Sponges, Mollusks, and Worms, Oh My!

Animals

General Features of Animals

• animals share many important characteristics, such as they
 are heterotrophs
 are multicellular and lack cell walls
 can move from place to place
 have diverse forms and habitats
 reproduce, mostly, by sexual reproduction
 have a common pattern of development
 unique tissues
The Animal Family Tree
• the multicellular animals 35 very different phyla
 to judge which phyla are more closely related, taxonomists compare anatomical
features and aspects of embryological development
 the end result are phlyogenies, which are basically like family trees
 the main branches of the phylogenies make possible the evolutionary history of
animals

• the kingdom Animalia is traditionally divided into two main branches based on tissue
presence

 Parazoa possess neither tissues nor organs and have no discernible symmetry
• they are represented mostly by the phylum Porifera, the sponges

 Eumetazoa have a definite shape and symmetry and, in most cases, tissues
organized into organs and organ systems

• although very different, the Parazoa and Eumetazoa are thought to have evolved from
a common ancestor

 the shared ancestor was probably a choanoflagellate

• the choanoflagellate lived over 700 million years ago and was a colonial,
flagellated protist

• within the Eumetazoan phylogeny, the family tree branches on the basis of the type of
embryological layering

 Radiata have two embryological layers, an outer ectoderm and an inner

endoderm
• this body plan is called diploblastic
  Bilateria have a third embryological layer, the mesoderm, that occurs between the
ectoderm and the endoderm
• this body plan is called triploblastic

• additional branches to the phylogenetic tree were assigned by identifying traits that
were important to the evolutionary history of phyla
 for example, the presence or absence of a body cavity
 the traditional phylogeny of taxonomists relies on the either-or-nature of categories
Figure 25.1 The animal family tree: the traditional viewpoint.

• the traditional animal phylogeny is being revised because some of the important
characters may not be conserved to the extent previously thought
 molecular systematics offers a means to construct phylogenic trees using clusters
of genes as means to detect relatedness
 this new approach has resulted into significant refinements of the traditional
phylogeny
• for example, the protostomes have a more complex evolutionary history

Figure 25.2 The animal family tree: A new look.

Five Key Transitions in Body Plan

• the evolution of animals is marked by five key transitions in body plan

1. evolution of tissues
2. bilateral symmetry
3. body cavity
4. deuterostome development
5. segmentation

• the presence of tissues is the first key transition in the animal body plan
 only the Parazoa, the sponges, lack defined tissues and organs
• these animals exist as aggregates of cells with minimal intercellular
coordination
 all other animals besides members of the Parazoa possess tissues
• they belong to the Eumetazoa

• virtually all animals other than sponges have a definite shape and symmetry
 radial symmetry is a body plan in which all parts of the body are arranged along a
central axis
• if a plane passing through the central axis divides the organism in halves, the
halves will be mirror images
  bilateral symmetry is body plan with distinct right and left halves that are mirror
images
• the plan allows for specialization among body regions

• the evolution of a body cavity was an important step in animal evolution

 this internal space allowed for the support of organs, distribution of materials, and
coordination of development
 for example, the digestive tract can be larger and longer

• bilateral animals can be divided into two groups based on differences in the basic
pattern of development

 protostomes include the flatworms, nematodes, mollusks, annelids, and arthropods

 deuterostomes include the echinoderms and the chordates

 deuterostomes evolved from protostomes more than 630 million years ago

• the subdivision of the body into segments is a key transition to the animal body plan
that occurs early on during development

• in highly segmented animals, each segment can develop a more or less complete set of
adult organ systems

• each segment can function as a separate locomotory unit

Figure 25.3 Evolutionary trends among the animals.

Sponges: Animals Without Tissues

• sponges, members of the phylum Porifera

 their bodies a little more than masses of specialized cells embedded in a gel-like
matrix

 clumps of cells disassociated from a sponge can give rise to new sponges

 the body of a sponge is perforated by many pores

• choanocytes are flagellated cells that line the body cavity of the sponge and
draw in water through the pores

 the sponge is a filter feeder which traps any food particles

Figure 25.4 Diversity in sponges.

Figure 25.5 Phylum Porifera: sponges.

 Cnidarians: Tissues Lead to Greater Specialization
• the Radiata include two phyla
 Cnidaria comprises the hydra, jellyfish, corals and anemones
 Ctenophora comprises the comb jellies

• the members of the Radiata have a body plan that allows them to interact with their
environment on all sides

• a major evolutionary advance in the Radiata is extracellular digestion of food

 digestion begins outside the body in a gut cavity called, the gastrovascular cavity
 this form of digestion allows animals to digest an animal larger than itself

Figure 25.6 Representative cnidarians.

• cnidarians (phylum Cnidaria) are carnivores that capture prey with tentacles that ring
their mouths
 these tentacles and, sometimes, the body surface, bear stinging cells called
cnidocytes
 within each cnidocyte is a harpoon-like barb, called a nematocyst, which
cnidarians use to spear their prey and they retract towards the tentacle
 the nematocyst can discharge so explosively that it is capable of piercing the hard
shell of a crab

Figure 25.7 Phylum Cnidaria: cnidarians.

• cnidarians have two basic body forms

 medusae are the floating form
 polyps are the sessile form

• medusae are often called “jellyfish,” because of their gelatinous interior, or “stinging
nettles,” because of their nematocyts

• polyps are pipe-shaped animals that usually attach to rock

 in corals, the polyps secrete a deposit of calcium carbonate in which they live
Figure 25.9 The life cycle of Obelia, a marine colonial hydroid.

Solid Worms: Bilateral Symmetry

• body symmetry differs among the Eumetazoa

 radial symmetry means that multiple planes cutting the organism in half will
produce mirror images

 bilateral symmetry means that only one plane can cut the organism in half to
produce mirror images
 Figure 25.10 How radial and bilateral symmetry differ.

• most bilaterally symmetrical animals have evolved a definitive head end

 this process is termed cephalization

• the bilaterally symmetrical eumetazoans produce three embryonic layers

 ectoderm will develop into the outer coverings of the body and the nervous system
 mesoderm will develop into the skeleton and muscles
 endoderm will develop into the digestive organs and intestine

• the solid worms are the simplest of all bilaterally symmetrical animals
 the largest phylum of these worms is the Phylum Platyhelminthes, which includes
the flatworms
• flatworms lack any internal cavity other than the digestive tract
– this solid condition is called acoelomate
• they have separate organs, including a uterus and testes

Figure 25.11 Body plan of a solid worm.

Figure 25.12 Flatworms.

• most flatworms are parasitic but some are free-living

 flatworms range in size from less than a millimeter to many meters long

• there are two classes of parasitic flatworms

 flukes
 tapeworms

• the parasitic lifestyle has resulted in the eventual loss of features not used or needed by
the parasite

 for example, the parasites lack cilia in the adult stage and do not need eye spots

 this loss of features that lack adaptive purpose for parasitism is sometimes called
degenerative evolution

Figure 25.13 Life cycle of the human liver fluke, Clonorchis sinensis.

• tapeworms are a classic example of degenerative evolution

 the body of a tapeworm has been reduced to two primary functions

• eating
 • reproduction

Figure 25.14 Phylum Platyhelminthes: solid worms.

 if flatworms have a digestive cavity, then it is incomplete

• the gut branches throughout the body and is involved in both digestion and
excretion
• they are capable of performing some extracellular digestion

 the parasitic flatworms lack a gut entirely and absorb food directly through their
body walls

• flatworms have an excretory system

 the system consists of a network of fine tubules that run through the body
 enlarged flame cells (cilia-lined bulbs) are located on the side branches of the
tubules
 the cilia move water and excretory substances into the tubules and then into exit
pores
 the primary function of the flame cells is in the regulation of water balance
 most of the metabolic wastes are excreted through the gut

Figure 25.15 Diagram of flatworm anatomy.

• flatworms lack a circulatory system and all cells must be within diffusion distance of
oxygen and food

• flatworms have a simple nervous system

 they use sensory pits or tentacles along the sides of the head to detect food,
chemical, and movement
 free-living forms have eyespots to distinguish light from dark

• reproduction in flatworms is complex

 most flatworms are hermaphroditic, meaning that each individual contains both
male and female reproductive structures
 some flatworms have a complex succession of distinct larval stages
 some flatworms are capable of asexual regeneration
Roundworms: The Evolution of a Body Cavity
• a key transition in the evolution of the animal body plan was the evolution of the body
cavity

• the evolution of an internal body cavity helped improve the animal body design in
three areas
  circulation
 movement
 organ function

• there are three basic kinds of body plans found in bilaterally symmetrical animals
 acoelomates have no body cavity

 pseudocoelomates have a body cavity located between the mesoderm and the
endoderm

 coelomates have a body cavity (called a coelom) that develops entirely within the
mesoderm

Figure 25.16 Three body plans for bilaterally symmetrical animals.

• seven phyla of bilaterally symmetrical animals have a pseudocoelom

 the pseudocoelom serves as a hydrostatic skeleton, a skeleton that gains its
rigidity from fluids kept under pressure
 all pseudocoelomates lack a circulatory system
 most pseudocoelomates have a complete digestive tract

• the phylum Nematoda contains the greatest number of species among the phyla that
are pseudocoelomates
 the members of this phylum include nematodes, eelworms, and other roundworms
 they are unsegmented, cylindrical worms covered by a flexible cuticle that is
molted as they grow
 nematodes move in a whip-like fashion
Figure 25.17 Pseudocoelomates. (a) Nematodes (phylum Nematoda)

• the mouth of a nematode is often equipped with piercing organs called stylets
 food passes through a muscular chamber called the pharynx

• reproduction in nematodes is usually sexual with, usually, separate sexes

 their development is simple and adults have very few cells
 Caenorhabditis elegans is a roundworm important to genetic and developmental
studies

Figure 25.18 Phylum Nematoda: roundworms.

• some nematodes are parasitic in humans, cats, dogs, and animals of economic
importance
 heartworm in dogs is caused by a nematode
  trichinosis is an infection caused by the nematode Trichinella and transmitted to
humans who eat undercooked pork
 intestinal roundworms, Ascaris lumbricoides, live in human intestines

• another phylum consisting of animals with a pseudocoelomate body plan is the

phylum Rotifera
 rotifers are small, aquatic organisms that have a crown of cilia at their heads
 the cilia help in both locomotion and feeding
Figure 25.17 Pseudocoelomates.

(b) Rotifers (phylum Rotifera)

Mollusks: Coelomates
• coelomate animals are more successful than pseudocoelomates because of the nature
of embryonic development
 primary induction is a process in animal development in which one of the three
primary embryonic tissues interacts with another
 the interaction requires physical contact
 in coelomates, contact is made possible between mesoderm and endoderm
• this interaction permits localized portions of the digestive tract to become highly
specialized

• the mollusks, members of the phylum Mollusca, are the only coelomates without
segmented bodies

• the basic body of a mollusk is comprised of three regions

 a head-foot
 a visceral mass containing the body’s organs
 a mantle that envelopes the visceral mass and is associated with the gills

• there three major groups of mollusks

 gastropods—include the snails and slugs

 bivalves—include clams, oysters, and scallops

 cephalopods—include the octopi and squids

Figure 25.19 Three major groups of mollusks.

• mollusks have a unique feeding structure, called a radula

 the radula is a rasping tongue-like organ that bears rows of pointed, backward-
curving teeth
• in most mollusks, the outer surface of the mantle secretes a protective shell
 the shell has multiple layers comprised of protein, calcium, and pearl
Figure 25.20 Phylum Mollusca: mollusks.

Annelids: The Rise of Segmentation

• one of the early innovations to body plan to arise among the coelomates was
segmentation
 segmentation is the building of a body from a series of similar segments
• this body plan offers a lot of flexibility in that small changes to segments can
produce a new kind of segment with different functions
 the first segmented animals to evolve were the annelid worms, phylum Annelida

• the basic body plan of an annelid is a tube within a tube

 the digestive tract is suspended within the tube of the coelom
 the tubes run from mouth to anus

• derived from this basic organization are three characteristics

 repeated segments
 specialized segments
 connections

Figure 25.22 Phylum Annelida: annelids.

Arthropods: Advent of Jointed Appendages

• the most successful of all animal groups is the phylum Arthropoda, comprising the
arthropods
 these animals have jointed appendages
 in addition to joints, arthropods have an exoskeleton made of chitin
• the muscles of arthropods attach to the interior of this outer shell
• the shell offers protection against predators and water loss

• chitin cannot support much weight

 arthropod size is limited as a result

• arthropod bodies are segmented like annelids

 segments often fuse into functional groups in the adult stage
Figure 25.24 Segmentation in insects.
Figure 25.25 Phylum Arthropoda: arthropods.

• chelicerates are arthropods that lack jaws

 they include spiders, mites and scorpions
• mandibulates are arthropods with jaws, called mandibles
 they include the crustaceans, insects, centipedes and millipedes
Figure 25.26 Chelicerates and mandibulates.

• the chelicerate fossil record goes back 630 million years

 a surviving type of chelicerate from this period is the horseshoe crab

• the class Arachnida has 57K named species and includes the spiders, ticks, mites,
scorpions, and daddy longlegs
 arachnids have a pair of chelicerae, a pair of pedipalps, and four pairs of walking
legs
Figure 25.28 Arachnids.

• crustaceans belong to the phylum Crustacea and comprise a diverse group of

mandibulates
 there a 35K species of crustacea described including species of crabs, shrimps,
lobsters, crayfish, water fleas, pillbugs, and sowbugs
 most crustaceans have two pairs of antennae, three pairs of chewing appendages,
and various numbers of legs
 all crustaceans pass through a larval stage called the nauplius

Figure 25.29 Crustaceans.

Figure 25.30 Body of a lobster, Homarus americanus.

• millipedes and centipedes have bodies that consist of a head region followed by
numerous similar segments

 centipedes have one pair of legs per segment while millipedes have two

 centipedes are all carnivorous while millipedes are herbivorous

Figure 25.31 Centipedes and millipedes.

• insects belong to the Class Insecta and are the largest group of arthropods
 they are the most abundant eukaryotes on the earth

• insects have three body sections

 head
 thorax
 abdomen
 Figure 25.33 Insect diversity.

Protostomes and Deuterostomes

• there are two major kinds of coelomate animals representing two distinct evolutionary
lines
 protostomes
• the mouth develops from or near the blastopore
 deuterostomes
• the anus forms from or near the blastopore; the mouth forms on another part of
the blastula
Figure 25.34 Embryonic development in protostomes and deuterostomes.

• deuterostomes also differ from protostomes in three other fundamental ways

 the pattern of cleavage
• protostomes have spiral cleavage while deuterostomes have radial cleavage
 fating of cells
• it occurs later in deuterostome cleavage than in protostome cleavage
 origin of the coelom
Echinoderms: The First Deuterostomes
• echinoderms belong to the phylum Echinodermata
 echinoderm means “spiny skin” and refers to the calcium-rich ossicles that protude
just beneath the echinoderm’s skin
 they are entirely marine animals and include sea stars, sea urchins, sand dollars,
and sea cucumbers
 all are bilaterally symmetrical as larvae but become radially symmetrical as adults
Figure 25.35 Diversity in echinoderms.

• a key adaptation of echinoderms is the water vascular system

 this system is a fluid-filled and composed of a central ring canal around which five
radial canals extend out into the arms
 from each radial canal short side branches extend to form thousands of tiny, hollow
tube feet

• most echinoderms reproduce sexually but asexual regeneration is also common

Figure 25.36 Phylum Echinodermata: echinoderms.

Chordates: Improving the Skeleton

• chordates belong to the phylum Chordata and are deuterostome coelomates
  they exhibit a truly internal endoskeleton with muscles attached to an internal rod,
called a notochord
 this innovation opened the door to large body sizes not possible in earlier animal
forms

• the approximately 56K species of chordates share four principal features

 notochord
 nerve cord
 pharyngeal pouches
 postanal tail

• all chordates have all four of these characteristics at some time in their lives.

Figure 25.37 Phylum Chordata: chordates.

• not all chordates are vertebrates

 tunicates and lancelets

• vertebrate chordates differ from tunicates and lancelets in two important respects

 vertebrates have a backbone

• this replaces the role of the notochord

 vertebrates have a distinct and well-differentiated head

Figure 25.39 A mouse embryo.