UNIVERSIDADE DE LISBOA FACULDADE DE CIÊNCIAS DEPARTAMENTO DE BIOLOGIA ANIMAL The European Fauna of Annelida Polychaeta (III) João Carlos Ferreira Gil DOUTORAMENTO EM BIOLOGIA (ESPECIALIDADE: BIODIVERSIDADE) Tese orientada por: Professor Associado Doutor Pedro Miguel Alfaia Barcia Ré (FCUL) Doutor Rafael Sardá Borroy (CEAB-CSIC) 2011 Present adress of the author: CEAB-CSIC Carrer d’accés a la Cala Sant Francesc, 14 E-17300 Blanes (Girona) Spain e-mail: [email protected] © João Gil (text and tables) Copyright and ownership of figures and maps as stated at figure legends. Printed by Stein Girona S.L. Plaça Pompeu Fabra, 11 E-17002 Girona (Catalonia, Spain) www.stein.cat July 2011 TABLE OF CONTENTS TABLE OF CONTENTS VOLUME I FOREWORD . xxiii RESUMO . xxv ABSTRACT . xxxi ACKNOWLEDGMENTS . xxxiii INTRODUCTION . 1 GENERAL INTRODUCTION . 3 TAXONOMY AND THE BIODIVERSITY AND TAXONOMIC CRISES . 6 The Biodiversity Crisis and the Holocene Extinction . 6 Taxonomy and the Taxonomic Crisis 8 Morphological versus Molecular Taxonomy? . 14 Natural History Institutions and their Role in the Study of Biodiversity . 16 The Future of Taxonomy . 19 THE STUDY SUBJECT: THE POLYCHAETA . 28 METHODS OF STUDY OF POLYCHAETA . 29 PHYSIOGRAPHY OF THE STUDIED AREAS . 30 a) The Portuguese Southwestern Continental Shelf and Coast . 30 b) The Continental Shelves of South Spain and Gulf of Cádiz . 32 SOME NOTIONS ON THE HYDROGRAPHY OF THE STUDIED REGIONS . 35 1) Mediterranean Sea . 35 2) Alborán Sea . 35 3) Gulf of Cádiz . 37 4) Southwestern Portugal . 39 THE PRESENT WORK: ORIGINAL PROJECT AND FINAL VERSION . 41 The original project . 42 The final version . 43 MATERIAL AND METHODS . 45 THE CAMPAIGNS . 47 a) The SEPLAT Cruises . 47 b) The Fauna 1 Campaign . 48 c) Used sampling gears . 49 d) Resume of SEPLAT and Fauna 1 campaigns . 51 TREATMENT OF THE SAMPLES . 52 a) SEPLAT Cruises . 52 b) Fauna 1 Campaign . 52 iii THE EUROPEAN FAUNA OF ANNELIDA POLYCHAETA IDENTIFICATION OF THE SPECIMENS . 54 BIBLIOGRAPHICAL RESEARCH AND RESOURCES . 61 STRUCTURE OF THE PRESENT WORK . 62 POLYCHAETE FAMILIES . 69 WORLDWIDE KEY TO THE FAMILIES OF POLYCHAETA . 71 FAMILY ABERRANTIDAE . 81 Genus Aberranta . 81 FAMILY ACOETIDAE . 83 Genus Acoetes . 86 Genus Euarche . 87 Genus Eupanthalis . 88 Genus Eupolyodontes . 89 Genus Panthalis . 89 Genus Polyodontes . 90 FAMILY ACROCIRRIDAE . 91 Genus Acrocirrus . 94 Genus Chauvinelia . 94 Genus Flabelligella . 96 Genus Macrochaeta . 98 FAMILY AMPHARETIDAE . 101 Genus Adercodon . 105 Genus Alkmaria . 105 Genus Amage . 106 Genus Amagopsis . 107 Genus Ampharete . 107 Genus Amphicteis . 110 Genus Amythasides . 111 Genus Anobothrus . 112 Genus Auchenoplax . 113 Genus Eclysippe . 115 Genus Glyphanostomum . 115 Genus Hypania . 116 Genus Hypaniola . 116 Genus Isolda . 117 Genus Lysippe . 118 Genus Lysippides . 118 Genus Melinna . 118 Genus Melinnopsis . 122 Genus Mugga . 123 Genus Neosabellides . 123 Genus Noanelia . 124 Genus Sabellides . 124 Genus Samytha . 125 Genus Samythella . 126 Genus Sosane . 126 Genus Sosanopsis . 127 Genus Uschakovius . 127 Genus Ymerana . 128 Genus Zatsepinia . 128 FAMILY AMPHINOMIDAE . 129 Genus Amphinome . 131 Genus Chloeia . 132 Genus Chloenopsis . 134 iv TABLE OF CONTENTS Genus Eurythoe . 134 Genus Hermodice . 135 Genus Hipponoe . 136 Genus Linopherus . 136 Genus Notopygos . 137 Genus Paramphinome . 137 Genus Pareurythoe . 138 FAMILY APHRODITIDAE . 141 Genus Aphrodita . 141 Genus Laetmonice . 144 Genus Pontogenia . 147 FAMILY APISTOBRANCHIDAE . 149 Genus Apistobranchus . 150 FAMILY ARENICOLIDAE . 151 Genus Abarenicola . 152 Genus Arenicola . 153 Genus Arenicolides . 157 Genus Branchiomaldane . 158 FAMILY CAPITELLIDAE . 159 Genus Baldia . 162 Genus Capitella . 162 Genus Capitellethus . 166 Genus Dasybranchus . 167 Genus Heteromastides . 169 Genus Heteromastus . 170 Genus Leiocapitella . 171 Genus Leiochrides . 173 Genus Mastobranchus . 174 Genus Mediomastus . 175 Genus Notomastus . 177 Genus Paracapitella . 182 Genus Peresiella . 182 Genus Pseudocapitella . 183 Genus Pseudonotomastus . 184 Genus Pulliella . 184 Doubtful or Indeterminable Species . 184 FAMILY CHAETOPTERIDAE . 187 Genus Chaetopterus . 188 Genus Mesochaetopterus . 191 Genus Phyllochaetopterus . 192 Genus Spiochaetopterus . 194 FAMILY CHRYSOPETALIDAE . 197 Genus Arichlidon . 198 Genus Bhawania . 199 Genus Chrysopetalum . 199 Genus Dysponetus . 200 Genus Paleanotus . 201 Genus Vigtorniella . 202 FAMILY CIRRATULIDAE . 205 Genus Aphelochaeta . 209 Genus Aphropharynx . 211 Genus Caulleriella . 212 Genus Chaetozone . 214 Genus Cirratulus . 220 Genus Cirriformia . 222 Genus Ctenodrilus . 223 Genus Dodecaceria . 224 Genus Fauvelicirratulus . 226 Genus Monticellina . 227 Genus Protocirrineris . 229 v THE EUROPEAN FAUNA OF ANNELIDA POLYCHAETA Genus Raphidrilus . 229 Genus Raricirrus . 230 Genus Tharyx . 230 Genus Timarete . 231 FAMILY COSSURIDAE . 233 Genus Cossura . 233 FAMILY DORVILLEIDAE . 237 Genus Apharyngtus . 240 Genus Arenotrocha . 241 Genus Dinophilus . 241 Genus Diurodrilus . 244 Genus Dorvillea . 245 Genus Marycarmenia . 248 Genus Ophryotrocha . 248 Genus Ougia . 261 Genus Parapodrilus . 262 Genus Parophryotrocha . 262 Genus Parougia . 263 Genus Pettiboneia . 264 Genus Protodorvillea . 265 Genus Pusillotrocha . 266 Genus Schistomeringos . 267 Genus Trilobodrilus . 268 FAMILY EULEPETHIDAE . 269 Genus Grubeulepis . 270 FAMILY EUNICIDAE . 271 Genus Eunice . 272 Genus Euniphysa . 284 Genus Lysidice . 284 Genus Marphysa . 287 Genus Nematonereis . 290 Genus Palola . 291 Genus Paramarphysa . 292 FAMILY EUPHROSINIDAE . 293 Genus Euphrosine . 293 Genus Palmyreuphrosyne . 297 FAMILY FAUVELIOPSIDAE . 299 Genus Fauveliopsis . 300 Genus Laubieriopsis . 303 FAMILY FLABELLIGERIDAE . 305 Genus Brada . 308 Genus Bradabyssa . 310 Genus Diplocirrus . 310 Genus Flabelliderma . 312 Genus Flabelligera . 312 Genus Ilyphagus . 313 Genus Pherusa . 314 Genus Piromis . 315 Genus Therochaeta . 316 Incertae sedis . 317 FAMILY GLYCERIDAE . 319 Genus Glycera . 321 Genus Glycerella . 329 FAMILY GONIADIDAE . 331 Genus Bathyglycinde . 331 Genus Glycinde . 333 Genus Goniada . 334 Genus Goniadella . 338 Genus Progoniada . 341 FAMILY HESIONIDAE . 343 vi TABLE OF CONTENTS Genus Amphiduros . 345 Genus Dalhousiella . 346 Genus Gyptis . 346 Genus Hesione . 347 Genus Hesionides . 350 Genus Hesiospina . 351 Genus Leocrates . 353 Genus Microphthalmus . 354 Genus Micropodarke . 357 Genus Nereimyra . 358 Genus Ophiodromus . 359 Genus Podarkeopsis . 361 Genus Psamathe . 362 Genus Syllidia . 363 Genus Wesenbergia . 364 Invalid or Indeterminable Species . 364 FAMILY HISTRIOBDELLIDAE . 367 Genus Histriobdella . 368 FAMILY ICHTHYOTOMIDAE . 369 Genus Ichthyotomus . 369 FAMILY LACYDONIIDAE . 371 Genus Lacydonia . 372 FAMILY LONGOSOMATIDAE . 375 Genus Heterospio . 376 FAMILY LUMBRINERIDAE . 379 Genus Abyssoninoe . 381 Genus Aotearia . 385 Genus Augeneria . 386 Genus Cenogenus . 387 Genus Eranno . 388 Genus Helmutneris . 388 Genus Hilbigneris . 389 Genus Lumbricalus . 391 Genus Lumbrinerides . 391 Genus Lumbrineris . 393 Genus Ninoe . 398 Genus Scoletoma . 399 Invalid Species . 402 FAMILY MAGELONIDAE . 403 Genus Magelona . 403 Genus Octomagelona . 411 FAMILY MALDANIDAE . 413 Genus Axiothella . 416 Genus Chirimia . 417 Genus Clymenella . 417 Genus Clymenopsis . 418 Genus Clymenura . 419 Genus Euclymene . 421 Genus Isocirrus . 424 Genus Johnstonia . 424 Genus Lumbriclymene . 424 Genus Macroclymene . 425 Genus Maldane . 426 Genus Maldanella . 428 Genus Metasychis . 429 Genus Microclymene . 429 Genus Micromaldane . 430 Genus Nicomache . 431 Genus Notoproctus . 433 Genus Petaloproctus . 434 vii THE EUROPEAN FAUNA OF ANNELIDA POLYCHAETA Genus Praxillella . 435 Genus Praxillura . 438 Genus Proclymene . 438 Genus Pseudoclymene . 438 Genus Rhodine . 439 FAMILY NAUTILINIELLIDAE . 441 Genus Natsushima . 441 FAMILY NEPHTYIDAE . 443 Genus Aglaophamus . 444 Genus Inermonephtys . 446 Genus Micronephthys . 448 Genus Nephtys . 450 FAMILY NEREIDIDAE . 463 Genus Alitta . 467 Genus Ceratocephale . 470 Genus Ceratonereis . 471 Genus Composetia . 472 Genus Eunereis . 474 Genus Hediste . 475 Genus Leonnates . 476 Genus Micronereis . 477 Genus Namalycastis . 479 Genus Namanereis . 479 Genus Neanthes . 481 Genus Nereis . 486 Genus Perinereis . 490 Genus Platynereis . 495 Genus Pseudonereis . 498 Genus Rullierinereis . 499 Genus Sinonereis . 499 Genus Websterinereis . 499 Addendum . 500 FAMILY NERILLIDAE . 503 Genus Afronerilla . ..