Scallops: Biology, Ecology and Aquaculture

Table of Contents

Dedication v
Preface vii
Preface from first edition ix
List of contributors xi

Chapter 1. New Phylogenies of the Pectinidae (Mollusca: Bivalvia): Reconciling Morphological and Molecular Approaches
Thomas R. Waller

1.1 Introduction 1

1.1.1 Molecular genetic studies 1

1.1.2 Methods and materials 7

1.2 New observations 10

1.2.1 Subfamily Camptonectinae Habe, 1977 10

1.2.2 Subfamily Palliolinae Korobkov in Eberzin, 1960 10

1.2.2.1 Tribe Pseudentoliini, new tribe 12

1.2.2.2 Tribe Adamussiini Habe, 1977 13

1.2.2.3 Tribe Eburneopectinini, new tribe 14

1.2.2.4 Tribe Serripectinini, new tribe 15

1.2.2.5 Tribe Palliolini Korobkov in Eberzin, 1960 17

1.2.2.6 Tribe Mesopeplini, new tribe 20

1.2.3 Origins of the Decatopecten and Pecten Groups 22

1.2.4 Eastern Atlantic Pecten 28

1.3 Conclusions 30
Acknowledgments 32
References 32
Appendix 43

Chapter 2. Development, Physiology, Behaviour and Ecology of Scallop Larvae
Simon M. Cragg

2.1 Introduction 45

2.2 Scallop life history characteristics 45

2.3 Larval development 55

2.3.1 Embryogenesis 55

2.3.2 Development of the larval shell 56

2.3.3 Organogenesis 59

2.3.3.1 Development of ciliation, the prototroch and the velum 59

2.3.3.2 Larval mantle 64

2.3.3.3 Musculature 64

2.3.3.4 Digestive tract 68

2.3.3.5 Sense organs 68

2.3.3.6 Apical organ 68

2.3.3.7 Statocysts 73

2.3.3.8 Eye spots 73

2.3.3.9 Mantle ciliation 74

2.3.3.10 Foot 77

2.3.3.11 Nervous system 78

2.3.3.12 Other organs 79

2.3.4 Metamorphosis 79

2.4 Comparative anatomy 81

2.5 Functional morphology 84

2.5.1 Locomotion 84

2.5.2 Feeding 85

2.5.3 Respiration 85

2.5.4 Sensory reception 86

2.6 Physiology and effects of environmental variables 87

2.6.1 Energy reserves and feeding 87

2.6.2 Respiration 88

2.6.3 Growth 89

2.6.4 Biochemical events during metamorphosis 90

2.7 Behaviour and larval distribution 90

2.7.1 Characteristics of locomotion during larval development 90

2.7.2 Responses to stimuli 91

2.7.3 Distribution of larvae 94

2.7.4 Development and the larval environment 98

2.7.5 Tolerance of environmental variables 99

2.8 Rearing methods 101

2.8.1 Spawning and manipulation of zygotes 101

2.8.2 Feeding 102

2.8.3 Control of disease in cultures 103

2.8.4 Collection of spat from wild stocks 104
References 105

Chapter 3. Structure and Function in Scallops
Peter G. Beninger and Marcel Le Pennec

3.1 Introduction 123

3.1.1 An overview of the scallop body 123

3.2 The mantle and its derivatives 126

3.2.1 Gross functional anatomy 127

3.2.2 Microanatomy and functions 130

3.3 Pallial organs and particle processing 132

3.3.1 Gills 132

3.3.1.1 Gill axis and arch 135

3.3.1.2 Principal filaments and dorsal expansion 135

3.3.1.3 Ordinary filament 137

3.3.1.4 Haemolymph circulation in the gill 141

3.3.1.5 Particle processing on the gill 141

3.3.1.6 Particle selection at the gill 141

3.3.1.7 Particle retention lower size limit 143

3.3.1.8 Ingestion volume regulation on the gill 143

3.3.2 Labial palps and lips 144

3.3.2.1 Labial palps 144

3.3.2.2 Particle processing on the labial palps 144

3.3.2.3 Lips 147

3.4 Digestive system and digestion 150

3.4.1 Mouth and oesophagus 150

3.4.2 Stomach, crystalline style and gastric shield 150

3.4.3 The digestive gland 154

3.4.4 Intestine, rectum and anus 157

3.4.5 Digestive sites and postingestive selection 159

3.5 Cardio-Vascular system 159

3.5.1 General circulation 160

3.5.1.1 The arterial system 160

3.5.1.1.1 Anterior aorta 160

3.5.1.1.2 Posterior aorta 160

3.5.1.2 The venous system 162

3.5.2 The heart 162

3.5.2.1 The ventricle 162

3.5.2.2 The auricles and their excretory structures 163

3.5.2.3 Structure and ultrastructure of heart cells 163

3.5.2.4 Contraction 165

3.5.2.4.1 Refilling 165

3.5.2.4.2 Coordination of alternate AV beat 166

3.5.2.4.3 Pacemaker mechanism 166

3.5.2.4.4 Regulation of pacemaker 167

3.5.3 Haemolymph 167

3.5.3.1 Plasma 168

3.5.3.2 Haemocytes 168

3.5.3.2.1 Haemocyte types 169

3.5.3.2.2 Functions of haemocytes 169

3.6 Excretory system 170

3.6.1 Pericardial (auricular) glands 170

3.6.2 Kidney 172

3.6.3 Functions of the kidney and pericardial glands 173

3.7 Reproductive system 173

3.7.1 Sexuality: Gonochory, hermaphroditism and their variants 173

3.7.2 Origin and formation of the gonad 176

3.7.3 Anatomy, histology and ultrastructure of the adult gonad 177

3.7.3.1 Outer epithelium 177

3.7.3.2 Perigonadal connective tissue 179

3.7.3.3 Inter-acinal connective tissue 179

3.7.3.4 Haemolymph sinuses 179

3.7.3.5 Acini 180

3.7.3.6 Evacuating ducts 180

3.7.4 Gametogenesis 180

3.7.4.1 Oogenesis 180

3.7.4.1.1 Premeiotic stage 180

3.7.4.1.2 Previtellogenic stage 181

3.7.4.1.3 Vitellogenesis and metabolite transport to the oocyte 181

3.7.5 Oocyte atresia 185

3.7.6 Spermatogenesis, spermatozoon ultrastructure and taxonomy 188

3.7.7 Fertilisation 190

3.8 Nervous and sensory systems 191

3.8.1 General organisation of the nervous system and functional
anatomy of principal ganglia 191

3.8.1.1 Cerebral and pedal ganglia 191

3.8.1.2 The parietovisceral ganglion and its nerves 191

3.8.1.3 Histology and neurosecretions of the ganglia 196

3.8.1.4 The circumpallial nerve 198

3.8.2 Sensory structures 198

3.8.2.1 Visual system 198

3.8.2.2 Epithelial sensory cells and tentacles 198

3.8.2.3 Abdominal sense organ 199

3.8.2.4 Osphradia 201

3.8.2.5 Statoreceptors 204

3.8.3 Neurotransmitters and neurohormones 204

3.9 Foot-byssal complex 205

3.9.1 External morphology and development of the foot-byssal complex 207

3.9.2 Anatomy and histology of the foot-byssal gland complex 208

3.9.2.1 The protein gland 208

3.9.2.2 The enzyme gland 208

3.9.2.3 The byssus 210

3.9.3 Functioning of the foot-byssal complex 210
Acknowledgments 210
References 211

Chapter 4. Scallop Adductor Muscles: Structure and Function
Peter D. Chantler

4.1 Introduction 229

4.2 Structure of the striated adductor muscle 231

4.2.1 Fibre microanatomy 231

4.2.2 Actin and thin filament structure 233

4.2.3 Thick filament structure 240

4.3 Structure of the smooth adductor muscle 243

4.3.1 Fibre microanatomy 243

4.3.2 Thin filament structure 245

4.3.3 The structure of paramyosin-rich thick filaments 247

4.4 Myosin 253

4.5 Function of the striated adductor 271

4.5.1 Mechanics 271

4.5.2 The interaction of myosin with actin 272

4.5.3 The crossbridge cycle 277

4.5.4 Myosin-linked regulation 282

4.6 Function of the smooth adductor 289

4.6.1 Physiology 289

4.6.2 Catch mechanism 292

4.7 Achievements and goals 296
Acknowledgments 298
References 298

Chapter 5. Neurobiology and Behaviour of the Scallop
Lon A. Wilkens

5.1 Introduction 317

5.2 The visual system 317

5.2.1 Functional anatomy of the eyes 317

5.2.2 Retinal physiology 322

5.2.3 Receptor potential biophysics 324

5.2.4 Anatomy and physiology of vision in the central nervous system 328

5.2.5 Visual behaviours 332

5.3 The locomotory system 335

5.3.1 Escape responses and swimming 335

5.3.2 Sensory and motor functions of the mantle 341

5.3.3 Innervation and neuromuscular physiology of the adductor muscle 345

5.3.4 Coordination of locomotory behaviour by the central nervous
system 348
Acknowledgments 352
References 352

Chapter 6. Reproductive Physiology
Bruce J. Barber and Norman J. Blake

6.1 Introduction 357

6.2 Gametogenic cycles 358

6.2.1 Definition 358

6.2.2 Means of assessment 358

6.2.2.1 Visual observation 360

6.2.2.2 Gonad mass and index 360

6.2.2.3 Histology 367

6.2.2.4 Abundance of larvae and spat 369

6.2.3 Variations in gametogenic cycles 370

6.2.3.1 Intra-specific variations 370

6.2.3.2 Inter-specific variations 375

6.3 Regulation of gametogenic cycles 375

6.3.1 Gametogenesis and fecundity 376

6.3.1.1 Exogenous regulation 376

6.3.1.2 Endogenous regulation 381

6.3.2 Spawning and spawning synchrony 385

6.3.2.1 Exogenous regulation 385

6.3.2.2 Endogenous regulation 389

6.4 Energy metabolism 390

6.4.1 Tissue weights and indices 391

6.4.2 Tissue biochemical composition 392

6.4.3 Physiological indices 395

6.4.4 Radiotracer experiments 398

6.4.5 Ultrastructure and histochemistry 398

6.4.6 Mechanisms 400

6.5 Applications to aquaculture 401

6.5.1 Broodstock conditioning 401

6.5.2 Spawning 403

6.5.3 Larval growth and survival 403

6.6 Summary 404

6.6.1 Gametogenic cycles 404

6.6.2 Regulation of gametogenesis 404

6.6.3 Energy metabolism 405

6.6.4 Applications to aquaculture 406
References 406

Chapter 7. Physiology: Energy Acquisition and Utilisation
Bruce A. MacDonald, V. Monica Bricelj and Sandra E. Shumway

7.1 Introduction 417

7.2 Energy acquisition 417

7.2.1 Food sources 417

7.2.2 Feeding currents and mechanisms of particle capture 420

7.2.3 Particle retention efficiency 421

7.2.4 Feeding rates 424

7.2.5 Clearance rate in relation to food concentration 427

7.2.6 Influence of temperature on feeding rates 430

7.2.7 Pseudofeces production, pre- and post-ingestive particle selection 432

7.2.8 Absorption efficiency 436

7.2.9 Effects of suspended sediments on feeding and growth 439

7.2.10 Effects of flow on feeding and growth 445

7.2.11 Effects of harmful and toxic algae 447

7.3 Energy utilisation: Metabolic expenditure 453

7.3.1 Metabolic rate and oxygen availability 455

7.3.2 Metabolic cost of reproduction 458

7.3.3 Metabolic rate in relation to temperature and latitude 460

7.3.4 Metabolic rate in relation to activity levels 461

7.3.5 Anaerobic metabolism 463

7.4 Energy utilisation 467

7.4.1 Excretion and byssus secretion 467

7.4.2 Growth 468
References 474

Chapter 8. Physiological Integrations and Energy Partitioning
Raymond J. Thompson and Bruce A. MacDonald

8.1 Introduction 493

8.2 Energy balance, physiological integrations and the partitioning of
energy between growth and reproduction 493

8.2.1 Energy budgets 493

8.2.2 Scope for growth 494

8.2.3 Growth efficiency and turnover ratio 494

8.2.4 Growth of shell and somatic tissue 495

8.2.5 Reproductive effort 496

8.3 Changes related to age of individuals 498

8.3.1 Scope for growth 498

8.3.2 Growth efficiency and turnover ratio 498

8.3.3 Somatic growth and maximum size 499

8.3.4 Somatic production and reproductive output 500

8.3.5 Reproductive effort 502

8.4 Environmental influence on production 504

8.4.1 Scope for growth and growth efficiency 504

8.4.2 Growth of shell and somatic tissue 506

8.4.3 Reproductive effort 509

8.5 Reproductive value and cost 511

8.5.1 Residual reproductive value 511

8.5.2 Reproductive cost 512

8.6 Population production 514
Acknowledgments 516
References 516

Chapter 9. Nutrition in Pectinids
Ana Farías and Iker Uriarte

9.1 Introduction 521

9.2 Pectinid feeding 521

9.3 Nutritional requirements during reproductive conditioning 528

9.4 Nutritional requirements of the larvae 529

9.5 Post-metamorphic nutritional requirements 533

9.6 Summary 536
References 536

Chapter 10. Genetics
Andy Beaumont

10.1 Introduction 543

10.2 Chromosomal genetics and ploidy manipulation 544

10.2.1 Chromosome numbers and karyotypes 544

10.2.2 Ploidy manipulation 548

10.3 Genetic markers and population genetics 550

10.3.1 Genetic markers 550

10.3.1.1 Allozymes 551

10.3.1.2 DNA methods 551

10.3.1.3 DNA based markers 552

10.3.1.3.1 Restriction fragment length polymorphism (RFLP) 552

10.3.1.3.2 Random amplified polymorhic DNA (RAPDs) 552

10.3.1.3.3 Microsatellites 553

10.3.1.3.4 Amplified fragment length polymorphism (AFLP) 553

10.3.1.4 Mitochondrial DNA in scallops 554

10.3.1.5 Types of data produced by different markers 555

10.3.1.6 Analysis of data 555

10.3.2 Overall genetic variation 557

10.3.3 Genetic differentiation of populations 559

10.3.3.1 Placopecten magellanicus 559

10.3.3.2 Patinopecten (= Mizuhopecten) yessoensis 561

10.3.3.3 Pecten maximus 562

10.3.3.4 Aequipecten opercularis 563

10.3.3.5 Chlamys islandica 564

10.3.3.6 Chlamys farreri 565

10.3.3.7 Mimachlamys varia 565

10.3.3.8 Euvola (Pecten) ziczac 565

10.3.3.9 Argopecten gibbus 565

10.3.3.10 The Argopecten irradians species complex 566

10.3.4 Genetic differences at species level and above 567

10.3.5 Heterozygote deficiency 568

10.3.6 Heterozygosity and growth 571

10.4 Quantitative genetics 575

10.4.1 Heritability and artificial selection 575

10.4.2 Inbreeding and hybridisation 577

10.5 Genome mapping and gene sequences 579

10.6 Conclusion 580
Acknowledgments 581
References 581

Chapter 11. Diseases and Parasites of Scallops
Sharon E. McGladdery, Susan M. Bower and Rodman G. Getchell

11.1 Introduction 595

11.2 Microbial diseases 595

11.2.1 Viruses 595

11.2.2 Prokaryota 605

11.2.2.1 Vibrionaceae 605

11.2.2.2 Intracellular prokaryotes (Rickettsiales; Chlamydiales
and Mycoplasma) 607

11.2.2.3 Other bacterial pathogens of scallops 610

11.2.2.4 Bacterial management under hatchery conditions 611

11.3 Mycota 612

11.4 Protista 614

11.4.1 Sarcomastigophorea (Amoebae and Flagellates) 614

11.4.2 Labyrinthomorpha (Thraustochytrids and Labythinuloids) 615

11.4.3 Apicomplexa 615

11.4.3.1 Perkinsorida 615

11.4.3.2 Eucoccidiia 617

11.4.3.3 Eugregarinida 618

11.4.4 Microspora 619

11.4.5 Ascetospora 620

11.4.5.1 Marteiliida 621

11.4.5.2 Balanosporida 621

11.4.6 Ciliates 621

11.5 Platyhelminths 623

11.5.1 Trematodes 623

11.5.2 Cestodes 626

11.5.3 Turbellaria 626

11.5.4 Nematodes 628

11.6 Polychaetes 629

11.7 Crustacea 632

11.7.1 Pinnotheriidae 632

11.7.2 Copepodidae 632

11.8 Gastropods 633

11.9 Algae 635

11.10 Foraminiferans 636

11.11 Porifera 636

11.12 Cnidaria 637

11.13 Non-Infectious diseases 638

11.14 Summary 638
Acknowledgments 639
References 639

Chapter 12. Scallop Ecology: Distributions and Behaviour
Andrew R. Brand

12.1 Introduction 651

12.2 Geographical distribution 653

12.2.1 Geographical distribution of commercially important species 653

12.2.1.1 North Atlantic species 653

12.2.1.1.1 Pecten maximus and P. jacobaeus 653

12.2.1.1.2 Aequipecten opercularis 655

12.2.1.1.3 Mimachlamys varia 655

12.2.1.1.4 Placopecten magellanicus 655

12.2.1.1.5 Argopecten irradians 657

12.2.1.1.6 Argopecten gibbus 659

12.2.1.2 Sub-arctic species 659

12.2.1.2.1 Chlamys islandica 659

12.2.1.3 North Pacific species 661

12.2.1.3.1 Patinopecten caurinus 661

12.2.1.3.2 Mizuhopecten yessoensis 661

12.2.1.3.3 Pecten albicans, Mimachlamys nobilis and
Chlamys farreri 662

12.2.1.4 Southern hemisphere species 662

12.2.1.4.1 Pecten fumatus 662

12.2.1.4.2 Mimachlamys asperrima and Equichlamys bifrons 663

12.2.1.4.3 Amusium balloti and A. pleuronectes 664

12.2.1.4.4 Pecten novaezelandiae 664

12.2.1.4.5 Argopecten purpuratus and A. ventricosus 665

12.2.1.4.6 Aequipecten tehuelchus 665

12.2.1.4.7 Zygochlamys patagonica 665

12.2.2 Factors affecting geographical distribution 666

12.3 Local distribution 669

12.3.1 Spatial distribution 669

12.3.2 Year-class separation 676

12.3.3 Factors affecting local distribution 677

12.3.3.1 Depth 678

12.3.3.2 Substrate type 678

12.3.3.3 Currents 679

12.3.3.4 Turbidity 680

12.3.3.5 Salinity 681

12.3.3.6 Competitors and predators 684

12.4 Scallop behaviour - aspects affecting distribution 689

12.4.1 Byssus attachment 689

12.4.2 Recessing 692

12.4.3 Orientation 695

12.4.4 Swimming 698

12.4.5 Reactions to predators 707

12.4.6 Dispersal and migrations 709

12.5 Further study 712
Acknowledgments 713
References 713

Chapter 13. Scallops and Marine Contaminants
Peter J. Cranford

13.1 Introduction 745

13.2 Trace metals 746

13.3 Trace organics 748

13.4 Offshore oil and gas operations 750

13.5 Scallops as sentinel organisms 754
Acknowledgments 759
References 759

Chapter 14. Dynamics, Assessment and Management of Exploited Natural Populations
J.M. (Lobo) Orensanz, Ana M. Parma, Teresa Turk and Juan Valero

14.1 Introduction 765

14.2 Spatial scales 765

14.3 Population structure and dynamics 769

14.3.1 Aggregate stocks 769

14.3.1.1 Patterns of fluctuation 769

14.3.1.2 Climatic forcing 771

14.3.1.3 Correlation between consecutive life-history stages 773

14.3.1.4 The relation between aggregate stock and settlement/
recruitment 774

14.3.2 Macroscale 777

14.3.2.1 Patterns of connectivity: metapopulation structure 777

14.3.2.2 Metapopulation structure and geographic genetic
differentiation 780

14.3.2.3 Larval retention/dispersal: mechanisms and oceanographic
scenarios 782

14.3.2.4 Metapopulation models 784

14.3.3 Mesoscale 785

14.3.3.1 Persistence, extinction and resurgence of subpopulations 785

14.3.3.2 Relation between abundance and area occupied by a
subpopulation 785

14.3.4 Microscale 786

14.3.4.1 Density, neighbourhoods and concentration 786

14.3.4.2 Concentration profiles 788

14.3.4.3 Density- vs. concentration-dependence; compensation vs.
depensation 789

14.4 The fishing process 795

14.4.1 Types of fishing gear used in scallop fisheries 796

14.4.2 Effort and fishing mortality 797

14.4.2.1 Effort data 797

14.4.2.2 Effort units 797

14.4.2.3 Components of fishing mortality under homogeneity
assumptions 798

14.4.2.4 Relative fishing power and standardisation of effort 799

14.4.3 Spatial patterns of effort allocation 799

14.4.4 The depletion process 803

14.4.5 Vulnerability and selectivity 805

14.4.6 Gear performance 806

14.4.7 Incidental fishing mortality and sub-lethal damage 808

14.5 Assessment 811

14.5.1 Macroscale 811

14.5.2 Mesoscale 812

14.5.2.1 Estimation of aggregated abundance: methods based on
sampling 812

14.5.2.2 Mapping 815

14.5.2.3 Methods based on modelling the depletion process 816

14.5.3 Microscale 819

14.5.3.1 Small-scale spatial data 819

14.5.3.2 Spatial pattern 820

14.5.3.3 Scaling problems 821

14.6 Management 823

14.6.1 Types of scallop fisheries 823

14.6.2 Overfishing 824

14.6.2.1 Growth overfishing: yield-per-recruit (Y/R) analysis 824

14.6.2.2 Recruitment overfishing 826

14.6.2.3 A special case: recruitment fisheries 829

14.6.3 Sustainability 830

14.6.3.1 The precautionary approach to fisheries management
(PAFM) and risk assessment (RA) 830

14.6.3.2 Marine protected areas (MPAs) 832

14.6.3.3 Ecosystem management concerns 834

14.6.3.4 Strategic structures: use and property rights 835

14.6.3.5 Diversification of the resource-base 837

14.6.4 Tactics 838

14.6.4.1 Size limits 838

14.6.4.2 Quota regulations 840

14.6.4.3 Escapement regulations 840

14.6.4.4 Direct effort regulations 840

14.6.4.5 Rotation 841

14.6.4.6 Direct interventions to enhance productivity 842

14.6.4.7 Experimental management 844
Acknowledgments 844
References 844
Endnotes 867

Chapter 15. Fisheries Sea Scallop, Placopecten magellanicus
K.S. Naidu and G. Robert

15.1 Introduction 869

15.2 Distribution 871

15.3 History of fishery 874

15.4 Population biology 878

15.5 Growth and yield per recruit 880

15.6 Gear and boats 883

15.7 Exploitation and resource management 886

15.8 Outlook 895
References 897

Chapter 16. Sea Scallop Aquaculture in the Northwest Atlantic
G. Jay Parsons and Shawn M. C. Robinson

16.1 Introduction 907

16.2 History of sea scallop culture 908

16.3 Hatchery spat production 909

16.3.1 Broodstock conditioning 909

16.3.2 Spawning methods 910

16.3.3 Egg and larval development 910

16.3.4 Settlement and metamorphosis 911

16.3.5 Spat husbandry 911

16.4 Wild spat collection 912

16.4.1 Spat collection techniques 912

16.4.2 Settlement intensity 913

16.4.3 Maximising spat collection 914

16.4.4 Spat growth 916

16.4.5 Spat sorting 916

16.5 Intermediate culture 916

16.5.1 Growth 917

16.5.2 Stocking density 917

16.5.3 Depth 919

16.5.4 Season 919

16.5.5 Location 919

16.5.6 Survival 920

16.5.7 Gear selection 920

16.5.8 Strategies for intermediate culture 921

16.6 Final grow-out 922

16.6.1 Suspension culture 922

16.6.2 Bottom culture 922

16.6.3 Growth 923

16.7 Scheduling and strategies 924

16.8 Site selection 924

16.8.1 Environmental criteria 925

16.8.1.1 Temperature and salinity regime 925

16.8.1.2 Ice coverage 925

16.8.1.3 Wind 925

16.8.1.4 Bottom type 925

16.8.1.5 Turbidity 926

16.8.2 Biological constraints 926

16.8.2.1 Predators 926

16.8.2.2 Fouling organisms 926

16.8.2.3 Phycotoxins 926

16.8.2.4 Diseases and parasites 927

16.9 Products and marketing 928

16.10 Economics 928

16.11 Social issues 929

16.12 Future prospects 930
Acknowledgments 930
References 930

Chapter 17. Bay Scallop and Calico Scallop Fisheries, Culture and Enhancement in Eastern North America
Norman J. Blake and Sandra E. Shumway

17.1 Introduction 945

17.2 Fisheries 945

17.2.1 Bay scallop, Argopecten irradians 945

17.2.1.1 Distribution 945

17.2.1.2 Biology 948

17.2.1.3 Fishery 950

17.2.2 Calico scallop, Argopecten gibbus 952

17.2.2.1 Distribution 952

17.2.2.2 Biology 953

17.2.2.3 Fishery 955

17.3 Aquaculture and enhancement 958

17.4 Future 961
References 961

Chapter 18. Scallops of the West Coast of North America
Raymond B. Lauzier and Neil F. Bourne

18.1 Introduction 965

18.2 Fisheries 967

18.2.1 Weathervane scallop 967

18.2.1.1 Biology 967

18.2.1.2 Fishery 971

18.2.1.3 Gear 973

18.2.1.4 Management 973

18.2.2 Rock scallop 973

18.2.2.1 Biology 974

18.2.2.2 Fishery 975

18.2.3 Pink and spiny scallops 975

18.2.3.1 Biology 975

18.2.3.2 Fisheries 977

18.2.3.3 Management 979

18.3 Aquaculture 979

18.3.1 Pink and spiny scallops 980

18.3.2 Weathervane scallop 980

18.3.3 Rock scallops 981

18.3.4 Japanese weathervane scallop 982

18.3.5 Pacific calico scallop 983

18.4 Future 983
Acknowledgments 984
References 984

Chapter 19. The European Scallop Fisheries for Pecten maximus, Aequipecten opercularis and Mimachlamys varia
Andrew R. Brand

19.1 Introduction 991

19.2 Biology and ecology 993

19.2.1 The great scallop, Pecten maximus (L.) 993

19.2.2 The queen scallop, Aequipecten opercularis (L.) 1006

19.2.3 The black or variegated scallop, Mimachlamys varia (L.) 1011

19.3 Fisheries 1014

19.3.1 Isle of Man 1017

19.3.2 Scotland 1023

19.3.3 England and Wales 1025

19.3.4 Northern Ireland 1027

19.3.5 Republic of Ireland 1027

19.3.6 France 1029

19.3.7 Spain 1031

19.4 The future 1031
Acknowledgments 1035
References 1035

Chapter 20. European Aquaculture
Mark Norman, Guillermo Román and Øivind Strand

20.1 Introduction 1059

20.2 Pecten maximus 1060

20.3 Aequipecten opercularis 1062

20.4 Chlamys varia 1063

20.5 Summary 1063
References 1063

Chapter 21. Scandinavia
Øivind Strand and G. Jay Parsons

21.1 Introduction 1067

21.2 Chlamys islandica 1067

21.2.1 Biology 1067

21.2.2 Fishery 1073

21.2.3 Aquaculture 1076

21.3 Pecten maximus 1077

21.3.1 Biology 1077

21.3.2 Fishery 1078

21.3.3 Aquaculture 1079

21.4 Aequipecten opercularis 1085
Acknowledgments 1086
References 1086

Chapter 22. Japan
Yoshinobu Kosaka and Hiroshi Ito

22.1 Introduction 1093

22.2 Patinopecten (Mizuhopecten) yessoensis 1099

22.2.1 Biology 1099

22.2.2 Fishery 1104

22.2.3 Culture 1111

22.2.4 Seed production 1111

22.2.5 Spat collection 1111

22.2.6 Intermediate culture 1116

22.2.7 Hanging culture 1116

22.2.8 Fouling 1118

22.2.9 Sowing culture 1120

22.2.10 Enemy clearance 1120

22.2.11 Seed sowing 1120

22.2.12 Care 1121

22.2.13 Recapture 1121

22.2.14 Value 1124

22.2.15 Processing 1127

22.2.16 Occurrence of shellfish poisoning 1130

22.3 Future 1130

22.3.1 Technology 1130

22.3.2 Control of culture 1133

22.3.3 Cost-reduction 1133

22.3.4 Concept 1133

22.4 Pecten albicans 1134

22.4.1 Biology 1134

22.4.2 Fishery 1134

22.4.3 Culture 1135

22.5 Chlamys (Mimachlamys) nobilis 1135

22.5.1 Biology 1135

22.5.2 Culture 1135
Acknowledgments 1136
References 1136

Chapter 23. Scallop Culture in China
Ximing Guo and Yousheng Luo

23.1 Introduction 1141

23.2 Zhikong scallop 1146

23.2.1 Shell morphology 1146

23.2.2 Distribution 1146

23.2.3 Growth 1147

23.2.4 Reproduction 1149

23.3 Other scallop species 1151

23.3.1 Huagui scallop 1151

23.3.2 Non-native species 1152

23.4 Fishery 1153

23.5 Aquaculture 1154

23.5.1 Species and history 1154

23.5.2 Collection of natural seed for Zhikong scallop 1155

23.5.3 Hatchery production of bay scallop 1155

23.5.4 Grow-out 1158

23.6 Harvest, processing and marketing 1160
Acknowledgments 1160
References 1161

Chapter 24. Scallops Fisheries and Aquaculture of Northwestern Pacific, Russian Federation
Victor V. Ivin, Vasily Z. Kalashnikov, Sergey I. Maslennikov and Vitaly G. Tarasov

24.1 Introduction 1163

24.2 Taxonomic status 1163

24.3 Biology and ecology 1164

24.3.1 Chlamys albida 1164

24.3.2 Chlamys asiatica 1166

24.3.3 Chlamys behringiana 1167

24.3.4 Chlamys chosenica 1168

24.3.5 Chlamys farreri 1170

24.3.6 Chlamys swifti 1172

24.3.7 Delectopecten randolphi 1174

24.3.8 Mizuhopecten yessoensis 1174

24.3.8.1 Total populations and biomass 1175

24.3.8.2 Distribution in Primorye 1176

24.3.8.3 Distribution over depths 1176

24.3.8.4 Age structure of scallop settlements 1176

24.3.8.5 Scallop growth 1177

24.3.8.6 Sex structure of settlements 1178

24.3.8.7 Replenishment 1181

24.3.8.8 Spawning 1181

24.3.8.9 Larvae morphology 1182

24.3.8.10 Development in plankton 1182

24.3.8.11 Migration behaviour 1184

24.3.8.12 Risk factors 1184

24.3.8.12.1 Abiotic factors 1184

24.3.8.12.2 Storms 1184

24.3.8.12.3 Predators 1185

24.3.8.12.4 Parasites 1187

24.3.8.12.5 Bacterial contamination 1189

24.3.8.12.6 Epibionts 1189

24.4 Fishing and aquaculture 1189

24.4.1 Fishing 1189

24.4.1.1 History 1189

24.4.1.2 Fishing gear 1191

24.4.1.3 Yesso scallop landings 1191

24.4.1.3.1 Primorsky territory 1191

24.4.1.3.2 Sakhalin-Kurile region 1192

24.4.1.4 Yesso scallop commercial stock 1192

24.4.1.4.1 Primorsky territory 1192

24.4.1.4.2 Sakhalin-Kurile region 1192

24.4.1.4.2.1 Aniva Bay 1194

24.4.1.4.2.2 Terpenie Bay 1194

24.4.1.4.2.3 Kuriles 1194

24.4.1.5 Commercial Chlamys scallops 1194

24.4.1.5.1 Primorye 1194

24.4.1.5.2 Kurile Islands 1195

24.4.1.5.3 In Bering Sea 1196

24.4.1.6 Other Chlamys species 1196

24.4.1.6.1 Chlamys farreri 1196

24.4.1.6.2 Chlamys swifti 1196

24.4.2 Aquaculture 1196

24.4.2.1 History 1196

24.4.2.2 Present situation 1197

24.4.2.3 Marketing 1198

24.4.2.4 The culture methods 1198

24.4.2.4.1 Spat collection 1198

24.4.2.4.2 Intermediate culture 1199

24.4.2.4.3 Transport of scallop seed 1199

24.4.2.4.4 Sowing or on-bottom culture 1200

24.4.2.4.5 Hanging or off-bottom culture 1201

24.4.2.4.6 Obstacles to mariculture development 1201

24.4.2.4.7 Ecological constraints associated with cultivation 1202

24.4.2.4.7.1 Predation 1202

24.4.2.4.7.2 Epibionts 1202

24.4.2.4.7.3 Biofouling of cultivation structures 1203

24.4.2.4.8 Effect of scallop mariculture on coastal ecosystems 1204

24.4.2.5 Future prospects 1211
Acknowledgments 1213
References 1213

Chapter 25. Scallop Aquaculture and Fisheries in Brazil
Guilherme S. Rupp and G. Jay Parsons

25.1 Introduction 1225

25.2 Nodipecten nodosus – biology, aquaculture and fisheries 1227

25.2.1 Taxonomy and distribution 1227

25.2.2 Ecology 1228

25.2.3 Reproduction 1231

25.2.4 Aquaculture 1232

25.2.4.1 Status 1232

25.2.4.2 Culture technology 1233

25.2.4.2.1 Wild seed collection 1233

25.2.4.2.2 Hatchery production 1234

25.2.4.2.3 Growout 1236

25.2.4.2.4 Constraints 1239

25.2.5 Fisheries and marketing 1239

25.2.6 Future prospects 1240

25.3 Euvola (Pecten) ziczac – biology, aquaculture and fisheries 1240

25.3.1 Taxonomy and distribution 1240

25.3.2 Ecology 1241

25.3.3 Reproduction 1241

25.3.4 Aquaculture 1242

25.3.4.1 Status 1242

25.3.4.2.Culture technology 1242

25.3.4.2.1 Wild seed collection 1242

25.3.4.2.2 Hatchery production 1242

25.3.4.2.3 Growout 1243

25.3.4.2.4 Constraints 1243

25.3.5 Fisheries 1243

25.3.6 Future prospects 1244
Acknowledgments 1244
References 1245

Chapter 26. Argentina
Néstor F. Ciocco, Mario L. Lasta, Maite Narvarte, Claudia Bremec, Eugenia Bogazzi, Juan Valero and J.M. (Lobo) Orensanz

26.1 Introduction 1251

26.2 The Tehuelche scallop, Aequipecten tehuelchus 1251

26.2.1 Biology 1251

26.2.2 Population dynamics 1257

26.2.3 Fisheries 1258

26.2.3.1 The inshore dredge fishery of San Matías Gulf 1258

26.2.3.2 The commercial diving fishery of San José Gulf 1261

26.2.4 Aquaculture 1263

26.2.4.1 Spat collection 1263

26.2.4.1.1 Vertical distribution of settlement 1263

26.2.4.1.2 Temporal variation in settlement 1263

26.2.4.1.3 Type of collector 1263

26.2.4.1.4 Location 1265

26.2.4.1.5 Influence of depth, type of collector and location
on the average size of the ‘seed’ 1265

26.2.4.2 Growth in suspended structures of spat obtained from
collectors 1266

26.2.4.3 Larval culture in the laboratory 1268

26.2.4.4 Prospects and problems 1268

26.3 The Patagonian scallop, Zygochlamys patagonica 1269

26.3.1 Biology and ecology 1269

26.3.2 Population dynamics and stock assessment 1273

26.3.3 The fishery 1275

26.3.3.1 Exploratory surveys and experimental fishing programs 1275

26.3.3.2 Management 1276

26.3.3.3 Development of the fishery 1277

26.3.3.4 Observers program 1280

26.3.3.5 Ecological effects of fishing 1281

26.3.3.6 Marketing 1283
Acknowledgments 1283
References 1283

Chapter 27. Scallop Fishery and Aquaculture in Chile
Elisabeth von Brand, German E. Merino, Alejandro Abarca and Wolfgang Stotz

27.1 Introduction 1293

27.2 Species description 1293

27.2.1 Argopecten purpuratus (Lamarck 1819) the northern scallop 1295

27.2.2 Zygochlamys patagonica (King and Broderip 1831) the
southern scallop 1295

27.2.3 Chlamys vitrea (King & Broderip, 1831) the southern scallop…. 1296

27.3 Fisheries 1296

27.3.1 Argopecten purpuratus, the northern scallop 1296

27.3.2 Chlamys vitrea and Zygochlamys patagonica, the southern
scallop 1298

27.4 How scallop aquaculture started in Chile 1299

27.5 Aquaculture production 1301

27.5.1 Production stages 1303

27.5.1.1 Hatchery broodstock conditioning and spawning 1303

27.5.1.2 Larval stage 1303

27.5.1.3 Settlement, metamorphosis and postlarval stage 1304

27.5.1.4 Seed supply 1305

27.5.1.5 Ongrowing stage 1305

27.5.2 Constraints and opportunities for northern scallop hatcheries 1306

27.5.3 Interaction between natural beds and aquaculture 1307

27.5.4 Recovery of a natural bed of Argopecten purpuratus 1308

27.5.5 Final overview and projections of Chilean scallop farming 1308
Acknowledgments 1311
References 1311

Chapter 28. Venezuela
César J. Lodeiros, Luis Freites, Maximiano Nuñez, Anibal Vélez and John H. Himmelman

28.1 Introduction 1315

28.2 Distribution, habitat and reproduction 1315

28.2.1 Euvola (Pecten) ziczac 1315

28.2.2 Argopecten nucleus 1319

28.2.3 Nodipecten (Lyropecten) nodosus 1319

28.2.4 Amusium papyraceum and Amusium laurenti 1320

28.3 Fisheries 1320

28.4 Aquaculture 1322

28.4.1 Euvola ziczac 1322

28.4.1.1 Gonad maturation and spawning 1322

28.4.1.2 Spat production 1322

28.4.1.3 Grow-out 1323

28.4.2 Nodipecten nodosus 1326

28.4.2.1 Gonad maturation and spawning 1326

28.4.2.2 Spat production 1326

28.4.2.3 Grow-out 1326

28.4.3 Argopecten nucleus 1329

28.5 Perspectives for culture 1330
References 1332

Chapter 29. Mexico
Esteban Fernando Félix-Pico

29.1 Fishery 1337

29.1.1 Introduction 1337

29.1.2 Species and distribution of scallops 1338

29.1.3 History of the fishery 1339

29.1.4 Status of the resource 1340

29.1.4.1 Reproductive biology 1340

29.1.4.1.1 Size at maturity 1340

29.1.4.1.2 Fecundity 1341

29.1.4.1.3 Maturity and spawning 1341

29.1.4.2 Population biology 1344

29.1.4.3 Abundance and density 1345

29.2 Exploitation of the resource 1347

29.2.1 Fishing areas 1347

29.2.2 Fishing seasons 1348

29.2.3 Fishing operations and results 1349

29.2.4 The market value-landings data from 1970 to 2000 1349

29.3 Constraints 1350

29.3.1 Pollution 1350

29.3.2 Biological constraints 1352

29.3.3 Predation 1353

29.3.4 Resource management constraints 1354

29.4 Harvesting and transporting 1354

29.4.1 Methods 1354

29.4.2 Marketing 1355

29.5 Expected future 1356

29.6 Aquaculture 1357

29.6.1 Introduction 1357

29.6.2 Species 1358

29.6.3 History 1358

29.7 Hatchery techniques 1359

29.7.1 Conditioning 1359

29.7.2 Induction of spawning and fertilisation 1360

29.7.3 Larval culture and metamorphosis 1361

29.7.4 Genetic enhancement 1366

29.7.4.1 Selection 1366

29.7.4.2 Crossbreeding 1366

29.7.4.3 Triploidy 1367

29.7.5 Antibiotics 1368

29.8 Natural spat 1368

29.8.1 Methods employed 1368

29.8.2 Spat collection 1369

29.9 Growth 1372

29.9.1 Suspended cultures 1372

29.9.2 Bottom cultures 1375

29.9.3 Factors influencing growth 1376

29.10 Mortality 1377

29.10.1 Suspended culture 1377

29.10.2 Bottom culture 1378

29.11 Constraints 1379

29.12 Future prospects 1379
References 1380

Chapter 30. Scallop Fisheries, Mariculture and Enhancement in Australia
Mike Dredge

30.1 Introduction 1391

30.2 History of the fisheries 1392

30.3 Biology of target species 1395

30.3.1 Pecten fumatus 1395

30.3.1.1 Distribution and life cycle 1395

30.3.1.2 Mortality 1396

30.3.1.3 Monitoring, abundance and population dynamics 1397

30.3.2 Amusium balloti 1398

30.3.2.1 Distribution and life cycle 1398

30.3.2.2 Mortality 1398

30.3.2.3 Monitoring, abundance and population dynamics 1399

30.4 Fisheries and their management 1400

30.4.1 Pecten fumatus 1400

30.4.1.1 Regulation 1400

30.4.1.2 Environmental management and sustainability issues 1402

30.4.2 Amusium balloti 1403

30.4.2.1 Regulation 1403

30.4.2.2 Environmental management and sustainability issues 1404

30.5 Culture of scallops in Australia 1405

30.5.1 Pecten fumatus 1405

30.5.1.1 Tasmania 1405

30.5.1.1.1 Spat production 1405

30.5.1.1.2 Culture operations 1405

30.5.1.2 Victoria 1406

30.5.1.3 New South Wales 1406

30.5.1.4 South Australia 1407

30.5.2 Amusium balloti 1407

30.6 Summary 1407
Acknowledgments 1408
References 1408

Chapter 31. New Zealand
Islay D. Marsden and Michael F. Bull

31.1 Introduction 1413

31.2 Biology 1413

31.2.1 Morphology 1413

31.2.2 Distribution 1413

31.2.3 Life cycle 1414

31.2.4 Growth rates 1415

31.2.5 Mortality 1415