Fishery Science: The Unique Contributions of Early Life Stages Lee A. Fuiman Robert G. Werner Blackwell Science 00 03/05/2002 08:37 Page i Fishery Science 00 03/05/2002 08:37 Page ii We dedicate this book to our good friend John Blaxter, the gentleman scientist. His scientific excellence and creativity as well as his personal charm and good humor have made permanent impressions on both of us. John’s scientific contributions permeate this book, which we hope will carry his legacy to many future generations of fishery scientists. 00 03/05/2002 08:37 Page iii Fishery Science The Unique Contributions of Early Life Stages Edited by Lee A. Fuiman Department of Marine Science, University of Texas at Austin, Marine Science Institute, Port Aransas, Texas, USA and Robert G. Werner College of Environmental Science and Forestry, State University of New York, Syracuse, New York, USA 00 03/05/2002 08:37 Page iv © 2002 by Blackwell Science Ltd, First published 2002 by Blackwell Science Ltd a Blackwell Publishing Company Editorial Offices: Library of Congress Osney Mead, Oxford OX2 0EL, UK Cataloging-in-Publication Data Tel: +44 (0)1865 206206 is available Blackwell Science, Inc., 350 Main Street, Malden, MA 02148-5018, USA ISBN 0-632-05661-4 Tel: +1 781 388 8250 Iowa State Press, a Blackwell Publishing A catalogue record for this title is available from Company, 2121 State Avenue, Ames, Iowa the British Library 50014-8300, USA Tel: +1 515 292 0140 Set in Times by Gray Publishing, Tunbridge Blackwell Science Asia Pty, 54 University Street, Wells, Kent Carlton, Victoria 3053, Australia Printed and bound in Great Britain by Tel: +61 (0)3 9347 0300 MPG Books, Bodmin, Cornwall Blackwell Wissenschafts Verlag, Kurfürstendamm 57, 10707 Berlin, Germany Tel: +49 (0)30 32 79 060 For further information on Blackwell Science, visit our website: The right of the Author to be identified as the www.blackwell-science.com 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. 00 03/05/2002 08:37 Page v Contents Contributors vii Preface ix 1 Special Considerations of Fish Eggs and Larvae 1 Lee A. Fuiman 2 Age and Growth 33 Cynthia M. Jones 3 Mortality 64 Edward D. Houde 4 Recruitment 88 James H. Cowan, Jr. and Richard F. Shaw 5 Population Analysis 112 Pierre Pepin 6 Cohort Identification 143 Karin E. Limburg 7 Habitat Requirements 161 Robert G. Werner 8 Assemblages, Communities, and Species Interactions 183 Thomas J. Miller 9 Fishery Management 206 Edward S. Rutherford 10 Human Impacts 222 G. Joan Holt 11 Case Studies Resurgence and Decline of the Japanese Sardine Population 243 Yoshiro Watanabe Cascading Effects of Human Impacts on Fish Populations in the Laurentian Great Lakes 257 James A. Rice Understanding Conservation Issues of the Danube River 272 Hubert Keckeis and Fritz Schiemer v 00 03/05/2002 08:37 Page vi Contents vi 12 Methodological Resources 289 Robert G. Werner and Lee A. Fuiman Appendix: List of Symbols 301 Literature Cited 303 Subject Index 316 Taxonomic Index 324 00 03/05/2002 08:37 Page vii Contributors JAMES H. COWAN, JR. Department of Oceanography and Coastal Sciences, Louisiana State University, Baton Rouge, Louisiana, USA LEE A. FUIMAN Department of Marine Science, University of Texas at Austin, Marine Science Institute, Port Aransas, Texas, USA G. JOAN HOLT Department of Marine Science, University of Texas at Austin, Marine Science Institute, Port Aransas, Texas, USA EDWARD D. HOUDE University of Maryland Center for Environmental Science, Chesapeake Biological Laboratory, Solomons, Maryland, USA CYNTHIA M. JONES Center for Quantitative Fisheries Ecology, Old Dominion University, Norfolk, Virginia, USA HUBERT KECKEIS Institute of Ecology and Conservation Biology, Department of Limnology, University of Vienna, Vienna, Austria KARIN E. LIMBURG College of Environmental Science and Forestry, State University of New York, Syracuse, New York, USA THOMAS J. MILLER University of Maryland Center for Environmental Science, Chesapeake Biological Laboratory, Solomons, Maryland, USA PIERRE PEPIN Department of Fisheries and Oceans, St. Johns, Newfoundland, Canada JAMES A. RICE Zoology Department, North Carolina State University, Raleigh, North Carolina, USA EDWARD S. RUTHERFORD School of Natural Resources and Environment, University of Michigan, Ann Arbor, Michigan, USA FRITZ SCHIEMER Institute of Ecology and Conservation Biology, Department of Limnology, University of Vienna, Vienna, Austria RICHARD F. S HAW Department of Oceanography and Coastal Sciences, Louisiana State University, Baton Rouge, Louisiana, USA YOSHIRO WATANABE Ocean Research Institute, University of Tokyo, Tokyo, Japan ROBERT G. WERNER College of Environmental Science and Forestry, State University of New York, Syracuse, New York, USA vii 00 03/05/2002 08:37 Page viii 00 03/05/2002 08:37 Page ix Preface There has been explosive growth in research on the early life of fishes over the past 30 years. A great deal of that research was undertaken in order to develop a more complete under- standing of the dynamics of fish populations. It is now clear that information derived from fish eggs and larvae makes a number of unique contributions to fishery science that are cru- cial for accurate assessment and management of fish populations. Nevertheless, very little of this information has found its way into textbooks and college-level courses in fishery sci- ence. After several decades of intensive research on early life stages of fishes, there is now a mature body of knowledge that is ready to be summarized and distilled for students and the non-specialist audience. Our intention with this book is to improve the training of young fishery scientists by demonstrating why fish eggs and larvae are important to con- sider, how the characteristics of early life stages require a somewhat different research approach, and how information on early life stages can be applied and interpreted to yield unique and important insights into fish populations. This is designed to be a supplemental textbook, to complement the material that is covered in existing textbooks and courses on fishery science. The chapters are written in a didactic and readable style by university instructors who are also respected specialists in larval fish research. The first 10 chapters focus on the concepts that are traditionally covered in such courses. Each chapter ends with a list of references that provide more information on the material covered. A selection of case studies follows, each of which demonstrates several specific applications of early life-history information to a fishery problem. The case studies represent three very different aquatic environments: the open ocean, a lake, and a river. The final chapter provides a starting point for learning more about the methods for working with fish eggs and larvae. Throughout the book we attempted to provide a balanced coverage of marine and freshwater environments and geographical regions, but uneven availability of information and personal biases were inevitable. We hope that this book will be useful not only for instruction of introductory fishery science courses, but also as a core text for graduate-level courses in larval fish biology. It may also serve well as a reference that brings to a wider fishery science audience an understanding of the unique contributions of early life stages. We gratefully acknowledge the strong support of the Early Life History Section of the American Fisheries Society, which generously provided funding and encouragement ix 00 03/05/2002 08:37 Page x x Preface for production of this book. We are grateful to our colleagues who enthusiastically agreed to contribute chapters and did a fine job of understanding and following our plan. We also appreciate the assistance of Darrel Snyder, who offered thoughtful advice, and Heather Alexander, who help with the graphics. Finally, we would like to acknowledge the thoughtful support of our wives, Linda and Jo, who made this effort much more pleasant than it otherwise would have been. L.A.F. R.G.W. 01 03/05/2002 08:39 Page 1 Chapter 1 Special Considerations of Fish Eggs and Larvae Lee A. Fuiman 1.1 Introduction Fishery science attempts to understand the dynamics of fish populations with the goal of optimizing some human benefit or value, such as the yield to a commercial fishery, hours of pleasurable angling, or conservation of a species, population, or ecological community. Virtually all management goals center on adult fishes and so the traditional tools of fishery science were forged in an understanding of the biology and ecology of adult fishes. The critical role that early life stages play in the dynamics of fish populations is easily recognized when we consider that every fish taken by commercial trawlers and long-liners, each trophy fish landed by the patient and skilled angler, and all of the spawners in the relic population of an endangered species were, at one time, inconspicuous embryos and larvae, and that only a tiny fraction of their cohort survived the first few months of life. With this realization it becomes clear that fishery science demands a broader understanding of fishes, one that encompasses the early life stages. This chapter highlights some of the important differences between early life stages of fishes and the more familiar juveniles and adults. It provides a general introduction to the developmental changes that take place during early life and some of the ways in which early life-history traits are interrelated. A comprehensive description of all developmental changes or the breadth of variation among species is beyond the scope of this chapter.