List of contributors xv
Series editor’s foreword xxi
Preface to the second edition xxv
Preface to the first edition xxvii
Acknowledgements xxix
1 Introduction 1
Robin N. Gibson
1.1 The fascination of flatfishes 1
1.2 A brief history of flatfish research and its contribution to fish biology and fisheries science 3
1.3 Scope and contents of the book 4
1.4 Nomenclature 9
Acknowledgements 10
References 10
2 Systematic diversity of the Pleuronectiformes 13
Thomas A. Munroe
2.1 Introduction 13
2.2 Systematic profile of the Pleuronectiformes 18
2.3 Intrarelationships of the Pleuronectiformes 19
2.4 Brief synopses of the suborders and families 22
2.5 Diversity of the Pleuronectiformes 26
2.5.1 Overview 26
2.5.2 Flatfish species diversity 27
2.5.3 Diversity of species within families 28
2.5.4 Standing diversity estimate for species of Pleuronectiformes 29
2.5.5 Relative diversity of the Pleuronectiformes 31
2.6 Patterns of species discovery among pleuronectiform families 32
2.6.1 History 32
2.6.2 Factors contributing to new species discovery among the Pleuronectiformes 35
2.6.2.1 Systematic activities 35
2.6.2.2 Geographic region 37
2.6.2.3 Depth 39
2.6.2.4 Size 40
2.7 Conclusions 42
Acknowledgements 44
References 44
3 Distributions and biogeography 52
Thomas A. Munroe
3.1 Introduction 52
3.2 Geographic distribution of pleuronectiform lineages 56
3.3 Global patterns of species richness for the Pleuronectiformes 61
3.3.1 Latitudinal gradients in species richness 61
3.3.2 Tropical and subtropical regions 61
3.3.3 Temperate regions 62
3.3.4 Species richness on continental shelves 63
3.3.5 Insular versus continental regions 64
3.3.6 Continental versus oceanic islands 66
3.4 Species richness in specific environments 66
3.4.1 Freshwater environments 66
3.4.2 Antarctic Ocean 67
3.4.3 Arctic Ocean 68
3.4.4 Shallow-water versus deep-sea habitats 69
3.5 Historical biogeography 71
3.5.1 Pleuronectidae 71
3.5.2 Achiridae 72
3.5.3 Paralichthyidae 72
3.5.4 New World tropical flatfishes 73
3.5.5 Indo-west Pacific region 73
Acknowledgements 76
References 76
4 Life-history traits in flatfishes 83
Catarina Vinagre and Henrique N. Cabral
4.1 Introduction 83
4.2 Diversity in life-history traits of flatfishes 85
4.3 Variation according to geographical area, habitat use patterns and functional guilds 86
4.4 Intraspecies variability 89
4.4.1 Phenotypic plasticity, local adaptation, cogradient variation and parental effects 93
4.5 Anthropogenic impacts on life-history traits 94
4.6 Future directions 95
References 96
5 Ecology of reproduction 101
Adriaan D. Rijnsdorp, Cindy J.G. van Damme and Peter R. Witthames
5.1 Introduction 101
5.2 Spawning 102
5.2.1 Spawning behaviour 102
5.2.2 Spawning mode 102
5.2.3 Egg size 102
5.2.4 Spawning season 103
5.2.5 Duration of spawning 106
5.3 Gonad development 106
5.3.1 Testis 106
5.3.2 Ovary 107
5.3.3 Fecundity 110
5.3.4 Geographical pattern in fecundity 112
5.3.5 Batch spawning 113
5.3.6 Egg and sperm quality: maternal and paternal effects 113
5.4 Age and size at first maturation 114
5.5 Energetics 115
5.5.1 Energetics of reproduction and growth 115
5.5.2 Nonannual spawning 117
5.5.3 Spawning fast 118
5.5.4 Sexual dimorphism in reproduction and growth 119
5.6 Fisheries-induced evolution in reproduction and growth 120
5.7 Reproductive potential 121
References 123
6 The planktonic stages of flatfishes: physical and biological interactions in transport processes 132
Janet T. Duffy-Anderson, Kevin M. Bailey, Henrique N. Cabral, Hideaki Nakata and Henk W. van der Veer
6.1 Introduction 133
6.2 Variations in time and space in the plankton 134
6.3 Physical mechanisms of transport and retention 136
6.3.1 Wind-forcing & Ekman transport 136
6.3.2 Estuarine circulation 137
6.3.3 Fronts and eddies 138
6.3.4 Influence of climate and oceanographic shifts 138
6.3.5 Behaviour 141
6.3.6 Models 141
6.4 Adaptations to transport conditions: geographical and species comparisons 146
6.4.1 Comparisons among species within a geographic region 148
6.4.2 Congeneric comparisons in different regions 151
6.4.3 Conspecific comparisons in different geographic areas 151
6.4.4 Local adaptations 153
6.5 Transitioning from the plankton 154
6.5.1 Criticality of timing 154
6.5.2 Fidelity to initial touchdown sites 155
6.5.3 Importance of initial settlement areas 155
6.6 Implications 156
6.6.1 Population genetics 156
6.6.2 Recruitment 157
6.6.3 Connectivity 158
6.6.4 Management 159
6.6.5 Research needs 160
Acknowledgements 161
References 161
7 Development and regulation of external asymmetry during flatfish metamorphosis 171
Tohru Suzuki and Masaru Tanaka
7.1 Introduction 171
7.2 Development and evolution of flatfish external asymmetry 172
7.3 Regulation of flatfish eye-sidedness 174
7.4 Pigmentation 177
7.5 Hormonal regulation 180
7.6 Summary and future work 181
Acknowledgements 182
References 182
8 Recruitment level and variability 185
Henk W. van der Veer, Vania Freitas and William C. Leggett
8.1 Introduction 185
8.2 Range of distribution 187
8.3 Average recruitment levels 189
8.4 Recruitment variability 192
8.4.1 Processes influencing recruitment variability 194
8.4.2 Recruitment variability in flatfishes relative to other marine fish species 198
8.5 Future perspectives 199
References 200
9 Age and growth 207
Richard D.M. Nash and Audrey J. Geffen
9.1 Introduction 207
9.2 Age estimation 209
9.2.1 Larvae and juveniles 209
9.2.2 Adults 211
9.3 Growth of larvae 211
9.3.1 Variation in growth 212
9.3.2 Factors affecting larval growth 212
9.4 Growth during metamorphosis 213
9.5 Growth on the nursery grounds 217
9.5.1 Growth models and growth experiments 218
9.5.2 Maximum achievable growth and evidence for deviations from maximum growth 218
9.5.3 Growth compensation and depensation 220
9.5.4 Nursery ground quality and the use of growth as an indicator of habitat quality 221
9.6 Growth of adults 222
9.6.1 Factors affecting adult growth rates 223
9.6.2 Tradeoff between growth and reproduction 223
9.7 Longevity 225
References 227
10 Distribution and dynamics of habitat use by juvenile and adult flatfishes 242
Kenneth W. Able and F. Joel Fodrie
10.1 Introduction 242
10.2 Distribution of habitat associations 243
10.2.1 Effects of spatial scale on habitat use and selection 245
10.3 Nursery role of juvenile habitats 247
10.4 Dynamics of habitat associations 252
10.4.1 Settlement 253
10.4.2 Ontogeny 253
10.4.3 Long-term changes 256
10.4.4 Tidal, diel and seasonal cycles 257
10.4.5 Migrations and site fidelity 259
10.4.6 Episodic events 261
10.5 Future emphasis 262
Acknowledgements 264
References 264
11 The trophic ecology of flatfishes 283
Jason S. Link, Brian E. Smith, David B. Packer, Michael J. Fogarty and Richard W. Langton
11.1 Introduction 283
11.2 Major flatfish feeding groups 284
11.2.1 Polychaete and crustacean eaters 292
11.2.2 Piscivores 293
11.2.3 Specialists 295
11.2.4 Other considerations 296
11.2.4.1 Anthropogenically produced food 296
11.2.4.2 Seasonality 296
11.2.4.3 Ontogeny 296
11.2.4.4 Spatial factors 297
11.3 Flatfish predators 297
11.4 Flatfish competitors 298
11.5 Flatfish trophic dynamics: a case study of Georges Bank 300
11.5.1 Shifts in abundance and species composition 300
11.5.2 Potential competitive interactions 301
11.5.3 Predation by flatfishes 302
11.5.4 Have changes in flatfish populations influenced the Georges Bank ecosystem? 304
11.6 Summary and conclusions 304
Acknowledgements 305
References 305
12 The behaviour of flatfishes 314
Robin N. Gibson, Allan W. Stoner and Clifford H. Ryer
12.1 Introduction 314
12.2 Locomotion and related behaviour 315
12.2.1 Locomotion 315
12.2.2 Burying 316
12.3 Reproduction 317
12.4 Feeding 317
12.4.1 Flatfish feeding types 317
12.4.2 Feeding behaviour 318
12.4.2.1 Search 318
12.4.2.2 Encounter and detection 320
12.4.2.3 Capture 320
12.4.3 External factors modifying feeding behaviour 321
12.5 Predation and reactions to predators 323
12.5.1 Burial and the role of sediment 323
12.5.2 Cryptic colouration and behaviour 324
12.5.3 Escape from predators following attack 325
12.5.4 Predator avoidance through habitat choice 325
12.5.5 Effect of size on vulnerability and avoidance of ingestion 326
12.6 Movements, migrations and rhythms 326
12.7 Behaviour in relation to fishing 329
12.7.1 Reactions to mobile fishing gear 329
12.7.2 Reactions to fixed gear 331
12.8 Behaviour in relation to aquaculture and stock enhancement 331
12.9 Conclusions 332
References 333
13 Atlantic flatfish fisheries 346
Stephen J. Walsh, Juan M. Diaz de Astarloa and Jan-Jaap Poos
13.1 Introduction 346
13.2 Main species and nature of the fisheries 348
13.2.1 North-west Atlantic 348
13.2.2 North-east Atlantic 351
13.2.3 Southern Atlantic 355
13.2.3.1 South-west Atlantic 355
13.2.3.2 South-east Atlantic 358
13.3 History of exploitation 360
13.3.1 North-west Atlantic 360
13.3.1.1 West Greenland 363
13.3.1.2 Canada 363
13.3.1.3 United States 365
13.3.2 North-east Atlantic 366
13.3.3 Southern Atlantic 367
13.3.3.1 Fishing fleets and fishing gears 367
13.3.3.2 The fisheries 368
13.4 Economic importance 371
13.4.1 North-west Atlantic 371
13.4.1.1 West Greenland 371
13.4.1.2 Canada 372
13.4.1.3 United States 372
13.4.1.4 Employment 372
13.4.2 North-east Atlantic 375
13.4.2.1 Employment 376
13.4.3 Southern Atlantic 377
13.5 Management 378
13.5.1 North-west Atlantic 378
13.5.1.1 International Commission for the Northwest Atlantic Fisheries (ICNAF) 378
13.5.1.2 Northwest Atlantic Fisheries Organization (NAFO) 378
13.5.1.3 Fisheries and Oceans Canada 380
13.5.1.4 United States National Marine Fisheries Service 381
13.5.2 North-east Atlantic 384
13.5.3 Southern Atlantic 386
13.6 Notes 387
Acknowledgements 388
References 388
14 Pacific flatfish fisheries 395
Thomas Wilderbuer, Bruce Leaman and Chang Ik Zhang
14.1 Introduction 395
14.2 Main species and nature of fisheries 396
14.3 History of exploitation 401
14.3.1 General account 401
14.3.2 Republic of Korea 403
14.3.3 Japan 403
14.3.4 Russia (including the former Soviet Union) 404
14.3.5 Canada 404
14.3.6 United States 405
14.3.7 New Zealand 406
14.3.8 Australia 407
14.4 Economic importance 407
14.5 Management 408
14.5.1 Western North Pacific 408
14.5.2 Eastern North Pacific 409
14.5.3 Australia and New Zealand 411
14.5.4 Data collection 412
14.5.5 Ecosystem-based fisheries management 413
References 414
15 Tropical flatfish fisheries 418
Thomas A. Munroe
15.1 Introduction 418
15.2 Main species and nature of the fisheries 420
15.2.1 Habitats 420
15.2.2 Commercially important species and/or taxa 422
15.2.3 Nature of the fisheries 425
15.2.4 Types of gear employed 428
15.2.5 Harvest on spawning concentrations, migrating stocks and impacts on recruitment 428
15.2.6 Industrial versus artisanal characteristics of the fisheries 429
15.3 History of exploitation 430
15.3.1 Commercial landings 430
15.3.2 Geographic occurrence and historical landings 433
15.4 Importance 441
15.4.1 Economic importance 441
15.4.2 Human importance 443
15.5 Management and conservation 443
15.5.1 Fishery conflicts, regulations and management 443
15.5.2 Conservation 448
Acknowledgements 450
References 450
16 Assessment and management of flatfish stocks 461
Steven X. Cadrin, William G. Clark and Daniel Ricard
16.1 Concepts and terms 461
16.2 Population dynamics, assessment, and management 464
16.2.1 Stock and recruitment 467
16.2.2 Recruitment, environment, assessment and management 475
16.2.3 Assessment, management, and uncertainty 477
16.3 Assessment and management summary 478
16.3.1 North-east Pacific 478
16.3.2 North-west Atlantic 478
16.3.3 North-east Atlantic 483
16.4 Conclusions 484
Acknowledgements 484
References 485
17 Synergies between aquaculture and fisheries 491
Audrey J. Geffen, Karin Pittman and Albert K. Imsland
17.1 Introduction 491
17.2 Species 492
17.3 Population structure and genomics 494
17.4 Life history stages 497
17.4.1 Egg and larval stages 498
17.4.2 Metamorphosis 499
17.4.3 Growth 502
17.4.4 Reproduction 502
17.5 Future directions for common goals and synergies between fisheries and aquaculture 505
References 508
Appendix A: List of scientific and common names of living flatfishes used in the book 519
Appendix B: Common synonyms of Pleuronectidae used in the text 523
Index of scientific and common names 525
Subject index 535
