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

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

Deuterostomia

Bilateria

Eumetazoa

Ancestral colonial choanoflagellate

Figure 33.2
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Chordata

• Exploring invertebrate diversity
PORIFERA (5,500 species) CNIDARIA (10,000 species)

A sponge PLACOZOA (1 species)
0.5 mm

A jelly KINORHYNCHA (150 species)

250 µm

A placozoan (LM) A kinorhynch (LM) ROTIFERA (1,800 species) PLATYHELMINTHES (20,000 species)

A marine flatworm ECTOPROCTA (4,500 species)

A rotifer (LM) PHORONIDA (20 species)

Figure 33.3

Ectoprocts Phoronids

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

A brachiopod ACANTHOCEPHALA (1,100 species)
5 mm

A ribbon worm CTENOPHORA (100 species)

An acanthocephalan

A ctenophore, or comb jelly

MOLLUSCA (93,000 species)

ANNELIDA (16,500 species)

An octopus LORICIFERA (10 species)
50 µm

A marine annelid PRIAPULA (16 species)

Figure 33.3

A loriciferan (LM)

A priapulan

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• Exploring invertebrate diversity
NEMATODA (25,000 species) A roundworm CYCLIOPHORA (1 species)
100 µm

ARTHROPODA (1,000,000 + species)

A scorpion (an arachnid) TARDIGRADA (800 species)

100 µm

A cycliophoran (colorized SEM) Tardigrades (colorized SEM) ONYCHOPHORA (110 species) HEMICHORDATA (85 species)

An onychophoran

An acorn worm CHORDATA (52,000 species)

ECHINODERMATA (7,000 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, the choanocytes create a current that draws water in through the porocytes. Flagellum Collar Food particles in mucus Choanocyte

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. 1 Mesohyl. The wall of this simple sponge consists of two layers of cells separated by a gelatinous matrix, the mesohyl (“middle matter”).

Osculum

Phagocytosis of food particles

Amoebocyte

Spicules

Water flow

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).

Figure 33.4

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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 Tentacle Gastrovascular cavity Gastrodermis Mesoglea Body stalk Epidermis Medusa

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

Figure 33.6

Cnidocyte

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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

(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.

(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.

Figure 33.7a–d

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Hydrozoans • Most hydrozoans
– Alternate between polyp and medusa forms
2 Some of the colony’s polyps, equipped with tentacles, are specialized for feeding. 3 Other polyps, specialized for reproduction, lack tentacles and produce tiny medusae by asexual budding. 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 ASEXUAL REPRODUCTION (BUDDING) FERTILIZATION Zygote Developing polyp Mature polyp Planula (larva) 1 mm 6 The planula eventually settles and develops into a new polyp. 5 The zygote develops into a solid ciliated larva called a planula. Key Haploid (n) Diploid (2n) Egg Sperm 4 The medusae swim off, grow, and reproduce sexually.

Portion of a colony of polyps

Figure 33.8

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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
Pharynx. The mouth is at the tip of a muscular pharynx that extends from the animal’s 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.

Ventral nerve cords. From the ganglia, a pair of ventral nerve cords runs 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 the light micrograph at right. Male

Female

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

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

Figure 33.11

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 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

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 Mantle Mantle cavity Anus The nervous system consists of a nerve ring around the esophagus, from which nerve cords extend. Figure 33.16
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Intestine Gonads Stomach Shell Radula 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.

Mouth

Gill

Foot

Nerve cords

Esophagus

Mouth

• 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 animal’s anus and mantle to end up above its head Stomach
Mantle cavity Anus Intestine

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

Palp Foot Water flow Mantle cavity Gill Incurrent siphon

Figure 33.21

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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

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
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. Many of the internal structures are repeated within each segment of the earthworm. Longitudinal muscle Dorsal vessel Intestine Tiny blood vessels are abundant in the earthworm’s skin, which functions as its respiratory organ. The blood contains oxygen-carrying hemoglobin. Circular muscle Coelom. The coelom of the earthworm is partitioned by septa. Cuticle Septum (partition between segments) Epidermis 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. Anus

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

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. Nephrostome Pharynx

Ventral vessel Clitellum Esophagus Crop

Metanephridium

Giant Australian earthworm

Intestine Gizzard Subpharyngeal ganglion

Mouth

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.

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.

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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 Head Thorax Antennae (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
Digestive gland Intestine Stomach Heart Brain Eyes Ovary Poison gland

Anus Book lung Spinnerets Figure 33.32 Silk gland Sperm Gonopore (exit for eggs) receptacle Chelicera Pedipalp

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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 of the thorax and abdomen are obvious, but the segments that form the head are fused. Abdomen Thorax Head Compound eye Antennae Heart. The insect heart drives hemolymph through an open circulatory system. Ovary 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

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.

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

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.

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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
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(e) Adult

• Insects are classified into about 26 orders
ORDER APPROXIMATE NUMBER OF SPECIES 4,000 MAIN CHARACTERISTICS EXAMPLES Blattodea 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 cockroach species live in houses; the rest exploit habitats ranging from tropical forest floors to caves and deserts. German cockroach

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 other membranous. They have an armored exoskeleton and mouthparts adapted for biting and chewing. Beetles undergo complete metamorphosis.

Japanese beetle

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 undergo 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 metamorphosis. Flies and mosquitoes are among the best-known dipterans, which live as scavengers, predators, and parasites.

Horsefly

Hemiptera

85,000

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.

LeafFooted bug

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 complete 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
ORDER APPROXIMATE NUMBER OF SPECIES 120,000 MAIN CHARACTERISTICS EXAMPLE Lepidoptera 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

Odonata

5,000

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 relativ es 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

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 mouthparts are adapted for biting or chewing.

Stick insect

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 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.

Human Body louse

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 together 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

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. Spine Gills Central disk. The central disk has a nerve ring and nerve cords radiating from the ring into the arms. Stomach Anus 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.

Digestive glands secrete digestive juices and aid in the absorption and storage of nutrients.

Ring canal

Gonads

Radial nerve

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.

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.

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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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