THE SHARKS OF NORTH AMERICA

JOSE I. CASTRO

COLOR ILLUSTRATIONS BY DIANE ROME PEEBLES

OXFORD UNIVERSITY PRESS CONTENTS

Foreword, by Eugenie Clark v Mosaic gulper shark, Centrophorus tesselatus 79 Preface vii Little gulper shark, Centrophorus uyato 81 Acknowledgments ix Minigulper, Centrophorus sp. A 84 Slender gulper, Centrophorus sp. B 85 Introduction 3 Birdbeak dogfish, Deania calcea 86 How to use this book 3 Arrowhead dogfish, Deaniaprofundorum 89 Description of species accounts 3 Illustrations 6 Family Etmopteridae, The Black Dogfishes Glossary 7 and Lanternsharks 91 Bibliography 7 Black dogfish, Centroscyllium fabricii 93 The knowledge and study of sharks 7 Pacific black dogfish, Centroscyllium nigrum 96 The shark literature 8 Emerald or blurred lanternshark, Etmopterus bigelowi 98 Lined lanternshark, Etmopterus bullisi 101 Broadband lanternshark, Etmopterus gracilispinis 103 A KEY TO THE FAMILIES OF Caribbean lanternshark, Etmopterus hillianus 105 NORTH AMERICAN SHARKS 11 Great lanternshark, Etmopterusprinceps 107 Fringefin lanternshark, Etmopterus schultzi 110 SPECIES ACCOUNTS 19 Green lanternshark, Etmopterus virens 112

Family Chlamydoselachidae, The Frill Shark 21 Family Somniosidae, The Sleeper Sharks 115 Frill shark, Chlamydoselachus anguineus 22 Portuguese shark, Centroscymnus coelolepis 117 Roughskin dogfish, Centroscymnus owstoni 120 Family Hexanchidae, The Cowsharks 26 Velvet dogfish, Zameus squamulosus \T1 Sharpnose sevengill, or perlon shark, Heptranchias Greenland shark, Somniosus microcephalus 124 perlo 28 Pacific sleeper shark, Somniosuspacificus 131 Sixgill shark, Hexanchus griseus 31 Little sleeper shark, Somniosus rostratus 135 Bigeye sixgill shark, Hexanchus nakamurai 36 Broadnose sevengill shark, Notorynchus cepedianus 38 Family Oxynotidae, The Rough Sharks 137 Caribbean roughshark, Oxynotus caribbaeus 138 Family Echinorhinidae, The Bramble Sharks 42 Bramble shark, Echinorhinus brucus 44 Family Dalatiidae, The Kitefin Shark, the Prickly shark, Echinorhinus cookei 47 Cookiecutter Sharks, and Dwarf Sharks 140 Kitefin shark, Dalatias licha 142 Family Squalidae, The Dogfishes 50 Cookiecutter shark, Isistius brasiliensis 145 Roughskin spiny dogfish, Cirrhigaleus asper 52 Largetooth cookiecutter shark, Isistius plutodus 151 Spiny dogfish, Squalus acanthias 55 Pygmy shark, Euprotomicrus bispinatus 153 Cuban dogfish, Squalus cubensis 63 Spined pygmy shark, Squaliolus laticaudus 156 Shortspine dogfish, Squalus mitsukurii 65 Family Pristiophoridae, The Sawsharks 159 Family Centrophoridae, The Gulper Sharks and American sawshark, Pristiophorus schroederi 160 Birdbeak Dogfishes 67 Blackfin gulper shark, Centrophorus isodon 71 Family Squatinidae, The Angel Sharks 162 Taiwan gulper shark, Centrophorus niaukang 73 Pacific angel shark, Squatina californica 163 Leafscale gulper shark, Centrophorus squamosus 76 Atlantic angel shark, Squatina dumerili 167

XI Gulf angel shark, Squatina heteroptera 170 Marbled catshark, Galeus arae 323 Mexican angel shark, Squatina mexicana 172 Springer's catshark, Galeus springeri 326 Peppered catshark, Galeuspiperatus 328 Family Heterodontidae, The Bullhead Sharks 174 Campeche catshark, Parmaturus campechiensis 330 Horn shark, Heterodontus francisci 175 Filetail catshark, Parmaturus xaniurus 332 Mexican horn shark, Heterodontus mexicanus 179 Narrowtail catshark, Schroederichthys maculatus 334 Cryptic horn shark, Heterodontus sp. X 182 Boa catshark, Scyliorhinus boa 336 Family Ginglymostomatidae, The Nurse Sharks 183 Western catshark, Scyliorhinus hesperius 338 Nurse shark, Ginglymostoma cirratum 184 Mead's catshark, Scyliorhinus meadi 340 Pacific nurse shark, Ginglymostoma sp. 191 Chain dogfish, Scyliorhinus retifer 342 Oakley's catshark, Scyliorhinus sp. X 346 Family Rhincodontidae, The Whale Shark 193 Dwarf catshark, Scyliorhinus torrei 347 Whale shark, Rhincodon typus 194 Family Proscyllidae, The Ribbontail Catsharks 349 Family Mitsukurinidae, The Goblin Shark 201 Cuban ribbontail catshark, Eridacnis barbouri 350 Goblin shark, Mitsukurina owstoni 202 Family Pseudotriakidae, The False Catshark 352 Family Odontaspididae, The Sand Tigers 206 False catshark, Pseudotriakis microdon 353 Sand tiger, Carcharias taurus 208 Ragged-tooth shark, Odontaspis ferox 215 Family Triakidae, The Smoothhound Sharks 357 Bigeye sand tiger, Odontaspis noronhai 218 Soupfin shark, Galeorhinus galeus 361 Whiterip smoothhound, Mustelus albipinnis 364 Family Pseudocarchariidae, The Crocodile Shark 221 Gray smoothhound, Mustelus californicus 366 Crocodile shark, Pseudocarcharias kamoharai 222 Smooth dogfish, Mustelus canis 369 Family Megachasmidae, The Megamouth Shark 225 Sharptooth smoothhound, Mustelus dorsalis 372 Megamouth shark, Megachasma pelagios 226 Brown smoothhound, Mustelus henlei 374 Sicklefin smoothhound, Mustelus lunulatus 377 Family Alopiidae, The Thresher Sharks 231 Florida smoothhound or Florida dogfish, Mustelus Smalltooth thresher, Alopiaspelagicus 233 norrisi 379 Bigeye thresher, Alopias superciliosus 236 Gulf smoothhound, Mustelus sinusmexicanus 382 Common thresher, Alopias vulpinus 241 Leopard shark, Triakis semifasciata 384

Family Cetorhinidae, The Basking Shark 248 Family Carcharhinidae, The Requiem Sharks 388 Basking shark, Cetorhinus maximus 249 Blacknose shark, Cttrcharhinus acronotus 393 Silvertip shark, Carcharhinus albimarginatus 397 Family Lamnidae, The Mackerel Sharks 256 Bignose shark, Carcharhinus altimus 400 White shark, Carcharodon carcharias 258 Narrowtooth or copper shark, Carcharhinus Shortfin mako, Isurus oxyrinchus 267 brachyurus 403 Longfin mako, Isuruspaucus 274 Spinner shark, Carcharhinus brevipinna 407 Salmon shark, Lamna ditropis 278 Pacific smalltail shark, Carcharhinus cerdale 410 Porbeagle, lamna nasus 284 Silky shark, Carcharhinus falciformis 413 Family Scyliorhinidae, The Catsharks 290 Galapagos shark, Carcharhinus galapagensis 417 Brown catshark, Apristurus brunneus 296 Finetooth shark, Cttrcharhinus isodon 421 Hoary catshark, Apristurus canutus 299 Bull shark, Carcharhinus leucas 426 White-edge or longnose catshark, Apristurus Blacktip shark, Carcharhinus limbatus 432 kampae 301 Oceanic whitetip shark, Carcharhinus longimanus 438 Icelandic catshark, Apristurus laurussoni 303 Dusky shark, Carcharhinus obscurus 444 Ghost catshark, Apristurus manis 305 Caribbean reef shark, Carcharhinus perezii 449 Black roughscale catshark, Apristurus melanoasper 307 Sandbar shark, Carcharhinusplumbeus 453 Smallfin catshark, Apristurusparvipinnis 309 Smalltail shark, Carcharhinusporosus 459 Deep-water catshark, Apristurus profundorum 311 Night shark, Carcharhinus signatus 463 Rivera's catshark, Apristurus riveri 314 Tiger shark, Galeoccrdo cuvier 466 Galbraith's catshark, Apristurus sp. X 316 Daggernose shark, Isogomphodon oxyrhynchus 475 Swell shark, Cephaloscyllium ventriosum 318 Whitenose shark, Nasolamia velox 41% Lollipop catshark, Cephalurus cephalus 321 Lemon shark, Negaprion brevirostris 481

XII CONTENTS Blue shark, Prionace glauca 486 Golden or smalleye hammerhead, Sphyrna tudes 525 Pacific sharpnose shark, Rhizoprionodon longurio 493 Smooth hammerhead, Sphyrna zygaena 529 Atlantic sharpnose shark, Rhizoprionodon Appendix 1 533 terraenovae 496 A primer on shark reproduction 533 Whitetip reef shark, Triaenodon obesus 501 Appendix 2 547 Family Sphyrnidae, The Hammerhead Sharks 504 An essay on the conservation of sharks 547 Scalloped bonnethead, Sphyrna corona 507 References Cited 551 Scalloped hammerhead, Sphyrna lewini 509 Glossary 601 Scoophead, Sphyrna media 514 Great hammerhead, Sphyrna mokarran 516 Scientific Names 607 Bonnethead, Sphyrna tiburo 521 Common Names 611

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