113-Intro Animal Diversity-Chapter 32

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    Chapter 32

    An Introduction to Animal

    DiversityEstimates of the number of animalspecies range from 10-20 million to100-200 million.

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    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 conductionand muscle cells for movement. Nervous and muscular tissues are unique to animals.

    Sexual reproduction (mostly), with diploid stage thatdominates the life cycle

    All animals and only animals have Homeobox-containing family of genes (Hoxgenes) whichcontrol development of body form. Produce a wide diversity ofanimal morphology

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    Reproduction and Development

    Most animals reproduce sexually

    With the diploid stage usually dominating thelife cycle

    After a sperm fertilizes an egg

    The zygote undergoes cleavage, leading to theformation of a blastula

    The blastula undergoes gastrulation

    Resulting in the formation of embryonic tissuelayers and a gastrula

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    Zygote

    Cleavage

    Eight-cell stage

    Cleavage

    Blastula Cross sectionof blastula

    Blastocoel

    Blastocoel

    Gastrula Gastrulation

    Endoderm

    Ectoderm

    Blastopore

    Early embryonic development in animals

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

    3

    The endoderm ofthe archenteron de-

    velops into the tissuelining the animals

    digestive tract.

    6

    The blind pouchformed by gastru-

    lation, calledthe archenteron,

    opens to the outsidevia the blastopore.

    5

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

    4

    Only one cleavagestagethe eight-cellembryois shown here.

    2The zygote of an animalundergoes a succession of mitoticcell divisions called cleavage.

    1

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    Reproduction and Development

    Some animals develop directly through

    transient stages into adults, but others havedistinct larval stages.

    Larva is a sexually immature stage that is

    morphological distinct from the adult

    Larvae undergo metamorphosis, transformingthe animal into an adult

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    Evolution of Animals

    The history of animals may span more than a billionyears

    The animal kingdom includes not only great diversity ofliving species

    But the even greater diversity of extinct ones as well

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    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 ofanimals

    Single cell

    Stalk

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    The common ancestor of living animals

    Was probably itself a colonial, flagellatedprotist

    Colonial protist,

    an aggregate ofidentical cells

    Hollow sphere ofunspecialized

    cells (shown incross section)

    Beginning of cellspecialization

    Infolding Gastrula-likeprotoanimal

    Somatic cells Digestivecavity

    Reproductive cells

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    (a) (b)

    Neoproterozoic Era (1 Billion524 Million Years Ago)

    Early members of the animal fossil record

    Include the Ediacaran fauna (From Ediacara Hills of Australia)

    Radial bodyform

    Segmentedbody form

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    Paleozoic Era (542251 Million Years Ago)

    The Cambrian explosion

    Marks the earliest fossil appearance of manymajor groups of living animals

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    Paleozoic Era (542251 Million Years Ago)

    What caused the Cambrian explosion

    New predator-prey relationships may have generated diversitythrough natural selection.

    Predators acquired adaptations to catch prey

    Prey acquired adaptations that helped them resist predation

    Rise in atmospheric oxygen may have provided opportunities foranimals with higher metabolic rates and larger body sizes

    Evolution of the Hoxcomplex provided developmental flexibilitythat resulted in variations in morphology

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    Mesozoic Era (25165.5 Million Years Ago)

    During the Mesozoic era

    Dinosaurs were the dominant terrestrialvertebrates

    Coral reefs emerged, becoming important

    marine ecological niches for other organisms

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    Cenozoic Era (65.5 Million Years Ago to the Present)

    The beginning of this era

    Followed mass extinctions of both terrestrialand marine animals

    Modern mammal orders and insects

    Diversified during the Cenozoic

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    Animals can be characterized by body plans

    One way in which zoologists categorize the diversity ofanimals is according to general features of

    morphology and development

    A group of animal species that share the same level oforganizational complexity is known as a grade

    The set of morphological and developmental traits thatdefine a grade

    Are generally integrated into a functional whole

    referred to as a body plan

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

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    Symmetry

    Some animals have radial symmetry

    Like in a flower pot

    Radial symmetry. The parts of aradial animal, such as a sea anemone(phylum Cnidaria), radiate from thecenter. Any imaginary slice throughthe central axis divides the animalinto mirror images.

    (a)

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    Symmetry

    Some animals exhibit bilateral symmetry

    Or two-sided symmetry

    Bilateral symmetry. A bilateralanimal, such as a lobster (phylum

    Arthropoda), has a left side and aright side. Only one imaginary cutdivides the animal into mirror-imagehalves.

    (b)

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    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

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    Tissues Animal body plans

    Also vary according to the organization of theanimals tissues

    Tissues

    Are collections of specialized cells isolatedfrom other tissues by membranous layers

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    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

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    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)

    Digestive tract(from endoderm)

    Tissue layerlining coelomand suspending

    internal organs(from mesoderm)

    Coelomate. Coelomates such asannelids have a true coelom, a bodycavity completely lined by tissue

    derived from mesoderm.

    (a)

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    Body cavities

    A pseudocoelom

    Is a body cavity derived from the blastocoel,rather than from mesoderm

    PseudocoelomMuscle layer(frommesoderm)

    Body covering

    (from ectoderm)

    Digestive tract(from ectoderm)

    Pseudocoelomate. Pseudocoelomatessuch as nematodes have a body cavity onlypartially lined by tissue derived frommesoderm.

    (b)

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    Body Cavities

    Organisms without body cavities

    Are considered acoelomates

    Body covering

    (from ectoderm) Tissue-filled region(frommesoderm)

    Digestive tract(from endoderm)

    Acoelomate. Acoelomates such asflatworms lack a body cavity betweenthe digestive tract and outer body wall.

    (c)

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    Protostome and Deuterostome Development Based on certain features seen in early

    development

    Many animals can be categorized as havingone of two developmental modes: protostomedevelopment or deuterostome development

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    Cleavage (spiral versus radial) In protostome development

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

    In deuterostome development

    Cleavage is radial and indeterminate (radial = thecleavage is either parallel or perpendicular to the vertical axis of the embryo)

    Protostome development(examples: molluscs, annelids,

    arthropods)

    Deuterostome development(examples: echinoderms,

    chordates)

    Eight-cell stage Eight-cell stage

    Spiral and determinate Radial and indeterminate

    (a) Cleavage. In general, protostomedevelopment begins with spiral,determinate cleavage.Deuterostome development ischaracterized by radial, indeterminatecleavage.

    Cl (d d )

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    Cleavage (determinant versus indeterminate) Determinate: Ultimate fate of each cell is

    determined (locked-in) early.

    Indeterminate: Each early cell retains thepotential to develop into a complete embryo!(this type of cleavage is what makes human identical twins possible)

    Protostome development(examples: molluscs, annelids,

    arthropods)

    Deuterostome development(examples: echinoderms,

    chordates)

    Eight-cell stage Eight-cell stage

    Spiral and determinate Radial and indeterminate

    (a) Cleavage. In general, protostomedevelopment begins with spiral,determinate cleavage.

    Deuterostome development ischaracterized by radial, indeterminatecleavage.

    C l F i

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    Coelom Formation

    In protostome development

    The splitting of the initially solid masses ofmesoderm to form the coelomic cavity is calledschizocoelous development

    In deuterostome development

    Formation of the body cavity is described asenterocoelous development

    Archenteron

    Blastopore MesodermCoelom

    BlastoporeMesoderm

    Schizocoelous: solidmasses of mesodermsplit and form coelom

    Enterocoelous:folds of archenteronform coelom

    Coelom (b) Coelom formation. Coelomformation begins in the gastrulastage. In protostome development,

    the coelom forms from splits in themesoderm (schizocoelousdevelopment). In deuterostomedevelopment, the coelom forms frommesodermal outpocketings of thearchenteron (enterocoelousdevelopment).

    F f h Bl

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    Fate of the Blastopore

    In protostome development

    The blastopore becomes the mouth

    In deuterostome development

    The blastopore becomes the anus

    Anus

    Anus

    Mouth

    Mouth

    Mouth developsfrom blastopore

    Anus developsfrom blastopore

    Digestive tube

    A i l h l ti t

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    Animal phylogenetic tree

    Zoologists currently recognize about 35 animalphyla

    The current debate in animal systematics

    Has led to the development of two

    phylogenetic hypotheses, but others exist aswell

    A i l h l ti t

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    One hypothesis of animal phylogeny basedmainly on morphological and developmentalcomparisons

    Animal phylogenetic tree

    Porifera

    Cnidaria

    Ctenophora

    Phoronida

    Ectoprocta

    Brachiopoda

    Echinodermata

    Chordata

    Platyhelminthes

    Mollusca

    Annelida

    Arthropoda

    Rotifera

    Nemertea

    Nematoda

    Radiata Deuterostomia Protostomia

    Bilateria

    Eumetazoa

    Metazoa

    Ancestral colonialflagellate

    A i l h l ti t

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    Animal phylogenetic tree

    One hypothesis of animal phylogeny basedmainly on molecular data

    Calcarea

    Silicarea

    Ctenophora

    Cnidaria

    Echinodermata

    Chordata

    Brachiopoda

    Phoronida

    Ectoprocta

    Platyhelminthes

    Nemertea

    Mollusca

    Annelida

    Rotifera

    Nematoda

    Arthropoda

    Radiata

    Porifera Deuterostomia Lophotrochozoa Ecdysozoa

    Bilateria

    Eumetazoa

    Metazoa

    Ancestral colonial

    flagellate

    P i t f A t

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    Points of Agreement All animals share a common ancestor

    Sponges are basal animals

    Eumetazoa is a clade of animals with truetissues

    Most animal phyla belong to the clade Bilateria

    Vertebrates and some other phyla belong to

    the clade Deuterostomia

    Di t th Bil t i

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    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 theprotostomes rather than one: the ecdysozoansand the lophotrochozoans

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    Ecdysozoans share a common characteristic

    They shed their exoskeletons through aprocess called ecdysis

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    Lophotrochozoans

    Have a lophophore, a ciliated feeding structure

    OR

    Go through a distinct larval stage called a

    trochophore larva Apical tuftof cilia

    Mouth

    Anus

    (a) An ectoproct, a lophophorate (b) Structure of trochophore larva