Chapter 32

An Introduction to Animal Diversity

Estimates of the number of animal species range from 10-20 million to 100-200 million.

What is an Animal (some characteristics of animals) Multicellular eukaryotic heterotrophs that ingest their food. Lack the cell walls found in plants and fungi Have two unique cell types: nerve cells for impulse conduction
and muscle cells for movement. Nervous and muscular tissues are unique to animals.

Sexual reproduction (mostly), with diploid stage that

dominates the life cycle

All animals and only animals have Homeoboxcontaining family of genes (Hox genes) which control development of body form. Produce a wide diversity of
animal morphology

Reproduction and Development Most animals reproduce sexually

With the diploid stage usually dominating the life cycle

After a sperm fertilizes an egg

The zygote undergoes cleavage, leading to the formation of a blastula

The blastula undergoes gastrulation

Resulting in the formation of embryonic tissue layers and a gastrula

Early embryonic development in animals

1 The zygote of an animal undergoes a succession of mitotic cell divisions called cleavage.

2 Only one cleavage stagethe eight-cell embryois shown here.

3 In most animals, cleavage results in the formation of a multicellular stage called a blastula. The blastula of many animals is a hollow ball of cells.

Blastocoel

Cleavage

Cleavage

6 The endoderm of the archenteron develops into the tissue lining the animals digestive tract.

Zygote

Eight-cell stage Blastocoel Endoderm

Blastula

Cross section of blastula

5 The blind pouch formed by gastrulation, called the archenteron, opens to the outside via the blastopore.

Ectoderm Gastrula
Blastopore

Gastrulation
4 Most animals also undergo gastrulation, a rearrangement of the embryo in which one end of the embryo folds inward, expands, and eventually fills the blastocoel, producing layers of embryonic tissues: the ectoderm (outer layer) and the endoderm (inner layer).

Reproduction and Development Some animals develop directly through transient stages into adults, but others have distinct larval stages. Larva is a sexually immature stage that is morphological distinct from the adult

Larvae undergo metamorphosis, transforming the animal into an adult

Evolution of Animals
The history of animals may span more than a billion years The animal kingdom includes not only great diversity of living species
But the even greater diversity of extinct ones as well

The common ancestor of living animals

May have lived 1.2 billion800 million years ago May have resembled modern choanoflagellates, protists that are the closest living relatives of animals

Single cell Stalk

The common ancestor of living animals

Was probably itself a colonial, flagellated protist
Somatic cells Digestive cavity

Reproductive cells Colonial protist, an aggregate of identical cells Hollow sphere of unspecialized cells (shown in cross section)

Beginning of cell specialization

Infolding

Gastrula-like protoanimal

Neoproterozoic Era (1 Billion524 Million Years Ago)

Early members of the animal fossil record

Include the Ediacaran fauna (From Ediacara Hills of Australia)
Segmented body form

Radial body form

(a)

(b)

Paleozoic Era (542251 Million Years Ago) The Cambrian explosion

Marks the earliest fossil appearance of many major groups of living animals

Paleozoic Era (542251 Million Years Ago) What caused the Cambrian explosion
New predator-prey relationships may have generated diversity through natural selection.

Predators acquired adaptations to catch prey

Prey acquired adaptations that helped them resist predation Rise in atmospheric oxygen may have provided opportunities for animals with higher metabolic rates and larger body sizes Evolution of the Hox complex provided developmental flexibility that resulted in variations in morphology

Mesozoic Era (25165.5 Million Years Ago) During the Mesozoic era
Dinosaurs were the dominant terrestrial vertebrates Coral reefs emerged, becoming important marine ecological niches for other organisms

Cenozoic Era (65.5 Million Years Ago to the Present)

The beginning of this era

Followed mass extinctions of both terrestrial and marine animals

Modern mammal orders and insects

Diversified during the Cenozoic

Animals can be characterized by body plans

One way in which zoologists categorize the diversity of animals is according to general features of morphology and development

A group of animal species that share the same level of organizational complexity is known as a grade
The set of morphological and developmental traits that define a grade
Are generally integrated into a functional whole referred to as a body plan

Animals can be categorized according to the symmetry of their bodies, or lack of it

Symmetry Some animals have radial symmetry

Like in a flower pot

(a) Radial symmetry. The parts of a radial animal, such as a sea anemone (phylum Cnidaria), radiate from the center. Any imaginary slice through the central axis divides the animal into mirror images.

Symmetry Some animals exhibit bilateral symmetry

Or two-sided symmetry
(b) Bilateral symmetry. A bilateral animal, such as a lobster (phylum Arthropoda), has a left side and a right side. Only one imaginary cut divides the animal into mirror-image halves.

Symmetry Bilaterally symmetrical animals have

A dorsal (top) side and a ventral (bottom) side

A right and left side

Anterior (head) and posterior (tail) ends

Cephalization, the development of a head

Tissues Animal body plans

Also vary according to the organization of the animals tissues

Tissues
Are collections of specialized cells isolated from other tissues by membranous layers

Embryonic tissues Animal embryos

Form germ layers, embryonic tissues, including ectoderm, endoderm, and mesoderm

Diploblastic animals
Have two germ layers

Triploblastic animals
Have three germ layers

Body Cavities In triploblastic animals a body cavity may be present or absent A true body cavity is called a coelom and is derived from mesoderm
Coelom Body covering (from ectoderm)

(a) Coelomate. Coelomates such as annelids have a true coelom, a body cavity completely lined by tissue derived from mesoderm.

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

Body cavities A pseudocoelom

Is a body cavity derived from the blastocoel, rather than from mesoderm
Body covering (from ectoderm)

(b) Pseudocoelomate. Pseudocoelomates such as nematodes have a body cavity only partially lined by tissue derived from mesoderm.

Pseudocoelom

Muscle layer (from mesoderm)

Digestive tract (from ectoderm)

Body Cavities Organisms without body cavities

Are considered acoelomates

Body covering (from ectoderm) (c) Acoelomate. Acoelomates such as flatworms lack a body cavity between the digestive tract and outer body wall.

Tissuefilled region (from mesoderm)

Digestive tract (from endoderm)

Protostome and Deuterostome Development Based on certain features seen in early development
Many animals can be categorized as having one of two developmental modes: protostome development or deuterostome development

Cleavage (spiral versus radial) In protostome development

Cleavage is spiral and determinate (spiral = cell division
diagonal to vertical axis of the embryo)

In deuterostome development
Cleavage is radial and indeterminate (radial = the
cleavage is either parallel or perpendicular to the vertical axis of the embryo)
Protostome development (examples: molluscs, annelids, arthropods) Eight-cell stage Deuterostome development (examples: echinoderms, chordates) Eight-cell stage (a) Cleavage. In general, protostome development begins with spiral, determinate cleavage. Deuterostome development is characterized by radial, indeterminate cleavage.

Spiral and determinate

Radial and indeterminate

Cleavage

(determinant versus indeterminate)

Determinate: Ultimate fate of each cell is determined (locked-in) early. Indeterminate: Each early cell retains the potential to develop into a complete embryo!
(this type of cleavage is what makes human identical twins possible)
Protostome development (examples: molluscs, annelids, arthropods)
Eight-cell stage Deuterostome development (examples: echinoderms, chordates) Eight-cell stage

(a) Cleavage. In general, protostome development begins with spiral, determinate cleavage. Deuterostome development is characterized by radial, indeterminate cleavage.

Spiral and determinate

Radial and indeterminate

Coelom Formation
In protostome development
The splitting of the initially solid masses of mesoderm to form the coelomic cavity is called schizocoelous development

In deuterostome development
Formation of the body cavity is described as enterocoelous development
Coelom Archenteron (b) Coelom formation. Coelom formation begins in the gastrula stage. In protostome development, the coelom forms from splits in the mesoderm (schizocoelous development). In deuterostome development, the coelom forms from mesodermal outpocketings of the archenteron (enterocoelous development).

Coelom Mesoderm Blastopore Mesoderm Blastopore Enterocoelous: Schizocoelous: solid folds of archenteron masses of mesoderm form coelom split and form coelom

Fate of the Blastopore In protostome development

The blastopore becomes the mouth

In deuterostome development
The blastopore becomes the anus
Anus Mouth

Digestive tube

Mouth Mouth develops from blastopore

Anus Anus develops from blastopore

Animal phylogenetic tree Zoologists currently recognize about 35 animal phyla The current debate in animal systematics
Has led to the development of two phylogenetic hypotheses, but others exist as well

Animal phylogenetic tree One hypothesis of animal phylogeny based mainly on morphological and developmental comparisons
Ctenophora Arthropoda Phoronida Cnidaria Ectoprocta Mollusca Chordata Platyhelminthes Brachiopoda Echinodermata Annelida Rotifera Nematoda Porifera Nemertea

Radiata

Deuterostomia

Protostomia

Bilateria

Eumetazoa

Metazoa Ancestral colonial flagellate

Animal phylogenetic tree One hypothesis of animal phylogeny based mainly on molecular data
Echinodermata Brachiopoda Chordata Platyhelminthes Ctenophora Cnidaria Phoronida Arthropoda Ectoprocta Nemertea Mollusca Silicarea Annelida Nematoda Calcarea Rotifera

Radiata

Porifera

Deuterostomia

Lophotrochozoa

Ecdysozoa

Bilateria

Eumetazoa

Metazoa

Ancestral colonial flagellate

Points of Agreement All animals share a common ancestor Sponges are basal animals

Eumetazoa is a clade of animals with true tissues Most animal phyla belong to the clade Bilateria
Vertebrates and some other phyla belong to the clade Deuterostomia

Disagreement over the Bilaterians The morphology-based tree

Divides the bilaterians into two clades: deuterostomes and protostomes

In contrast, several recent molecular studies

Generally assign two sister taxa to the protostomes rather than one: the ecdysozoans and the lophotrochozoans

 Ecdysozoans share a common characteristic

They shed their exoskeletons through a process called ecdysis

 Lophotrochozoans
Have a lophophore, a ciliated feeding structure OR Go through a distinct larval stage called a trochophore larva Apical tuft
of cilia

Mouth

(a) An ectoproct, a lophophorate

Anus (b) Structure of trochophore larva