Flatfishes

Robin N. Gibson is an Honorary Research Fellow of the Scottish Association for Marine Science, Scottish Marine Institute, Oban, Scotland. His lifetime research interests have been in the ecology and behaviour of intertidal and shallow water marine fishes. Richard D.M. Nash is a senior research scientist at the Institute of Marine Research, Bergen, Norway. His research is mainly concerned with recruitment processes and the early life history dynamics of marine fishes. He has particular research interests in the dynamics of nursery grounds. Audrey J. Geffen is a professor in the Department of Biology, University of Bergen, Norway. Her research is primarily concerned with fish growth in natural and culture environments. A significant aspect of her work involves theoretical and applied research on the growth and formation of fish otoliths, analysing the structure and composition as a record of life history. Henk W. van der Veer is a senior scientist at the Royal Netherlands Institute for Sea Research, The Netherlands. His main research topics are related to fish recruitment, concentrating on flatfishes and the functioning of coastal nursery areas. His work is strongly focussed on field observations supported by and embedded in ecological theory, especially the Dynamic Energy Budget theory.

• List of contributors xvSeries editor’s foreword xxiPreface to the second edition xxvPreface to the first edition xxviiAcknowledgements xxix1 Introduction 1Robin N. Gibson1.1 The fascination of flatfishes 11.2 A brief history of flatfish research and its contribution to fish biology and fisheries science 31.3 Scope and contents of the book 41.4 Nomenclature 9Acknowledgements 10References 102 Systematic diversity of the Pleuronectiformes 13Thomas A. Munroe2.1 Introduction 132.2 Systematic profile of the Pleuronectiformes 182.3 Intrarelationships of the Pleuronectiformes 192.4 Brief synopses of the suborders and families 222.5 Diversity of the Pleuronectiformes 262.5.1 Overview 262.5.2 Flatfish species diversity 272.5.3 Diversity of species within families 282.5.4 Standing diversity estimate for species of Pleuronectiformes 292.5.5 Relative diversity of the Pleuronectiformes 312.6 Patterns of species discovery among pleuronectiform families 322.6.1 History 322.6.2 Factors contributing to new species discovery among the Pleuronectiformes 352.6.2.1 Systematic activities 352.6.2.2 Geographic region 372.6.2.3 Depth 392.6.2.4 Size 402.7 Conclusions 42Acknowledgements 44References 443 Distributions and biogeography 52Thomas A. Munroe3.1 Introduction 523.2 Geographic distribution of pleuronectiform lineages 563.3 Global patterns of species richness for the Pleuronectiformes 613.3.1 Latitudinal gradients in species richness 613.3.2 Tropical and subtropical regions 613.3.3 Temperate regions 623.3.4 Species richness on continental shelves 633.3.5 Insular versus continental regions 643.3.6 Continental versus oceanic islands 663.4 Species richness in specific environments 663.4.1 Freshwater environments 663.4.2 Antarctic Ocean 673.4.3 Arctic Ocean 683.4.4 Shallow-water versus deep-sea habitats 693.5 Historical biogeography 713.5.1 Pleuronectidae 713.5.2 Achiridae 723.5.3 Paralichthyidae 723.5.4 New World tropical flatfishes 733.5.5 Indo-west Pacific region 73Acknowledgements 76References 764 Life-history traits in flatfishes 83Catarina Vinagre and Henrique N. Cabral4.1 Introduction 834.2 Diversity in life-history traits of flatfishes 854.3 Variation according to geographical area, habitat use patterns and functional guilds 864.4 Intraspecies variability 894.4.1 Phenotypic plasticity, local adaptation, cogradient variation and parental effects 934.5 Anthropogenic impacts on life-history traits 944.6 Future directions 95References 965 Ecology of reproduction 101Adriaan D. Rijnsdorp, Cindy J.G. van Damme and Peter R. Witthames5.1 Introduction 1015.2 Spawning 1025.2.1 Spawning behaviour 1025.2.2 Spawning mode 1025.2.3 Egg size 1025.2.4 Spawning season 1035.2.5 Duration of spawning 1065.3 Gonad development 1065.3.1 Testis 1065.3.2 Ovary 1075.3.3 Fecundity 1105.3.4 Geographical pattern in fecundity 1125.3.5 Batch spawning 1135.3.6 Egg and sperm quality: maternal and paternal effects 1135.4 Age and size at first maturation 1145.5 Energetics 1155.5.1 Energetics of reproduction and growth 1155.5.2 Nonannual spawning 1175.5.3 Spawning fast 1185.5.4 Sexual dimorphism in reproduction and growth 1195.6 Fisheries-induced evolution in reproduction and growth 1205.7 Reproductive potential 121References 1236 The planktonic stages of flatfishes: physical and biological interactions in transport processes 132Janet T. Duffy-Anderson, Kevin M. Bailey, Henrique N. Cabral, Hideaki Nakata and Henk W. van der Veer6.1 Introduction 1336.2 Variations in time and space in the plankton 1346.3 Physical mechanisms of transport and retention 1366.3.1 Wind-forcing & Ekman transport 1366.3.2 Estuarine circulation 1376.3.3 Fronts and eddies 1386.3.4 Influence of climate and oceanographic shifts 1386.3.5 Behaviour 1416.3.6 Models 1416.4 Adaptations to transport conditions: geographical and species comparisons 1466.4.1 Comparisons among species within a geographic region 1486.4.2 Congeneric comparisons in different regions 1516.4.3 Conspecific comparisons in different geographic areas 1516.4.4 Local adaptations 1536.5 Transitioning from the plankton 1546.5.1 Criticality of timing 1546.5.2 Fidelity to initial touchdown sites 1556.5.3 Importance of initial settlement areas 1556.6 Implications 1566.6.1 Population genetics 1566.6.2 Recruitment 1576.6.3 Connectivity 1586.6.4 Management 1596.6.5 Research needs 160Acknowledgements 161References 1617 Development and regulation of external asymmetry during flatfish metamorphosis 171Tohru Suzuki and Masaru Tanaka7.1 Introduction 1717.2 Development and evolution of flatfish external asymmetry 1727.3 Regulation of flatfish eye-sidedness 1747.4 Pigmentation 1777.5 Hormonal regulation 1807.6 Summary and future work 181Acknowledgements 182References 1828 Recruitment level and variability 185Henk W. van der Veer, Vania Freitas and William C. Leggett8.1 Introduction 1858.2 Range of distribution 1878.3 Average recruitment levels 1898.4 Recruitment variability 1928.4.1 Processes influencing recruitment variability 1948.4.2 Recruitment variability in flatfishes relative to other marine fish species 1988.5 Future perspectives 199References 2009 Age and growth 207Richard D.M. Nash and Audrey J. Geffen9.1 Introduction 2079.2 Age estimation 2099.2.1 Larvae and juveniles 2099.2.2 Adults 2119.3 Growth of larvae 2119.3.1 Variation in growth 2129.3.2 Factors affecting larval growth 2129.4 Growth during metamorphosis 2139.5 Growth on the nursery grounds 2179.5.1 Growth models and growth experiments 2189.5.2 Maximum achievable growth and evidence for deviations from maximum growth 2189.5.3 Growth compensation and depensation 2209.5.4 Nursery ground quality and the use of growth as an indicator of habitat quality 2219.6 Growth of adults 2229.6.1 Factors affecting adult growth rates 2239.6.2 Tradeoff between growth and reproduction 2239.7 Longevity 225References 22710 Distribution and dynamics of habitat use by juvenile and adult flatfishes 242Kenneth W. Able and F. Joel Fodrie10.1 Introduction 24210.2 Distribution of habitat associations 24310.2.1 Effects of spatial scale on habitat use and selection 24510.3 Nursery role of juvenile habitats 24710.4 Dynamics of habitat associations 25210.4.1 Settlement 25310.4.2 Ontogeny 25310.4.3 Long-term changes 25610.4.4 Tidal, diel and seasonal cycles 25710.4.5 Migrations and site fidelity 25910.4.6 Episodic events 26110.5 Future emphasis 262Acknowledgements 264References 26411 The trophic ecology of flatfishes 283Jason S. Link, Brian E. Smith, David B. Packer, Michael J. Fogarty and Richard W. Langton11.1 Introduction 28311.2 Major flatfish feeding groups 28411.2.1 Polychaete and crustacean eaters 29211.2.2 Piscivores 29311.2.3 Specialists 29511.2.4 Other considerations 29611.2.4.1 Anthropogenically produced food 29611.2.4.2 Seasonality 29611.2.4.3 Ontogeny 29611.2.4.4 Spatial factors 29711.3 Flatfish predators 29711.4 Flatfish competitors 29811.5 Flatfish trophic dynamics: a case study of Georges Bank 30011.5.1 Shifts in abundance and species composition 30011.5.2 Potential competitive interactions 30111.5.3 Predation by flatfishes 30211.5.4 Have changes in flatfish populations influenced the Georges Bank ecosystem? 30411.6 Summary and conclusions 304Acknowledgements 305References 30512 The behaviour of flatfishes 314Robin N. Gibson, Allan W. Stoner and Clifford H. Ryer12.1 Introduction 31412.2 Locomotion and related behaviour 31512.2.1 Locomotion 31512.2.2 Burying 31612.3 Reproduction 31712.4 Feeding 31712.4.1 Flatfish feeding types 31712.4.2 Feeding behaviour 31812.4.2.1 Search 31812.4.2.2 Encounter and detection 32012.4.2.3 Capture 32012.4.3 External factors modifying feeding behaviour 32112.5 Predation and reactions to predators 32312.5.1 Burial and the role of sediment 32312.5.2 Cryptic colouration and behaviour 32412.5.3 Escape from predators following attack 32512.5.4 Predator avoidance through habitat choice 32512.5.5 Effect of size on vulnerability and avoidance of ingestion 32612.6 Movements, migrations and rhythms 32612.7 Behaviour in relation to fishing 32912.7.1 Reactions to mobile fishing gear 32912.7.2 Reactions to fixed gear 33112.8 Behaviour in relation to aquaculture and stock enhancement 33112.9 Conclusions 332References 33313 Atlantic flatfish fisheries 346Stephen J. Walsh, Juan M. Díaz de Astarloa and Jan-Jaap Poos13.1 Introduction 34613.2 Main species and nature of the fisheries 34813.2.1 North-west Atlantic 34813.2.2 North-east Atlantic 35113.2.3 Southern Atlantic 35513.2.3.1 South-west Atlantic 35513.2.3.2 South-east Atlantic 35813.3 History of exploitation 36013.3.1 North-west Atlantic 36013.3.1.1 West Greenland 36313.3.1.2 Canada 36313.3.1.3 United States 36513.3.2 North-east Atlantic 36613.3.3 Southern Atlantic 36713.3.3.1 Fishing fleets and fishing gears 36713.3.3.2 The fisheries 36813.4 Economic importance 37113.4.1 North-west Atlantic 37113.4.1.1 West Greenland 37113.4.1.2 Canada 37213.4.1.3 United States 37213.4.1.4 Employment 37213.4.2 North-east Atlantic 37513.4.2.1 Employment 37613.4.3 Southern Atlantic 37713.5 Management 37813.5.1 North-west Atlantic 37813.5.1.1 International Commission for the Northwest Atlantic Fisheries (ICNAF) 37813.5.1.2 Northwest Atlantic Fisheries Organization (NAFO) 37813.5.1.3 Fisheries and Oceans Canada 38013.5.1.4 United States National Marine Fisheries Service 38113.5.2 North-east Atlantic 38413.5.3 Southern Atlantic 38613.6 Notes 387Acknowledgements 388References 38814 Pacific flatfish fisheries 395Thomas Wilderbuer, Bruce Leaman and Chang Ik Zhang14.1 Introduction 39514.2 Main species and nature of fisheries 39614.3 History of exploitation 40114.3.1 General account 40114.3.2 Republic of Korea 40314.3.3 Japan 40314.3.4 Russia (including the former Soviet Union) 40414.3.5 Canada 40414.3.6 United States 40514.3.7 New Zealand 40614.3.8 Australia 40714.4 Economic importance 40714.5 Management 40814.5.1 Western North Pacific 40814.5.2 Eastern North Pacific 40914.5.3 Australia and New Zealand 41114.5.4 Data collection 41214.5.5 Ecosystem-based fisheries management 413References 41415 Tropical flatfish fisheries 418Thomas A. Munroe15.1 Introduction 41815.2 Main species and nature of the fisheries 42015.2.1 Habitats 42015.2.2 Commercially important species and/or taxa 42215.2.3 Nature of the fisheries 42515.2.4 Types of gear employed 42815.2.5 Harvest on spawning concentrations, migrating stocks and impacts on recruitment 42815.2.6 Industrial versus artisanal characteristics of the fisheries 42915.3 History of exploitation 43015.3.1 Commercial landings 43015.3.2 Geographic occurrence and historical landings 43315.4 Importance 44115.4.1 Economic importance 44115.4.2 Human importance 44315.5 Management and conservation 44315.5.1 Fishery conflicts, regulations and management 44315.5.2 Conservation 448Acknowledgements 450References 45016 Assessment and management of flatfish stocks 461Steven X. Cadrin, William G. Clark and Daniel Ricard16.1 Concepts and terms 46116.2 Population dynamics, assessment, and management 46416.2.1 Stock and recruitment 46716.2.2 Recruitment, environment, assessment and management 47516.2.3 Assessment, management, and uncertainty 47716.3 Assessment and management summary 47816.3.1 North-east Pacific 47816.3.2 North-west Atlantic 47816.3.3 North-east Atlantic 48316.4 Conclusions 484Acknowledgements 484References 48517 Synergies between aquaculture and fisheries 491Audrey J. Geffen, Karin Pittman and Albert K. Imsland17.1 Introduction 49117.2 Species 49217.3 Population structure and genomics 49417.4 Life history stages 49717.4.1 Egg and larval stages 49817.4.2 Metamorphosis 49917.4.3 Growth 50217.4.4 Reproduction 50217.5 Future directions for common goals and synergies between fisheries and aquaculture 505References 508Appendix A: List of scientific and common names of living flatfishes used in the book 519Appendix B: Common synonyms of Pleuronectidae used in the text 523Index of scientific and common names 525Subject index 535