Teresa Audesirk • Gerald Audesirk • Bruce E. Byers Biology: Life on Earth Eighth Edition Lecture for Chapter 23 Animal Diversity I: Invertebrates Copyright © 2008 Pearson Prentice Hall, Inc. Chapter 23 Outline • 23.1 What Are the Key Features of Animals? p. 442 • 23.2 Which Anatomical Features Mark Branch Points on the Animal Evolutionary Tree? p. 442 • 23.3 What Are the Major Animal Phyla? p. 445 Section 23.1 Outline • 23.1 What Are the Key Features of Animals? • Animals possess all of the following characteristics – Multicellularity – Heterotrophic – Most reproduce sexually – Cells lack a cell wall – Are motile at some point in life cycle – Are able to respond rapidly to external stimuli Section 23.2 Outline • 23.2 Which Anatomical Features Mark Branch Points on the Animal Evolutionary Tree? – Lack of Tissues Separates Sponges from All Other Animals – Animals with Tissues Exhibit Either Radial or Bilateral Symmetry – Most Bilateral Animals Have Body Cavities Animal Evolution • Most evolutionists believe animal phyla currently populating the Earth were present by the Cambrian period (544 million years ago) • The scarcity of pre-Cambrian fossils led systematists to search for clues about the evolutionary history of animals by examining features of – Anatomy – Embryological development – DNA sequences Animal Evolution • It is believed certain features represent evolutionary milestones – The appearance of tissues – The appearance of body symmetry – Protostome and deuterostome development • These features mark major branching points on the animal evolutionary tree The Appearance of Tissues • Tissues are groups of similar cells that carry out a specific function (e.g. muscle) • The ‘earliest’ animals had no tissues • Sponges are the only ‘modern-day’ animals that lack tissues – Individual cells may be specialized, but they act independently • Evolutionists believe sponges & all remaining tissue-containing phyla arose from an ancient common ancestor without tissues The Appearance of Body Symmetry • Symmetrical animals have an upper (dorsal) surface and a lower (ventral) surface • Animals with tissues exhibit either radial or bilateral symmetry Radial Symmetry • Animals with radial symmetry can be divided into roughly equal parts by any plane that passes through the central axis • Note they can be divided into more than two parts (more than two halves) FIGURE 23-2a Body symmetry and cephalization (a) Animals with radial symmetry lack a welldefined head. Any plane that passes through the central axis divides the body into mirror-image halves. Animals with Radial Symmetry • Have two embryonic tissue (germ) layers – Ectoderm (outer layer which covers the body, lines its inner cavities, and forms the nervous system) – Endoderm (inner layer which lines most hollow organs) • Tend to be either sessile (fixed to one spot) or drift around on currents – e.g. cnidarians, ctenophors Bilateral Symmetry • Animals with bilateral symmetry can be divided into mirror-image halves only along one plane that runs down the midline FIGURE 23-2b Body symmetry and cephalization (b) Animals with bilateral symmetry have an anterior head end and a posterior tail end. The body can be split into two mirror-image halves only along a particular plane that runs down the midline. Animals with Bilateral Symmetry • Have an additional germ layer – Mesoderm (layer sandwiched between ectoderm and endoderm) • Forms muscle • Forms circulatory and skeletal systems when present Animals with Bilateral Symmetry • Exhibit cephalization (the concentration of sensory organs and a brain in a welldefined head) with definite anterior and posterior regions • Tend to move forward through environment – e.g. flatworms, roundworms, arthropods, annelids, mollusks, echinoderms, chordates Body Cavities • Most bilateral animals have a body cavity • Body cavities serve a variety of functions • Act as a skeleton (provides support for the body and a framework against which muscles can act) • Form a protective buffer between internal organs and the outside world • Allow organs to move independently of the body wall Body Cavities • Coelomate animals possess a coelom (a fluid-filled body cavity that is completely lined with mesoderm) – e.g. annelids, arthropods, mollusks, echinoderms, chordates Body Cavities • Pseudocoelomate animals possess a pseudocoelom (a fluid-filled body cavity that is not completely lined with mesoderm) – e.g. nematodes (roundworms) Body Cavities • Acoelomate animals lack a body cavity – e.g. flatworms Section 23.3 Outline • 23.3 What Are the Major Animal Phyla? – Sponges Have a Simple Body Plan – Cnidarians Are Well-Armed Predators – Flatworms Have Organs but Lack Respiratory and Circulatory Systems – Annelids Are Composed of Identical Segments – Most Mollusks Have Shells – Arthropods Are the Dominant Animals on Earth? Section 22.3 Outline • 23.3 What Are the Major Animal Phyla? (continued) – Roundworms Are Abundant and Mostly Tiny – Echinoderms Have a Calcium Carbonate Skeleton – The Chordates Include the Vertebrates Major Animal Phyla • Animals probably originated from ancestral colonial protists • Present day biologists recognize about 27 phyla of animals, some of which are summarized in Table 23-1, pp. 446-7 Invertebrates and Vertebrates • Most animals are invertebrates (lack a vertebral column) • Less than 3% of all known animals are vertebrates (possess a vertebral column) 3.2: The Sponges • • • • Phylum Porifera Asymmetrical body plan Lack true tissues and organs Body perforated by tiny pores FIGURE 23-5 The body plan of sponges Water enters through numerous tiny pores in the sponge body and exits through oscula. Microscopic food particles are filtered from the water. The Sponges • Three major types of cells – Epithelial cells (cover outer body surface) • Some are modified into pore cells (regulate flow of water through pores) – Collar cells (flagellated cells that maintain water flow through the sponge) – Amoeboid cells (motile cells that digest and distribute nutrients, produce reproductive cells, and secrete spicules) The Sponges • Water flows through the body – Water enters through pores and travels through canals • Oxygen is extracted • Nutrients are removed and digested intracellularly • Wastes are released – Water exits through large openings called oscula The Sponges • Internal skeleton made of spicules – Spicules are composed of calcium carbonate, silica, or protein • Aquatic (mostly marine) • Adults usually sessile; larvae are motile The Sponges • May reproduce – Asexually by budding (adult produces a bud that breaks off and becomes independent) – Sexually through fusion of sperm and eggs • Larvae develop in body of adult and escape through oscula • Larvae are dispersed by water currents • Come in a wide variety of sizes, shapes, and colors FIGURE 23-4a The diversity of sponges Sponges come in a wide variety of sizes, shapes, and colors. Some, such as (a) this fire sponge, grow in a free-form pattern over undersea rocks. FIGURE 23-4b The diversity of sponges (b) Tiny appendages attach this tubular sponge to rocks FIGURE 23-4b The diversity of sponges (c) this reef sponge with flared tubular openings attaches to a coral reef. 3.3: The Cnidarians • Phylum Cnidaria • Include jellyfish, sea anemones, corals, and hydrozoans FIGURE 23-6a Cnidarian diversity (a) A red-spotted anemone spreads its tentacles to capture prey. FIGURE 23-6b Cnidarian diversity (b) A small medusa. FIGURE 23-6c Cnidarian diversity: A close-up of coral reveals bright yellow polyps in various stages of tentacle extension. At the lower right, areas where the coral has died expose the calcium carbonate skeleton that supports the polyps and forms the reef. A strikingly patterned crab (an arthropod) sits atop the coral, holding tiny white anemones in its claws. Their stinging tentacles help protect the crab. FIGURE 23-6d Cnidarian diversity (d) A sea wasp, a cnidarian whose stinging cells contain one of the most toxic of all known venom The Cnidarians • Radial symmetry • Develop from two embryonic germ layers: ectoderm and endoderm • Jellylike mesoglea forms between ectoderm and endoderm • Two basic body plans – Polyp (tubular body; sessile) – Medusa (bell-shaped body; free-floating) FIGURE 23-7 Polyp and medusa (a) The polyp form is seen in hydra (see Fig. 23-8), sea anemones (Fig. 23-6a), and the individual polyps within a coral (Fig. 23-6c). (b) The medusa form, seen in the jellyfish (Fig. 23-6b), resembles an inverted polyp. The Cnidarians • Possess tissues – Contractile tissue (acts like muscle) – Nerve net (composed of nerve cells) • Lack true organs • Have tentacles equipped with cnidocytes (specialized cells that function in defense and the capture of prey) Cnidocytes • Contain a finely coiled filament that is explosively expelled when the trigger is touched – Some filaments inject poison into the prey – Others either stick to or entangle small prey • Venom of some can cause extreme pain or death in humans FIGURE 23-8 Cnidarian weaponry: the cnidocyte At the slightest touch to the trigger of a special structure in their cnidocytes, cnidarians, such as this hydra, violently expel a poisoned filament. Cnidarians • Have a gastrovascular cavity (sac-like digestive chamber with a single opening; serves as both mouth and anus) – Tentacles force prey through opening into the gastrovascular cavity – Undigested wastes are expelled from opening when digestion is completed • Aquatic (mostly marine) Corals • Form reefs in warm tropical waters – Colonial polyps secrete a hard external skeleton of calcium carbonate – Skeleton remains after polyp dies – New polyps build on skeletal remnants of earlier generations • Reefs provide undersea habitats that support a wealth of diversity 3.4: The Flatworms • Phylum Platyhelminthes • Many species are parasitic • Non-parasitic, free-living flatworms inhabit aquatic, marine, and moist terrestrial habitats FIGURE 23-9a Flatworm diversity (a) This fluke is an example of a parasitic flatworm. FIGURE 23-9b Flatworm diversity (b) Eyespots are clearly visible in the head of this freeliving, freshwater flatworm. FIGURE 23-9c Flatworm diversity (c) Many of the flatworms that inhabit tropical coral reefs are brightly colored. The Flatworms • Bilateral symmetry • Acoelomate • Distinct head with sensory organs – Eyespots of freshwater planarians detect light and dark • Tissues organized into organs • Lack circulatory and respiratory systems – Gas exchange occurs by diffusion between body cells and the environment The Flatworms • Nervous system consists of – Clusters of nerve cells called ganglia (singular, ganglion) in the head that function as a simple brain – Ladderlike nerve cords extending the length of the body that conduct nerve signals to and from ganglia • Digestive system is an elaborately branched gastrovascular cavity – Centrally located ventral pharynx has opening that functions as both mouth and anus The Flatworms • Reproduce both sexually and asexually • Most are hermaphroditic (have both male and female sexual organs) – Allows parasitic flatworms to reproduce by self-fertilization Parasitic Flatworms • Free-living flatworms do not live in close association with members of other species • Parasitic flatworms live in or on the body of another organism (the host) which is harmed Parasitic Flatworms • Tend to have (VERY) complex life cycles • Some can infect humans – Liver flukes (common in Asia) – Blood flukes (cause schistosomiasis) – Tapeworms infect people who eat undercooked beef, pork, or fish containing cysts (encapsulated larvae!!! AHHHH!) FIGURE 23-10 The life cycle of the human pork tapeworm Each reproductive unit, or proglottid, is a self-contained reproductive factory that includes both male and female sex organs. 3.5: The Annelids • • • • Phylum Annelida AKA the segmented worms Bilateral symmetry Coelomate – Fluid-filled coelom functions as a hydrostatic skeleton (pressurized fluid provides a framework against which muscles can act) The Annelids • Body divided into a series of repeating units (segmentation) – Allows for complex movement – Segments contain identical copies of nerves, excretory structures, and muscles • Closed circulatory system (blood confined to heart and blood vessels) – Distributes gases and nutrients throughout body FIGURE 2311 An annelid, the earthworm This diagram shows an enlargement of segments, many of which are repeating similar units separated by partitions. The Annelids • Nervous system consists of – Clusters of ganglia in the head (brain) – Paired segmental ganglia – Paired ventral nerve cords extending the length of the body The Annelids • Digestive system consists of a tubular gut with two openings (mouth and anus) • Digestion occurs in a series of compartments – Pharynx – draws in food – Esophagus – conducts food to crop – Crop – stores food – Gizzard – grinds food – Intestine – absorbs digested nutrients • Excretion occurs through nephridia FIGURE 2311 An annelid, the earthworm This diagram shows an enlargement of segments, many of which are repeating similar units separated by partitions. The Annelids • Many annelids reproduce sexually • Some species are hermaphroditic; others have separate sexes • Some annelids reproduce asexually by fragmentation – The body breaks into two pieces, each of which regenerates the missing part The Annelids • The Annelid species fall into three main subgroups – Oligochaetes – Polychaetes – Leeches Oligochaetes & Polychaetes • Oligochaetes – Live in moist terrestrial habitats – e.g. earthworms • Polychaetes – Most are marine • Some live in tubes from which they project feathery gills (exchange gases and filter water) • Others have segmental, paired fleshy paddles that function in locomotion FIGURE 23-12a Diverse annelids (a) A polychaete annelid projects brightly spiraling gills from a tube attached to rock. When the gills retract, the tube is covered by the trap door visible on the lower right. FIGURE 2312b Diverse annelids (b) This tiny polychaete (seen here through a microscope) lives among seaside rocks near the tide line. Leeches • Leeches – Live in fresh water or moist terrestrial habitats – Are either carnivorous (prey on small invertebrates) or parasitic (suck blood) FIGURE 23-12c Diverse annelids (c) This leech, a freshwater annelid, shows numerous segments. The sucker encircles its mouth, allowing it to attach to its prey. 3.6: The Roundworms • • • • • • Phylum Nematoda Bilateral symmetry Pseudocoelomate Abundant, ubiquitous, and diverse Most are microscopic Elongate body is protected by a cuticle that must be molted periodically (kinda like arthropod exoskeleton…) FIGURE 23-25 A freshwater nematode Eggs can be seen inside this female freshwater nematode, which feeds on algae The Roundworms • Tubular gut with separate mouth and anus • Sensory organs transmit information to a simple brain • Lack circulatory and respiratory systems – Gas exchange occurs by diffusion between body cells and the environment • Most reproduce sexually – Male places sperm in body of female The Roundworms • Most are free-living and break down organic matter (very important ecologically) • Some are parasitic – Hookworm larvae bore into human feet and travel to the intestine, where they cause continuous bleeding – Trichinella worms infect people who eat improperly cooked infected pork • Larvae invade blood vessels and muscles, causing bleeding and muscle damage – Heartworms can be transmitted to dogs by the bite of an infected mosquito FIGURE 23-26a Some parasitic nematodes (a) Encysted larva of the Trichinella worm in the muscle tissue of a pig, where it may live for up to 20 years. FIGURE 23-26b Some parasitic nematodes (b) Adult heartworms in the heart of a dog. The juveniles are released into the bloodstream, where they may be ingested by mosquitoes and passed to another dog by the bite of an infected mosquito. 3.7: The Mollusks • • • • Phylum Mollusca Bilateral symmetry Coelomate Have a mantle (extension of the body wall) – Forms gill chamber – Secretes shell (in shelled species) The Mollusks • Most have an open circulatory system (blood is not confined to heart and blood vessels) – Blood percolates through a hemocoel (blood cavity) bathing internal organs directly The Mollusks • Nervous system is similar to that of annelids (ganglia connected by ventral nerves); however, more of the ganglia are concentrated in the head • Reproduce sexually – Some species have separate sexes – Others are hermaphroditic FIGURE 23-13 A generalized mollusk The general body plan of a mollusk, showing the mantle, foot, gills, shell, radula, and other features that are seen in most (but not all) mollusk species The Mollusks • Three prominent classes of molluscs are – Gastropods – Bivalves – Cephalopods Gastropods • Gastropods – Have a muscular foot for locomotion – May possess a shell – Feed using a radula (ribbon of tissue that bears numerous teeth) • Gastropods – Most use their skin and gills for respiration • Terrestrial mollusks have a simple lung – Include snails and slugs FIGURE 23-14a The diversity of gastropod mollusks (a) A Florida tree snail displays a brightly striped shell and eyes at the tip of stalks that retract instantly if touched. FIGURE 23-14b The diversity of gastropod mollusks (b) Spanish shawl sea slugs prepare to mate. The brilliant colors of many sea slugs warn potential predators that they are distasteful. Bivalves • Bivalves – Live in fresh water and marine habitats – Possess two shells that can be clamped shut by a strong muscle – Are filter feeders • Use gills for feeding and respiration • Bivalves – Most have a muscular foot used for burrowing or attachment to rocks – Include scallops, oysters, mussels, and clams FIGURE 23-15a The diversity of bivalve mollusks (a) This swimming scallop parts its hinged shells. The upper shell is covered with an encrusting sponge. FIGURE 23-15b The diversity of bivalve mollusks (b) Mussels attach to rocks in dense aggregations exposed at low tide. White barnacles are attached to the mussel shells and surrounding rock. Cephalopods • Cephalopods – Marine – Predatory carnivores • Cephalopods – Tentacles with chemosensory abilities and suction disks • Used for locomotion (octopus) and capture of prey – Able to move rapidly by forcefully expelling water from the mantle cavity – Closed circulatory system Cephalopods • Cephalopods – Large, complex brain (capable of learning) – Highly developed sensory systems • Eyes are remarkably similar to vertebrate eyes – May have a shell (nautiluses) – Include octopuses, nautiluses, cuttlefish, and squids FIGURE 23-16a The diversity of cephalopod mollusks (a) An octopus can crawl rapidly by using its eight suckered tentacles, and it can alter its color and skin texture to blend with its surroundings. In emergencies, this mollusk can jet backward by vigorously contracting its mantle. Octopuses and squid can emit clouds of dark purple ink to confuse pursuing predators. FIGURE 23-16b The diversity of cephalopod mollusks (b) The squid moves entirely by contracting its mantle to generate jet propulsion, which pushes the animal backward through the water. FIGURE 23-16c The diversity of cephalopod mollusks (c) The chambered nautilus secretes a shell with internal, gas-filled chambers that provide buoyancy. Note the well-developed eyes and the tentacles used to capture prey. 3.8: The Echinoderms • Phylum Echinodermata • Larvae exhibit bilateral symmetry; adults show radial symmetry • Coelomate • Exclusively marine • Include sand dollars, sea urchins, sea stars (starfish), sea cucumbers, and sea lilies FIGURE 23-27a The diversity of echinoderms (a) A sea cucumber feeds on debris in the sand. FIGURE 23-27b The diversity of echinoderms (b) The sea urchin's spines are actually projections of the internal skeleton. FIGURE 23-27c The diversity of echinoderms (c) The sea star has reduced spines and typically has five arms. The Echinoderms • Possess an endoskeleton (internal skeleton) that sends projections through the skin • A unique water-vascular system – Consists of the sieve plate, a circular central canal, several radial canals, and numerous tube feet – Functions in locomotion, respiration, and food capture FIGURE 23-28a The water-vascular system of echinoderms (a) Changing pressure inside the seawater-filled water-vascular system extends or retracts the tube feet. FIGURE 23-28b The water-vascular system of echinoderms (b) The sea star often feeds on mollusks such as this mussel. A feeding sea star attaches numerous tube feet to the mussel's shells, exerting a relentless pull. Then, the sea star turns the delicate tissue of its stomach inside out, extending it through its centrally located ventral mouth. The stomach can fit through an opening in the bivalve shells which measures less than 1 millimeter. Once insinuated between the shells, the stomach tissue secretes digestive enzymes that weaken the mollusk, causing it to open further. Partially digested food is transported to the upper portion of the stomach, where digestion is completed. The Echinoderms • Primitive nervous system; no distinct brain – Consists of a nerve ring, radial nerves, and a nerve network running through the epidermis – Sea stars have simple light and chemical receptors – Some brittle star species have lens-containing light receptors • Lack a circulatory system The Echinoderms • Most reproduce sexually – Shed eggs and sperm into water – Fertilization is external • Many are able to regenerate lost body parts – A sea star arm with part of the central body attached is able to form a whole animal 3.9: The Arthropods • • • • • Phylum Arthropoda Bilateral symmetry Coelomate Paired, jointed appendages Most abundant and diverse organisms on Earth • Include insects, arachnids, myriopods, and crustaceans The Arthropods • Have an exoskeleton (external skeleton) – Secreted by the epidermis (outer layer of skin) – Composed primarily of protein and chitin (a polysaccharide) – Functions include • Conservation of water • Protection against predators • Attachment sites for muscles (allows precise movements) FIGURE 23-17 The exoskeleton allows precise movements A net-throwing spider begins to wrap a captured insect in silk. Such dexterous manipulations are made possible by the exoskeleton and jointed appendages characteristic of arthropods. The Arthropods • Exoskeleton must be molted (shed) periodically FIGURE 23-18 The exoskeleton must be molted periodically A newly emerged praying mantis (a predatory insect) hangs beside its outgrown exoskeleton (left). The Arthropods • Insect body divided into three distinct regions, each specialized for different functions – Head (front region) – feeding and sensing the environment – Thorax (middle region) – locomotion – Abdomen (rear region) – digestion FIGURE 23-19 Segments are fused and specialized in insects Insects, such as this grasshopper, show fusion and specialization of body segments into a distinct head, thorax, and abdomen. Segments are visible on the abdomen beneath the wings. The Arthropods • Efficient gas exchange – Aquatic arthropods (crustaceans) have gills (thin, external respiratory membranes) – Terrestrial arthropods have either: • Tracheae (singular, trachea) – network of narrow, branching respiratory tubes • Book lungs (arachnids) • Most have an open circulatory system The Arthropods • Complex nervous system – Responsible for finely coordinated movement and complex behaviors • Well-developed sensory structures – Chemical and tactile receptors – Compound eyes • Composed of many light-detecting subunits • Are image-forming and can detect color FIGURE 23-20 Arthropods possess compound eyes This scanning electron micrograph shows the compound eye of a fruit fly. Compound eyes consist of an array of similar lightgathering and sensing elements whose orientation gives the arthropod a wide field of view. Insects have reasonably good image-forming ability and good color discrimination. The Arthropods • Phylum Arthropoda includes the classes: – – – – Insects (Insecta) Arachnids (Aracnida Myriopods (Myriapoda) Crustaceans (Crustacea) Insects • Most abundant and diverse class of arthropods • Most have – One pair of antennae – Three pairs of legs – Two pairs of wings (are the only invertebrates capable of flight) FIGURE 23-21a The diversity of insects (a) The rose aphid sucks sugar-rich juice from plants. FIGURE 23-21b The diversity of insects (b) A mating pair of Hercules beetles. The large ‘horns’ are found only on males. FIGURE 23-21d The diversity of insects (d) Insects such as this locust can cause devastation of food crops and natural vegetation. Insects • The ability to fly has contributed to the enormous success of insects – Helps insects escape predators – Allows insects to find widely dispersed food FIGURE 23-21c The diversity of insects (c) A June beetle displays its two pairs of wings as it comes in for a landing. The outer wings protect the abdomen and the inner wings, which are relatively thin and fragile. Insects • Use tracheae for gas exchange • In many, the juvenile is a larva (has a worm-shaped body form) – e.g. maggot (housefly larva), caterpillar (moth or butterfly larva) FIGURE 23-21e The diversity of insects (e) Caterpillars are larval forms of moths or butterflies. This caterpillar larva of the Australian fruitsucking moth displays large eyespot patterns that may frighten potential predators, who mistake them for eyes of a large animal. Insects • Undergo metamorphosis (radical change from juvenile body form to adult body form) • In insects with complete metamorphosis, developmental stages include: egg, larva (feeding stage), pupa (non-feeding stage), adult • In insects with incomplete metamorphosis, developmental stages include: egg, nymph (feeding stage that resembles adult), adult Insects • Insect classes are further classified into orders • Three of the largest insect orders include – Butterflies and moths – Bees, ants, and wasps – Beetles Arachnids • Lack antennae • Eight walking legs • Most are carnivorous (feed on blood or predigested prey) • Include spiders, mites, ticks, and scorpions FIGURE 23-22a The diversity of arachnids (a) The tarantula is one of the largest spiders but is relatively harmless. FIGURE 23-22b The diversity of arachnids (b) Scorpions, found in warm climates such as the deserts of the southwestern United States, paralyze their prey with venom from a stinger at the tip of the abdomen. A few species can harm humans. FIGURE 23-22c The diversity of arachnids (c) Ticks before (left) and after (right) feeding on blood. The uninflated exoskeleton is flexible and folded, allowing the animal to become grotesquely bloated while feeding. Arachnids • Some inject paralyzing venom into prey (spiders, scorpions) • Breathe by using tracheae and/or book lungs • Have simple image-forming eyes, each with a single lens • Abdominal glands produce protein threads (silk), used to weave webs Myriapods • • • • One pair of antennae Simple light-detecting eyes Respire by means of tracheae Include the centipedes and millipedes – Centipedes have one pair of legs per body segment – Millipedes have two pairs of legs per body segment FIGURE 23-23a The diversity of myriapods (a) Centipedes and (b) millipedes are common nocturnal arthropods. Each segment of a centipede's body holds one pair of legs, while each millipede segment has two pairs. FIGURE 23-23b The diversity of myriapods (a) Centipedes and (b) millipedes are common nocturnal arthropods. Each segment of a centipede's body holds one pair of legs, while each millipede segment has two pairs. Myriapods • Inhabit terrestrial environments (soil, leaf litter) • Centipedes are usually carnivorous – Inject poison into prey • Millipedes are usually decomposers; feed on decaying vegetation – Secrete a foul-smelling, distasteful liquid when attacked Crustaceans • Include crabs, crayfish, lobsters, shrimp, and barnacles • Most are aquatic • Two pairs of antennae • Most have compound eyes • Most respire by means of gills FIGURE 23-24a The diversity of crustaceans (a) The microscopic waterflea is common in freshwater ponds. Notice the eggs developing within the body. FIGURE 23-24b The diversity of crustaceans (b) The sowbug, found in dark, moist places such as under rocks, leaves, and decaying logs, is one of the few crustaceans to invade the land successfully. FIGURE 23-24c The diversity of crustaceans (c) The hermit crab protects its soft abdomen by inhabiting an abandoned snail shell. FIGURE 23-24d The diversity of crustaceans (d) The gooseneck barnacle uses a tough, flexible stalk to anchor itself to rocks, boats, or even animals such as whales. Other types of barnacles attach with shells that resemble miniature volcanoes (see Fig. 23-15b). Early naturalists thought barnacles were mollusks until they observed barnacles' jointed legs (seen here extending into the water).