LOBSTERS: BIOLOGY, MANAGEMENT, AQUACULTURE AND FISHERIES Second Edition Edited by Bruce F. Phillips Department of Environment & Agriculture Curtin University, Western Australia A John Wiley & Sons, Ltd., Publication This edition fi rst published 2013 © 2006, 2013 by John Wiley & Sons, Ltd Wiley-Blackwell is an imprint of John Wiley & Sons, formed by the merger of Wiley’s global Scientifi c, Technical and Medical business with Blackwell Publishing. Registered offi ce: John Wiley & Sons, Ltd, The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK Editorial offi ces: 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 offi ces, 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. 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Limit of Liability/Disclaimer of Warranty: While the publisher and author(s) have used their best efforts in preparing this book, they make no representations or warranties with respect to the accuracy or completeness of the contents of this book and specifi cally disclaim any implied warranties of merchantability or fi tness for a particular purpose. It is sold on the understanding that the publisher is not engaged in rendering professional services and neither the publisher nor the author shall be liable for damages arising herefrom. If professional advice or other expert assistance is required, the services of a competent professional should be sought. Library of Congress Cataloging-in-Publication Data Lobsters : biology, management, aquaculture and fisheries / edited by Bruce F. Phillips, Department of Environment & Agriculture, Curtin University, Western Australia. pages cm Includes bibliographical references and index. ISBN 978-0-470-67113-9 (hardback : alk. paper) 1. Lobsters. I. Phillips, Bruce F. QL444.M33L62 2013 595.3'84–dc23 2012032890 A catalogue record for this book is available from the British Library. Wiley also publishes its books in a variety of electronic formats. Some content that appears in print may not be available in electronic books. Cover image: iStock/Dmitriy Pochitalin Cover design by Design Deluxe Set in 10/12 pt Times by Toppan Best-set Premedia Limited 1 2013 Contents Contributors xi Preface xiv Chapter 1 Lobsters as Part of Marine Ecosystems – A Review 1 Bruce F. Phillips, Richard A. Wahle and Trevor J. Ward 1.1 Introduction 1 1.2 Species overviews 3 1.2.1 Western rock (spiny) lobster Panulirus cygnus 3 1.2.2 The American lobster Homarus americanus 7 1.3 How far have we come in thinking about lobsters as part of the ecosystem? 11 1.3.1 Panulirus cygnus 11 1.3.2 Homarus americanus 15 1.4 Human role in ecosystem dynamics 18 1.4.1 Institutional structures 18 1.4.2 Direct effects of management 19 1.4.3 Indirect effects – top-down forcing by predator removal 20 1.4.4 Indirect effects – bottom-up forcing by bait subsidies 21 1.4.5 Climate change impacts 21 1.5 Single species to ecosystem management – how far have we come? 22 1.5.1 Panulirus cygnus 23 1.5.2 Homarus americanus 26 1.6 Implications for management and research 27 1.6.1 Top-down, bottom-up ecology 27 1.6.2 Inclusive governance systems 27 1.6.3 Stock rebuilding strategies 28 1.6.4 Environmental drivers of settlement patterns 28 1.6.5 Historical ecosystem structure 29 1.7 Conclusions 29 References 30 Chapter 2 Genetics of Wild and Captive Lobster Populations 36 W. Jason Kennington, Roy Melville-Smith and Oliver Berry 2.1 Introduction 36 2.2 Population structure 37 2.2.1 Chaotic genetic patchiness 39 2.2.2 Post-glaciation demographic expansions 39 2.3 Species identifi cation 40 2.4 Applications of species identifi cation and DNA barcoding to lobster research and management 41 2.4.1 DNA surveillance of product substitution and cross-contamination 41 2.4.2 Identifi cation of wild lobsters 41 iii iv Contents 2.4.3 Charactering lobster diets 42 2.4.4 DNA-based methods typically employed for species identifi cation 43 2.4.5 Polymerase chain reaction–restriction fragment length polymorphism 43 2.4.6 Melt-curve analysis 44 2.4.7 Species-specifi c polymerase chain reaction 44 2.5 Variation in reproductive success 45 2.6 Genetic effects of harvesting 46 2.6.1 Fishing-induced evolution 46 2.6.2 Impacting genetic diversity 50 2.7 Considerations for aquaculture and stock enhancement 50 2.7.1 Aquaculture 50 2.7.2 Stock enhancement 51 2.8 Conclusions and future directions 52 References 54 Chapter 3 Enhancement of Lobster Fisheries to Improve Yield and Value 64 Bridget S. Green, Caleb Gardner and Gro I. van der Meeren 3.1 Introduction 64 3.1.1 What is enhancement? 64 3.1.2 The motivation for enhancement 65 3.2 Monitoring the outcomes of enhancement operations 67 3.2.1 Growth and reproduction 68 3.2.2 Survival/changes to natural mortality 68 3.2.3 Displacement and integration of released lobsters 69 3.2.4 Habitat and predator change 69 3.3 Economic and policy issues 70 3.3.1 Economic feasibility 70 3.3.2 Optimal management of harvests with enhancement 71 3.3.3 Ownership, policy and risk 72 3.4 Case studies 73 3.4.1 Clawed lobster restocking 73 3.4.2 Range extension of American lobster to Europe 74 3.4.3 Improving yield in Southern rock lobster: translocating low quality adults 74 3.5 Conclusions 75 References 75 Chapter 4 The Impact of Climate Change on Exploited Lobster Stocks 84 Nick Caputi, Simon de Lestang, Stuart Frusher and Richard A. Wahle 4.1 Introduction 84 4.2 Case studies 85 4.2.1 Western rock lobster (Panulirus cygnus) 85 4.2.2 Southern rock lobster (Jasus edwardsii) 89 4.2.3 American lobster (Homarus americanus) 92 4.3 Other lobster species 98 4.3.1 Caribbean lobster (Panulirus argus) 98 4.3.2 Southern African west coast rock lobster (Jasus lalandii) 98 Contents v 4.3.3 Hawaiian lobster (Panulirus marginatus) 99 4.3.4 Japanese lobster (Panulirus japonicus) 99 4.3.5 Californian lobster (Panulirus interruptus) 100 4.3.6 Pronghorn spiny lobster (Panulirus pencillatus) 100 4.3.7 European lobster (Homarus gammarus) 100 4.3.8 Norway Lobster (Nephrops norvegicus) 101 4.4 Discussion 101 4.4.1 Climate change effect type 101 4.4.2 Sensitivity to environmental effects 101 4.4.3 Climate change effects 102 4.4.4 Implications for stock assessment 104 4.4.5 Implications for management 104 4.4.6 Risk assessment approach to assess vulnerability 105 4.5 Conclusions and future research 105 References 106 Chapter 5 Systems to Maximize Economic Benefi ts in Lobster Fisheries 113 Caleb Gardner, Sherry Larkin and Juan Carlos Seijo 5.1 Introduction 113 5.2 Economic concepts used in lobster fi shery management 114 5.2.1 Net economic benefi ts 114 5.2.2 Economic targets and performance measures 115 5.2.3 Economic modelling 116 5.3 Lobster fi shery management systems 118 5.3.1 Limited entry 118 5.3.2 Effort limitation 118 5.3.3 Quota management 121 5.3.4 Territorial use rights 123 5.4 Resource sharing and allocation of lobster stocks 124 5.4.1 Recreational and commercial sectors 124 5.4.2 Customary harvests 125 5.4.3 Non-extractive users 126 5.4.4 Dividing catches between nations 126 5.4.5 Managing confl icting objectives 127 5.5 Developing issues in lobster fi shery economics 128 5.5.1 Rights-based management systems 128 5.5.2 Multiple species and ecosystem interactions 128 5.5.3 Marine protected areas 129 5.5.4 Climate change 130 5.6 Conclusions 131 References 132 Chapter 6 Lobster Ecolabelling 139 Trevor J Ward and Bruce F Phillips 6.1 Introduction 139 6.1.1 What is ecolabelling? 142 6.1.2 Global context 146 vi Contents 6.2 Ecolabelling objectives 147 6.2.1 Consumers 147 6.2.2 Retailers and wholesalers 148 6.2.3 Fishing and aquaculture industry 149 6.2.4 Non-government organizations 149 6.2.5 Governments 150 6.2.6 Developing countries 151 6.3 Comparative performance assessment 152 6.3.1 Criteria 152 6.3.2 Assessing performance 155 6.4 Ecolabelled lobsters 158 6.4.1 Panulirus cygnus (Australia) 158 6.4.2 Homarus americanus (Canada) 161 6.4.3 Homarus gammarus (France, UK) 163 6.4.4 Panulirus interruptus (Mexico) 166 6.5 Conclusions 168 6.5.1 Management issues 169 6.5.2 Stock issues 170 6.5.3 Environmental impacts 171 6.5.4 Developing countries 173 6.5.5 Successful certifi cation and ecolabelling of lobsters 174 Appendix 175 Example Aquaculture Criteria and Grades: ecological performance and sustainability stringency 175 References 182 Chapter 7 Essential Habitats for Panulirus Spiny Lobsters 186 Patricia Briones-Fourzán and Enrique Lozano-Álvarez 7.1 Introduction 186 7.2 Essential habitats for ontogenetic shifters 189 7.2.1 Nursery and juvenile habitats 189 7.2.2 Adult and breeding habitats 192 7.2.3 Foraging habitats 193 7.3 Habitat specialists 194 7.4 Coexisting species: interspecifi c trade-offs 196 7.5 Global and local threats to habitats used by Panulirus lobsters 198 7.5.1 Fishing practices 198 7.5.2 Coastal development 199 7.5.3 Climate change and ocean acidifi cation 199 7.5.4 Potential effects of habitat loss 200 7.6 Habitat management 200 7.6.1 Marine protected areas 200 7.6.2 Corollary: marine reserves and invasive species 202 7.6.3 Habitat enhancement 202 7.6.4 Casitas and the controversy over their use 203 7.6.5 Corollary: habitat enhancement and disease transmission 204 7.7 Conclusions 205 References 206 Contents vii Chapter 8 Homarus 221 Richard A. Wahle, Kathleen M. Castro, Oliver Tully and J. Stanley Cobb 8.1 Introduction 221 8.2 Species overview 222 8.3 Life-history overview 224 8.4 Growth and age 225 8.4.1 Components of growth 225 8.4.2 Age determination 227 8.5 Maturation 228 8.5.1 Size at onset of maturity 228 8.5.2 Clutch size and fecundity 230 8.6 Larval and pre-settlement dynamics 230 8.7 Postlarval settlement and benthic recruitment 231 8.7.1 Settlement indices 231 8.7.2 Settler – fi shery recruit relationship 233 8.8 Meta-population structure 233 8.9 Post-settlement mortality sources 235 8.9.1 Predators and habitat-mediated survival bottlenecks 235 8.9.2 Evidence of density-dependence 236 8.9.3 Disease 236 8.10 Homarus in a changing ecosystem 239 8.10.1 Global climate change effects 239 8.10.2 Altered food webs 240 8.11 Harvest and management of wild populations 240 8.11.1 Gear type and methods 240 8.11.2 Commercial exploitation 241 8.11.3 Fishery management 242 8.12 Mariculture and population enhancement 244 8.12.1 Mariculture 244 8.12.2 Stock enhancement 244 8.12.3 Habitat enhancement 245 8.13 Conclusions and future research 246 References 247 Chapter 9 Jasus and Sagmariasus Species 259 Andrew G. Jeffs, Caleb Gardner and Andy Cockcroft 9.1 Species and distribution 259 9.2 Reproduction, life history and growth 261 9.3 Ecology and behaviour 265 9.4 Predators and disease 267 9.5 Population dynamics 267 9.6 Harvest of wild populations and their regulation 268 9.6.1 South Africa and Namibia 269 9.6.2 Australia 270 9.6.3 New Zealand 271 9.6.4 Other Jasus fi sheries 272 9.7 Aquaculture 274 9.8 Marine protected areas 275 viii Contents 9.9 Managing ecosystem effects of fi shing 276 9.10 Conclusions 277 References 277 Chapter 10 Panulirus Species 289 Bruce F. Phillips, Roy Melville-Smith Matthew C. Kay and Armando Vega-Velázquez 10.1 Species and distribution 289 10.2 Life history, growth and reproduction 290 10.3 Predators and diseases 291 10.4 Ecology and behaviour 292 10.5 Population dynamics and regulation 293 10.6 Harvest of wild populations and their regulations 293 10.6.1 Australia and Papua New Guinea 293 10.6.2 Cuba 300 10.6.3 USA (Florida) 302 10.6.4 Brazil 304 10.6.5 Baja California Mexico and California USA 305 10.6.6 India 310 10.6.7 Kenya and Somalia 311 10.6.8 Indonesia 311 10.6.9 Japan 311 10.7 Aquaculture and enhancement 312 10.7.1 Aquaculture 312 10.7.2 Enhancement 314 10.8 Management and conservation 314 10.9 Conclusions 315 References 315 Chapter 11 Palinurus Species 326 Johan C. Groeneveld, Raquel Goñi and David Díaz 11.1 Introduction 327 11.2 Species and distribution 327 11.2.1 Present-day distributions 327 11.2.2 Effects of climate change on past distribution patterns 327 11.3 Biology and ecology 329 11.3.1 Reproductive biology 329 11.3.2 Larval dispersal patterns and recruitment of pueruli 332 11.3.3 Moulting and growth 336 11.3.4 Population structure, size composition and sex ratios 338 11.3.5 Migrations and movement 339 11.3.6 Predators, defence and natural mortality 340 11.3.7 Diet 341 11.4 Molecular phylogeny and evolution 341 11.5 Harvest of wild populations 342 11.5.1 Palinurus elephas 342 11.5.2 Palinurus mauritanicus 343 11.5.3 Palinurus charlestoni 344 Contents ix 11.5.4 Palinurus gilchristi 344 11.5.5 Palinurus delagoae 344 11.5.6 Palinurus barbarae 346 11.6 Protection and enhancement 346 11.6.1 Marine Protected Areas 346 11.6.2 Enhancement 347 11.7 Monitoring and management 347 11.7.1 Northeast Atlantic and Mediterranean fi sheries 347 11.7.2 Southwest Indian Ocean fi sheries 348 11.8 Conclusions 349 References 349 Chapter 12 Nephrops Species 357 Mike Bell, Ian Tuck and Helen Dobby 12.1 Introduction 357 12.2 Species and distribution 357 12.3 Life history 361 12.3.1 Growth 361 12.3.2 Reproduction 364 12.3.3 Food and feeding 369 12.3.4 Activity patterns 371 12.4 Population dynamics 375 12.4.1 Population structure 375 12.4.2 Mortality 377 12.4.3 Population regulation 382 12.5 Harvest of wild populations 383 12.5.1 Fishing methods 383 12.5.2 Catchability 388 12.5.3 Catches 389 12.6 Monitoring and management 389 12.6.1 Management measures and structures 389 12.6.2 Stock assessments 391 12.6.3 Status of stocks 395 References 396 Chapter 13 Commercial Scyllarids 414 Ehud Spanier and Kari L. Lavalli 13.1 Introduction 414 13.2 Taxonomy and systematic hierarchy 415 13.2.1 Features of the genera 415 13.2.2 Species and distribution 416 13.3 Anatomy 426 13.4 Life history 429 13.4.1 Phyllosoma 430 13.4.2 Nistos 431 13.4.3 Juveniles 432 13.4.4 Adults 434 x Contents 13.5 Behaviour 436 13.5.1 Feeding behaviour 436 13.5.2 Sheltering behaviour and substrate preferences 440 13.5.3 Predators and antipredator behaviour 442 13.5.4 Mating behaviour 444 13.5.5 Movement patterns 445 13.6 Diseases 449 13.7 Harvest of wild populations 450 13.7.1 Thenus spp. fi shery 451 13.7.2 Ibacus alticrenatus fi shery 451 13.7.3 Ibacus brucei fi shery 451 13.7.4 Ibacus chacei fi shery 451 13.7.5 Ibacus ciliatus fi shery 452 13.7.6 Ibacus peronii fi shery 452 13.7.7 Parribacus spp. fi sheries 452 13.7.8 Scyllarides aequinoctialis fi shery 452 13.7.9 Scyllarides astori fi shery 452 13.7.10 Scyllarides deceptor fi shery 453 13.7.11 Scyllarides latus fi shery 453 13.7.12 Scyllarides nodifer fi shery 453 13.7.13 Scyllarides obtusus fi shery 454 13.7.14 Scyllarides squammosus fi shery 454 13.7.15 Fishery concerns 454 13.8 Aquaculture and restocking 455 13.9 Summary 456 References 456 Chapter 14 Conclusions 467 Bruce F. Phillips References 468 Index 471 Colour plate section between pp. 130 and 131