Animal Phylogeny

Saturday, June 09, 2007

2:01 PM

METAZOA or ANIMALIA
Eukaryotic
Multicellular (except
Myxozoa)
Heterotrophic
No cell wall
Collagen
Gastrulation
Character state terms you should be
familiar with:
1. Plesiomorphy- ancestral state
2. Apomorphy - derived state
3. Synapomorphy - shared derived state

How do zoologists categorize

animals?
Morphology and development
Molecular data

Cladograms illustrating phylogenetic groupings: Monophyly

(upper left and right), Polyphyly (lower left), Paraphyly (lower
right). Only monophyletic groups are considered true clades
(natural taxonomic group)
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Homology of forelimbs
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Body symmetry: Radial (a), Bilateral (b)

Convergent evolution
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Even though starfishes (P Echinodermata) are radially

symmetrical, they are in fact more closely related to you than to
a jellyfish. Their bilateral ancestry can be seen in their larvae.

E Aurellado Zoology 3 Page 1

Embryonic development
Sunday, June 10, 2007
11:59 PM

E Aurellado Zoology 3 Page 2

Body Cavities & Molecular Data

Monday, June 11, 2007
1:59 PM

Body plans according to coelom formation

Traditional animal phylogeny (A) and current (B).

Adoutte et al. (2000) Proc. Natl. Acad. Sci. USA 97, 4453-4456

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Do molecular data support the traditional or the current phylogeny? Recent analyses are now relying on
genomic data rather than single genes to reconstruct animal phylogeny. See the following articles supporting
either one of the hypothesis:
Coelomata: Rogozin et al. 2007. Ecdysozoan clade rejected by genome-wide analysis of rare amino acid
replacements. Mol Biol Evol 24:1080-1090. [Link]
Ecdysozoa: Irimia et al. 2007. Rare coding sequence changes are consistent with Ecdysozoa, not Coelomata.
Mol Biol Evol. Epub May 27, 2007 . [Link]

E Aurellado Zoology 3 Page 3

Kingdom Protista (protists)

Sunday, June 17, 2007
3:59 PM

Protista
1. Unicellular
2. Eukaryotic
3. Specialized organelles

Choanoflagellates:
sister group of animals

Protistan phyla (Traditional Classification)

Sarcodina

Pseudopodia

Mastigophora

Flagellum

Ciliophora

Cilia
Macro- and micronucleus

Apicomplexa

Apical complex
No obvious locomotory structures
Endoparasitic

What is the difference between cilia and flagellum?

Protista is a paraphyletic group

The phylum Myxozoa, once thought

to be protists, have been found to
be members of K Metazoa

Current "supergroups" of Eukaryotes

Simpson & Roger. 2004. The real "kingdoms" of Eukaryotes. Curr Biol 14:R693-R696.
Plantae

Plastids from primary endosymbiosis of a cyanobacterium

Volvox, land plants

Excavata

Distinctive feeding groove (cytostome) of excavate type

Trypanosoma, Trichonympha, Euglena

Rhizaria

Filopodia or axopodia
Forams & radiolarians

Chromalveolata

Plastids from 2ndary endosymbiosis of a red alga

Ciliates, dinoflagellates, apicomplexans, Opalina

Amoebozoa

Lobopodium
Amoeba, Arcella

Opisthokonta

Single posterior flagellum

Choanoflagellates, animals and fungi

E Aurellado Zoology 3 Page 4

Phylum Porifera (sponges)

Saturday, June 09, 2007
12:23 PM

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Porifera (L porus =pore, ferre = to bear)

Asymmetrical or radially symmetrical
Cellular level of organization
Body covered with pores (ostia)
Specialized cells , e.g., choanocytes
Sessile
Aquatic
Generalized sponge body plan
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Classes of Porifera
Hexactinellida

Glass sponges
6-rayed siliceous spicules
Syncitial cells
Syconoid or leuconoid

Demospongiae

Siliceous spicules (not 6-rayed)

May have spongin
Leuconoid

Calcarea

Calcareous spicules
Asconoid, syconoid, leuconoid

Not all sponges are filter-feeders

Class Sclerospongiae is no longer a valid class

Carnivorous sponge Asbestopluma

(Demospongiae)
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Sponge body forms

Hexactinellidae: Euplectella sp.
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Why do freshwater
sponges have
gemmules as part of
their life cycle?
Calcarea: Leucosolenia eleanor
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0000+0000+0605+1284>

Demospongiae: Spongilla sp.

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E Aurellado Zoology 3 Page 5

Phylum Cnidaria
Thursday, June 07, 2007
8:53 PM

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Cnidaria (Gk. cnidos = stinging nettle)

Cnidocytes
Planula larva
Radial symmetry
Tissue level of organization
Diploblastic
Nerve net & musculo-epithelial cells
Aquatic

Cnidarian polyp and medusa forms

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Classes of Cnidaria
Anthozoa

True corals, sea anemones, etc.

Polyp form with mesenteries

Scyphozoa

True jellyfishes
Medusa dominant, polyp reduced or
absent

Cubozoa

Box jellyfishes
Medusa dominant w/ velarium

Hydrozoa

Hydroids, siphonophores, etc.

Polyp w/o mesenteries and/or
medusa w/ velum

Cnidocytes
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Why shouldn't you use freshwater to treat

jellyfish stings?

Anthozoa: Acropora, staghorn coral

Photo : E Aurellado

Cubozoa: Carukia barnesi, Irukandji jellyfish

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Hydrozoa: Hydra w/ bud and ovary

Scyphozoa: left, jellyfish (photo: E Aurellado);
right, Aurelia scyphistoma
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What contributed to the diversity of cnidarians?

Polymorphism
Colony formation
Coral formation

E Aurellado Zoology 3 Page 6

Phylum Platyhelminthes (flatworms)

Tuesday, December 26, 2006
11:08 PM

BILATERIA

Bilaterally symmetrical -- evolution of actively hunting animals

Cephalization-- development of a head region
where sensory structures are concentrated

Triploblastic -- allows for development of organ systems

Ectoderm
Endoderm
Mesoderm
Protostomes: blastopore becoming the mouth
Lophotrochozoa: trochozoa & lophophorates

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General Characteristics of flatworms:

Dorsoventrally flattened
Acoelomate
Incomplete digetive system
Ladderlike nervous system with simple
brain
Protonephridial excretory system
No vascular system
Circular & longitudinal muscles
Mostly hermaphroditic
Tegument syncitial in parasitic forms

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[Link]

1.

Dugesia

Classes
1. Turbellaria

Free-living
Covered with ciliated epidermis
Well-developed sensory structures, e.g., eyespots

2. Monogenea

Ectoparasitic flukes
Attachment organs (prohaptor & opisthaptor)
Single host

3. Trematoda

Endoparasitic flukes
2 suckers: oral and ventral
Complex life history: intermediate & definitive hosts

4. Cestoda

Tapeworms, parasitic
Scolex, neck, strobila made of proglottids

2. Dactylogyrus
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[Link]

[Link]

4.
[Link]
3. Schistosoma is an example of a dioecious flatworm

E Aurellado Zoology 3 Page 7

Life cycles of parasitic flatworms

Wednesday, December 27, 2006
12:07 AM

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How are endoparasitic

flatworms adapted to their
lifestyle?
Organs for attachment
Can withstand host's defenses
High fecundity

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E Aurellado Zoology 3 Page 8

Phylum Nemertea (ribbon worms)

Thursday, December 28, 2006
11:50 AM

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Nemertea / Rhyncocoela
Rhyncocoel - cavity that operates the proboscis
"Acoelomate"
Complete gut
Blood system
Protonephridial excretory system
Serial gonads
Mostly marine

What are the advantages of a

complete gut?
Allows linear processing of food and
specialization of gut regions
Increases feeding efficiency
Could allow increase in body size

Pilidium larva of nemertines

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Classes (traditional classification)

Enopla

Proboscis armed with a stylet

Anopla

Unarmed proboscis

Anoplan nemertine: Lineus bilineatus

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Enoplan nemertine showing everted proboscis

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E Aurellado Zoology 3 Page 9

Phylum Nematoda (roundworms)

Wednesday, December 27, 2006
3:18 PM

Ecdysozoa: protostomes that molt (ecdysis)

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Nematoda (Gk. nema = thread):

Slender cylindrical body
Pseudocoelomate w/ high hydrostatic pressure
Complete digestive system w/ muscular pharynx
Collagenous cuticle
Mostly dioecious
Longitudinal muscles, no circular muscles
No circulatory system
Brain & longitudinal nerve cords
Sensory structures
a. anterior amphids
b. posterior phasmids

What are the advantages of a tube-within-tube

pattern of pseudocoelomates and coelomates?
Allows fluid circulation
Hydrostatic skeleton
More room for internal organs to develop

Classes (traditional classification)

Free-living nematodes

What is eutely?

1. Secernentea

With phasmids
e.g., Ascaris, Ancylostoma
(parasitic), C. elegans, T. aceti
(free-living)

2. "Adenophorea"

Without phasmids
e.g., Trichuris, Trichinella
(parasitic)

Caenorhabditis elegans

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Females larger than males

males with copulatory spicules, often
a bent tail or copulatory bursa

Ascaris

Ancylostoma

Vinegar Eel 1 (Turbatrix aceti)

[Link]
inegar_Eel_1/vinegar_eel_1.htm
Screen clipping taken: 12/27/2006, 2:11 PM

Trichuris

Trichinella

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E Aurellado Zoology 3 Page 10

Phylum Rotifera (rotifers)

Wednesday, December 27, 2006
4:15 PM

Rotifera (wheel bearers):

1. Head bears ciliated crown called corona
2. Pseudocoelomate
3. Modified muscular pharynx called
mastax containing trophi
4. Body comprised of head, trunk & foot
(bears pedal glands)
5. Microscopic
6. Dioecious, some species parthenogenetic
7. Maybe a lophotrochozoan
8. Mostly freshwater, planktonic or benthic

[Link]
Screen clipping taken: 12/27/2006, 6:25 PM

Classes
Seisonidea

Epizoic on gills of a marine crustacean,

Nebalia
Corona reduced
Paired gonads
Reproduce sexually

Bdelloidea

Monogononta

1 gonad
Reduced corona
Alternate parthenogenesis (amictic)
and sexual (mictic) reproduction

Seison (Seisonidea)
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Corona divided into 2 parts

Paired gonads
Parthenogenetic, no boys allowed
Swimming or crawling (like a leech)
Bdelloidea
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Brachionus quadridentatus
(Monogononta)
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Monogonont life cycle

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What is the advantage of cryptobiosis and embryonic

diapause in rotifers capable of asexual reproduction?
How are these adaptations related to their habitats?

E Aurellado Zoology 3 Page 11

Both bdelloid rotifers and

nematodes can undergo
cryptobiosis-- a state of suspended
animation under extreme
conditions, e.g., dessication,
freezing, and heating.
Fertilized eggs of monogonont
rotifers can also lie dormant in
adverse conditions (embryonic
diapause)

Phylum Mollusca (mollusks)

Thursday, December 28, 2006
2:43 PM

Trochozoa : coelomate protostomes w/ trochophore

larva
Mollusca (Gk. Molluscus = soft)
Radula (rasping tongue)
Ctenidium
Mantle covering visceral mass
Muscular foot
Hemocoel
Nephridial excretory system
Molluscan shell not a defining characteristic, although
it contributed to the diversification of the phylum

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Trochophore larva
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What advances are made possible by a true

coelom?
Independent movement of gut and body wall
Development of more complex respiratory,
circulatory, and excretory systems

Classes
1. Neomeniomorpha
2. Chaetodermomorpha

Vermiform bodies
Spiculate

3. Polyplacophora

8 dorsal plates
Spiculate

4. Monoplacophora

Calcareous shell
serial replication

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Calcareous shell
Reduced serial
replication

Scaphopoda
Bivalvia
Gastropoda
Cephalopoda

Basic molluscan body plan

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Chaetodermomorpha: no foot & pedal groove

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Polyplacophora (chitons)
Chiton - Wikipedia, the free encyclopedia
[Link]
Screen clipping taken: 12/28/2006, 6:24 PM

Neomeniomorpha: foot w/ pedal groove

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E Aurellado Zoology 3 Page 12

Aculifera: spiculate molluscs

Conchifera (shelled molluscs)

Thursday, December 28, 2006
5:27 PM

What are the advantages and disadvantages of a shell?

Monoplacophora (ventral view): single

shell; serial repetition of kidneys, gonads
& gills

V. Horigue

Scaphopoda: Captacula, no
ctenidium
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Bivalvia: Tridacna (giant clam)

2 lateral shells; no radula

A. Spring

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V. Horigue
A. Spring

Gastropoda: Left, queen conch; right, nudibranch

Single shell, absent in nudibranchs; gills or lung

Gastropods exhibit a condition called torsion. Why

did it evolve and what are its disadvantages?

Cephalopoda: top, Nautilus; bottom,

Octopus;
Siphon & tentacles derived from footmantle; well-developed nervous system;
closed circulatory system; shell
internal/absent in coleoids, external in
Nautilus

Internal anatomy of a squid (Cephalopoda)

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E Aurellado Zoology 3 Page 13

Phylum Annelida (segmented worms)

Friday, December 29, 2006
2:07 PM

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Annelida (L. annulus = little ring)

Segmented body
Coelom divided by septa
Chitinous setae
Circular and longitudinal muscles
Closed circulatory system
Complete gut
Solid ventral nerve cord
Respire through skin, gills or parapodia
Metanephridial excretory system

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Metamerism: serial repetition of segments

(somites) along the length of the animal
Criteria of true metamerism:
Mesodermal
Embryonic development
Penultimate budding
What are the advantages of metamerism?
More efficient locomotion
Functional independence among segments
Provides a framework for specialization

Classes (traditional classification)

Polychaeta
(bristleworms &
tubeworms)

Many setae in parapodia

Mostly marine
Separate sexes

Oligochaeta
(earthworms)

Few setae,no parapodia

Clitellum
Hermaphroditic

Hirudinea (leeches)

No setae nor parapodia

Clitellum
Hermaphroditic

Fanworm with radioles exposed

Photo by VVHilomen

x-s Polychaete showing parapodia w/ setae

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Ophryotrocha (Polychaeta)

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Basic Annelid Structure

Prostomium
Peristomium (1st true segment)
Segments
Pygidium (w/ anus)
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Clitellate annelids: top, Oligochaeta;

bottom, Hirudinea

Metanephridia
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E Aurellado Zoology 3 Page 14

Phylum Arthropoda
Friday, December 29, 2006
3:58 PM

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Arthropods are the largest phylum (>80%

species) in the animal kingdom .
Arthropoda (arthros = joint, podus = foot)
Jointed appendages
Chitinous exoskeleton
Segmented body, tagmatization
Special respiratory structures , e.g., gills,
tracheae or book lungs
Open circulatory system, hemocyanin
Well-developed sensory structures
Excretory system i.e., malpighian tubules and
coxal glands for terrestrial arthropods
Dioecious, fertilization external in aquatic
arthropods and internal in terrestrial
arthropods

Tagmatization - fusion of body segments into

specialized functional groups. Each specialized
group is called a tagma (pl. tagmata)
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Vertebrate (left) vs. Arthropod joint

(right).
Muscles attach to the exoskeleton and
the cuticle at the joints are thin and
flexible, allowing for rapid movement.

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Arthropod Subphyla
Chelicerata

Lack antennae
Tagmata: prosoma and opisthosoma
Chelicerae, pedipalps & 4 prs. of walking legs

"Myriapoda"

1 pair of antennae
Low level of tagmatization
Many segments

Crustacea

Biramous appendages, 2 prs. antennae

Mainly aquatic, nauplius larva
Head, thorax (or cephalothorax) & abdomen

Hexapoda

Uniramous appendages, 1 pr. antennae

Mainly terrestrial
Head, thorax, & abdomen
6 legs

Myriapods are often classified with hexapods under "Uniramia",

but myriapods may actually be allied with Chelicerata, thus forming
the clade "Paradoxopoda"
Myriapoda may be a polyphyletic group

E Aurellado Zoology 3 Page 15

Trilobites are an extinct group of

arthropods, distinguished by the
following characters:
a body built from a cephalon,
thorax, and pygidium
a body divided into three lobes,
running from head to tail
one pair of antennae
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Crustacea
Friday, February 16, 2007
4:48 PM

Nauplius larva

Daphnia: C. Branchiopoda, O.
Cladocera
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Fairy shrimp: C.
Branchiopoda, O.
Anostraca
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Argulus: Branchiura
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Copepod: C. Maxillopoda
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C. Ostracoda

Euphausia: C. Malacostraca

Balanus : Cirripedia
Photo by E Aurellado

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crab: C. Malacostraca,
O. Decapoda
Biramous appendage of crustaceans

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E Aurellado Zoology 3 Page 16

Cuticle of crustaceans is further

hardened by calcium salts to
add thickness and deter
predators. Because they live in
a buoyant medium, weight is
not a problem.

Paradoxopoda: Chelicerates and Myriapods

Friday, February 16, 2007
5:06 PM

Chelicerate Classes
Merostomata

Horseshoe crabs
Limulus, Tachypleus, Carcinoscorpius
Marine
Book gills

Pycnogonida

Sea spiders
Marine
No respiratory or excretory structures

Arachnida

Carcinoscorpius rotundicauda
C. Merostomata, O. Xiphosura

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Spiders, scorpions, mites & ticks

Terrestrial
Waxy cuticle
Book lungs

C. Pycnogonida
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Arachnid anatomy
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Typical myriapod body plan

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Myriapod Classes
Chilopoda

Centipedes
Poison claws
1 pair of legs/segment

Diplopoda

Millipedes
2 pairs of legs/segment

Symphyla

Resemble centipedes
Not all segments bears legs

Pauropods

Resemble millipedes
9 prs. of legs (> no. segments)

Although all myriapods are terrestrial, they are

confined to damp and humid environments.

E Aurellado Zoology 3 Page 17

Hexapoda (insects)
Friday, February 16, 2007
3:49 PM

What contributed to the diversity of insects?

Ability to invade terrestrial habitats
Co-evolution with angiosperms
Evolution of flight (in pterygote insects)

3 tagmata of insects
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Tracheal system of insects

Butterfly: Order Lepidoptera
Photo by E Aurellado
Advantages of flight:
1. Means of avoiding predation
2. Facilitates food access
3. Means of dispersal

Beetle: O. Coleoptera
Photo by E Aurellado

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What adaptations enabled insects to minimize desiccation

and thus, dominate land?
Epicuticular wax
Tracheal system with valves
Elimination of uric acid (insoluble in water)
Ability to reabsorb water in rectum

E Aurellado Zoology 3 Page 18

Phylum Echinodermata
Thursday, December 28, 2006
7:46 PM

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Deuterostomes
Blastopore forms anus
Coelom formed by outpuching (enterocoelous)
Radial cleavage
Echinodermata (echinos = spiny, derma = skin)
Water vascular system
Calcitic endoskeleton of ossicles
Mutable connective tissue
Pentaradial symmetry (bilateral larvae)
Decentralized nervous system (nerve ring)
No blood or excretory system
Marine

Water vascular system

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Classes
Crinoidea

Sea lilies & feather stars

Asteroidea

Sea stars

Ophiuroidea

Brittle stars & basket stars

Echinoidea

Sea urchins & sand dollars

Holothuroidea

Sea cucumbers

Pedicellariae

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Ophiuroidea: 5 arms sharply

marked off; no anus

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"Concentricycloidea" Sea daisies

Brachiolaria (Asteroid)
Ashley Spring

Crinoidea: mainly sessile; ciliated

ambulacral groove; mouth upward

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Asteroidea: typically 5 arms,

Mouth w/ anus

Echinoidea: globose;
long spines &
covered in test;
aristotle's lantern

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Holothuroidea:
Elongates; few
dermal ossicles;
respiratory tree

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Ophiopluteus (Ophiuroid)

Doliolaria (Holothuroid &

Crinoid)

Xyloplax n. sp.
[Link]. 208:
7780.(April2005)
[Link]
g/cgi/reprint/208/2/77

Echinopluteus (Echinoid)

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Concentricycloidea : Xyloplax medusiformis, X. turnerae,

and a new species from N. Pacific. They are named after
their WVS of 2 concentric rings; could be highly modified
sea stars.

E Aurellado Zoology 3 Page 19

Phylum Hemichordata
Saturday, March 03, 2007
5:25 PM

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Hemichordata ("half-string")
Body divided into 3 parts: proboscis, collar and trunk
Gill slits
Stomochord
Dorsal tubular nerve cord
Hemichordate Classes

Enteropneusta

Acorn worms

Pterobranchia

Sessile & colonial worms

With lophophore (hollow
tentacles derived from
extensions of the mesocoel)

Saccoglossus kowalevskii (Enteropneusta)

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What evidences support the close relationship between

hemichordates and echinoderms?
Similarity between the tornaria larva of hemichordates
and the bipinnaria of asteroidea
Trimerous body plan of hemichordates and echinoderms
Left coelomic pouches become dominant as the right
pouches are lost in both phyla
Molecular evidence

Cephalodiscus (Pterobranchia)
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What is the fate of the 3 coelomic sacs in hemichordates and

echinoderms?
Comparison between tornaria and
bipinnaria larvae
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Coelom

Hemichordate

Echinoderm

Protocoel

Proboscis

Axocoel

Mesocoel

Collar & collar pore

Hydrocoel (WVS &

madreporite)

Metacoel

Trunk

Somatocoel

Nature 444, 85-88 (2 November 2006)

Deuterostome phylogeny reveals monophyletic chordates and the new phylum
Xenoturbellida
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This article in Nature reports that the enigmatic Xenoturbella, a worm-like animal
without a brain, gut or gonads discovered in 1949 belongs to a phylum of its own,
not to Mollusca as previously thought. P Xenoturbellida may be closely related to
Echinodermata and Hemichordata.

E Aurellado Zoology 3 Page 20

Xenoturbella
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Phylum Chordata (chordates)

Friday, January 05, 2007
8:23 PM

1.
2.
3.
4.
5.

Chordata (chorda = string)

Notochord
Dorsal hollow nerve cord
Pharyngeal gill slits
Post-anal tail
Subpharyngeal endostyle or thyroid gland
Chordate Subphyla

4 main chordate characters

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Urochordata

Tunicates

Tunic (cellulose-like tunicin)

Notochord at the tail
Blood w/ high concentrations
of vanadium
Marine filter feeders

Cephalochordata

Lancelets

All chordate characters

persist in adult
Marine filter feeders

Vertebrata

Vertebrates Cranium
Vertebral column
Segmented muscles

Oikopleura (Larvacea)
Pelagic, solitary; forms mucus "house"
to trap food
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Salp (Thaliacea )
Pelagic, solitary or colonial, barrelshaped
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Inversion Hypothesis
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Rhopalaea crassa (Ascidiacea)

Benthic, solitary, colonial or
compound; barrel-shaped
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Adult (left) and "tadpole" larva (right) of an ascidian

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Branchiostoma (Amphioxus, subPhylum Cephalochordata)

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E Aurellado Zoology 3 Page 21

Megalodicopia hians (Sorberacea)

Solitary, deep sea benthic, predatory
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Vertebrate Classes
Myxini &
Jawless fishes
Cephalaspidomorphi Hagfishes and lampreys
Chondricthyes

Sharks, rays, and chimaeras

Cartilaginous endoskeleton
no operculum & swim bladder

Osteichthyes

Bony fishes
Operculum & swim bladder

Amphibia

Frogs & salamanders

Aquatic tadpole
Cutaneous respiration

Reptilia

Turtles, lizards, snakes &

crocodilians
Epidermal scales

Aves

Birds
Feathers

Mammalia

Mammals
Milk-producing mammary glands
Hair/fur

Tetrapods

Jawed vertebrates

Saturday, March 03, 2007

6:26 PM

Amniotes

Subphylum Vertebrata

Vertebrate phylogeny (modified from Brooks/Cole

2001)
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Evolution of jaws from anterior gill arches

Agnatha
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Amniote egg
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Chondrichthyes (photo: A. Spring)

Amphibia (photo : J Baril)

Aves
Osteichthyes (Photo: E Aurellado)

Reptilia (photo: A Spring)

Which invertebrate chordate group is the closest relatives of the vertebrates?

Many molecular studies strongly suggest the close affinity between
cephalochordates and vertebrates. However, in early 2006, an article in Nature
has shown molecular evidence claiming tunicates may actually be the closest
relatives to vertebrates. This debate is likely to continue in the near future.

E Aurellado Zoology 3 Page 22

Mammalia
Photos: E Aurellado