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Chapter 33

Invertebrates

PowerPoint Lectures for


Biology, Seventh Edition
Neil Campbell and Jane Reece

Lectures by Chris Romero


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• 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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• A review of animal phylogeny

Cnidaria
Porifera

Echinodermata

Chordata
Mollusca, and Annelida)
Other bilaterians (including
Nematoda, Arthropoda,
Deuterostomia

Bilateria

Eumetazoa

Ancestral colonial
choanoflagellate
Figure 33.2
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• Exploring invertebrate diversity

PORIFERA (5,500 species) CNIDARIA (10,000 species)

A sponge A jelly
PLACOZOA (1 species) KINORHYNCHA (150 species)
0.5 mm

250 µm

A placozoan (LM) A kinorhynch (LM)


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

A marine flatworm A rotifer (LM)


ECTOPROCTA (4,500 species) PHORONIDA (20 species)

Figure 33.3 Ectoprocts Phoronids

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• Exploring invertebrate diversity
BRACHIOPODA (335 species) NEMERTEA (900 species)

A brachiopod A ribbon worm


ACANTHOCEPHALA (1,100 species) 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


LORICIFERA (10 species) PRIAPULA (16 species)
50 µm

Figure 33.3 A loriciferan (LM) A priapulan

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• Exploring invertebrate diversity
NEMATODA (25,000 species) ARTHROPODA (1,000,000 + species)

A roundworm A scorpion (an arachnid)


CYCLIOPHORA (1 species) TARDIGRADA (800 species)
100 µm

100 µm

A cycliophoran (colorized SEM) Tardigrades (colorized SEM)

ONYCHOPHORA (110 species) HEMICHORDATA (85 species)

An onychophoran An acorn worm

ECHINODERMATA (7,000 species) CHORDATA (52,000 species)

Figure 33.3 A sea urchin A tunicate


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• 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, Flagellum
the choanocytes create a current that Food particles Choanocyte
draws water in through the porocytes. Collar in mucus

Azure vase sponge (Callyspongia Osculum


plicifera)
4 Spongocoel. Water
passing through porocytes
enters a cavity called the Phagocytosis of
spongocoel. food particles Amoebocyte
3 Porocytes. Water enters
the epidermis through 6 The movement of the choanocyte
channels formed by flagella also draws water through its
porocytes, doughnut-shaped Spicules collar of fingerlike projections. Food
cells that span the body wall. particles are trapped in the mucus
coating the projections, engulfed by
2 Epidermis. The outer
phagocytosis, and either digested or
layer consists of tightly
Water transferred to amoebocytes.
packed epidermal cells.
flow 7 Amoebocyte. Amoebocytes
1 Mesohyl. The wall of this transport nutrients to other cells of
simple sponge consists of the sponge body and also produce
two layers of cells separated materials for skeletal fibers (spicules).
by a gelatinous matrix, the
Figure 33.4 mesohyl (“middle matter”).

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• 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 Mouth/anus Medusa

Tentacle
Gastrovascular
cavity
Gastrodermis

Mesoglea
Epidermis
Body
stalk

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”

Discharge
Of thread
Nematocyst

Coiled thread Cnidocyte


Figure 33.6
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• The phylum Cnidaria is divided into four major
classes

Table 33.1
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– Hydrozoa, Scyphozoa, Cubozoa, and
Anthozoa

(b) Many species of jellies (class (c) The sea wasp (Chironex (d) Sea anemones and other
Scyphozoa), including the fleckeri) is a member of members of class Anthozoa
species pictured here, are class Cubozoa. Its poison, exist only as polyps.
bioluminescent. The largest which can subdue fish and
scyphozoans have tentacles other large prey, is more
more than 100 m long potent than cobra venom.
dangling from a bell-shaped
body up to 2 m in diameter.
(a) These colonial polyps are members of
class Hydrozoa.
Figure 33.7a–d

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Hydrozoans
• Most hydrozoans
– Alternate between polyp and medusa forms
3 Other polyps, specialized
2 Some of the colony’s for reproduction, lack 4 The medusae
polyps, equipped with tentacles, tentacles and produce tiny swim off, grow, and
are specialized for feeding. medusae by asexual budding. reproduce sexually.

Reproductive
polyp
Feeding
1 A colony of
polyp
interconnected
polyps (inset, Medusa
LM) results MEIOSIS
bud
from asexual Gonad
Medusa
reproduction
by budding. SEXUAL
Egg Sperm
REPRODUCTION

ASEXUAL
REPRODUCTION
Portion of (BUDDING)
a colony
of polyps FERTILIZATION

Zygote
Developing
polyp

Mature
polyp

Planula
(larva) Key

Haploid (n)
1 mm 6 The planula eventually settles 5 The zygote develops into a Diploid (2n)
Figure 33.8 and develops into a new polyp. solid ciliated larva called a planula.

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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
Digestion is completed within
the cells lining the gastro-
Pharynx. The mouth is at the vascular cavity, which has
tip of a muscular pharynx that three branches, each with
extends from the animal’s fine subbranches that pro-
ventral side. Digestive juices vide an extensive surface area.
are spilled onto prey, and the
pharynx sucks small pieces of Undigested wastes
food into the gastrovascular are egested
cavity, where digestion continues. through the mouth.

Gastrovascular
cavity

Eyespots

Ganglia. Located at the anterior end Ventral nerve cords. From


of the worm, near the main sources the ganglia, a pair of
of sensory input, is a pair of ganglia, ventral nerve cords runs
Figure 33.10 dense clusters of nerve cells. the length of the body.

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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 male’s body, as shown in
Male
the light micrograph at right.

Female

1 mm
5 These larvae penetrate
the skin and blood 2 Blood flukes reproduce
vessels of humans sexually in the human host.
working in irrigated The fertilized eggs exit the
fields contaminated host in feces.
with infected human
feces.
3 The eggs develop in
water into ciliated
larvae. These larvae
infect snails, the
intermediate hosts.
4 Asexual reproduction
within a snail results in
another type of motile
larva, which escapes from
the snail host. Snail host
Figure 33.11
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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 µm

Scolex Hooks
Sucker

Figure 33.12

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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

0.1 mm
Figure 33.13
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• 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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Heart. Most molluscs have an open circulatory
system. The dorsally located heart pumps
Nephridium. Excretory organs circulatory fluid called hemolymph through arteries
called nephridia remove metabolic into sinuses (body spaces). The organs of the
wastes from the hemolymph. mollusc are thus continually bathed in hemolymph.

Visceral mass The long digestive tract is


coiled in the visceral mass.
Coelom Intestine
Gonads
Mantle Radula. The mouth
Stomach
Mantle region in many
Shell
cavity Mouth mollusc species
Radula
Anus contains a rasp-like
feeding organ
The nervous Gill called a radula. This
system consists belt of backward-
of a nerve ring curved teeth slides
around the back and forth,
esophagus, from Foot Nerve Mouth
cords Esophagus scraping and
which nerve scooping like a
cords extend. backhoe.
Figure 33.16

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• 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

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


Figure 33.18a, b 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 animal’s anus and mantle to
end up above its head Stomach
Mantle Intestine
cavity

Anus

Mouth

Figure 33.19

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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
Hinge area Coelom
Mantle Gut
Heart
Shell Adductor
muscle
Mouth
Anus
Excurrent
siphon

Palp
Water
Foot flow
Mantle Incurrent
Figure 33.21 cavity Gill siphon

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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

(a) Octopuses are considered among the


Figure 33.22a most intelligent invertebrates.

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• Squids use their siphon
– To fire a jet of water, which allows them to
swim very quickly

(b) Squids are speedy carnivores with


Figure 33.22b beaklike jaws and well-developed eyes.

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

(c) Chambered nautiluses are the only living


Figure 33.22c 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
Each segment is surrounded by longitudinal muscle, which in Metanephridium. Each
of the earthworm is
turn is surrounded by circular muscle. Earthworms coordinate segment of the worm
partitioned by septa.
the contraction of these two sets of muscles to move (see contains a pair of
Figure 49.25). These muscles work against the noncompressible excretory tubes, called
coelomic fluid, which acts as a hydrostatic skeleton. metanephridia, with
Epidermis Cuticle
ciliated funnels, called
Circular Septum nephrostomes. The
Many of the internal metanephridia remove
muscle (partition
structures are repeated wastes from the blood
between
within each segment of and coelomic fluid
segments)
the earthworm. through exterior pores.

Longitudinal
Chaetae. Each segment Anus
muscle
has four pairs of Dorsal
chaetae, bristles that vessel
provide traction for
burrowing. Intestine
Tiny blood vessels are
abundant in the earthworm’s
skin, which functions as its
Nerve respiratory organ. The blood
Ventral
cords contains oxygen-carrying
vessel
Cerebral ganglia. The hemoglobin.
earthworm nervous system Nephrostome Clitellum
features a brain-like pair of Pharynx
cerebral ganglia above and Esophagus
Crop Metanephridium
in front of the pharynx. A ring
of nerves around the pharynx
Giant Australian earthworm
connects to a subpharyngeal Intestine
ganglion, from which a fused
pair of nerve cords runs
posteriorly. Gizzard
Mouth
Subpharyngeal
ganglion
Ventral nerve cords with segmental ganglia.
The circulatory system, a network of vessels,
The nerve cords penetrate the septa and run
is closed. The dorsal and ventral vessels are linked
the length of the animal, as do the digestive
by segmental pairs of vessels. The dorsal vessel
tract and longitudinal blood vessels.
and five pairs of vessels that circle the esophagus
of an earthworm are muscular and pump blood
Table 33.23 through the circulatory system.

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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 µm
Figure 33.26
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• Some species of nematodes
– Are important parasites of plants and animals
Encysted juveniles Muscle tissue 50 µm

Figure 33.27

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• 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
Head Thorax
(sensory
reception)

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 µm

(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 33.31a–c
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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
Intestine
Stomach
Digestive Brain
gland Heart

Eyes

Ovary Poison
gland

Anus
Book lung
Spinnerets
Gonopore Sperm Pedipalp
Chelicera
Figure 33.32 Silk gland (exit for eggs) receptacle
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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 Cerebral ganglion. The two nerve
of the thorax and abdomen are obvious, Heart. The cords meet in the head, where the
but the segments that form the head are fused. insect heart ganglia of several anterior segments
drives hemolymph are fused into a cerebral ganglion
Abdomen Thorax Head through an (brain). The antennae, eyes, and
Compound eye open circulatory other sense organs are concentrated
system. on the head.
Antennae
Ovary
Dorsal
artery Crop
Malpighian tubules. Anus
Metabolic wastes are
removed from the Vagina
hemolymph by excretory
organs called Malpighian
tubules, which are out-
pocketings of the
digestive tract.
Tracheal tubes. Gas exchange in insects is Nerve cords. The insect Insect mouthparts are formed from
accomplished by a tracheal system of branched, nervous system several pairs of modified appendages.
chitin-lined tubes that infiltrate the body and consists of a pair of The mouthparts include mandibles,
carry oxygen directly to cells. The tracheal ventral nerve cords which grasshoppers use for chewing.
system opens to the outside of the body with several In other insects, mouthparts are
through spiracles, pores that can control air segmental ganglia. specialized for lapping, piercing, or
Figure 33.35 flow and water loss by opening or closing. sucking.

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• 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
(d) Emerging adult

Figure 33.6a–e (e) Adult

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• Insects are classified into about 26 orders
APPROXIMATE
ORDER NUMBER OF MAIN CHARACTERISTICS EXAMPLES
SPECIES

Blattodea 4,000 Cockroaches have a dorsoventrally f lattened body, with legs


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

Coleoptera 350,000 Beetles comprise the most species-rich order of insects. They
have two pairs of wings, one of which is thick and leathery, the
Japanese
other membranous. They have an armored exoskeleton and
beetle
mouthparts adapted for biting and chewing. Beetles undergo
complete metamorphosis.

Dermaptera 1,200 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 un-
dergo incomplete metamorphosis. Earwig

Diptera 151,000 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 metamorpho- Horsefly
sis. Flies and mosquitoes are among the best-known dipterans,
which live as scavengers, predators, and parasites.

Hemiptera 85,000 Hemipterans are so-called “true bugs,” including bed bugs,
assassin bugs, and chinch bugs. (Insects in other orders are Leaf-
sometimes erroneously called bugs.) Hemipterans have two Footed
pairs of wings, one pair partly leathery, the other membranous. bug
They have piercing or sucking mouthparts and undergo
incomplete metamorphosis.

Hymenoptera 125,000 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 com-
plete metamorphosis.
Cicada-killer wasp

Isoptera 2,000 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. Termite

Figure 33.37
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• Insects are classified into about 26 orders
APPROXIMATE
ORDER NUMBER OF MAIN CHARACTERISTICS EXAMPLE
SPECIES

Lepidoptera 120,000 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

5,000 Dragonflies and damselflies have two pairs of large, membran-


Odonata
ous wings. They have an elongated abdomen, large, compound
eyes, and chewing mouthparts. They undergo incomplete meta-
morphosis and are active predators.
Dragonfly

Orthoptera 13,000 Grasshoppers, crickets, and their relativ es are mostly herbi-
vorous. 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 2,600 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 Stick insect
mouthparts are adapted for biting or chewing.

Phthiraptera 2,400 Commonly called sucking lice, these insects spend their entire
life as an ectoparasite feeding on the hair or feathers of a single
Human
host. Their legs, equipped with clawlike tarsi, are adapted for
Body
clinging to their hosts. They lack wings and have reduced eyes.
louse
Sucking lice undergo incomplete metamorphosis.

Siphonaptera 2,400 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.
Flea

Thysanura 450 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 The larvae of caddisflies live in streams, where they make houses
from sand grains, wood fragments, or other material held to-
gether by silk. Adults have two pairs of hairy wings and chewing
or lapping mouthparts. They undergo complete metamorphosis.
Caddisfly
Figure 33.37
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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

(b) Planktonic crustaceans


known as krill are
consumed in vast
Figure 33.38b quantities by whales.
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• Barnacles are a group of mostly sessile
crustaceans
– Whose cuticle is hardened into a shell

(c)The jointed appendages


projecting from the shells
of these barnacles capture
organisms and organic
particles suspended in
Figure 33.38c 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.
The surface of a sea star is
Central disk. The central Spine covered by spines that help
Stomach
disk has a nerve ring and defend against predators, as
Anus
nerve cords radiating from well as by small gills that
the ring into the arms. Gills
provide gas exchange.

Madreporite. Water can flow


in or out of the water vascular
Radial
Digestive glands secrete system into the surrounding
Gonads nerve
digestive juices and aid in Ring water through the madreporite.
the absorption and storage canal Ampulla
of nutrients. Podium
Tube
feet
Radial canal. The water vascular Branching from each radial canal are hundreds of hollow, muscular tube
system consists of a ring canal in the feet filled with fluid. Each tube foot consists of a bulb-like ampulla and
central disk and five radial canals, suckered podium (foot portion). When the ampulla squeezes, it forces
each running in a groove down the water into the podium and makes it expand. The podium then
entire length of an arm. contacts the substrate. When the muscles in the wall of the podium
Figure 33.39 contract, they force water back into the ampulla, making the podium
shorten and bend.
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• 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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