THE DIVERSITY OF FISHES Dedications: To our parents, for their encouragement of our nascent interest in things biological; To our wives – Judy, Sara, Janice, and RuthEllen – for their patience and understanding during the production of this volume; And to students and lovers of fishes for their efforts toward preserving biodiversity for future generations Front cover photo: A Leafy Sea Dragon, Phycodurus eques, South Australia Well camouflaged in their natural, heavily vegetated habitat, Leafy Sea Dragons are closely related to seahorses (Gasterosteiformes: Syngnathidae) “Leafies” are protected by Australian and international law because of their limited distribution, rarity, and popularity in the aquarium trade Legal collection is highly regulated, limited to one “pregnant” male per year See Chapters 15, 21, and 26 Photo by D Hall, www.seaphotos.com Back cover photos (from top to bottom): A school of Blackfin Barracuda, Sphyraena qenie (Perciformes, Sphyraenidae) Most of the 21 species of barracuda occur in schools, highlighting the observation that predatory as well as prey fishes form aggregations (Chapters 19, 20, 22) Blackfins grow to about m length, display the silvery coloration typical of water column dwellers, and are frequently encountered by divers around Indo-Pacific reefs Barracudas are fast-start predators (Chapter 8), and the pan-tropical Great Barracuda, Sphyraena barracuda, frequently causes ciguatera fish poisoning among humans (Chapter 25) Longhorn Cowfish, Lactoria cornuta (Tetraodontiformes: Ostraciidae), Papua New Guinea Slow moving and seemingly awkwardly shaped, the pattern of flattened, curved, and angular trunk areas made possible by the rigid dermal covering provides remarkable lift and stability (Chapter 8) A Silvertip Shark, Carcharhinus albimarginatus (Carcharhiniformes: Carcharhinidae), with a Sharksucker (Echeneis naucrates, Perciformes: Echeneidae) attached This symbiotic relationship between an elasmobranch (Chapter 12) and an advanced acanthopterygian teleost (Chapter 15) probably benefits both, the Sharksucker scavenging scraps from the shark’s meals and in turn picking parasitic copepods off the shark Remoras also attach to whales, turtles, billfishes, rays, and an occasional diver Remoras generate sufficient suction to hang on even at high speeds via a highly modified first dorsal fin A recently discovered 10 cm long Indonesian antennariid, nicknamed the Psychedelic Frogfish (Lophiiformes: Antennariidae) (Chapters 14, 18) Among its atypical traits are its shallow water habitat, a lack of an illicial lure, jet propulsion, and a bouncing method of movement, and its practice of hiding in holes, not to mention the spectacular head and body coloration A mating pair of Mandarinfish, Synchiropus splendidus (Perciformes: Callionymidae), Indonesia These small (6 cm), secretive dragonets live among coral branches or rubble, and usually emerge just after sunset to mate Recently extruded eggs can be seen just below the pair Lionfish, Pterois volitans (Scorpaeniformes: Pteroidae), are native to the Indo-Pacific region They have been introduced along the southeastern coast of the USA and the Bahamas, apparently due to aquarium releases In their native habitats they seldom reach high densities but have undergone a population explosion on Bahamian reefs Atlantic reef fishes are naive to lionfish predatory tactics, and predation rates by lionfish are high Photos by D Hall, www.seaphotos.com Gene S Helfman Bruce B Collette Douglas E Facey Brian W Bowen Second Edition THE DIVERSITY OF FISHES Biology, Evolution, and Ecology A John Wiley & Sons, Ltd., Publication This edition first published 2009, © 2009 by Gene S Helfman, Bruce B Collette, Douglas E Facey, and Brian W Bowen Blackwell Publishing was acquired by John Wiley & Sons in February 2007 Blackwell’s publishing program has been merged with Wiley’s global Scientific, Technical and Medical business to form Wiley-Blackwell Registered office: John Wiley & Sons Ltd, The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK Editorial offices: 9600 Garsington Road, Oxford, OX4 2DQ, UK The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK 111 River Street, Hoboken, NJ 07030-5774, USA For details of our global editorial offices, for customer services and for information about how to apply for permission to reuse the copyright material in this book please see our website at www.wiley.com/wiley-blackwell The right of the author to be identified as the author of this work has been asserted in accordance with the Copyright, Designs and Patents Act 1988 All rights reserved No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, except as permitted by the UK Copyright, Designs and Patents Act 1988, without the prior permission of the publisher Wiley also publishes its books in a variety of electronic formats Some content that appears in print may not be available in electronic books Designations used by companies to distinguish their products are often claimed as trademarks All brand names and product names used in this book are trade names, service marks, trademarks or registered trademarks of their respective owners The publisher is not associated with any product or vendor mentioned in this book This publication is designed to provide accurate and authoritative information in regard to the subject matter covered It is sold on the understanding that the publisher is not engaged in rendering professional services If professional advice or other expert assistance is required, the services of a competent professional should be sought Library of Congress Cataloguing-in-Publication Data The diversity of fishes / Gene Helfman [et al.] – 2nd ed p cm Rev ed of: The diversity of fishes / Gene S Helfman, Bruce B Collette, Douglas E Facey c1997 Includes bibliographical references ISBN 978-1-4051-2494-2 (hardback : alk paper) I Helfman, Gene S II Helfman, Gene S Diversity of fishes QL615.H44 2009 597.13′8–dc22 2008029040 A catalogue record for this book is available from the British Library Set in 9.5 on 12 pt Classical Garamond BT by SNP Best-set Typesetter Ltd., Hong Kong Printed in Malaysia 2009 Brief contents BRIEF CONTENTS Full contents vii Preface to the second edition xi Preface to the first edition xii Phylogenetic relationships among living and extinct fish groups xv Part I Introduction The science of ichthyology Systematic procedures 11 Part II Form, function, and ontogeny 21 10 Skeleton, skin, and scales 23 Soft anatomy 41 Oxygen, metabolism, and energetics 57 Sensory systems 75 Homeostasis 91 Functional morphology of locomotion and feeding 111 Early life history 129 Juveniles, adults, age, and growth 149 Part III Taxonomy, phylogeny, and evolution 167 11 12 13 “A history of fishes” 169 Chondrichthyes: sharks, skates, rays, and chimaeras 205 Living representatives of primitive fishes 231 14 15 Teleosts at last I: bonytongues through anglerfishes 261 Teleosts at last II: spiny-rayed fishes 291 Part IV Zoogeography, genetics, and adaptations 327 16 17 18 Zoogeography 329 Fish genetics 355 Special habitats and special adaptations 393 Part V Behavior and ecology 19 20 21 22 23 24 25 Fishes as predators 425 Fishes as prey 439 Fishes as social animals: reproduction 455 Fishes as social animals: aggregation, aggression, and cooperation 477 Cycles of activity and behavior 499 Individuals, populations, and assemblages 525 Communities, ecosystems, and the functional role of fishes 551 Part VI The future of fishes 26 423 583 Conservation 585 References 625 Index 693 v CONTENTS Preface to the second edition xi Preface to the first edition xii Phylogenetic relationships among living and extinct fish groups xv Part I Introduction Part II Form, function, and ontogeny 21 1 The science of ichthyology What is a fish? Superlative fishes A brief history of ichthyology Additional sources of information Summary Systematic procedures Collections 18 Summary 19 Supplementary reading 19 11 Species 11 Taxonomy versus systematics 12 Approaches to classification 12 Taxonomic characters 14 Vertebrate classes 15 Units of classification 16 International Code of Zoological Nomenclature 16 PhyloCode 17 Name changes 17 Skeleton, skin, and scales 23 Skeleton 23 Integumentary skeleton 36 Summary 40 Supplementary reading 40 Soft anatomy 41 Muscles 41 Cardiovascular system 45 Alimentary canal 48 Gas bladder 50 Kidneys 52 Gonads 52 Nervous system 54 Summary 56 Supplementary reading 56 vii viii Contents Oxygen, metabolism, and energetics 57 Respiration and ventilation 57 Gas transport 64 Metabolic rate 66 Energetics 68 Summary 73 Supplementary reading 73 Sensory systems 75 Summary 147 Supplementary reading 148 10 Juveniles, adults, age, and growth 149 Juveniles 149 Adults 153 Age and growth 157 The ontogeny and evolution of growth 162 Summary 164 Supplementary reading 165 Mechanoreception 75 Electroreception 80 Vision 84 Chemoreception 87 Magnetic reception 89 Summary 89 Supplementary reading 90 Homeostasis 91 Coordination and control of regulation 91 Temperature relationships 94 Osmoregulation, excretion, ion and pH balance 100 The immune system 105 Stress 106 Summary 108 Supplementary reading 109 Functional morphology of locomotion and feeding 111 Locomotion: movement and shape 111 Feeding: biting, sucking, chewing, and swallowing 119 Summary 127 Supplementary reading 128 Early life history 129 Complex life cycles and indeterminate growth 129 Early life history: terminology 130 Eggs and sperm 130 Embryology 137 Larvae 139 Getting from here to there: larval transport mechanisms 145 Part III Taxonomy, phylogeny, and evolution 11 “A history of fishes” 167 169 Jawless fishes 170 Gnathostomes: early jawed fishes 175 Advanced jawed fishes I: teleostomes (Osteichthyes) 178 Advanced jawed fishes II: Chondrichthyes 197 A history of fishes: summary and overview 200 Summary 203 Supplementary reading 204 12 Chondrichthyes: sharks, skates, rays, and chimaeras 205 Subclass Elasmobranchii 205 Subclass Holocephali 227 Summary 229 Supplementary reading 230 13 Living representatives of primitive fishes 231 Jawless fishes: lancelets, hagfishes, and lampreys 231 Contents ix Primitive bony fishes 241 Conclusions 258 Summary 258 Supplementary reading 259 17 Fish genetics 14 Teleosts at last I: bonytongues through anglerfishes 261 Teleostean phylogeny 261 A survey of living teleostean fishes 263 Neognathi 280 Neoteleostei 281 Acanthomorpha: the spiny teleosts 284 Summary 289 Supplementary reading 290 355 Fish genomics 355 Molecular ecology 360 Population genetics 365 Phylogeography 370 Molecular evolution 379 Conservation genetics 385 Summary 389 Supplementary reading 390 18 Special habitats and special adaptations 393 15 Teleosts at last II: spiny-rayed fishes 291 Superorder Acanthopterygii: introduction 291 Series Mugilomorpha 292 Series Atherinomorpha 293 Series Percomorpha: basal orders 296 Series Percomorpha, Order Perciformes: the perchlike fishes 300 Series Percomorpha: advanced percomorph orders – flatfishes and twisted jaws 322 Summary 325 Supplementary reading 326 The deep sea 393 The open sea 401 Polar regions 405 Deserts and other seasonally arid habitats 410 Strong currents and turbulent water 415 Caves 417 Summary 420 Supplementary reading 421 Part V Behavior and ecology 423 Part IV Zoogeography, genetics, and adaptations 16 Zoogeography 329 Marine fishes 329 Freshwater fishes 339 Summary 354 Supplementary reading 354 19 Fishes as predators 327 Search and detect 425 Pursuit 426 Attack and capture 429 Handling 433 Scavengers, detritivores, and herbivores 436 Optimally foraging fishes 437 Summary 437 Supplementary reading 438 425 706 Index prey crushing 123, 196 placoderm 178 protrusion 119–22, 195–6 sharks 217 salmoniforms 269 sharks 209–10, 216–17 suspension 29, 30 teleosts 193 see also Gnathostomata superclass; pharyngeal jaws jellynose fish 282 John Dory 297 Jordan, David Starr Jordan’s rule 138 junior synonym 17 juveniles 149–53 coral reef fishes 548, 549 food acquisition 149–50 habitat preferences 549 intermediate stages 144 transient 142 transitions/transitional stages 149–50, 548 karyotype 355, 356 keels 403 Kelp Bass 362 kelp beds 559 fish feces production 572 fish–urchin–algae interactions 562 zonation 540 kelpfishes 316 gene allele frequency 534–5 kidney tubules 101, 409 kidneys 52 aglomerular 52, 99, 409 urine formation 103–4 Antarctic notothenioids 409 elasmobranch 102 glomeruli reduction 409–10 nephrons 101, 102 osmoregulation 101, 103 killifishes 294, 295 salinity range tolerance 411 species flock 310 tidal flooding 507 trematode parasites 554 kinship, larvae 361–2 Kissimmee River (US) channelization 591 Kissing Gourami 320 Kitefin Shark 218 knifefishes, gymnotiform 274, 275 electrical communication 83–4, 274, 485 swimming 115 knotting eels 434–5 hagfishes 235 kraemeriids 317 Kryptolebias (cyprinodontiform) 458–9 kuhliids 306 Kuroshio current 617 kurtids, male care of eggs 318 Kurtoidei 318 Kyoga, Lake (East Africa) 599 kyphosids 305 Labridae 312, 383 labriform swimming 115 Labroidei 307, 308–12, 312–13 Labroides dimidiatus (Cleaner Wrasse) 458, 461, 492, 493 labyrinth fishes 320–1 lactariids 303 lactate 57 lactate dehydrogenase allozymes 95 Lactoria cornuta (Longhorn Cowfish) 20, 21 lagena 89 Lake Trout distribution 373–4 diversity 365 restoration 620 Lake Whitefish distribution 374 gill net fishery 611–12 lakes cannibalism 532 epilimnetic region 568 fecal material in nutrient budget 572 genetically distinct populations 534 hypolimnetic region 568 metalimnetic region 568 phosphorus impact on primary production 570 phytoplanktivory 559 plankton biomass 568 plant biomass 558–9 plant production 566, 567 primary production 570 proglacial 373 seasonal mixing depths 568 temperate 558, 559 temperature 568 thermocline placement 568 trophic cascades 566–9 tropical 556 whiting events 567–8 Lamna nasus (Porbeagle), exploitation 225 Lampetra minima (Miller Lake Lamprey) 588 lampreys 234, 237–40, 241 alimentary canal 49 anatomy 239 blood circulation 239 brook 238 control strategies 239 dentition 240 distribution 239–40 dwarf 238 evolution 163 evolutionary relationships with hagfish 234 fossil species 240 gills 59, 60 heterochronic shifts 163 lineage 381 muscles 41 nonparasitic species 240, 241 osmoregulation 101, 102 ovaries 54 paired species 240 parasitic species 238, 240, 241, 492 reproduction 237–8, 239 senescence 157 skull 25 smell organs 87 spawning 239 taxonomy 240 testis 53 Lampriomorpha 284, 285 Lanao, Lake (Philippines) 309–10 lancelets 171, 231–3 body structure 232–3 cladistic analysis 233 lancetfish 282–383 landscape conservation 619 riverscape 542 lanternfishes 284, 396 Antarctic species 408 lapillus 78 Largemouth Bass competition 532 introductions 599 nutrient cycling 573 Largetooth Sawfish 212 larvae 139–45 abundance 547 behavior 141–3 conditions for growth/survival 511 coral reef fishes 141, 142, 514 settlement 547–8 sound perception 146 demersal 139 development temperature 141 dispersal 141, 530 diversity 144–5 electric organ 143 eyes 143 feeding 139–41, 146 flatfish 152, 153 food availability 140 growth rate 528 habitat preferences 549 herring 160–1 interval length 528 kinship 361–2 leptocephalus 264, 266, 521 mimicry 144 morphology 143–5 mortality 140–1, 548 movement to nursery areas 145–6 navigation 361 non-dispersers 141 organ system functions 143 passive transport 145 pearlfishes 287 pelagic 139, 141, 361 duration 368, 369 physiology 141–3 pigmentation patterns 144 planktonic 405 point-of-no-return 140 predator avoidance 143, 144 prey detection ability 143 priority effects 547 scutatus 144–5 selective tidal stream transport 146 settling 150, 547–8 sibling groups 361–2 size 527 spiny structures 144 startle response 143 starvation risk 532 survival 139–41 Index swimming 142, 145–7, 361 taxonomy 143–5 timing of metamorphosis 163 transport mechanisms 145–7 vexillifer 287 vision 143 vulnerability 532 yolk sac 139 larval biology hypotheses of spawning 509 larval stage 129–30, 139 variability of period 150 lateral line 38, 75, 77 habitat use 78 hair cells 75, 76–7 neuromasts 400 lateral line scales 38 Lates cf niloticus (Nile Perch) 599, 600 latids 300 Latimeria chalumnae (Coelacanth) 242 Laurasia supercontinent 171, 333 Laurentian Great Lakes 310–11 leaffishes 306 Leafy Sea Dragon 2, leatherjackets 324 Leedsichthys problematicus (Pachycormiformes) 191 lekking 457 lens 85 Lepidogalaxias salamandroides (Salamanderfish) 279, 280, 413 Lepidosiren paradoxa (lungfish) 245–6, 247 lepisosteiforms 191 Lepisosteus (gar) 255 Lepisosteus platyrhincus (Florida Gar) 255 Lepomis (sunfish) 542–3 Lepomis macrochirus (Bluegill Sunfish) 474 leptocephalus larva 264, 266, 521 leptocercal tail 33 leptolepidiforms 191 Lessepsian migrants 338 Leuresthes tenuis (grunion) 507–8 Leydig’s gland 53 life cycle, complex 129–30 life history 525–9 characteristics 525–6, 610–11 dangerous stages 532 early 129–47 scale morphology 39 terminology 130, 131 traits 525–9 cave-dwelling fish 418 evolution impact 611 exploitation effects 526, 527 life history theory 610–11 life span elasmobranchs 215–16 life stage prejuveniles 144 thermal preference 99 see also adult fishes; juveniles; larvae life tables 529–30 ligaments 44 light 84–5 activity cycles 499, 500, 501–2, 503–4 attenuation with depth 442 bathypelagic zone 397, 398 decline at twilight 503 707 deepsea fishes 397–8 distribution in water 441–2 mesopelagic zone 398 polarized 86, 518 light–dark cycles 92 light emission, ponyfishes 478–9 light organs see photophores light production 6, 36 lightfishes 282 limestone caves 417 lineages of fish 381 linear hierarchies 486 Lingcod 300 Linnaeus Lionfish 300, 582, 583 lipids 70, 396 body density 70 deepsea fish 397 low-density in elasmobranchs 70, 396 storage 71 Lipophrys pholis (Shanny) 506 lips egg carrying 471 vascularized 412 littoral fishes 329, 331 live-bearers 133, 294 gynogenetic 136, 364, 456, 460 hybridogenetic 456, 460 parthenogenetic 156, 460 see also viviparity live birth 138 liver 50 deepsea fish 396, 397 low-density lipids in elasmobranchs 70, 396 sharks 396 lizardfish 282, 283 loaches 269, 271 Palearctic region 346 lobotids 304 locomotion activity 161 aerial 116 body shape 111–19, 161 carangids 304 coelacanths 243 elasmobranchs 212–14 palaeoniscoids 190 palindromic 538–9 shark swimming 117–19 specialized 116–17 terrestrial 116 types 113–16 locus 360 logging see deforestation longevity 526 adult fish 156 Longhorn Cowfish 2, Longnose Dace, habitat choice 544 lophiiforms 287–8 Lota lota (Burbot) 286 Louvar 12, 318 low-density lipids 70, 396 low-flow conditions 577–8 Luciocephalus pulcher (Pikehead) 121, 196 luminous organ Flashlight Fish 479, 483 see also photophores lumpfishes 300 lumping (taxonomic) 17 lunar patterns of activity 507–9 lungfishes air breathing 63, 182, 246, 248 ancestry 183 characteristics 182, 250 development 248 diphycercal tail 195 distribution 348 diversification 182 electrosensitivity 248 estivation 182, 246–7, 248, 412 evolution 181, 182 external gills 244, 248 living 245–8, 249 osmoregulation 102 pelvic appendages 184 teeth 246 toothplates 246 lungs 62 lungfish 63 lures 428 bioluminescent 399, 400 deepsea fishes 399 lutjanids 304 Lutjanus campechanus (Red Snapper) Luvaridae 318 Luvarus imperialis (Louvar) 12 lymphatic system 47 mackerels 319 distribution 330, 332, 333 geminate species 335 invading species 338 Maco Shark, food consumption 215 macrophages 106 aggregates 107 magnetic field detection 89 migration 518, 519 magnetic field of Earth 84 magnetic reception 89 magnetite 89 shark inner ear 221–2 mahimahi 303, 304 Makaira nigricans (Blue Marlin) 377 malacanthids 303 Malawi, Lake (East Africa) 384–5 male fish attachment to females 397 care-giving 471–3, 474 egg carrying 462 nest guarding 363, 470, 472, 473 parental 471–3, 474 pouch-brooding 364, 462, 469 satellite 474 sex-reversed 606 sexual dimorphism 396–7, 462 sneaker 363, 474, 475 management units 365 Mandarinfish 422, 423 mandibular arch 28, 29 mangroves 506 manta rays body form 206 feeding habits 216, 220 Marine Aquarium Council 615 388 708 Index marine fishes 329–38 air breathing 63–4 aquarium trade 614–15 biogeographic barriers 376–7, 378–9, 379 cold temperatures 98 ecological divisions 329–31 genetic analysis 534 habitats 329–31 herbivores 559 high-energy zones 415 osmoregulation 102, 103–4 phylogeography 376–7, 378–9, 379 seasonal activity 510–11 storm effects 578–9 vulnerable species 589 marine protected areas (MPAs) 569, 619, 620 marine zoogeographic regions 331–2, 333, 334–5, 336–7, 338 market force-free species 612 markings, ultraviolet light-reflective 86 marlins 319 spears 430 marshes coastal 617 intermittent 413 saltmarshes 506, 574 mass extinctions 202, 585 Match–Mismatch Hypothesis 140, 515 mate choice 461, 461–2, 463–5 mate location 88 maternity, multiple 361 mating frequency 455, 456 systems 456, 457, 473–5 tactile communication 485 matrotrophy 469 maturation adult fish 153, 154, 156 age at 527 salmon 162 size at 526 Mauthner cells 143 maxilla 121 maximum likelihood (ML) 380, 383 maximum sustainable yield (MSY), sharks 225 mechanoreception 75–80 compensation for poor visual environment 78 sharks 221 Meckel’s cartilage 28 Meda fulgida (Spikedace) 543 Medaka 295 transgenic 360 median fins 35 Mediterranean Sea 335, 338 medusafishes 320 Megalops atlanticus (Atlantic Tarpon) 265–6, 377 Megamouth Shark, feeding habits 216, 220 Megatooth Shark 209–10 melanophore-stimulating hormone 94 melanophores 94 melanotaeniids 295 melatonin 54, 92, 505 memory cells 106 menhadens 267 Menidia menidia (Atlantic Silverside) 155–6, 295 meristic characters 14, 15 variation 138–9 mermaid purses 224 Mesa Central lakes (Mexico) 310 mesencephalon 54 mesonephric ducts 53 mesonephric tubules 53 mesonephros 52 mesopelagic zone 87, 329, 393, 394 Antarctic fishes 408 diurnal vertical migration 396 gas bladders of deepsea fish 396 light 398 teleost taxa 395 mesorchia 52 mesovaria 53 Mesozoic era 169, 170 Messinian Salinity Crisis 335, 338 metabolic intensity 67 metabolic rate 66–8 hagfish 235–6 maximum 67 routine 67 sharks 214–15 standard 66 water temperature 616 metabolic scope 67 metalimnion 568 metamorphosis thyroxin 92 timing 163 metanephros 52 metapopulations 535 metencephalon 54, 55 Mexican Transition Province 339, 345 microbial fermentation, herbivorous fish 71 microcarnivores 566 microdesmids 318 Micropterus dolomieu (Smallmouth Bass) 473 Micropterus salmoides (Largemouth Bass) 532 micropyle 136 microsatellites 360, 365 genetic diversity 367 genotypes 386–7 Middle America, freshwater fishes 353 midges 567, 568 midshipmen 287 distribution 335 migration anadromy 519–20, 573 annual patterns 515, 516, 517–22 blocking 574 catadromous fishes 521 compass direction 518 complex population structure 370 daily by grunts 491 eels 509, 514 energy use 71 lamprey spawning 239 life histories 519–22 lunar pattern of activity 509 mechanisms 518–19 nutrient cycling 571, 572–4 oceanodromy 521–2 olfaction use 88 olfactory cues 519–20 orientation 518 osmoregulatory environments 104–5 pineal body 54 populations 530 reproductive 515 salmon 519–20 salmonids 509 seasonal 572 sharks 214, 215, 221–2 straying pattern 520 thyroxin 92 tidal 506 top-down processes 564 vertical 394, 502, 504–5 nutrient movement 571 sharks 214 zooplankton 560 see also diurnal vertical migration (DVM) migration rate (Nem) 365–6 migratory overlap 370 Milkfish 269 Miller Lake Lamprey 588 mimicry aggressive 428, 432–3 inedible objects 440–1 larvae 144 miniaturization 163, 164 minnows African region 346 alarm reaction 149 distribution 351 nest building 576 Palearctic region 346 mirror-sided fishes 442, 443–4 Mississippi River (US) channelization 591 mitochondria 44 elasmobranch gills 102 swimming muscles 95 mitochondria-rich cells 103, 104, 105 mitochondrial DNA 14, 15, 356, 359, 365 molecular clocks 371 nucleotide diversity decline 374 phylogeography 374 Mnemiopsis leidyi (ctenophore) 602 mobbing 445 mock turtle syndrome 388 mojarras 304–5 Mola mola (Ocean Sunfish) 324 molariform teeth 125 molas 324 molecular characters 14–15 molecular clocks 371, 372 molecular ecology 360–4 terminology 360 molecular evolution 379, 380, 381–5 Molidae 324 mollusks, feeding on 125 Monacanthidae 324 monkfish 288 monodactylids 305 monogamy 361, 456, 457 pouch-brooding 364 subversion 363 monophyly 13, 367, 368 Index monospondylous 32 moon orbit 507 see also lunar patterns of activity; semilunar patterns of activity moonfish 284, 285 Moorish Idol 318 moray eels 266 feeding 434 pharyngeal jaws 123–4 mormyrids 263, 264 electrical sensitivity 83, 263 elephantfish 83, 264 Morone saxatilis (Striped Bass) 530 morphometric characters 14, 15 morphospecies 11 mortality aquarium trade 614 exponentially declining 545 fishing 611, 612 larvae 140–1, 548 Mosquitofish 363, 414 high-flow conditions 577 Mottled Sculpin, habitat choice 544 mountain streams 416 mouth 70 deepsea fishes 394, 398–9 elasmobranch 205 extension speed 196 function 126 inferior 126 pipette 119, 195 position 126 predatory fish 430 sharks 216 size 123 suction feeding 121 suctorial 126 superior 126 terminal 126 water expulsion 122 mouth-brooders 133, 363–4, 470–1, 528 cardinalfishes 303 cichlids 363, 385 convergence 529 movement desert fishes 411, 412 elasmobranchs 212–14 selective tidal stream transport 507 sharks 212–14 vertical 214 see also locomotion; migration; swimming mucin 36 mucus secretion 106 captured prey 451 hagfish 36, 234–5, 451 nest construction 465 nest guarding males 470 shoals 489 sleeping cocoon 502 mudminnows, distribution 348–9 mudskippers, air breathing 64 Mugilomorpha 292–3, 294 Müllerian duct 54 mullets 292–3, 294 mullids 305 709 muscle 41–5 cardiac 41 cheek 41, 42 dorsal gill-arch 42 electric organ 45 eye 42–4 fin 42 gas bladder 466 myoglobin absence 409 pink 44 red 44, 95, 96, 402 sharks 117, 118 skeletal 41 smooth 44 sonic 44 swimming 95–7 white 44 yellow 45 museum collections 18 Muskellunge 280, 281 mutualism 492–5 cleaning 554 myctophiforms 283–4 myelencephalon 55 Myers, George S myoglobin 44 myomeres 41, 112 Myripristis berndti (soldierfish) 366 Myxiniformes 234–6, 237, 381 Myxinomorphi 234 myxopterygia elasmobranchs 205, 223 see also claspers Myxosoma cerebralis (whirling disease) 602 Na-K ATPase 95 Nabugabo, Lake (East Africa) 311–12 nandids 306 nasal openings, elasmobranch 205 nasal organ, flatfish 152 Naso unicornis (Unicornfish) 361 National Marine Fisheries Service (US) 608 natural selection navigation electroreception 82 larvae 361 neap tides 506 Nearctic region 339, 340–4, 344–5 neck, flexibility in Salamanderfish 279, 280 needlefishes 294, 295 distribution 330, 331 neighbor joining tree 366, 378, 380 Nemastistiidae 303 Nematistius pectoralis (Roosterfish) 303 Neoceratodus (lungfish) 247–8 neofunctionalization 357, 359 Neognathi 280, 281 neon gobies 492 Neopterygii 190–7 body size 208, 209, 210–11 living fish 255–6 phylogenetic relationships 192 Neoselachii 199 modern diversity 206–8, 209–10, 210–17, 218, 219–27 Neoteleostei 281–4 characteristics 281 neoteny 162–3, 163–4 Neotropical region 340–4, 345–6 nephridia, lancelet 233 nephrons 101 elasmobranch 102 nervous system 54–5 autonomic 94 peripheral 55 see also central nervous system (CNS) nest builders cichlids 385 ecosystem engineering 565–6, 576 nest construction 465 nest guarding 363, 470, 528 by another species 472 costs 473 nesting fishes 363 mate choice 461–2 neurocranium 23, 25–8 flatfish 152, 153 neuroendocrine control 505 neurohypophysis 91, 92 neuromasts 76, 77–8 canal 77–8 deepsea fish 400 superficial 77–8 nibbling 434 niche expansion hypothesis 97 niches 536–7 ontogenetic 531–2 nidamental gland 54 night coral reef prey resources/predation pressure 501 see also diel patterns; nocturnal fishes Nile Perch 300 introduction 599, 600 Nile River (Egypt) 593–4 Nimbochromis livingstonii (cichlid) 428 nitrogenous waste excretion 52 air breathing 64 energy loss 72 gills 52, 63, 100 urea 102 no-take areas 619 Nocomis leptocephalus (Bluehead Chub) 463 nocturnal fishes 499, 500, 501, 502 free-running rhythms 505 seasonal change 510 twilight changeover 503 nomeids 320 NONA program 13 noncatch waste 613 nonindigenous organisms 596 norepinephrine 92 North America 339, 344–5 deserts 413–15 freshwater fishes 353 endangered/threatened 587, 588–9 North American Paddlefish 253–4 North Atlantic food web disruption 569 overfishing 569 transarctic interchange with North Pacific 376–7, 378–9, 379 710 Index North Pacific, transarctic interchange with North Atlantic 376–7, 378–9, 379 Northern Pike 280 distribution 349 nostrils 87 nothobranchiids 296 notochord 31–2 Nototheniidae 315, 338 see also Antarctic notothenioids Notothenioidei 314–15, 338, 407–8 see also Antarctic notothenioids nuclear DNA 14, 15, 356, 357 nucleotides 356 diversity 365 nuptial tubercles 463 Nurse Shark food consumption 215 teeth 218 nurseries, intertidal zones 506 nurseryfishes, egg carrying by males 469, 471 nutrients cycling 570–4 marine-derived 573 mobilization 559 movement across thermocline 571 reservoirs 571, 572 retention by dams 593 transport 565, 570–4 oarfish 284, 285 ocean, sharks 211, 214 ocean currents 617 Ocean Sunfish 324 Ocean Surgeonfish 366 oceanodromy, migration 521–2 ocelli 481 oculomotor nerve 55 odacids 312 ogcocephalids 288 oilfishes 300, 309, 319 olfaction 87–8 sharks 220–1 olfactory cues migration 519–20 water currents 518–19 olfactory epithelium 87, 88 olfactory lobes 50, 54, 55 olfactory nerve 55 olfactory sac 87–8 olfactory system, deepsea fish 397 Oligocottus maculosus (Tidepool Sculpin) 487 Ommatokoita elongata (copepod) 553–4 Ompax spatuloides (lungfish hoax) 249 On the origin of species (Darwin) Oncorhynchus (trout/salmon) 279 anadromous species 375–6 Atlantic salmon invasion of Pacific 377 Oncorhynchus keta (Chum Salmon) 602 Oncorhynchus kisutch (Coho Salmon) 359, 474 Oncorhynchus mykiss (Rainbow Trout) 17, 277–8 Oncorhynchus nerka (Sockeye Salmon) 520 Oncorhynchus tshawytscha (Chinook Salmon) 519–20 ontogenetic niche 531–2 oocytes 132, 135 maturation 135 oogenesis 130, 132, 135 oogones 130 oophagy, sharks 223 Opah 284, 285 open sea fishes 401–5 adaptations 401–4 body shape/composition 402–3 countercurrent exchange 402 red muscle 402 see also pelagic fishes operational taxonomic units (OTUs) 13–14 opercular apparatus 29, 31 opercular chamber 58–9 operculum, pseudobranch 46, 47, 86 operculum pupillare 221 ophidiiforms 287 Ophioblennius atlanticus (Redlip Blenny) 366, 367 Ophiodon elongatus (Lingcod) 300 opisthocoelous vertebrae 256 Opsanus tau (Oyster Toadfish) 466 optic lobes 50, 54, 55 optic nerve 54–5 oral care see mouth-brooders Orange Roughy 297, 610 orbital region 25, 26, 27 flatfish 152, 153 Orestias 296 Oriental region 340–4, 346–7 Oryzias latipes (Medaka), transgenic 360 oscillation 113–14 Osmeridae 279 Osmeroidei 276 Osmerus eperlanus (Smelt) 542 osmoconformers 101 osmolytes 98 osmoregulation 92, 100–5 agnathans 101, 102 coelacanths 102 control 105 elasmobranchs 101–2, 205, 212 energetic costs 72 freshwater fishes 102–3 gills 100–1 kidneys 101, 103 lungfishes 102 marine fishes 102, 103–4 sarcopterygians 102 teleosts 102–5 vasopressin 92 osmotic stress 411 Osphronemidae 320–1 ostarioclupeomorphs 191 Ostariophysi 269 Osteichthyes 178–83, 184, 185, 186, 187, 188–97 osteoblasts 157 Osteoglossomorpha 191, 263, 264 distribution 348 osteolepidiforms 183, 184 osteolepidimorphs, diphycercal tail 195 Osteostracomorphi 170, 173–4, 175 ostraciids 324 ostraciiform swimming 114–15 ostracoderms 170, 172, 175 otic region 25, 26, 27 Otocephala 191, 263, 267, 268–74 otoliths 78, 79, 158 age of fish 158–9 iron content 89 lamellae 158–9 otophysans 79–80, 267, 268 distribution 349, 351, 352 hearing specialists 79–80 intense sound damage 80 Otophysi 269 outgroups 13, 380 oval 51 ovarian space 527 ovaries 53–4 sperm storage 136 overfishing 527, 574, 608–11 genotypic alterations 610 North Atlantic 569 phenotypic alterations 610 population genetic makeup 535 trophic cascade disruption 569 oviducts 54, 132 oviparity 54, 138, 363–4 elasmobranchs 224 nest guarding 363 retained 224 sharks 224 ovoviviparity 138 oxygen 57, 58 activity cycle constraint 507 debt 57 demand 97–8 ecosystem effects 577–8 hemoglobin affinity 64–5 transport 64–5 water temperature 58 oxygen consumption sharks 214–15 swimming velocity 67 oxygen dissociation curves 64, 65 oxygen partial pressure 51, 69–70 Oyster Toadfish 466 Pachycormiformes 191 Pacific Herring 377 shoaling 484 Pacific Pearlfish 287 Pacific Plate 332 Pacific salmon commercial exploitation 611 invasion by Atlantic salmon 377 senescence 156–7 Pacific Sardine 608–9 paddlefishes 253–5 ampullary receptors 82 characteristics 250 distribution 348 fossil record 255 vulnerability 255 Paedocypris progenetica (cyprinid) 163, 164 paedomorphosis 162, 163 pain detection 76 palaeoniscoids 188, 189–90 jaws 189–90 locomotion 190 scales 190 Palaeozoic era 169, 170 palatine arch 29 Index palatoquadrate elasmobranch 205, 216 synbranchiforms 299 Palearctic region 340–4, 346 palindromic locomotion 538–9 Panama barrier 377, 378–9 Panama Canal 335, 377 Panamanian Isthmus, elevation 335 pancreas 50, 92 pancreatic juice 70 Pangaea 333 panmixis 534 Paracanthopterygii 285–8, 289 characteristics 285–6 paradisefishes 320 Paralabrax clathratus (Kelp Bass) 362 Paralichthyidae 322 paraphyletic groups 13 parasite–host relationships 554 parasites biodiversity loss 602 farmed salmon 603 removal 493 parasitism 492, 553–4 anglerfish males 288 brood 472 cutthroat eel 266 fish behavior alteration 554 intermediate hosts 554 lampreys 238, 240, 241, 492 parental assignments 361 parental care 6, 143, 456, 468–73 activities 469 bearers 468 cichlids 471 costs 473 diversity 468–71 extent 468–71 gender of care-givers 471–3, 474 guarding 468, 469 males 471–3, 474 nonguarding 468 postzygotic 528 pouch-brooding 364, 462, 469 predation risks 473 prezygotic 528 touching 485 trophic provisioning 470 see also mouth-brooders; nest guarding parr 152 parrotfishes 312 algal grazing 569, 570 body size 459 evolutionary innovation 382–3 jaws 382–3 pharyngeal mill 576 phylogeny 383 parsimony 13, 380 Actinopterygii phylogenetic relationships 382 parsimony network 366, 372, 378, 380, 381 parsimony trees 380 parthenogenesis 359, 456, 460 automictic 364 partial consumers 492 partners, number 455, 456 711 Patagonian Toothfish 315 exploited stocks 527 paternity assurance 471 multiple 361, 363 patrotrophy 469 PAUP program 13 pearlfishes 287, 494 peck order 486 Pecos River Pupfish 536 pectoral appendages, internally supported 176–7 pectoral girdle 34 peduncle, keels 403 pelagic fishes 401–5 anticyclonic patterns 404 body shape/composition 402–3 characteristics 401 circulatory system 404 convergence 405 cyclonic patterns 404 deep 329, 330 evolution 405 flotsam-associated 405 foraging 404 gills 404 keels 403 life history patterns 404–5 planktonic larvae 405 population structure 369 ram ventilation 403 respiration 403–4 swimming 403 pelagic larval duration (PLD) 368, 369 pelvic appendages 183, 184, 185 internally supported 176–7 pelvic axillary process, Salmoniformes 276 pelvic fins 35 jugular 35 modified 53 suction disk 126 pelvic girdle 34–5 Perca flavescens (Yellow Perch) 301 Perca fluviatilis (Eurasian Perch) 302 perches 301–2 body depth 451 climbing 320 predator response 451 Percidae 301–3 Perciformes 300–7, 308–12, 312–21 Percina tanasi (Snail Darter) 592–3 Percomorpha 322–4 basal orders 296–300 Perciformes 300–7, 308–12, 312–21 phylogeny 382 percopsiforms 286 peripheral nervous system 55 periphyton consumption 555–6 perivitelline space 135 perleidiforms 188–9 personal space 486 pesticides 605 Petromyzon marinus (Sea Lamprey) 88, 238– 9, 598 Petromyzontiformes 237–40, 241, 381 Pfiesteria piscicida (dinoflagellate) 552, 553 pH levels 410–11 phallostethids 295 pharyngeal jaws 122–4 Acanthopterygii 291 cyprinids 269 moray eels 123–4 prey crushing 123, 196 wrasses 123 pharyngeal mills 436 parrotfishes 576 pharyngeal pads 123 teleosts 196 pharyngeal tooth patches 31 pharyngognathous condition 196 pharynx, food grinding 70 phenetics 13–14, 380 phenograms 14 phenological relationship decoupling 617 pheromones 88, 484, 485 pholidophoriforms 191 pholids 314 phosphorus 570 Photoblepharon (Flashlight Fish) 479, 483 photoinducible phase 512 photoperiod antifreeze production 98 reproductive cycle 511–12 photophores 14, 36, 282 deepsea fishes 393–4, 398 photoreceptors circadian rhythms 505 distribution on retina 86 ontogenetic shifts 86 photosensitivity, photoinducible phase 512 photosynthesis 507 whiting events in lakes 567–8 Phycodurus eques (Leafy Sea Dragon) 2, PhyloCode 17 phylogenetic constraints 529 phylogenetic techniques, comparison 380 phylogenetics 386, 387 phylogeny 202, 380 phylogeography 370–1, 372, 373–7, 378, 379 anadromous fishes 375–6 freshwater fishes 373–5 marine fishes 376–7, 378–9, 379 plate tectonics 370–1, 374–5 terminology 371 see also dispersal; vicariance biogeography phylograms 14 sperm use 134 physoclistous fish 50, 68, 69 deepsea fish 396 secondary 396 teleosts 195 physostomous fish 50, 68, 69 teleosts 195, 396 phytoplanktivorous fishes 556, 559 phytoplankton 404 pH regulation 568 temperature effects on production 569 pickerels, distribution 348, 349 pigmentation 477–8 embryos 138 larvae 144 melanophores 94 see also coloration 712 Index pigments rod 503, 504 visual in deepsea fishes 401 Pike Characin 271 Pikehead 121, 320–1 jaw protrusion 196, 321 pikeperches 302, 542 pikes 280 predation 432 pilchards 267 pineal body 54, 92 melatonin secretion 505 Pink Salmon 375, 376 disease 602 pipefishes 298 egg laying 469 genetic divergence 371, 373 nutrient transfer to young 469 pouch-brooding males 364, 462, 469 piranhas 271–2 Amazon Basin dam construction 591 group hunting 429 Pirate Perch 286 piscidins 470 piscivores 433, 553 crepuscular 501, 502 trophic cascades 566, 567 pituitary anterior 91, 92, 94 posterior 91, 92 placenta 469 shark 223, 224 placental structures placoderms 177–8 placoid scales 36–7, 38 sharks 213 plankton biomass 568 coral reef fish interactions 546–7 see also phytoplankton; zooplankton plankton feeders 430 plant production in lakes 566, 567 plants biomass in lakes 558–9 defensive responses 556 fish effects 554–9 gallery forests 595 germination 566 mechanical grinding 71 riparian trees 573, 595 seagrass grazing 382–3 seasonal availability 555 trophic cascade with fish and sea urchins 562 see also algae plate tectonics 332, 333, 336 freshwater fish impact 373 phylogeography 370–1, 374–5 plates, placoderms 177 Platichthyis stellatus (Starry Flounder) 152–3 plesiomorphies 12 pleuracanth dentition 198 Pleurogramma antarcticum (Cod Icefish) 405, 408, 409 pleuronectids 322 plunderfishes 408 pneumatic duct 50–1, 68, 69 Poecilia reticulata (Guppy) 600–1 poeciliids 296 Poeciliopsis occidentalis (Sonoran Topminnow) 577 poikilothermy point-of-no-return larvae 140 polar regions 405, 406–7, 407–10 see also Antarctic fishes; Antarctic notothenioids; Arctic fishes polarity 13 polarized light detection 86 migration 518 Pollock 286, 287 pollutants, thermal preference hazards 100 pollution 596, 605–7 environmental contamination assessment 607 reversal/restoration 620 salmon farming 603 polyandry 364, 456, 457 polycentrids 306 polygamy 456, 457 polygyny 364, 456, 457 polymerase chain reaction (PCR) 356–7, 361 forensic use 387–8 Polymixiomorpha 285 polynemids 305 Polyodon spathula (North American Paddlefish) 253–4 Polyodontidae 253–5 fossil record 255 polyphenism, cannibalistic 532 polyphyletic groups 13 polyploidization 357, 359 polyploidy 95, 357 Cyprinidae 269 polypterids characteristics 250 distribution 348 polypteriforms 382 Polypterus (bichir) 249, 251 Pomacanthidae 306 pomacentrids 307, 312 pomatomids 303 Pomatomus saltatrix (Bluefish) 303 pompanos 304 ponyfishes 450 light emission 478–9 pools, desert 413 population control, conservation 621 population density, cave-dwelling fish 418 population dynamics 529–32 cannibalism 532 population genetics 365, 365–7, 367–70 population structure 356, 360 complex 370 deep 365, 367 dispersal 367–8 genetic 533–5 habitat preference 369–70 pelagic larval duration 368, 369 shallow 365, 366 size structured 530–1 populations 385–6, 529–36 annual turnover 533 declines 585 demographically independent 385 genetic structure 533–5 genotype 534 hybridization 535–6 migration 530 production 532–3 recruitment 530 regulation 529–32 size structured 530–1 size variation 530 Porbeagle, exploitation 225 porgies 305 porkfish 305 Porolepimorpha 181–2 porphyropsins 85, 86 postcardinal vein 47 poster colored fish 481–2 pouch-brooding 364, 462, 469 power plants, cooling systems 100 preadaptation cave-dwelling fish 419 high-energy habitats 416–17 predation/predators 425–38, 544–6 aggressive mimicry 428, 432–3 attack 429–30, 430–1, 432–3 avoidance juveniles 149–50 larvae 143, 144 olfaction 88 body shape 119 camouflage 425–6, 428–9 capture 429–30, 430–1, 432–3 community interactions 552, 553 compensation 544 concealment in shoals 488 confusion by prey shoals 448 coral reefs 562, 569 crepuscular 503 density 545 density-dependent changes 544 detection 82 early by prey 444–5 olfaction 88 detritivores 436 direct effects 544 eavesdropping 484 egg mortality 532 exponentially declining mortality 545 eyes 481 fast start 114 foraging risk balance 452 gape limitation 433, 434–5 gene frequency impact 545 group formation 426 group hunting 429 head-on facial characteristics 429 human 607–8 indirect effects 544 intraspecific 532 introduced species 546, 598–9, 600 jaw protrusion 121 larval mortality 532 lie-in-wait 427–8 low water levels 578 lures 428 Index lurking 427–8 mouth 430 night time 501 non-fish 552, 553 parental care risks 473 piscivorous 433, 501, 502, 553 pressure 488 prey backlit 504 choice 433 detection 143, 425–6 handling 433, 434–5, 436 immobilization 430, 432 manipulation in mouth 196, 433 mimicry 440–1 nibbling 434 parasitized 554 searching for 425–6 pursuit 426–9 reaction distance 426 reef systems 482 refuge sites 482, 494 removal 569 resource partitioning 542 on salmon 572–3 scavenging 436 search speed 426 sharks 208, 210 size-selective 560 size-specific 546 spawning seasonality 513 tactics 6, 78 territoriality 437 top-down processes 564 trophic cascades 566–9 warding off 84 yields to 533 young age class exploitation 533 zooplankton 560–1 see also prey predator inspection visits 445 predator–prey interactions high latitude effects 406–7 reaction times 444 predictive cues, seasonal reproduction 514 prejuveniles 144 premaxilla, ascending process 195 pressure, deepsea fishes 394–6 pressure drag 112 prey 439–53 acceleration speed 446 aggregated 448 aggressive behavior 445 alarm chemicals 451 alarm signals 451 armored 449 attack prevention/deflection 447–8 backlit 504 body size increase 449 capture in bathypelagic zone 400 discouraging 448–52 choice by predator 433 coral reefs 501 crushing 123 defended 433, 446 713 dermal thickening 451 detection ability 143, 425–6 ampullary receptors 81–2 avoidance 439–46 distasteful substances 433 diurnally active 482 early detection of predator 444–5 eyes 481 fin size 449 frightening 429 handling by predators 433, 434–5, 436 immobilization 430, 432 invisibility 441–4 leaping to escape predators 447 maneuverability 446 manipulation in predator mouth 196, 433 mimicry of inedible objects 440–1 mucus exudation 451 nocturnal 482 oddity in appearance/behavior 433 parasitized 554 predator detection/avoidance with olfaction 88 predator inspection visits 445 protean behavior 448 pursuit evasion 446–7 refuge sites 482, 494 scales 451 searching for 425–6 size 123 sound production 451 spines 449, 450 stunning 84 terrestrial of archerfish 305 threat sensitivity 452 toxic chemical emission 446 young age class exploitation by predators 533 priapium 137, 138 pricklebacks 314 pricklefishes 297 primitive fish bony 241–58 living representatives 231–59 see also hagfishes; lampreys; lancelets principle components analysis (PCA) 14 Principle of Convergence 419 Principle of Priority 16, 17 priority effects 547 Pristis perotteti (Largetooth Sawfish) 212 Prochilodus mariae (Coporo) 515 production plant in lakes 566, 567 populations 532–3 production : biomass ratio 533, 563 productivity, freshwater 339 proglacial lakes 373 Project Piaba 615–16 prolactin 92, 105 smoltification 151 promiscuous breeders 456, 457 pronephros 52 protandry 156, 456, 458, 528 protective resemblance 425, 439–40 protein, storage 71 protein transporters 105 protocercal tail 33 protogyny 156, 456, 458, 528 Protopterus (lungfish) 246, 247 Psephurus gladius (Chinese Paddlefish) 253, 254–5 psettotids 322 pseudobranch 46, 47, 86 Psychedelic Frogfish 326, 327 Pterapogon kauderni (Banggai Cardinalfish) 303, 363–4 Pteraspidomorphi 170, 171, 172, 175 ptereleotrids 318 Pterois volitans (Lionfish) 582, 583 pterygiophores 282 Ptyctodontiforms 177–8 puffers (pufferfishes) 323, 324 genome 357 inflation 449–50 predation protection 509 Pumpkinseed Sunfish, resource use shifts 542–3 pupfishes 296 desert 413, 414 salinity range tolerance 411, 413 pupils 85 Four-eyed Fish 87 pursuit, predatory fish 426–9 pycnodontiforms 190–1 pygmy sunfishes 306–7 pyloric caeca 49–50 pyruvate 57 quiet period 503 rabbitfishes 318 see also chimaeras Rachycentridae 303 Ragfish 316 Rainbow Smelt 98, 279 Rainbow Trout 277–8 habitat choice 544 hybridization risk 601 name change 17 transgenic 359 rainbowfishes 295 rajiform swimming 115 ram feeding 220 ram ventilation 59 pelagic fishes 403 ratfish see chimaeras ray-fin 188 ray-finned fishes 381–2 see also Teleostei; teleosts rays ampullary receptors 82 body form 206 characteristics 205 electroreception 82 filter-feeding 220 manta 206, 216, 220 modern diversity 207 taxonomic distribution 208 torpediniform 207 torpedo 84, 207, 219–20 see also stingrays 714 Index razorfishes 312 recolonization pathways 374 recruitment coral reef fishes 546–7 populations 530 recruitment limitation hypothesis 547 rectal gland 102 rectum 50 rectus eye muscle, heat production 97 red blood cells 48 carbonic anhydrase 66 Red List 586 Red Sea, marine fish movement into Mediterranean 338 Red Shiners 543, 559 Red Snapper 388 red tides 552 Redlip Blenny 366, 367 Reedfish 249 refuge sites 482 availability 545 competition 554 invertebrate 494 refugia, glacial 374 Regan, C Tate 7, 11 regression analysis 14 regressive evolution 419 rehabilitation 619–20 release calls 483 remoras 166, 167, 303–4 cleaning 494 renal corpuscles 52 reproduction 6, 455–75 age-specific 529 cave-dwelling fish 418 circadian rhythms 512 climate change effects 616, 617 diadromy 517 elasmobranchs 205 embryology 137–9 energy required 135, 526 environmental cues 511 gametogenesis 130–6, 132–6, 528–9 seasonality 512, 515 hagfish 236 home range desertion 487 lampreys 237–8, 239 larvae 139–45 lifetime opportunities 455, 456, 457 lunar patterns of activity 507–8, 509 migrations 515 patterns 455, 456, 457–61 photoperiod 511–12 seasonal pattern of activity 511–15 sharks 208, 210–11, 222–4 temperature 511–12 timing 616, 617 water temperature 616 see also courtship; fertilization; mating; parental care; spawning reproductive allotment 223–4 reproductive ecology 525–9 reproductive effort 135 indices 135–6 reproductive guilds 468 reproductive interval 528 reproductive output measurement 607 reproductive system 52–4 reservoirs, sediment retention 593–4 resident species, intertidal 506–7 resource partitioning 542 temporal/trophic 543 resource use shifts 542–3 resources availability 565–6 sustainable use 621 respiration 57–64 aerial 62–3 air breathing 60, 61–2, 62–4 cutaneous 59 pelagic fishes 403–4 respiratory structures, accessory 410–11, 412 restoration projects 620 rete mirabile 47, 51, 69 function 70 heterothermic fish 96, 97 retina 86 photoreceptor distribution 86 retropinnids 279 Reynolds number 142, 161 Rhenaniforms 177 rheotaxis reversal 151 Rhinichthys cataractae (Longnose Dace) 544 rhizodonts 183 rhodopsins 85, 86 rhyacichthyids 317 ribosomal RNA 356 ribs 32–3 rice eels 299 ricefishes 295 ritualized combat 479 river continuum concept (RCC) 542 rivers channelization 591 debris dams 589 deforestation 590 downstream- (D-) links 541 multiple habitats 542 reduced flow 596 species diversity 541 stream order 540–1 vulnerable species 589 woody debris 595 removal 589–90 zones 539–42 riverscape 542 rivulines 296 rockbasses 301 rockfishes 299, 300 rocks 576 rods (vision) 85, 503 pigments 503, 504 role reversed species 462 Roosterfish 303 Root effect 51, 64, 65, 69 Root-off shift 70 Root-on shift 70 Ropefish see Reedfish Rosen, Donn Eric rostral organ, coelacanths 243 rostral region, armed 219 Rosyside Dace 544 rotational feeding 434, 435 routine metabolic rate 67 Rudd 602 saccopharyngiform fish 266 Sacramento Perch 601 sagitta 79 Sailfin Molly, respiration 58 sailfishes 319 group hunting 429 Salamanderfish 279, 280, 413 salinity 411 Messinian Salinity Crisis 335, 338 physiological barriers 411 pupfish tolerance 411, 413 salinization 596 Aral Sea 411 Salminus maxillosus (Dorado) 538 Salmo salar (Atlantic Salmon) 279 Salmo trutta (European Brown Trout) 279 salmon 279 anadromous 375–6, 573 commercial exploitation 611 conservation genetics 376 dam hazards 593, 594 escapes from farms 603–4 farmed 603–4 genetic diversity 534 maturation 162 migration 519–20 nomenclature 277–8 nutrient budget impact 572–3 predation on 572–3 smoltification 150–2, 162 salmonids migration 509 phylogeny 278 Salmoniformes 276 jaws 269 Salmoninae 277, 279 Salmonoidei 276 salt excretion, marine teleosts 103 salt storms 596 salting out effect 58, 69 saltmarshes creeks 506 primary productivity 574 Salvelinus alpinus (Arctic Char) 277, 279 Salvelinus fontinalis (Brook Trout) 149–50 Salvelinus namaycush (Lake Trout) distribution 374 diversity 365 sand 576 Sand Lances 315, 407 sand stargazers 316 Sandbar Shark, teeth 218 sanddivers 315 Sander (pikeperch) 302 Sander vitreus (Walleye) 301 Sander vitreus glaucus (Blue Pike) 588 sarcoplasmic reticulum, calcium pumping 97 Sarcopterygii 178, 179–83, 187 homocercal tail 195 living 242–8 osmoregulation 102 Index sardines 267 Californian fishery 608–9 dispersal 372 hearing specialists 79 vicariance biogeography 372 Sardinops (sardines) 372 Sardinops sagax caeruleus (Pacific Sardine) 608–9 Sargasso Sea 521 satellite lake hypothesis 312 satellite males 474 saturated salt (SED) buffer 362 saurichthyiforms 188 Saurida undosquamis (Brushtooth Lizardfish) 338 sauries 294 sawfish armed rostral region 219 habitat 212 life span 216 pristiform 207 vulnerability 212 sawsharks, armed rostral region 219 scales 36–9 annuli 39, 157–8 barrier function 106 caducous 38 circuli 39 cosmid 37 ctenoid 271 cycloid 37 deciduous 38, 451 development 157 developmental pattern 38, 139, 140 false annuli 158 ganoid 37, 188, 190, 451 geographic variation 38 growth ring validation 158 lateral line 38 mirror-sided fish 443–4 modifications 38 morphology 39 palaeoniscoids 190 phylogenetic significance 37–8 placoid 36–7, 38, 213 removing 492 sexual dimorphism 38, 463 size 38 spawning checks 157–8 spined 37 taxonomic use 37–8, 39 teleosts 193 tunas 403 types 36–7 scaling 160–1 Scaridae 312, 382–3 scatophagids 318, 571–2 scats 318 scavengers 436 schindleriids 318 schooling 468 spacing 489 tactics 445 see also shoals/shoaling Schreckstoff 269 sciaenids 305 715 sclera 86–7 Scleropages (arowanas) 374–5 sclerotic bones 86–7 Scomberomorus (Spanish mackerel) 15 distribution 332 taxonomy 17 vicariance biogeography 336–7 Scomberomorus regalis (Spanish mackerel) species group 336–7 scombrids 319 Scombroidei 12, 319, 320 Scombrolabracidae 319 Scopelomorpha 283–4 Scorpaemichthys marmoratus (Cabezon) 300 scorpaeniforms 299–300 scorpionfishes 300 scrapers 382–3 scrounging 494 sculling 116 sculpins 309 Antarctic species 408 antifreeze compounds 99 freshwater 300 habitat choice 544 habitat diversity 539 morphological adaptations for water velocity 67, 68 scutes 38 clupeomorphs 267 sea chubs 305 digestive tract microflora 437 sea dragons 298 Sea Lamprey 88, 238–9 introduction 598 sea level rise 579, 617 sea urchins coral reef overfishing 563 feeding on 122, 125, 433 fish–algae interactions 562 toadfish feeding 575–6 trophic cascade with fish and plants 562 sea water, freezing point 98 seabass 301 seagrass grazing 382–3 seahorses 297, 298–9 egg laying 469 nutrient transfer to young 469 pouch-brooding males 364, 462, 469 vulnerability 298–9 seamoths 298 seasonal pattern of activity 509–15 climate change 617 reproduction 511–15 seaweed see algae secretory region, anteroventral 51 SED buffer 362 sediment changes with dam construction 593–4 deforestation impact 595, 598 stream siltation 595 turnover 576 seed dispersal 556, 566 selective tidal stream transport 507 self-fertilization 458–9 semelparous fish 455, 456, 457, 515, 528 semicircular canals 78–9 flatfish 152 see also otoliths semilunar patterns of activity 507–9 seminal vesicle 53 semionotids, species flock 311 semionotiforms 190 Semionotus (neopterygian) 311 senescence, adult fish 156–7 senior synonym 17 sensory hair cells 75, 76–7 sensory organs 75 sensory systems 75–90 chemoreception 87–9 electroreception 80–4 magnetic reception 89 mechanoreception 75–80 sharks 220–2 vision 84–7 sequence divergence 371 Serranidae 301, 458 hermaphroditism 458, 460 settlement limitation hypothesis 547 settling, larvae 150 sex change 458, 459, 528–9 bristlemouths 397 postmaturational 156 prematurational 155 sex chromosomes 359 sex determination 153, 154, 155–6, 359 temperature effects 155 sex hormones disruption 93 estradiol 151 stress response 107 sex ratio variation 528–9 sex role reversal 364 sexual dimorphism 14, 456, 461, 462, 463–5 coloration 463, 478 deepsea fishes 396–7 electric organ discharge 485 scales 38, 463 sexual selection 6, 364, 456, 461, 461–2, 463–5 trade-off 462 shade hovering in 445 predator early detection by prey 444 predator hiding 444 veiling brightness 444–5 shads 267 seasonal movement 519 Shanny circadian rhythm 506 tidal activity cycle 506, 507 sharks 199–200 age at maturity 223 ampullary receptors 81–2 anticancer drugs 226 attacks on humans 569–70 bamboo 125 birth adaptations 224 body size 208, 209, 210–11 brain 222 buoyancy control 213–14 cancer cure 226 716 Index cartilage 226 central nervous system 222 characteristics 205 claspers 223 clutch sizes 223–4 compass orientation 221–2 conservation 224–7 convergence of deepsea species 401 copulation 224 courtship 224 dentition 125, 217, 218, 219 distant touch sensitivity 221 ear 221 ecotourism 227 egg laying 224 egg sacks 363 eggs 134 electroreception 81–2, 221 embryonic development 223–4 embryonic nutrition 223–4 energy conservation 215 exploitation 225 feeding 208, 210, 214, 219 habits 216–17, 218, 219–20 filter-feeding 214, 219, 220 fins 117, 118 exploitation 225, 226 international trade 388 fishing 225–6 food consumption 215 foraging times 220 fresh water 211–12 gestation period 224 gill netting of large species 569–70 ground 207 group hunting 429 growth rate 215–16 habitats 211–12 hearing 80, 221 heterocercal tail 118, 212–13 home ranges 214 Indian–Atlantic barrier 379 internal fertilization 137, 223, 363 jaws 209–10, 216–17 lamniform 207 livers 396 maximum sustainable yield 225 mechanoreception 221 metabolism 214–15 migration 214, 215, 221–2 orientation 518 modern groups 206 diversity 207 mouth 216 movement 212–14 muscle 117, 118 oceanic 211, 214 olfaction 220–1 oxygen consumption 214–15 parthenogenesis 364 placoid scales 213 predation 208, 210 reproduction 208, 210–11, 222–4 requiem 207 river 211–12 sensory physiology 220–2 skin 117–18 squaliform 207 swimming 117–19, 379 taxonomic distribution 208 teeth 217, 218, 219 tooth replacement 199–200 vertical movements 214 vision 221 viviparity 223, 224 Sharksucker 166, 167, 303–4 Sharpnose Puffer 509 shell gland 54 shoals/shoaling 426, 432, 468 alarm chemicals 451–2 antipredation benefits 445–6 attack abatement effects 448 attack response 448 coordinated flight behavior 451 dilution effect 448 fidelity 489 formation 484 group-separating tactics 432 home range size 487 hydrodynamic advantage 489 interspecific 495–6 lattice structure 489 maintenance 484 mucus secretion 489 non-fish species 488 predator confusion 448 resting 501 sound production 484 spacing 489 stability 489 tactics 445 visual advantage 444–5 water displacement 484 zooplanktivorous fishes 501 short-chained fatty acids (SCFAs) 71 Shortfin Mako, teeth 218 shrimp fishing, bycatch 613 shrimpfish 297 Siamese Fighting Fish 320 Siganidae 318 Siluriformes 272 distribution 351, 352 silversides 294, 295 sex determination 529 species flock 310 Silvertip Shark 166, 167 silvery sides 442, 443–4 Siniperca (Aucha Perch) 472 sister groups 13 sister species 371 size advantage model 459 skates ampullary receptors 82 characteristics 205 diversity 207 egg sacks 363 electroreception 82 life span 216 North Atlantic populations 569 rajiform 207 taxonomic distribution 208 skeletal muscle 41 skeleton 23, 24, 25–39 appendicular 34–5 cartilage 70 elasmobranch 205 external dermal 38 gar 255–6 integumentary 36–9 pectoral girdle 34 pelvic girdle 34–5 postcranial 31–4 teleost tail bones 192–3 see also skull; vertebrae skin 36 barrier function 106 elastic recoil 118 gas exchange 59 shark 117–18 skull 23, 24, 25–9, 30, 31 amphistylic suspension 30, 227, 299 branchiocranium 23, 25, 28–9, 30, 31 Neoteleostei 281 neurocranium 23, 25–8 flatfish 152, 153 pectoral girdle attachment 34 teleosts 192 sleep 502 sleepers 317 slime prey production 451 toxic 319 see also mucus secretion slime eels see hagfishes Sling-jaw Wrasse 121–2 small intestine, absorption 70–1 Smallmouth Bass 473 smell organs 87–8 smelts 279 resource partitioning 542 southern 279 Smith, J L B 243, 244 smoltification 150–2 age 151 lunar patterns 509 salmon 162 timing 152 smooth muscle 41, 44 snags 589–90 Snail Darter 592–3 snailfishes 300 snake mackerels 319 snakeheads 321 snappers 304 sneaker males 363, 474, 475 snooks 300 soapfishes 301 social behavior transmission 490–1 social interactions, ampullary receptors 82 social signaling 483 Sockeye Salmon 375, 376 life history 520 nest building 565–6 sodium–potassium ATPase 95 sodium–potassium exchange 104–5 soldierfishes 366 Soleidae 322 soles 322 Index soluble reactive phosphorus (SRP) 570 somatolactin 92 somatostatin 92 sonic ligament 484 sonic muscles 44 Sonoran Topminnow 577 sound, underwater 80 ambient 80 communication 80 hair cell damage 80 localization 446 sharks 221 sound production aggression reduction 484 agonistic displays 461–2 communication 483–4 courtship 461, 466, 483 eavesdropping by predators 484 gas bladder 396 group formation 484 mechanisms 80 Otocephala 268–9 prey 451 sciaenids 305 shoaling behavior 484 spawning 466, 483 territorial behavior 461–2 Soupfin Shark, exploitation 225 South America freshwater fishes 351–2, 353 ichthyological provinces 345–6 southern Africa, biogeographic pathway 379 space light 441 spadefishes 318 Spanish mackerels 15 distribution 330, 332, 333 geminate species 335 taxonomy 17 vicariance biogeography 336–7 Sparidae 305 Sparisoma (parrotfish) 383 Sparisoma radians (Bucktooth Parrotfish) 459 spawning 456, 461, 461–2, 463–8 act 466–8 aggregations 468 bottom 468 bouts per year 528 broadcast 528 checks 157–8 cichlid lunar pattern of activity 509 coral reef fishes 508–9, 514 estuaries 146 group 466, 467 home range desertion 487 intertidal 508 iteroparous fish 455, 456 lamprey 239 lancelets 232 larval biology hypotheses 509 locale 514 lunar patterns of activity 507–8, 509 migration interruption with dams 593 number per lifetime 528 nutrient cycle 573 pair formation 466 panmictic 534 717 patterns 141 season duration 528 seasonality 511–12, 513–14 semelparous fish 455, 456, 457 semilunar patterns 507–8 sound production 466, 483 spent females 132 sturgeon 253 substrates 467, 494 tidal patterns 507–8 turbidity impact 464 water column 466 spawning site 461 mate choice 462 selection 465 spawning stupor 468 spears 430–1 specialist species 575–6 speciation 163 allopatric 312 cichlids 311–12, 384–5 explosive 308–12 sympatric 312 species 11–12 allopatric 542 biological species concept 12 chemical recognition 464 cryptic 383–4, 385 diversity in flowing water 541 endemic 598, 599 evolutionary 12 generalist 575–6, 598 isolation mechanisms 464, 535 mislabelling in wildlife trade 388 sister 371 specialist 575–6 sympatric 542 see also introduced species species flocks 308–12 cichlids of Lake Malawi 384–5 species isolating mechanisms 455 Species Survival Commission (IUCN) 586 sperm drinking 467 sperm duct 53 spermatogenesis 130 spermatogonia 53 spermatozoa 130, 134 fertilization 136, 137 Sphyraena barracuda (Great Barracuda) 306, 307 Sphyraenidae 306, 307 Sphyrna mokarran (Great Hammerhead) 431 Sphyrna tiburo (Bonnethead Shark) 364 spiderfishes 282, 283 Spikedace 543 spillover effect 619, 620 spinal cord 55 spines 38, 449, 450 Spiny Dogfish 156 spiral valve 49, 50, 70–1 spleen, environmental stressor effects 107 splitting (taxonomic) 17 sponges 334 Spookfish 228 Spotfin Chub 596 sprats 267 Spraying Characin 469–70 spring tides 506, 508 squalamine 226 squalene 396, 401 Squalus acanthias (Spiny Dogfish) 156 squaretails 320 squirrelfishes 285, 297 standard metabolic rate 66 stanniocalcin 92 stargazers 315 Starry Flounder 152–3 startle calls 483 startle response, larval 143 static displays 478 Steelhead Trout 375 stenohaline fish 100 Stenopterygii 281–2 stephanoberyciforms 297 stereocilia 75, 77 stichaeids 314 sticklebacks 297, 298 egg thievery 363 morphs 534 photosensitivity 512 spines 450 UV reflectance 483 stingrays freshwater 211, 212 hammerhead shark prey 431 life span 216 myliobatiform 207 stochastic processes 546, 548–9 stock : recruitment relationship 530 stocks of fish 365, 385–6, 530 exploitation 526, 527, 607–16 world status 608 stomach 49, 70 food breakdown 70 stomiiforms 281–2, 396 stonefishes 300, 428 storms 578–9 Straits of Gibraltar 335 stream order 539 streams channelization 591 debris dams 589 deforestation 590 food webs 574, 575 Invertebrate response to fish predation 561–2 low-flow conditions 578 sediments 595 siltation 595 woody debris 595 removal 589–90 stress 106–7 immune system 107 indicators 107 primary responses 106 secondary responses 106–7 tertiary response 107 stress proteins 106 stressors, environmental 107 Striped Bass 530, 577 restoration 620 718 Index stromateids 320 Stromateoidei 320 sturgeons 252–3 characteristics 250 exploitation 253, 387–8 life history 253 mtDNA sequences 388 spawning 253 vulnerability 253, 388 subcarangiform swimming 114 subclavian artery 46 suckers 269, 270–1 distribution 351 sucking organ of remoras 303–4 suction inertial 121, 434 pressure 121 suction feeding 120–1, 434 evolution 122 phases 121 Suez Canal 338 sulfur detoxification 64 sun compass 518 sunfishes 301 distribution 350 pygmy 306–7 resource use shifts 542–3 supercooling 98 superfetation 136 surface-dwelling fish 87 see also pelagic fishes surfperches 307 surgeonfishes 12, 318–19 digestive tract microflora 437 survival larvae 139–41 see also mortality survivorship 529 curves 545 suspensorium 121 swallower eels 266 swallowers 315 swallowing deepsea fishes 399 prey manipulation 196, 433 teleosts 196 swamp eels 299 swamps, tidal 506 swim bladder see gas bladder swim-up stage 139 swimming anatomy 112 anguilliform 114, 116, 117–18, 419 carangiform 114, 304 cave-dwelling fish 418 cost 161 elasmobranchs 117–19 energetics 67, 68 labriform 115 larvae 142, 145–7, 361 metabolic cost 67, 68 modes 113–16 muscle fibers at low temperatures 95 pelagic fishes 403 rajiform 115 sculling 116 sharks 117–19, 379 subcarangiform 114 switching of modes 116 thunniform 96 swimming muscle, elevated temperature 95–7 swordfishes 319 bill use 430–1 swordtails 462 species recognition 464 symbiosis 492–6, 566 non-fish species 494 sympatric species 542 Symphodus melops (Corkwing Wrasse) 493 symplesiomorphies 380 synapomorphies 12 synbranchiforms 299 Synchiropus splendidus (Mandarinfish) 422, 423 synchronizing cues 514 Syngnathoidei 298–9 pouch-brooding males 364 Synodontis multipunctatus (Cuckoo Catfish) 472 systematics 11–18 evolutionary 14 molecular 14–15 tactile communication 485 cleaners 493 tail bones, teleost 192–3 tail forms 33–4 diphycercal 195 heterocercal 33 Bowfin 256, 257 shark 118, 212–13 high aspect ratio 114 homocercal 195 leptocercal 33 low aspect ratio 114 protocercal 33 shark 118 tails keels 403 prey capture in sharks 219 Tana, Lake (Ethiopia) 311 tangs 318–19 tapetum lucidum 86, 221 tarpon 265–6 taste 88–9 taste buds 78 taste receptors 88–9 taxonomic characters 14–15 taxonomic units 16 taxonomy 4, 11–17 name changes 17 numerical 13–14 tectonic plates 332, 333, 336 freshwater fish impact 373 phylogeography 370–1, 374–5 teeth 28–9 deepsea fishes 398, 399 elasmobranch 205 lungfish 246 placoderm 178 prey preparation for swallowing 433 196, replacement patterns in elasmobranchs 199–200 sharks 217, 218, 219 see also dentition telencephalon 54, 55 Teleostei 191–7 Acanthomorpha 284–8, 289 bony element reduction 192–3 classification diversification 191 Elopomorpha 191, 263, 264–6 Mesozoic radiation 190 Neoteleostei 281–4 Osteoglossomorpha 263, 264 Otocephala 191, 263, 267, 268–74 Percomorpha 322–4 phylogenetic relationships 191, 192 phylogeny 261, 262, 263 radiations 191–2 spiny-rayed 291–325 synapomorphies 262 trends 192–7 true 274–7, 277–8, 279, 280 see also Acanthopterygii Teleostomi 178–83, 184, 185, 186, 187, 188–97 living representatives 241–58 teleosts caudal fins 195 deepsea habitats 395 diadromous 103, 104 distribution 261 DNA sequences 381–2 dorsal fins 193, 194 feeding apparatus modifications 195–6 freshwater 102–3 gas bladder 68–70 modifications 195 hermaphroditism 282, 301 intestines 71 lipids in body density reduction 70 marine 103–4 osmoregulation 102–5 paired fins 194 physoclistous 195 physostomous 195, 396 pipette mouth 119 reproductive guilds 468 spawning seasonality 513–14 spiny 284–8, 289 Tellico Dam (Little Tennessee River, US) 592–3 temperate communities, herbivory 557–9 temperature, body 3, 6, 94–7 temperature, water 58 Arctic region 410 climate change 579, 616–17, 618 deepsea fish 396 deforestation impact 595–6 distribution of fish 616–17 diurnal vertical migration 394, 396 ecosystem effects 577–8 embryonic vertebrae sensitivity 138–9 extreme weather events 579 extremes 97–9 fluctuations 94–5 Index lakes 568 larval development 141 lunar rhythm of preference 509 meristic variation 138 metabolic rate 67, 72 physiological stress 98 phytoplankton production 569 preference lunar rhythm 509 ontogenetic shift 99 thermal 99–100 reproduction 511–12 sensitivity 616 seasonal movements 511, 519 seasonal patterns of activity 509–11 sex determination 155, 529 shallow water species 331 sudden shifts 579 weather patterns with climate change 618 temperature relationships 94–100 tenaculae 227 tenuis larvae 287 terapontids 306 terminating cues 514–15 terpenoids, halogenated 556 territorial behavior 486–7 electric organ discharge 485 sound production 461–2 territorial defense 486–7 territoriality 437, 486–7 damselfishes 557, 558 testes 52–3 tetragonurids 320 tetraodontiform swimming 115 Tetraodontiformes 323–4 Tetrapoda 181 ancestors 183, 184, 185, 186 cladogram 187 pelvic appendages 183, 184 tetrapodomorphs 183, 184, 185, 186 ancestral group to coelacanths 243 Thalassoma bifasciatum (Bluehead Wrasse) 458, 461 cultural traditions 490–1 thanatosis 428 Theldonti 170, 173, 175 thermal preference 99–100 thermocline nutrient movement across 571 placement in lakes 568 thermogenic tissues 97 Thoracopterus (Chondrostei) 188–9, 190 flying fish morphology 447 thread cells 36 threadfins 305 threat sensitivity 452 threatened species 585–6, 587, 588–9 see also International Union for the Conservation of Nature (IUCN) Three-spined Stickleback 297 thresher sharks, prey capture 219 thunniform swimming 96, 114, 115 Thunnus (tuna) 17 Thunnus alalunga (Albacore) 320, 519 Thunnus thynnus (Bluefin Tuna) 319 thymallines 277 719 thyroid-stimulating hormone (TSH) 92 thyroxin 92 smoltification 151 tidal patterns 506–7 spawning 507–8 tidal regions 506 tidal stream transport, selective 146 Tidepool Sculpin 487 Tiger Shark 218 Tiktaalik roseae (tetrapodomorph) 185, 186, 187 tilapia mouth-brooding 364 salinity range tolerance 411 tilefishes 303, 576 tiptoeing 116 tissue collection for genetic analysis 361, 362 Tjeukemeer, Lake (Netherlands) 542 toadfishes 287 distribution 335 feeding regime 575–6 tongue parasites 554 tonguefishes 322–3 toothplates, lungfish 246 top-down processes 563, 564 Topminnow 294, 414 high-flow conditions 577 topology 380 Torpedo (Electric Ray) 84 torpedo rays 84, 207 electric discharges 219–20 Torrentfish 315 cave-dwelling 419 torrents 415–17 Totoaba macdonaldi (Giant Totoaba) 610 toxotids 305 trachinids 315 Trachinoidei 315 trade-offs 161 Trafalgar effect 448 traits 525–9 transarctic interchange 376–7, 378–9, 379 transfer RNA 356 transgenic fishes 359–60 transitional juveniles 547 transparency 442, 444 transplant organisms 596 trees riparian 573, 595 see also deforestation triangular dentition 124 trichiurids 319 trichonotids 315 trigeminal nerve 55 triggerfishes 323, 324 blowing 122 lekking 457 spines 450 water expulsion from mouth 122 trimethylamine oxide (TMAO) 102 elasmobranch excretion 205 tripletails 304 tripodfish 282 Tripterygiidae 316 trochlear nerve 55 trophic cascades 545, 566, 566–9 application 569–70 direction 568 fish, sea urchins, and plants 562 human disruption 569 trophic provisioning 470 tropical communities climate change 617 herbivores 556–7 tropical submergence 338, 396 trout 279 anadromous 375, 376 dam construction impact 591–2 nomenclature 277–8 smoltification 150–2 trout zone of river 539 troutperches 285, 286 trumpetfishes 299, 377 dispersal barriers 378–9 tselfatiiforms 191 tuberous receptors 82–3 tunas 319, 320 circulatory system 404 cutaneous arteries 47, 48 distribution 330 gills 59 group hunting 429 life history patterns 404–5 migration 54, 521–2 ram ventilation 59 red muscle location 96 scales 403 seasonal movement 519 shoaling with dolphins 488 turbidity 444 spawning behavior reduction 464 turbots, spiny 322 turbulent water 415–17 turkeyfishes 300 Tursiops truncatus (Bottlenose Dolphins) 484 turtle exclusion devices (TEDs) 614 twilight changeover patterns 502, 503–4 see also crepuscular fishes type specimens 16–17 typhlosole 49 Tyrannochromis macrostoma (cichlid) 443 ultrasound detection 80 ultraviolet light 85–6 detection 483 harmful exposure 595 reduction by shade 595 reflectance 483 ultraviolet light-reflective markings 86 communication 483 ultraviolet radiation 595 umbilical cord, appendicula in sharks 223, 224 undulation 113–14 Unicornfish 318–19, 361 upwelling light 441 uranoscopids 315 urea elasmobranch excretion 205 retention 102 urinary bladder 52, 101, 104 720 Index urine 52, 101 teleost excretion 103, 104 urophysis 92 urotensins 92, 105 uterine milk 223 uterus 54 utricle 78, 79 vagus nerve 55 intestinal 52 Valenciennes, Achille validation, scale growth rings 158 Vandellia cirrhosa 273 variable number of tandem repeats (VNTR) 360 vasa efferentia 53 vasopressin 92, 105 veiling brightness 444–5 velum 59 ventilation 57, 58–64 agnathans 59, 60 gill 58–9 ram 59, 403 see also air breathing vertebrae 32 amphicoelous 256 opisthocoelous 256 teleosts 192 temperature sensitivity in embryos 138–9 vertebral column 32 vertebrates classification 15–16 jawed 381 radiations 381 vexillifer larvae 287 vibrations detection 76, 77 gas bladder 79 prey fish attraction 78 vicariance biogeography 13, 336–7, 370–1, 372, 373 Galaxias maculatus distribution 352 sardines 372 Victoria, Lake (East Africa) 464 Nile Perch introduction 599, 600 villiform dentition 124 viscous drag 112 visibility 478 vision 84–7 adaptations for special habitats 87 aerial 87 larvae 143 sharks 221 visiting species, tidal areas 506 visual agonistic displays 479 visual communication 477–9, 480–2, 483 visual cues, home range location 487 visual pigments, deepsea fishes 401 vitellogenesis 132 viviparity 54, 138, 363 placental 223, 224 sharks 223, 224 uterine 223 yolk sac 223 see also live-bearers Volga River (eastern Europe) 593–4 von Bertalanffy growth equation 159–60 Wahoo, systematics 15 walking 116 Walking Catfish 63 Wallace’s Line 332, 333, 347 Walleye 301 warts, bony 38 water clarity 482 competition for 596 depth increase with age 539 displacement for group maintenance sounds 484 flowing 541 hardening 137 sea water freezing point 98 see also currents; temperature, water water flow ecosystem effects 577–8 species diversity 541 water levels flood conditions 414, 577 high-flow conditions 414, 577 low-flow conditions 577–8 sea level rise 579, 617 seasonal cycles 578 water table lowering 596 watersheds, perturbation 594–6 wax esters 70, 400 weather climate change impact 617–18 extreme 578–9, 617–18 Weberian ossicles 79–80, 268 Weber’s Line 347 weeverfishes 315 Wegener, Alfred 333 Weitzman, Stanley H western Atlantic region 332, 334–5, 336 Whale Shark clutch size 223–4 feeding habits 216 ram feeding 220 size 209 whalefishes 297, 397 whirling disease 602 white muscle 44 White Seabass 143 White Shark energy consumption 215 feeding 215, 216 migration 214, 215 ritualized combat 479 size 209 whitefish 276–7 whiting events 567–8 whitings 286 wildlife trade 387–8, 586, 614–16 aquariums 590–1, 614–15 winter, activity of fish 510 wolf-eels 125, 314 Wolffian duct 53 wolffishes 314 wonder net see rete mirabile woody debris 595 removal from rivers 589–90 World Conservation Union 586 wormfishes 318 wrasses 312 cleaner 492, 493, 554 cultural traditions 490–1 pharyngeal jaws 123 prey handling 433 protogyny 458 scrounging 494 sex change 461 wreckfish 300–1 xenacanthimorphs 198 Xiphias gladius (Swordfish) 430–1 xiphiids 319 Xiphioidei 319 Xiphophorus (swordtails) 462 chemical recognition 464 year classes 530, 531 yellow muscle 45 Yellow Perch 301 Zanclidae 318 Zanclus canescens (Moorish Idol) 318 zebra mussel 602, 604 Zebrafish 269 genome 355, 359 transgenic 359 zeiforms 297 Zeitgeber 505, 507 Zeus faber (John Dory) 297 Zoarcoidei 313–14 zonation 539–42 longitudinal 539, 541 zoogeography 329–54 freshwater regions 339, 340–4, 344–7 marine fishes 329–38 marine regions 331–2, 333, 334–5, 336– 7, 338 see also vicariance biogeography zooplanktivorous fishes 566 aggregations 501 detection capabilities 426 diurnal 501, 502 feeding 430 guild 536–7 predator avoidance 446, 447 search tactics 426 shelter when pursued 446 trophic cascades 566, 567 zooplankton 404 coloration 560–1 detection 483 impact of fish 559–61 nocturnal predators 561 population size structure 561 predation 560–1 seasonal blooms 515 size 560 species composition 561 trophic cascades 566, 567 ZW heterogamety 155 ... the services of a competent professional should be sought Library of Congress Cataloguing-in-Publication Data The diversity of fishes / Gene Helfman [et al.] – 2nd ed p cm Rev ed of: The diversity. .. enough information on the general aspects of ichthyology to make informed, biologically sound judgments and decisions, and to gain a larger appreciation of the diversity of fishes beyond the. .. of fishes are as much anthropological as they are ichthyological, namely Johannes’ (1981) Words of the lagoon and Goulding’s (1980) The fishes and the forest A stroll through the shelves of any