Cephalopods Ecology and Fisheries Peter Boyle University of Aberdeen Paul Rodhouse British Antarctic Survey Blackwell Science Cephalopods Ecology and Fisheries Cephalopods Ecology and Fisheries Peter Boyle University of Aberdeen Paul Rodhouse British Antarctic Survey Blackwell Science © 2005 by Blackwell Science Ltd a Blackwell Publishing company Editorial offices: Blackwell Science Ltd, 9600 Garsington Road, Oxford OX4 2DQ, UK Tel: +44 (0) 1865 776868 Blackwell Publishing Professional, 2121 State Avenue, Ames, Iowa 50014-8300, USA Tel: +1 515 292 0140 Blackwell Science Asia Pty Ltd, 550 Swanston Street, Carlton, Victoria 3053, Australia Tel: +61 (0)3 8359 1011 The right of the Author to be identified as the Author of this Work has been asserted in accordance with the Copyright, Designs and Patents Act 1988. All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, except as permitted by the UK Copyright, Designs and Patents Act 1988, without the prior permission of the publisher. First published 2005 Library of Congress Cataloging-in-Publication Data Boyle, P. R. Cephalopods: ecology and fisheries / Peter Boyle, Paul Rodhouse.– 1st ed. p. cm. Includes bibliographical references and index. ISBN 0-632-06048-4 (hardback : alk. paper) 1. Cephalopoda. 2. Octopus fisheries. 3. Squid fisheries. I. Rodhouse, Paul. II. Title. QL430.2.B69 2004 594′.5–dc22 2004005858 ISBN-10: 0-632-06048-4 ISBN-13: 978-0-632-06048-1 A catalogue record for this title is available from the British Library Set in 10/13 pt Times by Graphicraft Ltd, Hong Kong Printed and bound in India by Gopsons Papers, Noida The publisher's policy is to use permanent paper from mills that operate a sustainable forestry policy, and which has been manufactured from pulp processed using acid-free and elementary chlorine-free practices. Furthermore, the publisher ensures that the text paper and cover board used have met acceptable environmental accreditation standards. For further information on Blackwell Publishing, visit our website: www.blackwellpublishing.com To Ann and Laura Contents Preface ix Acknowledgements xi 1 Introduction 1 2 Form and function 7 3 Origin and evolution 36 4 Nautilus: the survivor 50 5 Biodiversity and zoogeography 62 6 Life cycle 80 7 Growth 101 8 Physiological ecology 116 9 Reproduction 131 10 From egg to recruitment 146 11 Coastal and shelf species 161 12 Oceanic and deep-sea species 176 13 Population ecology 205 14 Cephalopods as predators 222 15 Cephalopods as prey 234 16 Fishing methods and scientific sampling 259 17 Fishery resources 277 18 Fisheries oceanography 296 19 Assessment and management 313 20 Conclusion 335 Appendix A: Classification of living cephalopod families 343 Appendix B: Synopsis of living cephalopod families 346 References 373 Index 439 The colour plate section falls after page 308 Preface Squid, cuttlefish and octopuses are a group of marine molluscs which have been recognised, illustrated and appreciated from ancient times, and are of great interest to fishermen, cooks and students of natural history. Most were described and named dur- ing the nineteenth century, but it was the technology of the late twentieth century that brought them to wider scientific attention, allowing a generation of anatomists and physiologists to use laboratory studies on cephalopods to investigate the principles of organisation and function in their nervous system. Now, underwater access by diving and submersible vehicles increasingly reveals the range and complexity of their normal life in the sea, oceanic and deepwater expeditions explore the full range of their habi- tats, and twenty-first-century techniques such as molecular biology and remote sensing are making new advances. Widening investigations of the ecology of the seas and oceans have progressively revealed the major importance of cephalopods in the trophic structure of the marine ecosystem – as predators on many different animal types and as prey themselves for the larger fish, birds and mammals at the top of the ocean foodweb. Coastal hand-fisheries for cephalopods have always had local human importance, but man has joined the list of major cephalopod predators through large-scale commercial fisheries operating globally. These international fisheries take a large and still increasing share of the global take of living marine resources. The variety, scale and range of the fisheries make a major contribution to scientific knowledge of cephalopods at sea, and we feel that this fully justifies the integration of fisheries studies with traditional ecological approaches. Both of us, over many years, have introduced students and researchers from differ- ent backgrounds to a variety of cephalopod topics, and we have felt the need for a gen- eral text covering the basis of cephalopod biology, the patterns of their life history, and their relevance in the changing ecology of the world ocean. Despite the wealth of infor- mation in the scientific literature, there are few viewpoints from which the student and marine biologist or fisheries scientist can see a broader picture and begin to navigate towards their own area of interest. We have written this book for readers from backgrounds in biology, zoology, eco- logy, biological oceanography or fisheries, to help them become familiar with the themes of cephalopod biology and ecology, and to promote their greater integration into the mainstream of marine science. A particular challenge for all of us is the fluid state of the taxonomy and systematics of the group, and the changes that are likely to arise from application of the emerging molecular techniques in evolutionary and pop- ulation biology. We have adopted a pragmatic approach to the problem of scientific x Preface names that we judge will suit the majority of readers, since it is a compromise between the terminology in general use and the more strictly correct forms. Generations of marine biologists have found interest and excitement in the study of cephalopods and their work has been our source material for the writing of this book. To acknowledge this foundation, and to take full account of the diversity of modern studies, we have used citations from all periods of work, and drawn information and examples from most areas of the world. In undertaking this review we have been acutely aware of the increasing volume of new scientific literature – some ground- breaking and some repetitive. Inevitably, as soon as this book is in the hands of its readers it will already appear out of date, but new literature is readily found through modern bibliographic methods and set into the framework we have tried to assemble. We hope to have reported our remit objectively, if selectively, and to have placed cephalopod ecology and fisheries into some overall perspective which will be useful as a basis for widening our understanding of the marine ecosystem as a whole. Peter Boyle and Paul Rodhouse Aberdeen and Cambridge Acknowledgements We gratefully acknowledge the sustained support and indulgence we have received from the University of Aberdeen and the British Antarctic Survey in the writing of this book, and the stimulating environment of a scientific cruise to the Southern Ocean on the research vessel RRS James Clark Ross during which much of the early planning of the volume took place. Specifically, we wish to acknowledge the following individuals and organisations for their help in preparing this book and for sources of information and illustra- tions. Colleagues who read individual chapters or sections of the work and directly provided very generous scientific input and comment were: David Agnew, Louise Allcock, Sigurd von Boletzky, Andy Clarke, Martin Collins, John Croxall, Natalie Moltschaniskyj, Uwe Piatkowski, Graham Pierce, Dick Young, Roger Villanueva and Jose Xavier. Broadly, we owe a great debt to the many scientific friends and colleagues worldwide with whom our scientific views have evolved through numerous sociable discussions as well as formal conferences over the last couple of decades. Among these friends and mentors, Sigurd von Boletzky, Malcolm Clarke, Katharina Mangold, Kir Nesis, Takashi Okutani, Clyde Roper, Gilbert Vass and Nancy Vass need special mention for their influence in establishing cephalopod ecology and fisheries as a field of endeavour. For their contribution of original illustrations, photographs or unpublished obser- vations we are grateful to: Stephen Craig for Fig. 10.5; Martin Collins for Fig. 12.10 and the photograph of the giant squid in Box 12.1; Françoise Dubas for the photo- graphs of chromatophores in Box 2.3; Aidan Emery for Fig. 9.6; Magnus Ngoile for the photograph of the statolith section in Box 7.1; Ron O’Dor for the photograph of squid tagging in Box 13.1; Claire Waluda for Fig. 19.5 and 18.2. For original colour plates we are grateful to Martin Gorman for the photograph of the Octopus prepared for sale at a Fijian open market (Plate 12), Martin Guard for the photograph of the Tanzanian fisherman with his catch of Octopus cyanea (Plate 10), Jaruwat Nabhitabhata for the photograph of the squid trap in Thailand (Plate 9) and Carmen Yamashiro for the photograph of the Humbolt Current squid, Dosidicus gigas, caught by a Peruvian jigging vessel (Plate 11). Ian Rendell (Aberdeen) and Juliet Corley (Cambridge) undertook the major work of preparing the line illustrations, Jianjun Wang (Aberdeen) provided all of the dis- tribution maps, and Lyndsay Brown (Aberdeen) and Nadine Johnston (Cambridge) provided considerable assistance with the manuscript and bibliography. PRB is particularly grateful to the Aberdeen Royal Infirmary for their timely medical intervention in his case and for the support of the Inverurie Medical Group, xii Acknowledgements without whose professional skills and attention, his part in this work would certainly not have been completed. The cover design, a mosaic from the House of the Faun, Pompeii, is reproduced by kind permission of the Soprintendenza per I Beni Archaologici delle province di Napoli e Caserta, Ministero per I Beni e le Attività Culturali, Piazza Museo, Napoli.