A

brief evolutionary history

of animals
Lecture 1
 Evolutionary history of animals
• Evolution is the unifying theme in “Nothing in Biology
biology because it connects observed makes sense except
patterns with processes. in the light of
• Organisms are modified descendants evolution”
of common ancestors.
• Evolution continually modifies
organisms
• The characteristics of organisms are
thus a mixture of ancestral traits
derived from their ancestors and more
recently derived traits
• By the light of evolution Dobzhansky is
referring to
• The evolutionary history of organisms
• The evolutionary relationship among
organisms
• The process of of evolution itself (e.g.
mutation, selection, etc.) Theodosius Dobzhansky
How to join the animal club?
Animal taxonomic diversity

To a rough approximation and setting aside

vertebrate chauvinism, it can be said that
essentially all organisms are insects

- R. M. May 1988 – Science 241-1146

The big picture
 ARI 28 October 2014 12:59

Ctenophora

Animals Porifera
Placozoa
Cnidaria
Xenacoelomorpha
Parahoxozoa
Ambulacraria Echinodermata
Planulozoa Hemichordata
Deuterostomia
Cephalochordata

Chordata Urochordata
Bilateria Craniata
Chaetognatha
Bryozoa
Entoprocta
Cycliophora
Nephrozoa
Annelida

Trochozoa Mollusca
Nemertea
Brachiopoda
Phoronida
Spiralia Gastrotricha
Protostomia Platyhelminthes
Gnathostomulida
Micrognathozoa
Nucleariida Gnathifera
Rotifera
Fungi
Orthonectida
thokonta Filasterea
Dicyemida
Ichthosporea
Priapulida
Holozoa Animals Scalidophora
Loricifera
Choanoflagellata Kinorhyncha
Nematoida Nematoda
Ecdysozoa Nematomorpha
Tardigrada
Panarthropoda Onychophora
Arthropoda

e1
othesis of animal phylogeny, compiled across multiple studies. Black dots denote clades that have
consensus across studies. Red dots denote clades that have poor or conflicting support or whose exact
sition is uncertain. The organism silhouettes were illustrated by Noah Schlottman and submitted to
Pic (www.phylopic.org). They are available for reuse under the Creative Commons Attribution-
Alike 3.0 Unported license.

ous approaches. Some of the relevant diversification events happened in quick succession
reds of millions of years ago, which resulted in relatively few informative characters for
relationships (Rokas & Carroll 2006). Though some of these hard problems may never
solved, there are several reasons to be optimistic that coming years will see continued
ess.
The first animal fossils: the Ediacaran fauna
 The Ediacaran fauna: the first animals

Kimberella. Earliest Bilateralia (mollusc??) (555-558 MYA)

Fedonkin and Waggoner 1997. Nature 388, 868-871

The first animal fossils: the Ediacaran fauna
• First complex multicellular organisms.
• Unclear exactly what they are and how they relate to living animals.
• For instance one species (Charnia) was described as an alga but also as a
coral, fungi, lichen, or a colony of myxobacteria…
• The ecology of ediacaran animals is largely unknown and still debated.
Many are suspected to be filter feeders /osmotrophic.
• High abundance indicating high productivity.
• The ediacaran animals (if they are indeed animals) look like nothing alive
today.
The big bang of animal evolution
The big bang of animal evolution
“There is another and allied difficulty, which
is much graver. I allude to the manner in
which numbers of species of the same
group, suddenly appear in the lowest known
fossiliferous rocks”

- Charles Darwin
 The Cambrian Fauna

Charles Walcott (1850-1927) Burgess Shale

The Cambrian Fauna

Marrella Aysheaia
Hallucigenia Anomalocaris
 ooding of 1. C. R. Ma
e of Earth Number of taxa (2006).
the exten- 20 40 60 80 100 120 140 2. D. H. Erw
443 3. D. H. Erw
weathered

Ord.
Construc
put of cal- lage, CO
485 Phyla Classes
and other Millions of years 4. C. J. Low

Cambrian
oncentra- 5. K. J. Pete
6. D. H. Erw
ost three- (2002).
541
this input EB
7. N. J. But
8. R. H. T. C
Ediacaran

origin of
(2009).
t of phos- 9. D. J. E. M
ent flux to 98 (201
10. R. Wood
635
11. S. E. Pet
e is a via-
The origins of vertebrates
• Haikouichtys
• 530 MYA (Cambrian)
• Eyes, segmented muscles, skull and a
backbone made out of cartilage
• Stem craniate

Haikouichtys
 The Cambrian Explosion: summary
• The Cambrian explosion refers to
• The sudden apparition of most animal phyla (including the chordates) in the fossil record in a span of 15-20 MYA
• Rapid diversification of many animal phyla
• Emergence of marine ecosystems with “modern” trophic structures
• Many morphological innovations: large body size, legs, skeletons, teeth, eyes.
• Potential causes of the Cambrian explosion include:
• O2 necessary for large body size and high metabolic rate but O2 levels may have been sufficient during the
Ediacaran period.
• Calcium. Spike in calcium concentration during the Cambrian may have facilitated the the development of
skeletons. Calcium segregation may have evolved as a detoxification mechanism.
• Predation and competition may have favored diversification of animals.
• Exploitation of the sea floor. The availability of organic matter combined with greater O2 availability opened the
sea floor niche. Burrowing organisms further created more ecological niche (ecosystems engineers) i.e. the
substrate revolution.
 Ordovician (488-443 Mya)
Diversity of Life through Time

3500 Modern
Palaeozoic
Cambrian
3000

2500
Number of general

2000

1500

1000

500

0
Cm O S D C P Tr J K Pg N

500 400 300 200 100 0

Time (Ma)

1 Global Phanerozoic diversity for marine genera (based on Sepkoski, 2002) showing the trajectories of the three evolutionary faunas. Only genera
 The Great Ordovician Biodiversification Event
• Arguably the most important and sustained increase of marine
biodiversity in Earth’s history during which marine paleobiodiversity
tripled.
• Rapid (over 25 ma) diversification at the order, family, genus, and
species level.
• Caused by a combination of factors including
• Greatest continental dispersal of the Paleozoic.
• Warm climates
• High sea levels
• The largest tropical shelf area of the Phanerozoic.
• Increased phytoplankton availability and high nutrient input to the
oceans driven by intense volcanic activity.
• “explosion” of both zooplankton and suspension feeding organisms.
Ordovidician: terrestrialization
Ordovician: mass extinction Diversity of Life through Time

3500 Modern
Palaeozoic
Cambrian
3000

2500
Number of general

2000

1500

1000

500

0
Cm O S D C P Tr J K Pg N

500 400 300 200 100 0

Time (Ma)
 Ordovidician: terrestrialization & extinction
• First evidence of multicellular land life and development of terrestrial
ecosystems but fossils of terrestrial animals are scant
• Molecular clock analysis suggest that terrestrial arachnids have originated
in the ordovician
• End of the Ordovician (439 Mya) is marked by the first of the big 5 mass
extinctions
• Eliminated 85% of marine species, 55% of genera, 22% of families, but
few orders and no classes or phyla.
• Occurred in 2 phases: a cooling phase followed by a warming phase
caused major environmental changes.
Devonian (420-350 Mya) Eophalangium

• First terrestrial ecosystems with

plants, fungi, and arthropods
• Increasing plant cover
• First deciduous forests
Figure 1 Eophalangium sheari gen. et sp. nov. Holotype (female), slide no. 3503. A harvestman (Opiliones) from
the Rhynie chert, Aberdeenshire, Scotland. Early Devonian (Pragian). (a) Lateral section close to the midline
showing the outline of the compact body. Note the mouthparts, anteriorly positioned optic tubercle and distinctly
‘hunchbacked’ appearance. The elongate internal structure is the ovipositor; (b) Reverse side of same specimen
at a more lateral plane showing a dark internal gut trace in the opisthosoma and the branching tracheal system
originating just behind the last leg coxa. The structures are labelled in Fig. 2 (scale bars=1 mm).

differences or muscle apodemes (Shultz 2000). Ventrally, the body surface of the fossil. There is some suggestion of a
cuticle is smooth and does not show division into sternites. swelling around the anal region, but individual sclerites –
Extant harvestmen are sometimes quite heavily armoured, but which can be quite complex around the anus in living taxa –
there is no ornament of spines or tubercles anywhere on the are not preserved.
Devonian: the age of fish
Devonian: Fins to limbs transition
 Access provided by Carleton University (Canada) on 07/15/15. For personal u
Annu. Rev. Ecol. Evol. Syst. 2008.39:571-592. Downloaded from www.annualr
a b c
10:36

10:36
Devonian: Fins to limbs transition
Humerus
Radius
Ulna

b c
b c
h i
d e f g
Humerus
Humerus
Radius
Radius
Ulna
Ulna

e e ff gg Figure 4
Eusthenopteron Tiktaalik Acanthostega
Fin and limb skeletons. (a) Sauripterus, a rhizod
Long (1989) and Garvey et al. (2005). (c) Tiktaa
adual carbon dioxide downdraw from the
due to biological and chemical weathering of
two (Figure 1). All three extinction pulses are estimated to
have occurred in the past 100 000–300 000 years of the
cks.
Frasnian (Walliser, 1996). A final extinction pulse occurred
End of Devonian mass extinction approximately 374.5 My bp in the earliest Famennian

Zone
tion 374.0

Early triangularis
Famennian
• Second of the big 5 mass
vonian extinction is one of the ‘Big Five’ mass
extinction
hat have occurred in the past 600 My of Earth
other four being the end-Ordovician, end- 5
d-Triassic• Loss of 70% and 82% of

Geological time (Ma BP)

and end-Cretaceous mass extinc- 374.5 4
marine species

Extinction pulse
ate Devonian extinction is unusual in that it 3

linguiformis
hin the last epoch of the Devonian Period, at
• Two most probable causes are 2
y between the Frasnian and Famennian
not at theanoxia (oxygen deprivation)
end of the geological period, as

Frasnian
r four massand global cooling (glaciation
extinctions. See also: Extinction: 375.0
and/or volcanism/meteorite)
n Mass Extinction; Extinction: End-Triassic

Late rhenana
1
a: Evolution & Diversity of Life

375.5
ge R (November 2012) Extinction: Late Devonian Mass
Figure 1 Temporal pattern of biotic diversity loss in the Late Devonian
 Carboniferous (359-299 Mya)
• Radiation of amphibians
and insects on land
• High levels of
atmospheric oxygen
• Spread of forests
• Beginning of
Meganeura
exploitation of plants by
insects (galls, resin)
• First amniotes Pederpes
Eucritta melanolimnetes
Carboniferous (359-299 Mya)
 Dimetrodon

Moschops

Dicynodon Gorgonops
End pf Permian Mass extinctions (251 Mya) Diversity of Life through Time

3500 Modern
Palaeozoic
Cambrian
3000

2500
Number of general

2000

1500

1000

500

0
Cm O S D C P Tr J K Pg N

500 400 300 200 100 0

Time (Ma)
Permian (298-252 Mya):summary
• Diversification of the amniotes: three groups of amniotes
are present: (anapsids, diapsids, synapsids)
• Synapsids become the dominant group of amniotes
• Diversification of herbivorous tetrapods
• Ends with the greatest mass extinction of all time: Permo-
triassic mass extinctions (85% of all species perished)
• Massive volcanic eruption that lasted several million years
caused extreme environmental changes
• Long recovery but resulted in the change from the paleozoic
fauna to the modern fauna in marine ecosystems
Triassic (250-200 mya)
• Radiation of the diapsids and take-over of the
terrestrial environment
• Predatory fishes and reptiles dominate oceans
•Origin and radiation of the dinosaurs. They will
dominate the terrestrial environment for 180 my.
• Origins of birds and mammals Silesaurus (early dinosaur ?)

Morganucodon (late
triassic mammaliaform)
Shonisaurus (late triassic reptile)
 Cretaceaous-Paleogene mass extinction
(65.5mya)
Diversity of Life through Time

• Asteroid impact and/or 3500 Modern

volcanism caused
Palaeozoic
Cambrian

extreme environmental
3000

changes 2500

Number of general
• Resulted in the 2000

extinction of dinosaurs 1500

(except birds) and 1000

provided an opportunity
for the radiation of
500

mammals 0
Cm O S D C P Tr J K Pg N

500 400 300 200 100 0

Time (Ma)