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Expanded Contents List of Research Focus Boxes iii Preface/Introduction ix Guide to the Major Animal Groups xiii The Geologic Time Scale xiv 1 Environmental Considerations 1 2 Invertebrate Classification and Relationships 5 Introduction 5 Classification by Cell Number, Embryology, and Body Symmetry 6 Classification by Developmental Pattern 7 Classification by Evolutionary Relationship 14 Deducing Evolutionary Relationships 17 Why Determine Evolutionary Trees? 21 How Evolutionary Relationships Are Determined 21 Uncertainty about Evolutionary Relationships 27 Time Machines: Help from the Fossil Record 27 Classification by Habitat and Lifestyle 28 3 The Protozoans 33 Introduction 33 General Characteristics 35 Phylum Ciliophora 39 Structure and Function of Cilia 39 Patterns of Ciliation 40 Other Morphological Features 41 Reproductive Characteristics 41 Ciliate Lifestyles 47 Amoeboid Protozoans-The Sarcodinids 50 Phylum Amoebozoa 51 Phylum Foraminifera 53 Phylum Radiozoa 54 Phylum Heliozoa 56 Flagellated Protozoans 57 The Phytoflagellated Protozoans 57 The Zooflagellated Protozoans- Free-Living Forms 58 The Zooflagellated Protozoans-Parasitic Forms 58 Some Transitional Forms and Misfits 61 Spore-Forming Protozoans 64 Phylum Sporozoa (" Apicomplexa) 64 Phylum Microsporidea 67 Phylum Myxozoa (= Myxospora) 67 4 The Poriferans and Placozoans 73 Phylum Porifera 73 Introduction 73 General Characteristics 75 Poriferan Diversity 80 Other Features of Poriferan Biology: Reproduction and Development 84 Phylum Placozoa 85 Introduction 85 General Characteristics 85 5 Introduction to the Hydrostatic Skeleton 91 6 The Cnidarians 95 Introduction and General Characteristics 95 Class Scyphozoa 98 Class Cubozoa 102 Class Hydrozoa 103 Order Hydroida 104 Order Siphonophora 107 The Hydrocorals 110 Class Anthozoa 110 Subclass Hexacorallia (= Zoantharia) 113 Subclass Octocorallia (= Alcyonaria) 117 7 The Ctenophores 127 Introduction and General Characteristics 127 Ctenophore Diversity 133 Class Tentaculata 133 Class Nuda 136 Other Features of Ctenophore Biology: Reproduction 136 8 The Platyhelminthes 139 Introduction and General Characteristics 139 Class Turbellaria 141 Class Cestoda 145 Class Monogenea 149 Class Trematoda 149 The Digeneans 150 The Aspidogastreans (= Aspidobothreans) 159 9 The Mesozoans: Possible Flatworm Relatives 169 The Mesozoans 169 Class Orthonectida 170 Class Rhombozoa 171 10 The Nemertines 173 Introduction and General Characteristics 173 Other Features of Nemertine Biology 180 Classification 180 Reproduction and Development 180 11 The Rotifers and Acanthocephalans 185 Introduction 185 Phylum Rotifera 186 Introduction and General Characteristics 186 Class Seisonidea 191 Class Bdelloidea 191 Class Monogononta 191 Other Features of Rotifer Biology 196 Digestive System 196 Nervous and Sensory Systems 196 Excretion and Water Balance 197 Phylum Acanthocephala 197 12 The Molluscs 203 Introduction and General Characteristics 203 Class Polyplacophora 205 Class Aplacophora 210 Class Monoplacophora 210 Class Gastropoda 213 Subclass Prosobranchia 216 Subclass Opisthobranchia 222 Subclass Pulmonata 226 Class Bivalvia (= Pelecypoda) 227 Subclass Protobranchia 228 Subclass Lamellibranchia 231 Subclass Septibranchia 239 Class Scaphopoda 240 Class Cephalopoda 244 Other Features of Molluscan Biology 252 Reproduction and Development 252 Circulation, Blood Pigments, and Gas Exchange 256 Nervous System 256 Digestive System 257 Excretory System 257 13 The Annelids and Sipunculans 277 Introduction 277 Phylum Annelida 277 General Annelid Characteristics 277 Class Polychaeta 279 Polychaete Reproduction 282 Class Clitellata 288 Subclass Oligochaeta 289 Oligochaete Reproduction 291 Subclass Hirudinea 292 Leech Reproduction 294 Class Pogonophora 294 General Characteristics 294 Pogonophoran Reproduction and Development 299 Class Echiura 300 Other Features of Annelid Biology 304 Digestive System 304 Nervous System and Sense Organs 305 Circulatory System 306 Phylum Sipuncula 307 14 The Arthropods 323 Introduction and General Characteristics 323 Subphylum Trilobitomorpha 329 Class Trilobita 329 Subphylum Chelicerata 330 Class Merostomata 330 Class Arachnida 331 Class Pycnogonida (= Pantopoda) 335 Subphylum Mandibulata 335 Class Myriapoda 336 Orders Chilopoda and Diplopoda 336 Class Insecta (= Hexapoda) 337 Class Crustacea 351 Subclass Malacostraca 352 Subclass Branchiopoda 354 Subclass Ostracoda 357 Subclass Copepoda 359 Subclass Pentastomida 365 Subclass Cirripedia 366 Other Features of Arthropod Biology 368 Reproduction and Development 368 Digestion 374 Excretion 374 Blood Pigments 377 15 Two Phyla of Likely Arthropod Relatives: Tardigrada and Onychophora 399 Introduction and General Characteristics 399 Phylum Tardigrada 400 Phylum Onychophora 402 16 The Nematodes 409 Introduction and General Characteristics 409 Body Coverings and Body Cavities 410 Musculature, Internal Pressure, and Locomotion 412 Organ Systems and Behavior 413 Reproduction and Development 415 Parasitic Nematodes 416 Beneficial Nematodes 421 17 Four Phyla of Likely Nematode Relatives: Nematomorpha, Priapulida, Kinorhyncha, and Loricifera 429 Introduction 429 Phylum Nematomorpha 430 Phylum Priapulida 430 Phylum Kinorhyncha (= Echinoderida) 432 Phylum Loricifera 432 18 Five Phyla (Plus One Species) of Uncertain Affiliation: Myxozoa, Gnathostomulida, Gastrotricha, Xenoturbella bocki, Chaetognatha, and Cycliophora 437 Introduction 437 Phylum Myxozoa (= Myxospora) 437 Phylum Gnathostomulida 438 Phylum Gastrotricha 439 Phylum Uncertain 441 Xenoturbella bocki 441 Phylum Chaetognatha 441 General Characteristics and Feeding 441 Chaetognath Reproduction 444 Chaetognath Lifestyles and Behavior 444 Chaetognath Relationships 447 Phylum Cycliophora 448 19 The Lophophorates (Phoronids, Brachiopods, Bryozoans) and Entoprocts 453 Introduction and General Characteristics 453 Phylum Phoronida 455 Phylum Brachiopoda 456 Phylum Bryozoa (= Ectoprocta; = Polyzoa) 460 Class Phylactolaemata 463 Class Gymnolaemata 465 Class Stenolaemata 468 Other Features of Lophophorate Biology 468 Reproduction 468 Digestion 469 Nervous System 470 Phylum Entoprocta (= Kamptozoa) 470 20 The Echinoderms 477 Introduction and General Characteristics 477 Class Crinoidea 480 Class Stelleroidea 483 Subclass Ophiuroidea 483 Subclass Asteroidea 485 Class Concentricycloidea 489 Class Echinoidea 490 Class Holothuroidea 493 Other Features of Echinoderm Biology 498 Reproduction and Development 498 Nervous System 500 21 The Hemichordates 509 Introduction and General Characteristics 509 Class Enteropneusta 510 Class Pterobranchia 512 22 The Nonvertebrate Chordates 517 Introduction and General Characteristics 517 Subphylum Urochordata 518 Class Ascidiacea 518 Class Larvacea (= Appendicularia) 520 Class Thaliacea 522 Other Features of Urochordate Biology 525 Reproduction 525 Excretory and Nervous Systems 525 Subphylum Cephalochordata (= Acrania) 526 23 Invertebrate Reproduction and Development- An Overview 533 Introduction 533 Asexual Reproduction 534 Sexual Reproduction 537 Patterns of Sexuality 537 Gamete Diversity 539 Getting the Gametes Together 539 Larval Forms 544 Dispersal as a Component of the Life-History Pattern 552 Glossary of Frequently Used Terms 563 Index 565
Library of Congress Subject Headings for this publication: Invertebrates