six species recorded
this genus from Central America, but there are museum in the region. Trigonomma is almost entirely endemic to
specimens from Costa Rica, Trinidad, and Venezuela. All the Neotropical Region, with a single species extending
specimens examined appear to belong to a single species, into the southernmost part of the Nearctic Region.
probably S. flavifrons (Duda).

Literature cited

Becker, T. 1912. Chloropidae. Eine monographische Studie. IV. Enderlein, G. 1911. Klassification der Oscinosominen. Sitzungber-
Teil. Nearktische Region. V. Teil. Neotropische Region. Annales ichten der Gesellschaft Naturforschender Freunde 4: 185–244.
Musei Nationalis Hungarici 10: 21–256. Ferrar, P. 1987. A guide to the breeding habits and immature stages
Brake, I. 2000. Phylogenetic systematics of the Milichiidae of Diptera Cyclorrhapha. Entomonograph 8(1). E.J.  Brill  /
(Diptera, Schizophora). Entomologica scandinavica, Supplement Scandinavian Science Press, Leiden, Copenhagen, 478 pp.
57: 1–120. Hibbert, A.C. & T.A. Wheeler. 2007. Systematics of Chaetochlorops
Buck, M. 2006. A new family and genus of acalypterate flies from Malloch, 1914 (Diptera: Chloropidae) and morphology of
the Neotropical region, with a phylogenetic analysis of Car- secondary sexual structures in the male abdomen. Studia
noidea family relationships (Diptera: Schizophora). Systematic dipterologica 14: 97–105.
Entomology 31: 377–404. Ismay, J.W. 1986. New Psilacrum from the Old World (Diptera:
Curran, C.H. 1928. Insects of Porto Rico and the Virgin Islands.
Chloropidae). Esakia 24: 19–35.
Diptera or two-winged flies. Scientific Survey of Porto Rico and
Ismay, J.W. 1993. Revision of Tricimba Lioy and Aprometopis
the Virgin Islands 11: 1–118.
Duda, O. 1929. Die Ausbeute der deutschen Chaco-Expedition Becker (Diptera: Chloropidae) from Australia and the Papuan
1925/1926 (Diptera). X.  Chloropidae. (Fortsetzung). Konowia region. Invertebrate Taxonomy 7: 297–499.
8: 165–169. Malloch, J.R. 1934. Acalyptrata (concluded), pp. 393–489. Diptera
Duda, O. 1930. Die neotropischen Chloropiden (Dipt.). Folia of Patagonia and South Chile, Part VI  – Fascicle 5. British
Zoologica et Hydrobiologica 2: 46–128. Museum of Natural History, London.
Duda, O. 1931. Die neotropischen Chloropiden (Dipt.). 1. Fortsetzung: McAlpine, J.F. 1989. Phylogeny and classification of the
Nachtrag, Ergänzungen, Berichtigungen und Index. Folia Muscomorpha, pp. 1397–1518. MND, Volume 3.
Zoologica et Hydrobiologica 3: 159–172. Mlynarek, J.J. & T.A. Wheeler. 2008. Revision of the Costa Rican
Duda, O. 1933. Die neotropischen Chloropiden (Dipt.). 2. Fortsetzung. species of Elachiptera (Diptera: Chloropidae). Zootaxa 1754:
Folia Zoologica et Hydrobiologica 5: 41–47. 41–51.
CHLOROPIDAE (frit flies, grass flies, eye gnats) 92 1153

Mlynarek, J.J. & T.A. Wheeler. 2009. Revision of the Neotropical Sabrosky, C.W. 1990. New synonymy and new species in Neotropical
genus Goniaspis Duda (Diptera: Chloropidae). Zootaxa 2033: Chloropidae. Memórias do Instituto Oswaldo Cruz 84 (Supplement
26–40. 4): 463–465.
Paganelli, C.H.M. 1985. Revisão dos Gêneros Neotropicais de Sabrosky, C.W. 1994. A synopsis of the genus Notaulacella Ender-
Chloropinae (Diptera, Chloropidae). D.Sc. thesis, Instituto de lein in Central America: The diversity of a canopy fauna
Biosciências, Universidade de São Paulo, 243 pp. (Diptera: Chloropidae). Proceedings of the Entomological
Paganelli, C.H.M.  & C.W. Sabrosky. 1993. Hippelates flies Society of Washington 96: 428–439.
(Diptera: Chloropidae) possibly associated with Brazilian Sabrosky, C.W. 1996. Two additional species of Notaulacella
Purpuric Fever. Proceedings of the Entomological Society of Enderlein (Diptera: Chloropidae) from Panama. Proceedings of
Washington 95: 165–174. the Entomological Society of Washington 98: 471–472.
Sabrosky, C.W. 1950. A synopsis of the chloropid genera
Sabrosky, C.W. & C.H.M. Paganelli. 1984. 81. Family Chloropidae.
Chaetochlorops and Eugaurax (Diptera). Journal of the
Washington Academy of Sciences 40: 183–188. In Papavero,  N. (editor). A Catalogue of the Diptera of the
Sabrosky, C.W. 1951. A revision of the Nearctic species of the Americas South of the United States. Departamento de
genus Gaurax (Diptera, Chloropidae). American Midland Zoologia, Secretaria da Agricultura, São Paulo, 63 pp.
Naturalist 45: 407–431. Sivinski, J., S.A. Marshall, & E. Petersson. 1999. Kleptoparasitism
Sabrosky, C.W. 1961. A new Nearctic species of Stenoscinis, with and phoresy in the Diptera. Florida Entomologist 82: 179–197.
key to the species of the Western Hemisphere (Diptera, Spencer, K.A. 1986. The Australian Chloropinae (Diptera:
Chloropidae). Entomological News 72: 19–23. Chloropidae). Journal of Natural History 20: 503–615.
Sabrosky, C.W. 1966. Three new Brazilian species of Pseudogaurax Wheeler, T.A. 2007. Two new genera of oscinelline Chloropidae
with a synopsis of the genus in the Western Hemisphere (Diptera, (Diptera) from Costa Rica. Zootaxa 1413: 47–53.
Chloropidae). Papeis Avulsos de Zoologia 19: 117–127.
Wheeler, T.A.  & J. Forrest. 2002. A new species of Elachiptera
Sabrosky, C.W. 1967. Two new and economically significant
Chloropidae, with the description of a new genus (Diptera). from the Galápagos Islands, Ecuador, and the taxonomic status
Journal of the Kansas Entomological Society 40: 151–156. of Ceratobarys Coquillett (Diptera: Chloropidae). Zootaxa 98:
Sabrosky, C.W. 1974. Eugaurax setigena (Diptera: Chloropidae), 1–9.
a new stem miner in water hyacinth. Florida Entomologist Wheeler, T.A. & J. Forrest. 2003. The Chloropidae (Diptera) of the
57: 347–348. Galápagos Islands, Ecuador. Insect Systematics and Evolution
Sabrosky, C.W. 1980. New genera and new combinations in 34: 265–280.
Nearctic Chloropidae (Diptera). Proceedings of the Entomological Wheeler, T.A. & J.J. Mlynarek. 2008. Systematics of Agrophaspidium,
Society of Washington 82: 412–429. a new genus of Neotropical Chloropidae (Diptera). Zootaxa
Sabrosky, C.W. 1987. Chloropidae, pp. 1049–1067. MND, 1926: 41–52.
Volume 2
1154
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
CHYROMYIDAE (chyromyid flies) 93
Terry A. Wheeler

Fig. 93.1. Male of Chyromya flava (Linnaeus), (Holarctic, MND, fig. 91.1).

Diagnosis sclerotized, with eggs clearly visible in gravid females. Male

postabdomen (Fig.  6) various, especially in Aphaniosoma.
Minute to small (body length 0.5–3.0 mm, rarely larger) Female postabdomen (Fig. 7) simple, not telescopic, cercus
yellow or yellow and gray flies, usually with all bristles short.
yellow (Fig. 1). Fronto-orbital bristles varied in number and The combination of small size and pale yellow color
orientation (Figs. 2–4); ocellar and inner and outer vertical (including most bristles) distinguishes chyromyids from
bristles usually strong, postocellar bristles usually present, most other acalyptrate flies. Chloropidae are distinguished
convergent; vibrissa weak. Antenna with first flagellomere by their reduced chaetotaxy, a vertical carina on the propleu-
round or reniform, arista bare or short pubescent. Proboscis ron, and the absence of cell cup in the wing. Specimens of
Zagonia Coquillett (Heleomyzidae — not found in Central
short, fleshy. Thoracic bristles various: one to two postpronotals;
America) have the ocellar bristles outside the ocellar triangle,
two notopleurals; zero to two presutural and one to five
lateral to the anterior ocellus; Zagonia and other small
postsutural dorsocentrals; 2–10 rows of acrostichal setulae;
yellow Heleomyzidae are distinguished by the presence of
four to six marginal scutellar bristles (sometimes with additional preapical dorsal tibial bristles and a spinose costa. Small
marginal or dorsal setulae); anepisternum with one posterior yellow Drosophilidae are distinguished by the arrangement
bristle and some setulae; katepisternum usually with one of the fronto-orbital bristles and usually plumose arista.
strong bristle; anepimeron bare. Wing broad with well-
developed anal angle, C not spinose, extending to M1, sub-
costal break present, humeral break absent, Sc complete but Biology
weak apically, joining C close to insertion of R1, crossveins
bm and dm-cu present, cell cup present, A1 present, ending Larvae of Chyromyidae are apparently saprophagous in
before wing margin (Fig. 5). Legs with weak bristles, dorsal a range of substrates. Adults have been reared from nests
preapical tibial bristles absent. Abdomen usually pale, weakly and dung of birds and mammals (including bats) and from

1155
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

rotting wood and other vegetation (Ferrar, 1987). Adults are tionships within this group (especially the first two families)
frequently collected on flowers and other vegetation, especially are poorly defined (D.K.  McAlpine, 1985; J.F.  McAlpine,
in saline areas, including ocean beaches. There are no 1989). There has been insufficient study of male genitalic and
economically important species. Sabrosky (1958) reported other characters among the Chyromyidae and Heleomyzidae
that Chyromya flava  (L.) can congregate in large numbers to make inferences on higher relationships (T.A.  Wheeler,
on windows, but this is a rare occurrence and would not 1998). The Heleomyzidae may be paraphyletic relative to
normally constitute a nuisance. the Chyromyidae, although a comprehensive phylogenetic
analysis would be required to resolve this question.
Classification Four genera of Chyromyidae are currently recognized,
of which Aphaniosoma is by far the largest, with over
The Chyromyidae are usually considered closely related to 60 species worldwide (Ebejer, 1998). This genus is heteroge-
Heleomyzidae sensu lato and Sphaeroceridae, although rela- neous in both external and genitalic characters and may not

i vt s poc s
o vt s

orb s

2 Gymnochiromyia 3 Aphaniosoma 4 Aphaniosoma

Sc R1 syntgst 6+7

R2+3
st 8
R4+5

M1
epand

A1+CuA2 CuA1 cerc

st 5
ph sur
5 Chyromya
6 Chyromya

tg 7
tg 8

st 7 st 8 cerc

7 Chyromya

Figs. 93.2–7. Heads, wing, and abdomens: anterodorsal view of head of (2) Gymnochiromyia concolor (Malloch), (Nearctic, MND, fig. 91.2);
(3) Aphaniosoma quadrivittatum Malloch, (Nearctic, MND, fig. 91.3); and (4) A. aldrichi Wheeler, (Nearctic, MND, fig. 91.4); dorsal view
of wing of (5) Chyromya flava (Linnaeus), (Holarctic, MND, fig. 91.5): abdomen of male (6) and female (7) of C. flava, (MND, figs. 91.6,
91.8).
Abbreviations: cerc, cercus; epand, epandrium; i vt s, inner vertical seta; orb s, orbital seta; o vt s, outer vertical seta; ph, phallus; poc s,
postocellar seta; st, sternite; syntgst, syntergosternite; sur, surstylus; tg, tergite.
CHYROMYIDAE (chyromyid flies) 93 1157

constitute a monophyletic group (Wheeler & Sinclair, 1994; T.A. Wheeler, 1998) will serve to identify the Central and
T.A.  Wheeler, 1998). Future phylogenetic analyses may South American genera. Identification of the genera as currently
result in Aphaniosoma being divided into multiple genera defined relies heavily on external characters, especially the
based primarily on male genitalic characters. chaetotaxy of the head and thorax, so identification to the generic
level does not usually present a problem with specimens in
good condition. Specimens of Chyromyidae are often min-
Identification ute, weakly sclerotized, and pale so that air dried specimens
shrivel badly and are extremely difficult to identify, even to
Three of the four described genera of Chyromyidae the generic level. Adult specimens should be preserved in
are found in most major zoogeographic realms, so keys ethanol and critical-point dried or chemically dried soon
to the fauna of other realms (e.g., J.F.  McAlpine, 1987; afterwards.

Key to the genera of Chyromyidae of Central America

1. Scutellum with 10–12 dorsal setulae and four marginal bristles; Nearctic Mexico. . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Chyromya Robineau-Desvoidy]
– Scutellum bare dorsally, with four to six marginal bristles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2. Frons with three pairs of fronto-orbital bristles, anterior pair inclinate (Fig. 5); face flat or concave;
proepisternal bristle absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gymnochiromyia Hendel
– Frons with zero to six pairs of fronto-orbital bristles, anterior pair not inclinate (Figs. 3–4); face usually
with median vertical carina; proepisternal bristle present but small. . . . . . . Aphaniosoma Becker

Synopsis of the fauna key to the species of the Galápagos Islands; these are the
only described Neotropical species of Chyromyidae. One of
There are no published records of chyromyid species from these species, A. rabida Wheeler, also occurs on the Pacific
Central America, and there are few specimens in museum coast of Costa Rica (Guanacaste Province); other Galápagos
collections. More Neotropical species undoubtedly await de- species may also be present in Central America. There are
scription, especially with additional collecting effort in dry unpublished records, based on museum specimens, of at
habitats such as marine beaches or other saline areas. The least six undescribed species of Aphaniosoma from Mexico
total number of species of Chyromyidae in Central America (Chiapas, Tampico), Belize, Costa Rica, Panama, Venezuela,
is unknown, but there are probably no more than 15–20 and the West Indies (Anguilla, Antigua, Bahamas, Curaçao,
species in the region. Dominica, St. Thomas, Turks and Caicos).

Aphaniosoma Becker. Species of this genus are usually Gymnochiromyia Hendel. The only described New
associated with dry and/or saline areas, especially near the World species are three Nearctic species (see Malloch,
margin of water bodies. Specimens of two species from 1914). Although one or more of these species may occur
the Galápagos Islands, Ecuador were collected on flowers in southern Mexico or Central America, most Neotropical
of beach morning-glories (Ipomoea pes-caprae (L.) R. Br., species are probably undescribed. There are unpublished
Convolvulaceae) in the littoral zone (Wheeler & Sinclair, records, based on museum specimens, of one or more un-
1994). M.R.  Wheeler (1961) provided a key to the three identified species of Gymnochiromyia from Mexico (Chia-
described Nearctic species, some of which may extend into pas, Nuevo Leon, San Luis Potosí), the West Indies (Jamaica,
southern Mexico. Wheeler & Sinclair (1994) provided a Dominica), and Peru.
Literature cited
Ebejer, M.J. 1998. A review of the Palaearctic species of Aphaniosoma McAlpine, J.F. 1989 Phylogeny and classification of the
Becker (Diptera, Chyromyidae), with descriptions of new Muscomorpha, pp. 1397–1518. MND, Volume 3.
species and a key for the identification of adults. Deutsche Sabrosky, C.W. 1958. An overlooked name in “Musca”. Proceedings
Entomologische Zeitschrift 45: 191–230. of the Entomological Society of Washington 60: 134–135.
Ferrar, P. 1987. A guide to the breeding habits and immature stages Wheeler, M.R. 1961. New species of southwestern acalyptrate
of Diptera Cyclorrhapha. Entomonograph 8(1). E.J.  Brill  / Diptera. The Southwestern Naturalist 6: 86–91.
Scandinavian Science Press, Leiden, Copenhagen, 478 pp. Wheeler, T.A. 1998. Family Chyromyidae, pp. 457–461 In
Malloch, J.R. 1914. Notes on the dipterous genus Chyromya R-D. Pro- Papp, L. & B. Darvas (editors). Contributions to a Manual
ceedings of the Entomological Society of Washington 16: 179–181. of Palaearctic Diptera. Volume 3, Higher Brachycera. Science
McAlpine, D.K. 1985. The Australian genera of Heleomyzidae Herald Publishers, Budapest, 880 pp.
(Diptera: Schizophora) and a reclassification of the family Wheeler, T.A. & B.J. Sinclair. 1994. Chyromyidae (Diptera)
into tribes. Records of the Australian Museum 36: 203–251. from the Galápagos Islands, Ecuador: three new species of
McAlpine, J.F. 1987. Chyromyidae, pp. 985–988. MND, Aphaniosoma Becker. Proceeding of the Entomological Society
Volume 2. of Washington 96: 440–453.
1158
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
HELEOMYZIDAE (heleomyzid flies) 94
David K. McAlpine and Norman E. Woodley

1 Trixoscelis

2 Neorhinotora

Figs.  94.1–2. Males of (1) Trixoscelis fumipennis Melander, (Nearctic, MND, fig.  90.1, as Trixoscelidae); and (2)  Neorhinotora diversa
(Giglio-Tos), (Nearctic, MND, fig. 92.1, as Rhinotoridae).

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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Diagnosis Biology

Flies of small to medium size (body length 2.0–11.0 mm), Many heleomyzid flies live in shaded places in cool
of slender (Fig. 1) to stout build (Fig. 2), their predominant forests. Some, particularly in tribes Cnemospathidini and
color often dull yellowish-brown or gray. Head with face Trixoscelidini, live in dry or sandy habitats, sometimes near
often uniformly sclerotized (with weakly sclerotized median the sea. Few live in tropical lowlands (mainly Rhinotorini).
panel in Heleomyzini), with sclerotized part not extending Adults of Rhinotorini are often seen on tree trunks where
onto ventral surface of head capsule; postocellar bristles they may be associated with wounds in the bark. Species in
convergent or crossed (absent in some Rhinotorini); with one several tribes have been found in bird nests, rodent burrows,
to three orbital bristles, variously directed but generally not or in association with bats. Garnett  & Foote (1967) give
incurved nor strongly proclinate; interfrontal bristles not in many details of the biology of Pseudoleria placata (Hutton)
(as P. crassata Garrett). Larvae of Suillia live in fructifica-
longitudinal series; vibrissae present. Pedicel of antenna not
tions of fungi, in decaying bulbs and other plant material,
dorsally slit nor forming hoodlike or cuplike structure over
and sometimes in mammalian feces. Those of Prosopantrum
base of first flagellomere (Figs. 3, 4). Mouthparts well devel-
Enderlein are probably associated with dung and carrion and
oped, not long-attenuated; labella sometimes short or vestigial. have been reported to breed in a breeding house for laboratory
Thorax with prosternum usually moderately narrow and mice. Larvae of Australian rhinotorines have been found in
isolated (generally broader and subtriangular in Heleomyzini; coleopterous and lepidopterous tunnels in trees.
connected to propleuron by precoxal bridge on each side in Among genera mentioned below, the egg is described
Cinderella); the following bristles generally present: three for Pseudoleria Garrett (Garnett & Foote, 1967), Cinderella
or more dorsocentrals (except in many Rhinotorini), two (Steyskal, 1969), Suillia (Janvier, 1963), Neorhinotora, and
notopleurals, three supra-alars, two subequal scutellars, one Rhinotora Schiner (Lopes, 1935). Eggs are elongate-ovoid
or more katepisternals. Legs usually moderately strong, or sausage-shaped, the surface smooth, reticulate or longitu-
not attenuated; forefemur usually with seriate posteroventral dinally ridged. Eggs of Neorhinotora spp. are more stoutly
bristles (Fig. 1); midfemur often with uniseriate anterior bris- ovoid, with a large concavity at the anterior end.
tles; tibiae generally with preapical dorsal bristles (Fig. 1), The larvae of heleomyzids are diverse in appearance
often solitary, particularly on foretibia (reduced or absent in and morphology. Those of most genera treated here are
some Rhinotorini); hind basitarsus generally not particularly undescribed. Larvae of Suillia species have been described
short and compact, but midbasitarsus often more elongate by Janvier (1963), Ferrar (1987), and Papp (1999a). They
than other basitarsi. Costa with subcostal break (Fig. 7), some- are of medium to stout build, anteriorly tapered, posteriorly
times also with humeral break, often with spaced anterior to truncate to broadly rounded. Papp (1999a) characterizes the
anteroventral spines extending beyond apex of R1 (Fig.  1; third instar larva of Suillia, as opposed to other heleo-
absent in many Neotropical genera); R1 without dorsal setu- myzids (Heleomyzini in sense of McAlpine, 1985), as
lae; cells bm and cup closed; vein A1+CuA2 well sclerotized, follows: “No spiracular hairs on posterior spiracle ... Basal
sometimes discontinued well short of margin. Sclerites of part of mouth hooks robust, but apical part comparatively
smaller [i.e. more slender] .... Anterior spiracles with 9–14,
male protandrium usually strongly asymmetrical; sternite 8
rather large, radially arranged lobes. Ventral cornu with a
usually well developed and distinct. Female postabdomen
subdorsal hole.”
without aculeus or egg guide; cerci usually separate.
Many heleomyzids are recognizable by the combination
Classification
of convergent ocellar bristles, orbital bristles in one to three
pairs that are not proclinate, vibrissa present, costa with sub- This treatment of Heleomyzidae includes taxa that are some-
costal break and spaced spines beyond R1, closed cells bm times included in the families Cnemospathidae, Rhinotoridae,
and cup, and presence of differentiated preapical dorsal tibial and Trixoscelididae. Classification of the heleomyzid-like
bristles. More specialized taxa of Rhinotorini lack postverti- flies has long been a major source of difficulty and disagree-
cal bristles, spaced costal spines, and preapical tibial bris- ment. Hence, in A Catalogue of the Diptera of the Americas
tles, but can be recognized by the deeply excavated vertex, South of the United States, some rhinotorine genera are listed
dorsally prominent eyes, and greatly thickened forefemora. under Heleomyzidae (Gill, 1968), and others are listed
Heleomyzidae of typical form are most likely to be confused under Rhinotoridae (Papavero, 1967). It is now certain that
with either Sciomyzidae or Lauxaniidae, but heleomyzids the division of the heleomyzid-like genera into eight fami-
have a subcostal break in the costa, which the sciomyids lies, distributed among three different schizophoran alliances
(prefamilies) and one of uncertain relationships, as proposed
and lauxaniids lack, as they have a complete costa. The
by Griffiths (1972) based mainly on male postabdominal
more atypical Rhinotorini might be mistaken for small Ro- characters, is untenable. McAlpine (1985) classified the
palomeridae; however, ropalomerids have veins R4+5 and M world heleomyzid-like genera into 22 tribes, which appear
convergent toward the wing apex, while these veins are par- to be natural (perhaps monophyletic) groups, but left these
allel or slightly divergent in rhinotorines. in a single family with no defined subfamilies, pending further
HELEOMYZIDAE (heleomyzid flies) 94 1161

information on their interrelationships. Papp (1999a) re- populations, which are thought to be parthenogenetic
turned to a multiple family classification for his treatment of (Cogan, 1971). It is yet to be demonstrated that the few
the Palearctic “Heleomyzoidea” (in his own special sense), recorded Afrotropical males are reproductively functional.
but this classification, like some earlier ones made from a The species of tribe Rhinotorini are distributed from
purely north temperate viewpoint, lacks phylogenetic bal- southwestern USA to southern Chile, and also in Australasia
ance and distorts some relationships. The taxa considered in (Sinclair  & McAlpine, 1995). The tropical taxa, e.g.,
this chapter that are likely present in Central America belong Neorhinotora and Rhinotora, are stout flies with unusu-
to seven tribes of McAlpine (1985): Heleomyzini, Suilliini, ally tall heads, no postocellar bristles, and no spaced costal
Cinderellini (Cinderella), Cnemospathidini (Prosopantrum), spines, in contrast with taxa from southern South America,
Gephyromyzini Paraneossos Wheeler, Rhinotorini, and which are more conventionally heleomyzid-like (Malloch,
Trixoscelidini. 1933). Some species have several crossveins between R2+3
The tribe Heleomyzini includes many Nearctic genera (Gill, and the costa. The tropical American taxa were reviewed by
1962), and some have been introduced in south temperate Guimarães & Papavero (1966).
countries, e.g., Oecothea Haliday (synonyms Aecothea
Haliday, Neoecothea Peterson & Gill), Pseudoleria, and Teph- Identification
rochlamys Loew. These and Scoliocentra (including Amoe-
baleria Garrett) extend into Mexico. McAlpine (1985) gave keys to the world tribes and the
The only included genus of the tribe Cnemospathidini, recorded Neotropical genera of Heleomyzidae sensu lato.
Prosopantrum, is usually considered to be native to temperate Little has been published at the species level for Central America.
South America. However, Papp (1999b) described three Several genera have been included in the key that have not
species from tropical Africa. Prosopantrum flavifrons yet been found in Central America, some of which have been
(Tonnoir & Malloch) has been introduced in several mainly collected in Nearctic Mexico. These have been introduced
temperate countries, although we are aware of no Central widely in other parts of the world and could well be collected
American records. Males are usually absent in these introduced in Central America at some point in the future.

poc s i vt s
i vt s
o vt s

orb s o vt s

oc s
orb plt

3 Suillia
4 Scoliocentra 5 Neorhinotora

Sc R1 R2+3

R4+5

M1

A1+CuA2 CuA1

6 Cinderella 7 Spilochroa
Figs. 94.3–7. Heads and wing: anterior view of head of (3) Suillia longipennis (Loew), (Nearctic, MND, fig. 89.2); (4) Scoliocentra helvola
Loew, (Nearctic, MND, fig. 89.3, as Amoebaleria helvola); (5) Neorhinotora diversa (Giglio-Tos), (Nearctic, MND, fig. 92.2, as Rhinotoridae);
anterodorsal view of head of (6) Cinderella lampra Steyskal, (Nearctic, MND, fig. 4.112); dorsal view of wing of (7) Spilochroa ornata
(Johnson), (Nearctic, MND, fig. 90.3, as Trixoscelididae).
Abbreviations: i vt s, inner vertical seta; oc s, ocellar seta; orb plt, orbital plate; orb s, orbital seta; o vt s, outer vertical seta; poc s, postocellar
seta.
1162
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Key to the genera of Heleomyzidae of Central America

1. Orbital plate short, sloping mesad away from eye margin, and bearing only one strong orbital
bristle (Fig. 3); dorsocentral bristles generally in five pairs; postpronotal bristle absent . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Suillia Robineau-Desvoidy
– Orbital plates, if differentiated, parallel with eye margin (Fig. 4); other characters various. . . . . 2

2. Anepisternal bristle present, large; ocellar bristles inserted lateral to anterior ocellus (Fig. 1);
usually two pairs of reclinate orbital bristles, one postpronotal bristle, and five pairs of dorsocentral
bristles present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
– Anepisternal bristle absent; ocellar bristles inserted between anterior and posterior ocelli (Figs. 3, 4);
other characters various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

3. Wing dark with hyaline spots (Fig. 7). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Spilochroa Williston

– Wing hyaline, or with infuscation along veins and/or bordering crossveins. . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trixoscelis Rondani

4. Face with relatively lightly sclerotized median panel; vein A1+CuA2 extending approximately to
wing margin (tribe Heleomyzini). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
– Face without such differentiated median panel; vein A1+CuA2 discontinued well before margin. . . . 8

5. Three long dorsocentral bristles present, any fourth (anterior) dorsocentral short; midtibia with
one ventral to subventral apical spurlike bristle; widespread, but not known from Central America.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Tephrochlamys Loew]
– Four long dorsocentral bristles present; midtibia with two or more apical, more or less ventral
spurlike bristles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

6. Prosternum with at least one pair of long bristles . . . . . . . . . . . . . . . . . . . . . . . Scoliocentra Loew

– Prosternum without bristles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

7. Scutellum setulose; anepimeron bare; Holarctic Region, widely introduced elsewhere. . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Oecothea Haliday]
– Scutellum bare, except for major bristles; anepimeron setulose; Nearctic Region, widely distributed
elsewhere. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Pseudoleria Garrett]

8. Prosternum joined to propleuron on each side by precoxal bridge; face with rounded median
tubercle (Fig. 6); hypofacial and anteclypeus deep. . . . . . . . . . . . . . . . . . . . . . Cinderella Steyskal
– Prosternum not joined to propleuron; face without distinct median tubercle; hypofacial and
anteclypeus various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

9. Orbital bristles in three pairs; genal region divided into two zones of different texture: upper zone
of dense, pale tomentum; lower dark zone of sparser tomentum; Arizona (USA), temperate South
America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Paraneossos Wheeler]
– Orbital bristles not more than two pairs; genal region not thus divided into two zones. . . . . . . 10

10. Head not excavated at vertex; arista subbasal on first flagellomere; midfemur with one or two
anterior bristles but no anteroventral bristles; all tibiae with preapical dorsal bristles; costa with
two (one dorsal, one ventral) costagial bristles; Africa, temperate South America, introduced
elsewhere. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Prosopantrum Enderlein]
– Head excavated at vertex (Fig. 5); arista inserted near midlength of first flagellomere; midfemur
without anterior bristles, sometimes with anteroventral or posteroventral bristles; tibiae without
distinct preapical dorsal bristles; costa with one (ventral) costagial bristle. . . . . . . . . . . . . . . . . 11
HELEOMYZIDAE (heleomyzid flies) 94 1163

11. Scutellum with marked median dorsal channel, without apical median tubercle (Fig. 2); face with
convexity on each side; eye rounded-oval in profile. . . . . . . . . . . . . . . . . . . . Neorhinotora Lopes
– Scutellum without dorsal channel, with apical median tubercle between apical setae; face without
pair of convexities; eye narrowly oblique in profile; South America . . . . . . . [Rhinotora Schiner]

Synopsis of the fauna patterned wings, a feature that is unusual for the genus. Species
of Scoliocentra have been collected from caves, where they
As the Heleomyzidae are more common in temperate apparently feed on bat guano, and from mammal burrows
regions, it is not surprising that fewer than 10 species, both (Ferrar, 1987).
described and undescribed, are known from Central America
and southern Mexico. It is unlikely that many additional Spilochroa Williston. This genus differs from Trixoscelis
species will be found here. by having patterned wings, a more strongly developed
anterior bristle on the hind femur, and a smaller genal region.
Cinderella Steyskal. This genus is widespread in North Some authors (e.g., Malloch, 1933) considered Spilochroa as
America and temperate (including high elevation) South a subgenus of Trixoscelis. The species of Spilochroa range
America. Morphology and species groups were discussed by from the extreme southern USA south through the West
McAlpine (1985). There are two specimens of an unidentified Indies to Peru, Chile, and Argentina (Gill, 1968). There are
Cinderella in INBio, but otherwise nothing is known of the undetermined specimens in USNM from Mexico (Oaxaca,
genus in Central America. Sonora, Veracruz), and specimens have been noted from
Guatemala (A. Norrbom, personal communication).
Neorhinotora Lopes. There are five described species in
this genus, mostly from South America (Papavero, 1967). Suillia Robineau-Desvoidy. This genus has a wide
Almeida  & Ale-Rocha (2008) recently reviewed the genus Holarctic, Afrotropical, and Oriental distribution and occurs
and presented a key to the described species. One species, as far south as Ecuador in the Neotropical Region (Woźnica,
N.  diversa (Giglio-Tos) is known from the southwestern 2006). Steyskal (1972, 1980) reviewed the species from south
United States and has been recorded from Veracruz, Mexico of the United States, including one from Colombia. Subse-
(Almeida  & Ale-Rocha, 2008). Neorhinotora aristalis quently, Woźnica (2006) described additional Neotropical
(Fischer) occurs in Panama (USNM specimens) as well as species and included a key to the 11 known species from the
in Brazil and Bolivia (USNM). There are specimens in the region. He noted seven species from Mexico, at least two of
USNM of an undescribed species from El Salvador, and some which occur in Chiapas, S. punctulata (Wulp) (one specimen
INBio specimens are probably conspecific with this species in USNM) and S. iniens (Giglio-Tos), the latter also recorded
as well. The genus Rhinotora has been recorded only from from Honduras. An apparently undescribed species found at
South America, but could possibly occur in Central America, high elevations in Costa Rica (USNM, INBio) is unusual in
and so it is included in the generic key. These genera, formerly that it has a large anepisternal bristle in addition to smaller
included in a separate family Rhinotoridae, are usually collected setulae. An unidentified species is also known from Guatemala
in traps baited with fermented fruits or by rubbing fruits on (CNC specimens). Larvae of Suillia are usually associated
the bark of trees (Guimarães & Papavero, 1966). with fungi (Ferrar, 1987), but one species is also a pest of
garlic and onion in Europe (Papp, 1999a).
Scoliocentra Loew. This genus includes species formerly
placed in the genus Amoebaleria (Gill, 1962; Gill & Peter- Trixoscelis Rondani. Trixoscelis has a wide distribution
son, 1987) that differ from Scoliocentra sensu stricto by but is apparently absent from the Australian and Oriental
having the anepimeron bare. Scoliocentra is widespread in Regions. Gill (1968) recorded species from Mexico and
the Nearctic and Palearctic Regions. There are some un- the Galápagos Islands. There are a few undetermined speci-
identified specimens of Scoliocentra in CNC from Mexico mens of Trixoscelis in the INBio collection collected by
and three specimens of a distinctive unnamed species from Malaise trap in a dry forest in Palo Verde National Park,
Mexico (Chiapas) and El Salvador (USNM) with boldly Costa Rica.

Literature cited

Almeida, J.C.  & R. Ale-Rocha. 2008. Taxonomic review of Garnett, W.B. & B.A. Foote. 1967. Biology and immature stages
Neorhinotora Lopes 1934 (Diptera, Heleomyzidae). Zootaxa of Pseudoleria crassata (Diptera: Heleomyzidae). Annals of the
1936: 40–58. Entomological Society of America 60: 126–134.
Cogan, B.H. 1971. The Heleomyzidae of the Ethiopian Region Gill, G.D. 1962. The heleomyzid flies of America north of Mexico
(Diptera). Annals of the Natal Museum 20: 627–695. (Diptera: Heleomyzidae). Proceedings of the United States
Ferrar, P. 1987. A guide to the breeding habits and immature stages National Museum 113: 495–603.
of Diptera Cyclorrhapha. Entomonograph 8(1). E.J.  Brill  / Gill, G.D. 1968. 85. Family Heleomyzidae (Helomyzidae) including
Scandinavian Science Press, Leiden, Copenhagen, 478 pp. the Trixoscelididae (Trichoscelidae). In Papavero, N. (editor).
1164
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

A Catalogue of the Diptera of the Americas South of the United States. Departamento de Zoologia, Secretaria da Agricultura,
States. Departamento de Zoologia, Secretaria da Agricultura, São Paulo, 4 pp.
São Paulo, 13 pp. Papp, L. 1999a. 3.41. Families of Heleomyzoidea, pp. 425–455.
Gill, G.D.  & B.V. Peterson. 1987. Heleomyzidae, pp. 973–980, In Papp, L. & B. Darvas (editors). Contributions to a Manual
MND, Volume 2. of Palaearctic Diptera. Volume 3, Higher Brachycera. Science
Griffiths, G.C.D. 1972. The phylogenetic classification of Diptera Herald Publishers, Budapest, 880 pp.
Cyclorrhapha, with special reference to the structure of the male
Papp, L. 1999b. The Afrotropical species of Prosopantrum
postabdomen. Series entomologica 8. Dr. W. Junk, The Hague,
340 pp. Enderlein (Diptera: Cneomospathidae). Annals of the Natal
Guimarães, J.H. & N. Papavero. 1966. A contribution to the knowledge Museum 40: 83–91.
of the Neotropical Rhinotoridae (Diptera, Acalyptratae). Papéis Sinclair, B.J. & D.K. McAlpine. 1995. Zinza, a new genus of rhino-
Avulsos do Departamento de Zoologia 18: 261–269. torine flies from northern Queensland, Australia (Diptera: He-
Janvier, H. 1963. La mouche de la truffe (Helomyza tuberiperda leomyzidae). Records of the Australian Museum 47: 225–230.
Rondani). Bulletin de la Société Entomologique de France Steyskal, G.C. 1969. The postabdomen of Cinderella, with a new
68: 140–147. species from Argentina (Diptera: Heleomyzidae). Journal of the
Lopes, H. de S. 1935. A sub-familia Rhinotorinae Williston (Dipt. Kansas Entomological Society 42: 80–83.
Rhopalomeridae). Archivos do Instituto de Biologia Vegetal Steyskal, G.C. 1972. Notes on the genus Suillia in Mexico, with the
2: 19–26.
description of a new species (Diptera: Heleomyzidae). Proceedings
Malloch, J.R. 1933. Acalyptrata. (Helomyzidae, Trypetidae,
Sciomyzidae, Sapromyzidae, etc.), pp.  177–392. Diptera of of the Entomological Society of Washington 74: 303–305.
Patagonia and South Chile, Part VI  – Fascicle  1. British Steyskal, G.C. 1980. The species of the genus Suillia found in the
Museum of Natural History, London. Americas south of the United States (Diptera: Heleomyzidae).
McAlpine, D.K. 1985. The Australian genera of Heleomyzidae Proceedings of the Entomological Society of Washington
(Diptera: Schizophora) and a reclassification of the family into 82: 401–404.
tribes. Records of the Australian Museum 36: 203–251. Woźnica, A.J. 2006. Three new species of the genus Suillia
Papavero, N. 1967. 87. Family Rhinotoridae. In Papavero, N. (editor). Robineau-Desvoidy, 1830 from the Neotropical Region
A Catalogue of the Diptera of the Americas South of the United (Diptera: Heleomyzidae). Annales zoologici 56: 657–665.
SPHAEROCERIDAE (small dung flies) 95
Stephen A. Marshall and Matthias Buck

Fig. 95.1. Male of Limomyza venia Marshall, (Nearctic, Roháček et al., 2001, fig. 16).

Diagnosis katepisternum with zero to three (usually one to two) dorsal

bristles. Wing with two costal breaks, reduced subcosta, and
Generally small acalypterate flies (body length 0.7–6.0 mm, in most species (most Limosininae and Homalomitrinae) with
usually 1–3  mm), but larger species occur outside Central cells bm and dm fused and an open cell cup. Wing patterned
America (Fig. 1). Body color black to brown, rarely brightly in some species, frequently reduced or lost in Limosininae
patterned. Head with zero to (usually) one or two pairs of and Copromyzinae, especially in those living at higher
lateroclinate orbital bristles, interfrontal bristles usually altitudes. Midtibia usually (except in Sphaerocerinae and
distinct, vibrissae present but sometimes small, postocel- Homalomitrinae) with several (at least two) bristles. First,
lar bristles, if present, convergent. Anepisternum bare and in some species also second, hind tarsomere shortened
(Limosininae, Copromyzinae) or setulose (Sphaerocerinae, and/or thickened.
Homalomitrinae), anepimeron usually bare (setulose in Sphaeroceridae can easily be distinguished from other
Homalomitrinae, with few setulae in some Sphaerocerinae), Acalyptratae families by the shortened and/or thickened first

1165
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

hind tarsomere, a reduced subcosta, and the presence of two Classification

costal breaks. The highly aberrant subfamily Homalomitrinae
can be recognized by the absence of ocelli (sometimes Sphaeroceridae are divided into five subfamilies, four
replaced by tubercles) and fused or absent wing cells bm and of which occur in Central America. The vast majority of
dm. the species are currently placed in the huge, diverse sub-
family Limosininae. The distinctive subfamily Sphaerocerinae,
Biology although much smaller than the Limosininae, contains
some abundant and relatively large Neotropical species.
Sphaerocerid larvae are microbial grazers in a variety of Another subfamily, the Copromyzinae, is primarily north
humid, bacteria-rich environments. They are abundant in and south temperate in the New World and has only a few
dung, carrion, and various types of decomposing vegetation, species in Central America. The species of the subfamily
including fungi. Although many species appear to be polysap- Homalomitrinae are extremely rare and are associated
rophagous and can be found in large numbers in a variety of with army ants (Roháček & Marshall, 1998). The higher
decomposing materials ranging from compost heaps to tree classification of the Sphaeroceridae is currently under revision,
falls, other species are highly specialized. One species known
and the monophyly of the Copromyzinae and Limosininae is
only from Costa Rica develops in the excretory glands of land
crabs (Gómez, 1977), and several Neotropical species are in question (Marshall, 1998). Two of the five subfamilies
associated with army ants, bromeliads or other plants with have only recently been described from South and Central
pockets of moist decay, caves, mud flats, seaweed, and fungi. America, and the Neotropical fauna includes a large number
Elsewhere in the world some Sphaeroceridae are associated of undescribed genera in the subfamily Limosininae. Rec-
with mammal burrows and dung-rolling scarab beetles, and ognized but undescribed genera that are known to occur in
it is likely that some Neotropical sphaerocerid lineages have Central America are indicated in the key as such.
similar associations.
Adults can be collected in a number of different ways:
Malaise traps and aerial sweep netting often yield large
numbers of specimens but usually collect only a small fraction Identification
of the fauna. Probably the most productive general collecting
methods are pan traps and flight intercept traps. Since most The key below is the first key to the Neotropical genera
Sphaeroceridae are specific to certain microhabitats, the species of Sphaeroceridae. For the identification of genera, genitalic
composition of trap samples depends greatly on the immediate characters are usually not required, but they are important
location where traps have been placed. In tropical regions, for species identification in most groups. The world Spha-
accumulations of fresh decaying vegetation (e.g., tree falls) eroceridae were recently catalogued (Roháček et al., 2001),
are especially productive. The fairly diverse dung fauna can including a complete bibliography to the family. The best
only be adequately sampled with dung traps. Trapping with method for preservation is to preserve specimens in alcohol
other baits (e.g., carrion, fungi) yields fewer specimens and prior to critical-point drying them, and mounting them with
fewer species. Many of the elusive brachypterous or apterous
white glue on points. Air-dried specimens usually shrivel
species that inhabit leaf litter at higher elevations require
sampling with Berlese/Tullgren extractors or sieving. and are difficult to dissect.

Key to the genera of Sphaeroceridae of the Neotropical Region and Nearctic Mexico
This key does not include the enigmatic genus Pycnopota Bezzi, which was tentatively treated as an aberrant member of
Copromyzinae by Roháček et al. (2001). Previously known only from Bolivia, Pycnopota was recently discovered in Costa
Rica. It differs from all other Sphaeroceridae in having a conspicuous tubular proboscis that reaches at least to the midcoxae.

1. Fully winged to slightly brachypterous; wings at least 0.75 times as long as abdomen and not
extremely narrowed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Apterous or strongly brachypterous; wings shorter than 0.75 times length of abdomen, or wings
greatly narrowed and straplike. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83

Note: Moderately brachypterous species can be keyed both ways.

SPHAEROCERIDAE (small dung flies) 95 1167

2 Mesosphaerocera

3 Archiborborus 4 Undescribed Genus B

orb s

5 Undescribed Genus lun

(key: couplet 55)
7 Sclerocoelus

6 Pterogramma

8 Homalomitra 9 Podiomitra 10 Sphaeromitra

Figs. 95.2–10. Heads and thoraces: anterolateral view of head of (2) Mesosphaerocera annulicornis (Malloch), (Nearctic, MND, fig. 93.11);
(3) Archiborborus sp.; (4) Undescribed Genus B; (5) Undescribed Genus (key: couplet 55); (6) Pterogramma aquatile Smith & Marshall,
(Florida, Caribbean, Smith  & Marshall, 2004, fig.  4); and (7)  Sclerocoelus caribensis Marshall; dorsal view of head and thorax of
(8) Homalomitra antiqua Roháček & Marshall, (Roháček & Marshall, 1998, fig. 2); (9) Podiomitra ostracotarsata Marshall & Roháček,
(Marshall & Roháček, 2003, fig. 5); (10) Sphaeromitra inepta Roháček & Marshall, (South America, Roháček & Marshall, 1998, fig. 53).
Figures 3–5 and 7 illustrated by K. Bethune-Leamen.
Abbreviations: lun, lunule; orb s, orbital seta.
1168
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

2. Ocelli absent; frontal bristles greatly reduced (Figs. 8–10); thorax small; scutum shorter to slightly
longer than head (Figs. 8–10); midtibia without bristles; wing venation often reduced in characteristic
manner (Figs. 12–13); rare flies associated with army ants; Homalomitrinae. . . . . . . . . . 3
– Ocelli present; usually at least some bristles of frons well developed (except in Sphaerocerinae);
thorax larger; scutum usually distinctly longer than head; midtibia usually with at least some
bristles (except in Sphaerocerinae); wing venation more or less complete, not as in Figs. 12–13;
common flies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

3. Antennal scapes enlarged, almost contiguous medially (Fig. 8); wing venation normal, vein R2+3
and cell bm+dm long (Fig. 11); scutum with well-developed macrosetae other than prescutellar
dorsocentral and scutellar bristles (Fig. 8). . . . . . . . . . . . . . . . . . . . . . . . . Homalomitra Borgmeier
– Scapes not enlarged, broadly separated medially (Figs. 9–10); wing venation more or less reduced,
vein R2+3 and cell bm+dm, if present, not extending beyond basal fifth of wing (Figs.  12–13);
scutum without macrosetae except sometimes prescutellar dorsocentral and scutellar bristles
(Figs. 9–10) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4. Ocelli replaced by prominent tubercles (Fig. 10); wing cell bm+dm absent, costa and vein R4+5
long, ending near wing apex (Fig. 13); Peru. . . . . . . . . . . . . [Sphaeromitra Roháček & Marshall]
– Frons without tubercles (Fig. 9); wing cell m present but small, costa and vein R4+5 extremely
shortened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Podiomitra Marshall & Roháček

5. Cells bm and dm separated; cell cup closed (Fig.  14); M and sometimes CuA1 reaching wing
margin; usually either bristles of scutum and scutellum reduced or replaced by short spines, teeth,
or warts, or ocellar bristles inserted slightly to distinctly before anterior ocellus (absent in some
species). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
– Cells bm and dm fused; cell cup usually open (Figs.  16–17, 19–25); M rarely, and CuA1 not
reaching wing margin; scutellum with at least four marginal bristles; scutum with bristles; ocellar
bristles inserted behind anterior ocellus; Limosininae (and some Copromyzinae). . . . 17

6. Bristles of scutellum reduced or replaced by short spines, teeth or warts, at least with pair of
subapical or basal marginal tubercles (e.g., Fig. 44); CuA1 reaching wing margin; anepisternum
with patch of short setae; lower face prominent (Fig. 2); arista bare; Sphaerocerinae. . . . 7
– Scutellum with at least four marginal bristles; CuA1 not reaching wing margin, but usually more
or less produced beyond crossvein dm-cu; anepisternum bare; lower face not prominent; arista
usually pubescent but sometimes almost bare (Tucma only). . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

7. Scutellum with six or more warts along posterior margin (Fig. 44); hind tibia with strong, curved
apicoventral bristle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ischiolepta Lioy
– Scutellum at most with pair of marginal teeth laterally; hind tibia at most with small apicoventral
spurlike seta. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

8. Lower margin of antennal fovea somewhat rounded at least in front of vibrissa; lower face flat,
more or less triangular, without groove below antennal fovea; abdominal syntergite 1+2 fused to
tergite 3 in male, separate or fused in female. . . . . . . . . . . . . . . . . . . . . . . . . Neosphaerocera Kim
– Lower margin of antennal fovea carinate along its whole length; lower face convex and/or grooved
just below antennal fovea; abdominal syntergite 1+2 and tergite 3 various . . . . . . . . . . . . . . . . . 9

9. Abdominal syntergite 1+2 separate from tergite 3. . . . . . . . . . . . . . . . . . . . Mesosphaerocera Kim

– Abdominal syntergite 1+2 fused with tergite 3. . . . . . . . . . . . . . . . . . . . . Parasphaerocera Spuler
SPHAEROCERIDAE (small dung flies) 95 1169

11 Homalomitra 12 Podiomitra

13 Sphaeromitra 14 Copromyza

15 Anomioptera 16 Archiceroptera (s. str.)

17 Archiceroptera 18 Archileptocera

Figs. 95.11–18. Wings: dorsal view of (11) Homalomitra antiqua Roháček & Marshall, (Roháček & Marshall, 1998, fig. 6); (12) Podiomitra
ostracotarsata Marshall & Roháček, (Marshall & Roháček, 2003, fig. 4); (13) Sphaeromitra inepta Roháček & Marshall, (South America,
Roháček & Marshall, 1998, fig. 56); (14) Copromyza equina Fallén, (Holarctic, MND, fig. 93.3); (15) Anomioptera quinquealba Marshall,
(South America, Marshall, 1998, fig. 111); (16) Archiceroptera (sensu stricto) sp.; (17) Archiceroptera (genus complex) sp., (Nearctic); and
(18) Archileptocera lutea Marshall, (South America, Marshall, 1998, fig. 112). Figures 16 and 17 illustrated by A. Cormier.
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

19 Bromeloecia 20 Leptocera

21 Poecilosomella 22 Pterogramma

23 Robustagramma 24 Spelobia

25 Spinilimosina

Figs. 95.19–25. Wings (concluded): dorsal view of (19) Bromeloecia sp.; (20) Leptocera fontinalis (Fallén), (Holarctic, MND, fig. 93.4);
(21)  Poecilosomella angulata (Thomson), (MND, fig.  93.9); (22)  Pterogramma palliceps (Johnson), (Nearctic, MND, fig.  93.8);
(23) Robustagramma flavistylum Cui & Marshall; (24) Spelobia (Bifronsina) bifrons (Stenhammar), (redrawn from Roháček, 1985, fig. 695);
and (25) Spinilimosina brevicostata (Duda), (redrawn from Roháček, 1985, fig. 807). Figures 19, 23–25 illustrated by A. Cormier.
SPHAEROCERIDAE (small dung flies) 95 1171

10. One pair of orbital bristles present; hind femur with two rows of stout ventral spinelike setae;
with one strong pair of presutural dorsocentral bristles; with four scutellar bristles, laterobasal pair
small and hairlike; katatergite protuberant; arista virtually bare; TUCMINAE; Argentina. . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Tucma Mourgués-Schurter]
– Two pairs of orbital bristles present; hind femur without ventral spinelike setae; dorsocentral
bristles various but not only in single presutural pair; scutellum at least with two well-developed
pairs of macrosetae, sometimes with additional marginal setulae; katatergite not prominent; arista
pubescent to long-pubescent; Copromyzinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

11. Katepisternum with well-developed dorsal bristle; ocellar bristles inserted slightly to distinctly
before level of anterior ocellus (Fig. 3), sometimes absent; head sometimes bright orange (especially
in Central American species); Archiborborus Duda (sensu lato), in part . . . . . . . . . . . . . . . . . . 12
– Katepisternum with dorsal bristle short or absent; ocellar bristles inserted behind level of anterior
ocellus; ground color of head black. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

12. Head with one to two rows of postocular hairs; female with three spermathecae. . . . . . . . . . . . 13
– Head with numerous rows of postocular hairs; female with two spermathecae; temperate South
America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

13. Head, thorax, and abdomen mostly glossy black, pruinose areas restricted; only two rows of
acrostichals; legs including tarsi black; wing not maculated; South America. . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Archiborborus orbitalis group (unnamed subgenus)]
– Head, thorax, and abdomen more or less extensively pruinose; head often partly reddish to
entirely bright orange; at least four rows of acrostichal setae present; legs sometimes in part
yellow or tarsi in part whitish; wing often maculated along veins . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Archiborborus (Archiborborus) Duda, in part

14. Eye small and flat, height at middle about half genal height; katepisternum with strong bristle and
few hairs dorsally, broadly separated from ventral group of setae; legs not banded; mid- and hind
tibia with strong bristles on dorsal surface; wing not maculated; Argentina, Chile. . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Archiborborus (Huapia) Richards]
– Eye large and convex, much higher than half genal height; katepisternum haired throughout; legs
sometimes prominently banded; mid- and hind tibia with long hair, but no bristles except near
apex; wing maculated along veins; Bolivia and Uruguay south . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Archiborborus hirtipes group (unnamed subgenus)]

15. Scutellum with two pairs of macrosetae only; hind tibia with anteroventral seta at about two-thirds
to three-quarters; male first foretarsomere with apicoventral blunt thornlike projection . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Norrbomia Papp
– Scutellum with at least three pairs of marginal setae (some may be small); hind tibia without
anteroventral seta; male first foretarsomere without apicoventral spurlike seta . . . . . . . . . . . . . 16

16. Hind tibia with apicoventral spurlike seta; margin of scutellum with several small hairs in addition
to the two pairs of scutellar bristles; frons completely pruinose. . . . . . . . . . . . . Copromyza Fallén
– Hind tibia without apicoventral spurlike seta; scutellum with four to five pairs of subequal marginal
bristles (apical pair larger); frontal triangle partially shiny; mountains of North and Central
Mexico. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Lotophila Lioy]

17. Scutellum with discal or marginal setulae or setae in addition to the usual four marginal bristles
(Figs. 45–47) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
– Scutellum with four marginal bristles only, with no additional discal or marginal setulae or setae
(Fig. 48). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

26 Archiceroptera 27 Archileptocera 28 Bitheca 29 Druciatus 30 Lepidosina

31 Minilimosina 32 Phthitia 33 Poecilosomella 34 Pterogramma 35 Robustagramma

36 Sclerocoelus 37 Spelobia 38 Telomerina 39 Minilimosina

40 Opacifrons 41 Pseudocollinella 42 Spelobia 43 Bitheca

Figs. 95.26–43. Legs of Limosininae: dorsal view of left midtibia of (26) Archiceroptera sp.; (27) Archileptocera lutea Marshall, (South
America); (28) Bitheca sp.; (29) Druciatus ovisternus Marshall, (Caribbean, redrawn from Marshall, 1995, fig. 17); (30) Lepidosina inaequalis
Marshall & Buck; (31) Minilimosina (Svarciella) sp.; (32) Phthitia antillensis Marshall, (Caribbean); (33) Poecilosomella angulata (Thomson);
(34) Pterogramma aquatile Smith & Marshall, (Florida, Caribbean, Smith & Marshall, 2004, fig. 10); (35) Robustagramma disjunctum Cui &
Marshall; (36) Sclerocoelus caribensis Marshall; (37) Spelobia (Bifronsina) bifrons (Stenhammar), (redrawn from Roháček, 1985, fig. 692);
(38) Telomerina flavipes (Meigen), (widespread temperate, absent from Central America, redrawn from Roháček, 1985, fig. 963); anterior
view of male midtibia of (39) M. (Amputella) bistylus Marshall; anterior view of female midtibia and first tarsomere of (40) Opacifrons
quarta Marshall & Langstaff; and (41) Pseudocollinella sp., (South America); anterior view of midtibia of (42) Sp. aciculata Marshall;
anterior view of hind tibia and tarsus of (43) Bitheca sp. Figures 26–33, 35–43 illustrated by A. Cormier.
SPHAEROCERIDAE (small dung flies) 95 1173

44 Ischiolepta
a dc s

pprn s

npl s
presut
spal s
spal s
dc s
pal s 45 Coproica 46 Leptocera
ial s

acr s

ial s

47 Rachispoda

48 Sclerocoelus 49 Druciatus

50 Opacifrons 51 Chespiritos 52 Opacifrons

Figs. 95.44–52. Thoraces, scutella, and terminalia: dorsal view of scutellum of (44) Ischiolepta ischnocnemis Buck & Marshall, (Nearctic,
MND, fig. 93.28, as I. micropyga of authors, not Duda); (45) Coproica ferruginata (Stenhammar), (MND, fig. 93.27); and (46) Leptocera
fontinalis (Fallén), (Holarctic, MND, fig. 93.24); dorsal view of thorax of (47) Rachispoda limosa (Fallén), (Holarctic, MND, fig. 93.25, as
Leptocera (Rachispoda) limosa); and (48) Sclerocoelus sp.; posterior view of male terminalia of (49) Druciatus angustus Marshall,
(Marshall & Totton, 1995, fig. 1); and (50) Opacifrons quarta Marshall & Langstaff, (Marshall & Langstaff, 1998, fig. 74); dorsal view
of female terminalia of (51)  Chespiritos sindecimus Marshall, (Marshall, 2000a, fig.  6); and (52)  O. quarta. Figure  48 illustrated by
K. Bethune-Leamen; Figure 52 illustrated by R. Langstaff.
Abbreviations: acr s, acrostichal seta; a dc s, anterior dorsocentral seta; dc s, dorsocentral seta; ial s, intra-alar seta; npl s, notopleural seta;
pal s, postalar seta; pprn s, postpronotal seta; presut spal s, preutural supra-alar seta; spal s, supra-alar seta.
1174
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

18. Scutellum with discal setulae (Fig. 45) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

– Scutellum with six or more marginal setae or setulae but no discal setulae (Fig. 46). . . . . . . . . 23

19. Cell cup closed (Figs. 15, 18); South America. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

– Cell cup open (as in Fig. 20); widespread. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

20. R2+3 strongly sinuate, forked at apex (Fig. 15); either two orbital bristles at same level (side by
side), or only one orbital bristle; at least some tarsomeres of fore- and hind legs white or yellow,
contrasting with dark brown tibiae; Venezuela to Bolivia and Brazil. . . . . [Anomioptera Schiner]
– R2+3 at most slightly sinuate, unforked; two orbital bristles in usual position, one inserted well in
front of other; coloration of fore- and hind tarsomeres not contrasting with tibiae. . . . . . . . . . . 21

21. Arista short, 1.5 times length of first flagellomere; M strongly pigmented right to wing margin;
Bolivia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Palaeolimosina Duda]
– Arista at least 2 times length of first flagellomere; M weak or not reaching wing margin; Colom-
bia to Argentina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Palaeocoprina Duda, in part]

22. Scutellar margin with only two long pairs of bristles (Fig. 45); first pair of dorsocentrals reclinate;
midtibia without preapical ventral bristle; first tarsomere of midleg with several slightly enlarged
bristles ventrally. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Coproica Macquart
– Scutellar margin with two long and two shorter pairs of bristles (similar to Fig. 46); first pair of
dorsocentrals inclinate (Fig. 47); midtibia with preapical ventral bristle; first tarsomere of midleg
with one distinctly enlarged ventral bristle in basal half (similar to Fig. 41); only species from
South America key here. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rachispoda Lioy, in part

23. Midtibia with preapical ventral bristle; first tarsomere of midleg with enlarged ventral bristle
(similar to Fig. 41). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
– Midtibia without preapical ventral bristle; first tarsomere of midleg without enlarged ventral
bristle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

24. First pair of dorsocentrals enlarged and inclinate (Fig. 47). . . . . . . . . . . . Rachispoda Lioy, in part
– First pair of dorsocentrals small, directed posteriorly. . . . . . . . . . . . . . . Leptocera Olivier, in part

25. Wing with cell cup closed (as in Fig. 18); three pairs of dorsocentral bristles present (anterior pair
may be small). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
– Wing with cell cup open (e.g., Figs. 16, 21); usually less than three pairs of dorsocentral bristles
present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27

26. Costa ending at apex of R4+5; cell cup strong; left and right surstyli similar, bilobed and without
stout bristles; female cerci dorsally shining, fused with tergite 10 and with stout apical bristles;
Colombia to Argentina. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Palaeocoprina Duda, in part]
– Costa extending far beyond apex of R4+5; cell cup weak; left and right surstyli different, each with
three stout bristles; female cerci long, free from tergite 10 and with slender apical setae; Chile
(rare). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Hellerella Duda]

27. Costa extending far (more than three 3 times costal width) beyond end of R4+5. . . . . . . . . . . . . 28
– Costa not or scarcely (no more than costal width) extending beyond end of R4+5. . . . . . . . . . . . 30

28. Scutellum with one to two setulae between long laterobasal and apical pair of bristles; only
species from Mexico key here. . . . . . . . . . . . . . . . . . Pullimosina (Dahlimosina) Roháček, in part
– Scutellum without setulae between the long laterobasal and apical bristles. . . . . . . . . . . . . . . . 29
SPHAEROCERIDAE (small dung flies) 95 1175

29. Two pairs of dorsocentral bristles, anterior pair large and close to suture; male with midventral
bristle on midtibia; female abdomen lacking tergite 10 (Fig. 51); frons broad, with two large and
one small pair of interfrontal bristles; lunule broad, at least somewhat bulging. . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chespiritos Marshall
– Three pairs of dorsocentral bristles, anterior two pairs small, middle pair close to posterior pair;
male without midventral bristle on midtibia; female tergite 10 present; frons normal, with three to
four small interfrontal bristles; lunule triangular. . . . Pullimosina (Dahlimosina) Roháček, in part

30. Midtibia without posterodorsal bristles in basal two-thirds. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

– Midtibia with one or more posterodorsal bristles in basal two-thirds (e.g., Fig. 33). . . . . . . . . . 32

31. Midtibia without midventral bristle; hind tibia without exserted preapical bristle; only one pair of
dorsocentral bristles; epandrium without long bristles; female cerci fused to tergite 10, each bearing
long, thick curved apical bristle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Undescribed Genus A
– Midtibia with midventral bristle; hind tibia with exserted preapical bristle; two large pairs of
dorsocentral bristles; epandrium dorsolaterally with pair of long bristles; female cercus not fused
to tergite 10, only with normal hairs; only species from Mexico and Guatemala key here. . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Spelobia (Spelobia) Spuler, in part

32. R2+3 sharply bent towards costa near apex, apical section in distinct brown spot (Fig. 21); tibiae
with alternating yellow and brown bands (Fig. 33); head and thorax with silvery spots; atypical
specimens (see also couplet 77). . . . . . . . . . . . . . . . . . . . . . . . . . . . . Poecilosomella Duda, in part
– R2+3 not abruptly curved forward; wing membrane without brown spots (Fig. 16); tibiae, head and
thorax not patterned as above. . . . . . . . . . . . . . . . . . Archiceroptera (Archiceroptera) Papp, in part

33. R4+5 strikingly curved forward to meet anterior edge of wing well before wing tip and costa not
extending beyond apex of R4+5 (Fig. 19); female cercus much larger than tergite 10, usually with
long, thick apical bristle; male sternite 5 usually simple, without paired posteromedial lobes;
usually small flies, 1–2 mm long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bromeloecia Spuler
– R4+5 usually straight or less strongly curved forward (e.g., Figs. 16–17, 21–25), and usually ending
closer to wing tip; if strongly upcurved then apex conspicuously bypassed by costa; female cercus
usually with only thin bristles but sometimes with stout, spinelike apical bristles; male sternite 5
various; size various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

34. One orbital bristle, anterior one absent or scarcely distinguishable from additional setulae on orbit
(e.g., Fig. 6) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
– Two orbital bristles, anterior one sometimes small but always distinctly longer than additional
setulae on orbit. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40

35. Three pairs of dorsocentral bristles (anterior pair may be small); midtibia with at least one
posterodorsal bristle in basal two-thirds (e.g., Fig.  27) and/or frons with distinctly enlarged
(almost cruciate) middle pair of interfrontal bristles; South America. . . . . . . . . . . . . . . . . . . . . 36
– One to two pairs of dorsocentral bristles; midtibia without posterodorsal bristles in basal two-
thirds (e.g., Fig. 34); middle pair of interfrontal bristles not distinctly enlarged. . . . . . . . . . . . . 38

36. R2+3 forked, with posterior stump vein; middle pair of interfrontal bristles distinctly enlarged and
almost cruciate; midtibia with single dorsal to posterodorsal bristle in basal two-fifths (Fig. 35);
only species from South America key here. . . . . . . . . . . Robustagramma Marshall & Cui, in part
– R2+3 simple; middle pair of interfrontal bristles not distinctly enlarged, subequal in size to other
pairs; midtibia with at least two more or less paired antero- and posterodorsal bristles in basal
two-fifths (e.g., Fig. 27). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

37. Cell cup open; costa ending at R4+5; scutum and scutellum velvety dark brown with silvery
pruinose pattern; midtibia with at least two posterodorsal bristles in basal two-thirds, first
tarsomere of midleg with enlarged ventral seta; southern South America . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Gyretria biseta (Duda)]
– Cell cup closed; costa extending far beyond R4+5; scutum and scutellum shiny black; midtibia with
one posterodorsal bristle in basal two-thirds (Fig. 27); first tarsomere of midleg without enlarged
ventral seta; high elevations from Ecuador to Bolivia. . . . . . . . . . . [Archileptocera Duda, in part]

38. First segment of R 4+5 (between Rs and r-m) longer than second segment of M (between
crossveins) (Fig.  22); wing almost always maculated (at least faintly); midtibia with isolated
anterodorsal bristle near middle (well separated from proximal bristle and distal pair of bristles)
(Fig. 34); maxillary palpus with single long apical bristle. . . . . . . . . Pterogramma Spuler, in part
– First segment of R4+5 shorter than second segment of M; wing not maculated; midtibia
without isolated anterodorsal bristle near middle (no isolated bristle between proximal one to two
bristle(s) and distal group of three to four bristles) (e.g., Figs. 29, 31); palpus apically only with
short setulae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39

39. Costa extending far beyond apex of R4+5 (~4 times costal width); two pairs of dorsocentral bristles
present; abdomen well sclerotized; male subanal plate with distinctive comblike row of long bristles
(similar to Fig. 49); female terminalia not telescoping; one atypical species from Ecuador. . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Druciatus Marshall, in part
– Costa extending slightly beyond apex of R4+5 (2 times costal width or less); one to two pairs of
dorsocentral bristles present; abdomen sometimes weakly sclerotized; male subanal plate without
comblike row of long bristles; female terminalia telescoping. . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Minilimosina Roháček (sensu lato), in part

40. Cell cup closed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41

– Cell cup open. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43

41. Costa extending far beyond end of R4+5; basal two-thirds of midtibia only with one pair of bristles
dorsally (Fig. 27); hind tibia without exserted preapical bristle; hind femur without anterodorsal
bristle near apex; high elevations from Ecuador to Bolivia. . . . . . . [Archileptocera Duda, in part]
– Costa ending at or little beyond apex of R4+5; basal two-thirds of midtibia at least with four bristles;
hind tibia with long exserted preapical bristle; hind femur with anterodorsal bristle near apex. . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42

42. Dorsocentral bristles present as single, prescutellar pair; head with inclinate bristle between upper
orbital and inner vertical bristles; hind tibia with enlarged anteroventral seta near middle; costa
ending at apex of R4+5. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Limomyza Marshall
– Three dorsocentral bristles present (anterior ones small); head without additional bristles between
upper orbital and inner vertical bristles; hind tibia with uniformly small ventral setulae; costa
extending beyond apex of R4+5; Argentina, Bolivia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Palaeoceroptera Duda (Copromyzinae)]

43. Dorsocentral bristles in three or more pairs (anterior pair often small, requires careful examination) .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
– Dorsocentral bristles in one to two pairs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53

44. Scutellum and scutum velvety dark brown, with conspicuous silvery pruinose pattern. . . . . . . 45
– Scutellum and scutum not velvety, uniformly pruinose, or with faint pruinose pattern . . . . . . . 46
SPHAEROCERIDAE (small dung flies) 95 1177

45. First tarsomere of midleg with strong ventral bristle in basal half; midtibia with proximal
posterodorsal bristles, midventral bristle present in female; southern South America . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Gyretria melanogaster (Thomson)]
– First tarsomere of midleg without ventral bristle; midtibia without midventral or proximal
posterodorsal bristle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetopodella Duda (C. tonsa group)
46. First tarsomere of midleg with large ventral bristle; midtibia without apicoventral bristle but with
preapical ventral bristle in female (Fig. 41); Baja California, high elevations in Ecuador, Venezuela.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Pseudocollinella Duda]
– First tarsomere of midleg usually without large ventral bristle, but if ventral bristle enlarged then
midtibia with apicoventral bristle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47

47. Anterior dorsocentral bristles inclinate, more than one presutural pair present (sometimes small);
coastal. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thoracochaeta Duda
– All dorsocentral bristles reclinate, at most one presutural pair present. . . . . . . . . . . . . . . . . . . . 48

48. Midtibia proximally without posterodorsal bristle (Fig.  37), midventral bristle present in both
sexes; R4+5 apically slightly recurved (Fig. 24) . . . . . . . . . . . . . . . . Spelobia (Bifronsina) Roháček
– Midtibia proximally with at least small posterodorsal bristle, if without proximal posterodorsal
then midventral absent in both sexes; R4+5 straight or slightly bent anteriorly . . . . . . . . . . . . . . 49

49. Katepisternum with two large, subequal bristles; frons anteriorly with additional row of small
setulae between interfrontal row and fronto-orbital plate; wing membrane distinctly whitish-brown
opaque; coastal Chile. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Archicollinella Duda]
– Katepisternum with at most one large bristle and one small bristle; frons without additional row
of setulae outside interfrontal row; wing membrane sometimes infuscated but not whitish opaque;
widespread, rarely coastal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
50. Midtibia proximally with isolated anterodorsal, dorsal or posterodorsal bristle (e.g., Fig. 35) (if
with pair of bristles then wing distinctly patterned and costal sector  2 less than half as long
as costal sector 3); midtibia without midventral bristle; R2+3 usually angulate (apically abruptly
curved forward), forked or straight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
– Midtibia proximally with one to two anterodorsal bristles, and one posterodorsal (e.g., Fig. 30;
small in some Lepidosina males); midtibia of female with midventral bristle; costal sector  2
slightly shorter, equal, or longer than costal sector 3; R2+3 gently curved forward apically (neither
angulate, forked, nor straight), wing unpatterned . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
51. Midtibia dorsally with at least five bristles (including smaller ones), proximal bristle dorsal or
posterodorsal (Fig. 35); R2+3 straight to angulate (e.g., Fig. 23), sometimes forked; wing membrane
sometimes distinctly patterned; interfrontals often with one pair cruciate and distinctly longer
than others . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Robustagramma Marshall & Cui, in part
– Midtibia dorsally with exactly four bristles, proximal bristle anterodorsal (as in Fig.  34); R2+3
angulate (apically abruptly curved forward), not forked (similar to Fig. 21); wing at most slightly
clouded (mainly around certain veins); interfrontals short, subequal . . . . . . Undescribed Genus D

52. Body heavily gray-pruinose, some acrostichals usually enlarged; dorsal surface of midtibia distally
only with pair of bristles (one of these bristles sometimes reduced or absent in males), without
additional smaller bristles (Fig. 30); male foretibia often clavate, foretarsus broadened and flattened;
male often with long scalelike bristles on sternite 5 and with syntergosternite six to eight forming
a complete ring, surstylus not divided; female cercus scoop-shaped with flattened apical bristle .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepidosina Marshall
– Body not heavily gray-pruinose, acrostichals not enlarged; dorsal surface of midtibia distally with
two to three larger and three to four smaller bristles (Fig. 32); male foretibia and tarsus normal;
surstylus divided into anterior and posterior sections; male and female genitalia not as above;
Caribbean, South America. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Phthitia Enderlein, in part]
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

53. Interfrontal bristles with one pair cruciate and distinctly larger than others (e.g., Fig. 5). . . . . . 54
– Interfrontal bristles in three or more subequal pairs (sometimes all small and difficult to see) . . . 58

54. Costa not or slightly extending beyond apex of R4+5 (at most 1.5 times costal width) . . . . . . . . 55
– Costa extending distinctly beyond apex of R4+5 (over 3 times costal width). . . . . . . . . . . . . . . . 56

55. Midtibia with strong midventral bristle in both sexes (Fig. 42); lunule narrow, triangular; scutum
and scutellum not flattened; tarsi slender; male sternite  5 posteromedially with conspicuous
spinelike bristles; wing not patterned. . . . . . . . . . . . . . . . . . . . . Spelobia (Spelobia) Spuler, in part
– Not with the above combination of characters: either midtibia without midventral bristle and/
or lunule broad and rounded or (if midventral present) thoracic scutum and scutellum some-
what flattened and tarsi relatively short and stout; male sternite 5 without conspicuous spinelike
bristles; wing sometimes patterned . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Undescribed Genera

56. Face bulging and prominent, often continuous with extremely broad, rounded lunule (Fig.  4);
frons distinctly shortened (distance between anterior ocellus and ptilinal suture shorter than
distance between enlarged interfrontals). . . . . . . . . . . . . . . . . . . . . . . . . . . . Undescribed Genus B
– Face at most slightly bulging, lunule broad to narrow; frons not distinctly shortened (distance between
anterior ocellus and ptilinal suture greater than distance between enlarged interfrontals). . . . . . . 57

57. Both katepisternal bristles minute; male subanal plate with distinctive comblike row of long
bristles (Fig. 49); midventral midtibial bristle sometimes present in either sex . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Druciatus Marshall, in part
– Posterior katepisternal bristle well developed; male subanal plate with at most one or two bristles;
midventral midtibial bristle present in female, absent in male; only one species from Mexico keys
here. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pullimosina (Pullimosina) Roháček, in part

58. Lunule rounded and prominent (Fig. 7); alula broad (circa half as broad as long); postsutural intra-
alar bristle well developed (Fig. 48); midtibia with small proximal posterodorsal bristle (Fig. 36);
first tarsomere of midleg with small ventral bristle; female with midventral midtibial bristle, male
without; male terminalia complex, especially genital pouch. . . . . . . . . . . . . Sclerocoelus Marshall
– Lunule narrow or triangular; alula usually narrow, if broad midtibia without proximal posterodorsal
bristle; intra-alars usually not developed, if present midtibia without proximal posterodorsal
bristle; midventral bristles of midtibia present or absent; male terminalia various, but genital
pouch usually simple, not consisting of several sclerites as in Sclerocoelus. . . . . . . . . . . . . . . . 59

59. Midtibia without apicoventral bristle in both sexes, first tarsomere of midleg with large ventral
bristle (distinctly longer than diameter of first tarsomere) (Fig. 40); dorsocentral bristles in two
well-developed pairs; costa with two subequal costagial bristles; male hypopygium almost always
with prominent, stout, ventrally directed cercal bristles (Fig. 50); female tergite 10 long and shiny,
longitudinally folded; cercus apically with two flat bristles (Fig. 52). . . . . . . . . . Opacifrons Duda
– Midtibia with apicoventral bristle (if rarely absent in male then either first tarsomere of midleg
without large ventral bristle or dorsocentral bristles in single, prescutellar pair); first tarsomere of
midleg usually without enlarged ventral bristle; female tergite 10 usually small, not shiny; cercus
usually without flattened bristles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
60. Costa extending far beyond apex of R4+5 (at least 4 times costal width). . . . . . . . . . . . . . . . . . . 61
– Costa ending at apex of R4+5 or slightly beyond (less than 4 times costal width). . . . . . . . . . . . 66

Note: Doubtful cases will key both ways.

SPHAEROCERIDAE (small dung flies) 95 1179

61. Two pairs of dorsocentral bristles present; frons without additional row of lateroclinate or incli-
nate setulae between orbital and interfrontal bristles; costagial bristles of wing subequal and not
unusually long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
– One pair of dorsocentral bristles present; frons with additional row (at least one seta) of laterocli-
nate or inclinate setulae between orbital and interfrontal bristles and/or wing with one costagial
bristle enlarged. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63

62. Crossveins r-m and dm-cu closely approximated, distance between them less than length of dm-
cu; midtibia with midventral bristle in female; epandrium dorsolaterally with outstanding pair of
(fine) bristles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Pullimosina Roháček (sensu stricto), in part
– Crossveins r-m and dm-cu well separated or less approximated (distance between them greater
than length of dm-cu); midventral bristle of midtibia various; epandrium dorsolaterally without
outstanding bristles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66

63. Small flies (less than 1 mm in body length); frons with additional row of inclinate setulae between
orbital and interfrontal bristles; costa with few enlarged setae projecting almost perpendicularly
above wing plane, but costagium without prominent bristle; crossveins almost at same level; Ca-
ribbean. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Elachisoma Rondani]
– Body size various; frons usually without additional row of inclinate setulae; if inclinate setu-
lae present then costagial bristle long and prominent; costa without setae projecting above wing
plane; crossveins clearly separated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64

64. Small flies, usually 1 mm or less in body length; frons with one or more (often a row) of
lateroclinate bristles between interfrontal and orbital bristles; R4+5 strongly upcurved to almost
straight; eye sometimes setose; first flagellomere often strongly pointed apically, with apical arista.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trachyopella Duda
– Body length usually over 1 mm; frons without additional lateroclinate bristles between interfrontal
and orbital bristles; R4+5 slightly curved forward to straight; eye bare; first flagellomere not pointed
apically, arista subapical. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65

65. Costa with costagial bristle much longer than surrounding bristles; alula broad, hind margin convex;
midtibia with midventral bristle in both sexes, apex with several bristles. . . Gonioneura Rondani
– Costagial bristle not long (but sometimes slightly enlarged); midventral bristle present or absent,
apex of midtibia usually with single apicoventral bristle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66

66. Abdomen weakly sclerotized to almost completely desclerotized . . . . . . . . . . . . . . . . . . . . . . . 67

– Abdomen normally to heavily sclerotized. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68

67. One pair of dorsocentral bristles present; tarsomere 2 of hind leg more or less swollen (Fig. 43);
hind tibia almost always with exserted preapical bristle; midtibia with posterodorsal bristle near
middle and/or one near proximal fourth (rarely missing) (Fig. 28); head often partly to completely
orange or pale yellow; first flagellomere often pointed and tapered apically, with apical or
subapical arista; two spermathecae present. . . . . . . . . . . . . . . . . . . . . . . Bitheca Marshall, in part
– Two pairs of dorsocentral bristles present (anterior pair sometimes small and difficult to see);
tarsomere 2 of hind leg not conspicuously swollen; hind tibia without exserted preapical bristle;
midtibia without posterodorsal bristles in basal half (Fig. 31); head (except sometimes antennae)
black; first flagellomere not pointed and tapered, arista not apical; three spermathecae present . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Minilimosina (Svarciella) Roháček, in part

68. Basal half of midtibia devoid of bristles; dorsal surface of tibia with two to three bristles (including
smaller ones); high elevations in Ecuador. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . [males of undescribed species of uncertain generic affinities]
– Basal half of midtibia with at least one bristle; dorsal surface of tibia at least with four bristles
(including smaller ones, if present). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
1180
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

69. Dorsal surface of midtibia with four well-developed bristles (no additional smaller bristles
present): the bristle near middle isolated and approximately equidistant from proximal bristle and
distal pair of bristles (as in Fig. 34). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
– Dorsal surface of midtibia either with more than four bristles (including smaller ones) or with four
bristles arranged in distal group of three and single proximal bristle (but without isolated bristle
near middle) (e.g., Fig. 31). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74

70. Frons somewhat shortened, with only two large pairs of interfrontal bristles (besides small
anterior pair); orbital bristles removed from eye margin; shortest distance between orbital and eye
margin subequal to shortest distance between orbital and closest interfrontal; first tarsomere of
midleg with strong ventral bristle that is longer than diameter of tarsomere; dorsocentral bristles
in one pair; sternite 5 short, with posterior pair of processes; female cerci simple, not fused to
tergite 10. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Undescribed Genus C
– Frons not shortened, with three pairs of interfrontal bristles (besides small anterior pair); orbital
bristles close to eye margin; shortest distance between orbital and eye margin circa half
shortest distance between orbital and closest interfrontal; first tarsomere of midleg sometimes
with slightly enlarged bristle that is shorter than diameter of tarsus; dorsocentral bristles in one to two
pairs; sternite 5 not short and without processes; female cerci simple or fused to tergite 10. . . . . . . 71

71. Anterior pair of dorsocentral bristles long, approximately three-quarters as long as prescutellar
pair; R2+3 apically abruptly curved anteriorly towards costa (apical section sometimes perpendicular
to costa); wing sometimes strongly patterned. . . . . . . . . . . . . . . . . . . . . . . . . Undescribed Genera
– Anterior pair of dorsocentral bristles short, less than half as long as prescutellar pair; R2+3 apically
not abruptly curved towards costa; wing without pattern but often evenly infuscated. . . . . . . . 72

72. With one or more of the following characters: only one pair of dorsocentral bristles present; only
one well-developed costagial bristle (sometimes extremely long) present; costal sector 2 distinctly
shorter than costal sector 3 (two-thirds as long or less) (Fig. 17); eye small, its height less than 2.5
times genal height; epandrium with outstanding bristles; female tergite 10 and cerci fused, shiny.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Archiceroptera Papp genus complex, in part
Note: Rudolfina Roháček and several other clades of uncertain affinities key out at this couplet.

– Two pairs of dorsocentral bristles present (anterior pair usually small); two subequal costagial
bristles present; costal sector 2 subequal to or longer than costal sector 3; eye large, its height
more than 2.5 times genal height; epandrium uniformly haired; female tergite 10 and cerci not
fused and shiny. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73

73. Male sternite 5 large, more strongly sclerotized than preceding sternites, with broadly rounded
posteromedial projection; midtibia with midventral bristle in female; female sternite 8 large,
subquadrate and well sclerotized; Argentina, Chile. . . . . . . [Parasclerocoelus Marshall & Dong]
– Male sternite 5 not unusually large, without posteromedial projection, not more strongly sclerotized
than preceding sternites; midtibia without midventral bristle; female sternite 8 reduced or small;
high elevations in Ecuador, Bolivia. . . . . . . . [undescribed species of uncertain generic affinities]

74. Midtibia with posterodorsal and/or dorsal bristle(s) in basal three-fifths (e.g., Figs. 26, 28,
33, 35) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
– Dorsal surface of midtibia only with anterodorsal bristle(s) in basal three-fifths (e.g.,
Figs. 37–38). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79

75. Basal three-fifths of midtibia without posterodorsal bristle besides proximal bristle that is dorsal
or posterodorsal (e.g., Fig. 35). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
– Basal three-fifths of midtibia with at least one posterodorsal bristle in addition to proximal bristle
that is anterodorsal, dorsal, or posterodorsal (e.g., Figs. 26, 28, 33). . . . . . . . . . . . . . . . . . . . . . 77
SPHAEROCERIDAE (small dung flies) 95 1181

76. Costal sector 2 short, less than two-thirds as long as costal sector  3 (usually half as long or
less) (Fig. 23); R2+3 short, usually straight; wing usually faintly patterned; two (rarely one) pairs
of dorsocentral setae present; midtibia without midventral bristle; female genitalia simple, cerci
separate from tergite 10, with simple hairs . . . . . . . . . . . Robustagramma Marshall & Cui, in part
– Costal sector 2 longer than costal sector 3 (Fig. 16); R2+3 longer, curved near apex; wing not
patterned; one pair of dorsocentral setae present; midtibia with midventral bristle in female; female
genitalia modified, tergite 10 and cerci fused, each cercus with stout flattened apical spine . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Archiceroptera Papp genus complex, in part
77. R2+3 angulate, sharply bent towards costa near apex, apical section in distinct brown spot (Fig. 21);
tibiae with alternating yellow and brown bands (Fig. 33); head and thorax with silvery spots. . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Poecilosomella Duda, in part
– R2+3 not angulate, gently curved forward near apex and unspotted; tibiae not prominently banded;
body without silvery spots. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78

78. Frons without additional row of inclinate setulae; interfrontal bristles uniformly small; hind tibia with
exserted preapical bristle, tarsomere 2 of hind leg more or less swollen (Fig. 43); first flagellomere
sometimes pointed, with apical arista; two spermathecae present. . . . . . Bitheca Marshall, in part
– Frons with additional row of inclinate setulae outside interfrontal row; interfrontals not strikingly
small; hind tibia usually without exserted preapical bristle; tarsomere 2 of hind leg not conspicuously
swollen; first flagellomere not pointed, arista subapical; three spermathecae present. . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Archiceroptera Papp genus complex, in part

79. Midtibia without midventral bristle, male sometimes with ventral row of spinules or short bristles
distally (Fig.  39); costa often extending distinctly beyond apex of R4+5; female abdomen
telescoping. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Minilimosina Roháček (sensu lato), in part
– Midtibia with midventral (or midanteroventral) bristle in both sexes (as in Fig.  42), male
without ventral row of spinules or short bristles; costa at most slightly extending beyond R4+5 (3
times costal width or less); female abdomen truncate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80

80. Only one dorsocentral bristle present; R4+5 strongly curved anteriorly, costa ending at apex of R4+5
(Fig. 25); epandrium with several thick, relatively short bristles . . . . . . . . Spinilimosina Roháček
– Two dorsocentral bristles present; R4+5 straight or only slightly curved anteriorly, if more strongly
so then costa usually extending beyond its apex; epandrium without strong bristles, sometimes
with enlarged slender setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81

81. Hind tibia with extremely long spurlike seta (length approximately half length of tibia) originating
well before apex of tibia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Opalimosina Roháček
– Hind tibia without spurlike seta. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82

82. Eye small, less than 2 times genal height; male sternite 5 with pair of posterior lobes; surstylus
elongate and pointed, without apical spine in Neotropical species; postgonites long and projecting
far below ventral margin of epandrium (easily visible without preparation); epandrium without
dorsolateral bristles; female cercus only with short straight hairs some of which are thickened;
Mexico, Caribbean, South America. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Telomerina Roháček]
– Eye large, more than 2.5 times genal height in Neotropical species; male sternite 5 not bilobate;
surstylus usually much wider than long or deeply bilobate, rarely elongate and pointed, but then with
blunt apical spine; postgonites not long and not conspicuously projecting; epandrium with pair of
dorsolateral bristles; female cercus with long sinuate hairs. . . . . . Spelobia (Spelobia) Spuler, in part
1182
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

83. Ocellar bristles inserted slightly to distinctly before level of anterior ocellus (Fig.  3), rarely
absent; if ocelli absent then with two orbital bristles present; midtibia usually without bristles on
dorsal surface of basal two-fifths, but if so then also with posteroventral bristle(s) in distal half;
hind tibia with exserted preapical bristle (sometimes small); four, rarely two scutellar bristles
(anterior pair often hairlike); COPROMYZINAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
– Ocellar bristles inserted between ocelli; if ocelli absent, with zero to one orbital bristles; midtibia
almost always with one or more bristles on dorsal surface of basal two-fifths, without posteroventral
bristles; hind tibia usually without exserted preapical bristle, if present scutellum with more than
four bristles (including smaller ones); LIMOSININAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87

84. Wing rudiment linear and cylindrical, about as long as width of scutellum, with distinct apical
bristle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
– Wing rudiment short and scalelike or elongate and strap-shaped, without distinct apical bristle, or
apterous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86

85. Anterior notopleural bristle long, 0.6–0.7 times width of thorax; scutum strikingly flattened;
dorsocentral bristles in one prescutellar pair which may be small; head, thorax, and legs yellow;
Falkland Islands. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Penola Richards]
– Anterior notopleural bristle not unusually long, its length at most 0.4 times width of thorax;
scutum slightly convex; dorsocentral bristles usually in three pairs but anterior two pairs
sometimes reduced; at most head yellow, thorax and legs (excluding tarsi and trochanters) black
to brown; Chile, western Argentina. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Frutillaria Richards]

Note: The relationships of these two genera require further study. Frutillaria might be a synonym of Penola.

86. Ocelli absent; first tarsomere of foreleg of male with apicoventral, blunt, thornlike projection;
Chile, Falkland Islands, South Georgia Island. . . . . . . . . . . . . . . . . . . . . . . . . [Antrops Enderlein]
– Ocelli present; first tarsomere of foreleg of male usually without apicoventral projection. . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Archiborborus (Archiborborus) Duda, in part

Note: The relationships of these two genera require further study.

87. Scutellum with eight bristles (anterior pair short) (Fig. 46); scutum with two pairs of greatly enlarged
acrostichal bristles; midtibia with ventral preapical but no apical bristle (similar to Fig. 41); first
tarsomere of midleg with strong ventral bristle; only species from Juan Fernández Islands key
here. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leptocera Olivier, in part
– Scutellum at most with six bristles; acrostichals not enlarged; midtibia without ventral preapical
bristle but sometimes with preapical antero- or posteroventral bristles; first tarsomere of midleg at
most with small ventral bristle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88

88. With at most one pair of orbital bristles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89

– With two pairs of orbital bristles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92

89. Orbital and outer vertical bristles absent; ocellar and/or inner vertical bristles sometimes also
reduced. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aptilotella Duda
– With one orbital bristle, outer verticals rarely missing (some Pterogramma); ocellar and inner
vertical bristles well developed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90

90. Midtibia proximally only with one bristle (Fig. 34) . . . . . . . . . . . . . Pterogramma Spuler, in part
– Midtibia proximally with two more or less paired bristles (one anterodorsal, one posterodorsal)
(e.g., Fig. 27) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
SPHAEROCERIDAE (small dung flies) 95 1183

91. Brachypterous; midtibia with midventral bristle; interfrontal bristles well developed; abdomen
not petiolate; fore- and hind tarsi without white tarsomeres; high elevations from Ecuador to
Bolivia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Archileptocera Duda, in part]
– Apterous; midtibia without midventral bristle; interfrontal bristles minute; abdomen petiolate,
habitus antlike; fore- and hind tarsi with contrasting black and white tarsomeres . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myrmolimosina Marshall

92. Midtibia without proximal posterodorsal bristle; apterous or brachypterous . . . . . . . . . . . . . . . 93

– Midtibia with proximal posterodorsal bristle; brachypterous . . . . . . . . . . . . . . . . . . . . . . . . . . . 94

93. Midtibia without midventral bristle; eye large; four scutellar bristles present; hind tibia without
exserted preapical bristle; foretibia not clavate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aptilotus Mik
– Midtibia with midventral bristle (Fig. 42); eye in one species from Mexico strongly reduced; one
species from Guatemala with six scutellar bristles (anterior pair small and hairlike), exserted preapical
bristle on hind tibia, and clavate male foretibia. . . . . . . . . . . . . . Spelobia (Spelobia) Spuler, in part

94. Two pairs of dorsocentral bristles present (anterior pair small); Mexico, Ecuador, Bolivia. . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [undescribed species of uncertain affinities]
– Three pairs of dorsocentral bristles present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95

95. Midventral midtibial bristle present in both sexes; female terminalia laterally compressed; midtibia
proximally only with one pair of bristles (Fig. 27); head and thorax black, in part shining; wing
reduction various but wing rudiment not narrow and elongate; high elevations from Ecuador to
Bolivia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Archileptocera Duda, in part]
– Midventral midtibial bristle absent in male; female genitalia not compressed; midtibia proximally
sometimes with two pairs of bristles; head and thorax sometimes pale brown, not shining; wing
rudiment either narrow and elongate or broad; only species from South America, Juan Fernández
Islands key here. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Phthitia Enderlein, in part]

Synopsis of the fauna species. Five species have been found in Costa Rica. At least
one of the Central American species is brachypterous, and
The approximately 100 species currently known from most species are associated primarily with dung. The
Central America is a small fraction of the actual fauna, and subgenera and species groups keyed out here should probably
there are at least another 500 occurring there. They are found be elevated to generic rank.
in all habitats, from the seashore to the highest elevations.
Archiceroptera Papp genus complex. This group compris-
Aptilotella Duda. This might be just an apterous clade of es several distinct lineages (including Rudolfina Roháček),
the large genus Pterogramma. Several undescribed species some of which might deserve generic rank. They are mostly
occur at higher elevations in Central America, and additional Neotropical, with some species in the Nearctic Region and
species occur in South America. Adults live in leaf litter and one species of Rudolfina in the Palearctic Region. One
can be collected using pan traps, pitfall traps, or Berlese described and numerous undescribed species occur in Cen-
funnels. tral America. All appear to be associated with dung and can
be collected in large numbers in dung traps.
Aptilotus Mik. This is a largely Holarctic genus that
reaches as far south as Panama (Marshall, 1997a). The Bitheca Marshall. This is a mostly Neotropical genus
Central American species (three described, at least one with six described (Marshall, 1987) and probably many
undescribed) are apterous and occur in leaf litter of montane undescribed species from Central America. At least some
forests or paramos, usually above 2500 m (rarely as low as species appear to be associated with fungi, but adults are also
1400 m). attracted to dung and carrion.

Archiborborus Duda. This is a Neotropical genus that Bromeloecia Spuler. This is a large, primarily Neotropical
reaches its highest diversity in South America (Richards, genus with one described and many undescribed species in
1931, 1961). The Central American fauna includes one Central America. It is one of the most prevalent sphaerocerid
described species from Mexico and at least 13 undescribed genera on dung.
1184
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Chaetopodella Duda (C. tonsa group). The only described L. inaequalis (Malloch) and L. evanescens Marshall & Buck,
species of this Neotropical clade (from Costa Rica) was occur in Costa Rica; L. angusticercus Marshall & Buck has
placed in the Old World genus Chaetopodella but actually been recorded from Panama (Marshall et al., 2007). Their
represents an undescribed genus. Few species occur biology is unknown, and most species are not attracted to
in Central America, and they appear to be associated with baits.
dung.
Leptocera Olivier. This is a moderately large genus of
Chespiritos Marshall. Described as a monotypic genus worldwide distribution. The New World species were re-
from Costa Rica (Marshall, 2000a), this genus is distributed cently revised (Buck  & Marshall, 2009), and include 10
throughout tropical parts of the Neotropical Region and contains Central American species, six of which have been recorded
many undescribed species. The type species is associated from Costa Rica. One of the Central American species is
with downed bromeliads, and other species probably have subcosmopolitan, one is widespread in the New World, and
specialized phytosaprophagous habits as well. two are widespread in the Neotropical Region. The remaining
species are either endemic to Central America (one species
Coproica Rondani. This genus is distributed throughout reaches the extreme southern United States) or occur in
all major biogeographical regions and contains several Panama and the northern half of South America (one spe-
widespread synanthropic species, including most of those cies). Except for the subcosmopolitan, polysaprophagous
in Central America. Five described (Richards, 1960) and L. caenosa (Rondani), most species are probably associated
at least two undescribed species occur in Central America. with decaying vegetation. Adults are rarely attracted to baits
They are associated with dung and to a lesser degree with (dung, carrion) but are well represented in Malaise trap and
carrion and decaying vegetable matter (e.g., compost). aerial sweep samples.
Copromyza Fallén. This genus is represented in Central Limomyza Marshall. This is a small, distinct genus restricted
America by the synanthropic, primarily Holarctic, and to the New World. Only one species is known from higher
probably introduced C.  equina Fallén (Norrbom  & Kim, elevations in Central America (Guatemala), but three further
1985), which breeds in dung. species occur in Mexico (Marshall, 1997b). Apparently
developing in dung, one Nearctic species lives in caves.
Druciatus Marshall. This is a relatively small and
uncommon genus restricted to warm parts of the Neotropical Mesosphaerocera Kim. See discussion under Parasphaero-
Region. Three species have been described from Central cera. Four of the five described species of Mesosphaerocera
America (Marshall  & Totton, 1995), and a few more occur in Central America, the other is known only from
undescribed species can be expected. Adults are associated Mexico (Kim, 1972; Papp, 1978). Two species are recorded
with decaying vegetation, such as treefalls, compost, and from Costa Rica.
leaf litter.
Minilimosina Roháček. The New World Minilimosina
Gonioneura Rondani. This is a small, primarily Holarc- were revised by Marshall (1985b). One species of the subgenus
tic genus that is restricted to medium and high elevations Allolimosina Roháček, M.  (A.)  rotundipennis (Malloch), is
in Central and northern South America (Marshall, 1982). known from southern Mexico, the Caribbean, and South
One of the two Central American species appears to be America. The primarily Neotropical subgenus Amputella
undescribed, and all are associated with dung. Marshall is represented in Central America by four species.
Homalomitra Borgmeier. Rarely collected, and apparently The nominate subgenus is mainly temperate Holarctic with
associated with army ants, this Neotropical genus has two one species from southern Mexico and one from South America.
described species from Costa Rica (Roháček  & Marshall, The subgenus Svarciella Roháček, which occurs in most
1998). biogeographical regions, is more diverse in tropical regions.
Two species have been recorded from Central America, and
Ischiolepta Lioy. This is a primarily Old World/Nearctic some undescribed species probably occur there. Several
genus with one endemic Central American species (Buck & undescribed species that will key to Minilimosina are of
Marshall, 2002). The subcosmopolitan species I.  pusilla uncertain subgeneric or even generic affiliation. The biology
(Fallén) might be discovered in Central America in the of Neotropical species is insufficiently known. Most species
future. Another widespread species, I. scabricula (Haliday), are attracted to dung and carrion, but some extralimital species
has been introduced to Brazil. The Costa Rican I. hyalophora are known to breed in decaying vegetation or fungi.
Buck  & Marshall was collected on compost, whereas
I. pusilla is primarily coprophagous. Myrmolimosina Marshall. Although described as a
monotypic genus from Chiapas, Mexico (Marshall, 2000b),
Lepidosina Marshall. This is a small and rarely collected one undescribed species from Guatemala probably belongs
genus with 11 species found mostly at lower elevations from here. The apterous adults live in leaf litter and have a
the southern United States to northern Argentina. Two species, strikingly antlike habitus.
SPHAEROCERIDAE (small dung flies) 95 1185

Neosphaerocera Kim. See discussion under Parasphaero- Pterogramma Spuler. This is one of the predominant
cera. Two of the nine species of Neosphaerocera are known Neotropical sphaerocerid genera, with only a few species in
from Central America, and both are recorded from Costa the Nearctic Region. Sixteen species are known from Central
Rica (Mourgués-Schurter, 1981). The group is otherwise America (Smith & Marshall, 2004), and a large number of
South American. species still await description. They are associated with fresh,
decaying vegetation (e.g., treefalls) and are ubiquitous in
Norrbomia Papp. Although Norrbomia is primarily an tropical forests from sea level to higher elevations. At higher
Old World/Nearctic genus, the widespread Holarctic species elevations many species are brachypterous or apterous.
N. sordida (Zetterstedt) ranges south to Guatemala (Marshall &
Norrbom, 1993). Four other species are known from Mexico, Pullimosina Roháček. This Old World/Nearctic genus
and all are associated with dung. is represented in Central America by P.  zayensis Marshall
(Marshall, 1986). The generic affiliation of Central and
Opacifrons Duda. This is a moderate-sized genus distributed South American species tentatively placed in the subgenus
throughout all major biogeographical regions. The Central Dahlimosina Roháček needs to be reviewed. They are as-
American fauna comprises seven species (Marshall  & sociated with decayed vegetation, including grass clippings
Langstaff, 1998). They are most common along streams and compost.
where the larvae probably develop in pockets of drift.
Pycnopota Bezzi. This genus includes a single described
Opalimosina Roháček. This primarily Old World genus species from Bolivia and at least two undescribed species,
is represented in Central America by the introduced, synan- one of which was recently collected in Costa Rica. Pycnopota
thropic, subcosmopolitan O. mirabilis (Collin) (redescription was treated as an aberrant member of Copromyzinae by
in Roháček, 1983, 1985). It is a polysaprophagous species, Roháček et al. (2001), although some character states,
predominantly on dung, less frequently on compost. including a spinose costa, suggest that the genus should be
placed in the Heleomyzidae. Examination of new material
Parasphaerocera Spuler. Although we have followed from Costa Rica confirms that Pycnopota is correctly placed
the recent catalog of world Sphaeroceridae in recogniz- in the Sphaeroceridae. Members of this genus appear to be
ing the genera Parasphaerocera, Mesosphaerocera, and nocturnal, which might explain their rarity in collections.
Neosphaerocera, these closely related genera are difficult
to distinguish. We expect that a careful reconsideration of Rachispoda Lioy. This genus is closely related to Lepto-
cera and probably renders the latter paraphyletic (Buck  &
the relationships of this clade will result in synonymy
Marshall, 2009). Worldwide in distribution, Rachispoda is
of Neosphaerocera and Mesosphaerocera under the older
the largest genus of Sphaeroceridae. Remarkably few (11)
name Parasphaerocera. Parasphaerocera as recognized
species have been reported from Central America (Wheeler,
here is probably rendered paraphyletic by the recogni-
1995; Wheeler  & Marshall, 1995), and few undescribed
tion of Neosphaerocera. This group of genera is restricted
species are expected. The genus is far more diverse in North
to the New World and is mainly Neotropical. Some species
and South America. Most species are associated with riparian
of Parasphaerocera and Mesosphaerocera extend northwards
habitats, especially muddy areas along streams and lakes,
into the Nearctic Region. Species of all three genera are and some species live in habitats influenced by salt.
associated with dung and decaying vegetation. Thirty-one
Neotropical species of Parasphaerocera are recognized, 24 Robustagramma Marshall & Cui. This group was recently
of which occur in Central America, including 16 known from separated from Pterogramma (Marshall & Cui, 2005). It is a
Costa Rica (Kim, 1972; Papp, 1978). large Neotropical genus with a few species whose range
extends into the Nearctic Region. Fourteen species have
Podiomitra Marshall & Roháček. This monotypic genus been described from Central America; most appear to be
was described from Costa Rica (Marshall & Roháček, 2003). associated with dung.
The type species is only known from the holotype, which
was collected in a Malaise trap. They are probably associ- Sclerocoelus Marshall. Along with Pterogramma, this
ated with army ants as are other species of the subfamily is one of the largest Neotropical genera of Sphaeroceridae
Homalomitrinae. (a few species extend north into the Nearctic Region). Only
four described species have been recorded from Central
Poecilosomella Duda. The only Central American America (Marshall, 1997c), but many more are recognized
species of this primarily tropical Old World/Australasian and await description. Most species appear to be associated
genus, P.  angulata (Thomson), was probably introduced with decayed vegetation, and adults are common in sweep
from the Afrotropical Region. It now occurs in many Central samples and pan traps.
and South American countries and is associated with dung.
Diagnosis and illustrations of genitalia are given by Papp Spelobia Spuler. This is a mostly Holarctic genus that
(1990, 1991). ranges along the mountain chains through Central America
1186
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

to Venezuela and Ecuador (Marshall, 1985a). The Central are recognized but undescribed, and most are associated
American fauna consists of four described species and as with decayed vegetation. Some extralimital species develop
many undescribed species, all belonging to the nominate in dung or carrion.
subgenus. The subcosmopolitan, synanthropic S. (Bifronsina)
bifrons (Stenhammar) was probably introduced to Central Undescribed Genus A. This is a small group of unde-
America. Most species appear to be associated with dung, scribed species often collected at dung in Central and South
but some extralimital species are known to breed in fungi America.
and carrion.
Undescribed Genus B. This small, distinctive, and rarely
Spinilimosina Roháček. This is a relatively small Old collected Neotropical genus has five undescribed species in
World/Australasian genus with one rare, introduced species Central America. Adults are attracted to fresh, decaying
in Central America: S.  brevicostata (Duda) (redescribed in vegetation and are often associated with treefalls.
Roháček, 1983, 1985). A second introduced species,
S.  rufifrons (Duda), known from South America, might be Undescribed Genus C. This is a small but well-defined
found in Central America in the future. These flies are group of undescribed species known from Central America
probably coprophagous. to Ecuador. Most specimens are either from Malaise traps or
Thoracochaeta Duda. These flies are widespread along collected in association with treefalls.
seashores of the world, mostly in temperate regions. The
Undescribed Genus D. This is a poorly defined group of
only Central American species T. brachystoma (Stenhammar)
has a subcosmopolitan distribution but appears to be rare undescribed species, superficially similar to Poecilosomella.
in tropical parts of the Neotropical Region (Marshall  & Species from Barbados to Argentina have been tentatively
Roháček, 2000). The larvae develop in wrack. placed in this group.

Trachyopella Duda. This is a moderately large genus of

worldwide distribution. Several Central American species

Literature cited

Buck, M.  & S.A. Marshall. 2002. A review of Ischiolepta Lioy Marshall, S.A. 1997a. A first record of Aptilotus Mik (Diptera:
(Diptera: Sphaeroceridae), including five new species and a key Sphaeroceridae) from the Neotropical region, with the descrip-
to New World species. Studia dipterologica 9: 249–281. tion of three new wingless species of the Aptilotus paradoxus
Buck, M. & S.A. Marshall. 2009. Revision of New World Leptocera group from high elevations in Costa Rica. Proceedings of the
Olivier (Diptera, Sphaeroceridae). Zootaxa 2039: 1–139. Entomological Society of Washington 99: 505–512.
Gómez, L.D. 1977. La mosca del cangrejo terrestre Cardisoma Marshall, S.A. 1997b. Limomyza, a new genus of primitive Limosininae
crassum Smith (Crustacea: Gecarcinidae) en la Isla del Coco, (Diptera: Sphaeroceridae), with five new species from United
Costa Rica. Revista de Biología Tropical 25: 59–63. States, Mexico and Central America. Proceedings of the Ento-
Kim, K.C. 1972. The New World genus Parasphaerocera and allied mological Society of Washington 99: 279–289.
groups, with descriptions of new genera and species (Diptera: Marshall, S.A. 1997c. A revision of the Sclerocoelus galapagensis
Sphaeroceridae). Miscellaneous Publications of the Entomological group (Diptera: Sphaeroceridae: Limosininae). Insecta Mundi
Society of America 8: 377–444. 11: 97–115.
Marshall, S.A. 1982. A revision of Halidayina Duda (Dip- Marshall, S.A. 1998. A revision of the Archileptocera group, includ-
tera: Sphaeroceridae). The Canadian Entomologist 114: ing Anomioptera Schiner, Palaeocoprina Duda and Archileptocera
841–847. Duda, with a key to sphaerocerid genera with similar wing venation
Marshall, S.A. 1985a. A revision of the genus Spelobia Spuler (Diptera: and a description of a new species of Palaeoceroptera Duda (Dip-
Sphaeroceridae) in North America and Mexico. Transactions of the tera: Sphaeroceridae). Journal of Natural History 32: 173–216.
American Entomological Society 111: 1–101. Marshall, S.A. 2000a. Chespiritos, a new genus of Limosininae
Marshall, S.A. 1985b. A revision of the New World species of (Diptera: Sphaeroceridae) from Costa Rica. Proceedings of the
Minilimosina Roháček (Diptera: Sphaeroceridae). Proceedings Entomological Society of Washington 102: 609–612.
of the Entomological Society of Ontario 116: 1–60. Marshall, S.A. 2000b. A new genus of ant-like sphaerocerid from Mex-
Marshall, S.A. 1986. A revision of the Nearctic species of the ico (Diptera, Sphaeroceridae). Studia dipterologica 7: 213–223.
genus Pullimosina (Diptera, Sphaeroceridae). Canadian Marshall, S.A.  & Y. Cui. 2005. Systematics of Robustagramma,
Journal of Zoology 64: 522–536. a new genus of New World Sphaeroceridae (Diptera). Zootaxa
Marshall, S.A. 1987. Systematics of Bitheca, a new genus of New 1026: 3–122.
World Sphaeroceridae (Diptera). Systematic Entomology 12: Marshall, S.A. & R. Langstaff. 1998. Revision of the New World
355–380. species of Opacifrons Duda (Diptera, Sphaeroceridae, Limosini-
Marshall, S.A. 1995. Sclerocoelus and Druciatus, new genera nae). Contributions in Science 474: 1–27.
of New World Sphaeroceridae (Diptera; Sphaeroceridae; Marshall, S.A. & A.L. Norrbom. 1993. A revision of the New World
Limosininae). Insecta Mundi 9: 283–289. species of Norrbomia (Diptera: Sphaeroceridae), including all
SPHAEROCERIDAE (small dung flies) 95 1187

American species previously placed in Borborillus. Insecta Richards, O.W. 1931. Sphaeroceridae (Borboridae), pp. 62–84.
Mundi 6: 151–181. (1992) Diptera of Patagonia and South Chile, Part VI – Fascicle 2.
Marshall, S.A.  & J. Roháček. 2000. A world revision of the British Museum of Natural History, London.
seaweed fly genus Thoracochaeta Duda (Diptera: Sphaeroceridae: Richards, O.W. 1960. On two N. American species of Leptocera
Limosininae). Part 1: extra-Holarctic species. Studia diptero- Oliv., subgenus Coproica Rdi., with a review of the subgenus
logica 7: 259–311. (Dipt., Sphaeroceridae). Annals and Magazine of Natural
Marshall, S.A.  & J. Roháček. 2003. Podiomitra, a new genus of History, Series 13, 2: 199–208.
Homalomitrinae (Diptera: Sphaeroceridae) from Costa Rica. Richards, O.W. 1961. Diptera (Sphaeroceridae) from south Chile.
Proceedings of the Entomological Society of Washington 105: Proceedings of the Royal Entomological Society of London (B)
708–714. 30: 57–68.
Marshall, S.A. & S. Totton. 1995. Seven new species of Druciatus Roháček, J. 1983. A monograph and re-classification of the previous
Marshall (Diptera: Sphaeroceridae; Limosininae). Insecta Mundi genus Limosina Macquart (Diptera, Sphaeroceridae) of Europe.
9: 291–299. Part II. Beiträge zur Entomologie 33: 3–195.
Marshall, S.A., M. Buck, & O. Lonsdale. 2007. Lepidosina, a new Roháček, J. 1985. A monograph and re-classification of the previous
genus of New World Limosininae (Diptera: Sphaeroceridae). genus Limosina Macquart (Diptera, Sphaeroceridae) of Europe.
European Journal of Entomology 104: 573–599. Part IV. Beiträge zur Entomologie 35: 101–179.
Mourgués-Schurter, L.R. 1981. Sobre as espécies neotropicais de Roháček, J.  & S.A. Marshall. 1998. Revision of Homalomitrinae
Neosphaerocera Kim, 1972 (Diptera, Sphaeroceridae). Papéis subfam.  n. (Diptera: Sphaeroceridae), with the description of
Avulsos de Zoologia 34: 179–188. a new genus and three new species. European Journal of
Norrbom, A.L.  & K.C. Kim. 1985. Taxonomy and phylogenetic Entomology 95: 455–491.
relationships of Copromyza Fallén (s.s.) (Diptera: Sphaero- Roháček, S.A., S.A. Marshall, A.L. Norrbom, M. Buck, D.I.
ceridae). Annals of the Entomological Society of America 78: Quiros,  & I.P. Smith. 2001. World catalog of Sphaeroceridae
331–347. (Diptera). Slezské zemské muzeum, Opava, 414 pp.
Papp, L. 1978. Sphaeroceridae (Diptera) in the collection of Smith, I.P.  & S.A. Marshall. 2004. A revision of the New World
the Hungarian Natural History Museum. IV. Sphaeroceri- genus Pterogramma. Contributions in Science 499: 1–163.
nae. Acta Zoologica Academiae Scientiarum Hungaricae 24: Wheeler, T.A. 1995. Systematics of the New World Rachispoda
371–395. Lioy (Diptera: Sphaeroceridae): morphology, key to species
Papp, L. 1990. A review of the Afrotropical species of Poecilo- groups, and revisions of the atra, fuscipennis, limosa and vespertina
somella Duda (Diptera, Sphaeroceridae). Annales Histori- species groups. Journal of Natural History 29: 159–230.
co-Naturales Musei Nationalis Hungarici 81: 133–151. Wheeler, T.A.  & S.A. Marshall. 1995. Systematics of the New
(1989) World Rachispoda Lioy (Diptera: Sphaeroceridae): revisions of
Papp, L. 1991. A review of the Oriental species of Poecilosomella the primarily Neotropical aequipilosa, divergens, fuscinervis,
Duda (Diptera, Sphaeroceridae). Acta Zoologica Hungarica 37: maculinea, marginalis, and m-nigrum species groups. Journal
101–122. of Natural History 29: 1209–1307.
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
Curtonotidae (hunchbacked flies) 96
Stephen A. Marshall, Ashley H. Kirk-Spriggs, and John Klymko

Fig. 96.1. Female of Curtonotum sp., (Zumbado, 2006: 181).

Diagnosis fused with long and curved phallapodeme (Fig. 3). Immature

stages of Neotropical species unknown.
Small to medium-sized (body length 4–12 mm), distinc- Species of Curtonotum superficially resemble some
tively robust, hunchbacked flies (Fig. 1). Usually grayish to Drosophilidae, from which they differ most obviously in the
yellow, often with spots, stripes, or speckles on thorax and presence of a complete subcosta, a setose anepisternum, and
pigment patterns on abdomen. Arista plumose, with long a spinose costal margin. Curtonotids also resemble some
dorsal and ventral rays. Wing pigmentation varying from Heleomyzidae, which are often similar in size, color, and
costal chaetotaxy, but lack the arched scutum and plumose
hyaline to lightly fumose or boldly patterned; subcosta com-
arista that characterize curtonotids.
plete, cell cup present, cells dm and bm confluent, costa with
humeral and subcostal breaks. Curtonotum, the only Central
American genus, with several spinelike bristles between apices Biology
of R1 and R2+3. All tibiae with preapical dorsal bristles.
Female abdomen with tubular ovipositor; cerci, sometimes Afrotropical species of Curtonotum have been reared
fused, often with stout upcurved spinelike setae (Fig.  2). from decaying locust egg pods (Greathead, 1959; Cuthbertson,
Male abdomen with large tergite 5 followed by narrow tergite 7 1936; Kirk-Spriggs, 2008), and the Nearctic Curtonotum
(or tergites 6 and 7); sternites 6 and 7 asymmetrical, often helvum (Loew) has been reared from decaying grasshopper
poorly defined. Sternite 8 present and well developed only egg pods under laboratory conditions (Meier et al., 1997).
in Cyrtona Séguy sensu lato (an Old World genus). Surstyli Adults of African curtonotids have been observed sheltering
freely articulated or fused with epandrium. Male cerci in mammal burrows (Tsacas, 1977; Wirth & Tsacas, 1980)
simple, separate, unornamented (most species) or elongate and under overhangs of riverbanks (Pollock, 2002) and have
and fused basally (Nearctic, Palearctic, and several Afrotropical also been collected from decaying fungi and fresh elephant
Curtonotum species). Phallus extremely large, curved, and dung. Adult Neotropical curtonotids have been found on

1189
1190
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

tg 6
tg 7
tg 8 cerc

spmth
st 10

st 7 st 8
st 6

2 Curtonotum

ej apod
phapod

cerc

epand

hypd

pgt

sur

distph
bph

3 Curtonotum

Figs. 96.2–3. Abdomen and terminalia: lateral view of female abdomen (2) and male terminalia (3) of Curtonotum sp. Figures 2–3 illustrated
by J. Klymko.
Abbreviations: bph, basiphallus; cerc, cercus; distph, distiphallus; ej  apod, ejaculatory apodeme; epand, epandrium; hypd, hypandrium;
pgt, postgonite; phapod, phallapodeme; spmth, spermatheca; sur, surstylus; st, sternite; tg, tergite.
Curtonotidae (hunchbacked flies) 96 1191

dung baits at higher elevations and in association with tree especially apparent in the structure of the pseudocepha-
falls near sea level, but oviposition has not been observed lon and cephaloskeleton. The greater length of abdomi-
and larval habits of these species remain unknown. Pollock nal segments 6 and 7 may represent an autapomorphy of the
(2002) found some species of Cyrtona sensu lato from family.
Africa to be viviparous. The relationships among the three genera Curtonotum,
Unlike the Nearctic Curtonotum helvum, which is asso- Cyrtona sensu lato, and Axinota Wulp are discussed by
ciated with sand dunes (especially near large lakes), adults Meier et al. (1997) and Pollock (2002). Meier et al. (1997)
of Neotropical curtonotid species are found in a variety tentatively treat Curtonotum and Cyrtona sensu lato as a
of humid habitats. We have observed them associated with clade based on their similar spinulose spermathecae. Pol-
riparian habitat in dry forests in Costa Rica; freshwater and lock (2002), in contrast, tentatively treated Cyrtona sensu
marine shorelines in primary rainforests of Costa Rica, Peru, lato as the sister group to Curtonotum plus Axinota, which
and Ecuador; forest openings in primary rainforests in Peru; have similar phallic asymmetry and reduced pregenital
and high-elevation cloud forest in Ecuador and Bolivia. Male sclerites.
and female adults sometimes perch near the tips of twigs on
isolated floodplain trees and shrubs, but mating has not been
observed in Neotropical species. Tsacas (1977) notes that the
Identification
African species of Curtonotum are also found in a range of
Curtonotum is the only genus recognized in the New
both dry and humid habitat types, and at least one species
World. This virtually cosmopolitan genus includes one
appears to be restricted to high altitudes in Angola, while
Nearctic species and 19 described Neotropical species. The
an undescribed species of Cyrtona sensu lato is associated
remaining two known genera in the family (Axinota and
with high elevations in Namibia.
Cyrtona sensu lato) occur chiefly in the Afrotropical and
Meier et al. (1997) suggest that the Nearctic C.  helvum
Oriental Regions. The endemic Malagasy species of
is crepuscular or nocturnal; however, it is also common on
Curtonotum were revised by Tsacas (1974) and those of
open sand on overcast days. African curtonotid species are
mainland Africa by Tsacas (1977). The Afrotropical species
also usually crepuscular or nocturnal; however, no records
of Cyrtona sensu lato were partially dealt with by Hackman
suggest that Neotropical curtonotids are restricted to crepuscular
(1960) and the Oriental species of Curtonotidae by Delfinado
or nocturnal activity periods, or shelter in mammal burrows
(1969). The Afrotropical species are currently under revision
as do African species.
(A.H. Kirk-Spriggs, in preparation), and at least two new en-
demic Afrotropical genera and numerous new species within
Classification species complexes await description.
The Neotropical species are currently under revision
Curtonotids have at various times been placed in the (Klymko and Marshall, in preparation), and the majority of
Diastatidae, Ephydridae, Drosophilidae, Heleomyzidae, Neotropical species await description. The only key to Neo-
and Opomyzidae (see McAlpine 1987). Duda (1934) placed tropical species is that of Hendel (1913), and the most recent
them in their own family (Curtonotidae), and this has been key to any part of the Neotropical fauna is Curran’s (1934)
widely accepted in later works (Delfinado, 1969; Hennig, key to species occurring in Guyana. No Neotropical species
1973; Hackman, 1960; Kirk-Spriggs, 2008; Kirk-Spriggs & have been described since 1934.
Freidberg, 2007; Tsacas, 1974, 1977). Curtonotidae have
been considered to be the sister group of the remainder of
the superfamily Ephydroidea (McAlpine, 1989) or the sister
Synopsis of the fauna
group of Drosophilidae (Grimaldi, 1990).
Although the immature stages of Neotropical species are Curtonotum Macquart. All 19 Neotropical species of
unknown, Afrotropical Curtonotum species were recently this genus were described from South America, mostly
reviewed by Kirk-Spriggs (2008), who provided a detailed Bolivia, Brazil, and Peru (Wirth, 1975). Although there are
description of the third-instar larva of one species and a no published records of Curtonotidae from Central America,
larval generic diagnosis. Larval morphology exhibits many at least six species (including C. vulpinum Hendel) occur in
putative groundplan features of the Ephydroidea, this being Costa Rica.
1192
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Literature cited

Curran, C.H. 1934. The Diptera of Kartabo, Bartica District, to other known Ephydroidea larvae. African Entomology 15:
British Guiana, with descriptions of new species from other 226–243.
British Guiana localities. Bulletin of the American Museum of Kirk-Spriggs, A.H. & A. Freidberg, 2007. The Palaearctic species
Natural History 66: 287–532. of Curtonotidae (Diptera: Schizophora), with special reference
Cuthbertson, A. 1936. Biological notes on some Diptera of Southern to the fauna of Israel. Bulletin de l’Institut royal des sciences
Rhodesia. Occasional Papers of the Rhodesian Museum 5: naturelles de Belgique (Entomologie) 77: 133–146.
46–63. McAlpine, J.F. 1987. Curtonotidae, pp. 1007–1010. MND, Volume 2.
Delfinado, M.D. 1969. The Oriental species of Curtonotidae (Diptera). McAlpine, J.F. 1989. Phylogeny and classification of the Musco-
Oriental Insects 3: 199–219. morpha, pp. 1397–1518. MND, Volume 3.
Duda, O. 1934. Curtonotidae. Vol. 6/1, pp. 1–5. In Lindner, E. (editor). Meier, R., M. Kotrba, & K. Barber. 1997. A comparative study of
Die Fliegen der palaearktischen Region. E. Schweizerbart’sche the egg, first-instar larva, puparium, female reproductive system
Verlagsbuchhandlung, Stuttgart. and natural history of Curtonotum helvum (Curtonotidae; Ephy-
Greathead, D.J. 1959. Notes on the larvae and life history of droidea; Diptera). American Museum Novitates 3219: 1–20.
Curtonotum cuthbertsoni Duda (Dipt., Drosophilidae), a fly Pollock, J.N. 2002. Observations on the anatomy of Curtonotidae
associated with the desert locust, Schistocerca gregaria (Diptera: Schizophora). Journal of Natural History 36:
(Forskål). Entomologist’s Monthly Magazine 94: 36–37. 1725–1745.
Grimaldi, D. 1990. A phylogenetic, revised classification of genera Tsacas, L. 1974. Études préliminaire des Curtonotidae de Madagascar
in the Drosophilidae (Diptera). Bulletin of the American [Diptera]. Annals de la Société Entomologique de France (n.s.)
Museum of Natural History 197: 1–139. 10: 703–719.
Hackman, W. 1960. Chapter XVIII. Diptera (Brachycera): Camillidae, Tsacas, L. 1977. Les Curtonotidae (Diptera) de l’Afrique: 1. Le
Curtonotidae and Drosophilidae, pp. 381–389. In Hanström, B., genre Curtonotum Macquart. Annals of the Natal Museum 23:
P.  Brinck,  & G.  Rudebeck (editors). South African animal 145–171.
life. Results of the Lund University Expedition in 1950–1951. Wirth, W.W. 1975. 78. Family Curtonotidae. In Papavero,  N.
7. Almqvest  & Wilsell, Göteborg, Stockholm, Uppsala, [ix]  + (editor). A Catalogue of the Diptera of the Americas South of
10–488 pp. the United States. Departamento de Zoologia, Secretaria da
Hendel, F. 1913. Neue amerikanische Diptera. 1. Beitrag. Deutsche Agricultura, São Paulo, 5 pp.
Entomologische Zeitschrift 1913: 617–636. Wirth, W.W.  & L. Tsacas, 1980. 73. Family Curtonotidae,
Hennig, W. 1973. Ordnung Diptera (Zweiflügler), pp.  1–337. In pp.  671–672. In Crosskey,  R.W. (editor). Catalogue of the
Helmcke, J.-G., D. Starck, & H. Wermuth (editors). Handbuch Diptera of the Afrotropical Region. British Museum (Natural
der Zoologie. 4(2) 2/31 (Lfg. 20). Walter De Gruyter, Berlin & History), London, 1437 pp.
New York. Zumbado, M.A. 2006. Dípteros de Costa Rica y la América
Kirk-Spriggs, A.H. 2008. A contribution to the knowledge of the tropical — Diptera of Costa Rica and the New World tropics.
immature stages of Curtonotum (Diptera: Curtonotidae), from Instituto Nacional de Biodiversidad, Santo Domingo de Heredia,
Africa and the Middle East, with a discussion of relationships Costa Rica, 272 pp.
CAMILLIDAE (camillid flies) 97
Terry A. Wheeler

Fig. 97.1. Male of Camilla glabra (Fallén), (Holarctic, MND, fig. 97.1).

Diagnosis and subapical anteroventral ctenidial bristle, midtibia with

or without dorsal preapical bristle. Abdomen narrow, shining,
Small (body length 2.0–4.0 mm), shining black, brown, often metallic, syntergite 1+2 sometimes (Afrocamilla
or metallic flies. Frons usually with strong proclinate anterior Barraclough) with long median marginal bristles (Fig.  4);
fronto-orbital bristle (rarely reduced or absent), strong tergite  4 usually long. Male genitalia (Fig.  6) with narrow
reclinate posterior fronto-orbital bristle, short, reclinate epandrium, simple surstylus, elongate postgonite; long,
fronto-orbital bristle sometimes present between stronger membranous distiphallus. Female postabdomen telescopic,
bristles (Figs.  1, 2); ocellar bristles long, proclinate; sclerites of segments 6–8 reduced, cerci distinct, elongate,
postvertical bristles convergent; face flat or slightly concave, setulose; two spermathecae, sometimes unsclerotized;
without carina, ventral facial margin sometimes projecting ventral receptacle well developed.
(Fig. 2); antenna with dorsal seam in pedicel, first flagellomere The combination of plumose arista, size and orientation of
strongly decumbent; arista plumose in New World genera the fronto-orbital bristles, setose anepisternum, and absence
with dorsal rays longer; gena with strong vibrissa and row of bm-cu and cell cup distinguishes New World camillids
of several strong subvibrissal or genal bristles below this; from other acalyptrate families in the region. They are most
palpus well developed, proboscis short. Thoracic chaetotaxy: easily confused with Drosophilidae, but can be distinguished
by the above characters.
one postpronotal; two notopleurals; one presutural and one
postsutural supra-alar; one postalar; one to two dorsocen-
trals; two pairs of scutellar bristles; one katepisternal bristle; Biology
anepisternum setose and setulose (Fig. 1). Wing narrow, C
with humeral and subcostal breaks, Sc incomplete, crossvein Larvae of Camillidae are saprophagous in a range of
bm-cu absent, A1 and cell cup absent (Fig.  3). Forefemur substrates. In the Palearctic and Afrotropical Regions, adults
with several posterodorsal and/or posteroventral bristles are most frequently collected on dung or in the nests or latrines

1193
1194
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

3 Afrocamilla

2 Afrocamilla

5 Camilla

4 Afrocamilla

epiph

pgt

distph

6 Camilla

Figs. 97.2–6. Head, wing, basal abdominal segments, and male terminalia: lateral view of head of (2) Afrocamilla bispinosa Barraclough &
Wheeler, (Nearctic, Barraclough & Wheeler, 1995, fig. 2); dorsal view of wing (3) and lateral view of basal three abdominal tergites (4) of
A. arizonica Barraclough & Wheeler, (Nearctic, Barraclough & Wheeler, 1995, figs. 3, 4); lateral view of basal three abdominal tergites (5)
and lateral view of male terminalia (6) of Camilla arnaudi Barraclough & Wheeler, (Nearctic, Barraclough & Wheeler, 1995, figs. 5, 10).
Abbreviations: distph, distiphallus; epiph, epiphallus; pgt, postgonite.
CAMILLIDAE (camillid flies) 97 1195

of small mammals (Barraclough, 1992, 1993, 1997a, 1997b, Afrocamilla may also be present in the region, because three
1998). Nearctic species are also apparently associated with species of that genus are known from the southwestern
dung in nests or latrines of small mammals (Barraclough & United States (Barraclough & Wheeler, 1995; Barraclough &
Wheeler, 1995; Barraclough  & Fitzgerald, 2001). Adults Fitzgerald, 2001).
remain in close proximity to the larval food source and can
be observed hovering over and displaying and mating on the
surface of the dung (Barraclough, 1997b; Barraclough  & Identification
Fitzgerald, 2001).
World genera of Camillidae may be identified using
the keys in Barraclough (1992, 1993). The described New
Classification World species may be identified using keys and descrip-
tions in Barraclough & Wheeler (1995) and Barraclough &
The Camillidae are closely related to the Ephydridae and Fitzgerald (2001). Identification of the genera relies on
Drosophilidae. McAlpine (1987) considered Camillidae
external characters, especially the antenna, chaetotaxy, and
the sister group to Drosophilidae, whereas Grimaldi (1990)
placed them as the sister group to Ephydridae plus Risidae wing characters, so identification to that level does not usu-
(the latter is now treated as a synonym of Ephydridae). ally present a problem with specimens in good condition.
Of the four recognized extant genera, Camilla is the only Because of their small size and the importance of genitalic
one recorded to date in Mexico or Central America. characters in species identification, adult specimens should
Barraclough & Wheeler (1995) described Camilla arnaudi be preserved in ethanol and critical-point dried or chemically
Barraclough & Wheeler from Mexico (Baja California Sur). dried soon afterwards.

Key to the genera of Camillidae of the New World

1. Wing clear; abdominal syntergite 1+2 unmodified, with short median marginal bristles (Fig. 5) .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Camilla Haliday
– Wing with dark band crossing humeral region (Fig.  3); abdominal syntergite 1+2 convex at
posterior margin, projecting above base of tergite 3, with long median marginal bristles (Fig. 4);
southwestern USA. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Afrocamilla Barraclough]

Synopsis of the fauna Camilla Haliday. Like most camillids, members of

this genus are usually found in association with decaying
Although only one species has been recorded from organic material, especially dung in mammal burrows or
Central America, collecting in appropriate microhabitats in latrines. Most described species of Camilla are Palearc-
other regions, especially Africa, has produced higher than tic or Afrotropical; Camilla arnaudi from Mexico (Baja
expected species richness in Camillidae. The same may be California Sur) is the only species recorded from Central
true of arid habitats in southern Mexico and western Central America.
America, and a small number of undescribed species may
await discovery.

Literature cited

Barraclough, D.A. 1992. Afrocamilla stuckenbergi, a new African bats, birds and hyraxes in Africa and the Arabian Peninsula.
genus and species of Camillidae (Diptera), with comments on its African Entomology 6: 159–176.
behaviour and biology. Annals of the Natal Museum 33: 37–49. Barraclough, D.A. & S.J. Fitzgerald. 2001. A new Afrocamilla from
Barraclough, D.A. 1993. Teratocamilla, a new genus of Camillidae Colorado, the third species of Camillidae (Diptera: Schizophora)
(Diptera) and two new species of Camilla, from southern Africa. from the United States. Tijdschrift voor Entomologie 144: 1–4.
Annals of the Natal Museum 34: 19–30. Barraclough, D.A.  & T.A. Wheeler. 1995. Three new species of
Barraclough, D.A. 1997a. New east and west African species of Camillidae (Diptera) from the southwestern Nearctic Region,
Afrocamilla Barraclough (Diptera: Schizophora: Camillidae). the first species of the family described from the New World.
African Entomology 5: 181–193. Proceedings of the Entomological Society of Washington 97:
Barraclough, D.A. 1997b. The South African species of Afrocamilla 737–745.
Barraclough, 1992 (Diptera: Schizophora), a genus of Camillidae Grimaldi, D.A. 1990. A phylogenetic, revised classification of
associated with rock hyrax latrines. Annals of the Natal Museum genera in the Drosophilidae (Diptera). Bulletin of the American
38: 21–53. Museum of Natural History 197: 1–139.
Barraclough, D.A. 1998. Katacamilla Papp, 1978, a genus of McAlpine, J.F. 1987. Camillidae, pp.  1023–1025. MND,
Camillidae (Diptera: Schizophora) associated with the dung of Volume 2.
1196
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
DROSOPHILIDAE (small fruit flies, pomace flies,
vinegar flies) 98
David Grimaldi

Fig. 98.1. Female of Drosophila melanogaster Meigen, (cosmopolitan, MND, fig. 95.1).

Diagnosis Biology
Greatly varied in size and color, Neotropical species ranging Although the lifestyles of this family are extremely diverse,
from just slightly more than 1  mm in body length (some they are known best on the basis of several cosmopolitan
Cladochaeta, Diathoneura) to 6–7 mm (some Drosophila); species thriving among humans and that breed in decaying
from yellow to brown, black, mottled, and banded, or metallic fruits and vegetables, in wineries, and other places. The
colors. Eyes often red, but can be brownish or gray. Antennal most ubiquitous species is Drosophila melanogaster
arista usually plumose (Figs. 1, 2, 3, 9, 10), sometimes with Meigen, the most intensively studied eukaryotic organism
just ventral branches lacking, occasionally with all branch- and a native of central Africa. Other “tramp” species
es of arista lacking. Frons with three pairs of fronto-orbital include D.  simulans Sturtevant (African), D.  funebris
setae: the anterior-most pair proclinate, then two pairs of (Fabricius) (European), D.  hydei Sturtevant (American),
reclinate setae, referred to as anterior and posterior recli- D.  malerkotliana Parshad  & Paika, D.  busckii Coquil-
nates (Figs. 4–8). Anterior reclinates often small, sometimes lett, and D. immigrans Sturtevant (the last three native to
reduced or lost (Figs. 2, 3). southeast Asia). Adults of these species, and wild species
Drosophilidae are superficially similar to the smaller, breeding in fallen rain forest fruits, are attracted to the
yellowish Lauxaniidae, some of which have a plumose arista aromas of fermentation; their larvae graze on yeasts and
(but usually with all fronto-orbital setae being reclinate). bacteria in the rotting fruits. Many fruit-breeding species
The inside surface of the forefemur of Drosophilidae lacks breed well on laboratory media and are used in experimental
ctenidial spines, which many species in the closely related research. Despite the common name, most species do not
families of Ephydroidea, such as Camillidae and Curtonotidae, breed in fruits. Central American species represent most
possess. Ephydridae commonly have a plumose arista, but of the ecological diversity found in this family through-
almost always just with dorsal rays (and they further have a out the Neotropical Region. Major larval breeding sites
bulging face). include:

1197
1198
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

omtr omtr

fc car
fc car

2 Microdrosophila
3 Mycodrosophila
i vt s
p rc orb s
i vt s
poc s
p o vt s

p rc orb s
a rc orb s
p
a rc orb s oc s
pc orb s

p rc orb s pc orb s

5 Leucophenga
fc car
p rc orb s i vt s
p rc orb s
vb
o vt s
4 Drosophila p rc orb s
a

pc orb s

6 Chymomyza

i vt s
o vt s
p rc orb s poc s

p
a rc orb s

fc car pc orb s
vb

7 Neotanygastrella

8 Zygothrica

Figs. 98.2–8. Heads: anterolateral view, left antenna and mouthparts omitted, of (2) Microdrosophila sp.; (3) Mycodrosophila sp.; anterolateral
view, arista omitted, of (4) Drosophila sp.; anterolateral view of frons of (5) Leucophenga sp.; and (6) Chymomyza sp.; anterolateral view,
arista omitted, of (7) Neotanygastrella sp.; and (8) Zygothrica sp. Figures 2–8 illustrated by D. Grimaldi.
Abbreviations: a rc orb s, anterior reclinate orbital seta; fc car, facial carina; i vt s, inner vertical seta; oc s, ocellar seta; omtr, ommatrichia;
o vt s, outer vertical seta; pc orb s, proclinate orbital seta; poc s, postocellar seta; p rc orb s, posterior reclinate orbital seta; vb, vibrissa.
DROSOPHILIDAE (small fruit flies, pomace flies, vinegar flies) 98 1199

– living flowers: the Drosophila dreyfusi group, – sap fluxes on trees: many Drosophila, Chymomyza,
D. onychophora group, D. tripunctata group, Drosophila Neotanygastrella, Amiota, Stegana, and
(Phloridosa), Drosophila (Siphlodora), Laccodrosophila, Microdrosophila.
Zapriothrica, and many Zygothrica (reviewed by
– leaf mining: Scaptomyza.
Brncic, 1983).
– decaying pockets of cacti (Cactaceae): Drosophila
– fresh fungi: mostly species in the related genera
repleta species group.
Hirtodrosophila, Mycodrosophila, Paramycodrosophila,
and Zygothrica. Though probably all species of Zygo- – larval predators of sessile Sternorrhyncha (Coccoidea,
thrica swarm over forest fungi, many actually breed in Aleyrodoidea, Aphidoidea): Pseudiastata, Rhinoleu-
flowers (Grimaldi, 1987). cophenga, and Pseudocacoxenus (Ashburner, 1981).

arista

ar 1

10 Gitona
scp
ped

flgm 1

9 Rhinoleucophenga

11 Amiota (Sinophthalmus)

acr s

12 Palmophila

a dc s

a dc s

p dc s

p dc s

psc s
b sctl s
ap sctl s b sctl s

ap sctl s

13 Leucophenga 14 Scaptomyza
Figs. 98.9–14. Antennae and thoraces: lateral view of antenna of (9) Rhinoleucophenga sp.; and (10) Gitona sp.; antenna, showing pubescent
arista, of (11) Amiota (Sinophthalmus) sp.; and (12) Palmophila sp.; dorsal view of thorax of (13) Leucophenga sp.; and (14) Scaptomyza sp.
Figures 9–14 illustrated by D. Grimaldi.
Abbreviations: acr s, acrostichal setula; a dc s, anterior dorsocentral seta; ap sctl s, apical scutellar seta; ar 1, first aristomere; b sctl s, basal
scutellar seta; flgm 1, first flagellomere; p dc s, posterior dorsocentral seta; ped, pedicel; psc s, prescutellar seta; scp, scape.
1200
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

– parasites of spittle bug (Cercopidae) nymphs: Cladochaeta useful for distinguishing a few closely related genera. It
(though some may breed in flowers) (Grimaldi  & is always preferable to have a series of males and females
Nguyen, 1999). when identifying drosophilids. Most of the useful charac-
ters are on the head, though the thorax, legs, wings, and
– one species in Central America, Drosophila (Hir-
female genitalia (oviscape or ovipositor) are also important.
todrosophila) batracida Grimaldi, is a predator of the
Number and position of branches of plumose microtrichia
embryos of glass frogs (Centrolenidae) (Grimaldi,
on the antennal arista varies greatly and can be highly
1994).
reduced, even with branches lost (Hyalistata, Pseudiastata,
Laccodrosophila, Palmophila, Palmomyia, and some
Zapriothrica and Cladochaeta) (e.g., Figs.  11, 12).
Classification Setation of the fronto-orbital setae is significant. Extremely
important is the carina on the face, varying from completely
The family is closely related to the Curtonotidae, and absent to occurring in a wide range of sizes and shapes
these to other families of the Ephydroidea (Diastatidae, (Figs. 2–4, 7, 8). Significant setation on the thorax involves
Ephydridae). There are approximately 3300 world species the dorsocentrals, acrostichals, prescutellars, and katepis-
and 60 genera, classified into two subfamilies, the Steganinae ternal setae. Forelegs of males may have sexual structures,
and Drosophilinae. Steganinae comprises species possessing such as the “sex comb” of introduced species of Droso-
a larger proportion of primitive features. phila (Sophophora) malerkotliana, melanogaster, and
The traditional classification of genera is based on Sturtevant simulans. Wing venation is useful, such as the shape and
(1921, 1939, 1942) and Duda (1924). Sturtevant’s work was position of some veins, particularly small ones at the base
far ahead of its time; his 1939 paper was one of the earliest of the wing (closed or open bm cell; cup cell absent or
uses of a matrix of derived characters for reconstructing present; A1 present, reduced, or absent) (Figs.  15–23).
relationships, far preceding the work of Willi Hennig. The wing tip is pointed in some genera, such as many
Sturtevant established a system of subgenera in Drosophila Stegana and most Microdrosophila (Figs.  15, 19), and
that is largely in use today. Patterson  & Stone (1952, and the costal lappet occurs in several genera that breed in
elsewhere) were the first to consistently use species groups fungi (Mycodrosophila (Fig.  21), Paramycodrosophila).
categories within the subgenera Drosophila and Sophophora The oviscape is extremely important: whether it is ab-
Sturtevant. Though not all species in these subgenera have sent or present, generally as a pair of laterally flattened
been classified into species groups, this informal category plates having a row of small pegs around the margin. All ste-
is in widespread use today among Drosophila biologists. ganines and some basal drosophilines (e.g., Cladochaeta,
Throckmorton (1962, 1975) extensively studied phylogeny Microdrosophila) lack the oviscape, or the oviscape prim-
of Drosophilidae (especially Drosophila), hypothesizing itively lacks pegs and has just fine setulae (Chymomyza,
widespread paraphyly in Drosophila. The study by Grimaldi some Neotanygastrella, Palmomyia, Palmophila). Some
(1990) was the first major cladistic study of genera; it pruned genera have a large tooth at the tip of the oviscape. Body
the genus Drosophila of several major subgenera (Scap- coloration contributes useful additional features, but is
todrosophila, Hirtodrosophila, Lordiphosa Basden), making rarely diagnostic in itself.
The following key will be useful for identifying genera and
the nominal genus monophyletic. The generic status of these
some subgenera, roughly from central Mexico and southern
former subgenera has been corroborated by several DNA se-
Florida south through Central America and the Caribbean,
quencing studies, though some taxonomists continue the use of
and South America north of Argentina and Chile. A useful
the traditional Drosophila subgenera (e.g., Vilela & Bächli, reference for the identification of species is the study of
1990; but see Bächli, 2001). The phylogeny and classifica- old types of Neotropical Drosophilidae by Vilela & Bächli
tion of the subfamily Drosophilinae will be on firm ground (1990). It is carefully done and heavily illustrated, reducing the
in the near future, but Steganinae remain neglected. Most need to resort to historical literature and old types for some spe-
taxonomic work is at the species level within particular spe- cies, though it does not replace the need for species-level revi-
cies groups of Drosophila. sions. Much of it is based on the work of Duda (1925, 1927),
whose papers on the Costa Rican fauna were the first significant
Identification studies of Neotropical drosophilids. The website by Gerhard
Bächli (2001), hosted by the University of Zürich (http://tax-
Drosophilidae can generally be identified to genus odros.unizh.ch), is an extremely useful and carefully prepared
using external features of the males or females, though database for the names and references of world species and
having both sexes (or the female in particular) is very updates the world catalogue by Wheeler (1981, 1986).
DROSOPHILIDAE (small fruit flies, pomace flies, vinegar flies) 98 1201

costal spinules
spinules

15 Stegana 16 Leucophenga

h Sc R1
R2+3

R4+5

A1 M1

17 Rhinoleucophenga 18 Amiota (Sinophthalmus)

19 Microdrosophila 20 Chymomyza

costal lappet

21 Mycodrosophila 22 Diathoneura

23 Drosophila

Figs. 98.15–23. Wings (photomicrographs): dorsal view of (15) Stegana sp.; (16) Leucophenga sp.; (17) Rhinoleucophenga sp.; (18) Amiota

(Sinophthalmus) sp.; (19) Microdrosophila sp.; (20) Chymomyza sp.; (21) Mycodrosophila sp.; (22) Diathoneura sp.; and (23) Drosophila sp.
Figures 15–23 illustrated by D. Grimaldi.
1202
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Key to the genera of Drosophilidae of the Neotropical Region

1. Pair of large prescutellar setae present (Fig. 13); two large katepisternal setae present; STEGANINAE.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– No prescutellar setae present; one large and one small katepisternal seta present; DROSOPHILINAE
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

2. Lower surface of wing tip with several (sometimes just one) minute, thornlike structures on costal
vein (Figs. 15, 16, costal spinules). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
– Without costal spinules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

3. Body lightly colored, no pleural bands; dorsum of abdomen with paired spots, or silvery; wings
not flexed down along sides; tips of veins R4+5 and M1 only slightly converging. . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leucophenga Mik
– Body generally dark brown, usually with pleural bands; abdomen usually homogeneously dark
brown; wings flexed down along sides, tips of veins R4+5 and M1 strongly converge (Fig. 15) . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stegana Meigen

4. Branches of plumose arista long, or short and numerous (e.g., Fig. 9). . . . . . . . . . . . . . . . . . . . . 5

– Arista pubsecent, without plumose branches (e.g., Fig. 11) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

5. Black or yellow bodied, with white postpronotal (humeral) lobes, without numerous interfrontal
setulae; aristal branches long, three to six in number dorsally . . . . . . . . . . Amiota (Amiota) Loew
– Yellow or gray bodied, postpronotal lobes not white; with numerous, fine interfrontal setulae;
aristal branches often short, but more than six dorsally. . . . . . . . . . . . . Rhinoleucophenga Hendel

6. Body gray or brown, with mottled darker patterns and generally with spot at base of each seta
and setula; face with prominent carina; ocellar setae typically large; males swarm around heads of
humans. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amiota (Sinophthalmus) Coquillett
– Body not mottled, without spots; face flat; ocellar setae small. . . . . . . . . . . . . . . . . . . . . . . . . . . 7

7. Scutellum, abdomen, and face shiny black . . . . . . . . . . . . . . . . . . . . . . . . Pseudocacoxenus Duda

– Scutellum, abdomen, and face usually yellowish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

8. Wings hyaline. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hyalistata Wheeler

– Wings with patterning . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

9. Frons and face narrow, width barely equal to distance between lateral ocelli; frons without numerous
interfrontal setulae; Caribbean. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Mayagueza Wheeler]
– Front wider, frons with numerous interfrontal setulae. . . . . . . . . . . . . . . . Pseudiastata Coquillett

10. Arista pubescent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

– Arista plumose, even if with just one dorsal branch at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

11. Dark flies, with narrow facial carina . . . . . . . . . . . . . . . . . . . . . . . . . Zapriothrica Wheeler, in part
– Light colored flies, with broad, flat carina; Ecuador. . . . . . . . . . . . [Palmophila Grimaldi, in part]

12. With three pairs of dorsocentral setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

– With two pairs of dorsocentral setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
DROSOPHILIDAE (small fruit flies, pomace flies, vinegar flies) 98 1203

13. With four rows of acrostichal setulae; Colombia. . . . . . . . . . . . . . . [Palmophila Grimaldi, in part]
– Lacking acrostichal setulae; Colombia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Apacrochaeta Duda]

14. Plumose arista with one ventral branch or none, exclusive of apical fork. . . . . . . . . . . . . . . . . . 15
– Plumose arista with two or more ventral branches, exclusive of apical fork. . . . . . . . . . . . . . . . 20

15. With costal lappet on wing (Fig. 21). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

– Without costal lappet. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

16. Mottled flies with spots at bases of setae. . . . . . . . . . . . . . . . . . . . . . . . Paramycodrosophila Duda

– Scutum entirely black, sometimes brown . . . . . . . . . . . . . . . . . . . . . . Mycodrosophila Oldenberg

17. Proboscis jutting, facial carina noselike and bulbous (Fig. 8). . . . . . . . . . Zygothrica Wiedemann
– Proboscis not protruding, facial carina small or face entirely flat. . . . . . . . . . . . . . . . . . . . . . . . 18

18. No oviscape; surstyli (male) without teeth. . . . . . . . . . . . . . . . . . . . . . . . . Cladochaeta Coquillett

– Oviscape present; surstyli with teeth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

19. Ocellar triangle large, shiny; with two dorsal aristal branches . . . . . . . . . Paraliodrosophila Duda
– Ocellar triangle small or average; arista with three or more dorsal branches. . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hirtodrosophila Duda

20. Anterior reclinate orbital setae far anterior (and lateral) to proclinates (Figs. 6, 7); foreleg dark
with lightly colored tarsus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
– Anterior reclinates posterior (as in Fig. 5), lateral, or slightly anterolateral (Fig. 4) to proclinates;
foreleg usually unicolorous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

21. Facial carina small, bulbous; male lacking femoral spines. . . . . . . . . . . . . Neotanygastrella Duda
– Face without carina; male fore- or midfemur with spines. . . . . . . . . . . . . . . . Chymomyza Czerny

22. Face flat, without carina; eyes bare; vein A absent . . . . . . . . . . . . . . . . . . . . . . Diathoneura Duda
– Face with small or large carina, eyes bare to pilose; vein A small to large. . . . . . . . . . . . . . . . . 23

23. Wing at apex of R4+5 slightly acute (Fig. 19); crossveins close. . . . . . . . Microdrosophila Malloch
– Wing tip rounded (Fig. 23); crossveins distant . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

24. With three long, nearly equal katepisternal setae. . . . . . . . . . . . . . . . . . . . Scaptodrosophila Duda
– With two katepisternal setae of unequal lengths. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

25. Acrostichals in two to four rows (Fig. 14). . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scaptomyza Hardy

– Acrostichals in six or more rows. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26

26. Facial carina large, often with flat surface; eyes pubescent (Fig. 4) . . . . . . . . . . Drosophila Fallén
– Facial carina thin; eyes bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27

27. With thick black spines on foretarsi. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Laccodrosophila Duda

– Without such spines. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Zapriothrica Wheeler, in part
1204
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Synopsis of the fauna Cladochaeta Coquillett. This is the largest lineage of

drosophilids endemic to the Western Hemisphere, and it
Approximately 300 described species are known in our is closely related to another genus restricted to this area,
region (slightly more than the whole Nearctic fauna), but Diathoneura. Prior to a recent monograph on the genus
several times this number actually exist. Many undescribed (Grimaldi  & Nguyen, 1999), there were 13 described
and undocumented (but described) species are predicted. species, and now there are 118 (most of them Neotropical),
Intensive, systematic sampling routinely yields more than though an estimated 500 species are thought to actually
100 species at any one location. At Las Alturas Biological exist. Most species are externally uniform, light yellow, and
Station, Costa Rica, nearly 130 species were found (and difficult to discern based on external features; male genitalia
probably more occur there). Forests at La Selva Biological are complex and clearly demarcate species. Costa Rica has
Station, Costa Rica, have an estimated 170 species, and in the best known fauna for this genus. Life histories were re-
cloud forests of adjacent areas like Braulio Carrillo National viewed (Grimaldi & Nguyen, 1999), and of the few that are
Park there is a completely different fauna (D.  Grimaldi, known most are parasites as larvae on the nymphs of spittle-
unpublished observations). Additionally, recent and compre- bugs (Auchenorrhyncha: Cercopidae). Presumably they feed
hensive revisions have revealed unexpected diversity (e.g., on the hemolymph of their host. One species, C. psychotria
Grimaldi, 1987; Grimaldi  & Nguyen, 1999). For example, Grimaldi & Nguyen, has been bred from flowers of Psychotria
just four species of Cladochaeta were known for Costa Rica (Rubiaceae) at Monteverde, Costa Rica.
prior to 1999; 24 additional ones have been described in a
subsequent monograph (Grimaldi & Nguyen, 1999). For the Diathoneura Duda. This is also a species-rich genus of
Neotropical Region, 13 species of Cladochaeta were known, New World drosophilids and more restricted to the tropics
with 105 new ones discovered, and many more thought to than is Cladochaeta. Like Cladochaeta they have complex
still occur, principally in forest canopies. Though undescribed male genitalia, but Diathoneura species are also colourful,
diversity of Cladochaeta may be exceptional, it is estimated often with patterned wings and bodies, so species are easier
that at least 500 species of Drosophilidae occur in Central to separate using just external features. Pipkin et al. (1966)
America. Costa Rica is by far the best studied country in bred various species from flowers in Panama (recorded under
Central America for drosophilids. Additionally, new genera are the name Clastopteromyia Malloch). Though not as diverse
still being discovered in the Neotropical Region (Grimaldi et as Cladochaeta, there are many more than the approximately
al., 2003). The family is diverse throughout the world’s trop- 25 described species in Central America, and probably well
ics, but by far the greatest diversity is Neotropical. Despite over 100 species throughout the Neotropical Region. There
all the attention paid to the phylogeny of the Drosophilinae, is no modern revision of this genus, but the careful and
field exploration and species-level revisions of various detailed redescription of types by Vilela & Bächli (1990) is
genera are needed most. the most important reference.

Amiota Loew. This is a cosmopolitan genus of over 100 Drosophila Fallén. Drosophila is a large, diverse, cos-
species, with fewer than 10 described Central American species. mopolitan genus best known for the few species used in lab-
Males have the peculiar habit of flying around the head of oratory research. Most species in the genus are grouped into
humans, perhaps attracted to eyes. Amiota (Amiota), which subgenera, and the subgenus Drosophila further into species
is worldwide, are entirely yellowish or black with white groups. Five subgenera occur in the New World. Dorsi-
postpronotal lobes, plumose arista, and a flat face; Amiota lopha Sturtevant is represented in this region by only one
(Sinophthalmus), which is North and Central American, are cosmopolitan tramp species, D.  (Dorsilopha) busckii. It is
mottled or spotted, the postpronotal lobes are not white, and a small, active, spotted fly that breeds in a great range of
they have a pubescent arista and a prominent carina. Male decaying materials and is a native of southeast Asia. Droso-
genitalia in the group are complex and homologies of phila is the largest subgenus, and though limits and mono-
structures obscure. Females are collected by sweeping along phyly are still not strictly defined, most have a broad, flat facial
tree trunks, and their life histories are virtually entirely carina. Species groups of the subgenus Drosophila were
unknown. There are many undescribed species. covered most thoroughly by Patterson & Stone (1952), but
unfortunately, diagnostically important features of wings,
Chymomyza Czerny. Like the closely related genus genitalia, and setation were not discussed, so it is of lim-
Neotanygastrella, this is a cosmopolitan genus, most diverse ited taxonomic use. An excellent review of the genus was
in tropical forests. Most of them have dark forelegs with done by Ashburner (1990). Phloridosa Sturtevant is a small
light tarsi, which they stroke outward while waving their Neotropical subgenus (introduced to Hawaii) with three de-
wings. They are primarily associated with trees, particularly scribed species in Costa Rica; they are small, dark flies that
ones that are wounded or decaying. The taxonomy of one breed in flowers. Siphlodora Patterson & Mainland likewise
Chymomyza group was covered by Grimaldi (1986); fewer is a small subgenus, restricted to the New World, which was
than 10 species are described for Central America, although recently revised by Vilela  & Bächli (2000). Sophophora is
there are approximately twice this number actually known. the second largest subgenus of Drosophila, the great propor-
DROSOPHILIDAE (small fruit flies, pomace flies, vinegar flies) 98 1205

tion of species occurring in central Africa and southeast Asia. Mycodrosophila Oldenberg. For biology, see Hirtodros-
Within the subgenus Sophophora, various species in the mel- ophila, above. The most important reference for Neotropical
anogaster group (including D. melanogaster) are introduced Mycodrosophila is by Wheeler & Takada (1963). There are
to the New World; all 10 species of the D. willistoni group five described species of this genus in Central America.
and 10 species of the D. saltans group are native to the New
World tropics. Most species of the D. obscura group are in Neotanygastrella Duda. Revised most recently by
the Holarctic Region; Neotropical species typically occur in Frota-Pessoa  & Wheeler (1951), these flies have similar
cooler, montane forests. distributions and lifestyles as the related Chymomyza (see
above). Five species of Neotanygastrella are described for
Hirtodrosophila Duda. With its close relatives My- Central America, though there are approximately twice this
codrosophila, Paramycodrosophila, and Zygothrica, these number actually present.
flies have the distinctive habit of swarming about forest fungi.
As many as 25 species in these genera can co-occur on one Paraliodrosophila Duda. This is a small genus of black,
bloom of fungi. They court, mate, and adults feed on the shiny flies comprising just four species, all Mesoamerican.
fungi, and many species oviposit there too. Mycodrosophila
prefer more dense, woody shelf fungi; species in the other Paramycodrosophila Duda. For biology, see Hirtodroso-
genera are most common on fleshy bracket fungi, usually phila, above. Three described species of this genus are known
white. Many of the species in all four genera have a dark for Central America.
scutum and tergites, contrasting with light pleura, but many
Zygothrica are colorful, having stripes on the thorax and Pseudocacoxenus Duda. Only one rare species of this
patterned wings. All four genera are circumtropical; only genus is described, P. lineatifrons Duda, from Costa Rica.
Hirtodrosophila and Mycodrosophila venture into cool
temperate regions. There are 13 described species of Pseudiastata Coquillett. This is a small Neotropical
Hirtodrosophila in Central America, although many further genus composed of six rare species, larvae of which are
new species exist. The most recent paper of importance on predators on mealybugs and other coccoids.
this genus is by Vilela & Bächli (2004).
Rhinoleucophenga Hendel. A New World genus with six
Hyalistata Wheeler. This is a small genus of five rare species described from Central America, Rhinoleucophena
Neotropical species (Grimaldi, 1993), formerly placed as includes bivisualis (Patterson), formerly placed in the Palearc-
a subgenus of Pseudiastata. Like closely related genera, tic genus Gitona Meigen. Like Pseudiastata, larvae are preda-
larvae are probably predacious on sessile Sternorrhyncha. tors of sessile Sternorrhyncha.

Laccodrosophila Duda. Like the closely related Scaptodrosophila Duda. This is a large genus of approximately
Zapriothrica (which may be paraphyletic with respect to 250 described species, which appear to all be native to the
Laccodrosophila), this is a small Neotropical genus whose Old World, where they breed in flowers and decaying plant
species breed in flowers. Undescribed species occur throughout material. Scaptodrosophila brooksae Pipkin, from North
montane forests of Central America and the South American America, is probably misclassified. Scaptodrosophila
Andes (D.  Grimaldi, unpublished observations). Eight of latifasciaeformis Duda is originally from Africa, but has
the 10 described species in both genera occur in the páramo been introduced and is widespread throughout southern parts
of the Andes of Colombia, Ecuador, and Venezuela; only of North America to South America.
two named species (L.  flavescens Wheeler and Z.  nudiseta
Wheeler) are from Central America, both recorded from Scaptomyza Hardy. This is also a large genus, of approx-
southern Mexico. The main reference is Duda (1927). imately 300 described species, and is cosmopolitan. Species
are most diverse in temperate areas of the Holarctic and
Leucophenga Mik. This is a large, cosmopolitan genus
Austral Regions. In tropical regions they are most commonly
of over 160 species, the greatest proportion occurring in the
encountered in cool montane habitats, particularly open ones
Old World tropics. There are 13 species in Central America,
like fields and paramo, where they mine stems and leaves.
described by various authors but mostly from Mexico and
Costa Rica. They are commonly believed to breed in fungi, There are nearly 20 described species from Central America,
but are rarely encountered there, so their life histories are but the genus as a whole is in serious need of careful revision.
obscure. The most important reference for the Western Hemisphere
species is Wheeler and Takada (1966).
Microdrosophila Malloch. All the Neotropical species
(approximately 10; D. Grimaldi, unpublished) of this genus Stegana Meigen. This is a cosmopolitan genus of ap-
are undescribed; virtually all described species are Paleotropical. proximately 70 species, with 23 described species known
They are most commonly encountered by sweeping over for Central America, 19 of those based on species described
decaying tree trunks or the decaying sawdust of cut forest only from Costa Rica (mostly in Malloch, 1924). They are
trees, where they are probably breeding. collected by sweep netting quickly along tree trunks and
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

beneath large branches, particularly of trees in incipient stages have bizarre, anvil-shaped heads and eyes; they are ag-
of decay. Larvae probably feed on fungi and subcortical gressive and defend territories on mushrooms. Revi-
mycelia, but life histories are essentially unknown. There are sions of Neotropical Zygothrica are by Burla (1956) and
many undescribed Neotropical species. Grimaldi (1987), though more work is needed. Neorhi-
noleucophenga Duda (four species described from Costa
Zapriothrica Wheeler. For biology and general discussion, Rica) appears to be a synonym of Zygothrica. There
see Laccodrosophila, above. Wheeler (1959) is the main ref- are nearly 40 Zygothrica described for Central Ameri-
erence for this genus. ca, though many more exist (D.  Grimaldi, unpublished
observations).
Zygothrica Wiedemann. For biology, see Hirtodrosophila,
above. Males of many species in the Zygothrica dispar group

Literature cited

Ashburner, M. 1981. Entomophagous and other bizarre Drosophili- Malloch, J.R. 1924. Descriptions of Neotropical two-winged flies
dae, pp. 395–429. In Ashburner, M., H.L. Carson, & J.N. Thomp- of the family Drosophilidae. Proceedings of the United States
son (editors). The Genetics and Biology of Drosophila, vol. 3a. National Museum 66: 1–11.
Academic Press, New York, London, xvi + 429 + Ii-Ilxx pp. Patterson, J.T. & W.S. Stone. 1952. Evolution in the genus Droso-
Ashburner, M. 1990. Drosophila: A laboratory handbook. Cold phila. MacMillan & Co., New York, 610 pp.
Spring Harbor Press, New York, 434 pp. Pipkin, S.B., R.L. Rodríguez,  & J. León. 1966. Plant host speci-
Bächli, G. 2001. Taxodros. A taxonomic database of the Drosophil- ficity among flower-feeding Neotropical Drosophila (Diptera:
idae (online). Available from http://taxodros.unizh.ch. Univer- Drosophilidae). American Naturalist 100: 135–156.
sity of Zürich. Sturtevant, A.H. 1921. The North American species of Drosophila.
Brncic, D. 1983. Ecology of flower-breeding Drosophila, pp. 333– Publication 301. Carnegie Institution of Washington, Washing-
382. In Ashburner, M., H.L. Carson, & J.N. Thompson (editors). ton, D.C., 150 pp.
The Genetics and Biology of Drosophila, vol.  3d. Academic Sturtevant, A.H. 1939. On the subdivision of the genus Drosophila.
Press, New York, London, 474 pp. Proceedings of the National Academy of Sciences 25: 137–141.
Burla, H. 1956. Die Drosophiliden-Gattung Zygothrica und ihre Sturtevant, A.H. 1942. The classification of the genus Drosophila,
Beziehung zur Drosophila-Untergattung Hirtodrosophila. with descriptions of nine new species. University of Texas Pub-
Mitteilungen aus dem Zoologischen Museum in Berlin 32: lications 4213: 5–51.
190–321. Throckmorton, L.H. 1962. The problem of phylogeny in the genus
Duda, O. 1924. Beitrag zur Systematik der Drosophiliden unter Drosophila. University of Texas Publications 6205: 297–343.
besonderer Berücksightigung der Paläarktischen u. Oriental- Throckmorton, L.H. 1975. The phylogeny, ecology, and geography
ischen Arten (Dipteren). Archiv für Naturgeschicthe 90(A) 3: of Drosophila, pp. 421–469. In King, R.C. (editor). Handbook
172–259. of Genetics, vol. 3. Plenum Press, New York, 874 pp.
Duda, O. 1925. Die costaricanischen Drosophiliden des Ungarisch- Vilela, C.R.  & G. Bächli. 1990. Taxonomic studies on Neotropi-
en National-Museums zu Budapest. Annales Historico-Natu- cal species of seven genera of Drosophilidae (Diptera). Mittei-
rales Musei Nationalis Hungarici 22: 149–229. lungen der Schwizerischen Entomologischen Gesellschaft 63,
Duda, O. 1927. Die südamerikanischen Drosophiliden (Dipteren) supplement: 1–332.
unter Berücksichtigung auch der anderen neotropischen sowie Vilela, C.R. & G. Bächli. 2000. Morphological and ecological notes
der nearktischen Arten. Archiv für Naturgeschichte 91: 1–228. on the two species of Drosophila belonging to the subgenus
[1925] Siphlodora Patterson and Mainland, 1944 (Diptera: Drosophili-
Frota-Pessoa, O. & M.R. Wheeler. 1951. A revision of the genus dae). Mitteilungen der Schwizerischen Entomologischen Gesell-
“Neotanygastrella” Duda (Diptera, Drosophilidae). Revista schaft 73: 23–47.
Brasileira de Biologia 11: 145–151. Vilela, C.R. & G. Bächli. 2004. On the identities of nine Neotro-
Grimaldi, D.A. 1986. The Chymomyza aldrichii species-group pical species of Hirtodrosophila (Diptera, Drosophilidae). Mit-
(Diptera: Drosophilidae): Relationships, new Neotropical spe- teilungen der Schwizerischen Entomologischen Gesellschaft 77:
cies, and the evolution of some sexual traits. Journal of the New 161–195.
York Entomological Society 94: 342–371. Wheeler, M.R. 1959. Three new species of Zapriothrica Wheel-
Grimaldi, D.A. 1987. Phylogenetics and taxonomy of Zygothrica er (Diptera, Drosophilidae). The Southwestern Naturalist 4:
(Diptera: Drosophilidae). Bulletin of the American Museum of 83–87.
Natural History 186: 103–268. Wheeler, M.R. 1981. 1. The Drosophilidae: A taxonomic overview,
Grimaldi, D.A. 1990. A phylogenetic, revised classification of gen-
era in the Drosophilidae (Diptera). Bulletin of the American Mu- pp.  1–97. In Ashburner,  M., H.L.  Carson,  & J.N.  Thompson
seum of Natural History 197: 1–139. (editors). The Genetics and Biology of Drosophila, vol. 3a. Aca-
Grimaldi, D.A. 1993. Amber fossil Drosophilidae (Diptera), Part demic Press, New York, London, xvi + 429 + Ii-Ilxx pp.
II. Review of the genus Hyalistata, new status (Steganinae). Wheeler, M.R. 1986. Additions to the catalog of the world’s Dros-
American Museum Novitates 3084: 1–15. ophilidae, pp.  395–409. In Ashburner,  M., H.L.  Carson,  &
Grimaldi, D.A. 1994. Description and immature stages of Hir- J.N. Thompson (editors). The Genetics and Biology of Droso-
todrosophila batracida sp.n. (Diptera: Drosophilidae), a preda- phila, vol.  3e. Academic Press, New York, London, xviii  +
tor of frog embryos. Entomologica scandinavica 25: 129–136. 548 pp.
Grimaldi, D.  & T. Nguyen. 1999. Monograph on the spittlebug Wheeler, M.R. & H. Takada. 1963. A revision of the American spe-
flies, genus Cladochaeta (Diptera: Drosophilidae: Cladochae-
tini). Bulletin of the American Museum of Natural History 241: cies of Mycodrosophila (Diptera; Drosophilidae). Annals of the
1–326. Entomological Society of America 56: 392–399.
Grimaldi, D., F. Ervik, & R. Bernal. 2003. Two new Neotropical Wheeler, M.R. & H. Takada. 1966. The Nearctic and Neotropical
genera of Drosophilidae (Diptera) visiting palm flowers. Jour- species of Scaptomyza Hardy (Diptera: Drosophilidae). Univer-
nal of the Kansas Entomological Society 76: 109–124. sity of Texas Publications 6615: 37–78.
DIASTATIDAE (diastatid flies) 99
Wayne N. Mathis

Fig. 99.1. Male of Diastata vagans Loew, (Nearctic, MND, fig. 96.1).

Diagnosis vibrissal setae well developed; two well-developed fronto-

orbital setae present, proclinate seta laterad of reclinate
Adults small, grayish-brown, drosophilid-like flies (body seta present (Fig. 2). Antenna decumbent; arista elongate,
length 2.5–4.0  mm); usually with patterned wings and macropubescent or short plumose. Anepisternum setose
weakly spinose costa (Fig. 1). Postocellar setae convergent (Diastata) or bare (Campichoeta); katepisternum bearing
(Fig. 2); both inner and outer verticals present; ocellar and 12 setae, posterior seta stronger. C extending to M; C with

1207
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

rc orb s i vt s
poc s

o vt s
pc orb s

vb sbvb s

2 Diastata 3 Campichoeta
Sc R1
R2+3

R4+5

M1

CuA1

4 Diastata

tg 5

phapod
st 5

hypd
ej apod
ph gd tg 6
pgt

ph
pregt syntgst
sur 7-8
epand

cerc

5 Diastata 6 Diastata

Figs. 99.2–6. Heads, wing, and male terminalia: anterodorsal view of head of (2) Diastata vagans Loew, (Nearctic, MND, fig. 96.2); lateral
view of head of (3) Campichoeta latigena McAlpine, (Nearctic, McAlpine, 1962, fig. 4); dorsal view of wing of (4) D. vagans, (MND,
fig. 96.3); ventral view (5) and lateral view (6) of male terminalia of D. vagans, (MND, figs. 96.8–9).
Abbreviations: cerc, cercus; ej apod, ejaculatory apodeme; epand, epandrium; hypd, hypandrium; i vt s, inner vertical seta; o vt s, outer
vertical seta; pc orb s, proclinate orbital seta; pgt, postgonite; ph, phallus; ph gd, phallic guide; phapod, phallapodeme; poc s, postocellar
seta; pregt, pregonite; rc orb s, reclinate orbital seta; sbvb s, subvibrissal seta; st, sternite; sur, surstylus; syntgst, syntergosternite; tg, tergite;
vb, vibrissa.
DIASTATIDAE (diastatid flies) 99 1209

either just subcostal break (Campichoeta) or both humeral classifications have been advocated to account for these
and subcostal breaks (Diastata; Fig. 4); Sc incomplete, not differences (Chandler, 1998a, 1998b). The classifications
reaching C, often fusing with R1 distally; R1 bare; both CuA2 differ primarily in whether Diastata alone is recognized as
and A1 present; cell cup closed, crossvein bm-cu present. a family (Chandler, 1987, 1998b) or as an included lineage
Legs moderately elongate, slender. Abdominal segments 1–5 within the Ephydridae (Griffiths, 1972), and thus with Cam-
unmodified; male with segments 6–8 reduced (Figs. 5–6), pichoeta, Euthychaeta Loew, and Pareuthychaeta Hennig
although tergite 7 well developed, sternite 7 lacking; abdominal as a separate family, the Campichoetidae (Griffiths, 1972;
spiracles (6 or 7) in membrane. Chandler, 1987, 1998a). Another alternative is to recog-
Diastatidae are similar to Drosophilidae, especially the nize all four genera in a single family (Barraclough, 1992;
genus Diathoneura Duda, and Opomyzidae but are distin- McAlpine, 1987, 1989). The classification adopted here is
guished from either of these two families by having two large deliberately conservative, with Campichoeta, Euthychaeta,
fronto-orbital setae (one in Opomyzidae), by the dorsolateral Diastata, and Pareuthychaeta as a single family, awaiting
position of the large proclinate fronto-orbital seta relative to more definitive resolution. Grimaldi’s (1990) cladogram of
the reclinate seta, and by the weakly spinose costal margin. the Ephydroidea suggests that the Diastatidae are the sister
In addition, Diastata has a setulose anepisternum and two group to the clade Camillidae + Ephydridae (I also include
costal breaks. the Risidae in the Ephydridae).
Even nomenclatural issues are problematic although stable.
The type species of Diastata, which is the type genus for
Biology the family-group name, is based on a misidentification
(Sabrosky, 1999).
The immature stages are unknown with the possible ex-
ception of a third-instar larva of Campichoeta (Hennig, 1952,
as Thryptocheta tristis Fallén  = Campichoeta basalis (Mei- Identification
gen)). Adults of Diastata occur on low deciduous shrubs
along the margins of marshes, bogs, damp woodlands, and Chandler’s papers (1987, 1998a, 1998b) are essential
other lush habitats. Campichoeta likewise frequents lush, for understanding the family and identifying the included
low vegetation, especially where such habitats occur in genera. Most keys to the family are difficult, as they do not
otherwise dry to arid environments, such as found in the recognize that the costa of Diastata is broken twice (h and
American southwest. Sc) whereas that of Campichoeta is broken just once (Sc).
Moreover, McAlpine’s key (1987) to genera is faulty in that
Classification the anepisternum is either setulose or bare, not the katepis-
ternum. The New World species of Diastata need revision.
The phylogenetic relationships among taxa here in- Adults are best preserved dry, and given their relatively small
cluded in the Diastatidae remain unresolved, and varying size, they should be double mounted.

Key to the genera of Diastatidae of the New World

1. Arista plumose with longer hairs dorsally (Figs. 1–2); large reclinate fronto-orbital seta inserted
close to anterior margin of fronto-orbital plate, lacking shorter reclinate fronto-orbital seta anterior
to it (Fig. 2); anepisternum setulose; costa with both humeral and subcostal breaks (Fig. 4). . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diastata Meigen
– Arista with short pubescence only (Fig. 3); large reclinate fronto-orbital seta inserted well posterior
from anterior margin of fronto-orbital plate, usually bearing shorter reclinate fronto-orbital seta
anterior to it (Fig. 3); anepisternum bare; costa with subcostal break only; Holarctic, Afrotropical
Regions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Campichoeta Macquart]
1210
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Synopsis of the fauna Diastata Meigen. Worldwide there are 35 described

species in Diastata, and others, especially from the New
The Diastatidae now include 47 valid species that are World, remain to be described. In the Neotropical Region,
placed in four genera (Chandler, 1987; Mathis  & Bar- Prado (1984) reported three species of which one, D. costa-
ricensis Hennig, is from Costa Rica. The other two species
raclough, in press). The New World fauna comprises two
are from Argentina and Bolivia (D. argentina (Malloch)) and
genera and 11 species (McAlpine, 1987; Prado, 1984). One Peru (D. dimidiata (Hendel)). Identification of the New World
genus, Pareuthychaeta Hennig with two species, is known species of Diastata is difficult, as there is no recent revision,
from Baltic amber (Hennig, 1965). Examination of collec- and several species remain undescribed. A combination
tions has revealed only a few species of Diastata in Central of Chandler (1987), Melander (1913), and recent catalogs
America. (McAlpine, 1965; Prado, 1984) should be consulted.

Literature Cited

Barraclough, D.A. 1992. A synopsis of the Afrotropical Diastatidae Entwicklung dieser Dipteren-Gruppe. Stuttgarter Beiträge zur
(Diptera), with the description of five new species from southern Naturkunde 145: 1–215.
and east Africa and first record of the Campichoetinae. Annals Mathis, W.N.  & D.A. Barraclough. in press. A catalog and
of the Natal Museum 33: 13–36. conspectus on the family Diastatidae (Diptera: Schizophora).
Chandler, P.J. 1987. The families Diastatidae and Campichoetidae Myia.
(Diptera, Drosophiloidea) with a revision of Palaearctic and McAlpine, J.F. 1962. A revision of the genus Campichoeta
Nepalese species of Diastata Meigen. Entomologica scandinavica Macquart (Diptera, Diastatidae). The Canadian Entomologist
18: 1–50. 94: 1–10.
Chandler, P.J. 1998a. 3.46. Campichoetidae, pp.  515–522. In McAlpine, J.F. 1965. Diastatidae, pp.  772–773. In Stone,  A.,
Papp, L. & B. Darvas (editors). Contributions to a Manual of C.W.  Sabrosky, W.W.  Wirth, R.H.  Foote,  & J.R.  Coulson
Palaearctic Diptera. Volume  3, Higher Brachycera. Science (editors). A catalog of the Diptera of America north of Mexico.
Herald Publishers, Budapest, 880 pp. United States Department of Agriculture, Agriculture Handbook
Chandler, P.J. 1998b. 3.47. Diastatidae, pp. 523–530. In Papp, L. & 276, Washington, D.C., iv + 1696 pp.
B. Darvas (editors). Contributions to a Manual of Palaearctic McAlpine, J.F. 1987. 96. Diastatidae, pp.  1019–1022. MND,
Diptera. Volume 3, Higher Brachycera. Science Herald Publishers, Volume 2.
Budapest, 880 pp. McAlpine, J.F. 1989. 116. Phylogeny and classification of the
Griffiths, G.C.D. 1972. The phylogenetic classification of Diptera Muscomorpha, pp. 1397–1518. MND, Volume 3.
Cyclorrhapha, with special reference to the structure of the male Melander, A.L. 1913. A synopsis of the dipterous groups
postabdomen. Series entomologica 8. Dr. W. Junk, The Hague, Agromyzinae, Milichiinae, Ochthiphilinae and Geomyzinae.
340 pp. Journal of the New York Entomological Society 21: 219–273,
Grimaldi, D.A. 1990. A phylogenetic, revised classification of genera 283–300.
in the Drosophilidae (Diptera). Bulletin of the American Museum Prado, A.P. do. 1984. 80. Family Diastatidae. In Papavero,  N.
of Natural History 197: 1–139. (editor). A Catalogue of the Diptera of the Americas South of
Hennig, W. 1952. Die Larvenformen der Dipteren. 3. Teil. Akademie- the United States. Departamento de Zoologia, Secretaria da
Verlag, Berlin, 628 pp. Agricultura, São Paulo, 1 p.
Hennig, W. 1965. Die Acalyptratae des Baltischen Bernsteins und Sabrosky, C.W. 1999. Family-group names in Diptera. An annotated
ihre Bedeutung für die Erforschung der phylogenetischen catalog. Myia 10: 1–360.
EPHYDRIDAE (shore flies) 100
Wayne N. Mathis

Fig. 100.1. Female of Ephydra (Ephydra) riparia (Fallén), (Nearctic, MND, fig. 981).

Diagnosis Eggs (Fig.  60) white to grayish-white, 0.30–1.30  mm

long, 0.14–0.49  mm wide, usually ellipsoidal ovoid, often
Adults minute to medium-sized flies (body length with flattened surface, although contours generally symmet-
0.6–11.0 mm), usually dull and dark colored (Fig. 1), often ric except near ends. Ends bluntly rounded to sub-bulbous;
invested with whitish, grayish, or brownish microtomentum; micropylar end sometimes drawn out, forming elongate
but unusually diverse in body structure, vestiture, and orna- process or horn of varying length (some species of Ochthera)
mentation; female usually larger than male. Head, including and usually with small, narrowly campanulate nipple. Chorion
subcranial cavity, large; one to three, rarely up to five microreticulate to deeply rugose, sometimes with longitudinal
lateroclinate or reclinate/proclinate fronto-orbital setae; pseudo- ridges; ridges sometimes forked, as in Hydrellia and Discocerina
postocellar setae divergent or lacking. Arista pectinate on obscurella (Fallén).
dorsum only or bare to pubescent; face flat to conspicuously First-instar larvae metapneustic, as small as 0.30 mm in
protruding. Scutum usually bearing numerous acrostichal length (Scatophila); second and third-instar larvae amphip-
setulae, one to five dorsocentral setae, two to three supra-alar neustic; third-instar larvae (Figs.  61–62) reaching 18  mm
setae, one postalar seta; anepisternum setulose. Wing usually (Ochthera); some earlier instars more spinulose, but most
well developed, hyaline but sometimes maculate; C with instars generally similar except for size.
humeral and subcostal breaks; Sc incomplete; vein R1 merging Puparia roughly similar to third-instar larvae, but more com-
with C before middle of wing; cells  bm and dm confluent, pact, darker in color (light yellow to grayish-brown), hardened,
crossvein bm-cu absent; cell cup absent. Preapical dorsal seta integumental spinules more prominent than on larvae and
lacking on fore- and hind tibiae. Males commonly with five anterior respiratory processes fully extended and with some
exposed tergites; female with six to seven; abdominal spiracles distinctive features, such as dorsal cephalic cap; in species
1–5 located in tergites, 6th in pleural membrane near genital with more maggotlike larvae, puparium normally shorter and
capsule. Male terminalia (Fig. 58) symmetrical. Female usually broader than larva, but in species with elongate larvae such as
with two spermathecae, rudimentary and probably nonfunc- those of Ephydra, with relatively little change of form; larval
tional; ventral receptacle (Fig.  59) heavily sclerotized, respiratory tube and spiracles extended fully at pupariation; in
functioning in place of spermathecae. Immature stages mostly Ephydra anal proleg particularly large and curved forward at
aquatic or associated with aquatic habitats. pupariation to grip the substrate (usually an aquatic plant).

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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Although similar to some species of the families typical microphagous habit. These habits have been reported in
Drosophilidae, Diastatidae, and Chloropidae, shore flies are part or totally for adults of Hydrellia (Deonier, 1971, 1998),
distinguished by the non-plumose arista (pectinate on dorsal Notiphila (Mathis, 1979a), and Hecamede Haliday (Ardö,
surface only), costal wing breaks (a humeral and subcostal 1957).
break), convergent cells bm and dm, only having five abdominal Although the mating behavior of ephydrids has been
tergites in males, and an internal female reproductive system widely studied, generalization is difficult. Dahl (1959)
that includes two rudimentary spermathecae and a well- distinguished six phases while studying the mating behavior
sclerotized ventral receptacle. of Scandinavian shore flies. Deonier (1971) found it more
flexible simply to recognize courtship and mating behavior for
describing the mating habits of various Hydrellia species.
Biology The basic work on shore fly ecology is Dahl’s (1959)
study of the Scandinavian fauna. Other papers since then
Aquatic and semiaquatic habitats are typical for this include Deonier (1965), Collins (1975), Simpson (1976b),
family, although several genera have evolved structural and and Foote (1995). The papers by Scheiring & Foote (1973)
behavioral modifications that adapt them to remarkably and Simpson (1976b) provide excellent summaries for
divergent habitats. One of these evolutionary specializations several species. Oldroyd (1964) aptly summarized shore fly
is the adaptation to extremes in saline concentrations. biology when he wrote “Evidently we are seeing in the
Although many shore flies frequent the muddy margins of Ephydridae a family of flies in the full flower of its evolution,
freshwater environments, their abundance and/or biotic and as such they offer attractive material for study, not only
importance do not compare with those occurring in maritime to the dipterist, but also to students of insect physiology and
marshes, tidal salt pools (Simpson, 1976a), or the saline and behavior.”
alkaline lakes of arid regions. The relative isolation and
reduced interspecific competition in such habitats have
resulted in the evolution of highly distinctive and specialized Classification
genera and species.
Eggs are deposited singly or in groups over the substrate The characterization of Ephydridae here excludes Nan-
(surface of algae, rocks vegetation, surface of plants). Ovi- nodastiidae and Diastatidae but includes the genus Risa
position by a female commonly continues for 7–10  days, Becker (formerly the type genus of a separate family, Risidae).
and documented observations in laboratory conditions of The family is divided into five subfamilies: Discomyzinae,
time from oviposition to emerging adults takes 13–44 days, Hydrelliinae, Gymnomyzinae, Ilytheinae, and Ephydrinae
development of egg 1–4  days, each larva 3–17  days, and (Zatwarnicki, 1992). The classification differs considerably
puparium 2–9 days. from those used in various regional catalogs (including the
Most shore fly larvae feed by filtering microorganisms Neotropical Region by Wirth, 1968), being supported
from the surrounding semiliquid medium. The food sources predominantly by characters of the male terminalia. Previous
vary considerably, depending on the feeding habit. Most systems with four subfamilies were developed by Cresson
larvae are microphagous on bacteria, unicellular algae, and between 1942 and 1949. Mathis (1991) and Mathis  &
yeasts, although some prefer dead and decaying animal tis- Zatwarnicki (1995) published brief histories on the development
sue or excrement (Foote, 1995). An exception is Ochthera, of shore fly classifications.
whose cephalopharyngeal skeleton, including the pointed
mouthhooks, is robust and well suited for piercing exo-
skeletons of prey. Hennig (1943, 1952) and Ferrar (1987)
Identification
summarized data then available, and the former attempted
There is no key to genera on a worldwide basis, but
a partial key.
regional keys exist for the Nearctic (Wirth et al., 1987) and
Feeding habits of adults likewise vary considerably,
Palearctic (Mathis & Zatwarnicki, 1998b) Regions that can
but most feed on microscopic algae (Foote, 1977, 1981a, be used to identify most genera found in the tropics and
1981b), dinoflagellates, and other unicellular forms. A major south temperate zones. References to existent keys to species
digression from the microphagous feeding habit occurs in are found in Mathis & Zatwarnicki’s (1995) world catalog.
Ochthera (Simpson, 1975). Prey of Ochthera varies from There are a few faunal treatments that may be useful for
bloodworms captured by excavating the substrate to adults identifying Central American taxa: Argentina (Lizaralde de
of most small insects that can be overpowered. Species of Grosso, 1989), Belize (Mathis, 1997b), and southern Chile
many other genera are polyphagous, sometimes even and Patagonia (Cresson, 1931). Although unfinished,
cannibalistic, scavenging on dead or dying insects, and Cresson (1946a, 1947) began a series of synopses on the
others feed on nectar and leaf epidermis, besides having the Neotropical shore flies, which included keys to species.
EPHYDRIDAE (shore flies) 100 1213

Key to the genera of Ephydridae of Central America

1. Fronto-orbital setae reclinate and proclinate or absent (Figs. 2–24, 26–31); median facial area and
lower facial margin without setae; facial setae inserted in more or less vertical series, parallel with
parafacial setae; subcranial cavity small to large. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Fronto-orbital setae lateroclinate and usually conspicuous (Figs. 32–42); median facial area and
lower facial margin setose, the latter often with long setae (if not, costa short, extended to R4+5);
insertions of facial series of setae convergent above; subcranial cavity large and cavernous;
EPHYDRINAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58

2. Presutural or sutural dorsocentral seta present, conspicuous (Figs. 46, 51) . . . . . . . . . . . . . . . . . 3

– Presutural or sutural dorsocentral seta inconspicuous or absent (Fig. 50). . . . . . . . . . . . . . . . . . 16

3. Posterior notopleural seta inserted conspicuously above level of anterior seta; ILYTHEINAE, in
part. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
– Posterior notopleural seta at same level as anterior seta; HYDRELLINAE. . . . . . . . . . . . . . . . . 8

4. Fronto-orbital setae greatly reduced, at most with tiny proclinate setulae; PHILYGRINI, in part.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
– Fronto-orbital setae well developed; ILYTHEINI. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

5. Outer vertical seta lacking; fronto-orbital setae lacking . . . . . . . . . . . . . . . . . . . . Garifuna Mathis

– Outer vertical seta usually present; fronto-orbital seta present but often greatly reduced,
proclinate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

6. Arista appearing bare, length of dorsal hairs less than basal aristal width; dorcentral setae 2,
anterior seta presutural or sutural (1+1). . . . . . . . . . . . . . . . . . . . . . . . . . . . Philygria Stenhammar
– Arista bearing conspicuous dorsal hairs, length of hairs much longer than bgasal aristal width . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nostima Coquillett

7. Vein R2+3 long, subparallel to C; costal sector 2 over twice length of sector 3; head as in Fig. 29.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ilythea Haliday
– Vein R2+3 short, running almost straight to C; costal sector 2 subequal in length to sector  3
(Fig. 53). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Zeros Cresson

8. Eye bearing short, dense setulae; ocellar setae seldom as strong as pseudopostocellar setae,
usually much weaker; postsutural supra-alar seta usually short, not longer than posterior
notopleural seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
– Eye bare; ocellar setae stronger than weak pseudopostocellar setae; postsutural supra-alar setae
longer than posterior notopleural seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

9. Ocellar setae well developed (Fig. 13); anterior and posterior notopleural setae inserted at about
same level. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrellia Robineau-Desvoidy
– Ocellar setae greatly reduced or absent (Fig.  16); posterior notopleural seta inserted distinctly
above level of anterior seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lemnaphila Cresson

10. Midtibia lacking dorsal, spinelike setae; shiny black to dark green species. . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Typopsilopa Cresson
– Midtibia with one to four erect, spinelike setae along dorsal surface; usually dull, microtomentose
species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
1214
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

oc s
infr s
lc orb s
pc orb s

pgn

2 Clasiopella 3 Paratissa 4 Rhysophora

oc s

5 Cressonomyia 6 Psilopa 7 Psilopa

psdpoc s

flgm 1 ped

8 Ceropsilopa 9 Leptopsilopa 10 Notiphila (Dichaeta)

Figs. 100.2–10. Heads: anterodorsal view of (2) Clasiopella uncinata Hendel, (Nearctic, MND, fig. 98.14); (3) Paratissa semilutea (Loew),
(Nearctic, MND, fig. 98.6); (4) Rhysophora robusta Cresson, (Nearctic, MND, fig. 98.7); (5) Cressonomyia aciculata (Loew), (Nearctic,
MND, fig. 98.11); (6) Psilopa flavida Coquillett, (Nearctic, MND, fig. 98.15); (7) Psilopa sp., (Nearctic, MND, fig. 98.16); (8) Ceropsilopa
coquilletti Cresson, (Nearctic, MND, fig. 98.13); (9) Leptopsilopa varipes (Coquillett), (Nearctic, MND, fig. 98.12); and (10) Notiphila
(Dichaeta) caudata (Fallén), (Holarctic, MND, fig. 98.30).
Abbreviations: flgm 1, first flagellomere; infr s, interfrontal seta; lc orb s, lateroclinate orbital seta; oc s, ocellar seta; pc orb s, proclinate
orbital seta; ped, pedicel; pgn, postgena; psdpoc s, pseudopostocellar seta.
EPHYDRIDAE (shore flies) 100 1215

11 Notiphila (Agrolimna) 12 Atissa 13 Hydrellia

14 Paralimna (Phaiosterna) 15 Ptilomyia 16 Lemnaphila

i vt s

o vt s

pgn

17 Gastrops 18 Athyroglossa 19 Mosillus

Figs.  100.11–19. Heads (continued): anterodorsal view of (11)  Notiphila (Agrolimna) scalaris Loew, (Nearctic, MND, fig.  98.31, as
N. (Dichaeta) scalaris); (12) Atissa atlantica Cresson, (Nearctic, MND, fig. 98.22, as A. pygmaea); (13) Hydrellia griseola (Fallén),
(Palaearctic, MND, fig.  98.29); (14)  Paralimna (Phaiosterna) decipiens Loew, (Nearctic, MND, fig.  98.32); (15)  Ptilomyia mabelae
(Cresson), (Nearctic, MND, fig. 98.20); (16) Lemnaphila scotlandae Cresson, (Nearctic, MND, fig. 35); (17) Gastrops nebulosus Coquillett,
(Nearctic, MND, fig. 98.3); (18) Athyroglossa glaphyropus Loew, (Nearctic, MND, fig. 98.4); and (19) Mosillus tibialis Cresson, (Nearctic,
MND, fig. 98.27).
Abbreviations: i vt s, inner vertical seta; o vt s, outer vertical seta; pgn, postgena.
1216
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

11. Costa extended to vein R4+5 (Fig. 54); with three dorsocentral setae (1+2). . . . . . . . . . . . . . . . . 12
– Costa extended to vein M (as in Fig. 55); with four dorsocentral setae (1+3) or one (0+1). . . . 13

12. Midtibia bearing four dorsal erect setae along dorsum; two small, proclinate fronto-orbital setae
inserted laterad of large, reclinate fronto-orbital seta (Figs. 10–11); forecoxa with two distinctly
larger setae on lower half of anterolateral margin; midfemur of male lacking posteroventral comb
of setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Notiphila (Agrolimna) Meigen
– Midtibia bearing three dorsal erect setae along dorsum; zero to one small, proclinate fronto-orbital
setae inserted laterad of large, reclinate fronto-orbital seta; forecoxa lacking distinctly larger setae
on lower half of anterolateral margin; midfemur of male bearing comblike row of setae along
posteroventral margin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Notiphila (Notiphila) Fallén

13. Katepisternal seta absent; with three to five aristal hairs. . . . . . . . . . . . . . . . . . . Oedenops Becker
– Katepisternal seta present; arista bearing eight or more dorsal hairs; Paralimna Loew. . . . . . . 14

14. Gena high, height about one-third or more or eye height; first flagellomere shorter, length at most
1.5 times width. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paralimna (Paralimna) Loew, in part
– Gena short, height about one-quarter eye height (Fig. 14); first flagellomere elongate, length more
than twice width. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

15. Scutum (brown medially and through dorsocentral track, between yellowish- to brownish-gray
margins) and anepisternum (brown dorsally and ventrally, gray between) conspicuously vittate;
crossvein dm-cu and sometimes adjacent veins with halo of brown. . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paralimna (Paralimna) Loew, in part
– Scutum and anepisternum mostly unicolorous and grayish-brown to black; crossvein and veins
and rest of wing essentially unicolorous . . . . . . . . . . . . . . . . . . Paralimna (Phaiosterna) Cresson

16. Arista pectinate, with branches sometimes pale and difficult to discern. . . . . . . . . . . . . . . . . . . 17
– Arista bare to macropubescent; if pectinate, hairs short, about equal to aristal width at base. . . 31

17. Prescutellar acrostichal setae large (subequal to posterior dorsocentral seta), inserted widely apart
(distance between them subequal to that between either prescutellar and the posterior dorsocentral
seta on the same side) and usually in front of intra-alar seta; reclinate fronto-orbital seta usually
inserted behind larger, proclinate fronto-orbital seta; DISCOMYZINAE. . . . . . . . . . . . . . . . . . 18
– Prescutellar acrostichal setae small (about one-half to two-thirds length of posterior dorsocentral
seta), inserted close together (distance between about one-half that between either prescutellar
and posterior dorsocentral seta on the same side) and behind or aligned with intra-alar seta;
reclinate fronto-orbital seta usually inserted slightly anterior to larger, proclinate fronto-orbital
seta; GYMNOMYZINAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

18. Vein R2+3 well separated from costal vein; crossvein dm-cu nearly straight or shallowly arched,
not angulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
– Vein R2+3 close to costal vein beyond end of vein R l; crossvein dm-cu with sharp angle;
DISCOMYZINI, in part. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

19. Face strongly and coarsely sculptured on at least lower one-half; facial setae short, the longest
at most three-quarters as long as its distance from opposite seta; base of R4+5 bearing two to four
setulae on dorsum; DISCOMYZINI, in part. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
– Face usually smooth, if finely striate the longest facial seta at least as long as its distance from
opposite seta; base of R4+5 lacking setulae on dorsum; PSILOPINI . . . . . . . . . . . . . . . . . . . . . . 26
EPHYDRIDAE (shore flies) 100 1217

20. Postsutural supra-alar seta much reduced (no larger than surrounding setulae) or absent . . . . . 21
– Postsutural supra-alar seta present, size subequal to presutural seta. . . . . . . . . . . . . . . . . . . . . . 24

21. Pseudopostocellar setae well developed, length about one-half that of ocellar setae, orientation
divergent at usually less than 90° (Fig. 2). . . . . . . . . . . . . . . . . . . . . . . . . . . . . Clasiopella Hendel
– Pseudopostocellar setae weakly developed, length considerably less than one-half that of ocellar
setae, orientation various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

22. Face conspicuously and deeply transversely rugose; only reclinate fronto-orbital seta well
developed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Discomyza Meigen
– Face at most with shallowly impressed transverse striae; at least one proclinate fronto-orbital seta
in addition to reclinate seta well developed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

23. Eye appearing bare; with one well-developed proclinate fronto-orbital seta (second seta greatly
reduced), inserted anterior to reclinate seta; presutural supra-alar seta weakly developed, length
less than anterior notopleural seta (except in M. cressoni Lizarralde de Grosso); legs bicolored. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mimapsilopa Cresson
– Eye conspicuously setulose; with two well-developed proclinate fronto-orbital setae, anterior
proclinate seta at about same level as large, reclinate seta, posterior proclinate seta inserted posterior
of reclinate seta; presutural supra-alar seta well developed, length longer than anterior notopleural
seta; legs unicolorous, blackish-brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Helaeomyia Cresson

24. Interfrontal seta present; with four fronto-orbital setae (anterior two setae proclinate, third
lateroclinate, fourth lateroreclinate; Fig. 3). . . . . . . . . . . . . . . . . . . . . . . . . . . Paratissa Coquillett
– Interfrontal seta absent; with two to three fronto-orbital setae . . . . . . . . . . . . . . . . . . . . . . . . . . 25

25. Supra-alar seta well developed, length subequal to postalar seta; wing usually mostly hyaline, at
most with anterior margin faintly infumate; head as in Fig. 4. . . . . . . . . . . . . Rhysophora Cresson
– Supra-alar seta moderately well developed, length about half postalar seta; wing usually with at
least anterior margin infumate, wing sometimes mostly brown and with white spots . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Guttipsilopa Wirth

26. Base of wing blackish, contrasting with remainder of wing; head as in Fig. 5. . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cressonomyia Arnaud
– Base of wing concolorous with rest of wing, usually hyaline or lightly tinged; head as in
Figs. 6–9. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27

27. Scutellum greatly enlarged, extended far over and above abdomen. . . . . . . Peltopsilopa Cresson
– Scutellum normally developed, not extended far over abdomen. . . . . . . . . . . . . . . . . . . . . . . . . 28

28. Antenna with pedicel conical, broader apically, without dorsoapical lobe, with dorsoapical spine
weak and at most one-third as long as first flagellomere; first flagellomere from two to four times
as long as wide (Fig. 8) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceropsilopa Cresson
– Antenna with pedicel short and subtriangular, with dorsoapical lobe, with dorsoapical spine strong
and at least half as long as first flagellomere; first flagellomere at most two times as long as wide.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

29. Vein R2+3 close to costa beyond end of vein R1; crossvein dm-cu with sharp angle at middle . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Clanoneurum Becker
– Vein R2+3 well separated from costa; crossvein dm-cu mostly straight or shallowly arched, not
angulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
1218
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

20 Hecamede 21 Allotrichoma (Pseudohecamede) 22 Diphuia

23 Glenanthe 24 Lipochaeta 25 Discocerina

fc s

gn pafc s

26 Polytrichophora 27 Hydrochasma 28 Discocerina

Figs. 100.20–28. Heads (continued) and notopleuron: anterodorsal view of head of (20) Hecamede albicans (Meigen), (Holarctic, MND,
fig. 98.19); (21) Allotrichoma (Pseudohecamede) abdominale (Williston), (Nearctic, MND, fig. 98.23, as Pseudohecamede abdominalis);
(22)  Diphuia nitida Sturtevant  & Wheeler, (Nearctic, MND, fig.  98.24); (23)  Glenanthe interior Chillcott, (Nearctic, MND, fig.  98.25);
(24) Lipochaeta slossonae Coquillett, (Nearctic, MND, fig. 98.26); lateral view of notopleuron of (25) Discocerina obscurella (Fallén),
(Holarctic, MND, fig.  98.45); anterodorsal view of head of (26)  Polytrichophora agens Cresson, (Nearctic, MND, fig.  98.8);
(27) Hydrochasma leucoproctum (Loew), (Nearctic, MND, fig. 98.9); and (28) Dis. obscurella, Nearctic, MND, fig. 98.10).
Abbreviations: fc s, facial seta; gn, gena; pafc s, parafacial seta.
EPHYDRIDAE (shore flies) 100 1219

30. Face smooth, lacking transverse striae (Figs. 6–7) . . . . . . . . . . . . . . . . . . . . . . . . . . Psilopa Fallén

– Face with transverse striae (Fig. 9) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leptopsilopa Cresson

31. Posterior notopleural seta inserted near ventral margin and at same level as anterior seta;
HYADININI. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
– Posterior notopleural seta inserted at much higher level than anterior seta; LIPOCHAETINI, in
part. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53

32. Tergite 4 subequal in length to tergite 5, neither conspicuously punctate; some species with both
vertical setae (Fig. 31); tergite 5 of male with more posterior orientation. . . . . . Hyadina Haliday
– Tergite 4 long, 1.3–2.0 times as long as tergite 5, both conspicuously punctata; outer vertical seta
absent; tergite 5 of male with more ventral orientation. . . . . . . . . . . . . . . . . . . . Lytogaster Becker

33. Arista bare to macropubescent, if pectinate, rays shorter than one-half height of first
flagellomere. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
– Arista pectinate dorsally. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37

34. Posterior margin of gena broadly rounded onto occiput; body densely invested with microtomentum,
generally dull colored; usually with two notopleural setae, posterior seta inserted at level above
anterior seta; LIPOCHAETINI, in part. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
– Posterior margin of gena sharply angulate; body extensively shiny black, although with some gray
to whitish microtomentum on dorsum; with single notopleural seta, inserted near posterior angle;
GYMNOMYZINI, in part. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54

35. Eye pyriform, distinctly narrowed ventrally; gena short, less than one-fourth eye height (Fig. 23);
katepisternal seta present along posterior margin, moderately well developed. . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Glenanthe Haliday
– Eye generally oval or round, not distinctly narrowed ventrally; gena high, one-third or more of
eye height; katepisternal seta greatly reduced or lacking. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36

36. Antenna reduced, inserted in well-separated cavity, arista atrophied, budlike (Fig. 24); face short,
height subequal to length of reduced antenna; clypeus bandlike; oral opening gaping; body setae
and setulae pale; costa distinctly thickened at merger of vein R1 . . . . . . . . . Lipochaeta Coquillett
– Antenna normally developed, not in deep cavity, arista as long as first flagellomere; face well
developed, height much more than length of antenna; clypeus various; oral opening narrow or
gaping; setae and setulae largely black; costa only slightly thickened at merger of vein R1. . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paraglenanthe Wirth

37. Foreleg raptorial, with forefemur greatly enlarged and foretibia with large apical spur (Fig. 49);
OCHTHERINI. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ochthera Latreille
– Foreleg normal, forefemur slender, foretibia without large apical spur. . . . . . . . . . . . . . . . . . . . 38

38. Abdominal tergites 3 and 4 enlarged, wider than second tergite, rugose or pitted; face trigonal;
GASTROPSINI. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
– Abdominal tergites 2–4 subequal in width, microtomentose, but more or less smooth; face
normal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40

39. Face with gently rounded, broad swelling on dorsal half (Fig. 17); apex of vein R2+3 merging with
C at less than 45° angle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gastrops Williston
– Face flat or concave; apex of vein R2+3 merging with C at nearly a right angle. . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Beckeriella Williston
1220
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

i vt s
o vt s

29 Ilythea 30 Philygria 31 Hyadina

oc s psdpoc s

i vt s

o vt s
lc orb s

32 Parydra 33 Brachydeutera 34 Dagus

35 Dagus 36 Diedrops 37 Diedrops

Figs. 100.29–37. Heads (continued): anterodorsal view of (29) Ilythea spilota (Curtis), (Holarctic, MND. fig. 98.28); (30) Philygria debilis
Loew, (Nearctic, MND, fig. 98.36); (31) Hyadina binotata (Cresson), (Nearctic, MND, fig. 98.37); (32) Parydra aquila (Fallén), (Holarctic,
MND, fig. 98.33); (33) Brachydeutera argentata (Walker), (Nearctic, MND, fig. 98.2); lateral view (34) and anterior view (35) of Dagus
rostratus (Cresson), (Mathis, 1982, figs. 56, 57); lateral view (36) and anterior view (37) of Diedrops hitchcocki Mathis & Wirth, (Peru,
Mathis 1982, figs. 11, 12) .
Abbreviations: i vt s, inner vertical seta; lc orb s, lateroclinate orbital setae; oc s, ocellar seta; o vt s, outer vertical seta; psdpoc s,
pseudopostoccellar seta.
EPHYDRIDAE (shore flies) 100 1221

40. Posterior margin of postgena meeting occiput at obtuse angle, broadly rounded; postgena usually
with coarse black setulae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
– Posterior margin of gena sharply angulate; postgena with fine pale setulae. . . . . . . . . . . . . . . . 49

41. Anterior and posterior notopleural setae inserted equidistant from notopleural suture;
DISCOCERININI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
– Posterior notopleural seta inserted much farther dorsad from notopleural suture than anterior
seta. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
42. Notopleuron lacking setulae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
– Notopleuron bearing setulae in addition to two large setae (Fig. 25) . . . . . . . . . . . . . . . . . . . . . 45

43. Forefemur lacking row of short, stout setae along posteroventral surface . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gymnoclasiopa Hendel
– Forefemur bearing distinct row of stout, short setae along apical half of posteroventral surface. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44

44. Face metallic, shiny, bearing white microtomentose spots laterally; forefemur slightly enlarged .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pectinifer Cresson
– Face either wholly shiny or microtomentose; forefemur normally developed . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ditrichophora Cresson

45. Face with secondary series of dorsolaterally inclined setae laterad to primary series (Fig. 26). . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polytrichophora Cresson
─ Face with secondary of setae lacking or suggested only by medially inclined setulae. . . . . . . . 46

46. Gena and lower part of parafacial broad (Fig. 27); lateral margin of abdomen usually with gray to
whitish microtomentose areas, these usually wedge shaped. . . . . . . . . . . . . Hydrochasma Hendel
─ Gena and parafacial narrow; abdomen lacking wedge-shaped, light-colored areas laterally . . . 47

47. Parafacial bearing setulae (Fig. 28). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Discocerina Macquart

─ Parafacial lacking setulae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48

48. With two setae in facial series, these well separated, distance between them subequal to length
of first flagellomere; parafacial narrow at anteroventral margin of eye; postsutural supra-alar and
prescutellar acrostichal setae greatly reduced or lacking. . . . . . . . . . . . . . Lamproclasiopa Hendel
─ With three to four setae in facial series, distance between setae conspicuously less than length
of first flagellomere; parafacial of even width throughout length; postsutural supra-alar and
prescutellar acrostichal setae present. . . . . . . . . . . . . . . . . . . . . . Orasiopa Zatwarnicki & Mathis

49. Scutellum with three pairs of marginal setae; body mostly dull, whitish to grayish. . . . . . . . . . 50
– Scutellum with two pairs of marginal setae; body mostly shiny black; GYMNOMYZINI, in part.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54

50. Ocellar setae behind level of anterior ocellus; wing pointed at apex of vein R4+5; crossvein dm-cu
clouded; ATISSINI. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
– Ocellar seta before level of anterior ocellus; wing rounded at apex of vein R4+5; crossvein dm-cu
usually not clouded; HECAMEDINI. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
1222
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

51. Face concave in lateral view, most prominent at ventral facial margin, lacking dorsally curved
setae (Fig. 12). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Atissa Haliday
– Face convex in lateral view, protruding, prominent just ventrad of antennal bases, bearing large,
dorsally-curved setae near facial prominence (Fig. 15). . . . . . . . . . . . . . . . . . Ptilomyia Coquillett

52. Scutellum bearing three marginal setae; postgenal margin sharp; gena high, over one-half eye
height (Fig. 20). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hecamede Haliday
– Scutellum bearing two marginal setae; postgenal margin rounded; gena short, less than one-half
eye height. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53

52. Color generally black; microtomentum sparse, subshiny to shiny; head as in Fig. 22 . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diphuia Cresson
– Color generally gray to brown; microtomentum dense, generally appearing dull; head as in
Fig. 21 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Allotrichoma Becker

54. Anterior and posterior notopleural setae present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55

– Anterior notopleural seta lacking. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57

55. Alula wide, width greater than subcostal cell, and auriculate; face below antennal grooves evenly
convex and completely transversely wrinkled to form series of depressions . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cerometopum Cresson
– Alula narrow, width less than subcostal cell; face usually with midfacial prominence or if convex
not wrinkled as above (Fig. 18); Athyroglossa Loew . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56

56. Mesonotal setulae in regular rows; forefemur without posteroventral, spinelike setulae; aristal
rays relatively short, length of longest rays about one-half or less height of first flagellomere . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Athyroglossa (Parathyroglossa) Hendel
– Mesonotal setulae in irregular rows; forefemur bearing three to seven, short, posteroventral,
spinelike setulae; aristal rays relatively long, length of longest rays equal to or greater than half
height of first flagellomere (Fig. 18) . . . . . . . . . . . . . . . . . . . . Athyroglossa (Athyroglossa) Loew

57. Forefemur unarmed, lacking row of stout setae along posteroventral surface at apical one-quarter;
outer vertical seta absent; scutum with several setae in oblique row between postalar seta and base
of scutellum. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Placopsidella Kertész
– Forefemur bearing 5–10 stout setae along posteroventral surface at apical one-third; inner and
outer vertical setae present (Fig.  19); scutum lacking setae between postalar seta and base of
scutellum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mosillus Latreille

58. Median facial area and lower facial margin setose, the latter often with long setae; insertions of
facial series of setae convergent above; subcranial cavity large and cavernous . . . . . . . . . . . . . 59
– Median facial area and lower facial margin without setae; facial setae inserted in more or less
vertical series, parallel with parafacial; subcranial cavity small to large. . . . . . . . . . . . . . . . . . . 63

59. Two postsutural dorsocentral setae present, both equally strong; disc of proepisternum usu-
ally bare, sometimes with few fine setulae; SCATELLINI. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
– With three to four postsutural dorsocentral setae (Fig.  46), anterior three sometimes short and
weak; disc of proepisternum with few to many fine setulae; EPHYDRINI . . . . . . . . . . . . . . . . 67

60. With one lateroclinate fronto-orbital seta. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61

– With two lateroclinate fronto-orbital setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
EPHYDRIDAE (shore flies) 100 1223

infr s

38 Physemops 39 Physemops 40 Ephydra (Hydropyrus)

41 Cirrula 42 Cirrula 43 Cirrula

44 Dimecoenia 45 Dimecoenia 46 Dimecoenia

Figs.  100.38–46. Heads (continued), leg, and thorax: lateral view  (38), with offset of ommatidia, and anterior view  (39) of Physemops
nemorosus (Cresson), (Mathis, 1982, figs. 28, 29); anterodorsal view of head of (40) Ephydra (Hydropyrus) hians (Say), (Nearctic, MND,
fig. 98.38); lateral view (41) and anterior view (42) of head of Cirrula austrina (Coquillett), (Nearctic, Mathis & Simpson, 1981, figs. 1, 2);
anterior view of hind leg of (43) C. austrina, (Nearctic, Mathis & Simpson, 1981, fig. 4): lateral view (44) and anterior view (45) of head of
Dimecoenia spinosa (Loew), (Nearctic, Mathis & Simpson, 1981, figs. 62, 63); dorsal view of thorax of (46) D. spinosa, (Nearctic, Mathis &
Simpson, 1981, fig. 64).
Abbreviation: infr s, interfrontal seta.
1224
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

47 Scatella 48 Haloscatella

49 Ochthera 50 Allotrichoma

presut
dc s

psut spal s
p npl s
pal s
presctl s

a npl s

52 Haloscatella

51 Hydrellia

Figs. 100.47–52. Heads (concluded), leg, thoraces, and notopleuron: lateral view of head of (47) Scatella marinensis (Cresson), (Mathis &
Shewell, 1978, fig. 55); anterodorsal view of head of (48) Haloscatella muria (Mathis), (Nearctic, Mathis, 1979b, fig. 14); posterior view of
foreleg of (49) Ochthera mantis (De Geer), (Holarctic, MND, fig. 98.63); dorsal view of thorax of (50) Allotrichoma simplex (Loew), (Nearc-
tic, MND, fig. 98.48); and (51) Hydrellia griseola (Fallén), (Holarctic, MND, fig. 98.49); lateral view of notopleuron of (52) Ha. muria,
(Nearctic, MND, fig. 98.49, as Lamproscatella).
Abbreviations: a npl s, anterior notopleural seta; pal s, postalar seta; p npl s, posterior notopleural seta; presctl s, prescutellar seta; presut dc s,
presutural dorsocentral seta; psut spal s, postsutural supra-alar setae.
EPHYDRIDAE (shore flies) 100 1225

61. Costa short, extended to apex of vein R4+5 (Fig. 57) . . . . . . . . . . . . . . . . . . . . . . Scatophila Becker
– Costa long, extended to apex of vein M. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Limnellia Malloch

62. Arista pectinate with short, dorsally branching rays (Fig.  47); genal seta large, conspicuous;
posterior notopleural seta inserted at approximately same level as anterior seta; wing frequently
spotted (Fig. 56). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scatella Robineau-Desvoidy
– Arista at most macropubescent, lacking prominent, dorsally branching rays (Fig. 48); genal seta
greatly reduced or lacking; posterior notopleural seta inserted at distinctly higher level than
anterior seta (Fig. 52); wing hyaline or slightly infuscate, not spotted. . . . . . Haloscatella Mathis

63. Presutural or sutural dorsocentral seta present, conspicuous; head as in Fig. 32; PARYDRINI. . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parydra Stenhammar
– Presutural or sutural dorsocentral seta inconspicuous or absent; head various; DAGINI. . . . . . 64

64. Costal vein short, extending only to apex of vein R4+5; face with distinct, mediovertical carina
(Fig.  33), extended from ptilinal suture to oral margin; arista distinctly pectinate, longest rays
subequal to length of first flagellomere. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachydeutera Loew
– Costal vein long, extending to apex of vein M; face evenly transversely arched, not carinate; arista
essentially bare to subpectinate with longest rays not more than one-half length of first flagellomere.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65

C sector 2 C sector 3

R4+5

53 Zeros 54 Notiphila (Agrolimna)

R1

r-m

55 Parydra 56 Scatella

R4+5

57 Scatophila

Figs. 100.53–57. Wings: dorsal view of (53) Zeros flavipes (Williston), (Nearctic, MND, fig. 98.52); (54) Notiphila (Agrolimna) aenigma
Cresson, (Nearctic, MND, fig. 98.53, as N. (Dichaeta) aenigma); (55) Parydra fossarum (Haliday), (Holarctic, MND, fig. 98.54, as
P. (Chaetoapnea) parasocia); (56) Scatella marinensis (Cresson), (Mathis & Shewell, 1978, fig. 53); (57) Scatophila cribrata (Stenhammar),
(Holarctic, MND, fig. 98.59).
1226
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

65. Ventral one-half to two-thirds of face distinctly projected anteriorly (Figs.  34–35), arched
transversely and vertically (best seen in profile); with three fronto-orbital setae, anterior seta
smaller; pulvilli lacking; postpronotum with one to few setulae. . . . . . . . . . . . . . . Dagus Cresson
– Face shieldlike, broadly but more shallowly projected over most of height (Figs. 37, 39); usually
with one to two fronto-orbital setae, if two, these subequal; pulvilli present, conspicuous;
postpronotum bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66

66. Gena high, equal to or greater than eye height (Figs. 36–37); genal seta well developed and
conspicuous; prescutellar acrostichal setae well developed; propleuron setulose; tarsomere 5 with
dorsoapical process extended beyond base of tarsal claws. . . . . . . . . . . Diedrops Mathis & Wirth
– Gena short, usually not more than one-half eye height (Fig. 38); genal seta, if present, weakly
developed and inconspicuous; prescutellar acrostichal setae not evident; propleuron lacking
setulae; tarsomere 5 not as above. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Physemops Cresson

67. Three to four well-developed fronto-orbital setae present; anterior presutural supra-alar seta
present, subequal to posterior notopleural seta; Ephydra Fallén. . . . . . . . . . . . . . . . . . . . . . . . . 68
– Two well-developed fronto-orbital setae present; anterior presutural supra-alar seta absent or
much reduced, much smaller than posterior notopleural seta . . . . . . . . . . . . . . . . . . . . . . . . . . . 70

68. Two presutural dorsocentral setae present, anterior sometimes short; face thickly setulose; arista
short, weakly haired, thicker on its basal half; usually two posteriorly directed rows of
well-developed cruciate interfrontal setae present, with these rows closer to orbital setae than to
each other; male with tarsomere 1 of foreleg bearing ventral tuft of long setulae near tip; head as
in Fig. 40 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ephydra (Hydropyrus) Cresson
– Only one presutural dorsocentral setae present; face with one well-developed row of facial setae
and one row of oral setae, otherwise thinly short-haired; arista without markedly swollen basal
region; usually not more than one interfrontal seta situated as close to each other as to orbital
setae; male with tarsomere 1 of foreleg without ventral tuft of setulae; head as in Fig. 1. . . . . . 69

69. One well-developed interfrontal seta present (Fig. 1); crossvein dm-cu making nearly right angle
with vein CuA1; palpus well developed. . . . . . . . . . . . . . . . . . . . . . . . . Ephydra (Ephydra) Fallén
– Interfrontal seta weak or absent; crossvein dm-cu making acute angle with vein  CuA1; palpus
small. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ephydra (Halephydra) Wirth

70. Cruciate interfrontal setae present; South America, Caribbean. . . . . . . . . . [Paraephydra Mathis]
– Cruciate interfrontal setae absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71

71. Aristal rays long, length subequal to width of pedicel (Figs. 44–45); hind femur of male not
differing markedly from fore- or midfemur, lacking stout setae as below; hind tibia of male
lacking tuft of setulae; hind tarsus of male cylindrical, normal. . . . . . . . . . . . Dimecoenia Cresson
– Aristal rays short, length approximately one-half width of pedicel (Figs. 41–42); hind femur of
male conspicuously swollen, bearing short row of four to five stout setae along anteroventral
surface toward base; hind tibia of male with ventroapical tuft of setulae (Fig. 43); hind tarsus of
male variously modified. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cirrula Cresson
EPHYDRIDAE (shore flies) 100 1227

v rep

pregt

pgt ph st 6
sur
hypd tg 6
st 7

ph tg 7
pregt
epand st 8
pgt tg 8
cerc
hypd cerc

58 Hydrellia 59 Paralimna

60 Hydrellia

61 Notiphila

62 Ephydra

Figs. 100.58–62. Terminalia, egg, and larvae: ventral view of apex of male abdomen and details of male terminalia (inset) of (58) Hydrellia
griseola (Fallén), (Nearctic, MND, fig. 98.66); lateral view of female terminalia of (59) Paralimna punctipennis (Wiedemann), (Nearctic,
MND, fig. 98.67); lateral view of egg of (60) H. bergi Cresson, (Nearctic, MND, fig. 98.68); lateral view of third instar larva, with inset of
enlarged apex of respiratory tube, of (61) Notiphila sp., (Nearctic, MND, fig. 98.69); and (62) Ephydra sp., (Nearctic, MND, fig. 98.70).
Abbreviations: cerc, cercus; epand, epandrium; hypd, hypandrium; pgt, postgonite; ph, phallus; pregt, pregonite; st, sternite; sur, surstylus;
tg, tergite; v rep, ventral receptacle.
1228
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Synopsis of the fauna lentic aquatic habitats, especially small, undisturbed pools.
The species are seemingly able to tolerate considerable
The Ephydridae include 1842 valid species that are placed pollution, such as the pooled effluent from coffee processing
in 127 genera (Mathis  & Zatwarnicki, 1995, and updates). plants.
The New World fauna comprises 86 genera and 743 species,
and we estimate that there are 280 species in Central Ameri- Cerometopum Cresson. This genus includes two Neotro-
ca. Two named fossil species in Ephydra and undescribed pical species, one of which is apparently endemic to Central
representatives of Beckeriella Williston are known (Evenhuis, America. Adult specimens are rare, and nothing is known
1994). about the immature stages of the only Central American spe-
cies, C. lacunosum Cresson.
Allotrichoma Becker. This genus occurs worldwide and
includes 52 species of which four are found in Central America. Ceropsilopa Cresson. Among shore fly genera, Ceropsilopa
One subgenus, Pseudohecamede Hendel, primarily occurs is moderately diverse, with 18 species worldwide. The
in maritime saline habitats, and the nominate subgenus, Central American fauna includes four species that occur
Allotrichoma, is mostly associated with freshwater environ- most often on the coastal plain. Adults are found on low-lying
ments. Larvae are probably scavengers, and adults have been vegetation and are sometimes common adjacent to beaches
reared from dead and decaying arthropods. near vegetated sand dunes.

Athyroglossa Loew. Worldwide there are 31 species of Cirrula Cresson. This genus occurs only in North and
which six have been recorded from Central America. At Central America where there are three species. One species,
least three others are known and remain to be described. The C. austrina (Coquillett), is found in Central America, usually
larvae of A. granulosa (Cresson) and A. glabra (Meigen), associated with algal mats in coastal marine habitats
which occur in North America, are associated with skunk (Mathis, 1997b). The larvae feed on the algae, and adults are
cabbage (Grimaldi  & Jaenike, 1983) and as scavengers in most often encountered on the surface of the algal mats.
decaying vertebrates (Krivosheina & Ozerov, 1989), respec- Clasiopella Hendel. A single species, C. uncinata Hendel,
tively. Adults are associated with both lotic and lentic aquatic is found in the New World, including Central America, and
systems. was probably introduced (Mathis, 1994) from the Australasian
Atissa Haliday. Like other genera of the tribe Atissini, and/or Oceanian Regions, where a second species occurs.
Atissa is found almost exclusively where saline aquatic Adults occur along maritime coasts and are sometimes
systems occur, primarily along maritime coasts but also common along the margins of mangrove swamps where
inland. Two species are known from Central America, and there is emergent vegetation.
worldwide there are 14. Adult specimens are tiny, frequently Cressonomyia Arnaud. Among Psilopine genera,
less than 2  mm in body length, and are found directly on Cressonomyia is easily recognized by the darkened wing
exposed sand or mud with little or no vegetation. base and brownish-black halteres. The genus and its sister
Beckeriella Williston. Beckeriella was recently syn- genus, Peltopsilopa Hendel, occur only in the New World,
and four species comprise the Central American fauna,
onymized with Gastrops (Lizarralde de Grosso, 1998), but
which Mathis  & Zatwarnicki (2004) recently revised. The
I prefer to recognize both genera. The two genera are sis-
immature stages are not known, but there is some indication
ter groups, however (Mathis, 1977). Beckeriella, which cur-
that the species are phytophagous.
rently includes 10 species worldwide, has a peculiar and
disjunct distribution, occurring in the New World tropics, Dagus Cresson. A single species, D. rostratus, is known
Madagascar (Mathis  & Grimaldi, 2000), and the Philip- from Central America, but in the West Indies, especially the
pines (Lizarralde de Grosso, 1994). Moreover, Beckeriella Dominican Republic, the genus has radiated, with over eight
is one of the few shore fly genera for which fossils are species (Stark, 1993). The genus is only known from the
known (from Dominican amber; Evenhuis, 1994). There New World tropics. Adults and presumably the immatures
are records of three species in Central America, although occur in the splash zone and along the sides of waterfalls
others remain to be described. The immature stages are and cascades.
unknown.
Diedrops Mathis & Wirth. This genus is found only in
Brachydeutera Loew. Although easily recognized, the the New World tropics, where four species are known. A
phylogenetic relationships of this genus are not fully resolved single species, D. steineri Mathis, occurs in Central America.
and placement in Dagini remains tentative. There are 15 Adults are relatively uncommon, and most are found on
species worldwide, and from Central America there are two, exposed rocks in well-oxygenated, fast-flowing streams and
B.  neotropica Wirth and B.  longipes Hendel (Mathis  & associated with waterfalls. Mathis  & Hogue (1986) reared
Winkler, 2003). The latter species was introduced from the adults of D. roldanorum Mathis & Hogue from puparia that
Oriental Region. Adults and immature stages are found in were collected in a lotic aquatic system in Colombia.
EPHYDRIDAE (shore flies) 100 1229

Dimecoenia Cresson. This genus is represented by a Glenanthe Haliday. Worldwide there are 15 described
single species in Central America, D. spinosa (Loew), that species and four are recorded from Central America. The
occurs in coastal habitats where there are protected embayments genus is easily recognized by the pyriform eye and only
with algal mats, such as in some mangrove swamps (Mathis, occurs in coastal or inland saline habitats. Adults from
1997b). Worldwide there are 18 species, with most occurring Central America can be identified using Mathis (1995). The
in South America. The larvae breed in the algal mats immature stages are unknown.
(Mathis & Simpson, 1981).
Hecamede Haliday. A single species, H.  brasiliensis
Diphuia Cresson. Although superficially resembling Cresson, occurs in Central America, and worldwide there are
Athyroglossa, being somewhat small, robust and shiny 12 species with greater species diversity in the Old World
black, this genus is more closely related to genera of the tropics (Mathis, 1993b). The genus is associated only with
tribe Hecamedini. The genus occurs only in the New World, maritime coastal habitats, where the adults are found on bare
where there are four species, of which two occur in Central sand and/or muddy shores. The immatures are scavengers/
America, D.  anomala Cresson and D.  nitida Sturtevant  & saprophytic and can occur in great numbers on dead fish,
Wheeler (Mathis, 1997a). Adults are mostly found in coastal horseshoe crabs, and other marine carrion (Norrbom, 1983).
marine habitats, although sometimes also in freshwater
aquatic environments. The immature stages are unknown. Hyadina Haliday. Worldwide there are 40 species in
this genus, and only two, H.  bulbosa Clausen and H.  pal-
Discocerina Macquart. Worldwide this genus includes lipes (Cresson), are recorded from Central America. Others
28 species, of which seven occur in Central America. There are known, however, and await description. Foote (1993)
are numerous undescribed species from the region, however. discovered larvae of H.  albovenosa Coquillett on soil-
Adults are found on the sandy to muddy banks of various inhabiting colonies of Anabaena, a genus of Cyanobacteria.
aquatic systems. Foote & Eastin (1974) reared the larva of
D.  obscurella (Fallén) on a medium of wet peat moss and Hydrellia Robineau-Desvoidy. This is the largest genus
decaying lettuce and suggested that the larvae were probably of shore flies, with over 210 species worldwide. Six species
utilizing microorganisms. are known from Central America, but numerous others
remain to be named. Larvae are miners in the leaves and
Ephydra Fallén. This is the nominate genus for the family stems of various plants, mostly monocots that are associated
and is relatively species rich, with 33 species worldwide. The with aquatic environments. A few species can be serious
Neotropical fauna of Ephydra, however, is relatively depau- pests in cereal crops, especially rice.
perate, comprising four species, of which three, E. mexicana
Cresson, E.  millibrae Jones, and E.  packardi Wirth, occur Hydrochasma Hendel. This genus occurs only in the New
in central and northern Mexico. The occurrence of the World, where there are six described species, of which two
latter two species in northern Mexico are extensions of more occur in Central America. Numerous undescribed species are
widespread distributions in the United States. Ephydra known, however. Adults occur on sandy to muddy shores of both
gracilis Packard is widespread throughout the Caribbean and freshwater and brackish water habitats but are more diverse and
is expected along the Atlantic coast of Central America. Wirth abundant in the latter. The immature stages are unknown.
(1971) revised the North American species and included the
Ilythea Haliday. Although primarily a north temperate
four species that are known from the northern Neotropical
genus, a single species, I. caniceps Cresson, extends into the
Region.
northern part of the Neotropical Region, including Central
Garifuna Mathis. This is a monotypic genus that America. Worldwide there are 12 species.
is known only from the New World and was described
Lamproclasiopa Hendel. Worldwide there are 14 species,
from specimens collected in Belize and Panama (Mathis,
and in Central America three have been recorded, but others
1997b). The only species, G.  sinuata Mathis, is rare and
are known and remain to be described. Adults are not
has been collected along the coast. The immature stages
common and are found along the banks of small streams.
are unknown.
The immature stages are unknown.
Gastrops Williston. This genus includes eight species, all
in the New World, and five are found in Central America. Most Lemnaphila Cresson. Only one species of the seven
of the species can be identified using Lizarralde de Grosso’s species of this genus, L. neotropica Lizarralde de Grosso, is
review (1998), but there are a few undescribed Central known from Central America (Mathis  & Edmiston, 2000).
American species that are not included. The known larvae It is apparently widespread in most of the Neotropical
are egg predators in the foam nests of frogs, especially frogs Region, although there are large gaps in the known distribution.
of the family Leptodactylidae and the genus Hyla. Gastrops Specimens are tiny, often less than 1 mm in body length, and
niger Williston, however, also occurs on islands of the are associated only with plants of the famililes Lemnaceae
Belizian barrier reef, where no frogs are found (Mathis, and Hydrocharitaceae. Larvae of Lemnaphila are miners in
1997b). the leaflets of these plants.
1230
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Leptopsilopa Cresson. Five species comprise the Central Ochthera Latreille. This genus is easily recognized by
American fauna, and worldwide there are 17. Adults are the large, raptorial forelegs. Worldwide there are 37 species,
collected by sweeping vegetation, especially grasses, and and eight are known from Central America. Both adults and
some species are fairly common on urban lawns that have not larvae are predatory (Simpson, 1975) on small terrestrial
been recently mowed. The immature stages are unknown. insects and on midge and mosquito larvae. The New World
species can be identified using Clausen (1977, 1980).
Limnellia Malloch. This is mostly a temperate genus,
and worldwide there are 18 species, of which one, L. huachuca Oedenops Becker. In the New World, this genus is
Mathis, is known from Central America (Mathis, 1978). represented by a single species, O. nudus (Coquillett), which
Specimens, which are easily recognized by the spotted occurs primarily in subtropical and tropical areas (Mathis &
wing, shiny black abdomen, and a single, lateroclinate Zatwarnicki, 2002). Two additional species are found in the
fronto-orbital seta, are from higher elevations in Central Old World. Adults are found on mostly bare sandy to muddy
America, usually above 2000 m. The immature stages are margins of rivers and streams and occasionally along
unknown. maritime beaches. The immature stages are unknown.

Lipochaeta Coquillett. Mathis  & Trautwein (2003) Orasiopa Zatwarnicki  & Mathis. A single species,
recently revised this genus, which occurs only in the New O.  mera (Cresson), occurs in Central America and is
World and includes two species, L. slossonae Coquillett and apparently adventive (Zatwarnicki, 1991) and now fairly
L.  ranica Mathis  & Trautwein. Both species are found in widespread. Adults are associated with coastal marine
Central America, where they are associated with bare and habitats, especially the wrack zone. The immature stages are
exposed sand and sometimes muddy shorelines in pro- unknown.
tected embayments along the coast. The immature stages are
unknown. Paralimna Loew. This is one of the largest shore fly
genera with over 85 species worldwide. In Central America
Lytogaster Becker. This is primarily a north temper- there are nine described species, but many others are known
ate genus with 10 species worldwide. A single species, in collections and undoubtedly numerous others remain to be
L.  granulosus Cresson, was described from specimens discovered. Apart from the subgenus Phaiosterna Cresson,
collected in Costa Rica. which was recently revised (Mathis  & Zatwarnicki, 2002),
the genus is greatly in need of revision, and species are
Mimapsilopa Cresson. Two species groups are evident presently difficult to determine from the scattered records
in Mimapsilopa (Mathis  & Zatwarnicki, 1998a), and the and species descriptions.
two species found in Central America both belong to the
metatarsata group, which has a shiny face. There is at least Paratissa Coquillett. Three extremely similar species
one new species in the collection of INBio. comprise the Central American fauna, and reference to
structures of the male terminalia is needed to distinguish
Mosillus Latreille. This genus was fairly recently revised among them (Mathis, 1993a). Worldwide there are four spe-
(Mathis et al., 1993), and worldwide there are 12 species. cies. Adults, and presumably the immature stages, are found
Only M. stegmaieri Wirth occurs in Central America, where in wrack, and specimens are sometimes abundant where
it is found in a variety of habitats, including the brine resulting large windrows of wrack have accumulated in protected
from processing coffee beans or brackish-water habitats embayments.
along the coast. The immature stages are unknown.
Parydra Stenhammar. This is primarily a north temperate
Nostima Coquillett. Thirty species are presently known genus and is relatively diverse with more than 70 species.
for this genus, of which 12 are reported from Central Only three species are known from Central America, and
America. The New World fauna is diverse and was recently one, P.  nigripes (Cresson), was described from specimens
revised (Edmiston & Mathis, 2005). Foote (1983) discovered collected in Costa Rica (Clausen  & Cook, 1971). The
larvae of N. approximata Sturtevant & Wheeler feeding on immature stages are in muddy to sandy substrates associated
the algal genus Oscillatoria and certain other related genera with aquatic environments.
of blue-green algae in Ohio, USA.
Pectinifer Cresson. This monotypic genus includes only
Notiphila Fallén. This genus is diverse worldwide with P. aeneus (Cresson), which was first described from specimens
about 150 species. In Central America there are only three collected on the slopes of Volcán Irazú, Costa Rica. Although
known species, although others are known and await description. the species is more widespread in Central America, the
The larvae are detritivores in highly organic mud and sand, immature stages are still not known.
where they obtain their oxygen by tapping the rootlets of
various aquatic plants with their rear spiracles (Houlihan, Physemops Cresson. This genus is well represented in
1969). Adults and immatures are found where emergent Central America, where five of the six described species
vegetation, especially monocots, occur. occur. The genus is exclusively Neotropical and is in need
EPHYDRIDAE (shore flies) 100 1231

of revision, as numerous species, especially in the P. nem- species, R. ardeoceras Mathis and R. griseola Rao & Mathis,
orosus group, are known in collections. Adults are found on occur in Central America. Of these two species, R. griseola
exposed rocks along margins and in fast-flowing and cascading is more widespread, being found as far south as Bolivia;
streams. R. ardeoceras has been only found near Tilaran (Costa Rica:
Guanacaste). The immature stages are unknown, but adults
Placopsidella Kertész. Two species occur in Central of R. griseola are associated with lotic aquatic systems, often
America, P. cynocephala Kertész and C. grandis (Cresson), occurring on exposed rocks in and along the margins of
and both are apparently adventive to the New World. fast-flowing streams.
Elsewhere, especially in the Oriental, Australasian, and
Oceanian Regions, there are eight species (Mathis, 1986). Scatella Robineau-Desvoidy. This is a diverse genus
Tenorio (1980) and Kirk-Spriggs (1986) reported on the worldwide with 135 species, of which only three are reported
biology of P.  marquesana (Malloch) and P.  cynocephala from Central America. The Neotropical fauna includes many
Kertész, respectively. Larvae of both species apparently feed undescribed species, however, especially from higher elevations
as scavengers on dead marine invertebrates that have become such as mountainous wetlands in Central and South America.
Adults occur in a diversity of habitats (Foote, 1995), and the
stranded on the beach. Adults often occur on seaweed and
larvae feed on algae.
other debris on maritime beaches.
Scatophila Becker. The Neotropical fauna of Scatophila
Polytrichophora Cresson. Worldwide there are 18 species,
of which three are reported to occur in Central America. The is largely unknown and numerous species await description.
Neotropical fauna is more diverse, however, and is now Worldwide there are 50 species, and at least three from Costa
being revised (W.N.  Mathis, in preparation). Adults occur Rica have been identified but are undescribed. In Central
in coastal marine habitats as well as in freshwater environ- America specimens generally occur at higher elevations and
ments. The immature stages are unknown. are usually associated with sandy to muddy shoreline habitats
that are largely bare of vegetation.
Psilopa Fallén. Psilopa is one of the most species-rich
genera of shore flies, with over 70 species worldwide. The Typopsilopa Cresson. Three species are known from
Old World fauna is considerably more diverse, and conversely Central America, and worldwide there are 19. Adults occur
the Central American fauna is relatively depauperate with on emergent vegetation, and Keiper et al. (2001) discovered
only two species, P. girschneri Röder and P. pulchripes Loew. that one species that occurs in Central America, T.  nigra
Of these two species, P. girschneri is probably adventive. The Williston, is a secondary invader of bulrush and cattails that
immature stages are virtually unknown, but phytophagy is had first been mined by larvae of the noctuid moth, Bellura
suspected. obliqua gargantua (Dyar). Although the genus is distinctive,
the species are homogeneous in external structures and, to
Ptilomyia Coquillett. This genus is associated with saline an extent, color. Thus, dissection and study of structures of
habitats and to a lesser degree with freshwater environments. the male terminalia are frequently necessary to distinguish
The Central American fauna is relatively diverse with five among the species.
species, and worldwide there are 12. The dorsal rays on the
arista distinguish this genus from Atissa, and both genera fre- Zeros Cresson. This genus appears to be pantropical.
quently occur together. The immature stages are unknown. Worldwide there are 13 species, of which four are known
from Central America. The natural history of the species
Rhysophora Cresson. Species of Rhysophora are occurring in the tropics, including those in Central America,
only known from the New World and five species are is essentially unknown. A Nearctic species, Z. flavipes (Wil-
now included in the genus (Rao  & Mathis, 1995). Two liston), is found in sedge-meadow and mud-shore habitats.

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excavata, a grazer of blue-green algae (Diptera: Ephydridae). 1–111.
Proceedings of the Entomological Society of Washington 83: Mathis, W.N. 1979b. Studies of Ephydrinae (Diptera: Ephydridae),
304–315. II:  Phylogeny, classification, and zoogeography of Nearctic
Foote, B.A. 1981b. Biology and immature stages of Pelina Lamproscatella Hendel. Smithsonian Contributions to Zoology
truncatula, a consumer of blue-green algae (Diptera: Ephydridae). 295: 1–41.
Proceedings of the Entomological Society of Washington 83: Mathis, W.N. 1982. Studies of Ephydrinae (Diptera: Ephydridae),
607–619. VI:  Review of the tribe Dagini. Smithsonian Contributions to
Foote, B.A. 1983. Biology and immature stages of Nostima Zoology 345: 1–30.
approximata (Diptera: Ephydridae), a grazer of blue-green algal Mathis, W.N. 1986. Studies of Psilopinae (Diptera: Ephydridae), I: A
genus Oscillatoria. Proceedings of the Entomological Society of revision of the shore fly genus Placopsidella Kertész. Smithsonian
Washington 85: 472–484. Contributions to Zoology 430: 1–30.
Foote, B.A. 1993. Biology of Hyadina albovenosa (Diptera: Mathis, W.N. 1991. Classification of the shore flies (Diptera:
Ephydridae), a consumer of cyanobacteria. Proceedings of the Ephydridae). Past, present, and future, pp.  209–227. In Weis-
Entomological Society of Washington 95: 377–382. mann, L., I. Országh, & A.C. Pont (editors). Proceedings of the
Foote, B.A. 1995. Biology of shore flies. Annual Review of Second International Congress of Dipterology. SPB Academic
Entomology 40: 417–442. Publishing bv, The Hague, The Netherlands, 367 pp.
Foote, B.A. & W.C. Eastin. 1974. Biology and immature stages of Mathis, W.N. 1993a. A revision of the shore-fly genera Hostis Cresson
Discocerina obscurella (Diptera: Ephydridae). Proceedings of and Paratissa Coquillett (Diptera: Ephydridae). Proceedings of
the Entomological Society of Washington 76: 401–408. the Entomological Society of Washington 95: 21–47.
EPHYDRIDAE (shore flies) 100 1233

Mathis, W.N. 1993b. Studies of Gymnomyzinae (Diptera: Ephydri- Mathis, W.N. & T. Zatwarnicki. 2002. A phylogenetic study of the
dae), IV: A revision of the shore-fly genus Hecamede Haliday. tribe Dryxini Zatwarnicki (Diptera: Ephydridae). Smithsonian
Smithsonian Contributions to Zoology 541: 1–46. Contributions to Zoology 617: 1–101.
Mathis, W.N. 1994. A revision of the genus Clasiopella Hendel Mathis, W.N. & T. Zatwarnicki. 2004. A revision of the shore-fly
(Diptera: Ephydridae). Proceedings of the Entomological genus Cressonomyia Arnaud (Diptera: Ephydridae), with com-
Society of Washington 96: 454–465. ments on species that have been excluded. Proceedings of the
Mathis, W.N. 1995. Studies of Gymnomyzinae (Diptera: Ephydridae), Entomological Society of Washington 106: 249–279.
VI: A revision of the shore-fly genus Glenanthe Haliday from Mathis, W.N., T. Zatwarnicki, & M.G. Krivosheina. 1993. Studies
the Western Hemisphere. Smithsonian Contributions to Zoology of Gymnomyzinae (Diptera: Ephydridae), V: A revision of the
567: 1–26. shore-fly genus Mosillus Latreille. Smithsonian Contributions
Mathis, W.N. 1997a. A review of the shore-fly genus Diphuia to Zoology 548: 1–38.
Cresson (Diptera: Ephydridae). Proceedings of the Entomological Norrbom, A.L. 1983. Four acalyptrate Diptera reared from dead
Society of Washington 99: 28–36. horseshoe crabs. Entomological News 94: 117–121.
Mathis, W.N. 1997b. The shore flies of the Belizean Cays (Diptera: Oldroyd, H. 1964. The Natural History of Flies. Weidenfeld  &
Ephydridae). Smithsonian Contributions to Zoology 592: 1–77. Nicholson, London, 324 pp.
Mathis, W.N.  & J.F. Edmiston. 2000. A revision of the shore-fly Rao, N. & W.N. Mathis. 1995. A revision of the genus Rhysophora
genus Lemnaphila Cresson (Diptera: Ephydridae). Proceedings Cresson (Diptera: Ephydridae). Proceedings of the Entomologi-
of the Entomological Society of Washington 102: 652–677. cal Society of Washington 97: 666–688.
Mathis, W.N. & D.A. Grimaldi. 2000. Chapter 4. The first Beckeriella Scheiring, J.F. & B.A. Foote. 1973. Habitat distribution of the shore
Williston from the Afrotropical Region: two new species from flies of northeastern Ohio (Diptera: Ephydridae). Ohio Journal
Madagascar (Diptera: Ephydridae), pp. 105–115. In Goodman, S.M. of Science 73: 152–166.
(editor). A floral and faunal inventory of the Parc National de Simpson, K.W. 1975. Biology and immature stages of three species
Marojejy, Madagascar: with reference to elevational variation. of Nearctic Ochthera (Diptera: Ephydridae). Proceedings of the
Fieldiana: Zoology 97: viii + 1–286. Entomological Society of Washington 77: 129–155.
Mathis, W.N. & C.L. Hogue. 1986. Description of a new species Simpson, K.W. 1976a. Shore flies and brine flies (Diptera: Ephydri-
of the shore fly genus Diedrops (Diptera: Ephydridae) from dae), pp. 465–495. In Cheng, L. (editor). Marine Insects. North-
Colombia. Contributions in Science 377: 21–26. Holland Publishing Company, Amsterdam, xii + 581 pp.
Mathis, W.N.  & G.E. Shewell. 1978. Studies of Ephydrinae Simpson, K.W. 1976b. The mature larvae and puparia of Ephydra
(Diptera: Ephydridae), I: Revisions of Parascatella Cresson and (Halephydra) cinerea Jones and Ephydra (Hydropyrus) hians
the triseta group of Scatella Robineau-Desvoidy. Smithsonian Say (Diptera: Ephydridae). Proceedings of the Entomological
Contributions to Zoology 285: 1–44. Society of Washington 78: 263–269.
Mathis, W.N.  & K.W. Simpson. 1981. Studies of Ephydrinae Stark, J. 1993. A revision of the Neotropical genus Dagus Cresson
(Diptera: Ephydridae), V:  The genera Cirrula Cresson and (Diptera: Ephydridae). American Museum Novitates 3080: 1–21.
Dimecoenia Cresson in North America. Smithsonian Contribu- Tenorio, J.A. 1980. Family Ephydridae, pp. 251–351. In Har-
tions to Zoology 329: 1–51. dy,  D.E.  & M.D.  Delfinado (editors). Insects of Hawaii. Vol-
Mathis, W.N. & M.D. Trautwein. 2003. A revision and phyloge- ume 13. Diptera: Cyclorrhapha III, Series Schizophora section
netic study of Lipochaeta Coquillett (Diptera: Ephydridae). Acalypterae, exclusive of family Drosophilidae. The University
Proceedings of the Entomological Society of Washington Press of Hawaii, Honolulu, vii + 451 pp.
105: 30–49. Wirth, W.W. 1968. 77. Family Ephydridae. In Papavero, N. (editor).
Mathis, W.N. & I.S. Winkler. 2003. A review of the Neotropical A Catalogue of the Diptera of the Americas South of the United
species of Brachydeutera Loew (Diptera: Ephydridae). Proceedings States. Departamento de Zoologia, Secretaria da Agricultura,
of the Entomological Society of Washington 105: 406–423. São Paulo, 43 pp.
Mathis, W.N. & T. Zatwarnicki. 1995. A world catalog of the shore Wirth, W.W. 1971. The brine flies of the genus Ephydra in North
flies (Diptera: Ephydridae). Memoirs on Entomology, Interna- America (Diptera: Ephydridae). Annals of the Entomological
tional 4: vi + 1–423. Society of America 64: 357–377.
Mathis, W.N.  & T. Zatwarnicki. 1998a. A review of the West Wirth, W.W., W.N. Mathis, & J.R. Vockeroth. 1987. 98. Ephydri-
Indian species of Mimapsilopa Cresson (Diptera: Ephydridae). dae, pp. 1027–1047. MND, Volume 2.
Proceedings of the Entomological Society of Washington 100: 7–24. Zatwarnicki, T. 1991. Changes in nomenclature and synonymies
Mathis, W.N.  & T. Zatwarnicki. 1998b. Chapter 49. Family of some genera and species of Ephydridae (Diptera). Deutsche
Ephydridae, pp.  537–570. In Papp, L.  & B.  Darvas (edi- Entomologische Zeitschrift 38: 295–333.
tors). Contributions to a Manual of Palaearctic Diptera. Zatwarnicki, T. 1992. A new classification of Ephydridae based on
Volume  3, Higher Brachycera. Science Herald Publishers, phylogenetic reconstruction (Diptera: Cyclorrhapha). Genus 3:
Budapest, 880 pp. 65–119.
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
NANNODASTIIDAE (nannodastiid beach flies) 101
Wayne N. Mathis

Fig. 101.1. Male of Azorastia mediterranea Papp, (Palearctic, Papp, 1980, fig. 19).

Diagnosis sclerotized medially; clypeus small; three well-developed,

slightly dorsoclinate, parafacial setae present; vibrissae and
Adults minute to small flies (body length 0.70–1.25 mm); vibrissal angle lacking. Scutum with one dorsocentral seta
mostly brown and dull colored, sparsely to moderately present (0+1); one postalar seta present; apical scutellar seta
microtomentose (Fig. 1). Three reclinate fronto-orbital setae dorsoclinate; posterior notopleural seta inserted at much
present (anterior seta small). Antenna (Fig. 2) with pedicel higher level than anterior seta; four to five anepisternal setae
lacking suture, bearing long setae dorsally and ventrally, present, two to three dorsoclinate; one katepisternal seta pres-
with process apically or ventroapically; first flagellom- ent. Wing hyaline (Figs. 1, 3); C with humeral and subcostal
ere with posterior cavity, bearing long setae apically; arista breaks; subcosta rudimentary; M thin and weak compared
short, bearing sparse, short rays. Face short, narrow, weakly with R4+5; crossvein dm-cu present or absent; anal cell, cell

1235
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

3 Nannodastia

2 Nannodastia

4 Nannodastia

5 Nannodastia

Figs. 101.2–5. Antenna, wing, and abdomen: lateral view of antenna of (2) Nannodastia horni Hendel, (Australasian/Oceanian, Papp & Mathis,
2001, fig. 16); dorsal view of wing of (3) N. horni, (Papp & Mathis, 2001, fig. 17); lateral view (4) and ventral view (5) of abdomen of male
of N. horni, (Papp & Mathis, 2001, figs. 18, 19).
NANNODASTIIDAE (nannodastiid beach flies) 101 1237

cup, CuA2, A1, and alula absent; wing margin bearing long he, followed by Frey (1958) and Sabrosky & Wirth (1959),
fringe. Abdominal spiracles absent. Male with six exposed treated these genera as members of the family Ephydridae,
tergites (Figs. 4–5); postabdomen symmetrical (Figs. 6–7); tribe Atissini. Papp (1980) described Nannodastiinae as a
cercus weakly sclerotized, bearing long setae; hypandrium subfamily within Ephydridae but qualified his classification
elongate, fused anteriorly with lateral extensions of with the statement that “The majority of the above differences
phallapodeme; phallapodeme fused with elongate phallus; [of Nannodastiinae and other ephydrid subfamilies] separate
pregonites and postgonites absent. Female tergites 7–8 superfamilies in Griffiths’ (1972) system; they seem sufficient
undivided but more strongly sclerotized laterally (Fig.  8); to separate subfamilies here.” Pitkin (1989), as part of the
spermathecae and ventral receptacle apparently absent. Australasian/Oceanian catalog, recognized Nannodastiinae
Nannodastiidae are similar to some Ephydridae (genera as a subfamily in the family Chyromyidae. No synapomorphies
of the tribe Atissini) and Chyromyidae but are distinguished linking these two families have been elaborated, however,
from either family as follows: three reclinate fronto-orbital as the number and position of the fronto-orbital setae are
setae (anterior seta small), none proclinate; Sc incomplete; regarded as a convergence, shared by other families, such
vein M thin and weak compared with R4+5; and lacking cell as the Tethinidae and Acartophthalmidae. Thus, familial
cup, CuA2, A1, and an alula. affiliations remain undocumented except that both genera
lack the synapomorphies that characterize Ephydroidea
(Zatwarnicki, 1992; Mathis  & Zatwarnicki, 1995), and an
Biology affiliation with Chyromyidae seems doubtful.
The two genera included in Nannodastiidae, Azorastia and
Adults, which are extremely small, are usually associated
Nannodastia, are similar externally, as are structures of the
with maritime beaches in tropical and subtropical zones.
male terminalia. Their close resemblance led Sabrosky & Wirth
Specimens were often collected in caves or under cliff
overhangs where the substrate is rich in organic matter from (1959) and Papp (1980) to question their status as distinct
the droppings of bats, small mammals, and birds. Nothing is genera. Certainly the two groups are sister taxa, but until
known about the immature stages, although presumably they their familial placement is resolved, recognition of them as
are in the richly organic substrate. distinct genera is preferred. Although similar, the two genera
are easily diagnosed by external characters and structures of
the male terminalia.
Classification
The two genera comprising this family remain enigmatic Identification
with unknown familial affiliations within the Schizophora
(Opomyzoidea/Carnoidea). The type genus, Nannodastia, Papp & Mathis (2001) provide background for understanding
was first described in the Ephydridae, and Azorastia, the the family and identifying the included genera and species.
only other included genus, in the Asteiidae. Sabrosky (1956) Adults are best preserved dry, and because of their minute
first noted a close relationship between these genera, and size, they should be double mounted.

Key to genera of Nannodastiidae of the world

1. Crossvein dm-cu lacking (Fig. 3); vein M attenuate apically, not extended to wing margin; costal
vein continued past apex of vein R2+3 but not to vein M; male syntergosternite 7+8 much reduced
(Figs. 6–7); epandrium with posteroventral process, bearing thick thornlike structure apically;
anterior portion of surstylus bearing three long setae; posterior portion with digitiform process
with three thornlike setae apically; hypandrium less dorsally placed on epandrium, shorter,
anterior part robust with ventral dilation; female cercus bearing short setulae only (Fig. 8) . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nannodastia Hendel
– Crossvein dm-cu present (Fig. 1); vein M distinct to wing margin; costal vein extended to vein M;
male syntergosternite 7+8 distinct; epandrium lacking posteroventral process; surstyli without
any processes or thorns, medial surface with numerous short setae; hypandrium relatively dorsally
placed on epandrium; hypandrium and phallus long and slender; female cercus bearing at least
one long seta; Palearctic Region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Azorastia Frey]
1238
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Synopsis of the fauna Nannodastia Hendel. Worldwide there are two species in
Nannodastia, one of which, N. atlantica Papp & Mathis, was
The Nannodastiidae now include five species in two genera recently described from specimens collected in Belize and
(Papp  & Mathis, 2001). The New World fauna comprises Barbados (West Indies).
one genus with a single species. No fossils are known.

6 Nannodastia

7 Nannodastia

8 Nannodastia

Figs. 101.6–8. Terminalia: posterolateral view (6) and lateral view (7) of male terminalia of Nannodastia atlantica Papp & Mathis, (Papp &
Mathis, 2001, figs. 12, 13); dorsal view of terminal segments of female abdomen of (8) N. atlantica, (Papp & Mathis, 2001, fig. 15).
NANNODASTIIDAE (nannodastiid beach flies) 101 1239

Literature cited

Frey, R. 1958. Zur Kenntnis der Diptera brachycera p.p. der Pitkin, B.R. 1989. 95. Family Chyromyidae, p. 600. In Evenhuis, N.L.
Kapverdischen Inseln. Societas Scientiarum Fennica, Commen- (editor). Catalog of the Diptera of the Australasian and
tationes Biologicae 18: 1–61. Oceanian Regions. Bishop Museum Press & E.J. Brill, Honolulu &
Griffiths, G.C.D. 1972. The phylogenetic classification of Diptera
Leiden, 1155 pp.
Cyclorrhapha, with special reference to the structure of the male
postabdomen. Series entomologica 8. Dr. W. Junk, The Hague, Sabrosky, C.W. 1956. Additions to the knowledge of the Old
340 pp. World Asteiidae (Diptera). Revue Française d’Entomologie
Mathis, W.N. & T. Zatwarnicki. 1995. A world catalog of the shore 23: 216–243.
flies (Diptera: Ephydridae). Memoirs on Entomology, International Sabrosky, C.W. & W.W. Wirth. 1959. A Formosan ephydrid new
4: vi + 1–423. to Hawaii (Diptera: Ephydridae). Notulae Entomologicae
Papp, L. 1980. New taxa of the acalyptrate flies (Diptera: Tuni- 38: 109–110. (1958)
simyiidae fam.  n., Risidae, Ephydridae: Nannodastiinae
Zatwarnicki, T. 1992. A new classification of Ephydridae based
subfam. n.). Acta Zoologica Hungarica 26: 415–431.
Papp, L.  & W.N. Mathis. 2001. A review of the family Nanno- on phylogenetic reconstruction (Diptera: Cyclorrhapha).
dastiidae (Diptera). Proceedings of the Entomological Society Genus 3: 65–119.
of Washington 103: 337–348.
1240
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
HIPPOBOSCIDAE (louse flies) 102
D. Monty Wood

Fig. 102.1. Female of Ornithomya anchineura Speiser, (MND, fig. 111.1).

Diagnosis larger reclinate bristle. Scutum broad, flat, with scattered

bristles arranged more in transverse patches than in lon-
Small to medium-sized flies (body length 4–12  mm). gitudinal rows (not conforming to usual position in other
Head and body broad, strongly flattened dorsoventally. calyptrates). Notopleuron with single posterior bristle, usu-
Eyes widely separated by broad parallel-sided frons, usu- ally continuous with, and scarcely differentiated from, scutum.
ally large, occupying most of side of head (Figs. 1–3, 5, 6). Ventral surface of thorax flattened; all three thoracic sternites
Ocellar triangle usually distinct; ocelli small or absent. Fron- much wider than their counterparts in other Diptera (except
tal vitta leathery, flanked on each side by reclinate frontal Nycteribiidae and Streblidae). Claws on each tarsus equal,
bristles arising along edge of narrow, shiny, frontal plate. except claw more strongly curved in Lipoptena. Each claw
Inner vertical bristle well developed. Outer vertical bristle bifid in Lipoptena, Melophagus, and Ornithoica (Fig. 22), or
absent. Lunule prominent, bare, usually separated by fissure trifid in remaining genera, because of secondary subdivision
from face, or continuous with face in Olfersia. Parafacial of main claw (Fig. 23). Abdomen of tough, leathery, membra-
and gena reduced. Vibrissa undeveloped. Antenna reduced nous cuticle, capable of considerable extension to accommo-
in size and highly modified, consisting almost entirely date blood meals and/or (in female adults) developing larva
of pedicel, each in socket at side of face, separated medi- but usually shriveled in dried specimens. Both tergites and
ally, or, in Ornithoica, contiguous medially in common pit. sternites of fused first two abdominal segments usually pres-
Scape greatly reduced. Pedicel flattened above, expanded ent, sometimes bearing long bristles. Remaining tergites and
laterally, and hollowed out ventrally to receive minute first sternites greatly reduced, varied in size and shape. Terminalia
flagellomere visible only in anterior view. Antennal seam of of male, unlike those of other calyptrates, fully retracted
pedicel usually prominent in lateral view. Coxae widely sep- within apex of abdomen except during copulation, greatly
arated by expanded katepisternum, legs appear to arise later- modified and simplified in comparison with male terminalia
ally, crablike, rather than ventrally. Postpronotum strongly of most other flies, and consisting almost entirely of median
flattened dorsally, with scattered bristles and at least one tubular phallus, with well-developed phallapodeme, supported

1241
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

i vt s

vrt

fr

scp

ped ped

2 Ornithoica 3 Olfersia 4 Myophthiria

frorb
plt
fr vit

lun
fc
fr s
ped
plp

5 Ornithoctona 6 Icosta 7 Lipoptena

8 Icosta 9 Microlynchia

Figs. 102.2–9. Heads: anterodorsal view of (2) Ornithoica vicina (Walker), (MND, fig. 111.2); (3) Olfersia fumipennis (Sahlberg), (MND,
fig. 111.3); (4) Myophthiria fimbriata (Waterhouse), (Nearctic, MND, fig. 111.4); (5) Ornithoctona erythrocephala (Leach), (MND, fig. 111.5);
(6) Icosta americana (Leach), (MND, fig. 111.6); and (7) Lipoptena depressa (Say), (Nearctic, MND, fig. 111.7, as ssp. pacifica Maa); lateral
view of (8) I. americana, (MND, fig. 111.8); and (9) Microlynchia pusilla (Speiser), (MND, fig. 111.9).
Abbreviations: fc, face; fr, frons; fr s, frontal bristles; frorb plt, fronto-orbital plate; fr vit, frontal vitta; i vt s, inner vertical seta; lun, lunule;
ped, pedicel; plp, palpus; scp, scape; vrt, vertex.
HIPPOBOSCIDAE (louse flies) 102 1243

by troughlike hypandrium bearing paired lateral shaftlike ground, and return to the host, although this has never been
postgonites. Epandrium, cercus, and surstylus rudimentary, actually observed.
and in some species not recognizable. Both males and females of hippoboscids are obligate
Egg and larval stages all passed within female, and blood feeders. Their only free-living existence is as a puparium
dissection is necessary to observe them. Larva without head on the ground, or in some species, as an adult searching for a
capsule and mouthparts. Mouth opening anteriorly on small suitable place to deposit that puparium, or trying to locate a
papilla. Anus opening posteriorly, between and ventral to host. In Central America there are two mammalian ectopara-
posterior spiracles, presumably functional throughout sites, Melophagus ovinus (Linnaeus), the sheep ked, and
larval life. Lateral spiracles (two thoracic and six abdominal) Lipoptena mazamae (Rondani), the deer ked. The latter species
rudimentary in early instars, apparently disappearing by can fly after emergence from the puparium, but after attaining
third instar. Posterior abdominal spiracles well developed, a host it sheds most of its wings and remains permanently
on that host. Adults of Melophagus never develop wings,
on prominent spiracular plate at posterior end of larva.
and the puparia are formed in the host wool. Because sheep
Puparium similar to last larval stage, but more strongly
spend much of their time herded together in close contact,
sclerotized, especially on the spiracular plate. Anterior lines
adult flies can spread easily from host to host as do lice. The
of dehiscence more strongly delineated. remaining species of hippoboscids are obligate ectoparasites
Highly specialized for an ectoparasitic existence in the of birds. They can slip quickly through the feathers by means
feathers or fur of its bird or mammalian hosts, respectively, of their trifid claws and more flattened bodies. By retaining
members of the Hippoboscidae are most likely to be confused their wings throughout life, most species can transfer readily
only with bat flies, Nycteribiidae and Streblidae. Although from host to host by flying. However, species of Myophthi-
species of these two families are much smaller than most ria and Crataerina, parasites of swifts and swallows, which
hippoboscids, the smallest hippoboscid is not much larger should be expected in Central America but have not yet been
than the largest streblid. The nature of the host itself is reported from there, have reduced or aborted wings and must
diagnostic, for hippoboscids do not occur on bats, and bat remain permanently on their hosts.
flies do not attack other mammals or birds, except perhaps Whether birds or mammals were the first hosts of the
bat collectors. Hippoboscids (except Melophagus, with earliest hippoboscids is a subject of controversy, summarized
a relatively large robust species) and most streblids have by Bequaert (1954). His conclusion was that birds served
wings, or remnants thereof, and the head is directed forward, as the original hosts, on the assumption that the genus
while nycteribiids lack all traces of wings, and the head is Ornithoica, an ectoparasite of birds, possesses the simplest
carried bent back over the thorax in a characteristic pose. claws and many other character states that suggest that it is
Hippoboscids have relatively large compound eyes with the most generalized hippoboscid.
hundreds of ommatidia, while those of streblids and nycteribiids Because of the migratory habits of their avian hosts,
consist of only a few ommatidia. The claws of bat flies are many bird-infesting species tend to have wide ranges. Raptors
bifid, like those of Lipoptena and Melophagus, but unlike capturing infested smaller birds may themselves become
other hippoboscids, which have trifid claws. When wings are infested with the hippoboscid ectoparasites of their prey and
subsequently carry them for great distances. Whether these
present, the wing veins of streblids are evenly spaced over
transferred flies continue to feed and produce larvae on the
the wing surface, while those of hippoboscids are crowded
raptors or leave to seek new passerine hosts again is a
into the anterior half or less. subject needing further research. The lack of records south
of Mexico of Ornithomya anchineuria Speiser, one of the
Biology most common Nearctic species on passerines, suggests that
the flies do not remain on their hosts after migration begins,
Immature stages of Hippoboscidae are passed within the or that they do not live long enough to accompany their
female’s vagina, which is highly modified to serve as a uterus migrating hosts. Some species that occur widely in both
North and South America (for example the two genera
for supporting the developing larva. Modified accessory
mentioned above on swifts and swallows), but that have not
glands, the “milk glands”, provide sufficient nourishment,
been recorded in Central America, may actually pass through
and the uterine wall is extensively provided with tracheae to Central America annually on their hosts but have not yet
provide oxygen. A more or less steady diet of blood from the been intercepted. Whether they can and do breed in Central
host is presumably essential for the larva’s nutritional needs America is a matter for further study, for transfer to new
as well as that of the adult. hosts may depend on how gregarious their hosts are while
Members of the Hippoboscoidea are best known, however, overwintering.
for supporting complete larval development entirely within Most species of hippoboscids tend to have some sort of
the female. Only a single larva develops at a time. In some host specificity, which is summarized in the synopses
hippoboscids, the fully developed larva is presumably ejected below. The smallest species, of Ornithoica and Ornithomya,
to fall to the ground to pupate. In Melophagus, the puparium tend to infest passerines, while species of Ornithoctona and
forms in the sheep’s wool. In most bird-feeding species, the Stilbometopa are on larger birds, such as raptors and gallina-
female remains fully winged throughout her life and presumably ceous birds, respectively. Hippoboscids do not appear to be
has the capability to leave the host, deposit her larva on the pests of poultry.
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Classification Lipopteninae (Maa, 1969c, which Bequaert called Melopha-

ginae, a junior synonym of Lipopteninae), with four genera
Between 150 and 200 species, in 19 to 21 genera, are on cervids and other Artiodactyla, most diverse in the Old
known worldwide (Maa, 1969e, 1989; Maa  & Peterson, World; and Ornithomyinae, encompassing all the remaining
1987). Although the monophyly of the Hippoboscidae has genera, distributed worldwide on a variety of birds.
never been challenged, and its sister-group relationship with Maa (1969e) reduced the number of subfamilies to three,
the Glossinidae is now firmly established, the relationships Ornithomyinae, Hippoboscinae, and Lipopteninae, and
among these two families and the Streblidae and Nycteribiidae increased the number of genera to 21. Two of these sub-
have had a long controversial history, outlined in detail families, Ornithomyinae and Lipopteninae, and 12 of the 13
by Bequaert (1954). The four families share intrauterine New World genera, are known from the Neotropical Region.
development of the larva to full maturity and have therefore Both subfamilies, comprising 10 of the 13 genera and
been traditionally grouped as the informal taxon “Pupipara”. encompassing a total of about 25 species, have been recorded
However, Hennig (1952) after a careful examination of all from Central America. Ornithomya has not yet been recorded
relevant details, concluded that the Streblidae and Nycteribi- south of Mexico, but it is also known from Venezuela and
idae, which are presumably sister groups, developed their probably will be found between these points. The existence
“pupipary” independently from the Glossinidae and Hippo- of a species of Crataerina in Colombia and one of Myophthiria
boscidae and that the Pupipara do not form a monophyletic in Venezuela, both from infrequently collected hosts (swallows
group. and swifts, respectively), justifies their inclusion in the
Bequaert (1954) recognized five subfamilies of Hippo- following key.
boscidae: Ornithoicinae (which he considered the most
primitive) with one genus, Ornithoica, distributed world- Identification
wide on a variety of birds; Hippoboscinae, with one Old
World genus, Hippobosca Linnaeus, on mammals (horses, Keys to all of the New World species of Hippoboscidae
bovids, camels, and carnivores) and the ostrich; Alloboscinae, were provided by Bequaert (1954, 1955, 1957). Maa (1969a,
with one species, Allobosca crassipes Speiser, on lemurs in 1969b) has provided more recent treatments of the spe-
Madagascar; Ortholfersiinae, with one genus, Ortholfersia cies of Ornithoctona and Icosta (as Lynchia in Bequaert’s
Speiser, on marsupials, particularly kangaroos, in Australia; treatment).

Key to the genera of Hippoboscidae of Central America

1. Wings of normal shape (2–3 times as long as wide), used for flight (Figs. 11–14) . . . . . . . . . . . 2
– Wings absent, or nonfunctional (aborted, vestigial, or reduced to stumps by the fly itself)
(Figs. 15, 17) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

2. Apical tooth of claw simple, entire claw appearing bifid (Fig. 22); ventral surface of thorax, at
least of mesosternum, with stout posteriorly projecting setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
– Apical tooth of claw deeply cleft, entire claw appearing trifid (Fig. 23); ventral surface of thorax
essentially bare and smooth or with soft recumbent setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

3. Wing with seven distinct, darkly pigmented, longitudinal veins (Fig. 11); eye occupying entire
side of head; inner vertical bristles convergent, meeting at midline, each arising from cylindrical
tubercle (Fig. 2). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ornithoica Rondani
– Wing with only three distinct longitudinal veins (R1, R4+5, and CuA1), the rest colorless and
indistinct (as in Fig.  10); eye small, confined to anterior half of head; inner vertical bristles
convergent but widely separated medially; each arising flush with vertex (Fig. 7). . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lipoptena Nitzsch, in part

4. Wing with vein CuA2 enclosing cell cup (Figs. 11, 14, 16). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
– Wing lacking cell cup (Figs. 12, 13) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

5. Scutellum fringed with ctenidium of 40 or more long, straight, parallel setae (Fig. 18); laterotergite
with foot-shaped lateral projection (Fig. 18, ltg). . . . . . . . . . . . . . . . . . . . Stilbometopa Coquillett
– Scutellum with fewer shorter bristles, not arranged comblike; laterotergite without prominent
projection (Fig. 19). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
HIPPOBOSCIDAE (louse flies) 102 1245

6. Pedicel, as viewed from above, broadened, leaflike, 2–3 times as long as wide; antennae separated
from each other by less than width of one antenna (Fig. 5). . . . . . . . . . . . . . Ornithoctona Speiser
– Pedicel, as viewed from above, narrower, tapering, more triangular than leaflike; antennae
separated from each other by more than width of single antenna (Fig. 1; as in Fig. 6). . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ornithomya Latreille

7. Face (between antennae) greatly enlarged, appearing as forward extension of lunule (from which
it is not clearly demarcated), broader than width of antenna, and shallowly notched apically,
center of notch at level of apices of antennae (Fig. 3). . . . . . . . . . . . . . . . . . . Olfersia Wiedemann
– Face distinctly demarcated from lunule, projecting anteriorly as two slender arms resembling pair
of tusks, each longer than antennae (Fig. 6) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

8. Wing with dm-cu crossvein at least partly present as stub perpendicular to vein CuA1 (Fig.12);
vibrissal angle rounded anteriorly (Fig. 8). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Icosta Speiser
– Wing without dm-cu crossvein (Fig.  13); vibrissal angle produced anteroventrally as angulate
process (Fig. 9). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

Sc R1 R4+5 R2+3 R4+5

M1+2
A1
CuA1 A1+CuA2 CuA1

10 Neolipoptena 11 Ornithoica

12 Icosta 13 Pseudolynchia

dm-cu

14 Ornithomya
bm br
15 Myophthiria
cup
CuA2

A1+CuA2
16 Stilbometopa 17 Crataerina

Figs. 102.10–17. Wings: dorsal view of (10) Neolipoptena ferrisi (Bequaert), (Nearctic, MND, fig. 111.10); (11) Ornithoica vicina (Walker),
(MND, fig. 111.11); (12) Icosta americana (Leach), (MND, fig. 111.12); (13) Pseudolynchia canariensis (Macquart), (MND, fig. 111.13);
(14)  Ornithomya anchineuria Speiser, (MND, fig.  111.14); (15)  Myophthiria fimbriata (Waterhouse), (Nearctic, MND, fig.  111.15);
(16)  Stilbometopa impressa (Bigot), (Nearctic, MND, fig.  111.16); and (17)  Crataerina seguyi Falcoz, (South America, Bequaert, 1954,
fig. 34c).
1246
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

9. Scutellum truncate posteriorly, with row of hair-bearing teeth at each corner (Fig. 20). . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudolynchia Bequaert
– Scutellum rounded posteriorly, without hair-bearing teeth (Fig. 21). . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microlynchia Lutz, Neiva, & Costa Lima

10. Wing absent, represented by small tubercle on side of thorax; halter absent; on domestic sheep. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Melophagus Latreille
– Wing present, but incomplete or aborted; halter present; on deer, swallows, or swifts. . . . . . . . 11

11. Wing represented by its basal veins and ragged bits of membrane, flight no longer possible; claw
bifid (Fig. 22); on deer. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lipoptena Nitzsch, in part
– Wing represented by small rounded pad, or long and narrow, 4 or more times as long as broad
(Figs. 15, 17); claw trifid (Fig. 23); on swallows or swifts, expected but not yet recorded from
Central America. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

12. Wing represented by small rounded pad (Fig. 15); eye small, centered on side of head (suggesting
a streblid) (Fig. 4). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myophthiria Rondani
– Wing of normal length, but too narrow for flight (Fig. 17); eye small, located on anterior half of
head . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Crataerina Olfers

ltg ltg

18 Stilbometopa 19 Ornithoctona

sct

ltg

20 Pseudolynchia 21 Microlynchia

emp

clw
pulv

22 Lipoptena 23 Icosta

Figs. 102.18–23. Scutella and tarsal claws: dorsal view of scutellum of (18) Stilbometopa impressa (Bigot), (Nearctic, MND, fig. 111.19);
(19)  Ornithoctona erythrocephala (Leach), (MND, fig.  111.18); (20)  Pseudolynchia canariensis (Macquart), (MND, fig.  111.24);
(21) Microlynchia pusilla (Speiser), (MND, fig. 111.22); lateral view of fifth tarsomere and tarsal claws of (22) Lipoptena depressa (Say),
(Nearctic, MND, fig. 111.25, as ssp. pacifica Maa); and (23) Icosta americana (Leach), (MND, fig. 111.26).
Abbreviations: clw, claw; epm, empodium; ltg, laterotergite; pulv, pulvillus; sct, scutum.
HIPPOBOSCIDAE (louse flies) 102 1247

Synopsis of the fauna in the New World, in western USA and Venezuela, but not yet
in Central America. They probably have been overlooked
The Central American fauna is poorly collected, although because their hosts are difficult to collect. The two species
at least 27 species have been recorded out of a possible total may be identified using Bequaert’s (1954) key.
of about 40.
Olfersia Wiedemann. All seven known species occur in
Crataerina Olfers. Of a total world fauna of six species the New World, and all are found in Central America. Most
(Maa, 1969e), only one is known from the New World. have been collected from marine fish-eating birds, including
Crataerina seguyi Falcoz occurs on swallows from Colombia frigatebirds, boobies, pelicans, albatrosses, petrels, cormo-
to Argentina; although it has not yet been recorded from rants, and tropicbirds. The pantropical species, O. spinifera
Central America one of its hosts, the Blue and White Swal- (Leach) is specific to seabirds, especially frigatebirds.
low, Notiochelidon cyanoleuca (Vieillot), occurs there and Another widespread species, also found in Europe and
migrates to and from South America. Africa, O.  fumipennis (Sahlberg), is specific to the osprey,
and a third, O. bisulcata (Macquart), is found on New World
Icosta Speiser. This is by far the largest and most vultures.
taxonomically difficult genus of louse flies, with over 50
species (Maa, 1969e). Nine species occur in the New World Ornithoctona Speiser. Of a total of 12 species of this
(Maa, 1969b), and seven have either been recorded in genus worldwide, five have been recorded from the New World.
Central America or in both North and South America and Widespread, they are found on various families of birds; adults
could thus be expected there. This genus is widespread and are perhaps most common on hawks. Maa (1969a) provided
a key to the species.
found mostly on grouse and quail, hawks, and owls, but
I. albipennis (Say) is known only from herons. Maa (1969b) Ornithoica Rondani. This is a large genus of 23 species
provided keys to subgenera and species, as well as to other worldwide, only two of which have been recorded from
closely related genera. the New World, and at least one from Central America
(Bequaert, 1965). They occur on a variety of small birds,
Lipoptena Nitzsch. This is another large genus of nearly mostly passerines. Bequaert (1954) provided a key to these
30 species worldwide, on various ungulates of the families two species.
Bovidae and Cervidae (Maa, 1969e). Only two are known
from the Neotropical Region and only one from Central Ornithomya Latreille. This is another large genus of
America, L.  mazamae Rondani, which occurs from 21 species worldwide that is poorly represented in the New
southeastern USA to Argentina (Maa, 1969d) on deer, World by only three species. Two occur in North America,
Odocoileus spp. and Mazama sp. The genus is most diverse and at least one of these, the common O. anchineuria, extends
in the Old World tropics. south to southern Mexico; the third is confined to South
America. Perhaps the Nearctic flies do not live long enough
Melophagus Latreille. A single species in the New to accompany their hosts all the way south to their wintering
World, M.  ovinus, which was introduced from Europe, is ranges in Central America.
now found wherever domestic sheep are raised. The species
is probably uncommon in Central America because of Pseudolynchia Bequaert. This is a small genus of five
the scarcity of its host, which does not thrive under tropical species with two in the New World, both in Central America
conditions, but has been recorded from Mexico and Panama (Bequaert, 1965). Pseudolynchia canariensis (Macquart),
(Maa, 1969e). Two other species of the genus are known, the cosmopolitan pigeon louse fly, is an immigrant from
both in the Palearctic Region. Europe on the rock dove and is now also found on native
Microlynchia Lutz, Neiva,  & Costa Lima. This is a doves, and P. brunnea (Latreille), specific to caprimulgids, is
small genus of only four species, all occurring in the New widespread in the New World.
World. Three have been recorded in Central America, taken
Stilbometopa Coquillett. There are five species of
mostly from tinamous, doves, and perching birds. Bequaert
(1955: p. 364) provided a key to species. this genus, found only in the New World, four of which
are recorded from Central America; the fifth, the Nearctic
Myophthiria Rondani. A small genus of 10 species in the S. impressa (Bigot), extends south into Sonora, Mexico, on
world, these flies are highly modified as ectoparasites of swifts California quail. They live on various families of birds,
(Apodidae) and distinctive in the family because of their small including doves, quail, Cracidae (Galliformes), trogons,
eyes and vestigial wings. Only two species have been collected toucans, and perching birds.
1248
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Literature cited

Bequaert, J.C. 1954. The Hippoboscidae or louse-flies (Diptera) of Maa, T.C. 1969a. Synopses of the genera Ornithophila and Orni-
mammals and birds Part II. Taxonomy, evolution and revision of thoctona with remarks on their habitat diversification (Diptera:
American genera and species [part]. Entomologica Americana Hippoboscidae). Pacific Insects Monograph 20: 1–23.
34 (new series): 1–232. Maa, T.C. 1969b. Revision of Icosta (= Lynchia auctt.) with erection
Bequaert, J.C. 1955. The Hippoboscidae or louse-flies (Diptera) of of a related genus Phthona (Diptera: Hippoboscidae). Pacific
mammals and birds Part II. Taxonomy, evolution and revision of Insects Monograph 20: 25–203.
American genera and species [part]. Entomologica Americana Maa, T.C. 1969c. Further notes on Lipopteninae (Diptera: Hippo-
35 (new series): 233–416. boscidae). Pacific Insects Monograph 20: 205–236.
Bequaert, J.C. 1957. The Hippoboscidae or louse-flies (Diptera) of Maa, T.C. 1969d. Notes on the Hippoboscidae (Diptera). II. Pacific
mammals and birds Part II. Taxonomy, evolution and revision of Insects Monograph 20: 237–260.
American genera and species [part]. Entomologica Americana Maa, T.C. 1969e. A revised checklist and concise host index
36 (new series, 1956): 417–611. of Hippoboscidae (Diptera). Pacific Insects Monograph 20:
Bequaert, J.C. 1965. Family Hippoboscidae. In Stone, A. et al. 261–299.
(editors). A catalog of the Diptera of America north of Mexico. Maa, T.C. 1989. Family Hippoboscidae, pp. 785–789. In Evenhuis,
United States Department of Agriculture, Agriculture handbook N.L. (editor). Catalog of the Diptera of the Australasian and
276, 1696 pp. Oceanian Regions. Bishop Museum Press & E.J. Brill, Honolulu &
Hennig, W. 1952. Die Larvenformen der Dipteren. Eine Übersicht Leiden, 1155 pp.
über die bisher bekannten Jugendstandien der zweiflügeligen Maa, T.C. & B.V. Peterson. 1987. Hippoboscidae, pp. 1271–1281.
Insekten. 3. Teil. Adademie-Verlag, Berlin, 628 pp. MND, Volume 2.
STREBLIDAE (bat flies) 103
Carl W. Dick and Jeremy A. Miller

Fig. 103.1. Female of Speiseria ambigua Kessel, (Jobling, 1939a, fig. 1A, as Synthesiostrebla amorphochili Townsend).

1249
1250
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Diagnosis perspicillata (Linnaeus), vacate their roosts to forage (Fritz,

1983). Although bat flies feed frequently, some authors have
Small, tan to light brown flies (Fig.  1), body length found little evidence of disturbance to the host (Wenzel et al.,
generally 1.5–2.5  mm, but some exceptional species as 1966; Overal, 1980). However, Linhares & Komeno (2000)
small as 0.73 mm (e.g., Mastoptera minuta Costa Lima) or correlated significant weight loss in Sturnira lilium (Geoffroy)
as large as 5.50 mm (e.g., Joblingia schmidti Dybas & Wenzel). with parasitism by Megistopoda proxima (Séguy).
Females generally slightly larger than males, especially in Streblid bat flies tend to be host specific, with one fly
Mastoptera; sexual dimorphism in other, nonreproductive species typically parasitizing a single host species or a few
structures rare. Head, body, and legs generally densely closely related host species (Wenzel et al., 1966; Wenzel,
hirsute with body form varying from strongly laterally 1976; Dick  & Patterson, 2007). Distributional evidence
compressed (e.g., Nycterophiliinae) to dorsoventrally flattened suggests that they cospeciate with host species of bat,
(e.g., Streblinae) to uncompressed (e.g., most Trichobiinae). producing parallel phylogenies of host and parasite (Patterson
Eyes generally reduced. Ocelli absent. Head of some et al., 1998). It is notable that streblids are often absent from
species (e.g., Streblinae) with ventral ctenidium. Thorax bat species that roost solitarily or in temporary roosts
varied in shape, longitudinal median suture and transverse (Wenzel & Peterson, 1987). Conversely, bat fly species richness,
scutal suture various. Legs varied in length, thickness, setal along with the intensity and prevalence of parasitism, seems
form, and chaetotaxy, ranging from short and stout to elongate. generally to increase with roost size, duration of roost sites,
Tibiae without apical spurs and claws usually large and and the number of bats roosting there (Komeno & Linhares,
simple. Abdomen extensively membranous, corresponding 1999; Patterson et al., 2007). The dynamics of bat roosts as
to a reduction or loss of sclerites. Macropterous, stenopterous, they pertain to the biology, ecology, and host specificity of
brachypterous, or apterous. streblid bat flies is a critically understudied area. Bat host
Streblids are distinguished from hippoboscids by their species often harbor more than one species of streblid. This
reduced compound eyes, with number of facets ranging from may be related to the observation that some bat flies prefer
1 to 36. Most Hippoboscidae have well-developed, horizontally particular areas (microhabitats; e.g., fur or wing membranes)
elongate compound eyes with ocelli present. Streblids are on the host’s body (Ross, 1961; Wenzel et al., 1966; Fritz,
distinguished from nycteribiids by the latter having their 1983; A.X.  Linhares  & C.A.  Komeno, 2000; personal
head folded back when not feeding and by the presence of a observations). For more information regarding the ecology,
ctenidium on the ventral surface of the abdomen. Furthermore, distribution, and host associations of streblid bat flies, see
all streblids except Paradyschiria possess wings and a largely papers by Wenzel & Tipton (1966), Kunz (1976), Reisen et
sclerotized thoracic dorsum. al. (1976), Marshall (1980, 1981, 1982), Gannon & Willig
(1995), Komeno & Linhares (1999), Guerrero (1997), Lin-
hares & Komeno (2000), Graciolli & Dick (2004), Dick &
Biology Patterson (2006, 2007), and Patterson et al. (2007).
Streblids are obligate blood-feeding ectoparasites of bats.
Adult females of the Old World genus Ascodipteron Adensamer Classification
are the only exception in that they embed into the skin of
their host and lose certain morphological structures, becoming Streblids belong to the monophyletic superfamily
effectively endoparasitic (Audy et al., 1972). The life cycle Hippoboscoidea, along with the families Glossinidae, Hippo-
of bat flies seems well adapted to facilitate their ectoparasitic boscidae, and Nycteribiidae (McAlpine, 1989; Yeates  &
habits. Like all hippoboscoid flies, streblids are viviparous. Wiegmann, 1999). Five subfamilies of Streblidae are rec-
The life cycle of Megistopoda aranea (Coquillett) (Tricho- ognized (Wenzel et al., 1966; Wenzel, 1976; Dick & Patterson,
biinae), a parasite of Artibeus jamaicensis Leach (Phyllos- 2006). Exclusive to the Old World are the Brachytarsininae
tomidae), has been described by Overal (1980). Larvae of (4 genera, 52 species) and the Ascodipterinae (3 genera, 21
M. aranea develop one at a time, nourished by intrauterine species). Exclusive to the New World are the subfamilies
“milk” glands within the female (adenotrophy sensu Hagan, Nycterophiliinae (2 genera, 6 species), Trichobiinae (20 gen-
1951). Female M.  aranea produce no more than one larva era, 115 species), and Streblinae (4 genera, 35 species). Some
every 10 days. In order to larviposit, the female leaves the workers suggest that one subfamily (Nycterophiliinae) as
host briefly and finds a suitable location to attach her larva recognized currently exhibits morphological variation that
(most often within the host’s roost). Almost immediately after warrants familial status (R.L.  Wenzel, personal communi-
the larva is deposited, the cuticle hardens and a puparium is cation), while others argue that streblids should be placed
formed. After some 22–24 days, the teneral adult emerges within the Hippoboscidae (F.C.  Thompson, personal com-
and must locate a host and take a blood meal before breed- munication). Nonetheless, a robust and comprehensive phy-
ing (Ross, 1961; Overal, 1980). Larviposition in Trichobius logeny is lacking for the Streblidae. Although earlier workers
joblingi Wenzel, Speiseria ambigua Kessel, and Strebla suggested that the Streblidae was monophyletic (McAlpine,
guajiro (Garcia & Casal) normally occurs between 1730 and 1989), recent molecular analysis suggests paraphyly for the
2300 h, coinciding with the time when their hosts, Carollia Streblidae, divisible into an Old World clade allied with the
STREBLIDAE (bat flies) 103 1251

family Nycteribiidae, and a separate New World clade (Ditt- can be made using a dissecting microscope. Species-level
mar et al., 2006). Important references concerning streblid identifications may often be made using low magnification;
taxonomy, classification, and systematics include Jobling however, some species (especially some Trichobius) are best
(1929, 1936), Theodor (1964), Hennig (1965, 1971), Wen- identified using slide-mounted specimens and a compound
zel et al. (1966), Schlein (1970), Griffiths (1972), Wenzel microscope. For techniques on slide-mounting streblids,
(1976), McAlpine (1989), Yeates & Wiegmann (1999), Nir- refer to Wenzel et al. (1966: pp. 411–414).
mala et al. (2001), and Dittmar et al. (2006). The key below will facilitate identification of all New
World genera of Streblidae, including the 23 genera known
from Central America and three genera known only from
Identification South America. Other generic keys are available in Wenzel
(1976), Guerrero (1993), and Graciolli & Carvalho (2001).
The adult taxonomy of New World streblid genera relies
primarily on overall body form, the presence or absence of The latter contains a key to the 10 genera known from Paraná
a ctenidium along the posteroventral margin of the head, state in Brazil. The single, most comprehensive key to New
gross characteristics of the wings, leg chaetotaxy, details of World streblid species is Wenzel (1976). For species-level
the thorax, especially the mesonotal and transverse sutures, keys in Spanish see Guerrero (1993, 1994a, 1994b, 1995a,
and male reproductive structures. Generic identification 1995b, 1996).

Key to the genera of Streblidae of the New World

1. Body laterally compressed, flealike; wings, if present, with most veins weakly developed. . . . . 2
– Body not laterally compressed; compressed dorsoventrally or not at all; wings, if present, with
well-developed longitudinal veins. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

2. Macropterous; eye consists of single large facet. . . . . . . . . . . . . . . . . . . . . . . Nycterophilia Ferris

– Wings reduced to small flaps; eye absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . Phalconomus Wenzel

3. Head with comb of closely spaced, thick, spinelike setae (ctenidium) along posteroventral margin
(Figs. 2–3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
– Head without ctenidium along posteroventral margin (Figs. 4–7) . . . . . . . . . . . . . . . . . . . . . . . . 8

4. Ctenidium consisting of only 18–19 spinelike setae, restricted to posteroventral margin of head
(Fig. 2). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eldunnia Curran
– Ctenidium consisting of numerous spinelike setae (at least 30), extending around sides to
dorsolateral parts of head (Figs. 3, 8–11). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

5. Broad scale present on side of head behind ctenidium (Figs. 8–9). . . . . . . . . . . . . . . . . . . . . . . . 6

– Broad scale absent from side of head behind ctenidium (Figs. 10–11). . . . . . . . . . . . . . . . . . . . . 7

6. Brachypterous (as in Figs. 15–20); hind legs shorter than or subequal to body . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Metelasmus Coquillett
– Macropterous (as in Figs. 1, 21–23); hind legs longer than body. . . . . . . . . . . Anastrebla Wenzel

7. Hind tibia with two to eight macrosetae conspicuously longer than others on dorsal surface
(Figs. 12a–f); hind leg shorter than or subequal to body. . . . . . . . . . . . . . . . . . Strebla Wiedemann
– Hind tibia with numerous short setae, lacking conspicuous macrosetae longer than others on
dorsal surface; hind leg conspicuously longer than body . . Paraeuctenodes Pessôa & Guimarães

8. Brachypterous, stenopterous (Figs. 13b, 14), or apterous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

– Macropterous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
1252
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

pgn
plp
premnt plp
fc

fr

pgn
eye
2 Eldunnia
lbl

vrt ocp

3 Metelasmus
fr ocp
4 Trichobius
ped
flgm

ar
premnt fr

plp pgn
eye
rst memb

6 Trichobius
5 Trichobius
ocp vrt
plp gn
7 Trichobius
fc
ant gn pgn
eye
pgn
cten cten
pgn
brd sc

brd sc
ocp

9 Anastrebla
8 Metelasmus

10 Strebla

c d e
b f

a
11 Strebla 12 Strebla
Figs. 103.2–12. Heads and hind tibiae: ventral view of hind margin of head, showing ctenidium, of (2) Eldunnia breviceps Curran, (Jobling,
1949a, fig.  3B); ventral view of head of (3)  Metelasmus pseudopterus Coquillett, (Jobling, 1936, fig.  3B); dorsal  (4), ventral  (5),
and lateral (6) views of head of Trichobius sphaeronotus Jobling (Zeve & Howell, 1962, figs. 3, 9, 6); dorsal view, setae omitted, of head
of (7) Trichobius sp., (Jobling, 1936, fig. 1E); dorsal view of head of (8) M. pseudopterus, (Jobling, 1936, fig. 3A); dorsal view, palpi and
face (as frontoclypeal area) omitted, of head of (9) Anastrebla sp., (Wenzel et al., 1966, fig. 39D); dorsal view of head of (10) Strebla
guajiro (Garcia & Casal), (Wenzel et al., 1966, fig. 122A, as S. carolliae); and (11) S. hertigi Wenzel, (Wenzel et al., 1966, fig. 122B); dorsal
view of hind tibia of (12a) S. consocia Wenzel, (South America, Trinidad), (Wenzel et al., 1966, fig. 125A); (12b) S. wiedemanni Kolenati,
(Wenzel et al., 1966, fig. 125B, as S. vespertilionis), (12c) S. diaemi Wenzel, (Wenzel et al., 1966, fig. 125C); (12d) S. galindoi Wenzel,
(Wenzel et al., 1966, fig. 125D); (12e) S. mirabilis (Waterhouse), (Wenzel et al., 1966, fig. 125E); and (12f) S. hertigi Wenzel, (Wenzel et al.,
1966, fig. 125F).
Abbreviations: ant, antenna; ar, arista; brd sc, broad scale; cten, ctenidium; fc, face (frontoclypeus); flgm, flagellomere; fr, frons; gn, gena;
lbl, labella; ocp, occiput; ped, pedicel; pgn, postgena; plp, palpus; premnt, prementum; rst memb, rostral membrane; vrt, vertex.
STREBLIDAE (bat flies) 103 1253

b
b
a
a

13 Neotrichobius 14 Megistopoda 15 Mastoptera

a a
a

16 Joblingia 17 Aspidoptera 18 Noctiliostrebla

Figs. 103.13–18. Adults and detached wings: dorsal view of adult (a) and right wing (b) of (13) of Neotrichobius stenopterus Wenzel &
Aiken, (Wenzel et al., 1966, figs. 97, 99C); right wing of (14) Megistopoda proxima Séguy, (Séguy, 1926, fig. 5); dorsal view of adult (a)
and right wing  (b) of (15)  Mastoptera guimaraesi Wenzel, (Jobling, 1949b, figs.  4A, C); dorsal view of adult  (a) and right wing  (b) of
(16) Joblingia schmidti Dybas & Wenzel, (Dybas & Wenzel, 1947, figs. 25A, B); dorsal view of adult (a) and left wing (b) of (17) Aspidoptera
phyllotomatis (Perty), (Jobling, 1949b, figs. 1A, C); and (18) Noctiliostrebla aitkeni Wenzel, (South America, Jobling, 1949b, figs. 3A, C, as
Aspidoptera megastigma Speiser).
1254
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

a
c

19 Exastinion 20 Anatrichobius 21 Paratrichobius

fr plp

22 Synthesiostrebla 23 Stizostrebla 24 Parastrebla

Figs. 103.19–24. Adults and detached wings (concluded) and apex of female abdomen: dorsal view of female with extruded larva (a), right
wing (b), and dorsal view of head (c), setae omitted, of (19) Exastinion clovisi Pessôa & Guimarães, (Jobling, 1949b, figs. 2A, D; Wenzel et
al., 1966, fig. 105E); dorsal view of female (a) and right wing (b) of (20) Anatrichobius scorzai Wenzel, (Wenzel et al., 1966, fig. 76); dorsal
view of adult of (21) Paratrichobius longicrus Miranda-Ribeiro, (Jobling, 1939a, fig. 2C); (22) Synthesiostrebla amorphochili Townsend,
(South America, Jobling, 1947, fig. 1A); and (23) Stizostrebla longirostris Jobling, (South America, Jobling, 1939b, fig. 2A); ventral view of
apex of abdomen of (24) Parastrebla handleyi Wenzel, (Wenzel et al., 1966, fig. 116B).
Abbreviations: plp, palpus; fr, frons.
STREBLIDAE (bat flies) 103 1255

9. Wings absent; thorax with membranous dorsal regions. . . . . . . . . . . . . . . . Paradyschiria Speiser

– Brachypterous or stenopterous, occasionally wings minute; thorax fully sclerotized. . . . . . . . . 10

10. Stenopterous; thorax shieldlike in ventral view, posterior margin broadly rounded; hind leg more
than 1.5 times length of body. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
– Brachypterous; thorax lacking broadly rounded posterior margin in ventral view, approximately
as broad as anterior margin; hind leg at most 1.2 times longer than body. . . . . . . . . . . . . . . . . . 13

11. Forefemur with inner face armed with stout spinelike setae (Fig. 13a); wing with only two or three
longitudinal veins (Fig. 13b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neotrichobius Wenzel & Aitken
– Forefemur with inner face lacking stout spinelike setae; wing with four or more longitudinal veins
(Fig. 14). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

12. Wing broad, with seven veins; thorax not laterally compressed, dorsal surface only slightly
convex, transverse suture distinct at lateral margins; males with postgonites massive, more than
one-half body length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Megistapophysis Dick & Wenzel
– Wing narrow, with four to six veins; thorax laterally compressed; dorsal surface strongly convex;
transverse suture, if complete, indistinct at lateral margins; males with postgonites of normal size,
less than one-quarter body length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Megistopoda Macquart

13. Median mesonotal suture extending posteriorly beyond transverse suture to scutellum (Figs. 15a,
16a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
– Median mesonotal suture not extending posteriorly beyond transverse suture (Figs. 17a, 18a, 19a,
20a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

14. Minute flies, body less than 1.5 mm long; wing venation indistinct (Fig. 15b). . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mastoptera Wenzel
– Large flies, body more than 3.5 mm long; wing venation distinct (Fig. 16b). . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Joblingia Dybas & Wenzel

15. Median mesonotal suture meeting transverse suture at obtuse angle to form inverted Y (Fig. 18a).
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Noctiliostrebla Wenzel
– Median mesonotal suture meeting transverse suture at right angle to form inverted T (Figs. 17a,
19a, 20a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

16. Posterior margin of occipital lobes flaplike (Fig. 19c); median mesonotal suture bifurcate anteriorly
(Fig. 19a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Exastinion Wenzel
– Posterior margin of occipital lobes rounded (Fig.  7); median mesonotal suture not bifurcate
anteriorly (Figs. 17a, 20a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

17. Abdomen clothed with short setae dorsally (Fig. 17a); palpus nearly vertical. . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aspidoptera Coquillett
– Abdomen clothed with long setae dorsally (Fig. 20a); palpus nearly horizontal. . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anatrichobius Wenzel

18. Forefemur with inner face armed with stout spinelike setae (Figs. 13a, 21); hind leg conspicuously
longer than body (Fig. 21); thorax shieldlike in ventral view, posterior margin broadly rounded,
but not smoothly so . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paratrichobius Costa Lima
– Forefemur with inner face lacking stout spinelike setae (Fig. 1, 23); hind leg may or may not be
conspicuously longer than body; thorax lacking broadly rounded margins in ventral view,
approximately as broad as anterior margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
1256
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

19. Anterodorsal margin of thorax with two anteriorly directed teeth; complimentary grooves present
on posterior margin of head (Fig. 22); Peru. . . . . . . . . . . . . . . . . . . . [Synthesiostrebla Townsend]
– Anterodorsal margin of thorax flat or with pair of blunt projections, without sharp teeth; without
complimentary grooves on posterior margin of head. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

20. Hind leg conspicuously longer than body (Fig. 1). . . . . . . . . . . . . . . . . . . . . . . . . Speiseria Kessel
– Hind leg shorter than or subequal to length of body. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

21. Hind tibia with all setae nearly uniform in length. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

– Hind tibia with some setae conspicuously longer than others on dorsal surface. . . . . . . . . . . . . 23

22. Costa and R1 with strong macrosetae to point where they join distally; palpus with marginal setae
only, ventral surface bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trichobioides Wenzel
– Costa and R1 lacking such strong macrosetae; palpus with setae along margins and on ventral
surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trichobius Gervais

23. Posterior margins of occipital lobes extending posteriorly, partially covering anterolateral region
of thorax in dorsal view; transverse suture incomplete; mesonotal suture absent (Fig. 23); South
America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Stizostrebla Jobling]
– Posterior margins of occipital lobes not partially covering thorax in dorsal view; transverse suture
complete; mesonotal suture present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

24. CuA1 united with dm-cu in arc, uniting with M1 before wing apex; transverse suture distinctly
recurved; South America. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Xenotrichobius Wenzel]
– CuA1 not united with dm-cu; transverse suture nearly straight. . . . . . . . . . . . . . . . . . . . . . . . . . 25

25. Head more or less round, narrower than anterior margin of thorax; venter of female abdomen with
subapical plate bearing transverse row of blunt setae (Fig. 24). . . . . . . . . . . . Parastrebla Wenzel
– Head distinctly broader than long, about as wide as anterior margin of thorax; venter of female
abdomen without transverse row of blunt setae. . . . . . . . . . . . . . . . . . . Pseudostrebla Costa Lima

Synopsis of the fauna detailed the fauna of Venezuela. This treatment of the bat flies
of the Smithsonian Venezuelan Project was based on more
Ninety-nine described species of Streblidae are known than 36 000 streblid specimens and included descriptions of
to occur in Central American countries, including tropical two new genera and 45 new species. Wenzel et al. (1966)
regions of southern Mexico. The occurrence of streblid and Wenzel (1976) often noted additional material collected
species corresponds closely with that of their host bats. Given from Central American countries. Guerrero (1997) sum-
the lack of sampling in many Central American regions, the marized a great deal of information regarding New World
presence of several undescribed species in museum collections, streblids, including a list of countries and the streblid species
and the fact that some currently recognized species probably that have been reported from each. Tschapka and co-workers
represent complexes, we estimate the total number of assembled a collection of 492 records (each record constitutes
Streblidae in Central America and southern Mexico to be one species of bat fly collected from one host individual) from
approximately 125 species. Costa Rica (Miller & Tschapka, 2001). Finally, the first au-
The following information regarding the 23 streblid thor extracted a great deal of information from the streblid
genera known from Mexico and Central America has been collection housed at the Field Museum of Natural History,
compiled from six major sources. Wenzel et al. (1966) Chicago. Those collections (not including the Panamanian ma-
provided a comprehensive catalog of the Panamanian fauna, terial treated by Wenzel et al., 1966) include almost 3350
based on approximately 12  000 specimens, with descrip- records of Streblidae from Mexico and Central America.
tions of many new taxa including one subfamily, eight
genera, and 51 species. Wenzel’s (1970) catalog treated all Anastrebla Wenzel. This genus is composed of five
New World streblids known to occur south of the United described species, three of which occur in Central Ameri-
States. Wenzel (1976) was similar to Wenzel et al. (1966), but ca (Wenzel et al., 1966). Species of Anastrebla typically
STREBLIDAE (bat flies) 103 1257

are associated with phyllostomid bats of the subfamily Megistapophysis Dick & Wenzel. This monotypic genus
Glossophaginae. is known only from Costa Rican Sturnira mordax (Goodwin)
(Phyllostomidae: Stenodermatinae), a species of bat known
Anatrichobius Wenzel. This genus is composed of two from higher elevations in Costa Rica and Panama (Matson &
described species. Anatrichobius scorzai Wenzel has been McCarthy, 2004). Currently, no specimens of Megistapophysis
reported from Costa Rica, Honduras, and Panama (Wenzel, mordax Dick & Wenzel are known from Panama, but it is
1976; Guerrero, 1997). Graciolli (2003) described a second
likely that more collecting effort will document the species
species from Brazil, and more undescribed species are known
there.
to exist (B.V. Peterson, personal communication; Guerrero,
1995b). Anatrichobius is hypertrichous, more so than any Megistopoda Macquart. This genus is composed of two
other streblid, and is associated with vespertilionid bats of described species, both known from most of Central America.
the genus Myotis Kaup. Species are almost wholly restricted to phyllostomid bats of
Aspidoptera Coquillett. Aspidoptera is composed of the genera Artibeus (sensu stricto) and Sturnira. The taxonomy
three described species, all of which are known from Mexico of the genus is confused and in need of revision, and
and most Central American countries (Wenzel et al., 1966; undescribed species have been recognized (Wenzel, 1976).
Wenzel, 1970; Guerrero, 1997). Aspidoptera is restricted to The undescribed species of Megistopoda from Costa Rican
phyllostomid bats of the genera Artibeus Leach (sensu stricto) Sturnira mordax noted by Timm et al. (1989) are in fact
and Sturnira Gray (Stenodermatinae). specimens of Megistapophysis.

Eldunnia Curran. The single described species, Eldunnia Metelasmus Coquillett. Two described species belong
breviceps Curran, is known only from Panama (Wenzel et al., to this genus, and one, Metelasmus pseudopterus Coquillett,
1966) and Costa Rica (Miller & Tschapka, 2001). The typical has been recorded from Mexico to Panama (Wenzel et al.,
host appears to be the phyllostomid Lonchophylla robusta 1966) and south to northern Argentina and southern Brazil
Miller (Glossophaginae). It is noteworthy that the range of (Graciolli & Dick, 2004). The second species, Metelasmus
E. breviceps is much more restricted than that of L. robusta, wenzeli Graciolli  & Dick, a parasite of Sturnira lilium, is
which ranges from southeastern Nicaragua (Reid, 1997) to known only from southern Brazil and eastern Paraguay
Ecuador, Colombia, and Venezuela (Koopman, 1982). This (Graciolli  & Dick, 2004). Undescribed species also are
cannot be wholly attributed to undersampling bias because known from Guatemala and northern South America, associated
Venezuela has been sampled extensively for bat ectoparasites, with species of Sturnira.
and yet E.  breviceps has not been reported from there
(Wenzel, 1976). Neotrichobius Wenzel & Aitken. Four described species
belong to this genus, three of which are known from Central
Exastinion Wenzel. This genus is composed of three America. An undescribed species of Neotrichobius has been
described species, two of which are known from Central collected from Mesophylla macconnelli Thomas at La Selva,
America (Wenzel et al., 1966; Guerrero, 1997). Flies of this
Costa Rica (Miller  & Tschapka, 2001) and in Ecuador.
genus typically are associated with bats of the phyllostomid
genus Anoura Gray (Glossophaginae). Moreover, Neotrichobius delicatus Machado-Allison, known
from Central America, may be a complex of species (Wen-
Joblingia Dybas  & Wenzel. The single described species, zel, 1976). Neotrichobius  spp. are associated typically with
Joblingia schmidti Dybas & Wenzel, has been reported from phyllostomid bats of the subfamily Stenodermatinae and
Guatemala (Dybas & Wenzel, 1947), Panama (Wenzel et al., with Rhinophylla pumilio Peters (Rhinophyllinae).
1966), and Costa Rica. The largest of all streblids (3.96–
5.50  mm total length; Wenzel et al., 1966), it is exclusively Noctiliostrebla Wenzel. Noctiliostrebla is composed
associated with vespertilionid bats of the genus Myotis. of four recognized species, two of which are known from
An undescribed species is known from southern Brazil Central America (Wenzel et al., 1966). Species of this genus
(G. Graciolli, personal communication). are wholly restricted to bats of the genus Noctilio Linnaeus
(Noctilionidae).
Mastoptera Wenzel. Two described species belong to
this genus, and both are recorded from Central America. The Nycterophilia Ferris. This genus is composed of five
smallest of all streblids (0.73–1.29 mm total length; Wenzel described species, which are characteristic parasites of bats
et al., 1966), Mastoptera is restricted to phyllostomid bats of of the family Mormoopidae. Species are known from Mexico
the subfamily Phyllostominae. According to Wenzel (1976), and as far north as Sonora, south to Colombia and Venezuela
Mastoptera is complex taxonomically, particularly those (Wenzel, 1976). Perhaps the most widespread and commonly
forms referred to M.  minuta Costa Lima. An undescribed collected species is Nycterophilia parnelli Wenzel, a
species is associated with Tonatia brasiliense (Peters) at characteristic parasite of the widely distributed bat Pteronotus
La Selva, Costa Rica (Miller & Tschapka, 2001). parnellii (Gray) (Wenzel et al., 1966; Guerrero, 1997).
1258
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Paradyschiria Speiser. This genus is composed of six Pseudostrebla Costa Lima. This genus is composed of
recognized species, four of which are known to occur in three described species, two of which are known from Cen-
Central America (Wenzel, 1976; Guerrero, 1997). Species tral America (Wenzel et al., 1966). Species of Pseudostrebla
of this genus are completely restricted to bats of the genus are associated with phyllostomid bats of the genus Tonatia
Noctilio. Paradyschiria spp., the only wingless streblids, co- Gray (Phyllostominae).
occur with vestigial-winged Noctiliostrebla   spp. on hosts
that possess the shortest hair of any New World bat species. Speiseria Kessel. Speiseria is composed of three de-
scribed species, two of which are known from Central
Paraeuctenodes Pessôa  & Guimarães. Two described America. In Central America, flies of this genus typically
species belong to this genus. Paraeuctenodes longipes are associated with phyllostomid bats of the genus Carollia
Pessôa  & Guimarães is known from Central America and Gray (Carolliinae).
typically is associated with the phyllostomid bat Glossopha-
ga soricina (Pallas) (Glossophaginae). A single specimen of Strebla Wiedemann. This genus is composed of 25 de-
Paraeuctenodes similis Wenzel has been collected in Puntar- scribed species, second in number only to Trichobius. Four-
enas, Costa Rica, although this species appears to be more teen species of Strebla are known from Central America and
common in Brazil, Colombia, and Venezuela (Wenzel, 1976; southern Mexico. An undescribed species from Costa Rica
Guerrero, 1997). has been collected from Hylonycteris underwoodi Thomas
(R.L. Wenzel, personal communication).
Parastrebla Wenzel. The single described species Paras-
trebla handleyi Wenzel has been reported only from Pana- Trichobioides Wenzel. The single described species
ma. The typical host species, Trinycteris nicefori (Sanborn) Trichobioides perspicillatus (Pessôa  & Galvão) is wide-
(Wenzel et al., 1966), is widespread in Central America. It is spread in Central and South America and typically is asso-
likely that more sampling will broaden the known range of ciated with Phyllostomus discolor Wagner (Phyllostomidae:
P. handleyi. Phyllostominae).

Paratrichobius Costa Lima. Five described species be- Trichobius Gervais. The genus Trichobius contains 68
long to this genus, all of which occur in Central America. All described species and is by far the most diverse streblid ge-
are associated with phyllostomid bats of the subfamily Steno- nus (Guerrero, 1994a, 1995a, 1998; Guerrero  & Morales-
dermatinae. Several undescribed species are known (Wenzel, Malacara, 1996). Forty described Trichobius species occur
1970), and a generic revision is needed (Wenzel, 1976). in Central America. Undescribed species have been reported
from Costa Rica (Timm et al., 1989; Miller  & Tschapka,
Phalconomus Wenzel. Although there is only one described 2001). Members of this genus parasitize a wide variety of
species within this genus, Wenzel (1976) mentioned the pres- bats, including representatives of the families Emballonu-
ence of two undescribed species. Phalconomus puliciformis ridae, Furipteridae, Molossidae, Mormoopidae, Natalidae,
(Wenzel) is known only from Lonchophylla robusta (Phyllos- Phyllostomidae, and Vespertilionidae. The genus Trichobi-
tomidae: Glossophaginae) in Venezuela (Wenzel, 1976; Guer- us may not represent a monophyletic group (Wenzel et al.,
rero, 1997). One of the undescribed species (“Phalconomus 1966; G. Graciolli, personal communication) and is in need
species A”; Wenzel, 1976) is known from Guatemala. of revision.

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Griffiths, G.C.D. 1972. The phylogenetic classification of Diptera Streblidae of Brazil. Arbeiten über morphologische und taxono-
Cyclorrhapha, with special reference to the structure of the male mische Entomologie 6: 268–275.
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parásitos de murciélagos (Mammalia: Chiroptera) del nuevo Koopman, K.F. 1982. Biogeography of the bats of South America.
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15: 1–27. Kunz, T.H. 1976. Observations on the winter ecology of the bat fly
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Reisen, W.K., M.L. Kennedy, & N.T. Reisen. 1976. Winter ecology Wenzel, R.L. 1976. The streblid batflies of Venezuela (Diptera:
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Schlein, Y. 1970. A comparative study of the thoracic musculature of the V.J. Tipton (editors). Ectoparasites of Panama. Field Museum
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Séguy, E. 1926. Diptères exotiques peu connus. Encyclopédie
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Theodor, O. 1964. On the relationships between the families of the
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Parasitology 2: 999–1000. Panama. Field Museum of Natural History, Chicago, 861 pp.
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C.S. Vaughan. 1989. Mammals of the La Selva–Braulio Carrillo toward a higher-level phylogeny of the Diptera. Annual Review
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Wenzel, R.L. 1970. 100. Family Streblidae. In Papavero,  N. Zeve, V.H.  & D.E. Howell. 1962. The comparative external
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Agricultura, São Paulo, 25 pp. Entomological Society of America 55: 685–694.
NYCTERIBIIDAE (bat flies, spider bat flies) 104
Gustavo Graciolli

Fig. 104.1. Female of Basilia forcipata Ferris, (Nearctic, MND, fig. 112.1).

Diagnosis Biology
Small light brown flies (body length 1.5–5.5 mm). Head Development of immature nycteribiids is by adenotrophic
small, laterally compressed, rounded, or flattened ventrally, viviparity. The egg and all three larval instars are retained
folded back over the thorax during resting (non-feeding) pe- in the uterus, and the larvae are nourished by secretions
riods (Fig. 1). Eyes with one to four facets, or absent. Ocelli produced by accessory glands (milk glands). The female
absent. Thorax compressed dorsoventrally with few sclero- deposits a single mature larva in the immediate vicinity of
tized plates and ridges (Figs. 1, 2, 10). Mesonotum small, un- the bat host (usually the wall of the roost), where it pupates.
sclerotized. Pleuron and insertion of legs displaced to dorsal Normally, nycteribiids have a balanced sex ratio (Marshall,
surface. Scutellum absent (Figs.  1, 2). Thoracic ctenidium 1981). The adults are obligate blood-feeding ectoparasites of
present on each side between fore- and midcoxae (Figs. 2, bats (Marshall, 1981) and have been observed living in fur
10). Wings absent. Halter situated in grooves of metatho- on the host (Marshall, 1970; ter Hofstede et al., 2004).
racic pleural plates (Figs.  2, 10). Femora and tibiae com- There are few studies about biology and behavior of nyc-
pressed laterally. Empodia absent. Thoracic sterna broad and teribiids, the most important contributions being those of
flat sclerotized plates with oblique (inverted V-shaped) and Leong & Marshall (1968) and Marshall (1970, 1971), who
median sutures (Fig. 11). Female abdomen largely membra- studied Oriental species. Hase (1931) studied the behavior
nous (except Hershkovitzia) (Figs.  12, 13). Male abdomen and digestive physiology of the American species Basilia
sclerotized, with large clasperlike surstyli that fold anteriorly ferrisi Schuurmans-Stekhoven (as Basilia bellardii (Ron-
under apex of abdomen (Figs. 9, 14). Both sexes of most spe- dani)). So far the life cycle is best described in Basilia his-
cies with abdominal ctenidium present at posterior margin of pida Theodor by Marshall (1970). Below follows a summary
sternite 2 (Figs. 7–9, 13, 14). of its natural history.
Nycteribiids are easily distinguished from streblids and Males and females of B. hispida become sexually mature
hippoboscids by the small head that is folded back over the within 5–6 days of their emergence from puparia. Mating can
thorax when not feeding and by the positions of the pleuron last from 30 minutes to 2 hours. After copulation, a female
and leg insertions on the dorsal side of thorax. will deposit a mature larva every 12 to 17 days on the wall

1261
1262
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

of roost. The majority of mature larvae are deposited during (1971) and Reckardt & Kerth (2006) have demonstrated that
the day, between 09:00 and 18:00. After about 25  days the emergence of the adult is stimulated by presence of the
of development, the teneral adult emerges from the pupari- host. In laboratory studies, the longevity of females is longer
um. For other genera of Nycteribiidae, the pupal stage has a than that of males. Females can live up to 156 days, while
duration of about 17 to 20 days (Marshall, 1970). Marshall males can live 97 days.

tg 1

tsm 1

tg 2

tb

tg 5
fem

th cten
a spr
3 Basilia 4 Basilia
npl rg

pn
hlt

tg 1

tg 2

tg 5

trm abd sg

2 Basilia 5 Basilia 6 Basilia

st 2

abd ctn
st 3

st 2
st 4

st 5

st 5
st 6
sur
st 7

7 Basilia 8 Basilia 9 Basilia

Figs. 104.2–9. Abdomens: dorsal view of adult of (2) B. antrozoi (Townsend), (Nearctic, MND, fig. 112.3); dorsal view of abdomen of
(3) B. anomala Guimarães & D’Andretta, (Guimarães, 1977, fig. 1); (4) B. handleyi Guimarães, (Guimarães, 1966, fig. 35A); (5) B. costaricensis
Guimarães & D’Andretta, (Theodor, 1967, fig. 473); and (6) B. juquiensis Guimarães, (South America, Theodor, 1967, fig. 477); ventral view
of abdomen of female of (7) B. costaricensis, (Guimarães & D’Andretta, 1956, fig. 170); (8) B. juquiensis, (Graciolli & Carvalho, 2001,
fig. 9); ventral view of abdomen of male of (9) B. sp., (illustrated by G. Graciolli).
Abbreviations: a  spr, anterior spiracle; abd  cten, abdominal ctenidium; fem,  femur; hlt,  halter; npl  rg, notopleural ridge; pn,  postnotum;
st, sternite; sur, surstylus; tb, tibia; tg, tergite; th cten, thoracic ctenidium; trm abd sg, terminal abdominal segment; tsm 1, first tarsomere.
NYCTERIBIIDAE (bat flies, spider bat flies) 104 1263

Most species of Basilia show low levels of host specific- Similar data are scarce for Central America (ter Hofstede et
ity. Excluding accidental host associations by Graciolli et al. al., 2004).
(2007), 17 species of Basilia are restricted to a single host Nycteribiids are vectors of haemosporidian parasites of
species, 18 species are found on two or more species of a sin- bats; Old World species of Nycteribia Latreille are the main
gle genus, and eight species are recorded with more than two vectors of Polychromophilus dionisi to vespertilionid bats,
genera. At this time, the host is unknown for five species. The whereas in the New World, Basilia is the main vector of
species of Hershkovitzia are highly specific parasites; there Bioccala deanei to species of Myotis Kaup and Eptesicus
is one species for each species of Thyroptera Spix. In the Rafinesque (Garnham, 1973; Marinkelle, 1995). Parasites of
New World, data about prevalence and abundance or inten- nycteribiids recorded in the Old World are parasitoid wasps
sity of infestation of nycteribiids on bats are available mainly (Chalcidoidea) (Urich et al., 1922) and Laboulbeniales fun-
for the Nearctic Region (for example, Lewis, 1996; Ritzi et gae (Blackwell, 1980); none have been recorded for American
al., 2001) and for South America (for example, Graciolli et nycteribiids. Information about nycteribiid symbionts was
al., 2006; Graciolli & Bianconi, 2007; Bertola et al., 2005). summarized by Marshall (1982).

th cten
npl rg

msn

hlt

10 Hershkovitzia 11 Hershkovitzia

st 2 st 2
syntg1+2
abd cten

tg 3 st 3
st 3

tg 4 st 4
st 4

tg 5 st 5

st 5

tg 6
st 6

st 7 sur
tg 7

12 Hershkovitzia 13 Hershkovitzia 14 Hershkovitzia

Figs. 104.10–14. Thorax and abdomen: dorsal view of thorax of (10) Hershkovitzia primitiva Guimarães & D’Andretta, (South America,
Guimarães & D’Andretta, 1956, fig. 217); ventral view of thorax of (11) H. inaequalis Theodor, (South America, Theodor, 1967, fig. 596);
dorsal view (12) and ventral view (13) of abdomen of female H. primitiva and (14) ventral view of male H. primitiva, (South America,
Guimarães & D’Andretta, 1956, figs. 223–224, 227).
Abbreviations: abd cten, abdominal ctenidium; hlt, halter; msn, mesonotum; npl rg, notopleural ridge; st, sternite; syntg, syntergite;
sur,  surstylus; tg, tergite; th cten, thoracic ctenidium.
1264
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Classification Basilia. In the scheme of Guimarães & D’Andretta (1956),

Basilia may have been introduced to the New World by ves-
Nycteribiidae belong to the superfamily Hippoboscoidea, pertilionid bats, twice to North America during the early Ter-
along with the Glossinidae, Hippoboscidae, and Streblidae tiary and several times to South America during the Pliocene.
(McAlpine, 1989). The monophyly of the Hippoboscoidea According to Theodor (1967), New World Basilia would be
has been corroborated by morphological (Griffiths, 1972; closely related to the Oriental and Ethiopian B. bathybothyra
McAlpine, 1989) and molecular (Dittmar et al., 2006; Nir- group. However, Maa (1971) considered an Australian spe-
mala et al., 2001; Petersen et al., 2007) data. However, there cies Basilia transversa Maa closest to South American Basil-
is little agreement about relationships among the families. ia. Recently, Dittmar et al. (2006) corroborated Theodor’s
For example, Griffiths (1972) placed Nycteribiidae and (1967) ideas, with dispersal of Basilia spp. from the Oriental
Streblidae as the subfamily Nycteribiinae within the Hippo- Region to the Neotropical and the Nearctic Regions.
boscidae. In the more accepted scheme, McAlpine (1989)
considered Nycteribiidae the sister group of Streblidae and Identification
Hippoboscidae the sister group of Glossinidae. In a molec-
ular phylogenetic study, Nirmala et al. (2001) found Nyc-
The adult taxonomy of New World nycteribiid genera re-
teribiidae and Streblidae did not form a monophyletic group.
lies primarily on morphology of the mesonotum, legs, and
In contrast, Dittmar et al. (2006) and Petersen et al. (2007)
abdomen. Species-level taxonomy is based on female char-
corroborated the monophyly of Nycteribiidae and Strebli-
acteristics, especially those of the abdomen; for 12 species
dae, although the first authors considered Streblidae to be
of Basilia and two of Hershkovitzia, only the female sex is
paraphyletic relative to Nycteribiidae.
known. All generic and most species-level identifications
Nycteribiidae is composed of about 274 species divided
can be made using a stereo microscope, but for species-level
into three subfamilies (Graciolli  & Dick, 2006): Archynyc-
identifications it is often necessary to use slide-mounted spec-
teribiinae (one genus, three species) and Cyclopodiinae (four
imens examined with a high-power compound microscope.
genera, 60 species), both primarily parasites of bats of the sub-
Techniques for slide-mounting Nycterbiidae are the same as
order Megachiroptera and exclusively Old World in distribu-
those for the Streblidae (Wenzel et al., 1966: pp. 411–414).
tion; and Nycteribiinae (six genera, 212 species) primarily
Generic keys for American nycteribiids are available in
parasites of Megachiroptera bats with Old and New World
Guimarães & D’Andretta (1956) and Theodor (1967).
distributions. Basilia (Nycteribiinae), the largest genus of
The key below will allow generic identification and fa-
Nycteribiidae, contains about 114 described species, 48
of which exclusively occur in the Americas. All American cilitate the identification of species groups of Basilia. There
species of this genus are included in Basilia sensu stricto are no keys for immature stages of American Nycteribiidae.
(Guimarães & D’Andretta, 1956; Theodor, 1967), although Immature stages of Hershkovitzia  spp. are unknown, and
some authors disagree with this point of view. Miranda- only the larva of five species of American Basilia have been
Ribeiro (1907) erected the genus Pseudelytromyia to include described by Guimarães & D’Andretta (1956), and the pu-
American nycteribiids with the posterior margin of abdomi- paria of two Nearctic species have been described by Ferris
nal tergite  2 rounded, straight, or concave, whereas Maa (1930) and Reeves  & Reeves (2000). General information
(1965) considered Pseudelytromyia as a subgenus of Basilia. concerning immature nycteribiids is found in Ferrar (1987)
Moreover, there is no agreement as to the origin of American and Peterson & Wenzel (1987).

Key to the genera of Nycteribiidae of the New World and to the species groups of Basilia
(females only)
1. Anterior region of head membranous; notopleural ridges diverging (Fig.  10); two to four
notopleural setae present; basitarsi all shorter than remaining tarsomeres together; abdomen with
seven tergites, of which first and second fused (Fig. 12); female anal sclerite absent; posterior
margin of abdominal sternite 5 deeply concave (Fig. 14) in male. . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hershkovitzia Guimarães & D’Andretta
– Anterior region of head sclerotized; notopleural ridges parallel (Fig. 2); six or more notopleural
setae present (Fig.  2); basitarsi all longer than remaining tarsomeres together (Fig.  2); female
abdomen with two to three tergites (Figs. 2–6); female anal sclerite present; posterior margin of
abdominal sternite 5 rounded, straight, or convex (Fig. 9) in male; Basilia Miranda-Ribeiro (only
females keyed to species groups hereafter). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2. Posterior margin of abdominal tergite 2 rounded, straight, or concave (Figs. 2, 5, 6). . . . . . . . . . 3

– Posterior margin of abdominal tergite 2 with two lobes (Figs. 3, 4). . . . . . . . . . . . . . . . . . . . . . . 5
NYCTERIBIIDAE (bat flies, spider bat flies) 104 1265

3. Three abdominal tergites present; tergite 3 divided into lateral sclerites (Fig. 2); North America.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [B. antrozoi group]
– Two abdominal tergites present (Figs. 5, 6). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4. Abdominal sternite 2 extremely long, extending over sternite 3 (Fig. 8) . . . . . B. juquiensis group
– Abdominal sternite 2 longer than wide, but not extending over sternite 3 (Fig. 7). . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. speiseri group

5. Three abdominal tergites present (Fig. 3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. forcipata group

– Two abdominal tergites present (Fig. 4). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. ferruginea group

Synopsis of the fauna known from La Selva, Costa Rica, from Myotis elegans Hall
(Timm et al., 1989) and from Belize from Rhogeessa ae-
There are 16 described species of Nycteribiidae in Cen- neus Goodwin (ter Hofstede et al., 2004). Species of Basilia
tral America, but this represents about one-half of the fauna. usually are associated with vespertilionid bats, especially
New records of described and undescribed species may be of the genus Myotis; however, Basilia mimoni Theodor &
found by sampling bats of the genera Myotis, Lasiurus Gray, Peterson and B. tiptoni Guimarães are associated with the
and Rhogeessa H. Allen. All Central American nycteribiid phyllostomid bat Mimon crenulatum (E. Geoffroy).
bat flies are from the tropical zone. For species of Basilia, the most complete keys are
The following information regarding nycteribiid spe- available in Guimarães  & D’Andretta (1956) and Theodor
cies and their distribution throughout Central America has (1967), and the latter author provided a tentative key using
been compiled from Graciolli & Dick (2009), Graciolli et al. male characteristics. However, 16 species are not included in
(2007), Guimarães & D’Andretta (1956), Guimarães (1966, these keys. Other useful keys for Central American species
1972, 1977), Peterson (1971), Peterson  & Lacey (1985), are available in Peterson (1971) (including a key to males)
ter Hofstede et al. (2004), Theodor (1967), and Timm et al. and Guimarães (1966, 1972) for Panamanian and Venezu-
(1989). elan species, respectively. All of these keys are somewhat
incomplete, with some species excluded.
Basilia Miranda-Ribeiro. The species of American
Basilia are divided into five morphological groups based on Hershkovitzia Guimarães & D’Andretta. This genus is
female characters. These groups were originally proposed composed of four described species that are associated with
by Guimarães & D’Andretta (1965) and modified by Theodor New World endemic bats of the family Thyropteridae. Pe-
(1967) and Guerrero (1996). Basilia flava (Weyenbergh) and terson & Lacey (1985) provided a key to species. Hershko-
B. rugosa Schuurmans-Stekhoven are not included in any vitzia primitiva Guimarães & D’Andretta has been recorded
group because the females are unknown. At present, 15 from Costa Rica (Miller & Tschapka, 2001). An undescribed
species have been recorded for Central America, none species is known from Peruvian Thyroptera lavali Pine
belonging to the B. antrozoi group. Undescribed species are (C.W. Dick, personal communication).

Literature cited

Bertola, P.B., C.C. Aires, S.E. Favorito, G. Graciolli, M. Amaku, & Ferris, G.F. 1930. The puparium of Basilia corynorhini (Ferris)
R. Pinto-da-Rocha. 2005. Bat flies (Diptera: Sterblidae, Nycte- (Diptera: Nycteribiidae). Entomological News 41: 295–297.
ribiidae) at Parque Estadual de Cantareira, São Paulo, Brazil: Garnham, P.C.C. 1973. The zoogeography of Polychromophilus
parasitism rates and host–parasite associations. Memórias do and description of a new species of a gregarine. Annales de Pa-
Instituto Oswaldo Cruz 100: 25–32. rasitologie 48: 231–242.
Blackwell, M. 1980. Incidence, host specificity, distribution, Graciolli, G., A.G. Autino, & G.L. Claps. 2007. Catalogue of Ame-
and morphological variation in Arthrorhynchus nycteribiae rican Nycteribiidae (Diptera, Hippoboscoidea). Revista Brasi-
and A.  eucampsidae (Laboulbeniomycetes). Mycologia 72: leira de Entomologia 51: 142–159.
143–158. Graciolli, G. & G.V. Bianconi. 2007. Moscas ectoparasitas (Dipte-
Dittmar, K., M.L. Potter, S. Muray,  & M.F.  Whiting. 2006. Mo- ra, Streblidae e Nycteribiidae) em morcegos (Mammalia, Chi-
lecular phylogenetic analysis of nycteribiid and streblid bat flies roptera) em área de Floresta com Araucária no estado do Paraná,
(Diptera: Brachycera, Calyptratae): implications for host asso- sul do Brasil. Revista Brasileira de Zoologia 24: 246–249.
ciations and phylogeographic origins. Molecular Phylogenetics Graciolli, G.  & C.J.B. De Carvalho. 2001. Moscas ectoparasitas
and Evolution 38: 155–170. (Diptera, Hippoboscoidea, Nycteribiidae) de morcegos (Mam-
Ferrar, P. 1987. A guide to the breeding habits and immature stages malia, Chiroptera) do Estado do Paraná. I. Basilia, taxonomia e
of Diptera Cyclorrhapha. Entomonograph 8(1). E.J.  Brill  / chave pictórica para as espécies. Revista Brasileira de Zoologia
Scandinavian Science Press, Leiden, Copenhagen, 478 pp. 18 (Supplement 1): 33–49.
1266
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Graciolli, G. & C.W. Dick. 2006. Checklist of World Nycteribiidae McAlpine, J.F. 1989. Phylogeny and classification of the Musco-
(Diptera: Hippoboscoidea). Available from http://fm1.fieldmuseum. morpha, pp. 1397–1518. MND, Volume 3.
org/aa/Files/cdick/Nycteribiidae_Checklist_2oct06.pdf. Miller, J.A. & M. Tschapka. 2001. The Bat Flies of La Selva (Dip-
Graciolli, G. & C.W. Dick. 2009. A new species of Basilia Miran- tera: Nycteribiidae, Streblidae). Beta Version 0.96. Available
da-Ribeiro (Diptera: Nycteribiidae) from Honduras, parasite of from http://www.biologie.uni-ulm.de/bio3/Batfly/index.html.
Bauerus dubiaquercus (Van Gelder) (Chiroptera: Vespertilioni- Miranda-Ribeiro, A. 1907. Alguns dípteros interessantes. Archivos
dae: Antrozoinae). Zootaxa 1972: 59–64. do Museu Nacional 14: 231–239.
Graciolli, G., C.W. Dick, & D. Gettinger. 2006. A faunal survey of Nirmala, X., V. Hypsa, & M. Zurovec. 2001. Molecular phylog-
nycteribiid flies (Diptera: Nycteribiidae) associated with bats in eny of Calyptratae (Diptera: Brachycera): the evolution of
Paraguay. Zootaxa 1220: 35–46. 18S and 16S ribosomal rDNAs in higher dipterans and their
Griffiths, G.C.D. 1972. The phylogenetic classification of Diptera Cyclo- use in phylogenetic inference. Insect Molecular Biology 10:
rrhapha, with special reference to the structure of the male postabdo- 475–485.
men. Series entomologica 8. Dr. W. Junk, The Hague, 340 pp. Petersen, F.T., R. Meier, S.N. Kutty, & B.M. Wiegmann. 2007.
Guerrero, R. 1996. The Basilia juquiensis species-group (Diptera: The phylogeny and evolution of host choice in the Hippo-
Nycteribiidae) with description of a new species from Pakitza, boscoidea (Diptera) as reconstructed using four molecular
Perú, pp.  665–674. In Wilson,  D.E.  & A.  Sandoval (editors). markers. Molecular Phylogenetics and Evolution 45: 111–122.
MANU: La biodiversidad del sureste del Perú. Smithsonian In- Peterson, B.V. 1971. Notes on the bat flies of Costa Rica (Diptera:
stitution, Washington, D.C., 679 pp. Nycteribiidae). Contributions in Science 212: 1–8.
Guimarães, L.R. 1966. Nycteribiid batflies from Panama (Díptera: Peterson, B.V. & L.A. Lacey. 1985. A new species of Hershkovitzia
Nycteribiidae), pp. 393–404. In Wenzel, R.L. & V.J. Tipton (edi- (Diptera: Nycteribiidae) from Brazil, with a key to the described
tors). Ectoparasites of Panama. Field Museum of Natural His- species of the genus. Proceedings of the Entomological Society
tory, Chicago, xii + 861 pp. of Washington 87: 578–582.
Guimarães, L.R. 1972. Venezuelan nycteribiid batflies (Diptera: Peterson, B.V. & R.L. Wenzel. 1987. Nycteribiidae, pp. 1283–1291.
Nycteribiidae). Brigham Young University Science Bulletin, MND, Volume 2.
Biological Series 17: 1–11. Reckardt, K.  & G. Kerth. 2006. The reproductive success of
Guimarães, L.R. 1977. Supplementary note on Venezuelan bat- the parasitic bat fly Basilia nana (Diptera: Nycteribiidae)
flies (Diptera: Nycteribiidae). The Great Basin Naturalist 37: is affected by the low roost fidelity of its host, the Bech-
221–224. stein’s bat (Myotis bechsteinii). Parasitological Research
Guimarães, L.R. & M.A.V. D’Andretta. 1956. Sinopse dos Nycte- 98: 237–243.
ribiidae (Diptera) do Novo Mundo. Arquivos de Zoologia 10: Reeves, W.K.  & M.H. Reeves. 2000. The puparium of Basilia
1–184. boardmani (Diptera: Nycteribiidae) from eastern North Amer-
Hase, A. 1931. Über die Lebensgewohnheiten einer Fledermaus- ica. Studia dipterologica 7: 507–509.
fliege in Venezuela; Basilia bellardii Rondani (Fam. Nycteri- Ritzi, C.M., L.K. Ammerman, M.T. Dixon, & J.V. Richerson. 2001.
biidae  – Diptera Pupipara). Zeitschrift für Parasitenkunde 3: Bat ectoparasites from the Trans-Pecos Region of Texas, includ-
220–257. ing notes from Big Bend National Park. Journal of Medical En-
Leong, M.C. & A.G. Marshall. 1968. The breeding biology of the tomology 38: 400–404.
bat-fly (Eucampsipoda sundaicum Theodor, 1955) (Diptera: ter Hofstede H.M., M.B. Fenton, & J.O. Whitaker, Jr. 2004. Host
Nycteribiidae). Malayan Nature Journal 21:171–180. and host-site specificity of bat flies (Diptera: Streblidae and Nyc-
Lewis, S.E. 1996. Low roost-site fidelity in pallid bats: associated teribiidae) on Neotropical bats (Chiroptera). Canadian Journal
factors and effect on group stability. Behavioral Ecology and of Zoology 82: 616–626.
Sociobiology 39: 225–344. Theodor, O. 1967. An illustrated catalogue of the Rothschild collec-
Maa, T.C. 1965. An interim world list of batflies (Diptera: Nyc- tion of Nycteribiidae (Diptera) in the British Museum (Natural
teribiidae and Streblidae). Journal of Medical Entomology 1: History) with keys and the short descriptions for the identifi-
337–386.
cation of subfamilies, genera, species and subspecies. British
Maa, T.C. 1971. Revision of the Australian batflies (Diptera: Strebli-
dae and Nycteribiidae). Pacific Insects Monograph 28:1–118. Museum (Natural History) Publication 665: 1–506.
Marinkelle, C.J. 1995. The haemoproteid parasite, Bioccala deanei, Timm, R.M., D.E. Wilson, B.L.  Clauson, R.K.  La  Val,  &
C.S. Vaughan. 1989. Mammals of the La Selva-Braulio Carrillo
from a Colombian bat, Eptesicus fuscus (Vespertilionidae). An-
complex, Costa Rica. United States Department of the Interior,
nals of Tropical Medicine and Parasitology 89: 89–91.
Fish and Wildlife Service, 162 pp.
Marshall, A.G. 1970. The life cycle of Basilia hispida Theodor, 1967
Urich, F.W., H. Scott, & J. Waterston. 1922. Note on the dipterous
(Diptera: Nycteribiidae) in Malaysia. Parasitology 61: 1–18. bat-parasite Cyclopodia greeffi Karsch, and on a new species of
Marshall, A.G. 1971. The Ecology of Basilia hispida (Diptera: Nycteribi- hymenopterous (chalcid) parasite bred from it. Proceedings of
idae) in Malaysia. The Journal of Animal Ecology 40: 141–154. the Zoological Society of London 32: 471–477.
Marshall, A.G. 1981. The sex ratio in ectoparasitic insects. Ecologi- Wenzel, R.L., V.J. Tipton, & A. Kiewlicz. 1966. The streblid bat-
cal Entomology 6: 155–174. flies of Panama (Diptera: Calyptera: Streblidae), pp. 405–675. In
Marshall, A.G. 1982. Ecology of insects parasitic on bats. In Wenzel, R.L. & V.J. Tipton (editors). Ectoparasites of Panama.
Kunz, T.H. (editor). Ecology of bats. Plenum, New York, 450 pp. Field Museum of Natural History, Chicago, xii + 861 pp.
SCATHOPHAGIDAE (DUNG FLIES) 105
J. Richard Vockeroth

Fig. 105.1. Male of Scathophaga stercoraria (Linnaeus), (Holarctic, MND, fig. 103.1).

Diagnosis frons of male as wide as that of female and without cruciate

interfrontal bristles; scutellum bare below, without fine short
Slender, medium-sized (body length 6.5–13 mm), strongly erect hairs near apex; vein M with apical section straight, not
bristled and sometimes setose flies (Fig.  1), subshining to bent or curved forward; hind basitarsus without short erect
densely pruinose. Wing clear or with crossveins clouded. ventral bristle near base.
Frons about as wide as head in both sexes, without cruciate
interfrontal bristles. Arista nearly bare to plumose. Palpus Biology
slightly broadened toward apex (Figs. 2–4), with long slender
apical bristle in Cordilura (Fig.  2). Head and thorax with Larvae of Scathophaga feed on mammal dung, rotting
bristles as normally present in Calyptratae (Fig. 1); anepis- seaweed, carrion (in the Arctic), or on terrestrial egg capsules
ternum extensively bare on anteroventral portion (Fig. 6) or of Trichoptera. Larvae of Scathophaga stercoraria (Linnaeus)
with abundant fine setae on anterior one-quarter (Fig.  5); and S. furcata (Say) breed in dung; the third regional species,
katepisternum with one strong bristle posterodorsally. S. reses Giglio-Tos, is related to an Asian species, S. ampli-
Postmetacoxal bridge present (Fig.  7) or absent (Fig.  8). pennis Portschinsky, whose adults frequent horse and yak
Vein R1 bare or setose above; A1 reaching wing margin. Male dung. Larvae of other genera probably feed on insect larvae
and female terminalia varied in details of structure, the latter in water or wet soil or in nests of coots (Fulica spp.) or eider
often obviously adapted to method of oviposition. Female ducks (Somateria spp.). Larvae of Cordilura feed on plants,
abdomen with seven pairs of spiracles. in the stems of Juncus or the culms of Scirpus and Carex.
The family Scathophagidae is similar to Anthomyiidae Larvae of six other genera of Scathophaginae feed on
and Muscidae. The Neotropical species of Scathophagidae various parts of seven genera of vascular plants of five fam-
may be distinguished from species of the other two families ilies (Gorodkov, 1986; Vockeroth, 1987). All known larvae
by two characters: only one katepisternal bristle present, and of the small subfamily Delininae are leafminers in Liliaceae,
posterior margin of the lower calypter straight rather than Orchidaceae, and Commelinaceae. Adults of several genera
slightly to strongly convex. The following characters together of scathophagids, and perhaps of all, are predaceous on
also separate most species from those of the other two families: insects.

1267
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Classification ginae and Delininae, with only the former occurring in the
Neotropical Region (Delininae are north temperate). Males
Scathophagidae belong to the calyptrate superfam- of Scathophaginae have tergite 6 with bristles and hairs, and
ily Muscoidea. The family appears to be the sister group only rarely is it fused with syntergosternites  7 and 8. The
to a lineage composed of the remaining muscoid families, species of Delininae have tergite 6 bare and fused with syn-
namely Anthomyiidae, Faniidae, and Muscidae. Two sub- tergosternites 7 and 8.
families of Scathopahagidae are recognized, the Scathopha-

plp

2 Cordilura 3 Scathophaga 4 Scathophaga

a spr

anepst

5 Scathophaga 6 Cordilura

h cx h cx

pmtcx brg
mtepm mtepm

7 Scathophaga 8 Scathophaga

Figs. 105.2–8. Heads and thoracic structures: lateral view of head of (2)  Cordilura rufipes Meigen, (Holarctic, MND, fig.  103.7);
(3) Scathophaga furcata (Say), (Holarctic, MND, fig. 103.2); and (4) S. reses Giglio-Tos, (Nearctic, MND, fig. 103.3); anepisternum of
(5) S. obscura (Fallén), (Holarctic, MND, fig. 103.20); and (6) C. pudica Meigen, (Holarctic, MND, fig. 103.21); posteroventral surface of
thorax and hind coxae of (7) S. reses Giglio-Tos, (Nearctic, MND, fig. 103.23); and (8) S. aldrichi (Malloch), (Nearctic, MND, fig. 103.24).
Abbreviations: a  spr, anterior thoracic spiracle; anepst, anepisternum; h  cx,  hind coxa; mtepm, metepimeron; plp,  palpus; pmtcx  brg,
postmetacoxal bridge.
SCATHOPHAGIDAE (DUNG FLIES) 105 1269

Identification Central American species. The catalog of Neotropical

species by Albuquerque (1984) is incomplete; two omitted
James (1950), in his paper on western Nearctic papers, by Giglio-Tos (1893) and Wulp (1897), are listed in
Scathophaga (as Scopeuma Meigen), included the three the Literature cited.

Key to the genera of Scathophagidae of Central America and the West Indies
1. Palpus with one strong apical bristle subequal in length to palpus (Fig. 2); anepisternum with most
of anteroventral one-half bare (Fig. 6); postcoxal bridge absent (as in Fig. 8); scutellum with two
bristles; vein R1 setose dorsally on about apical one-quarter; posteroventral apical bristle of hind
tibia long and strong, subequal to anteroventral apical bristle. . . . . . . . . . . . . . . . Cordilura Fallén
– Palpus with several apical bristles about one-quarter as long as palpus (Figs. 3, 4); anepisternum
almost completely haired (as in Fig. 5); postcoxal bridge present (Fig. 7) or absent (Fig. 8); scutellum
with four bristles: vein R1 bare; posteroventral apical bristle of hind tibia at most one-quarter as
long as anteroventral apical birstle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scathophaga Meigen

Synopsis of the fauna lobes, occurs in Guerrero and Chiapas, Mexico, and in Costa
Rica. Both species are varied in color.
Only two of the 40 New World genera are known from
Central America; the diagnosis is based on these two. Scathophaga Meigen. Three named species are known
Scatogera Albuquerque occurs in Ecuador and the other 37 from Mexico, Guatemala, Costa Rica, and Haiti. They can
genera occur north of Mexico. A more detailed description of be identified using James (1950). Scathophaga stercoraria
the Nearctic fauna is given by Vockeroth (1987). (Fig.  1) has fine hairs on the anepimeron; S.  furcata has
short aristal hairs (Fig. 3) and no postmetacoxal bridge (as
Cordilura Fallén. An undescribed species of Cordilura, in Fig. 8); S. reses has a plumose arista (Fig. 4) and an entire
with the lobes of male sternite 5 black, occurs in Arizona and postmetacoxal bridge (Fig. 7).
Durango, Mexico. Cordilura brevicornis Wulp, with yellow

Literature cited

Albuquerque, D. de O. 1984. 96B. Family Scatophagidae. In James, M.T. 1950. The genus Scopeuma in the western United States
Papavero, N. (editor). A Catalogue of the Diptera of the Americas and southwestern Canada. Annals of the Entomological Society of
south of the United States. Departamento de Zoologia, Secre- America 43: 343–353.
taria da Agricultura, São Paulo, 4 pp. Vockeroth, J.R. 1987. 103. Scathophagidae, pp.  1085–1097. MND,
Giglio-Tos, E. 1893. Diagnosi di nuovi generi e di nuove specie Volume 2.
di Ditteri. IX.  Bollettino dei Musei di Zoologia ed Anatomia Wulp, F.M. van der. 1897. Insecta. Diptera [part], pp. 345–360. In God-
Comparata della Reale Università di Torino 8: 1–14. man, F.D. & O. Salvin (editors). Biologia Centrali-Americana, or,
Gorodkov, K.B. 1986. Family Scathophagidae, pp.  11–41. In contributions to the knowledge of the fauna and flora of Mexico
Soós, Á. & L. Papp (editors). Catalogue of Palaearctic Diptera, and Central America. Zoologia. Class Insecta. Order Diptera.
Volume 11. Akadémiai Kiadó, Budapest, 346 pp. Volume 2. London, 489 pp.
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
ANTHOMYIIDAE (ANTHOMYIID FLIES) 106
Verner Michelsen

Fig. 106.1. Anthomyia pluvialis (Linnaeus), (Holarctic, MND, fig. 104.1).

Diagnosis Presence of a projecting (nonlinear) lower calypter and/or

a pair of crossed setae or setulae on frontal vitta will separate
Small to medium-sized muscoid flies (Fig. 1) with wing anthomyiids from Scathophagidae. Presence of an outstand-
length ranging from 2.2–10.0  mm. Most species strongly ing ventrobasal setula on hind basitarsus will distinguish
sexually dimorphic. Male abdomen usually slender and them from Fanniidae. Presence of hairs on the ventral tip of
parallel-sided, cylindrical to strongly depressed with promi- scutellum (Fig. 4) and/or the faint extension of wing vein A1
nent U-shaped sternite 5. Ground color of head, body, and to the wing margin (Fig. 5) will separate most anthomyiids
appendages often wholly or predominantly brownish-black, from Muscidae. However, the anthomyiid genus Coenos-
but shades of amber, orange–red or yellow occur frequently, opsia resembles Muscidae of the tribe Coenosiini in having
especially on frontal vitta, antenna, palpus, legs, and abdo- both a ventrally bare scutellum and a short vein A1, but can
men. More or less intense layer of whitish-gray to dark brown be easily distinguished by the presence of a pair of setae on
pruinosity covers most body parts, often resulting in pattern the frontal vitta.
of dark stripes, markings, or bands on dorsum of thorax and
abdomen; patterns often only clearly visible from certain
angles of view. Pair of inclinate frontal setae usually present Biology
on frontal vitta of females and broad-fronted males. Katepis-
ternal setae 0–2 + 1–3; meron bare, exceptionally with some The adults of many anthomyiid species are anthophilous
irregular setulae, but never with row of setae. Ventral tip and play a significant role as pollinators, at least in the north
of scutellum usually but not always with ventral hairs (fine temperate zones. As well as feeding on nectar, certain spe-
setae). Basitarsus of hind leg with outstanding ventrobasal cies also eat pollen. Other species are more attracted to
setula. M1 not strongly curved or angled forward on distal honeydew or fermenting sap from wounded trees and fruits
part; vein A1 usually traceable to margin of wing. Upper and or may feed on dung and carrion. Males of many species
lower calypteres usually forming subequal, modest-sized tend to engage in premating activities like aerial swarming
lobes, but exceptionally, lower calypter strongly reduced. or station-taking.

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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Eggs (or sometimes first stage larvae) are laid singly or Classification
in small clusters, normally onto or buried within the larval
food item. The larvae are highly varied in their feeding hab- Anthomyiids are usually grouped together with
its, including scavengers as well as herbivores and fungivores. Scathophagidae, Fanniidae, and Muscidae in the superfamily
Larval exploitation of decaying substrates like dung, carrion,
Muscoidea, although this taxon is now considered as a grade
and debris in vertebrate nests and borrows may hold true for
rather than a clade. Morphological evidence (Michelsen,
all Neotropical representatives of the genera Phaonantho,
Coenosopsia, Hydrophoria, Fucellia, Calythea, Pegoplata, 1991) supports a sister-group relationship between the An-
Anthomyia, Lasiomma, Eutrichota, Emmesomyia, and Tae- thomyiidae and Muscidae, while current molecular analyses
niomyia. Specialized cleptoparasitism has developed in Leu- suggest that Anthomyiidae are paraphyletic without inclusion
cophora and Eustalomyia, where the larvae feed on larval of a monophyletic Scathophagidae (Kutty et al., 2007) and
provision (pollen balls and arthropod prey, respectively) in the that a sister-group relationship exists between Oestroidea
nests of solitary bees and sphecoid wasps. Feeding on fresh and Anthomyiidae + Scathophagidae (Kutty et al., 2008).
herbal or fungal tissues are important habits among anthomyiid Most of the approximately 40 anthomyiid genera recog-
larvae of the temperate zones, but in tropical Central America nized worldwide appear well defined morphologically and
herbivory occurs only in Delia and some Pegomya, and fungi- are presumably monophyletic. A practical problem is, how-
vory may be restricted to one species group of Pegomya. ever, that most of the characters diagnostic for the genera

2 Zaphne 3 Strobilomyia

ap sctl s
ap sctl sta
Sc R1
R2+3

br r-m
R4+5
dm
bm-cu
dm-cu
M1
CuA2

A1+CuA2 CuA1

v hairs
5 Botanophila

4 Paradelia
Figs. 106.2–5. Heads, scutellum, and wing: lateral view of head of (2) Zaphne divisa (Meigen), (Holarctic, MND, fig. 104.13, as Hydrophoria);
and (3)  Strobilomyia neanthracina Michelsen, (Nearctic, MND, fig.  104.14, as Lasiomma anthracina); posteroventral view of scutellum
of (4) Paradelia intersecta (Meigen), (Holarctic, MND, fig. 104.23, as Pseudonupedia); dorsal view of wing of (5) Botanophila spinidens
(Malloch), (Nearctic, MND, fig. 104.29).
Abbreviations: ap sctl s, apical scutellar seta; ap sctl sta, apical scutellar setula; v hairs, ventral hairs.
ANTHOMYIIDAE (ANTHOMYIID FLIES) 106 1273

are found in the male genitalia and tend to be inaccessible Asia, are poorly sampled, which makes it plausible that there
without dissection. exists a total of 2500–3000 species worldwide.
A formal higher classification of the anthomyiid genera There is no general revision or identification guide to the
is not given here, because the existing attempts are neither genera and species of Neotropical Anthomyiidae. A useful
in good agreement nor founded on stringent cladistic argu- but somewhat outdated treatment of the anthomyiid species
mentation. Some natural groups of genera can of course be of southern South America is given by Malloch (1934). Pont
identified. Of particular interest here is the possibility that (1972, 1974) compiled the genera and species of Anthomyii-
there exists a sister-group relationship between the New dae of the Neotropical Region (including Nearctic parts of
World genera Phaonantho + Coenosopsia and the remaining Mexico) reported in the pre-1970 literature. More recent
Anthomyiidae (see Michelsen, 1991, 1996). contributions to the knowledge of Neotropical anthomyiids
are remarkably scanty, but see Albuquerque & Couri (1979,
1981), Couri (1979), Griffiths (1987, 1998, 2001, 2003), Mi-
Identification chelsen (1991), Pamplona (1991), Pamplona & Couri (1993),
Nihei & Carvalho (2004), and Pont & Ackland (2009). Pam-
The family Anthomyiidae is primarily confined to the plona (1992) attempted to construct a key to Neotropical
cooler, temperate zones of the Northern Hemisphere. Less genera of Anthomyiidae, but it does not work in many cases
than 2000 species in around 40 genera have been described, (species of Delia, Eutrichota, Taeniomyia, etc.) and omits
but many species-rich areas, not the least in Central and East some genera from consideration.

Key to the genera of Anthomyiidae of the Neotropical Region

The name of the endemic South American genus Myopella Malloch, 1934 (type species Myopina appendiculata Stein, 1904,
by original designation) is preoccupied by Myopella Robineau-Desvoidy, 1853 (Diptera, Conopidae). I hereby propose
Myopinella (new replacement name) for Myopella Malloch.

1. Prosternum setulose. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Prosternum bare, rarely with pair of setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

2. Lower calypter wider than upper calypter; hind tibia with one posterodorsal seta. . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calythea Schnabl & Dziedzicki
– Lower calypter reduced to linear fold narrower than upper calypter; hind tibia with three to six
posterodorsal setae; southern South America . . . . . . . . . . . . . . . . . . . . . . [Myopinella Michelsen]

3. Hind tibia with one posterodorsal seta; arista long-plumose. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

– Hind tibia with two to five posterodorsal setae; arista various . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

4. Scutellum bare ventrally; hind tibia with posterodorsal seta shorter than hind tarsomere  2 and
inserted on distal half of tibia; body small, wing length 2.2–4.0 mm; male head dichoptic. . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Coenosopsia Malloch
– Scutellum with hairs (fine setae) ventrally at tip (Fig. 4); hind tibia with posterodorsal seta longer
than hind tarsomere 2 and inserted at midlength of tibia; larger, wing length exceeding 5 mm;
male head holoptic. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phaonantho Albuquerque

5. Scutellum bare ventrally; vein C with scattered series of long ventral spinules; male head dichoptic;
worldwide coastal. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Fucellia Robineau-Desvoidy]
– Scutellum with some fine or even microscopic hairs ventrally at tip (Fig. 4); vein C short-setulose
or bare on ventral surface; male head various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

6. Arista long-plumose (Fig. 2), its longest rays far exceeding width of flagellomere. . . . . . . . . . . 7
– Arista pubescent to short-plumose (Fig.  3), its longest rays at most equal to width of
flagellomere. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
1274
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

7. Lower calypter longer than upper calypter; palpus and legs brownish-black. . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrophoria Robineau-Desvoidy
– Lower calypter shorter than upper calypter; palpus and legs in part or entirely ochreous or
yellow. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

8. Antenna black; scutellum light-gray pruinose with pair of large black lateral markings . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eustalomyia Kowarz
– Antenna ochre-yellow; scutellum with thin brown pruinosity over amber to black ground color .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leucophora Robineau-Desvoidy, in part

9. Vein C bare on ventral surface. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

– Vein C with short setulae on ventral surface. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

10. Head enlarged, parafacial and gena much wider than comparatively small flagellomere; female
cercus with strong, upcurved spines. . . . . . . . . . . . . . . . .Leucophora Robineau-Desvoidy, in part
– Head not modified as above; female cercus lacking strong, upcurved spines. . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Delia Robineau-Desvoidy

11. Middle of proepisternum with fine setae; light-gray pruinose with conspicuous pattern of dark
markings or bands on scutum and abdomen . . . . . . . . . . . . . . . . . . . . . Anthomyia Meigen, in part
– Middle of proepisternum bare; body pattern various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

12. Hind tibia with three to five posterodorsal setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

– Hind tibia with two posterodorsal setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

13. Frontal vitta bare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eutrichota Kowarz, in part

– Frontal vitta with pair of interfrontal setae or fine setae (may need careful observation in
holoptic males). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

14. Prealar seta shorter than anterior notopleural seta. . . . . . . . . . . . . . . . . . . . . . . . . Lasiomma Stein
– Prealar seta same length or longer than anterior notopleural seta . . . . . Anthomyia Meigen, in part

15. Apex of scutellum, between pair of strong setae, with fine setae no larger than ventral setae at
apex of scutellum; female palpus strongly dilated apically. . . . . Pegoplata Schnabl & Dziedzicki
– Apex of scutellum (if not bare) with apical setulae of same size as those on discal and lateral parts;
female palpus normal. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

16. Anepimeron with one (to three) setae near upper margin; basal node of vein R2+3 with short, fine
setulae on dorsal and ventral surfaces. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Emmesomyia Malloch
– Anepimeron and basal node of vein R2+3 bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

17. Arista short-plumose, its longest rays exceeding greatest width of palpus. . . . . Taeniomyia Stein
– Arista pubescent, its longest rays barely exceeding greatest width of arista. . . . . . . . . . . . . . . . 18

18. Prealar seta much longer than anterior notopleural seta . . . . . . . . . . . . Eutrichota Kowarz, in part
– Prealar seta same length or shorter than anterior notopleural seta. . Pegomya Robineau-Desvoidy
ANTHOMYIIDAE (ANTHOMYIID FLIES) 106 1275

Synopsis of the fauna tropics. Griffiths (1984) recorded a couple of species from
the Nearctic frontier zone of Mexico, while four different
There are few published records of Anthomyiidae from species have been identified from Costa Rica. This suggests
Central America. The following summary of the known an- that the genus occurs throughout Central America.
thomyiid diversity of Central America is based primarily
on an unpublished, preliminary examination of 609 speci- Eustalomyia Kowarz. One species has been seen from
mens from Costa Rica, most of which belong to the INBio Costa Rica, which could be identical with the widespread
Collection (Heredia, Costa Rica). Some 32 morphospecies Holarctic E.  vittipes (Zetterstedt), but safe identification is
in 14 genera have been identified. I expect that there are not possible in the absence of males. The genus is presently
at least 50 species of anthomyiids to be found in Central recorded for the first time from the Neotropical Region.
America. Eutrichota Kowarz. Specimens of this genus are known
At least 14 out of a total of 16 anthomyiid genera known from South America (Bolivia), but have up to the present
from the entire Neotropical Region also occur in Central been wrongly assigned to Pegomya. Two unidentified spe-
America. One of the missing genera, Fucellia Robineau- cies have been found in Costa Rica. Previously, it was only
Desvoidy, which includes the well-known kelp flies devel- recorded from the Mexican border zone northward in the
oping in seashore beds of brown algae, will probably also Nearctic Region (Griffiths, 1984).
be found on coastal sites in southern Mexico or elsewhere
in Central America. The other missing genus, Myopinella, Hydrophoria Robineau-Desvoidy. Only H. plumosa van
consists of two species that are endemic to southern South der Wulp, a widespread New World species, is known from
America and the high Andes of Peru and Bolivia. Central America. It was recorded from Neotropical parts
of Mexico by Griffiths (1998) and has also been seen from
Anthomyia Meigen (synonyms: Craspedochaeta Mac- Costa Rica.
quart; Hylemyioide Albuquerque). This is the largest an-
thomyiid genus in the Neotropical Region (Pont, 1974). Lasiomma Stein. The first Neotropical species, L.  zor-
One species, A.  confusa (Albuquerque), apparently ranges rense, was described from the southern tip of Baja California,
from Brazil to southern United States. Griffiths (2001) re- Mexico, by Griffiths (2003). A second species, represented
corded A. confusa and A. ophiata Griffiths from Neotropical by females only, has been seen from Costa Rica. It resembles
Mexico. Two species have further been identified from Costa the Holarctic L. seminitidum (Zetterstedt) and allies by the
Rica. One is A. oculifera (Bigot), an eastern Palearctic and presence of ventral setulae on vein C, a short and pruinose
widespread Nearctic species. The other is an unidentified prementum, and absence of an apical posteroventral seta on
species with dichoptic males belonging to the “A. punctipen- hind tibia.
nis section” of Griffiths (2001).
Leucophora Robineau-Desvoidy (synonym: Neohyl-
Calythea Schnabl & Dziedzicki. Ackland (1968) record- emyia Malloch). One distinctive species has been seen from
ed C.  micropteryx (Thomson) from the West Indies, while Costa Rica. It belongs to a new species in the “L. nigricauda
Griffiths (1987) recorded C.  crenata (Bigot) from Neotro- section” of Griffiths (1996) not previously recorded south of
pical Mexico. Both species have now been identified from Mexico. More species probably occur in Central America, at
Costa Rica and may be widespread in the Central American least the widespread Nearctic and South American species
highlands. L. johnsoni (Stein).

Coenosopsia Malloch. This New World genus revised Pegomya Robineau-Desvoidy (synonym: Petrmyia Al-
by Michelsen (1991) and Nihei  & Carvalho (2004) ranges buquerque). This genus has not previously been recorded
from Texas and Florida to Rio de Janeiro. Two species are from Central America, but I have seen six species of Pego-
presently recorded from Central America: C. prima Malloch mya from Costa Rica. Three of them belong to the “P. win-
(Mexico, Belize, Costa Rica, Panama) and C. mexicana Mi- themi superspecies” of Griffiths (1983), characterized by the
chelsen (Mexico). The genus was still assigned to the family disc-shaped distal tarsomeres of the female mid- and hind
Muscidae by Pont (1972). legs. Larvae of this mainly Holarctic group feed expectedly
on mushrooms. The remaining species, as judged from the
Delia Robineau-Desvoidy (synonym: Erioischia Lioy). structure of the female ovipositor, have leaf-mining larvae,
Only two species have been seen from Costa Rica. One and at least two of them belong to the “P.  dorsimaculata
appears identical with the common seed-corn maggot, superspecies” of Griffiths (1983) recorded previously from
D.  platura (Meigen, 1826). More species can certainly be North America and Brazil.
expected from Central America, especially from high alti-
tude grasslands. Pegoplata Schnabl  & Dziedzicki. I have examined a
couple of species of Pegoplata belonging to the “P. palposa
Emmesomyia Malloch. This is one of the few anthomyiid subsection” of Griffiths (1986) from Costa Rica. The occur-
genera with a focal distribution in the Old and New World rence of species of the “P. palposa subsection” in the Nearctic
1276
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

frontier zone of Mexico (see Griffiths, 1986) suggests that Taeniomyia Stein (synonym: Anthojuba Albuquerque &
the group is widespread in Central America. Couri, based on species with feminized male head; new syn-
onymy). This is an important genus in the New World tropics,
Phaonantho Albuquerque. This is a small genus endemic but a few species reach into the Nearctic Region according
to the New World tropics. One species, P. mallochi (Curran),
to Griffiths (1984), who incidentally treated Taeniomyia as
recorded from Panama by Curran (1934), is also found in
Costa Rica. The genus was still assigned to the family a subgenus of Emmesomyia. At least five different species
Muscidae by Pont (1972). have been identified from Costa Rica.

Literature cited

Ackland, D.M. 1968. The World species of Calythea Schnabl and Kutty, S.N., M.V. Bernasconi, F. Šifner, & R. Meier. 2007. Sensitiv-
Dzied. (Dipt., Anthomyiidae) with notes on Bigot’s types. Ento- ity analysis, molecular systematics and natural history evolution
mologist’s Monthly Magazine 104: 135–144. of Scathophagidae (Diptera: Cyclorrhapha: Calyptratae). Cla-
Albuquerque, D. de O.  & M.S.  Couri. 1979. Sobre Emmesomyia distics 21: 258–271.
Malloch, 1917 e Taeniomyia Stein, 1918 com descrição de uma Kutty, S.N., T. Pape, A. Pont, B.M. Wiegmann, & R. Meier. 2008.
espécie nova (Diptera, Anthomyidae). Revista Brasileira de The Muscoidea (Diptera: Calyptratae) are paraphyletic: Evi-
Biologia 39: 493–498. dence from four mitochondrial and four nuclear genes. Molecu-
Albuquerque, D. de O. & M.S. Couri. 1981. Considerações sobre lar Phylogenetics and Evolution 49: 639–652.
Taeniomyia Stein, 1918 e descrição de Anthojuba, gen. n. (Dip- Malloch, J.R. 1934. Muscidae, pp. 171–346. Diptera of Patagonia
tera, Anthomyidae). Papéis Avulsos de Zoologia 34: 155–160. and South Chile, Part VII. British Museum of Natural History,
Couri, M.S. 1979. Sobre o gênero Phaonantho Albuquerque, London.
1957 com descrição de uma espécie nova e notas sinonímicas Michelsen, V. 1991. Revision of the aberrant New World genus
(Diptera, Anthomyidae). Revista Brasileira de Biologia 39: Coenosopsia (Diptera: Anthomyiidae), with a discussion of an-
525–528. thomyiid relationships. Systematic Entomology 16: 85–104.
Curran, C.H. 1934. The Diptera of Kartabo, Bartica District, Brit- Michelsen, V. 1996. First reliable record of a fossil species of An-
ish Guiana, with descriptions of new species from other British thomyiidae (Diptera), with comments on the definition of recent
Guiana localities. Bulletin of the American Museum of Natural and fossil clades in phylogenetic classification. Biological Jour-
History 66: 287–532. nal of the Linnean Society 58: 411–451.
Griffiths, G.C.D. 1983. Anthomyiidae 2. In Griffiths, G.C.D. (edi- Nihei, S.S. & C.J.B. de Carvalho. 2004. Taxonomy, cladistics and
tor). Flies of the Nearctic Region. 8: 161–288. E. Schweizerbart, biogeography of Coenosopsia Malloch (Diptera, Anthomyii-
Stuttgart. dae) and its significance to the evolution of anthomyiids in the
Griffiths, G.C.D. 1984. Anthomyiidae 3. In Griffiths, G.C.D. (edi- Neotropics. Systematic Entomology 29: 260–275.
tor). Flies of the Nearctic Region. 8: 289–408. E. Schweizerbart, Pamplona, D. 1991. Descrição das genitálias de dois holótipos de
Stuttgart. Craspedochoeta Macquart (Diptera, Anthomyiidae). Revista
Griffiths, G.C.D. 1986. Anthomyiidae 5. In Griffiths, G.C.D. (edi- Brasileira de Zoologia 7: 657–661.
tor). Flies of the Nearctic Region. 8: 601–728. E. Schweizerbart, Pamplona, D. 1992. Gêneros Neotropicais de Anthomyiidae  —
Stuttgart. chave para adultos (Insecta, Diptera). Revista Brasileira de En-
Griffiths, G.C.D. 1987. Anthomyiidae 6. In Griffiths, G.C.D. (edi- tomologia 36: 569–574.
tor). Flies of the Nearctic Region. 8: 729–952. E. Schweizerbart, Pamplona, D. & M.S. Couri. 1993. Descrição de Phaonantho sor-
Stuttgart. dilloae, sp.  n. (Diptera, Anthomyiidae). Revista Brasileira de
Griffiths, G.C.D. 1996. Anthomyiidae 11. In Griffiths, G.C.D. (edi- Entomologia 37: 555–558.
tor). Flies of the Nearctic Region. 8: 1633–1872. E. Schweizer- Pont, A.C. 1972. 97. Family Muscidae. In Papavero, N. (editor). A
bart, Stuttgart. Catalogue of the Diptera of the Americas South of the United
Griffiths, G.C.D. 1998. Anthomyiidae 12. In Griffiths, G.C.D. (edi- States. Departamento de Zoologia, Secretaria da Agricultura,
tor). Flies of the Nearctic Region. 8: 1873–2120. E. Schweizer- São Paulo, 111 pp.
bart, Stuttgart. Pont, A.C. 1974. 96a. Family Anthomyiidae. In Papavero, N. (edi-
Griffiths, G.C.D. 2001. Anthomyiidae 13. In Griffiths, G.C.D. (edi- tor). A Catalogue of the Diptera of the Americas South of the
tor). Flies of the Nearctic Region. 8: 2121–2288. E. Schweizer- United States. Departamento de Zoologia, Secretaria da Agri-
bart, Stuttgart. cultura, São Paulo, 21 pp.
Griffiths, G.C.D. 2003. Anthomyiidae 14. In Griffiths, G.C.D. (edi- Pont, A.C.  & D.M. Ackland. 2009. The types of Anthomyiidae
tor). Flies of the Nearctic Region. 8: 2289–2484. E. Schweizer- (Diptera) in the Museum für Naturkunde Berlin, Germany.
bart, Stuttgart. Zoosystematics and Evolution 85: 5–56.
fANNIIDAE (FANNIID FLIES, LATRINE FLIES) 107
Jade Savage

Fig. 107.1. Male of Fannia canicularis (Linnaeus), (illustrated by A. Brenes).

Diagnosis projecting beyond anterior level of frons. Males generally

holoptic in Fannia, dichoptic in Euryomma. Holoptic males
Small to medium-sized flies (body length 3.5–7.5  mm). with up to 14 fronto-orbital bristles and with zero to one or-
Generally dark in color (rarely metallic), but occasionally bital bristle; all female fanniids and males of Euryomma with
with partially to completely yellow legs, head and abdomen. two distinct orbital bristles. Parafacial in females usually
Sexual dimorphism common in the genus Fannia, mostly ap- broad, completely tomentose to shiny black. Arista almost
parent in head, legs, and abdomen. Eyes large, setulose or bare, bare to short pubescent, not plumose in Central American
with gena at most one-eighth of eye height. Oral margin rarely species (only plumose in Piezura Rondani, a Holarctic genus

1277
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

not recorded from Central America). Wing vein Sc diverging 1993). Not much is known of the biology of Euryomma, but
from R1 at point near base and with apical half almost straight two species are known to be associated with refuse heaps of
(Figs. 2, 3). In all species of Fannia, imaginary extensions army ants (Chillcott, 1958).
of wing veins A1 and A2 meeting well before wing margin
(Fig. 2). Lower calypter normally subequal to upper one, but
occasionally reduced and linear or projecting beyond level
Classification
of upper calypter. Femora and tibiae of the mid- and hind
legs often modified in males of Fannia, bearing mats of short Fanniids were long treated as a subfamily of the Muscidae
setae or tufts of long setae. In both sexes, hind coxa often (Chilcott, 1961; Hennig, 1965; Huckett & Vockeroth, 1987),
bearing few posterodorsal setae; hind tibia with true middor- but arguments presented by Roback (1951), Griffiths (1972),
sal bristle at midlength in addition to preapical dorsal bristle and Pont (1977) eventually led to the ranking of this group
(Fig.  5). Larvae dorsoventrally flattened, bearing various as a full family. Michelsen (1991) pointed out that fanniids
fleshy processes on the bodysurface (Fig. 4). exhibit many plesiomorphic characters believed to be part of
Fanniids can easily be separated from most other mus- the ground plan of the Muscoidea sensu McAlpine (1989),
coids by the distinctive curvature of vein Sc and the presence suggesting that it might be the most primitive muscoid fam-
of a middorsal bristle on the hind tibia. Females can further ily. However, two analyses of molecular data have recently
be separated from muscids by the presence of abdominal challenged the monophyly of the Muscoidea (Bernasconi et
spiracles  6 and 7. The peculiar shape of the larvae is also al., 2000; Kutty et al., 2008), and in both of these studies the
distinctive and not found in any other muscoid groups, but Fanniidae were placed as the sister group to a clade contain-
similar adaptations are present in some platypezids, phorids ing the Oestroidea and the remaining Muscoidea.
(Chillcott, 1961), and calliphorids. Worldwide, there are only four valid genera recognized:
Fannia (syn. Homalomyia Bouché, Coelomyia Haliday), Eu-
ryomma, Piezura (syn. Platycoenosia Strobl), and Australo-
Biology fannia Pont. Only the first two are found in Central America,
with Piezura being restricted to the Holarctic Region and
Fanniid larvae feed as scavengers and have been recorded Australofannia found only in southeastern Australia. There
from a wide array of decaying organic matter, including rot- are no generic-level phylogenies available for this family.
ting plant material, latrines, dung, and carrion (Pont, 1973;
Ferrar, 1987). Many species are synanthropic and considered
to be urban pests. Fannia canicularis (Linnaeus), the lesser
Identification
house fly, has an almost cosmopolitan distribution and is the
most common species of fanniid involved in cases of uro- Vockeroth’s (1996) key to Central American muscids
genital myiasis, and to a lesser extent intestinal myiasis. It is including fanniids can be used to separate the two genera.
also of hygienic concern, as it is a synanthropic species known Chillcott’s (1961) key to the Nearctic fauna is more detailed
to be attracted to feces and carrion. Fannia incisurata (Zetter- at the generic level and also applies to the Central American
stedt), not as broadly distributed as F. canicularis, has been fauna.
involved in cases of urogenital, intestinal, and aural myia- Fanniids are often collected from Malaise and bait traps
sis. The latrine fly Fannia scalaris (Fabricius) has a biology (feces, carrion, rotting vegetables) or by sweeping over flow-
similar to that of F. incisurata, but this species seems to favor ers and decaying matter. The larvae are easy to identify at the
a more temperate climate, as it is common in the Nearctic family level and can be collected directly from a variety of
Region and in southern South America, but has never been often semi-liquid media, such as cesspools, dung, and decay-
recorded from Central America. Fannia benjamini Malloch, ing organic matter. Smaller adult specimens collected in al-
present in Mexico, Cuba, and the southern USA, pesters cohol should be either chemically or critical-point dried and
humans and livestock by feeding on body secretions. This then mounted on a cardboard point or glued to a pin, while
species is also a carrier of eye nematodes of the genus Thela- larger ones can be air-dried or chemically dried using ethyl
zia Bosc. Males of many Fannia species can be observed acetate (Vockeroth, 1966). Freshly killed specimens should
hovering in aerial swarms, while females will generally be be pinned after a relaxing period (24 hours), depending on
found resting on vegetation. A few species are known to be size, and either directly mounted on a regular pin or double-
attracted to sweat and body secretions (Carvalho et al., mounted using a minuten pin.
fANNIIDAE (FANNIID FLIES, LATRINE FLIES) 107 1279

Key to the genera of Fanniidae of Central America

1. Imaginary extension of wing veins A1 and A2 meeting well before wing margin (Fig. 2); males
generally holoptic, not with more than one orbital bristle present; first presutural dorsocentral
bristle various, but usually over half as long as second; size various . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fannia Robineau-Desvoidy
– Imaginary extensions of wing veins A1 and A2 meeting at or near wing margin (Fig. 3); males
dichoptic, with two orbital bristles present; first presutural dorsocentral bristle weak, generally
less than half as long as second; small flies, normally less than 4.5 mm long.. . . Euryomma Stein

Synopsis of the fauna Central America, but there is at least one undescribed
species from Chiapas, Mexico, in the holdings of the Canadian
The family contains over 270 described species world- National Collection of Insects.
wide, most of which belong to Fannia. Euryomma is an
Fannia Robineau-Desvoidy. This genus is relatively
exclusive New World genus, while Fannia species can be diverse in the Neotropical Region, consisting of about 67
found in all realms, being more diverse in the Holarctic and species. Only nine species have officially been recorded
the Neotropical Regions. About 13 species are found in from Central America, but at least eight more can be found in
Central America. Nearctic Mexico. There is no comprehensive key to species
Species numbers for each genus were compiled from for Neotropical Fannia. Albuquerque et al. (1981), provided
Carvalho et al. (2003). a key to major species groups as well as a partial key includ-
ing 31 Neotropical species. Pont & Carvalho (1994) wrote
Euryomma Stein. This is a New World genus, with nine a key to all Neotropical species of the Fannia anthracina
out of ten described species found in the Neotropical realm. group. The Nearctic fauna of Fanniidae has been revised by
Carvalho & Pamplona (1979) wrote a key to all species of Chillcott (1961) and can be used to confirm the identification
Euryomma in Portuguese. Three species are known from of a number of species also present in Central America.

A1
A2

2 Fannia 3 Euryomma

ds

4 Fannia

5 Fannia
Figs. 107.2–5. Wings, larva, and hind tibia: dorsal view of wing of (2) Fannia canicularis (Linnaeus), (MND, fig. 105.25); (3) Euryomma
peregrinum (Meigen), (MND, fig. 105.26); lateral view of larva of (4) F. canicularis, (MND, fig. 105.50); dorsal view of hind tibia of
(5) F. scalaris (Fabricius), (cosmopolitan but not recorded from Central America, MND, fig. 105.31).
Abbreviation: d s, dorsal setae.
1280
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Literature cited

Albuquerque, D de O., D. Pamplona, & C.J.B. de Carvalho. 1981. Hennig, W. 1965. Vorarbeiten zu einem phylogenetischen System
Contribuçiãdo ao conhecimento dos Fannia  R.-D., 1830 da der Muscidae (Diptera: Cyclorrhapha). Stuttgarter Beiträge zur
região neotropical (Diptera, Fanniidae). Archivos Museu Nacio- Naturkunde 14: 1–100.
nal 56: 9–34. Huckett, H.  & J.R. Vockeroth. 1987. Muscidae, pp.  1115–1131.
Bernasconi, M.V., C. Valsangiacomo, J.-C. Piffaretti, & P.I. Ward. MND, Volume 2.
2000. Phylogenetic relationships among Muscoidea (Diptera: Kutty, S.N., T. Pape, A.C. Pont, B.M. Wiegmann, & R. Meier. 2008.
Calyptratae) based on mitochondrial DNA sequences. Insect The Muscoidea are paraphyletic: Evidence from four mitochon-
Molecular Biology 9: 67–74. drial and four nuclear genes. Molecular Phylogenetics and Evo-
Carvalho, C.J.B. de & D. Pamplona. 1979. Sobre uma nova espécie lution 49: 639–652.
de Euryomma Stein, 1899 (Diptera: Fanniidae). Revista Brasi- McAlpine, J.F. 1989. Phylogeny and classification of the Musco-
leira de Biologia 39: 601–604. morpha, pp. 1397–1518. MND, Volume 3.
Carvalho, C.J.B. de, A.C. Pont, M.S. Couri, & D. Pamplona. 1993. Michelsen, V. 1991. Revision of the aberrant New World genus
Part I. Fanniidae, 29 pp. In Carvalho, C.J.B. de (editor). A Cata- Coenosopsia (Diptera: Anthomyiidae), with a discussion of an-
logue of the Fanniidae and Muscidae (Diptera) of the Neotropi- thomyiid relationships. Systematic Entomology 16: 85–104.
cal Region. Sociedade Brasileira de Entomologia, Sao Paulo, Pont, A.C. 1973. Muscidae (house-flies, stable-flies, etc.). Chap-
29 + 201 pp. ter 5b, pp. 251–269. In Smith, K.G.V. (editor), Insects and other
Carvalho, C.J.B. de, A.C. Pont, M.S. Couri, & D. Pamplona. 2003. arthropods of medical importance. British Museum (Natural
A catalogue of the Fanniidae (Diptera) of the Neotropical Re- History), London, xiv + 561 pp.
gion. Zootaxa 219: 1–32. Pont, A.C. 1977. A revision of Australian Fanniidae (Diptera: Ca-
Chillcott, J.G. 1958. Two new species of Euryomma (Stein) (Mus- lyptrata). Australian Journal of Zoology Supplement Series 51:
cidae: Diptera) from Panama. The Canadian Entomologist 90: 1–60.
725–731. Pont, A.C.  & C.J.B. de Carvalho. 1994. Neotropical Fanniidae
Chillcott, J.G. 1961. A Revision of the Nearctic species of Fannii- (Diptera): a key to the Fannia anthracina-group. Entomologist’s
nae (Diptera: Muscidae). The Canadian Entomologist Supple- Monthly Magazine 130: 229–238.
ment 14: 1–295. (1960) Roback, S.S. 1951. A classification of the muscoid calyptrate
Ferrar, P. 1987. A guide to the breeding habits and immature stag- Diptera. Annals of the Entomological Society of America 44:
es of Diptera Cyclorrhapha. Entomonograph  8(1). E.J.  Brill  / 327–361.
Scandinavian Science Press, Leiden, Copenhagen, 478 pp. Vockeroth, J.R. 1966. A method of mounting insects from alcohol.
Griffiths, G.C.D. 1972. The phylogenetic classification of Diptera The Canadian Entomologist 98: 69–70.
Cyclorrhapha, with special reference to the structure of the male Vockeroth, J.R. 1996. Key to the genera of Muscidae (Diptera) of
postabdomen. Series entomologica 8. Dr. W. Junk, The Hague, Mexico, Central America and the West Indies. Memoirs of the
340 pp. Entomological Society of Washington 18: 280–288.
MUSCIDAE (house flies, stable flies) 108
Jade Savage and J.R. Vockeroth

Fig. 108.1. Female of Musca domestica Linnaeus, (MND, fig. 105.1, as male).

Diagnosis posterodorsal bristle (calcar) at about two-thirds from base.

First hind tarsomere without sub-basal ventral seta except
Small to large flies (body length 2–20 mm) (Fig. 1). Stout in some species of Schoenomyza and Drymeia. Abdominal
or slender, generally dull grey, black or yellow but occa- sclerites 6–8 in males usually asymmetric.
sionally metallic blue or green. Male frons narrow to broad, Muscidae can easily be separated from other muscoids by
with one to many frontal bristles. Female frons at least one- a combination of characters. Vein A1 is present and usually
fourth as wide as head, with or without pair of cruciate in- extended halfway to wing margin, not reaching completely
terfrontal bristles. Eyes bare or setulose. Antennal pedicel to wing margin as in most Scathophagidae and Anthomyii-
with distinct dorsal seam, arista almost bare to long plumose dae. The curvature of vein Sc separates most Muscidae from
or pectinate. Proboscis generally with prementum broad Fanniidae: Sc lies close to R1 over most of its length in the
and labella fleshy, but with prementum tapering to slender majority of muscids, while it diverges from R1 at a point near
apex and labella reduced in some, including the Stomoxyini. its base in fanniids and some members of the muscid tribe
Thoracic bristles usually distinct and long, meron without Azeliini. They can further be distinguished from fanniids by
row of strong bristles. Wings well developed, with vein A1 the absence of a mid-dorsal bristle on the hind tibia. Both
incomplete, not reaching wing margin, occasionally with calypters are well developed, but the upper one is reduced in
setulae above or below R1 and R4+5. Both calypters generally Cordiluroides. The first hind tarsomere generally is without
well developed, with upper one reduced in Cordiluroides a sub-basal ventral seta, such as the one seen in the Anthomyii-
Albuquerque. Legs usually slender, but occasionally with dae. The ventral tip of the scutellum is usually lacking the
marked modifications or tufts of fine setae in males. Hind group of soft setulae that is present in Anthomyiidae. A
tibia without true dorsal bristle on apical half, but often with unique synapomorphy of the family is the loss of abdominal

1281
1282
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

spiracles 6 and 7 in females of all genera except in the all families of calyptrate flies without meral bristles, namely
Palearctic genus Achanthiptera Rondani and in the South the Muscidae, Anthomyiidae, Scathophagidae, and Fannii-
American genus Cariocamyia Snyder, in which spiracle dae. However, Kutty et al. (2008) recently presented strong
six is present; Fanniidae, Anthomyiidae, and Scathophagi- support for a paraphyletic Muscoidea caused by the place-
dae all have seven pairs of spiracles in the female abdomen. ment of the Oestroidea as sister group to Anthomyiidae  +
Scathophagidae. While fanniids were long considered to be
a subfamily of muscids, they are now treated as a separate
Biology family, and the Anthomyiidae, rather than the Fanniidae, had
been suggested as a potential sister group to muscids (Mi-
Muscids are adapted to a variety of habitats and lifestyles chelsen, 1991). That relationship was recently challenged by
and can be found almost everywhere except in the most arid Kutty et al. (2008), who placed the Muscidae as the sister
environments. Larval habitats include dung, carrion, garbage, group to Oestroidea  + (Anthomyiidae  + Scathophagidae).
rotting fungi, fresh or decomposing vegetal matter, bird and While much phylogenetic attention has recently been de-
mammal nests, sewage, mud, and running water. The larvae voted to this family (Hennig, 1965; Skidmore, 1985; Pont,
are saprophagous, coprophagous, predaceous, or more rarely 1986a; Carvalho, 2002; Schuehli et al., 2007), the higher
phytophagous, as in the genus Atherigona, which attack the classification of the Muscidae is still not resolved. The most
shoots of a variety of cereal crops. A few species, especially widely accepted classification (Carvalho, 2002) divides
in the genus Philornis, are hematophagous in the larval stages the Neotropical fauna into seven subfamilies, namely the
and develop as subcutaneous parasites of birds. The biology Atherigoninae (with a single genus), Muscinae, Azeliinae,
of adults is also highly varied. Many species are anthophilous Phaoniinae, Cyrtoneurininae, Mydaeinae, and Coenosiinae,
and can be found in large numbers on flowers, especially at but the monophyly of many of these subfamilies was recently
higher elevations and in boreal regions. Adults of the subfam- challenged by Schuehli et al. (2007). At the generic level, the
ily Coenosiinae are generally predaceous, feeding on a vari- large genera Phaonia and Helina are groups in great need of
ety of small insects and arthropods. All members of the tribe revision, as their generic limits are poorly defined and most
Stomoxyini and a few Muscinae are blood feeders and often a of the research done on these taxa has been conducted at a
great annoyance to both humans and livestock. Other groups regional scale.
associated with humans feed as adults on sweat and other body
secretions and/or act as mechanical vectors of diseases, trans-
porting bacteria between contaminated feeding or egg-laying Identification
sites to human dwellings. Economically important species in
Central America include the highly synanthropic house fly Carvalho (2002) provides keys to species for most gen-
Musca domestica Linnaeus, the horn fly Haematobia irritans era found in the Neotropical Region. Unfortunately, many
(Linnaeus), and the stable fly Stomoxys calcitrans (Linnaeus) taxa were left unplaced in his keys, and original descrip-
(see below under each genus for details). tions will often have to be consulted in order to confirm
Although the family is generally known for its pest spe- species-level identifications. Muscids can be collected in
cies, it should be remembered that muscid larvae play an es- large numbers from Malaise and bait traps (feces, carrion,
sential role as decomposers and are intimately involved with rotting vegetables) or by sweeping over flowers and decay-
the breakdown and recycling of organic matter in most eco- ing matter. Minimum handling of the specimens is recom-
systems. Predaceous adults are important contributors to the mended, as they will easily lose bristles, which are often
natural regulation of populations of agricultural and veterinary important for generic and specific level identifications.
pests such as aphids, whitefly, black flies (Simuliidae), and Small specimens (less than 4  mm) collected in alcohol
mosquitoes (Culicidae). should be either chemically or critical-point dried and then
mounted on a cardboard point or glued to a pin, while larg-
er ones can be dried using ethyl acetate (Vockeroth, 1966).
Classification Freshly killed specimens should be pinned after a relaxing
period (24 hours, preferably in a freezer), and depending on
For the last three decades, muscids have been placed in the size, either directly mounted on a regular pin or double-
the Muscoidea sensu McAlpine (1989), a group that includes mounted using a minuten pin.

Key to the genera of Muscidae of Central America

1. Anepimeron at least with few distinct setulae or/and body coloring metallic blue, green, or
purple. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Anepimeron bare or with short and dense golden fuzz (some Hydrotaea Robineau-Desvoidy),
without setulae, and body coloring not strongly metallic blue, green, or purple. . . . . . . . . . . . . 26
MUSCIDAE (house flies, stable flies) 108 1283

2. Proboscis strongly sclerotized, tapered from broad base to slender apex and with labella reduced,
not fleshy (Fig. 2); arista with long microtrichia above, bare or with single microtrichium below
(Fig. 2). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
– Proboscis moderately to weakly sclerotized, not strongly tapered at apex and with labella well
developed and often fleshy (Figs. 3–9); arista various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

3. Palpus about one-third as long as labrum (Fig. 2). . . . . . . . . . . . . . . . . . . . . . . Stomoxys Geoffroy

– Palpus about three-fourths as long as labrum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4. Notopleuron with setulae in addition to the two notopleural bristles; South America, Trinidad. . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Neivamyia Pinto & Fonseca]
– Notopleuron without setulae in addition to notopleural bristles. . . Haematobia Lepeletier & Serville

5. Parafacial with short setae along its length (Fig. 8); palpus moderately to strongly broadened from
base to apex (Fig. 8); frons of male about one-third as wide as head . . . . . . . . . . . . Lispe Latreille
– Parafacial with at most few setae near base of antenna; palpus slender or gradually broadened
from base to apex; frons of male at most one-fourth as wide as head and usually much narrower
(if one-fourth, then first flagellomere about 6 times as long as wide). . . . . . . . . . . . . . . . . . . . . . 6

6. Lower calypter broad, subtruncate posteriorly and with its anteromedian corner extended slightly
under scutellum (Figs. 20, 21). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
– Lower calypter narrow, rounded posteriorly and with its anteromedian corner distinctly lateral to
scutellum (Fig. 19). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

7. First flagellomere about 6 times as long as wide; M vein nearly straight apically, not curved for-
ward; male with frons about one-fourth as wide as head. . . . . . . . . . . . . . . . . . . Charadrella Wulp
– First flagellomere at most 3 times as long as wide; M vein curved at least slightly forward apically
(Figs. 12–14); male with frons at most one-fifth as wide as head. . . . . . . . . . . . . . . . . . . . . . . . . 8

8. Postalar wall setulose (Fig. 23) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Philornis Meinert

– Postalar wall bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

9. General body pruinose or only slightly metallic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

– General body strongly metallic blue, green, or purple. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

10. Suprasquamal ridge setulose (Fig. 21); apex of abdomen often golden yellow; prosternum bare .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sarcopromusca Townsend
– Suprasquamal ridge bare; apex of abdomen not golden yellow; prosternum bare or setulose . . 11

11. Dorsocentrals 2+4; prosternum setulose.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Musca Linnaeus

– Dorsocentrals 0+1, 0+2, 0+3, or 1+2; prosternum usually bare.. . . . . . . . . . . . . . . . . . . . . . . . . 12

12. Katepisternal bristles 0+1; vestiture of scutum, scutellum, katepisternum, and abdomen consisting
mainly of dense and fine setulae, with fringe of 8–10 pairs of long and strong marginal bristles on
scutellum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mesembrina Meigen
– Katepisternal bristles 1+1–2; vestiture of scutum, scutellum, katepisternum, and abdomen not as
above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Morellia (Morellia) Robineau-Desvoidy, in part

13. Suprasquamal ridge and prosternum setulose; dorsocentrals 2+3; general body coloration metallic
green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neomyia Walker
– Suprasquamal ridge bare, prosternum bare or with fine setae; dorsocentrals various; general body
coloration dark metallic blue or green. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
1284
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

pc fr s

inc fr s

2 Stomoxys 3 Azelia 4 Atherigona

rc orb s
oc tr

pc orb s

5 Hydrotaea 6 Hydrotaea 7 Potamia

pafc

8 Lispe 9 Schoenomyza 10 Agenamyia

Figs. 108.2–10. Heads: lateral view of (2) Stomoxys calcitrans (Linnaeus), (illustrated by J. Savage); anterolateral view of (3) Azelia sp.,
(MND, fig. 105.5, as A. cilipes); lateral view of (4) Atherigona orientalis Schiner, (MND, fig. 105.6) and (5) Hydrotaea ignava (Harris),
(Holarctic and Oriental, MND, fig. 105.11); anterolateral view of (6) H. anxia (Zetterstedt), (Holarctic, MND, fig. 105.12); lateral view of
(7) Potamia scabra (Giglio-Tos), (MND, fig. 105.13) and (8) Lispe canadensis Snyder, (Nearctic, MND, fig. 105.3); anterolateral view of
(9) Schoenomyza dorsalis Loew, (Nearctic, MND, fig. 105.8); lateral view of (10) Agenamyia sp., (Couri & Pont, 1999, fig.29).
Abbreviations: inc fr s, inclinate frontal seta; oc tr, ocellar triangle; pafc, parafacial; pc fr s, proclinate frontal seta; pc orb s, proclinate orbital
seta; rc orb s, reclinate orbital seta.
MUSCIDAE (house flies, stable flies) 108 1285

14. Apical portion of stem vein setulose dorsally; prosternum setulose; general body coloration dark
metallic green; Mexico, West Indies, South America.. . . . . . [Morellia (Parapyrellia) Townsend]
– Apical portion of stem vein bare dorsally (may be setulose ventrally); prosternum bare or with
fine setae; general body coloration dark metallic blue or green. . . . . . . . . . . . . . . . . . . . . . . . . . 15

15. Anterior katepisternal bristle absent or weak; prosternum bare with anterior portion notably
narrow; males with anteromedial ommatidia enlarged, similar in diameter to anterior ocellus . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Biopyrellia Townsend
– Anterior katepisternal bristle present and strong; prosternum bare or with fine setae, more or less
parallel-sided; males with anteromedial ommatidia not enlarged as described above . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Morellia (Morellia) Robineau-Desvoidy, in part

C Sc R1

R2+3
R4+5

CuA2 M1

A2 A1
CuA1
11 Neomuscina 12 Neomyia

r-m

dm-cu M1

13 Synthesiomyia 14 Philornis

15 Graphomyia 16 Morellia

17 Muscina 18 Morellia

Figs.  108.11–18. Wings: dorsal view of (11)  Neomuscina tripunctata (Wulp), (MND, fig.  105.21); (12)  Neomyia cornicina (Fabricius),
(MND, fig. 105.22); (13) Synthesiomyia nudiseta (Wulp), (MND, fig. 105.28); (14) Philornis sp., (MND, fig. 105.24); (15) Graphomyia
idessa (Walker), (Holarctic and Oriental, MND, fig. 105.27); (16) Morellia podagrica (Loew), (Holarctic, MND, fig. 105.23); (17) Muscina
levida (Harris), (Holarctic, MND, fig. 105.29); dorsal view of wing base of (18) Mo. (Xenomorellia) sp., (illustrated by J. Savage).
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

16. Anepimeron bare; apical section of M curved strongly forward, ending at or slightly before wing
apex (as in Fig. 16); general body coloration metallic blue. . . . . . . . . Callainireinwardtia Savage
– Anepimeron at least with few setulae; apical section of M various; general body coloration
various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

17. Stem vein, in dorsal view, with long seta near insertion point of humeral vein (Fig. 18). . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Morellia (Xenomorellia) Malloch
– Stem vein, in dorsal view, without long bristle near insertion point of humeral vein but occasionally
with weak setulae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

18. Hind tibia with strong posterodorsal bristle (calcar), at least 1.5 times as long as tibial diameter,
placed at about two-thirds of tibial length (Fig.  25); female with anteriormost orbital bristle
proclinate (Fig. 6). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
– Hind tibia usually without distinct posterodorsal bristle, if such bristle is present, then weak and
no longer than tibial diameter, and apical section of M not strongly curved forward; female with
anteriormost orbital bristle reclinate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

19. Postalar wall setulose (as in Fig. 23); notopleuron with three bristles; apical section of M straight
or nearly so, ending far behind wing apex; prosternum setulose laterally. . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polietina Schnabl & Dziedzicki
– Postalar wall bare; notopleuron with two bristles; apical section of M curved strongly forward,
ending at or slightly before wing apex (as in Fig. 16); prosternum bare. . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Morellia (Trichomorellia) Stein

20. All veins except costa bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemichlora Wulp

– Some part of stem vein of R1 and/or of R4+5 setose dorsally and/or ventrally. . . . . . . . . . . . . . . 21

21. Lower calypter with fine, erect, dark setulae on posterior one-third of upper surface; postalar wall
setulose (as in Fig. 23). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudoptilolepis Snyder
– Lower calypter without setulae above; postalar wall usually bare, but occasionally with one to
three weak setulae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

22. M curved strongly forward apically and ending at or near wing apex (Fig. 11); stem vein usually
with few ventral setulae on apical half; prosternum bare . . . . . . . . . . . . . . Neomuscina Townsend
– M curved only slightly forward apically and ending well behind wing apex (as in Figs. 14, 17);
stem vein without ventral setulae on apical half but occasionally with setulae on basal half;
prosternum setulose or bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

23. All veins except costa bare dorsally, R4+5 with two to five setulae ventrally; anterior pair of frontal
bristles in females much stronger than other frontal bristles, almost as strong as vibrissae. . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Helina Robineau-Desvoidy, in part
– R1 and/or R4+5 setulose on both dorsal and ventral surfaces; anterior pair of frontal bristles
various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

24. Arista pubescent to short plumose, with longest microtrichium no longer than width of
first flagellomere; intra-alars 0+2. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chortinus Aldrich
– Arista long plumose, with longest microtrichium at least 1.5 times width of first flagellomere;
intra-alars 0+1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
MUSCIDAE (house flies, stable flies) 108 1287

25. R1 setulose on ventral surface; ovipositor medium or short and not delicate, without microtrichia
on the membrane between segments 6–7 and 7–8; tergites 6 and 7 sclerotized; cerci short and
flat. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyrtoneuropsis Malloch
– R1 bare on ventral surface; ovipositor long, with microtrichia on the membrane between segments
6–7 and 7–8; tergites 6 and 7 not sclerotized; cerci long and not flat . . . Cyrtoneurina Giglio-Tos

26. Hind coxa with few setulae on posterodorsal surface (Fig. 31). . . . . . . . . . . . . . . . . . . . . . . . . . 27

– Hind coxa bare on posterodorsal surface. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28

27. One medioclinate frontal bristle present (Fig. 3); A1 abbreviated, not reaching more than halfway
between apex of cell cup and wing margin; hind tibia without posterodorsal bristle on apical half
(calcar). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Azelia Robineau-Desvoidy
– More than one medioclinate frontal bristle present; A1 reaching more than halfway to wing
margin; hind tibia with strong posterodorsal bristle on apical half (calcar) (as in Fig. 25); adjacent
Mexico. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Thricops Rondani]

28. Node of Rs (Fig. 15) or extreme base of R4+5 with at least one setula above. . . . . . . . . . . . . . . . 29
– Node of Rs and base of R4+5 bare above (Figs. 13, 17) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

u calyp l calyp
u calyp

l calyp l calyp
sctl spsq rg

19 Neomuscina 20 Musca 21 Sarcopromusca

l prepm s spsq rg
kepst
cx 2
cx 1 l calyp
pal
wall

22 Coenosia 23 Philornis

Figs. 108.19–23. Thoracic structures: dorsal view of calypters and associated structures of (19) Neomuscina tripunctata (Wulp), (MND,
fig.  105.15) and (20)  Musca domestica Linnaeus, (MND, fig.  105.16); dorsal view of lower calypter and suprasquamal ridge of
(21) Sarcopromusca arcuata Townsend, (illustrated by J. Savage); lateral view of anterior part of pleuron of (22) Coenosia tigrina (Fabricius),
(Holarctic, MND, fig. 105.20); lateral view of thorax and scutellum of (23) Philornis sp., (MND, fig. 105.17).
Abbreviations: cx 1, forecoxa; cx 2, midcoxa; kepst, katepisternum; l calyp, lower calypter; l prepm s, lower proepimeral seta; pal wall,
postalar wall; sctl, scutellum, spsq rg, suprasquamal ridge; u calyp, upper calypter.
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

29. Prosternum setulose laterally; first postsutural supra-alar (prealar) bristle absent . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Limnophora Robineau-Desvoidy
– Prosternum bare; first postsutural supra-alar present or absent. . . . . . . . . . . . . . . . . . . . . . . . . . 30

30. Hind tibia with strong posterodorsal bristle at about three-fourths of length (calcar) (Fig. 25) . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phaonia Robineau-Desvoidy, in part
– Hind tibia without calcar. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

31. M not curved forward apically. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mydaea Robineau-Desvoidy

– M slightly to strongly curved forward apically. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32

32. Anterior katepisternal bristle absent, with one posterior katepisternal bristle; katepimeron and
meron with coarse black setulae . . . . . . . . . . . . . . . . . . . . . . . . . Graphomya Robineau-Desvoidy
– One anterior and two posterior katepisternal bristles present; katepimeron and meron bare . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myospila Rondani

33. M curved strongly forward apically and ending well before wing apex (Fig. 13); aristal microtrichia
shorter than basal diameter of arista; lower calypter enlarged and truncated posteriorly; prosternum
setulose.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Synthesiomyia Brauer & Bergenstamm
– M usually straight or curved only slightly forward, but if curved strongly forward and ending near
wing apex, then arista plumose; lower calypter narrow, rounded posteriorly; prosternum bare or
setulose.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

34. Pedicel with dorsal surface flattened (Fig. 29); arista short, only slightly longer than first flagellomere
(Fig. 29); katepisternal bristles 2+2. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
– Pedicel with dorsal surface rounded; arista and katepisternal bristles various . . . . . . . . . . . . . . 36

35. Dorsocentral bristles 2+3; USA, Bahamas, South America. . . . . . . [Pachyceramyia Albuquerque]
– Dorsocentral bristles 2+4; Nearctic, Bahamas, South America. . . . . . . . . . . . . [Tetramerinx Berg]

36. Presutural dorsocentral bristles short, scarcely distinguishable from surrounding setulae, longest
at most one-fourth as long as notopleural bristles; head subquadrate in profile (Fig. 4); antenna
inserted little below level of upper margin of eye (Fig. 4). . . . . . . . . . . . . . . . Atherigona Rondani
– At least one presutural dorsocentral bristle moderately strong, clearly distinguishable from
surrounding setulae, and at least half as long as notopleural bristles; head varied in shape; antenna
usually inserted well below level of upper margin of eye. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37

37. Labella reduced; prestomal teeth developed; first presutural dorsocentral bristle absent or weak
and less than half as long as second and/or katepisternal bristles arranged in nearly equilateral
or isosceles triangle, with lower bristle almost equidistant from the two upper bristles (Fig. 22)
(rarely closer to anterior upper katepisternal bristle in Bithoracochaeta); lower proepimeral bristle
usually curved ventrally or anteroventrally (Fig. 22); first postsutural supra-alar (prealar) bristle
absent; male frons at least one-fifth as wide as head. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
– Labella fleshy or reduced; prestomal teeth present or absent; first presutural dorsocentral bristle
more than half as long as second, but if absent, then labella fleshy, prestomal teeth absent and male
holoptic (aberrant Helina spp.); katepisternal bristles, if more than two in number, with lower or
lower posterior one always closer to upper posterior one than to upper anterior one; lower proepimeral
bristle curved dorsally; first postsutural supra-alar (prealar) present or absent; male frons varied in
width. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
MUSCIDAE (house flies, stable flies) 108 1289

38. Scutum with two pairs of postsutural dorsocentral bristles; first flagellomere with apex pointed;
hind tibia with long posteroventral bristle at midlength and without midventral bristle. . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bithoracochaeta Stein
– Scutum with three pairs of postsutural dorsocentral bristles; first flagellomere with apex usually
rounded, rarely pointed; hind tibia without long posteroventral bristle, but sometimes with
midventral bristle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39

39. Frons wider than long (Fig. 9); midtibia with anteroventral bristle. . . . . . . . Schoenomyza Haliday
– Frons longer than wide; midtibia without anteroventral bristle. . . . . . . . . . . . . . . . . . . . . . . . . . 40

40. Hind femur with two preapical bristles on anterodorsal and posterodorsal surfaces.. . . . . . . . . 41
– Hind femur with three preapical bristles on anterodorsal, dorsal, and posterodorsal to posterior
surfaces (Fig. 30). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43

pd s

ad s

pd s
(calcar)

ds ddss
ad s

24 Drymeia 25 Phaonia 26 Brontaea 27 Spilogona 28 Hydrotaea

ds
cx 3
depressed
area
ps
trc 3
ps

30 Neodexiopsis
31 Thricops

29 Tetramerinx
32 Lispoides 33 Brontaea
Figs. 108.24–33. Leg segments, antenna, and abdominal sternite 1: dorsal view of hind tibia of (24) Drymeia quadrisetosa (Malloch), (Nearctic,
MND, fig. 105.32); (25) Phaonia bysia (Walker), (Nearctic, MND, fig. 105.34); anterior view of hind tibia of (26) Brontaea humilis
(Zetterstedt), (Holarctic, MND, fig. 105.40) and (27) Spilogona almquisti Holmgren, (Holarctic, MND, fig. 105.41); posterior view of femur
and tibia of foreleg of (28) Hydrotaea anxia (Zetterstedt), (Holarctic, MND, fig. 105.42); lateral view of antenna of (29) Tetramerinx littoralis
(Malloch), (Nearctic, MND, fig. 105.14); posterior view of hind femur of (30) Neodexiopsis ovata Stein, (Nearctic, MND, fig. 105.43, as
Coenosia); posterolateral view of hind coxa and trochanter of (31) Thricops innocuus (Zetterstedt), (Holarctic, MND, fig. 105.45); ventral
view of abdominal sternite 1 of (32) Lispoides aequifrons (Stein), (Nearctic, MND, fig. 105.44) and (33) B. humilis (Zetterstedt), (Holarctic,
MND, fig. 105.46).
Abbreviations: ad s, anterodorsal seta; cx 3, hind coxa; d s, dorsal seta; pd s, posterodorsal seta; p s, posterior seta; trc 3, hind trochanter.
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

41. Apical scutellar bristle reduced, much shorter than basal bristle. . . . . . . . . Pilispina Albuquerque
– Apical scutellar bristle as long or longer than basal bristle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42

42. Gena greatly reduced, almost linear (Fig. 10); ovipositor of medium length; tergites and sternites
broad. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Agenamyia Albuquerque
– Gena of moderate width; ovipositor long; tergites and sternites long and narrow . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Coenosia Meigen

43. One postsutural intra-alar bristle present; ocellar bristles much shorter than postocellar bristles. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cordiluroides Albuquerque
– Two postsutural intra-alar bristles present and/or ocellar bristles at least as strong as postocellar
bristles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neodexiopsis Malloch

44. Hind tibia with three or more posterodorsal bristles that are more or less uniformly distributed
along tibial length and are subequal in length and strength to anterodorsal bristles (Fig. 24); legs
entirely black; Holarctic and Oriental Regions. . . . . . . . . . . . . . . . . . . . . . . . . [Drymeia Meigen]
– Hind tibia usually without posterodorsal bristles or with at most two bristles; if three or more are
present, they are restricted to basal half of tibia and are much shorter than anterodorsal bristles;
legs sometimes partly yellow. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45

45. Hind tibia with strong posterodorsal bristle on apical half (calcar) (Fig. 25), if calcar present but
weak, then M vein curved forward (Fig. 17). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
– Hind tibia with calcar absent, at most with one to two short bristles on basal half, curvature of
M various. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51

46. Eye extending below level of major vibrissa; gena narrow, at most one-sixth as high as eye
(Figs. 5, 6); aristal microtrichia not longer than basal diameter of arista (Figs. 5, 6); male sometimes
with apical part of forefemur and basal part of foretibia excavated ventrally, the former with one
to two acute processes near base of excavation (Fig. 28); dorsocentral bristles 2+4; female with
interfrontal bristles present and with anteriormost orbital bristle proclinate (Fig. 6) . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrotaea Robineau-Desvoidy
– Eye usually not extending below level of major vibrissa; gena broader, at least one-fourth as
high as eye; aristal microtrichia varied in length; male with forefemur and foretibia normal, not
excavated; dorsocentral bristles 2+3–4; female with interfrontal bristles present or absent and
anteriormost orbital bristle proclinate or reclinate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47

47. Gena anteriorly with one to three strong upcurved bristles slightly above level of weaker bristles
on lower margin of gena (Fig.  7); dorsocentral bristles 2+3; female with anteriormost orbital
bristle proclinate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
– Gena without strong upcurved bristles; dorsocentral bristles 2+3–4; female with anteriormost
orbital bristle reclinate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49

48. Aristal microtrichia about twice as long as basal diameter of arista (Fig.  7); large species,
4.5–6.5 mm long; Nearctic Mexico, Peru . . . . . . . . . . . . . . . . . . . . [Potamia Robineau-Desvoidy]
– Aristal microtrichia no longer than basal diameter of arista; small species, 2.0–4.0 mm long. . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Micropotamia Carvalho

49. Anterior postsutural intra-alar bristle situated slightly behind level of supra-alar bristle;
M moderately curved apically (Fig. 17) . . . . . . . . . . . . . . . . . . . . . . Muscina Robineau-Desvoidy
– Anterior postsutural intra-alar bristle situated in front of level of supra-alar bristle; curvature of M
various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
MUSCIDAE (house flies, stable flies) 108 1291

50. Postalar wall setulose (as in Fig. 23). . . . . . . . . . . . . . . . . . . . . Chaetophaonia Carvalho & Nihei

– Postalar wall bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phaonia Robineau-Desvoidy, in part

51. Katepimeron and prosternum setulose. . . . . . . . . . . . . . . . . . . Helina Robineau-Desvoidy, in part

– Katepimeron bare; prosternum bare or setulose. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52

52. Sternite 1 setulose (Figs. 32, 33); first postsutural supra-alar (prealar) bristle usually absent, but if
visible, then less than half as long as second notopleural; hind tibia with anterodorsal apical bristle
various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
– Sternite 1 bare; first postsutural supra-alar bristle present or absent; hind tibia with anterodorsal
apical bristle strong (Fig. 27). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54

53. Sternite 1 with few strong setae near posterior margin (Fig. 33); frons of male narrow, frontal
vitta obliterated on most of upper half of frons; hind tibia with anterodorsal apical bristle weak
(Fig. 26). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brontaea Kowarz
– Sternite 1 with fine setulae on most of its surface (Fig. 32); frons of male wide, frontal vitta
distinct throughout; hind tibia with anterodorsal apical bristle strong (as in Fig. 27). . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lispoides Malloch

54. Arista pubescent, longest microtrichium no longer than basal diameter of arista; first postsutural
supra-alar (prealar) bristle absent; legs entirely black; found in all biogeographic realms but no
published records for Central America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Spilogona Schnabl]
– Either arista with longest microtrichium much longer than basal diameter of arista or first postsutural
supra-alar (prealar) bristle present or legs partly to entirely yellow . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Helina Robineau-Desvoidy, in part

Synopsis of the fauna decaying plant material or animal matter, and A. orientalis is
known to produce secondary infestations in a few crop plants
Muscids currently account for more than 4000 species such as sorghum and maize.
worldwide. There are 843 species recognized in the Neotro-
pical Region (Carvalho et al., 2005), and approximately 150 Azelia Robineau-Desvoidy. This small genus occurs
in most geographical regions. There is at least one species
of these have so far been recorded in Central America as defined
found in Middle America (Neotropical Mexico), and the
in this manual. Unless stated otherwise, species numbers
single male specimen examined here partially matches the
listed below have been taken from the most recent catalog of original description of A. neotropica Snyder (Snyder, 1957),
Muscidae of the Neotropical region (Carvalho et al., 2005). but differs in a few characters, suggesting that it might be
Except where indicated, Carvalho (2002) is the best a new species. The larvae of Azelia are found in vertebrate
source of keys for all Central American species and will excrement (horses and cattle) and in the final instar will prey
not be repeated. Similarly, Skidmore (1985) and Ferrar upon small dipteran larvae.
(1987) were the main references on the biology of adults
and immatures, respectively, and are therefore not cited Biopyrellia Townsend. This genus was treated as a syn-
in the following section. The number of species recorded onym of Morellia by Pont (1972), but is now believed to de-
from Central America is likely to increase substantially for serve full generic status (Pamplona, 1986, Nihei & Carvalho,
most genera as our knowledge of the fauna of this region 2007). The genus has a single species, B. bipuncta (Wiede-
improves. mann), broadly distributed in the Neotropical Region.
Agenamyia Albuquerque. This genus is strictly Neotro- Bithoracochaeta Stein. This exclusively New World ge-
pical and contains two described species. Only one of them, nus has three species found in Neotropical Mexico.
A. exotica Carvalho & Couri, is known from Central Amer-
ica (Panama). Brontaea Kowarz. This genus is found nearly worldwide
with the exception of New Zealand and Oceania (Couri  &
Atherigona Rondani. This genus has over 195 described Carvalho, 2002). There are four Neotropical species, two
species worldwide (Pont, 1986b), but only one of them, the present in Neotropical Mexico. The larvae are found in mam-
introduced A. orientalis Schiner, occurs in the New World. malian (and rarely avian) dung and are highly carnivorous in
Often called shoot-flies, Atherigona species will breed in the final instar.
1292
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Callainireinwardtia Savage. The single species of this Afrotropical Regions (Vockeroth, 1972). Two species are
recently described genus, C.  woodi Savage, is only known recorded from Central America. The larvae are found in liq-
from Costa Rica, where all specimens have been collected uid or semi-liquid substrates and are highly predaceous, es-
above 1000 m (Savage, 2009). pecially on syrphid and ptychopterid larvae. The adults are
mainly anthophilous.
Chaetophaonia Carvalho  & Nihei. This recently de-
scribed Nicaraguan genus was known until 2005 under the Haematobia Lepeletier & Serville. This genus is present
preoccupied name Maesia (Carvalho et al., 2005). Car- in all biogeographical regions, with a single widespread spe-
valho & Nihei (2002) state that the two known species are cies in Middle America, the almost cosmopolitan horn fly,
the only Neotropical species of Phaoniinae with a setulose H.  irritans. Adults are a great nuisance to cattle on whose
postalar wall, but Helina rubripalpis Wulp, found in Costa blood they feed and will only leave their hosts to oviposit
Rica, also has setulae on the postalar wall. or sip moisture from fresh cowpats. Often abundant on hot
days, this species can cause a decrease in milk production,
Charadrella Wulp. This exclusively Neotropical genus but it is not believed to be involved in the transmission of
has three described species, but only one of them, C. mac- important livestock diseases (Greenberg & Povolný, 1971).
rosoma Wulp, is found in Central America. Nothing is
known of the biology of C. macrosoma, but C. malacophaga Helina Robineau-Desvoidy. Helina is most probably
Lopes is viviparous, depositing late second or third instar not a monophyletic assemblage of species (Carvalho  &
larvae, and is believed to parasitize and/or breed in dead Couri, 2002). This large and diverse genus is present in all
gastropods. biogeographical regions, with more than ten species occur-
ring in Central America. The genus Xenothoracochaeta Mal-
Chortinus Aldrich. Chortinus bequaerti Malloch, one loch, whose members usually have a setulose anepimeron,
of the two species belonging to this Neotropical genus, is has recently been synonymized with Helina (Carvalho et
found in Mexico, Panama, and Costa Rica. Vockeroth (1996) al., 2005). This taxon was formerly known only from South
suggested this genus might be a synonym of Cyrtoneurina, America, but we have seen specimens of at least two dif-
but Pamplona & Couri (1998) treated it as a separate genus. ferent species from Costa Rica and Panama with a setulose
Chortinus bequaerti is a larviparous species (Albuquerque & anepimeron. In warm climates, most species of Helina typi-
Lopes, 1983). cally breed in decaying trees or other plant material, where
the larvae are predaceous on other insect larvae.
Coenosia Meigen. This large cosmopolitan genus con-
tains over 360 described species. Surprisingly, in spite of Hemichlora Wulp. The only described species,
there being 39 species present in the Neotropical Region, this H. scordalus (Walker), is restricted to Mexico, Guatemala,
genus had not been previously recorded in Central America. Costa Rica, and Brazil. Little is known of the biology of this
We have seen at least two species from Costa Rica. Both group but a dry female specimen with a larva extending from
larvae and adults of Coenosia are predaceous. The larvae are the tip of the abdomen was recently reported by D.M. Wood
found in humus soil and damaged plants infested with other
(personal communication).
fly larvae, while the adults hunt other insects in grassy habi-
tats. Some species are being used in biocontrol programs in Hydrotaea Robineau-Desvoidy (including Ophyra Rob-
greenhouses. ineau-Desvoidy). This is a diverse genus found in all bio-
geographical regions. There has been some disagreement on
Cordiluroides Albuquerque. Three of the six known spe-
the status of Ophyra, treated as a full genus by Pamplona &
cies of this Neotropical genus are found in Central America
Couri (1989), Carvalho et al. (1993), Carvalho  & Couri
(Couri et al., 2006).
(2002), Schuehli et al. (2004), and Carvalho et al. (2005).
Cyrtoneurina Giglio-Tos. This is a Neotropical genus However, Pont (1986a, 1989), Vockeroth (1996), and Sav-
with ten described species, three of which are known from age  & Wheeler (2004) consider Ophyra a junior synonym
Central America. They are viviparous, with larvae (if feeding of Hydrotaea. A modified forefemur in the male, along with
at all) apparently coprophagous in the final instar. a strong upcurved genal bristle and a short pruinose fron-
tal triangle in females, have often been used as characters
Cyrtoneuropsis Malloch. This Neotropical genus was of- to separate Hydrotaea from Ophyra. While these characters
ten treated as a synonym of Cyrtoneurina Giglio-Tos (Vock- seem to hold when dealing exclusively with the Neotropical
eroth, 1996; Pont, 1972) but is now considered a valid genus fauna, they do not apply to the Nearctic fauna, where at least
mainly based on characters of the female ovipositor (Pam- two species of Hydrotaea do not have the front legs modi-
plona, 1999). It contains over 30 described species, about fied. The females of a number of species of Hydrotaea also
half of which occur in Central America. do not have strong upcurved genal bristles, while others have
a well-developed shiny frontal triangle and are otherwise
Graphomya Robineau-Desvoidy. Essentially a world- indistinguishable from Ophyra females (e.g., the Palearc-
wide genus, found in temperate and tropical areas except tic H. glabricula (Fallén)). It is for these reasons, then, that
New Zealand and Oceania (one introduced species in we treat Ophyra as a junior synonym of Hydrotaea. While
Hawaii), Graphomya is most diverse in the Oriental and Hydrotaea sensu lato is relatively diverse in the Neotropi-
MUSCIDAE (house flies, stable flies) 108 1293

cal Region, only two species are found in Central America Subgenus Morellia sensu stricto. This subgenus is present
and only one has modified forelegs in the male. The larvae in most biogeographical regions, with over 15 Neotropical
are found in a variety of substrates, including, among oth- species, at least two of which are found in Central America.
ers, dung, carrion, decaying plant material, and bird nests. Larvae are saprophagous in dung of ungulates, especially
Hydrotaea dentipes (Fabricius) breeds in mammalian dung cattle, and the adults of many species are sweat flies.
and feeds on other dipteran larvae in the last instar. It is also Subgenus Trichomorellia (Stein) (=Dasymorellia Mal-
capable of decimating local populations of Stomoxys calci- loch). Known until 2005 under the name Dasymorellia (see
trans larvae (Greenberg  & Povolný, 1971). In Guatemala, Pont et al., 2005 for details), this Neotropical taxon con-
H. aenescens (Wiedemann), which often breeds in human re- tains eight described species, one of which occurs in Central
fuse sites, has been recorded in great abundance in kitchens, America (Carvalho et al., 2005; Nihei & Carvalho, 2007).
where it acted as a major contaminant of food (Shannon & Subgenus Xenomorellia Malloch. This is a small Neo-
del Ponte, 1926). tropical subgenus with two described species. Although
there are no published records of this group for Central
Limnophora Robineau-Desvoidy (including Syllimno- America, we identified specimens from Honduras and
phora Speiser (see Vockeroth, 1996 for comments on synony- Costa Rica.
my) and Neotropical “Heliographa” Malloch species). This is
a large genus, well represented in all biogeographical regions. Musca Linnaeus. This is an Old World genus with two
There are six species found in Central America. The larvae introduced species in the New World. These are the cosmo-
are generally aquatic and found in running water, where they politan house fly, M. domestica, widespread in the Nearctic
feed on oligochaetes and small insect larvae. Adults are pre- and Neotropical Regions, and M.  autumnalis De Geer, the
daceous, but rarely observed taking prey. face fly, having now reached Mexico from the north. Most
Musca species will favor dung as a breeding medium, but
Lispe Latreille. Two of the twelve Neotropical species of the house fly will just as easily develop in almost any kind of
this almost cosmopolitan group are found in Central Ameri- decaying animal or vegetal matter. Highly synanthropic, the
ca. Larvae develop in mud with high organic content and are adults will move readily between garbage, dung, or carrion to
obligate carnivores. Adults are highly predaceous on other human food, carrying a variety of micro-organisms, particu-
insects and often exhibit striking sexual dimorphism. larly those associated with diseases of the human alimentary
tract. The larvae of the face fly are essentially coprophagous,
Lispoides Malloch. This is a New World genus, contain- favoring cow dung, and the adults are mainly sweat-feeding
ing 19 Neotropical species. A single species has so far been pests of livestock.
recorded from Central America, L.  guatemala Snyder. The
larvae are aquatic, and both the immature and adult stages Muscina Robineau-Desvoidy. This genus is found in the
are presumably predaceous. New World and the Palearctic Region. At least one of the
three Neotropical species are found in Central America. The
Mesembrina Meigen. There is a single Neotropical spe- larvae are facultative carnivores and highly polyphagous.
cies of this mainly Holarctic and Oriental group (Kuchta & The cosmopolitan M. stabulans (Fallén), widespread in Cen-
Savage, 2008). Mesembrina nigribasis Kuchta and Savage tral America, can cause serious ulcerations on living tissues
is so far known only from a number of high elevation sites of chordates, as the larvae will feed on both healthy and ne-
in Costa Rica. The larvae of most Mesembrina species are crotic tissues. On the other hand, the same species will also
found in mammalian dung, where the larvae are facultatively heavily feed on larvae of Musca domestica, thus contributing
saprophagous or predaceous. to a reduction in housefly populations.
Micropotamia Carvalho. This genus was originally Mydaea Robineau-Desvoidy. Found mainly in the Hol-
proposed for three South American species, but Phorbia arctic, Neotropical, and Oriental Regions, there are between
fuscisquama Wulp, known only from Mexico, was recently 17 and 20 species of this genus recognized from the Neotro-
transferred to the group (Carvalho & Couri, 2002). We have pical Region, with at least three found in Central America.
also seen one undescribed species from Costa Rica. Adults The larvae are found in excrement, fungi, or rotting vege-
of Micropotamia amazonica (Albuquerque  & Lopez) were table matter and are probably predaceous on other larvae in
collected using a Shannon trap baited with decomposing or- these media.
ganic matter (Carvalho, 1993).
Myospila Rondani. A widespread genus, most diverse
Morellia Robineau-Desvoidy. Many recent changes in the Oriental Region, there are six Neotropical species of
have been brought to the taxonomy of Morellia, and a Myospila, two of which are widespread in Central America.
number of recent synonymies have widened the scope of this Breeding media of the larvae include dung and rotting fruit
group (Nihei & Carvalho, 2007). As all four subgenera key and vegetables.
out in different parts of the key, we have decided to divide
the synopsis section of Morellia to follow these subdivisions Neodexiopsis Malloch. This is a New World genus with 87
(only three subgenera have a confirmed Central American of the 96 described species found in the Neotropical Region.
distribution). There are at least seven species found in Central America.
1294
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

The biology of Neodexiopsis is poorly known, but larvae of Pilispina Albuquerque. This exclusively Neotropical
N. basalis (Stein) have been found in the galleries of scolytid genus contains six described species, all restricted to South
beetles under the bark of pine trees. America (Brazil), with the exception of P. medinai (Snyder),
also recorded from Puerto Rico (Couri, 2000). We have seen
Neomuscina Townsend. This New World genus has over undetermined material of Pilispina from Costa Rica.
30 described species, about eight of which are found in Cen-
tral America. Their biology is poorly known, but the larvae Polietina Schnabl  & Dziedzicki. This is a Neotropical
will breed in human feces, and the adults of a few species genus with 18 described species. No species are recorded for
can be found inside houses in Central America; thus, they are Central America, but we have seen specimens from Hondu-
potentially of hygienic concern. ras and Costa Rica. The genus was presumed viviparous by
Skidmore (1985). We have examined an undetermined dry
Neomyia Walker. This genus has a worldwide distribu- female specimen with a larva extending from the tip of the
tion, but only N. cornicina (Fabricius) has been introduced abdomen.
into the New World and is known from Central America and
Brazil. The coprophagous larvae are found exclusively in bo- Pseudoptilolepis Snyder. This is a Neotropical genus
vine dung and are prone to predation by other muscid larvae. with six described species, four of which are found in Cen-
Adults are anthophilous and especially attracted to yellow tral America.
flowers, and they will also be found feeding on human and
animal excrement. Sarcopromusca Townsend. Treated as a synonym of
Morellia by Pont (1972) and Vockeroth (1996), this group
Phaonia Robineau-Desvoidy (including Dolichopha- was reinstated by Pamplona (1992). It is a Neotropical genus
onia Carvalho). Phaonia is a large cosmopolitan genus with with only two described species, both of which occur in Cen-
more than 760 described species worldwide. It is highly tral America (Nihei, 2005).
heterogeneous and probably paraphyletic (Hennig, 1965;
Huckett  & Vockeroth, 1987; Carvalho, 1989). Vockeroth Schoenomyza Haliday. This genus is found in all
(1996) synonymized the genus Dolichophaonia with Pha- biogeographical regions except Australasia, and most
onia, on the basis that only females could be properly identi- species (18) are described from the Neotropical Region.
fied to genus. Carvalho & Couri (2002) rejected Vockeroth’s Only one species, S.  aurifrons Malloch, occurs in Cen-
tral America (Mexico, Costa Rica).
synonymy, arguing that characters of the female proboscis
and the ovipositor are enough to support the monophyly of Stomoxys Geoffroy. This is an Old World genus with
Dolichophaonia. We agree here with Vockeroth (1996), treat- only the cosmopolitan stable fly, S.  calcitrans, wide-
ing Dolichophaonia as a synonym of Phaonia. The genus in- spread in Central and South America, undoubtedly as
cludes more than 50 Neotropical species, seven known from an immigrant. All larval instars are saprophagous in
Central America. The larvae are obligate carnivores on other manure heaps and decaying vegetable matter. Adults
insect larvae and are found in sap runs, decaying trees, fungi, of both sexes are obligate blood feeders on a variety of
vertebrate and invertebrate carrion, decaying vegetal matter, mammals and are a great annoyance to both livestock
and humus soil. In the Palearctic Region, many adults are and man. In addition to causing decreased milk produc-
anthophilous, especially at higher elevations (Pont, 1993). tion in cows, S.  calcitrans also transmits many species
of trypanosomes.
Philornis Meinert. This is a New World genus but with
the great majority of species restricted to the Neotropical Re- Synthesiomyia Brauer  & Bergenstamm. The single
gion. There are about 50 described species, 11 of which are species of this pantropical genus, Synthesiomyia nudiseta
found in Central America. Larvae are usually associated with (Wulp), is widespread in Central and South America. Larvae
bird nests, where they either act as scavengers in nest mate- are apparently facultatively saprophagous or carnivorous,
rial, or more frequently, suck the blood of nestlings. Some found in rotting meat, decaying vegetables, garbage, carri-
are subcutaneous endoparasites, forming boils under the skin on, feces, and dead insects. Adults are sun-lovers and rarely
of host birds. found indoors.

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tion of Muscini (Diptera, Muscidae). Zoological Journal of the
Vockeroth, J.R. 1972. A review of the World Genera of My-
Linnean Society 149: 493–532.
Pamplona, D.M. 1986. Revalidição de Biopyrellia Townsend, daeinae, with a Revision of the Species of New Guinea and
1932 com redescrição de B. bipuncta (Wiedemann, 1830) (Dip- Oceania (Diptera: Muscidae). Pacific Insects Monograph
tera–Muscidae–Muscinae). Revista Brasileira de Biologia 46: 29: 1–134
49–54. Vockeroth, J.R. 1996. Key to the genera of Muscidae (Diptera)
Pamplona, D.M. 1992. Revalidição de Sarcopromusca Townsend, of Mexico, Central America, and the West Indies. Memoirs
1927 com redescrição de S. pruna (Shannon & Del Ponte, 1926) of the Entomological Society of Washington 18: 280–288.
1296
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
CALLIPHORIDAE (BLOW FLIES) 109
John Vargas and D. Monty Wood

Fig. 109.1. Female of Calliphora terraenovae Macquart, (Nearctic, MND, fig. 106.1).

Diagnosis row, sometimes preceded by fourth smaller bristle; or with

the middle bristle displaced anteriorly, in Mesembrinellinae;
Medium-sized to large flies (body length 5–15  mm) rarely, inner bristle lacking. Meron with row of well-developed
(Fig.  1), usually shiny, metallic green, blue, or purple, at bristles. Wing with bend in vein M.
least on abdomen. Frons of male narrower, and eye larger, Calliphorids share with Sarcophagidae and almost all Ta-
than that of female of the same species, and with few excep- chinidae the presence of a bend in vein M, combined with
tions lacking proclinate orbital bristles (Fig. 2); chaetotaxy a row of well-developed meral bristles. Some Muscidae in
otherwise similar in both sexes. Lowest frontal bristle arising the subfamily Muscinae also have a bend in vein M and may
at level of base of antenna (Fig. 3). Females of Mesembrinel- even be metallic green (Neomyia Walker) or blue (Morellia
linae with single pair of erect crossed bristles on frontal vitta Robineau-Desvoidy) and thus are frequently mistaken for
anterior to ocellar triangle, between frontal bristles. Gena blow flies, but they lack meral bristles. Most calliphorids,
with well-developed genal dilation. Vibrissa usually aris- especially the common species, are easily distinguished
ing above level of lower facial margin, accompanied by four from nearly all Sarcophagidae and Tachinidae by their shiny
or more short subvibrissal bristles in Chrysomyinae, Calli- metallic blue or green color, especially on the abdomen. A
phorinae, and Melanomyinae, or by only one or two large few Tachinidae (Chlorohystricia Townsend, Chrysotachina
bristles in Mesembrinellinae. Facial ridge usually bristled Brauer & Bergenstamm) and Sarcophagidae (some subgenera
more than halfway to base of antenna. Pedicel with single of Lepidodexia Brauer & Bergenstamm) also have metallic
elongate bristle among few shorter setae. First flagellomere green or blue abdomens and therefore could be misidentified
similar in length in both sexes, or slightly longer in female. as calliphorids, but tachinids have a large convex subscutellum
Arista long-plumose dorsally and ventrally. Palpus clavate, while metallic sarcophagids have four notopleural bristles.
reduced in length in Cochliomyia. Notopleuron with two Non-metallic calliphorids, including some Mesembrinella
setae. Postpronotum with three major bristles in transverse and all Angioneura, are superficially similar to some muscids

1297
1298
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

or tachinids. The presence of erect hairlike setae under the anteriorly and the posterior lappet is the larger and forms an
scutellum is diagnostic for Mesembrinella. operculum over the spiracular opening. In the majority of
In almost all Calliphoridae (except Lucilia cuprina (Wie- species of Mesembrinella, the anterior spiracle has an oval
demann) and members of Angioneura), the posterior tho- or droplet-shaped opening near its upper end (Fig. 9), while
racic spiracle is larger, relative to body size, than that in the posterior spiracle is exceptionally large, kidney-shaped,
Sarcophagidae and Tachinidae. It is rounded at its anterior and is as long as the meron, extending anteriorly nearly to
end and the anterior lappet is larger than the posterior lappet the anterior margin of the katatergite (Fig.  7). Its opening
(Fig.  6), while in Sarcophagidae, in most Tachinidae, and is nearly occluded by a continuous fringe arising from the
in the above-mentioned calliphorids (Fig.  8), it is angular anterior, ventral, and posterior spiracular margins.

rc orb s i vt s
oc s
o vt s
pc orb s

fr s

pafc

gn grv
vb
gn
plp

lbl
lbl

2 Cochliomyia 3 Pollenia

a spal s Sc
R1
ial s R2+3
stm vn
npl s R4+5
M

CuA1
CuA2

tym pit A1+CuA2

A2

l calyp

4 Compsomyiops 5 Calliphora

Figs. 109.2–5. Heads and wing bases: lateral view of head of (2) Cochliomyia macellaria (Fabricius), (MND, fig. 106.2) and (3) Pollenia sp.,
(Holarctic, MND, fig. 106.4); dorsal view of right side of thorax and wing base of (4) Compsomyiops callipes (Bigot), (MND, fig. 106.4, as
Paralucilia wheeleri) and (5) Calliphora sp., (Nearctic, MND, fig. 106.12).
Abbreviations: a spal s, anterior supra-alar seta; fr s, frontal seta; gn, gena; gn grv, genal groove; ial s, intra-alar seta; i vt s, inner vertical seta;
lbl, labellum; l calyp, lower calypter; npl s, notopleural seta; oc s, ocellar seta; o vt s, outer vertical seta; pafc, parafacial; pc orb s, proclinate
orbital seta; plp, palpus; rc orb s, reclinate orbital seta; stm vn, stem vein; tym pit, tympanal pit; vb, vibrissa.
CALLIPHORIDAE (BLOW FLIES) 109 1299

Biology other vertebrate scavengers. Adult calliphorids are the first

insects to discover and colonize vertebrate remains, includ-
The larval habits of Calliphoridae vary considerably from ing human bodies. For the past 20  years, forensic studies
one subfamily to another. The most common and well-known have used them to determine time of death, movement of
species, known as blow flies, belong to the Chrysomyinae the body, presence of toxic substances in putrefying corpses,
and Calliphorinae and develop in carrion. Eggs are laid soon postmortem traumas, assessment of improper care or aban-
after the demise of an animal and hatch within hours. The donment of disabled persons, and inadequate management
quickly developing larval aggregations, which may consist or storage of food products, among other applications. In
of various species, undertake a rapid and efficient consump- Costa Rica, calliphorids were first used in 1988 in forensic
tion of tissues through a combination of enzymatic and ex- applications to determine time of death and movement of
ternal mechanical activity (Hobson, 1932a,  b). Adults of bodies. Eight species of calliphorids, all belonging to Chry-
both sexes feed on virtually all forms of carrion and feces somyiinae and Calliphorinae, viz. Cochliomyia macellaria
as well as on honeydew and nectar. Adults have been shown (Fabricius), Hemilucilia segmentaria (Fabricius), H.  semi-
by Jirón  & Hedström (1985) to be important pollinators diaphana (Rondani), Chrysomya megacephala (Fabricius),
of mango, Mangifera indica, and in mango orchards when C.  rufifacies (Macquart), Lucilia eximia (Wiedemann),
these trees are in bloom, adults of both sexes of several spe- L. purpurescens (Walker), Calliphora irazuana (Townsend),
cies may be encountered in large numbers. and Chloroprocta idioidea (Robineau-Desvoidy), are the
These flies are extremely important as consumers of dead species most frequently collected from human bodies.
vertebrates and recyclers of nutrients (Byrd & Castner, 2001). It is probably not a large step from feeding on carrion to
Females of most species oviposit on carrion, while a few can wound parasitism, and Cochliomyia hominivorax (Coquer-
survive on feces and decomposing vegetation. In most parts el), the infamous primary screwworm, is an obligate para-
of the world they are among the most important agents dis- site that has caused more economic losses than any other
posing of carrion, their chief competitors being vultures and calliphorid in the New World. Females of this species are

a lappet
p lappet p spr

hlt

mr s

6 Calliphora 7 Mesembrinella

pprn s

anepst

eye
a spr

prepm s

hd

8 Lucilia 9 Mesembrinella
Figs. 109.6–9. Lateral view of posterior thoracic spiracle of (6) Calliphora irazuana Townsend, (7) Mesembrinella bicolor (Fabricius), and
(8) Lucilia cuprina (Wiedemann); lateral view of anterior thoracic spiracle of (9) M. bicolor. Figures 6–9 illustrated by A. Brenes.
Abbreviations: a lappet, anterior lappet of posterior thoracic spiracle; anepst, anepisternum; a spr, anterior thoracic spiracle; hd, head; hlt, halter;
mr s, meral seta; p lappet, posterior lappet; p spr, posterior thoracic spiracle; prepm s, proepimeral seta; pprn s, postpronotal seta.
1300
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

attracted to fresh cuts or scratches on larger mammals, and mystery. In spite of various tests using decomposing animal
the newly hatched larvae penetrate and enlarge these lesions, and vegetable substrates, Guimarães (1977) and co-workers
soon invading healthy tissue. As bacterial infection sets in, were able to rear only a single adult of one South American
females of other calliphorids, such as the secondary screw- species, Laneella nigripes Guimarães, from dead snails. The
worm, Cochliomyia macellaria, are attracted and begin flies’ use of an obscure host such as snails is worthy of further
ovipositing, eventually leading to the death of the host if investigation, and they may prefer living hosts to dead ones,
left untreated. Since early settlement of the New World, the which were not investigated by these researchers. Although
primary screwworm has inflicted such enormous damage on adult Mesembrinellinae are readily attracted to dung, so are
livestock that a technique was developed in Texas for rear- many other muscoid flies, which do not necessarily breed
ing, sterilizing, and releasing males in vast numbers, thus there. The possibility of their being predacious on muscid
outnumbering the wild male population (Knipling, 1955). or sarcophagid larvae that inhabit dung appears not to have
Female flies mating with these sterile males produced no been investigated.
offspring, and the population declined precipitously. The Snails of the genus Succinia are the hosts of larvae of the
species was eliminated from the USA by 1966, from Mexico obscure parasitoid genus Angioneura (subfamily Melano-
by 1991, from Belize and Guatemala by 1999, and from Costa myinae), at least in the Nearctic Region (Reinhard, 1929).
Rica by 2000. Efforts to eradicate the species continue in After completing their development, larvae form puparia
Panama and the West Indies. However, the species continues inside the shell of the snail.
to thrive in South America. In the absence of C. hominivorax, Although not occurring in Central America, the larvae of
secondary screwworms no longer attack livestock but other calliphorid groups are known to suck blood. In Africa,
develop only in carrion. larvae of various species of Auchmeromyia Brauer  &
Eggs are fertilized as they pass from the ovaries to the Bergenstamm suck the blood of man, warthogs, and other
vagina and are not stored (at least not in New World species) mammals. In the Holarctic Region, species of Protocalliphora
in an ovisac or “uterus” (as they are in all Sarcophagidae Hough living in the nests of birds suck blood from the nest-
and most Tachinidae). A few may begin embryonic develop- lings, while species of Trypocalliphora Peus even invade the
ment in the vagina, but as they usually are laid soon after tissues as subdermal parasites.
they enter the vagina, most embryonation takes place after
oviposition. Embryonation is rapid, however, requiring only
a few hours, and competition among larvae must be severe.
Classification
As a result of such intense competition, it is perhaps not
surprising that a few species have become predaceous on The Calliphoridae have had a long history of misidenti-
fication at the species level and oversplitting at the generic
other blow fly larvae. When sufficient food is not available,
level. Some common species that were repeatedly sent back
however, smaller adults are produced. The introduction of
from the New World to Europe by early collectors were de-
species of the genus Chrysomya to the New World in the
scribed over and over in various genera by the early authors,
1970s, beginning in South America (Prado  & Guimarães,
based on specimens that were far from perfect. The result
1982) and subsequently spreading north to Mexico and the
was numerous subsequent misidentifications and a long list
USA (Whitworth, 2006), has probably had a profound effect of unplaced species (James, 1970). To complicate matters,
on the faunal composition of Central America. Their high Townsend (1937), following a pattern established as early
adaptability, greater powers of dispersal as adults, and, in as 1830 by Robineau-Desvoidy, described numerous addi-
particular, the ability of their larvae to become predatory on tional genera based on slight differences, some also based
other blow fly larvae under conditions of scarce resources on misidentified type species. Some of these misconceptions
has been a factor in the reduction of many populations of along with additional oversplit genera were set in stone by an
native species (Baumgartner & Greenberg, 1984). influential textbook, The Blowflies of North America (which
The biology of species of the Mesembrinellinae is entirely included Central America) (Hall, 1948). Unfortunately, Hall
different. The reproductive system of the female is highly misidentified the type species of some genera, resulting in a
modified for the retention and nutrition of a single larva until realignment of generic concepts (Dear, 1985; Rognes, 1991).
it is ready to molt to the second instar (Guimarães, 1977). Dear’s (1985) treatment of the Chrysomyini forms the basis
Vivipary is not unusual among muscoid flies; it is also found for that part of the classification adopted here. For the
in all Sarcophagidae and also in the Old World calliphorid Calliphorinae, Hall’s treatment is still the only one available
genus Bellardia Robineau-Desvoidy, whose eggs are retained for Central America.
and hatch in utero and first instar larvae are deposited on an In his revision of the Mesembrinellinae, Guimarães (1977)
appropriate substrate. Among Calliphoridae, the Ameniinae recorded nine species occurring north of South America, out
of Australia and Eurychaeta Brauer & Bergenstamm of of a total of 29. He raised the rank of the subfamily to fam-
Europe also retain in utero a single well-grown larva, although ily status and proposed three subfamilies, the Mesembrinel-
there is otherwise no indication that these groups are closely linae, with metallic abdomens, and two new subfamilies,
related to Mesembrinellinae. However, where females of this the Souzalopesiellinae and Laneelinae, with non-metallic,
latter subfamily deposit their larvae is still somewhat of a brown abdomens. These new subfamilies each embraced a
CALLIPHORIDAE (BLOW FLIES) 109 1301

single species, Souzalopesiella facialis (Aldrich) and Laneella Calliphoridae. To this must be added the small, neglected
nigripes Guimarães. The latter was described only from Brazil, subfamily Melanomyinae. Reinhard (1929) and Downes
while the former, with 3+2 acrostical bristles and 3+3 dorso- (1986) have provided revisions of the North American spe-
central bristles, was recorded from Central America and cies, but there is apparently no information available for
Venezuela, but to date no specimen fitting the description has Central and South America (James, 1970 did not include
been found among the extensive holdings of Mesembrinel- mention of them), and none of the melanomyine species
linae in the INBio collection. Guimarães further subdivided found in Central America appear to be named.
his Mesembrinellinae into two tribes: Mesembrinellini, with For the Nearctic fauna, Shewell (1987) recognized only
three postpronotal bristles, and Eumesembrinellini, with two subfamilies, Chrysomyinae (based on the presence of
only two postpronotal bristles. The two genera encompassing a row of hairs along the trailing edge of the base of vein R
all but one of those recorded from Central America were (Fig.  4) with three tribes: Chrysomyini, Phormiini, and
Mesembrinella, with three postpronotal bristles, 2+2 acrostical Rhiniini) and Calliphorinae (which lack these hairs, with
bristles, but no discal bristles on tergite  5, and Huascaro- four tribes: Calliphorini, Luciliini, Polleniini, and Melano-
musca Townsend, with two postpronotals, 0+1 acrostichals, myini (for which he used the junior synonym Angioneurini)).
and a transverse row of discal bristles on tergite 5 (according Three of these tribes also occur in Central America; the
to statements given in either keys or diagnoses). Only two fourth tribe, Polleniini, has New World representatives in the
of the nearly 20 Costa Rican morphospecies in the INBio western USA and is also represented by several species of
collection have two postpronotal bristles; one appears to be Pollenia, whose larvae attack earthworms and which were
the sister species of another with three postpronotals, and a presumably introduced from Europe, along with the worms,
third male has two on one side and three on the other. Only by early settlers to the east coast of North America. Some
one species, M. abaca (Hall), has all three of the character of the species have spread more widely over the continent,
states of Mesembrinella, while only one other has all three
including Nearctic Mexico, but have apparently not yet
of the character states of Huascaromusca. All the remaining
reached Central America.
species clash in at least one character state; most have ei-
Four subfamilies are recognized here for Central America:
ther 2+1 acrosticals or 2+3 acrosticals. It is difficult to know
Mesembrinellinae, with only one genus, Mesembrinella (includ-
which of these characters is more reliable for splitting the
ing Huascaromusca and Eumesembrinella Townsend — see
group, or indeed if any can be used, or if splitting is even
advisable. No system of identification based on such variable discussion above); Chrysomyinae, with six genera as revised
or uncorrelated character states is likely to survive the cur- by Dear (1985); Calliphorinae, with two genera, Calliphora
rent push to supplant morphology-based identification with and Lucilia (with Phaenicia Robineau-Desvoidy in synony-
DNA sequencing, so until the Central American species can my); and Melanomyinae, with a single genus, Angioneura,
be studied more thoroughly the single genus Mesembrinella as treated by Downes (1986) as a subgenus of Melanomya
is used here for all of them. Rondani.
Another problem in the classification of the Calliphoridae
is the presence of many diverse groups across the globe that Identification
are probably not closely related to one another, but have been
grouped together on shared plesiomorphic character states or Guimarães (1977) provided keys to genera and species of
on the basis of what they are not, i.e., they are not muscids, all the Mesembrinellinae (as the family Mesembrinellidae),
rhinophorids, sarcophagids, or tachinids. This situation is including the nine Central American species known to him.
particularly vexing when reviewing the world fauna, and a
Problems recognizing the genera notwithstanding (discussed
cladistic analysis of the entire assemblage, i.e., Calliphori-
above), this revision provides information for species iden-
dae in the traditional sense, by Rognes (1997) suggested that
tification, although the Central American fauna is evidently
the family was not monophyletic. However, the New World
species, except for Mesembrinellinae, do appear to be mono- larger than previously realized. For the Chrysomyini, Dear
phyletic. These two New World lineages did not include the (1985) has given keys to all species in the New World. For
Old World subfamily Rhiniinae, present in the New World the Calliphorinae, Hall’s (1948) keys to Calliphora and
only by virtue of a single species introduced to Bermuda, but Lucilia (as the genus Phaenicia) of North America (which
were included in Shewell’s (1987) Chrysomyinae, a lineage included Central America) should be used in conjunction
defined by shared possession of a row of setae on the base with Whitworth’s (2006) keys to all the Nearctic blow flies,
of vein R1. including several that extend south to Central America, as
Dear (1985) provided a key to five subfamilies (Mesem- well as those for Northern Europe (Rognes, 1991) for mod-
brinellinae, Toxotarsinae, Chrysomyinae, Rhiniinae, and ernization of the names. The Melanomyinae of the Nearctic
Calliphorinae) that he believed comprised the New World Region were revised by Downes (1986).
1302
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Key to the genera of Calliphoridae of Central America

1. Stem vein (basal section of vein R) with row of hairs along posterior margin dorsally (Fig. 4);
posterior margin of hind coxa haired; CHRYSOMYINAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Stem vein bare (Fig. 5); posterior margin of hind coxa bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

2. Palpus reduced, slender, filiform, in profile no more than 1.5 times as wide as base (Fig. 2).. . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cochliomyia Townsend
– Palpus club shaped and of normal length, in profile more than twice as wide as base (Fig. 3). . . 3

3. Lower calypter entirely covered with dark appressed hair dorsally (as in Fig. 5), its hind margin
relatively straight and its outer corner angular; greater ampulla covered with fine hairs longer than
height of ampulla. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chrysomya Robineau-Desvoidy
– Lower calypter bare, or with sparse erect pale hair on basal half and along inner margin, with
rounded hind margin and broadly rounded outer corner (Fig. 4); greater ampulla covered with
microtomentum only, lacking pubescence. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4. Apex of wing transparent; both upper and lower calypters with long dark sparse hairs on upper
surfaces, either laterally or on basal half (as in Fig. 4); katepisternum with three bristles. . . . . . 5
– Wing apex, beyond r-m crossvein, tinted brown, best seen without magnification; neither upper
nor lower calypters with long hairs on surfaces; katepisternum with two bristles . . . . . . . . . . . . 6

5. Parafacial with fine dark hairs; presutural dorsocentral and intra-alar bristles absent or scarcely
differentiated from adjacent hairs; scutum boldly marked with three broad black longitudinal
stripes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Compsomyiops Townsend
– Parafacial bare; presutural dorsocentral and intra-alar bristles distinct; scutum with indistinct dark
longitudinal stripes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paralucilia Brauer & Bergenstamm

6. Posterior margin of hind coxa haired only on posterolateral corner (often hidden by abdominal
sternite which itself is haired); male eye with upper facets distinctly enlarged and sharply
delineated from smaller ventral facets. . . . . . . . . . . . . . . . . . . . . . . . . Chloroprocta van der Wulp
– Posterior margin of hind coxa haired along its entire length (as in tachinid Fig. 113.97); upper
facets of male eye neither distinctly enlarged nor delineated from lower facets. . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemilucilia Brauer

7. Anterior, ventral, and posterior margins of posterior thoracic spiracle supporting dense fringe
of uniformly long dense hairs (pale yellowish in nearly all species), overlaid with more widely
spaced, coarser, usually darker hairs, that form kidney-shaped, feltlike cover over spiracle, leaving
only small circular rounded opening near middle of dorsal margin (Fig. 7); MESEMBRINELLINAE.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mesembrinella Giglio-Tos
– Posterior thoracic spiracle closed by larger crescent-shaped or triangular, anterior feltlike cover,
or lappet, and smaller, more circular posterior lappet (Figs. 6, 8). . . . . . . . . . . . . . . . . . . . . . . . . 8

8. Body gray or brown, non-metallic, resembling a small tachinid; MELANOMYINAE. . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Angioneura Brauer & Bergenstamm
– Body shiny, metallic blue or green; CALLIPHORINAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

9. Lower calypter haired dorsally (Fig. 5); thorax dull gray pruinose; abdomen shining dark metallic
blue. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calliphora Robineau-Desvoidy
– Lower calypter bare; both thorax and abdomen shining metallic green, greenish-blue, or dark
purplish. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lucilia Robineau-Desvoidy
CALLIPHORIDAE (BLOW FLIES) 109 1303

Synopsis of the fauna tralasian region, including C. megacephala and C. rufifacies

(Kurahashi, 1989).
These flies are found throughout Central America in all
habitats. They are represented by about 25 described and at Cochliomyia Townsend. Three species are present in Central
least 15 undescribed species. America; C.  hominivorax, the primary screwworm, extir-
pated intentionally by sterile male release from North and
Angioneura Brauer & Bergenstamm. This is an inter- Central America, but still extant in southeastern Panama and
esting but rare and economically unimportant group. In the South America; C.  macellaria, the secondary screwworm,
Nearctic Region, some are parasitic on land snails of the common and widespread from southern Canada to southern
genus Succinia. At least five species have been collected in South America; and C. aldrichi Del Ponte, recorded from El
the highlands of Mexico and Costa Rica, but none can be Salvador as well as in the Florida Keys and the West Indies.
identified to species. Usage of the name Angioneura follows Only two other species, C. minima Shannon from the West
that of Shewell (1987) rather than that of Downes (1986), Indies and C. fontanai Garcia from Argentina, are present in
but a comprehensive comparative study of the world fauna is the genus (James, 1970).
needed to clarify generic concepts.
Compsomyiops Townsend. Also confined to the New
Calliphora Robineau-Desvoidy. This is a worldwide World, this small genus of six species is predominantly
genus, particularly species-rich in Australia, with several montane, occurring from southern Arizona to Chile. Three
Holarctic and Afrotropical species but only two native species have been recorded from Mexico and Central
Neotropical species and a third, Nearctic species, C. vicina America.
Robineau-Desvoidy, introduced into South America.
Females prefer to oviposit on carrion, especially of larger Lucilia Robineau-Desvoidy. This is a widespread genus
animals. Only one species, C. irazuana Townsend, is known that includes the common greenbottle flies that oviposit on
in Central America, from high elevations in Costa Rica and a wide variety of decomposing substances. As treated here,
Phaenicia Robineau-Desvoidy is regarded as a synonym of
Guatemala. Although this name was tentatively included in
Lucilia, in accordance with present European usage. The
the synonymy of the Nearctic C. terraenovae Macquart by
Central American species, of which there are perhaps a half
Hall (1948), the terminalia of Central American males is
dozen (Rognes, 1991), are badly in need of revision.
unlike any found in North America.
Hemilucilia Brauer. This is a small New World genus
Chloroprocta Wulp. This New World genus with only a
of six species occurring from Mexico to Argentina. Three
single species, C.  idioidea (synonym fuscanipennis (Mac- species have been recorded from Central America.
quart)), occurs from Texas to Paraguay. It is perhaps the least
common species in Central America, suggesting it might Mesembrinella Giglio-Tos. Nine species of Mesem-
have specialized habits, although larvae have been reared on brinella were recorded from Central America by Guimarães
meat. (1977) from a total of 29, all restricted to rain forests and
cloud forests of the Neotropical Region, where they feed on
Chrysomya Robineau-Desvoidy. Three species of this dung, fruit, and carrion, and skulk about in shaded situations.
genus, C.  albiceps (Wiedemann), C.  megacephala, and There appear to be at least 20 species collected to date in
C.  rufifacies, all recently introduced from the Old World Costa Rica. Males of one of these were observed congregating
tropics to South America, are now found in Central Amer- on a tree trunk at a hilltop.
ica (Baumgartner  & Greenberg, 1984). The last two have
recently spread to the northern USA (Whitworth, 2006, Paralucilia Brauer  & Bergenstamm. Also confined to
and personal observations) and could even be expected in the Neotropical Region from Mexico to Chile (including
southern Canada. There are 11 Afrotropical species (Pont, Easter Island), this genus contains only three species. This
1980), including C. megacephala, 10 species in the Oriental is not Paralucilia as used by Hall (1948), who erected
Region, including C.  megacephala and C. rufifacies, two the genus Myiolucilia for one of its species, nor as used by
shared with Africa (James, 1977), and 15 species in the Aus- Shewell (1987).

Literature cited

Baumgartner, D.L. & B. Greenberg. 1984. The genus Chrysomya Dear, J.P. 1985. A revision of the New World Chrysomyini (Diptera:
(Diptera: Calliphoridae) in the New World. Journal of Medical Calliphoridae). Revista Brasileira de Zoologia 3: 109–169.
Entomology 21: 105–113. Downes, W.L. 1986. The Nearctic Melanomyinae and relatives
Byrd, J.H.  & J.L. Castner (editors). 2001. Forensic Entomology. (Diptera: Calliphoridae), a problem in calyptrate classification.
The utility of arthropods in legal investigations. CRC Press,
Boca Raton, Florida, 440 pp. Bulletin of the New York State Museum, 460: iii–v + 35 pp.
1304
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Guimarães, J.H. 1977. A systematic revision of the Mesembrinel- asian and Oceanian Regions. Bishop Museum Press & E.J. Brill,
lidae, stat. nov. (Diptera, Cyclorrhapha). Arquivos de Zoologia Honolulu & Leiden, 1155 pp.
29: 1–109. Pont, A.C. 1980. 90. Calliphoridae, pp. 779–800. In Crosskey, R.W.
Hall, D.G. 1948. The blowflies of North America. Thomas Say (editor). Catalogue of the Diptera of the Afrotropical Region.
Foundation, Baltimore, Maryland, 477 pp. British Museum (Natural History), London, 1437 pp.
Hobson, R.P. 1932a. Studies on the nutrition of blow-fly larvae II. Prado, A.P. do & J.H. Guimarães. 1982. Estado atual de dispêrsão
Role of the intestinal flora in digestion. Journal of Experimental e distribuicão do gênero Chrysomya Robineau-Desvoidy na
Biology 9: 128–129. região neotropical (Diptera, Calliphoridae). Revista Brasileira
Hobson, R.P. 1932b. Studies on the nutrition of blow-fly larvae III. de Entomologia 26: 225–231.
The liquifaction of muscle. Journal of Experimental Biology 9: Reinhard, H.J. 1929. Notes on the muscoid flies of the genera
359–365. Opelousia and Opsodexia with the description of three new
James, M.T. 1970. 102. Calliphoridae. In Papavero, N. (editor). A species. Proceedings of the United States National Museum 76:
Catalogue of the Diptera of the Americas South of the United 1–9.
States. Departamento de Zoologia, Secretaria da Agricultura, Rognes, K. 1991. Blowflies (Diptera, Calliphoridae) of Fennoscan-
São Paulo, 28 pp. dia and Denmark. Fauna Entomologica Scandinavica 24:
James, M.T. 1977. Family Calliphoridae, pp.  526–556. In Delfi- 1–272.
nado, M.D. & D.E. Hardy (editors). A catalog of the Diptera of Rognes, K. 1997. The Calliphoridae (blowflies) (Diptera: Oestroidea)
the Oriental Region, Volume  III. University Press of Hawaii, are not a monophyletic group. Cladistics 13: 27–66.
Honolulu, x + 854 pp. Shewell, G.E. 1987. Calliphoridae, pp.  1133–1145. MND,
Jirón, L.F.  & I. Hedström 1985. Pollination ecology of mango Volume 2.
(Mangifera indica L.) (Anacardiaceae) in the Neotropic region. Townsend, C.H.T. 1937. Manual of Myiology, in twelve parts. Pt. 5.
Turrialba 35: 267–277. Muscoid generic diagnoses and data. Glossinini to Agriini.
Knipling, E.F. 1955. Possibilities of insect control or eradication Charles Townsend & Filhos, Itaquaquectuba, São Paulo, Brazil,
through the use of sexually sterile males. Journal of Economic 232 pp.
Entomology 48: 459–462. Whitworth, T.L. 2006. Keys to the genera and species of blow flies
Kurahashi, H. 1989. 109. Family Calliphoridae, pp.  702–718. In (Diptera: Calliphoridae) of America north of Mexico. Proceed-
Evenhuis, N.L. (editor). Catalog of the Diptera of the Austral- ings of the Entomological Society of Washington 108: 689–725.
OESTRIDAE (BOT FLIES) 110
Thomas Pape

Fig. 110.1. Male of Cuterebra emasculator Fitch, (Nearctic, MND, fig. 107.2).

Diagnosis First instar larvae usually with one or two bands of ro-
bust spines anteriorly on each segment. Posterior segment
Large, robust flies (body length 15–25 mm) with reduced with adhesive sucker (Cuterebra), with numerous crochet-
mouthparts and body clothed with pilelike setosity (Fig. 1). like spines (Cephenemyia, Oestrus), or carrying posterior
Body color blackish, yellowish, or metallic blue. Antenna spiracles on tubular extensions (Gasterophilus). Second and
third instar larva (Figs. 2–6) often with strong spines encircl-
usually short but first flagellomere elongate in Cuterebra
ing segments anteriorly, but dorsal surface bare in Oestrus,
hassleri (Guimarães). Arista bare, pectinate or plumose
and Cuterebra larvae have segments covered armorlike with
(Figs.  17–19). Vein  M either almost straight (Fig.  13) or more or less platelike or scalelike spines. Posterior spiracle
weakly to strongly bent (Figs. 11–12, 14–16), vein R1 with- with slits almost straight (Fig. 7), curved (Fig. 9), intertwined
out knob at level where subcosta joins costa. Legs short (Fig. 8), or forming porous plate (Fig. 10). During puparia-
and stout (e.g., Cuterebra) or slender (e.g., Gasterophilus). tion, larvae of Dermatobia and Cuterebra evert felt chamber
Abdomen globose or elongate oval. Male terminalia with of the anterior spiracles.
small and simple sternite  5, cerci and surstyli relatively Adult bot flies can hardly be confused with any other fam-
unmodified or shortened, phallus short and stout (almost ily, except perhaps a few large Tachinidae, which are always
stublike in Gasterophilus), with or without epiphallus. more setose and with a strongly swollen postscutellum. Some
Female terminalia usually short, but telescopic and retract- oestrids resemble bumblebees or carpenter bees; for in-
ed in Hypoderma, elongated and folded under the abdomen stance, Cuterebra hassleri looks so bumblebee-like that
in Gasterophilus. even a trained entomologist may be deceived.

1305
1306
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Biology leaves the host, backing out of the warble, passing through
the anus or exiting via the nostrils, and pupariates in the
Bot flies are parasites of mammals (records of Dermato- ground. Good accounts of bot fly biology may be found in
bia hominis Linnaeus, Jr. from birds apparently refer to the Zumpt (1965), Grunin (1965, 1966, 1969), Catts (1982), Sa-
muscid Philornis Meinert (Guimarães & Papavero, 1999)). brosky (1986), and Guimarães & Papavero (1999).
The main position of the larvae is subdermal (Cuterebrinae,
Hypodermatinae), intestinal (Gasterophilinae), or in the Classification
naso-pharyngeal area (Oestrinae). Cuterebra species glue
eggs to objects on the ground (C. baeri Shannon & Greene, Bot flies are members of the Oestroidea, and Rognes (1997)
a parasite of howler monkeys, may be an exception but its recently provided convincing evidence that they may be nest-
oviposition is still unknown), and Dermatobia hominis glues ed within a non-monophyletic Calliphoridae. A solution to the
eggs to mosquitoes or zoophilous calyptrate flies that act as related ranking problem has not been explicitly addressed.
carriers. When such carriers land on potential hosts, radiat- Bot flies are conveniently divided into four morphologi-
ing heat triggers eclosion of mature larvae, which wriggle cally and biologically distinct subgroups, here given rank as
down and bury into the skin (Catts, 1982; Sancho, 1988). subfamilies: Cuterebrinae, Gasterophilinae, Hypodermati-
Gasterophilinae and Hypodermatinae glue eggs to single nae, and Oestrinae (Pape, 2001).
hairs of the host. Species of Oestrinae are ovi-larviparous The Neotropical fauna of bot flies is still poorly known,
and eject newly hatched larvae onto the muzzle of the host. and the taxonomy is made difficult by considerable intraspe-
First instar larvae of some species migrate for short or cific variation in some species, which has resulted in a large
long distances in the host body before taking up their main number of species-group names (Guimarães  & Papavero,
position, often at specific sites. When the larva is mature, it 1999).

2 Cuterebra 3 Gasterophilus

4 Hypoderma 5 Cephenemyia

6 Oestrus

7 Dermatobia 8 Cuterebra 9 Gasterophilus 10 Hypoderma

Figs. 110.2–10. First instar larvae and posterior spiracles of third instar larvae: lateral view of first instar larva of (2) Cuterebra tenebrosa
Coquillett, (Nearctic, Baird & Graham, 1973, fig. 5A); ventral view of (3) Gasterophilus intestinalis (De Geer), (Holarctic, Grunin, 1969,
fig. 106); lateral view of (4) Hypoderma tarandi (Linnaeus), (Holarctic, Grunin, 1965, fig. 276); ventral view of (5) Cephenemyia trompe
(Modeer), (Holarctic, Grunin, 1966, fig. 106); and (6) Oestrus ovis Linnaeus, (Grunin, 1966, fig. 178); posterior spiracles of third instar
larva of (7) Dermatobia hominis (Linnaeus, Jr.), (modified from Kremer et al., 1978, fig. 1); (8) Cu. tenebrosa, (Nearctic, modified from
Baird & Graham, 1973, fig. 7D); (9) G. intestinalis, (Holarctic, MND, fig. 107.19); and (10) H. lineatum (Villers), (Holarctic, Grunin, 1965,
fig. 324).
OESTRIDAE (BOT FLIES) 110 1307

Identification Hypoderma, and Gasterophilus can be identified using

Grunin (1965, 1966, 1969).
The Neotropical (or at least South American) bot flies Routine pinning is recommended for adults. Puparia
are monographed by Guimarães  & Papavero (1999), should be mounted dry: for example, glued to cardboard tri-
which for Central America should be supplemented with angles with water-soluble glue, with explicit cross-reference
Bennett  & Sabrosky (1962) and Sabrosky (1986) for to the particular adult that emerged. Immature stages are best
Cephenemyia and Cuterebra, respectively. The synan- preserved in 80% alcohol or, for first instars and eggs, as
thropic and almost cosmopolitan species of Oestrus, permanent slides.

Key to the genera of Oestridae of Central America

Adults
1. Suprasquamal ridge setose. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Suprasquamal ridge bare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

11 Dermatobia 12 Cuterebra

M1

13 Gasterophilus 14 Hypoderma

r4+5

dm-cu

15 Cephenemyia 16 Oestrus

Figs. 110.11–16. Wings: dorsal view of (11) Dermatobia hominis (Linnaeus, Jr.); (12) Cuterebra sp., (Nearctic); (13) Gasterophilus intestinalis
(De Geer), (Holarctic); (14) Hypoderma bovis (Linnaeus), (Holarctic/cosmopolitan); (15) Cephenemyia trompe (Modeer), (Holarctic); and
(16) Oestrus ovis Linnaeus. Figures 11–16 photoscanned and adjusted by E. Binkiewicz.
1308
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

17 Dermatobia 18 Cuterebra 19 Gasterophilus

20 Hypoderma 21 Cephenemyia 22 Oestrus

Figs. 110.17–22. Heads: anterior (upper) and lateral (lower) view of (17) Dermatobia hominis (Linnaeus, Jr.), (Pape, 2001, fig. 1A) and
(18) Cuterebra semiatra (Wiedemann), (South America, Pape, 2001, fig. 1B); lateral (upper) and anterior (lower) view of (19) Gasterophilus
ternicinctus Gedoelst, (Afrotropical, Pape, 2001, fig.  2B); anterior  (upper) and lateral  (lower) view of (20)  Hypoderma actaeon Brauer,
(Palearctic, Pape, 2001, fig. 3A); (21) Cephenemyia rufibarbis (Meigen), (Pape, 2001, fig. 3D); and (22) Oestrus ovis Linnaeus. Figure 22
illustrated by E. Binkiewicz.
OESTRIDAE (BOT FLIES) 110 1309

2. Ground color of abdomen metallic blue.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dermatobia Brauer

– Ground color of abdomen dark brown, blackish or metallic black. . . . . . . . . . . . Cuterebra Clark

3. Wing vein M running practically straight to the margin (Fig. 13). . . . . . . . . . Gasterophilus Leach
– Wing vein M with smooth or abrupt bend at or distal to crossvein dm-cu (Figs. 11–12, 14–16). . . . . 4

4. Body with short clothing setae; frons and vertex with small pits (Fig. 22), often with central
tubercle carrying seta; wing cell r4+5 closed and with short petiole (Fig. 16) . . . Oestrus Linnaeus
– Body with long clothing setae; frons and vertex with neither pits nor tubercles; wing cell  r4+5
open . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

5. Antennal pedicel shining; facial plate swollen and broadening below antennae (Fig. 20); vein M
bent at blunt angle, bent part appearing as continuation of crossvein dm-cu (Fig. 14). . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hypoderma Latreille
– Antennal pedical with at least sparse microtomentum, not shining; facial plate not swollen,
narrowing ventrally (Fig. 21); vein M bent at right angle distal to crossvein dm-cu and with short
extension from bend (Fig. 15). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cephenemyia Latreille

First instar larvae

1. Terminal segment with several stout, hook-shaped spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Terminal segment with or without spines, but if present spines short and not hook-shaped. . . . . 3

2. Segments dorsally with at most four rows of spines; spines of last row on each abdominal segment
only hairlike laterally. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oestrus Linnaeus
– Most segments dorsally with seven or more rows of spines; all spines of the last row on each
abdominal segment hairlike. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cephenemyia Latreille

3. Nine abdominal segments, last three without spines; posterior spiracles carried on tubular
projections; labrum elongate, tapering; mandibles (mouth hooks) slender, with small terminal
hook and usually directed upwards.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gasterophilus Leach
– Eight abdominal segments; posterior spiracles level with terminal segment; labrum and mandibles
short and stout, directed downwards. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4. Keilin’s organ (paired groups of three setae ventrally on thoracic segments) absent; sclerotized
plate present anterodorsally on first thoracic segment above the head . . . . . . Hypoderma Latreille
– Keilin’s organ present; no sclerotized plate anterodorsally on first thoracic segment. . . . . . . . . . 5

5. Terminal segment unmodified, without adhesive sack.. . . . . . . . . . . . . . . . . . . Dermatobia Brauer

– Terminal segment with adhesive sack (Fig. 2).. . . . . . . . . . . . . . . . . . . . . . . . . . . Cuterebra Clark

Second and third instar larvae and puparia

1. Posterior spiracles composed as pair of porous plates with no trace of tripartition (Fig. 10). . . . 2
– Posterior spiracles either shaped as three distinct, cross-barred slits (Figs.  7, 9) or each slit
complexly intertwined but with tripartition still recognizable (Fig. 8). . . . . . . . . . . . . . . . . . . . . 4
1310
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

2. Antennomaxillary sense organs not situated on fleshy lobes; mandibles (mouth hooks) small,
fused to remaining cephalopharyngeal skeleton, and usually not visible without dissection. . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hypoderma Latreille
– Antennomaxillary sense organs situated on fleshy lobes; mandibles (mouth hooks) strong,
movably articulating with remaining cephalopharyngeal skeleton, and protruding from mouth
opening.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3. Dorsal surface without spines, ventral surface with strong spines anteriorly on every segment.. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oestrus Linnaeus
– Dorsal and ventral surface with about equal numbers of spines. . . . . . . . . Cephenemyia Latreille

4. Antennomaxillary lobe sclerotized on sides; posterior spiracles with single ecdysial scar and three
distinct, slightly arched slits in more or less vertical position (Fig. 9) . . . . . . Gasterophilus Leach
– Antennomaxillary lobe entirely unsclerotized; posterior spiracles each with their own ecdysial
scar and three slits or partitions slanting relative to median plane (Figs. 7, 8). . . . . . . . . . . . . . . 5

5. Strong spines in one to two rows on several segments; posterior spiracles shaped as three distinct
slits (Fig. 7) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dermatobia Brauer
– Numerous, more or less platelike or scalelike, spines on most segments; posterior spiracles with
each slit complexly intertwined, yet with tripartition still recognizable (Fig. 8). . Cuterebra Clark

Synopsis of the fauna One species, C.  baeri, specifically infests howler mon-
keys (Zeledón et al., 1957; Milton, 1996; Colwell  & Mil-
Around 30 species of this family are expected to occur in ton, 1998). The remaining species are rodent and lagomorph
Central America. parasites.

Dermatobia Brauer. Only one species, D. hominis, Gasterophilus Leach. There are eight Old World spe-
which is widespread in the Neotropical Region and in Cen- cies of which three have been introduced to the New World
tral America, occurs practically everywhere from sea level to and occur widespread in the Neotropical Region, including
above 1400 m (Sancho, 1988). This species seems to prefer Central America (Guimarães, 1967; Guimarães & Papavero,
humid, or at least evergreen tropical forests. High relative 1999).
humidity is required for eggs to survive, and moist soil for
the pupa to develop (Sancho, 1988; see also Janzen, 1976). Hypoderma Latreille. There are nine Palearctic species
Dermatobia hominis is a common parasite of domestic cat- of which H. bovis (Linnaeus) and H. lineatum (Villers) have
tle, but larvae may also be found in sheep, pigs, cats, dogs, become almost cosmopolitan, but apparently without estab-
humans, and more rarely horses and donkeys (Costa et al., lished populations in the Neotropical Region (Guimarães &
1990; Guimarães & Papavero, 1999). Among wildlife, the ta- Papavero, 1999). Both species occur in Central America,
pir is apparently not parasitized (Janzen, 1983) and, although however, as larvae in imported cattle.
the whitetailed deer is a suspected host (Janzen, 1976), there Cephenemyia Latreille. The eight species of this genus
is a remarkable absence of such records from the literature. are mainly restricted to the Holarctic Region, but one (C. jel-
Jaguars, pumas, agoutis, and armadillos have been recorded lisoni Townsend) has been captured in Costa Rica at 300 m
as hosts (Guimarães & Papavero, 1999). altitude in Guanacaste province.
Cuterebra Clark. Sixty New World species exist, of Oestrus Linnaeus. There are six Old World species, one
which 13 are known from Central America (Sabrosky, of which is the almost cosmopolitan O. ovis Linnaeus that
1986; Guimarães & Papavero, 1999; T. Pape, unpublished; infests domestic sheep. It occurs in Central America, but the
D.M.  Wood, unpublished), but several more are expected. distribution of this species is poorly documented.
OESTRIDAE (BOT FLIES) 110 1311

Literature cited

Baird, C.R. & C.L. Graham. 1973. Cuterebra tenebrosa: descrip- Guimarães, J.H.  & N. Papavero. 1999. Myiasis in Man and Ani-
tion of immature stages and a redescription of the adult (Diptera: mals in the Neotropical Region; bibliographic database. Edi-
Cuterebridae). The Canadian Entomologist 105: 1281–1293. tora Plêiade, São Paulo, 308 pp.
Bennett, G.F. & C.W. Sabrosky. 1962. The Nearctic species of the Janzen, D.H. 1976. The occurrence of the human warble fly (Der-
genus Cephenemyia (Diptera, Oestridae). Canadian Journal of matobia hominis) in the dry deciduous forest lowlands of Costa
Zoology 40: 431–448. Rica. Biotropica 8: 210.
Catts, E.P. 1982. Biology of New World bot flies: Cuterebridae. An- Janzen, D.H. 1983. Tapirus bairdii (Danto, Danta, Baird’s Tapir),
nual Review of Entomology 27: 313–338. pp. 496–497. In Janzen, D.H. (editor). Costa Rican Natural His-
Colwell, D.D.  & K. Milton. 1998. Development of Alouattamyia tory. The University of Chicago Press, Chicago & London, xi +
baeri (Diptera: Oestridae) from howler monkeys (Primates: Ce- 816 pp.
bidae) on Barro Colorado Island, Panama. Journal of Medical Kremer, M., C. Rebholtz,  & J.-P. Rieb. 1978. Iconographie des
Entomology 35: 674–680. plaques stigmatiques de Dermatobia hominis Linné Jr (= D. cy-
Costa, J.O., W.S. Lima, M.P. Guimarães,  & N.M.  Lima. 1990. aniventris Macquart 1843). Annales de Parasitologie 53:
Frequência de endo e ecto parasitos de cães capturados nas ruas 439–440.
de Vitoria, E.S. Brasil. Arquivo Brasileiro de Medicina Veter- Milton, K. 1996. Effects of bot fly (Alouattamyia baeri) parasitism
inária e Zootecnia 42: 451–452. on a free-ranging howler monkey (Alouatta palliata) population
Grunin, K.J. 1965. Hypodermatidae. In Lindner, E. (editor). Die in Panama. Journal of Zoology 239: 39–63.
Fliegen der palaearktischen Region 8: 1–154, 41  pls., adden- Pape, T. 2001. Phylogeny of Oestridae (Insecta, Diptera). System-
dum 155–160. E.  Schweizerbart’sche Verlagsbuchhandlung, atic Entomology 26: 133–171.
Stuttgart [pp.  1–40 published 31.xii.1964; pp.  155–160 pub- Rognes, K. 1997. The Calliphoridae (blowflies) (Diptera: Oestroi-
lished 30.ix.1969]. dea) are not a monophyletic group. Cladistics 13: 27–66.
Grunin, K.J. 1966. Oestridae. In Lindner, E. (editor). Die Fliegen Sabrosky, C.W. 1986. North American species of Cuterebra, the
der Paläarktischen Region 8: 1–96. E. Schweizerbart’sche Ver- rabbit and rodent bot flies (Diptera: Cuterebridae). Thomas Say
lagsbuchhandlung, Stuttgart. Foundation Monographs 11: 1–240.
Grunin, K.J. 1969. Gasterophilidae. In Lindner, E. (editor). Die Flie- Sancho, E. 1988. Dermatobia, the Neotropical warble fly. Parasi-
gen der Paläarktischen Region 8: 1–61. E. Schweizerbart’sche tology Today 4: 242–246.
Verlagsbuchhandlung, Stuttgart. Zeledón, R., Q.O. Jiménez,  & M.R.R. Brenes. 1957. Cuterebra
Guimarães, J.H. 1967. Family Gasterophilidae. In Papavero, N. baeri Shannon  & Greene en el mono aullador de Costa Rica.
(editor). A Catalogue of the Diptera of the Americas South of Revista de Biología Tropical 5: 129–134.
the United States. Departamento de Zoologia, Secretaria da Ag- Zumpt, F. 1965. Myiasis in Man and animals in the Old World. But-
ricultura, São Paulo, 4 pp. terworth & Co. Ltd., London, xv + 267 pp.
1312
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
SARCOPHAGIDAE (FLESH FLIES) 111
Thomas Pape and Gregory A. Dahlem

Fig. 111.1. Male of Peckia gulo (Fabricius), (illustrated by T. Sjöberg).

Diagnosis but entire distiphallus mostly strongly modified (Fig. 54);

phallotreme often bi- or tripartite.
Generally medium-sized to large (body length 10–25 mm), Larvae typical maggots. Posterior spiracles of larvae I–III
robust flies (Fig. 1), but a few species smaller (5–10 mm), placed in recession or cavity (Fig.  55); this cavity flat and
and some species produce grossly undersized individuals almost indiscernible in some species. Spiracular slits set at
given sparse larval resources. Body color generally distinc- acute angle relative to median plane, often almost vertical,
tive, thorax with three strong black stripes (Figs. 1, 13) and and in many species with “kink” near base; peritreme in sec-
abdomen speckled or checkerboard pattern (although some ond and third instar larva incomplete and without ecdysial
species not pigmented in this way). Male dichoptic or with scar.
semi-narrow frons, not truly holoptic; female dichoptic. Sub- Species of the subfamily Miltogramminae possess neither
scutellum concave. Male often with fringe of hairlike setae the three dorsal stripes on the thorax nor the checkerboard
ventrally on hind tibia (Fig. 36). Abdominal sternites with- pattern on the abdomen. They superficially resemble small
out sensilla trichodea (alpha setae). Female ovoviviparous Tachinidae, quite often even displaying similar behavior.
(common oviduct with a bilobed incubatory pouch) and with Some species, especially of Lepidodexia (sensu stricto) and
short, non-telescopic terminalia; common oviduct with Lepidodexia (Johnsonia), are darker and more bristly than
bilobed incubatory pouch. Bacilliform sclerites (processi usual in the Sarcophaginae and may be taken for Tachinidae,
longi; divided male sternite 10) shortened and widely diverg- but all these species lack the greatly swollen subscutellum
ing; cercus and surstylus highly species-specific (Figs. 48– characteristic of the latter family. However, the slender-
53); phallus with undivided dorsomedian sclerotization, bodied and somewhat tachinid-like Lepidodexia (Johnsonia)

1313
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

2 Amobia 3 Eumacronychia 4 Macronychia

6 Metopia
5 Metopia 7 Oebalia

8 Opsidia 9 Senotainia 10 Sphenometopa

Figs. 111.2–10. Heads of Miltogrammini: lateral view of (2) Amobia floridensis (Townsend), (MND, fig. 108.85); (3) Eumacronychia
polita (Townsend), (Nearctic, MND, fig. 108.90, as Gymnoprosopa); (4) Macronychia aurata (Coquillett), (Nearctic, MND, fig. 108.86);
and (5) Metopia sinipalpis Allen, (MND, fig. 108.95, as Allenanicia); dorsal view of (6) Me. argyrocephala (Meigen), (MND, fig. 108.96);
lateral view of (7) Oebalia aristalis (Coquillett), (Nearctic, MND, fig. 108.88); (8) Opsidia diademata Pape, (South America, Pape, 2002,
fig. 1); (9) Senotainia trilineata (Wulp), (MND, fig. 108.87, as Sphixapata); anterolateral view of (10) Sphenometopa tergata (Coquillett),
(Nearctic, MND, fig. 108.92, as Euaraba).
SARCOPHAGIDAE (FLESH FLIES) 111 1315

11 Sarcophaga 12 Lepidodexia (Johnsonia) 13 Sarcophaga

pal wall

anepm

14 Sarcophaga 15 Amobia

16 Titanogrypa

19 Lepidodexia (Duckemyia)
17 Helicobia 18 Lepidodexia (Notochaeta)

20 Peckia 21 Emdenimyia
22 Peckia

Figs. 111.11–22. Heads and thoracic structures: lateral view of head of (11)  Sarcophaga aldrichi Parker, (Nearctic, MND, fig.  108.5, as
Arachnidomyia); (12) Lepidodexia (Johnsonia) elegans (Coquillett), (MND, fig. 108.21); dorsal view of thorax, vestiture omitted on left
side (13) and posterolateral view, with lower calypter cut away, of thorax (14) of S. aldrichi, (Nearctic, MND, figs. 108.23–24); lateral view
of thorax of (15) Amobia floridensis (Townsend), (MND, fig. 108.98); scutellum, enlarged inset showing white setae, of (16) Titanogrypa
placida (Aldrich); post-sutural dorsocentral bristles of (17) Helicobia sp.; (18) L. (Notochaeta) sp.; (19) L. (Duckemyia) latifrons (Kano &
Lopes); (20) Peckia chrysostoma (Wiedemann); and (21) Emdenimyia meyersi (Curran); dorsal view of notopleuron of (22) P. intermutans
(Walker). Figures 16–22 illustrated by E. Binkiewicz.
Abbreviations: anepm, anepimeron; pal wall, postalar wall.
1316
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

woodorum (Pape) has a moderately swollen subscutellum, seem to be specialized predators of turtle eggs (Vogt, 1981;
a tongue-shaped lower calypter, and a small metathoracic T. Pape, unpublished). Blaesoxipha contains numerous para-
spiracle like the Rhinophoridae. This, in combination with sitoids, especially of acridid grasshoppers and tenebrionid
a notopleuron showing no trace of subprimary bristles and beetles, but also cockroaches, mantids, and other Saltatoria
a hind trochanter without posterior setae, makes this species and Coleoptera serve as hosts (Pape, 1994). Emblemasoma
difficult to key out with other Sarcophaginae. The plumose contains parasitoids of Cicadidae (Homoptera), with females
arista, absence of coxopleural streak, and exposed abdominal locating their male hosts by the song (Soper et al., 1976), and
sternites in the male reveal the subfamilial position, and the the genus Spirobolomyia has been bred exclusively from spi-
setose wing vein CuA1 is shared with other species of Lepi- robolid millipedes (Pape, 1990). Dexosarcophaga contains
dodexia (Johnsonia). Bright metallic green or blue species, species with larvae living in termite mounds and ant nests
like Lepidodexia (Chlorosarcophaga), L. (Chloronesia), and (Lopes, 1939a, 1968). Certain Sarcophaga are predators of
even a few L. (Notochaeta), may be taken for Calliphoridae, spider egg sacs (Lopes, 1989), with at least one species spe-
but they have a more elongated abdomen and differ by hav- cializing on colonial orb webs of Metepeira incrassata F.O.
ing subprimary notopleural bristles (Fig. 22), normal-sized Pickard-Cambridge and showing an intricate prey mimicry
posterior spiracle, setae posteriorly on the hind trochanter, ploy to lure the spiders off their egg sacs (Hieber et al., 2002).
and different male and female terminalia. Some Blaesoxipha (Gigantotheca) and many Sarcophaga
are scavengers of small carrion, such as dead insects, snails,
and smaller vertebrates, and a few species breed in larger
Biology vertebrate carcasses (Pape, 1994, 1996). A large group of
Neotropical Sarcophaginae, which is here combined under
Flesh flies are necrophagous, coprophagous, and parasi- Lepidodexia (sensu lato), is biologically very superficially
toids/predators on insects, snails, reptiles, and amphibians. known. Rearing records (e.g., myiasis in frogs, predation of
Females are ovoviviparous, and larvae often develop to ma- earthworms, and breeding in snails (Lopes, 1973a; Crump &
turity within 4–5 days. Pounds, 1985)) are remarkably sparse, and most species are
Species of Miltogramminae are generally kleptoparasites presumably highly specialized.
of solitary bees and wasps, feeding on the food stored for Several species of Comasarcophaga, Malacophagomyia,
the host progeny. An important exception is Eumacronychia, Udamopyga, Helicobia, and Lepidodexia (Johnsonia) have
whose species are scavengers or predators of lizard and turtle been bred from snails and may be specialized parasitoids
eggs (e.g., Lopes, 1982a; Mullen et al., 1984; Trauth & Mul- (Lopes, 1966, 1969a, 1973a; Neck & Lopes, 1973; Reeves
len, 1990; Gautreau, 2007). Female miltogrammines may et al., 2000).
trail wasps returning with paralyzed prey to their nest in The large majority of adult Sarcophagidae are sun-
flight or ride the abdomen of large prey items being dragged loving, but preferences may be narrow, such as small sun-
over the ground, or they may deposit larvae directly into the spots on the ground deep in humid forest, insolated twigs
host nest entrance in the ground or in hollow twigs (Spof- at a certain height, or sun-baked stones on river beaches.
ford & Kurczewski, 1990). Such spots may be used as meeting sites, with males taking
The Neotropical fauna of Sarcophaginae is remarkably stations and chasing away other males and “jumping” at
biologically diverse. Most species have narrow diets, but females in order to mate. Other notable meeting sites are
Sarcodexia lambens (Wiedemann) is diverse in its food carrion, where males make up a conspicuous part of the
choice and has been bred from nestling birds, snails, dead visitors (or they will take stations low in the nearby vegeta-
and living (but probably often injured or otherwise weak- tion), and elevated parts of the landscape, like hilltops and
ened) insects and scorpions, and vertebrate carrion, and it hillcrests (Downes, 1994).
is known from cases of human myiasis (Townsend, 1893;
Hoffmann, 1927; Almeida, 1933; Callan, 1946; Parker et al.,
1953; Harrison, 1963; Stegmaier, 1972; Fessl et al., 2001). Classification
Oxysarcodexia and Ravinia are associated with dung, mostly
as coprophages, but some species may be partly predatory A division into three subfamilies (Miltogramminae, Para-
(Pickens, 1981). One species of Sarcofahrtiopsis has special- macronychiinae, and Sarcophaginae) appears phylogeneti-
ized on dung from ephemeral leaf roosts of the disk-winged cally sound (Pape, 1996), but hypotheses beyond that are
bat (Pape et al., 2002), and another species of this genus is a speculative and provisional (e.g., Lopes, 1983; Pape, 1994,
scavenger in dead crabs, together with Villegasia postunci- 1998). Definitions and circumscriptions of genus-group
nata (Hall) (Pape & Méndez, 2002; Méndez & Pape, 2003). taxa are improving (e.g., Pape, 1990, 1994; Mello-Patiu &
Peckia gulo (Fabricius) appears to be an obligate scavenger of Pape, 2000), yet are still preliminary for the large majority
land crabs, Cardisoma spp. (Méndez & Pape, 2003). Several of genera and, in some cases, are absent. Little consensus
species produce myiasis in turtles, lizards, and amphibians has emerged on formal ranking, and changes are expected
(e.g., Blake, 1955; Dodge, 1955), and species of Tripanurga following better supported phylogenies.
SARCOPHAGIDAE (FLESH FLIES) 111 1317

Identification as far as possible, though this has not been fully attainable.
During identification, it is recommended to use a fine pin
The Neotropical fauna of Sarcophagidae is strongly dom- or other tool (a matchstick with a minuten pin inserted at
inated by the subfamily Sarcophaginae, which has diversi- the end is perfect) to gently push back the alula of the wing
fied considerably in external morphology as compared with from the postalar wall when looking for the postalar setae
their Old World relatives. Still, few males and even fewer (Fig. 14). These are often obscured by the wing, especially
females can be identified on external characters alone, and a the alula that folds up along the postalar wall when the wing
study of the male and female genitalia remains the only reli- is in resting position. If done gently, this does not damage
able means of identification for the vast majority of species. the specimen.
Spreading the male genitalia during field work or shortly af- Routine pinning is the best way to preserve adult Sar-
ter return (see below) facilitates identification tremendous- cophagidae, but as examination of male terminalia is the only
ly. Studies of female genitalia are lagging, yet these often means of a reliable species-level identification for species of
provide useful diagnostic features and important progress Sarcophaginae  — and in general is a much more effective
is being made (e.g., Tibana  & Mello, 1983; Tibana, 1985; way of sorting at the generic level for the more experienced
Mello-Patiu & Santos, 2001). Gravid females may be identi- student — it is strongly recommended to pull out the termi-
fied through rearing of their male progeny, which is not too nalia at least for members of this subfamily. This is best done
complicated for those species where larvae squeezed from on specimens shortly after collecting and after disappearance
the mother can be bred on liver, feces, or other readily avail- of rigor mortis. Specimens collected around noon and stored
able media (Bänziger & Pape, 2004). in a humid (not wet!) container, e.g., a film canister with a
Genus-level and species-level keys, such as Hall’s rolled-up leaf, will usually be ready for preparation later the
(1933a) and Dodge’s (1968) papers on the Sarcophagidae same day, but they will be even easier on the following day.
from Panama, exist for only a fraction of the fauna. Such Terminalia can be pulled out (e.g., with a crooked insect pin)
keys, however, are generally difficult to use without exten- and ideally should be kept in position until dry. This may be
sive knowledge of the fauna as well as of the intricacies of accomplished by setting the specimen at an angle and insert-
an ever-changing nomenclature and the bewildering gener- ing a minuten or fine insect pin behind the cerci to prevent
ic concepts. Species-level identifications for those genera terminalia from sliding back. When processing large catches
where no recent revision is available can only be done by during field work, such preparation may be prohibitively
consulting the most recent catalog (Pape, 1996) and tracking time consuming or might exhaust storage facilities. Speci-
relevant illustrations in the literature — in itself an intimidat- mens should still have their terminalia pulled out, as they
ing task — or by consulting an extensive and authoritatively will usually not retract fully, allowing a rough sorting and
identified reference collection. significantly facilitating processing if specimens are relaxed
The present key to Central American genera of Sar- at a much later time. Pulling down the legs immediately after
cophagidae is our “best attempt so far”. It is certainly not pinning, so that they are not tightly folded under the speci-
easy to use and we already know of undescribed species that men, will provide a much better view of diagnostic bristles
will lead the user astray. We expect the number of excep- and setae. Immature stages should be preserved in 80% al-
tions to grow steadily with increasing knowledge of the fau- cohol or as permanent slides. For slide preparations of the
na, and we strongly recommend that non-experienced users cephalopharyngeal skeleton, separating and unfolding the
confirm their results against reliably identified material or two halves (e.g., with a sharpened insect pin) will allow a
by consulting an expert. We have tried to key out females much better view.

Key to the genera of Sarcophagidae of Central America

Note that the Nearctic Cistudinomyia Townsend is not included, as the record of a Cistudinomyia  sp. from Venezuela
(Marmels, 1994) is a misidentification of undersized specimens of Blaesoxipha plinthopyga (Wiedemann) (specimens at the
University of Maracay examined by T. Pape). Extra-regional genera are noted in brackets.

1. Antennal arista bare (Figs. 2–10) or at most with trichia about as long as greatest aristal diameter,
not plumose; notopleuron without subprimary bristles; metasternal area bare; hind coxa bare
posteriorly; coxopleural streak present; male abdominal sternites 2–4 partly hidden by overlapping
margins of corresponding tergites (Fig. 46). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Antennal arista usually distinctly plumose (Figs. 11–12); notopleuron usually with subprimary
bristles (Fig. 22); metasternal area usually setose; hind coxa usually setose posteriorly; coxopleural
streak absent; male abdominal sternites 2–4 exposed and overlapping margins of corresponding
tergites (Fig. 47); SARCOPHAGINAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
1318
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

2. Head profile with flat or even slightly concave occipital region (Figs.  2–10); eyes distinctly
(Figs. 2, 5, 7–10) or moderately (Figs. 3–4) enlarged; postocular setae not alternating, of about
equal size; male midfemur without apical posteroventral ctenidium; male terminalia not or only
slightly protruding and if tergite 6 fused to syntergosternite 7+8 then these structures not forming
right angle; MILTOGRAMMINAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
– Head profile with convex occipital region (Figs. 11–12); eyes not enlarged (Fig. 11) or at most
slightly so (Fig. 12); postocular setae alternating in size; male midfemur with apical posteroventral
ctenidium of stout, flattened bristles; male terminalia usually protruding and with tergite 6
distinctly fused to syntergosternite 7+8 at more or less right angle; PARAMACRONYCHIINAE.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

3. Wing cell r4+5 short petiolate or closed at wing margin (Fig. 23). . . . . . . . . . . . Taxigramma Perris
– Wing cell r4+5 open. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4. More than three proclinate fronto-orbital bristles present (Fig. 2); anterior margin of anepimeron
with distinct swelling below ampulla (Fig. 15). . . . . . . . . . . . . . . . . . Amobia Robineau-Desvoidy
– At most two proclinate fronto-orbital bristles present; anterior margin of anepimeron without
distinct swelling below ampulla. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

5. Setae of pedicel stout; first flagellomere at most slightly longer than 2 times length of pedicel;
eye moderately enlarged (Fig. 4); upper part of proepisternum swollen (or at least not particularly
depressed). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macronychia Rondani
– Setae of pedicel not modified; first flagellomere usually more than 2.5 times length of pedicel; eye
distinctly (Figs. 5–10) or moderately (Fig. 3) enlarged; upper part of proepisternum unmodified,
not swollen. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

6. Parafacial plate with row of bristly setae along anterior margin close to facial ridge (Fig. 5) . . . 7
– Parafacial plate bare or setose, but setae not arranged in row along anterior margin close to facial
ridge (the facial ridge itself may be equipped with a row of strong setae) . . . . . . . . . . . . . . . . . . 8

7. Two reclinate and two proclinate fronto-orbital bristles arranged in partly overlapping rows with
anterior reclinate orbital bristle positioned anterior to posterior proclinate bristle (Fig. 6); lunule
often setose. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Metopia Meigen
– Only one reclinate fronto-orbital bristle positioned posterior to row of proclinate bristles; lunule
bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Opsidia Coquillett

8. Facial ridge with row of bristly setae on more than basal half (Fig. 10); basicosta black, concolorous
with tegula. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sphenometopa Townsend
– Facial ridge bare or with some supravibrissal setae on less than basal 0.3; basicosta orangish,
contrasting with black or dark brown tegula. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

9. Setae of lower postcranium (i.e., below the cervix) white. . . . . . . Phrosinella Robineau-Desvoidy
– Setae of lower postcranium black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

10. Arista thickened on more than proximal half (Fig.  7); male cercus with sickle-shaped prong;
female cercus with setae distinctly erect or slanting anteriorly; female hypoproct with slightly
swollen rim. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oebalia Robineau-Desvoidy
– Arista thickened on less than proximal half; male cercus with almost straight or evenly curved
prong; female cercus with decumbent setae; female hypoproct not modified. . . . . . . . . . . . . . . 11
SARCOPHAGIDAE (FLESH FLIES) 111 1319

11. Scape slightly raised above lunule; vibrissal angle not, or barely, developed, facial ridge receding
throughout (Fig. 3); abdominal pattern of distinct, silvery, transverse bands. . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eumacronychia Townsend
– Scape flush with lunule; vibrissal angle small but distinct, head with trapezoidal or almost square
appearance (Fig.  9); abdominal pattern speckled, spotted, or rather uniform; without distinct,
transverse bands. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Senotainia Macquart

12. Palpus distinctly clavate and practically bare dorsally; vibrissa short, about as long as distance
between vibrissal sockets; small species (5–7 mm); southern USA, Mexico. . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Cattasoma Reinhard]
– Palpus more cylindrical and only slightly swollen apically, setose dorsally; vibrissa distinctly
longer than distance between vibrissal sockets; larger species (greater than 7 mm). . . . . . . . . . 13

23 Taxigramma 24 Sarcophaga

C sector 2 C sector 3

26 Argoravinia
25 Lepidodexia (Johnsonia)
R2+3 C sector 2 C sector 3
R4+5

27 Boettcheria

28 Argoravinia
R2+3

R4+5

29 Boettcheria

30 Boettcheria 31 Peckia
Figs. 111.23–31. Wings and calypters: dorsal view of right wing of (23) Taxigramma heteroneura (Meigen), (Holarctic, MND, fig. 108.103);
(24) Sarcophaga aldrichi Parker, (Nearctic, MND, fig. 108.32, as Arachnidomyia); and (25) Lepidodexia (Johnsonia) elegans (Coquillett),
(MND, fig. 108.30); ventral view of left wing showing part of costal apices of veins Sc and R1 of (26) Argoravinia rufiventris (Wiedemann)
and (27) Boettcheria cimbicis (Townsend); ventral view of node at base of radial sector of left wing of (28) A. rufiventris and (29) B. cimbicis
(Townsend); dorsal view of left upper and lower calypters of (30) B. cimbicis and (31) Peckia intermutans (Walker). Figures 26–31 illustrated
by E. Binkiewicz.
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

13. Parafacial plate with setae not reaching below centre of eye; abdominal microtomentum silvery
gray with distinct black spots that may coalesce posteriorly (black spots rarely covering larger
part of abdomen); Nearctic Region . . . . . . . . . . . . . . . . . . . [Wohlfahrtia Brauer & Bergenstamm]
– Parafacial plate setose to, or almost to, lower margin of eye; abdominal pattern with irregular
transverse bands or ill-defined pattern that changes with incidence of light, at most ill-defined
spots. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

14. Dark, blackish species; palpus black; abdominal microtomentum silvery, arranged in irregular
transverse bands leaving broad atomentose posterior margin; terminalia black. . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachicoma Rondani
– Grayish species; palpus orange; abdominal microtomentum greyish, not arranged in transverse
bands, extending to or almost to posterior margin; terminalia orange; southern USA, northern
Mexico. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Erythrandra Brauer & Bergenstamm]

15. Midtibia without anterodorsal bristles except for apicals (Fig. 45). . . . . . . . . . . . . . . . . . . . . . . 16

– Midtibia with one or more anterodorsal bristles (Fig. 44). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

16. Vein R1 bare dorsally. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tricharaea Thomson, in part

– Vein R1 setose dorsally. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

17. Subprimary notopleurals present (Fig. 22); metasternal area with several setae; wing usually with
brownish infuscated, subapical costal spot; male midfemur with apical posteroventral ctenidium.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nephochaetopteryx Townsend
– Subprimary notopleurals absent, at most represented by minute setulae; metasternal area bare or
with one to two setulae; wing hyaline; male midfemur without apical posteroventral ctenidium .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sarcofahrtiopsis Hall, in part

18. Prosternum greatly widened anteriorly, anterior width twice or more posterior width and anterolateral
corners tending to be pointed, not separated from propleuron by any great length of membrane
(Fig. 33). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Emblemasoma Aldrich
– Prosternum parallel sided or spatulate (Fig. 32), not greatly widened anteriorly . . . . . . . . . . . . 19

19. Male with one or more proclinate orbital bristles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

– Male without proclinate orbital bristles, or female . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

20. Vein R1 setose dorsally. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

– Vein R1 bare dorsally. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

21. Proepisternum setose (sometimes only few setulae in anterior part); vein R1 setose ventrally near
humeral crossvein; one or few setae on ventral side of base of Rs elongated (Fig. 28); margin of
lower calypter with long trichia extending to posterolateral corner (Figs. 30–31); genitalia with
sternite  7+8 and epandrium enlarged, bulging from abdominal segment  5; medium-sized flies,
body length about 10–12 mm. . . . . . . . . . . . . . . . . . . . Tripanurga Brauer & Bergenstamm, in part
– Proepisternum bare; vein R1 bare ventrally; setae on ventral side of of base of Rs not elongated
(Fig. 29); margin of lower calypter with long trichia no further than middle; genitalia not bulging;
generally small flies, body length about 4–6 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

22. Sternite 5 without incision, posterior margin often sinuous; surstylus apically without short, spiny
setae and usually with one to two long setae; phallic vesica drawn out into often long projection.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sarcofahrtiopsis Hall, in part
– Sternite 5 with deep, V-shaped incision; surstylus apically at least with few short, stout, or spiny
setae; phallic vesica at most with short, curved projection. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
SARCOPHAGIDAE (FLESH FLIES) 111 1321

32 Sarcophaga 33 Emblemasoma 34 Blaesoxipha 35 Boettcheria

37 Sarcodexia
36 Boettcheria

38 Oxysarcodexia 39 Dexosarcophaga

43 44

40 Blaesoxipha 41 Helicobia 42 Helicobia 43,44 Dexosarcophaga 45 Nephochaetopteryx

Figs. 111.32–45. Prosterna and leg structures: ventral view of prosternum and left forecoxa of (32) Sarcophaga aldrichi Parker, (Nearctic,
MND, fig. 108.25, as Arachnidomyia) and (33) Emblemasoma erro Aldrich, (MND, fig. 108.28); posterior view of hind coxa, trochanter,
and base of femur of (34) Blaesoxipha plinthopyga (Wiedemann), (MND, fig. 108.26, as Kellymyia) and (35) Boettcheria latisterna Parker,
(Nearctic, MND, fig. 108.27); posterior view of hind femur and tibia of (36) Bo. latisterna, (MND, fig. 108.38); anterior view of hind femur
of (37) Sarcodexia lambens (Wiedemann); posterior view of midfemur, inset showing enlargement of distal spine of comb and its shape in
cross-section, of (38) Oxysarcodexia timida (Aldrich) and (39) Dexosarcophaga transita Townsend; posterior view of midtibia of
(40) Bl. costaricensis (Lopes) and (41) Helicobia pilipleura Lopes; posterior view of hind tibia of (42) H. pilipleura; posterior view (43)
and anterior view  (44) of midtibia of D. transita; anterior view of midtibia of (45)  Nephochaetopteryx  sp. Figures  37–45 illustrated by
E. Binkiewicz.
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

23. Phallus long, distiphallus longer than cercus and with two sclerotized winglike plates in basal
part. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Panava Lopes
– Phallus not particularly elongated, distiphallus shorter than cercus and without platelike or
winglike sclerotizations in basal part. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Wulpisca Lopes

24. Postalar wall bare on middle; hind coxa bare posteriorly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

– Postalar wall setose on middle (Fig. 14); hind coxa setulose posteriorly . . . . . . . . . . . . . . . . . . 26

25. Midfemur with apical posteroventral ctenidium.. . . . . . . . . . . . . . . . . . . . . . . . . . Pacatuba Lopes

– Midfemur without apical posteroventral ctenidium. . . . . . . . . . . . . . Tricharaea Thomson, in part

26. With whitish setae anterior to genal sulcus; phallus short, compact. . . . . . . . . . . . . . . . . . . . . . 27
– With only black setae anterior to genal sulcus; phallus not particularly short or compact. . . . . 28

27. Wing with third costal sector setose ventrally; terminalia black or dark brown; postgonites
(parameres) directed anteriorly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tapacura Lopes
– Wing with costal sector 3 setose ventrally (as in Fig. 27); terminalia reddish; postgonites
(parameres) directed laterally. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Retrocitomyia Lopes, in part

28. First flagellomere at least 4 times as long as pedicel; facial ridge straight and densely setose on
lower 0.8; male cercus bifurcate apically. . . . . . . . . . . . Lepidodexia (Duckemyia) Kano & Lopes
– First flagellomere about 3 times as long as pedicel; facial ridge weakly concave and setose on
about lower 0.2; male cercus gradually tapering and without apical bifurcation . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rettenmeyerina Dodge

29. Two postsutural dorsocentrals present, spaced as for three but the anterior bristle absent or
reduced in size and as close or closer to middle bristle than to transverse suture (Fig. 18) . . . . 30
– Two or more postsutural dorsocentrals present, if two, then not spaced as above (e.g., Figs. 19–
21) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36

30. Vein R1 setose dorsally. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepidodexia (Abacantha) Hall

– Vein R1 bare dorsally. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
31. Terminalia orange. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
– Terminalia black or dark blackish-red. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

32 With whitish setae extending anterior to genal sulcus; fronto-orbital and parafacial plates with
almost whitish microtomentum, giving head light-colored appearance . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Retrocitomyia Lopes, in part
– With only black setae anterior to genal sulcus; fronto-orbital and parafacial plates with usual
silvery or golden microtomentum, head not appearing particularly light-colored. . . . . . . . . . . . 33

33. Facial ridge densely setose on lower 0.75. . . . . . . . . . . . Lepidodexia (Dexomyophora) Townsend
– Facial ridge setose on lower 0.5 and not particularly dense . . . . . . . . . . . . . . Thomazomyia Lopes

34. Facial ridge densely setose in lower 0.8; proepisternum bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepidodexia (Duckemyia) Kano & Lopes
– Facial ridge setose in about lower 0.2 (at most); proepisternum setose. . . . . . . . . . . . . . . . . . . . 35

35. Proepisternum with few setae (fewer than five); lateral styli of phallus inserted close to one
another. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepidodexia (Eufletcherimyia) Townsend
– Proepisternum with several, scattered setae (more than 10); lateral styli of phallus more widely
separated. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepidodexia (Notochaeta) Aldrich, in part
SARCOPHAGIDAE (FLESH FLIES) 111 1323

36. Two postsutural dorsocentrals, spaced as for three but middle bristle missing (Fig. 21) . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Emdenimyia Lopes, in part
– Two or more postsutural dorsocentrals, if two not spaced as above. . . . . . . . . . . . . . . . . . . . . . 37

37. With both three large, more or less evenly spaced marginal scutellars and frontoclypeal sutures
approximated at vibrissae, with distance between sutures at level of vibrissae at most 1.5 times
greatest width of second antennal segment in frontal view, usually much less; arista at most short
plumose; apical posterodorsal bristle of hind tibia larger than apical dorsal bristle . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepidodexia (Neophyto) Townsend
– Usually two marginal scutellars or, if more, additional bristles smaller than and situated much
closer to large posterior marginal than to anterior marginal; distance between frontoclypeal sutures
at vibrissae usually twice or more width of second antennal segment; arista usually medium to
long plumose; apical posterodorsal bristle of hind tibia seldom differentiated. . . . . . . . . . . . . . 38

38. Lunule large, as long as broad; parafacial row of strong setae present; proanepisternum setose;
three postsutural dorsocentral bristles spaced for three (as in Fig. 17); wing vein R1 setose . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Malacophagula Bequaert
– Without this combination of characters. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39

39. With both whitish setae anterior to genal sulcus, and posterior bristle of midtibia (located at about
two-thirds length of tibia from base) notably larger than posterodorsal bristle (Fig. 41); abdomen
not metallic blue or green. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
– Either whitish setae not anterior to genal sulcus or posterior bristle of midtibia much smaller than
or at most subequal to posterodorsal bristle (Fig. 40), or both; if neither, abdomen metallic. . . 53

40. Presutural dorsocentrals reduced, usually one small bristle near transverse suture, none larger than
subprimary notopleurals. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
– Presutural dorsocentrals differentiated, at least one larger than subprimary notopleurals. . . . . . 43

41. Costal sector 3 with ventral setulae (Fig. 27). . . . . . . . . . . . . . . . . . . . . . Malacophagomyia Lopes

– Costal sector 3 without ventral setulae (Fig. 26). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42

42. Lateral margins of scutellum with dense patch of whitish hairlike setae between lateral scutellar
bristles (Fig. 16). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Titanogrypa Townsend, in part
– Lateral margins of scutellum without patch of whitish hairlike setae. . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Peckia Robineau-Desvoidy, in part

43. Vein R1 bare dorsally. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44

– Vein R1 setose dorsally. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47

44. Male without (or with weak and hairlike) apical scutellar bristles; midfemur with apical
posteroventral ctenidium; female with tergite  6 divided in two halves that close off remaining
terminalia tightly, leaving only narrow vertical seam. . . . . . . . . . . . . . . . . . . . . . Villegasia Dodge
– Male with apical scutellar bristles; midfemur without apical posteroventral ctenidium; female
with abdominal tergite 6 unmodified or partly desclerotized dorsally, arching over remaining
terminalia and not closing them off. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45

45. Male hind femur with black patch of setae near apex of anterior surface (Fig. 37); female abdominal
sternite 2 with pair of erect or semi-erect discal as well as marginal bristles . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sarcodexia Townsend, in part
– Male hind femur without black patch of setae; female abdominal sternite 2 with pair of marginal
bristles but no discals. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

46. With two or at most three postsutural dorsocentral bristles as strong as or stronger than
subprimary notopleurals. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Retrocitomyia Lopes, in part
– With four postsutural dorsocentral bristles as strong as or stronger than subprimary notopleurals.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hallina Lopes, in part

47. With strong costal spinelike seta at junction with Sc. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48

– Without strong costal spinelike seta at junction with Sc. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49

48 Presutural acrostichals well developed; abdominal segments 4–5 reddish or orangish. . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rafaelia Townsend, in partt
– Presutural acrostichals not developed; abdominal segments unicolorous, not orangish. . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Titanogrypa Townsend, in part

49. Male with apical scutellar setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sarcodexia Townsend, in part

– Male without apical scutellar setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50

50. One or few setae on ventral side of Rs elongated (Fig. 28). . . . . . . Argoravinia Townsend, in part
– Setae on ventral side of Rs not elongated (Fig. 29). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51

51. Abdomen ventrally exclusively with white setae in male, in female with white setae except for
pair of black median marginal bristles on sternites. . . . . . . . . . . . . Argoravinia Townsend, in part
Note: Species keying here are sometimes separated as Raviniopsis Townsend.

– Abdomen ventrally with mainly black setae in both male and female . . . . . . . . . . . . . . . . . . . . 52

52. Third antennal segment and palpus light brown to orange . . . . . . . . . . Rafaelia Townsend, in part
– Third antennal segment and palpus dark brown to black. . . . . . . . . Titanogrypa Townsend, in part

53. Postalar wall setose on middle (Fig. 14) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54

– Postalar wall without setae on middle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79

54. Apical posteroventral bristle of hind tibia well differentiated, subequal in size to apical anteroven-
tral bristle (Fig. 43) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
– Apical posteroventral bristle of hind tibia not differentiated or only slightly so (Fig. 42) . . . . . 59

55. Vein R1 setose dorsally. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Helicobia Coquillett, in part

– Vein R1 without setae dorsally. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56

56. Male midfemur without apical posteroventral ctenidium, stout bristles sometimes present but
usually pointed and not so closely set; cerci with posterior profile in lateral view straight or nearly
so on basal two-thirds, curved evenly forward distally, and spinelike setae, if present, directed
dorsally, anterodorsally, or anteriorly (cerci vertical); tergite 6 of female almost invariably incised
or divided on midline, entire in species with longitudinal clear streak without microtrichia in
cell cu1, or in some species with ventral setulae on costal sector 3 (between tips of Sc and R1);
sternites 7–8 not fused, if so, sternite 8 divided, latter commonly in form of narrow transverse
strip, sometimes membranous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sarcophaga Meigen
– Male midfemur with apical posteroventral ctenidium; distal half to two-thirds of cerci bent
posteriorly from basal part (but recurved anteriorly towards tip), almost always with small to minute
spinelike setae directed posterodorsally on posterior apical half (as the terminalia are spread, with
the general longitudinal axis of the phallus and cerci vertical); tergite 6 of female entire, rarely
bent rooflike on midline; sternites 7–8 fused and sclerotized . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
SARCOPHAGIDAE (FLESH FLIES) 111 1325

57. Palpus black (yellow only in some Blaesoxipha plinthopyga; note spinelike setae on posterior surface
of hind trochanter); phallus with juxta flattened or absent, not beaklike; median stylus tubelike
and usually broad; lateral styli variously modified into flattened platelike or flaplike structures
or reduced, not tubelike; vesica not particularly well developed or absent (but often difficult
to homologize); both sexes usually with row of spinelike setae on posterior surface of hind
trochanter (Fig. 34). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Blaesoxipha Loew, in part
– Palpus yellow; phallus different, median and lateral styli shaped as curved, slender tubes; vesica
distinctly developed; hind trochanter without spinelike setae in both sexes. . . . . . . . . . . . . . . . 58

58. Phallus with juxta bilobed (posterior view), blunt, and distinctly separated from distiphallic tube;
female abdominal tergite  6 with posterior margin unmodified and without median tonguelike
projection.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Comasarcophaga Hall, in part
– Phallus with juxta beaklike, tapering and fused with distiphallic tube; female abdominal tergite 6
with the median (= dorsal) part of posterior margin devoid of setae and produced into tonguelike
projection. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Spirobolomyia Townsend

59. Tegula orangish or light brown, sometimes partly infuscated. . . . . . . . . . . . . . . . . . . . . . . . . . . 60

– Tegula blackish, similar in color to adjacent scutum. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62

60. All genal setae white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Retrocitomyia Lopes, in part

– At least some black setae anteriorly on gena. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61

61. Male with apical scutellars present; female with frontal vitta at midpoint at most slightly wider
than parafrontal; female abdominal spiracle  6 generally in tergite  6; sternites  7–8 fused and
usually formed into a trough- or scoop-shaped larvipositor . . . . . . . . . Blaesoxipha Loew, in part
– Male without apical scutellars; female with frontal vitta at midpoint generally twice or more parafrontal
width at that point; female abdominal spiracle 6 in membrane anterior to tergite; sternites 7 and 8
not usually fused, not modified into larvipositor. . . . . . . . . . . . . . . . . Ravinia Robineau-Desvoidy

62. Vein R1 setose dorsally. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63

– Vein R1 without setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64

63. Male midfemur without apical posteroventral ctenidium; posterior bristle of midtibia (located at
about two-thirds length of tibia from base) larger than posterodorsal bristle (Fig. 41); terminalia
reddish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Helicobia Coquillett, in part
– Male midfemur with apical posteroventral ctenidium (Fig. 39); posterior bristle of midtibia smaller
than posterodorsal bristle or absent (Fig. 43); terminalia blackish . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dexosarcophaga Townsend, in part

64. Presutural dorsocentrals reduced, usually only one small bristle near transverse suture, none larger
than subprimary notopleurals; hind trochanter without spinelike setae on posterior surface . . . 65
– Presutural dorsocentrals well differentiated, if reduced, posterior surface of hind trochanter with
row of spinelike setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67

65. Costal sector 3 with ventral setulae (as in Fig. 27) . . . . . . . . . . . . . . . . . . . . . . Peckiamyia Dodge
– Costal sector 3 without ventral setulae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66

66. Long hairlike setae of axillary cord of lower calypter abundant and extending at least to outer hind
corner of calypter (Fig. 31); whitish setae of posterior head region extending anterior to genal sulcus;
abdominal sternites 3 and 4 without erect black setae. . . . . . . . Peckia Robineau-Desvoidy, in part
– Long marginal hairlike setae of lower calypter not extending to outer hind corner of calypter;
whitish setae not anterior to genal sulcus; abdominal sternites 3 and 4 with dense erect black setae
at least near apical margin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Engelimyia Lopes
1326
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

67. Proepisternum setose; facial ridge with long dense setae almost on full length. . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Emdenimyia Lopes, in part
– Proepisternum bare, rarely setose (some Blaesoxipha spp.); facial ridge setose on at most basal
0.75 and with upper setulae short and scattered. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68

68. Costal sector 3 with ventral setulae (as in Fig. 27) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69

– Costal sector 3 without ventral setulae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72

69. Prementum of proboscis bare or almost bare of pruinosity. . . . . . . Microcerella Macquart, in part
– Prementum of proboscis almost completely pruinose. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70

70. Metallic species; presutural acrostichals not differentiated . . . . Lepidodexia (Chloronesia) Townsend
– Non-metallic species; presutural acrostichals moderately to strongly differentiated . . . . . . . . . 71

71. Ocellar bristles well developed, stronger than weakest frontals; male cerci simple, straight, not
particularly long, and fused at midline to or almost to apex. . . . . . . . . . . . . . . . . Udamopyga Hall
– Ocellar bristles weakly developed and at most as strong as weakest frontals; male cerci strong and
with distinct flaplike swelling distal to middle, prongs separate and diverging for considerable
length. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Halliosca Lopes

72. Posterior surface of head almost entirely with black setae, at most few whitish setae around neck;
male midfemur with apical posteroventral ctenidium. . . . . . . Dexosarcophaga Townsend, in part
– Posterior surface of head with some whitish setae distinctly visible in lateral view; male midfemur
with or without apical posteroventral ctenidium. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73

73. Whitish setae of posterior surface of head extending anteriorly on ventral part of head up to genal
sulcus; costal spinelike seta present; arista pubescent or short plumose, longest trichia at most
about half as long as width of first flagellomere . . . . . . . . . . . . . . . Comasarcophaga Hall, in part
– Without above combination of characters. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74

74. Males . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
– Females . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78

75. Distal 0.5–0.7 of cercus bent posteriorly from basal part and, on distal half, with small spinelike
setae in most cases; phallus distinctly divided into basal and distal segment; presutural acrostichals
usually strong. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Blaesoxipha Loew, in part
– Cercus usually with distal part not bent relative to base, without small spinelike setae; phallus not
or only weakly segmented; presutural acrostichals weak or absent. . . . . . . . . . . . . . . . . . . . . . . 76

76. Scutellum with apical setae; midfemur without apical posteroventral ctenidium. . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chrysagria Townsend
– Scutellum without apical setae; midfemur with apical posteroventral ctenidium. . . . . . . . . . . . 77

77. Ctenidium with normal bristles (circular cross-section: Fig. 39). . . . . . . . . . . . . . Oxyvinia Dodge
– Ctenidium with flattened bristles (oval or rectangular cross-section: Fig. 38) . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oxysarcodexia Townsend

78. Hind tibia almost invariably with single bristle in anteroventral row; tergite 6 usually angularly
arched rooflike, or divided, or concealed under tergite 5 at midline; spiracle 6 in membrane
ahead of tergite, or contiguous with tergite. . . . . . . . . . . . . . . . . . . . . . . . . Chrysagria Townsend,
Oxysarcodexia Townsend,
Oxyvinia Dodge
– Hind tibia usually with two bristles in anteroventral row; tergite 6 entire, only curved slightly or
straight at midline; spiracle 6 usually in tergite 6 . . . . . . . . . . . . . . . . . Blaesoxipha Loew, in part
SARCOPHAGIDAE (FLESH FLIES) 111 1327

79. Costal sector 3 with ventral setulae (as in Fig. 27); subprimary notopleurals differentiated. . 80
– Third costal section without ventral setulae or subprimary notopleurals not differentiated. . . . 83

80. Long trichia of axillary cord of lower calypter extending to posterolateral corner of calypter
(Fig. 30); theca of proboscis almost bare of pruinosity; presutural acrostichals usually differentiated;
hind trochanter anteroventrally with brushlike clump of short, stubby setae (Fig. 35). . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Boettcheria Parker
– Long trichiae of lower calypter restricted to anterior half of outer edge; theca of proboscis
pruinose; presutural acrostichals not differentiated; hind trochanter without clump of short, stubby
setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81

81. Vein R1 setose dorsally (Fig. 25). . . . . . . . . . . . . . . . . . . . . . . Lepidodexia (Johnsonia) Coquillett

– Vein R1 bare dorsally (as in Fig. 24) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82

82. Ocellar bristles absent or slightly differentiated . . . . . . . . . . Lepidodexia (Harpagopyga) Aldrich

– Ocellar bristles differentiated, but may be small. . . . . . . . . Lepidodexia (Chloronesia) Townsend,
L. (Chlorosarcophaga) Townsend,
L. (Notochaeta) Aldrich, in part

83. Subprimary notopleurals not differentiated. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84

– Subprimary notopleurals distinctly differentiated (Fig. 22). . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85

84. Vein R1 without dorsal setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tricharaea Thomson (females),

Pacatuba Lopes (females)
– Vein R1 with dorsal setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sarcofahrtiopsis Hall, in part,
Wulpisca Lopes (females),
Panava Lopes (females)

85. Arista bare to short plumose on basal half, length of trichia seldom approaching width of second
antennal segment, but if so, long trichia of margin of lower calypter restricted to anterior half of
outer edge. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microcerella Townsend, in part
– Aristal trichia as long or longer than greatest width of second antennal segment in frontal view,
if approaching width of antennal segment, long trichia of margin of lower calypter extending to
outer hind corner of calypter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86

86. Long trichia of lateral edge of lower calypter extending to posterolateral corner of calypter; one
or few setae on ventral side of base of Rs elongated (as in Fig. 28); posterior and posterodorsal
bristles of foretibia (located at about two-thirds length of tibia from base) usually present; presutural
acrostichals almost invariably differentiated and strong; male terminalia large and bulging. . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tripanurga Brauer & Bergenstamm, in part
– Long trichiae of lateral edge of calypter restricted to anterior half; setae on ventral side of base
of Rs not elongated (as in Fig. 29); posterodorsal bristle of foretibia present or absent; presutural
acrostichals absent or differentiated; male terminalia normal, not enlarged. . . . . . . . . . . . . . . . 87

87. Four postsutural dorsocentrals. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hallina Lopes, in part

– Three postsutural dorsocentrals. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88

88. Ocellar bristles absent or only slightly differentiated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89

– Ocellar bristles well differentiated. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
1328
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

48 Sarcophaga 49 Dexosarcophaga

46 Metopia 47 Oxysarcodexia

52 Blaesoxipha 53 Blaesoxipha

50 Udamopyga 51 Blaesoxipha

55 Peckia

54 Sarcophaga

56 Sarcofahrtiopsis

Figs. 111.46–56. Abdomens, male terminalia, larval spiracles, and puparium: ventral view of abdomen of (46) Metopia campestris (Fallén),
(Nearctic) and (47) Oxysarcodexia timida (Aldrich); lateral view of cercus and surstylus of (48) Sarcophaga johnsoni Aldrich, (Nearctic,
MND, fig. 108.47, as Wohlfartiopsis); (49) Dexosarcophaga transita Townsend, (MND, fig. 108.41); (50) Udamopyga niagarana (Parker),
(Nearctic, MND, fig. 108.55); (51) Blaesoxipha hunteri (Hough), (MND, fig. 108.52, as Protodexia); (52) B. prohibita (Aldrich), (Nearctic,
MND, fig. 108.53, as Acanthodotheca); and (53) B. atlanis (Aldrich), (MND, fig. 108.51); lateral (left) and anterior (right) views of distiphal-
lus of (54) Sarcophaga triplasia Wulp; posterior view of larva (right) and enlargement of larval spiracles (left) of (55) Peckia gulo (Fabricius),
(Méndez & Pape, 2003, fig. 2); lateral view of puparium (right) and dorsal view of operculum (left) of (56) Sarcofahrtiopsis thyropteronthos
Pape, Dechmann, & Vonhof, (Pape et al., 2002, figs. 4–5). Figures 46–47 illustrated by E. Binkiewicz; Figure 54 illustrated by M. Meldert.
SARCOPHAGIDAE (FLESH FLIES) 111 1329

89. Two postsutural intra-alar setae present. . . . . . . . . . . . . Lepidodexia (Gymnocamptops) Townsend

– One postsutural intra-alar seta present. . . . . . . . . . . . . . . . . . . . . . . Lepidodexia (Petriana) Lopes

90. Setulae of scutum adjacent to presutural dorsocentrals sparse and much smaller than dorsocentrals,
about 0.2 or less length of latter; scutellum with two large marginal scutellars (sometimes a small
apical pair present); prementum of proboscis pollinose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepidodexia (Lepidodexia) Brauer & Bergenstamm
– Setulae of scutum not so different in size from dorsocentrals; scutellum with three or four
marginal scutellars; prementum of proboscis largely bare of pollen. . . . . . . Archimimus Reinhard

Synopsis of the fauna of katydids and praying mantids. Subgenus Acanthodotheca

Townsend contains parasitoids of mainly scarabaeid beetles
Some 300 species of Sarcophagidae have been recorded but also Tenebrionidae and Cerambycidae. Blaesoxipha
from Central America, but we have seen numerous addi- (Acanthodotheca) dampfi (Hall), however, has been bred
tional species, many of which are probably undescribed, and from a dead rodent in Costa Rica (Reeves et al., 2000). Sub-
the fauna is likely to at least double when thorough studies genus Gigantotheca Townsend contains saprophages and
are undertaken. We are here treating the Central American predators of wounded or otherwise weakened insects. Blaes-
Sarcophagidae in 51 genera, largely following Pape (1996) oxipha (G.) plinthopyga (Wiedemann) appears to be mainly
and with the few changes annotated in the generic synopsis synanthropic in the tropics, and it has been bred from a hu-
below. We prefer to recognize broad (user-friendly) generic man corpse (Jirón et al., 1983).
concepts using a strict phylogenetic argumentation. Future
changes are expected, especially when the phylogeny of the Boettcheria Parker. There are 27 New World species of
large and morphologically diverse Lepidodexia (sensu lato) this genus, of which nine have been recorded from Central
becomes better known. America. The biology of the Neotropical species is poor-
ly known, but several Nearctic species have been reared
Amobia Robineau-Desvoidy. This is a widely distributed from live (possibly wounded or otherwise weakened)
genus with a total of 13 species, two of which have been insects, including species of Coleoptera, Hymenoptera,
recorded from Central America. Species of Amobia are in Lepidoptera, and Orthoptera (Dahlem & Downes, 1996).
general kleptoparasites of mud-daubers and twig-nesting There is no complete key, but Dahlem & Downes (1996)
aculeate hymenopterans. Adult flies occur on vegetation as combined with Pape & Dahlem (1998) will cover several
well as on trunks and twigs of dead trees (Spofford et al., species, and the latter authors provide references to fig-
1989). ures of male and female terminalia of all Central Ameri-
can species. While Nearctic and Central American species
Archimimus Reinhard. There are five New World spe- of Boettcheria may appear reasonably homogeneous,
cies of this genus, of which only one, A.  turbata (Wulp), the South American fauna contains several species that
has been recorded in Central America. A key to species is may appear as reminiscent of Microcerella as they are of
provided by Lopes  & Tibana (1988). Nothing is currently Boettcheria.
known of the biology of these flies.
Brachicoma Rondani. This genus is found mainly in the
Argoravinia Townsend. Six New World species of this Holarctic Region and is known from eight species, with at
genus are known, two of which have been recorded from least one species, B.  setosa Coquillett, extending into the
Central America. Two species have been associated with northernmost parts of5 Central America. Larvae are preda-
nests of Hymenoptera, one with Sphecidae and one with tors of the progeny, especially last instar larvae, of social
Bembicidae (Lopes, 1988a). Argoravinia rufiventris Hymenoptera, and in particular bumble bees (Plath, 1934;
(Wiedemann) seems to have broad food preferences, ranging Verves, 1984).
from urban waste to damaged eggs and dead hatchlings in
turtle nests (Lopes, 1973a; T. Pape, unpublished). Chrysagria Townsend. Three New World species have
been described within this genus, and two of these have been
Blaesoxipha Loew. As redefined by Pape (1994), this is a recorded from Central America. One species has been bred
large genus with 245 species worldwide. Forty-eight species from larvae of a chrysomelid beetle (Lopes & Achoy, 1986),
have been recorded from Central America, but several more but the biology is otherwise unknown.
are to be expected. For identification see Pape (1994). Spe-
cies of subgenera Blaesoxipha (sensu stricto), Acridiophaga Comasarcophaga Hall. There are four southern Nearctic
Townsend, Servaisia Robineau-Desvoidy, and Tephromyia species of this genus, of which one extends into Neotropi-
Brauer & Bergenstamm are parasitoids of acridid grasshop- cal Mexico. One species, Comasarcophaga texana Hall, has
pers, with some species of the latter being parasitoids also been bred from a snail (Neck & Lopes, 1973).
1330
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Dexosarcophaga Townsend. This is a large, predomi- Halliosca Lopes. Only one Central American species
nantly Neotropical genus with currently 45 New World of this genus, H.  declinata (Hall), is known. Its biology is
species, of which 16 have been recorded from Central America. unknown. Like Hallina, this genus was considered by Pape
We expect to discover many undescribed species from the (1996) to be a subgenus under the broad Lepidodexia sensu
Neotropical Region. Some species seem to be associated lato.
with either ants or termites, but biological observations are
fragmentary. Helicobia Coquillett. This is a large genus with 33 New
World species. Eight species have been recorded from Cen-
Emblemasoma Aldrich. This New World genus contains tral America. The generic monophyly seems reasonably well
16 species, four of which have been recorded from Central corroborated when species earlier assigned to Punaphyto
America. Several of these species are parasitoids of male Townsend are included. However, it remains an open ques-
cicadas. The flies have a prosternal acoustic organ, which tion if Helicobia is a subset (e.g., subgenus) of the large
is most developed in the female (Lopes, 1981; Soper et al., Sarcophaga sensu lato. For identification, Lopes (1939b)
1976) and which allows them to locate cicadas by their song. is useful but needs to be supplemented with additional spe-
A key to the species of Emblemasoma is provided by Lopes cies. The widespread H. rapax (Walker) and H. morionella
(1988b). We do not recognize Colcondamyia Reinhard and (Aldrich) have been bred from dead and injured insects and
Pessoamyia Lopes as being distinct from Emblemasoma (cf. snails (e.g., Plank, 1929; Stegmaier, 1972; Lopes, 1973a;
Lopes, 1971; Shewell, 1987). Reeves et al., 2000).

Emdenimyia Lopes. There are 11 Neotropical species of Lepidodexia Brauer  & Bergenstamm. There are 185
this genus, three of which have been recorded from Central validly named New World species of this genus, of which 34
America. The monophyly seems well corroborated, but Pape have been recorded from Central America. Many additional
(1996) mentioned the possibility that Emdenimyia could be species are currently undescribed and the taxonomy is ex-
nested within Boettcheria. One species, Emdenimyia koryt- ceedingly complex. This large taxon is morphologically di-
kowskii Lopes, has been reared from an arctiid larva (Lepi- verse and many groups are poorly circumscribed. As we learn
doptera) (Lopes, 1969b), but no other biological information more about this diverse genus, dramatic changes in the clas-
exists. Lopes (1969b) provides a key to the species of this sification are expected. Duckemyia Kano  & Lopes (known
genus. from Costa Rica, T. Pape & G.A. Dahlem, unpublished) is
here considered as a subtaxon under Lepidodexia. Several
Engelimyia Lopes. There are four Neotropical species of bright metallic taxa occur, such as all Lepidodexia (Chloro-
this small genus, two of which occur in Central America. sarcophaga) Townsend and L. (Chloronesia) Townsend, but
The South American Engelimyia inops (Walker) has been also some species of Lepidodexia (Notochaeta) Aldrich. The
collected on human feces and fermented bananas (Lopes, latter subgenus is rich in species, most of which are blackish
1973a, 1975), and the Costa Rican E. bosqi Pape & Mello is and morphologically similar. Species of Lepidodexia (John-
attracted to dead fish (Pape & Mello-Patiu, 2006). sonia) Coquillett are tachinid-like and have long frontal
bristles. The biology is remarkably poorly known consider-
Eumacronychia Townsend. With a total of 22 species, ing the large number of species, and a high fraction of these
this is the largest of the few endemic New World miltogram- may have specialized life habits. Lepidodexia (Notochaeta)
mine genera. Nine species have been recorded from Central bufonivora Lopes & Vogelsang, for example, is an obligate
America, mainly from the northernmost part. Most Eumac- parasitoid of various amphibians (Crump & Pounds, 1985),
ronychia are found in dry areas, where they are apparently and it may be a pest in small-scale farming of harlequin frogs
predators on turtle and lizard eggs. Eumacronychia sternalis (T. Pape, unpublished). The presence of numerous small lar-
Lopes has been bred from nests and hatchlings of the green vae in the females of several species and their inability to de-
turtle (Chelonia mydas) in Mexico (Lopes, 1982a) and from velop in feces and carrion (T. Pape, unpublished) is a further
eggs of the leatherback turtle (Dermochelys coriacea) in indication of a specialized diet.
Costa Rica (T.  Pape, unpublished). This species is also at-
tracted to carrion (Cornaby, 1974; T. Pape, unpublished). Macronychia Rondani. This is a widespread genus with
a total of 16 species, but only one has so far been record-
Hallina Lopes. Three species are found in Central Amer- ed from Central America (T.  Pape, unpublished). Males
ica and the northern parts of South America. Pape (1996) of Macronychia are commonly found hilltopping and on
considered this genus as a subgenus under Lepidodexia sen- firetowers at the level of the surrounding canopy (Dodge &
su lato, but we prefer to treat it here as a valid genus awaiting Seago, 1954). Several Nearctic species are associated with
better phylogenetic resolution of the sarcophagine genera. the nests of aculeate Hymenoptera (Spofford et al., 1989),
Also, we are here treating Hallina as a senior synonym of and an unidentified species has been reported to be a para-
Hallimyia Lopes. Nothing is currently known of the biology sitoid of adult tabanids (Thompson, 1978a, b; Thompson &
of members of this genus. Love, 1979).
SARCOPHAGIDAE (FLESH FLIES) 111 1331

Malacophagomyia Lopes. This is a small genus with two recommended to confirm identification by comparing with
named Neotropical species, neither of which have been re- published illustrations. Dodge (1966) also provides a key to
corded from Central America, but material of a single un- species and includes genitalia drawings of a variety of spe-
identified species has been found in Costa Rica. Members cies. Lopes (1946) includes many additional genitalia fig-
of this genus have been recorded as parasitoids of snails ures. Many species are known to be dung-breeders.
(Lopes, 1966).
Oxyvinia Dodge. There are 10 Neotropical species of
Malacophagula Bequaert. There are five Neotropical this genus, one of which, O. panamensis Lopes, has been re-
species of this genus, none of which have been recorded corded from Central America. Little is known of the biology
from Central America, but like the preceding genus, material of members of this genus, but adults are attracted to feces
of a single unidentified species has been found in Costa Rica. and at least one species has been bred from carrion (T. Pape,
Species of Malacophagula have been recorded as parasitoids unpublished).
of snails (Bequaert, 1925; Lopes, 1969a).
Pacatuba Lopes. This is a monotypic Neotropical genus
Metopia Meigen. There are 34 species worldwide of this with its single species P. matthewsi Lopes known from Cen-
genus, of which six have been recorded from Central Amer- tral America. Males have been taken from carrion near the
ica, but several more are represented in various collections. beach (T. Pape, unpublished).
Many of these species are associated with and have been
reared from nests of a variety of bees and wasps (Wcislo et Panava Dodge. Four Neotropical species are known of
al., 1985; Wcislo 1984, 1986; Spofford et al., 1989). this genus, one of which, P. inflata Dodge, is known from
Central America. Males of P. inflata are attracted to carrion
Microcerella Macquart. In the broad definition of Pape and have been collected over swarming ants (Dodge, 1968;
(1990), this genus contains 69 New World species, of which Lopes, 1978).
only a single undescribed species is known from Central
America (G.A. Dahlem & T. Pape, unpublished). The genus Peckia Robineau-Desvoidy. There are 70 New World
has its center of diversity in the South American highlands species of this genus of mainly large and robust species, and
and it is remarkable not only by a striking diversity in male 23 have been recorded from Central America. Cocos Island
genitalia but also for having species with green eyes. The ge- harbors the partly synanthropic P. chrysostoma (Wiedemann)
nus is still poorly circumscribed, and especially the separa- as well as an endemic, undescribed species. Most species of
tion from Boettcheria may seem problematic. Little is known Peckia are carrion breeders and important decomposers in
of the biology of the included species (e.g., Lopes, 1982b). tropical forest ecosystems. Peckia chrysostoma and what is
probably P. intermutans (Walker) have been bred from hu-
Nephochaetopteryx Townsend. There are 31 New World, man corpses (Jirón et al., 1983). The large P.  gulo breeds
mainly Neotropical, species of this genus. Eight species have in dead semi-terrestrial crabs in coastal habitats (Méndez &
been recorded from Central America, but several additional Pape, 2003). Pape & Andersson (2001) provide a key to spe-
undescribed species are known, and the genus is probably cies of Peckia known from the Guanacaste region of Costa
rich in species. Females of some species are attracted in Rica.
great numbers to fermenting fruit, while males are much
more rarely encountered. At least some species breed in Peckiamyia Dodge. There are five known species of this
dung and others have been reared from vertebrate carrion Neotropical genus, with two species recorded from Central
(Lopes, 1973a; T. Pape, unpublished). America. At least some species breed in carrion (T.  Pape,
unpublished).
Oebalia Robineau-Desvoidy. This genus contains 12
species worldwide, one of which, O.  costarica Pape, has Phrosinella Robineau-Desvoidy. A total of 23 species
been recorded from Central America. Species of Oebalia have been described in this genus, which is mainly of Hol-
are kleptoparasites of twig-nesting aculeate hymenopterans arctic distribution. A single species, P. aldrichi Allen, extends
(Pape, 1989a). For identification, see Pape (1989a, b). into northern Central America. Several species of Phrosinella
are associated with the nests of aculeate Hymenoptera (Spof-
Opsidia Coquillett. There are 10 New World species ford et al., 1989).
of this genus, of which one, O.  metopioides Allen, has
been recorded from Central America. The Nearctic species Rafaelia Townsend. There are eight New World species
O. gonioides Coquillett is associated with the nests of Halic- of this genus, three of which have been recorded from Cen-
tidae and Sphecidae (Hymenoptera) (Spofford et al., 1989). tral America. The definition of this genus-group taxon is par-
ticularly problematic. Rafaelia aurigena (Lopes) from Chile
Oxysarcodexia Townsend. This is a large genus with 79 and Peru has been reared from snails (Lopes 1969a).
New World species, of which 28 have been recorded from
Central America. Lopes  & Tibana (1987) provide a useful Ravinia Robineau-Desvoidy. This is a large genus with
key, but few species have their genitalia illustrated, and it is 34 described species. Except for one Palearctic species, all
1332
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

have a New World distribution, and 16 species are known Senotainia Macquart. With a total of 64 species, this ge-
from Central America. Species whose biology is known are nus is one of the largest and most widespread of the milto-
dung-breeders, with larvae of at least some species prey- grammine genera. Four species have been recorded from the
ing (possibly facultatively) on other invertebrate larvae northern parts of Central America. Many species of Seno-
(Coffey, 1966; Pickens, 1981). One species, R.  floridensis tainia are typical “satellite flies”, with adult females trailing
(Aldrich), has been reared from dead crabs (G.A. Dahlem, their hymenopteran host during flight at a very exact distance
unpublished). (Spofford et al., 1989).

Retrocitomyia Lopes. There are nine Neotropical species Sphenometopa Townsend. There are 47 species of this
of this genus, two of which are recorded from Central Amer- genus, mostly Nearctic and Palearctic, with two species ex-
ica. Nothing is known of the biology of these flies except that tending into northern Central America. One Nearctic spe-
males are attracted to carrion (T. Pape, unpublished). cies, S. tergata (Coquillett), is associated with the nests of
Pompilidae and Sphecidae (Hymenoptera) (Spofford et al.,
Rettenmeyerina Dodge. The single Neotropical species of 1989).
this genus, R. serrata Dodge, is known from Central America.
Males have been caught over army ant raids (Dodge, 1968). Spirobolomyia Townsend. This genus contains five
mainly Nearctic species, but two of these extend into Neo-
Sarcodexia Townsend. There are two named species in tropical Mexico. Pape (1990) provides illustrations for all
this New World genus, both of which are known from Cen- species. The only biological information available suggests
tral America. The widespread and common S. lambens (Wie- that the species are parasitoids of large spirobolid millipedes
demann) is biologically versatile and has been recorded as an (Aldrich, 1916; Pape, 1990), but they are also attracted to
insect parasitoid, a predator on wounded insects, a myiasis vertebrate carrion (T. Pape, unpublished).
producer in birds and mammals, and as breeding in verte- Tapacura Tibana  & Lopes. There are two New World
brate and invertebrate carrion (Townsend, 1893; Hoffmann, species of this genus, one of which is known from Baja Cali-
1927; Almeida, 1933; Callan, 1946; Parker et al., 1953; Har- fornia Sur in Mexico. Nothing is known of the biology.
rison, 1963; Stegmaier, 1972; Fessl et al., 2001). A third,
undescribed species of Sarcodexia occurs on Cocos Island, Taxigramma Perris. A total of 18 species of this genus
Costa Rica (T. Pape, unpublished). are known. One species, Taxigramma hilarella (Zetterstedt),
extends into the northern parts of Central America, but so far
Sarcofahrtiopsis Hall. The 12 species of this genus are no representatives are known from South America. The New
mainly Central American and Antillean, and seven have been World species are associated with aculeate Hymenoptera
recorded from Central America. A key to all species is given nests (Spofford et al., 1989).
by Méndez et al. (2008). The genus is biologically diverse, Thomazomyia Lopes. Five Neotropical species of this
with larvae of one species found exclusively in the leaf roosts genus have been described, one of which, T. pilipes Lopes,
of the short-tailed bat (Pape et al., 2002), while other species is known from Costa Rica and southern Mexico. Most of the
have been bred from dead fish and semi-terrestrial crabs in species are illustrated by Lopes (1976a).
mangrove habitats (Pape & Méndez, 2002, 2004; Méndez et
al., 2008). Titanogrypa Townsend. There are 18 New World species
of this genus, of which five have been recorded from Central
Sarcophaga Meigen. There are over 800 species world- America. Some species have been reared from snails, and
wide of this genus in the broad definition of Pape (1996). one species from South America, T. larvicida, has been re-
While this genus is well represented over most of the world, corded to have a first instar larva that produces a toxin that
surprisingly few species are known from the Neotropical suppresses other larvae in the same food source (Lopes,
Region. Only 18 species have been recorded from Central 1976b). Lopes (1976b) provides a key to most of the species
America, mostly from the northernmost parts. We consider of the subgenus Cuculomyia Lopes, which we consider as
Sarcophaga in a broad sense, but we realize that a rigorous part of Titanogrypa.
definition and well-supported monophyly is still lacking (see Tricharaea Thomson. The 15 species of this genus are
further in Pape, 1996), and changes are anticipated. While primarily New World, but a few species have been spread by
most species probably are associated with small vertebrate humans to Australia and the Pacific. Seven species have been
and invertebrate carrion, Sarcophaga polistensis Hall has recorded from Central America. Lopes (1956, 1973b, 1986)
been bred from nests of paper wasps (Hall, 1933b), and other will key out most species, but a modern taxonomic revision
species are known to be predators of spider egg-sacs (Lopes, is strongly needed. Mostly saprophagous and coprophagous,
1989). Some synanthropic species breed in larger carrion some species are restricted to coastal areas where they breed
(e.g., S.  crassipalpis Macquart, S.  ruficornis Fabricius) or in fish and invertebrate carrion and damaged turtle eggs
feces (e.g., S. africa (Wiedemann)). (T. Pape, unpublished).
SARCOPHAGIDAE (FLESH FLIES) 111 1333

Tripanurga Brauer & Bergenstamm. There are 14 New Villegasia Dodge. There are three Neotropical species of
World species of this genus, of which six have been recorded this genus, two of which have been recorded from Central
from Central America. Biologically they are still superfi- America. Both of these appear to be coastal species, and
cially known, but most species may be obligate predators on V. postuncinata larvae are in dead crabs (Reeves et al., 2000;
freshwater turtle eggs (Vogt, 1981; T. Pape, unpublished). Méndez  & Pape, 2003) and possibly other carrion. Lopes
(1973a) reported the other species, V.  almeidai (Lopes), to
Udamopyga Hall. Of the 20 New World species of this be frequently found on sea beaches.
genus, two have been recorded from Central America. Dodge
(1965) provides the only key to species, but many illustra- Wulpisca Lopes. There are two Neotropical species of
tions are lacking or poorly drawn. Several species of Udam- this genus, both of which appear to be restricted to Central
opyga are known to breed in dead snails (Lopes, 1973a). America. Nothing is known of their biology.

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(Diptera: Sarcophagidae). Proceedings of the Entomological So- Wcislo, W.T. 1986. Host nest discrimination by a cleptoparasitic fly,
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Thompson, P.H. 1978b. An infestation of miltogrammine Sar- minae). Journal of the Kansas Entomological Society 59: 82–88.
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373–379. 11: 115–125.
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
RHINOPHORIDAE (WOODLOUSE FLIES) 112
Thomas Pape

Fig. 112.1. Male of Bezzimyia bisecta Pape & Arnaud, (Pape & Arnaud, 2001, fig. 3E).

Diagnosis i.e., mostly parallel to vein R1. Vein M various, ending free-

ly in wing membrane (Figs.  5, 6), reaching costa at wing
Small to medium-sized (body length 3.5–8.0 mm) calyp- margin (Figs.  7, 8), or joining vein  R4+5 well before wing
trate flies (Fig.  1). Body color mostly brown or blackish- tip. One case of female aptery known, with wings reduced
brown, more rarely partly or almost entirely yellow. Antenna to halterlike stumps (Fig. 4). Legs with foretarsus unmodi-
with first flagellomere  varying from short (Fig. 11) to longer fied (Fig. 15) or laterally compressed in one or both sexes
than eye height (Fig. 12); concolorous with body or strongly (Fig.  16). Subscutellum moderately swollen. Abdomen
contrasting orange; sometimes lobate in male (Fig. 9). Male oval or elongate oval. Male terminalia with surstylus fused
dichoptic and with neither proclinate nor reclinate orbital to epandrium in native species, cerci with separate prongs,
bristles, or semi-holoptic with fronto-orbital plates almost phallus with dorsolateral processes that often have free tips.
entirely occluded at vertex but frontal stripe only moderately Female terminalia telescopic, cerci elongate.
narrowed. Female without proclinate orbital bristles (some Eggs fusiform with blunt ends, flattened dorsal side and
species with setulae that may be reduced orbitals). Head dorsomedian plastron running full length of egg. Chorion
with receding vibrissal angle (i.e., head moderately or dis- with rugged longitudinal ridges running full length of egg
tinctly shorter at level of vibrissa than at level of antennal and entire surface pitted with small hexagons. Plastron
insertion). Vibrissa indistinct (Figs.  9, 10) or accompanied with somewhat irregular, delicate hexagonal meshwork.
by long genal setae (Figs.  11, 13). Proboscis short; length First instar larvae (Figs.  2, 3) with distinct segmenta-
at most slightly more than greatest diameter of forecoxa. tion, head with long and apparently two-segmented antenna,
Wing membrane hyaline, darkened along veins, fumose with cephalopharyngeal skeleton with long anterior rod, mouth
white wing tip, or with various patterns of fumose bands. hooks toothed, labrum absent. Most segments encircled by
Vein  Sc normal or with indistinct or no median concavity, setae, and integument encrusted with small, scalelike spines.

1337
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

2 Bezzimyia

3 Bezzimyia

4 Bezzimyia
R1
R2+3

R4+5
M1

5 Bezzimyia 6 Bezzimyia

7 Shannoniella 8 Undescribed Genus A

Figs. 112.2–8. Larvae, adults, and wings: lateral view of first instar larva (2) and pseudocephalon and anterior part of thorax (3) of
Bezzimyia yepezi Pape & Arnaud, (South America, Pape & Arnaud, 2001, figs. 3A, 3C); lateral view of apterous female of B. hansoni Pape &
Arnaud (4); dorsal view of wing of (5) B. lapidicina Pape & Arnaud, (Pape & Arnaud, 2001, fig. 9G); (6) B. bisecta Pape & Arnaud, (Pape &
Arnaud, 2001, fig. 9B); (7) Shannoniella cuspidata Townsend, (South America); and (8) Undescribed Genus A. Figure 4 photographed by
T. Pape. Figures 7–8 illustrated by E. Binkiewicz.
RHINOPHORIDAE (WOODLOUSE FLIES) 112 1339

9 Bezzimyia 10 Bezzimyia 11 Bezzimyia

12 Shannoniella 13 Trypetidomima 14 Undescribed Genus A

15 Bezzimyia

16 Bezzimyia

Figs. 112.9–16. Heads, tibiae, and tarsi: lateral view of head of (9) Bezzimyia bisecta Pape & Arnaud, (Pape & Arnaud, 2001, fig. 7D);
(10) B. sternothrix Pape & Arnaud, (Pape & Arnaud, 2001, fig. 8G); (11) B. lapidicina Pape & Arnaud, (Pape & Arnaud, 2001, fig. 8A);
(12)  Shannoniella cuspidata Townsend, (South America); (13)  Trypetidomima lutea Townsend, (South America); and (14)  Undescribed
Genus A; posterior view of left foreleg, enlarged inset showing tarsomere 2 with complete vestiture, of (15) B. ramicornis Pape & Arnaud,
(South America, Pape & Arnaud, 2001, fig. 11C); and (16) B. setifax Pape & Arnaud, (Pape & Arnaud, 2001, fig. 11D). Figures 12–14
illustrated by E. Binkiewicz.
1340
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Abdominal segment  8 equipped with terminal pair of pro- Classification

cesses having broad, vesicular bases and tapering into long,
slender filaments. The only known Neotropical larva is the Rhinophoridae are members of the Tachinidae family
first instar from a Venezuelan species of Bezzimyia (Pape & group (or Oestroidea) and have been considered a full family
Arnaud, 2001). (Crosskey, 1977) or a subfamily of either Tachinidae (Sa-
While larvae of Rhinophoridae are truly unique, adults brosky  & Arnaud, 1965) or Calliphoridae (Rognes, 1991).
have no distinctive features that easily separate them Recently Pape  & Arnaud (2001) proposed a sister-group
from other tachinoid families. The combination of oval relationship to the Rhiniinae (Calliphoridae). Note that the
or tongue-shaped lower calypteres, small, triangular overwhelming evidence for calliphorid non-monophyly pre-
sented by Rognes (1997) necessitates some additional rank-
metathoracic spiracles, and the moderately (not greatly)
ing, but this need not involve the Rhinophoridae.
swollen subscutellum is a good shortcut but will include The Rhinophoridae is a small family with 142 named species
a few Tachinidae. The presence of separate dorsolateral worldwide, 15 of which have been recorded from the Neotropi-
phallic processes in the male and telescopic terminalia cal Region. Several unnamed Neotropical species are known
with elongate cerci in the female will exclude them from (T. Pape, unpublished) and more certainly await discovery.
these Tachinidae.
Identification
Biology The majority of New World Rhinophoridae are treated by
Pape  & Arnaud (2001), who monographed Bezzimyia, but
All species for which the biology is known are internal several species show moderate to considerable sexual dimor-
parasitoids of woodlice (Crustacea: Isopoda, Oniscoidea). phism, and most species are known from the male sex only.
Eggs are dispersed freely on the surfaces of objects like The monotypic Shannoniella and Trypetidomima are hardly
withered leaves and bark, with a preference for surfaces con- treated in the literature apart from their short original descrip-
taminated by woodlice. Eggs hatch after about a week, and tion (Townsend, 1935, 1939). Melanophora roralis (Linnaeus)
first instar larvae attach to passing woodlice. The puparium is treated in Herting’s (1961) and Crosskey’s (1977) treatments
is formed inside the completely empty host exoskeleton. of the Palearctic and Afrotropical faunas, respectively.
Host specificity is generally high (R. Bedding, unpublished). Routine pinning is the recommended way of preserving
adult Rhinophoridae. While known species may be recog-
Nothing is known of larval host relations for native New
nized without examination of male terminalia, species com-
World Rhinophoridae, but the morphology of the first instar plexes should be anticipated and it is recommended that newly
larva strongly suggests habits similar to European species caught male specimens have their terminalia extended. This
(Pape & Arnaud, 2001). is most easily done with an insect pin on relaxed specimens
Most specimens of Bezzimyia have been caught at mid- after disappearance of rigor mortis. Immature stages are best
elevations (500–2300 m). preserved in 80% alcohol or as permanent slides.

Key to the genera of Rhinophoridae of the Neotropical Region

1. Wing vein M complete and reaching costa at wing margin (Figs. 7, 8) . . . . . . . . . . . . . . . . . . . . 2
– Wing vein M not reaching wing margin; either straight, incomplete and ending freely in wing
membrane (Figs. 5, 6), or curved abruptly anteriorly and fused to vein R4+5 well before wing
margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

2. Veins R1 and R4+5 with row of several, long setulae (Fig.  7); male vibrissal angle drawn out
(Fig. 12); male antenna with first flagellomere about as long as eye height (Fig. 12), female
first flagellomere at most as long as 0.5 times eye height; Brazil . . . . . . [Shannoniella Townsend]
– Veins R1 and R4+5 bare or with one to two short setulae (Fig. 8); male vibrissal angle not drawn out;
male and female antenna with first flagellomere at most as long as 0.5 times eye height (Figs. 13,
14) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
RHINOPHORIDAE (WOODLOUSE FLIES) 112 1341

3. Head, body and legs extensively yellow; frontal bristles long, many as long as smallest diameter
of eye (Fig. 13); lower calypter with long, blackish setulae along margin; Brazil . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Trypetidomima Townsend]
– Head and body black, legs yellow; frontal bristles short, at most 0.5 times smallest diameter of eye
(Fig. 14); lower calypter without setulae along margin. . . . . . . . . . . . . . . . . Undescribed Genus A

4. Wing vein M straight and ending freely in wing membrane (Figs. 5, 6); both sexes without or with
minute proclinate orbital bristles; female with patterned or unpatterned wings, but not with white
wing tip. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bezzimyia Townsend
– Wing vein M fused to vein R4+5 well before wing margin; both sexes with four to six proclinate
orbital bristles; female with white wing tip; Argentina, Brazil, Chile, West Indies. . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Melanophora Meigen]

Synopsis of the fauna Species of this genus are collected in humid forests and seem
most common at mid-elevations (500–2300 m). Both sexes
Twelve species of Rhinophoridae are known from Central are active during the day and may be seen sunning themselves
America, distributed in two genera, one of which is still on leaves close to the ground. A few specimens have been
unnamed. Most specimens have been taken in humid forests taken at light.
at mid-elevations.
Undescribed Genus A. There is only a single, undescribed
Bezzimyia Townsend. There are 17 named species, 11 species of this genus known, collected from Costa Rica and
of which occur in Central America (Pape & Arnaud, 2001). Nicaragua (T. Pape, unpublished).

Literature cited

Crosskey, R.W. 1977. A review of the Rhinophoridae (Diptera) and Rognes, K. 1997. The Calliphoridae (Blowflies) (Diptera: Oestroi-
a revision of the Afrotropical species. Bulletin of the British Mu- dea) are not a monophyletic group. Cladistics 13: 27–66.
seum of Natural History (Entomology) 36: 1–66. Sabrosky, C.W. & P.H. Arnaud, Jr. 1965. Rhinophorinae, pp. 963–
Herting, B. 1961. 64e. Rhinophorinae. In Lindner,  E. (edi- 964. In Stone, A., C.W. Sabrosky, W.W. Wirth, R.H. Foote, &
tor). Die Fliegen der palaearktischen Region 9: 1–36. E. J.R.  Coulson (editors). A catalog of the Diptera of America
Schweizerbart’sche Verlagsbuchhandlung, Stuttgart. north of Mexico. United States Department of Agriculture,
Pape, T.  & P.H. Arnaud, Jr. 2001. Bezzimyia — a genus of New Agriculture Handbook 276, Washington, D.C., iv + 1696 pp.
World Rhinophoridae (Insecta, Diptera). Zoologica Scripta 30: Townsend, C.H.T. 1935. New muscoid genera, mainly from the
257–297. Neotropical Region. Revista de Entomologia 5: 68–74.
Rognes, K. 1991. Blowflies (Diptera, Calliphoridae) of Fenno- Townsend, C.H.T. 1939. New striking melanophorid tribe and gym-
scandia and Denmark. Fauna entomologica scandinavica 24: nosomatid genera from Rio de Janeiro (Diptera, Oestromusca-
1–272. ria). Revista de Entomologia 10: 249–254.
1342
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113
D. Monty Wood and Manuel A. Zumbado

Fig. 113.1. Male of Juriniopsis badiiventris (Wulp), (Zumbado, 1999, p. 111).

Diagnosis ally present, the latter sometimes directed laterally (laterocli-

nate) or posteriorly (reclinate). Upper orbital bristles usually
Small to large flies (body length 3–25 mm), usually bris- present, anterior one(s) reclinate, posteriormost sometimes
tly (Fig. 1), extremely varied in shape and proportions. Most lateroclinate. Occipital setae usually long and fine, delin-
genera of usual calyptrate shape, but some slender and wasp- eated laterally by evenly spaced vertical row of stouter pos-
like in appearance. Frons of male either narrower than that of toccipital bristles, either extending ventrally to meet lower
female of same species and usually lacking proclinate orbital cranial margin (Figs.  14, 30) or curving forward on genal
bristles, or about as wide as that of female and usually also dilation (a setose strip or patch between eye and lower cra-
with proclinate orbital bristles; first flagellomere of latter nial margin; Figs.  20, 27, 28), that may also have one or
type usually much longer in male than in female, in some more large bristles (Fig. 4). Vibrissa usually well developed,
species bifurcate, trifurcate, or multibranched (fissicorn). usually accompanied by one or more smaller bristles both
Arista usually bare, or short to long plumose in most Dexii- below (on subvibrissal ridge) and above it (on facial ridge).
nae. Eye usually with ommatrichia, which may be long and Face concave or convex, often protruding between and/or
dense or so short and sparse as to make the eye appear bare; below vibrissae. Facial ridge, especially in species with long
ommatrichia usually longer in male than in female and often antennae, usually with row of erect bristles. Proboscis usu-
longer dorsally than ventrally. Ocelli and ocellar bristles usu- ally short and straight, with padlike labella, but may be long

1343
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

and curved, with small labella. Prosternum pilose in most a scorpion, and two genera of centipedes are also parasitized
Goniinae, bare in Phasiinae and most Dexiinae and Tachini- (Stireman et al., 2006). At least eight orders of insects are
nae. Proepisternum occasionally pilose. Postpronotum with afflicted, although the great majority of tachinids attack lar-
two (Fig.  93) or usually three major bristles in transverse vae of Lepidoptera. Larvae and adults of Coleoptera come
row (Fig. 95), sometimes preceded by fourth smaller bristle; second in the list of most favored hosts, followed by bugs
or with middle bristle displaced anteriorly (Fig. 94). Postsu- (Hemiptera: Heteroptera) of several families, then larvae
tural supra-alar bristles typically three, middle bristle larg- of sawflies and social wasps (Hymenoptera). Adults and
est (unless first is lacking), anterior one usually smaller and nymphs of the orthopteroid orders, including Blattaria, Phas-
thinner than first postsutural dorsocentral bristle (Figs. 103, mida, and Mantodea, as well as Orthoptera themselves are
104), except larger than first postsutural dorsocentral and not immune from tachinid attack, and grasshoppers have
nearly as large as middle bristle (Fig. 102) in tribe Tachinini been adopted as hosts by several different lineages. Larvae
of subfamily Tachininae and tribes Goniini, Eryciini, and of Tabanidae (Diptera) are parasitized by a subgroup of
most Winthemiini of subfamily Exoristinae. Scutellum with tachinids that normally develop on beetle grubs in soil or
one to four pairs of bristles (basal, lateral, subapical, and rotten wood, while at least three unrelated lineages attack
apical), lateral bristles sometimes duplicated. Subscutellum larvae of Tipulidae. Even Dermaptera and Embioptera serve
well developed, evenly rounded in profile. Posterior thoracic as hosts for a few species. Most tachinid species are either
spiracle small, confined to region behind uppermost meral host specific or are associated with a limited number of dif-
bristles, and well behind katepisternum, separated from latter ferent species of hosts (Janzen  & Hallwachs, 2005), but a
by rectangular katepimeron. Meral bristles present. Vein M few have the capacity to attack many species.
with distinct bend. A tachinid larva begins its life as a parasite, growing
All Tachinidae of Central America (except for the rare along with the host as it continues to feed; then at some point
genus Cinochira) share with all Sarcophagidae and Calli- in its development, it usually kills the host and thus becomes
phoridae the presence of a row of well-developed meral bristles a parasitoid. There are, however, a few instances of hosts
(Fig. 105), combined with a bend in vein M (Fig. 157). This surviving to maturity after the tachinid larvae have depart-
bend may vary considerably, from being broadly obtuse to ed (Heitzmann-Fontenelle  & Guimarães, 1966; DeVries,
forming a right angle as in Exoristini, (Fig. 157) or even an 1984).
acute angle (Fig. 154). The bend is also found in some mem- During early development, tachinid larvae of most species
bers of the subfamily Muscinae of the Muscidae, but not in become enshrouded in a sac of host tissue, the respiratory
other muscids, nor in Anthomyiidae or Scathophagidae, and funnel, laid down by the host around them in an effort to
meral bristles are lacking in all Muscidae, Anthomyiidae, encapsulate them. The larva’s posterior end, on which the
and Scathophagidae. Meral bristles are present in all Central posterior spiracles are located, is anchored by hooks either
American Tachinidae, but are lacking in the Brazilian phasi- to the skin of the host or to a tracheal trunk, providing the
ine genus Tarassus Aldrich, not yet recorded from Central tachinid larva with access to air, while the anterior end pro-
America. The single feature that distinguishes all Tachinidae trudes from the respiratory funnel free to probe and consume
from most members of the other families is the presence of an host tissues. As the tachinid larva grows, the funnel continues
evenly convex subscutellum. A few calliphorids, especially to enlarge around it. A few species, in contrast, develop free
some members of the genus Mesembrinella Giglio-Tos, also in the haemocoel (Thompson, 1915), presumably absorbing
have a convex subscutellum, but have an enlarged posterior oxygen from the host’s haemolymph. At least one species,
spiracle that extends anteriorly nearly to the posterodorsal Compsilura concinnata (Meigen), develops between the gut
corner of the katepimeron almost obliterating the katepimeron. wall and the peritrophic membrane, where it must escape the
Central American Rhinophoridae have a weakly convex host’s defensive reaction (Ichiki & Shima, 2003). Compsilura
subscutellum but lack the apex of vein M, as the vein ends concinnata was introduced into North America in the early
in the membrane where the bend otherwise occurs. A few twentieth century to control gypsy moth, but has now spread
acalyptrates also have a moderately developed subscutellum widely over the continent where it attacks a wide variety
but lack meral bristles. When alive, most tachinids hold their of caterpillars. It has even been accused of decimating native
wings at a wider angle to the body than do sarcophagids and saturniid moths. Many species of Eucelatoria, close relatives
calliphorids, and most are also more active, seldom stay- of Compsilura Bouché, occur in Central America, and some
ing in one place for more than a few seconds. Tachinids are may be capable of similar development.
widely believed to be more bristly than members of the other It is probably safe to say that development in the body
families, but this is not a reliable criterion. of another arthropod is characteristic of all members of the
family. This type of development is not confined to the
Biology Tachinidae among the calyptrates, however, but is also found
in Rhinophoridae (Thompson, 1934) and perhaps also some
All species of Tachinidae whose life cycles are known Sarcophagidae. Larvae of the former develop only in isopods
develop as larvae inside the bodies of other arthropods. and can also stimulate the host to form a respiratory funnel
Hosts are almost exclusively other insects, although a spider, around them. Larvae of Sarcophagidae, on the other hand,
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1345

tend to treat the host as a source of carrion, although a successful groups of tachinids have evolved strategies to
parasitic existence is undoubtedly true for some species. overcome these challenges by leaving host finding to chance
The majority of female Tachinidae retain their eggs af- or to their offspring. Because of the greater risks involved,
ter fertilization in a greatly elongate and expanded vagina, females must consequently lay a larger number of eggs than
called the uterus or ovisac. Members of subfamilies Tachini- those that can oviposit directly.
nae and Dexiinae, and the great majority of the Exoristinae, One strategy is the placement of minute eggs, each ready
share this feature. However, members of subfamily Phasii- to hatch, on the food of the host. Females of Goniini lay
nae, as well as several genera of Exoristinae, including all “microtype” eggs; tiny limpetlike structures with a thick-
Exoristini and Winthemiini, deposit unembryonated eggs on ened upper shell to resist desiccation, while females of the
the host, and Herting (1963) has argued convincingly that blondeliine genus Anisia lay extremely small, transparent
this must represent the ancestral condition within Tachinidae eggs that are eaten by their orthopteroid hosts. Some goni-
from which the uterus arose. There is no reason to suppose ine females actually find the host and deposit eggs near it
that the retention of eggs until the first instars are ready to on the chance they will be eaten. Other species oviposit on
hatch has arisen in the Tachinidae more than once, although damaged leaves. Hosts of Goniini are mostly caterpillars, but
it also frequently occurs in other calyptrates. Sarcophagidae adult beetles and larvae of Tipulidae are also parasitized by
retain their eggs within a special bilobed diverticulum of the eating microtype eggs.
vagina until they hatch and larviposit rather than oviposit; Another way to circumvent a host’s hiding place is to
several lineages of Calliphoridae also retain their first instars deposit embryonated eggs in a suitable habitat, leaving their
inside the vagina; and Oestrinae have an elaborate uterus fate to the searching ability of the mobile first instars. Prob-
with separate pouches for each developing egg. However, ably all members of two unrelated tribes, Dexiini and Mega-
the ancestral condition of the calyptrates must have been prosopini, drop their embryonated eggs on the ground, on
oviposition of unembryonated eggs, as occurs in almost rotten wood, or at the mouths of host tunnels; the hatching
all Diptera, as well as almost all Muscidae, Anthomyiidae, larvae burrow directly down into the substrate or frass and,
Scathophagidae, most Calliphoridae and Oestridae, and some if they are lucky, find a suitable beetle grub and penetrate
Tachinidae. Members of subfamily Phasiinae, as well as sev- it. It is not known to what extent a gravid female fly can
eral groups in the subfamily Exoristinae, including all mem- detect the presence of grubs buried beneath it or whether it
bers of the tribes Exoristini, Ethillini, and Winthemiini, have chooses the larval habitat on surface features. Females of
a short vagina that can contain only a single fertilized egg at tribe Myiophasiini detect weevil grubs in acorns and other
a time. Females of these taxa must deposit unembryonated fruits by searching for the holes the female weevil made to
eggs on the host (unless a single fertilized egg was carried oviposit in the fruit. Some Blondeliini (Lixophaga) as well
about in the vagina for a day or two before being laid). This as a few Eryciini (Lydella) have also adopted this strategy to
type of oviposition must surely be the ancestral condition in attack larvae in stem borings and in webs.
the Tachinidae. First instar development is completed on the A third strategy involves leaving embryonated eggs in
outside of the host, where it is subject to being eaten, rubbed places where the host is likely to pass later. Probably all
off, or lost as a result of host ecdysis, leading to selection members of the tribes Tachinini and Polideini, and probably
pressure for a speedier method of host penetration. some other members of the subfamily Tachininae as well,
One method of speedier penetration is by direct injec- drop their eggs where their hosts might pass; the first instars,
tion, and several genera of Phasiinae have evolved elabo- which are armored with thickened platelets to resist desicca-
rate piercing and grasping structures to hold and inject their tion, simply wait out their lives in hope of attaching them-
bug hosts with unembryonated eggs, which continue their selves to a passing arthropod. It is in the Polideini where
development and hatch inside the host. Some genera of the greatest range of different types of host occur, including
Blondeliini, likewise, have the seventh sternite of the female those that attack scorpions, spiders, centipedes, cockroaches,
modified into a sickle-shaped hook for guiding the remain- earwigs, and web-spinners (Embioptera), as well as a wide
ing abdominal segments inside the host and depositing their range of species of caterpillars.
embryonated eggs where the larvae are immediately able to One of the most sophisticated strategies is used by females
take up a position for feeding. of Ormia and perhaps by all members of the tribe Ormiini.
Female tachinids are essentially diurnal, although a few, Using tympanic membranes on each half of their enlarged
particularly those that parasitize adult scarabs, are crepuscu- prosternum, they can locate singing male orthopterans, in-
lar, and at least one species of Ormia that attacks stridulat- cluding katydids, crickets, and mole crickets, and by firing
ing nocturnal Orthoptera can apparently function in darkness eggs in the direction of the sound they can place their first
(Hoy & Robert, 1996). For those females that must find their instars in close proximity to a suitable host (Hoy & Robert,
host in order to oviposit directly on it, this limits the choice 1996).
of hosts to those that are accessible during the day or eve- When larval development has been completed, tachinid
ning. Nocturnal hosts, and those hidden in soil or wood or larvae usually leave the carcass of the host and enter the soil
that have surrounded themselves with webbing, do not escape to form a puparium. Some beetle parasitoids, however, take
parasitism, however, for several large and presumably advantage of the protective shell of the beetle and form their
1346
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

plp

2 Peleteria 3 Copecrypta 4 Bombyliomyia

oc s

oc s

comp eye

pafc
pafc

pafc s
gn
gn

5 Winthemia 6 Chrysoexorista 7 Carcelia

oc s
oc s

pafc s
pafc s

fc s pafc
pafc s fc s
vb

l fc mg

8 Distichona 9 Gaediopsis 10 Mystacella

Figs. 113.2–10. Heads: lateral view of (2)  Peleteria anaxias (Walker), (Nearctic, MND, fig.  110.62); (3)  Copecrypta ruficauda (Wulp),
(Nearctic, MND, fig. 110.63); (4) Bombyliomyia sp.; (5) Winthemia rufopicta (Bigot), (Nearctic, MND, fig. 110.5); (6) Chrysoexorista sp.,
(Nearctic, MND, fig. 110.6); (7) Carcelia reclinata (Aldrich & Webber), (Nearctic, MND, fig. 110.7); (8) Distichona autumnalis (Townsend),
(Nearctic, MND, fig. 110.2); (9) Gaediopsis sierricola (Townsend), (Nearctic, MND, fig. 110.9); and (10) Mystacella frioensis (Reinhard),
(Nearctic, MND, fig. 110.10). Figure 4 illustrated by A. Brenes.
Abbreviations: comp eye, compound eye; fc s, facial setae; gn, gena; l fc mg, lower facial margin; oc s, ocellar seta; pafc, parafacial; pafc s,
parafacial setae; plp, palp; vb, vibrissa.
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1347

oc s

rc orb s

fr s

ar 2
ptil fis spvb s
pafc s

l fc mg

11 Gonia 12 Chaetogaedia 13 Ametadoria

spvb s
l fc mg pafc s

14 Argyrochaetona 15 Chetogena 16 Paradidyma

17 Anisia 18 Borgemeiermyia 19 Thelairodoria

Figs. 113.11–19. Heads (continued): lateral view of (11) Gonia frontosa Say, (Nearctic, MND, fig. 110.4); (12) Chaetogaedia analis (Wulp),
(Nearctic, MND, fig. 110.17); (13) Ametadoria harrisinae (Coquillett), (Nearctic, MND, fig. 110.32); (14) Argyrochaetona cubana Townsend;
(15) Chetogena tachinomoides (Townsend), (Nearctic, MND, fig. 110.35); (16) Paradidyma affinis Reinhard, (Nearctic, MND, fig. 110.33);
(17) Anisia sp.; (18) Borgmeiermyia sp.; and (19) Thelairodoria sp. Figures 14 and 17–19 illustrated by A. Brenes.
Abbreviations: ar 2, second aristomere; fr s, frontal setae; l fc mg, lower facial margin; oc s, ocellar setae; pafc s, parafacial setae; ptil fis,
ptilinal fissure; rc orb s, reclinate orbital setae; spvb s, supravibrissal setae.
1348
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

oc s

pc orb s

pafc s

spvb s
gn dil
vb
sbvb s

20 Myiopharus 21 Admontia 22 Celatoria

pc orb s

spvb s

vb
sbvb s gn s

23 Eribella 24 Anoxynops 25 Phyllophilopsis

pafc s
pafc s vb
gn
sbvb s dil gn
dil

26 Vibrissina 27 Cryptomeigenia 28 Trigonospila

Figs. 113.20–28. Heads (continued): lateral view of (20)  Myiopharus doryphorae (Riley), (Nearctic, MND, fig.  110.41); (21)  Admontia
degeerioides (Coquillett), (Nearctic, MND, fig. 110.42); (22) Celatoria diabroticae (Shimer), (Nearctic, MND, fig. 110.43); (23) Eribella
exilis (Coquillett), (Nearctic, MND, fig. 110.44); (24) Anoxynops aldrichi (Curran), (Nearctic, MND, fig. 110.45); (25) Phyllophilopsis nitens
(Coquillett), (Nearctic, MND, fig. 110.46); (26) Vibrissina hylotomae (Coquillett), (Nearctic, MND, fig. 110.38); (27) Cryptomeigenia theutis
(Walker), (Nearctic, MND, fig. 110.40); and (28) Trigonospila verticalis (Reinhard), (Nearctic, MND, fig. 110.49).
Abbreviations: gn dil, genal dilation; gn s, genal setae; oc s, ocellar setae; pafc s, parafacial setae; pc orb s, proclinate orbital setae; sbvb s,
subvibrissal setae; spvb s, supravibrissal setae; vb, vibrissa.
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1349

29 Athrycia 30 Voria 31 Cyrtophleba

32 Metaplagia 33 Ebenia 34 Periscepsia

pafc s

35 Kirbya 36 Muscopteryx 37 Eulasiona

Figs. 113.29–37. Heads (continued): lateral view of (29)  Athrycia cinerea (Coquillett), (Nearctic, MND, fig.  110.65); (30)  Voria ruralis
(Fallén), (MND, fig.  110.71); (31)  Cyrtophleba coquilletti Aldrich, (Nearctic, MND, fig.  110.72); (32)  Metaplagia  sp.; (33)  Ebenia  sp.;
(34)  Periscepsia helymus (Walker), (Nearctic, MND, fig.  110.90); (35)  Kirbya setosa (Townsend), (Nearctic, MND, fig.  110.69);
(36)  Muscopteryx  sp.; and (37)  Eulasiona comstocki Townsend, (Nearctic, MND, fig.  110.37). Figures  32, 33, and 36 illustrated by
A. Brenes.
Abbreviation: pafc s, parafacial setae.
1350
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

fc

38 Trafoia 39 Linnaemya 40 Panzeria

fc car

41 Euthera 42 Penthosia 43 Gnadochaeta

44 Microphthalma 45 Pyrrhodexia 46 Acronacantha

Figs. 113.38–46. Heads (continued): lateral view of (38) Trafoia arctica (Sack), (Nearctic, MND, fig. 110.116); (39) Linnaemya tessellata
(Brooks), (Nearctic, MND, fig.  110.101); (40)  Panzeria ampelus (Walker), (Nearctic, MND, fig.  110.102);  (41) Euthera tentatrix Loew,
(Nearctic, MND, fig. 110.122); (42) Penthosia satanica (Bigot), (MND, fig. 110.98); (43) Gnadochaeta clistoides (Townsend), (Nearctic,
MND, fig. 110.92); (44) Microphthalma sp.; (45) Pyrrhodexia sp.; and (46) Acronacantha sp. Figures 44–46 illustrated by A. Brenes.
Abbreviations: fc, face; fc car, facial carina.
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1351

fr s

ped
ped s
ar
fc car

sbvb s

47 Ptilodexia 48 Thelairodes 49 Strongygaster

50 Leskia 51 Eutrixa 52 Cholomyia

53 Clausicella 54 Beskia 55 Crocinosoma

Figs. 113.47–55. Heads (continued): (47)  Ptilodexia sp., (Nearctic, MND, fig.  110.64, as Estheria); (48)  Thelairodes  sp.;
(49)  Strongygaster triangulifer (Loew), (Nearctic, MND, fig.  110.99); (50)  Leskia similis (Townsend), (Nearctic, MND, fig.  110.120);
(51) Eutrixa exilis (Coquillett), (Nearctic, MND, fig. 110.121); (52) Cholomyia sp.; (53) Clausicella politura (Reinhard), (Nearctic, MND,
fig. 110.113); (54) Beskia aelops (Walker), (MND, fig. 110.111); and (55) Crocinosoma cornuale Reinhard, (MND, fig. 110.115). Figures 48
and 52 illustrated by A. Brenes.
Abbreviations: ar, arista; fc car, facial carina; fr s, frontal setae; ped, pedicel; ped s, pedicellar seta; sbvb s, subvibrissal setae.
1352
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

rc orb s

u orb s

o fr s

56 Spallanzania 57 Atacta 58 Neobrachelia

rc orb s rc orb s
rc orb s
pc orb s

59 Pseudochaeta 60 Pseudochaeta 61 Pseudochaeta

oc s

oc s
rc orb s
rc orb s

62 Lydella 63 Drino 64 Calodexia

pc orb s

pc orb s

65 Euhalidaya 66 Actinodoria 67 Medina

Figs. 113.56–67. Heads (continued): anterodorsal view of frons of (56) Spallanzania hesperidarum (Williston), (Nearctic, MND, fig. 110.21);
(57)  Atacta brasiliensis Schiner, (MND, fig.  110.31); (58)  Neobrachelia  sp.; (59)  Pseudochaeta (Pseudochaeta) argentifrons Coquillett,
(Nearctic, MND, fig.  110.23); (60)  P.  (Metopiops) pyralidis Coquillett, (Nearctic, MND, fig. 110.24); (61)  P.  (Phaenopsis) venusta
(Reinhard), (Nearctic, MND, fig. 110.22); (62) Lydella thompsoni Herting, (Nearctic, MND, fig. 110.29); (63) Drino rhoeo (Walker), (MND,
fig. 110.30); (64) Calodexia sp., (Wood, 1985, fig. 2); (65) Euhalidaya genalis (Coquillett), (Nearctic, MND, fig. 110.52); (66) Actinodoria
argentea Thompson, (Wood, 1985, fig. 3); (67) Medina barbata (Coquillett), (Nearctic, MND, 110.53). Figure 58 illustrated by A. Brenes.
Abbreviations: o fr s, outer frontal setae; pc orb s, proclinate orbital setae; rc orb s, reclinate orbital setae; u orb s, upper orbital seta.
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1353

strap-like
ext of oc tr

lun

68 Hemyda 69 Oestrophasia 70 Zelia

fc car

71 Myiomima 72 Gymnosoma 73 Gymnoclytia

i vt s

fc car
ptil fis
fc rg

spvb s

74 Billaea 75 Admontia 76 Campylocheta

Figs. 113.68–76. Heads (continued): anterodorsal view of frons of (68) Hemyda aurata Robineau-Desvoidy, (Nearctic, MND, fig. 110.58);
(69) Oestrophasia clausa Brauer & Bergenstamm, (Nearctic, MND, fig. 110.59); and (70) Zelia vertebrata (Say), (Nearctic, MND, fig. 110.55);
anterodorsal view of head of (71)  Myiomima sarcophagina Brauer  & Bergenstamm; (72)  Gymnosoma canadensis (Brooks), (Nearctic,
MND, fig. 110.87); (73) Gymnoclytia immaculata (Macquart), (Nearctic, MND, fig. 110.88); (74) Billaea satisfacta (West), (Nearctic, MND,
fig.  110.83); (75)  Admontia degeerioides (Coquillett), (Nearctic, Wood, 1985, fig.  1); and (76)  Campylocheta semiothisae (Brooks),
(Nearctic, MND, fig. 110.109). Figure 71 illustrated by A. Brenes.
Abbreviations: ext of oc tr, extension of ocellar triangle; fc car, facial carina; fc rg, facial ridge; i vt s, inner vertical seta; lun, lunule; ptil fis,
ptilinal fissure; spvb s, supravibrissal setae.
1354
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

vb

l fc mg paclyp
scl

77 Mystacomyia 78 Leschenaultia 79 Blepharipa 80 Clausicella

l fc mg
paclyp
scl

lbl

81 Siphona 82 Ginglymia 83 Adejeania

84 Masiphya 85 Prosenoides 86 Plagiomima

Figs. 113.77–86. Heads (concluded): anterior view of face and clypeus of (77)  Mystacomyia rubriventris (Wulp), (MND, fig.  110.149,
as M.  scordala); right half of face and vibrissal area of (78)  Leschenaultia leucophrys (Wiedemann); left half of face and vibrissal area
of (79)  Blepharipa albicauda (Townsend); anterior view of face and clypeus of (80)  Clausicella politura (Reinhard), (Nearctic, MND,
fig.  110.152): lateral view of lower half of head and proboscis of (81)  Siphona intrudens (Curran), (Nearctic, MND, fig.  110.143);
(82) Ginglymia acrirostris Townsend, (Nearctic, MND, fig. 110.144); (83) Adejeania vexatrix (Osten Sacken), (Nearctic, MND, fig. 110.145);
(84) Masiphya sp., (Nearctic, MND, modified from fig. 110.141); (85) Prosenoides flavipes Coquillett, (Nearctic, MND, fig. 110.147); and
(86) Plagiomima similis (Townsend), (Nearctic, MND, fig. 110.147). Figures 78, 79, and 84 illustrated or modified by A. Brenes.
Abbreviations: l fc mg, lower facial margin; lbl, labellum; paclyp scl, paraclypeal sclerite; vb, vibrissa.
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1355

puparium within it. Tachinid larvae in caterpillars that have Genera, like all taxa, are hypotheses and are thus subject to
delayed development until their host pupates may also form rigorous study. Choice of taxa to use, as well as their breadth
their puparium within the empty host pupa. Most temperate or narrowness, are admittedly a matter of individual per-
region tachinids overwinter in their puparium, unless they ception, preference, and tradition, and unfortunately an ex-
are multivoltine, but some overwinter as first instars, tremely narrow interpretation of genera has persisted to the
concealed from the host’s immune responses by hiding in a present in several muscoid families studied by Townsend.
fat body or the brain until the host pupates. They then resume This narrowness is gradually being supplanted by more in-
development in spring. As the host pupa presumably has no clusive and thus more meaningful, informative genera. On
defences, this strategy probably permits a wider host choice. the other hand, genera can be too heterogeneous (the use of
“Phorocera” in the early years of the twentieth century, for
example), and their information content thus declines with
Classification increasing diversity. A correct balance has yet to be achieved
in many tribes of Tachinidae.
An important goal of systematics is the discovery of In this chapter, genera treated in the key are based on an
relationships among taxa. The history of tachinid system- overall similarity, or “gestalt”, of the constituent species. Ide-
atics, however, particularly at the generic level, has been ally a concept, once perceived, is immediately recognizable
almost entirely preoccupied with emphasis on differences at a glance. Although this gestalt can be difficult to charac-
in chaetotaxy to the neglect of similarities and relationships terize, once learned it can be more reliable for grouping spe-
based on other anatomical structures. The family is admit- cies into genera than presence or absence of bristles. Once
tedly large, diverse, and complex, the dipteran equivalent this appearance is appreciated, new members of the genus
of the Asteraceae, Poaceae, or Orchidaceae among vascu- may be recognized more readily, and the diagnosis of the
lar plants. This huge diversity of superficially similar forms genus can be adapted or altered if necesary to accommodate
within the family is partly to blame for the family’s confused them. Character states found only in one sex, for example,
classification and subsequent unpopularity, but this may be branched male antennae or peculiarities of the terminalia,
as much a reflection of the personalities of various special- have been popular in the past. Although useful for indicating
ists of the group, as of any special features of the flies them- relationships among species, they have no value in character-
selves. It has been said that the Tachinidae have so recently izing the opposite sex and thus are of limited usefulness and
radiated that there has been little extinction, but there is no are therefore not used here for generic recognition. In many
evidence for this belief beyond the lack of fossils. Begin- instances, single character states of the chaetotaxy are used
ning with Robineau-Desvoidy (1830, 1863a,  b), followed in the key, but the genera are not based solely on these differ-
by Brauer  & Bergenstamm (1889, 1891, 1893, 1895), and ences, and as new species are found the character states used
brought to its most extreme state by Townsend (1936–1942), to separate them ought to be altered to accommodate them.
genera were split nearly to the level of one genus per spe- In the past, chaetotaxy was used to characterize genera when
cies. Generic names each containing only one or two species it is probably more suited to separating species. Positions of
reduces to a minimum the amount of biological information bristles relative to one another often seem more reliable than
provided by the classification, as well as increasing memory the presence or absence of a particular bristle.
work and creating confusion among numerous similar names All authors today agree that the family is monophyletic.
and long lists of synonyms. In the end, Townsend described The same cannot be said for the subfamilies, for many of
992 generic names to accompany 1086 specific names for the tribes, or even for many genera, which seem to be inter-
the New World fauna on top of the many proposed by his connected to other tribes or genera to form a complex web,
predecessors. He further complicated the task of interpret- joined by taxa that cannot be readily included satisfactorily
ing his work by extensively using abbreviations for various in either. In-depth studies of the male and female termina-
morphological structures and not illustrating them. Subse- lia have clarified relationships among the Australian and
quent authors faced with this multiplicity of generic names Palearctic taxa (Cantrell, 1988; Tschorsnig, 1985; Shima,
have had little choice, when their specimens did not fit the 1989), but such studies have yet to be undertaken for most
existing narrow definitions, but to describe yet more genera. Neotropical taxa.
This trend can be illustrated by the genus Calodexia, one
Subfamily Phasiinae used to comprise only those species
of the most biologically and morphologically uniform and
that parasitized bugs (Hemiptera), but recent discoveries
distinctive genera, comprising at least 40 species, in which
of a few bug parasitoids that share character states of other
the females accompany army ant raiding swarms to para-
sitize orthopteroid insects flushed out by them. For various subfamilies have shown that reliance on host type is errone-
species of Calodexia, Townsend proposed 10 generic names, ous. Males of many typical phasiines have an asymmetrical
and two more were subsequently added by Curran and Rein- distiphallus, but this is apparently not universal among all
hard. This tendency of splitting across the family has created genera. Their common lack of a uterus and consequent de-
such an impenetrable thicket of generic names that dipterists position of unembryonated eggs is itself a probable symple-
attempting to identify their specimens have met with disil- siomorphy. At the moment, the subfamily lacks supporting
lusionment and frustration. synapomorphies, although most genera are better supported.
1356
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Subfamily Dexiinae is perhaps the only subfamily based have been dissected possess a uterus. In males of many taxa
on a shared derived character state, namely the presence of a there is a strong connection between the fifth sternite and
hinge in the phallus accompanied by close association of the the fourth, which is weak and membranous in the other sub-
postgonites with the base of the phallus. Dexiinae possess a families. Presumably the most plesiomorphic taxa deposit
uterus and deposit embryonated eggs; presumably the most their embryonated eggs directly on the host, but those in
derived group, the tribe Dexiini deposit their eggs, not on the many, perhaps most, other tribes, particularly members of
host, but on a likely substrate where their characteristic elon- the tribe Tachinini, search out a potentially suitable sub-
gate, long-haired first instars can locate a host (see above). strate on which to deposit their eggs. These hatch imme-
Subfamily Exoristinae (Goniinae of Sabrosky & Arnaud, diately and the first instars simply wait for a passing host.
1965; Wood, 1987; and most other authors except O’Hara & Their integument is reinforced with platelets that, as the
Wood, 2004) includes both tribes that lack a uterus (Ethillini, larva desiccates, form a suit of armor to prevent further
Exoristini, Masiphyini, Neothelairini, Winthemiini, as well desiccation.
as several genera of Blondeliini and one of Eryciini, and
probably others not yet investigated) and those that possess
a uterus and deposit embryonated eggs. The New World
Identification
Neothelairini, abundant in the Neotropical Region, may be
the sister group of the Old World Ethillini. Blondeliini and No treatment exists for any genus of Tachinidae that is
Eryciini lacking a uterus are evidently not present in the aimed specifically at identification of the Central American
Neotropical Region, however. Either the uterus has evolved species, although some revisions of Nearctic or South Amer-
more than once, or the classification is in need of restructuring. ican faunas include the species of Central America known to
Other than common possession of setae on the prosternum their authors. It is a certainty that many undescribed species
(not found universally among all taxa), there appears to be remain to be collected. In addition, so many unknown spe-
no synapomorphy of the subfamily. However, the large cies have been discovered, belonging to nearly every genus,
cosmopolitan tribe Goniini is distinguished by oviposition during the last decade by the parataxonomists of the Instituto
of minute, limpetlike, “microtype” eggs that are deposited Nacional de Biodiversidad, Costa Rica, and by the rearing
on foliage and subsequently eaten by the host. efforts of D.H.  Janzen and W.  Hallwachs in Guanacaste,
Likewise, the diverse subfamily Tachininae is not based Costa Rica, that any existing key, including the following
on any synapomorphy, although females of all members that key to genera, will almost certainly be incomplete.

Key to the genera of Tachinidae of Central America

The following key is biased toward the Costa Rican fau- the Dexiini in couplets 167 to 181; the Voriini sensu stricto
na, both for the character states used and for the choice of in couplets 183 to 189; the Megaprosopini in couplets 196 to
genera included, because this fauna has been better docu- 198; and the bulk of the Phasiinae in couplets 208 to 222.
mented than that of other Central American countries. Un- Keys to Palearctic and Nearctic genera have tradition-
doubtedly genera not included here are to be found in other ally begun with separation on the basis of setose versus bare
parts of Central America, particularly in eastern Panama and prosternum. In the Neotropical fauna, there are so many ex-
in Guatemala. ceptions within genera and tribes that this choice has been
If the reader thinks that a specimen belongs to a particular relegated to couplet 102. No known character divides the
tribe, all of the Tachinini may be found in couplets 3 to 30, Tachinidae into two sizeable units in which exceptions do
most of the Goniini and Eryciini (intermingled) in couplets not occur, but the condition of the first postsutural supra-alar
31 to 33 and 41 to 87; the Winthemiini in couplets 37 to 40; bristle seems to be the most useful, separating about 30%,
the Polideini in couplets 88 to 97; the Nemoraeini in couplets including all members of the tribe Tachinini (subfamily Ta-
99 to 101; the Exoristini in couplet 113; the Blondeliini with chininae) as well as all members of Goniini and Eryciini
setose prosternum in couplets 115 to 119 and 125 to 165, (subfamily Exoristinae). Bristles are simply large thick se-
and those with bare prosternum at the end of the key after tae, while hairs are small, thin ones, but there is every grada-
couplet 265; the Siphonini in couplets 120 to 124; most of tion between the two extremes.

1. Three stout postsutural supra-alar bristles, the first at least as long and stout as first postsutural
dorsocentral bristle (Fig. 102) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Two to five postsutural supra-alar bristles, if only three then the first shorter and usually finer than
first postsutural dorsocentral bristle (Figs. 103, 104), or absent (in this case, the second postsutural
supra-alar, the largest in the series, will appear to be the first, as in Fig.  99, and only two
postsutural supra-alar bristles will be present). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1357

2. Hind coxa setose with group of small setae on posterolateral corner, these setae often extending
along entire coxal margin (Fig. 97) (hairs arising on first abdominal segment may, in profile,
appear to arise on coxa); first flagellomere bean-shaped or kidney-shaped (Figs. 2, 3, 88); prosternum
bare; inner vertical bristles crossed at midlength; TACHININI . . . . . . . . . . . . . . . . . . . . . . . . . . 3
– Hind margin of hind coxa bare, except for few genera in which one to three straight black hairs
are present near posteromedial corner of coxa (Fig. 6); first flagellomere parallel-sided in profile
(Fig.  7); prosternum haired (as in Figs.  90, 91); inner vertical bristles more or less parallel;
WINTHEMIINI, GONIINI, and ERYCIINI. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

3. Parafacial with one or two bristles in addition to smaller hairs; these bristles distinctly larger than
surrounding hairs and situated near lower margin of eye (Figs. 2, 3). . . . . . . . . . . . . . . . . . . . . . 4
– Parafacial with hairs only or bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

4. Palpus present, sometimes reduced to short bristle-tipped stub. . . . . . . . . . . . . . . . . . . . . . . . . . . 5

– Palpus absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

5. Palpus present, long and filiform, not enlarged apically (Fig.  2); parafacial bristles widely
separated from each other, about equal in size, or upper one slightly larger. . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Peleteria Robineau-Desvoidy
– Palpus reduced to bristle-tipped stub shorter than width of first flagellomere; parafacial bristles
arising close together, near lower end of parafacial, lower bristle larger of two. . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Itacuphocera Townsend

6. Parafacial with two stout bristles; these more or less equal in size, separated from each other by
about half distance between upper parafacial bristle and lowest frontal bristle (as in Fig. 2); frontal
bristles arising in two rows, lowest bristle at level of lower fourth of eye; abdominal tergites
mostly gray pruinose, with narrow apical bands of shiny cuticle . . . . . . . . . Deopalpus Townsend
– Parafacial usually with single stout bristle, sometimes with smaller bristle immediately below it
(Fig. 3); frontal bristles arising in single row, lowest bristle at level of middle of eye; abdominal
tergites with pruinosity only on basal half of each tergite, or less, or lacking pruinosity. . . . . . . 7

7. Thorax and abdomen slender; bluish-gray species . . . . . . . . . . . . . Copecrypta Townsend, in part

– Thorax and abdomen broad; yellow, reddish, or black species. . . . . . . . . . Trichophora Macquart

8. Palpus well developed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

– Palpus absent or reduced to minute tubercle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

9. Palpus longer than height of head, flattened from side to side, parallel-sided in profile, wider than
proboscis, extending forward far beyond lower facial margin (Fig. 83). . . . . . . . . . . . . . . . . . . 10
– Palpus shorter than height of head, usually distinctly clubbed apically, but if apparently parallel-sided
(as in Protodejeania), then narrower than proboscis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

10. Eye covered with conspicuous dense hairs . . . . . . . . . . . . . . . . . . . . . . . . . . Dejeaniops Townsend

– Eye apparently bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

11. Palpus fringed with long setae along entire length . . . . . . . . . . . . . . . . . Eulasiopalpus Townsend
– Palpus covered with short appressed setae that are much shorter than width of palpus, except
ventrally at base (Fig. 83). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

12. Body and legs orange. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adjeania Townsend

– Body black, legs orange or yellow. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eudejeania Townsend
1358
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

13. Eye covered with conspicuous dense hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

– Eye apparently bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

14. Abdominal tergites 3 and 4 each with only one pair of median discal bristles. . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Jurinia Robineau-Desvoidy
– Abdominal tergites 3 and 4 almost completely covered with discal bristles (as in Fig. 185). . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Jurinella Brauer & Bergenstamm

15. Abdominal tergite 3 with only one pair of median marginal bristles; abdominal tergites 2–4 bluish
pruinose, contrasting with non-pruinose tergite 5. . . . . . . . . . . . . . . . . . . . . . . Archytas Jaennicke
– Abdominal tergite 3 with more than one pair (usually three or more pairs) of median marginal
bristles (Figs. 192, 193); pruinosity of tergites, if present, uniformly distributed. . . . . . . . . . . . 16

16. Abdominal tergites  3 and 4 each with complete transverse row of marginal bristles displaced
anteriorly at midline enclosing small patch of bristles between it and posterior margin of tergite
(Fig. 192). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paradejeania Brauer & Bergenstamm
– Marginal bristles all more or less equidistant from posterior margin of tergite (Fig. 193). . . . . 17

17. Abdominal tergites 3 and 4 without median discal bristles; tergite 3 with three to five pairs of short
stout median marginal bristles; longest of these no more than twice as long as shortest (Fig. 193).
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Juriniopsis Townsend
– Abdominal tergites 3 and 4 each with several pairs of discal bristles forming median patch or
irregular transverse row; tergite 3 with single pair of median marginal bristles and several much
shorter ones less than one-third their length (as in Fig. 194). . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

18. Palpus clavate or spatulate, shorter than length of head at lower cranial margin, fringed with setae
that are shorter than height of palpus at widest point; abdomen dark brown or black . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eujuriniodes Townsend
– Palpus long and slender, longer than length of head at level of lower cranial margin and fringed
with long setae, especially along ventral side of palpus, many of them longer than height of palpus
at widest point; abdomen either yellow with black apex or entirely jet black. . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Protodejeania Townsend

19. Eye covered with conspicuous dense hairs; syncercus of male cone-shaped, truncate apically, and
flattened dorsally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cryptopalpus Rondani
– Eye apparently bare; syncercus of male, if cone-shaped, either pointed apically or with notched
apex, and rounded dorsally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

20. Both abdominal tergites 3 and 4 lacking median discal bristles . . . . . . . . . . . . . . . . . . . . . . . . . 21

– Either abdominal tergites 3 and/or 4 with median discal bristles . . . . . . . . . . . . . . . . . . . . . . . . 23

21. Head and postpronotum pale translucent yellowish; occipital bristles black, descending in nearly
straight row to lower cranial margin, contrasting with pale hair of gena. . Phosocephala Townsend
– Head dark in ground color, covered with uniformly yellowish or whitish pruinosity; occipital
bristles black, not descending below lower margin of eye, followed below by paler hairs. . . . . 22

22. Abdominal tergite 4 with closely set row of stout, spikelike marginal bristles extending across
tergite; male with yellow, black-tipped abdomen; tergite 5 black, entirely covered dorsally with
dense black, appressed hairs, with narrow pale area above terminalia; female with dark brown,
yellow-tipped abdomen; tergite 5 dark basally proximal to transverse row of bristles, yellowish
apically distal to this row of bristles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Euepalpus Townsend
– Abdominal tergite 4 with single median pair of marginal bristles of normal dimensions, separated
by wide gap from lateral marginal bristles; both sexes with polished bluish-black abdomen;
tergite 5 with small area of silvery pruinosity laterally. . . . . . . . . . . . . . . . . . . . Gymnomma Wulp
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1359

23. Abdominal tergite 3 with single pair of median marginal bristles; intersegmental lines between
abdominal tergites weakly impressed, in places scarcely discernible; abdominal sternites
narrow, overlapped by ventral edges of tergites, and supporting only one or two pairs of bristles
(Fig. 197). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
– Abdominal tergite  3 with transverse row of median marginal bristles; segmental lines deeply
impressed and readily discernible; abdominal sternites broad, not overlapped by ventral edges of
tergites, with three or more pairs of long stout bristles (Fig. 196). . . . . . . . . . . . . . . . . . . . . . . . 25

24. Parafacial clothed with erect hairs along entire length; parafrontal yellowish pruinose; abdominal
tergite 3 with median discal bristles; abdomen black, or orange with black markings. . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parepalpus Coquillett
– Parafacial with small patch of appressed dark hairs below, continuing dorsally along orbit;
parafrontal dark, shiny; abdominal tergite 3 lacking discal bristles; abdomen yellow with black
tergite 5. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xanthozona Townsend

25. Abdominal tergite 4 lacking median discal bristles; apex of abdomen notched (as in Figs. 192,
196) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
– Abdominal tergite 4 with patch of median discal bristles; apex of abdomen usually rounded or
pointed (as in Figs. 193, 194) (notched apically in Xanthoepalpus). . . . . . . . . . . . . . . . . . . . . . 27

26. Abdominal tergite 3 lacking median discal bristles, except occasionally one to three erect bristles
immediately in front of marginal row; syncercus of male heart-shaped, pointed apically, rounded
anterodorsally. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Quadratosoma Townsend
– Abdominal tergite 3 with median patch or transverse row of discal bristles, usually along anterior
edge of tergite; syncercus of male notched apically, with bilobed anterodorsal extension and oval
median area densely studded with short, clubbed hairs. . . . . . . . . . . . . Rhachoepalpus Townsend

27. Abdomen yellow with black apex; tergite 3 with transverse row of five or more pairs of median
marginal bristles, median ones arising slightly farther forward than lateral ones (Fig. 195),
longest bristles of marginal row near outer edge; tergite 3 with discal bristles forming transverse
row across anterior edge of tergite. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xanthoepalpus Townsend
– Abdomen red, brown, bluish-gray, or black; bristles of tergite 3 not arranged as above. . . . . . . 28

28. Subvibrissal bristles lacking, and prementum longer than height of head; abdomen black, covered
with bristles and scutum yellowish-brown; legs and base of wing orange–yellow. . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Austeniops Townsend
– Two or more pairs of subvibrissal bristles present, and prementum usually shorter than height of
head; abdomen, scutum, legs, and wing base as above in only few species of Epalpus. . . . . . . 29

29. Syncercus of male with notched apex; surstylus awl-shaped, its pointed apex directed medially;
abdomen red, orange, brown, bluish-gray, or black, often with median gold or silver spot on
tergite 5. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epalpus Rondani
– Syncercus of male with single pointed apex; surstylus U-shaped, its apex directed ventrally;
abdomen of Central American species either orange–red or dark brown without light-colored
median spot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

30. Median discal bristles of tergites 3 and 4 confined to median third of abdomen; abdomen orange–red
in Central American species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lindigepalpus Townsend
– Median discal bristles of tergite 4, and in some species of tergite 3 as well, distributed across the
tergite; abdomen dark brown to black. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anepalpus Townsend
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

87 Chaetona 88 Archytas 89 Ceracia

prst prst prst

fore cx
fore cx

90 Macromya 91 Microchaetina 92 Ormia

93 Erythromelana 94 Myiopharus 95 Eucelatoria

pmtcx brg

h cx h cx

96 Carcelia 97 Epalpus 98 Xanthomelanodes

Figs. 113.87–98. Antennae, prosterna, postpronota, and hind coxae: lateral view of left antenna of (87) Chaetona sp., (Wood, 1985, fig. 23);
(88) Archytas apicifer (Walker), (Nearctic, MND, fig. 110.130); and (89) Ceracia dentata (Coquillett), (Nearctic, MND, fig. 110.126); ventral
view of prosternum of (90) Macromya crocata Reinhard, (Nearctic, MND, fig. 110.153); (91) Microchaetina valida Townsend, (Nearctic,
MND, fig.  110.154); and (92)  Ormia dominicana Townsend, (Nearctic, MND, fig.  110.155); dorsal view of left postpronotum
of (93) Erythromelana sp.; (94) Myiopharus chrysocephalus (Bigot); and (95) Eucelatoria sp.; posterior view of hind coxae and base of
abdomen of (96) Carcelia reclinata (Aldrich & Webber), (Nearctic, MND, fig. 110.228); (97) Epalpus signifer (Walker), (Nearctic, MND,
fig. 110.229); and (98) Xanthomelanodes arcuatus (Say), (Nearctic, MND, fig. 110.230). Figures 93–95 illustrated by A. Brenes.
Abbreviations: fore cx, fore coxa; h cx, hind coxa; pmtcx brg, postmetacoxal bridge; prst, prosternum.
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1361

31. Ocellar bristles reclinate or lateroclinate (Figs. 8, 11). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32

– Ocellar bristles proclinate or absent (as in Figs. 5–7). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

32. Eye covered with conspicuous hairs; lower part of parafacial with patch of fine hairs (Fig. 8) in
some species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Distichona Wulp
– Eye apparently bare; parafacial with coarse hairs or bristles or both over most of length (Fig. 11).
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

33. Parafacial with even row of four or more bristles adjacent to ptilinal fissure, isolated from smaller
remaining parafacial setae (Fig. 11); second aristomere usually ten or more times as long as wide
and usually more than half as long as third aristomere; male with proclinate orbital bristles. . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gonia Meigen
– Parafacial with scattered setae; if those nearest ptilinal fissure appear larger, then they are distant
from fissure by at least half their length and do not form even row; second aristomere usually less
than 5 times as long as wide and usually less than half as long as third aristomere; male lacking
proclinate orbital bristle (Fig. 56) . . . . . . . . . . . . . . . . . . . . . . . Spallanzania Robineau-Desvoidy

34. Postpronotum with middle basal bristle displaced anterolaterally forming nearly right-angled
triangle with outer and inner basal bristles (as in Fig. 94); middle basal bristle also arising lateral
to line between outer basal and outer anterior bristles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
– Postpronotum with middle basal bristle more or less in line with outer and inner basal bristles
(Fig. 95), or inner basal absent (Fig. 93); if middle basal apparently forms angle with outer and
inner basals, this angle at least 160°; middle basal usually medial to line between outer basal and
outer anterior (latter not always present). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50

35. Eye covered with ommatrichia (conspicuous dense hairs, each hair longer than combined
diameter of four or more eye facets) (Figs. 5–8). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
– Eye apparently bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45

36. Katepimeron haired, at least on its anterior two-thirds (Figs. 105, 106); WINTHEMIINI. . . . . 37
– Katepimeron bare, or with at most three to four hairs near anterior end; GONIINI, ERYCIINI. . 41

37. Parafacial with hairs on upper half or more (Fig. 5) . . . . . . . . . . . Winthemia Robineau-Desvoidy
– Parafacial bare on lower two-thirds or more (as in Figs. 6, 7). . . . . . . . . . . . . . . . . . . . . . . . . . . 38

38. Facial ridge with bristles on at least lower third or more (as in Figs. 8, 9). . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemisturmia Townsend
– Facial ridge bare on all but lowest fifth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39

39. Scutum with median longitudinal stripe pale, flanked by two pairs of dark lateral stripes. . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Avibrissosturmia Townsend
– Scutum with median stripe dark, flanked by one dark lateral stripe on each side (suggesting a
sarcophagid fly). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40

40. Abdominal tergites 3 and 4 each with one pair of median discal bristles (as in Fig. 186); usually
small flies. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nemorilla Rondani
– Abdominal tergites 3 and 4 without median discal bristles; usually medium-sized flies . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Triodontopyga Townsend

41. Facial ridge with row of long erect bristles on lowest two-thirds or more (as in Fig. 15). . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Patelloa Townsend
– Facial ridge at most with row of hairlike decumbent setae on lower half or less . . . . . . . . . . . . 42
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

42. Gena reduced to narrow strip about one-twelfth height of head (Fig. 7). . . . . . . . . . . . . . . . . . . 43
– Gena higher, at least one-tenth height of head (as in Figs. 9, 10) . . . . . . . . . . . . . . . . . . . . . . . . 44

43. Hind coxa with one to three small black hairs on posteromedial margin (Fig. 96). . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carcelia Robineau-Desvoidy, in part
– Hind coxa bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Houghia Coquillett, in part

44. Pruinosity of frons and most of thorax and abdomen, in life, vivid metallic green or gold, fading
soon after death to yellowish-brown color that may be temporarily restored to its metallic luster
by application of alcohol or petroleum solvent; abdomen with transverse distal black bands; gena
exceptionally high, distance between eye and lower cranial margin more than one-ninth height of
head (as in Figs. 9, 10). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chrysoexorista Townsend
– Pruinosity of body mostly or entirely gray, without metallic green color; gena narrower, less than
one-fifth height of head . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hyphantrophaga Townsend

45. Parafacial with single stout bristle, sometimes with smaller bristle immediately below it, on lower
part of parafacial (Fig. 2) (aberrant Tachinini with bare hind coxa) . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Copecrypta Townsend, in part
– Parafacial bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46

46. Lower facial margin sloping evenly forward from lower part of face, thus visible in profile below
and in front of vibrissal angle (Fig. 84); palpus reduced or absent in several species. . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Masiphya Brauer & Bergenstamm
– Lower facial margin, if visible in profile, bent forward abruptly at level of vibrissal angle (as in
Fig. 12); palpus well developed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47

47. Facial ridge with row of stout erect bristles along most of length (as in Figs. 12, 15) . . . . . . . . 48
– Facial ridge bare, with at most few small decumbent setae above vibrissa. . . . . . . . . . . . . . . 49

48. Gena reduced to narrow strip about one-tenth height of head; postpronotum with only three
bristles; body and abdomen dull brown, lacking tessellated pattern; abdomen broad, flattened
dorsally. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Casahuiria Townsend
– Gena higher, about one-fifth height of head; postpronotum with small bristle medial to outer basal
bristle; body tessellated gray pruinose; abdomen slender and tapering, usually with reddish apex.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prospherysa Wulp

49. Hind coxa with one to three small black hairs on posteromedial margin (Fig. 96). . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carcelia Robineau-Desvoidy, in part
– Hind coxa bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Houghia Coquillett, in part

50. Eye with ommatrichia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51

– Eye bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67

51. Facial ridge bristled on more than lower half . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52

– Facial ridge setose on lower third or less. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59

52. Parafacial setose along its entire length, from lowest frontal bristle to lower margin, setae usually
more bristlelike medially (Fig. 9). . . . . . . . . . . . . . . . . . . . . . Gaediopsis Brauer & Bergenstamm
– Parafacial bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1363

53. Distance between eye and subcranial margin, in profile, less than one-tenth height of head (as
in Figs. 7, 14); frons with two or more large reclinate orbital bristles arising distinctly lateral to
frontal row, with anteriormost arising near midlength of frons (Figs. 59–61); ocellar setae minute
or absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudochaeta Coquillett, in part
– Distance between eye and subcranial margin more than one-sixth height of head; reclinate upper
orbital bristles confined to posterior third of frons, usually arising behind uppermost frontal;
ocellar bristles well developed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54

54. Arista thickened nearly to its abruptly tapered apex; frons in dorsal view wider than eye when
viewed from above in both sexes; three katepisternal bristles confined to area behind pleural
suture . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Madremyia Townsend
– Arista thickened on basal half or less; frons in dorsal view narrower than eye when viewed from
above, or if apparently as wide, then four katepisternal bristles present; anterior katepisternal
bristle arising in front of pleural suture. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55

55. Apex of scutellum with small pair of crossed apical bristles arising between much larger subapical
bristles and at same level (as in Fig. 135) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
– Apex of scutellum lacking apical bristles (Figs. 125, 126). . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58

56. Katepisternum with four bristles (Fig. 116). . . . . . . . . . . . . . Lespesia Robineau-Desvoidy, in part

– Katepisternum with three bristles, two anterior and single posterior bristle (as in Fig. 115) . . . 57

57. Wing darkened on anterior half (Fig. 148); body metallic green and gold pruinose in life, fading
to gold or brownish-yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myothyriopsis Townsend, in part
– Wing clear; body black and gray, lacking metallic green or gold pruinosity. . . Acantholespesia Wood

58. Discal scutellar bristles arising closer to one another than subapical bristles (Fig. 125). . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyzenis Robineau-Desvoidy
– Discal scutellar bristles more widely separated than subapical bristles (Fig. 126). . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Frontiniella Townsend

59. Gena reduced to narrow strip about one-twelfth height of head (Fig. 7); hind coxa with one to
three small black hairs on posteromedial margin (Fig. 96). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carcelia Robineau-Desvoidy, in part
– Gena higher, at least one-tenth height of head; hind coxa bare. . . . . . . . . . . . . . . . . . . . . . . . . . 60

60. Vibrissa arising far above lower facial margin (Fig. 77), at least half distance between vibrissal
bases, and nearly twice length of pedicel; ocellar bristles absent; katepisternum with two bristles;
abdominal tergites 1–4 of male lacking median bristles; body reddish-brown, whitish pruinose,
without any trace of green or gold coloring. . . . . . . . . . . . . . . . . . . . . . . . Mystacomyia Giglio-Tos
– Vibrissa arising at most only slightly above lower facial margin, much less than half distance
between vibrissae and less than length of pedicel; ocellar bristles well developed; katepisternum
usually with three bristles; abdominal tergites with median bristles; body either yellowish or gold
(green in life), or black with grayish pruinosity. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61

61. Parafacial with few inconspicuous hairs or setae below lowest frontal bristle (Fig. 10). . . . . . . 62
– Parafacial bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64

62. Parafacial exceptionally wide (Fig.  10); vibrissa arising above lower facial margin; scutum or
abdomen in life usually metallic green or blue, fading to yellow or brown (as in Chrysoexorista).
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mystacella Wulp, in part
– Parafacial narrower, about as wide as width of first flagellomere; body black in ground color with
grayish pruinosity. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
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99 Ischyrophaga 100 Chrysotachina

first psut dc s
first psut ial s first psut dc s

trn sut psut spal s

npl s first psut spal s

101 Xanthophyto 102 Atacta

first psut dc s
first psut dc s

first psut spal s

first psut spal s

103 Chetogena 104 Lixophaga

Figs. 113.99–104. Thoracic chaetotaxy: lateral view of left side of postsutural scutum of (99) Ischyrophaga sp.; (100) Chrysotachina sp.;
(101)  Xanthophyto labis (Coquillett), (Nearctic, MND, fig.  110.162); (102)  Atacta  sp., (MND, fig.  110.163, as Atactopsis);
(103) Chetogena sp., (MND, fig. 164, as Exorista); and (104) Lixophaga parva Townsend, (Nearctic, MND, fig. 110.165). Figures 99 and
100 illustrated by A. Brenes.
Abbreviations: npl s, notopleural setae; psut dc s, postsutural dorsocentral seta; psut ial s, postsutural intra-alar seta; psut spal s, postsutural
supra-alar seta; trn sut, transverse suture.
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1365

63. Katepisternum with three bristles (as in Fig. 116); palpus pale; western Nearctic Region, southern
South America. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . [Eunemorilla Townsend]
– Katepisternum with three bristles (as in Fig. 115); palpus black. . . . . Eumea Robineau-Desvoidy

64. Integument of body and legs mostly yellow, pruinosity of body metallic green in life fading
to yellow; prosternum with pale hairs; subvibrissal area with only two bristles; gena with single
black bristle; transverse pruinose band on abdominal tergite 3 with pair of V-shaped notches . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eulobomyia Woodley & Arnaud
– Integument of body and legs black with gray, or brown and gold pruinosity; prosternum with
black hairs; subvibrissal area with three or more bristles; transverse pruinose band on abdominal
tergite 3 uniform in width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65

65. Hind tibia with comblike row of closely spaced anterodorsal setae of uniform length in male (as
in Fig. 172) (female unknown but probably with single longer bristle (as in Fig. 173); abdominal
tergite 4 of male with large dense ventral patch of appressed black hair (as in Fig. 164). . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Zygozenillia Townsend
– Hind tibia lacking row of closely spaced setae; abdominal tergite 4 with dense black hair laterally
or not at all. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66

66. Abdominal tergites 4 and 5 with basal gray pruinose bands and shiny black apical areas laterally,
with small patch of denser black hair laterally on tergite 4 in male. . . . . . Aplomyopsis Townsend
– Abdominal tergites with mostly gold–green pruinose in life, fading to golden brown. . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mystacella Wulp, in part

67. Facial ridge with row of stout erect bristles on more than its lower half (as in Figs. 12, 15). . . 68
– Facial ridge bare on all but lowest third (as in Figs. 13, 14). . . . . . . . . . . . . . . . . . . . . . . . . . . . 76

68. Parafacial entirely covered with setae (as in Figs. 11, 12). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69

Parafacial bare below lowest frontal bristles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70

69. Largest setae on parafacial forming even vertical row of at least ten small bristles continuous with
lowest frontal bristles, flanked by smaller hairs. . . . . . . . . . . . . . . . . . . . Choeteprosopa Macquart
– Parafacial setae mostly few and small, if bristlelike, then confined to lower part of parafacial, far
removed from lowest frontal bristles, which diverge laterally toward eye margin (Fig. 12). . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetogaedia Brauer & Bergenstamm

70. Distance between eye and subcranial margin, in profile, less than one-tenth height of head (as
in Figs. 7, 14); frons with two or more large reclinate orbital bristles arising distinctly lateral to
frontal row, anteriormost arising near midlength of frons (Figs. 59–61); ocellar setae minute or
absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudochaeta Coquillett, in part
– Distance between eye and subcranial margin more than one-sixth height of head; reclinate upper
orbital bristles confined to posterior third of frons, usually arising behind uppermost frontal seta;
ocellar bristles well developed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
71. Anterodorsal setae of hind tibia in both sexes forming even, closely spaced row (Figs. 172, 173),
each bristle separated from its neighbors by width of one bristle or less, at most, only one bristle
slightly longer than others. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72
– Anterodorsal setae of hind tibia irregularly arranged and more widely separated from one another,
at least one bristle distinctly longer than the others (Fig. 174) . . . . . . . . . . . . . . . . . . . . . . . . . . 75
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72. Area of facial ridge, lateral to vibrissa and row of facial bristles narrow, at most with single row
of small setae (Fig. 78); facial ridge fairly straight in frontal aspect. . . . . . . . . . . . . . . . . . . . . . 73
– Area of facial ridge, lateral to vibrissa and facial bristles, with more extensive patch of small
setae, usually in two or three irregular rows (Fig. 79); facial ridge strongly bowed outward above
vibrissa. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74

73. Katepisternum with three bristles; sexual hair patch on tergite 4 of male, if present, extending
from ventral surface around side of abdomen to dorsal surface; abdomen black or dark reddish,
with at most sparse brownish pruinosity; gravid female with microtype eggs . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leschenaultia Robineau-Desvoidy
– Katepisternum with four bristles; tergite 4 of male lacking sexual hair patch; abdomen of most
species with basal pruinose bands, often entirely gray or yellowish pruinose, rarely all black;
gravid female with transparent, fully embryonated eggs. . . . Lespesia Robineau-Desvoidy, in part

74. Abdomen globose or cylindrical in cross section, as wide as high; tergite 4 of male ventrolaterally
with patch of short dense appressed black hairs (Fig. 164). . . . . . . . . . . . . . . Blepharipa Rondani
– Abdomen dorsoventrally flattened, wider than high; tergite 4 of male lacking patch of appressed
hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Atactosturmia Townsend, in part

75. Medium to large robust species, 10 mm or more in length; facial ridge wide and flat, its bristles
separated from ptilinal suture by at least 3 times diameter of their basal sockets; vibrissa arising
above lower facial margin; in most species, abdominal tergites 4 and 5, or 5 alone, yellow, gold,
or orange pruinose, contrasting with remaining black abdomen; ocellar setae minute or absent. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Belvosia Robineau-Desvoidy
– Medium to small species, less than 9 mm in body length; facial ridge narrow, its bristles close to
ptilinal suture; vibrissa arising level with lower facial margin; abdomen unicolorous except for
few species in which last tergite contrastingly paler; ocellar setae well developed . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lespesia Robineau-Desvoidy, in part

76. Bifurcation of R2+3 and R4+5 with single long seta on dorsal surface (Fig. 156). . . . . . . . . . . . . . 77
– Bifurcation of R2+3 and R4+5 with two or more small setae on dorsal surface . . . . . . . . . . . . . . . 79

77. Katepisternum with three bristles; fronto-orbital plate with single row of frontal bristles;
anterior reclinate orbital bristle larger than posterior one, bristles well separated from each other
(Fig. 13). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ametadoria Townsend
– Katepisternum with four bristles; fronto-orbital plate with extra medioclinate bristles outside
frontal row, usually forming second row (Figs. 62, 63); two or three reclinate orbital bristles usually
present but if only two, then anterior one usually smaller than posterior one (Fig. 63) . . . . . . . 78

78. Ocellar bristles long, divergent (Fig. 62). . . . . . . . . . . . . . . . . . . . . . . Lydella Robineau-Desvoidy

– Ocellar bristles short, curving toward each other apically (Fig. 63), or absent. . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Drino Robineau-Desvoidy

79. Gena reduced to narrow strip about one-twelfth height of head (Figs. 7, 14). . . . . . . . . . . . . . . 80
– Gena higher, at least one-tenth height of head (as in Figs. 9, 10) . . . . . . . . . . . . . . . . . . . . . . . . 82

80. Reclinate orbital bristles arising distinctly outside frontal row (as in Figs. 59–61); ocellar setae
minute or absent; frons with bulging aspect in profile. . . . . Argyrophylax Brauer & Bergenstamm
– Reclinate orbital bristles arising behind and more or less continuous with frontal row (as in
Figs. 62, 63); ocellar setae usually well developed; frons not protruding in profile. . . . . . . . . . 81
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1367

81. Hind coxa with one to three black hairs on posteromedial area of hind margin (Fig. 96); abdominal
tergites with transverse pruinose bands with evenly curved posterior margins, or abdomen entirely
pruinose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carcelia Robineau-Desvoidy, in part
– Hind coxa bare; transverse pruinose bands on abdominal tergite  4 with undulating posterior
margin (Fig. 183). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Argyrochaetona Townsend

82. Vibrissa arising above lower facial margin, protruding below and in front of vibrissal angle when
viewed in profile. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83
– Vibrissa arising at level of lower facial margin, either protruding at this level or not visible in
lateral view. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85

83. Notopleuron with one or two smaller bristles in addition to the usual two large ones (Fig. 102);
parafrontal of female with three to six small proclinate orbital bristles in row (Fig. 57); frons of
male inflated, interocular distance at level of antennae of male at least 4 times width of vertex. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Atacta Schiner
– Notopleuron with two large bristles only (as in Figs. 101, 103, 104); parafrontal of female with
two pairs of proclinate orbital bristles; frons of male not unusually bulging or inflated, interocular
distance at level of antennae less than 3 times width of vertex. . . . . . . . . . . . . . . . . . . . . . . . . . 84

84. Facial ridge strongly bowed laterally above vibrissa, area of facial ridge lateral to vibrissa broad
with extensive patch of small setae, forming two to three irregular rows (as in Fig. 79); gravid
female with microtype eggs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Atactosturmia Townsend, in part
– Facial ridge fairly straight in frontal aspect, area of facial ridge lateral to vibrissa narrow, with at
most single row of small setae (as in Fig. 78); gravid female with transparent fully embryonated
eggs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Siphosturmia Coquillett, in part

85. Lower facial margin protruding in front of vibrissal angle, visible in profile (as in Fig. 12); vein R1
setose dorsally; male with two pairs of proclinate orbital bristles. . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Siphosturmia Coquillett, in part
– Lower facial margin not visible in profile; vein R1 bare; male lacking proclinate orbital bristles .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86

86. Wing darkened on anterior half (Fig. 148); body metallic green and gold pruinose in life, fading
to gold or brownish-yellow; hind coxa with one to three black hairs on posteromedial area of hind
margin (as in Fig. 96). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myothyriopsis Townsend, in part
– Wing transparent; body black and gray or silvery, not changing after death; hind coxa bare on hind
margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87

87. Male with abdominal tergites 4 and 5 each with extensive sexual hair patch, extending from entire
ventral surface of tergite up onto lateral margin of dorsal surface; apex of female palpus mostly
covered with appressed setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Zizyphomyia Townsend
– Male without well-defined sexual hair patches, although tergite 4 may have slightly more dense
hair ventrally; apex of female palpus swollen, virtually bare . . . . . . . . . . Sturmiomima Townsend

88. With following two characters present simultaneously: metathoracic spiracle with almost equal
outgrowths of hairs (lappets) along each side of opening, with V-shaped opening between them
(Fig.  105), and anepimeral bristle exceptionally large, extending posteriorly beyond middle of
lower calypter (Figs. 110, 111); syntergosternite 7+8 of male characteristically enlarged, bulbous
and shiny, wider than hypandrium; POLIDEINI. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
– Metathoracic spiracle covered with nearly circular flap arising from posterior edge, separated
from hairs along anterior edge by crescent-shaped opening (Fig. 106), or if both lappets appear
more nearly equal in size, leaving U- or V-shaped central opening, then anepimeron at most with
short bristle that does not extend back beyond apex of upper calypter; syntergosternite  7+8
usually less prominent (except in Linnaemya and Panzeria), and usually as pruinose as adjacent
segments. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98
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89. Dorsal surface of lower calypter covered with long erect hairs (Fig. 110). . . . Hystricia Macquart
– Dorsal surface of lower calypter bare (Fig. 111). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90

90. Parafacial setose along its entire length (as in Fig. 4). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91

– Parafacial bare or at most with few hairs below lowest frontal bristle . . . . . . . . . . . . . . . . . . . . 93

91. Gena uniformly covered with appressed hairs; frons with purplish metallic reflections. . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudobombyliomyia Townsend
– Gena with three or more stout bristles arising from midregion (Fig. 4); frons gray, without metallic
reflections. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92

92. Scutellum densely covered with at least 50 erect bristles; syncercus of male of most species with
bifurcate or trifurcate flaglike process arising from dorsal surface. . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bombyliomyia Brauer & Bergenstamm
– Scutellum with at most four to five pairs of weak bristles among long hairs; syncercus of male
without dorsal process. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phobetromyia Reinhard

93. Abdominal tergite 4 with transverse row of discal bristles (Fig. 184). . . . . . . . . . . . . . . . . . . . . 94

– Abdominal tergite 4 with single pair of median discal bristles. . . . . . . . . . . . . . . . . . . . . . . . . . 95

94. Body black with metallic greenish or purplish reflections. . . . . . . . . . . Chlorohystricia Townsend
– Body without metallic reflections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eucheirophaga James

95. Middle katepisternal bristle located closer to posterior bristle than to anterior bristle, and both
middle and posterior bristles lower than anterior bristle, below level of katepimeron (Fig. 121);
scutellum with two pairs of discal bristles, one ahead of the other (Fig. 144); abdominal tergite 3
with two pairs of discal bristles, one larger pair in front of smaller pair (as in Fig. 188). . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lypha Robineau-Desvoidy
– Middle katepisternal bristle, when present, closer to anterior bristle than to posterior bristle (as in
Fig. 122), or lacking; scutellum with single pair of discal bristles, or if two pairs, then abdominal
tergite 3 with single pair of discal bristles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96

96. Vein M ending in R4+5 before reaching wing margin (as in Fig. 154). . . . . . Mauromyia Coquillett
Vein M ending in wing margin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97

97. Katepisternum with three bristles, middle bristle closer to anterior bristle than to posterior bristle
but distinctly behind it (Fig. 122); body often green metallic, otherwise bluish or purplish with
white pruinosity, or gray tessellated. . . . . . . . . . . . . . . . . . Chrysotachina Brauer & Bergenstamm
– Katepisternum with two bristles, occasionally with hairlike third located directly below anterior
bristle; body not metallic, brown or gray tessellated. . . . . . . . . . . . . . . . . . Exoristoides Coquillett

98. Four or five postsutural supra-alar bristles, first arising close to suture, separated from suture by
no more than twice its basal diameter, followed by small bristle more medially situated (lacking
in some Xanthophyto), followed by major supra-alar bristle, followed by one small and lastly one
large bristle; first postsutural intra-alar bristle almost as close to suture as first postsutural supra-alar,
separated from suture by no more than 4 times its basal diameter (Fig. 101); upper surface of
lower calypter with soft recumbent hairs on narrow strip along lateral margin and sometimes also
on posterolateral corner (except in Xanthophyto); segments 6–8 of male longer than epandrium in
most species; NEMORAEINI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
– Three, or rarely two, pairs of postsutural supra-alar bristles, first usually farther away from suture
than 2 times its own basal diameter (as in Figs. 99, 102) (in Chetogena a small fourth bristle may
be present between last and second last bristles, Fig. 103); first postsutural intra-alar farther away
from suture than 4 times its own basal diameter; lower calypter bare except at margin; segments 6–8
of male usually smaller than epandrium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1369

99. Abdominal tergites 3 and 4 each with transverse row of median marginal bristles and row or patch
of numerous median discal bristles; epandrium of male at least as wide as segments 6–8. . . . 100
– Abdominal tergites 3 and 4 at most with single pair of well-developed median marginal and discal
bristles; epandrium narrower than segments 6–8. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101

100. Large species, usually over 10 mm in length, with broad flattened orange, brown, black, or yellow
and black abdomen; abdominal tergite 3 with triangular median area of erect bristles; abdominal
tergite 4 with single transverse row of marginal bristles and irregular transverse row or patch of
discal bristles; facial ridge bare; gena with one or more isolated bristles; medial surface of
forecoxa entirely covered with appressed setae (Fig. 90); male syncercus lacking dorsal processes.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macromya Robineau-Desvoidy
– Medium-sized species at most 10  mm in length, with head, body, and abdomen blue–gray
pruinose; tergites 3–5 each with wide transverse band of dense erect marginal bristles on posterior
half of tergite, continuous medially with triangular patch of erect discal bristles; anterior half of
tergite pruinose and devoid of bristles; facial ridge with row of fine erect dark hairs; genal dilation
with small recumbent hairs; medial surface of forecoxa setose only on anterior half (as in Figs. 91,
92); male syncercus with pair of dorsally directed setose processes. . . . . . . . . . . . . Lasiona Wulp

101. Medial surface of forecoxa entirely covered with appressed setae (as in Fig. 90); facial ridge with
erect bristles on lower half or more, well above lower level of eye; wing transparent; bend of
vein M with stub as long as crossvein r-m; two katepisternal bristles. . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hypotachina Brauer & Bergenstamm
– Medial surface of forecoxa setose only on anterior half (as in Figs. 91, 92); facial ridge setose only
up to lower level of eye; wing of most species with prominent dark area around crossvein r-m;
bend of M at most with short stub, usually only with dark shadow; three katepisternal bristles . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xanthophyto Townsend

102. Prosternum setose (as in Figs. 90, 91) (if setation reduced to a single seta or pair of setae, try also
second half of this couplet) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
– Prosternum bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166

103. Longest bristle of scutellum situated closer to basal bristle than to apex of scutellum (Fig. 129) .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104
– Longest bristle of scutellum situated closer to apex of scutellum than to basal bristle
(Figs. 130–132, 140) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108

104. Inner margin of parafacial with vertical row of long stout erect proclinate bristles (Fig. 16), sometimes
arising so close to ptilinal suture that, in profile, bristles could be interpreted as being on facial
ridge. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paradidyma Brauer & Bergenstamm, in part
– Parafacial bare or with single bristle near its upper extremity below lowest frontal bristle. . . 105

105. Eye covered with long conspicuous ommatrichia, each one longer than combined diameter of four
or more eye facets; with three katepisternal bristles; apex of first flagellomere evenly rounded . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
– Eye bare or with sparse minute ommatrichia; with two katepisternal bristles; apex of first flagellomere
mucronate (with sharp point at apex) (Fig. 89). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107

106. Veins R1, R4+5, and CuA1 setose dorsally for most of their length; ocellar bristles arising in front of
anterior ocellus; upper part of parafacial with single reclinate bristle separated from lowest frontal
by twice distance between two lowest frontals . . . . . . . . . . . . . . . Minthoplagia Townsend, in part
– All wing veins bare, except for base of R4+5 up to crossvein r-m, and in few species, R1; ocellar
bristles arising behind anterior ocellus; parafacial without setae. . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neomintho Brauer & Bergenstamm
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

kepm p spr
p spr
kepm

105 Chrysotachina 106 Winthemia

prepst

prepm s

107 Billaea 108 Neaera 109 Telothyria

anepm s

l calyp anepm s

110 Hystricia 111 Lypha 112 Leucostoma

Figs. 113.105–112. Thoracic structures: lateral view of katepimeron and posterior spiracle of (105) Chrysotachina alcedo (Loew), (Nearctic,
MND, fig. 110.169); and (106) Winthemia sp., (Nearctic, MND, fig. 110.170); lateral views of anterior portion of thorax of (107) Billaea sp.;
(108) Neaera leucoptera (Johnson), (Nearctic, MND, fig. 110.167); and (109) Telothyria sp., (Nearctic, MND, fig. 110.168, as Leskiopsis);
dorsal views of left calypter of (110) Hystricia abrupta (Wiedemann), (Nearctic, MND, fig. 110.215); (111) Lypha setifacies (Brooks), (Nearctic,
MND, fig. 110.218); and (112) Leucostoma simplex (Fallén), (Holarctic, MND, fig. 110.217). Figure 107 illustrated by A. Brenes.
Abbreviations: anepm s, anepimeral seta; kepm, katepimeron; l calyp, lower calypter; p spr, posterior spiracle; prepm s, proepimeral setae;
prepst, proepisternum.
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1371

a kepst s
p kepst s
m kepst s a kepst s

pv kepst s
av kepst s

113 Phasia 114 Anoxynops 115 Myiopharus 116 Lespesia

117 Actia 118 Pseudosiphona 119 Trafoia 120 Thelairaporia

121 Lypha 122 Chrysotachina 123 Thelairodes 124 Xanthodexia

Figs. 113.113–124. Katepisterna: lateral view of (113) Phasia diversa (Coquillett), (Nearctic, MND, fig. 110.171); (114) Anoxynops aldrichi
(Curran), (Nearctic, MND, fig. 110.172); (115) Myiopharus dorsalis (Coquillett), (Nearctic, MND, fig. 110.173); (116) Lespesia archippivora
(Riley), (Nearctic, MND, fig. 110.177); (117) Actia sp.; (118) Pseudosiphona sp.; (119) Trafoia sp.; (120) Thelairaporia sp.; (121) Lypha sp.;
(122) Chrysotachina sp.; (123) Thelairodes sp.; and (124) Xanthodexia sp. Figures 117–124 illustrated by A. Brenes.
Abbreviations: a kepst s, anterior katepisternal seta; av kepst s, anteroventral katepisternal seta; m kepst s, middle katepisternal seta; p kepst s,
posterior katepisternal seta; pv kepst s, posteroventral katepisternal seta.
1372
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

107. Genal and subvibrissal bristles distinctly differentiated, anteriormost genal bristles arising
distinctly lateral to lowermost subvibrissal bristle and curving more strongly forward and outward.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceracia Rondani
– Genal and subvibrissal bristles in more or less continuous row, anteriormost genal bristles arising
behind subvibrissal bristles and appearing continuous with them. . . Acemya Robineau-Desvoidy

108. Ventral proepimeral bristle directed anteroventrally (as in Fig. 108), at least half as long as dorsal
proepimeral bristle; base of R4+5 with single large bristle (rarely with smaller additional bristle);
and vein M ending at apex of wing (Fig. 159); katepimeron usually with single seta . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phytomyptera Rondani
– Ventral proepimeral bristle directed anterodorsally, parallel to dorsal proepimeral bristle (except
in Neaera and some Chaetostigmoptera, in which ventral bristle is less than one-third as long
as dorsal bristle); vein M ending before apex of wing (except in all Chaetostigmoptera); if both
of these character states present then base of R4+5 with several short bristles (Fig.  158) and
katepimeron bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109

109. At most only one frontal bristle arising below upper end of pedicel (as in Fig. 47); pedicel with
one to two long, slender, curved setae more than half as long as length of plumose arista; male
with all frontal bristles proclinate (Fig. 47); lateral scutellar bristles lacking; DEXIINI . . . . . 110
– Frontal bristles usually descending below upper end of pedicel (as in Figs. 15, 20–28), but if not,
then either pedicel without such long bristles or arista bare; male usually with some reclinate
frontal bristles; lateral scutellar bristles present (Fig. 135). . . . . . . . . . . . . . . . . . . . . . . . . . . . 112

110. M terminating in R4+5 well before wing margin, with long extension beyond bend (Fig. 163); arista
short plumose, with bulbous base; lower half of parafacial with four to five proclinate bristles;
vibrissa arising well above lower facial margin, with four to five subvibrissal bristles . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microchaetina Wulp, in part
– Vein M ending in wing margin; arista long plumose, tapering evenly from base; parafacial bare;
vibrissa arising at level of lower facial margin, with only one to two subvibrissal bristles . . . 111

111. Basal scutellar bristles subparallel, standing out from sides of scutellum; lower calypters dark
brown, widely separated medially; scutum entirely dark brown. . . . . . . Ebenia Macquart, in part
– Basal scutellar bristles convergent, partially overlying edges of scutellum (Fig. 138); lower
calypters yellow, those of male enlarged and closely approximated medially; scutum yellow in
front of transverse suture, black behind it. . . . . . . . . . . . . . . Pseudodexia Brauer & Bergenstamm

112. Bend of M forming angle of 90º or less; portion beyond bend sinuous, curving toward apex
(Fig. 157); basal scutellar bristle nearly as long as subapical bristle (Fig. 135); extra smaller bristle
usually present between second and third postsutural supra-alar bristles (Fig. 103); cerci of male fused
into syncercus with pointed apex; surstyli reduced or vestigial; female vagina short, not modified into
ovisac for retention and embryonation of eggs, containing only single large planoconvex egg that
usually does not contain fully formed first instar larva; EXORISTINI. . . . . . . . . . . . . . . . . . . 113
– Bend of M obtuse; portion beyond bend straight or convex (Figs. 158, 160, 161); basal scutellar
bristle no more than two-thirds as long as subapical bristle (Figs. 130–132, 140); no extra small
bristle between second and third postsutural supra-alar bristles; cerci of male free at their apices,
fused only at base; surstyli elongate; female vagina, in most genera, elongated into ovisac for
retention of eggs; those eggs toward apex of abdomen usually containing fully developed first instar
larvae; BLONDELIINI, in part, plus one dexiine genus (Eulasiona), and SIPHONINI. . . . . . . 114
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1373

113. Lower margin of face protruding below vibrissal angle (scarcely evident in one species), visible
in profile (Fig. 15), and apical scutellar bristles inclined at about 40º to the horizontal (less,
perhaps 30º in another species, however, that has a prominent lower facial margin); wing
membrane crease along anterior edge of vein M extending well beyond bend of M, longer than
length of crossvein r-m, appearing from above as stub or continuation of M (Fig. 157) . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chetogena Rondani
– Lower margin of face not protruding, not visible in profile; apical scutellar bristles nearly level
with subapical scutellars in profile, not raised more than 10º above the horizontal; wing membrane
crease anterior to vein M extending only slightly beyond bend of M, appearing from above as
smudge or rarely minute stub, shorter than crossvein r-m. . . . . . . . . . Austrophorocera Townsend

114. Vein R4+5 setose on dorsal surface halfway or more from its base at junction of R2+3 and R4+5 to
crossvein r-m (Figs. 158, 160, 161). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115
– Vein R4+5 dorsally with few setae at base only, not extending halfway to crossvein r-m. . . . . . 129

115. Eye with conspicuous ommatrichia, each longer than combined diameter of four or more eye
facets (as in Fig. 20). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116
– Eye apparently bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120

116. Facial ridge bristled on lower half or more, with row of erect bristles along most of length
(Figs. 21–24) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
– Facial ridge bare except for few small recumbent bristles above vibrissa. . . . . . . . . . . . . . . . . 118

117. Lateral scutellar bristles divergent, about four-fifths as long as subapical bristles (as in Fig. 127);
basal scutellar bristles convergent (as in Fig.  125), longer than subapical bristles; frons wide,
nearly one-third as wide as width of head; posteriormost reclinate orbital bristle as large as or
larger than ocellar bristle; predominant color of pruinosity of head and body brown. . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodoria Townsend
– Lateral and basal scutellar bristles at most two-thirds as long as subapical bristles, parallel to each
other or convergent; frons narrow, less than one-fifth width of head in female and narrower in
male; posteriormost reclinate orbital bristle much smaller than ocellar bristle; predominant color
of pruinosity of head and body silvery gray. . . . . . . . . . . . . . . . . . . .Thelairodoriopsis Thompson

118. Lateral scutellar bristles parallel to one another and shorter than subapical bristles (as in Fig. 130);
ventral surfaces of abdominal tergites 4 and 5 of male each with patch of appressed black hair (sex
patch, Fig. 165) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leptostylum Macquart
– Lateral scutellar bristles divergent and about as long as subapical bristles (Fig. 127); ventral
surfaces of abdominal tergites 4 and 5 of male with or without patches of appressed hair. . . . 119

119. Ocellar bristle well developed, in male as long and as thick as posterior reclinate orbital bristle
(Fig. 19); in female about equal in length and thickness to posteriormost proclinate orbital bristle;
katepisternum with three bristles; male lacking proclinate orbital bristles or patches of appressed
hair on ventral surfaces of tergites; abdominal tergite 5 shining black, or in one species with
narrow transverse basal band of whitish pruinosity. . . . . . . . . . . . . . . . . . Thelairodoria Townsend
– Ocellar bristle hairlike, no larger than posteriormost reclinate orbital bristle; male with single
proclinate orbital bristle, corresponding to anterior one in female (Fig. 24) (a male of A. aldrichi
(Curran) is atypical in having an irregular row of hairlike proclinate orbital setae); katepisternum
with two bristles (Fig. 114); male with ventral surfaces of abdominal tergites 3–5 each with patch
of appressed black hair (as in Fig. 165); abdominal tergite 5 gold pruinose dorsally on at least
basal half . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anoxynops Townsend
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

120. Ventral katepisternal bristle as large as, or larger than, anterodorsal katepisternal bristle (rarely
only slightly thinner) and situated close to upper margin of midcoxa, within no more than twice its
diameter from coxal margin (Fig. 118); vein A1 ending at wing margin (Fig. 160), although apex
of vein may be thin and easily overlooked without transmitted light or light reflected from upper
surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121
– Ventral katepisternal bristle absent or distinctly smaller than anterodorsal katepisternal bristle and
usually situated closer to anterodorsal bristle than to midcoxa (intermediate or closer to coxa in a
few Actia and Ceromya), but not as close to coxa as twice its diameter (Fig. 117); vein A1 ending
in membrane before reaching margin of wing (Fig. 161) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122

121. Labella slightly longer than prementum, and both longer than height of eye (Fig.  81); second
aristomere at least 2.5 times as long as wide; abdomen uniformly tan pruinose, except for darker
ring surrounding base of each median marginal bristle on abdominal tergites 3–5. . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Siphona Meigen
– Labella shorter than prementum; prementum shorter than height of eye (in one species nearly
equal); second aristomere less than twice as long as wide; median marginal bristles of tergites 3–5
encompassed by larger dark areas that blend together, especially medially . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudosiphona Townsend

122. Vein R4+5 setulose dorsally from base to well beyond crossvein r-m (Fig. 161). . . . . . . . . . . . 123
– Vein R4+5 without setulae beyond crossvein r-m. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124

123. Katepisternum with row of fine hairs extending from midventral margin of katepisternum up to
or almost to ventral katepisternal bristle (Fig. 117); anterodorsal corner of anepisternum usually
with two bristles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Actia Robineau-Desvoidy
– Katepisternum bare below ventral katepisternal bristle; anterodorsal corner of anepisternum
usually with single bristle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceromya Robineau-Desvoidy

124. Scutellum lacking both lateral and discal bristles (as in Fig.  132); basal portion of proboscis
when extended longer than prementum (Fig. 82), and membrane between lower genal margin and
clypeus thickened, forming convex paraclypeal sclerite (as in Fig. 80) (not visible if proboscis is
retracted into base of head); labella extending forward. . . . . . . . . . . . . . . . . Ginglymia Townsend
– Scutellum with lateral and discal bristles; basal portion of proboscis shorter than prementum, and
membrane between lower genal margin and clypeus without sclerite; labella either padlike or
extending posteriorly. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125

125. Facial ridge with row of erect bristles on basal half or more. . . . . . . . . . . . . . . . . . . . . . . . . . . 126
– Facial ridge bare except for few small setae above vibrissa . . . . . . . . . . . . . . . . . . . . . . . . . . . 127

126. Vein R1 setose on dorsal surface; antenna of male nearly as long as height of head, that of female
three-quarters or more; both sexes with proclinate orbital bristles, often more than two pairs;
anterior reclinate orbital bristle about as long as posterior reclinate orbital bristle and both about
as long as ocellar bristle (Fig. 18); abdominal tergites each with basal band of golden pruinosity.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Borgmeiermyia Townsend
– Vein R1 bare; antenna of male less than three-quarters height of head, of female even less; male
lacking proclinate orbital bristles, female with at most two pairs; anterior reclinate orbital bristle
much longer and thicker than either posterior reclinate orbital bristle or ocellar bristle; abdomen
uniformly black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oxynops Townsend

127. Veins R4+5 and M ending separately on either side of wing apex relatively far apart (Fig. 158). . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetostigmoptera Townsend, in part
– Veins R4+5 and M both ending before wing apex (as in Fig. 148) . . . . . . . . . . . . . . . . . . . . . . . 128
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1375

128. Both lateral and subapical scutellar bristles long, stout, divergent (as in Fig. 131); vibrissa
subtended by one or more subvibrissal bristles below it (as in Figs.  20–22); three postsutural
supra-alar bristles present, middle one largest. . . . . . . . . . . . . . . . . . . . . . . . Italispidea Townsend
– Lateral scutellar bristles either lacking or short, thin, convergent; subapical bristles also convergent,
crossed medially; vibrissa arising from anteroventral corner of head without subvibrissal bristles
below it (as in Fig. 25); postsutural supra-alar bristles reduced to two: the true first bristle absent;
the apparent first, therefore, the larger of the two (Fig. 99). . . . . . . . . . . . Ischyrophaga Townsend

129. Eye with conspicuous ommatrichia, each longer than combined diameter of four or more eye
facets (as in Fig. 20). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130
– Eye apparently bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134

130. Parafacial with row of stout erect bristles along entire length (Fig. 37); base of vein R4+5 with
single large bristle (as in Figs. 156, 159). . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eulasiona Townsend
– Parafacial lacking row of erect bristles; base of vein R4+5 with more than one small bristle. . . 131

131. Vibrissa arising at level of lower margin of head (as in Fig. 25); with two postpronotal bristles
(as in Fig.  93); middorsal depression on abdominal syntergite  1+2 not extending back to hind
margin of syntergite. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Erythromelana Townsend, in part
– Vibrissa arising above level of lower margin of head, with at least one subvibrissal bristle (Fig. 20);
three or more postpronotal bristles present; middorsal depression on abdominal syntergite 1+2
extending back to hind margin of syntergite (as in Figs. 186, 188). . . . . . . . . . . . . . . . . . . . . . 132

132. Integument of entire fly pale; antenna, legs, and abdomen pale orange with sparse pale pruinosity;
palpus nearly white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Angustia Sellers
– Integument, including antenna, legs, and abdomen, dark; palpus yellow to black. . . . . . . . . . 133

133. Male with two pairs of proclinate orbital bristles; postpronotum with three bristles arranged in
triangle (Fig.  94); abdomen of female rounded or flattened ventrally, not keeled; if sternite  7
modified into piercer (few species), piercer opposed by sternite 6, not ventral carina of abdomen;
surstylus long and slender, usually exceeding cerci in length. . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myiopharus Brauer & Bergenstamm, in part
– Male lacking proclinate orbital bristles; postpronotum, in almost all species, with three to four
bristles, with posteriormost three arranged in arc or line (Fig. 95), in some species with small
fourth anteromedial bristle; abdomen of female keeled midventrally, lower margin usually with
short stout bristles, and sternite  7 modified into curved piercer visible in profile (Fig.  178) or
concealed between ventral edges of abdominal tergites (Fig. 177); surstylus shorter than cerci,
notched on posterior edge at base (as in Fig. 168). . . . . . . . . . . . . . Eucelatoria Townsend, in part

134. Facial ridge setose on lower half or more, with row of erect bristles or hairs or both along most of
length. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
– Facial ridge bare except for few small recumbent bristles above vibrissa. . . . . . . . . . . . . . . . . 150

135. Eye extremely large, covering almost entire side of head, distance between eye and lower cranial
margin no more than twice thickness of palpus (as in Fig. 25); setae on facial ridge mostly
hairlike; usually with three or more proclinate orbital setae in both sexes. . . . . . . . . . . . . . . . 136
– Eye smaller, distance between eye and lower cranial margin more than twice thickness of palpus;
setae on facial ridge coarser; not more than two pairs of proclinate orbital setae present. . . . . 137
1376
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

ds sctl s
l sctl s
b sctl s

sbap sctl s

127 Thelairodoria 128 Micronychiops

125 Cyzenis 126 Frontiniella

b sctl s b sctl s

l sctl s
l sctl s
ap sctl s
ap sctl s
l sctl s sbap sctl s
sbap sctl s ap sctl s

129 Neomintho 130 Eucelatoria 131 Eucelatoria 132 Phyllophilopsis

preap
sctl s

133 Voria 134 Kirbya 135 Austrophorocera 136 Chaetonopsis

Figs. 113.125–136. Scutellar chaetotaxy: dorsal view of scutellum of (125)  Cyzenis albicans (Fallén), (Holarctic, MND, fig.  110.179);
(126)  Frontiniella mitis (Curran), (Nearctic, MND, fig.  110.180, as Cyzenis); (127)  Thelairodoria  sp.; (128)  Micronychiops  sp.;
(129) Neomintho celeris (Townsend), (Nearctic, MND, fig. 110.183); (130) Eucelatoria armigera (Coquillett), (Nearctic, MND, fig. 110.184);
(131) E. leucophaeata (Reinhard), (Nearctic, MND, fig. 110.185); (132) Phyllophilopsis nitens (Coquillett), (Nearctic, MND, fig. 110.186);
(133)  Voria ruralis (Fallén), (Holarctic, MND, fig.  110.187); (134)  Kirbya pacifica (Curran), (Nearctic, MND, fig.  110.188);
(135)  Austrophorocera complicata (Aldrich  & Webber); and (136)  Chaetonopsis spinosa (Coquillett), (Nearctic, MND, fig.  110.190).
Figures 127, 128, and 135 illustrated by A. Brenes.
Abbreviations: ap  sctl  s, apical scutellar seta; b  sctl  s, basal scutellar seta; ds  sctl  s, discal scutellar seta; l  sctl  s, lateral scutellar seta;
preap sctl s, preapical scutellar seta; sbap sctl s, subapical scutellar seta.
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1377

a. b.

138 Pseudodexia

137 Cordyligaster

b sctl
s

sbap
sctl s
l sctl s
ap sctl s

140 Actia 142 Spathidexia

139 Ichneumonops

141 Polygaster

144 Lypha 145 Phasia

143 Neosolieria

146 Gymnosoma 147 Xanthomelanodes

Figs. 113.137–147. Scutellar chaetotaxy (concluded): dorsal view of scutellum, base of abdominal syntergite 1+2, and adjacent structures
of (137)  Cordyligaster sp.; female (a) and male (b) of (138)  Pseudodexia  sp.; and (139)  Ichneumonops  sp.; dorsal view of scutellum of
(140) Actia interrupta (Curran), (Nearctic, MND, fig. 110.182); (141) Polygaster egregia Wulp; (142) Spathidexia sp.; (143) Neosolieria sp.;
(144)  Lypha  sp.; (145)  Phasia  sp; (146)  Gymnosoma fuliginosum Robineau-Desvoidy, (Nearctic); and (147)  Xanthomelanodes  sp.
Figures 137–139 and 141–147 illustrated by A. Brenes.
Abbreviations: ap sctl s, apical scutellar seta; b sctl s, basal scutellar seta; l sctl s, lateral scutellar seta; sbap sctl s, subapical scutellar seta.
1378
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

136. Frontal vitta extremely broad, more than two-thirds width of distance between eyes at midpoint
between antennae and ocellar triangle (Fig. 66) and concolorous with remainder of frons, either
silvery, coppery, or brown; frontal and proclinate orbital bristles displaced laterally nearly to
margin of eye, nearly in line with each other. . . . . . . . . . . . . . . . . . . . . . . . Actinodoria Townsend
– Frontal vitta narrower, about one-third width of frons, darker brown than parafrontal; proclinate
orbital bristles distinctly separated from frontal bristles (Fig. 65). . . . . Euhalidaya Walton, in part

137. Parafacial with setae, sometimes pale or minute and inconspicuous, below lowest frontal bristle
(Figs. 21, 26, 27) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138
– Parafacial bare below lowest frontal bristle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143

138. Subvibrissal ridge with row of four or more bristles that exceeds length of row of setae above
vibrissa (Fig. 27) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
– Subvibrissal ridge with row of three or fewer bristles, shorter than row of setae above vibrissa. . . 140

139. Median discal bristles present on abdominal tergite 3 and usually also on tergite 4. . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cryptomeigenia Brauer & Bergenstamm
– Median discal bristles absent. . . . . . . . . . . . . . . . . . . . . . . . . . . Zaira Robineau-Desvoidy, in part

140. Parafacial setae more numerous on lower half of parafacial and usually recumbent (Fig. 26) (in
some species minute and difficult to see, but may be detected in profile); middorsal depression
on abdominal syntergite 1+2 extending back at least to level of median marginal bristles (as in
Fig.  186); abdomen of female with well-developed midventral carina, usually reinforced with
short stout bristles, and sternite 7 developed into stout curved piercer, usually concealed between
ventral edges of tergites (as in Fig. 177) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Vibrissina Rondani
– Parafacial setae confined to upper half of parafacial; middorsal depression on abdominal
syntergite  1+2 not extending back to level of median marginal bristles; abdomen of female
without midventral carina, and sternite 7 not modified into piercer . . . . . . . . . . . . . . . . . . . . . 141

141. Abdominal tergites 3 and 4 lacking median discal bristles. . . . . . . . . Lixadmontia Wood & Cave
– Abdominal tergites 3 and 4 each with one pair of discal bristles. . . . . . . . . . . . . . . . . . . . . . . . 142

142. Parafacial setae usually adjacent to ptilinal suture, and at least some of them erect (Figs. 21, 75);
lower facial margin concave, not visible between vibrissae in profile. . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Admontia Brauer & Bergenstamm
– Parafacial setae small and recumbent, scattered in middle of parafacial as far from ptilinal suture
as from eye margin; lower facial margin convex, visible between vibrissae in profile. . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudoviviania Brauer & Bergenstamm

143. Medial discal bristles present on abdominal tergites  3 and  4; male lacking proclinate orbital
bristles in Anisia, Blondelia, and Eribella. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144
– Medial discal bristles absent on abdominal tergites 3 and 4; male with two pairs of proclinate
orbital bristles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148

144. Midtibia with at least three well-developed anterodorsal bristles; discal scutellar bristles stout
and inclined at angle of about 45º, arising behind middle of scutellum, and as widely separated as
subapical bristles; syncercus keeled middorsally and surstylus with notch at posterolateral corner
(Fig.  168); abdomen of female keeled midventrally, the lower margin with short stout bristles
and sternite 7 modified into sharp, hooklike piercer, usually concealed between ventral edges of
tergites (as in Fig. 177). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Blondelia Robineau-Desvoidy
– Midtibia with single anterodorsal bristle (as in Fig.  175); discal scutellar bristles weak or
undifferentiated, if present arising closer together than subapical bristles in anterior part of
scutellum; syncercus flattened dorsally and surstylus lacking notch at posterolateral corner;
abdomen of female either with extremely large, sickle-shaped, exposed piercer or rounded
ventrally, without ventral keel or sharp, hooklike piercer. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1379

145. Parafacial parallel-sided and strongly curved to match contour of eye (Figs. 17, 22). . . . . . . . 146
– Parafacial broader dorsally than at level of vibrissa, facial ridge straighter (Fig. 23). . . . . . . . 147

146. Base of R4+5 with single long seta, usually as long as one-third or more distance to crossvein r-m
(as in Figs. 156, 159); sternite 7 of female forming enormous sickle-shaped piercer, opposed by
seta-studded peglike extension of ventral margin of tergite 3 (Fig. 179) . . . . Celatoria Coquillett
– Base of R4+5 with two or more shorter setae; abdomen of female unmodified for piercing host. . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anisia Wulp, in part

147. Facial ridge setose for most of length, setae erect (Fig. 23); discal scutellar bristles long, thin, and
erect, arising at or in front of middle of scutellum, closer together than subapical bristles; abdominal
tergite 4 with single pair of discal bristles; male without proclinate orbital bristles, female with
two pairs; male fifth sternite lobes without tuft of bristles; sternite 7 of female small, triangular,
pointed apically . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eribella Mesnil
– Facial ridge setose only on lower half, with setae recumbent; discal scutellar bristles not
differentiated; abdominal tergite 4 with scattered discal bristles often appearing to be in two
transverse rows; both sexes with two pairs of proclinate orbital bristles crowded against frontal
row (Fig. 67); male fifth sternite lobes each with tuft or clump of medially directed curved bristles
(Fig. 167); sternite 7 of female broad, truncate or obtusely pointed apically (Fig. 189). . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Medina Robineau-Desvoidy

148. Postpronotum with three bristles arranged in triangle (Fig.  94); subapical scutellar bristles
divergent (as in Fig. 130). . . . . . . . . . . . . . . . . . . . . . . Myiopharus Bauer & Bergenstamm, in part
– Postpronotum with two bristles, or with three arranged in arc, with middle bristle displaced
anteriorly not more than its own basal diameter (as in Fig. 95); subapical scutellar bristles parallel
or convergent (as in Fig. 140) (although often displaced from this position in pinned specimens).
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149

149. Pedicel above level of middle of eye (as in Figs. 21, 22); scutellum with discal bristles . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thelyoxynops Townsend
– Pedicel below level of middle of eye (as in Figs. 25, 28); scutellum lacking discal bristles. . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetonodexodes Townsend

150. Median discal bristles present on tergites 3 and 4. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151

– Median discal bristles absent from tergites 3 and 4. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160

151. Eye exceptionally large, covering most of side of head; distance between eye and lower margin
of head less than twice width of palpus (as in Fig. 14); anterior ocellus larger than two posterior
ocelli (Fig. 64); ocellar bristles closer together than width of anterior ocellus; female abdomen
globose, rounded apically, segments  6–9 forming tubular, telescopic ovipositor projecting
ventrally unless fully withdrawn (Fig. 182). . . . . . . . . . . . . . . . . . . . . . . . Calodexia Wulp, in part
– Eye smaller, distance between lower margin of eye and lower margin of head greater than twice
width of palpus; anterior ocellus not appreciably larger than other ocelli; ocellar bristles usually
more widely spaced (except in some species of Chaetona); female abdomen ovoid, with
segments 6–9 projecting terminally when not withdrawn. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152

152. Subvibrissal ridge with row of four or more bristles that exceeds length of row of setae above
vibrissa (as in Fig. 27). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153
– Subvibrissal ridge with row of three or fewer bristles, shorter than row of setae above vibrissa. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154

153. Wing infuscated, usually darker beyond crossvein r-m; arista usually plumose (Fig. 87) . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetona Wulp
– Wing transparent; arista bare. . . . . . . . . . . . . . . . . . . . . . . . . . . Zaira Robineau-Desvoidy, in part
1380
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

154. Vein M ending in R4+5 well before wing margin (as in Fig. 153, but without pattern); female
abdomen somewhat globose, segments 6–9 forming telescopic ovipositor projecting posteroventrally
from apex (as in Fig.  180); prosternum with single large seta on each side (as in Fig.  91);
katepimeron with single small seta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Steleoneura Stein
– Vein M ending in R4+5 at wing margin or ending separately from R4+5; female abdomen either
carinate ventrally (Figs. 177, 178) or somewhat flattened from side to side (Fig. 181); prosternum
with small fine setae; katepimeron usually bare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 155

155. Middorsal depression of syntergite 1+2 extending back at least to level of median marginal bristles
(as in Fig. 186); abdomen of female keeled midventrally, lower margin with short stout bristles,
and sternite 7 forming small, sharp, hooked piercer, usually concealed within ventral edges of
tergites (Fig. 177). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eucelatoria Townsend, in part
– Middorsal depression of syntergite 1+2 not extending back to level of median marginal bristles;
abdomen of female not keeled ventrally and sternite 7 of female not forming piercer. . . . . . . 156

156. Postpronotal bristles arranged in triangle (as in Fig. 94), middle bristle displaced anteriorly by
more than twice width of its base. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157
– Postpronotal bristles in line or arc, middle bristle not displaced anteriorly more than width of its
own base (as in Fig. 95). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159

157. Scutellum lacking lateral bristles (as in Fig. 132); abdomen of female flattened from side to side
(Fig. 181); body uniformly pale brown or gray with inconspicuous dark markings; abdomen
without transverse dark apical bands. . . . . . . . . . . . . . . . . . . . . . . . . Dolichotarsus Brooks, in part
– Scutellum with distinct lateral bristles; abdomen of female more or less cylindrical, as broad as
high; scutum usually with distinct dark stripes; abdominal tergites with apical dark bands. . . 158

158. Scutum with three pairs of presutural acrostichal bristles; sternite  5 of male with single long
bristle (Fig. 169); sternite 6 of female rounded or truncate apically. . . . . . . Lixophaga Townsend
– Scutum with only two pairs of presutural acrostichal bristles, anterior two pairs present,
posteriormost pair absent; sternite 5 of male lacking long bristle; sternite 6 of female large, heavily
sclerotized, bluntly pointed apically (as in Fig. 180). . . . . . . . . . . . . . . . . . Calolydella Townsend

159. Facial ridge curved, parafacial conforming to shape of eye (Fig.  17); antenna inserted above
middle of eye; abdomen globose. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anisia Wulp, in part
– Facial ridge straight, parafacial tapering ventrally (as in Figs.  23, 24); antenna inserted below
middle of eye; abdomen more parallel-sided, tapering apically . . . . . . . . Lydinolydella Townsend

160. Eye exceptionally large, covering almost all of side of head; distance between eye and lower
margin of head less than twice width of palpus (as in Fig. 14); ocellar triangle not raised to form
tubercle; ocellar bristles arising beside or in front of anterior ocellus, their bases about as far apart
as posterior ocelli. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sphaerina Wulp
– Eye smaller, distance between eye and lower margin of head greater than twice width of palpus;
ocellar triangle raised; ocellar bristles arising behind anterior ocellus, their bases closer together
than posterior ocelli. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161

161. Vibrissa arising from anteroventral corner of head (Fig. 25), with at most one subvibrissal bristle
below it; lateral scutellar bristle small or lacking (Fig. 132); postsutural supra-alar bristles usually
reduced to two, true first bristle absent (as in Fig. 99). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162
– Vibrissa arising above anteroventral corner of head (Fig. 20), usually subtended by more than
one subvibrissal bristle; lateral scutellar bristle well developed (as in Figs. 130, 131); postsutural
supra-alar bristles three or more, middle one largest (as in Figs. 100–104) . . . . . . . . . . . . . . . 163
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1381

162. Arista plumose (Fig. 25); genal dilation extending forward to abut vibrissal angle, anterior genal
seta thus arising close to base of vibrissa; midtibia at most with small anterodorsal seta scarcely
longer than width of tibia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phyllophilopsis Townsend, in part
– Arista bare; genal dilation distinctly separated from vibrissal angle by gap of membrane, so that
single subvibrissal seta distinctly separated from genal setae; midtibia with well-developed
anterodorsal seta. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Erythromelana Townsend, in part

163. Lateral scutellar bristles at least four-fifths as long and as straight as subapical scutellar bristles,
strongly divergent (as in Fig.  131); parafacial extremely narrow; with two reclinate orbital
bristles, markedly different from each other in size (as in Fig. 19) . . . Italispidea Townsend, in part
– Lateral scutellar bristles about two-thirds as long as subapical scutellar bristle (as in Fig. 130);
parafacial broader; reclinate orbital bristles more numerous or more uniform in size . . . . . . . 164

164. Ocellar setae minute, shorter than length of ocellar triangle; frontal and reclinate orbital bristles
forming single even row, increasing in size toward vertex usually regularly (as in Figs. 65, 66), or
with abrupt increase in some species; body pale ochreous brown. . . . . . . . . . Ophirion Townsend
– Ocellar setae present, longer than ocellar triangle; frontal and reclinate orbital bristles, if arising
in single row, usually varying in size, with largest frontal bristles in middle of row (as in Figs. 63,
64); body color usually brown or black, except on sides of abdomen. . . . . . . . . . . . . . . . . . . . 165

165. Veins M and R4+5 each ending separately on either side of wing apex (Fig. 158). . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetostigmoptera Townsend, in part
– M and R4+5 both ending anterior to wing apex (as in Fig. 156). . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myiopharus Brauer & Bergenstamm, in part

166. All four of the following character states present simultaneously (Fig. 47): only one frontal bristle
arising below upper margin of pedicel; pedicel with one or two exceptionally long slender setae,
which are more than half as long as arista; arista pubescent or plumose; subvibrissal ridge with
five or more slender bristles usually continuous with bristles of subcranial margin; male with
upper frontal bristles proclinate (Figs. 47, 70, 74). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167
– Frontal bristles usually descending below upper margin of pedicel (Figs. 29–32, 39, 40, 53), but
if not, then either setae on pedicel all shorter than half aristal length or arista minutely pubescent
or apparently bare; subvibrissal ridge usually with fewer than five bristles, usually not continuous
with bristles along subvibrissal ridge; male with upper frontal bristles reclinate. . . . . . . . . . . 182

167. Proepisternum setose (Fig. 107). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168

– Proepisternum bare (as in Fig. 108), rarely with single hair. . . . . . . . . . . . . . . . . . . . . . . . . . . 171

168. Proboscis long, slender, anteriorly curved at apex, more than twice as long as head height
(Fig. 85). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prosenoides Brauer & Bergenstamm
– Proboscis straight, at most 1.5 times as long as head height. . . . . . . . . . . . . . . . . . . . . . . . . . . 169

169. Face with prominent, wide, flat-topped carina that extends beyond antenna in profile (Fig. 71). .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myiomima Brauer & Bergenstamm
– Face with at most small carina that is not visible in profile (Fig. 74) . . . . . . . . . . . . . . . . . . . . 170

170. Genal dilation distinct (as in Figs. 39, 40); vibrissal angle not extending as far forward as antennal
angle; abdomen short and stout, almost as wide as long. . . . . . . . . . . Billaea Robineau-Desvoidy
– Genal dilation undeveloped, genal margin curving forward evenly to lower cranial margin (as in
Fig. 55); vibrissal angle prominent, extending as far forward as antennal angle; abdomen slender,
tapering, more than twice as long as wide. . . . . . . . . . . . . . . . . . . . . . . . . Dolichocodia Townsend
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

171. Middorsal depression of abdominal syntergite 1+2 extending back to hind margin of that segment
(as in Figs. 186, 188). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 172
– Middorsal depression of syntergite  1+2 smaller and shallower, not extending back to hind
margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 179

172. Proboscis long and slender, longer than height of head (as in Figs. 53–55). . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mochlosoma Brauer & Bergenstamm
– Proboscis shorter and stouter, not as long as head height . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173

173. Fronto-orbital plate outside of row of frontal bristles almost devoid of setae except near ocellar
triangle (Fig. 70). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 174
– Small setae lateral to frontal bristles extending forward to base of antenna (as in Figs. 69, 72, 74).
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175

148 Myothyriopsis 149 Phasia

150 Bibiomima 151 Euantha

152 Catharosia 153 Euthera

154 Euscopoliopteryx 155 Oestrophasia

Figs. 113.148–155. Wings: dorsal view of (148)  Myiothyriopsis picta (Wulp), (Nearctic, MND,  fig. 110.208); (149)  Phasia  sp.;
(150) Bibiomima handlirschi Brauer & Bergenstamm; (151) Euantha litturata (Olivier), (Nearctic, MND, fig. 110.209); (152) Catharosia lustrans
(Reinhard), (Nearctic, MND, fig. 110.211); (153) Euthera tentatrix Loew, (Nearctic, MND, fig. 110.212); (154) Euscopoliopteryx sp.; and
(155) Oestrophasia clausa Brauer & Bergenstamm, (Nearctic, MND, fig. 110.214). Figures 149, 150, and 154 illustrated by A. Brenes.
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1383

174. Abdomen strongly pointed apically, especially in male; yellow or reddish basally, with median
black stripe on basal segments and tessellated patterns laterally. . . . . . . Zelia Robineau-Desvoidy
– Abdomen black or dark brown; tergites 3, or 3 and 4, of some species each with pair of conspicuous
rounded, silvery or bluish spots. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scotiptera Macquart

175. Tergite 3 with one pair of median marginal bristles and no discals. . . . . . . Oberonomyia Reinhard
– Tergite  3 with at least one pair of discal bristles and more than one pair of median marginal
bristles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176

176. Tergites 3 and 4 each with two pairs of discal bristles, one in front of other (Fig. 188). . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ptilodexia Brauer & Bergenstamm
– Tergite  3 with triangular patch of discals; tergite  4 with at least one transverse row of discal
bristles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177

s
R2+3
R4+5

M1

bend of M

156 Drino 157 Chetogena

R1 s
R4+5

r-m
R4+5

M
bend of M

158 Chaetostigmoptera 159 Phytomyptera

R1

r-m
R4+5
R4+5

A1
A1

160 Pseudosiphona 161 Actia

R4+5

M
extension of M

162 Cyrtophleba 163 Microchaetina

Figs. 113.156–163. Wings (concluded): dorsal view of (156)  Drino antennalis (Reinhard), (Nearctic, MND, fig.  110.193); (157)  Chetogena
gelida (Coquillett), (Nearctic, MND, fig.  110.197); (158)  Chaetostigmoptera crassinervis (Walton), (Nearctic, MND, fig.  110.195);
(159)  Phytomyptera tarsalis (Coquillett), (Nearctic, MND, fig.  110.196); (160)  Pseudosiphona flavipes (Coquillett), (Nearctic, MND,
fig. 199, as Ceranthia); (161) Actia interrupta (Curran), (Nearctic, MND, fig. 110.200); (162) Cyrtophleba horrida (Giglio-Tos); and
(163) Microchaetina sinuata (Townsend), (Nearctic, MND, fig. 202). Figure 162 illustrated by A. Brenes.
Abbreviation: s, seta (nodal seta) at bases of R2+3 and R4+5.
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

177. Discal bristles covering all of tergite 3 except for small anterolateral region overlapped by wing;
those on tergite 4 entirely covering dorsal surface; abdomen shiny black. . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tropidopsiomorpha Townsend
– Discal bristles on tergite 3 arranged in median patch occupying one-third to one-half of tergite;
those on tergite 4 arranged in one or more transverse rows across tergite; abdomen orange, red,
brown or dull black, not shiny. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 178

178. Discal bristles on tergite 4 forming single irregular row across tergite; abdomen brown, reddish,
or dull black. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hystricodexia Roeder
– Discal bristles on tergite 4 arranged in at least three transverse rows; abdomen orange. . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eudexia Brauer & Bergenstamm

179. M ending in R4+5 before reaching wing margin (Fig. 163). . . . . . . . . . Microchaetina Wulp, in part
– M ending separately in wing margin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180

180. Face with well-developed carina separating antennae and visible in profile (as in Fig. 71); anterior
katepisternal bristle arising distinctly anterior to pleural suture (as in Fig. 122); dorsal surface of
costa lacking long bristle; posterior thoracic spiracle covered with operculum, distinctly larger
than anterior lappet (as in Fig. 106); abdominal tergite 5 of male prolonged behind genitalia into
slender cylindrical process, studded with long bristles. . . . . . . . . . . . . . . . . Trichodura Macquart
– Face with at most low carina that is not visible in profile; anterior katepisternal bristle arising below
pleural suture, or absent; base of costa with long bristle on dorsal surface (Fig. 151); anterior and
posterior lappets of posterior thoracic spiracle nearly equal in size, each composed of widely
spaced, hairy, fingerlike outgrowths with V-shaped opening between the two; male abdominal
tergite 5 truncate or rounded apically. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181

181. Frontal vitta distinct; scapes distinctly separated medially; wing strongly patterned, veins
emphasized with dark brown (Fig. 151); two katepisternal bristles present. . . . . . . Euantha Wulp
– Frontal vitta virtually obliterated, frontal bristles of two sides nearly confluent medially; scapes
adjacent to one another; wing transparent; only one katepisternal bristle present, anteriormost
absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neosophia Guimarães

182. With following three character states occurring simultaneously: scutellum with one to three
straight, nearly erect preapical bristles arising on dorsal surface just anterior to apex (Figs. 133,
134); anterior margin of postgena at most with slight genal dilation (Figs. 29–32); last section of
CuA1 (beyond crossvein dm-cu) more than half as long as preceding section (Fig. 162). . . . . 183
– Scutellum without group of one to three long, straight preapical bristles (except some Trafoia);
anterior margin of postgena usually with well-developed genal dilation; last section of CuA1
usually less than half as long as preceding section . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190

183. Anepimeron with exceptionally long stout bristle, extending beyond apex of lower calypter (as in
Fig. 111); Holarctic Region, including adjacent Nearctic Mexico. . . [Athrycia Robineau-Desvoidy]
– Anepimeron with at most short bristle not extending beyond middle of lower calypter (as in
Fig. 112). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184

184. Parafacial with single large proclinate bristle below lowest frontal bristle (Fig. 30) . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Voria Robineau-Desvoidy
– Parafacial with row of stout bristles or patch of small decumbent setae (Figs. 29, 31, 32). . . . 185

185. Ocellar bristles arising in front of anterior ocellus; lowest frontal bristle reclinate and separated
from remaining frontals by distance greater than that separating them; veins R1, R4+5, and CuA1
each bristled from base to well beyond middle. . . . . . . . . . . . . . . Minthoplagia Townsend, in part
– Ocellar bristles arising behind anterior ocellus; lower frontal bristles all equally spaced, lowest
not more widely separated than others; only vein R4+5 with few bristles at base. . . . . . . . . . . . 186
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1385

186. Vein M ending in R4+5 well before wing margin (as in Fig. 163); frontal and parafrontal bristles
each arising from shiny black spot (Fig. 35). . . . . . . . . . . . . . . . . . . . Kirbya Robineau-Desvoidy
– Vein M ending separately in wing margin (as in Figs. 156–160); frontal and parafacial bristles
each surrounded by silvery pruinosity (Figs. 31, 32). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187

187. Parafacial with vertical row of three or more stout erect proclinate bristles (Fig. 31) . . . . . . . 188
– Parafacial with patch of small recumbent setae, one or more of which may be larger than the
others, but none appearing as long erect bristle (Fig. 32) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189

188. Eye covered with conspicuous ommatrichia (Fig. 31).. . . . . . . . . . . . . . . . . Cyrtophleba Rondani

– Eye apparently bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Meleterus Aldrich

189. Prementum slender, straight and as long as or longer than height of head; labella reduced, pointed
apically (Fig. 86) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plagiomima Brauer & Bergenstamm
– Prementum shorter than head height; labella enlarged (Fig. 32). . . . . . . . . . Metaplagia Coquillett

190. Parafacial setose on upper half or more. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 191

– Parafacial mostly bare, at most with few setae on upper third or less. . . . . . . . . . . . . . . . . . . . 207

191. M ending in R4+5 well before wing margin; united petiole at least half as long as that part of M
beyond bend (as in Figs. 149, 153) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 192
– M ending separately in wing margin or in R4+5 close to margin . . . . . . . . . . . . . . . . . . . . . . . . 194

192. Ocellar bristles reclinate (as in Fig. 11); antennal bases separated from one another (as in Fig. 74);
calypter exceptionally large, white, extending over basal third of abdomen (Fig. 112); apex of
female abdomen with pincerlike cerci opposing one another horizontally, each with sharp apex
and inner margin armed with peglike setae (Fig. 166). . . . . . . . . . . . . Leucostoma Meigen, in part
– Ocellar bristles proclinate (Fig. 34) or absent; antennal bases contiguous; calypter not exception-
ally large; abdomen of female without modified pincerlike cerci. . . . . . . . . . . . . . . . . . . . . . . 193

193. Postpronotum with four bristles, the outer three arranged in triangle (as in Fig. 94) . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Periscepsia Gistel
– Postpronotum with two bristles, or with third small inner bristle arranged in nearly straight row.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paradidyma Brauer & Bergenstamm, in part

194. Forecoxa nearly as long as femur, extending nearly to midcoxa; foretarsus thickened dorsoventrally,
flattened laterally, and elongate; scutellum with small basal bristle, long subapical bristle, and
small crossed apicals (as in Fig. 132 except that apicals are crossed); postpronotum with only two
bristles (as in Fig. 93). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Actinochaeta Brauer & Bergenstamm
– Forecoxa less than half as long as femur; foretarsus cylindrical; scutellum with three or more pairs of
bristles, basals longer and lateral bristles usually present; postpronotum with three bristles. . . . . . 195

195. Median surface of forecoxa almost entirely covered with short appressed black setae (as in
Fig.  90); antenna small, recessed into small, deep oval facial cavity; lower part of face long
and narrow, with several pairs of large subvibrissal bristles above level of lower facial margin
(Figs. 44–46) (some species of Cholomyia with part or all of median surface of forecoxa haired,
but antenna longer and lower part of face not long and narrow). . . . . . . . . . . . . . . . . . . . . . . . 196
– Median surface of forecoxa with setae confined to anterolateral edge; antenna and face not modified
as above. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 199
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

196. Abdomen stout, unicolorous; tergite 3 with more than one pair of median marginal bristles. . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Megaprosopus Macquart
– Abdomen less stout, tessellated; tergite 3 with one pair of median marginal bristles. . . . . . . . 197

197. Arista plumose; uppermost subvibrissal bristle nearly as large as vibrissa and separated from it by
more than twice distance between it and remaining subvibrissal bristles (Fig. 45); katepisternum
with two bristles, anterior bristle located below and in line with pleural suture (as in Fig. 123) . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pyrrhodexia Townsend
– Arista pubescent or bare; uppermost subvibrissal bristle separated from vibrissa less than twice
distance between it and remaining subvibrissal bristles (Fig. 46); katepisternum usually with three
bristles, rarely two, anteriormost located in front of pleural suture, middle bristle located below
suture (as in Fig. 122). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 198

198. Arista pubescent; scape without projecting hood over pedicel; crossveins r-m and m-cu and
vein M beyond bend, and anterior margin of wing darkened . . . . . . . . . . . . . Acronacantha Wulp
– Arista bare; scape with dorsal flange projecting beyond base of pedicel (Fig. 44); wing veins not
emphasized by dark markings. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microphthalma Macquart

199. All three following character states occurring simultaneously: vibrissa not distinctly differentiated
from remaining subvibrissal bristles; genal groove exceptionally large; postgenal margin vertical,
ending below eye (Fig. 51). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 200
– Vibrissa distinctly differentiated as single, large, erect bristle; genal groove not unusually
extensive; postgenal margin extending forward below eye as distinct genal dilation (as in Figs. 39,
40) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 201

200. Scutellum with three pairs of bristles: basal, lateral, and apical; lateral bristles curve laterally. . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eutrixa Coquillett
– Scutellum with four pairs of bristles: basal, lateral, subapical, and apical (as in Fig. 131); lateral
and subapical bristles straight or curve medially . . . . . . . . . . . . . . . . . . . . . . . . Isidotus Reinhard

201. Parafacial setae of two sizes: single row of larger bristles along anterior margin, sometimes even
arising from membrane anterior to it, flanked by smaller and finer setae laterally (Fig.  43);
integument of body shiny black with bluish or greenish luster; robust species, abdomen more
than half as wide as long; tergite 6 and syntergosternite 7+8 of male exposed; eye facets of male
enlarged on upper part of eye; ommatrichia, when present, longer above than below. . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gnadochaeta Macquart
– Parafacial setae not differentiated into two series; more slender species, with transversely banded,
pruinose, or partly yellow abdomen; more slender species, abdomen less than half as wide as
long; tergite 6 and syntergosternite 7+8 of male mostly concealed beneath tergite 5; eye facets of
male not enlarged dorsally; ommatrichia uniform in length. . . . . . . . . . . . . . . . . . . . . . . . . . . 202

202. Parafacial with single row of conspicuous, erect bristles (Figs. 16, 36). . . . . . . . . . . . . . . . . . 203
– Parafacial with decumbent, often inconspicuous hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 204

203. Aristomere 3 strongly swollen on basal fourth or less, tapering abruptly to apex (Fig. 36); first
flagellomere no longer than 1.5 times length of pedicel; vibrissa arising above level of lower facial
margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Muscopteryx Townsend
– Aristomere 3 thickened on basal third or more before tapering to apex; first flagellomere 2 or more
times length of pedicel; vibrissa arising at level of lower facial margin (Fig. 16). . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paradidyma Brauer & Bergenstamm, in part
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1387

204. Palpus greatly reduced, shorter than scape, with single long terminal bristle more than 4 times as
long as palpus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Linnaemya Robineau-Desvoidy, in part
(L. comta (Fallén) species complex)
– Palpus of normal length, clavate, longer than pedicel, lacking long terminal bristle . . . . . . . . 205

205. Scutellum with long, stout, divergent lateral bristles (as in Fig.  132) and small, crossed apical
bristles (as in Fig. 144). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stenodexia Wulp
– Scutellum with small, convergent lateral bristles and stout, convergent subapical bristles (as in
Figs. 136, 145). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 206

206. Parafacial wide and convex, with scattered hairs, often pale and inconspicuous (Fig. 52); males of
some species with largely yellow abdomen and extremely long midfemur and tibia, other species
darker and with more normal leg proportions; some species with medial surface of forecoxa bearing
appressed black hairs, others with downwardly directed hairs on underside of scutellum. . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cholomyia Bigot
– Parafacial flat, with single row of small black bristles; body essentially black in color; midleg not
modified; medial surface of forecoxa bare; scutellum without hairs on underside. . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phasiophyto Townsend

207. With following two characters occurring together: scutum with single postsutural supra-alar bristle
(probably corresponding to largest, or middle, bristle in usual group of three, although relatively far
forward for this bristle), and vibrissa raised far above lower margin of face (as in Fig. 73) . . . . 208
– Scutum with two or three postsutural supra-alar bristles, or if only one, then vibrissae level with
lower facial margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215

208. Scutellum with basal bristles longer than apical bristles, widely divergent from one another, and
arising in lateral position, midway between anterolateral corner of scutellum and apical bristles
(Fig. 147). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xanthomelanodes Townsend
– Scutellum with basal bristles shorter than or about equal in length to apical bristles, directed posteriorly,
and inserted nearer anterolateral corner of scutellum than base of apical bristle (Fig. 145). . . 209

209. M ending in R4+5 well before wing margin; united petiole at least twice as long as crossvein r-m
(Figs. 149, 154) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 210
– M ending separately in wing margin or in R4+5 close to wing margin (as in Fig. 150). . . . . . . 211

210. Apex of vein M evenly curved, meeting R4+5 at distance from wing apex greater than length of
crossvein r-m (Fig. 149); wing clear or evenly infuscated, or with wide, weakly defined, darker
crossband that does not include apex of vein M; arista bare. . . . . . . . . . . . . . . . . Phasia Latreille
– Vein M bent sharply at right angles, with small stump vein at bend; wing patterned, with costal
area, including vein R2+3, crossveins, and apex of M infuscated (Fig. 154); arista sparsely short
plumose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Euscopoliopteryx Townsend

211. Hind tibia with dorsal longitudinal row of flattened bladelike setae (Figs. 170, 171). . . . . . . . 212
– Hind tibia with usual row of bristles but lacking flattened setae. . . . . . . . . . . . . . . . . . . . . . . . 213

212. Postmetacoxal area membranous, paler than adjacent metepimera, and ridges or ripples, if any,
vertical; entire length of dorsal edge of hind tibia with long flattened bladelike setae (Fig. 170). .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trichopoda Berthold
– Postmetacoxal area sclerotized, i.e., posterior edges of metepimera extending posteromedially
to meet each other at midline between hind coxae and base of first abdominal sternite, forming
postmetacoxal bridge (as in Fig. 98), recognizable even when sclerotization is pale, by presence
of several transverse ridges or ripples; apical two-thirds or less of dorsal edge of hind tibia with
short flattened setae (Fig. 171). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Homogenia Wulp
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

164 Blepharipa 165 Leptostylum 166 Leucostoma

167 Medina

168 Blondelia 169 Lixophaga

170 Trichopoda 172 Leschenaultia 174 Chaetogaedia 176 Trochiloleskia

171 Homogenia 173 Blepharipa 175 Leskia

Figs. 113.164–176. Apical abdominal segments, male genitalic structures, and tibiae: ventral view of segments  4 and  5 of male of
(164) Blepharipa pratensis (Meigen), (Holarctic, MND, fig. 110.247); (165) Leptostylum leuconotum (Wulp) (terminalia omitted), (Wood,
1985, fig. 42); dorsal view of segments 4 and 5 of female of (166) Leucostoma simplex (Fallén), (Holarctic, MND, fig. 110.249); ventral
view of sternite 5 of male of (167) Medina sp., (Nearctic, Wood, 1985, fig. 41); posterodorsal view of male terminalia of (168) Blondelia
hyphantriae (Tothill), (Nearctic, Wood, 1985, fig. 39); ventral view of sternite 5 of (169) Lixophaga variabilis (Coquillett), (Nearctic, MND,
fig.  110.262); posterior view of left hind tibia of (170)  Trichopoda pennipes (Fabricius), (MND, fig.  110.224); (171)  Homogenia  sp.;
(172)  Leschenaultia reinhardi (Toma  & Guimarães), (Nearctic, MND, fig.  110.225, as L.  leucophrys); and (173)  Blepharipa albicauda
(Townsend); dorsal view of hind tibia of (174) Chaetogaedia sp.; dorsal view of mid tibia of (175) Leskia sp.; and (176) Trochiloleskia loriola
(Reinhard), (Nearctic). Figures 171 and 173–176 illustrated by A. Brenes.
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1389

st 7

177 Eucelatoria 178 Eucelatoria

st 6

st 7

179 Celatoria 180 Phyllophilopsis

181 Dolichotarsus 182 Calodexia

Figs. 113.177–182. Female abdomens of Blondeliini: lateral view of (177) Eucelatoria leucophaeata (Reinhard), (Nearctic, Wood, 1985,
fig. 53); (178) E. texana (Reinhard), (Nearctic, MND, fig. 110.256); (179) Celatoria diabroticae (Shimer), (Nearctic, MND, fig. 110.257);
(180) Phyllophilopsis nitens (Coquillett), (Nearctic, MND, fig. 110.258); (181) Dolichotarsus griseus Brooks, (Nearctic, MND, fig. 110.259);
and (182) Calodexia sp., (Wood, 1985, fig. 49).
Abbreviation: st, sternite.
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

213. Wing infuscated only along veins, leaving larger cells mostly transparent (Fig. 150); head and
body almost entirely velvety black . . . . . . . . . . . . . . . . . . . . . Bibiomima Brauer & Bergenstamm
– Wing uniformly infuscated on anterior half or more, sometimes with central paler area; head and
body colored in various shades of yellow, orange, and brown . . . . . . . . . . . . . . . . . . . . . . . . . 214

214. Scutum with one presutural dorsocentral bristle and one postsutural dorsocentral. . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acaulona Wulp
– Scutum with two presutural dorsocentral bristles and three postsutural dorsocentrals. . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xanthomelanopsis Townsend

215. Postmetacoxal area sclerotized, i.e., posterior edges of metepimera extending posteromedially
to meet each other at midline between hind coxae and base of first abdominal sternite, forming
postmetacoxal bridge (as in Fig. 98); abdomen slender, wasplike, much longer than wide. . . 216
– Postmetacoxal area membranous, distinctly paler in color than adjacent metepimera; abdomen no
more than twice as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219

216. Hind margin of eye indented (as in Fig. 54); ocellar triangle extending forward to lunule as
polished brown straplike sclerite (Fig. 68); palpus present; vibrissa minute; tibiae and most of
femora yellowish.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemyda Robineau-Desvoidy
– Hind margin of eye straight or evenly convex; ocellar triangle normal in shape, not dividing
frontal vitta; palpus absent; vibrissa distinct; legs entirely dark . . . . . . . . . . . . . . . . . . . . . . . . 217

217. Abdomen, although slender, widening evenly from base to middle, then tapering to apex; scutellum
with basal, lateral, and apical bristles (as in Fig. 128); vibrissa clearly differentiated . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cylindromyia Meigen
– Abdomen wasplike, syntergite 1+2 strongly constricted at intersegmental junction, narrower
than segment 1; scutellum with one to two pairs of bristles (Fig. 139); vibrissa not clearly
differentiated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218

218. Scutellum short and truncate, with single pair of bristles (Fig.  139); subscutellum weakly
developed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ichneumonops Townsend
– Scutellum with two pairs of bristles, basal and apical; subscutellum well developed. . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polistiopsis Townsend

219. M ending in R4+5 well before wing margin (Figs. 152, 153) . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
– M ending in wing margin or in R4+5 near margin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 228

220. Ocellar bristles lateroclinate or slightly reclinate (as in Fig.  76); calypter exceptionally large,
extending beyond middle of syntergite 1+2 (Fig. 112); apex of female abdomen with pincerlike
cerci, each with sharp apex and peglike setae on inner margin (Fig. 166) . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leucostoma Meigen, in part
– Ocellar bristles proclinate or absent; calypter not extending back to middle of syntergite 1+2; apex
of female without pincerlike cerci. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221

221. Wing slender, usually darkened with whitish apex; M beyond bend meeting R4+5 at right angles;
R4+5+M twice as long as portion of vein M between crossvein r-m and bend of M (Fig. 152). . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Catharosia Rondani
– Wing not as described above; vein M not so abruptly bent anteriorly. . . . . . . . . . . . . . . . . . . . 222

222. Large species, over 10 mm in length, slender, black, resembling pompilid wasps, with entirely
black wings; facial ridge with recumbent hairs (Fig.  42); upper part of parafacial with several
anteromedially directed setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Penthosia Wulp
– Smaller species, less than 10 mm in length, not entirely black, nor with black wings, not
resembling pompilid wasps; facial ridge and parafacial bare . . . . . . . . . . . . . . . . . . . . . . . . . . 223
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1391

223. Wing patterned with brown spots, bars, and triangles (Figs. 153, 155). . . . . . . . . . . . . . . . . . . 224
– Wing transparent, unpatterned. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225

224. Face with prominent oval central tubercle or carina (Fig. 41); antenna long, cylindrical; pedicel
and first flagellomere each as long as arista; lunule without setae; thorax black; apex of cell r4+5
dark (Fig. 153). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Euthera Loew
– Face with at most low indistinct keel-like carina between antennae; antenna short; length of
pedicel and first flagellomere together no longer than arista; lunule, in some species, with minute
setae (Fig. 69); thorax yellow or pale brown; apex of cell r4+5 transparent (Fig. 155). . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oestrophasia Brauer & Bergenstamm

225. Dorsum of abdomen lacking erect bristles; median marginal setae, if present, recumbent and
scarcely differentiated from other setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 226
– Dorsum of abdomen with erect median marginal bristles on tergites 3 and 4. . . . . . . . . . . . . . 227

226. Apex of scutellum without bristles; apical pair so widely separated and short that they appear as
lateral bristles and cannot cross at their apices (Fig. 146); abdomen shiny, orange–red with black
markings, with short hairs each arising from raised base, giving abdomen somewhat rugose
appearance; intertergal sutures indistinct, their positions most easily recognizable by discontinuities
in orange and black pattern; pedicel and first flagellomere elongate, extending to level of vibrissa
(Fig. 72). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gymnosoma Meigen
– Apex of scutellum with usual crossed pair of apical bristles (as in Fig. 145); abdomen usually
golden pruinose (in males) or gray pruinose (in females); intertergal sutures distinct; pedicel and
first flagellomere short, not extending to level of vibrissa (Fig. 73) . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gymnoclytia Brauer & Bergenstamm

227. Eye large, occupying almost all of side of head; gena reduced to narrow band less than one-tenth
height of head (Fig. 49); lunule with minute setae. . . . . . . . . . . . Strongygaster Macquart, in part
– Eye smaller; gena much wider, one-sixth height of head or more; lunule bare. . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Vanderwulpia Townsend

228. Eye almost entirely covered with conspicuous ommatrichia. . . . . . . . . . . . . . . . . . . . . . . . . . . 229

– Eye apparently bare. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 241

229. With following four character states together: facial ridge with row of stout erect evenly spaced
bristles (Fig. 76); ocellar bristle laterally or posterolaterally inclined; genal dilation with isolated
bristle at anterior margin; and proepisternum with pale hairs (as in Fig. 107) . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Campylocheta Rondani
– With no more than one of the above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 230

230. Palpus reduced, parallel-sided, usually shorter than pedicel (Fig. 39). . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Linnaemya Robineau-Desvoidy, in part
– Palpus clavate, normal in length, longer than pedicel (Fig. 40). . . . . . . . . . . . . . . . . . . . . . . . . 231

231. Postpronotum, proepimeron, most of mesopleuron, and hind coxa with dense tufts of pale
plumose hairs (Fig. 109); meral setae forming tuft of pale plumose hairs, sometimes accompanied
by or interspersed with black bristles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Telothyria Wulp, in part
– Thoracic hairs not plumose and not especially long or dense, may be pale or dark; meral bristles
black, unaccompanied by plumose hairs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 232

232. Vibrissa arising above lower facial margin, which protrudes anteroventrally, visible in profile (as
in Figs. 38–40). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 233
– Vibrissa arising at same level as lower facial margin, which is not visible in profile.. . . . . . . . 237
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 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

233. Scutellum with only two pairs (basal and subapical) of bristles, subapicals divergent (Fig. 143) .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neosolieria Townsend
– Scutellum with three or more pairs (basal, lateral, and subapical) of bristles, and usually also with
small pair of crossed apical bristles (as in Fig. 131) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 234

234. Katepimeral bristles confined to area behind pleural suture, anterior bristle arising directly
below ventral end of pleural suture (Fig. 119); scutellum with single lateral bristle on each side
(Fig. 128). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 235
– Anterior katepisternal bristle arising anterior to ventral end of pleural suture (as in Fig.  124);
scutellum with two to three pairs of lateral bristles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 236

235. Scutellum with several long erect discal bristles; male with ocellar bristles proclinate and nearly
parallel to each other (Fig. 38); female with ocellar bristles lateroclinate to slightly reclinate. . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trafoia Brauer & Bergenstamm
– Scutellum with discal bristles decumbent (Fig. 128); ocellar bristles of both sexes proclinate and
moderately divergent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Micronychiops Townsend

236. Both sexes with frontal bristles extending to vertex, and in addition three pairs of orbital bristles
oriented in different directions, anterior bristle proclinate (lacking in some males of one species),
middle bristle lateroclinate, and posterior bristle reclinate, arising beside uppermost frontal bristle
(Fig. 58); tergite 6 and syntergosternite 7+8 of male much shorter than tergite 5, and together with
epandrium, largely withdrawn inside tergite. . . . . . . . . . . . . . . . . . . . . . . Neobrachelia Townsend
– Male lacking orbital bristles, although with proclinate frontal bristles (Fig. 40); female with two
pairs of proclinate orbital bristles and small reclinate orbital bristle; tergite 6 and syntergosternite 7+8
of male as extensive as tergite 5, and epandrium bulbous and conspicuous. . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Panzeria Robineau–Desvoidy

237. Arista long plumose (Fig. 33) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ebenia Macquart, in part

– Arista bare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 238

238. Posterior thoracic spiracle with fringe of short hairs leaving most of spiracle open. . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hyalurgus Brauer & Bergenstamm
– Posterior thoracic spiracle with comblike anterior and posterior fringes (lappets) of plumose hairs
that cover most of spiracle, leaving V-shaped middorsal opening (as in Fig. 105). . . . . . . . . . 239

239. Middorsal depression on syntergite  1+2 extending back at least to level of median marginal
bristles, and usually to hind margin of segment 2; eyes of male closely approximated medially,
nearly obliterating frontal vitta, which is narrower than ocellar triangle; vertical bristles of male
short, weak, and proclinate (as in Fig. 37). . . . . . . . . . . . . . . . . . . . . Uramya Robineau-Desvoidy
– Middorsal depression on syntergite 1+2 extending back to middle of segment 2; frontal vitta of
male, at its narrowest point, wider than ocellar triangle; inner vertical bristles of male well
developed and reclinate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 240

240. Facial ridge with four to five small, erect setae above vibrissa; scutellum with pair of widely separated
discal bristles; abdominal tergites lacking median discal bristles. . . . . . . . . . . . Itaplectops Townsend
– Facial ridge with few recumbent hairs above vibrissa; scutellum without discal bristles; abdominal
tergites 3 and 4 each with pair of median discal bristles. . . . . . . . . . . . . Thelairaporia Guimarães

241. Palpus minute, shorter than pedicel, or absent (Fig. 54). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 242

– Palpus well developed, longer than pedicel. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 243
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1393

242. Hind margin of eye indented (Fig. 54); prementum slender, elongate, more than twice as long as
height of head; palpus absent; abdomen orange–red, slender, contrasting with dark wing. . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Beskia Brauer & Bergenstamm
– Hind margin of eye straight or evenly convex; prementum shorter than height of head; palpus
minute; head and body pale reddish-brown pruinose . . . . . . . . . . . . . . Icelia Robineau-Desvoidy

243. Ocellar setae inclined laterally or posterolaterally (as in Fig. 76) . . Clairvillia Robineau-Desvoidy
– Ocellar setae proclinate or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 244

244. Vein M nearly straight, extending to wing margin without abrupt bend, thus resembling that of
muscids or acalyptrates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinochira Zetterstedt
– Vein M with distinct angular bend. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245

245. Lower proepimeral bristle well developed, directed anteroventrally, in opposite direction to upper
proepimeral bristle (Fig. 108) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neaera Robineau-Desvoidy
– Lower proepimeral bristle absent, or reduced in size and directed dorsally, parallel to upper
proepimeral (as in Fig. 107). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 246

246. All three following character states occurring together: vibrissa not distinctly differentiated from
remaining subvibrissal bristles; genal groove exceptionally large; postgenal margin vertical,
ending below eye (Fig. 51). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . return to couplet 200
– Vibrissa distinctly differentiated as single large erect bristle; genal groove not so extensive; postgenal
margin extending forward below eye as distinct genal dilation (as in Figs. 39, 40). . . . . . . . . 247

247. Prosternum inflated (Fig. 92). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ormia Robineau-Desvoidy

– Prosternum normal, flat or concave (as in Figs. 90, 91) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 248

248. With following three character states occurring together: anterior katepisternal bristle arising
directly ventral to pleural suture (Fig. 123); lateral scutellar bristles convergent (Figs. 141, 142);
vibrissa arising at anteroventral corner of head subtended by at most single subvibrissal bristle (as
in Fig. 25). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 249
– Anterior katepisternal bristle arising distinctly anterior to pleural suture (as in Fig. 124); lateral
scutellar bristles divergent or parallel; vibrissa usually arising above anteroventral corner of head,
subtended by more than one subvibrissal bristle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251

249. Frons of both sexes with two pairs of proclinate orbital bristles and two pairs of reclinate
orbital bristles, anterior pair of latter convergent (inclinate), posterior pair divergent (lateroclinate);
scutellum trapezoidal, with small crossed apical bristles enclosed by long convergent subapical
bristles, lateral bristle absent (Fig. 141); veins R1, R4+5, and CuA1 setose dorsally along most of
their lengths; with two to three meral bristles. . . . . . . . . . . . . . . . . . . . . . . . . . . . Polygaster Wulp
– Frons of male lacking proclinate orbital bristles; female as above; scutellum semicircular, rounded
apically, with crossed apical and subapical bristles, small convergent lateral bristles arising close
to subapicals, and large basal bristles (Fig. 142); vein R1 setose or bare, vein R4+5 setose only to
crossvein r-m, and CuA1 bare, except for single bristle basally in some species; with three or more
meral bristles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 250

250. First flagellomere of both sexes parallel-sided in profile; frontal bristles all reclinate, continuous
with reclinate orbital bristle; inner orbital bristles more or less parallel, not crossed at their apices;
outer orbital bristles absent; with three to four meral bristles. . . . . . . . . . . Spathidexia Townsend
– First flagellomere of male somewhat swollen apically, narrowed in middle (Fig. 48); of female
parallel-sided, as above; frontal bristles of both sexes proclinate, contrasting with reclinate
upper orbital bristles; inner orbital bristles convergent, crossing at their apices; outer orbital
bristles present; usually with more than five meral bristles. . . . . . . . . . . . . . . . . Thelairodes Wulp
1394
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

m mg s

ds s
m ds s

183 Argyrochaetona 184 Chlorohystricia 185 Bombyliomyia

midd dp

midd dp

m ds s

186 Blondelia 187 Argyromima 188 Ptilodexia

189 Medina 190 Spanthidexia 191 Phasia

Figs. 113.183–191. Abdomens: dorsal view of abdomen of (183) Argyrochaetona cubana Townsend; (184) Chlorohystricia cyaneiventris
(Wulp), (MND, fig. 110.233); (185) Bombyliomyia soror (Williston), (Nearctic, MND, fig. 110.234); (186) Blondelia hyphantriae (Tothill),
(Nearctic, MND, fig.  110.231); (187)  Argyromima mirabilis Brauer  & Bergenstamm; (188)  Ptilodexia rufipennis (Macquart), (MND,
fig. 110.239); ventrolateral view of (189) Medina barbata (Coquillett), (Nearctic, MND, fig. 110.252); ventral view of (190) Spathidexia
clemonsi Townsend, (Nearctic, MND, fig. 110.251); ventrolateral view of (191) Phasia fenestrata (Bigot), (Nearctic, MND, fig. 110.253).
Figures 183 and 187 illustrated by A. Brenes.
Abbreviations: ds s, discal setae; m ds s, median discal setae; m mg s, median marginal setae; midd dp, middorsal depression.
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1395

m mg s m mg s
tg 3

192 Paradejeania 193 Juriniopsis

194 Epalpus 195 Xanthoepalpus

st 4 st 4

196 Rhachoepalpus 197 Parepalpus

Figs. 113.192–197. Abdomens (concluded): dorsal view of (192) Paradejeania rutilioides (Jaennicke), (MND, fig. 110.240); (193) Juriniopsis
adusta (Wulp), (MND, fig. 110.241, as J. floridensis); (194) Epalpus signifer (Walker), (Nearctic, MND, fig. 110.242); (195) Xanthoepalpus
bicolor (Williston), (MND, fig.  110.243); ventral view of (196)  Rhachoepalpus olivaceus Townsend, (MND, fig.  110.244); and
(197) Parepalpus flavidus Coquillett, (Nearctic, MND, fig. 110.245).
Abbreviations: m mg s, median marginal setae; st, sternite; tg, tergite.
1396
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

251. Abdominal syntergite 1+2 elongate and strongly constricted to form cylindrical petiole; scutellum
with two pairs of long divergent bristles in the lateral and subapical positions (Fig. 137); lower
calypter rudimentary, at least in Central American species. . . . . . . . . . . . Cordyligaster Macquart
– Base of abdomen not petiolate; scutellum usually with three or more pairs of bristles, or if only
two then at least one pair convergent; lower calypter well developed. . . . . . . . . . . . . . . . . . . . 252

252. Median discal bristles of abdominal tergite 3 situated far forward on median lobe that projects into
posterior margin of tergite 1+2 and that superficially appears to be median marginal bristles of
tergite 1+2 (Fig. 187); frons and body metallic bluish-green or greenish-gold, overlaid with thin
whitish pruinosity; legs yellow from coxae to tibiae; tarsi black. . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Argyromima Brauer & Bergenstamm
– Median discal bristles, if present, not displaced forward; coloration of frons and body yellow,
black, or brown, not as above. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 253

253. Face convex, visible in lateral view, at least on lower half (Fig. 50). . . . . . . . . . . . . . . . . . . . . 254
– Face concave at least on upper half, not visible in profile except sometimes at vibrissal angle
(Fig. 53). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 257

254. Midtibia with two or more anterodorsal bristles (Fig. 176). . . . . . . . . . . Trochiloleskia Townsend
– Midtibia with only one anterodorsal bristle (Fig. 175) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 255

255. Postpronotum, proepimeron, most of mesopleuron, and hind coxa with dense tufts of long white
plumose hairs (Fig. 109); meral setae of most species consist of mass of white plumose hairs, or
with white plumose hairs and black bristles . . . . . . . . . . . . . . . . . . . . . . . Telothyria Wulp, in part
– Thoracic hairs not plumose and not especially long or dense. . . . . . . . . . . . . . . . . . . . . . . . . . 256

256. R4+5 setose more than halfway to crossvein r-m (as in Figs. 158, 160); postpronotum with small
anterolateral bristle in addition to three basal bristles. . . . . . . . . . . . . . . . . . . . . . . Genea Rondani
– R4+5 with few setae at base only; postpronotum lacking anterolateral bristle . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leskia Robineau-Desvoidy

257. Prementum long and slender, often curved posteriorly, at least as long as height of eye, and in
most species longer than height of head (Figs. 53, 55); labella minute, pointed, scarcely wider
than apex of prementum. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 258
– Prementum shorter than height of eye; labella normal, padlike. . . . . . . . . . . . . . . . . . . . . . . . . 259

258. Membrane between lower facial margin and clypeus with pair of convex subtriangular sclerites
(Fig. 80); aristomere 2 elongate, 2 or more times as long as wide; first flagellomere truncate,
widest beyond middle (Fig. 53). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Clausicella Rondani
– Membrane between lower facial margin and clypeus without above such sclerites; aristomere 2
minute, no longer than wide; first flagellomere rounded apically, widest before middle (Fig. 55).
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Crocinosoma Reinhard

259. Arista plumose, with longest hairs longer than greatest width of arista (as in Figs. 47, 48, 87). . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 260
– Arista bare, or nearly so, with longest hairs shorter than greatest width of arista. . . . . . . . . . . 261

260. Anterior katepisternal bristle small, less than half as long as posterior katepisternal (Fig. 124);
lateral scutellar bristles long and divergent, larger than apical bristles (as in Fig. 128); abdominal
tergites 1–4 yellow or orange, tergite 5 black . . . . . . . . . . . . . . . . . . . . . . . . . . Xanthodexia Wulp
– Anterior katepisternal bristle more than half as long as posterior katepisternal (as in Fig. 120);
lateral scutellar bristles parallel to each other, shorter than apical bristles (Fig.  136); abdomen
mostly or entirely black, or if yellow basally, most of tergite 4 and all of tergite 5 dark . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetonopsis Townsend
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1397

261. Eye, viewed in profile, occupying almost entire side of head, more than half as long (front to back)
as high, thus reducing parafacial and gena to narrow bands (Fig. 49). . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Strongygaster Macquart, in part
– Eye, viewed in profile, less than half as long (front to back) as high; parafacial, gena and postgena
more extensive (Figs. 25, 28). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 262

262. Scutum with only one, posteriormost, pair of acrostical bristles, situated anterior to scutellum;
lateral bristles of scutellum lacking or scarcely differentiated; male predominantly yellow, with
darker scutum; female mostly yellowish with abdomen black dorsally; antenna of both sexes
orange. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Euclytia Townsend
– Scutum with more than one pair of acrostical bristles; lateral scutellar bristles well differentiated;
both sexes predominantly black. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 263

263. Scutellum nearly as broad at apex as at base, truncate apically, with long, stout lateral bristles and
crossed apical bristles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arrhinactia Townsend
– Scutellum rounded or pointed at apex, with divergent subapical bristles; apical bristles parallel to
one another, or lacking; BLONDELIINI, in part (species with bare prosternum). . . . . . . . . . . 264

264. Lateral scutellar bristles weak or lacking (Fig. 132). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 265

– Lateral scutellar bristles well developed (as in Figs. 130, 131). . . . . . . . . . . . . . . . . . . . . . . . . 268

265. Vibrissa arising at lower corner of head, level with lower cranial margin, which appears straight
(Fig.  25); arista usually pubescent or plumose; abdominal sternite  7 of female sharp, slightly
curved piercer sheathed in spoutlike sternite 6 (Fig. 180). . . . . Phyllophilopsis Townsend, in part
– Vibrissa arising above level of lower cranial margin, which appears convex (Fig. 28); arista bare;
abdomen of female arched ventrally, with telescopic ovipositor directed ventrally, lacking piercer
(usually retracted into abdomen) (Figs. 181, 182). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 266

266. Eye exceptionally large, covering most of side of head; distance between eye and lower cranial
margin less than twice width of palpus (as in Fig. 14); female abdomen globose, rounded apically;
ovipositor projecting ventrally or anteroventrally (Fig. 182) . . . . . . . . . . Calodexia Wulp, in part
– Eye smaller, separated from lower cranial margin by greater than twice width of palpus (Fig. 28);
abdomen of female pointed or truncate apically; ovipositor, when not retracted, directed
posteroventrally. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 267

267. Eye kidney-shaped, more than twice as high as wide, its posterior margin distinctly concave be-
low (Fig. 28); antenna arising below middle of eye; male with 10 or more reclinate frontal bristles;
inner vertical bristles of male undifferentiated; abdomen of female pointed apically. . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trigonospila Pokorny, in part
– Eye more rounded, less than twice as high as wide, its posterior margin scarcely concave (as in
Fig.  21); antenna arising at level of middle of eye (as in Fig.  13); male with fewer than 10,
relatively straight frontal bristles (as in Fig. 25); inner vertical bristles of male well developed (as
in Fig. 26); abdomen of female flattened side to side, truncate apically, dorsal profile of tergite 5
nearly twice as long as tergite 4 (Fig. 181) . . . . . . . . . . . . . . . . . . . . Dolichotarsus Brooks, in part

268. Frons with three or more proclinate orbital bristles (Fig. 65) . . . . . . . . Euhalidaya Walton, in part
– Frons with not more than two proclinate orbital bristles (Fig. 17) . . . . . . . . . Anisia Wulp, in part
1398
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Synopsis of the fauna Adejeania Townsend. The species of this distinctive

Neotropical genus of 38 described species, are mostly read-
Only about 130 species of tachinids have been recorded ily identifiable thanks to the works of Curran (1947) and
from Central America, including 45 described from Costa Guimarães (1966a). A single species, A.  vexatrix (Osten
Rica (Guimarães, 1971b), but this represents a small fraction Sacken), extends north to southern British Columbia, Cana-
of the actual fauna of the region. Nearly 900 were recorded da, while five more (as well as several undescribed species)
from Mexico; the actual fauna of Central America might have been found in Central America. The majority occur in
number as high as 2000. Tachinids are found throughout the Andes and in southern Brazil.
Central America in all habitats, but seem to be most diverse
in cloud forests between 800 and 1500 m. Admontia Brauer & Bergenstamm. This is a widespread
genus, with 10 species in the Palearctic Region (Herting, 1984),
Acantholespesia Wood. This is a New World genus of two Oriental species (Crosskey, 1976), and 11 Nearctic spe-
only three species north of Mexico (O’Hara & Wood, 2004) cies (Wood & Cave, 2006), including two shared with the Old
and two in Guanacaste, Costa Rica, reared from larvae of World. The Neotropical Region has a surprisingly rich fauna
Crambidae (Lepidoptera) (Janzen & Hallwachs, 2005). of more than 17 species (including some currently classified
in other genera), the majority in Patagonia (Aldrich, 1934); at
Acaulona Wulp. Related to Trichopoda Berthold, but least seven species occur in Costa Rica. Hosts of some north-
lacking flattened setae on the hind tibia, this Neotropical ern species are larvae of Tipulidae, although Arnaud (1978)
genus contains three species (Toma, 2003). At least one of has recorded some lepidopterous larvae as well.
these, A.  costata Wulp, ranges from Mexico to Argentina.
Females deposit unembryonated eggs on bugs (Heteroptera) Ametadoria Townsend. This is a small New World ge-
of the families Largidae, Pentatomidae, and Pyrrhocoridae nus of seven described species ranging from USA to Bra-
(Guimarães, 1977c). zil, although only one, A. harrisinae (Coquillett), is known
from the Nearctic Region (O’Hara  & Wood, 2004). Three
Acemya Robineau-Desvoidy. Five Palearctic species morphospecies have been reared from larvae of Lacturidae,
(Herting, 1984), five species north of Mexico (O’Hara  & Zygaenidae, and Papilionidae (Lepidoptera) in Guanacaste,
Wood, 2004), and a single species in Costa Rica, which may Costa Rica (Janzen & Hallwachs, 2005).
be one of these, comprise the rather peculiar genus Acemya.
This genus, together with Ceracia, make up an isolated tax- Angustia Sellers. This genus has three named species:
on, the tribe Acemyini, tentatively assigned to the Tachininae A. pallidipalpis (Wulp) from Mexico and Costa Rica, A. pal-
(O’Hara & Wood, 2004). Hosts are acridid grasshoppers. lens (Reinhard) from Ohio, and the type species A. angus-
tivitta (Aldrich & Webber) from New York, all of which may
Acronacantha Wulp. This is a Neotropical genus related be conspecific. Too few specimens exist to form an opinion,
to Microphthalma and Pyrrhodexia, with a single species, and a male would be helpful. Its placement in Blondeliini is
A. nubilipennis Wulp, known from Costa Rica. problematic.

Actia Robineau-Desvoidy. Actia is a cosmopolitan ge- Anepalpus Townsend. A small Neotropical genus of
nus with eight described species recorded north of Mexico two species, one in Costa Rica, one in Peru, neither read-
(O’Hara, 1991), three from South America, but only two from ily distinguishable from Epalpus, except by male geni-
Central America, probably a small fraction of the fauna there. talia, which are of a type shared with Trichophora and
Hosts include some well-known pests, such as the spruce Xanthoepalpus rather than with the huge genus Epalpus.
budworm (O’Hara, 2005) and other microlepidoptera. Further study of the South American fauna is needed
to find means of separating females and to clarify their
Actinochaeta Brauer  & Bergenstamm. This Neotropical interrelationships.
genus has only four described species, including A. columbiae
Brauer & Bergenstamm, recorded from Panama and Costa Rica Anisia Wulp. Found only in the New World, Anisia is
(Aldrich, 1927b) as well as from Colombia. The total number of a species-rich genus in the Neotropical Region. Their hosts
species in Central America is unknown but probably not large. are various orthopteroid insects. Four species have been re-
corded north of Mexico (O’Hara  & Wood, 2004). At least
Actinodoria Townsend. Wood (1985) listed three spe- 15 morphospecies have been segregated from the INBio
cies of Actinodoria from Mexico and Central America and collections in Costa Rica that could be assigned to Anisia,
a fourth from Trinidad. Males are remarkable for having although lack of distinctive external adult characters makes
the frontal bristles crowded to the edge of the exceptionally this assignment tenuous for some species. Two species have
wide, often metallic frons. The genus does not occur north of been reared from grasshoppers (Orthoptera: Acrididae) (Al-
Mexico. One species was reared from the larva of a skipper drich, 1927a, 1928). One Nearctic species, A.  optata (Re-
of the genus Urbanus (Hesperiidae) in Guanacaste, Costa inhard) has been reared from a cockroach (Gemeno et al.,
Rica (Janzen & Hallwachs, 2005). 2002), and a Neotropical species (as Nepophasmophaga
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1399

Townsend) was recorded by Guimarães (1977c) as a parasi- Atacta Schiner. This distinctive Neotropical genus con-
toid of an earwig (Forficulidae). Females of another Nearctic tains only four species, ranging from southern USA to Bra-
species, A. flaveola (Coquillett), deposit minute, transparent, zil. One of these, A.  brasiliensis Schiner, was reared from
fully embryonated eggs, not on the camel cricket host itself larvae of Noctuidae (Lepidoptera) in Guanacaste, Costa Rica
(Rhaphidophoridae), but on its food, where they are presum- (Janzen & Hallwachs, 2005).
ably eaten by the host (D.M. Wood, personal observation).
This type of oviposition is analogous to the deposition of Atactosturmia Townsend. Four Neotropical species of
microtype eggs on the leaves of food plants found in Goniini, this genus have been recorded from Costa Rica to Paraguay.
and when further investigated this may prove to be the most An additional four unidentified morphospecies have been
convincing shared derived trait of the genus. reared from larvae of Noctuidae and Arctiidae (Lepidoptera)
in Guanacaste, Costa Rica (Janzen & Hallwachs, 2005).
Anoxynops Townsend. A New World genus with three
nominal species, Anoxynops is intimately related to Lepto- Austeniops Townsend. A single species, A.  truncati-
stylum and may prove to be inseparable when further species cornis (Wulp), is known only from the high mountains of
are studied. Six morphospecies have been segregated in the Costa Rica and Chiriqui, Panama. Further study of the South
INBio collection. American species of Trichophora may show that Austeniops
may represent merely an unusual abdominal shape in the ge-
Aplomyopsis Townsend. This small, poorly known New nus Trichophora.
World genus has a single species, A.  polita (Coquillett), from
north of Mexico, and two more unnamed species in Costa Rica. Austrophorocera Townsend. This is a large genus rep-
resented in the Australian and Oriental Regions (Crosskey,
Archytas Jaennicke. This is a large genus found from 1973, 1976) as well as in the New World. Ten species were
southern Canada to Patagonia. Well over 100 names have recorded north of Mexico (O’Hara  & Wood, 2004), and at
been proposed, although many will probably prove to be least 15 species have been described from the Neotropical
synonyms. Guimarães (1960, 1961a,  b, 1963b,  c) revised Region. Many morphospecies have been reared from Guana-
the Brazilian species, while the 12 species occurring north caste, Costa Rica, from the larvae of a variety of limacodid
of Mexico were treated by Ravlin & Stehr (1984). No esti- moths (Janzen & Hallwachs, 2005).
mate of the number of species in Central America has been
attempted. Fourteen morphospecies have been reared from Avibrissosturmia Townsend. This New World genus
larvae of Lepidoptera in the families Noctuidae, Pyralidae, is closely related to Winthemia and ranges from Mary-
Hesperiidae, Notodontidae, Lymantriidae, and Nymphalidae land, USA, to Brazil, with five species in South America
in Guanacaste, Costa Rica (Janzen & Hallwachs, 2005). (Guimarães, 1983a). Four species were reared from larvae
of Megalopygidae (Lepidoptera) in Guanacaste, Costa Rica
Argyrochaetona Townsend. Only two nominal species of (Janzen & Hallwachs, 2005).
this genus have been described, one from Cuba and one from
Peru, but they may be conspecific. An additional species has Belvosia Robineau-Desvoidy. Belvosia is a large
been found in Costa Rica. In Guanacaste, one was reared from New World genus of some 70 described species, ranging
a moth larva of Crambidae (Janzen & Hallwachs, 2005). from Canada to Argentina, including 15 north of Mexico
(O’Hara & Wood, 2004). Nineteen morphospecies have been
Argyromima Brauer  & Bergenstamm. This is a rare reared from lepidopterous larvae of the families Sphingidae,
Neotropical genus known only from high elevations in Ecua- Saturniidae, Notodontidae, Noctuidae, Eupterodidae, and
dor and Costa Rica. The type species, A. mirabilis Brauer & Arctiidae in Guanacaste, Costa Rica (Janzen & Hallwachs,
Bergenstamm, was described from “South America”, and 2005). Guimarães (1977c) listed a similarly wide range of
nothing is known about its biology. lepidopterous families as hosts for 14 more species.

Argyrophylax Brauer & Bergenstamm. This is a small Beskia Brauer  & Bergenstamm. This distinctive
Neotropical genus of six described species ranging from Neotropical genus contains only a single species, B.  ael-
southern Florida to Peru. Four species have been reared from ops (Walker), which ranges from eastern USA to Brazil
Lepidoptera of the families Nymphalidae, Crambidae, Thy- and which parasitizes bugs of the family Pentatomidae
rididae, and Arctiidae in Guanacaste, Costa Rica (Janzen & (Guimarães, 1977c).
Hallwachs, 2005), and Guimarães (1977c) listed Sphingidae
as a host of A. albincisa (Wiedemann). Bibiomima Brauer & Bergenstamm. This rare Neotro-
pical genus apparently has a single species, B.  handlirschi
Arrhinactia Townsend. A small Neotropical genus rang- Brauer  & Bergenstamm, recorded by Guimarães (1971b)
ing from southern Arizona, USA (O’Hara & Wood, 2004) to from Panama to Brazil and recently collected in Costa Rica.
Brazil, Arrhinactia consists of two nominal species. The num- It is notable for its superficial similarity to some species of
ber of species in the Central American fauna is unknown. Plecia Wiedemann (Bibionidae).
1400
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Billaea Robineau-Desvoidy. This widespread genus oc- 1971b). Cockroaches of the genus Periplaneta were listed as a
curs on all continents except Australia, with 17 species in host by Guimarães (1977c). While waiting for hosts to appear,
the Palearctic Region (Herting, 1984), nine north of Mex- they flit from leaf to leaf just above the ants, avoiding attack
ico (O’Hara  & Wood, 2004), five in the Oriental Region while at the same time chasing fleeing cockroaches and prob-
(Crosskey, 1976), and at least 11 nominal species in South ably other orthopterans as well. They are usually accompanied
America, treated in a revision (as the genus Paratheresia by conopids of the genus Stylogaster Macquart as well as by
Townsend) by Guimarães (1977b). The number of species in both sexes of the tachinid Phasia ecitonis (Townsend). Male
the Central American fauna is unknown. Hosts include lar- Calodexia have no association with the ants, but have been
vae of both Lepidoptera, including the sugar cane borer, and found aggregating near streams. Keys to the species were pro-
Coleoptera, mostly Cerambycidae tunneling in dead wood, vided by Curran (1934) and Rettenmeyer (1961).
as well as Curculionidae in plant stems, where they can only
be attacked by the first instar larvae burrowing in the host’s Calolydella Townsend. This is another essentially Neo-
tunnel. tropical genus, with a single Nearctic species, C.  lathami
(Curran), ranging north to southern Canada. The remain-
Blepharipa Rondani. This nearly cosmopolitan genus, ing 14 described species are known from Mexico to Brazil,
most diverse in the Old World, contains nine described spe- but the actual number of species in Central America is not
cies in the New World ranging from USA to Argentina. In known. Females of C. lathami have been observed oviposit-
the Nearctic Region there are two native species plus one Eu- ing on arctiid moth caterpillars, and Guimarães (1977c) listed
ropean species, B. pratensis (Meigen), introduced to combat both Arctiidae and Noctuidae as hosts for a Neotropical spe-
the gypsy moth. Ten morphospecies have been reared from cies. Five morphospecies have been reared from Guanacaste,
larvae of a variety of families of Lepidoptera in Guanacaste, Costa Rica, on larvae of various families of Lepidoptera and
Costa Rica (Janzen & Hallwachs, 2005). an argid sawfly (Janzen & Hallwachs, 2005).

Blondelia Robineau-Desvoidy. Six nominal species of Campylocheta Rondani. This is a medium-sized Holarc-
Blondelia have been listed for the Nearctic Region, but the tic genus also well represented in the Neotropical Region.
genus has not been revised. One or two species extend south O’Hara & Wood (2004) recorded 11 species north of Mexi-
to Brazil, and a single species appears to be present in Costa co, many reared from Pyralidae, Geometridae, and other mi-
Rica. Hosts are larvae of Lepidoptera. crolepidoptera. Nine morphospecies have been reared from
Guanacaste, Costa Rica, on a variety of families of Lepi-
Bombyliomyia Brauer & Bergenstamm. This is a New doptera (Janzen & Hallwachs, 2005).
World genus of mostly black, bristly flies most diverse in
Costa Rica, where more than 20 morpospecies have been Carcelia Robineau-Desvoidy. Carcelia is a large cosmo-
segregated from the INBio collection. Only one, B.  soror politan genus, with about 25 Neotropical species distributed
(Williston), is known north of Mexico, and eight nominal from Mexico to Chile and Argentina and 13 Nearctic species
species have been described from the Neotropical Region. (O’Hara & Wood, 2004), including one intentionally introduced
Although O’Hara (2002) did not include the genus in his from Europe to combat the gypsy moth. Fifteen morphospe-
tribe Polideini, the genus is regarded here as being a mem- cies have been reared from various families of Lepidoptera in
ber of that tribe in spite of autapomorphic modifications of Guanacaste, Costa Rica (Janzen & Hallwachs, 2005).
the male cerci. Some specimens were reared in Guanacaste,
Costa Rica, from saturniid and geometrid moth larvae (Jan- Casahuiria Townsend. This monotypic South American
zen & Hallwachs, 2005). genus, described from Peru, is included here on the basis of a
few specimens collected at Monteverde, Costa Rica.
Borgmeiermyia Townsend. This is a small Neotropical
genus of only five described species (Arnaud, 1963a; Seh- Catharosia Rondani. This distinctive, widespread genus
nal, 1998). Males of the type species B. brasiliana Townsend of small to minute tachinids is most diverse north of Mexico,
and the other named species have elaborate basketlike first where O’Hara  & Wood (2004) recorded five species. Four
flagellomeres, presumably for more effective detection of species are known from the Palearctic Region, where hosts
female pheromones, and the genus is traditionally based are bugs of the family Lygaeidae. Crosskey (1984) stated that
on species having these elaborate male antennae. However, three species were recorded from the Afrotropical Region. A
females (and males of other unnamed species) lack these single unidentified species occurs in Costa Rica.
adornments and instead have simple elongate antennae.
Celatoria Coquillett. Females of one of the two Nearctic
Calodexia Wulp. Confined to the Neotropical Region, fe- species of Celatoria capture their chrysomelid beetle host
males of this distinctive genus of over 40 species have exploit- while it is in flight and puncture it with their gigantic piercer
ed the raiding columns of army ants that chase their cockroach (Bussart, 1937, as the genus Chaetophleps Coquillett). Al-
hosts from the leaf litter (Rettenmeyer, 1961). At least 15 spe- though members of the genus occur from the USA to Argen-
cies have been recorded from Central America (Guimarães, tina, only five have been named, specific characters have not
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1401

been elucidated, and the number of species in the Central well represented in Central America. Similar to Lixophaga,
American fauna is unknown. the females possess a globose abdomen, suggesting that they,
too, may deposit fully embryonated eggs in webbing, frass,
Ceracia Rondani. This is a cosmopolitan genus but with leaf mines, or similar situations where host contact can only
only a few species recorded from each continent. Four spe- be accomplished by the newly hatched first instars.
cies have been described from the Neotropical Region, but
the number of species in Central America is not known. Chetogena Rondani. A large cosmopolitan genus (if one
includes Spoggosia Rondani and Stomatomyia Brauer  &
Ceromya Robineau-Desvoidy. Ceromya is a small cos- Bergenstamm), Chetogena has 16 nominal species north of
mopolitan genus in the tribe Siphonini, with eight species
Mexico (O’Hara & Wood, 2004) and at least 25 nominal spe-
north of Mexico (O’Hara, 1994) and two (described in Actia)
cies in the Neotropical Region. Eight morphospecies were
in Chile (Aldrich, 1934), in addition to many undescribed
reared from larvae of a variety of families of Lepidoptera in
Neotropical species. An undetermined number of species
Guanacaste, Costa Rica (Janzen & Hallwachs, 2005).
have been reared from larvae of noctuid and notodontid
moths in Guanacaste, Costa Rica (Janzen  & Hallwachs, Chlorohystricia Townsend. This is a Neotropical genus
2005). with only two described species ranging from Mexico to
Chaetogaedia Brauer  & Bergenstamm. A New World Brazil. A third species was reared from larvae of large Hes-
genus of 14 described species ranging from southern Canada periidae at high elevations in Guanacaste, Costa Rica (Jan-
to Argentina, Chaetogaedia includes seven north of Mexico zen & Hallwachs, 2005). Unpublished studies of its genetic
(O’Hara & Wood, 2004) and an undetermined number from sequences suggest that it may be merely a large spiny mem-
Central America. Five morphospecies have been reared from ber of the genus Chrysotachina.
larvae of Lepidoptera of the families Geometridae, Noctui- Choeteprosopa Macquart. This is a small Neotropical
dae, Arctiidae, and Crambidae in Guanacaste, Costa Rica genus of nine described species ranging from Mexico to Bra-
(Janzen & Hallwachs, 2005). zil, probably closely allied to and perhaps congeneric with
Chaetodoria Townsend. A single undetermined species Actactosturmia Townsend. At least one unidentified species
of this small Neotropical genus of two nominal species has is common in Costa Rica.
been collected in southern Costa Rica. Cholomyia Bigot. This predominantly Neotropical ge-
Chaetona Wulp. Another genus confined to the Neotropi- nus has a single Nearctic species, C.  inaequipes Bigot,
cal Region, most specimens of Chaetona have been collected and several Central American species, including C.  lutei-
at lights at night, along with their close relatives Zaira and pennis (Curran) from Panama. Hosts are larvae of weevils
Cryptomeigenia. Females also share with these two genera (Curculionidae) embedded in living plant material, parasit-
large, hard-shelled, fully embryonated eggs that are probably ized by searching first instar larvae that must enter from out-
attached to crepuscular beetles. Although only six species side the plant and penetrate the frass blocking the entrance
have been named, the genus is much larger, judging from the hole. This behavior is probably homologous to that found
INBio collection. Males have been collected in Costa Rica in Gnadochaeta Macquart, whose species also attack weevil
aggregating near a stream during light rain (N. Woodley, per- larvae in acorns and other nuts.
sonal communication), an unusual habit among Tachinidae. Chrysoexorista Townsend. This is a New World genus of
Chaetonodexodes Townsend. This is a New World genus 11 nominal species ranging from southwestern USA to Ar-
with three described species: (i) C. vanderwulpi Townsend, gentina. Five were described from North America and the re-
north of Mexico, (ii) C. marshalli Aldrich, in Trinidad, and mainder from the Neotropical Region, where there are many
(iii) C. rafaeli Townsend, from Mexico. An additional unde- more undescribed species, but the genus has never been
termined species has been found in Central America. Hosts revised so the number of species in the Central American
are chrysomelid beetles (Guimarães, 1977c). fauna is unknown. The characteristic metallic green or gold
color of living and freshly killed specimens gradually turns
Chaetonopsis Townsend. This is a small genus with to brown as they dry out. Six morphospecies were reared
a single named species, C.  spinosa (Coquillett), in North from lepidopterous larvae of Crambidae, Lymantriidae,
America, as well as an unnamed species extending south into and Geometridae in Guanacaste, Costa Rica (Janzen  &
Central and South America. Hallwachs, 2005).
Chaetostigmoptera Townsend. Members of this genus Chrysotachina Brauer & Bergenstamm. This large New
are among the smallest Tachinidae and are collected almost World genus has about 30 nominal species, most of them in
exclusively using Malaise traps. Three species have been de- the Neotropical Region. Nunez et al. (2002) and Nunez  &
scribed from the Nearctic Region, but over a dozen have been Couri (2002) treated the South American species. At least 20
segregated from INBio material, suggesting that the genus is morphospecies have been reared in Guanacaste, Costa Rica,
1402
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

mostly from larvae of Hesperiidae, but with some from larvae reared from crepuscular or nocturnal scarab beetles, and the
of other Lepidoptera of the families Nymphalidae, Riodini- Central American species might also attack scarabs. Females
dae, and Mimallonidae. O’Hara (2002) recorded Hesperii- deposit hard-shelled, fully embryonated eggs on the beetles,
dae as hosts for two of the five Nearctic species. Although as females of species of Zaira do on carabid beetles. Females
some species, including the type species, are bright metallic of several Old World genera, including Istochaeta aldrichi
green, many are non-metallic and may be scarcely separable (Mesnil) introduced into North America from Japan, deposit
from Exoristoides and other polideine genera. similar eggs on beetles. The seventh sternites of some Cryp-
tomeigenia females suggest that they are adapted for prying
Cinochira Zetterstedt. Known only from a single species up the elytra of their hosts.
in Europe, another in Texas, and a third undetermined spe-
cies in Costa Rica, these tiny flies have probably been widely Cryptopalpus Rondani. Although Guimarães (1971b)
overlooked. Females share with Leucostoma and Clairvillia listed nearly a dozen species in this South American genus,
the pincerlike cerci that they presumably use to grasp the all proposed by nineteenth century authors, all but two or
edge of their host’s abdomen. three have been assigned to other tachinine genera. Only one
undetermined species occurs in Central America.
Clairvillia Robineau-Desvoidy. With three Neotropical
species (including one in Costa Rica), two Palearctic spe- Cylindromyia Meigen. This is a large, uniform, eas-
cies, three Nearctic ones, and a single Afrotropical species, ily distinguished cosmopolitan genus of over 100 species,
this small genus is closely related to, and in the Neotropi- of which 17 have been recorded north of Mexico (Aldrich,
cal Region perhaps not separable from, Leucostoma. Bugs 1926a; O’Hara & Wood, 2004), 9 from the Oriental Region,
of the family Coreidae have been recorded as the host of the 28 from the Palearctic Region (Herting, 1984), 22 from
European species. the Afrotropical Region (Crosskey, 1984), 16 from Austra-
lia (Crosskey, 1973), and 24 from the Neotropical Region
Clausicella Rondani. With only three Neotropical spe-
(Guimarães, 1976b). Hosts are bugs of the family Pentatomi-
cies and eight species north of Mexico (O’Hara  & Wood,
dae (Guimarães, 1977c). The number of species in the Cen-
2004), this small genus is nevertheless widespread with
tral American fauna, however, is unknown.
four species recorded from the Palearctic Region (Herting,
1984). A single unidentified species is known from Central Cyrtophleba Rondani. This is a small genus of three
America. Nearctic species and nine Neotropical ones, occurring
Copecrypta Townsend. A sister taxon of Deopalpus, this from Canada to Chile (as the genus Opsophagus Aldrich).
genus also ranges from southern Canada to Argentina, with Guimarães (1977c) recorded Noctuidae (Lepidoptera) as a
a single Nearctic species, C.  ruficauda (Wulp), that may host. A single species, C. horrida Giglio-Tos, occurs in Cen-
also extend south into Central America. Only three other tral America.
names, all Neotropical, may be assigned to Copecrypta, and
Cyzenis Robineau-Desvoidy. This genus is included here
the genus is doubtfully distinct from Deopalpus. A species
on the basis of a single female that appears to be related to
in Guanacaste has been reared from larvae of noctuid and
C. ustulata (Reinhard) collected at 2700 m in the Talamanca
geometrid moths (Janzen & Hallwachs, 2005).
Range of Costa Rica. O’Hara & Cooper (1992) revised the
Cordyligaster Macquart. Cordyligaster is a New World five Nearctic species and listed larvae of both geometrid and
genus with only seven species. The single Costa Rican spe- tortricid moths as hosts.
cies, probably C. petiolata (Wiedemann), was reared from a
Dejeaniops Townsend. This is a small Neotropical genus
larva of a crambid moth (Janzen & Hallwachs, 2005), and a
of three nominal species ranging from Central America to
pyralid was listed as a host by Guimarães (1977c). The most
Peru. The Peruvian species, D. ollachea Townsend, is appar-
recent systematic treatment is that of Guimarães (1971a).
ently also found in Central America.
Crocinosoma Reinhard. The single species of this New
Deopalpus Townsend. Deopalpus is a large genus ex-
World genus, C. cornuale Reinhard, ranges from Florida to
Costa Rica. tending from Canada to Patagonia, with 11 Nearctic species
(O’Hara & Wood, 2004) and more than 30 nominal species
Cryptomeigenia Brauer  & Bergenstamm. Although south of the USA that ought to be assigned to it, although
there are 14 described species of this New World genus this number includes some apparent synonyms. A clear dis-
north of Mexico (O’Hara & Wood, 2004) but only four from tinction between Deopalpus and Copecrypta (both proposed
south of the USA, material in the INBio collection sug- by Townsend in the same paper) may not really exist, but this
gests that many more remain to be described from Central must await a study of all species. Earlier treatments of the
America. Furthermore, separation from Chaetona and Zaira complex are those of Reinhard (1934a) for the Nearctic Re-
may prove to be difficult or impossible, although their hosts gion and of Guimarães (1963a, 1964) for Brazil. The number
may be different. Several of the Nearctic species have been of species in the Central American fauna is unknown.
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1403

Distichona Wulp. The eight Neotropical and three Nearc- from Mexico to Peru. Nothing is known about their biol-
tic species of this genus range from southern Canada to Peru ogy. The number of species in the Central American fauna
and Brazil. One named species, D. petiolata (Townsend), and is unknown.
at least four other morphospecies occur in Central America.
Euantha Wulp. This is a small New World genus of
Dolichocodia Townsend. This is a small New World ge- spectacularly patterned flies, with a single Nearctic species,
nus of six species ranging from southwestern USA to Chile. E. litturata (Olivier), that extends south to Central and South
The Costa Rican fauna consists of four morphospecies, of America (O’Hara & Wood, 2004), along with at least three
which one, D. pretiosa Curran, was previously placed in the others south of the USA.
genus Prorhynchops Brauer & Bergenstamm. The remainder
are probably unnamed. Eucelatoria Townsend. One of the largest genera of
Blondeliini, this genus is confined to the New World, with
Dolichotarsus Brooks. This is a small New World ge- 52 nominal species listed from North and Central America
nus of only three species north of Mexico (O’Hara & Wood, and the West Indies (Wood, 1985). A comprehensive revi-
2004) and an additional one, D.  stipatus Reinhard, which sion will certainly add many more as well as create more
may be conspecific with D. livescens Reinhard, from Mexi- synonymy. Only 11 species have been recorded north of
co to Costa Rica. An undescribed species was reared from a Mexico (O’Hara & Wood, 2004). A small part of the genus,
weevil, Sciopithes obscurus Horn (Wood, 1985). Eucelatoria sensu stricto, was revised by Sabrosky (1981).
At least 25 more nominal species have been recorded from
Drino Robineau-Desvoidy. Drino is a large cosmopoli- South America, and there are a huge number of undescribed
tan genus particularly diverse in the Old World and with at species. Females of all species have sternite  7 modified to
least 30 species in the New World ranging from Canada to form a sharp curved piercer for injecting their embryonated
Brazil. Synonymy of the New World species, of which seven eggs into the host, a character state shared with Blondelia,
were recorded by O’Hara & Wood (2004), is complex and Vibrissina, the Nearctic genus Opsomeigenia Townsend, and
the infamous Compsilura concinnata (a European species
confused. All 17 morphospecies reared in Guanacaste, Costa
introduced to North America to counter the gypsy moth and
Rica, were reared exclusively from larvae of sphingid moths
which is now accused of decimating the native saturniid moth
(Janzen & Hallwachs, 2005). populations). Hosts of Eucelatoria, in most known cases, are
Ebenia Macquart. This is an entirely Neotropical genus larvae of Lepidoptera, although a few species also parasitize
chrysomelid beetle larvae (Guimarães, 1977c). Hosts of the
of small black dexiine flies, with at least 10 undetermined
great majority are unrecorded. From Guanacaste, Costa Rica,
species in Costa Rica. One species was reared in Guanacaste 11 species have been reared from larvae of both Lepidoptera
from a larva of chrysomelid beetle, subfamily Hispinae (Jan- and Chrysomelidae, as well as one from an exposed, leaf-
zen & Hallwachs, 2005). feeding larva of Curculionidae (Janzen & Hallwachs, 2005).
Epalpus Rondani. One of the best known Neotropical Eucheirophaga James. Although only one Mexican spe-
representatives of the Tachinini, and also one of the largest cies, E.  lugubris James, is named, others that apparently
Neotropical genera, Epalpus has at least 30 morphospecies belong to this Neotropical genus have been found in Costa
in Costa Rica and over 50 nominal species described from Rica.
South America. Many of these names were proposed by
early authors whose type material has not been located or Euclytia Townsend. Euclytia is a distinctive, predomi-
studied. Three species range north of the USA border: E. sig- nantly Nearctic genus with apparently a single described
nifer (Walker), a common, well-known species distributed species, E. flava (Coquillett). The genus ranges from south-
across USA and southern Canada, E. rufipes (Brooks) known ern Canada to Costa Rica.
from British Columbia and western USA, and E. albomacu-
latus (Jaennicke) in the southwestern USA ranging south to Eudejeania Townsend. Confined to the Neotropical Re-
Guatemala and perhaps to Costa Rica. The genus reaches its gion, most of the 21 species of this genus are found in cloud
greatest diversity in the Andes at mid-elevations. In spite of forests of the high Andes and are readily identifiable using
their abundance, only one species has been reared in Costa the key of Sabrosky (1947). Two, possibly three, species
Rica from a larva of a notodontid moth in Guanacaste (Jan- occur in Central America. Adults are black with orange or
zen & Hallwachs, 2005). silvery legs and are among the largest tachinids. They may
be found from southern Mexico to northern Argentina. An
Eribella Mesnil. With two Nearctic and one Mexican early record from Mexico, E. pallida Robineau-Desvoidy, is
species, this small New World genus is also present in the doubtfully a member of this genus.
highlands of Costa Rica. Hosts include larvae of pterophorid
moths (Sabrosky & Arnaud, 1965). Eudexia Brauer & Bergenstamm. This small Neotropi-
cal genus is closely related to Hystricodexia and Ptilodexia
Erythromelana Townsend. The six described species of and perhaps not separable from them. A single species occurs
this genus are restricted to the Neotropical Region, occurring in Central America, probably E. dreisbachi Reinhard, which,
1404
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

like the species of Ptilodexia, is presumably parasitic on Chrysops Meigen. The genus has long been segregated in
larvae of scarabaeid beetles in the soil. a tribe of its own, Eutherini, with a few species on every
continent and at least one in Central America that may be
Euepalpus Townsend. This small genus consists of conspecific with one of the three Nearctic species. Hosts of
only two, perhaps only a single, Neotropical species, one the few species that have been reared are bugs of the fam-
of which, E. flavicauda Townsend, has been reared from a ily Pentatomidae (Sabrosky, 1965), and because of this habit
noctuid moth larva in Guanacaste, Costa Rica (Janzen  & they were formerly classified with the Phasiinae. However,
Hallwachs, 2005). the male terminalia and retention of embryonated eggs in a
uterus indicate that they belong in Dexiinae (Cantrell, 1983;
Euhalidaya Walton. This small New World genus has a Tschorsnig, 1985; Shima, 1989).
single described species, E. genalis (Coquillett), found north
of Mexico. An additional species is found in Chile, another in Eutrixa Coquillett. This small crepuscular New World
Brazil, a third in Mexico, and a fourth in Peru. Three uniden- genus has only two species described from north of Mexico
tified species have been discovered in the INBio collection (O’Hara & Wood, 2004) and a single species in Puerto Rico.
in Costa Rica. The Nearctic species has been reared from The specimens from Costa Rica in the INBio collection have
the walking stick, Diapheromera femorata (Say). Females not been identified. Females have a small delicate piercer
run over the dorsal surface of the walking stick, depositing that they must use to insert their eggs into adult scarab bee-
embryonated eggs on the cuticle. Larvae hatch immediately tles. Along with Isidotus, they are the only North American
and burrow into the host. representatives of the cosmopolitan tribe Palpostomatini.
Eujuriniodes Townsend. This New World genus is Exoristoides Coquillett. This small New World genus
doubtfully separable from Archytas and contains only a has only three species north of Mexico (O’Hara, 2002) but a
single described species, E.  eva Townsend, from Trini- much larger number of undescribed or unidentified Neotro-
dad. Two species apparently belonging to this genus were pical species, including some from Central America. O’Hara
reared from butterfly larvae of families Nymphalidae and (2002) recorded a cockroach and a cricket as hosts of two
Hesperiidae in Guanacaste, Costa Rica (Janzen & Hallwachs, Nearctic species. Exoristoides and Chrysotachina appear to
2005). be sister taxa and may not be separable in the Neotropical
Region.
Eulasiona Townsend. This is a small New World genus
of perhaps eight species north of Mexico and two more un- Frontiniella Townsend. This is a small New World genus,
identified species from Central and South America. mostly confined to North America. O’Hara (1993) recorded
an unnamed specimen of this New World genus from Chia-
Eulasiopalpus Townsend. A distinctive Neotropical ge- pas and listed larvae of a variety of families of Lepidoptera
nus of 11 species ranging from Mexico to Peru, Eulasiopal- as hosts of the 10 Nearctic species of this genus. A single
pus is found mostly in the Andes at mid-elevations. At least male from Monteverde, Costa Rica, appears to be a member
one species, E. gertschi Curran, occurs in Central America. of the F. festinans species group, although the scutellum ap-
pears to have four discal setae in a transverse row.
Eulobomyia Woodley & Arnaud. This is a recently de-
scribed genus with several species extending from Central Gaediopsis Brauer  & Bergenstamm. This New World
America to southern Brazil (Woodley & Arnaud, 2008a, b). genus of 15 nominal species ranges from southern Canada
One species has been reared from a geometrid moth (Wood- to Peru, with the majority occurring in Mexico. Nine spe-
ley & Arnaud, 2008a). cies are known from the Nearctic Region (O’Hara & Wood,
2004). At least a dozen species occur in Central America,
Eumea Robineau-Desvoidy. This small Holarctic genus including G. rufescens Aldrich.
is included on the basis of a single rearing of an unidenti-
fied species from Guanacaste, Costa Rica, from a larva of Genea Rondani. Genea is a small New World genus of
the family Crambidae (Janzen & Hallwachs, 2005). In North 10 species in the Neotropical Region and seven species north
America, E. caesar (Aldrich) is a well-known parasitoid of of Mexico (O’Hara  & Wood, 2004). Three morphospecies
the spruce budworm (O’Hara, 2005) and other notorious de- have been reared from moth larvae of families Thyrididae,
foliating species in the genus Choristoneura (Lepidoptera: Gelechiidae, and Crambidae in Guanacaste, Costa Rica (Jan-
Tortricidae). zen & Hallwachs, 2005).

Euscopoliopteryx Townsend. This is another distinctive Ginglymia Townsend. This small New World genus has
Neotropical genus of Phasiinae, apparently with a single spe- two species north of Mexico (O’Hara  & Wood, 2004) and
cies, E. externa (Fabricius), that ranges north to Costa Rica. four species from the Neotropical Region. Based on differ-
ences in the fifth sternite of the male, however, there appear
Euthera Loew. Euthera is a small, distinctive, cosmo- to be six morphospecies in Costa Rica, all belonging to the
politan genus, superficially resembling tabanids of the genus subgenus Lasioneura Coquillett.
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1405

Gnadochaeta Macquart. This New World genus of Homogenia Wulp. Homogenia is a Neotropical genus of
parasitoids of larvae of weevils (Curculionidae) is repre- 10 species, closely related to and perhaps doubtfully distinct
sented in the Nearctic Region by 17 species (O’Hara  & from Acaulona. It is differentiated from Acaulona by posses-
Wood, 2004), while only 14 have been recorded from south sion of a row of bladelike setae on the hind tibia, but in some
of Mexico. The number of species in Central America is species these setae are small to lacking. The presence of
unknown. these setae is a character state also shared with Trichopoda.
This fringe simulates the comb on the hind tibia of a melo-
Gonia Meigen. Gonia is a large predominantly Hol-
ponine bee as well as the leaflike flange on the hind tibia
arctic genus with 28 Nearctic species (O’Hara  & Wood,
of a coreid bug (Hemiptera). Males of some species of Ho-
2004) and 22 Palearctic species (Herting, 1984). Only
seven nominal species (probably representing only two or mogenia have conspicuous orange abdomens and fly slowly
three actual species) extend south into Central America about in forest clearings, where they appear to resemble co-
and thence to Patagonia, mostly in open or arid areas. reids. The presence of eggs on males collected while flying
A single species is also known from Africa (Crosskey, about in clearings in Guanacaste, Costa Rica, which appear
1984). Hosts include larvae of various noctuid moths to be identical to eggs extracted from conspecific females,
(Guimarães, 1977c). suggests that males behave this way to attract the attention
of prospective mates by imitating their hosts, thus arousing
Gymnoclytia Brauer  & Bergenstamm. This is a New their host-seeking behavior, even risking being accidentally
World genus with five species recorded north of Mexico oviposited on in the process (D.M. Wood, personal obser-
(O’Hara & Wood, 2004) and nine in the Neotropical Region. vations). Females normally deposit unembryonated eggs on
Central American specimens have been collected but not yet bugs of the family Pyrrhocoridae (Guimarães, 1977c).
identified.
Houghia Coquillett. A complex New World genus of
Gymnomma Wulp. This Neotropical genus of 11 nominal about 30 named Neotropical species, Houghia is distrib-
species ranging from Mexico to Argentina has the habitus of uted from Mexico to Brazil, mostly confined to lowlands.
Archytas, but lacks palpi. At least two unidentified species Five species occur in the Nearctic Region, three of which
occur in Central America. reach southern Canada (O’Hara & Wood, 2004). Twenty-six
morphospecies have been reared from larvae of a wide vari-
Gymnosoma Meigen. This nearly cosmopolitan genus ety of Lepidoptera, the majority on Hesperiidae and Nymph-
is included on the basis of the occurrence of G.  ruficornis alidae, in Guanacaste, Costa Rica (Janzen  & Hallwachs,
(Wulp) in Tabasco, southern Mexico. Only one other spe- 2005).
cies has been recorded from the Neotropical Region, from
Chile (Cortés  & Campos, 1971), suggesting that members Hyalurgus Robineau-Desvoidy. This small Holarctic ge-
of the genus may occur elsewhere in South America. Two nus of sawfly parasitoids has two native and one introduced
species have been recorded from the Afrotropical Region European species in North America (O’Hara & Wood, 2004).
(Crosskey, 1984), five species from the Oriental Region Its presence in Central America is based on a single speci-
(Crosskey, 1976), and 18 species from the Palearctic Region men collected in Costa Rica.
(Herting, 1984). Six species are listed from north of Mexico
(O’Hara  & Wood, 2004). Females deposit unembryonated Hyphantrophaga Townsend. A New World genus of
eggs on pentatomid bugs. about 15 species occurring from Canada to Peru and Brazil,
Hyphantrophaga includes nine species in the Nearctic Re-
Hemisturmia Townsend. This small New World ge- gion (O’Hara & Wood, 2004). Some 18 morphospecies have
nus has only seven species, two in the Nearctic and five
been reared in Guanacaste, Costa Rica (Janzen & Hallwachs,
in the Neotropical Region (Guimarães, 1983a), distrib-
2005). Two of these, H. blanda (Osten Sacken) and H. viri-
uted from Mexico to Brazil. The most common Nearctic
lis (Aldrich & Webber), also found in North America, attack
species, H. parva (Bigot) (synonym H. tortricis (Coquil-
larvae of a wide variety of Lepidoptera. The other 16 Guana-
lett)), is a parasitoid of the spruce budworm (O’Hara,
caste species were reared from larvae of Nymphalidae, Noc-
2005) as well as other colonial web-spinning tortricids.
In Guanacaste, Costa Rica, H.  tenuipalpis (Wulp) was tuidae, Sphingidae, Apatelodidae, and Crambidae (Janzen &
reared from larvae of a variety of Lepidoptera (Janzen & Hallwachs, 2005).
Hallwachs, 2005). Hypotachina Brauer & Bergenstamm. This New World
Hemyda Robineau-Desvoidy. This small genus has a genus, with apparently only a single species, H. chrysophora
single Nearctic species, H. aurata Robineau-Desvoidy, that (Wiedemann), has been reared from several different fami-
probably extends south to Central America and three addi- lies of Lepidoptera in Guanacaste, Costa Rica (Janzen  &
tional Neotropical species. Herting (1984) has also included Hallwachs, 2005). Although Hypotachina was regarded by
in Hemyda two European species that were formerly placed Herting (1984) as a synonym of the Palearctic genus Nemo-
in Evibrissa Rondani. The number of species in Central raea Robineau-Desvoidy, the two are here considered con-
America is unknown but there might be only one. tribal but not congeneric.
1406
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Hystricodexia Röder. This is a small genus of large, Itacuphocera Townsend. A Neotropical genus of eight
mostly undescribed dexiine flies, confined to the Neotropical nominal species (Guimarães, 1963a, 1964), they range from
Region, closely related to Ptilodexia. Like that genus they Central America to southern Brazil. Several species appar-
presumably also parasitize grubs of scarabaeid beetles in the ently occur in Central America but have not yet been studied;
soil. One species, H. echinata (Wulp), was described from adults hold their wings more tightly closed than most tachin-
Costa Rica, but there are several more unnamed species in ids and thus resemble beetles to some extent.
Central America.
Italispidea Townsend. Restricted to the Neotropical Re-
Hystricia Macquart. This New World genus has over gion, Italispidea is a small obscure group of only four named
30 named species, most in the Andes. In Guanacaste, Costa species, still awaiting a more precise definition delineating
Rica, two species, among a total of 10 known from Costa it from Lixophaga and other small blondeliines. Nothing is
Rica, were reared from moth larvae of families Saturniidae known about its hosts or habits, nor the number of species in
and Arctiidae (Janzen & Hallwachs, 2005). O’Hara (2002) Central America.
listed the Neotropical species that had been assigned to Hys-
tricia and also recorded Saturniidae and Arctiidae as hosts of Itaplectops Townsend. A single described species, I. an-
H. abrupta (Wiedemann), one of the two species found north tennalis Townsend, is known from Brazil. One unidentifed
of Mexico. Although not related to the tribe Tachinini (they species was reared from larvae of Limacodidae (Lepi-
are in the tribe Polideini), adults of Hystricia resemble those doptera) in Guanacaste, Costa Rica (Janzen  & Hallwachs,
2005), supporting its suspected relationship to Uramya and
of Epalpus and other large bristly cloud forest tachinine gen-
Thelairaporia, which are also limacodid parasitoids.
era in size, shape, and color patterns.
Jurinella Brauer  & Bergenstamm. A large predomi-
Icelia Robineau-Desvoidy. Icelia is a small New World
nantly Neotropical genus, with over 50 names assigned to
genus of only six species in the Neotropical Region. The Bra-
it, Jurinella ranges from western USA to Argentina. North
zilian species were treated by Guimarães (1976a). Only one of Mexico it is represented by a single species, J. lutzi Cur-
species, I. triquetra (Olivier), ranges north of Mexico. Hosts ran, in southwestern USA, but there are an undetermined
of the genus include larvae of pyralid moths (Guimarães, number of species in Mexico and Central America. Curran
1977c). The number of species in Central America is (1947) treated some of the South American species. Seven
unknown. morphospecies have been reared from lepidopterous larvae
of Noctuidae, Hesperiidae, Notodontidae, and Geometridae,
Ichneumonops Townsend. A single rare species, I. mira- in Guanacaste, Costa Rica (Janzen & Hallwachs, 2005).
bilis Townsend, described from New Mexico, may also range
south to Costa Rica. Like its apparent sister genus, Polisti- Jurinia Robineau-Desvoidy. This is a Neotropical genus
opsis, it is a convincing mimic of vespid wasps of the ge- of about 20 named species, most of whose identities are un-
nus Polistes. Inclusion of Ichneumonops in Cylindromyia certain because of inconsistent early use or misuse of the
by O’Hara  & Wood (2004) was based only on superficial name Jurinia. One species, J.  pompalis (Reinhard), ranges
similarity. north to southern Canada, a second species is found in Flori-
da, while the remainder range from Mexico to southern Bra-
Isidotus Reinhard. This genus has only one species, I. in- zil. The number of species in Central America has not been
canus Reinhard, found from Arizona to Oaxaca (O’Hara & ascertained.
Wood, 2004); the same or a similar species extends south
to Costa Rica. It appears to be crepuscular, as is its clos- Juriniopsis Townsend. Occurring from eastern USA to
est relative, Eutrixa, and like members of that genus may Peru, this is a small genus of only 10 species. Four are re-
also parasitize adult scarabs. These two genera, along with corded from the Nearctic Region (O’Hara  & Wood, 2004)
the Patagonian genus Xanthobasis Aldrich (and some gen- and only one from Peru; the rest are found in Mexico and
era proposed by Blanchard that are probably congeneric Central America. Some of the species were reviewed by Cur-
with Xanthobasis), are the only New World representatives ran (1960) and Sabrosky (1969). Juriniopsis adusta (Loew)
currently assigned to a more extensive Old World tribe, was reared from moth larvae of Arctiidae and Saturniidae in
Palpostomatini. Guanacaste, Costa Rica (Janzen & Hallwachs, 2005).
Ischyrophaga Townsend. This is a rare Neotropical ge- Kirbya Robineau-Desvoidy. This Holarctic genus has
nus of only two nominal species: I. ischyri (Couillett) from five species in North America (O’Hara  & Wood, 2004). A
Cuba and I. polita (Townsend) (formerly in Pseudochaetona single specimen is known from Guanacaste, Costa Rica.
Townsend) from Panama. They appear to be related to Phyl-
lophilopsis, which has been reared from Endomychidae Lasiona Wulp. This genus was proposed for a single dis-
(Coleoptera). The type specimen of I.  ischyri was reared tinctive species, L. multisetosa Wulp, known only from the
from an erotylid beetle in Cuba. cloud forest of the Cordillera Central, Costa Rica.
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1407

Leptostylum Macquart. An entirely Neotropical genus, also been recorded as hosts (Guimarães, 1977c), this record
seldom collected (many species in Costa Rica are repre- needs confirmation.
sented by single males), but apparently quite diverse, Lep-
tostylum is arbitrarily separated for convenience here from Lindigepalpus Townsend. Another poorly understood
Anoxynops only by length and position of the lateral scutellar South American genus, with a single described species
bristles. Leptostylum leuconotum (Wulp) ranges north as far based on a misidentified type specimen, members of this
as the cloud forests of Sinaloa, Mexico. At least 10 species, genus are superficially similar to those of Epalpus, and only
including L. leuconotum (which may be conspecific with the males can be separated with certainty. Its relationships with
Brazilian type species L. pulchellum Macquart), have been Anepalpus and Austeniops, both genera that share nearly
found in the collection of INBio. In Guanacaste, Costa Rica, identical male terminalia, require further study. One spe-
six species have been reared, mostly from larvae of Saturnii- cies occurs in the Talamanca Mountains of southern Costa
dae, Nymphalidae, and some other families of Lepidoptera Rica.
(Janzen & Hallwachs, 2005).
Linnaemya Robineau-Desvoidy. Linnaemya is a
Leschenaultia Robineau-Desvoidy. This is a New World large cosmopolitan genus, particularly diverse in the Old
genus of 32 named species (Toma  & Guimarães, 2002), World, where Crosskey (1984) recorded 60 species from
ranging from southern Canada to Brazil, including 12 spe- the Afrotropical Region and Herting listed 52 species in
cies north of Mexico (O’Hara  & Wood, 2004). Twenty-six six subgenera from the Palearctic Region. By comparison,
morphospecies have been reared from lepidopterous larvae the New World is depauperate; O’Hara  & Wood (2004)
of Sphingidae, Saturniidae, Megalopygidae, Arctiidae, Noc- cataloged only six Nearctic species and there are only five
tuidae, Nymphalidae, and Lymantriidae in Guanacaste, Cos- named Neotropical species. One species was reared from
ta Rica (Janzen & Hallwachs, 2005). a notodontid caterpillar in Guanacaste, Costa Rica (Jan-
zen & Hallwachs, 2005).
Leskia Robineau-Desvoidy. Leskia is a Holarctic and
Neotropical genus with three species in the Palearctic Re- Lixadmontia Wood  & Cave. Intermediate between Ad-
gion (Herting, 1984), three north of Mexico (O’Hara  & montia and Lixophaga Townsend, this genus was recently
Wood, 2004), and as many as 18 in the Neotropical Re- created for a parasitoid of a weevil, Metamasius quadrilin-
gion, depending on the outcome of a revision in progress
eatus Champion, that infests bromeliads in tropical montane
by E.  Nunez. In Guanacaste, Costa Rica, one species
cloud forests of Honduras and that was accidentally intro-
(among a much larger but unresolved total) was reared
from larvae of Thyrididae and Crambidae (Janzen  & duced into Florida (Wood & Cave, 2006).
Hallwachs, 2005).
Lixophaga Townsend. This is the third largest New World
Lespesia Robineau-Desvoidy. This is one of the largest genus of Blondeliini after Myiopharus and Eucelatoria and
New World genera, ranging from Canada to Chile, attacking will probably prove to be the most difficult taxonomically
the exposed larvae of a variety of caterpillars of several fami- because most of the species, especially females, resemble
lies, particularly Noctuidae, Notodontidae, Saturniidae, and one another closely. With 48 nominal species in North and
Sphingidae. At least 50 species in South America should be Central America and the West Indies (Wood, 1985), 16 of
included in the genus but are now dispersed among smaller these were listed north of Mexico (O’Hara & Wood, 2004).
genera. Guimarães (1983b) assembled some of these in his The number of species in South America is impossible to
treatment of the Brazilian species. O’Hara & Wood (2004) estimate at this time, but appears to be large. There are also
recorded 32 species north of Mexico. In Guanacaste, Costa three species in Japan and one, L. sphenophori (Villeneuve),
Rica, 35 morphospecies have been reared from larvae of a in New Guinea that has been widely introduced in the Pa-
wide array of families of Lepidoptera (Janzen & Hallwachs, cific and established in Hawaii (Crosskey, 1973). Females of
2005). Lixophaga whose life cycle has been investigated attack hid-
den hosts, usually larvae of Lepidoptera, e.g., the sugar cane
Leucostoma Meigen. This small but nearly cosmopolitan
borer, as well as larvae of Coleoptera, including Curculioni-
genus is characterized by development of the female cerci
into a pincerlike structure for grasping the heteropteran host, dae (Guimarães, 1977c) and Cerambycidae. They deposit
concealing a sword-like piercer for egg insertion. This char- their embryonated eggs on the frass or sawdust that plugs the
acter state is found also in Clairvillia and Cinochira, which, entrance to the host burrow, leaving the task of locating the
with Leucostoma, comprise the tribe Leucostomatini. Re- host to the first instar. Newly hatched larvae of some species
inhard (1956) revised the nine Nearctic species of Leucos- can also successfully penetrate the webbing of communal
toma, some of which appear to extend into Central America. tortricid and pyralid caterpillars. In Guanacaste, Costa Rica,
Females inject unembryonated eggs into bugs of the family seven morphospecies have been reared from lepidopterous
Lygaeidae using their sword-like piercer to open the host’s larvae of Thyrididae, Pyralidae, Crambidae, and Riodinidae
integument. Although beetles of the family Meloidae have (Janzen & Hallwachs, 2005).
1408
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Lydella Robineau-Desvoidy. This small genus is known Mauromyia Coquillett. This is an essentially Nearctic
from the Palearctic Region as well as the New World, in- genus of six species (O’Hara, 2002), ranging as far south as
cluding three species north of Mexico (including species Costa Rica.
introduced against the cabbage butterfly), one (L.  jalisco
Woodley) from Mexico, another (L.  minensis (Townsend)) Medina Robineau-Desvoidy. A moderately large ge-
from Brazil, and an undescribed species from Costa Rica. nus, Medina is found worldwide except in Australia and
Neither the Nearctic nor the European species were resolved is best represented in Africa, where 13 species have been
prior to introduction into North America of various popula- recorded (Crosskey, 1984). O’Hara & Wood (2004) listed
tions from Europe; hence, the taxonomy of the New World four nominal species north of Mexico, and three more were
fauna is unclear. Woodley (1994) has listed the world species recorded from Mexico by Wood (1985). Two additional
and provided a phylogenetic interpretation of the group. species have been segregated from the INBio collection in
Costa Rica. Hosts of at least one Nearctic species are flea
Lydinolydella Townsend. This is a small nondescript beetles of the genus Altica. The female fly lands on the
Neotropical genus, perhaps indistinguishable from Euthel- beetle and pries up one of its elytra with its broad, spade-
yconychia, with four species distributed from Mexico to like seventh sternite and deposits an unembryonated egg
Brazil. One species, L. rasilis (Wulp), was reared (as Para- on the lower surface of the elytron (D.M. Wood, personal
dexodes epilachnae Aldrich) from leaf-feeding larvae of the observation).
bean beetle, Epilachna corrupta Mulsant (Coccinellidae)
(Aldrich, 1923). The species occurring in Central America Megaprosopus Macquart. This is a small genus closely
have not been resolved. related to Microphthalma, with a single species in Arizona
and at least one in Costa Rica. Like that genus, females
Lypha Robineau-Desvoidy. As recently redefined by presumably deposit eggs ready to hatch on soil where their
O’Hara (2002), this genus is chiefly Holarctic, reaching its hosts, the larvae of Scarabaeidae, are buried. Although their
southern limit in the highlands of the Talamanca Range in life cycle is analogous to that of several genera of Dexiini
Costa Rica, above 2000  m, where two species have been (for example, Ptilodexia), Megaprosopus and Microphthal-
collected. O’Hara (2002) recorded several families of mi- ma are not related to Dexiinae.
crolepidoptera as hosts of Nearctic species.
Meleterus Aldrich. This is a small New World genus
Macromya Robineau-Desvoidy. This genus is essen- with two species north of Mexico and three morphospecies
tially Neotropical, with a single species, M.  crocata Rein-
in Costa Rica.
hard, in southwestern USA and four species in Costa Rica.
Guimarães (1977c) recorded larvae of both Arctiidae and Metaplagia Coquillett. This predominantly Nearctic ge-
Megalopygidae as hosts of M. crocata. Macromya dorsalis nus has six species; a single species, which may be one of
(Wulp) was reared in Guanacaste, Costa Rica, from larvae of the six, has been reared from a noctuid moth larva in Guana-
Apatelodidae (Janzen & Hallwachs, 2005). Reinhard (1968) caste, Costa Rica (Janzen & Hallwachs, 2005). Two others
provided a key to the species.
were reared from Sphingidae (Lepidoptera).
Madremyia Townsend. Basically a Holarctic genus with
Microchaetina Wulp. Microchaetina is a small New
a single widespread Nearctic species, M.  saundersii (Wil-
liston), and another in northern Europe that may be con- World genus with eight species recorded from the Nearctic
specific with it, this genus is another example of a northern Region (O’Hara & Wood, 2004) and four from the Neotropi-
group that has successfully penetrated the tropics as far as cal Region, including a single species in Costa Rica.
the Talamanca Mountains of southern Costa Rica.
Micronychiops Townsend. This is a small Neotropical
Masiphya Brauer  & Bergenstamm. Masiphya is a genus of only two nominal species found from Costa Rica
New World genus of mantid parasitoids, ranging from to Bolivia.
USA to Brazil, with 26 named species that are currently
assigned to several genera in the Masiphyini (Guimarães, Microphthalma Macquart. This is a nearly cosmo-
1971b ). Generic limits are still close to the one species politan genus with few species scattered about the world: a
per genus range, and the entire assemblage, with the ex- single species in the Palearctic and Oriental Regions (Cross-
ception of Mystacomyia Giglio-Tos, recognized here, key, 1976; Herting, 1984), four species in Africa (Crosskey,
may be congeneric. Heitzmann-Fontinelle  & Guimarães 1984), three species in North America, and seven species in
(1966) reported survival of mantids after parasitism. Central and South America. The New World species were
Guimarães (1966b) added six new species and some new first treated by Aldrich (1926b). First instar larvae hatch from
genera to the Brazilian fauna based on permutations and eggs deposited on soil and burrow into the ground to locate
combinations of a few variable character states. O’Hara  & larvae of Scarabaeidae (Coleoptera), analogous to those of
Wood (2004) listed five species north of Mexico. the tribe Dexiini.
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1409

Minthoplagia Townsend. Minthoplagia setifrons (Wulp) majority of Mystacella species occur in Mexico. Three spe-
is the only species of this Neotropical genus, occurring from cies are known north of Mexico (O’Hara  & Wood, 2004),
Mexico to Venezuela. Its biology is apparently unknown. and several undescribed species are present in the INBio
collection.
Mochlosoma Brauer  & Bergenstamm. Mochlosoma
is a predominantly Neotropical genus, with 16 species in Mystacomyia Giglio-Tos. This is a small Neotropical ge-
Mexico, one each in Costa Rica and Brazil, and five north nus of only three described species ranging from southern
of Mexico, all but one in western USA (Reinhard, 1958). A Texas to Brazil. Guimarães (1977c) recorded a mantid of the
close relative of Ptilodexia, hosts are presumably also larvae genus Zoolea as a host of M. rubriventris (Wulp).
of Scarabaeidae buried in soil.
Neaera Robineau-Desvoidy. A small genus known from
Muscopteryx Townsend. This New World genus is most the Holarctic and Neotropical Regions, Neaera has three spe-
diverse in the Nearctic Region, where O’Hara  & Wood cies north of Mexico, one from the Bahamas, and one from
(2004) recorded 10 species north of Mexico. Three spe- Mexico. One species was reared from Guanacaste, Costa
cies were reared from lepidopterous larvae of Noctuidae, Rica, from larvae of a species of Ethmiidae (Lepidoptera)
Geometridae, and Apatelodidae in Guanacaste, Costa Rica (Janzen & Hallwachs, 2005).
(Janzen & Hallwachs, 2005).
Nemorilla Rondani. This is a small genus with only
Myiomima Brauer  & Bergenstamm. Myiomima is a three Nearctic and three Neotropical species, distributed
Neotropical genus of three nominal species ranging from from Canada to Brazil; it is also represented in the Palearctic
Costa Rica to Paraguay. Region. Two species were reared from larvae of Lacturidae,
Crambidae, and Gelechiidae (Lepidoptera) in Guanacaste,
Myiopharus Brauer & Bergenstamm. This is the larg- Costa Rica (Janzen & Hallwachs, 2005).
est genus of Blondeliini, with 54 nominal species listed for
North and Central America and the West Indies (Wood, 1985) Neobrachelia Townsend. This small enigmatic Neotropi-
and at least that many more in South America. O’Hara  & cal genus, the only described New World genus in the tribe
Wood (2004) recorded 14 from USA and Canada, and about Parerigonini, has only five described species and three un-
40 species based on males only have been segregated in the described species in Central America. One Patagonian spe-
INBio collection from Costa Rica. Because of substantial cies was reared from a pentatomid bug of the genus Edessa
sexual dimorphism, many females have not yet been associ- (Guimarães, 1977c).
ated with their respective males. The Nearctic species that
have been reared all parasitize chrysomelid beetles, some Neomintho Brauer  & Bergenstamm. This is a moder-
attacking larvae, others adult beetles, and Chrysomelidae ately large New World genus most diverse in Central and
are also listed by Guimarães (1977c) as hosts. Females of South America, with over 20 nominal species that could be
a few species, including those placed in Doryphorophaga assigned to the genus, but that are now dispersed among nu-
Townsend by Sabrosky & Arnaud (1965), attack larval hosts merous monotypic genera. Only two species are known north
by injecting their embryonated eggs into their hosts. The sev- of Mexico (O’Hara  & Wood, 2004). Although well repre-
enth sternite is modified into a small sharp piercer, analogous sented in Central America, the number of species has not
to that found in Eucelatoria, but opposed by the sixth sternite been ascertained. Arnaud (1963b) described several South
in front of it rather than the ventral carina of the abdomen. A American species with elaborately branched “fissicorn” an-
few other species, including M. dorsalis (Coquillett), parasit- tennae in the male but normal antennae in the female.
ize adults, and their seventh sternite is flattened side to side
Neosolieria Townsend. This small Neotropical genus has
like a blunt knife, presumably for prying up an elytron. The
three species that range from Baja California to Peru and
majority of females of Myiopharus, however, have simple tu-
Brazil.
bular ovipositors for depositing eggs directly on their host’s
cuticle. Two species were reared from Chrysomelidae in Neosophia Guimarães. This obscure Neotropical genus
Guanacaste, Costa Rica (Janzen & Hallwachs, 2005). of two species described from Brazil (Guimarães, 1982) is
also represented by a few specimens of two species collected
Myothyriopsis Townsend. This is a small Neotropical ge-
in Costa Rica.
nus of only five nominal species, of which only one, M. pic-
ta (Wulp), extends north to Texas. One or two species have Oberonomyia Reinhard. A Neotropical genus related
been reared in Costa Rica and Trinidad from larvae of the to Ptilodexia, Oberonomyia has a single described species,
nymphalid butterfly genus Heliconius (Janzen & Hallwachs, O. palpalis Reinhard, ranging from Mexico to Costa Rica.
2005; Thompson, 1966, as Heliconiophaga Thompson).
Oestrophasia Brauer & Bergenstamm. This is a small
Mystacella Wulp. A small New World genus of 11 de- distinctive New World genus with 10 species in the Neotro-
scribed species ranging from southern Canada to Brazil, the pical Region (Guimarães, 1977a). One, O. signifera (Wulp),
1410
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

is shared with the Nearctic Region, occurring from southern greatest diversity is apparently in Mexico and Central Amer-
Canada to Costa Rica, and three more are found north of ica, although the actual number of species is not known.
Mexico (O’Hara & Wood, 2004). Hosts are adult beetles of
the family Scarabaeidae (Spangler & Burger, 1999). Parepalpus Coquillett. This New World genus of 10
nominal species ranges from western USA to Peru. Three
Ophirion Townsend. This Neotropical genus of nine morphospecies are known from Central America.
nominal species is distributed from Mexico to Peru and Bra-
zil, with a single named species, O. tersum (Townsend), re- Patelloa Townsend. A genus confined to the New World,
corded from Guatemala. Additional species occur in Costa recorded from southern Canada to Patagonia, Patelloa has 10
Nearctic species and 15 nominal species recorded from the
Rica. Hosts are larvae of vespid wasps. Since the ovipositing
Neotropical Region distributed from Mexico to Argentina.
female is unlikely to be allowed to enter the wasp’s nest,
The best known Nearctic species, P. pachypyga (Aldrich &
she may deposit her eggs at the entrance to the nest in late Webber), is a common parasitoid of the forest tent caterpillar.
evening or at night, leaving the task of locating host larvae to In Guanacaste, the commonest species, P. xanthura (Wulp),
her first instar larvae. has been reared from larvae of a wide array of families of
Lepidoptera (Janzen & Hallwachs, 2005).
Ormia Robineau-Desvoidy. Ormia is a New World,
mostly tropical and subtropical genus with 34 species de- Peleteria Robineau-Desvoidy. This is a large widespread
scribed from the Neotropical Region but only eight occur- genus found in the Palearctic, Nearctic, and Neotropical Re-
ring north of Mexico, half of these shared with Mexico gions, with 85 nominal species in the New World, 28 spe-
(O’Hara & Wood, 2004). Ormia depleta (Wiedemann) was cies north of Mexico, almost all in the west (Richards, 1973;
introduced to Florida from Brazil for biocontrol of mole O’Hara & Wood, 2004), and 16 recorded from Mexico and
crickets. Tavares (1962, 1964, 1965a, b) dealt with the South Central America. The genus seems more diverse at higher el-
American species (under the generic names Ormia, Ormio- evations, and one undetermined species is known only from
phasia Townsend, and Euphasiopteryx Townsend). Although Mt. Chirripo, the highest peak in Costa Rica, where it is evi-
well represented in Central America, the number of species dently common. Guimarães (1977c) listed several species of
there has not been determined. noctuid moths as hosts.
Oxynops Townsend. Only one species of Oxynops, Penthosia Wulp. A single species, P. satanica (Bigot), a
O.  anthracinus (Bigot), is known from North and Central large black pompilid wasp mimic, occurs from southern Ari-
America. A second species, O. macrocera (Townsend), was zona to Costa Rica.
described from Brazil. Hosts are larvae of pterophorid moths
(Sabrosky & Arnaud, 1965), and O. anthracinus was reared Periscepsia Gistel. This small genus is predominantly
from a species of Pterophoridae in Guanacaste, Costa Rica northern in distribution, and eight species occur in the
(Janzen & Hallwachs, 2005). Nearctic Region, with some species or species pairs occur-
ring in both northern North America and Eurasia. Only two
Panzeria Robineau-Desvoidy. This is a large Holarctic species have been found in Costa Rica, both at high eleva-
genus with 29 species north of Mexico (O’Hara  & Wood, tions in the Talamanca Range.
2004) and about the same number in the Palearctic Region,
under several different generic names, including Ernes- Phasia Latreille. Phasia is a large cosmopolitan genus,
with 75 species distributed over every continent of the Old
tia Robineau-Desvoidy and Eurithia Robineau-Desvoidy
World and North America (Sun & Marshall (2003). O’Hara &
(Herting, 1984). Only a single species, probably P. ampelus
Wood (2004) listed 16 species north of Mexico. Although
(Walker), occurs in Costa Rica.
represented in Central and South America, the number of
Paradejeania Brauer & Bergenstamm. This distinctive Neotropical species is unknown. Like most phasiines, spe-
genus of only three species includes some of the largest ta- cies of Phasia are parasitoids of bugs of the families Pentato-
chinids. One is Central American, P. rutilioides (Jaennicke), midae and Pyrrhocoridae (Guimarães, 1977c). One species
which occurs from Colorado to Costa Rica, with a subspe- (belonging to the subgenus Paraphorantha Townsend) was
reared from a bug of the family Coreidae in Guanacaste,
cies, nigrescens Arnaud, from British Columbia to Califor-
Costa Rica (Janzen & Hallwachs, 2005).
nia, while two others are found in Colombia and Hispaniola.
Arnaud (2001) recorded P. rutilioides as a parasitoid of the Phasiophyto Townsend. This Neotropical genus, which
larva of an arctiid moth, Hemihylea sp. Woodley (1993) pro- may not be separable from Ebenia, has a single named spe-
vided a key to the three species. cies, P. fumifera Townsend, described from Peru. The Costa
Rican species may be conspecific.
Paradidyma Brauer & Bergenstamm. This New World
genus has 16 species recorded north of Mexico (O’Hara & Phobetromyia Reinhard. Known only from Mexico and
Wood, 2004). The most recent species treatment for North Central America, this genus contains only a single species,
and Central America was given by Reinhard (1934b). The P. dumalis Reinhard.
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1411

Phosocephala Townsend. The only species of the ge- doptera in Guanacaste, Costa Rica (Janzen  & Hallwachs,
nus, P.  metallica Townsend, is known from Costa Rica. It 2005), and Guimarães (1977c) recorded Pyralidae as a host.
has been reared from larvae of a species of noctuid moth in
Guanacaste (Janzen & Hallwachs, 2005). Pseudodexia Brauer & Bergenstamm. There is appar-
ently only one species of this genus, P. eques (Wiedemann),
Phyllophilopsis Townsend. This New World genus has occurring from Costa Rica to Brazil. Wood (1985) incor-
only two species north of Mexico (O’Hara & Wood, 2004) rectly included this genus in the Blondeliini because of the
but 20 nominal species in the Neotropical Region, includ- presence of prosternal setae, but the male terminalia indicate
ing an undetermined number from Central America. Known that its correct placement is in the Dexiinae.
hosts are larvae of endomychid beetles.
Pseudosiphona Townsend. O’Hara (1989) placed all
Phytomyptera Rondani. This is a large cosmopolitan ge- members of the tribe Siphonini, in which vein  A reached
nus of small flies with 23 species recorded north of Mexico the wing margin, in several subgenera of Siphona. Siphona
(O’Hara & Wood, 2004) and about an equal number from the sensu stricto is fairly easily recognized by the elongate la-
Neotropical Region. In Guanacaste, Costa Rica, two species bella and has been recognized here as a full genus, while the
were reared from moth larvae of the families Crambidae and other subgenera cannot be reliably separated from one an-
Elachistidae (Janzen & Hallwachs, 2005). other except by a detailed study of male genitalia. Although
there are many species in this category in Central America,
Plagiomima Brauer & Bergenstamm. Although not yet nearly all of them seem to belong to Pseudosiphona, which
known from Costa Rica, two species of this genus have been
can be easily recognized by the position of the katepisternal
recorded from southern Mexico.
bristles. Pseudosiphona is consequently recognized here as
Polistiopsis Townsend. Species of this distinctive Neo- a full genus to include all those species that were included in
tropical genus mimic vespid wasps of the genus Polistes. Siphona by O’Hara (1989) and O’Hara & Wood (2004) but
There are only two described species (Arnaud, 1966), one of that are not Siphona sensu stricto.
which, P. mima Townsend, occurs in Central America.
Pseudoviviania Brauer & Bergenstamm. This small, sel-
Polygaster Wulp. This Neotropical genus with three de- dom collected Neotropical genus of the high Andean páramo
scribed species is probably the sister genus of Spathidexia consists of three nominal species, ranging from Costa Rica
Townsend. (where only a single specimen has been collected) to Peru.

Prosenoides Brauer  & Bergenstamm. This is a small Ptilodexia Brauer & Bergenstamm. This is a large genus
New World genus with three species north of Mexico re- of uniformly brown, medium-sized dexiines, most diverse in
stricted to the southernmost states of USA and seven nomi- Mexico and the southwestern USA. The 22 Nearctic species
nal species in the Neotropical Region. were revised by Wilder (1979). Guimarães (1971b) recorded
29 species south of Mexico. Females deposit eggs ready to
Prospherysa Wulp. This New World genus has 10 de- hatch on soil where the first instars must burrow down to
scribed species ranging from USA to Brazil. Two species find larvae of Scarabaeidae (Coleoptera). The genus grades
have been recorded north of Mexico (O’Hara  & Wood, imperceptibly into Hystricodexia, Eudexia, and others, but
2004). members of Ptilodexia are rather uniform reddish-brown to
blackish, without enlarged abdomens and few discal bristles.
Protodejeania Townsend. This is a small, essentially Members of Mochlosoma are similar but have a much longer
Mexican and Central American genus of six nominal spe- proboscis.
cies, two of which have been recorded from British Colum-
bia to Mexico (O’Hara & Wood, 2004). Two more, P. major Pyrrhodexia Townsend. This is a small Neotropical genus
Curran and P.  pachecoi Curran, have been described from of only three nominal species, related to Acronacantha and
Guatemala. Microphthalma and with presumably a similar life history.

Pseudobombyliomyia Townsend. This New World genus Quadratosoma Townsend. This Neotropical genus of 11 spe-
has a single species, P.  linellii Townsend, occurring from cies ranges from Costa Rica to Peru in mid-elevation cloud for-
Costa Rica to Venezuela. ests. A single unidentified species occurs in Central America.

Pseudochaeta Coquillett. This is a New World genus of Rhachoepalpus Townsend. This New World genus of
26 described species ranging from southern Canada to Peru. about 20 nominal species, plus several unnamed ones, rang-
Fifteen of these are Neotropical and 11 are known north of es from Arizona to Argentina. Most occur in cloud forests
Mexico (O’Hara & Wood, 2004). Six were described from high in the Andes. Four nominal species have been recorded
Trinidad alone (Thompson, 1964). Three morphospecies from Central America; R. olivaceus Townsend extends north
have been reared from larvae of a variety of families of Lepi- to southern Arizona, while R. immaculatus (Macquart) has
1412
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

been reared from larvae of two species of saturniid moths in Strongygaster Macquart. One Nearctic species, S. trian-
Guanacaste, Costa Rica (Janzen & Hallwachs, 2005). gulifera (Loew), of this small cosmopolitan genus has been
reared from various insects, mostly adult beetles of more
Scotiptera Macquart. This is a small genus of five nomi- than 10 families (Arnaud, 1978; Reeves  & O’Hara, 2004)
nal species confined to forested lowlands of the Neotropical and a pyrrhocorid bug (Guimarães, 1977c). A European spe-
Region, with a single undescribed species extending up to cies, S. globula (Meigen), has been reared from ants, while
mid-elevations in Costa Rica. an unidentified species from South America was reared from
an adult moth, suggesting that this fly is not very discriminat-
Siphona Meigen. Siphona is a large cosmopolitan genus ing in its choice of host.
(O’Hara, 1983, 1989), with 16 species in Siphona sensu stricto
north of Mexico (O’Hara & Wood, 2004) and 11 more in five Sturmiomima Townsend. This small featureless genus
additional subgenera that may not occur in Central America. consists of two species, which may be conspecific; one was
The type species S. geniculata (De Geer) is a European para- reared in Guanacaste, Costa Rica, from a larva of Nymph-
sitoid of tipulid larvae and has been introduced into Canada as alidae (Janzen  & Hallwachs, 2005  — as Pseudosturmia
a biocontrol agent. Other species may attack larvae of Lepi- Thompson).
doptera, particularly Geometridae (see also Pseudosiphona).
Telothyria Wulp. This is a moderate-sized Neotropical
Siphosturmia Coquillett. This New World genus of 12 genus with 20 nominal species in South America, but only
species ranges from Canada to Brazil and includes eight a single species, T.  thecata (Coquillett), north of Mexico
recorded from America north of Mexico (O’Hara & Wood, (O’Hara  & Wood, 2004, as Leskiopsis Townsend). Five
2004). Siphosturmia baccharis (Reinhard), S.  rafaeli morphospecies were reared from larvae of crambid moths
(Townsend), and one other species have been reared from in Guanacaste, Costa Rica (Janzen  & Hallwachs, 2005).
larvae of skippers (Lepidoptera: Hesperiidae), while a fourth Guimarães (1977c) recorded a noctuid moth of the genus
species was reared from a larva of Lycaenidae in Guanacaste, Spodoptera as a host.
Costa Rica (Janzen & Hallwachs, 2005).
Thelairaporia Guimarães. A single species of this genus
Spallanzania Robineau-Desvoidy. A widespread Ho- was reared from larvae of Limacodidae and Dalceridae (Lep-
larctic and Neotropical genus, Spallanzania has six North idoptera) in Guanacaste, Costa Rica (Janzen & Hallwachs,
American species and about 20 nominal species (presently 2005).
assigned to nearly as many genera) at high elevations in the
Andes and at high latitudes in Patagonia. Hosts are larvae of Thelairodes Wulp. One species of this genus was reared
noctuid moths (Guimarães, 1977c). from a larva of a nymphalid butterfly in Guanacaste, Costa
Rica (Janzen & Hallwachs, 2005).
Spathidexia Townsend. This New World genus of nearly
20 species has seven species north of Mexico (O’Hara  & Thelairodoria Townsend. This is a New World genus
Wood, 2004) and 12 in the Neotropical Region. A key to nine of only three species, one of which, T.  setinervis (Coquil-
New World species was provided by Arnaud (1960). Five lett) occurs in eastern USA, and apparently extends south
morphospecies were reared from Hesperiidae in Guanacaste, to Costa Rica. Two additional species have been segregated
Costa Rica (Janzen & Hallwachs, 2005). from INBio material.
Sphaerina Wulp. Only five nominal species of this Neo- Thelairodoriopsis Thompson. Thelairodoriopsis mara-
tropical genus are known, occurring from southern Florida to casi Thompson, described from Trinidad, has been reared
Brazil. Hosts are lepidopterous larvae, mostly of Hesperiidae from a noctuid moth larva in Guanacaste, Costa Rica (Jan-
and Crambidae. Seven morphospecies have been reared from zen & Hallwachs, 2005). An additional morphospecies also
Hesperiidae, Crambidae, and Notodontidae in Guanacaste,
occurs in Costa Rica.
Costa Rica (Janzen & Hallwachs, 2005).
Thelyoxynops Townsend. This is a small Neotropical
Steleoneura Stein. This is a small genus of four widely
genus of two or three species with a similar general habi-
dispersed species, known from the Canary Islands, northern
Russia, northern Canada, and Costa Rica. Apparently, it is tus to that of Myiopharus, except that the subapical scutellar
the sister genus of two Chilean genera, Clastoneura Aldrich bristles are convergent instead of divergent. Their hosts are
and Embiomyia Aldrich; the latter may be congeneric with apparently not known.
Steleoneura. Their rarity, small size, and unknown host may
Trafoia Brauer & Bergenstamm. This small, widespread
retard the gathering of further information.
genus has a single circumpolar species, T. arctica (Sack), in
Stenodexia Wulp. In addition to two unnamed species in the high arctic of Canada and Russia, two more species in
Costa Rica, this obscure Neotropical genus consists of two Europe, and six nominal species ranging from southern Ari-
more nominal species, one in Mexico and one in Jamaica. zona to southern Brazil.
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1413

Trichodura Macquart. A single Costa Rican specimen is cies recorded from Mexico. One of these may occur in Costa
the only evidence of this otherwise South American genus in Rica.
Central America. Guimarães (1972) has revised the Brazilian
species. Vibrissina Rondani. A single species of this genus was
reared from a sawfly larva of the family Argidae in Guana-
Trichophora Macquart. Perhaps the sister taxon to Deo- caste, Costa Rica (Janzen & Hallwachs, 2005).
palpus and Copecrypta, Trichophora is entirely Neotropi-
cal. It could encompass about a dozen species, ranging from Voria Robineau-Desvoidy. One species of this genus was
Mexico to southern Brazil, with three or four found in Cen- reared from a noctuid moth larva in Guanacaste (Janzen &
tral America, including T. rufina (Wulp) from Costa Rica and Hallwachs, 2005).
Guatemala. Guimarães (1977c) recorded T. melas (Bigot) (as
Siphoniomyia melaena Bigot) from a larva of a geometrid Winthemia Robineau-Desvoidy. Winthemia is an easily
moth. recognized cosmopolitan genus of over 110 species, perhaps
most diverse in the New World, where at least 80 nominal
Trichopoda Berthold. This large, predominantly Neotro- species have been described. O’Hara  & Wood (2004) list-
pical genus has six Nearctic species, of which one, T. pen- ed 29 species north of Mexico. There are over 40 nominal
nipes (Fabricius), ranges north to southern Canada and 39 species recorded from the Neotropical Region. Some 48
nominal Neotropical species that range as far south as Chile. morphospecies have been reared from larvae of a variety of
Hosts include bugs of the families Coreidae and Pentatomi- families of Lepidoptera, chiefly Hesperiidae, Notodontidae,
dae (Guimarães, 1977c). The most common Nearctic spe- Noctuidae, Saturniidae, Nymphalidae, and Sphingidae in
cies, T. pennipes, which also may range south into Central Guanacaste, Costa Rica (Janzen  & Hallwachs, 2005). Fe-
America, has been introduced to Hawaii. males deposit unincubated eggs on caterpillars, and the first
instars, of at least one species, W. rufopicta (Bigot), emerge
Trigonospila Pokorny. This small cosmopolitan genus
from the egg shell and penetrate the skin of the host in front
has three species recorded north of Mexico (O’Hara & Wood,
2004), one species across Eurasia, several in Australia, and of the egg shell (Danks, 1975). This is in contrast with first
four in South America. Two species were reared from Cram- instars of the Exoristini, which penetrate the host beneath the
bidae and Pyralidae (Lepidoptera) in Guanacaste, Costa Rica egg shell (Wood, 1972).
(Janzen & Hallwachs, 2005).
Xanthodexia Wulp. A single species of this genus was
Triodontopyga Townsend. This small genus of 11 nomi- reared in Guanacaste, Costa Rica, from a larva of Hesperii-
nal species confined to the Neotropical Region occurs from dae (Lepidoptera) (Janzen & Hallwachs, 2005).
Mexico to Brazil. Seven species were treated by Guimarães
(1983a). Triodontopyga flavolimbata (Bigot) has been reared Xanthoepalpus Townsend. This small New World genus
from lepidopterous larvae of a megalopygid and an apatelo- has only seven nominal species, one of which, X.  bicolor
did in Guanacaste, Costa Rica (Janzen & Hallwachs, 2005). (Williston), occurs in western North America from British
Columbia to Mexico and probably also in Central America.
Trochiloleskia Townsend. This small New World ge- Five species were described from South America, but the ac-
nus consists of only three nominal species distributed from tual number remains unknown.
southern Texas to Brazil.
Xanthomelanodes Townsend. This is a New World genus
Tropidopsiomorpha Townsend. A single species of this with five species north of Mexico (O’Hara & Wood, 2004)
genus, T. tropica Townsend, occurs from Costa Rica to Bra- and 12 south of the USA (Sabrosky, 1950).
zil and is presumably closely related to Hystricodexia and
Eudexia. Its biology is presumably also similar, with embry- Xanthomelanopsis Townsend. This small Neotropical
onated eggs being laid on soil or rotten wood and the first in- genus has only three species, one of which extends north to
star larvae finding the larval scarab beetle host on their own Mexico. Females deposit unembryonated eggs on bugs of the
by burrowing into the substrate. families Largidae and Pyrrhocoridae (Guimarães, 1977c).

Uramya Robineau-Desvoidy. Guimarães (1980) treated Xanthophyto Townsend. This New World genus has three
12 species of this genus from South America and O’Hara & nominal species north of Mexico and an unknown number
Wood (2004) listed seven from north of Mexico. Four addi- from the Neotropical Region. Guimarães (1977c) recorded a
tional names are known from Mexico. Eight morphospecies rearing from an arctiid moth (subfamily Ctenuchinae).
were reared from larvae of limacodid moths in Guanacaste,
Costa Rica (Janzen & Hallwachs, 2005). Xanthozona Townsend. Only one species of this genus,
X. melanopyga (Wiedemann), is known, occurring from Cos-
Vanderwulpia Townsend. This small New World genus ta Rica to Brazil. Hosts include lepidopterous larvae of Arc-
has two species north of Mexico and two Neotropical spe- tiidae, Brassolidae, and Notodontidae (Guimarães, 1977c).
1414
 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Zaira Robineau-Desvoidy. A small genus in Central Zizyphomyia Townsend. This genus, which needs redefi-
America, Zaira has at least 14 species in the Nearctic Region nition, is based on three Nearctic species plus a fourth from
and a single species, Z. cinerea (Fallén), in the Palearctic Re- Brazil that probably belongs to another genus. Four addi-
gion. A few species, including Z. cinerea, have been reared tional species that appear to belong here were reared from
from carabid beetles. Like their apparently close relatives, Lepidoptera of families Thyrididae, Nymphalidae, Papil-
Chaetona and Cryptomeigenia, species of Zaira are crepus- ionidae, and Pyralidae in Guanacaste, Costa Rica (Janzen &
cular or nocturnal and are usually collected at light. Males of Hallwachs, 2005).
at least one species in Florida hilltop after sunset.
Zygozenillia Townsend. Further study may show that
Zelia Robineau-Desvoidy. Zelia is the New World sis- the six New World species now included in Sisyropa
ter genus of Dexia, with a dozen species north of Mexico Brauer  & Bergenstamm, a genus based on an Oriental
(O’Hara & Wood, 2004) and more than 20 nominal species type species, may actually be congeneric with this Brazil-
south of the USA, some now in other genera. Hosts include ian genus. One species, which may be conspecific with
larvae of Coleoptera, including Scarabaeidae, that are buried the Brazilian type species, has been reared in Guanacaste,
in rotten wood. Costa Rica.

Literature cited

Aldrich, J.M. 1923. A new parasitic fly bred from bean beetle. Pro- Wissenschaften in Wien, Mathematisch-Naturwissenschaftliche
ceedings of the Entomological Society of Washington 25: 95–96. Classe 56: 69–180.
Aldrich, J.M. 1926a. North American two-winged flies of the genus Brauer, F.  & J.E. von Bergenstamm. 1891. Die Zweiflügler des
Cylindromyia Meigen (Ocyptera of authors). Proceedings of the Kaiserlichen Museums zu Wien. V. Vorarbeiten zu einer Mono-
United States National Museum 68: 1–27. graphie der Muscaria Schizometopa (exclusive Anthomyidae).
Aldrich, J.M. 1926b. American two-winged flies of the genus Mi- Pars II. F. Tempsky, Wien, 142 pp.
crophthalma Macquart, with notes on related forms. Proceed- Brauer, F.  & J.E. von Bergenstamm. 1893. Die Zweiflügler des
ings of the United States National Museum 69: 1–8. Kaiserlichen Museums zu Wien. VI. Vorarbeiten zu einer Mono-
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Reeves, W.K. & J.E. O’Hara. 2004. First report of Strongygaster Stireman, J.O., J.E. O’Hara, & D.M. Wood. 2006. Tachinidae: Evo-
triangulifera (Diptera: Tachinidae) as a parasitoid of a cantharid lution, behavior, and ecology. Annual Review of Entomology 51:
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dae). The Canadian Entomologist 136: 661–662. Sun, X.  & S.A. Marshall. 2003. Systematics of Phasia Latreille
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belonging to the genus Cuphocera. Proceedings of the United Tavares, O. 1962. Contribuição ao conhecimento da tribo Ormiini I.
States National Museum 83: 45–70. Gênero Ormia Robineau-Desvoidy, 1830 (Diptera, Tachinidae).
Reinhard, H.J. 1934b. American muscoid flies of the genera Cer- Memórias do Instituto Oswaldo Cruz 60: 347–363.
atomyiella and Paradidyma. Proceedings of the United States Tavares, O. 1964. Contribuição ao conhecimento da tribo Ormii-
National Museum 83: 9–43. ni II. Gênero Ormiophasia Townsend. 1919 (Diptera, Tachini-
Reinhard, H.J. 1956. A synopsis of the tachinid genus Leucostoma dae). Memórias do Instituto Oswaldo Cruz 62: 37–52.
(Diptera). Journal of the Kansas Entomological Society 29: Tavares, O. 1965a. Contribuição ao conhecimento da tribo
155–168. Ormiini  III. Gênero Euphasiopteryx Townsend, 1915 (Dip-
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(Tachinidae, Diptera). The Canadian Entomologist 90: 13–25.
98–110. Tavares, O. 1965b. Contribuição ao conhecimento da tribo
Reinhard, H.J. 1968. A review of the genus Macromya (Tachini- Ormiini IV. Gênero Ormia Robineau-Desvoidy, 1830 (Dipte-
dae: Diptera). The Canadian Entomologist 100: 1280–1287. ra, Tachinidae). Revista Brasileira de Biologia 25: 211–216.
Rettenmeyer, C.W. 1961. Observation on the biology and tax- Thompson, W.R. 1915. Sur le cycle évolutif de Fortisia foeda, Dip-
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Tachinidae, Conopidae). University of Kansas Science Bulletin des Séances de la Société de Biologie, Paris 78: 413–416.
42: 993–1066. Thompson, W.R. 1934. The tachinid parasites of woodlice. Parasi-
Richards, L.L.E. 1973. A revision of the Peleteria (Diptera: Tachin- tology. 26: 378–448.
idae) of America north of Mexico. Unpublished Ph.  D. thesis, Thompson, W.R. 1964. The tachinids of Trinidad. VI. The larvipa-
Washington State University, Pullman, xvii + 255 pp. rous Goniines of the Carceliine type (Diptera, Tachinidae). Stu-
Robineau-Desvoidy, J.B. 1830. Essai sur les Myodaires. Mémoires dia Entomologica 7: 97–151.
présentés par divers Savants a l’Académie Royale des Sciences Thompson, W.R. 1966. The tachinids of Trinidad. VII. The larvipa-
de L’Institut de France. Sciences, Mathématiques et Physiques, rous Goniines with broad cheeks (Diptera, Tachinidae). Studia
Serie 2 2: 1–813. Entomologica 8: 353–434.
Robineau-Desvoidy, J.B. 1863a. Histoire naturelle des diptères Toma, R. 2003. Estudo das espécies do “complexo Acaulona” sen-
des environs de Paris, Vol.  1. V.  Masson et fils, Paris, xvi  + su Sabrosky (Diptera, Tachinidae). Revista Brasileira de Ento-
1143 pp. mologia 47: 267–282.
Robineau-Desvoidy, J.B. 1863b. Histoire naturelle des diptères des Toma, R.  & J.H. Guimarães. 2002. Estudo taxonômico de Les-
environs de Paris, Vol. 2. V. Masson et fils, Paris, 920 pp. chenaultia Robineau-Desvoidy (Diptera, Tachinidae). Revista
Sabrosky, C.W. 1947. A synopsis of the larvaevorid flies of the Brasileira de Entomologia 46: 33–70.
genus Eudejeania. Proceedings of the United States National Townsend, C.H.T. 1936a. Manual of Myiology, in twelve parts.
Museum 97: 141–156. Part  III: Oestroid classification and habits (Gymnosomatidae
Sabrosky, C.W. 1950. Notes on Trichopodini (Diptera, Larvaev- to Tachinidae). Charles Townsend  & Filhos, Itaquaquecetuba,
oridae), with description of a new parasite of cotton stainers in São Paulo, 255 pp.
Puerto Rico. Journal of the Washington Academy of Sciences Townsend, C.H.T. 1936b. Manual of Myiology, in twelve parts.
40: 361–371. Part  VI. Oestroid classification and habits (Dexiidae and Ex-
Sabrosky, C.W. 1965. Host of the tachinid tribe Eutherini (Diptera). oristidae). Charles Townsend  & Filhos, Itaquaquecetuba, São
Proceedings of the Entomological Society of Washington 67: Paulo, 303 pp.
61. Townsend, C.H.T. 1938. Manual of Myiology, in twelve parts. Part
Sabrosky, C.W. 1969. A review of the genus Juriniopsis Townsend VII. Oestroid generic diagnoses and data (Gymnosomatini to
(Diptera: Tachinidae). Florida Entomologist 52: 79–90. Senostomatini). Charles Townsend & Filhos, Itaquaquecetuba,
Sabrosky, C.W. 1981. A partial revision of the genus Eucelatoria São Paulo, 434 pp.
(Diptera, Tachinidae), including important parasites of Helio- Townsend, C.H.T. 1939a. Manual of Myiology, in twelve parts.
this. United States Department of Agriculture, Technical Bul- Part VIII. Oestroid generic diagnoses and data (Microtropesini
letin 1635: i–iv + 1–18. to Voriini). Charles Townsend & Filhos, Itaquaquecetuba, São
Sabrosky, C.W.  & P.H. Arnaud, Jr. 1965. Tachinidae, pp.  961– Paulo, 408 pp.
1108. In Stone, A., C.W. Sabrosky, W.W. Wirth, R.H. Foote, & Townsend, C.H.T. 1939b. Manual of Myiology, in twelve parts.
J.R. Coulson (editors). A catalog of the Diptera of America north Part  IX. Oestroid generic diagnoses and data (Thelairini to
of Mexico. United States Department of Agriculture, Agriculture Clythoini). Charles Townsend  & Filhos, Itaquaquecetuba, São
Handbook 276, Washington, D.C., iv + 1696 pp. Paulo, 270 pp.
Sehnal, P. 1998. A new species of Borgmeiermyia Townsend, 1935, Townsend, C.H.T. 1940. Manual of Myiology, in twelve parts.
from Paraguay (Insecta: Diptera: Tachinidae). Annalen des Part X. Oestroid generic diagnoses and data (Anacamptomyini
Naturhistorischen Museums in Wien 100B: 349–354. to Frontinini). Charles Townsend  & Filhos, Itaquaquecetuba,
Shima, H. 1989. Parasitic way of life in tachinid flies. Insectarium São Paulo, 335 pp.
26: 4–9, 46–51, 88–94, 120–126. [In Japanese.] Townsend, C.H.T. 1941. Manual of Myiology, in twelve parts.
Spangler, H.G. & J.F. Burger. 1999. Oestrophasia clausa (Diptera: Part XI. Oestroid generic diagnoses and data (Goniini to Try-
Tachinidae), a parasite of adult Diplotaxis moerens (Coleoptera: pherini). Charles Townsend  & Filhos, Itaquaquecetuba, São
Scarabaeidae). Entomological News 110: 123–124. Paulo, 342 pp.
TACHINIDAE (TACHINID FLIES, PARASITIC FLIES) 113 1417

Townsend, C.H.T. 1942. Manual of Myiology, in twelve parts. Part Wood, D.M. & R.D. Cave. 2006. Description of a new genus and
XII. General consideration of the Oestromuscaria. Charles species of weevil parasitoid from Honduras (Diptera: Tachini-
Townsend & Filhos, Itaquaquecetuba, São Paulo, 365 pp. dae). Florida Entomologist 82: 239–244.
Tschorsnig, H.-P. 1985. Taxonomie forstlich wichtiger Parasiten: Woodley, N.E. 1993. A remarkable new species of Paradejea-
Untersuchungun zur Struktur des männlichen Postabdomens nia from the Dominican Republic (Diptera: Tachinidae).
der Raupenfliegen (Diptera, Tachinidae). Stuttgarter Beiträge Proceedings of the Entomological Society of Washington 95:
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Wilder, D.D. 1979. Systematics of the Nearctic Ptilodexia Brauer Woodley, N.E. 1994. A new species of Lydella (Diptera: Tachini-
and Bergenstamm (Diptera: Tachinidae). Proceedings of the dae) from Mexico with a discussion of the definition of the
California Academy of Sciences 42: 1–55. genus. Bulletin of Entomological Research 84: 131–136.
Wood, D.M. 1972. A revision of the New World Exoristini (Dip- Woodley, N.E.  & P.H. Arnaud. 2008a. Lobomyia neotropica, a
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The Canadian Entomologist 104: 471–503. tropical Region. Zootaxa 1783: 31–39.
Wood, D.M. 1985. A taxonomic conspectus of the Blondeliini of Woodley, N.E.  & P.H. Arnaud. 2008b. Eulobomyia, a new re-
North and Central America and the West Indies (Diptera: Ta- placement name for Lobomyia Woodley  & Arnaud (Diptera:
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Wood, D.M. 1987. Tachinidae, pp. 1193–1269. MND, Volume 2. Biodiversidad, Santo Domingo de Heredia, 143 pp.
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TAXONOMIC INDEX TO VOLUME 2  1419

TAXONOMIC INDEX TO VOLUME 2

Brian V. Brown
This index includes all names below the level of family, agromyzina Hennig, Shewellia, 1054
including genus and species names, mentioned in the text of Agromyzinae, 1060, 1063, 1070, 1210
Volume 2. Agrophaspidium Wheeler & Mlynarek, 1146, 1149–1150, 1153
aibonito Curran, Minettia, 992
abaca (Hall), Mesembrinella 1301 alani Steyskal, Procecidochares, 939, 950
Abacantha Hall, 1322 albeto Gaimari & Silva, Physegeniopsis, 972, 992
Abaristophora Schmitz, 728, 732, 753 albeto Gaimari, Helgreelia, 1053
abnormalis (Malloch), Amauromyza, 1065, 1068, 1070 albibasis (Enderlein), Taeniaptera, 806
abrupta (Wiedemann), Hystricia, 1370, 1406 albiceps (Wiedemann), Chrysomya, 1303
Acanthodotheca Townsend, 1328–1329 albimana (Hennig), Pseudopomyza, 824
Acantholespesia Wood, 1363, 1398 albincisa (Wiedemann), Argyrophylax, 1399
Acanthopeltastes Enderlein, 1139–1141, 1148–1149 albipennis (Say), Icosta, 1247
Acanthophorides Borgmeier, 733, 735, 753 albipes (Coquillett), Siphonophysa, 993
Acaulona Wulp, 1390, 1398, 1405, 1416 albomaculatus (Jaennicke), Epalpus, 1403
Acemya Robineau-Desvoidy, 1372, 1398 albosetulosa (Strobl), Tethina, 1112
Acemyini, 1398 albostriata Hendel, Megalaemyia, 893
Achias Fabricius, 905, 907 albovenosa Coquillett, Hyadina, 1229, 1232
Acidogona Loew, 914, 926, 934, 947 aldrichi (Mesnil), Istochaeta, 1373, 1402
Acinia Robineau-Desvoidy, 926, 943, 946–947, 952 aldrichi Allen, Phrosinella, 1331
Aciurina Curran, 923, 926, 932, 942, 947, 953 aldrichi Del Ponte, Cochliomyia, 1303
Aciuroides Hendel, 881–882, 892, 894, 897–898 Aldrichiomyza Hendel, 1128–1129, 1132
Acompha Hendel, 877 aliena (Malloch), Gibbolauxania, 991
Acontistoptera Brues, 751–753 alini (Shatalkin), Procerosoma, 1043, 1047
Acridiophaga Townsend, 1329 Alipumilio Shannon, 772, 775, 782, 786, 792
Acrometopia Schiner, 1001, 1006 alleni Thompson, Eristalis, 787
Acronacantha Wulp, 1350, 1386, 1398, 1411 Allobosca Speiser, 1244
Acrosticta Loew, 883, 887–888, 890, 898–900 Alloboscinae, 1244
Acrotaenia Loew, 926, 938, 947 Allochaeta Borgmeier, 749, 753
Acrotaeniacantha Hering, 926, 938 Allograpta Osten Sacken, 766, 771, 773, 775, 777–779, 782,
Actia Robineau-Desvoidy, 1371, 1374, 1377, 1383, 1398, 1401, 1415 786, 791
Actinochaeta Brauer & Bergenstamm, 1385, 1398 Allolimosina Roháček, 1184
Actinodoria Townsend, 1352, 1378, 1398 Allomethus Hardy, 794, 798, 800, 802
Adelopteromyia Schmitz, 750, 752–753, 755, 760 Allometopon Kertész, 1043
Adenophora Borgmeier, 752–753 Allominettia Hendel, 974, 981–983, 991
Adjeania Townsend, 1357 Allotrichoma Becker, 1218, 1222, 1224, 1228
Admontia Brauer & Bergenstamm, 1348, 1353, 1378, 1398, 1407 almeidai (Lopes), Villegasia, 1333
Adramini, 910 Alujamyia Norrbom, 909, 922, 947, 954
adusta (Loew), Juriniopsis, 1395, 1406 alvarengai Borgmeier, Eurycnemis, 755
Aecothea Haliday, 1161 alvarengai Prado, Lopesiodinia, 1053
aelops (Walker), Austeniops, 1399 Amauromyza Hendel, 1060, 1065–1071
aenescens (Wiedemann), Hydrotaea, 1293 amazonica (Albuquerque & Lopez), Micropotamia, 1293
aeneus (Cresson), Pectinifer, 1230 ambigua Kessel, Speiseria, 1249–1250, 1259
Aenigmatiinae, 727, 758–759 Ameniinae, 1300
aestiva (Fabricius), Somatia, 833–835 americanus Wiedemann, Eupeodes, 787
aethiops Malloch, Milichia, 1134 Ametadoria Townsend, 1347, 1366, 1398
africa (Wiedemann), Sarcophaga, 1332, 855 Amiota Loew, 1199, 1201–1202, 1204
Afrocamilla Barraclough, 1193–1195 Amobia Robineau-Desvoidy, 1314–1315, 1318, 1329
Afrodinia Cogan, 1051–1052, 1054 Amoebaleria Garrett, 1161, 1163
Agallamyia Norrbom, 911, 914, 920, 923, 926, 947 amoena (Enderlein), Apterocerina, 898
Agathomyia Verrall, 716–719 ampelus (Walker), Panzeria, 1350, 1410
Agenamyia Albuquerque, 1284, 1290–1291 Amphicnephes Loew, 904–907
Agnitrena Korneyev, 910, 915, 922, 926 amplipennis Hendel, Leptopyrgota, 960
Agriphoneura Hendel, 981, 991 amplipennis Portschinsky, Scathophaga, 1267
Agrolimna Meigen, 1215–1216, 1225 Amputella Marshall, 1172, 1184
Agromyza Fallén, 1057, 1059, 1061, 1063, 1067 Amygdalops Lamb, 1075, 1079, 1081
1420 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

anacrostichalis Grimaldi & Mathis, Stenomicra, 1092 approximata Sturtevant & Wheeler, Nostima, 1230, 1232
Anagnota Becker, 1074, 1079 Apterella Borgmeier, 751, 753
Anapyrgota Steyskal, 956–961 Apterocerina Hendel, 892, 894, 897–900
Anarista Papp, 1094 Apterophora Brues, 729, 750, 752–753
Anastrebla Wenzel, 1251–1252, 1256 Apterosepsis Richards, 1075
Anastrepha Schiner, 909–910, 912, 919–920, 926, 944, 947, Aptilotella Duda, 1182–1183
953–954 Aptilotus Mik, 1183, 1186
Anatrichobius Wenzel, 1254–1255, 1257–1258 aranea (Coquillett), Megistopoda, 1250, 1259
anchineuria Speiser, Ornithomya, 1243, 1245, 1247 Archiborborus Duda, 1167, 1171, 1182–1183
Anchioleucopis Tanasijtshuk, 997–998, 1000, 1003 Archiceroptera Papp, 1169, 1172, 1175, 1180–1181, 1183
Anepalpus Townsend, 1359, 1398, 1407 Archicollinella Duda, 1177
Anevrina Lioy, 731, 753 Archileptocera Duda, 1169, 1172, 1176, 1183, 1186
Angioneura Brauer & Bergenstamm, 1297–1298, 1300–1303 Archimimus Reinhard, 1329
Angioneurini, 1301 Archisepsis Silva, 1033–1039
Angitulinae, 905 Archynycteribiinae, 1264
angulata (Thomson), Poecilosomella, 1170, 1172, 1185 Archytas Jaennicke, 1358, 1360, 1399, 1404–1405, 1415
angustata Brown, Cootiphora, 732, 736, 754 arctica (Sack), Trafoia, 1350, 1412
Angustia Sellers, 1375, 1398 Arctophila Schiner, 768–769, 790
angusticercus Marshall & Buck, Lepidosina, 1184 arcuatum Hendel, Chondrometopum, 893
angustivitta (Aldrich & Webber), Angustia, 1398 ardeoceras Mathis, Rhysophora, 1231
animata Spencer, Agromyza, 1067 argentina (Malloch), Diastata, 958, 1210
Anisia Wulp, 1345, 1347, 1378–1380, 1397–1398, 1415 Argentinomyia Lynch Arribálzaga, 767, 776, 779, 786
ankhoidea Gaimari & Silva, Physegeniopsis, 992 Argoravinia Townsend, 1319, 1324, 1329
Annimyzella Spencer, 1067, 1069, 1071 Argyrochaetona Townsend, 1347, 1367, 1394, 1399
annulata (Fabricius), Taeniaptera, 806 Argyromima Brauer & Bergenstamm, 1394, 1396, 1399
anomala Cole, Coelopina, 1014–1016 Argyrophylax Brauer & Bergenstamm, 1366, 1399
anomala Cresson, Diphuia, 1229, 1262 aristalis (Fischer), Neorhinotora, 1163
Anomioptera Schiner, 1169, 1174, 1186 Aristosyrphus Curran, 769, 775, 787, 792
anonyma (Williston), Monochaetoscinella, 1151 arnaudi Barraclough & Wheeler, Camilla, 1194–1195
Anoxynops Townsend, 1348, 1371, 1373, 1399, 1407 Arrhinactia Townsend, 1397, 1399
antennaepes (Say), Rainieria, 806, 808 arthriticus (Wiedemann), Plocoscelus, 806
antennalis Townsend, Itaplectops, 1406 Ascodipterinae, 1250
antennata Sabrosky, Asteia, 1095 Ascodipteron Adensamer, 1250
Anthojuba Albuquerque & Couri, 1276 Asemosyrphus Bigot, 767–768, 787, 791
Anthomyia Meigen, 1271–1272, 1274–1275 ashleyi Barraclough, Ramuliseta, 965
Anthomyza Fallén, 1074–1075, 1080 asiatica Spencer, Pseudonapomyza, 1070
Anthomyzinae, 1074 Asilostoma Hendel, 976, 991
anthracinus (Bigot), Oxynops, 1410 Asiopsila Shatalkin, 839
Anticheta Haliday, 1019 Aspidoptera Coquillett, 1253, 1255, 1257, 1259
Antillonerius Hennig, 817–819 Aspistomella Hendel, 900
Antineuromyia Hendel, 876, 878 assimilis (Malloch), Allominettia, 991
Antrops Enderlein, 1182 Asteia Meigen, 1093–1096
Apacrochaeta Duda, 1203 Asteiinae, 1094
Apallates Sabrosky, 1139, 1143, 1145, 1147, 1149–1150 Asteiini, 1094
Apetaenus Eaton, 1112 Asteimyia Sabrosky, 1095
apex Hendel, Setellia, 872, 878 Atacta Schiner, 1352, 1364, 1367, 1399
Aphaniosoma Becker, 1155–1157 Atactosturmia Townsend, 1366–1367, 1399
apicalis Loew, Chyliza, 838, 841–842 Atherigona Rondani, 1282, 1284, 1288, 1291, 1295
Apiochaeta Czerny, 1043, 1045–1048 Atherigoninae, 1282
Aplomyopsis Townsend, 1365, 1399 Athrycia Robineau-Desvoidy, 1349, 1384
Apocephalus Coquillett, 726, 731, 733–734, 737–738, 740–741, Athyroglossa Loew, 1215, 1222, 1228–1229, 1232
744–745, 749, 753–754, 759 Atissa Haliday, 1215, 1222, 1228, 1231
Apodicrania Borgmeier, 735, 748, 753 Atissini, 1221, 1228, 1237
Apophorhynchus Williston, 1026, 1029 atlantica Papp & Mathis, Nannodastia, 1215, 1238
Apops McAlpine, 823 atomaria (Wiedemann), Automola, 873, 878
Apotropina Hendel, 1137, 1139–1140, 1149 atriceps Malloch, Asilostoma, 991
appendiculata (Loew), Neoidiotypa, 875, 878 Atrichomelina Cresson, 1019, 1021–1023
appendiculata Stein, Myopina, 1273 atrichosa McAlpine, Protopiophila, 869
appretiata Schmitz, Diocophora, 738, 755 atrimana Meigen, Pseudopomyza, 823
approximata (Malloch), Allominettia, 991 Auchmeromyia Brauer & Bergenstamm, 1300
TAXONOMIC INDEX TO VOLUME 2  1421

aurata Robineau-Desvoidy, Hemyda, 1353, 1405 bigoti Melander, Chyliza, 841

auricaudata Williston, Mutillimyia, 788 bilineata (Foote), Dyseuaresta, 934
aurifrons Malloch, Schoenomyza, 1294 Billaea Robineau-Desvoidy, 1353, 1370, 1381, 1400
aurigena (Lopes), Rafaelia, 1331 bimaculata Malloch, Deutominettia, 991
Austeniops Townsend, 1359, 1399, 1407 bimaculata Williston, Tropidomyia, 851, 854
Australimyza Harrison, 1103–1104 Biopyrellia Townsend, 1285, 1291, 1295
Australofannia Pont, 1278 Biorbitella Sabrosky, 1142, 1149
austrina (Coquillett), Cirrula, 1223, 1228 bipuncta (Aczél), Xanthaciura, 931
Austrophorocera Townsend, 1373, 1376, 1399 bipuncta (Wiedemann), Biopyrellia, 1291, 1295
Automola Loew, 871–874, 878 bipunctata Hendel, Ophthalmoptera, 899
autumnalis De Geer, Musca, 1293 bipuncticeps Malloch, Sepedomerus, 1020
Auxanommatidia Borgmeier, 740, 744, 753–754, 759–760 bisignata Melander, Milichiella, 1134
Avibrissosturmia Townsend, 1361, 1399 bisulcata (Macquart), Olfersia, 1247
Axinota Wulp, 1191 Bitheca Marshall, 1172, 1179, 1181, 1183, 1186
Axiologina Hendel, 886, 889–890, 898, 902 Bithoracochaeta Stein, 1288–1289, 1291
Azelia Robineau-Desvoidy, 1284, 1287, 1291 bivisualis Patterson, Rhinoleucophena, 1205
Azeliinae, 1282, 1294–1295 bivittata (Malloch), Leucopina, 1005
Azorastia Frey, 1235, 1237 Blaesoxipha Loew, 1316–1317, 1321, 1325–1326,
azteca McAlpine, Prochyliza, 756, 868 1328–1329, 1334
aztecarum Wheeler, Nothomicrodon, 765, 784 blanda (Osten Sacken), Hyphantrophaga, 1405
baccharis (Reinhard), Siphosturmia, 1412 Blepharipa Rondani, 1354, 1366, 1388, 1400
Bactrocera Macquart, 910, 919, 926, 947 Blepharolauxania Hendel, 984, 991
Bactropalpus Borgmeier, 739, 758 Blepharoneura Loew, 910–911, 914–915, 920, 924, 927, 948,
bakeri Coquillett, Chaetoclusia, 1047 952–953
Baliopteridion Papp & Silva, 978 Blepharoneurinae, 910, 948, 952, 954
balli Brown, Platydipteron, 746, 757 Blondelia Robineau-Desvoidy, 1378, 1388, 1394, 1400, 1403
Baryplegma Wulp, 926–927, 938, 943, 945, 948 Blondeliini, 1345, 1356, 1372, 1389, 1397–1398, 1403, 1407,
basalis (Meigen), Campichoeta, 1209 1409, 1411, 1417
basalis (Stein), Neodexiopsis, 1294 Bocainamyia Albuquerque, 866, 868
Basileunculus Rafael, 794, 799–802 Boettcheria Parker, 1319, 1321, 1327, 1329–1331, 1333–1334
Basilia Miranda-Ribeiro, 1261, 1263–1266 boletina (Zetterstedt), Odinia, 1051
basilica Snow, Polymorphomyia, 950 boliviensis Sasakawa, Amauromyza, 1068
basistriga (Walker), Odontomera, 878 Bolopus Enderlein, 716
Batrachophthalmum Hendel, 877 Bombyliomyia Brauer & Bergenstamm, 1346, 1368, 1394, 1400
batracida Grimaldi, Drosophila, 1200 Boreothrinax Steyskal, 958, 960
beckeri Czerny, Hendelia, 1047 borgmeieri Hendel, Lochmostylia, 963–966, 968
Beckeriella Williston, 1219, 1228, 1232–1233 Borgmeieriphora Prado, 735, 745, 747, 751
Beebeomyia Curran, 877–878 Borgmeiermyia Townsend, 1347, 1374, 1400, 1414, 1416
bella Williston, Callomyia, 717, 719 borinquenensis Wheeler, Puliciphora, 726, 761
Bellardia Robineau-Desvoidy, 1300 Borophaga Enderlein, 730, 754, 756
bellardii (Rondani), Basilia, 1261, 1266 borrichia (Bush & Huettel), Dracontomyia, 930, 937, 949
bellula (Williston), Leucopina, 1005 bosqi Pape & Mello, Engelimyia, 1330
bellum Giglio-Tos, Melanostoma, 788 Bothrometopa Hendel, 892–893, 896
Belobackenbardiinae, 839 bovis (Linnaeus), Hypoderma, 1307, 1310
Belvosia Robineau-Desvoidy, 1366, 1399 Brachicoma Rondani, 1320, 1329
belzebul (Schiner), Scipopus, 806 Brachycephaloptera Borgmeier, 733, 740, 754
benjamini Malloch, Fannia, 1278 Brachydeutera Loew, 1220, 1225, 1228, 1233
bequaerti Malloch, Chortinus, 1292, 1294–1295 brachystoma (Stenhammar), Thoracochaeta, 1186
bergi Valley, Dictya, 1022 Brachytarsininae, 1250
Bertamyia Kessel, 716–719 brasiliana Townsend, Borgmeiermyia, 1400
Beskia Brauer & Bergenstamm, 1351, 1393, 1399 brasiliensis Cresson, Hecamede, 1229
Bezzimyia Townsend, 1337–1341 brasiliensis Schiner, Atacta, 1352, 1399
Bibiomima Brauer & Bergenstamm, 1382, 1390, 1399 brasiliensis (Schiner), Protodictya, 1023
bicasa Kung & Brown, Tapantia, 732, 738, 757 Braula Nitzsch, 1097–1099
bicolor (Williston), Xanthoepalpus, 1395, 1413 breviceps Curran, Eldunnia, 1252, 1257
bifenestratum Kertész, Chondrometopum, 892, 898 brevicornis Wulp, Cordilura, 1269
bifrons (Stenhammar), Spelobia, 1170, 1172, 1186 brevicostata (Duda), Spinilimosina, 1170, 1186
Bifronsina Roháček, 1170, 1172, 1177, 1186 brevihirta Malloch, Schildomyia, 1054
bifurcata (Fallén), Lonchoptera, 721–722 Bricinnia Walker, 907
bifurcata Sabrosky, Loewimyia, 1095 Bricinniella Giglio-Tos, 907
1422 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Bromeloecia Spuler, 1170, 1175, 1183 Carpomya Costa, 920, 925, 927, 948, 954
Brontaea Kowarz, 1289, 1291 Carposcalis Enderlein, 776, 789
brooksae Pipkin, Scaptodrosophila, 1205 Carrerapyrgota Aczél, 958, 960–961
Brownphora Disney, 732, 735, 754 Carreriamyia van Doesburg, 772
brunnea Cresson, Syringogaster, 843, 845 Casahuiria Townsend, 1362, 1400
brunnea (Latreille), Pseudolynchia, 1247 casei (Linnaeus), Piophila, 865, 867–868
brunneina Marshall & Buck, Syringogaster, 844–845 Cataclinusa Schmitz, 731, 734, 750, 754
brunnescens Collin, Platypeza, 716 catharinensis Rafael, Allomethus, 800–801
bufonivora Lopes & Vogelsang, Lepidodexia, 1330 Catharosia Rondani, 1382, 1390, 1400
bulbifacies Papp & Silva, Minilauxania, 992 Cattasoma Reinhard, 1319
bulbosa Clausen, Hyadina, 1229 caucensis Steyskal, Melanagromyza, 1069
busckii Coquillett, Drosophila, 1197, 1204 caudatus Borgmeier, Dinocercus, 754
Butomomyza Nowakowski, 1058, 1067–1068 caviceps Borgmeier, Cyphocephalus, 754
Cacoceria Hull, 773, 787 Cecidochares Bezzi, 910, 912, 918, 931, 937, 948
Cadrema Walker, 1139–1140, 1149 Celatoria Coquillett, 1348, 1379, 1389, 1400
Caenoriata Foote, 918, 927, 948 celeste (Curran), Trivialia, 994
caenosa (Rondani), Leptocera, 1184 cellularis Blanchard, Ortalidina, 1005
caerula (Malloch), Melanagromyza, 1069 Celypholauxania Hendel, 983, 991
caerulea (Macquart), Euphara, 883, 890, 898 Cenchrometopa Hendel, 888, 890
caerulescens Hendel, Setellida, 875, 878, 904 Cepa Thompson & Vockeroth, 764, 773, 787
caesar (Aldrich), Eumea, 1404 Cephalella Malloch, 976, 991, 995
Calamiscus Borgmeier, 733, 737, 754, 759 Cephaliini, 882, 888, 892
calcitrans (Linnaeus), Stomoxys, 1282, 1284, 1293–1294 Cephalops Fallén, 796–797, 801–803
californica Capelle, Loxocera, 841 Cephalopsini, 794
californicum Blanc, Chetostoma, 944, 948 Cephalosphaera Enderlein, 796–798, 801–802
Callainireinwardtia Savage, 1286, 1292 Cephenemyia Latreille, 1305–1311
Callicera Panzer, 766, 768, 785, 787, 792 Ceracia Rondani, 1360, 1372, 1398, 1401
Calliopum Strand, 972, 977–978 Cerantichir Enderlein, 816, 818–819, 823
Calliphora Robineau-Desvoidy, 1297–1299, 1301–1303 Ceratitis MacLeay, 909–910, 914, 920, 925, 927, 948
Calliphorinae, 1297, 1299–1302 Ceratodacus Hendel, 909, 914–915, 927
Calliphorini, 1301 Ceratophya Wiedemann, 770
Calliscia Steyskal, 1022 Ceratusa Borgmeier, 728, 730, 754
Callomyia Meigen, 717–719 Ceriana Rondani, 764, 768, 785, 787
Callomyiinae, 716 Ceriogaster Shannon, 769, 785, 790–791
Calodexia Wulp, 1352, 1355, 1379, 1389, 1397, 1400, 1414 Cerioides Rondani, 787–790
Calolydella Townsend, 1380, 1400 Cerodontha Rondani, 1058–1059, 1061, 1063, 1066–1068, 1070
caloptera (Bigot), Automola, 878 Cerometopum Cresson, 1222, 1228
caloptera Hendel, Chaetocoelia, 991 Ceromya Robineau-Desvoidy, 1374, 1401, 1415
Calosphen Hennig, 810–812 Ceropsilopa Cresson, 1214, 1217, 1228
Calotarsa Townsend, 717–719 Chaetaspidia Borgmeier, 732, 734–735, 754
Calycomyza Hendel, 1062–1063, 1066–1068, 1070–1071 Chaetochlorops Malloch, 1141–1142, 1149–1150, 1152–1153
Camilla Haliday, 1193–1195 Chaetoclusia Coquillett, 1043, 1047–1048
Campichoeta Macquart, 1207–1210 Chaetocnemistoptera Borgmeier, 730, 754
Campiglossa Rondani, 914, 923–924, 927, 945, 948 Chaetocoelia Giglio-Tos, 979, 991
Camptoprosopella Hendel, 972, 977, 980, 984, 991, 995 Chaetodoria Townsend, 1373, 1401
Campylocheta Rondani, 1353, 1391, 1400 Chaetogaedia Brauer & Bergenstamm, 1347, 1365, 1388, 1401
Canacea Cresson, 1115–1119 Chaetolonchaea Czerny, 858, 862
Canaceoides Cresson, 1116, 1118–1119 Chaetominettia Malloch, 979, 991
Canacinae, 1118 Chaetona Wulp, 1360, 1379, 1401–1402, 1414
Canacini, 1118–1119 Chaetonodexodes Townsend, 1379, 1401
canariensis (Macquart), Pseudolynchia, 1245–1247 Chaetonopsis Townsend, 1376, 1396, 1401
caniceps Cresson, Ilythea, 1229 Chaetophaonia Carvalho & Nihei, 1291–1292
canicularis (Linnaeus), Fannia, 1277–1279 Chaetopleurophora Schmitz, 754, 756
canospila Hendel, Melanoloma, 873, 878 Chaetopodella Duda, 1177, 1184
capitata (Wiedemann), Ceratitis, 909–910, 914, 920, 927, 948 Chaetopsis Loew, 881–884, 887, 890, 898, 900
carambolae Drew & Hancock, Bactrocera, 910, 926, 947 Chaetostigmoptera Townsend, 1372, 1374, 1381, 1383, 1401
Carcelia Robineau-Desvoidy, 1346, 1360, 1362–1363, 1367, 1400 Chaetotylus Hendel, 807
Cardiacephala Macquart, 806–807 Chalarinae, 793–795
Carninae, 1127 Chalarus Walker, 794–796, 801–802
Carnus Nitzsch, 1101–1104, 1108 Chalcosyrphus, 767, 774, 785, 787
TAXONOMIC INDEX TO VOLUME 2  1423

Chamaebosca Speiser, 1074–1076, 1079 Clausicella Rondani, 1351, 1354, 1396, 1402
Chamaemyia Meigen, 997, 1000–1001, 1004, 1006–1007 clavatus Fabricius, Pseudodoros, 766, 789
Chamaemyiinae, 1001–1002 clavipes (Fabricius), Ropalomera, 1026
Chamaemyiini, 1001–1002 clavipes Hendel, Coscinum, 898
chapmani (Curran), Scutops, 1092 Cliobata Enderlein, 808, 810–812
charactophalla Tanasijtshuk, Leucopis, 1005 Clusia Haliday, 1042–1044, 1047–1048
Charadrella Wulp, 1283, 1292 Clusiinae, 1041, 1043, 1045, 1048
Cheilosia Meigen, 763, 765, 767, 772, 774, 787, 791 Clusiodes Coquillett, 1041–1045, 1047–1048
Chespiritos Marshall, 1173, 1175, 1184, 1186 Clusiodinae, 1041, 1043–1044, 1048
Chetogena Rondani, 1347, 1364, 1368, 1373, 1383, 1401 Cnemospathidini, 1160–1161
Chetostoma Rondani, 914, 927, 929, 944, 948 coccidarum Stackelberg, Turanodinia, 1051
chilensis Cresson, Nocticanace, 1119 cocciphila (Brues), Syneura, 736, 757
Chlorohystricia Townsend, 1297, 1368, 1394, 1401 cochenillivorus Guérin-Méneville, Salpingogaster, 789
Chloronesia Townsend, 1316, 1326–1327, 1330 Cochliomyia Townsend, 1297–1300, 1302–1303
Chloropinae, 1138–1139, 1149, 1151–1153 coeca Nitzsch, Braula, 1097–1098
Chloroprocta van der Wulp, 1299, 1302–1303 Coelomyia Haliday, 1278
Chlorops Meigen, 1137–1138, 1148–1149, 1151 Coelopa Meigen, 1013–1016
Chlorosarcophaga Townsend, 1316, 1327, 1330 Coelopina Malloch, 1014–1016
Choeteprosopa Macquart, 1365, 1401 Coelopinae, 1015
Cholomyia Bigot, 1351, 1385, 1387, 1401 Coelopini, 1015
Chondrometopum Hendel, 889, 892–893, 896, 898 Coenosia Meigen, 1287, 1289–1290, 1292
Chonocephalus Wandolleck, 727, 734, 750–751, 754, 760 Coenosiinae, 1282, 1295
Chortinus Aldrich, 1286, 1292, 1294–1295 Coenosopsia Malloch, 1271–1273, 1275–1276, 1280, 1295
Choryeuromyia Gaimari & Silva, 975, 990–991 coeruleithorax Williston, Habromyia, 787
Chrysagria Townsend, 1326, 1329, 1334 Coilometopia Macquart, 876–878
chryserea Prado, Lenkokroeberia, 1026–1027, 1029 collaris Loew, Loxocera, 838, 841
Chrysidimyia Hull, 770 columbiae Brauer & Bergenstamm, Actinochaeta, 1398
Chrysoexorista Townsend, 1346, 1362–1363, 1401 Comasarcophaga, 1316, 1325–1326, 1329
Chrysomya Robineau-Desvoidy, 1299–1300, 1302–1304 Commoptera Brues, 736, 750, 753–754, 761
Chrysomyinae, 1297, 1299, 1301–1302 compressiceps Borgmeier, Lecanocerus, 733, 756
Chrysomyini, 1300–1301, 1303 Compsomyiops Townsend, 1298, 1302–1303
Chrysops Meigen, 1404 concinnata (Meigen), Compsilura, 1344, 1403, 1415
chrysostoma (Wiedemann), Peckia, 1315, 1331 confusa (Albuquerque), Anthomyia, 1275
Chrysotachina Brauer & Bergenstamm, 1297, 1364, 1368, Coniceps Loew, 872, 874
1370–1371, 1401, 1404, 1415 Conicera Meigen, 716, 726, 730, 754
Chrysotoxum Meigen, 767, 777–778, 780, 787 Conicerinae, 727
Chydaeopeza Shatalkin, 716 Coniceromyia Borgmeier, 730, 754, 758, 761
Chyliza Fallén, 837–842 conifer (Hendel), Plocoscelus, 806
Chylizinae, 837, 839 Conioscinella Duda, 1126, 1138, 1143, 1146, 1149
Chymomyza Czerny, 1198–1201, 1203–1206 connexa (Macquart), Cecidochares, 910, 948
Chymophila Macquart, 770 Conopinae, 848–849, 855, 955
Chyromya Robineau-Desvoidy, 1155–1157 Conops Linneaus, 849–850, 854–855
ciliata (Williston), Lenkokroeberia, 1029 consanguinea Schiner, Chyliza, 838, 842
cimiciforme Loew, Notogramma, 889–890, 899 contigua (Fabricius), Neominettia, 992
Cinderella Steyskal, 1160–1164 contigua Macquart, Sphaerophoria, 790
Cinderellini, 1161 convexifrons (Borgmeier), Chaetaspidia, 732, 754
cinerea (Fallén), Zaira, 1414 Cootiphora Brown, 732, 735–736, 754
Cinochira Zetterstedt, 1344, 1393, 1402, 1407 Copecrypta Townsend, 1346, 1357, 1362, 1402, 1413
Cipomyia Norrbom & Prado| Cipomyia, 938 Copestylum Macquart, 763, 769, 779, 783–785, 787, 791
Cirrula Cresson, 1223, 1226, 1228, 1233 Coproica Macquart, 1173–1174, 1184, 1187
Citrago Schmitz, 758 Copromyza Fallén, 1169, 1171, 1184, 1187
Cladiscophleps Enderlein, 877 Copromyzinae, 1165–1166, 1168, 1171, 1176, 1182, 1185
Cladochaeta Coquillett, 1197, 1200, 1203–1204, 1206 coquilletti Hendel, Physoclypeus, 973, 992
Clairvillia Robineau-Desvoidy, 1393, 1402, 1407 Cordilura Fallén, 1267–1269
Clanoneurum Becker, 1217 Cordiluroides Albuquerque, 1281, 1290, 1292, 1295
Claraeola Aczél, 801 Cordyligaster Macquart, 1377, 1396, 1402, 1415
Clasiopella Hendel, 1214, 1217, 1228, 1233 cornicina (Fabricius), Neomyia, 1285, 1294
Clastoneura Aldrich, 1412 cornuale Reinhard, Crocinosoma, 1351, 1402
clathratus (Hendel), Neotraginops, 1053 cornuta (Bigot), Dohrniphora, 726, 732, 736, 758, 760
clausa (Macquart), Physiphora, 883, 885, 899 corollae (Fabricius), Chaetominettia, 991
1424 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

coronata (Broadhead), Sciosapromyza, 993 cynocephala Kertész, Placopsidella, 1231–1232

coronata (Loew), Pelomyia, 1109–1110 Cyphocephalus Borgmeier, 750, 754
coronata Sabrosky, Odinia, 1053 Cyrtomostoma Hendel, 886, 894
corrigiolata (Linnaeus), Micropeza, 806 Cyrtona Séguy, 1189, 1191
cortesi Spencer, Liriomyza, 1069 Cyrtoneurina Giglio-Tos, 1287, 1292, 1295
Coryphisoptron Enderlein, 1146–1147, 1149, 1151 Cyrtoneurininae, 1282, 1294–1295
Coscinum Hendel, 894, 898 Cyrtoneuropsis Malloch, 1287, 1292
Costalima Sabrosky, 1125–1126, 1130–1132, 1134 Cyrtophleba Rondani, 1349, 1383, 1385, 1402
costalis (Curran), Xeniconeura, 973, 985, 994 cyrtorchidis Spencer, Melanagromyza, 1069
costalis Gerstaecker, Diacrita, 891, 898 Cyzenis Robineau-Desvoidy, 1363, 1376, 1402, 1415
costalis Hendel, Coilometopia, 878 czernyi Collin, Odinia, 878, 1051
Costarica Mengual & Thompson, 1331, 779, 786 czernyi Hendel, Oedematella, 878
costaricensis Hennig, Diastata, 1262, 1210 Dacinae, 910
costaricensis Spencer, Phytoliriomyza, 1070 Dacnophora Borgmeier, 740–741, 754
costaricensis Steyskal, Teutoniomyia, 1023 Dactylissa Fischer, 1026, 1029
costata Wulp, Acaulona, 1398 Dagini, 1225, 1228, 1232
Craspedochaeta Czerny, 1041, 1043–1044, 1046–1048 Dagus Cresson, 1220, 1226, 1228, 1233
Craspedochaeta Macquart, 1275 Dahlimosina Roháček, 1174–1175, 1185
crassata Garrett, Pseudoleria, 1160, 1163 Dahliphora Schmitz, 749, 754
crassipalpis Macquart, Sarcophaga, 1332 Dalmanniinae, 848–849
crassipes Curran, Pyrgotoides, 923, 951 dampfi (Hall), Blaesoxipha, 1329
crassipes Speiser, Allobosca, 1244 dampfi Sabrosky, Pholeomyia, 1126
crassitibia Aczél, Stenopyrgota, 960 Dasiopinae, 859
Crataerina Olfers, 1243–1247 Dasiops Rondani, 857–863
Cremersia Schmitz, 737–739, 754 Dasydorylas Skevington, 796, 801
Cremifania Czerny, 997–1002, 1006 Dasymetopa Hendel, 892–894, 897–898, 900
Cremifaniinae, 1001–1002 Dasymorellia Malloch, 1293
crenata (Bigot), Calythea, 1275 Dasyrhicnoessa Hendel, 1110, 1112–1113
crenulata (Wulp), Dictyotrypeta, 948 Dasysyrphus Enderlein, 778, 780, 787, 792
Crepidomyia Shannon, 774, 790 Deceia Malloch, 988
cressoni Hull, Cacoceria, 787 decempunctata Hendel, Rhyparella, 892, 896, 900
cressoni Lizarralde de Grosso, Mimapsilopa, 1217 declinata (Hall), Halliosca, 1330
Cressonomyia Arnaud, 1214, 1217, 1228, 1233 decrepita Hendel, Melanoloma, 871
cribratus (Enderlein), Sphecomyiella, 960 Dejeaniops Townsend, 1357, 1402
crinicornis Borgmeier, Ceratusa, 754 Delia Robineau-Desvoidy, 1272–1275
Criorhina Meigen, 774, 785, 787 delicatus Machado-Allison, Neotrichobius, 1257
crocata Reinhard, Macromya, 1360, 1408 Delininae, 1267–1268
crocea Borgmeier, Styletta, 757 Delphinia Robineau-Desvoidy, 882, 887–888, 891, 898, 902
Crocinosoma Reinhard, 1351, 1396, 1402 demetrius McAlpine, Librella, 1123
Cryogonus Cresson, 807, 812 dentipes (Fabricius), Hydrotaea, 1293
Cryptochetum Rondani, 1121–1123 Deopalpus Townsend, 1357, 1402, 1413
Cryptodacus Hendel, 912, 914, 922, 927–928, 944, 948, 954 depleta (Wiedemann), Ormia, 1410
Cryptomeigenia Brauer & Bergenstamm, 1348, 1378, Dermatobia Brauer, 1305–1311
1401–1402, 1414 Descoleia Aczél, 953, 955, 958
Cryptopalpus Rondani, 1358, 1402 Desmometopa Loew, 1125–1127, 1130–1131, 1134–1136
Cryptotreta Blanc & Foote, 930, 938 Dete McAlpine, 823
Cuculomyia Lopes, 1332, 1334 Deutominettia Hendel, 979, 991
cuprina (Wiedemann), Lucilia, 1298–1299 devia Spencer, Calycomyza, 1066
Curiosimusca Rung et al., 1084–1085 devians Hennig, Silba, 862
cursor (Kieffer), Chamaebosca, 1076, 1079 Dexiinae, 1343–1345, 1356, 1404, 1408, 1411
curtifrons Borgmeier, Macrocerides, 739, 756 Dexiini, 1345, 1356, 1372, 1408
Curtonotum Macquart, 1189–1192 Dexomyophora Townsend, 1322
Cuterebra Clark, 1305–1311 Dexosarcophaga Townsend, 1316, 1321, 1325–1326, 1328, 1330,
Cuterebrinae, 1306 1333–1334
Cyamops Melander, 1083, 1085, 1088, 1090–1092 Diacrita Gerstaecker, 881–882, 889, 891, 898, 901
Cyclopidiinae, Diastata Meigen, 1207–1210
cylindrica Say, Loxocera, 838 Diathoneura Duda, 1197, 1201, 1203–1204, 1209
Cylindromyia Meigen, 1390, 1402, 1406, 1414–1415 dichaetus Steyskal, Boreothrinax, 960
Cymatosus Enderlein, 893, 896, 898–899, 901 dichromata (Snow), Acidogona, 947
cynipsea (Linneaus), Sepsis, 1032, 1039 Dictya Meigen, 1019, 1021–1022, 1024
TAXONOMIC INDEX TO VOLUME 2  1425

Dictyodes Malloch, 1019, 1023 Echidnocephalodes Sabrosky, 1085

Dictyotrypeta Hendel, 930, 938, 948 echinata (Wulp), Hystricodexia, 1406
Dideomima Vockeroth, 775, 779, 787, 792 Echinoleucopis Gaimari & Tanasijtshuk, 1000–1003
Diedrops Mathis & Wirth, 1220, 1226, 1228, 1233 Ecitomyia Brues, 742, 751–752, 755, 757
Dimecoenia Cresson, 1223, 1226, 1229, 1233 ecitonis (Townsend), Phasia, 1400
dimidiata (Hendel), Diastata, 1210 Ecitophora Schmitz, 751, 753, 755
diminuta Marinoni & Zumbado, Neuzina, 1018–1019, 1022 Ecitoptera Borgmeier & Schmitz, 751–752, 755, 758
Dinocercus Borgmeier, 746–747, 754 Ecituncula Schmitz, 751, 755, 757
Diocophora Borgmeier, 738, 741, 744, 755, 757 Ectecephala Macquart, 879, 1148–1149, 1152
Diopsosoma Malloch, 1090–1091 egregia (Gerstaecker), Euolena, 875, 878
Dioxyna Frey, 914, 930, 932, 945, 948–949, 953 Eibesfeldtphora Disney, 744, 756, 759
Diphuia Cresson, 1218, 1222, 1229, 1233 Elachiptera Macquart, 1140, 1142, 1149–1150, 1152–1153
Diplonevra Lioy, 728–729, 743, 755, 760 Elachisoma Rondani, 1179
Diplotoxa Loew, 1148–1149, 1151 Eldunnia Curran, 1251–1252, 1257
Discocerina Macquart, 1211, 1218, 1221, 1229, 1232 electa (Say), Zonosemata, 952
Discocerinini, 1221 electra Hennig, Protoaulacigaster, 1085
Discogastrella Enderlein, 1152 electrus Grimaldi & Mathis, Planinasus, 1092
discolor (Bigot), Archisepsis, 1033, 1035, 1038 elegans Hendel, Ophthalmoptera, 899
Discomyza Meigen, 1217 Elliponeura Loew, 1149
Discomyzinae, 1212, 1216 Elmohardyia Rafael, 794, 796, 798–802
Distichona Wulp, 1346, 1361, 1403 eluta Loew, Euxesta, 882, 890, 899, 901
distinctissima (Schiner), Chaetocoelia, 991 Embiomyia Aldrich, 1412
Ditaeniella Sack, 1019, 1021–1022 Emblemasoma Aldrich, 1316, 1320–1321, 1330, 1334
Ditrichophora Cresson, 1221 Emdenimyia Lopes, 1315, 1323, 1326, 1330, 1334
diversa (Giglio-Tos), Neorhinotora, 1159, 1161, 1163 Emmesomyia Malloch, 1272, 1274–1276
diversa Prado, Lopesiodinia, 1053 enderleini (Hennig), Cerantichir, 816, 819
Dizygomyza Hendel, 1058, 1066–1068, 1070 Enderleiniella Becker, 1146, 1149–1150
Dohrniphora Dahl, 726, 728, 732, 736, 755, 758–761 Enderleinphora Disney, 731, 755
Dolichocodia Townsend, 1381, 1403 Engelimyia Lopes, 1325, 1330, 1334
Dolichophaonia Carvalho, 1294 Enigmocarnus Buck, 1103–1104
Dolichotarsus Brooks, 1380, 1389, 1397, 1403 ensifer Bezzi, Maenomenus, 956
dolini Kameneva, Vladolinia, 900 Ensina Robineau-Desvoidy, 916, 930, 945, 948
domestica Linnaeus, Musca, 1281–1282, 1287, 1293 Eoloxozus Aczél, 818
dorsalis (Coquillett), Myiopharus, 1371, 1408–1409 Eoneria Aczél, 817–818
dorsalis (Loew), Cerodontha, 1058–1059, 1061, 1068 Eopseudopomyza Hennig, 823–824
dorsalis (Wulp), Macromya, 1408–1409 Eosalpingogaster Hull, 778, 789
Dorsilopha Sturtevant, 1218, 1229–1230, 975, 995, 1110, 1112, Eoseristalis Kanervo, 769, 787
1197, 1200, 1204, 1206 Epalpus Rondani, 1359–1360, 1395, 1398, 1403, 1406–1407
Dorylomorpha Aczél, 796–798, 800–802 Ephydra Fallén, 1211, 1223, 1226–1229, 1233
Dorylomyia Albrecht, 797–798, 800–801 Ephydrinae, 1212–1213, 1232–1233
Doryphorophaga Townsend, 1409 epilachnae Aldrich), Paradexodes, 1408
Dracontomyia Becker, 930, 937, 949 Epiplatea Loew, 872–874, 878
dreisbachi Reinhard, Eudexia, 1403 Epiplateinae, 871, 873–874
Drino Robineau-Desvoidy, 1352, 1366, 1383, 1403 Epischnomyia Roháček, 1074
Drosophila Fallén, 1197–1201, 1203–1204, 1206 Epochrinopsis Hering, 916, 930, 938, 943
Drosophilinae, 1200, 1202, 1204 eques (Wiedemann), Pseudodexia, 1411
Druciatus Marshall, 1172–1173, 1176, 1178, 1184, 1186–1187 equina Fallén, Copromyza, 1169, 1184
Drymeia Meigen, 1281, 1289–1290 Eremophora Borgmeier, 744, 755
Dryomyzothea Hendel, 979, 991 Eribella Mesnil, 1348, 1378–1379, 1403
Dryosapromyza Hendel, 981 Eribolus Becker, 1146
Duckemyia Kano & Lopes, 1315, 1322, 1330 Erioischia Lioy, 1275
dumalis Reinhard, Phobetromyia, 1410 Eristalinae, 764–765, 779
durangoensis Kessel & Young, Calotarsa, 719 Eristalis Latreille, 769, 786–787, 792
Dynomiellini, 1118–1119 Eriurgus Hendel, 983
Dyscrasis Aldrich, 881, 887–888, 891, 901 Ernestia Robineau-Desvoidy, 1410
Dyseuaresta Hendel, 930, 932, 934, 949 Eryciini, 1344–1345, 1356–1357, 1361
Earomyia Zetterstedt, 858–859, 862 Erythrandra Brauer & Bergenstamm, 1320
Earomyiini, 859 erythrocephalum (Schiner), Sepsisoma, 873
Ebenia Macquart, 1349, 1372, 1392, 1403, 1410 Erythromelana Townsend, 1360, 1375, 1381, 1403
Eccoptomma Becker, 1127 Ethillini, 1345, 1356
1426 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Euacaina Steyskal, 882, 884, 889 Euxestina Curran, 898

Euantha Wulp, 1382, 1384, 1403 Euxestinae, 882
Euaresta Loew, 916, 924, 931, 933, 945, 949, 954 eva Townsend, Eujuriniodes, 1404
Euarestoides Benjamin, 933, 946, 949, 952 evanescens Marshall & Buck, Lepidosina, 1184
Euarestopsis Hering, 918, 933, 949 Evibrissa Rondani, 1405
Eucelatoria Townsend, 1344, 1360, 1375–1376, 1380, 1389, 1403, Exastinion Wenzel, 1254–1255, 1257
1407, 1409, 1416 exigua Wulp, Psila, 841
Eucelatoria Townsend, 1344, 1360, 1375–1376, 1380, 1389, 1403, eximia (Wiedemann), Lucilia, 1299
1407, 1409, 1416 Exoristinae, 1344–1345, 1356
eucephala (Loew), Pseudocalliope, 993 Exoristini, 1344–1345, 1356, 1372, 1413, 1417
Eucheirophaga James, 1368, 1403 Exoristoides Coquillett, 1368, 1402, 1404
Euclytia Townsend, 1397, 1403 exotica Carvalho & Couri, Agenamyia, 1291
Eudejeania Townsend, 1357, 1403, 1416 externa (Fabricius), Euscopoliopteryx, 1404
Eudexia Brauer & Bergenstamm, 1384, 1403, 1411, 1413 exul (Williston), Neodeceia, 992
Eudorylas Aczél, 796, 799–803 exul Buck, Inbiomyia, 1107–1108
Eudorylini, 794, 797, 801–802 facialis (Aldrich), Souzalopesiella, 1301
Euepalpus Townsend, 1358, 1404 fairmanorum Fairman, Sericomyia, 790
Eufletcherimyia Townsend, 1322 Fannia Robineau-Desvoidy, 1277–1280
Eugaurax Malloch, 1142, 1150, 1153 farinosus Hendel, Physoclypeus, 972, 995
Euhalidaya Walton, 1352, 1378, 1397, 1404 fasciata (Wiedemann), Pterotaenia, 882
Eujuriniodes Townsend, 1358, 1404 fascipennis Coquillett, Scutops, 1092
Eulasiona Townsend, 1349, 1372, 1375, 1404 Fazia Shannon, 778, 786
Eulasiopalpus Townsend, 1357, 1404 femorata (Fabricius), Ropalomera, 1025–1029
Eulobomyia Woodley & Arnaud, 1365, 1404, 1417 ferrisi Schuurmans-Stekhoven, Basilia, 1261
eulophus (Coquillett), Elachiptera, 1150 ferruginea (Lamb), Dasyrhicnoessa, 1113
Eumacronychia Townsend, 1314, 1316, 1319, 1330, 1334 ferruginea Macquart, Odontomera, 872, 875, 878
Eumea Robineau-Desvoidy, 1365, 1404 ferrumequinum Hendel, Axiologina, 889–890, 898
Eumecosomyia Hendel, 882, 884–885, 898, 900, 902 festinans (Aldrich & Webber), Frontiniella, 1404
Eumerus Meigen, 765, 772, 775 ficulnea (Broadhead), Sciosapromyza, 993
Eumesembrinella Townsend, 1301 Fiebrigella Duda, 1140, 1142, 1149–1150
Eumesembrinellini, 1301 figurata Marshall & Buck, Syringogaster, 845
Eumetopiella Hendel, 882, 884–885, 898 flava (Coquillett), Euclytia, 1403
Eumyiolepta Shannon, 788 flava (Linnaeus), Chyromya, 1155–1156
Eunemorilla Townsend, 1365 flavescens Wheeler, Laccodrosophila, 1205
Euolena Loew, 875–878 flavicauda Townsend, Euepalpus, 1404
Eupeodes Osten Sacken, 778, 780, 787, 792 flavicornis Aldrich, Leucopis, 1005
Euphara Loew, 883, 888, 890, 898 flavida (Wiedemann), Pachyopella, 992
Euphasiopteryx Townsend, 1410, 1416 flavifacies Malloch, Asilostoma, 991
Euphranta Loew, 919, 921, 933 flavifrons (Bigot), Glyphidops, 816, 819
Eurithia Robineau-Desvoidy, 1410 flavifrons (Duda), Steleocerellus, 1152
Eurybatinae, 805–807, 812–813 flavifrons (Tonnoir & Malloch), Prosopantrum, 1161
Eurycephalomyia Hendel, 882 flavigenis (Hendel), Tethina, 1112
Eurychaeta Brauer & Bergenstamm, 1300 flavipennis (Fabricius), Xenochaetina, 994
Eurychoromyia Hendel, 975, 990 flavipes (Williston), Zeros, 1225, 1231
Eurychoromyiinae, 971–972, 975–976, 993–994 flaviseta (Aldrich), Peltodasia, 956
Eurycnemis Borgmeier, 729, 750, 755 flavissima Foote, Procecidocharoides, 950
Euryhendelimyia Gaimari & Silva, 990 flavolimbata (Bigot), Triodontopyga, 1413
Euryomma Stein, 1277–1280 floridensis (Aldrich), Ravinia, 1332
Eurystratiomyia Gaimari & Silva, 990 floridensis (Aldrich), Rhytidops, 1026–1027
Euscopoliopteryx Townsend, 1382, 1387, 1404 Floriminettia Broadhead, 993–994
Eusiphona Coquillett, 1125, 1127–1129, 1134, 1136 foliata Krivosheina, Odinia, 1051
Eustalomyia Kowarz, 1272, 1274–1275 footei Korytkowski, Polymorphomyia, 950
Euthera Loew, 1350, 1382, 1391, 1404, 1414 fraterculus (Wiedemann), Anastrepha, 947
Euthycera Latreille, 1018–1019, 1021, 1024 Frendelia Collin, 972, 981, 983, 992
Euthychaeta Loew, 1209 Freyia Malloch, 984, 991
Eutreta Loew, 910, 916, 931–933, 938, 946, 949, 954 frigida (Fabricius), Coelopa, 1013–1015
Eutretopsis Norrbom & Prado, 938 frontalis Borgmeier, Neophora, 733, 748, 756
Eutrichota Kowarz, 1272–1275 frontalis Macquart, Sapromyza, 994
Eutrixa Coquillett, 1351, 1386, 1404, 1406 Frontiniella Townsend, 1363, 1376, 1404, 1415
Euxesta Loew, 881–883, 886, 890, 898–902 Frutillaria Richards, 1182
TAXONOMIC INDEX TO VOLUME 2  1427

Fucellia Robineau-Desvoidy, 1272–1273, 1275 Gonioneura Rondani, 1179, 1184

fuliginosa Lindner, Kroeberia, 1026–1029 gracilis Packard, Ephydra, 1229
fulvida Bigot, Megamerina, 845 Grallipeza Rondani, 810–812
fumicosta Malloch, Chamaemyia, 1004 Grallomyia Rondani, 809, 812
fumifera Townsend, Phasiophyto, 1410 grandis (Cresson), Placopsidella, 1231
fumipennis (Sahlberg), Olfersia, 1242, 1247 granulosa (Cresson), Athyroglossa, 1228
fumipennis Malloch, Cephalella, 991 granulosus Cresson, Lytogaster, 1230
funebris (Fabricius), Drosophila, 1197 Graphomya Robineau-Desvoidy, 1288, 1292
Fungomyza Roháček, 1074, 1081 Grimalantha Roháček, 1074
furcata (Fabricius), Centrophlebomyia, 866–867, 869 Griphoneura Schiner, 979, 985, 991
furcata (Fallén), Lonchoptera, 721–722 grisea (Fallén), Tethina, 1112
furcata (Say), Scathophaga, 1267–1269 griseola Rao & Mathis, Rhysophora, 1231
Furciseta Aczél, 963–968 grisescens Hendel, Procanace, 1117
fusca Gaimari, Neoschildomyia, 1053 Grossovena Kessel & Maggioncalda, 716
fusca Spencer, Agromyza, 1063, 1067 grunini Ozerov, Protothyreophora, 867
fuscanipennis (Macquart), Chloroprocta, 1303 guajiro (Garcia & Casal), Strebla, 1250, 1252
fuscisquama Wulp, Phorbia, 1293 guatemala Snyder, Lispoides, 1293
fuscohalterata (Borgmeier), Enderleinphora, 755 guatemalana Steyskal, Dictya, 1022
Gaediopsis Brauer & Bergenstamm, 1346, 1362, 1404 guatemalana (Steyskal), Sepedonea, 1023
gaimarii Filho, Esposito & Santos, Helgreelia, 1049, 1053–1054 guatemalensis Sasakawa, Phytobia, 1069
Galiomyza Spencer, 1067, 1070 Guatemalia Steyskal, 1022, 1024
Garifuna Mathis, 1213, 1229 guerinii (Bigot), Macrostenomyia, 875, 878
Gasterophilinae, 1306 gulo (Fabricius), Peckia, 1313, 1316, 1328, 1331, 1334
Gasterophilus Leach, 1305–1310 Guttipsilopa Wirth, 1217
Gastrops Williston, 1215, 1219, 1228–1229 Gymnocamptops Townsend, 1329
Gastropsini, 1219 Gymnocarena Hering, 916, 931, 933, 943, 949, 953–954
gatesi Thompson, Eristalis, 787 Gymnochiromyia Hendel, 1156–1157
Gaurax Loew, 1144, 1146–1147, 1150, 1153 Gymnoclasiopa Hendel, 1221
Gayomyia Malloch, 1011 Gymnoclytia Brauer & Bergenstamm, 1353, 1391, 1405
geminata (Fabricius), Setulina, 993 Gymnomma Wulp, 1358, 1405
geminatus Camras, Conops, 854 Gymnomyzinae, 1212, 1216, 1233
genalis (Coquillett), Euhalidaya, 1352, 1404 Gymnomyzini, 1219, 1221
Genea Rondani, 1396, 1404 Gymnophora Macquart, 747, 749, 755, 759
geniculata (De Geer), Siphona, 1412 Gymnoptera Lioy, 730, 738, 748
geniseta (Malloch), Allominettia, 974, 982, 991 Gymnosoma Meigen, 1353, 1377, 1391, 1405
geometroides (Cresson), Calobatina, 806, 813 Gyretria Enderlein, 1176–1177
Gephyromyzini, 1161 Habromyia Williston, 768, 786–787, 792
Geratomyza Spencer, 1068 hadrocara Gaimari & Silva, Physegeniopsis, 992
gertschi Curran, Eulasiopalpus, 1404 hadromera Gaimari, Pradomyia, 1054
Gibbolauxania Papp & Silva, 988, 991 Haematobia Lepeletier & Serville, 1282–1283, 1292
Gigantotheca Townsend, 1316, 1329 haemorrhous Meigen, Paragus, 767, 777, 789
gilvimana (Hennig), Metasphen, 811–812 Halephydra Wirth, 1226, 1233
Ginglymia Townsend, 1354, 1374, 1404 Hallina Lopes, 1324, 1327, 1330
girschneri Röder, Psilopa, 1231 Halliosca Lopes, 1326, 1330
Gitona Meigen, 1199, 1205 Haloscatella Mathis, 1224–1225
glabra (Meigen), Athyroglossa, 1228, 1232 halteralis (Coquillett), Leptometopa, 1134
glabricula (Fallén), Hydrotaea, 1292 hambletoni Steyskal, Eumecosomyia, 898
Glenanthe Haliday, 1218–1219, 1229, 1233 handleyi Wenzel, Parastrebla, 1254, 1258
Globomyia Hennig, 812–813 handlirschi Brauer & Bergenstamm, Bibiomima, 1382, 1399
Globopeza Marshall, 809, 811, 845–846 hansoni Disney, Enderleinphora, 755
globula (Meigen), Strongygaster, 1412 hansruttneri Huttinger, Braula, 1098–1099
Glumini, 1015 Haplopeodes Steyskal, 1060, 1065, 1068, 1071
Glyphidops Enderlein, 816–819 harenosus Cresson, Plocoscelus, 806, 811
Gnadochaeta Macquart, 1350, 1386, 1401, 1405 Harpagopyga Aldrich, 1327
Goedenia Freidberg & Norrbom, 916, 919, 924 harrisinae (Coquillett), Ametadoria, 1347, 1398
Gonia Meigen, 1347, 1361, 1405 Haywardina Aczél, 912, 925, 928, 944, 954
Goniaeola Hendel, 885, 889 Hecamede Haliday, 1212, 1218, 1222, 1229, 1233
Goniaspis Duda, 1139, 1150, 1153 Hecamedini, 1221, 1229
Goniinae, 1344, 1356 Helaeomyia Cresson, 1217
Goniini, 1344–1345, 1356–1357, 1361, 1399, 1416 Heleomyzini, 1160–1162
1428 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Helgreelia Gaimari, 1049, 1053–1054 hyalophora Buck & Marshall, Ischiolepta, 1184
Helicobia Coquillett, 1315–1316, 1321, 1324–1325, 1330, Hyalurgus Brauer & Bergenstamm, 1392, 1405
1333, 1335 hydei Sturtevant, Drosophila, 1197
Heliconiophaga Thompson, 1409 Hydrellia Robineau-Desvoidy, 1211–1213, 1215, 1224, 1227,
Helina Robineau-Desvoidy, 1282, 1286, 1288, 1291–1292 1229, 1232
Heliographa Malloch, 1293 Hydrelliinae, 1212
Hellerella Duda, 1174 Hydrochasma Hendel, 1218, 1221, 1229
Heloclusia Malloch, 821, 823–824 Hydrophoria Robineau-Desvoidy, 1272, 1274–1275
Helosciomyzinae, 1020 Hydropyrus Cresson, 1223, 1226, 1233
helvum (Loew), Curtonotum, 1189, 1191–1192 Hydrotaea Robineau-Desvoidy, 1282, 1284, 1289–1290,
Hemeromyia Coquillett, 1102–1104 1292–1293
Hemichlora Wulp, 1286, 1292 Hylemyioide Albuquerque, 1275
Hemilucilia Brauer, 1299, 1302–1303 Hypagoga Hendel, 972, 986, 991
Hemisturmia Townsend, 1361, 1405 Hyphantrophaga Townsend, 1362, 1405
hemithorax Frey, Mesoconius, 812 Hypocerides Schmitz, 728, 736, 755
Hemixantha Loew, 875–876, 878 Hypocerinae, 727
Hemyda Robineau-Desvoidy, 1353, 1390, 1405 Hypoderma Latreille, 1305–1310
hendeli Mello & Silva, Physoclypeus, 992 Hypodermatinae, 1306
Hendelia Czerny, 1041, 1043–1044, 1046–1048 Hypotachina Brauer & Bergenstamm, 1369, 1405
heringi Norrbom, Tetreuaresta, 941, 943 Hypselosyrphus Hull, 772
Hershkovitzia Guimarães & D’Andretta, 1261, 1263–1266 Hystricia Macquart, 1368, 1370, 1406
Heteromeringia Czerny, 1041–1044, 1046–1048 Hystricodexia Roeder, 1384, 1403, 1406, 1411, 1413
Hetschkomyia Hendel, 932–933 icarus Kameneva, Cymatosus, 898
Hexachaeta Loew, 915, 935, 949, 953 Icelia Robineau-Desvoidy, 1393, 1406
hilarella (Zetterstedt), Taxigramma, 1332 iceryae (Williston), Cryptochetum, 1121–1123
Himeroessa Loew, 905–906 Ichneumonops Townsend, 1377, 1390, 1406
Hippelates Loew, 1138–1139, 1143, 1145, 1150, 1152–1153 Icosta Speiser, 1242, 1244–1248
Hippelatinae, 1138 Icteromyza Hendel, 1067–1068
Hippobosca Linnaeus, 1244 Idana Loew, 882
Hippoboscinae, 1244 idioidea (Robineau-Desvoidy), Chloroprocta, 1299, 1303
Hirtodeceia Shewell, 978 Idiopyrgota Aczél, 956–960
Hirtodrosophila Duda, 1199–1200, 1203, 1205–1206 ignyodactyla Buck, Loxocera, 838, 840–841
hispida Theodor, Basilia, 1261, 1266 iguassu Steyskal, Protodictya, 1022
Holopterina Borgmeier, 752, 755 illota Haliday, Tethina, 1112
Homalocephala Zetterstedt, 882, 900–901 illustris Spencer, Calycomyza, 1066
Homalomitra Borgmeier, 1167–1169, 1184 Ilythea Haliday, 1213, 1220, 1229
Homalomitrinae, 1165–1166, 1168, 1185, 1187 Ilytheinae, 1212–1213
Homalomyia Bouché, 1278 Ilytheini, 1213
Homalophora Borgmeier, 750, 752, 755 Imantimyia Frey, 837–841
Homaluroides Sabrosky, 1148, 1150 immaculatus (Macquart), Rhachoepalpus, 1411
hominis Linnaeus, Jr., Dermatobia, 1306–1308, 1310–1311 immigrans Sturtevant, Drosophila, 1197
hominivorax (Coquerel), Cochliomyia, 1299–1300, 1303 imox (Bates), Neotaracia, 936, 950
Homoeothrix Hering, 935, 946 impressa (Bigot), Stilbometopa, 1245–1247
Homogenia Wulp, 1387–1388, 1405 inaequalis (Malloch), Lepidosina, 1172, 1184
Homoneura Wulp, 972, 975–977, 980, 982, 985, 989–990, 995 inaequipes Bigot, Cholomyia, 1401
Homoneurinae, 971–972, 974–976, 987, 990 Inbiomyia Buck, 1105-1108
hondurana Steyskal, Epiplatea, 873 inca McAlpine, Prochyliza, 868
Hoplocheiloma Cresson, 807 incanus Reinhard, Isidotus, 1406
Hoplodictya Cresson, 1019, 1022–1024 Incertella Sabrosky, 1146, 1150
horrida Giglio-Tos, Cyrtophleba, 1383, 1402 incisa (Wulp), Dictyotrypeta, 930, 938, 948
Houghia Coquillett, 1362, 1405 incisurata (Zetterstedt), Fannia, 1278
Huapia Richards, 1171 infumata Wheeler, Latheticomyia, 824
Huascaromusca Townsend, 1301 iniens (Giglio-Tos), Suillia, 1163
hubbardii (Coquillett), Minettia, 992 innominata (Williston), Stomosis, 1135
huidobrensis (Blanchard), Liriomyza, 1059–1060, 1069 innuba (Giglio-Tos), Scutominettia, 993
Huttonininae, 1020 inops (Walker), Engelimyia, 1330
Hyadina Haliday, 1219–1220, 1229, 1232 insignis Aldrich, Calotarsa, 718–719
Hyadinini, 1219 insularis (Aldrich), Dasyrhicnoessa, 1110, 1113
hyalina Borgmeier, Eremophora, 755 interjecta (Walker), Setulina, 993
Hyalistata Wheeler, 1200, 1202, 1205–1206 intermedius Kameneva, Plagiocephalus, 899
TAXONOMIC INDEX TO VOLUME 2  1429

intermutans (Walker), Peckia, 1315, 1319, 1331 Latheticomyia Wheeler, 821–825

Lipochaetini, 1219 Latiborophaga Brown, 730, 748, 756
irazuana (Townsend), Calliphora, 1299, 1303 latifasciaeformis Duda, Scaptodrosophila, 1205
irritans (Linnaeus), Haematobia, 1282, 1292 latiforceps Ramírez-García & Hernández-Ortiz,
irroratus Coquillett, Traginops, 1050–1051 Ropalomera, 1029
Ischiolepta Lioy, 1168, 1173, 1184, 1186 latifrons (Hardy), Microcephalops, 801
Ischnomyia Loew, 1075 latifrons (Hendel), Plagiocephalus, 895, 899
ischyri (Couillett), Ischyrophaga, 1406 latipes (Meigen), Desmometopa, 1126
Ischyrophaga Townsend, 1364, 1375, 1406 latipes (Meigen), Leptometopa, 1126, 1130, 1133–1134
Ischyropteron Bigot, 910, 921–922, 935, 952 latipes (Meigen), Protopiophila, 865–866, 869
Isidotus Reinhard, 1386, 1404, 1406 Lauxania Meigen, 974–975, 984, 987, 995
isthmi (Steyskal), Sepedonea, 1023–1024 Lauxaniella Malloch, 977, 984, 986, 992
Itacuphocera Townsend, 1357, 1406 Lauxaniinae, 974–975, 995
Italispidea Townsend, 1375, 1381, 1406 Lecanocerus Borgmeier, 733, 740, 744, 756
Itaplectops Townsend, 1392, 1406 Leiomyza Macquart, 1093–1095
jalisco Woodley, Lydella, 1408 Leiomyzinae, 1094
jamaicensis Spencer, Japanagromyza, 1064, 1068 Lejops Rondani, 767–768, 787
Japanagromyza Sasakawa, 1057, 1061, 1063–1064, 1068, Lemnaphila Cresson, 1213, 1215, 1229, 1233
1070–1071 Lenkokroeberia Prado, 1025–1027, 1029
Jassidophaga Aczél, 795–796, 801 Lepidodexia Brauer & Bergenstamm, 1297, 1313, 1315–1316,
jellisoni Townsend, Cephenemyia, 1310 1319, 1322–1323, 1326–1327, 1329–1330
joblingi Wenzel, Trichobius, 1250, 1259 Lepidomyia Loew, 767, 769, 785, 788
Joblingia Dybas & Wenzel, 1250, 1253, 1255, 1257 Lepidosina Marshall, 1172, 1177, 1184, 1187
Johnrichardia Perez-Gelabert & Thompson, 874 Leptocera Olivier, 1170, 1173–1174, 1182, 1184–1187
johnsoni (Stein), Leucophora, 1275 Leptometopa Becker, 1126–1127, 1130–1131, 1133–1134, 1136
Johnsonia Coquillett, 1313, 1315–1316, 1319, 1327, 1330 Leptopsilopa Cresson, 1214, 1219, 1230
Johowia Silva Figuero, 726, 737, 755 Leptopyrgota Hendel, 956, 958–961, 964
Jurinella Brauer & Bergenstamm, 1358, 1406 Leptostylum Macquart, 1373, 1388, 1399, 1407
Jurinia Robineau-Desvoidy, 1358, 1406 Leschenaultia Robineau-Desvoidy, 1354, 1366, 1388, 1407, 1416
Juriniopsis Townsend, 1343, 1358, 1395, 1406, 1414, 1416 Leskia Robineau-Desvoidy, 1351, 1388, 1396, 1407
kallima (Frost), Phytobia, 1070 Leskiopsis Townsend, 1370, 1412
kerteszi (Enderlein), Pseudohypocera, 726, 748, 757, 759, 761 Lespesia Robineau-Desvoidy, 1363, 1366, 1371, 1407, 1415
kinetrolicros (Caloren & Marshall), Hendelia, 1043, 1047 leuconotum (Wulp), Leptostylum, 1388, 1407
Kirbya Robineau-Desvoidy, 1349, 1376, 1385, 1406 Leucopella Malloch, 1000, 1002, 1006
knudseni (Mayer), Sphecomyiella, 960 Leucophenga Mik, 1198–1199, 1201–1202, 1205
knutsoni Vala et al., Sepedon, 1019, 1024 Leucophora Robineau-Desvoidy, 1272, 1274–1275
korytkowskii Lopes, Emdenimyia, 1330 Leucopina Malloch, 998, 1000–1001, 1003, 1005
Kroeberia Lindner, 1025–1030 Leucopini, 1001–1002
Laccodrosophila Duda, 1199–1200, 1203, 1205–1206 Leucopis Meigen, 998–1007
Laciniomyia Kung & Brown, 735, 742, 755 Leucopodella Hull, 776, 788
lacteipennis (Loew), Milichiella, 1129, 1134 Leucopomyia Malloch, 998, 1000, 1003, 1006
lacunosum Cresson, Cerometopum, 1228 Leucostoma Meigen, 1370, 1385, 1388, 1390, 1402, 1407, 1416
laevifrons (Hendel), Neogriphoneura, 992 leucozona Bilimek, Pholeomyia, 1126
lagoa (Steyskal), Sepedonea, 1018–1019, 1023 leve Hendel, Chondrometopum, 893, 896, 898
Laishania Kung & Brown, 742, 751, 756 Lewinsohnia Norrbom & Prado, 935, 943, 946
lambens (Wiedemann), Sarcodexia, 1316, 1321, 1332 Librella McAlpine, 1123
Lamproclasiopa Hendel, 1221, 1229 Lilloaciura Aczél, 932
Lamprolonchaea Bezzi, 858, 861 limbativertex Enderlein, Scipopus, 806
Lamproxyna Hendel, 921, 931, 935 limitatum Duda, Rhopalopterum, 1152
Lamproxynella Hering, 932, 934–935, 949 Limnellia Malloch, 1225, 1230, 1232
Laneelinae, 1300 Limnophora Robineau-Desvoidy, 1288, 1293
lanei Curran, Sobarocephala, 1048 Limomyza Marshall, 1165, 1176, 1184, 1186
lantanae (Froggatt), Ophiomyia, 1069 Limosininae, 1165–1166, 1168, 1172, 1182, 1186–1187
larvicida (Lopes), Titanogrypa, 1332 Lindigepalpus Townsend, 1359, 1407
lasciva (Fabricius), Taeniaptera, 806 Lindneromyia Kessel, 716–719
Lasiomma Stein, 1272, 1274–1275 lineata Wheeler, Latheticomyia, 823
Lasiona Wulp, 1369, 1406 lineatum (Villers), Hypoderma, 1306, 1310
Lateosepsis Ozerov, 1032, 1034, 1037 linellii Townsend, Protodejeania, 1411
lateralis (Malloch), Gibbolauxania, 991 liniaris Sabrosky, Tucumyia, 1095
lathami (Curran), Calolydella, 1400 Linnaemya Robineau-Desvoidy, 1350, 1367, 1387, 1391, 1407
1430 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Liohippelates Duda, 1138–1139, 1143, 1145, 1150, 1152 Lydella Robineau-Desvoidy, 1345, 1352, 1366, 1408, 1417
Lipara Meigen, 1074, 1151 Lydinolydella Townsend, 1380, 1408
Lipochaeta Coquillett, 1218–1219, 1230, 1233 Lypha Robineau-Desvoidy, 1368, 1370–1371, 1377, 1408
Lipochaetini, 1219 Lytogaster Becker, 1219, 1230, 1232
Lipoleucopis Meijere, 997–999 Macalpinella Papp, 823–824
Lipoptena Nitzsch, 1241–1244, 1246–1247 macateei Malloch, Canacea, 1115–1117, 1119
Lipopteninae, 1244, 1248 macellaria (Fabricius), Cochliomyia, 1298–1300, 1303
Lipsanini, 882–883, 900–901 macrocera (Townsend), Oxynops, 1410
Liriomyza Mik, 1058–1062, 1065, 1067–1069, 1071 Macrocerides Borgmeier, 728, 739–741, 756
Lispe Latreille, 1283–1284, 1293 Macromya Robineau-Desvoidy, 1360, 1369, 1408, 1416
Lispoides Malloch, 1289, 1291, 1293 Macronychia Rondani, 1314, 1318, 1330
Listriomastax Enderlein, 1112 macropus (Walker), Sepedomerus, 1019, 1023
litigata Bonduriansky, Protopiophila, 869 macrosoma Wulp, Charadrella, 1292
littoreus (Hutton), Apetaenus, 1112 Macrostenomyia Hendel, 874–876, 878
litturata (Olivier), Euantha, 1382, 1403 maculata Giglio-Tos, Paragorgopis, 899
livescens Reinhard, Dolichotarsus, 1403 maculata (Spencer), Japanagromyza, 1068, 1070
Lixadmontia Wood & Cave, 1378, 1407 maculineura Borgmeier, Rhyncophoromyia, 757, 759
Lixophaga Townsend, 1345, 1364, 1380, 1388, 1401, 1406–1407 maculosa (Malloch), Nemorimyza, 1069
Lochmostylia Hendel, 963–966, 968 Madiza Fallén, 1130–1132
Lochmostyliinae, 957, 961, 965, 969 Madizinae, 1127
loewii Hendel, Phytomyza, 1070 Madremyia Townsend, 1363, 1408
Loewimyia Sabrosky, 1094–1095 Maenomenus, 956
Lonchaea Fallén, 857, 859–862 Maerorichardia Hennig, 877
Lonchaeinae, 859 Maesia Carvalho & Nihei, 1292
Lonchaeini, 859 magnifica Duda, Metasiphonella, 1151
longicornis (Coquillett), Odontoloxozus, 815–817, 819 magnipalpus (Williston), Neophyllomyza, 1134
longicornis Hendel, Ceratodacus, 909, 914, 927 major Curran, Protodejeania, 1411
longicornis Hendel, Coilometopia, 878 major Hennig, Neopyrgota, 960
longicornis Hendel, Poecilomyia, 876, 878 major (Wulp), Chaetopsis, 882, 887, 890, 898
longicornis Hendel, Siopa, 883, 890, 900 malacophaga Lopes, Charadrella, 1292
longicornis Perty, Thecomyia, 1020 Malacophagomyia Lopes, 1316, 1323, 1331, 1333
Longina Wiedemann, 816, 818–819, 823 Malacophagula Bequaert, 1323, 1331
longipalpis Borgmeier, Holopterina, 755 malerkotliana Parshad & Paika, Drosophila, 1197, 1200
longipalpis sedula Borgmeier, Holopterina, 755 mallochi (Curran), Phaonantho, 1276
longipennis (Cogan), Ortalidina, 1005 mallochi Hendel, Scutominettia, 993
longipes Hendel, Brachydeutera, 1228 mallochi (Sturtevant), Pelomyiella, 1276, 1110, 1112
longipes Hendel, Pyrgota, 960 Mallochianamyia Santos-Neto, 1011
longipes Pessôa & Guimarães, Paraeuctenodes, 1258 Mallochomyza Hendel, 974, 979, 985, 987, 995
Lopa McAlpine, 1015 Malloewia Sabrosky, 1147, 1150–1151
Lopadops Enderlein, 960 Mallota Meigen, 768, 786, 788, 792
lopesi Keiser, Lochmostylia, 963–966 manihoti Lima, Anastrepha, 947
Lopesiodinia Prado, 1053 maracasi Thompson, Thelairodoriopsis, 1412
Lopinae, 1015 Marbenia Malloch, 1090–1091
Lotophila Lioy, 1171 Margdalops Roháček & Barraclough, 1074, 1080–1081
Loxobathmis Enderlein, 1141, 1144, 1147, 1150 Marmarodeceia Shewell, 989, 991
Loxocera Meigen, 837–842 marmorata (Malloch), Marmarodeceia, 991
Loxocerosoma Verbeke, 839 marquesana (Malloch), Placopsidella, 1231
Loxozus Enderlein, 818 marshalli Aldrich, Chaetonodexodes, 1401
Lucilia Robineau-Desvoidy, 1298–1299, 1301–1303 Masiphya Brauer & Bergenstamm, 1354, 1362, 1408
Luciliini, 1301 Masiphyini, 1356, 1408, 1415
lucipetum Roháček & Barber, Stiphrosoma, 1075, 1079 Masoniella Vockeroth, 1110–1114
ludens (Loew), Anastrepha, 920, 947 Mastoptera Wenzel, 1250, 1253, 1255, 1257
lugens Wulp, Pyrgota, 956–957, 959–960 Mauromyia Coquillett, 1368, 1408
lugubris James, Eucheirophaga, 1403 Mayagueza Wheeler, 1202
luteipennis (Curran), Cholomyia, 1401 mazamae (Rondani), Lipoptena, 1243, 1247
lutzi Curran, Jurinella, 1406 mazorca Steyskal, Euxesta, 882, 899, 901–902
Lycastrirhyncha Bigot, 767–768 mcalpinei Marshall, Notenthes, 812
Lyciella Collin, 972, 978 mcalpineorum Buck, Inbiomyia, 1105–1108
Lycopale Hull, 767–768, 786, 788, 792 Medeventor Wheeler, 1140, 1142, 1151
lycopersella Smyth, Rhagoletis, 951 Medina Robineau-Desvoidy, 1352, 1379, 1388, 1394, 1408
TAXONOMIC INDEX TO VOLUME 2  1431

medinai (Snyder), Pilispina, 1294 Metadorylas Rafael, 801–803

medleri Cogan, Afrodinia, 1051 metallica Townsend, Phosocephala, 1411
Megabraula Grimaldi & Underwood, 1098–1099 Metaplagia Coquillett, 1349, 1385, 1408
megacephala (Fabricius), Chrysomya, 1299, 1303 Metasiphonella Duda, 1142, 1151
megaepistoma Sasakawa, Agromyza, 1063, 1067 Metasphen Frey, 806, 810–813, 845
Megalaemyia Hendel, 885, 893, 899 Metasyrphus Matsumura, 778, 792
Megalothoraca Hendel, 877 Metelasmus Coquillett, 1251–1252, 1257–1259
Megamerina Rondani, 845 Metopia Meigen, 1314, 1318, 1328, 1331, 1335
Megaprosopini, 1345, 1356 Metopina Macquart, 736, 750–751, 754, 756
Megaprosopus Macquart, 1386, 1408 Metopininae, 727, 758–759
Megaselia Rondani, 726–727, 737–739, 741, 743–744, 749, 753, Metopocampta Enderlein, 900
755–761 mexicana Cresson, Ephydra, 1229
Megistapophysis Dick & Wenzel, 1255, 1257 mexicana Kessel & Young, Calotarsa, 719
Megistopoda Macquart, 1250, 1253, 1255, 1257, 1259 mexicana Macquart, Mixogaster, 788
meijerei Collin, Odinia, 1051, 1054 mexicana Malloch, Stenopyrgota, 960
Meiosimyza, 972, 978 mexicana Michelsen, Coenosopsia, 1275
melaena Bigot, Siphoniomyia, 1413 mexicana Steyskal, Mexicoa, 1026–1029
Melaleucopis Sabrosky, 998, 1000, 1002, 1005 mexicana Steyskal, Paraleucopis, 1009, 1011
Melaloncha Brues, 726, 731, 756, 759–761 mexicana (Wiedemann), Dyseuaresta, 949
Melanagromyza Hendel, 1057, 1059, 1063–1064, 1067–1071 Mexicoa Steyskal, 1026–1029
Melanderomyia Kessel, 716 meyersi Fluke, Sericomyia, 790
Melanderomyiinae, 716 Microcephalops De Meyer, 797, 801
Melanoclusia Lonsdale & Marshall, 1043, 1048 Microcephalopsini, 794
melanogaster Meigen, Drosophila, 1197, 1200, 1205 Microcerella Macquart, 1326–1327, 1329, 1331
melanoleuca Hennig, Aulacigaster, 1085 Microcerella Townsend, 1326–1327, 1329, 1331
Melanoloma Curran, 871, 873, 876–879 Microdon Meigen, 766, 770–772, 775, 781, 788
Melanomya Rondani, 1301 Microdontinae, 763, 765, 779, 790
Melanomyinae, 1297, 1300–1303 Microdrosophila Malloch, 1198–1201, 1203, 1205
Melanomyini, 1301 Microlynchia Lutz, Neiva, & Costa Lima, 1242, 1246–1247
Melanomyza Malloch, 972–973, 977, 980, 984, 986–987, 992 Micronychiops Townsend, 1376, 1392, 1408
Melanophora Meigen, 1340–1341 Micropeza Meigen, 806–808, 812
melanopyga (Wiedemann), Xanthozona, 1413 Micropezinae, 805–807, 812
Melanostoma Schiner, 767, 771, 776, 787–788 Microphthalma Macquart, 1350, 1386, 1398, 1408, 1411, 1414
melanura (Loew), Acidogona, 914, 947 Micropotamia Carvalho, 1290, 1293–1294
melas (Bigot), Trichophora, 1413 microptera Speiser, Chamaebosca, 1075, 1079
Meleterus Aldrich, 1385, 1408 Micropterocerus Hendel, 900
Melittophora Brues, 731, 756 micropteryx (Thomson), Calythea, 1275
Melophaginae, 1244 Microsania Zetterstedt, 716–719
Melophagus Latreille, 1241, 1243, 1246–1247 Microsaniinae, 716
Menozziola Schmitz, 740, 742, 756 Microsepsis Silva, 1032–1034, 1036–1039
mentula Sasakawa, Phytobia, 1069 Microsimus Aldrich, 1127, 1130–1134
Meoneura Rondani, 1102–1104, 1108 Milesia Latreille, 774–775, 785, 788, 791
mera (Cresson), Orasiopa, 1230 Milichia Meigen, 1126–1128, 1134–1135
meridiana (Hendel), Calycomyza, 1062, 1066 Milichiella Giglio-Tos, 1126–1129, 1134
meridionalis Hennig, Pseudodinia, 1005 Milichiinae, 1127, 1135, 1210
meridionalis Townsend¸ Nausigaster, 788 millibrae Jones, Ephydra, 1229
Merodontini, 764 Miltogramminae, 1313, 1316, 1318, 1335
Meromacrus Rondani, 763, 769, 783, 786, 788, 790–791 mima Townsend, Polistiopsis, 1411
Meromyza Meigen, 1138, 1145, 1148, 1151 Mimapsilopa Cresson, 1217, 1230, 1233
Meropliosepsis Duda, 1032–1034, 1037–1039 Mimocalla Hull, 764, 777, 788, 792
Meroplius Rondani, 1033–1034, 1037, 1039 minensis (Townsend), Lydella, 1408
Meroscinis de Meijere, 1152 Minettia Robineau-Desvoidy, 971–972, 980–983, 985–986, 990,
Mesembrina Meigen, 1283, 1293, 1295 992, 994
Mesembrinella Giglio-Tos, 1297–1299, 1301–1303, 1344 Minettiinae, 974
Mesembrinellinae, 1297, 1300–1302 Minilauxania Papp & Silva, 986, 992
Mesembrinellini, 1301 Minilimosina Roháček, 1172, 1176, 1179, 1181, 1184, 1186
Mesoconius Enderlein, 806, 809, 811–813 minor Marques & Ale-Rocha, Kroeberia, 1029
Mesosphaerocera Kim, 1167–1168, 1184–1185 Minthoplagia Townsend, 1369, 1384, 1409
Metaclythia Kessel, 716 minuta Costa Lima, Mastoptera, 1250, 1257
Metacosmus Coquillett, 793 minuta Hennig, Aulacigaster, 1085
1432 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

mira Coquillett, Eusiphona, 1129, 1134 Neaspilota Osten Sacken, 925

mirabilis Brauer & Bergenstamm, Argyromima, 1394, 1399 nebulifera Sasakawa, Japanagromyza, 1064, 1068
mirabilis (Collin), Opalimosina, 1185 Neivamyia Pinto & Fonseca, 1283
mirabilis (Frey), Hendelia, 1043, 1047 nelsoni Kondratieff & Fitzgerald, Sphecomyiella, 960
mirabilis Giglio-Tos, Ostracocoelia, 936, 950 Nemo McAlpine, 1085
mirabilis Townsend, Ichneumonops, 1406 Nemoraea Robineau-Desvoidy, 1405
miristigma Borgmeier, Megaselia, 757 Nemoraeini, 1356, 1368
Mixogaster Macquart, 769, 771, 775, 788, 791 Nemorilla Rondani, 1361, 1409
mixteca Hernández-Ortiz, Aciurina, 926, 947 Nemorimyza Frey, 1062, 1065–1069
Mochlosoma Brauer & Bergenstamm, 1382, 1409, 1411, 1416 Neoalticomerus Hendel, 1050–1051, 1053
molinai Borgmeier, Apodicrania, 753 Neobrachelia Townsend, 1352, 1392, 1409
molliventris Borgmeier, Apterella, 753 Neocephalosphaera De Meyer, 797–798
Molynocoelia Giglio-Tos, 912, 916, 922, 928, 949, 954 Neocoelopa Malloch, 1014–1016
Monoceromyia Shannon, 768, 788 Neodeceia Malloch, 973, 988, 992
Monochaetoscinella Duda, 1140, 1142, 1151 Neodexiopsis Malloch, 1289–1290, 1293–1294
mordax Dick & Wenzel, Megistapophysis, 1257 Neodiplotoxa Malloch, 1148, 1151
Morellia Robineau-Desvoidy, 1283, 1285–1286, 1291, Neodohrniphora Malloch, 733, 739, 741, 744, 746, 756, 759
1293–1294, 1297 Neodyscrasis Kameneva & Korneyev, 888, 891, 899
morionella (Aldrich), Helicobia, 1330 Neoecothea Peterson & Gill, 1161
Mosillus Latreille, 1215, 1222, 1230, 1233 Neogriphoneura Malloch, 974, 977–978, 980, 985, 992, 995
multisetosa Wulp, Lasiona, 1406 Neohylemyia Malloch, 1275
Mumetopia Melander, 1073–1077, 1079 Neoidiotypa Osten Sacken, 875, 877–878
Musca Linnaeus, 1281–1283, 1287, 1293, 1157 Neoleucopis Malloch, 998, 1000–1004, 1006
muscina (Meigen), Cerodontha, 1058, 1068 Neoliminiinae, 1020
Muscina Robineau-Desvoidy, 1285, 1290, 1293 Neomeoneurites Hennig, 1101, 1103–1104
Muscinae, 1282, 1295, 1297, 1344 Neominettia Hendel, 983, 992
Muscopteryx Townsend, 1349, 1386, 1409 Neomintho Brauer & Bergenstamm, 1369, 1376, 1409
Mutillimyia Hull, 774, 788 Neomuscina Townsend, 1285–1287, 1294
Mycetaulini, 867–868 Neomyennis Hendel, 893, 897
Mycodrosophila Oldenberg, 1198–1201, 1203, 1205–1206 Neomyia Walker, 1283, 1285, 1294, 1297
Mydaea Robineau-Desvoidy, 1288, 1293 Neomyopites Freidberg & Norrbom, 919, 935, 949–950
Mydaeinae, 1282, 1295 Neopachycerina Malloch, 988
Myennidini, 882, 888, 901 Neopelomyia Hendel, 1110, 1112–1114
Myennis Robineau-Desvoidy, 882, 901 Neophora Borgmeier, 733, 737, 740, 748, 756
Myiomima Brauer & Bergenstamm, 1353, 1381, 1409 Neophyllomyza Melander, 1126–1127, 1132, 1134
Myiopharus Brauer & Bergenstamm, 1348, 1360, 1371, 1375, Neophyto Townsend, 1323
1379, 1381, 1407, 1409, 1412 Neopleurophora Brown, 730, 743, 756
Myodris Lioy, 1087–1088, 1091 Neopyrgota Hendel, 958, 960
Myolepta Newman, 769, 782, 785, 788 Neorhagoletis Hendel, 935, 937
Myopa Fabricius, 848, 851, 854–855 Neorhinoleucophenga Duda, 1206
myopaeformis (Röder), Tetanops, 882 Neorhinotora Lopes, 1159–1161, 1163
Myopella Malloch, 1273 Neortalotrypeta Norrbom, 910, 916, 925, 935
Myopella Robineau-Desvoidy, 1273 Neoschildomyia Gaimari, 1053
Myophthiria Rondani, 1242–1247 Neoscutops Malloch, 1090–1091
Myopina Robineau-Desvoidy, 1273 Neosilba McAlpine, 857–860, 862–863
Myopinae, 848–849 Neosolieria Townsend, 1377, 1392, 1409
Myopinella Michelsen, 1273, 1275 Neosophia Guimarães, 1384, 1409
Myospila Rondani, 1288, 1293 Neosphaerocera Kim, 1168, 1185, 1187
Myothyriopsis Townsend, 1363, 1367, 1409 Neotanygastrella Duda, 1198–1200, 1203–1206
Myriophora Brown, 735, 746, 756 Neotanypeza Hendel, 827–831
myrmex (Schiner), Cardiacephala, 806 Neotaracia Foote, 918, 936, 943, 950, 952
myrmicola Sabrosky, Costalima, 1125, 1134 Neotephritis Hendel, 916, 921, 936, 943, 946, 950, 954
Myrmolimosina Marshall, 1183–1184 Neothelairini, 1356
Myrmosicarius Borgmeier, 733, 737, 744–745, 756, 760 Neotraginops Prado, 1053
Mystacella Wulp, 1346, 1363, 1365, 1409 Neotrichobius Wenzel & Aitken, 1253, 1255, 1257
Mystacomyia Giglio-Tos, 1354, 1363, 1408–1409 neotropica Lizarralde de Grosso, Lemnaphila, 1229
nana Verbeke, Sepedonella, 1019 neotropica Snyder, Azelia, 1291, 1417
Nannodastia Hendel, 1236–1238 neotropica Wirth, Brachydeutera, 1228
Nausigaster Williston, 771–772, 785, 788 neotropicanus Ozerov, Palaeosepsioides, 1038
Neaera Robineau-Desvoidy, 1370, 1372, 1393, 1409 Neottiophilinae, 865–867
TAXONOMIC INDEX TO VOLUME 2  1433

Neoxangelina Hendel, 984, 992 Nycterophilia Ferris, 1251, 1257

Nepaliseta Barraclough, 963, 965, 969 Nycterophiliinae, 1250, 1259
Nephochaetopteryx Townsend, 1320–1321, 1331, 1334 Oberonomyia Reinhard, 1383, 1409
Nephrocerinae, 793–795 obliqua (Macquart), Anastrepha, 912, 947
Nephrocerus Zetterstedt, 793–796, 801–802 obliqua Malloch, Xenopterella, 994
Neplas Porter, 767, 774, 785, 787 obliquum Enderlein, Loxobathmis, 1150
Neriinae, 815, 817 obscura Barber, Pseudodinia, 1005
Neriocephalus Enderlein, 807 obscura Spencer, Calycomyza, 1066
Nerius Fabricius, 816–819 obscura (Wiedemann), Pseudopterocalla, 899
Neuzina Marinoni & Knutson, 1018–1019, 1021–1022 obscurella (Fallén), Discocerina, 1211, 1218, 1229, 1232
Nicarao Silva, 1038 obscuriventris (Loew), Tetreuaresta, 941, 951
niger Williston, Gastrops, 1229 obtecta Borgmeier, Placophorina, 757
nigra Fluke, Milesia, 788 occidentalis Williston, Pelomyia, 1112
nigra (Hendel), Terpnomyennis, 1231, 892, 896, 900 occipitalis Melander, Mumetopia, 1075, 1079
nigra Hernández-Ortiz, Oedicarena, 919, 922, 936 Oceanicia Enderlein, 874
nigra Macquart, Rhingia, 766, 789 Ocellominettia Hendel, 983, 992
nigribasis Kuchta & Savage, Mesembrina, 1293 Ochthera Latreille, 1211–1212, 1219, 1224, 1230–1231, 1233
nigriceps (Meigen), Stearibia, 866, 869 Ochtherini, 1219
nigricoxa Ozerov, Pseudopalaeosepsis, 1032, 1035, 1038 Ochthiphilinae, 1006, 1010, 1210
nigrimana Coquillett, Mumetopia, 1073–1074, 1076, 1079 oculifera (Bigot), Anthomyia, 1275
nigrimana (Hennig), Pseudopomyza, 822, 824 Ocyptamus Macquart, 764, 776–777, 780–782, 788, 791–792
nigrimana (Meigen), Prochyliza, 868 Odinia Robineau-Desvoidy, 1049, 1051–1055
nigripalpis (Cogan), Ortalidina, 1005 Odiniinae, 1049, 1052
nigripes Camras, Thecophora, 854 Odontoloxozus Enderlein, 815–819
nigripes (Cresson), Parydra, 1230 Odontomera Hendel, 872, 875–876, 878
nigripes Guimarães, Laneella, 1300–1301 Odontomera Macquart, 845
nigrita Malloch, Freyia, 991 Odontomerella Hendel, 876
nigritarsis Marinoni, Guatemalia, 1022 Oebalia Robineau-Desvoidy, 1314, 1318, 1331, 1334
nigriventris (Curran), Protopiophila, 869 Oecothea Haliday, 1161–1162
nigroapicata (Malloch), Trivialia, 994 Oedematella Hendel, 874, 878
nigrocoeruleus Vockeroth, Alipumilio, 786 Oedenops Becker, 1216, 1230
Ningulus McAlpine, 1085 Oedicarena Loew, 916, 919, 922, 924, 929, 936
nitens Bigot, Lycastirhyncha, 788 Oestrinae, 1306, 1345
nitens (Loew), Paramyia, 1126, 1132 Oestrophasia Brauer & Bergenstamm, 1353, 1382, 1391,
nitens (Schiner), Odontomera, 878 1409, 1416
nitida Johnson, Heteromeringia, 1042, 1047 Oestrus Linnaeus, 1305–1310
nitida Sturtevant & Wheeler, Diphuia, 1218, 1229 Olcanabates Enderlein, 1150
nobilis (Walker), Conops, 850, 854 Olcella Enderlein, 1140–1142, 1146, 1150–1151
Nocticanace Malloch, 1116, 1118–1119 oleae (Rossi), Bactrocera, 910, 947
Nocticanacinae, 1118–1119 Olfersia Wiedemann, 1241–1242, 1245, 1247
Noctiliostrebla Wenzel, 1253, 1255, 1257–1258 olivaceus Townsend, Rhachoepalpus, 1395, 1411
Norrbomia Papp, 1171, 1185–1186 ollachea Townsend, Dejeaniops, 1402
Nostima Coquillett, 1213, 1230, 1232 Omegasyrphus Loew, 772, 788
notata (Loew), Bertamyia, 717–719 Omomyia Coquillett, 874
Notaulacella Enderlein, 1143–1144, 1151, 1153 Oncodometopus Shewell, 972–973, 979–980, 988, 992
Notenthes Marshall, 807, 812–813 Oncopsia Enderlein, 816–819
Nothomicrodon Wheeler, 765, 784 Onychaspidium Enderlein, 1141, 1147, 1149, 1151
Notiphila Fallén, 1212, 1214–1216, 1225, 1227, 1230, 1232 Opacifrons Duda, 1172–1173, 1178, 1185–1186
Notochaeta Aldrich, 1315–1316, 1322, 1327, 1330, 1333 opacula Fedoseeva, Meromyza, 1151
Notogramma Loew, 882, 884, 889–890, 899, 901 Opalimosina Roháček, 1181, 1185
nova (Schiner), Rhagoletis, 940, 951 Opetiophora Loew, 1143, 1151
nubecula (Malloch), Stenolauxania, 993 ophiata Griffiths, Anthomyia, 1275
nubila (Wiedemann), Eumecosomyia, 882, 898 Ophiomyia Braschnikov, 1057–1058, 1063–1064, 1069, 1071
nubilipennis Wulp, Acronacantha, 1398 Ophirion Townsend, 1381, 1410
nubilipennis (Wulp), Protodictya, 1018, 1022 Ophthalmoptera Hendel, 892–893, 896, 899
nudiseta (Wulp), Synthesiomyia, 1285, 1294 Ophyra Robineau-Desvoidy, 1292, 1295
nudiseta Wheeler, Laccodrosophila, 1205 Opsidia Coquillett, 1314, 1318, 1331, 1334
nudus (Coquillett), Oedenops, 1230 Opsomeigenia Townsend, 1403
Nycteribia Latreille, 1263 Opsophagus Aldrich, 1402
Nycteribiinae, 1264 optata (Reinhard), Anisia, 1398, 1415
1434 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Orasiopa Zatwarnicki & Mathis, 1221, 1230 pallipes Malloch, Asilostoma, 991
orbitalis (Frost), Japanagromyza, 1064 Palmomyia Grimaldi, 1200
orientalis Malloch, Stenomicra, 1088 Palpada Macquart, 765, 769, 773, 775, 786, 789
orientalis Schiner, Atherigona, 1284, 1291 palpalis Malloch, Asilostoma, 991
Ormia Robineau-Desvoidy, 1345, 1360, 1393, 1410, 1416 palpalis Reinhard, Oberonomyia, 1409
Ormiophasia Townsend, 1410, 1416 Panava Lopes, 1322, 1327, 1331, 1334
ornata (Melander), Pachyopella, 992 Pandasyophthalmus Stuckenberg, 767, 777, 789
ornata (Schiner), Neoxangelina, 992 Panermya Wulp, 877
ornata (Zetterstedt), Odinia, 1051 Panzeria Robineau-Desvoidy, 1350, 1367, 1392, 1410
Ornidia Lepeletier & Serville, 763, 765–766, 769, 781, 785, 789, Paracanace Mathis & Wirth, 1118–1119
791–792 Paracantha Coquillett, 917, 921, 936, 938, 950, 952
Ornithoctona Speiser, 1242–1248 Paracestrotus Hendel, 979
Ornithoica Rondani, 1241–1245, 1247 Paradejeania Brauer & Bergenstamm, 1358, 1395, 1410,
Ornithoicinae, 1244 1414, 1417
Ornithomya Latreille, 1241, 1243–1245, 1247 Paradidyma Brauer & Bergenstamm, 1347, 1369, 1385–1386,
Ornithomyinae, 1244 1410, 1416
orpheus Hull, Mixogaster, 788 Paradyschiria Speiser, 1250, 1255, 1258
Ortalidina Blanchard, 998–1002, 1004–1005 Paraephydra Mathis, 1226
Ortalis Meigen, 882 Paraeuctenodes Pessôa & Guimarães, 1251, 1258
Ortholfersia Speiser, 1244 Paraglenanthe Wirth, 1219
Ortholfersiinae, 1244 Paragodon Thompson, 770, 789
Orthonevra Macquart, 766, 772, 782, 789 paragoides Thompson, Paragodon, 789
Orygma Meigen, 1034 Paragorgopis Giglio-Tos, 886, 892, 894, 899
Oscinella Becker, 1138, 1147, 1151 Paragus Latreille, 767, 777–778, 780, 789, 792
Oscinellinae, 1138–1139, 1149 Paralimna Loew, 1215–1216, 1227, 1230
Oscinicita Wheeler, 1147, 1151 Paraliodrosophila Duda, 1203, 1205
Ostracocoelia Giglio-Tos, 929, 936, 947, 950 parallela (Walker), Ditaeniella, 1022
Otitinae, 882, 888–889, 891, 900–902 Paralucilia Brauer & Bergenstamm, 1298, 1302–1303
Otitini, 882 Paramacronychiinae, 1316, 1318, 1334
ovinus (Linnaeus), Melophagus, 1243, 1247 Paramicrodon de Meijere, 766, 770, 789
ovis Linnaeus, Oestrus, 1306–1308, 1310 Paramycodrosophila Duda, 1199–1200, 1203, 1205
Oxyna Robineau-Desvoidy, 945 Paramyia Williston, 1126–1129, 1132, 1134, 1136
Oxynops Townsend, 1374, 1410 Paraneossos Wheeler, 1050, 1161–1162
Oxysarcodexia Townsend, 1316, 1321, 1326, 1328, 1331, Paranthomyza Czerny, 1074, 1081
1333–1335 Parapamecia Cogan, 1001, 1004
Oxyvinia Dodge, 1326, 1331 Paraphorantha Townsend, 1410
Ozaenina Enderlein, 874 Paraphyola Hendel, 900
Pacatuba Lopes, 1322, 1327, 1331 Parapipunculus Rafael, 801
pachecoi Curran, Protodejeania, 1411 Parapterocalla Aczél, 900
Pachyceramyia Albuquerque, 1288 Parapyrellia Townsend, 1285
pachycericola Mangan & Baldwin, Odontoloxozus, 817, 819 Parasclerocoelus Marshall & Dong, 1180
pachycondylae (Brues), Cataclinusa, 734, 754 Parascutops Mathis & Papp, 1091
Pachyopella Shewell, 976–977, 992 Parasphaerocera Spuler, 1168, 1184–1186
pachypyga (Aldrich & Webber), Patelloa, 1410 Parasphen Enderlein, 809, 811
packardi Wirth, Ephydra, 1229 Parastenopa Hendel, 917, 922, 928, 936, 950, 952
Palaeoceroptera Duda, 1176, 1186 Parastrebla Wenzel, 1254, 1256, 1258
Palaeocoprina Duda, 1174, 1186 Parathyroglossa Hendel, 1222
Palaeolimosina Duda, 1174 Paratissa Coquillett, 1214, 1217, 1230, 1232
Palaeomyopinae, 849 Paratraginops Hendel, 1053–1054
Palaeosepsioides Ozerov, 1034, 1037–1038 Paratrichobius Costa Lima, 1254–1255, 1258
Palaeosepsis Duda, 1033–1035, 1037–1039 Parazodion Kröber, 849–850, 853–854
palans Giglio-Tos, Chaetocoelia, 991 Parectecephala Becker, 1140, 1148, 1151
pallens (Reinhard), Angustia, 1398 Parectinocera Becker, 1022
pallida (Loew), Cadrema, 1140, 1149 Parepalpus Coquillett, 1359, 1395, 1410
pallida Malloch, Xenochaetina, 994 Pareuthychaeta Hennig, 1209–1210
pallida McAlpine, Protopiophila, 869 Pareuxesta Coquillett, 886, 890, 899
pallida Robineau-Desvoidy, Eudejeania, 1403 parkeri Gaimari, Helgreelia, 1053
pallidipalpis (Wulp), Angustia, 1398 parnelli Wenzel, Nycterophilia, 1257
pallipes (Cresson), Hyadina, 1229 Parochthiphila Czerny, 1000–1001, 1004–1007
pallipes (Loew), Calotarsa, 719 Parophthalmoptera Hendel, 894, 896, 899
TAXONOMIC INDEX TO VOLUME 2  1435

Paroxyna Hendel, 948, 953 Phasiinae, 1344–1345, 1355–1356, 1404

parva (Bigot), Hemisturmia, 1405 Phasiophyto Townsend, 1387, 1410
Parydra Stenhammar, 1220, 1225, 1230 Pherbellia Robineau-Desvoidy, 1019, 1021–1023
Parydrini, 1225, 1231 Philornis Meinert, 1282–1283, 1285, 1287, 1294, 1306, 1333
patagonensis (Macquart), Ditaeniella, 1022 Philygria Stenhammar, 1213, 1220, 1232
Patelloa Townsend, 1361, 1410 Philygrini, 1213
paupera Hering, Euarestopsis, 949 Phlebosotera Duda, 1093–1095
Paurophora Silvestri, 742, 751, 756 Phloridosa Sturtevant, 1199, 1204
Peckia Robineau-Desvoidy, 1313, 1315–1316, 1319, 1323, 1325, Phobetromyia Reinhard, 1368, 1410
1328, 1331, 1334 Pholeomyia Bilimek, 1126–1129, 1135–1136
Peckiamyia Dodge, 1325, 1331 Phora Latreille, 725, 730, 757, 759, 761
pectinata Hendel, Procrita, 993 Phorinae, 727, 758
Pectinifer Cresson, 1221, 1230 Phormiini, 1301
peculiaris Malloch, Marbenia, 1091 Phosocephala Townsend, 1358, 1411
Pegomya Robineau-Desvoidy, 1272, 1274–1275 Phrosinella Robineau-Desvoidy, 1318, 1331
Pegoplata Schnabl & Dziedzicki, 1272, 1274–1275 Phthitia Enderlein, 1172, 1177, 1183
Peleteria Robineau-Desvoidy, 1346, 1357, 1410, 1416 Phylloclusia Hendel, 1043
Pelidnoptera Rondani, 1020 Phyllomyza Fallén, 1125–1127, 1130–1131, 1133, 1135
Pelomyia Williston, 1109–1110, 1112–1114 Phyllomyzinae, 1127
Pelomyiella Hendel, 1110, 1112–1113 Phyllophilopsis Townsend, 1348, 1376, 1381, 1389, 1397,
Pelomyiinae, 1112–1113 1406, 1411
Peltodasia, 956 Phymatopterella Brues, 733, 735–736, 744, 748, 757
Peltopsilopa Cresson, 1217, 1228 Physegeniopsis Gaimari & Silva, 972, 975, 990, 992
penelope (Osten Sacken), Procecidocharoides, 950 Physegenua Macquart, 984, 992
Penesymmetria Kessel & Maggioncalda, 716 Physemops Cresson, 1223, 1226, 1230
penicillatum Hendel, Platygraphium, 992 Physiphora Fallén, 883, 885, 899
pennipes (Fabricius), Trichopoda, 1388, 1413 Physocephala Schiner, 850–852, 854–855
Penola Richards, 1182 Physoclypeus, 972–973, 984, 992, 995
Pentanotaulax Enderlein, 1152 Physoclypeus Hendel, 972–973, 984, 992, 995
Penthosia Wulp, 1350, 1390, 1410 Physoconops Szilady, 847, 849–852, 854
perezi (Romero & Ruppel), Neosilba, 859, 862 Physoptera Borgmeier, 736–737, 745, 757
Perilimnia Becker, 1019, 1023 Phytalmyiinae, 910
Periscelidinae, 1090 Phytobia Lioy, 1060, 1062–1065, 1069–1070
Periscelis Loew, 1055, 1087–1092 Phytoliriomyza Hendel, 1062, 1066–1068, 1070–1071
Periscepsia Gistel, 1349, 1385, 1410 Phytomyptera Rondani, 1372, 1383, 1411
personta (Lutz & Lima), Anapyrgota, 960 Phytomyza Fallén, 1058–1059, 1062, 1067, 1070–1071
perspicillatus (Pessôa & Galvão), Trichobioides, 1258 Phytomyzinae, 1057, 1060, 1063
pertinax (Bates), Caenoriata, 927, 948 picea Hendel, Parophthalmoptera, 899
peruanus (Hennig), Cerantichir, 819 picea Hennig, Neopyrgota, 960
peruanus Hennig, Odontoloxozus, 819 picta (Fabricius), Delphinia, 887, 891, 898, 902
peruvianus (Korytkowski), Pyrgotoides, 940, 951 picta (Wulp), Myothyriopsis, 1409
petiolata (Townsend), Distichona, 1403 picticornis (Curran), Sapromyza, 993
petiolata (Wiedemann), Cordyligaster, 1402 picturata (Snow), Acinia, 926, 947
Petriana Lopes, 1329 Piezura Rondani, 1277–1278
Petrmyia Albuquerque, 1275 pilicornis Cresson, Paratraginops, 1054
Phaenicia Robineau-Desvoidy, 1301, 1303 pilipes Lopes, Thomazomyia, 1332
Phaeomyiinae, 1020 Pilispina Albuquerque, 1290, 1294–1295
Phaeopterina Frey, 810 Piophila Fallén, 865, 867–868
Phaiosterna Cresson, 1215–1216, 1230 Piophilinae, 865, 867–868
Phaiosterna Cresson, 1215–1216, 1230 Piophilini, 868
Phalacrotophora Brues, 729, 739, 744–745, 756, 761 pipinna Sasakawa, Phytobia, 1069
Phalconomus Wenzel, 1251, 1258 Pipiza Meigen, 772
Phanerochaetum Hennig, 1123 Pipunculina Albrecht, 797–798, 800
Phaonantho Albuquerque, 1272–1273, 1276 Pipunculinae, 793–795
Phaonia Robineau-Desvoidy, 1282, 1288–1289, 1291, 1294 Pipunculini, 794
Phaoniinae, 1282, 1292, 1295 Pipunculus Latreille, 793, 795–796, 798, 801–803
phaseoli (Tryon), Ophiomyia, 1069 placata (Hutton), Pseudoleria, 1160
phaseoli Spencer, Japanagromyza, 1061, 1068 Placophorina Borgmeier, 752, 757
phaseolivora Spencer, Melanagromyza, 1069 Placopsidella Kertész, 1222, 1231–1232
Phasia Latreille, 1371, 1377, 1382, 1387, 1394, 1400, 1410, 1416 Plagiocephalus Wiedemann, 889, 894–895, 899–901
1436 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Plagiomima Brauer & Bergenstamm, 1354, 1385, 1411 praeustum (Meigen), Neottiophilum, 866
Plagiostenopterina Hendel, 905–906 pratensis (Meigen), Blepharipa, 1388, 1400
Planes Rondani, 787 pretiosa Curran, Dolichocodia, 1403
Planinasus Cresson, 1085, 1090–1091 prima Malloch, Coenosopsia, 1275
Planipeza Marshall, 809–810 prima Malloch, Setulina, 993
Plastotephritinae, 905, 907, 910 prima Osten Sacken, Pseudeuxesta, 882
platura (Meigen), Delia, 1275 primitiva Guimarães & D’Andretta, Hershkovitzia, 1263, 1265
Platycheirus Lepeletier & Serville, 771, 776, 782, 789 proana (Curran), Melanomyza, 992
Platycoenosia Strobl, 1278 Procanace Hendel, 1117–1118
Platydipteron Borgmeier & Prado, 744, 746, 757 Procecidochares Hendel, 918, 921, 937, 939, 950, 953
Platygraphium Hendel, 984, 992 Procecidocharoides Foote, 918, 939, 950, 952
Platypeza Meigen, 716–719 Procerosoma Lonsdale & Marshall, 1041, 1043–1044, 1047–1048
platypeza Papp & Silva, Pseudominettia, 993 Prochyliza Walker, 865, 868
Platypezinae, 715–716 Procrita Hendel, 983, 993
Platystoma Meigen, 905, 907 Prorhynchops Brauer & Bergenstamm, 1403
Platystomatinae, 905 Prosenoides Brauer & Bergenstamm, 1354, 1381, 1411
plaumanni Borgmeier. Travassophora, 758 Prosopantrum Enderlein, 1160–1162, 1164
plaumanni Hennig, Teutoniomyia, 1023 Prospherysa Wulp, 1362, 1411
plaumanni Shewell, Paratraginops, 1054 Protanthomyza Hennig, 1074
Plaumannimyia Hering, 934, 939, 945, 950–951 Protearomyia McAlpine, 858–859
Plesioclythia Kessel & Maggioncalda, 715–716, 718 Proterpnomyia Blanchard, 900
pleuropunctata (Wiedemann), Willistoniella, 1026–1030 Proteseia Korneyev & Hernández, 888, 891–892, 899
plinthopyga (Wiedemann), Blaesoxipha, 1317, 1321, 1325, 1329 Protocalliphora Hough, 1300
Plocoscelus Enderlein, 806–807, 811–812 Protoclythia Kessel, 719
plumosa Loew, Tetanocera, 1023 Protodejeania Townsend, 1357–1358, 1411
plumosa van der Wulp, Hydrophoria, 1275 Protodictya Malloch, 1018–1019, 1021–1024
Plunomia Malloch, 998, 1000, 1002, 1004 Protolepidostola Hull, 788, 792
plurivittatus Bigot, Nerius, 816, 819 Protopiophila Duda, 865–866, 868–869
Podiomitra Marshall & Roháček, 1167–1169, 1185, 1187 Protothyreophora Ozerov, 867
poecilogastra Becker, Ulia, 1135 proxima (Séguy), Megistopoda, 1250, 1253, 1259
Poecilolycia Shewell, 972, 974, 980, 987, 992 Pseudacteon Coquillett, 726, 737, 747–748, 757, 761
Poecilominettia Hendel, 972, 980–981, 983, 992, 994 Pseudeurina de Meijere, 1151
Poecilomyia Hendel, 876–878 Pseudeurybata Hennig, 808, 810, 812
Poecilosomella Duda, 1170, 1172, 1175, 1181, 1185–1187 Pseudeutreta Hendel, 932, 939
Poecilotylus Hennig, 805, 809, 812–813 Pseudiastata Coquillett, 1199–1200, 1202, 1205
Poemyza Hendel, 1058, 1067–1068 Pseudobombyliomyia Townsend, 1368, 1411
Pogonortalis Hendel, 905–906 Pseudocacoxenus Duda, 1199, 1202, 1205
Polideini, 1345, 1356, 1367, 1400, 1406, 1415 Pseudocalliope Malloch, 972–973, 981, 993
Polietina Schnabl & Dziedzicki, 1286, 1294 Pseudochaeta Coquillett, 1352, 1363, 1365, 1411
Polionota Wulp, 918, 939, 950, 953 Pseudochaetona Townsend, 1406
polistensis Hall, Sarcophaga, 1332 Pseudocollinella Duda, 1172, 1177
Polistiopsis Townsend, 1390, 1406, 1411, 1414 Pseudodexia Brauer & Bergenstamm, 1372, 1377, 1411
polita (Coquillett), Aplomyopsis, 1399 Pseudodinia Coquillett, 998, 1000–1002, 1004–1006
polita (Townsend), Ischyrophaga, 1406 Pseudodoros Becker, 766, 779, 782, 789, 791
politus (Say), Toxomerus, 790 Pseudodyscrasis Hernández, 888, 891, 899, 901
Polleniini, 1301 Pseudoedaspis Hendel, 932, 934, 939, 949
Polphopeza Enderlein, 828 Pseudogaurax Malloch, 1137, 1141, 1146, 1151, 1153
Polybiomyia Shannon, 766, 768, 771, 785, 789 Pseudogriphoneura Hendel, 972, 988, 993
Polygaster Wulp, 1377, 1393, 1411 Pseudohecamede Hendel, 1218, 1228
polygoni Spencer, Japanagromyza, 1064 Pseudohypocera Enderlein, 726, 731, 748, 757, 759–761
Polymorphomyia Snow, 931, 939, 950, 953 Pseudoleria Garrett, 1160–1163
Polypathomyia Krivosheina, 823 Pseudolynchia Bequaert, 1245–1247
Polytrichophora Cresson, 1218, 1221, 1231 Pseudomicrodon Hull, 770
pompalis (Reinhard), Jurinia, 1406 Pseudomilichia Becker, 1126–1129, 1133, 1135
posticata (Meigen), Nemorimyza, 1062, 1069 Pseudominettia Papp & Silva, 981, 993
postuncinata (Hall), Villegasia, 1316, 1333 Pseudonapomyza Hendel, 1059, 1065, 1070
Potamia Robineau-Desvoidy, 1284, 1290 Pseudopachychaeta Strobl, 1149
poultoni Verrall, Callicera, 787 Pseudopalaeosepsis Ozerov, 1032, 1034–1035, 1037–1038
Pradomyia Gaimari, 1053–1054 Pseudophorellia Lima, 909–910, 912, 915, 940, 951, 954
praeceps Hendel, Chaetopsis, 898 Pseudopolionota Lima, 940, 943
TAXONOMIC INDEX TO VOLUME 2  1437

Pseudopomyza Strobl, 821–825 radiata Hendel, Megalaemyia, 893

Pseudopomyzella Hennig, 823–824 rafaeli (Townsend), Siphosturmia, 1412
Pseudopterocalla Hendel, 894, 899–900 rafaeli Townsend, Chaetonodexodes, 1401
pseudopterus Coquillett, Metelasmus, 1252, 1257, 1259 Rafaelia Townsend, 1324, 1331
Pseudoptilolepis Snyder, 1286, 1294 Rainieria Rondani, 806, 808, 810–813
Pseudoseioptera Stackelberg, 882, 901 Rainieriella Hennig, 811–812
Pseudosiphona Townsend, 1371, 1374, 1383, 1411–1412 Ramuliseta Keiser, 963, 965, 969
Pseudostrebla Costa Lima, 1256, 1258–1259 ranchograndensis (Spencer), Nemorimyza, 1069
Pseudosturmia Thompson, 1412 ranica Mathis & Trautwein, Lipochaeta, 1230
Pseudotephritis Johnson, 882, 901 rapax (Walker), Helicobia, 1330
Pseudoviviania Brauer & Bergenstamm, 1378, 1411 rasilis (Wulp), Lydinolydella, 1408
psidii Capoor, Pterocerina, 900 rattii Munari, Pseudorhicnoessa, 1112
Psila Meigen, 837–842 Ravinia Robineau-Desvoidy, 1316, 1325, 1331, 1334
Psilacrum Becker, 1143, 1151–1152 Raviniopsis Townsend, 1324, 1334
Psilinae, 837, 839 Receptrixa Roháček, 1074
Psilopa Fallén, 1214, 1219, 1231 Renocerinae, 1020
psychotria Grimaldi & Nguyen, Cladochaeta, 1204 reses Giglio-Tos, Scathophaga, 1267–1269
Pterocalla Rondani, 886, 889, 893, 899–900 reticulata Prado, Schildomyia, 1054
Pterocallinae, 882, 900–901 Retrocitomyia Lopes, 1322, 1324–1325, 1332
Pterocallini, 882, 889, 892, 900 Rettenmeyerina Dodge, 1322, 1332
Pterocerina Hendel, 886–887, 894, 897, 900 Rhabdomyia Borgmeier, 742, 751–752, 757
Pterogramma Spuler, 1167, 1170, 1172, 1176, 1182–1183, Rhachoepalpus Townsend, 1359, 1395, 1411
1185, 1187 Rhagoletis Loew, 910, 917, 923, 925, 928–929, 940, 942, 944,
Pteromicra Lioy, 1022 951–954
Ptilodexia Brauer & Bergenstamm, 1351, 1383, 1394, 1403–1404, Rhagoletotrypeta Aczél, 913, 928, 942, 944, 951, 954
1406, 1408–1409, 1411, 1417 Rhingia Scopoli, 766, 773, 782, 789
Ptilomyia Coquillett, 1215, 1222, 1231 Rhingiini, 764
Ptilosphen Enderlein, 806, 809, 811–812 Rhiniinae, 1301, 1340
pubera (Loew), Atrichomelina, 1019, 1022–1023 Rhiniini, 1301
puella (Williston), Trivialia, 994 Rhinoleucophenga Hendel, 1199, 1201–1202, 1205
pulchellum Macquart, Leptostylum, 1407 Rhinopomyzella Hennig, 822–824
pulchra Williston, Milesia, 775, 788 Rhinoprosopa Hull, 766, 779, 786
pulchripes Loew, Psilopa, 1231 Rhinotora Schiner, 1160–1161, 1163
puliciformis (Wenzel), Phalconomus, 1258 Rhinotorini, 1160–1161
Puliciphora Dahl, 726–727, 742, 749–750, 752–753, 755, 757, Rhithrum Hendel, 932, 940
760–761 Rhodesiella Adams, 1143–1144, 1152
Pullimosina Roháček, 1174–1175, 1178–1179, 1185–1186 Rhodesiellinae, 1138
pullum Roháček & Barber, Stiphrosoma, 1079 Rhoga Walker, 770
punctifrons Malloch, Schildomyia, 1054 Rhopalopterum Duda, 1147, 1152
punctulata Hendel, Megalaemyia, 893, 899 Rhopalosyrphus Giglio-Tos, 766, 770, 789, 792
punctulata (Wulp), Suillia, 1163 Rhyncophoromyia Malloch, 740–741, 746, 748, 755, 757, 759
purpuratum Cole, Notogramma, 899 Rhynencina Johnson, 919, 951, 954
purpurescens (Walker), Lucilia, 1299 Rhyparella Hendel, 892, 894, 896, 900
purpurops Steyskal, Traginops, 1051 Rhysophora Cresson, 1214, 1217, 1231, 1233
pusilla (Fallén), Ischiolepta, 1184 Rhytidops Lindner, 1026–1028
Pycnopota Bezzi, 1166, 1185 Richardia Robineau-Desvoidy, 871, 873–874, 878
Pyrgota Wiedemann, 955–961 Richardiinae, 873–874, 879
Pyrgotella Curran, 960 Richardiodes Hendel, 877
Pyrgotinae, 957–958 Risa Becker, 1212
Pyrgotini, 958 Ritaemyia Frey, 987, 993
Pyrgotoides Curran, 911, 917, 923, 940, 951 Rivellia Robineau-Desvoidy, 903–907
Pyrgotosoma Malloch, 957 Robustagramma Marshall & Cui, 1170, 1172, 1175, 1177, 1181,
Pyrrhodexia Townsend, 1350, 1386, 1398, 1411 1185–1186
Quadratosoma Townsend, 1359, 1411 roldanorum Mathis & Hogue, Diedrops, 1228
quadriseriata Hendel, Tomoplagia, 928 Ropalomera Wiedemann, Ropalomera, 1026, 1029
Quichuana Knab, 766, 768, 775, 782, 784, 786, 789, 791 roralis (Linnaeus), Melanophora, 1340
rabelloi Spencer, Phytobia, 1070 Roryeuchomyia Gaimari & Silva, 990
rabida Wheeler, Aphaniosoma, 1157 rosales Woodley, Melanagromyza, 1069
Rachiptera Bigot, 917, 938, 940 rossi MacGowan & Rotheray, Odinia, 1051
Rachispoda Lioy, 1173–1174, 1185, 1187 rostratus Cresson, Dagus, 1220, 1228
1438 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

rotundicornis Hennig, Latheticomyia, 824 Sciadocerinae, 716, 726–727

rotundipennis (Malloch), Minilimosina, 1184 Sciomyza Fallén, 1020
rotundipennis Malloch, Neoscutops, 1091 Sciomyzinae, 1020, 1024
rubella (Wulp), Notenthes, 812 Sciomyzini, 1018–1022
rubripalpis Wulp, Helina, 1292 Sciosapromyza Hendel, 981, 983, 993
rubriventris (Wulp), Mystacomyia, 1354, 1409 Scipopeza Enderlein, 828–830
Rudolfina Roháček, 1180, 1183 Scipopus Enderlein, 806, 810, 812–813
rufa Cresson, Automola, 872–873 Sclerocoelus Marshall, 1167, 1172–1173, 1178, 1185–1186
rufa Cresson, Syringogaster, 845–846 Scoliocentra Loew, 1161–1163
rufescens Aldrich, Gaediopsis, 1404 scoliostylus Buck, Inbiomyia, 1108
ruficauda Hendel, Pterocerina, 897, 900 Scopeuma Meigen, 1269
ruficauda (Wulp), Copecrypta, 1346, 1402 scordalus (Walker), Hemichlora, 1292
ruficornis Fabricius, Sarcophaga, 1332 Scotiptera Macquart, 1383, 1412
ruficornis (Wulp), Gymnosoma, 1332, 1405 scutellaris (Wiedemann), Pseudodyscrasis, 891, 899
rufifacies (Macquart), Chrysomya, 1299, 1303 scutellata (Schiner), Pseudopterocalla, 899
rufifrons (Duda), Spinilimosina, 1186 Scutolauxania Hendel, 987
rufina (Wulp), Trichophora, 1413 Scutominettia Hendel, 979, 993
rufipes (Brooks), Epalpus, 1403 Scutops Coquillett, 1090–1092
rufipes Meigen, Phytomyza, 1070 seamansi Shewell, Neoalticomerus, 1050–1051
rufiventris (Wiedemann), Argoravinia, 1319, 1329 secunda (Melander & Spuler), Sepsis, 1032, 1034, 1038
rufopicta (Bigot), Winthemia, 1346, 1413, 1415 segmentaria (Fabricius), Hemilucilia, 1299
rufula (Frey), Stomosis, 1135 seguyi Falcoz, Crataerina, 1245, 1247
rufum Macquart, Ctenostylum, 964–966, 968 Seiopterini, 882
rutilioides (Jaennicke), Paradejeania, 1395, 1410, 1414 Selachops Wahlberg, 1052
rutilioides nigrescens Arnaud, Paradejeania, 1395, 1410, 1414 Semicephalops De Meyer, 797
sabroskyi Grimaldi & Mathis, Stenomicra, 1092 semidiaphana (Rondani), Hemilucilia, 1299
Salpingogaster Schiner, 775, 777–778, 789 seminitidum (Zetterstedt), Lasiomma, 1275
saltans Townsend, Dasiops, 859 Senopterina Macquart, 904–907
salti Brues, Melittophora, 756 Senotainia Macquart, 1314, 1319, 1332
Salticellinae, 1020 Sepedomerus Steyskal, 1019–1021, 1023
Santhomyza Roháček, 1080 Sepedon Latreille, 1019–1020, 1023–1024
Sapromyza Fallén, 972, 986–987, 990, 993–994 Sepedonea Steyskal, 1018–1021, 1023–1024
Sarcodexia Townsend, 1316, 1321, 1323–1324, 1332 Sepedonella Verbeke, 1019–1020
Sarcofahrtiopsis Hall, 1316, 1320, 1327–1328, 1332, 1334 Sepedoninae, 1020
Sarcophaga Meigen, 1315–1316, 1319, 1321, 1324, 1328, 1330, Sepedoninus Verbeke, 1020
1332–1334 Sepsis Fallén, 1031–1032, 1034, 1037–1039
Sarcophaginae, 1313, 1316–1317, 1333–1334 Sepsisoma Johnson, 871, 873, 875–876, 878
Sarcopromusca Townsend, 1283, 1287, 1294–1295 Seri Kessel & Kessel, 716
satanica (Bigot), Penthosia, 1350, 1410 Sericomyia Meigen, 768–769, 786, 790
sativae Blanchard, Liriomyza, 1058, 1060, 1069 Servaisia Robineau-Desvoidy, 1329
saundersii (Williston), Madremyia, 1408 Setellia Robineau-Desvoidy, 871–874, 878–879
scabricula (Haliday), Ischiolepta, 1184 Setellida Hendel, 875–876, 878
scalaris (Fabricius), Fannia, 1278–1279 setifrons (Wulp), Minthoplagia, 1409
scalaris (Loew), Megaselia, 726, 756, 760 setinervis (Coquillett), Thelairodoria, 1412
Scaptodrosophila Duda, 1200, 1203, 1205 Setisquamalonchaea Morge, 862
Scatella Robineau-Desvoidy, 1224–1225, 1231, 1233 setiventris Aczél, Idiopyrgota, 956, 960
Scathophaga Meigen, 1267–1269 setosa Coquillett, Brachicoma, 1329
Scathophaginae, 1267–1268 Setulina Malloch, 979, 993
Scatogera Albuquerque, 1269 sexsetosa Duda, Meropliosepsis, 1032–1033, 1038
Scatophila Becker, 1211, 1225, 1231 Shannonia Malloch, 1019, 1023
schildi Steyskal, Somatia, 835 Shannoniella Townsend, 1338–1340
Schildomyia Malloch, 1053–1054 shewelli Papp & Silva, Tauridion, 972, 993
Schizostomyia Malloch, 1011 Shewellia Hennig, 1049, 1053–1054
schmidti Borgmeier, Zikania, 758 shorae Fluke, Syrphus, 763
schmidti Dybas & Wenzel, Joblingia, 1250, 1253, 1257 Sigaloessa Loew, 1094–1095
schmidti Kröber, Parazodion, 850, 853–854 Sigaloessinae, 1094
schmitzi Örösi-Pal, Braula, 1098–1099 Sigaloessini, 1094
Schnusimyia Hendel, 873–875, 878, 882 signifer (Walker), Epalpus, 1360, 1395, 1403
Schoenomyza Haliday, 1281, 1284, 1289, 1294 signifera (Wulp), Oestrophasia, 1409
Scholastinae, 905 Silba Macquart, 862
TAXONOMIC INDEX TO VOLUME 2  1439

silvai (Lima), Cryptodacus, 948 Spirobolomyia Townsend, 1316, 1325, 1332

similis Wenzel, Paraeuctenodes, 1258 Spoggosia Rondani, 1401
simmondsi Sabrosky, Melaleucopis, 1005 spreta Wulp, Tetanocera, 1023
simulans Sturtevant, Drosophila, 1197, 1200 stabulans (Fallén), Muscina, 1293
sinclairi Wheeler, Meoneura, 1104 stackelbergi Krivosheina & Krivosheina, Turanodinia, 1051
sinefurca (Borgmeier), Brownphora, 732, 754 stackelbergi Krivosheina, Polypathomyia, 823, 901
Sinolochmostylia Yang, 965 stata (Giglio-Tos), Setulina, 993
Sinophthalmus Coquillett, 1199, 1201–1202, 1204 Stearibia Lioy, 866, 868–869
sinuata Mathis, Garifuna, 1229 Stegana Meigen, 1199–1202, 1205
Siopa Hendel, 883, 888, 890, 900 Steganinae, 1200, 1202, 1206
Siphlodora Patterson & Mainland, 1199, 1204, 1206 Steganolauxania Frey, 976–977, 985
Siphona Meigen, 1354, 1374, 1411–1412, 1415 stegmaieri Wirth, Mosillus, 1230
Siphonellopsinae, 1138–1139, 1149 steineri Mathis, Diedrops, 1228
Siphonini, 1356, 1372, 1401, 1411, 1415 Steleocerellus Frey, 1148–1149, 1152
Siphonophysa Hendel, 979, 993 Steleoneura Stein, 1380, 1412
Siphosturmia Coquillett, 1367, 1412 stellatus Wulp, Amphicnephes, 904, 907
Siphunculina Rondani, 1138–1139, 1144, 1152 Steneretma Loew, 900
Sisyropa Brauer & Bergenstamm, 1414 Stenodexia Wulp, 1387, 1412
slossonae Coquillett, Lipochaeta, 1218, 1230 Stenolauxania Malloch, 976, 993
slossonae Coquillett, Sapromyza, 993 Stenomicra Coquillett, 1074, 1083, 1085, 1088, 1090–1093
Sobarocephala Czerny, 1041–1043, 1045–1048 Stenomicrinae, 1090
Sobarocephalinae, 1041, 1043, 1045, 1048 Stenomyia Hendel, 884, 889, 898, 900
sobrinata (Wulp), Dyseuaresta, 930, 949 Stenopa Loew, 937, 940
Somatia Schiner, 833–835, 1090 Stenopterina Macquart
sophiston Steyskal, Somatia, 835 Stenopyrgota Malloch, 956–960
Sophophora Sturtevant, 1200, 1204–1205 Stenoscinis Malloch, 1143–1144, 1152–1153
sordida (Wiedemann), Neogriphoneura, 974, 977, 980, 985, stercoraria (Linnaeus), Scathophaga, 1267, 1269
992 sternalis Lopes, Eumacronychia, 1330, 1334
sordida (Zetterstedt), Norrbomia, 1185 Sterphus Philippi, 769, 774, 785, 790–792
soror (Williston), Bombyliomyia, 1394, 1400 steyskali (Hernández-Ortiz), Neodyscrasis, 899
sororcula (Wiedemann), Euxesta, 882, 899 steyskali Hernández-Ortiz & Kameneva, Proteseia, 891–892, 899
sororia Williston, Sapromyza, 994 Stichillus Enderlein, 728, 743, 757
sororium Roháček & Barber, Stiphrosoma, 1079 stictica Wiedemann, Ropalomera, 1026
Souzalopesiellinae, 1300 Stictomyia Bigot, 882
spadix (Spencer), Japanagromyza, 1068 stigmatias Loew, Euxesta, 881–883, 899, 901
Spallanzania Robineau-Desvoidy, 1352, 1361, 1412 stigmaticalis Zetterstedt, Microsania, 719
Spathidexia Townsend, 1377, 1393–1394, 1411–1412, 1414 Stilbometopa Coquillett, 1243–1247
Speiseria Kessel, 1249–1250, 1256, 1258–1259 stimulans Thompson, Sterphus, 790
Spelobia Spuler, 1170, 1172, 1175, 1177–1178, 1181, 1183, stipatus Reinhard, Dolichotarsus, 1403
1185–1186 Stiphrosoma Czerny, 1074–1075, 1078–1081
Sphaerina Wulp, 1380, 1412 Stipomorpha Hull, 772
Sphaerocerinae, 1165–1166, 1168, 1187 Stirothrinax Enderlein, 958, 960
Sphaeromitra Roháček & Marshall, 1167–1169 Stizostrebla Jobling, 1254, 1256
Sphaerophoria Lepeletier & Serville, 778, 790–791 Stomatomyia Brauer & Bergenstamm, 1401
Sphecomyiella Hendel, 957–961 Stomosis Melander, 1130–1131, 1135
Spheneuolena Hendel, 877 Stomoxys Geoffroy, 1282–1284, 1293–1294
Sphenometopa Townsend, 1314, 1318, 1332 straminata (Wulp), Guatemalia, 1022
sphenophori (Villeneuve), Lixophaga, 1407 straminea Loew, Opetiophora, 1151
Sphiximorpha Rondani, 768, 790 Strebla Wiedemann, 1250–1252, 1258
Spilochroa Williston, 1161–1163 Streblinae, 1250, 1258–1259
Spilogona Schnabl, 1289, 1291 striata Malloch, Stenolauxania, 993
Spilomyia Meigen, 774, 785, 790, 792 striatella Wulp, Rhagoletis, 940, 951
spinicornis (Loew), Hoplodictya, 1019, 1022, 1024 striatifrons Hendel, Neogriphoneura, 992
spinifera (Leach), Olfersia, 1247 striga Curran, Neogriphoneura, 992
Spinilimosina Roháček, 1170, 1181, 1186 striolata (Wiedemann), Siphunculina, 1144, 1152
spinitibia Malloch, Chaetominettia, 991 Strongygaster Macquart, 1351, 1391, 1397, 1412, 1416
spinosa (Coquillett), Chaetonopsis, 1376, 1401 Sturmiomima Townsend, 1367, 1412
spinosa (Loew), Diedrops, 1223, 1229 Styletta Borgmeier, 747, 757
spinosa Sabrosky, Asteia, 1095 Stylogaster Macquart, 1400, 847–849, 851–855
spiracularis Borgmeier, Gymnophora, 755 Stylogastrinae, 848–849
1440 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

Stylusa Borgmeier & Prado, 756 terraenovae Macquart, Calliphora, 1297, 1303
subnearctica Feijen, Syringogaster, 845 tersum (Townsend), Ophirion, 1410
Suffomyia Freidberg| Suffomyia, 1118 tertia Curran, Neogriphoneura, 992
Suillia Robineau-Desvoidy| Suillia, 1162–1164 Tetanocera Duméril, 1019–1020, 1022–1023
Suilliini, 1161 Tetanocerinae, 1020
suspensa (Loew), Anastrepha, 947 Tetanocerini, 1018–1022
Svarciella Roháček, 1172, 1179, 1184 Tetanoceroides Malloch, 1022
Syllimnophora Speiser, 1293 Tethina Haliday, 1109–1110, 1112–1114
Symmetricella Kessel, 716 Tethininae, 1112–1114
Sympaectria Hendel, 900 Tetrameringia McAlpine, 1043
Synclinusa Borgmeier, 737, 745, 757 Tetramerinx Berg, 1288–1289
Syneura Brues, 735–737, 757 Tetrapleura Schiner, 894, 897, 900
syngenesiae (Hardy), Phytomyza, 1070 Tetreuaresta Hendel, 924, 941, 943, 945, 951, 954
Synthesiomyia Brauer & Bergenstamm, 1285, 1288, 1294 Teutoniomyia Hennig, 1022–1023
Synthesiostrebla Townsend, 1249, 1254, 1256, 1259 texana Hall, Comasarcophaga, 1329
Syringogaster Cresson, 843–846 Thalloptera Borgmeier & Schmitz, 753, 757
Syritta Lepeletier & Serville, 771, 773–774 Thaumatomyia Zenker, 1141, 1145, 1148, 1152
Syrphinae, 764–765, 779, 791 thecata (Coquillett), Telothyria, 1412
Syrphipogon Hull, 770 Thecomyia Perty, 1017, 1019–1021, 1023–1024
Syrphus Fabricius, 763, 775, 777–779, 781–782, 790–791 Thecophora Rondani, 852, 854–855
syssema Hendel, Dictyotrypeta, 948 Thelairaporia Guimarães, 1371, 1392, 1406, 1412
Tabelliphora Kung & Brown, 734, 736, 757 Thelairodes Wulp, 1351, 1371, 1393, 1412
Tachininae, 1344–1345, 1356, 1398 Thelairodoria Townsend, 1347, 1373, 1376, 1412
Tachinini, 1344–1345, 1356–1357, 1362, 1403, 1406 Thelairodoriopsis Thompson, 1373, 1412
Tachinisca Kertész, 909–910, 913, 915, 940 Thelazia Bosc, 1278
Tachiniscinae, 910, 953 Thelyoxynops Townsend, 1379, 1412
Taeniaptera Macquart, 806–809, 811–813 Themira Robineau-Desvoidy, 1034, 1037–1039
Taeniapterinae, 805–807, 811, 813, 846 Thomazomyia Lopes, 1322, 1332, 1334
Taeniomyia Stein, 1272–1274, 1276 Thoracochaeta Duda, 1177, 1186–1187
Tanypeza Fallén, 827–828, 831 Thricops Rondani, 1287, 1289
Tapacura Lopes, 1322, 1332 Thryptocheta, Rondani, 1209
Tapantia Brown & Kung, 730, 732, 738, 757 Thyreophorini, 865, 868
Tarassus Aldrich, 1344 tibialis Schmitz, Conicera, 754
tarsata Wiedemann, Grallomyia, 812 tica Norrbom, Carpomya, 920, 927, 948
Tarsohomoneura Hendel, 972, 977, 980, 982, 995 timida Curran, Neogriphoneura, 992
Tauridion Papp & Silva, 972, 975, 990, 993 tinctinervis (Malloch), Sciosapromyza, 993
Tauroscypson Curran, 964–966 tisciae Papp, Turanodinia, 1051, 1055, 1092
Taxigramma Perris, 1318–1319, 1332 Titanogrypa Townsend, 1315, 1323–1324, 1332
telescopica Gerstaecker, Richardia, 873 tomaterae Steyskal, Melanagromyza, 1069–1070
Telomerina Roháček, 1172, 1181 tomatis Foote, Rhagoletis, 951
Telostylinae, 817 Tomoplagia Coquillett, 910, 913, 928–929, 943, 951–952, 954
Telothyria Wulp, 1370, 1391, 1396, 1412 Tomosvaryella Aczél, 796–797, 801–802
tenax Linnaeus, Eristalis, 787 Tomosvaryellini, 794
Tenthes Cresson, 806, 808–809 Toropamecia Cogan, 999, 1001–1002, 1005
Tenuia Malloch, 823 tortricis (Coquillett), Hemisturmia, 1405
tenuipalpis (Wulp), Hemisturmia, 1405 Toxomerus Macquart, 766, 776, 782, 790–791
Tephritinae, 910–911, 951–953 Toxopyrgota Hendel, 964
Tephritis Latreille, 917, 923, 940–941, 945–946, 950–952 Toxotarsinae, 1301
Tephrochlamys Loew, 1161–1162 Toxotrypana Gerstaecker, 909–910, 919, 941, 947, 951–954
Tephromyia Brauer & Bergenstamm, 1329 Toxurinae, 957
Terellia Robineau-Desvoidy, 917, 923–925 Toxurini, 957–958
teretrura Aczél, Descoleia, 958 trabeculata (Loew), Pherbellia, 1022
Teretrura Bigot, 957–958 Trachyopella Duda, 1179, 1186
Teretrurinae, 957 Trafoia Brauer & Bergenstamm, 1350, 1371, 1384, 1392, 1412
Teretrurini, 957 Traginopinae, 1049, 1052
tergata (Coquillett), Sphenometopa, 1314, 1332 Traginops Coquillett, 1050–1051, 1053, 1055
terminalis (Loew), Mumetopia, 1079 Tranomeringia Sasakawa, 1047–1048
termitophila (Borgmeier), Apodicrania, 753 transversa Maa, Basilia, 1264
Terpnomyennis Kameneva, 892–893, 896, 900 Trapherinae, 905
Terpnomyia Hendel, 894, 897, 900 Travassophora Borgmeier, 749–750, 758
TAXONOMIC INDEX TO VOLUME 2  1441

triangulifera (Loew), Strongygaster, 1412, 1416 Ubristes Walker, 772

Tricharaea Thomson, 1320, 1322, 1327, 1332, 1334 Udamopyga Hall, 1316, 1326, 1328, 1333
Trichobiinae, 1250, 1259 Ulia Becker, 1128–1129, 1135
Trichobioides Wenzel, 1256, 1258 Ulidiinae, 882–883, 889, 901
Trichobius Gervais, 1250–1252, 1256, 1258–1260 Ulidiini, 882–883
Trichodura Macquart, 1384, 1413, 1415 umbrarum (Linnaeus), Dictya, 1022
Tricholauxania Hendel, 972 umbrosus (Loew), Oncodometopus, 973, 980, 992
Trichomorellia Stein, 1286, 1293, 1295 uncinata Hendel, Clasiopella, 1214, 1228
Trichophora Macquart, 1357, 1398–1399, 1413 undata Wiedemann, Pyrgota, 955–956, 961
trichopleura Borgmeier, Brachycephaloptera, 733, 754 undulata Becker, Leptopyrgota, 956, 959–960
Trichopoda Berthold, 1387–1388, 1398, 1405, 1413 unica Spencer, Phytobia, 1064, 1069–1070
Trichopsomyia Williston, 767, 772, 780, 790 unimaculatus Mello & Silva, Physoclypeus, 992
Tricimba Lioy, 1141, 1146, 1151–1152 uniseta Curran, Lonchoptera, 722
tricolor Wheeler, Latheticomyia, 821–824 unispinosa Bigot, Setellia, 873
Triconopsis Hendel, 976 unispinosa (Zetterstedt), Anevrina, 753, 873
trifolii (Burgess), Liriomyza, 1069 Uramya Robineau-Desvoidy, 1392, 1406, 1413
Trigonometopus Macquart, 972–973, 983, 990, 993 ustulata (Reinhard), Cyzenis, 1402
Trigonomma Enderlein, 1144, 1148, 1152 utilis Stone, Procecidochares, 950
Trigonospila Pokorny, 1348, 1397, 1413 Valentibulla Foote & Blanc, 932
trimaculata (Fabricius), Coilometopia, 876, 878 valida Harris, Sphecomyiella, 957, 959–960
Trineurocephala Schmitz, 730, 758 vanderwulpi Townsend, Chaetonodexodes, 1401
Triodontopyga Townsend, 1361, 1413 Vanderwulpia Townsend, 1391, 1413
Tripanurga Brauer & Bergenstamm, 1316, 1320, 1327, 1333 vanduzeei Cresson, Coelopa, 1014–1016
Triphleba Rondani, 729–731, 758 varipalpis Malloch, Desmometopa, 1126–1127
Tripolphopeza Enderlein, 828 varipes Wulp, Chyliza, 841
triquetra (Olivier), Icelia, 1406 venezolana Spencer, Agromyza, 1067
Trisapromyza Shewell, 974, 980, 987, 993 Verrallia Mik, 795–796, 801
tristis Fallén, Thryptocheta, 1209 verticalis Borgmeier, Borophaga, 754
Tritanypeza Enderlein, 828 Veruanus Schmitz, 753, 759–760
Tritoxa Loew, 882, 900 vexatrix (Osten Sacken), Adejeania, 1354, 1398
Trivialia Malloch, 973, 988, 994 viatrix Hendel, Melanoloma, 873, 878–879
trivittata Macquart, Taeniaptera, 806, 808 Vibrissina Rondani, 1348, 1378, 1403, 1413
trivittata Malloch, Rhyncophoromyia, 740, 757 vicina Robineau-Desvoidy, Calliphora, 1303
Trixoscelidini, 1160–1161 vigilans Frey, Metasphen, 812
Trixoscelis Rondani, 1159, 1162–1163 Villegasia Dodge, 1316, 1323, 1333
Trochiloleskia Townsend, 1388, 1396, 1413 vinnula Giglio-Tos, Sapromyza, 993
Tropeopsila Shatalkin, 839 virilis (Aldrich & Webber), Hyphantrophaga, 1405
Trophithauma Schmitz, 732, 741, 744, 748, 758 viriolatus Enderlein, Ptilosphen, 806
Trophodeinus Borgmeier, 734, 739, 750–751, 758 vittata Sabrosky, Eusiphona, 1134
tropica Townsend, Tropidopsiomorpha, 1413 vittigera (Coquillett), Trisapromyza, 974, 980, 994
Tropidomyia Williston, 849, 851, 854 vittipes (Zetterstedt), Eustalomyia, 1275
Tropidopsiomorpha Townsend, 1384, 1413 vittithorax Malloch, Schildomyia, 1054
Tropidothrinax Enderlein (Bolivia), 960 Vladolinia Kameneva, 884, 900
truncaticornis (Wulp), Austeniops, 1399 vockerothi Hardy & Delfinado, Dasyrhicnoessa, 1112
Trupanea Schrank, 934, 941–942, 951–952 vockerothi Thompson, Dideomima, 787
Trypanaresta Hering, 950 Voria Robineau-Desvoidy, 1349, 1376, 1384, 1413
Trypeta Meigen, 929, 931, 941, 944, 951, 953 Voriini, 1356, 1416
Trypetinae, 910, 953–954 vulpinum Hendel, Curtonotum, 1191
Trypetisoma Malloch, 976, 985, 987, 990, 994–995 watsoni Hardy, Apetaenus, 1112
Trypocalliphora Peus, 1300 wenzeli Graciolli & Dick, Metelasmus, 1257
tuberculata Barber, Pseudodinia, 1005 williamsi (Hardy), Microcephalops, 801
Tubiceroides Borgmeier, 757 willistoni Coquillett, Lycastirhyncha, 788
Tucma Mourgués-Schurter, 1168, 1171 Willistoniella Mik, 1025–1030
Tucminae, 1171 Winthemia Robineau-Desvoidy, 1346, 1361, 1370, 1399,
Turanodinia Stackelberg, 1051–1052, 1054–1055, 1092 1413, 1415
turbata (Wulp), Archimimus, 1329 Winthemiini, 1344–1345, 1356–1357, 1361, 1415
tuxeni Hennig, Phanerochaetum, 1123 Wohlfahrtia Brauer & Bergenstamm, 1320
Typhamyza Roháček, 1074 woodi Savage, Callainireinwardtia, 1292
Typhlophorina Silvestri, 756 Woodiphora Schmitz, 749, 758, 760
Typopsilopa Cresson, 1213, 1231–1232 woodorum (Pape), Lepidodexia, 1313
1442 MANUAL OF CENTRAL AMERICAN DIPTERA — VOLUME 2

wulpi (Hendel), Neophyllomyza, 1126 Xenophytomyza Frey, 1066, 1068, 1070

Wulpisca Lopes, 1322, 1327, 1333 Xenopsila Buck, 837, 839–841
wygodzinskyi Thompson, Lycopale, 788 Xenopterella Malloch, 978, 985, 994–995
Xanionotum Brues, 728, 743, 752, 758 Xenothoracochaeta Malloch, 1292
Xanthaciura Hendel, 913, 917, 931, 952 Xenotrichobius Wenzel, 1256
Xanthacrona Wulp, 893, 895, 900, 902 Xylota Meigen, 774, 785, 790, 792
Xanthandrus Verrall, 776, 782, 790 zacatecasense Arnaud & Gelhaus, Trypetisoma, 994
Xanthobasis Aldrich, 1406 Zacompsia Coquillett, 884, 889, 898, 900, 902
xanthocera Collin, Odinia, 1051, 1055 Zaira Robineau-Desvoidy, 1378–1379, 1401–1402, 1414
xanthochaeta Aldrich, Eutreta, 910, 949 Zaleinae, 1112, 1118–1119
Xanthodexia Wulp, 1371, 1396, 1413 Zapriothrica Wheeler, 1199–1200, 1202–1203, 1205–1206
Xanthoepalpus Townsend, 1359, 1395, 1398, 1413 Zealantha Roháček, 1074–1075, 1081
Xanthomelanodes Townsend, 1360, 1377, 1387, 1413 zebra Hendel, Neomyennis, 893
Xanthomelanopsis Townsend, 1390, 1413 zebrinus Mello & Silva, Physoclypeus, 992
Xanthophyto Townsend, 1364, 1368–1369, 1413 Zelatractodes Enderlein, 809, 813
xanthostoma Walker, Prochyliza, 865 Zelia Robineau-Desvoidy, 1353, 1383, 1414
Xanthozona Townsend, 1359, 1413 Zeros Cresson, 1213, 1225, 1231
xanthura (Wulp), Patelloa, 1410 Zikania Borgmeier, 751, 758
Xeniconeura Shewell, 973, 978, 985, 994 Zizyphomyia Townsend, 1367, 1414
xenisma Gaimari & Silva, Choryeuromyia, 991 Zodion Latreille, 848–852, 854–855
Xenochaeta Snow, 947, 953 Zonosemata Benjamin, 910, 913, 917, 924, 952, 954
Xenochaetina Malloch, 973, 982, 984–985, 987, 994 zorrense Griffiths, Lasiomma, 1275
Xenoleucopis Malloch, 998, 1004–1005 Zygothrica Wiedemann, 1198–1199, 1203, 1205–1206
Xenomorellia Malloch, 1285–1286, 1293 Zygozenillia Townsend, 1365, 1414
 Vol. 2
Manual of Central
National Research Conseil national
Council Canada de recherches Canada

American Diptera
True flies, or Diptera, are an exceedingly diverse and

Manual of Central
American Diptera
tremendously important group of animals, not only
because of their impact on human health, but also
because of their varied roles as decomposers, herbivores,
predators, and parasites of other animals. This two-
volume book, a collaboration of over 70 experts on
Diptera, is the first-ever resource for the identification and
understanding of Central American flies. Together, the two
Volume 2
volumes treat all 106 families of flies found in the region,
including discussions of biology, economic importance,
classification, identification to the genus level, as well as
a synopsis of each genus. In a time of major ecosystem
destruction, this work provides the basis for understanding
the biodiversity of one of the major orders of insects in a edited by:
large tropical region. It will be the foundation for future B.V. Brown (head editor)
taxonomic studies for decades to come.
A. Borkent
While Volume 1 includes several introductory chapters and J.M. Cumming
treats 42 families of flies in the Lower Diptera, Volume 2​ ​
covers the remaining 64 families of flies that make
D.M. Wood
up the Higher Diptera (or Cyclorrhapha). These include N.E. Woodley
families of house flies, fruit flies, bot flies, flower flies and
M.A. Zumbado
many other lesser-known groups. The text is accompanied
by over 1660 line drawings and photographs.
Brown
Borkent
Cumming
Wood
Woodley
Zumbado

ISBN-13 978-0-660-19958-0
ISBN 0 - 660 - 19958 - 0

9 780660 199580