Ecology of Teleost Fishes
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1 Ecology of Teleost Fishes
2 Ecology of Teleost Fishes Robert J. Wootton Lecturer. Department of Biological Sciences. University College of Wales. Aberystwyth London New York Chapman and Hall
3 First published in 1990 by Chapman and Hall Ltd 11 New Fetter Lane, London EC4P 4EE Published in the USA by Chapman and Hall 29 West 35th Street. New York NY R. J. Wootton Softcover reprint of the hardcover 1 st edition 1990 Typeset in 10/12 Photina by Thomson Press (India) Ltd, New Delhi ISBN-13: e-isbn-13: DOI: / This title is available in both hardback and paperback editions. The paperback edition is sold subject to the condition that it shall not, by way of trade or otherwise, be lent, resold, hired out, or otherwise circulated without the publisher's prior consent in any form of binding or cover other than that in which it is published and without a similar condition including this condition being imposed on the subsequent purchaser. All rights reserved. No part of this book may be reprinted or reproduced, or utilized in any form or by any electronic, mechanical or other means, now known or hereafter invented, including photocopying and recording, or in any information storage and retrieval system, without permission in writing from the publisher. British Library Cataloguing in Publication Data Wootton, Robert J. Ecology of teleost fishes. 1. Teleostei. Ecology I. Title II. Series 597'.5045 Library of Congress Cataloging-in-Publication Data Wootton, R. J. (Robin Jeremy) Ecology of teleost fishes / Robin J. Wootton. p. cm.-(fish and fisheries series: 1) Bibliography: p. Includes index. 1. Osteichthyes-Ecology. I. Title. II. Series. QL615.W '.5045-dc CIP
4 For Maureen
5 Contents Acknowledgements Series Foreword Preface x xi xii 1 Introduction The diversity of teleost fishes Defining the problem Organization of the text Adaptive response to environmental change Summary and conclusions 14 2 Environmental and organismic constraints Introduction Body form and locomotion Respiration and gill structure Sensory capacities Summary and conclusions 30 3 Feeding Introduction Trophic categories in fishes Morphological adaptations for feeding Diet composition Temporal changes in diet composition Factors that determine the rate of food consumption Flexibility in the feeding ecology of fishes Summary and conclusions 71
6 viii Contents 4 Bioenergetics Introduction Structure of an energy budget Effects of environmental factors on metabolism Examples of energy budgets Summary and conclusions 95 5 Use of time and space Introducton Methods of studying the use of time and space Temporal patterns of fish movement Patterns and movement and the use of space Summary and conclusions Growth Introduction Definition and measurement of growth Patterns of growth Factors affecting growth rates Endogenous control of growth rates Modelling the growth of fish Summary and conclusions Reproduction Introduction Timing of reproduction The site of reproduction Allocation of resources to reproduction Allocation to individual progeny Unusual reproductive strategies Summary and conclusions Biotic interactions: I. Predation and parasitism Introduction Predation Pathogens Summary and conclusions Biotic interactions: II. Competition and mutualism Introduction Competition 216
7 Contents ix 9.3 Mutualism Summary and conclusions Dynamics of population abundance and production Introduction Defining the population Estimation of fish abundance Measures of the rate of population change Measurement of mortality rates Patterns of mortality in fish populations Regulation of fish populations and the stockrecruitment relationship Models of population growth Concept of production Measurement of production Bioenergetic basis of production Magnitude of population production Summary and conclusions Life-history strategies Introduction Evolution of life-history patterns Cost of reproduction and the consequences Bioenergetics of life... history patterns Phenotypic plasticity of life-history traits Life-history patterns and exploitation Summary and conclusions Fish assemblages Introduction General patterns of species diversity Historical factors in species diversity Contemporary determinants of species diversity Examples of fish assemblages Characteristics of fish assemblages Community-wide properties Summary and conclusions 342 References 344 Species Index 387 Subject Index 396
8 Acknowledgements I wish to thank Paul Hart. Tony Pitcher and Fred Whoriskey (and his postgraduates) for their comments on the whole text. Don Kramer. Richard Mann. Yibo Cui. John Gee and two North American referees made comments on sections ofthe text. Charles Hollingsworth was a painstaking. constructive and creative editor. Not all of these readers agreed with my approach to teleost ecology. but all offered valuable critisisms. These were not always accepted in the revision of the text. but nevertheless contributed to its final form. The structure of the text. the approach and the errors are mine. Most figures have been redrawn. but I am grateful to the publishers and authors who have freely given me permission to use material and are acknowledged in the appropriate figures and tables.
9 Series Foreword Among the fishes, a remarkably wide range of biological adaptations to diverse habitats has evolved. As well as living in the conventional habitats of lakes, ponds, rivers, rock pools and the open sea, fish have solved the problems of life in deserts, in the deep sea, in the cold antarctic, and in warm waters of high alkalinity or of low oxygen. Along with these adaptations, we find the most impressive specializations of morphology, physiology and behaviour. For example we can marvel at the high-speed swimming of the marlins, sailfish and warm-blooded tunas, air-breathing in catfish and lungfish, parental care in the mouth-brooding cichlids and viviparity in many sharks and toothcarps. Moreover, fish are of considerable importance to the survival of the human species in the form of nutritious and delicious food of numerous kinds. Rational expoitation and management of our global stocks of fishes must rely upon a detailed and precise insight of their biology. The Chapman and Hall Fish and Fisheries Series aims to present timely volumes reviewing important aspects of fish biology. Most volumes will be of interest to research workers in biology, zoology, ecology and physiology but an additional aim is for the books to be accessible to a wide spectrum of nonspecialist readers ranging from undergraduates and postgraduates to those with an interest in industrial and commercial aspects of fish and fisheries. Dr Wootton's book, which I am confident will become a classic text, reflects his longstanding interest in the link between the modem experimental discipline of ecology, and the lifetime reproductive success: the very stuff of evolutionary change which has put before us the stunning adaptations of teleost fishes. An up-to-date treatment in this area has been long overdue and the book complements other volumes in the series on behaviour and fisheries ecology. Dr Tony J. Pitcher Editor, Chapman and Hall Fish and Fisheries Series Reader in Biology, Bangor
10 Preface This book grew out of a series of lectures on the ecology of teleosts which I give to final year honours students in zoology and aquatic biology. It has been written because there is no recent text on the ecology of teleosts that reflects the advances made in the past two decades in area such as feeding ecology, bioenergetics or life-history strategies. The intended audience includes advanced undergraduates, post-graduates and research workers. It assumes a knowledge of general ecology such as that given in the recent text-books by Krebs (1985) and Begon, Harper and Townsend (1986) and offish biology such as that provided in Bond (1979) or Moyle and Cech (1982). The text, though referring where relevant to applied problems in fish ecology, is intended to complement rather than compete with volumes exemplified by Pitcher and Hart (1981), which have as their central theme the biological problems posed by fisheries or aquaculture. The present volume is intended not as a self-contained and self-sufficient account of teleost ecology but as an instrument to provoke its readers into their own explorations and interpretations of the literature and of contemporary research programmes. It emphasizes my belief that the key to understanding the ecology of any group of animals is an understanding of how individual animals react to environmental conditions by altering their allocations of time and resources to their different activities, and the consequences of these alterations for the reproductive success of individuals. It also reflects my belief that natural selection operates predominantly at the level of individuals and is a pervasive and powerful process. Thirdly, it emphasizes that teleosts are not 'primitive' or 'lower' vertebrates, a fallacy commonly held by undergraduates. The text, perhaps inevitably, reflects my research experience with freshwater, north temperate fishes. It is also heavily biased toward studies published in North America and the U.K., a restriction imposed by ease of access to journals and the author's lack of knowledge of languages other than English. Perhaps the best fate for this volume is that it provokes another and more comprehensive account of the ecology of this wonderfully diverse group of animals - the teleost fishes. R. J. Wootton Aberystwyth, 1989
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