Scallops: Biology, Ecology and Aquaculture

Dr. Sandra L. Shumway is known as THE shellfish expert. She has more than 30 years of research experience, and is active on research panels and several international advisory boards. She has been the President-elect of the National Shellfisheries Association for five years, and is Editor-in-Chief of four top journals in fisheries and shellfish. She has been on the Steering Committee and served as Program Chair of the World Aquaculture Society and Fish Culture Section of the American Fisheries. She received the APEX Award for Publication Excellence for Journal of Shellfish Research in 2008. 2009, 2010, and 2011 and won the Bronze Award (2009) and Silver Award (2010) for the Association Trends, Journal of Shellfish Research. 4392 total citations by 2871 documents in Scopus Dr. Jay Parsons has been involved in the aquaculture sector for 30 years and has extensive experience in shellfish aquaculture research and management. Since 2003, Dr. Parsons has been with the Aquaculture Science Branch of Fisheries and Oceans Canada where he is Branch Director responsible for national aquaculture R&D programs and aquaculture research coordination. From 1995-2003 he was a researcher and faculty members at Memorial University (St. John’s, Newfoundland, Canada) where he taught graduate courses in shellfish aquaculture and directed several national and international projects, graduate students and publications on culture, feeding and reproduction in scallops, oysters, mussels, sea urchins and shrimp. He is also a past president of the World Aquaculture Society (WAS) and has served on the WAS Board since 2001. He was also President of the National Shellfisheries Association (NSA) and twice President of the Aquaculture Association of Canada (AAC). And he still maintains an active involvement in these professional societies through the promotion and dissemination of aquaculture science nationally and internationally.

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