Chapter 31

Animal Phylogeny, Acoelomates,

and Protostomes

Russell, Biology: The Dynamic Science, 5th edition. © 2021 Cengage. All Rights Reserved. May not be scanned, copied or
duplicated, or posted to a publicly accessible website, in whole or in part.
Animals
• Zoologists have described nearly 2 million living
species in the kingdom Animalia
• Most are invertebrates, animals without a
backbone
• About 65,000 are vertebrates, animals with a
backbone

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31.1 What Is an Animal?
• Animals are eukaryotic, multicellular organisms
• Unlike plants and fungi, their cells lack cell walls
• In large animals, specialized tissues and organ
systems deliver nutrients and oxygen to cells and
carry wastes away
• All animals are heterotrophs

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What Is an Animal? (cont'd.)
• Animals use oxygen to metabolize food
• All animals are motile at some time in their lives –
in some species, the young are motile and adults
are sessile
• Most animals have sensory and nervous systems

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What Is an Animal? (cont'd.)
• Animals reproduce either asexually or sexually
• Sexually reproducing species produce haploid
gametes, which fuse to form diploid zygotes
• A zygote develops into a multicelled embryo, then
into an immature juvenile or free-living larva,
which develops into a reproductively mature adult

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Animal Origins
• The common ancestor of all animals was probably
a colonial, flagellated protist that lived at least 700
million years ago
• Ernst Haeckel proposed that
• (1) the colonial, flagellated ancestor was a hollow,
ball-shaped organism with unspecialized cells,
• (2) its cells became specialized for particular
functions, and
• (3) developmental reorganization produced a
double-layered, sac-within-a-sac body plan

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Animal Origins

A. Hypothesized evolution of a two-layered animal body plan

Digestive cavity

Feedingcells

1 Colonialflagellated 2 Certain cells became 3 Adevelopmental reorganizationproduced

ancestor with specialized for feeding a two-layered animal with a sac-within-a-sac
unspecializedcells and otherfunctions. bodyplan.

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31.2 Key Innovations in Animal Evolution
• Several key morphological innovations were used
to develop hypotheses about the evolutionary
relationships of the major animal groups:
• Tissue structure
• Body symmetry
• Body cavities
• Developmental patterns
• Segmentation

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Tissues and Tissue Layers
• Tissues (groups of cells with a common structure
and function) divide animals into two branches:
• Sponges (Parazoa) lack tissues
• All other animals (Eumetazoa) have tissues
• In eumetazoan development, embryonic tissues
form either two or three primary cell layers
• A diploblastic body plan
• Most animals are triploblastic

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Primary Cell Layers
• Endoderm, the innermost layer, develops into the
lining of the gut.
• Ectoderm, the outermost layer, forms the external
covering and nervous system.
• Mesoderm, the middle layer in tribloblastic
animals, forms the muscles of the body wall.

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Body Symmetry
• Most sponges have irregular shapes and are
asymmetrical
• Most animals are symmetrical – their bodies can
be divided by a plane into mirror-image halves
• Eumetazoans exhibit one of two body symmetry
patterns:
• Radiata have radial symmetry
• Bilateria have bilateral symmetry

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Bilateral Symmetry
• Bilaterally symmetrical animals show
cephalization.
• A cut along the midline from head to tail divides
them into left and right sides
• Bilaterally symmetrical animals have anterior
(front) and posterior (back) ends, and dorsal
(upper) and ventral (lower) surfaces

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Patterns of Body Symmetry

Radialsymmetry Bilateral symmetry

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Body Cavities
• Acoelomate animals have no body cavity
• Pseudocoelomate animals have a
pseudocoelom.
• Coelomate animals have a body cavity (coelom)
completely lined by a membrane (peritoneum)
derived from mesoderm

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Figure 32.8
(a) Coelomate (Eg Earthworm)
Coelom
Body covering
(from ectoderm)

Tissue layer
lining coelom
Digestive tract and suspending
(from endoderm) internal organs
(from mesoderm)

(b) Pseudocoelomate (Eg roundworms)

Body covering
(from ectoderm)

Pseudocoelom Muscle layer

(from
mesoderm)
Digestive tract
(from endoderm)

(c) Acoelomate (Eg Flatworm)

Body covering
(from ectoderm) Tissue-
filled region
(from
mesoderm)

Wall of digestive cavity

(from endoderm)
Body Cavities (cont'd.)
• Body plans of pseudocoelomate and coelomate
animals form a “tube within a tube”
• The digestive system forms the inner tube
• The body wall forms the outer tube
• The fluid-filled body cavity, containing the internal
organs, lies between them
• Incompressible fluid within the body serves as a
hydrostatic skeleton

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Developmental Patterns
• Bilaterally symmetrical animals are divided into
two lineages
• Protostomes includes most phyla of invertebrates
• Deuterostomes includes the vertebrates and their
nearest invertebrate relatives

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Cleavage
• Cleavage (early mitotic divisions of the fertilized
egg):
• Protostomes- spiral cleavage
• Deuterostomes- radial cleavage
• Many protostomes undergo determinate
cleavage
• Many deuterostomes have indeterminate
cleavage

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Cleavage Patterns

Protostomes Deuterostomes
A. Cleavage
Four-cell embryo Four-cell embryo

Top Top
view view

Eight-cell embryo Eight-cell embryo

Axis Axis

Top Side Top Side

view view view view

Spiral cleavage Radial cleavage

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Origin of Mouth and Anus
• After cleavage, some cells migrate inward through
the blastopore which forms the developing gut
(archenteron)
• A second opening at the opposite end of the
embryo turns the archenteron into a digestive tube
• In protostomes, the blastopore develops into the
mouth and the second opening forms the anus
• In deuterostomes, the blastopore develops into the
anus and the second opening forms the mouth

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The Mouth and Anus
C. Origin of mouth andanus
Anus Mouth

Coelom

Gut

Mouth Anus

Blastopore develops into Blastopore develops into

mouth; anus forms later. anus; mouth forms later.

KEY
Derivatives of ectoderm Derivatives of endoderm
Derivatives of mesoderm Coelom (body cavity)

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Segmentation
• Some protostomes and deuterostomes exhibit
segmentation.
• In humans and other vertebrates, segmentation is
seen in mesodermal tissues.
• In some animals, such as earthworms,
segmentation.

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Molecular Phylogeny of Animals

Platyhelminthes
Brachiopoda

Echinodermata
Onychophora
Cnidaria

Ctenophora

Annelida
Ectoprocta
Porifera

Chordata
Rotifera
Phoronida

Mollusca

Arthropoda
Nemertea

Hemichordata
Nematoda
Segmentation Segmentation

Pseudo- Pseudo-
coelom coelom

No body Trochophore Segmentation

cavity larva

Lophophore

Spiral cleavage Radialcleavage

Ecdysis

LOPHOTROCHOZOA ECDYSOZOA

Schizocoelom Enterocoelom
Segmentation?

PROTOSTOMIA DEUTEROSTOMIA

Diploblastic Triploblastic

Radialsymmetry Bilateralsymmetry

RADIATA BILATERIA

No tissues Tissues

PARAZOA EUMETAZOA

COLONIAL
CHOANOFLAGELLATE
ANCESTOR

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Phylogenetic Tree of Animals
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

segmentation
commonancestor deuterostome

Deuterostomia
development Chordates

bilateral
symmetry Echinoderms
3 tissue layers
body cavity segmentation
moltingof
cuticle Arthropods

Ecdysozoa
tissue
layers Roundworms

segmentation

Bilateria
Annelids

Protostomia
trochophore Molluscs
multicellularity

Eumetazoa
choanoflagellate

Lophotrochozoa
ancestor
Flatworms

protostome
development Rotifers

“Lophophorans”

lophophore ctenophore

Ctenophores

Radiata
Cnidarians
radial symmetry
2 tissue layers

Parazoa
Sponges 37
31.4 Animals without Tissues: Parazoa
• Sponges (phylum Porifera):
• Lack true tissues.
• Mature sponges are sessile

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Asymmetry in Sponges

Watkins, Bruce/Animals Animals

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Sponges

A. Spongebody plan B. Sponge bodywall D. Venus’ flowerbasket (Euplectella species)

Water out
through osculum
Spicules

Osculum Mesohyl
Amoeboid
cell

Donald Fawcett/Visuals Unlimited, Inc.

Pore
Porocyte

Flattened
Spongocoel cells of
pinacodem

C. Choanocyte Food particles

trapped inmucus
Water in
through Colla
pores r

Flagellum Nucleus
Microvilli Phagocytosis
of collar of foodparticles

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Reproduction in Sponges
• Most sponges are hermaphroditic.
• Flagellated larvae are released into the
environment, where they attach to substrates
and undergo metamorphosis into sessile
adults.

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31.5 Eumetazoans with Radial Symmetry
• Eumetazoans have true tissues
• Eumetazoans with radial symmetry (Radiata)
can sense stimuli from all directions, an
adaptation for life in open water
• Two phyla: Cnidaria and Ctenophora

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Cnidarians
• Examples:
o Hydrozoans
o Scyphozoans (Jellyfish)
o Cubozoans (Box Jellyfish)
o Anthozoans (Corals and Sea Anemones)

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Cnidarian Body Plans

A. Cnidarian body plans B. Cnidarianpolyp C. Body wall cellsand tissues

Epidermis Tentacle with

Medusa Mesoglea
Mesoglea nematocysts Oral end of
Mouth Gastrodermis Epidermis
Gastrodermis body
Gastrovascular Nutritive
cavity cell
Cnidocyte
Mouth forming
food
vacuole
Polyp Epidermis
Nematocyst
Gastrodermis Gastro-
Mouth
Epidermis Gastrovascular cavity vascular
(extends to base ofpolyp) cavity
Gastrodermis Receptor
Mesoglea Cilium
Gastrovascular cell
(beneathepidermis) Contractile
cavity
Secretions
Mesoglea Nerve net cell Nerve cell
(beneathepidermis) Glandcell
Longitudinal muscle
Nucleus fibrils
Aboral endof
body Circular muscle
fibrils

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Predation by Cnidarians
A. Hydra consuming a crustacean

Kim Taylor/Bruce Coleman Ltd.

Kim Taylor/Bruce Coleman Ltd.

B. Cnidocytes Discharged
Operculum nematocyst
(capsule's lid Barbs
at cnidocyte's
free surface)
Trigger
(modified
cilium)

Nematocyst
coiled inside
capsule

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Cnidarian Life Cycle
• Many cnidarians exist in only the polyp or the
medusa form.
• The polyp often produces new individuals
asexually.
• The medusa is often the sexual stage
• The four lineages of Cnidaria differ in the form that
predominates in the life cycle

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Scyphozoan
A. Scyphozoan (Chrysaora quinquecirrha)

© Laitr Keiows/Shutterstock.com

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Cubozoan
B. Cubozoan (Chironex fleckeri)

Mesoglea-
filled bell

Tentacles

Visual&Written SL/Alamy

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 Anthozoans
© Cecilia Lim H M/Shutterstock.com

A. Staghorn coral B. Sea anemone (Urticina Iofotensis) escape behavior

(Acropora cervicornis)

Mark_Doh/iStock/Getty Images Plus

Tentacle of one polyp

F. S. Westmorland

F. S. Westmorland

F. S. Westmorland
Interconnected
skeletons of polyps
of a colonial coral

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Ctenophores: Comb Jelly

Combs

blickwinkel/Alamy
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31.6 Lophotrochozoan Protostomes
• The group Bilateria have bilateral symmetry
and a greater variety of tissues than the Radiata
• Bilaterians have organ systems, structures
that include two or more tissue
• Most have a coelom or pseudocoelom

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Molecular Phylogeny of Animals

Platyhelminthes
Brachiopoda

Echinodermata
Onychophora
Cnidaria

Ctenophora

Annelida
Ectoprocta
Porifera

Chordata
Rotifera
Phoronida

Mollusca

Arthropoda
Nemertea

Hemichordata
Nematoda
Segmentation Segmentation

Pseudo- Pseudo-
coelom coelom

No body Trochophore Segmentation

cavity larva

Lophophore

Spiral cleavage Radialcleavage

Ecdysis

LOPHOTROCHOZOA ECDYSOZOA

Schizocoelo Enterocoelom
m
Segmentation
?
PROTOSTOMIA DEUTEROSTOMIA

Diploblastic Triploblastic

Radialsymmetry Bilateralsymmetry

RADIATA BILATERIA

No tissues Tissues

PARAZOA EUMETAZOA

COLONIAL
CHOANOFLAGELLATE
ANCESTOR

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The Lophophorate Phyla
• Lophophorates (phyla Ectoprocta,
Brachiopoda, and Phoronida) have a
lophophore.
• The lophophore serves as a site for gas exchange
and waste elimination as well as for food capture
• Most lophophorates are sessile suspension-
feeders as adults.

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Lophophorate Animals: Ecotprocta

blickwinkel/Alamy

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Lophophorate Animals: Brachiopoda

© Scott Leslie/AgeFotostock
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Lophophorate Animals: Phoronida

Marevision/AGE Fotostock
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Phylum Platyhelminthes: Flatworms
Digestive system
Branching gastrovascular cavity
Eg of an acoelomate

Pharynx
Ocelli Mouth (protruded)
Nervous system
Ganglion

Ventral nerve cords

Reproductive system
Ovary Testis Oviduct Genital pore

Penis

Excretory system
Flame cells

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Phylum Platyhelminthes:Turbellaria

© Cory Gray
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Phylum Platyhelminthes:Trematode Liver Fluke

Yolk Gastrovascular Oral

Testes Ovary glands cavity sucker

E. R. Degginger/Science Source
Uterus Ventral Pharynx
sucker

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Phylum Platyhelminthes: Cestoda: Tapeworm
and Scolex
A. Tapeworm B. Scolex

Scolex

Photo Researchers/Science Source/Getty

Proglotti ds

Andrew Syred/Science Source

Images

Scolex

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Rotifers

A. Rotifer (Philodina roseola) body plan B. Rotifer (Philodina species)

Corona
Mouth
Mastax
(food-
grinding
organ)

Excretor
y
system

James W. Evarts/Science Source

Stomach

Intestine
Cloaca
(a storage
chamber for
digestive and
excretory
wastes)
Anus

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Ribbon Worms Phylum Nemertea

A. Ribbon worm (Lineus species) B. Ribbon worm anatomy

Proboscis Rhynchocoel

Mouth Intestine Proboscis Anus

retractor muscle

Marevision/Age Fotostock
Everted
proboscis

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Polyplacophora: Chiton

George Wood/Dreamstime.com
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Gastropoda

B. Terrestrial snail C. Marine nudibranch

(Helix pomatia) (Flabellina iodinea)

Thornberry/iStockphoto.com
KOO/Shutterstock.com

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Bivalves

B. Giant clam (Tridachnagigas) C. Geoduck (Panope generosa)

Tom McHugh/Science Source

treetstreet/iStock/Getty Images

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Three Lineages of Annelida
• Oligochaetes worms (Clitellata;
formerly Oligochaeta):
• Primarily terrestrial, restricted to moist
habitats
• Example: earthworms
• Leeches (Clitellata; formerly Hirudinea)
• Mostly freshwater, blood-sucking parasites
• Secrete hirudin to prevent blood
coagulation

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Oligochaetes: Earthworms
Eg of a coelomate

© mashe/Shutterstock.com
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Hirudinea: Parasitic Leeches

Leech before feeding Leech after feeding

J. A. L. Cooke
J. A. L. Cooke

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31.7 Ecdysozoan Protostomes
• Ecdysozoa have a protective external covering
that they shed periodically
• Three phyla:
• Nematoda
• Arthropoda

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Nematoda: Roundworms
Eg of a Pseudocoelomate

Alistair Dove/Image Quest Marine

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Phylum Arthropoda
• Arthropods:
• Include more than half the animal species on Earth
– insects, spiders, crustaceans, millipedes,
centipedes, extinct trilobites, and their relatives
• Segmented body is encased in a rigid exoskeleton
• Muscles attached to the inside of the exoskeleton
move individual body parts
• Body segments are fused in various ways,
reducing the overall number of segments

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Ecdysis in Insects

Mitsuhiko Imamori/Minden Pictures

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Phylum Arthropoda (cont'd.)
• Three body regions (head, thorax, and abdomen)
are specialized with highly modified paired
appendages
• Heart pumps hemolymph from the hemocoel
through an open circulatory system
• Gas exchange mechanisms vary by group
• Highly organized central nervous system, touch
receptors, chemical sensors, compound eyes
with multiple image-forming units, and in some,
hearing organs

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Chelicerata
• Arachnids and horseshoe crabs:
• First pair of appendages (chelicerae) are fanglike
structures.
• Second pair of appendages (pedipalps)
• Two major body regions: prosoma (fused head
and thorax) and opisthosoma

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Chelicerata (cont'd.)
• Arachnida (spiders, scorpions, mites, and ticks):
• Scorpions and some spiders have book lungs
• Spiders inject poisons and digestive enzymes into
prey –some weave webs from silk secreted by
spinnerets
• Mites feed on plants or animals; ticks feed on blood
• Merostomata (horseshoe crabs):
• Carnivorous bottom feeders in shallow coastal
waters

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Spider

A. Wolf spider (Lycosa species) B. Spider anatomy

Prosoma Opisthosoma

Cathy Keifer/Shutterstock.com
Digestive system
Eye Brain Heart Excretory organ

Poison
gland

Chelicera
Anus
Pedipalp

Mouth Book lung Ovar Spinnerets

Leg y Silk gland
s

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Arachnids

C. Scorpion D. House dust mite E. Spider (Argiope species)

(Tityus species) (Dermatophagoides on web
pteronyssinus)

Cathy Keifer/Shutterstock.com
Andrew Syred/Science Source
Chelicerae
Ingo Arndt/Getty Images

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Marine Chelicerates

DARLYNE A. MURAWSKI/National Geographic Creative

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Myriapoda
• Centipedes and millipedes:
• Two body regions: a head with one pair of
antennae, and a trunk with one (in centipedes) or
two (in millipedes) pairs of walking legs on most of
its many segments
• Centipedes are fast predators with powerful toxins
• Millipedes are slow but powerful herbivores or
scavengers

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Myriapods
A. Millipede (Spirobolus species)

Julian Money-Kyrle/Alamy
B. Centipede (Scolopendra species)

© Audrey Snider-Bell/Shutterstock
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Crustaceans
• Shrimps, lobsters, and crabs:
• Many have a fused cephalothorax and separate
abdomen
• Exoskeleton may include a carapace
• Five pairs of appendages on the head: two pairs of
antennae, one pair of mandibles, two pairs of
maxillae
• Chemical and touch receptors in antenna,
compound eyes, statocysts on the head, sensory
hairs embedded in the exoskeleton throughout the
body

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Decapod Crustaceans

A. Crab (Ocypode species) B. Lobster (Homarus americanusC). Lobster external anatomy

Herve Chaumeton/Agence Nature Eyes (one pair) Fused segments Segmented
of cephalothorax abdomen
Antennae
© EcoPrint/Shutterstock

(two pairs) Carapac

e
Telson
Maxillipeds
(three pairs)

Swimmerets

Cheliped
Four pairs of Uropods
walking legs Sperm transfer
appendage

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Hexapoda
• Insects and their relatives:
• The most diverse and successful animals on Earth
• Head includes multiple mouthparts, a pair of
compound eyes, and one pair of sensory
antennae
• Thorax bears three pairs of walking legs and often
one or two pairs of wings made of chitin and
sclerotin
• Abdomen includes much of the digestive system
• Gases exchange through a tracheal system
• Wastes are excreted through Malpighian tubules

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Insect Diversity

Steven Russell Smith Photos/Shutterstock.com

Stephen Dalton/Minden Pictures/Getty Images

Henrik Larsson/Shutterstock
Biosphoto/Bartomeu Borrell
A. Silverfish (Thysanura, B. Dragonflies (Odonata, C. Male praying mantids D. This rhinoceros beetle
Lepisma saccharina) are Epitheca cynosura) have (Mantodea, Mantis religiosa) (Coleoptera, Lucanus cervus) is
wingless, an ancestral trait aquatic larvae that are active are often eaten by the larger one of more than 250,000 beetle
within insects. predators; adults capture females during or immediately species that have been
other insects in mid-air. after mating. described.

Michael Durham/Minden Pictures

Cabezonication/istockphoto.com
Nigel Cattlin/Science Source

bpperry/istockphoto.com
E. Fleas (Siphonoptera, F. Crane flies (Diptera, Tipula G. The luna moth H. Like many other ant species,fire
Ctenophalides canis) have species) look like giant mosquitoes, (Lepidoptera, Actias luna), ants (Hymenoptera, Solenopsis
strong legs with an elastic but their mouthparts are not useful like other butterflies and invicta) live in large cooperative
ligament that allows these for biting other animals; the adults moths, has wings that are colonies. Fire ants—named for their
parasites to jump on and off of most species live only a few covered with colorful painful sting—were introduced into
their animal hosts. days and do not feed at all. microscopic scales. southeastern North America, where
they are now serious pests.

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Insect Development: No Metamorphosis

Egg Young Adult

Some wingless insects, like silverfish (Thysanura), do not

undergo a dramatic change in form as they grow.

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Incomplete Metamorphosis

Egg Nymphs Adult

Other insects, such as true bugs (Hemiptera), have

incomplete metamorphosis; they develop from nymphs into
adults with relatively minor changes in form.

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Complete Metamorphosis

Egg Larvae Pupa Adult

Fruit flies (Diptera) and many other insects have

complete metamorphosis; they undergo a total
reorganization of their internal and external anatomy
when they pass through the pupal stage of the life cycle.

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