Marine Bivalve Molluscs Marine Bivalve Molluscs Second Edition Elizabeth Gosling This edition first published 2015 © 2015 by John Wiley & Sons, Ltd First edition published 2003 © Fishing News Books, a division of Blackwell Publishing Registered Office John Wiley & Sons, Ltd, The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK Editorial Offices 9600 Garsington Road, Oxford, OX4 2DQ, UK The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK 111 River Street, Hoboken, NJ 07030‐5774, USA For details of our global editorial offices, for customer services and for information about how to apply for permission to reuse the copyright material in this book please see our website at www.wiley.com/wiley‐blackwell. The right of the author to be identified as the author of this work has been asserted in accordance with the UK Copyright, Designs and Patents Act 1988. All rights reserved. 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Cover image: (Scallop background) iStockphoto © DHuss (Oyster opening) iStockphoto © Riorita (Rough cockle sea shell out of its armor) iStockphoto © xbrchx (Other images) Courtesy of the author Set in 10/12pt Times by SPi Global, Pondicherry, India 1 2015 Contents Preface viii Acknowledgements x 1 Phylogeny and evolution of bivalve molluscs 1 Phylogeny of the Mollusca 1 Phylogeny and evolution of Bivalvia 3 Evolution and adaptive radiation in Bivalvia 7 Notes 10 References 10 2 Morphology of bivalves 12 Introduction 12 Shell 12 Mantle 21 Gills 24 The foot 27 Labial palps and alimentary canal 31 Gonads 34 Heart and haemolymph vessels 34 Excretory organs 35 Nerves and sensory receptors 36 References 38 3 Ecology of bivalves 44 Introduction 44 Global and local distribution patterns 44 Factors affecting geographic distribution 52 Factors affecting local distribution 55 Climate change 78 Note 84 References 84 4 How bivalves feed 99 Introduction 99 Filtration rate 100 Particle processing on the gills, labial palps and in the stomach 110 The alimentary canal and digestive process 128 Absorption efficiency 139 Effects of bivalve suspension‐feeders on the ecosystem 141 Notes 144 References 144 vi Contents 5 Reproduction, settlement and recruitment 157 Introduction 157 Sexuality 157 Gametogenesis 159 Reproductive cycles 159 Factors controlling reproduction 165 Annual storage cycle 170 Reproductive effort and fecundity 171 Deleterious effects on the reproductive cycle 175 Fertilization 175 Larval development 177 Factors affecting larval growth 178 Larval dispersal and population connectivity 181 Larval identification and abundance estimation 184 Settlement, metamorphosis and post‐larval dispersal 185 References 189 6 Growth 203 Introduction 203 Methods of measuring absolute growth 203 Allometric growth 213 Scope for growth and dynamic energy budget models 216 Factors affecting growth 219 Note 232 References 232 7 Circulation, respiration, excretion and osmoregulation 243 Circulation 243 Respiration 246 Excretion and osmoregulation 260 Note 264 References 264 8 Fisheries and management of natural populations 270 Introduction 270 Population dynamics 271 Fisheries assessment and management 282 Scallop fisheries 287 Oyster fisheries 301 Mussel fisheries 309 Clam fisheries 314 References 319 9 Bivalve culture 325 Introduction 325 Fundamentals of bivalve culture 326 Mussel culture 340 Oyster culture 348 Scallop culture 355 Clam culture 362 Contents vii Bivalve culture and the environment 369 Ecosystem approach to bivalve culture 371 Note 374 References 375 10 Genetics in aquaculture 383 Introduction 383 Quantitative genetics 384 Selective breeding 386 Protein and DNA markers 394 Bivalve genomics 403 Chromosomal genetics and ploidy manipulation 408 Transgenics 417 Notes 419 References 419 11 Diseases and parasites 429 Introduction 429 Viruses 429 Bacteria 431 Fungi 434 Protistans 434 Porifera 447 Helminths 448 Annelids 451 Crustaceans 452 Neoplasia 454 Defence mechanisms 455 Perspectives 463 References 464 12 Public health 478 Introduction 478 Bacterial infections 478 Viral infections 480 Biotoxins 481 Industrial pollutants 488 Controls on the production and processing of bivalves 490 Elimination of pathogens and other hazards 495 Monitoring and quality control 499 HAACP system 502 Notes 508 References 508 Subject index 514 Species index 520 Preface The phylum Mollusca is one of the largest, most diverse and important groups in the animal kingdom. There are at least 50 000 described species, and about 30 000 of these are found in the sea. The class Bivalvia is one of the eight classes of molluscs and comprises animals enclosed in two shell valves. Examples are mussels, oysters, scallops and clams. Although this class contains a relatively small number of species, about 10 000, it elicits substantial interest chiefly because these species are widely consumed by people. In 2011, production of marine bivalves from fisheries and aquaculture was almost 14 million metric tonnes worldwide with more than 90% of this figure coming from aquaculture. Some years ago while teaching a course on bivalve biology to aquaculture students, I realized that although the students could access much relevant information from several texts, a single book covering all aspects of the biology, ecology and culture of bivalve molluscs did not exist. Thus the idea to write such a book was conceived. What started off as a small undertaking very quickly snowballed into a substantial task, which took several years to complete, primarily due to my having to read and digest the wealth of published information. The first edition of Bivalve Molluscs, published in 2003, was very well received by reviewers, did well in terms of sales, and in 2011 Wiley‐Blackwell invited me to prepare a second edition. In the intervening years, there have been many significant advances, along with the emergence of new areas of research, one reason being the range of new techniques being applied in bivalve biology and ecology. For example, DNA genetic markers play an increas- ingly important role in disease diagnosis (Chapter 11), identification and tagging of larvae (Chapters 5 and 10), in detecting different strains of pathogenic bacteria (Chapter 12), and in the completely new area of bivalve genomics (Chapter 10). Also, isotope ratios and fatty acid signatures are being used to identify dietary food sources in bivalves (Chapter 4), while measurements of stable oxygen isotopes in fossil and contemporary bivalve shells are allowing reconstruction of past sea surface temperatures. Trace elemental fingerprinting, based on naturally occurring elements in bivalve shells, is being used to assess larval origins and trajectories, while fluorescence in situ hybridization (FISH) combined with cell sorting can identify and rapidly count and sort larvae of several bivalve species in diverse plankton samples (Chapter 5). Immunosensors – devices that use specific biochemical reactions mediated by antibodies to detect chemical compounds – may soon replace liquid chromatography, which has long been the official method for detection and quantification of algal toxins in shellfish (Chapter 12). Another significant development has been the increasing use of sophisticated mathematical models to describe and predict growth and bioenergetics of individual animals (Chapters 4 and 6), to track larval movement and recruitment (Chapter 5) and to predict geographic range of species in global warming scenarios (Chapter 3). Models are also being employed to simulate disease and forecast outbreaks (Chapters 11 and 12), to describe carrying capac- ity and environmental effects in aquaculture areas (Chapter 9). Also, several model‐based approaches are in use to assess fisheries stocks (Chapter 8). Preface ix Another area that has changed is the management of bivalve fisheries and aquaculture systems, where the emphasis is now more on the use of legislation and control measures, particularly in Europe, North America, Australia and New Zealand (Chapters 8 and 9). This also pertains