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Chapter 33

Invertebrates

PowerPoint Lectures for


Biology, Seventh Edition
Neil Campbell and Jane Reece

Lectures by Chris Romero


Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Overview: Life Without a Backbone


Invertebrates
Are animals that lack a backbone
Account for 95% of known animal species

Figure 33.1
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Chordata

Echinodermata

Other bilaterians (including


Nematoda, Arthropoda,
Mollusca, and Annelida)

Porifera

Cnidaria

A review of animal phylogeny

Deuterostomia

Bilateria

Eumetazoa

Ancestral colonial
choanoflagellate

Figure 33.2
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Exploring invertebrate diversity


PORIFERA (5,500 species)

A sponge
PLACOZOA (1 species)

CNIDARIA (10,000 species)

A jelly
KINORHYNCHA (150 species)

0.5 mm

250 m

A placozoan (LM) A kinorhynch (LM)


ROTIFERA (1,800 species)
PLATYHELMINTHES (20,000 species)

A marine flatworm
ECTOPROCTA (4,500 species)

Figure 33.3

A rotifer (LM)
PHORONIDA (20 species)

Ectoprocts Phoronids

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Exploring invertebrate diversity


BRACHIOPODA (335 species)

NEMERTEA (900 species)

A brachiopod
ACANTHOCEPHALA (1,100 species)

A ribbon worm
CTENOPHORA (100 species)

5 mm

An acanthocephalan

A ctenophore, or comb jelly

MOLLUSCA (93,000 species)

ANNELIDA (16,500 species)

An octopus

A marine annelid
PRIAPULA (16 species)

LORICIFERA (10 species)


50 m

Figure 33.3

A loriciferan (LM)

A priapulan

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Exploring invertebrate diversity

ARTHROPODA (1,000,000 + species)

NEMATODA (25,000 species)

A roundworm

A scorpion (an arachnid)

CYCLIOPHORA (1 species)

TARDIGRADA (800 species)


100 m

100 m

A cycliophoran (colorized SEM) Tardigrades (colorized SEM)


HEMICHORDATA (85 species)

ONYCHOPHORA (110 species)

An onychophoran

An acorn worm

ECHINODERMATA (7,000 species)

Figure 33.3

A sea urchin

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CHORDATA (52,000 species)

A tunicate

Sponges are sessile and have a porous body


and choanocytes
Sponges, phylum Porifera
Live in both fresh and marine waters
Lack true tissues and organs

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Sponges are suspension feeders


Capturing food particles suspended in the
water that passes through their body
5 Choanocytes. The spongocoel
is lined with feeding cells called
choanocytes. By beating flagella,
the choanocytes create a current that
draws water in through the porocytes.

Azure vase sponge (Callyspongia


plicifera)
4 Spongocoel. Water
passing through porocytes
enters a cavity called the
spongocoel.
3 Porocytes. Water enters
the epidermis through
channels formed by
porocytes, doughnut-shaped
cells that span the body wall.
2 Epidermis. The outer
layer consists of tightly
packed epidermal cells.

Figure 33.4

1 Mesohyl. The wall of this


simple sponge consists of
two layers of cells separated
by a gelatinous matrix, the
mesohyl (middle matter).

Flagellum
Collar

Food particles
in mucus

Choanocyte

Osculum

Phagocytosis of
food particles

Spicules

Water
flow

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Amoebocyte

6 The movement of the choanocyte


flagella also draws water through its
collar of fingerlike projections. Food
particles are trapped in the mucus
coating the projections, engulfed by
phagocytosis, and either digested or
transferred to amoebocytes.
7 Amoebocyte. Amoebocytes
transport nutrients to other cells of
the sponge body and also produce
materials for skeletal fibers (spicules).

Choanocytes, flagellated collar cells


Generate a water current through the sponge
and ingest suspended food

Most sponges are hermaphrodites


Meaning that each individual functions as both
male and female

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Concept 33.2: Cnidarians have radial


symmetry, a gastrovascular cavity, and
cnidocytes
All animals except sponges
Belong to the clade Eumetazoa, the animals
with true tissues

Phylum Cnidaria
Is one of the oldest groups in this clade

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Cnidarians
Have diversified into a wide range of both
sessile and floating forms including jellies,
corals, and hydras
But still exhibit a relatively simple diploblastic,
radial body plan

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The basic body plan of a cnidarian


Is a sac with a central digestive compartment,
the gastrovascular cavity

A single opening
Functions as both mouth and anus

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There are two variations on this body plan


The sessile polyp and the floating medusa
Polyp

Medusa

Mouth/anus
Tentacle
Gastrovascular
cavity
Gastrodermis
Mesoglea

Body
stalk

Epidermis

Tentacle
Mouth/anus
Figure 33.5

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Cnidarians are carnivores


That use tentacles to capture prey

The tentacles are armed with cnidocytes


Unique cells that function in defense and the
capture of prey
Prey
Tentacle

Trigger

Nematocyst

Figure 33.6

Coiled thread

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Discharge
Of thread

Cnidocyte

The phylum Cnidaria is divided into four major


classes

Table 33.1
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Hydrozoa, Scyphozoa, Cubozoa, and


Anthozoa

(a) These colonial polyps are members of


class Hydrozoa.

(b) Many species of jellies (class


Scyphozoa), including the
species pictured here, are
bioluminescent. The largest
scyphozoans have tentacles
more than 100 m long
dangling from a bell-shaped
body up to 2 m in diameter.

Figure 33.7ad

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(c) The sea wasp (Chironex


fleckeri) is a member of
class Cubozoa. Its poison,
which can subdue fish and
other large prey, is more
potent than cobra venom.

(d) Sea anemones and other


members of class Anthozoa
exist only as polyps.

Hydrozoans
Most hydrozoans
Alternate between polyp and medusa forms
2 Some of the colonys
polyps, equipped with tentacles,
are specialized for feeding.

3 Other polyps, specialized


for reproduction, lack
tentacles and produce tiny
medusae by asexual budding.

4 The medusae
swim off, grow, and
reproduce sexually.

Reproductive
polyp
1 A colony of
interconnected
polyps (inset,
LM) results
from asexual
reproduction
by budding.

Feeding
polyp
Medusa
bud

MEIOSIS
Gonad
Medusa
SEXUAL
REPRODUCTION

Sperm

Egg

ASEXUAL
REPRODUCTION
(BUDDING)

Portion of
a colony
of polyps

FERTILIZATION
Zygote
Developing
polyp
Mature
polyp
Planula
(larva)

Figure 33.8

1 mm

6 The planula eventually settles


and develops into a new polyp.

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5 The zygote develops into a


solid ciliated larva called a planula.

Key
Haploid (n)
Diploid (2n)

Scyphozoans
In the class Scyphozoa
Jellies (medusae) are the prevalent form of the
life cycle

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Cubozoans
In the class Cubozoa, which includes box
jellies and sea wasps
The medusa is box-shaped and has complex
eyes

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Anthozoans
Class Anthozoa includes the corals and sea
anemones
Which occur only as polyps

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Concept 33.3: Most animals have bilateral


symmetry
The vast majority of animal species belong to
the clade Bilateria
Which consists of animals with bilateral
symmetry and triploblastic development

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Flatworms
Members of phylum Platyhelminthes
Live in marine, freshwater, and damp terrestrial
habitats
Are flattened dorsoventrally and have a
gastrovascular cavity

Although flatworms undergo triploblastic


development
They are acoelomates

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Flatworms are divided into four classes

Table 33.2
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Turbellarian
Turbellarians
Are nearly all free-living and mostly marine

Figure 33.9
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The best-known turbellarians, commonly called


planarians
Have light-sensitive eyespots and centralized
nerve nets
Pharynx. The mouth is at the
tip of a muscular pharynx that
extends from the animals
ventral side. Digestive juices
are spilled onto prey, and the
pharynx sucks small pieces of
food into the gastrovascular
cavity, where digestion continues.

Digestion is completed within


the cells lining the gastrovascular cavity, which has
three branches, each with
fine subbranches that provide an extensive surface area.
Undigested wastes
are egested
through the mouth.

Gastrovascular
cavity

Eyespots

Figure 33.10

Ganglia. Located at the anterior end


of the worm, near the main sources
of sensory input, is a pair of ganglia,
dense clusters of nerve cells.

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Ventral nerve cords. From


the ganglia, a pair of
ventral nerve cords runs
the length of the body.

Monogeneans and Trematode


Monogeneans and trematodes
Live as parasites in or on other animals
Parasitize a wide range of hosts

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Trematodes that parasitize humans


Spend part of their lives in snail hosts
1 Mature flukes live in the blood vessels of the human
intestine. A female fluke fits into a groove running
the length of the larger males body, as shown in
the light micrograph at right.

Male

Female

1 mm

5 These larvae penetrate


the skin and blood
vessels of humans
working in irrigated
fields contaminated
with infected human
feces.

2 Blood flukes reproduce


sexually in the human host.
The fertilized eggs exit the
host in feces.

3 The eggs develop in


water into ciliated
larvae. These larvae
infect snails, the
intermediate hosts.

Figure 33.11

4 Asexual reproduction
within a snail results in
another type of motile
larva, which escapes from
the snail host.

Snail host

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Most monogeneans
Are parasites of fish

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Tapeworm
Tapeworms
Are also parasitic and lack a digestive system

Proglottids with
reproductive structures
200 m

Scolex

Figure 33.12
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Hooks
Sucker

Rotifers
Rotifers, phylum Rotifera
Are tiny animals that inhabit fresh water, the
ocean, and damp soil

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Rotifers are smaller than many protists


But are truly multicellular and have specialized
organ systems

Figure 33.13
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0.1 mm

Rotifers have an alimentary canal


A digestive tube with a separate mouth and
anus that lies within a fluid-filled pseudocoelom

Rotifers reproduce by parthenogenesis


In which females produce more females from
unfertilized eggs

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Lophophorates: Ectoprocts, Phoronids, and Brachiopods

Lophophorates have a lophophore


A horseshoe-shaped, suspension-feeding
organ bearing ciliated tentacles

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Ectoprocts
Are colonial animals that superficially resemble
plants
Lophophore

(a) Ectoprocts, such as this sea


mat (Membranipora
membranacea), are colonial
Figure 33.14a
lophophorates.
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Phoronids
Are tube-dwelling marine worms ranging from
1 mm to 50 cm in length
Lophophore

(b) In phoronids such as


Phoronis hippocrepia, the
lophophore and mouth
are at one end of an
Figure 33.14b
elongated trunk.
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Brachiopods superficially resemble clams and


other hinge-shelled molluscs
But the two halves of the shell are dorsal and
ventral rather than lateral, as in clams

Lophophore

(c) Brachiopods have a hinged shell.


The two parts of the shell are
Figure 33.14c
dorsal and ventral.
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Nemerteans
Members of phylum Nemertea
Are commonly called proboscis worms or
ribbon worms

Figure 33.15
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The nemerteans unique proboscis


Is used for defense and prey capture
Is extended by a fluid-filled sac

Nemerteans also have a closed circulatory


system
In which the blood is contained in vessels
distinct from fluid in the body cavity

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Concept 33.4: Molluscs have a muscular foot, a


visceral mass, and a mantle
Phylum Mollusca
Includes snails and slugs, oysters and clams, and
octopuses and squids

Most molluscs are marine


Though some inhabit fresh water and some are
terrestrial

Molluscs are soft-bodied animals


But most are protected by a hard shell
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All molluscs have a similar body plan with three


main parts
A muscular foot
A visceral mass
A mantle

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Nephridium. Excretory organs


called nephridia remove metabolic
wastes from the hemolymph.

Heart. Most molluscs have an open circulatory


system. The dorsally located heart pumps
circulatory fluid called hemolymph through arteries
into sinuses (body spaces). The organs of the
mollusc are thus continually bathed in hemolymph.
The long digestive tract is
coiled in the visceral mass.

Visceral mass
Coelom

Intestine
Gonads

Mantle
Mantle
cavity

Stomach
Shell
Radula

Anus
The nervous
system consists
of a nerve ring
around the
esophagus, from
which nerve
cords extend.

Gill

Foot

Nerve
cords

Esophagus

Figure 33.16
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Mouth

Mouth

Radula. The mouth


region in many
mollusc species
contains a rasp-like
feeding organ
called a radula. This
belt of backwardcurved teeth slides
back and forth,
scraping and
scooping like a
backhoe.

Most molluscs have separate sexes


With gonads located in the visceral mass

The life cycle of many molluscs


Includes a ciliated larval stage called a
trochophore

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There are four major classes of molluscs

Table 33.3
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Chitons
Class Polyplacophora is composed of the
chitons
Oval-shaped marine animals encased in an
armor of eight dorsal plates

Figure 33.17
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Gastropods
About three-quarters of all living species of
molluscs
Belong to class Gastropoda

(a) A land snail

Figure 33.18a, b

(b) A sea slug. Nudibranchs, or sea slugs, lost their shell


during their evolution.

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Most gastropods
Are marine, but there are also many
freshwater and terrestrial species
Possess a single, spiraled shell

Slugs lack a shell


Or have a reduced shell

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The most distinctive characteristic of this class


Is a developmental process known as torsion,
which causes the animals anus and mantle to
end up above its head
Stomach
Mantle
cavity
Anus

Mouth

Figure 33.19

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Intestine

Bivalves
Molluscs of class Bivalvia
Include many species of clams, oysters,
mussels, and scallops
Have a shell divided into two halves

Figure 33.20
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The mantle cavity of a bivalve


Contains gills that are used for feeding as well
as gas exchange
Mantle

Hinge area

Gut

Coelom
Heart

Shell

Adductor
muscle

Mouth

Anus
Excurrent
siphon

Palp
Water
flow

Foot

Figure 33.21

Mantle
cavity

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Gill

Incurrent
siphon

Cephalopods
Class Cephalopoda includes squids and
octopuses
Carnivores with beak-like jaws surrounded by
tentacles of their modified foot

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Most octopuses
Creep along the sea floor in search of prey

Figure 33.22a

(a) Octopuses are considered among the


most intelligent invertebrates.

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Squids use their siphon


To fire a jet of water, which allows them to
swim very quickly

Figure 33.22b

(b) Squids are speedy carnivores with


beaklike jaws and well-developed eyes.

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`
One small group of shelled cephalopods
The nautiluses, survives today

Figure 33.22c

(c) Chambered nautiluses are the only living


cephalopods with an external shell.

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Concept 33.5: Annelids are segmented worms


Annelids
Have bodies composed of a series of fused
rings

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The phylum Annelida is divided into three


classes

Table 33.4
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Oligochaetes
Oligochaetes (class Oligochaeta)
Are named for their relatively sparse chaetae,
or bristles made of chitin
Include the earthworms and a variety of
aquatic species

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Earthworms eat their way through the soil,


extracting nutrients as the soil moves through
the alimentary canal
Which helps till the earth, making earthworms
valuable to farmers

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Anatomy of an earthworm
Coelom. The coelom
of the earthworm is
partitioned by septa.

Each segment is surrounded by longitudinal muscle, which in


turn is surrounded by circular muscle. Earthworms coordinate
the contraction of these two sets of muscles to move (see
Figure 49.25). These muscles work against the noncompressible
coelomic fluid, which acts as a hydrostatic skeleton.

Epidermis

Septum
(partition
between
segments)

Circular
muscle

Many of the internal


structures are repeated
within each segment of
the earthworm.

Chaetae. Each segment


has four pairs of
chaetae, bristles that
provide traction for
burrowing.

Longitudinal
muscle
Dorsal
vessel

Anus

Intestine

Nerve
cords
Cerebral ganglia. The
earthworm nervous system
features a brain-like pair of
cerebral ganglia above and
in front of the pharynx. A ring
of nerves around the pharynx
connects to a subpharyngeal
ganglion, from which a fused
pair of nerve cords runs
posteriorly.

Cuticle

Metanephridium. Each
segment of the worm
contains a pair of
excretory tubes, called
metanephridia, with
ciliated funnels, called
nephrostomes. The
metanephridia remove
wastes from the blood
and coelomic fluid
through exterior pores.

Nephrostome
Pharynx

Tiny blood vessels are


abundant in the earthworms
skin, which functions as its
respiratory organ. The blood
contains oxygen-carrying
hemoglobin.

Ventral
vessel
Clitellum
Esophagus

Metanephridium

Crop

Giant Australian earthworm

Intestine
Gizzard

Mouth
Subpharyngeal
ganglion

Table 33.23

The circulatory system, a network of vessels,


is closed. The dorsal and ventral vessels are linked
by segmental pairs of vessels. The dorsal vessel
and five pairs of vessels that circle the esophagus
of an earthworm are muscular and pump blood
through the circulatory system.

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Ventral nerve cords with segmental ganglia.


The nerve cords penetrate the septa and run
the length of the animal, as do the digestive
tract and longitudinal blood vessels.

Polychaetes
Members of class Polychaeta
Possess paddlelike parapodia that function as
gills and aid in locomotion

Parapodia

Figure 33.24
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Leeches
Members of class Hirudinea
Are blood-sucking parasites, such as leeches

Figure 33.25
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Concept 33.6: Nematodes are nonsegmented


pseudocoelomates covered by a tough cuticle
Among the most widespread of all animals,
nematodes, or roundworms
Are found in most aquatic habitats, in the soil,
in moist tissues of plants, and in the body
fluids and tissues of animals

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The cylindrical bodies of nematodes (phylum


Nematoda)
Are covered by a tough coat called a cuticle

25 m
Figure 33.26
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Some species of nematodes


Are important parasites of plants and animals
Encysted juveniles

Figure 33.27
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Muscle tissue

50 m

Concept 33.7: Arthropods are segmented


coelomates that have an exoskeleton and
jointed appendages
Two out of every three known species of
animals are arthropods
Members of the phylum Arthropoda
Are found in nearly all habitats of the
biosphere

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General Characteristics of Arthropods


The diversity and success of arthropods
Are largely related to their segmentation, hard
exoskeleton, and jointed appendages

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Early arthropods, such as trilobites


Showed little variation from segment to
segment

Figure 33.28
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As arthropods evolved
The segments fused, and the appendages
became more specialized

The appendages of some living arthropods


Are modified for many different functions
Cephalothorax Abdomen

Antennae
(sensory
reception)

Head Thorax

Swimming
appendages

Walking legs

Figure 33.29

Pincer (defense) Mouthparts (feeding)

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The body of an arthropod


Is completely covered by the cuticle, an
exoskeleton made of chitin

When an arthropod grows


It molts its exoskeleton in a process called
ecdysis

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Arthropods have an open circulatory system


In which fluid called hemolymph is circulated
into the spaces surrounding the tissues and
organs

A variety of organs specialized for gas


exchange
Have evolved in arthropods

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Molecular evidence now suggests


That living arthropods consist of four major
lineages that diverged early in the evolution of
the phylum

Table 33.5
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Cheliceriforms
Cheliceriforms, subphylum Cheliceriformes
Are named for clawlike feeding appendages
called chelicerae
Include spiders, ticks, mites, scorpions, and
horseshoe crabs

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Most of the marine cheliceriforms are extinct


But some species survive today, including the
horseshoe crabs

Figure 33.30
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Most modern cheliceriforms are arachnids


A group that includes spiders, scorpions, ticks,
and mites
50 m

(a) Scorpions have pedipalps that are pincers (b) Dust mites are ubiquitous scavengers in (c) Web-building spiders are generally
specialized for defense and the capture of
human dwellings but are harmless except
most active during the daytime.
food. The tip of the tail bears a poisonous
to those people who are allergic to them
stinger.
(colorized SEM).

Figure 33.31ac
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Arachnids have an abdomen and a


cephalothorax
Which has six pairs of appendages, the most
anterior of which are the chelicerae
Digestive
gland

Intestine

Stomach
Heart

Brain
Eyes
Poison
gland

Ovary

Anus
Book lung
Spinnerets
Figure 33.32

Silk gland

Sperm
Gonopore
(exit for eggs) receptacle

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Chelicera

Pedipalp

Myriapods
Subphylum Myriapoda
Includes millipedes and centipedes

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Millipedes, class Diplopoda


Have a large number of legs

Each trunk segment


Has two pairs of legs

Figure 33.33
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Centipedes, class Chilopoda


Are carnivores with jaw-like mandibles
Have one pair of legs per trunk segment

Figure 33.34
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Insects
Subphylum Hexapoda, insects and their
relatives
Are more species-rich than all other forms of
life combined
Live in almost every terrestrial habitat and in
fresh water

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The internal anatomy of an insect


Includes several complex organ systems
The insect body has three regions: head,
thorax, and abdomen. The segmentation
of the thorax and abdomen are obvious,
but the segments that form the head are fused.
Abdomen

Thorax

Head
Compound eye
Antennae

Figure 33.35

Malpighian tubules.
Anus
Metabolic wastes are
removed from the
Vagina
hemolymph by excretory
organs called Malpighian
tubules, which are outpocketings of the
digestive tract.
Tracheal tubes. Gas exchange in insects is
accomplished by a tracheal system of branched,
chitin-lined tubes that infiltrate the body and
carry oxygen directly to cells. The tracheal
system opens to the outside of the body
through spiracles, pores that can control air
flow and water loss by opening or closing.

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Heart. The
insect heart
drives hemolymph
through an
open circulatory
system.
Ovary

Nerve cords. The insect


nervous system
consists of a pair of
ventral nerve cords
with several
segmental ganglia.

Cerebral ganglion. The two nerve


cords meet in the head, where the
ganglia of several anterior segments
are fused into a cerebral ganglion
(brain). The antennae, eyes, and
other sense organs are concentrated
on the head.

Dorsal
artery

Crop

Insect mouthparts are formed from


several pairs of modified appendages.
The mouthparts include mandibles,
which grasshoppers use for chewing.
In other insects, mouthparts are
specialized for lapping, piercing, or
sucking.

Flight is obviously one key to the great success


of insects
An animal that can fly
Can escape predators, find food, and disperse
to new habitats much faster than organisms
that can only crawl

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Many insects
Undergo metamorphosis during their
development

In incomplete metamorphosis, the young,


called nymphs
Resemble adults but are smaller and go
through a series of molts until they reach full
size

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Insects with complete metamorphosis


Have larval stages specialized for eating and
growing that are known by such names as
maggot, grub, or caterpillar

The larval stage


Looks entirely different from the adult stage

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Metamorphosis from the larval stage to the


adult stage
Occurs during a pupal stage

(a) Larva (caterpillar)

(b) Pupa

(c) Pupa

Figure 33.6ae
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(d) Emerging adult

(e) Adult

Insects are classified into about 26 orders


ORDER

Blattodea

Coleoptera

Dermaptera

Diptera

Hemiptera

Hymenoptera

Isoptera

APPROXIMATE
NUMBER OF
SPECIES
4,000

350,000

1,200

151,000

85,000

125,000

2,000

MAIN CHARACTERISTICS

EXAMPLES

Cockroaches have a dorsoventrally flattened body, with legs


modified for rapid running. Forewings, when present, are
leathery, whereas hind wings are fanlike. Fewer than 40 cockroach species live in houses; the rest exploit habitats ranging
from tropical forest floors to caves and deserts.

German
cockroach

Beetles comprise the most species-rich order of insects. They


have two pairs of wings, one of which is thick and leathery, the
other membranous. They have an armored exoskeleton and
mouthparts adapted for biting and chewing. Beetles undergo
complete metamorphosis.

Japanese
beetle

Earwigs are generally nocturnal scavengers. While some


species are wingless, others have two pairs of wings, one of
which is thick and leathery, the other membranous. Earwigs
have biting mouthparts and large posterior pincers. They undergo incomplete metamorphosis.

Dipterans have one pair of wings; the second pair has become
modified into balancing organs called halteres. Their head is
large and mobile; their mouthparts are adapted for sucking,
piercing, or lapping. Dipterans undergo complete metamorphosis. Flies and mosquitoes are among the best-known dipterans,
which live as scavengers, predators, and parasites.

Hemipterans are so-called true bugs, including bed bugs,


assassin bugs, and chinch bugs. (Insects in other orders are
sometimes erroneously called bugs.) Hemipterans have two
pairs of wings, one pair partly leathery, the other membranous.
They have piercing or sucking mouthparts and undergo
incomplete metamorphosis.

Ants, bees, and wasps are generally highly social insects. They
have two pairs of membranous wings, a mobile head, and
chewing or sucking mouthparts. The females of many species
have a posterior stinging organ. Hymenopterans undergo complete metamorphosis.

Termites are widespread social insects that produce enormous


colonies. It has been estimated that there are 700 kg of
termites for every person on Earth! Some termites have two
pairs of membranous wings, while others are wingless. They
feed on wood with the aid of microbial symbionts carried in
specialized chambers in their hindgut.

Figure 33.37
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Earwig

Horsefly

LeafFooted
bug

Cicada-killer wasp

Termite

Insects are classified into about 26 orders


ORDER

Lepidoptera

Odonata

APPROXIMATE
NUMBER OF
SPECIES
120,000

5,000

EXAMPLE

MAIN CHARACTERISTICS

Butterflies and moths are among the best-known insects. They


have two pairs of wings covered with tiny scales. To feed, they
uncoil a long proboscis. Most feed on nectar, but some species
feed on other substances, including animal blood or tears.

Swallowtail
butterfly

Dragonflies and damselflies have two pairs of large, membranous wings. They have an elongated abdomen, large, compound
eyes, and chewing mouthparts. They undergo incomplete metamorphosis and are active predators.
Dragonfly

Orthoptera

13,000

Grasshoppers, crickets, and their relatives are mostly herbivorous. They have large hind legs adapted for jumping, two
pairs of wings (one leathery, one membranous), and biting or
chewing mouthparts. Males commonly make courtship sounds
by rubbing together body parts, such as a ridge on their hind
leg. Orthopterans undergo incomplete metamorphosis.
Katydid

Phasmida

Phthiraptera

Siphonaptera

Thysanura

2,600

2,400

2,400

450

Stick insects and leaf insects are exquisite mimics of plants. The
eggs of some species even mimic seeds of the plants on which the
Insects live. Their body is cylindrical or flattened dorsoventrally.
They lack forewings but have fanlike hind wings. Their
mouthparts are adapted for biting or chewing.

Commonly called sucking lice, these insects spend their entire


life as an ectoparasite feeding on the hair or feathers of a single
host. Their legs, equipped with clawlike tarsi, are adapted for
clinging to their hosts. They lack wings and have reduced eyes.
Sucking lice undergo incomplete metamorphosis.

Fleas are bloodsucking ectoparasites on birds and mammals.


Their body is wingless and laterally compressed. Their legs are
modified for clinging to their hosts and for long-distance
jumping. They undergo complete metamorphosis.

Stick insect

Human
Body
louse

Flea

Silverfish are small, wingless insects with a flattened body and


reduced eyes. They live in leaf litter or under bark. They can also
infest buildings, where they can become pests.
Silverfish

Trichoptera

7,100

Figure 33.37
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The larvae of caddisflies live in streams, where they make houses


from sand grains, wood fragments, or other material held together by silk. Adults have two pairs of hairy wings and chewing
or lapping mouthparts. They undergo complete metamorphosis.
Caddisfly

Crustaceans
While arachnids and insects thrive on land
Crustaceans, for the most part, have remained
in marine and freshwater environments

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Crustaceans, subphylum Crustacea


Typically have biramous, branched,
appendages that are extensively specialized
for feeding and locomotion

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Decapods are all relatively large crustaceans


And include lobsters, crabs, crayfish, and
shrimp

(a) Ghost crabs (genus Ocypode) live on sandy ocean


beaches worldwide. Primarily nocturnal, they take
Figure 33.38a
shelter in burrows during the day.
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Planktonic crustaceans include many species


of copepods
Which are among the most numerous of all
animals

Figure 33.38b

(b) Planktonic crustaceans


known as krill are
consumed in vast
quantities by whales.

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Barnacles are a group of mostly sessile


crustaceans
Whose cuticle is hardened into a shell

Figure 33.38c

(c) The jointed appendages


projecting from the shells
of these barnacles capture
organisms and organic
particles suspended in
the water.

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Concept 33.8: Echinoderms and chordates are


deuterostomes
At first glance, sea stars and other
echinoderms, phylum Echinodermata
May seem to have little in common with
phylum Chordata, which includes the
vertebrates

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Chordates and echinoderms share


characteristics of deuterostomes
Radial cleavage
Development of the coelom from the
archenteron
Formation of the mouth at the end of the
embryo opposite the blastopore

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Echinoderms
Sea stars and most other echinoderms
Are slow-moving or sessile marine animals

A thin, bumpy or spiny skin


Covers an endoskeleton of hard calcareous
plates

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Unique to echinoderms is a water vascular


system
A network of hydraulic canals branching into
tube feet that function in locomotion, feeding,
and gas exchange
A short digestive tract runs from the
mouth on the bottom of the central
disk to the anus on top of the disk.

Central disk. The central


disk has a nerve ring and
nerve cords radiating from
the ring into the arms.

Digestive glands secrete


digestive juices and aid in
the absorption and storage
of nutrients.

Figure 33.39

Radial canal. The water vascular


system consists of a ring canal in the
central disk and five radial canals,
each running in a groove down the
entire length of an arm.

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Stomach
Anus

Spine
Gills

Ring
canal

Gonads

Radial
nerve

The surface of a sea star is


covered by spines that help
defend against predators, as
well as by small gills that
provide gas exchange.
Madreporite. Water can flow
in or out of the water vascular
system into the surrounding
water through the madreporite.

Ampulla
Podium
Tube
feet
Branching from each radial canal are hundreds of hollow, muscular tube
feet filled with fluid. Each tube foot consists of a bulb-like ampulla and
suckered podium (foot portion). When the ampulla squeezes, it forces
water into the podium and makes it expand. The podium then
contacts the substrate. When the muscles in the wall of the podium
contract, they force water back into the ampulla, making the podium
shorten and bend.

The radial anatomy of many echinoderms


Evolved secondarily from the bilateral
symmetry of ancestors

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Living echinoderms are divided into six classes

Table 33.6
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Sea Stars
Sea stars, class Asteroidea
Have multiple arms radiating from a central
disk

The undersurfaces of the arms


Bear tube feet, each of which can act like a
suction disk

Figure 33.40a

(a) A sea star (class Asteroidea)

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Brittle Stars
Brittle stars have a distinct central disk
And long, flexible arms

Figure 33.40b (b) A brittle star (class Ophiuroidea)

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Sea Urchins and Sand Dollars


Sea urchins and sand dollars have no arms
But they do have five rows of tube feet that
function in movement

Figure 33.40c (c) A sea urchin (class Echinoidea)

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Sea Lilies and Feather Stars


Sea lilies
Live attached to the substrate by a stalk

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Sea Cucumbers
Feather stars
Crawl about using their long, flexible arms

Figure 33.40d

(d) A feather star (class Crinoidea)

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Sea Cucumbers
Sea cucumbers
Upon first inspection do not look much like
other echinoderms
Lack spines, and their endoskeleton is much
reduced

Figure 33.40e

(e) A sea cucumber (class Holothuroidea)

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Sea Daisies
Sea daisies were discovered in 1986
And only two species are known

Figure 33.40f

(f) A sea daisy (class Concentricycloidea)

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Chordates
Chordates
Phylum Chordata
Consists of two subphyla of invertebrates as
well as the hagfishes and the vertebrates
Shares many features of embryonic
development with echinoderms

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A summary of animal phyla

Table 33.7
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