Towards a Science of Recruitment in Fish Populations
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OTTO KINNE Editor David H. Cushing TOWARDS A SCIENCE OF RECRUITMENT IN FISH POPULATIONS Introduction (Otto Kinne) David H. Cushing: A Laudatio (John D. Costlow) O L O G C Y Publisher: Ecology Institute E I Nordbünte 23, D-21385 Oldendorf/Luhe N E S T Germany T I T U David H. Cushing 198 Yarmouth Rd. Lowestoft Suffolk NR32 4AB United Kingdom ISSN 0932-2205 Copyright © 1996, by Ecology Institute, D-21385 Oldendorf/Luhe, Germany All rights reserved No part of this book may be reproduced by any means, or transmitted, or translated without written permission of the publisher Printed in Germany Typesetting by Ecology Institute, Oldendorf Printing and bookbinding by Westholsteinische Verlagsdruckerei Boyens & Co., Heide Printed on acid-free paper Contents Introduction (Otto Kinne) . VII David H. Cushing: A Laudatio (John Costlow) . XVII Preface . XX I HISTORY OF THE FISHERIES . 1 The Long History of the Fisheries . 1 Variability and Management . 8 II CLIMATE AND FISHERIES . 23 Faunal Movements and the Recent Period of Warming . 23 Evidence of Climate Effects . 25 Responses of Five Pelagic Stocks to Climate Changes . 28 The Climate of the North Atlantic . 37 Russell Cycle . 41 West Greenland Cod Stock . 45 Great Salinity Anomaly of the Seventies . 50 Zooplankton in the North Sea . 53 Gadoid Outburst . 55 A Broader View . 58 III A SIMPLE VIEW OF RECRUITMENT . 65 When Is the Magnitude of Recruitment Determined? . 65 Lack of Food . 70 Field Studies . 78 Predation . 83 Variability of Recruitment . 86 Match/Mismatch Hypothesis . 90 A Model of Match . 95 Conclusion . 103 IV NATURE OF THE STOCK RECRUITMENT RELATIONSHIP . 105 General Aspects . 105 Stock Recruitment Relationships of the Nine Well-Known Stocks 108 Two Investigations . 110 Density-Dependent Processes . 115 Ricker-Foerster thesis . 115 Cannibalism . 116 Density-dependent mortality . 119 Density-dependent growth . 126 Recruitment Overfishing . 129 Two Examples of Recruitment Collapse . 131 Collapse of the East Anglian herring fishery . 131 Collapse of the Northern cod . 133 Comparison between the two stocks . 137 Conclusion . 140 V THE SCIENCE OF RECRUITMENT . 143 General Aspects . 143 The Troubled Fisheries . 144 Recruitment Science . 151 Conclusion . 158 References . 161 Introduction Otto Kinne Ecology Institute, Nordbünte 23, D-21385 Oldendorf/Luhe, Germany This introduction addresses essential messages of David Cushing’s book. It then refers to the aims, activities and awards of the Ecology Institute, and lists ECI prize winners and OKF Fellows. Finally it presents the names of the 1996 ECI staff, and OKF trustees. About this book Knowledge on stock recruitment is the backbone of fisheries research and the basic prerequisite for sustaining the fishing industry. In this Excellence in Ecology book, David H. Cushing presents and weighs essential informa- tion on the conditions, known or hypothetical, that determine reproduction, growth and abundance of commercially valuable fish populations in oceans and coastal waters. He takes a close look at the history of fisheries and con- siders in detail environmental and human influences on fish production, and the autecological rules and dynamics of recruitment processes. Cushing’s intimate knowledge and solid expertise allow him to outline fascinating details on the birth of a new science—the science of recruitment—and to draft guidelines for developing that science in the years to come. David H. Cushing has dedicated most of his life, more than half a century, to fisheries research. No wonder, then, that his book comprises a unique wealth and diversity of information. To a considerable extent, he has pro- duced this information himself. The reader is presented with facts, thoughts and conclusions that have ripened over a long period of time. Human involvement with reproductive events in commercially exploited fish populations has ancient and multiple roots. These reach far into fields outside of scientific research. In fact, the total body of information available is enormous. But even more enormous is the complex web of interacting fac- tors involved and the size of the space in which all this happens: the seas and coastal waters which cover some two-thirds of the earth’s surface. Large sums of money, much ship space and numerous highly trained people are required for conducting modern ocean-wide fisheries investigations. Not surprisingly then, David Cushing concludes that we still do not know enough about the essential ecological phenomena of fish recruitment. And VIII INTRODUCTION Ecology Institute Prize 1992 in the field of marine ecology. Reproduction of the prize awarding document ECOLOGY INSTITUTE IX that our means for evaluating specific situations, for predicting future devel- opments, or for formulating detailed measures of protection and control remain, more often than not, inadequate. However, things are beginning to change: “Many have considered the study of the Stock Recruitment Rela- tionship a dead end, but it is now starting to yield information” (p. 140). Most fish populations are heavily exploited, often overexploited. What- ever is left is very vulnerable to impacts—natural or human-made. This is especially so if early offspring are overfished. Hence there is a great need for stricter fishing rules, protection and control. However, fishermen resist such interference with their business. They must make a profit to survive and they face increasing equipment costs and growing competition. Here, in the conflict between ecological and economical needs, lies the crux of it all. What can be done? True management of the vast areas of oceans and coastal waters is impossible. Besides, our oceans are more than production areas for human food and utilities. We must strive to accomplish three things: Reduction of the overall human impact on the seas. Enlargement and deepening of our knowledge on fish ecology, especially on larval stages. Enforcement of international cooperation in research, protection, politics and economics. About the Ecology Institute The international Ecology Institute (ECI) was founded in 1984. It is a non- governmental institution, sponsored by Inter-Research Science Publisher. ECI’s aims and activities have been described in detail in Excellence in Ecol- ogy Book 3 (Gene E. Likens, The Ecosystem Approach: Its Use and Abuse, 1992). In part, the ECI strives to achieve its aims by setting out awards to honor outstanding scientists: the ECI Prize and the IRPE Prize. The Institute also supports postgraduates in eastern European countries via the Otto Kinne Foundation (OKF). ECI and IRPE Prizes. The ECI Prize honors the sustained high perfor- mance of outstanding research ecologists. It is awarded annually, in a rotat- ing pattern, for the fields of marine, terrestrial and limnetic ecology. Laure- ates are selected by a jury of seven ECI members appointed by the director. The IRPE (International Recognition of Professional Excellence) Prize honors a young (not more than 40 years of age) research ecologists who has published uniquely independent and/or challenging papers representing an important scientific breakthrough and/or who must work under particularly difficult conditions. XINTRODUCTION OKF. The Otto Kinne Foundation supports promising postgraduates in environmental sciences in eastern European countries—especially in the fields of ecology, microbiology, disease studies, and climate research. The foundation aids individuals—without distinction of race, religion, national- ity, or sex—by providing financial assistance for research, for professional travel, and/or for acquiring scientific equipment or published scientific information. Details are available from the President of the Foundation: Dr. J. Lom, Institute of Parasitology, Academy of Sciences of the Czech Repub- lic, Branisˇovská 31, 370 05 Cˇ eské Budeˇjovice, Czech Republic; tel. (+42) 38 41158, fax (+42) 38 47743. EE Books. Excellence in Ecology books are authored by the winners of the ECI Prize. In these books, the laureates are offered the possibility of pub- lishing their personal views on the state of the art in their fields of expertise and of bringing to the attention of a world-wide audience their insights into the knowledge, problems and realities that form the biological basis for human existence. EE books address fellow scientists, teachers, students and decision makers who must translate ecological information into practicable rules and laws for the benefit of nature and mankind. EE books are made available at cost price.* A large number are donated to libraries in Third-World countries. Nominations. Nominations for ECI and IRPE Prizes (accompanied by CV, list of publications, and a statement why, in the opinion of the nomina- tor, the nominee qualifies for the prize) are invited from research ecologists on a global scale. They should be sent to the chairperson of the respective ECI Jury (in 1996: Prof. I. Hanski, Dept of Zoology, P. Rautatiekatu 13, FIN-00100 Helsinki, Finland; fax +358 191 1917492) or, alternatively, to the ECI’s director, who will then forward them to the chairperson. Eligible are all ecologists engaged in scientific research (except the ECI’s director, the Jury’s chairperson, and previous Laureates; Jury members nominated will be replaced by other ECI members). The Jury selects prize winners using the nominations received as well as their own knowledge of top performers and their own professional judgement. *For details see information at end of book. Please address orders for EE books to: *Inter-Research,