Extended Contents

Editors and Contributors xiv 2: Ecology and Behaviour 37 Echinoderm habitats 37 Contributors xiv Intertidal rock platforms 38 Illustrators and Photographers xv Embayments 39 Foreword xxi Reefs 42 Continental shelf and slope 44 Seamounts, vents and the abyssal plain 45 Part 1: Introduction to Echinoderms 1 Interactions 48 1: Introduction 2 Feeding 48 Echinoderms, what are they? 2 Predators 48 How to recognise the five classes 4 Diseases 50 Their place in the scheme of things 6 Defence against predators 51 Anatomy 7 Echinoderm parasites and associates 58 Body wall and skeleton 7 Parasites 58 Feeding and digestive system 13 Commensal associations: Asteroidea 14 excluding Crustacea 61 Ophiuroidea 15 Commensal associations: Crustacea 64 Echinoidea 19 Burrowing and boring 70 Holothuroidea 22 Ecological engineers 71 Crinoidea 24 Box 2.1 Urchin barrens and abalone 46 Water vascular system 25 Box 2.2 Bioluminescence 55 Respiration and excretion 27 Haemal system 30 3: Echinoderm Life Histories 75 Nervous system 30 Sexual reproduction 78 Sensory structures 34 Reproductive anatomy 80 Locomotion 35 Asexual reproduction 84 Box 1.1 Mutable connective tissues: Echinoderm gametes 86 autotomy and catch 10 Reproductive cycles and temporal cues 86 Box 1.2 Ophiarachna incrassata, Development 89 the fishing ophiuroid 17 Box 3.1 Life history, evolution and speciation Box 1.3 Regeneration 29 76 Box 1.4 Sensory receptors and Box 3.2 Northern Pacific Seastar — colour change 31 seasonal reproduction in the Box 1.5 Swimming, hitchhiking and Northern and Southern Hemispheres 87 pelagic echinoderms 32

ix Australian Echinoderms: Biology, Ecology & Evolution

4: Management: Fisheries, Ferals and Table 5.1 Species richness and Conservation 95 endemicity of shallow‑water Fisheries 95 echinoderms around Australia 140 Sea cucumbers – Trepang 95 Box 5.1 Rafting to remote Sea urchins 105 Macquarie Island 142 Introduced species 112 New Zealand Cushion Star 113 6: Phylogeny and Geological History 155 New Zealand Seven-armed Seastar 113 The phylogenetic relationships Northern Pacific Seastar 113 of the echinoderm classes 160 Box 4.4 Crown‑of‑thorns seastar 117 A brief overview of the fossil record Crown‑of‑thorns seastar 119 of echinoderms with reference Discovery 120 to Australia 164 Spread of the outbreaks 122 Box 6.1 Carpoids and their importance Causes 123 to echinoderm phylogeny 158 Current situation 125 Conservation issues 126 Introduced species 126 Part 2: Echinoderm Diversity 169 Trawling and dredging 131 7: Class CRINOIDEA 171 Over-exploitation 132 General Introduction 171 Water quality 133 Fossils 188 Climate Change 134 Phylogeny 188 Table 4.1 Commercial holothuroids Order HYOCRINIDA 189 in the waters of tropical Australia 98 Family Hyocrinidae 189 Table 4.2 Standard names used for Order CYRTOCRINIDA 191 the three commercial species of Order ISOCRINIDA 192 sea urchins 105 Family Isocrinidae 193 Box 4.1 The changing names of Family Cainocrinidae 193 sea cucumbers 96 Family Isselicrinidae 194 Box 4.2 New Zealand cushion star 108 Family Proisocrinidae 195 Box 4.3 Northern Pacific seastar 110 Order COMATULIDA 198 Box 4.5 Tasmanian cushion star 127 Family Comatulidae 199 Intertidal habitat destruction 129 Superfamily HIMEROMETROIDEA 202 Box 4.6 Seamount biological Family Himerometridae 203 communities and fishing 130 Family Colobometridae 204 Family Mariametridae 206 5: Biogeography 137 ‘Superfamily TROPIOMETROIDEA’ 208 Biogeographical patterns 138 Family Tropiometridae 209 Northern Australia 138 Family Calometridae 209 Southern Australia 140 Family Ptilometridae 210 Subantarctic Islands and Antarctica 141 Family Asterometridae 211 Diversity and endemism 141 Family Charitometridae 212 Historical explanations 141 Family Thalassometridae 212 Origin of the shallow-water Superfamily NOTOCRINOIDEA 213 Australian marine fauna 145 Family Notocrinidae 213 Diversity of the Australian Fauna 151 Family Aporometridae 214 x Contents

‘Superfamily ANTEDONOIDEA’ 215 Family Ophidiasteridae 265 ‘Family Antedonidae’ 215 Family Oreasteridae 267 Family Zenometridae 218 Family Podosphaerasteridae 272 Family Pentametrocrinidae 220 Family Poraniidae 272 Superfamily Incertae sedis 220 Family Solasteridae 274 Family Atopocrinidae 220 Order VELATIDA 275 Family Eudiocrinidae 221 Family Caymanostellidae 275 ‘Bourgueticrinines’ 221 Family Korethrasteridae 277 Family Bourgueticrinidae 223 Family Myxasteridae 277 Family Bathycrinidae 223 Family Pterasteridae 278 Family Phrynocrinidae 224 Order SPINULOSIDA 279 Family Septocrinidae 224 Family Echinasteridae 279 Family Atelecrinidae 225 Order FORCIPULATIDA 281 Family Guillecrinidae 226 Family Asteriidae 282 Incertae sedis 227 Family Heliasteridae 285 Family Caledonicrinidae 227 Family Pedicellasteridae 286 Table 7.1 Classification of living Crinoidea 185 Family Stichasteridae 286 Table 7.2 Recent taxonomic Family Zoroasteridae 289 arrangements of order Isocrinida 194 Order BRISINGIDA 290 Box 7.1 Associates and symbionts 182 Family Brisingidae 290 Box 7.2 The curious case of Aporometra 216 Family Freyellidae 292 Incertae sedis 292 8: Class ASTEROIDEA 231 Family Xyloplacidae 292 General Introduction 231 Table 8.1 Classification of the Asteroidea 237 Order PAXILLOSIDA 238 Box 8.1 Two common, but easily Family Astropectinidae 239 confused, little biscuit stars 260 Family Benthopectinidae 241 Box 8.2 Odontaster validus, the Antarctic Family Ctenodiscididae 242 seastar 264 Family Goniopectinidae 243 Box 8.3 The black and white seastar 287 Family Luidiidae 243 Family Porcellanasteridae 245 9: Class OPHIUROIDEA 295 Family Pseudarchasteridae 245 General description 295 Family Radiasteridae 246 Australian Fossil Record 302 Order VALVATIDA 247 EURYALIDA 304 Family Acanthasteridae 248 Family Euryalidae 305 Family Archasteridae 249 Family Gorgonocephalidae 310 Family Asterinidae 250 Family Asteronychidae 312 Family Asterodiscididae 253 Other Families 314 Family Asteropseidae 254 Families Ophiuridae and Family Chaetasteridae 255 Astrophiuridae 314 Family Ganeriidae 256 Ophiomusium and relatives 317 Family Goniasteridae 257 Ophioscolex, Ophiomyces and Family Mithrodiidae 262 relatives 320 Family Odontasteridae 263 Family Ophiacanthidae 321

xi Australian Echinoderms: Biology, Ecology & Evolution

Family Ophiocomidae 326 Family Toxopneustidae 389 Family Ophiodermatidae Family Echinometridae 394 (including Ophiomyxa) 328 Order Echinoneoida 403 Family Ophioleucidae 331 Family Echinoneidae 403 Family Ophionereididae 332 NEOGNATHOSTOMATES 405 Ophiopsila 334 Family Apatopygidae 407 Amphilimna 335 Family Cassidulidae 409 Family Ophiolepididae 336 Family Neolampadidae 410 Family Amphiuridae 338 Family Echinolampadidae 411 Family Ophiactidae, Ophiothamnus Family Clypeasteridae 413 and relatives 344 Family Arachnoididae 415 Family Ophiotrichidae 347 Family Fibulariidae 416 Table 9.1 A comparison of the Family Echinocyamidae 417 Ophiuroidea classification 299 Family Laganidae 418 Table 9.2. Fossil ophiuroids recorded Family Astriclypeidae 419 from Australia 302 ATELOSTOMATES 420 Box 9.1 The grasping snake star, Family Corystusidae 422 Astrobrachion constrictum 308 Family Micrasteridae 423 Box 9.2 The brooding brittle star, Family Hemiasteridae 424 Amphipholis squamata 342 Family Schizasteridae 425 Family Pericosmidae 430 10: Class ECHINOIDEA 351 Family Brissidae 433 General introduction 351 Family Asterostomatidae 434 Fossils 361 Family Spatangidae 436 CIDAROIDS 363 Family Eupatagidae 437 Family Histocidaridae 364 Family Maretiidae 439 Family Cidaridae 365 Family Loveniidae 440 Family Psychocidaridae 367 Table 10.1 Classification of echinoid ECHINOTHURIOIDS 368 families in Australian waters 357 Family Phormosomatidae 368 Box 10.1 Tetramerous and hexamerous Family Echinothuriidae 371 symmetry in regular sea urchins 352 DIADEMATOIDS and PEDINOIDS 372 Box 10.2 Canopy dwelling sea urchins 384 Family Micropygidae 372 Box 10.3 The Echinometra ‘mathaei’ Family Aspidodiadematidae 373 species complex 396 Family Diadematidae 374 Box 10.4 A tale of two urchins: Family Pedinidae 378 Heliocidaris erythrogramma and STIRODONTS 379 H. tuberculata 400 Family Stomopneustidae 379 Box 10.5 Family Arbaciidae 380 Abatus from the Australian Family Saleniidae 381 Antarctic Territory 428 CAMARODONTS 383 Box 10.6 The common heart urchin, Family Temnopleuridae 383 Echinocardium cordatum 442 Family Trigonocidaridae 388 Family Echinidae 389

xii Contents

11: Class HOLOTHUROIDEA 447 General description 447 Order DENDROCHIROTIDA 455 Family Paracucumidae 456 Family Psolidae 457 Family Phyllophoridae 459 Family Sclerodactylidae 461 Family Cucumariidae 462 Family Ypsilothuriidae 465 Order ASPIDOCHIROTIDA 466 Family Holothuriidae 467 Family Stichopodidae 469 Family Synallactidae 472 Order ELASIPODIDA 473 Family Laetmogonidae 474 Family Elpidiidae 475 Family Psychropotidae 477 Family Pelagothuriidae 478 Family Deimatidae 479 Order SYNAPTULIDA 480 Family Chiridotidae 481 Family Synaptidae 483 Family Myriotrochidae 485 Order MOLPADIDA 486 Family Molpadiidae 487 Family Caudinidae 489 Family Gephyrothuriidae 489 Table 11.1 Classification of the Holothuroidea 449 Box 11.1 Neocnus bimarsupiis 464

References 491 Index 577

xiii