DYNAMIC STATE VARIABLE MODELS IN ECOLOGY OXFORD SERIES IN ECOLOGY AND EVOLUTION Edited by Robert M. May and Paul H. Harvey The Comparative Method in Evolutionary Biology Paul H. Harvey and Mark D. Pagel The Causes of Molecular Evolution John H. Gillespie Dunnock Behaviour and Social Evolution N.B. Davies Natural Selection: Domains, Levels, and Challenges George C. Williams Behaviour and Social Evolution of Wasps: The Communal Aggregation Hypothesis Yosiaki Ito Life History Invariants: Some Explorations of Symmetry in Evolutionary Ecology Eric L. Charnov Quantitative Ecology and the Brown Trout J.M. Elliott Sexual Selection and the Barn Swallow Anders Pape M011er Ecology and Evolution in Anoxic Worlds Tom Fenchel and Bland J. Finlay Anolis Lizards of the Caribbean: Ecology, Evolution, and Plate Tectonics Jonathan Roughgarden From Individual Behaviour to Population Ecology William J. Sutherland Evolution of Social Insect Colonies: Sex Allocation and Kin Selection Ross H. Crozier and Pekka Pamilo Biological Invasions: Theory and Practice Nanako Shigesada and Kohkichi Kawasaki Cooperation among Animals: An Evolutionary Perspective Lee Alan Dugatkin Natural Hybridization and Evolution Michael L. Arnold Evolution of Sibling Rivalry Douglas Mock and Geoffrey Parker Asymmetry and Evolutionary Biology Anders Pape Mo11er and John P. Swaddle Dynamic State Variable Models in Ecology: Methods and Applications Colin W. Clark and Marc Mangel DYNAMIC STATE VARIABLE MODELS IN ECOLOGY Methods and Applications Colin W. Clark Marc Mangel New York Oxford OXFORD UNIVERSITY PRESS 2000 Oxford University Press Oxford New York Athens Auckland Bangkok Bogota Buenos Aires Calcutta Cape Town Chennai Dar es Salaam Delhi Florence Hong Kong Istanbul Karachi Kuala Lumpur Madrid Melbourne Mexico City Mumbai Nairobi Paris Sao Paulo Singapore Taipei Toronto Warsaw and associated companies in Berlin Ibadan Copyright © 2000 by Oxford University Press, Inc. Published by Oxford University Press, Inc. 198 Madison Avenue, New York, New York 10016 Oxford is a registered trademark of Oxford University Press 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, without the prior permission of Oxford University Press. Library of Congress Cataloging-in-Publication Data Clark, Colin Whitcomb, 1931- Dynamic state variable models in ecology: methods and applications / Colin W. Clark and Marc Mangel. p. cm. — (Oxford series in ecology and evolution) Includes bibliographical references and index. ISBN 0-19-512266-6; ISBN 0-19-512267-4 (pbk) 1. Animal behavior—Mathematical models. 2. Animal ecology— Mathematical models. I. Mangel, Marc. II. Title. III. Series. QL751.65.M3C58 1999 577'.01'5118—dc21 99-12265 1 3 5 7 9 8 6 42 Printed in the United States of America on acid-free paper Preface For discussions and comments about the book, we thank Peter Bednekoff, Jane Brockman, Anders Brodin, Tim Collier, Reuven Dukas, Shea Gardner, Paul Hart, Pat Kennedy, Don Ludwig, Barney Luttbeg, Bernie Roitberg, Jay Rosenheim, and Ron Ydenberg. Many young scientists contributed to the development of the book by participating in courses based on early drafts. We thank at the University of California at Santa Cruz, Edgar Becerra, Melanie Bojanowski, Bret Eldred, Samantha Forde, Eric Danner, Jennifer Brown, Karen Crow, Diana Steller, Matt Kauffman, Kim Heinemeyer, Josh Livni, Diane Thomson, Chris Wilcox, and especially Angie Shelton; at Princeton, Jon Cline, Leila Hadj-Chikh, Tim Kailing, David Smith, Thomas Valqui, and Erika Zavaleta; at the University of British Columbia, Marty Anderies, Chris Fonnesbeck and Edward Gregr; and at Simon Fraser University, Sigal Blay, Julian Christians, and Nick Hughes. A special vote of thanks goes to Janet Clark, who typeset in T K many drafts, as E well as the final published text. The original research reported on here has been supported by grants to CWC from the Natural Sciences and Engineering Research Council of Canada, and to MM from the California Sea Grant Program, the National Science Foundation, and the U. S. Department of Agriculture. We were fortunate to be able to review the final draft on the Big Island of Hawaii, uninterrupted except by omao, yellow tang, and spinner dolphins. Vancouver, British Columbia C. W. C. Santa Cruz, California M. M. August 1999 This page intentionally left blank Contents 1 The Basics, 3 1.1 Introduction: Biology, constraints and trade-offs, 3 1.2 An example: Formalizing constraints and trade-offs, 4 1.3 A brief digression on probability, 5 1.4 The algorithm of stochastic dynamic programming, 7 1.5 Computer implementation, 9 1.6 Evaluating the fitness of alternative strategies, 18 1.7 Developing a dynamic state variable model, 20 1.8 A model for feeding by the three-spined stickleback, 30 1.9 The role of dynamic state variable models in ecology, 38 1.10 Examples of projects, 40 Appendix: Rate-maximizing theories of diet selection and habitat choice, 45 2 Some Details of Technique, 49 2.1 Linear interpolation, 49 2.2 Sequential coupling, 52 2.3 Forward iteration and Monte Carlo simulation, 54 2.4 The critical level of reserves, 57 2.5 Environmental state variables, 57 2.6 Errors in decisions, 60 2.7 Variable constraints, 61 2.8 Functional forms, 67 3 Using the Model, 71 3.1 Levels of detail in model predictions, 71 3.2 Comparing models and data, 72 3.3 Obtaining and analyzing predictions, 78 3.4 Sensitivity analysis, 80 3.5 Suboptimality analysis, 81 4 Oviposition Behavior of Insect Parasitoids, 82 4.1 Parasitoid life histories, 82 4.2 Fixed clutch models, 86 viii • Contents 4.3 A dynamic but state-independent model, 89 4.4 The proovigenic parasitoid, 95 4.5 The synovigenic parasitoid: Eggs and reserves, 99 4.6 Discussion, 102 5 Winter Survival Strategies, 108 5.1 Foraging and fattening strategies for willow tits, 108 5.2 Social interaction, 118 5.3 Experiments suggested by the model, 121 5.4 Models of hoarding behavior, 122 5.5 Long-term hoarding, 124 5.6 Overwinter foraging by juvenile salmon, 132 5.7 A state variable model for overwinter lipid dynamics, 133 5.8 Empirical test of the predictions, 136 6 Avian Migration, 139 6.1 The energetics of bird migration, 140 6.2 Optimal fuel loads, 144 6.3 A dynamic state variable model of single-flight migration, 147 6.4 Spring migration of western sandpipers, 149 7 Human Behavioral Ecology, 161 7.1 Dynamic discarding in fisheries, 161 7.2 Optimal reproductive strategies in !kung bushmcn females, 167 8 Conservation Biology, 173 8.1 Assessing the fitness consequences of environmental change, 173 8.2 Using fire rotation schemes to maximize the persistence of midsuccessional species, 176 8.3 Managing a metapopulation, 180 8.4 Some other applications, 188 9 Agroecology, 192 9.1 Behavioral origins of the functional response, 192 9.2 Behavior and population regulation: The behavioral stabilization of host—parasitoid interactions, 199 9.3 Weed control: Predicting flowering in thistles and other monocarpic perennials, 206 9.4 Discussion, 209 Contents • ix 10 Population-Level Models, 212 10.1 Ideal free distributions, 212 10.2 Patch choice with frequency-dependent predation risk, 214 10.3 Genetic algorithms as optimization tools, 223 11 Stochasticity, Uncertainty, and Information as a State Variable, 232 11.1 Ephemeral patches and uncertain food, 232 11.2 A Bayesian foraging model, 238 11.3 Tracking a changing environment, 244 11.4 Discussion, 245 Appendix: Bayesian updating, 246 12 Measures of Fitness, 248 12.1 Nonoverlapping generations, 248 12.2 Overlapping generations, 253 12.3 Fitness measures in dynamic state variable models, 259 Appendix: Programs available at the OUP Web site, 265 References, 267 Index, 287
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