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Chapter 33: Invertebrates 33 of the 34 phyla of animals are invertebrates Eumetazoa Ancestral colonial choanoflagellate Chordata Echinodermata Other bilaterians (including Nematoda, Arthropoda, Mollusca, and Annelida) Cnidaria Porifera Figure 33.2 Review of animal phylogeny Deuterostomia Bilateria Invertebrate taxa to know for BIO 111 2007 Total # of phyla: # presented in textbook: # discussed here: (# classes you need to know: 34 24 9 24) Invertebrate taxa to know for BIO 111 2007 Porifera – sponges Arthropoda Cnidaria Hexapoda - insects Hydrozoa - Hydra Crustacea – crabs. shrimp, barnacles Scyphozoa - jellyfish Myriapods – centipedes, millipedes Cubozoa – sea wasps Cheliceriforms – spiders, mites, ticks Anthozoa – sea anemones & corals Platyhelminthes – flatworms Echinodermata Turbellaria Asteroidea – sea stars Trematoda Opiuroidea – brittle stars Cestoda Echinoidea – sea urchins, sand $’s Nematoda Crinoidea – sea lilies Rotifera Holothuroidea – sea cucumbers Mollusca Concentricycloidea – sea daisies Polyplacophora – chitons Gastropoda – snails Bivalvia – clams & mussels Cephalopodia – squids & octopi Annelida Polychaeta – more worms Oligochaeta – earthworms Hirudinea - leeches Phylum Porifera - sponges Adam Laverty 5 Choanocytes. The spongocoel is lined with feeding cells called choanocytes. By beating flagella, the choanocytes create a current that draws water in through the porocytes. Azure vase sponge (Callyspongia plicifera) Flagellum Food particles Collar in mucus Choanocyte Osculum 4 Spongocoel. Water passing through porocytes enters a cavity called the spongocoel. Phagocytosis of food particles 3 Porocytes. Water enters 6 the epidermis through channels formed by porocytes, doughnut-shaped cells that span the body wall. Spicules 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”). Amoebocyte Water flow 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). Phylum Cnidaria Phylum Cnidaria (basal metazoan) Polyp Medusa Mouth/anus Tentacle Gastrovascular cavity Gastrodermis Mesoglea Body stalk Epidermis Tentacle Mouth/anus A cnidocyte with a nematocyst Prey Tentacle “Trigger” Nematocyst Discharge Of thread Coiled thread Cnidocyte Cnidarians (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. 33.7 Jelly Swimming corals of Class Anthozoa Phylum Platyhelminthes A Turbellarian 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 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. Schistosomiasis (bilharzia) is caused by a trematode 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 1 mm 5 These larvae penetrate the skin and blood vessels of humans working in irrigated fields contaminated with infected human feces. 2 Blood flukes reproduce sexually in the human host. The fertilized eggs exit the host in feces. 3 The eggs develop in water into ciliated larvae. These larvae infect snails, the intermediate hosts. 4 Asexual reproduction within a snail results in another type of motile larva, which escapes from the snail host. Snail host Class Cestoda: tapeworms red meat fish Proglottids with reproductive structures 200 µm Scolex Hooks Sucker Phylum Rotifera 0.1 mm Lophophorates (several phyla) Lophophore Lophophore Lophophore (a) Ectoprocts, such as this sea (b) In phoronids such as mat (Membranipora Phoronis hippocrepia, the membranacea), are colonial lophophore and mouth lophophorates. are at one end of an elongated trunk. (c) Brachiopods have a hinged shell. The two parts of the shell are dorsal and ventral. A ribbon worm, phylum Nemertea, is a lophophorate Phylum Mollusca 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 The nervous system consists of a nerve ring around the esophagus, from which nerve cords extend. Intestine Gonads Stomach Shell Radula Anus Gill Foot Nerve cords Esophagus Mouth Mouth Radula. The mouth region in many mollusc species contains a rasp-like feeding organ called a radula. This belt of backwardcurved teeth slides back and forth, scraping and scooping like a backhoe. A chiton, class Polyplacophora Class Gastropoda (limpets, too) (a) A land snail (b) A sea slug. Nudibranchs, or sea slugs, lost their shell during their evolution. The results of torsion in a gastropod Stomach Mantle cavity Anus Mouth Intestine Class Bivalvia Figure 33.21 Anatomy of a clam Hinge area Mantle Gut Coelom pearls Heart Shell Adductor muscle Mouth Anus Excurrent siphon Palp Water flow Foot Mantle cavity Gill Incurrent siphon glochidia of a bivalve Class Cephalopoda (b) Squids are speedy carnivores with beaklike jaws and welldeveloped eyes. (a) Octopuses are considered among the most intelligent invertebrates. “head-foot” shell is internal (beak) reproduction camouflage & ink brains (c) Chambered nautiluses are the only living cephalopods with an external shell. show the movies Phylum Annelida – the segmented worms Chaetae. Each segment has four pairs of chaetae, bristles that provide traction for burrowing. 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. Epidermis Cuticle Circular muscle Many of the internal structures are repeated within each segment of the earthworm. Metanephridium. Each segment of the worm contains a pair of excretory tubes, called metanephridia, with ciliated funnels, called Septum nephrostomes. The (partition metanephridia remove between wastes from the blood segments) and coelomic fluid through exterior pores. Coelom. The coelom of the earthworm is partitioned by septa. Each segment is surrounded by longitudinal muscle, which in turn is surrounded by circular muscle. Earthworms coordinate the contraction of these two sets of muscles to move (see Figure 49.25). These muscles work against the noncompressible coelomic fluid, which acts as a hydrostatic skeleton. Longitudinal muscle Dorsal vessel Anus Intestine Nerve cords Tiny blood vessels are abundant in the earthworm’s skin, which functions as its respiratory organ. The blood contains oxygen-carrying hemoglobin. Ventral vessel Nephrostome Clitellum Pharynx Esophagus Metanephridium Crop Giant Australian earthworm Intestine 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 the length of the animal, as do the digestive linked by segmental pairs of vessels. The dorsal tract and longitudinal blood vessels. vessel and five pairs of vessels that circle the esophagus of an earthworm are muscular and pump blood through the circulatory system. Darwin again: “1 acre of British farmland contained about 50,000 earthworms that produced 18 tons of castings per year” (Campbell & Reese) Invasive species? A polychaete Annelid Parapodia A leech, class Hirudinea mostly freshwater, but watch out for those terrestrial forms. predators, parasites, scavengers reduced anatomy Some parasites highly specialized hirudin anaesthesia vasodilators HOWARD LIPIN / Union-Tribune Phylum Nematoda 25 µm Trichinosis is caused by parasitic nematodes Encysted juveniles Muscle tissue 50 µm Trilobites were early arthropods, >250 mya Cephalothorax Antennae (sensory reception) Head Abdomen Thorax Swimming appendages Walking legs Pincer (defense) Mouthparts (feeding) Horseshoe crabs are Cheliceriforms Arachnids are cheliceriforms 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). Intestine Digestive gland Stomach Brain Heart Eyes Poison gland Ovary Anus Book lung Spinnerets Silk gland Gonopore (exit for eggs) Sperm receptacle Chelicera Pedipalp millipedes are myriapods Centipedes are myriapods 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 Cerebral ganglion. The two nerve Heart. The cords meet in the head, where the insect heart ganglia of several anterior segments drives hemolymph are fused into a cerebral ganglion through an (brain). The antennae, eyes, and open circulatory other sense organs are concentrated system. on the head. Antennae Ovary 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. Dorsal artery Crop Insect mouthparts are formed from several pairs of modified appendages. The mouthparts include mandibles, which grasshoppers use for chewing. In other insects, mouthparts are specialized for lapping, piercing, or sucking. Metamorphosis (ecydsis) (a) Larva (caterpillar) (b) Pupa (c) Pupa (d) Emerging adult (e) Adult ORDER Blattodea Coleoptera Dermaptera Diptera Hemiptera Hymenoptera APROXIMATE NUMBER OF SPECIES 4,000 350,000 1,200 151,000 85,000 125,000 MAIN CHARACTERISTICS EXAMPLES Cockroaches have a dorsoventrally flattened body, with legs modified for rapid running. Forewings, when present, are leathery, whereas hind wings are fanlike. Fewer than 40 cockroach species live in houses; the rest exploit habitats ranging from tropical forest floors to caves and deserts. German cockroach Beetles comprise the most species-rich order of insects. They have two pairs of wings, one of which is thick and leathery, the other membranous. They have an armored exoskeleton and mouthparts adapted for biting and chewing. Beetles undergo complete metamorphosis. Japanese beetle Earwigs are generally nocturnal scavengers. While some species are wingless, others have two pairs of wings, one of which is thick and leathery, the other membranous. Earwigs have biting mouthparts and large posterior pincers. They undergo incomplete metamorphosis. Dipterans have one pair of wings; the second pair has become modified into balancing organs called halteres. Their head is large and mobile; their mouthparts are adapted for sucking, piercing, or lapping. Dipterans undergo complete metamorphosis. Flies and mosquitoes are among the best-known dipterans, which live as scavengers, predators, and parasites. Hemipterans are so-called “true bugs,” including bed bugs, assassin bugs, and chinch bugs. (Insects in other orders are sometimes erroneously called bugs.) Hemipterans have two pairs of wings, one pair partly leathery, the other membranous. They have piercing or sucking mouthparts and undergo incomplete metamorphosis. Earwig Horsefly LeafFooted bug 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 ORDER Lepidoptera Odonata APROXIMATE NUMBER OF SPECIES 120,000 5,000 EXAMPLE MAIN CHARACTERISTICS Butterflies and moths are among the best-known insects. They have two pairs of wings covered with tiny scales. To feed, they uncoil a long proboscis. Most feed on nectar, but some species feed on other substances, including animal blood or tears. Swallowtail butterfly Dragonflies and damselflies have two pairs of large, membranous wings. They have an elongated abdomen, large, compound eyes, and chewing mouthparts. They undergo incomplete metamorphosis and are active predators. Dragonfly Orthoptera 13,000 Grasshoppers, crickets, and their relatives are mostly herbivorous. They have large hind legs adapted for jumping, two pairs of wings (one leathery, one membranous), and biting or chewing mouthparts. Males commonly make courtship sounds by rubbing together body parts, such as a ridge on their hind leg. Orthopterans undergo incomplete metamorphosis. Katydid Phasmida Phthiraptera Siphonaptera 2,600 2,400 2,400 Stick insects and leaf insects are exquisite mimics of plants. The eggs of some species even mimic seeds of the plants on which the Insects live. Their body is cylindrical or flattened dorsoventrally. They lack forewings but have fanlike hind wings. Their mouthparts are adapted for biting or chewing. Commonly called sucking lice, these insects spend their entire life as an ectoparasite feeding on the hair or feathers of a single host. Their legs, equipped with clawlike tarsi, are adapted for clinging to their hosts. They lack wings and have reduced eyes. Sucking lice undergo incomplete metamorphosis. Stick insect Human Body louse 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 Class Crustacea (a) Ghost crabs (genus Ocypode) live on sandy ocean beaches worldwide. Primarily nocturnal, they take shelter in burrows during the day. (b) Planktonic crustaceans known as krill are consumed in vast quantities by whales. (c) The jointed appendages projecting from the shells of these barnacles capture organisms and organic particles suspended in the water. Phylum Echinodermata A short digestive tract runs from the mouth on the bottom of the central disk to the anus on top of the disk. Central disk. The central disk has a nerve ring and nerve cords radiating from the ring into the arms. Spine 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. Gills 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 Ampulla Podium Tube feet 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. 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. (a) A sea star (class Asteroidea) (b) A brittle star (class Ophiuroidea) (c) A sea urchin (class Echinoidea) (d) A feather star (class Crinoidea) (e) A sea cucumber (class Holothuroidea) (f) A sea daisy (class Concentricycloidea)