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.