Flatfishes (2^nd Ed.)
Biology and Exploitation
Fish and Aquatic Resources Series

Fascinating and instantly recognizable, flatfishes are unique in their asymmetric postlarval body form. With over 800 extant species recognized and a distribution stretching around the globe, these fishes are of considerable research interest and provide a major contribution to commercial and recreational fisheries worldwide. This second edition of Flatfishes: Biology and Exploitation has been completely revised, updated and enlarged to respond to the ever-growing body of research. It provides:

? Overviews of systematics, distribution, life history strategies, reproduction, recruitment, ecology and behaviour
? Descriptions of the major fisheries and their management
? An assessment of the synergies between ecological and aquaculture research of flatfishes.

Carefully compiled and edited by four internationally-known scientists and with chapters written by many world leaders in the field, this excellent new edition of a very popular and successful book is essential reading for fish biologists, fisheries scientists, marine biologists, aquaculture personnel, ecologists, environmental scientists, and government workers in fisheries and fish and wildlife departments. Flatfishes: Biology and Exploitation, Second Edition, should be found in all libraries of research establishments and universities where life sciences, fish biology, fisheries, aquaculture, marine sciences, oceanography, ecology and environmental sciences are studied and taught.

Reviews of the First Edition

? A solid, up-to-date book that advanced students and research scientists with interests in fish biology will find interesting and useful. Aquaculture International
? A data-rich book that outlines much of what you might ever want to know about flatfishes. Fish & Fisheries
? Well presented with clear illustrations and a valuable source of information for those with a general interest in fish ecology or for the more specialist reader. You should make sure that your library has a copy. J Fish Biology
? An excellent and very practical overview of the whole, global flatfish scene. Anyone interested in flatfish at whichever stage of the economic food chain should invest in a copy immediately. Ausmarine
? Because of the high quality of each chapter, written by international experts, it is a valuable reference. Reviews in Fish Biology and Fisheries

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

2.7 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

11.3 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

12.4.3 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. Díaz 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

13.3 History of exploitation 360

13.3.1 North-west Atlantic 360

13.3.2 North-east Atlantic 366

13.3.3 Southern Atlantic 367

13.4 Economic importance 371

13.4.1 North-west Atlantic 371

13.4.2 North-east Atlantic 375

13.4.3 Southern Atlantic 377

13.5 Management 378

13.5.1 North-west Atlantic 378

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