Scallops: Biology, Ecology and Aquaculture: Volume 40

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 Farias 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 Roman and Oivind 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
Oivind 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
Nestor 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 Matias Gulf 1258
26.2.3.2 The commercial diving fishery of San Jose 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
Cesar J. Lodeiros, Luis Freites, Maximiano Nunez, Anibal Velez 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 Felix-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