Lecture 9A

Origin and evolution of Paleozoic animals

Chapter 13
Questions:

• When did the earliest animals evolve?

• What did the earliest animals look like? Did they have
skeletons?

• How were the earliest animals preserved?

• Was the rise of animals rapid (geologically speaking)?

• How did animals change the world (e.g., their impact on

sedimentation)?
Origin of animals: When?

First fossil evidence of multicellular animals during the

Ediacaran Period (635 to 541 million years ago).

In fact the name of the period comes from a group of

these organisms: Ediacara fossils.
Ediacara biota

The Ediacara biota appeared globally around 575-540

million years ago. They did not have skeletons and are
preserved as molds and casts.

Dickinsonia Charniodiscus Reconstruction of Ediacaran organisms

Choanoflagellates Sponges Cnidarians Bilaterians

Ediacara biota

Some of the Ediacara fossils resemble

modern animal groups; however the
relationship between most Ediacaran
fossils and modern organisms is
(Xiao, unpublished) unknown.
 First animal trace fossils

The first trace fossils (structures made by the activities of

organisms), made by animals occur around 560 million
years ago.

~550 million years old, South China

Before then, the sediment is largely undisturbed

First skeletonized animal fossils

The first animals that made skeletons occur right at the

end of the Ediacaran Period around 550 million years ago.
- Vase-shaped, calcium carbonate, unknown identity
- Most other fossils are of soft bodied animals
Questions:

• When did the earliest animals evolve?

• What did the earliest animals look like? Did they have
skeletons?

• How were the earliest animals preserved?

Molds and casts

Followed by small
mineralized fossils
Questions:

• When did the earliest animals evolve?

• What did the earliest animals look like? Did they have
skeletons?

• How were the earliest animals preserved?

• Was the rise of animals rapid (geologically speaking)?

• How did animals change the world (e.g., their impact on

sedimentation)?
The Phaneozoic Eon is the most recent
of the four eons that make up the
geological time scale. It runs from 541
Ma to the present day.
The Phaneozoic Eon is the most recent
of the four eons that make up the
geological time scale. It runs from 541
Ma to the present day.

The Phanerozoic is divided into three

eras, the:
Paleozoic,
Mesozoic and
Cenozoic.
 These three eras are separated by major mass extinctions

Remember that there are a fair amount of “squiggles” in this curve. These “squiggles”
more or less form the basis for subdivision of the Phanerozoic geological time scale.

This high-resolution subdivision is possible in the Phanerozoic because (1) the rock record
is more complete than the “Precambrian” one (fewer and shorter unconformities in
younger strata), and (2) the fossil record is more complete thanks to the advent of
widespread biomineralization.
 The Paleozoic Era is
the first in the
Paleozoic and runs
from 541 to 252 Ma.

It consists of the:
Cambrian,
Carboniferous
Ordovician,
Silurian,
Devonian,
Carboniferous and
Permian periods.
 The Cambrian
Period is the first
period in the
Paleozoic Era. It runs
from 541 to 485 Ma.

Key events &

Carboniferous
concepts
• Large animals evolve
and radiate (i.e.,
Cambrian explosion)
• Cambrian substrate
revolution
• Trilobites dominate;
form basis for
stratigraphic
subdivision of
Cambrian
Early Cambrian fossils are small, weird and bear little resemblance to
modern animals. They are commonly referred to as the “small shelly
fossils”

Experimentation in body plan design was intense, with spicular or

discontinuously mineralized skeletons common.

“Pre-trilobite” fossils: Small, shelly and weird

If one fossil type defines the Cambrian, it is the trilobite
(Arthropoda). Trilobites underwent extreme evolutionary
diversification and faunal turnover in the Cambrian,
leading to distinct trilobite “biozones” (i.e., stratigraphic
units with distinct fossil content). These biozones form the
basis for stratigraphic subdivision of the Cambrian Period.

Body (thorax)

Tail (pygidium)
Head (cephalum)

Trilobite (“three lobes”: left, right, and middle)

The Cambrian Explosion
 The Cambrian Explosion of
Animals
• Introduction of major animal groups
• The fossil record of the Cambrian Explosion:
Chengjiang and Burgess Shale
• Consequences of the Cambrian Explosion on
the sedimentary record – substrate revolution
 Animalia

Echino- Brachio-
Porifera Arthropoda Mollusca
dermata poda
Cnidaria Chordata Annelida Bryozoa

Deuterostomata

Proterostomata

Bilateria – 3 tissues

Metazoa - multicellular
 The Cambrian Explosion

• Most of the 35 animal phyla first appeared in the Cambrian

• The (geologically) abrupt appearance of most animal
Dry valley phyla inA modern
in Antarctica: the
Cambrian led to the name Cambrian Explosionanalog to an ancient snowball?
 Fossil Record of the Cambrian Explosion:
Chengjiang and Burgess Shale Fossils

Burgess Shale: Middle Cambrian, ~515 Ma

Chengjiang: Early Cambrian, ~525 Ma

 Chengjiang and the origin of chordates

**
 Living chordates

Sea squirts Amphioxus

 Chengjiang fossils
Early
chordates
Didazoon

Gill slits,
Segments
Haikouella – Chengjiang fossils
 Pr
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Ambulacaria

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Origin of vertebrates

Deuterostomatia

Ve
Chordata

rte
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 Chengjiang Fossils
Myllokunmingia fengjiaoa Dorsal fin
ca. 5 mm

Segmented muscles

Gill pouch Courtesy of Degan Shu

Courtesy of Degan Shu

Myllokunmingia from the Chengjiang is interpreted as a vertebrate chordate

 Chengjiang fossils

Microdictyon
The ~515 Ma Burgess Shale, British Columbia, is one of the world’s best
known sites for Cambrian fossils. Discovered by Smithonian director
Charles Walcott in 1909, it contains an amazing record of Cambrian life.
Walcott worked there until close to his death in 1927, amassing over
65,000 fossils.
Yoho National Park, BC

Charles Walcott
US paleontologist
1850-1927
 The Burgess Shale
• Stephen Formation, Yoho National Park and
Kootenay National Park
The Burgess Shale
 Burgess Shale Fossils

Anomalocaris
Once biomineralization started, skeleton evolution was likely promoted by
the appearance of new kinds of large predators, like Anomalocaris
(“abnormal shrimp”) pictured below, which could reach lengths of up to 2
m.

Up to 2 m!
Anomalocaris was jawless. Here are its “pineapple ring” mouth parts.
(Many have argued that trilobites were Anonomalocaris’ main food
source, whereas others have argued such a mouth would have been
ineffective at biting through a trilobite’s exoskeleton, thus restricting
trilobite predation to times when trilobites moulted.)
 Burgess Shale Fossils

Hallucigenia
https://www.youtube.com/watch?v=p5LMG0fEBb8
Source: kyoichi sasazawa
 Burgess Shale Fossils

Laggania

Sarotrocercus
Source: kyoichi sasazawa
Burgess Shale Fossils

Opabinia Marrella
Source: kyoichi sasazawa
 Burgess Shale Fossils

Aysheaia pedunculata:
An onychophoran

Canadia: An annelid worm

Ottoia: A priapulid worm

Ottoia (http://burgess-shale.rom.on.ca)

Olenoides serratus:
A trilobite, arthropod Echmatocrinus: An echinoderm?
What makes the Burgess Shale so amazing is that the organisms’ soft
parts are exquisitely preserved, in addition to their hard parts. (This is
rarely the case because of bacterial degradation.)
Carbonaceous compression
 Burgess Shale Fossils

Pikaia from the Burgess Shale is interpreted as a fossil chordate

Conodonts are also thought to be early vertebrates that evolved in the
Late Cambrian. Their teeth are widespread in marine rocks and they have
been used extensively to perform biostratigraphy (even before it was
known what they actually were). Well preserved fossils of conodont soft
parts have been recently found in younger strata, confirming that they
were indeed swimming animals.
 Cambrian substrate revolution

The Cambrian substrate revolution involved a transition from more

stable, mat-dominated substrates in the Precambrian to less stable,
bioturbated substrates in the Cambrian (and throughout Phanerozoic).

In the Ediacaran Period, just prior

to the Cambrian, sediment on the
marine seafloor tended to be
stabilized by microbial mats.
Trace fossils were rare and tended
to be confined to the surface of
the substrate (e.g., tracks and
trails). Bioturbation* levels were
nil to very low.

*Bioturbation: Reworking
Ediacaran Cambrian of sediment by organisms.
(Precambrian) (Phanerozoic)
 Cambrian substrate revolution

The Cambrian substrate revolution involved a transition from more

stable, mat-dominated substrates in the Precambrian to less stable,
bioturbated substrates in the Cambrian (and throughout Phanerozoic).

In the Cambrian, more

penetrative traces started to
appear (e.g., burrows). Some
beds in Cambrian strata are
intensely bioturbated. This
indicates that the new
Cambrian organisms were
exploiting the substrate in a
different way than previous
organisms; they were digging
into it and filtering through it
in their search for food, and
they were also living in it. In
particular, bioturbation
intensity really took off with
Ediacaran Cambrian the appearance of trilobites.
(Precambrian) (Phanerozoic)
In fact, the base of the Cambrian is defined by the appearance of
complex, penetrative trace fossils (burrows). Remember, trace fossils
record an organism’s behaviour. Presumably these new complex
burrows record an increase in behavioural complexity
associated with the diversification of
animals at the start of the Cambrian.

In this photo, we’re

looking at the bottom of
the sedimentary bed.

Treptichnus pedum
 Early Cambrian Trace Fossils
Treptichnus pedum: The trace fossil Precambrian-Cambrian Boundary
whose first appearance define the Reference Section, Fortune Head,
Precambrian-Cambrian boundary Newfoundland, Canada

Precambrian-Cambrian Boundary

• Neoproterozoic traces mostly horizontal

• Early Cambrian traces:
• both horizontal and vertical
• much more diverse
• Reveals Cambrian Explosion was also behavioral breakthrough
• Sedimentary consequences of vertical burrowers
Cambrian oceans

Archaeocyathid (sponge) reefs

 The Cambrian Explosion: Summary
• Cambrian Explosion: Explosive evolution of almost all
major animal phyla in the first ~20 million years of
the Cambrian
• Cambrian Explosion as a skeletalization event
• Trace fossils: behavioral breakthrough during the
Cambrian Explosion?
• Ecological and sedimentological consequences of the
Cambrian Explosion: substrate revolution
• Reefs evolving but made of sponges primarily
There are two types, conservation Lagerstätten, which exhibit
The Ordovician
exquisite preservation of in situ (non-transported) hard and soft parts
Period follows the
(e.g., Burgess Shale), and concentration Lagerstätten, which exhibit a
Cambrian Period.
high concentration of hard parts (e.g., bone bed), in some cases
It runs from 485 to
transported largely from elsewhere and concentrated, commonly by
the action of water. 444 Ma.

Carboniferous Hudson-Meng
bison bone bed

Key events &

concepts
• Significant continued
radiation
• Stromatolites decline
• Stratigraphic
subdivision based on
Conservation Lagerstätte Concentration Lagerstätte
graptolites and
conodonts
Ordovician
 The Ordovician Period was marked by another great evolutionary
radiation. Species diversity increased dramatically. Some claim that
Paleozoic diversity reached a peak during the Ordovician (although
according to the graph below, maybe not quite). Pretty much all life was
marine, with the possible exception of some fungi.

**
 Corals appear for the first time...
Tabulata (Ord.-Perm.) Rugosa (Ord.-Perm.)
 Corals establish reefs
• Tabulate corals become important reef
building organisms
• Stromatoporoid sponges also important reef
components
• Together these are known as coral-strome
reefs
• Replace archaeocyathids of the Cambrian
 First Bryozoans
Trepostomata Fenestrata Cryptostomata
(Ord. – Triass.) (Ord. – Perm.) (Ord. – Perm.)
 First Bivalves

Tunnicliff, 1982 Cope, 1999

Floating and swimming animals, in particular nautiloids and graptolites,
became more common in the Ordovician.

Nautiloid: type of cephalopod mollusc

The base of the Ordovician is defined

(approximately) by the appearance of a new type
of organism, the graptolites, in addition to the
appearance of a specific type of conodont teeth.
Most graptolites floated as zooplankton. They tend to be preserved in
marine shale. Marine shale (mudstone) is typically deposited below wave
base in low energy environments, far offshore. They are considered
animals, related to chordates.
Nautiloids (a type of cephalopod mollusk) were swimmers. Like the living
nautilus and other cephalopod mollusks, they likely pursued their prey by
means of jet propulsion and caught them with their tentacles. Cambrian
nautiloids are only known from deposits in China. They diversified rapidly
in the Ordovician and spread throughout the global ocean.
Nautiloids (ord. – recent)
2500 fossil species, 2 or 3 living Nautilus species
Nautiloids (ord. – recent)
2500 fossil species, 2 or 3 living Nautilus
Eurypterids (sea scorpions) (Ord. – Perm.)
Arthropods, most closely related to horseshoe crabs
 Ordovician Trilobites

Isotelus rex
Trilobites underwent a significant extinction near the Cambrian-
Ordovician boundary, but rebounded somewhat, remaining the most
abundant members of many marine communities throughout the
Ordovician.

Permian-Triassic
boundary (approx)

Cambrian-Ordivician
boundary (approx)
Crinoids (a type of echinoderm) are another animal that first appeared
and radiated rapidly in the Ordovician. Modern crinoids still exist today,
though they are much reduced in geographic extent and diversity
compared to the Ordovician.
Modern crinoid

Crinoid
Ordovician limestone is common around Ottawa. It is commonly exposed
along road cuts and is a common building material, especially in older
buildings.

Vittoria
Trattoria
(Byward Market)

Piccolo Grande (Byward Market)

Common fossils in these limestones include trilobites, crinoid stem
fragments, and brachiopods, in addition to rugose corals (horn corals).
Crinoid stem fragments are particularly abundant, and particularly easy to
identify. Next time you see a gray coloured rock, see if you can see any.

Crinoid stem fragments in limestone

Stromatolites declined in diversity in the Ordovician as animals diversified.
This was likely in response to increased grazing pressure, as attested by
the increasingly disrupted layering in stromatolites from animals that
bored through them. These modified stromatolites referred to as
thrombatolites.

Stromatolite

Thromabatolite
Ordovician oceans
 Extinctions through Earth History

1)End
1) LateOrdovician
Ordovician

2)Late
2) LateDevonian
Devonian

3)End
3) EndPermian
Permian

4)End
4) EndTriassic
Triassic

5)End
5) EndCretaceous
Cretaceous
At the end of the Ordovician, Gondwanaland migrated to the south pole and
became glaciated. Climate had switched to “ice-house” conditions. As water
became incorporated into the ice sheets, global sea level fell, generating a
widespread unconformity. The ice house world was short lived—it only lasted a 0.5
to 1 million years—and the sea level fluctuation was correspondingly rapid and
short lived. The trigger for this Late Ordovician glaciation is poorly understood, but
may have been related to a brief reduction in greenhouse warming.

Glacial striations in the Libyan Desert

Many believe that the glaciation initiated Late Ordovician
mass extinction, during which ~50% of the organisms died.

What happens to shallow marine life when glaciers are extensive?

 Main Late Ordovician victims

90% of trilobites Most Nautiloids

Some Tabulate corals