Chapt 30

617
Animals: Part I
Chapter Concepts
30.1 Evolution and Classication of Animals
Animals are multicellular heterotrophs exhibiting
at least some mobility. 618
Animals are grouped according to level of
organization, symmetry, body plan, pattern of
embryonic development, and presence or
absence of segmentation. 619
30.2 Introducing the Invertebrates
Sponges are multicellular, with limited mobility
and no symmetry. 621
Cnidarians are radially symmetrical, with two
tissue layers. 622
Planarians are bilaterally symmetrical, with a
denite head region. 624
Roundworms have a pseudocoelom and the
tube-within-a-tube body plan. 626
30.3 Mollusks
Mollusks have a muscular foot (variously
modied), and a visceral mass enveloped by a
mantle. 628
30.4 Annelids
Annelids are segmented, with a well-developed,
true coelom. 631
30.5 Arthropods
Arthropods have jointed appendages and an
exoskeleton that must be periodically shed. 634
Sea anemones, such as these strawberry sea anemones,
Corynachtis californica, are called the owers of the sea, but
they are animals, not plants. The projections you see are tentacles,
which capture small prey and stuff it down their mouths.
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30.1 Evolution and Classication
of Animals M
Although there are many different types of animals, they
have characteristics in common. Animals
1. are heterotrophic and usually acquire food by ingestion
followed by digestion.
2. typically have the power of motion or locomotion by
means of muscle bers.
3. are multicellular, and most have specialized cells that
form tissues and organs.
4. have a life cycle in which the adult is typically diploid.
5. usually practice sexual reproduction and produce an
embryo that undergoes developmental stages.
There are approximately 34 animal phyla, but we will
consider only the 9 phyla listed in the classication table. All
9 phyla contain invertebrates, which are animals without
backbones. The phylum Chordata also contains vertebrates,
which are animals with backbones.
Animals evolved from unicellular protozoans. The ad-
vent of multicellularity, which allows specialization of cells
to occur, seems to have been a requirement for the evolution
of animals.
Sponges have the cellular level of organization, meaning
that they have no tissues. One of the main events during the
development of the rest of the animal groups is the
establishment of germ layers from which all other structures
are derived. Although a total of three germ layers is seen in
most animals, the cnidarians have only two germ layers
(ectoderm and endoderm) and the tissue level of organization.
Animals with three germ layersectoderm, mesoderm, and
endodermhave an organ level of organization.
618 Part 6 Evolution and Diversity 30-2
W
hile Oscar watches a spider close in on a dragon-
y trapped by its web, a darting buttery distracts
the young boy. He chases the winged creature
down the path to the ocean. At the water's edge, he notices
several sea sponges and shells that have washed up and
dried out in the hot sun. Oscar then spots his sister who has
swum out to a small coral reef to snorkel. She yelps as an
unseen jellysh stings her, causing her brother to laugh. "Os-
car! Time to set the table, the lobsters are boiling," shouts
out a motherly voice from the porch of the nearby beach
house.
Oscar's brief period of play, and his approaching dinner,
has brought him into contact with a large crowd of inverte-
brates, the amazingly diverse group of animals that includes
more than a million species of arthropods alone. This chap-
ter will introduce you to some of them, and explain how sci-
entists classify this rich zoo of creatures.
Kingdom Animalia
Multicellular organisms with well-developed tissues; usually
motile; heterotrophic by ingestion, generally in a digestive cavity;
diplontic life cycle.
Approximate
Some Number of
Phylum Representatives Described Species
Porifera Glass, chalk, 5,000
(sponges) bath sponges
Cnidarians Hydrozoans, jellyshes,
(cnidarians) sea anemones, corals 9,000
Platyhelminthes Planarians, ukes,
(atworms) tapeworms 13,000
Nematoda Pinworms,
(roundworms) hook worms 500,000
Mollusca Snails, clams, squids,
(mollusks) octopuses 110,000
Annelida Clam worms,
(annelids) earthworms,leeches 12,000
Arthropoda Craysh, insects,
(arthropods) millipedes, spiders One million plus
Echinodermata Starsh, sea urchins,
(echinoderms) sand dollars, sea 6,000
cucumbers
Chordata (chordates)
Cephalochordata Lancelet 23
(cephalochordates)
Urochordata Tunicates 1,250
(urochordates)
Vertebrata (vertebrates)
Agnatha Lampreys, hagshes 63
(jawless shes)
Chondrichthyes Sharks, skates, rays 850
(cartilaginous shes)
Osteichthyes Herring, salmon, 20,000
(bony shes) cod, eel
Amphibia Frogs, toads, 3,900
(amphibians) salamanders
Reptilia Snakes, lizards, 6,000
(reptiles) turtles
Aves Sparrows, 9,000
(birds) penguins,
ostriches
Mammalia Cats, dogs, 4,500
(mammals) horses, rats,
humans
Primates Prosimians, monkeys, apes
Anthropoidea Monkeys, apes, humans
Hominidae Apes, humans
Homo Humans
Sponges are asymmetrical. Asymmetry means that the
animal has no particular symmetry. Other animals are either
radially symmetrical or bilaterally symmetrical. Radial
symmetry means that the animal is organized circularly,
and, just as with a wheel, two identical halves are obtained
no matter how the animal is sliced longitudinally. Bilateral
symmetry means that the animal has denite right and left
halves; only one longitudinal cut down the center of the an-
imal will produce two equal halves.
Radially symmetrical animals are sometimes attached to
a substrate; that is, they are sessile. This type of symmetry is
useful to these animals since it allows them to reach out in
all directions from one center. Bilaterally symmetrical ani-
mals tend to be active and to move forward with an anterior
end. During the evolution of animals, bilateral symmetry is
accompanied by cephalization, localization of a brain and
specialized sensory organs at the anterior end of an animal.
Two body plans are observed in the animal kingdom:
the sac plan and the tube-within-a-tube plan. Animals with the
sac plan have an incomplete digestive system. It has only
one opening, which is used both as an entrance for food and
an exit for undigested material. Animals with the
tube-within-a-tube plan have a complete digestive system,
with a separate entrance for food and exit for undigested
material. Having two openings allows specialization of
parts to occur along the length of the tube.
A true coelom is an internal body cavity completely
lined by mesoderm, where internal organs are found. Flat-
worms are acoelomatesthey have mesoderm but no body
cavity. The roundworms have a pseudocoelom, a body cav-
ity incompletely lined by mesoderm because it develops be-
tween the mesoderm and endoderm. There is a layer of
mesoderm beneath the body wall but not around the gut.
The rest of the phyla in the classication table are true coelo-
matesthey have a coelom that is completely lined with
mesoderm. Coelomates are either protostomes or deutero-
stomes. When the rst embryonic opening becomes the
mouth, the animal is a protostome. When the second open-
ing becomes the mouth, the animal is a deuterostome.
Among coelomates, mollusks and echinoderms are non-
segmented, while annelids, arthropods, and chordates are
segmented. Segmentation, which is the repetition of body
parts along the length of the body, leads to specialization of
parts because the various segments can become differenti-
ated for specic purposes.
Classication of animals is based on type of level
of organization, symmetry, body plan, type of
coelom, and presence of segmentation.
Chapter 30 Animals: Part I 619 30-3
Figure 30.1 Animal diversity.
How do hydras differ from craysh and humans? They are all multicellular heterotrophic organisms but differ according to their level of
organization (hydras have only tissues but crayshes and humans have organ systems); symmetry (hydras are radially symmetrical but craysh
and humans are bilaterally symmetrical with cephalization); and body plan (hydras have a sac body plan while craysh and humans have a tube-
within-a-tube body plan).
a. Hydra, Hydra b. Green craysh, Barbi c. Human being, Homo
Figure 30.2 Evolutionary tree.
All animals are believed to be descended from protists; the Porifera (sponges) with the cellular level of organization may have evolved separately.
multicellularity
Ancestral protists
radial symmetry
diploblastic
(two germ layers)
acoelomates
(no coelom)
pseudocoelomates
(body cavity not completely
lined by mesoderm)
coelomates
(body cavity completely
lined by mesoderm)
protostomes
(first embryonic
opening is mouth)
bilateral symmetry
triploblastic
(three germ layers)
deuterostomes
(second embryonic
opening is mouth)
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Sponges are asymmetricalhave no denite symmetry
while cnidarians are radially symmetrical. Flatworms and
roundworms are bilaterally symmetrical, but differ in that
atworms are acoelomates while roundworms are pseudo-
coelomates.
Sponges
Sponges (phylum Porifera,
about 5,000 species) are
aquatic, largely marine ani-
mals, that vary greatly in
size, shape, and color. Their
saclike bodies are perforated
by many pores; the phylum
name, Porifera, means pore
bearing. Sponges are multi-
cellular with the cellular level
of organization. As such, they
are believed to be out of the
mainstream of animal evolution. Most likely, they evolved
separately from protozoan ancestors and represent a dead-
end branch of the evolutionary tree.
The outer layer of the wall contains attened epidermal
cells, some of which have contractile fibers; the middle
layer is a semiuid matrix with wandering amoeboid cells,
and the inner layer is composed of agellated cells called
collar cells (or choanocytes) (Fig. 30.3). The beating of the
flagella produces water currents that flow through the
pores into the central cavity and out through the osculum,
the upper opening of the body. Even a sim-
ple sponge only 10 cm tall is estimated to
lter as much as 100 liters of water each day.
It takes this much water to supply the needs
of the sponge. A sponge is a sessile filter
feeder, an organism that lters its food from
the water by means of a straining devicein
this case, the pores of the walls and the mi-
crovilli making up the collar of collar cells.
Microscopic food particles that pass between
the microvilli are engulfed by the collar cells
and digested by them in food vacuoles, or
are passed to the amoeboid cells for digestion. The amoe-
boid cells also act as a circulatory device to transport nutri-
ents from cell to cell, and they produce the sex cells (the egg
and the sperm) and spicules.
Sponges can reproduce asexually by fragmentation or
by budding. During budding, a small protuberance appears
and gradually increases in size until a complete organism
forms. Budding produces colonies of sponges that can
become quite large. During sexual reproduction, eggs and
sperm are released into the central cavity and the zygote de-
velops into a agellated larva that may swim to a new loca-
tion. If the cells of a sponge are mechanically separated, they
will reassemble into a complete and functioning organism!
Like all less specialized organisms, sponges are also capable
of regeneration, or growth of a whole from a small part.
Sponges are classied on the basis of their skeleton.
Some sponges have an internal skeleton composed of
spicules, small needle-shaped structures with one to six rays.
Chalk sponges have spicules made of calcium carbonate;
glass sponges have spicules that contain silica. Most
sponges have bers of spongin, a modied form of collagen.
But some sponges contain only spongin bers; a bath
sponge is the dried spongin skeleton from which all living
tissue has been removed. Today, however, commercial
sponges are usually synthetic.
Sponges have a cellular level of organization and
most likely evolved independently from protozoa.
They are the only animals in which digestion
occurs within cells.
collar cell
(choanocyte)
flagellum
collar
amoeboid cell
epidermal cell
spicule
sponge
wall
central
cavity
pore
Sponge Organization
central
cavity
water in
through
pores
water out
osculum
Figure 30.3 Sponge.
In a sponge, the wall contains two layers of cells:
the outer epidermal cells and the inner collar cells.
The collar cells (enlarged) have agella that beat,
moving the water through pores as indicated by
the arrows. Food particles in the water are trapped
by the collar cells and digested within their food
vacuoles. Amoeboid cells transport nutrients from
cell to cell; spicules form an internal skeleton in
some sponges.
multicellularity
Ancestral protists
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Cnidarians
Cnidarians (phylum Cnidaria,
about 9,000 species) are tubu-
lar or bell-shaped animals
that reside mainly in shallow
coastal waters, except for the
oceanic jellyshes. During
development, cnidarians have
only two germ layers (ecto-
derm and endoderm), and as
adults they have the tissue
level of organization. Cnidari-
ans are radially symmetrical,
meaning that any longitu-
dinal cut produces two iden-
tical halves.
fertilization
planula
polyp
zygote
sperm
medusa
egg
mesoglea
gastrovascular cavity
tentacle
mouth
mesoglea
a.
b. Sea anemone, Apitasia c. Coral, Tubastrea d. Portuguese man-of-war,
Physalia
e. Jellyfish, Aurelia
Figure 30.4 Cnidarian diversity.
a. The life cycle of a cnidarian; in some cnidarians there is both a polyp and a medusa stage; in others, one form is dominant, and in still others,
one form is absent altogether. b. The anemone, which is sometimes called the ower of the sea, is a solitary polyp. c. Corals are colonial polyps
residing in a calcium carbonate or proteinaceous skeleton. d. Portuguese man-of-war is a colony of modied polyps and medusae. e. True
jellysh undergo a complete life cycle; the photo shows the medusan stage.
Unique to cnidarians are specialized stinging cells,
called cnidocytes, which give the phylum its name. Each
cnidocyte has a capsule called a nematocyst, which contains
a long, spirally coiled hollow thread. When the trigger of the
cnidocyte is touched, the nematocyst is discharged. Some
threads merely trap a prey or predator; others have spines
that penetrate and inject paralyzing toxins.
Two basic body forms are seen among cnidarians. The
mouth of a polyp is directed upward from the substrate,
while the mouth of a jellysh or medusa is directed down-
ward. A medusa has more mesoglea than a polyp, and the
tentacles are concentrated on the margin of the bell. At one
time, both body forms may have been a part of the life cycle
of all cnidarians (Fig. 30.4a). When both are present, the ses-
sile polyp stage produces medusae and the motile medusan
stage produces egg and sperm; the zygote develops into a
ciliated larva that is capable of dispersal. In some cnidarians,
one stage is dominant and the other is reduced; in other
species one form is absent altogether.
Ancestral protists
radial symmetry
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cnidocyte
before discharge
cnidocyte
after discharge
nematocyst
barb
spines
lid
trigger
coiled
thread
nucleus
mouth
gastrovascular
cavity
gastrodermis
mesoglea
nerve net
epidermis
circular
muscle fibers
interstitial
cell
sensory
cell
cnidocyte
longitudinal
muscle fibers
epitheliomuscular cell
epidermis
gastrodermis
gastrovascular
cavity
nutritive -
muscular cell
cross section of Hydra
gland cell
mesoglea
tentacle
Cnidarians are quite diverse (Fig. 30.4be). Sea
anemones are solitary polyps, often very large and with
thick walls. They can be brightly colored, resembling beauti-
ful owers. Corals are similar to sea anemones, but they have
calcium carbonate skeletons. Some corals are solitary, but
most are colonial, with either at and rounded or upright
and branched colonies. The slow accumulation of coral skele-
tons forms coral reefs, which are areas of biological abun-
dance in warmer waters. The Portuguese man-of-war is a
colony of polyp and medusa types of individuals. One polyp
becomes a gas-lled oat and the other polyps are special-
ized for feeding. The medusae are specialized for reproduc-
tion. In jellyshes, the medusa is the primary stage of the life
cycle, and the polyp remains quite small and inconspicuous.
Hydra
Hydra is a freshwater cnidarian (Fig. 30.5); it is likely to be
found attached to underwater plants or rocks in most lakes
and ponds. The body is a small tubular polyp about 7.5 mm
in length. Like all cnidarians, a hydra has the sac body plan;
that is, there is only one opening that serves as both a mouth
and anus. The outer tissue layer is a protective epidermis de-
rived from ectoderm. The inner tissue layer, derived from
endoderm, is called a gastrodermis. The two tissue layers
are separated by a jellylike packing material called mesoglea.
There are both circular and longitudinal muscle bers.
Nerve cells located below the epidermis near the mesoglea
interconnect and form a nerve net that communicates with
sensory cells throughout the body. The nerve net allows
transmission of impulses in several directions at once. Hav-
ing both muscle bers and nerve bers, cnidarians are capa-
ble of directional movement; the body can contract or
extend, and the tentacles that ring the mouth can reach out
and grasp prey.
Digestion begins within the central cavity but is com-
pleted within the food vacuoles of gastrodermal cells. Nutri-
ent molecules are passed by diffusion to the other cells of the
body. The large central cavity allows gastrodermal cells to ex-
change gases directly with a watery medium. Because the cen-
tral cavity carries on digestion and acts as a circulatory system
by distributing food and gases, it is called a gastrovascular
cavity. All cnidarians have a gastrovascular cavity.
Although hydras exist only as polyps, and there are no
medusae, still they can reproduce sexually or asexually.
When sexual reproduction is going to occur, an ovary or a
testis develops in the body wall. Like sponges, cnidarians
can regenerate from a small piece. When conditions are fa-
vorable, hydras produce small outgrowths, or buds, that
pinch off and begin to live independently.
Figure 30.5 Hydrozoan, hydra.
The wall of a hydra, which lines a gastrovascular cavity, has two tissue layers. Special stinging cells (cnidocytes) contain nematocysts that assist
in capturing prey. The tentacles deliver the prey to the mouth.
Flatworms
Flatworms (phylum Platyhel-
minthes, about 13,000 species)
also have a sac body plan. Flat-
worms, however, have three
germ layers. The presence of
mesoderm in addition to
ectoderm and endoderm
gives bulk to the animal and
leads to greater complexity.
Free-living atworms have
muscles and excretory, repro-
ductive, and digestive organs.
The worms lack respiratory and circulatory organs
because the body is at and thin, diffusion alone is ade-
quate for the passage of oxygen and other substances from
cell to cell.
Planarians
Freshwater planarians (Fig. 30.6) are small (several mm to
several cm), literally at worms. Some tend to be colorless;
others have brown or black pigmentation. Planarians live in
lakes, ponds, streams, and springs, where they feed on small
living or dead organisms, such as worms and crustaceans.
Planarians live in fresh water and have an excretory or-
gan that serves primarily to rid the body of excess water. A
network of interconnecting canals extends through much of
the body. The beating of cilia in the ame cells (so named be-
cause the beating of the cilia reminded some early investiga-
tor of the ickering of a ame) keeps the water moving
toward the excretory pores.
Planarians have a ladder-type nervous organ. Asmall ante-
rior brain and two lateral nerve cords are joined by cross-
branches. Planarians exhibit cephalizationaside from a
brain, there are light-sensitive organs (the eyespots) and
Figure 30.6 Planarian.
a. The micrograph of Dugesia shows that this atworm is bilaterally symmetrical and has a head region with eyespots. b. When the pharynx is
extended as shown, food is sucked up into a gastrovascular cavity that branches throughout the body. c. The excretory system with ame cells is
shown in detail. d. The reproductive system has both male and female organs, and the digestive system has a single opening. e. The nervous
system has a ladderlike appearance.
Ancestral protists
acoelomates
bilateral symmetry
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chemosensitive organs located on the auricles. They have
well-developed muscles. There are three kinds of muscle
layersan outer circular layer, an inner longitudinal layer,
and a diagonal layerthat allow for quite varied move-
ment. Their ciliated epidermis allows planarians to glide
along a lm of mucus.
The animal captures food by wrapping itself around the
prey, entangling it in slime, and pinning it down. Then a
muscular pharynx is extended, and by a sucking motion the
food is torn up and swallowed. The pharynx leads into a
three-branched gastrovascular cavity in which digestion is
both extracellular and intracellular. The digestive tract is in-
complete because it has only one opening.
Planarians are hermaphrodites; they possess both male
and female sex organs. The worms practice cross-fertilization:
the penis of one is inserted into the genital pore of the other,
and there is a reciprocal transfer of sperm. The fertilized eggs
hatch in 23 weeks as tiny worms. Like sponges and cnidari-
ans, planarians can regenerate. If a worm is cut crosswise,
each piece grows a new head or a new tail, as appropriate.
Parasitic Flatworms
Flukes and tapeworms are two classes of parasitic flat-
worms. The anterior end of these animals carries suckers
and sometimes hooks for attachment to the host. The par-
asite absorbs nutrients from the digestive tract of the host,
and in tapeworms the digestive system is essentially ab-
sent. The tegument, a specialized body wall resistant to
host digestive juices, is covered by the glycocalyx, a mu-
copolysaccharide coating. The extensive development of
the reproductive system, with the production of millions
of eggs, may be associated with difculties in dispersing
offspring. Both parasites utilize a secondary host to trans-
port an intermediate stage from primary host to primary
host. The primary host is infected with the sexually ma-
ture adult; the secondary host contains the larval stage or
stages.
Both ukes and tapeworms cause serious illnesses in
humans. The uke body tends to be oval to elongate. At the
anterior end surrounded by sensory papilla there is an oral
sucker and at least one other sucker for attachment to the
host. Different uke species infect the digestive tract, the bile
duct, blood, and the lungs. Schistosomes are blood ukes
that enter the body by active penetration of the skin (Fig.
30.7). Schistosomiasis, a serious infection caused by blood
ukes, is seen predominantly in the Middle East, Asia,
Africa, and South America. About 200 million people are
infected worldwide.
adult worms
live and copulate
in blood vessels
of human gut;
eggs migrate into
digestive tract
eggs
passed
in feces
ciliated larvae
(miracidia) hatch
in water and
enter snail
mother sporocyst
encloses many
developing daughter
sporocysts
daughter sporocyst
encloses many
developing
larvae (cercariae)
larvae (cercariae)
break out of daughter
sporocysts, escape snail,
and enter water
Figure 30.7 Schistosomiasis.
This infection of humans, caused by blood ukes, Schistosoma, is an extremely prevalent disease in Egyptespecially since the building of the
Aswan High Dam. Standing water in irrigation ditches, combined with unsanitary practices, has created the conditions for widespread infection.
Atapeworm has an anterior region, called a scolex, con-
taining hooks and suckers for attachment to the intestinal
wall of the host. Behind the scolex, there is a long series of
proglottids, segments that contain a full set of both male and
female sex organs and little else. Mature proglottids, which
are nothing but bags of eggs, break off, and as they pass out
with the feces, the eggs are released. If feces-contaminated
food is fed to pigs or cattle, the larvae escape when the cov-
ering of the eggs is digested away. They burrow through the
intestinal wall and travel in the bloodstream to nally lodge
and encyst in muscle. Here a cyst means a small, hard-
walled structure that contains a larval worm. When humans
eat infected meat, the larvae break out of the cyst, attach
themselves to the intestinal wall, and grow to adulthood.
Then the cycle begins again.
Also interesting to humans is the fact that eas, which as
larvae have fed on the feces of an infected host, can transmit
some types of tapeworms between cats and/or dogs.
Roundworms
Roundworms (phylum Nematoda, 500,000 species) are non-
segmentedthey have a smooth outside body wall. These
worms, which are generally
colorless and less than 5 cm
in length, occur almost any-
wherein the sea, in fresh
water, and in the soilin
such numbers that thousands
of them can be found in a
small area.
Roundworms possess two
anatomical features not seen
before: a tube-within-a-tube
body plan and a body cavity.
With a tube-within-a-tube body
plan, the digestive tract is complete; there is both a mouth
and an anus. The body cavity is a pseudocoelom, or a body
cavity incompletely lined with mesoderm (Fig. 30.8b). The
uid-lled pseudocoelom provides space for the develop-
ment of organs, substitutes for a circulatory system by al-
lowing easy passage of molecules, and provides a type of
skeleton. Worms in general do not have an internal or exter-
nal skeleton, but they do have a hydrostatic skeleton, a
uid-lled interior that supports muscle contraction and en-
hances exibility.
ectoderm
endoderm
mesoderm
gut
gut
body wall
Acoelomate
flatworms
a. Pseudocoelomate
roundworms
b. True coelomate
mollusks, annelids, arthropods,
echinoderms, chordates
c.
ectoderm
ectoderm
true
coelom
gut
mesoderm
endoderm
mesoderm
mesentery
endoderm
pseudocoelom
body wall
body wall
Figure 30.8 Coelom structure and function.
a. Flatworms have no body cavity, and mesodermal tissue lls the interior space. b. Roundworms have a pseudocoelom, and the body cavity is
incompletely lined by mesodermal tissue. In animals that have no other skeleton, a uid-lled coelom acts as a hydrostatic skeleton. c. Humans
are true coelomates, the body cavity is completely lined by mesodermal tissue, and mesentery holds the internal organs in place.
Ancestral protists
pseudocoelomates
bilateral symmetry
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Ascaris
In the roundworm Ascaris lumbricoides (Fig. 30.9), females
tend to be larger (2035 cm in length) than males. Both sexes
move by means of a characteristic whiplike motion because
only longitudinal muscles and no circular muscles lie next to
the body wall.
The Ascaris life cycle usually begins when larvae within
a protective covering are swallowed. The worms then es-
cape from the covering and burrow through the hosts in-
testinal wall. Making their way through the organs of the
host, they move from the intestine to the liver, the heart, and
then the lungs. While in the lungs for about ten days, they
grow in size. The larvae then migrate up the windpipe to the
throat, where they are swallowed, allowing them to once
again reach the intestine. Once the worms are mature they
mate, and the female produces larva-containing eggs, which
pass out with the feces.
Other Roundworms
Trichinosis is a fairly serious infection caused by Trichinella
spiralis, a roundworm that rarely infects humans in the
United States. Humans contract the disease when they eat
rare pork containing encysted larvae. After maturation, the
female adult burrows into the wall of the hosts small intes-
tine and produces live offspring, which are carried by the
mouth
a. Ascaris b. Trichinella
cyst
20m
ventral
nerve
cord
excretory
pore
sperm
seminal
vesicle
cloaca
spicules that
aid in sperm
transfer
pharynx
brain
dorsal
nerve cord
cuticle sperm
duct
gut
lateral nerve cord
testis
pseudocoelom
muscle
layer
anus
bloodstream to the skeletal muscles, where they encyst (Fig.
30.9). The symptoms of trichinosis include muscular pain,
weakness, fever, and anemia.
Elephantiasis is caused by a roundworm called the lar-
ial worm, which utilizes the mosquito as a secondary host.
Because the adult worms reside in lymphatic vessels, uid
return is impeded, and the limbs of an infected human can
swell to an enormous size, even resembling those of an ele-
phant. When a mosquito bites an infected person, it trans-
ports larvae to a new host.
Other roundworm infections are more common in the
United States. Children frequently acquire a pinworm infec-
tion, and hookworm is seen in the southern states, as well as
worldwide. A hookworm infection can be very debilitating
because the worms attach to the intestinal wall and feed on
blood. Good hygiene, proper disposal of sewage, and cook-
ing meat thoroughly usually protect people from parasitic
roundworms.
Roundworms have bilateral symmetry, a tube-within-
a-tube body plan, three germ layers, the organ level
of organization, and a pseudocoelom. The rest of the
animal phyla have these features, except there is a
true coelom instead of a pseudocoelom.
Figure 30.9 Roundworm diversity.
a. Note that roundworms such as Ascaris lumbricoides have a pseudocoelom and a complete digestive tract with a mouth and an anus. Therefore,
roundworms have a tube-within-a-tube body plan. The sexes are separate; this is a male roundworm. b. The larvae of the roundworm Trichinella
spiralis encyst as larvae in skeletal muscle bers where they coil in a sheath formed from a muscle ber. This infection in humans is called trichinosis.
30.3 Mollusks
Mollusks, along with an-
nelids and arthropods, are
protostomes (Fig. 30.10). In
protostomes, the rst embry-
onic opening becomes the
mouth. Because the coelom
forms by splitting of the
mesoderm, protostomes are
also schizocoelomates. In cer-
tain (but not all) protostomes,
there is a trochophore larva
(top-shaped with a band of
cilia at the midsection).
Characteristics of Mollusks
Mollusks (phylum Mollusca, about 110,000 species) are a
very large and diversied group; however, they all have a
body composed of at least three distinct parts.
1. Visceral mass: the soft-bodied portion that contains
internal organs.
2. Foot: the strong, muscular portion used for locomotion.
3. Mantle: the membranous or sometimes muscular
covering that envelops but does not completely enclose
the visceral mass. The mantle cavity is the space
between the two folds of the mantle. The mantle may
secrete a shell.
In addition to these three parts, many mollusks have a head
region with eyes and other sense organs.
The division of the body into distinct areas may have
contributed to diversication of animals in this phyla. There
are many different types of mollusks adapted to various
ways of life (Fig. 30.11). Molluscan groups can be distin-
guished by a modication of the foot. In the gastropods
(meaning stomach-footed), including nudibranchs, conchs,
and snails, the foot is ventrally attened, and the animal
moves by muscle contractions that pass along the foot.
While nudibranchs, also called sea slugs, lack a shell, conchs
and snails have a coiled shell in which the visceral mass spi-
rals. Some types of snails are adapted to life on land. For ex-
ample, their mantle is richly supplied with blood vessels
and functions as a lung when air is moved in and out
through respiratory pores.
In cephalopods (meaning head-footed), including octo-
puses and squids, the foot has evolved into tentacles about
the head. Aside from the tentacles, which seize prey,
cephalopods have a powerful beak and a radula (toothy
tongue) to tear prey apart. Cephalization aids these animals
in recognizing prey and in escaping enemies. The eyes are
supercially similar to those of vertebratesthey have a lens
and a retina with photoreceptors. However, its construction
is so different from the vertebrate eye that we believe the
Figure 30.10 Protostomes versus deuterostomes.
In protostomes, the rst embryonic opening called the blastopore
becomes the mouth, the coelom forms by splitting of the mesoderm
(they are schizocoelomates), and the trochophore larva is typical. In
deuterostomes, the blastopore becomes the anus, the coelom forms
by outpocketing of the primitive gut (they are enterocoelomates), and
the dipleurula larva is found among some.
Protostomes Deuterostomes
mollusks
annelids
arthopods
echinoderms
chordates
blastopore mouth blastopore anus
primitive
gut
primitive
gut
anus
mouth
fate of blastopore
schizocoelom enterocoelom
coelom forms by splitting
of the mesoderm
coelom formation
mouth
coelom forms by out-
pocketing of primitive gut
mouth
anus
trochophore
larva
anus
larval form
dipleurula
larva
Ancestral protists
coelomates
protostomes
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so-called camera-type eye actually evolved twiceonce in
the mollusks and once in the vertebrates. In cephalopods, the
brain is formed from a fusion of ganglia, and nerves leaving
the brain supply various parts of the body. An especially
large pair of nerves controls the rapid contraction of the man-
tle, allowing these animals to move quickly by a jet propul-
sion of water. Rapid movement and the secretion of a brown
or black pigment from an ink gland help cephalopods to es-
cape their enemies. Octopuses have no shell, and squid have
only a remnant of one concealed beneath the skin.
In bivalves, such as clams, oysters, and scallops, the foot
is laterally compressed. They are called bivalves because
there are two parts to the shell. Notice in Table 30.1 that a
clam is adapted to a less active life and a squid is adapted to
a more active life.
Feature Clam Squid
Food Gathering Filter feeder Active predator
Skeleton Heavy shell for No external
protection skeleton
Circulation Open Closed
Cephalization None Marked
Locomotion Hatchet foot Jet propulsion
Nervous System 3 separate ganglia Brain and nerves
Table 30.1 Comparison of Clam to Squid
Figure 30.11 Molluscan diversity.
a. A chiton has a attened foot and a shell that consists of eight articulating valves. b. A chambered nautilus achieves buoyancy by regulating the
amount of air in the chambers of its shell. c. A scallop has sensory tentacles extended between the valves. d. A nudibranch (sea slug) lacks a
shell, gills, and a mantle cavity. Dorsal projections function in gas exchange.
a. Chiton, Tonicella b. Chambered nautilus, Nautilus
c. Scallop, Pecten d. Spanish shawl nudibranch, Flabellina
Bivalves
In a clam, such as the freshwater clam Anodonta, the shell, se-
creted by the mantle, is composed of protein and calcium
carbonate, with an inner layer of mother-of-pearl. If a foreign
body is placed between the mantle and the shell, pearls form
as concentric layers of shell are deposited about the particle.
The adductor muscles hold the valves of the shell to-
gether. Within the mantle cavity, the gills, an organ for gas
exchange in aquatic forms, hang down on either side of the
visceral mass, which lies above the foot. The heart of a clam
lies just below the hump of the shell within the pericardial
cavity, the only remains of the coelom. Therefore, the coelom
is reduced. The heart pumps blood into a dorsal aorta that
leads to the various organs of the body. Within the organs,
however, blood ows through spaces, or sinuses, rather than
through vessels. This is an open circulatory systembecause the
blood is not contained within blood vessels all the time. This
type of circulatory system can usually be associated with a
relatively inactive animal because it is an inefcient means
of transporting blood throughout the body.
The nervous system of a clam (Fig. 30.12) is composed of
three pairs of ganglia (anterior, foot, and posterior), which are
all connected by nerves. Clams lack cephalization. The foot
projects anteriorly from the shell, and by expanding the tip
of the foot and pulling the body after it, the clam moves
forward.
The clam is a lter feeder. Food particles and water enter
the mantle cavity by way of the incurrent siphon, a posterior
opening between the two valves. Mucous secretions cause
smaller particles to adhere to the gills, and ciliary action
sweeps them toward the mouth. This method of feeding
does not require rapid movement.
The digestive system of the clam includes a mouth with
labial palps, an esophagus, a stomach, and an intestine,
which coils about in the visceral mass and then is sur-
rounded by the heart as it extends to the anus. The anus emp-
ties at an excurrent siphon, which lies just above the incurrent
siphon. There is also an accessory organ of digestion called a
digestive gland. The two excretory kidneys in the clam (Fig.
30.12), which lie just below the heart, remove waste from the
pericardial cavity for excretion into the mantle cavity.
The sexes are usually separate. The gonad (e.g., ovary or
testis) is located around the coils of the intestine. While all
clams have some type of larval stage, only marine clams
have a trochophore larva. The presence of the trochophore
larva (see Fig. 30.10) among some mollusks indicates a rela-
tionship to the annelids, some members of which also have
this type of larval stage.
gonad
kidney
anterior aorta pericardial cavity
heart
posterior retractor muscle
posterior adductor muscle
posterior aorta
intestine
anus
excurrent siphon
incurrent siphon
gill
posterior ganglion
intestine
mantle
shell
mantle (cut)
intestine foot ganglion
foot
labial palpi
mouth
anterior ganglion
anterior adductor
muscle
esophagus
stomach
digestive gland
Figure 30.12 Clam.
The shell and the mantle have been removed from one side. Trace the path of food from the incurrent siphon past the gills, to the mouth, the
stomach, the intestine, the anus, and the excurrent siphon. Locate the three ganglia: anterior, foot, and posterior. The heart lies in the reduced
coelom.
View 1 View 2
30.4 Annelids
Annelids (phylum Annelida,
about 12,000 species) are seg-
mented, as is externally evi-
denced by the rings that
encircle the body. Internally,
partitions called septa (sing.,
septum) divide the well-
developed, uid-lled coe-
lom, which acts as a hydro-
static skeleton.
Segmentation is a subdivi-
sion of the body along its length into repeating units, called
segments. Annelids have a hydrostatic skeleton, and parti-
tioning of the coelom permits each body segment indepen-
dence of movement. Also, each segment of an earthworm has its
own set of longitudinal and circular muscles and its own nerve
supply, so each segment or group of segments may function in-
dependently. When circular muscles contract, the segments be-
come thinner and elongate. When longitudinal muscles
contract, the segments become thicker and shorten. By alter-
nating circular muscle contraction and longitudinal muscle
contraction, the animal moves forward.
In annelids, the tube-within-a-tube body plan has led to
specialization of the digestive tract. For example, the digestive
system may include a pharynx, esophagus, crop, gizzard, in-
testine, and accessory glands. They have an extensive closed
circulatory systemwith blood vessels that run the length of the
body and branch to every segment. The nervous system con-
sists of a brain connected to a ventral solid nerve cord, with ganglia
in each segment. The excretory system consists of nephridia in
most segments. Anephridium is a tubule that collects waste
material and excretes it through an opening in the body wall.
Marine Worms
Most annelids are marine; their class name, Polychaeta,
refers to the presence of many setae. Setae are bristles that
anchor the worm or help it move. In polychaetes, the setae are
in bundles on parapodia, which are paddlelike appendages
found on most segments. These are used not only in swim-
ming but also as respiratory organs where the expanded
surface area allows for increased exchange of gases. Clam
worms (Fig. 30.13) such as Nereis are predators. They prey
on crustaceans and other small animals, which are captured
by a pair of strong, chitinous jaws that extend with a part of
the pharynx when the animal is feeding. Associated with its
way of life, Nereis has a well-dened head region with eyes
and other sense organs.
Other polychaetes are sedentary (sessile) tube worms,
with tentacles that form a funnel-shaped fan. Water cur-
rents, created by the action of cilia, trap food particles that
are directed toward the mouth. They are sessile lter feeders; a
sorting mechanism rejects large particles, and only the
smaller ones are accepted for consumption.
Polychaetes have breeding seasons, and only during
these times do the worms have functional sex organs. In
Nereis, many worms concurrently shed a portion of their
bodies containing either eggs or sperm, and these oat to
the surface, where fertilization takes place. The zygote
rapidly develops into a trochophore larva, just as in marine
clams. The existence of this larva in both the annelids and
mollusks is evidence that these two groups of animals are
related.
Polychaetes are marine worms with bundles of
setae attached to parapodia.
a. Clam worm, Nereis
jaw
pharynx
(extended)
sensory projections
eyes
parapodia
b. Fan worm, Spirobranchus
Figure 30.13 Polychaete diversity.
a. Nereis is a predaceous polychaete that has a head region. Note the parapodia, which are used for swimming and as respiratory organs.
b. Fan worms (a type of tube worm) are sessile lter feeders whose cilia-lined tentacles funnel food particles to the mouth.
Ancestral protists
coelomates
protostomes
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Earthworms
The oligochaetes, which include earthworms, have few setae
per segment. Earthworms (e.g., Lumbricus) do not have a
well-developed head or parapodia (Fig. 30.14). Their setae
protrude in pairs directly from the surface of the body. Lo-
comotion, which is accomplished section by section, utilizes
muscle contraction and the setae. When longitudinal mus-
cles contract, segments bulge and their setae protrude into
the soil; then, when circular muscles contract, the setae are
withdrawn, and these segments move forward.
Earthworms reside in soil where there is adequate mois-
ture, which keeps the body wall moist for gas exchange pur-
poses. They are scavengers that do not have an obvious
head and feed on leaves or any other organic matter, living
or dead, which can conveniently be taken into the mouth
along with dirt. Food drawn into the mouth by the action of
the muscular pharynx is stored in a crop and ground up in a
thick, muscular gizzard. Digestion and absorption occur in a
long intestine whose dorsal surface has an expanded region
called a typhlosole that increases surface for absorption.
Earthworm segmentation, which is so obvious externally,
is also internally evidenced by septa. The long ventral solid
nerve cord leading from the brain has ganglionic swellings
and lateral nerves in each segment. The paired nephridia in
most segments have two openings: one is a ciliated funnel
that collects coelomic uid, and the other is an exit in the
body wall. Between the two openings is a convoluted region
where waste material is removed from the blood vessels
about the tubule. Red blood moves anteriorly in the dorsal
blood vessel, which connects to the ventral blood vessel by
ve pairs of connectives called hearts. Pulsations of the
dorsal blood vessel and the ve pairs of hearts are responsi-
ble for blood ow. As the ventral vessel takes the blood to-
ward the posterior regions of the worms body, it gives off
branches in every segment. Altogether, segmentation is evi-
denced by
body rings
coelom divided by septa
setae on most segments
ganglia and lateral nerves in each segment
nephridia in most segments
branch blood vessels in each segment
The worms are hermaphroditic; the male organs are the
testes, the seminal vesicles, and the sperm ducts, and the fe-
male organs are the ovaries, the oviducts, and the seminal
receptacles. Two worms lie parallel to each other facing in
opposite directions. The fused midbody segment, called a
clitellum, secretes mucus, protecting the sperm from drying
anterior end
anterior end
clitellum
clitellum
b.
mouth
a.
setae
ganglion
nerve cord
seminal receptacle
seminal vesicle
ovary
sperm duct
nephridium
coelom
muscle layers
septum
clitellum
intestine
blood
vessels
gizzard
anus
crop
esophagus
hearts
pharynx
brain
Figure 30.14 Earthworm.
a. Internal anatomy of the anterior part of an earthworm. Notice that
each body segment bears four pairs of setae and that internal septa
divide the coelom into compartments. b. When earthworms, such as
Lumbricus, mate, they are held in place by a mucus secreted by the
clitellum. The worms are hermaphroditic but practice cross-
fertilization; sperm pass from the seminal vesicles of each to the
seminal receptacles of the other.
Parasitic leeches attach themselves to the body of their hosts
and suck blood or other body uids from shes, turtles, snails,
or mammals. The European species of Hirudo medicinalis was
used in the 1700s and 1800s to let blood in feverish patients in
order to withdraw any poisons and excess blood. This proce-
dure, which is most debilitating for the patient, is no longer
practiced, but leeches are still used on occasion to remove blood
from bruised skin (Fig. 30A). They are also applied to small
body parts like ngers and toes that have just been reattached to
the body. The sucking by the leech unclogs small blood vessels,
causing blood to ow normally again through the part. The an-
terior sucker of the medicinal leech has three jaws that
saw through the skin, producing a Y-shaped incision through
which blood is drawn up by the sucking action of the muscular
pharynx. The salivary glands secrete an anticoagulant called
hirudin, which keeps the blood owing while the leech is feed-
ing. Other salivary ingredients dilate the hosts blood vessels
and act as an anesthetic. A medicinal leech can take up to ve
times its body weight in blood because the crop has pouches
where the blood can be stored as the animal expands in size.
When it has taken its ll, the leech drops off and digests its meal.
Complete digestion takes a long time, and its been suggested
that a leech needs to feed only once a year.
Not all leeches suck blood or uids; some are even preda-
ceous and eat a variety of small invertebrates.
633
Medicinal Bloodsuckers
Figure 30A Medicinal leeches, Hirudo medicinalis.
They are sometimes used medically to remove blood that has
accumulated in damaged tissues.
out as they pass between the worms. After the worms sepa-
rate, the clitellum of each produces a slime tube, which is
moved along over the anterior end by muscular contrac-
tions. As it passes, eggs and the sperm received earlier are
deposited and fertilization occurs. The slime tube then
forms a cocoon to protect the worms as they develop. There
is no larval stage.
It is interesting to compare the anatomy of marine clam
worms to terrestrial earthworms because it highlights the
manner in which earthworms are adapted to life on land.
Marked cephalization is seen in the nonpredatory earth-
worms that extract organic remains from the soil they eat.
The lack of parapodia helps reduce the possibility of water
loss and facilitates burrowing in soil. The clam worm
makes use of external water, while the earthworm provides
a mucous secretion to aid fertilization. It is the water form
that has the swimming, or trochophore larva, and not the
land form.
Earthworms, which burrow in the soil, lack obvious
cephalization and parapodia. They are
hermaphroditic, and there is no larval stage.
Leeches
Leeches are usually found in fresh water, but some are ma-
rine or even terrestrial. They have the same body plan as
other annelids but they have no setae, and each body ring
has several transverse grooves. Most leeches are only 26 cm
in length, but some, including the medicinal leech, are as
long as 20 cm.
Among their modications are two suckers, a small oral
one around the mouth and a large posterior one. While some
leeches are free-living predators, most are uid feeders that
attach themselves to open wounds. Some bloodsuckers,
such as the medicinal leech, are able to cut through tissue.
Leeches are able to keep blood owing and prevent clotting
by means of a substance in their saliva known as hirudin, a
powerful anticoagulant. This has added to their potential
usefulness in the eld of medicine today, as discussed in the
accompanying reading.
Leeches are modifed in a way that lends itself to
the parasitic way of life. Some are external
parasites known as bloodsuckers.
30.5 Arthropods
Arthropods (phylum Arthro-
poda) are extremely diverse.
Over one million species
have been discovered and de-
scribed, but some experts
suggest that as many as 30
million arthropods could
existmostly insects.
What characteristics ac-
count for the success of arthro-
pods? Arthropod literally
means jointed foot, but actu-
ally they have freely movable jointed appendages. The exoskele-
ton of arthropods is composed primarily of chitin, a strong,
exible, nitrogenous polysaccharide. The exoskeleton serves
many functions such as protection, attachment for muscles, lo-
comotion, and prevention of desiccation. However, because
this exoskeleton is hard and nonexpandable, arthropods must
undergo molting, or shedding of the exoskeleton, as they
grow larger. Before molting, the body secretes a new, larger
exoskeleton, which is soft and wrinkled, underneath the old
one. After enzymes partially dissolve and weaken the old exo-
skeleton, the animal breaks it open and wriggles out. The new
exoskeleton then quickly expands and hardens.
Arthropods are segmented, but some segments are fused
into regions, such as a head, a thorax, and an abdomen. In
trilobites, an early and now extinct arthropod, there was a
pair of appendages on each body segment. In modern arthro-
pods, appendages are specialized for such functions as walking,
swimming, reproducing, eating, and sensory reception.
These modications account for much of the diversity of
arthropods. Each of the ve major groups of arthropods (Fig.
30.15) contains species that are adapted to terrestrial life.
Arthropods have a well-developed nervous system. There is
Figure 30.15 Arthropod diversity.
a. A millipede has only one pair of antennae, and the head is followed by a series of segments, each with two pairs of appendages. b. The hairy
tarantulas of the genus Aphonopelma are dark in color and sluggish in movement. Their bite is harmless to people. c. A crab is a crustacean with
a calcied exoskeleton, one pair of claws and four other pairs of walking legs. d. A wasp is an insect with two pairs of wings, both used for ying,
and three pairs of walking legs. e. A centipede has only one pair of antennae, and the head is followed by a series of segments, each with a
single pair of appendages.
c. Dungeness crab, Cancer e. Stone centipede, Lithobius
a. Flat-backed millipede, Sigmoria
b. Tarantula, Aphonopelma
d. Paper wasp, Polistes
Ancestral protists
coelomates
protostomes
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a brain and a ventral solid nerve cord. The
head bears various types of sense organs,
including antennae (or feelers) and eyes of
two typescompound and simple. The
compound eye is composed of many com-
plete visual units, each of which operates
independently. The lens of each visual unit
focuses an image on the light-sensitive
membranes of a small number of photore-
ceptors within that unit. Vision is not acute,
but it is much better for details of move-
ment than with our eyes.
Crustaceans
Crustaceans are a group of largely marine
arthropods that include barnacles,
shrimps, lobsters, and crabs. There are also
some freshwater crustaceans, including the
craysh, and some terrestrial ones, includ-
ing the sowbug or roly-poly bug.
Crustaceans are named for their hard
shells; the exoskeleton is calcied to a
greater degree in some forms than in oth-
ers. Although crustacean anatomy is ex-
tremely diverse, the head usually bears a
pair of compound eyes and ve pairs of ap-
pendages. The rst two pairs, called anten-
nae, lie in front of the mouth and have
sensory functions. The other three pairs are
mouthparts used in feeding.
In craysh such as Cambaris, the thorax
bears ve pairs of walking legs. The rst
walking leg is a pinching claw (Fig. 30.16).
The gills are situated above the walking
legs. The head and thorax are fused into a cephalothorax, which
is covered on the top and sides by a nonsegmented carapace.
The abdominal segments are equipped with swimmerets,
small paddlelike structures. The last two segments bear the
uropods and the telson, which make up a fan-shaped tail to
propel the craysh backward.
Ordinarily, a craysh lies in wait for prey. It faces out
from an enclosed spot with the claws extended and the an-
tennae moving about. The claws seize any small animal, dead
or alive, that happens by and carry it to the mouth. When a
craysh moves about, it generally crawls slowly, but it may
swim rapidly by using its heavy abdominal muscles and tail.
The digestive system includes a stomach, which is di-
vided into two main regions: an anterior portion called the
gastric mill, equipped with chitinous teeth to grind coarse
food, and a posterior region, which acts as a lter to prevent
coarse particles from entering the digestive glands where ab-
sorption takes place. Green glands lying in the head region,
anterior to the esophagus, excrete metabolic wastes through
a duct that opens externally at the base of the antennae. The
coelom, which is so well developed in the annelids, is reduced
in the arthropods and is composed chiey of the space about
the reproductive system. A heart within a pericardial cavity
pumps blood containing the respiratory pigment hemo-
cyanin into a hemocoel consisting of sinuses (open spaces),
where the hemolymph ows about the organs. (Whereas he-
moglobin is a red iron-containing pigment, hemocyanin is a
blue copper-containing pigment.) This is an open circulatory
system because blood is not contained within blood vessels.
The nervous system is quite similar to that of the earth-
worm. There is a brain, as well as a ventral nerve cord that
passes posteriorly. Along the length of the nerve cord, gan-
glia in segments give off 8 to 19 paired lateral nerves.
The sexes are separate in the craysh, and the gonads
are located just ventral to the pericardial cavity. In the male,
a coiled sperm duct opens to the outside at the base of the
fth walking leg. Sperm transfer is accomplished by the rst
two pairs of swimmerets, which are enlarged and quite
strong. In the female, the ovaries open at the bases of the
third walking legs. A stiff fold between the bases of the
fourth and fth pairs of walking legs serves as a seminal re-
ceptacle. Following fertilization, the eggs are attached to the
swimmerets of the female.
Figure 30.16 Male craysh.
a. Externally, it is possible to observe the jointed appendages. Note the swimmerets and the
walking legs, which include the claws. These appendages, plus a portion of the carapace,
have been removed from the right side so that the gills are visible. b. Internally, the parts of
the digestive system are particularly visible. The circulatory system can also be clearly seen.
Note the ventral nerve cord.
cephalothorax
first walking
leg (modified
as a pincerlike
claw)
carapace
a.
b.
compound
eye
excretory pore
stomach
mouth
antennae
abdomen
gills
brain
green
gland
mouth
pericardial cavity dorsal abdominal
artery
heart
anus
ventral
nerve cord
sperm
duct testis digestive gland
opening of
sperm duct
anus
telson
uropods
swimmerets
fifth walking leg
claspers
fourth walking leg
third walking leg
second
walking leg
Insects
There are more different kinds of insects than any other type
animal. Examples of insects include crickets and grasshop-
pers, dragonies, water striders, beetles, moths and butter-
ies, ies, bees, and wasps (Fig. 30.17).
Insects have certain features in common. The body is di-
vided into a head, a thorax and an abdomen. The head usu-
ally bears a pair of sensory antennae, a pair of compound eyes,
and several simple eyes. The mouthparts are adapted to the
way of life: a grasshopper has mouthparts that chew, and a
buttery has a long tube for siphoning the nectar of owers.
The thorax bears three pairs of legs and no pairs, one pair, or
two pairs of wings, and the abdomen contains most of the
internal organs. Wings enhance an insects ability to survive
by providing a way of escaping enemies, nding food, facil-
itating mating, and dispersing the offspring. The exoskele-
ton of an insect is lighter and contains less chitin than that of
many other arthropods.
In the grasshopper (Fig. 30.18), the third pair of legs is
suited to jumping. There are two pairs of wings. The
forewings are tough and leathery, and when folded back at
rest, they protect the broad, thin hindwings. On the lateral
surface, the rst abdominal segment bears a large tympanum
on each side for the reception of sound waves. The posterior
region of the exoskeleton in the female has two pairs of pro-
jections that form an ovipositor, which is used to dig a hole
in soil where eggs are laid.
The digestive system is suitable for a herbivorous diet.
In the mouth, food is broken down mechanically by mouth-
Figure 30.17 Insect diversity.
a. Walking sticks are herbivorous, with biting and chewing mouthparts. b. Bees have four translucent wings and a thorax separated from the
abdomen by a narrow waist. c. Flies have a single pair of wings and lapping mouthparts. d. Dragonies have two pairs of similar wings. They
catch and eat other insects while ying. e. Butteries have forewings larger than hindwings. Their mouthparts form a long tube for siphoning
up nectar from owers.
a. Walking stick, Diapheromera
b. Bee, Apis
c. Housefly, Musca
d. Dragonfly, Aeshna
e. American copper butterfly, Lycaena
parts and enzymatically by salivary secretions. Food is
temporarily stored in the crop before passing into a gas-
tric mill, where it is nely ground before digestion is
completed in the stomach. Nutrients are absorbed into
the hemocoel from outpockets called gastric ceca; a ce-
cum is a cavity open at one end only. The stomach is fol-
lowed by an intestine and rectum, which empties by
way of an anus. The excretory system consists of
Malpighian tubules, which extend into the hemocoel
and collect nitrogenous wastes that are concentrated
and excreted into the digestive tract. The formation of a
solid nitrogenous waste, namely uric acid, conserves
water.
The respiratory system begins with openings in the
exoskeleton called spiracles. From here, the air enters
small tubules called tracheae (Fig. 30.18a). The tracheae
branch and rebranch until they end intracellularly,
where the actual exchange of gases takes place. The
movement of air through this complex of tubules is not
a passive process; air is pumped by alternate contraction
and relaxation of the body wall through a series of sev-
eral bladderlike structures, which are attached to the
tracheae near the spiracles. Air enters the anterior four
spiracles and exits by the posterior six spiracles. Breath-
ing by tracheae may account for the small size of insects
(most are less than 60 mm in length), since the tracheae
are so tiny and fragile that they would be crushed by
any signicant amount of weight.
The circulatory system contains a slender, tubular
heart that lies against the dorsal wall of the abdominal ex-
oskeleton and pumps hemolymph into an aorta that leads
to a hemocoel, where it circulates before returning to the
heart again. The hemolymph is colorless and lacks a respi-
ratory pigmentthe tracheal system exchanges gases.
Reproduction is adapted to life on land. The male has
a penis, and sperm passed to the female are stored in a
other colonial insects. Insects are so numerous and so di-
verse that the study of this one group is a major specialty in
biology called entomology.
It is of interest to compare the adaptations of a grasshop-
per to land with the adaptations of a craysh to an aquatic
environment. In craysh, gills take up oxygen from water,
while in the grasshopper, tracheae allow oxygen-laden air to
enter the body. Appropriately, the craysh has an oxygen-
carrying pigment, but a grasshopper has no such pigment in
its blood. Aliquid nitrogenous waste (ammonia) is excreted
by a craysh, while a solid nitrogenous waste (uric acid) is
excreted by a grasshopper. Only in grasshoppers (1) is there
a tympanum for the reception of sound waves and (2) do
males have a penis for passing sperm to females without
possible desiccation and females an ovipositor for laying
eggs in soil. Craysh utilize their uropods when they swim;
a grasshopper has legs for hopping and wings for ying.
rectum
Head Thorax Abdomen
simple eye tympanum
antenna
compound eye
forewing
hindwing
ovipositor
spiracles
air sac
spiracle tracheae
heart
ovary
Malpighian tubules
crop
aorta
vagina
seminal
receptacle
oviduct
intestine
nerve
ganglion
stomach gastric
ceca
salivary
gland
brain
mouth
b.
a.
Figure 30.18 Female grasshopper.
a. Externally, the tympanum receives sound waves, and the hopping legs
and the wings are for locomotion. b. Internally, the digestive system is
specialized. The Malpighian tubules excrete a solid nitrogenous waste (uric
acid). A seminal receptacle receives sperm from the male, which has a penis.
seminal receptacle. Internal fertilization protects both ga-
metes and zygotes from drying out. The female deposits the
fertilized eggs in the ground with her ovipositor.
Metamorphosis is a change in form and physiology that
occurs as an immature stage, called a larva, becomes an
adult. Grasshoppers undergo gradual metamorphosis, or a
gradual change in form, as the animal matures. The imma-
ture grasshopper, called a nymph, is recognizable as a
grasshopper, even though it differs somewhat in shape and
form from the adult. Other insects, such as butteries, un-
dergo complete metamorphosis, involving drastic changes in
form. At rst, the animal is a wormlike larva (caterpillar)
with chewing mouthparts. It then forms a case, or cocoon,
about itself and becomes a pupa. During this stage, the body
parts are completely reorganized; the adult then emerges
from the cocoon. This life cycle allows the larvae and adults
to typically make use of different food sources. Most eating
by insects occurs during the larval stage.
Insects also show remarkable behavior adaptations, ex-
emplied by the social systems of bees, ants, termites, and
By using data from many sources, the evolution, and thus the
classication, of an animal group can be claried. Agood exam-
ple comes from the spiders, members of the class Arachnida, or-
der Araneae. Spiders are distinguished from other similar
animals by their ability to weave webs of silk. The silk is pro-
duced from glands in their abdomens, and it emerges from
modied appendages called spinnerets.
Arachnologists (scientists who study spiders) are largely
convinced that spider webs were originally used to line a cavity
or a burrow in which an early spider hid. Many spiders that still
live in such burrows put a collar of silk around their burrow en-
trances to detect prey over a wider area than they could other-
wise easily search. From this array of threads evolved the basic
sheet web, made by a wide variety of primitive spider families.
The sheet of closely woven threads is useful not only in signal-
ing the presence of prey, but also in slowing the prey as the in-
sects legs tangle in the matted silk. However, the appearance of
the sheet web in many clearly unrelated families of spiders sug-
gests that it evolved numerous times and cannot be used to an-
swer questions about the true relationships of spider families.
Only if some advanced feature of a sheet web is shared by two
or more families does it indicate a common ancestry.
The geometric orb web probably evolved from a sheet web. The
orb web, which has threads placed in a regular fashion, uses less
silk and thus costs less. Until recently, the orb web was thought
to have arisen at least twice because it is made by two groups of
spiders that look very different. The dinopoids (superfamily
Dinopoidea) includes spiders with a special spinning apparatus,
the cribellum, which produces extremely ne bers. The araneoids
(superfamily Araneoidea), which also make orb webs, lacks the
cribellum, and thus are called ecribellates. If the cribellum is a spe-
cialization that arose only in the dinopoid lineage, then it seems
most logical that the araneoids and dinopoids are not closely re-
lated and their orb webs are quite separate developments.
Arachnologists noticed, however, the great similarities of the
orbs made by the two familieseven extending to the specic
movements made by the spiders legs while weaving them. Per-
haps then the two families are closely related despite the lack of a
cribellum in the araneoid line. The Finnish biologist Pekka Lehti-
nen made a sweeping study of all sorts of spiders in 1967 and
found numerous examples of spiders that were
nearly identical except for the presence or absence of
a cribellum. The mass of evidence he accumulated
convinced arachnologists that the cribellum could
easily have been lost in the araneoid line of descent.
By considering the new data regarding the
cribellum, together with observations of orb-web
building behavior, most arachnologists are con-
vinced that the orb web originated only once, and
that the dinopoids and araneoids do share a com-
mon ancestor (Fig. 30B).
638
Spider Webs and Spider Classication
primitive spiders tarantulas araneoids dinopoids
cribellum lost
cribellum and
orb web
silk and spinnerets
= orb web
Figure 30B Evolution of orb web.
a. Evolutionary tree of spiders. b. Orb web of the garden spider Araneus diadematus differs only in detail from (c) the orb of a cribellate
spider, the New Zealand species Waitkera waitkerensis. The leg movements used by both during web construction are very similar, making
it likely that there is a close evolutionary relationship between them, despite their anatomical differences.
a.
b. c.
Arachnids
The arachnids (class Arachnida, about 50,000 species) in-
clude terrestrial spiders, scorpions, ticks and mites
(Fig. 30.19). In this group, the cephalothorax bears six pairs
of appendages: the chelicerae and the pedipalps and four
pairs of walking legs. The cephalothorax is followed by an
abdomen that contains internal organs.
Scorpions are the oldest terrestrial arthropods. Today,
they occur in the tropics, subtropics, and temperate regions
of North America. They are nocturnal and spend most of the
day hidden under a log or a rock. In scorpions the pedipalps
are large pincers, and the long abdomen ends with a stinger
that contains venom. Ticks and mites are parasites. Ticks
suck the blood of vertebrates and are sometimes transmit-
ters of diseases, such as Rocky Mountain spotted fever or
Lyme disease. Chiggers, the larvae of certain mites, feed on
the skin of vertebrates.
Spiders, the most familiar arachnids, have a narrow
waist that separates the cephalothorax from the abdomen.
Each chelicera consists of a basal segment and a fang that de-
livers poison to paralyze or kill prey. Two venom glands are
located in the chelicerae or in the head. The pedipalps assist
in sensing and holding the prey. Digestive juices from the
mouth liquefy the tissues and the resulting broth is sucked
into the stomach. Spiders use silk threads for all sorts of
purposes, from lining their nests to catching prey. The read-
ing on the previous page tells how biologists have used
web-building behavior as a way to discover how spiders are
related.
The internal organs of spiders also show how they are
adapted to a terrestrial way of life. Malpighian tubules work
in conjunction with rectal glands to reabsorb ions and water
before a relatively dry nitrogenous waste (uric acid) is ex-
creted. Invaginations of the inner body wall form lamellae
(pages) of their so-called book lungs. Air owing into the
folded lamellae on one side exchanges gases with blood
owing in the opposite direction on the other side.
The arthropods are a diverse group of animals, of
which some members are adapted to an aquatic
existence and others are adapted to a terrestrial
way of life. Jointed appendages and a water
repellent exoskeleton assists locomotion and
lessens the threat of drying out on land.
pedipalp chelicera
with fang
ventral view
of mouthparts
blood flow
between
lamellae
lamellae of lung
body
wall
air flowing in through spiracle
stinger
pedipalp
chelicera
d. Book lung anatomy
b. Black widow spider, Latrodectus
c. Wood ticks, Ixodes
a. Kenyan giant scorpion, Pandinus
Figure 30.19 Arachnid diversity.
a. A scorpion has pincerlike chelicerae and pedipalps, and there is a long abdomen ending with a stinger that contains venom. b. Most spiders
are harmless, but the venom of the black widow spider is dangerous to humans. c. In the western United States, the wood tick carries a
debilitating disease called Rocky Mountain spotted fever. d. Arachnids breathe by means of book lungs, in which the pages are double sheets
of thin tissue (lamellae).
Summarizing the Concepts
30.1 Evolution and Classication of Animals
Animals are multicellular heterotrophs that ingest their food. The clas-
sication of animals is based on symmetry, number of germ layers,
type of coelom, and body plan, among other features. An evolutionary
tree based on these features depicts a possible evolutionary relation-
ship between the animals.
Sponges are asymmetrical, and cnidarians have radial symmetry. All
other phyla contain bilaterally symmetrical animals (Table 30.2). Flat-
worms have three germ layers but no coelom. Roundworms have a
pseudocoelom and a tube-within-a-tube body plan.
30.3 Mollusks
Mollusks, along with annelids and arthropods, are protostomes because
the rst embryonic opening becomes the mouth. The body of a mollusk
typically contains a visceral mass, a mantle, and a foot. Predaceous
squids display marked cephalization, move rapidly by jet propulsion,
and have a closed circulatory system. Bivalves (e.g., clams), which have
a hatchet foot, are lter feeders. Water enters and exits by siphons, and
food trapped on the gills is swept toward the mouth.
30.4 Annelids
Annelids are segmented worms; segmentation is seen both externally
and internally. Polychaetes are marine worms that have parapodia. A
clam worm is a predaceous marine worm with a dened head region.
Earthworms are oligochaetes that scavenge for food in the soil and do
not have a well-dened head region.
30.5 Arthropods
Arthropods are the most varied and numerous of animals. Their suc-
cess is largely attributable to a exible exoskeleton and specialization
of body regions. Like many other arthropods, crustaceans have a head
that bears compound eyes, antennae, and mouthparts. The craysh, a
crustacean, also illustrates features such as an open circulatory system,
respiration by gills, and a ventral solid nerve cord. Like most other in-
sects, grasshoppers have wings and three pairs of legs attached to the
thorax. Grasshoppers also have many other features that illustrate
adaptation to a terrestrial life, such as respiration by tracheae.
Spiders are arachnids with chelicerae, pedipalps, and four pairs of
walking legs attached to a cephalothorax. Spiders, too, are adapted to
life on land, and they spin silk which is used in various ways. Some spi-
ders spin webs, and the type of web can be used to discover the evolu-
tionary relationship among spiders.
T
he many types of animals in a coral
reef form a complex community that is
admired by both snorkelers and scuba
divers. The various types of sh and shell-
sh in a coral reef are sources of food for
millions of people. Like a tropical forest,
coral reefs are most likely sources of medi-
cines yet to be discovered. And a reef serves
as a storm barrier that protects the shoreline
and provides a safe harbor for ships.
Reefs around the globe are being de-
stroyed. Tons of soil from deforested tracts
of land bring nutrients that stimulate the
growth of all kinds of algae. This has con-
tributed to a population explosion of the
crown-of-thorn starsh that are devouring
Australias 1,200 mile-long Great Barrier
Reef. Reefs are also being damaged by pol-
lutants that seep into the sea from facto-
ries, farm elds, and sewers. Stress, com-
bined with unusually warm sea water, has
caused the corals to expel their colorful,
symbiotic algae, which carry on photosyn-
thesis and help sustain the corals. So-called
coral bleaching has been noticed in reefs of
the Pacic Ocean and the Caribbean.
Might worldwide global warming also
contribute to coral bleaching and death?
Marine scientist Edgardo Gomez esti-
mates that 90% of coral reefs of the Philip-
pines are dead or deteriorating due to
pollution, but especially due to overshing.
The methods are sinister, including the use
of dynamite to kill the sh, making it easier
to scoop them up, use of cyanide to stun the
sh to capture them alive, and using satel-
lite navigation systems to home in on areas
where mature sh are spawning to repro-
duce. If all large herbivores are killed, sea-
weed overgrows and kills the coral.
Paleobiologist Jeremy Jackson of the
Smithsonian Tropical Research Institute
near Panama City, Republic of Panama,
wonders if he is doing enough to warn the
public that reefs around the world are in
danger. He estimates that we may lose
60% of all coral reefs by the year 2050.
Questions
1. Do you think it would be possible to
make the public care about the loss of
coral reefs? Explain.
2. When and under what circumstances do
dire predictions help preserve the
environment?
3. Considering what is causing the loss of
coral reefs, would it be possible to save
them? How?
Sponges Cnidarians Flatworms Roundworms
Symmetry Radial or none Radial Bilateral Bilateral
Type of body plan Sac Sac Tube-within-a-tube
Tissue layers Two Three Three
Level of organization Cell Tissue Organ Organ
Body cavity Acoelomate Pseudocoelomate
Table 30.2 Comparison of Animals without a True Coelom
Go To Student OLC
Studying the Concepts
1. What does the evolutionary tree (see Fig. 30.2) tell you about
the evolution of the animals studied in this chapter? 620
2. List the types of cells found in a sponge, and describe their
functions. 621
3. What are the two body forms found in cnidarians? Explain
how they function in the life cycle of various types of cnidari-
ans. 622
4. Describe the anatomy of hydra, pointing out those features
that typify cnidarians. 623
5. Describe the anatomy of a free-living planarian, pointing out
those features that typify nonparasitic atworms. 624-25
6. Describe the parasitic atworms, and describe the life cycle of
a pork tapeworm. 625-26
7. Describe the anatomy of Ascaris, pointing out those features
that typify roundworms. 627
8. What are the general characteristics of mollusks? Contrast the
anatomy of the clam and the squid, indicating how each is
adapted to its way of life. 628-29
9. What are the general characteristics of annelids? Contrast the
anatomy of the clam worm and the earthworm, indicating
how each is adapted to its way of life. 631-32
10. What are the general characteristics of arthropods? Describe
the specic features of craysh, the grasshopper, and a spider,
indicating how each is adapted to its way of life. 634-39
Testing Yourself
Choose the best answer for each question.
1. Label the following diagram of the cnidarian polyp:
Chapter 30 Animals: Part 1 641 30-25
a.
b.
c.
d.
2. Which of these is not a characteristic of animals?
a. heterotrophic
b. diplontic life cycle
c. usually practice sexual reproduction
d. have contracting bers
e. single cells or colonial
3. The evolutionary tree of animals shows that
a. cnidarians evolved directly from sponges.
b. atworms evolved directly from roundworms.
c. both sponges and cnidarians evolved from protista.
d. coelomates gave rise to the acoelomates.
e. All of these are correct.
4. Which of these sponge characteristics is not typical of
animals?
a. They have a type of skeleton.
b. They practice sexual reproduction.
c. They have the cellular level of organization.
d. They are asymmetrical.
e. Both c and d are correct.
5. Which of these is mismatched?
a. spongesspicules
b. tapewormsproglottids
c. cnidariansnematocysts
d. crayshcompound eyes
e. roundwormscilia
6. Flukes and tapeworms
a. show cephalization.
b. have well-developed reproductive systems.
c. have well-developed nervous systems.
d. have a tube-within-a-tube body plan.
e. are plant parasites.
7. The presence of mesoderm
a. is necessary to mesoglea formation.
b. restricts the development of a coelom.
c. is associated with the organ level of organization.
d. is associated with the development of muscles.
e. Both b and c are correct.
8. Ascaris is a parasitic
a. roundworm. d. sponge.
b. atworm. e. mollusk.
c. hydra.
9. Which of these best shows that annelids and arthropods are
closely related? In both,
a. the rst embryonic opening becomes the mouth.
b. there is a complete digestive tract.
c. there is a ventral solid nerve cord.
d. there are segments.
e. All of these are correct.
a. clamgills
b. lobstergills
c. grasshopperbook lungs
d. polychaeteparapodia
e. All of these are matched correctly.
11. Which of these is an incorrect statement?
a. Planarians are heterotrophs.
b. Spiders are carnivores in the phylum Arthropoda.
c. Clams are lter feeders in the phylum Mollusca.
d. Earthworms are scavengers in the phylum
Arthropoda.
e. Squids are predators in the phylum Mollusca.
12. Aradula is a unique organ for feeding found in
a. mollusks. d. arthropods.
b. roundworms. e. All of these are correct.
c. annelids.
a. roundwormshydrostatic skeleton
b. crayshwalking legs
c. clamhatchet foot
d. grasshopperwings
e. earthwormmany cilia
Thinking Scientically
1. Considering the characteristics of animals (page 618):
a. Compare corresponding characteristics of plants and ani-
mals.
b. What one animal characteristic can be associated with an
animals need to acquire food?
c. Why is it reasonable to assume that animals arose from
protozoans?
d. Some animals can regenerate from a small part of the
whole. Why would you expect only simple animals, and
not those with highly differentiated tissues, to be able to
do this?
2. Some animals have a complete (both mouth and anus) diges-
tive tract, and some have an incomplete (only one opening)
digestive tract.
a. In which type of animal would you expect the digestive
tract to have specialized parts? Why?
b. Why is it consistent that a coelomate animal would have a
complete digestive tract with specialized parts?
c. In which type of animal would you expect the animal to be
a discontinuous (eat at intervals) feeder?
d. In more complex animals, blood vessels deliver the prod-
uct of digestion to the cells. How does a planarian make do
without a circulatory system?
a. atwormsacoelomate
b. molluskreduced coelom
c. insectshemocoel
d. crayshcoelom divided by septa
e. clam wormtrue coelom
15. Label this diagram.
Understanding the Terms
annelid 631
arachnid 639
arthropod 634
asymmetry 619
bilateral symmetry 619
bivalve 629
cephalization 619
chitin 634
cnidarian 622
coelom 619
coral 623
crustacean 635
deuterostome 619
lter feeder 621
uke 625
gastrovascular cavity 623
hemocoel 635
hermaphrodite 625
hydrostatic skeleton 626
insect 636
invertebrate 618
leech 633
Malpighian tubule 637
metamorphosis 637
mollusk 628
molting 634
nematocyst 622
nephridium 631
protostome 619
pseudocoelom 619
radial symmetry 619
segmentation 619
sessile 619
tapeworm 625
trachea 637
vertebrate 618
Match the terms to these denitions:
a. Air tube in insects located between the spiracles
and the tracheoles.
b. Body cavity lying between the digestive tract and
body wall that is completely lined by mesoderm.
c. Body plan having two corresponding or comple-
mentary halves.
d. Periodic shedding of the exoskeleton in arthro-
pods.
e. Segmentally arranged, paired excretory tubules of
many invertebrates, as in the earthworm.
a. e. g.
h.
i.
j.
n.
m.
l.
k.
d.
c.
b.
f.
Using Technology
Your study of animals is supported by these available
technologies:
Essential Study Partner CD-ROM
Evolution & Diversity Invertebrates
Visit the Mader web site for related ESP activities.
Exploring the Internet
The Mader Home Page provides resources and tools as
you study this chapter.
http://www.mhhe.com/biosci/genbio/mader

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