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LECTURE PRESENTATIONS For CAMPBELL BIOLOGY, NINTH EDITION Jane B. Reece, Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Robert B. Jackson Chapter 33 An Introduction to Invertebrates Lectures by Erin Barley Kathleen Fitzpatrick © 2011 Pearson Education, Inc. Overview: Life Without a Backbone • Invertebrates are animals that lack a backbone • They account for 95% of known animal species • They are morphologically diverse – For example, the Christmas tree worm (Spirobranchus giganteus) has tentacles for gas exchange and feeding © 2011 Pearson Education, Inc. Figure 33.1 Figure 33.3a Porifera (5,500 species) Placozoa (1 species) 0.5 mm A sponge Cnidaria (10,000 species) A placozoan (LM) Ctenophora (100 species) A jelly Acoela (400 species) 1.5 mm Acoel flatworms (LM) A ctenophore, or comb jelly Figure 33.3b Platyhelminthes (20,000 species) Rotifera (1,800 species) 0.1 mm Ectoprocta (4,500 species) Ectoprocts A marine flatworm Acanthocephala (1,100 species) Nemertea (900 species) Brachiopoda (335 species) A brachiopod A rotifer (LM) Annelida (16,500 species) Cycliophora (1 species) Curved hooks 100 m An acanthocephalan (LM) Mollusca (93,000 species) A ribbon worm An octopus A cycliophoran (colorized SEM) Lophotrochozoa A marine annelid Figure 33.3c Loricifera (10 species) Priapula (16 species) Onychophora (110 species) 50 m A loriciferan (LM) A priapulan An onychophoran Nematoda (25,000 species) Tardigrada (800 species) Arthropoda (1,000,000 species) 100 m A roundworm (colored SEM) Ecdysozoa Tardigrades (colorized SEM) A scorpion (an arachnid) Figure 33.3d Hemichordata (85 species) Chordata (52,000 species) A tunicate Echinodermata (7,000 species) An acorn worm Deuterostomia A sea urchin Concept 33.1: Sponges are basal animals that lack true tissues • Animals in the phylum Porifera are known informally as sponges • They are sedentary and live in marine waters or fresh water © 2011 Pearson Education, Inc. • Sponges are suspension feeders, capturing food particles suspended in the water that passes through their body • Water is drawn through pores into a cavity called the spongocoel and out through an opening called the osculum • Sponges lack true tissues and organs © 2011 Pearson Education, Inc. Figure 33.4 Azure vase sponge (Callyspongia plicifera) Food particles in mucus Flagellum Choanocyte Collar Choanocyte Osculum Phagocytosis of food particles Amoebocyte Spongocoel Pore Spicules Epidermis Mesohyl Water flow Amoebocytes • Choanocytes, flagellated collar cells, generate a water current through the sponge and ingest suspended food • Sponges consist of a gelatinous noncellular mesohyl layer between two cell layers • Amoebocytes are found in the mesohyl and play roles in digestion and structure • Most sponges are hermaphrodites: Each individual functions as both male and female © 2011 Pearson Education, Inc. • Cnidarians have diversified into a wide range of both sessile and motile forms including jellies, corals, and hydras • They exhibit a relatively simple radial body plan • The basic body plan of a cnidarian is a sac with a central digestive compartment, the gastrovascular cavity • A single opening functions as mouth and anus © 2011 Pearson Education, Inc. • There are two variations on the body plan: the sessile polyp and motile medusa • A polyp adheres to the substrate by the aboral end of its body • A medusa has a bell-shaped body with its mouth on the underside • Medusae do not attach to the substate but move freely © 2011 Pearson Education, Inc. Figure 33.5 Mouth/anus Tentacle Gastrovascular cavity Gastrodermis Mesoglea Body stalk Epidermis Tentacle Polyp Mouth/anus Medusa • Cnidarians are carnivores that use tentacles to capture prey • The tentacles are armed with cnidocytes, unique cells that function in defense and capture of prey • Nematocysts are specialized organelles within cnidocytes that eject a stinging thread © 2011 Pearson Education, Inc. Figure 33.6 Tentacle Cuticle of prey Thread Nematocyst “Trigger” Thread discharges Cnidocyte Thread (coiled) • Phylum Cnidaria is divided into four major classes – – – – Hydrozoa Scyphozoa Cubozoa Anthozoa © 2011 Pearson Education, Inc. Figure 33.7 (b) Scyphozoa (a) Hydrozoa (c) Cubozoa (d) Anthozoa Hydrozoans • Most hydrozoans alternate between polyp and medusa forms • Hydra, a freshwater cnidarian, exists only in polyp form and reproduces asexually by budding © 2011 Pearson Education, Inc. Figure 33.8-3 Feeding polyp Reproductive polyp Medusa bud MEIOSIS Gonad Medusa Egg ASEXUAL REPRODUCTION (BUDDING) Portion of a colony of polyps SEXUAL REPRODUCTION FERTILIZATION Developing polyp 1 mm Sperm Mature polyp Zygote Planula (larva) Key Haploid (n) Diploid (2n) Scyphozoans • In the class Scyphozoa, jellies (medusae) are the prevalent form of the life cycle © 2011 Pearson Education, Inc. Cubozoans • In the class Cubozoa, which includes box jellies and sea wasps, the medusa is box-shaped and has complex eyes • Cubozoans often have highly toxic cnidocytes © 2011 Pearson Education, Inc. Anthozoans • Class Anthozoa includes the corals and sea anemones, and these cnidarians occur only as polyps • Corals often form symbioses with algae and secrete a hard external skeleton © 2011 Pearson Education, Inc. • Lophotrochozoa develop a lophophore for feeding, others pass through a trochophore larval stage, and a few have neither feature • Lophotrochozoa includes the flatworms, rotifers, ectoprocts, brachiopods, molluscs, and annelids © 2011 Pearson Education, Inc. Flatworms • Members of phylum Platyhelminthes live in marine, freshwater, and damp terrestrial habitats • Flatworms are acoelomates • They are flattened dorsoventrally and have a gastrovascular cavity with one opening • Gas exchange takes place across the surface, and protonephridia regulate the osmotic balance © 2011 Pearson Education, Inc. • Flatworms are divided into two lineages – Catenulida, or “chain worms,” reproduce asexually by budding – Rhabditophora are more diverse and include both free-living and parasitic species © 2011 Pearson Education, Inc. Figure 33.9 5 mm Free-Living Species • The best-known rhabditophorans are planarians • Planarians live in fresh water and prey on smaller animals • Planarians have light-sensitive eyespots and centralized nerve nets © 2011 Pearson Education, Inc. • The planarian nervous system is more complex and centralized than the nerve nets of cnidarians • Planarians are hermaphrodites and can reproduce sexually, or asexually through fission © 2011 Pearson Education, Inc. Figure 33.10 Gastrovascular cavity Pharynx Gastrovascular cavity Mouth Eyespots Ventral nerve cords Ganglia Parasitic Species • Parasitic rhabditophorans live in or on other animals • Two important groups of parasitic rhabditophorans are the trematodes and the tapeworms © 2011 Pearson Education, Inc. Trematodes • Trematodes parasitize a wide range of hosts, and most have complex life cycles with alternating sexual and asexual stages • Trematodes that parasitize humans spend part of their lives in snail hosts • They produce surface proteins that mimic their host and release molecules that manipulate the host’s immune system © 2011 Pearson Education, Inc. Figure 33.11 Male Human host Female 1 mm Mature flukes Ciliated larva Motile larva Snail host Tapeworms • Tapeworms are parasites of vertebrates and lack a digestive system • Tapeworms absorb nutrients from the host’s intestine © 2011 Pearson Education, Inc. • The scolex contains suckers and hooks for attaching to the host • Proglottids are units that contain sex organs and form a ribbon behind the scolex • Fertilized eggs, produced by sexual reproduction, leave the host’s body in feces © 2011 Pearson Education, Inc. Figure 33.12 100 m Hooks Sucker Proglottids with reproductive structures Scolex Rotifers • Rotifers, phylum Rotifera, are tiny animals that inhabit fresh water, the ocean, and damp soil • Rotifers are smaller than many protists but are truly multicellular and have specialized organ systems © 2011 Pearson Education, Inc. Figure 33.13 Jaws Anus Stomach Crown of cilia 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 offspring from unfertilized eggs • Some species are unusual in that they lack males entirely © 2011 Pearson Education, Inc. Lophophorates: Ectoprocts and Brachiopods • Lophophorates have a lophophore, a crown of ciliated tentacles around their mouth • Lophophorates have a true coelom • Lophophorates include two phyla: Ectoprocta and Brachiopoda © 2011 Pearson Education, Inc. • Ectoprocts (also called bryozoans) are sessile colonial animals that superficially resemble plants • A hard exoskeleton encases the colony, and some species are reef builders © 2011 Pearson Education, Inc. Figure 33.14 Lophophore (a) Ectoprocts, colonial lophophorates Lophophore (b) Lampshell, a brachiopod • 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 • Brachiopods are marine and attach to the seafloor by a stalk © 2011 Pearson Education, Inc. Figure 33.14b Lophophore (b) Lampshell, a brachiopod Molluscs • Phylum Mollusca includes snails and slugs, oysters and clams, and octopuses and squids • Most molluscs are marine, though some inhabit fresh water and some snails and slugs are terrestrial • Molluscs are soft-bodied animals, but most are protected by a hard shell © 2011 Pearson Education, Inc. • All molluscs have a similar body plan with three main parts – Muscular foot – Visceral mass – Mantle • Many molluscs also have a water-filled mantle cavity and feed using a rasplike radula © 2011 Pearson Education, Inc. Figure 33.15 Nephridium Heart Visceral mass Coelom Digestive tract Intestine Gonads Mantle Stomach Shell Radula Mantle cavity Mouth Anus Gill Foot Nerve cords Esophagus Mouth Radula • Most molluscs have separate sexes with gonads located in the visceral mass, but many snails are hermaphrodites • The life cycle of many molluscs includes a ciliated larval stage called a trochophore © 2011 Pearson Education, Inc. • Four of the major classes of molluscs are – – – – Polyplacophora (chitons) Gastropoda (snails and slugs) Bivalvia (clams, oysters, and other bivalves) Cephalopoda (squids, octopuses, cuttlefish, and chambered nautiluses) © 2011 Pearson Education, Inc. Chitons • Chitons are oval-shaped marine animals encased in an armor of eight dorsal plates • They use their foot like a suction cup to grip rock, and their radula to scrape algae off the rock surface © 2011 Pearson Education, Inc. Figure 33.16 Gastropods • About three-quarters of all living species of molluscs are gastropods © 2011 Pearson Education, Inc. Figure 33.17 (a) A land snail (b) A sea slug (nudibranch) • Most gastropods are marine, but many are freshwater and terrestrial species • The most distinctive characteristic of gastropods is torsion, which causes the animal’s anus and mantle to end up above its head; torsion is different from the coiling of a shell • Most have a single, spiraled shell • Slugs lack a shell or have a reduced shell © 2011 Pearson Education, Inc. Figure 33.18 Mantle cavity Anus Mouth Stomach Intestine Bivalves • Bivalves are marine and include many species of clams, oysters, mussels, and scallops • They have a shell divided into two halves drawn together by adductor muscles • Some bivalves have eyes and sensory tentacles along the edge of their mantle © 2011 Pearson Education, Inc. Figure 33.19 • The mantle cavity of a bivalve contains gills that are used for feeding as well as gas exchange © 2011 Pearson Education, Inc. Figure 33.20 Mantle Hinge area Coelom Gut Heart Digestive gland Adductor muscle (one or two) Anus Mouth Excurrent siphon Shell Palp Foot Mantle cavity Gonad Gill Water flow Incurrent siphon Cephalopods • Cephalopods are carnivores with beak-like jaws surrounded by tentacles of their modified foot • Most octopuses creep along the sea floor in search of prey © 2011 Pearson Education, Inc. Figure 33.21 Squid Octopus Chambered nautilus • Squids use their siphon to fire a jet of water, which allows them to swim very quickly © 2011 Pearson Education, Inc. Figure 33.21b Octopus • One small group of shelled cephalopods, the nautiluses, survives today © 2011 Pearson Education, Inc. Figure 33.21c Chambered nautilus • Cephalopods have a closed circulatory system, well-developed sense organs, and a complex brain • Shelled cephalopods called ammonites were common but went extinct at the end of the Cretaceous 65.5 million years ago © 2011 Pearson Education, Inc. Protecting Freshwater and Terrestrial Molluscs • Molluscs are the animal group with the largest number of recent extinctions • The most threatened groups are – Freshwater bivalves, including pearl mussels – Terrestrial gastropods, including Pacific island land snails • These molluscs are threatened by habitat loss, pollution, and non-native species © 2011 Pearson Education, Inc. Figure 33.22 Other invertebrates Molluscs Amphibians Insects Birds Fishes Mammals An endangered Pacific island land snail, Partula suturalis Recorded extinctions of animal species (Data: International Union for Reptiles (excluding birds) Conservation of Nature, 2008) Workers on a mound of pearl mussels killed to make buttons (ca. 1919) Annelids • Annelids have bodies composed of a series of fused rings • Annelids are coelomates • The phylum Annelida is divided into two groups – Polychaeta (polychaetes) – Oligochaeta (earthworms and their relatives, and leeches) © 2011 Pearson Education, Inc. Polychaetes • Members of class Polychaetes have paddle-like parapodia that work as gills and aid in locomotion • Most polychaetes are marine © 2011 Pearson Education, Inc. Figure 33.23 Parapodia Oligochaetes • Oligochaetes (class Oligochaeta) are named for relatively sparse chaetae, bristles made of chitin © 2011 Pearson Education, Inc. Earthworms • Earthworms eat through soil, extracting nutrients as the soil moves through the alimentary canal • Earthworms are hermaphrodites but cross-fertilize • Some reproduce asexually by fragmentation © 2011 Pearson Education, Inc. Figure 33.24 Cuticle Coelom Epidermis Septum (partition between segments) Circular muscle Metanephridium Longitudinal muscle Anus Dorsal vessel Chaetae Intestine Skin Ventral vessel Fused nerve cords Nephrostome Metanephridium Clitellum Esophagus Crop Pharynx Intestine Giant Australian earthworm Gizzard Cerebral ganglia Mouth Subpharyngeal ganglion Circulatory system vessels Ventral nerve cords with segmental ganglia Leeches • Most species of leeches live in fresh water; some are marine or terrestrial • Leeches include predators of invertebrates, and parasites that suck blood • Leeches secrete a chemical called hirudin to prevent blood from coagulating © 2011 Pearson Education, Inc. Figure 33.25 Concept 33.4: Ecdysozoans are the most species-rich animal group • Ecdysozoans are covered by a tough coat called a cuticle • The cuticle is shed or molted through a process called ecdysis • The two largest phyla are nematodes and arthropods © 2011 Pearson Education, Inc. Nematodes • Nematodes, or roundworms, are found in most aquatic habitats, in the soil, in moist tissues of plants, and in body fluids and tissues of animals • They have an alimentary canal, but lack a circulatory system • Reproduction in nematodes is usually sexual, by internal fertilization © 2011 Pearson Education, Inc. Figure 33.26 25 m • Caenorhabditis elegans is a model organism in research • Some species of nematodes are important parasites of plants and animals • Thrichinella spiralis can be acquired by humans from undercooked pork © 2011 Pearson Education, Inc. Figure 33.27 Encysted juveniles Muscle tissue 50 m Arthropods • 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 © 2011 Pearson Education, Inc. Arthropod Origins • The arthropod body plan consists of a segmented body, hard exoskeleton, and jointed appendages • This body plan dates to the Cambrian explosion (535–525 million years ago) • Early arthropods show little variation from segment to segment © 2011 Pearson Education, Inc. Figure 33.28 General Characteristics of Arthropods • The appendages of some living arthropods are modified for functions such as walking, feeding, sensory reception, reproduction, and defense © 2011 Pearson Education, Inc. Figure 33.30 Cephalothorax Antennae (sensory reception) Abdomen Thorax Head Swimming appendages (one pair per abdominal segment Pincer (defense) Mouthparts (feeding) Walking legs • The body of an arthropod is completely covered by the cuticle, an exoskeleton made of layers of protein and the polysaccharide chitin • When an arthropod grows, it molts its exoskeleton © 2011 Pearson Education, Inc. • Arthropods have eyes, olfactory receptors, and antennae that function in touch and smell • Arthropods have an open circulatory system in which hemolymph is circulated into the spaces surrounding the tissues and organs • A variety of organs specialized for gas exchange have evolved in arthropods © 2011 Pearson Education, Inc. • Molecular evidence suggests that living arthropods consist of four major lineages that diverged early in the phylum’s evolution – Chelicerates (sea spiders, horseshoe crabs, scorpions, ticks, mites, and spiders) – Myriapods (centipedes and millipedes) – Hexapods (insects and relatives) – Crustaceans (crabs, lobsters, shrimps, barnacles, and many others) © 2011 Pearson Education, Inc. Chelicerates • Chelicerates, subphylum Chelicerata, are named for clawlike feeding appendages called chelicerae • The earliest cheliceriforms were eurypterids (water scorpions) • Most marine cheliceriforms (including eurypterids) are extinct, but some species survive today, including horseshoe crabs © 2011 Pearson Education, Inc. Figure 33.31 • Most modern cheliceriforms are arachnids, which include spiders, scorpions, ticks, and mites © 2011 Pearson Education, Inc. Figure 33.32 50 m Scorpion Dust mite Web-building spider • Arachnids have an abdomen and a cephalothorax, which has six pairs of appendages: the chelicerae, the pedipalps, and four pairs of walking legs • Gas exchange in spiders occurs in respiratory organs called book lungs • Many spiders produce silk, a liquid protein, from specialized abdominal glands © 2011 Pearson Education, Inc. Figure 33.33 Stomach Intestine Heart Brain Digestive gland Eyes Ovary Poison gland Anus Book lung Spinnerets Silk gland Gonopore (exit for eggs) Sperm receptacle Chelicera Pedipalp Myriapods • Subphylum Myriapoda includes millipedes and centipedes • Myriapods are terrestrial, and have jaw-like mandibles • Millipedes eat decaying leaves and plant matter • Millipedes have many legs, with two pairs per trunk segment © 2011 Pearson Education, Inc. Figure 33.34 (a) Millipede (b) Centipede • Centipedes are carnivores • Centipedes have one pair of legs per trunk segment © 2011 Pearson Education, Inc. Insects • Subphylum Hexapoda, insects and relatives, has more species than all other forms of life combined • They live in almost every terrestrial habitat and in fresh water • The internal anatomy of an insect includes several complex organ systems © 2011 Pearson Education, Inc. Figure 33.35 Abdomen Thorax Head Compound eye Antennae Heart Dorsal artery Crop Anus Vagina Malpighian tubules Ovary Tracheal tubes Nerve cords Mouthparts Cerebral ganglion • Insects diversified several times following the evolution of flight, adaptation to feeding on gymnosperms, and the expansion of angiosperms • Insect and plant diversity declined during the Cretaceous extinction, but has been increasing in the 65 million years since © 2011 Pearson Education, Inc. • Flight is 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 © 2011 Pearson Education, Inc. Figure 33.36 • 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 © 2011 Pearson Education, Inc. • Insects with complete metamorphosis have larval stages known by such names as maggot, grub, or caterpillar • The larval stage looks entirely different from the adult stage © 2011 Pearson Education, Inc. Figure 33.37 (a) Larva (caterpillar) (b) Pupa (c) Later-stage (d) Emerging pupa adult (e) Adult • Most insects have separate males and females and reproduce sexually • Individuals find and recognize members of their own species by bright colors, sound, or odors • Some insects are beneficial as pollinators, while others are harmful as carriers of diseases, or pests of crops • Insects are classified into more than 30 orders © 2011 Pearson Education, Inc. Figure 33.38 Archaeognatha (bristletails; 350 species) Thysanura (silverfish; 450 species) Winged insects (many orders) Complete metamorphosis Coleoptera (beetles; 350,000 species) Diptera (151,000 species) Hymenoptera (125,000 species) Proboscis Lepidoptera (120,000 species) Incomplete metamorphosis Hemiptera (85,000 species) Orthoptera (13,000 species) Crustaceans • While arachnids and insects thrive on land, crustaceans, for the most part, have remained in marine and freshwater environments • Crustaceans, subphylum Crustacea, typically have branched appendages that are extensively specialized for feeding and locomotion • Small crustaceans exchange gases through the cuticle; larger crustaceans have gills © 2011 Pearson Education, Inc. • Most crustaceans have separate males and females • Isopods include terrestrial, freshwater, and marine species – Pill bugs are a well known group of terrestrial isopods • Decapods are all relatively large crustaceans and include lobsters, crabs, crayfish, and shrimp © 2011 Pearson Education, Inc. Figure 33.39 (a) Ghost crab (b) Krill (c) Barnacles • Planktonic crustaceans include many species of copepods, which are among the most numerous of all animals © 2011 Pearson Education, Inc. • Barnacles are a group of mostly sessile crustaceans • They have a cuticle that is hardened into a shell © 2011 Pearson Education, Inc. Concept 33.5: Echinoderms and chordates are deuterostomes • Echinoderms (phylum Echinodermata) include sea stars and sea urchins • Chordates (phylum Chordata) include the vertebrates • Echinoderms and chordates constitute the clade Deuterostomia © 2011 Pearson Education, Inc. • Deuterostomes share developmental characteristics – Radial cleavage – Formation of the anus from the blastopore • However, some other animals also share these developmental characteristics • Deuterostomes are defined primarily by DNA similarities © 2011 Pearson Education, Inc. Echinoderms • Sea stars and most other echinoderms are slow-moving or sessile marine animals • A thin epidermis covers an endoskeleton of hard calcareous plates • Echinoderms have a unique water vascular system, a network of hydraulic canals branching into tube feet that function in locomotion and feeding • Males and females are usually separate, and sexual reproduction is external © 2011 Pearson Education, Inc. Figure 33.40 Short digestive tract Stomach Anus Spine Gills Central disk Madreporite Digestive glands Radial nerve Ring canal Radial canal Gonads Ampulla Podium Tube feet • Most adult echinoderms have radial symmetry with multiples of five • Echinoderm larvae have bilateral symmetry © 2011 Pearson Education, Inc. • Living echinoderms are divided into five classes – Asteroidea (sea stars and sea daisies) – Ophiuroidea (brittle stars) – Echinoidea (sea urchins and sand dollars) – Crinoidea (sea lilies and feather stars) – Holothuroidea (sea cucumbers) © 2011 Pearson Education, Inc. Asteroidea: Sea Stars and Sea Daisies • Sea stars have multiple arms radiating from a central disk • The undersurface of each arm bears tube feet, which grip substrate with adhesive chemicals • Sea stars feed on bivalves by prying them open with their tube feet, everting their stomach, and digesting their prey externally with digestive enzymes • Sea stars can regrow lost arms © 2011 Pearson Education, Inc. • Sea daisies were discovered in 1986, and only three species are known • Sea daisies live on submerged wood and absorb nutrients through a membrane that surrounds their body © 2011 Pearson Education, Inc. Figure 33.41 Ophiuroidea: Brittle Stars • Brittle stars have a distinct central disk and long, flexible arms, which they use for movement • Some species are suspension feeders, while others are predators or scavengers © 2011 Pearson Education, Inc. Figure 33.42 Echinoidea: Sea Urchins and Sand Dollars • Sea urchins and sand dollars have no arms but have five rows of tube feet • Their spines are used for locomotion and protection • Sea urchins feed on seaweed using a jaw-like structure on their underside © 2011 Pearson Education, Inc. Figure 33.43 Crinoidea: Sea Lilies and Feather Stars • Sea lilies live attached to the substrate by a stalk • Feather stars can crawl using long, flexible arms • Both are suspension feeders © 2011 Pearson Education, Inc. Figure 33.44 Holothuroidea: Sea Cucumbers • Sea cucumbers lack spines, have a very reduced endoskeleton, and do not look much like other echinoderms • Sea cucumbers have five rows of tube feet; some of these are developed as feeding tentacles © 2011 Pearson Education, Inc. Figure 33.45 Chordates • Phylum Chordata consists of two subphyla of invertebrates as well as hagfishes and vertebrates • Chordates are bilaterally symmetrical coelomates with segmented bodies • Chordates share many features of embryonic development with echinoderms, but have evolved separately for at least 500 million years © 2011 Pearson Education, Inc. Figure 33.UN06