Aquatic Food Webs This page intentionally left blank Aquatic Food Webs An Ecosystem Approach EDITED BY Andrea Belgrano National Center for Genome Resources (NCGR), Santa Fe, NM, USA Ursula M. Scharler University of Maryland Center for Environmental Science, Chesapeake Biological Laboratory (CBL), Solomons, MD, USA and Smithsonian Environmental Research Center, Edgewater, MD, USA Jennifer Dunne Pacific Ecoinformatics and Computational Ecology Lab, Berkeley, CA USA; Santa Fe Institute (SFI) Santa FE, NM, USA; Rocky Mountain Biological Laboratory, Crested Butte, CO USA AND Robert E. Ulanowicz University of Maryland Center for Environmental Science, Chesapeake Biological Laboratory (CBL), Solomons, MD, USA 1 1 Great Clarendon Street, Oxford OX2 6DP Oxford University Press is a department of the University of Oxford. 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No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, without the prior permission in writing of Oxford University Press, or as expressly permitted by law, or under terms agreed with the appropriate reprographics rights organization. Enquiries concerning reproduction outside the scope of the above should be sent to the Rights Department, Oxford University Press, at the address above You must not circulate this book in any other binding or cover and you must impose this same condition on any acquirer British Library Cataloging in Publication Data (Data available) Library of Congress Cataloging-in-Publication Data Aquatic food webs : an ecosystem approach / edited by Andrea Belgrano ... [et al.]. p. cm. Includes bibliographical references and index. ISBN 0-19-856482-1 (alk. paper) – ISBN 0-19-856483-X (alk. paper) 1. Aquatic ecology. 2. Food chains (Ecology) I. Belgrano, Andrea. QH541.5.W3A68225 2004 577.6016–dc22 2004027135 ISBN 0 19 856482 1 (Hbk) 9780198564829 ISBN 0 19 856483 X (Pbk) 9780198564836 10987654321 Typeset by Newgen Imaging Systems (P) Ltd., Chennai, India Printed in Great Britain on acid-free paper by Antony Rowe, Chippenham FOREWORD CURRENT AND FUTURE PERSPECTIVES ON FOOD WEBS Michel Loreau Food webs have been approached from two basic on food webs that is complementary to the energetic perspectives in ecology. First is the energetic view approach (DeAngelis 1992). Material cycles are articulated by Lindeman (1942), and developed by among the most common of the positive feedback ecosystem ecology during the following decades. loops discussed by Ulanowicz in his concluding In this view, food webs are networks of pathways remarks, and may explain key properties of eco- for the flow of energy in ecosystems, from its systems (Loreau 1998). The stoichiometry of ecolo- capture by autotrophs in the process of photo- gical interactions may further strongly constrain synthesis to its ultimate dissipation by hetero- food-web structure (Sterner and Elser 2002; Elser trophic respiration. I would venture to say that the and Hessen’s chapter). There has also been con- ecological network analysis advocated by siderable interest in the relationship between bio- Ulanowicz and colleagues in this book is heir to diversity and ecosystem functioning during the last this tradition. A different approach, rooted in decade (Loreau et al. 2002). Merging the theories that community ecology, was initiated by May (1973) bear upon food webs and the maintenance of species and pursued by Pimm (1982) and others. This diversity is urgently needed today, and may provide approach focuses on the dynamical constraints new insights into food-webs structure and ecosys- that arise from species interactions, and empha- tem functioning (Hillebrand and Shurin’s chapter). sises the fact that too much interaction (whether in The structure and functioning of ecological sys- the form of a larger number of species, a greater tems is determined not only by local constraints connectance among these species, or a higher and interactions, but also by larger-scale processes. mean interaction strength) destabilises food webs The importance of regional and historical influ- and ecological systems. The predictions resulting ences has been increasingly recognised in com- from this theory regarding the diversity and con- munity ecology (Ricklefs and Schluter 1993). The nectance of ecological systems led to a wave of extent to which they shape food webs, however, comparative topological studies on the structure of has been relatively little explored. The recent food webs. Thus, the two traditions converge in development of metacommunity theory (Leibold the search for patterns in food-web structure et al. 2004) provides a framework to start exam- despite different starting points. This book results ining spatial constraints on the structure and from the confluence of these two perspectives, functioning of local food webs (Melian et al.‘s which are discussed in a number of chapters. chapter). At even larger time scales, food webs are Patterns, however, are generally insufficient to the result of evolutionary processes which deter- infer processes. Thus, the search for explanations of mine their current properties. Complex food webs these patterns in terms of processes is still very much may readily evolve based on simple ecological alive, and in this search the energetic and dynamical interactions (McKane 2004). The evolution of food- perspectives are not the only possible ones. Bio- web and ecosystem properties is a fascinating geochemical cycles provide a functional perspective topic for future research. v vi FOREWORD This book provides a good synthesis of recent Loreau, M. 1998. Ecosystem development explained by research into aquatic food webs. I hope this competition within and between material cycles. Pro- synthesis will stimulate the development of new ceedings of the Royal Society of London, Series B 265: 33–38. approaches that link communities and ecosystems. Loreau, M., S. Naeem, and P. Inchausti. Eds. 2002. Bio- diversity and ecosystem functioning: synthesis and per- spectives. Oxford University Press, Oxford. References May, R. M. 1973. Stability and complexity in model ecosys- tems. Princeton University Press, Princeton. DeAngelis, D. L. 1992. Dynamics of nutrient cycling and McKane, A. J. 2004. Evolving complex food webs. The food webs. Chapman & Hall, London. European Physical Journal B 38: 287–295. Leibold, M. A., M. Holyoak, N. Mouquet, P. Amarasekare, Pimm, S. L. 1982. Food webs. Chapman & Hall, London. J. M. Chase, M. F. Hoopes, R. D. Holt, J. B. Shurin, R. Law, Ricklefs, R. E., and D. Schluter. Eds 1993. Species diversity D. Tilman, M. Loreau, and A. Gonzalez. 2004. The in ecological communities: historical and geographical per- metacommunity concept: a framework for multi-scale spectives. University of Chicago Press, Chicago. community ecology. Ecology Letters 7: 601–613. Sterner, R. W., and J. J. Elser. 2002. Ecological stoichiometry: Lindeman, R. L. 1942. The trophic-dynamic aspect of the biology of elements from molecules to the biosphere. ecology. Ecology 23: 399–418. Princeton University Press, Princeton. Contents Foreword v Michel Loreau Contributors ix Introduction 1 Andrea Belgrano PART I Structure and function 5 1 Biosimplicity via stoichiometry: the evolution of food-web structure and processes 7 James J. Elser and Dag O. Hessen 2 Spatial structure and dynamics in a marine food web 19 Carlos J. Melia´n, Jordi Bascompte, and Pedro Jordano 3 Role of network analysis in comparative ecosystem ecology of estuaries 25 Robert R. Christian, Daniel Baird, Joseph Luczkovich, Jeffrey C. Johnson, Ursula M. Scharler, and Robert E. Ulanowicz 4 Food webs in lakes—seasonal dynamics and the impact of climate variability 41 Dietmar Straile 5 Pattern and process in food webs: evidence from running waters 51 Guy Woodward, Ross Thompson, Colin R. Townsend, and Alan G. Hildrew PART II Examining food-web theories 67 6 Some random thoughts on the statistical analysis of food-web data 69 Andrew R. Solow 7 Analysis of size and complexity of randomly constructed food webs by information theoretic metrics 73 James T. Morris, Robert R. Christian, and Robert E. Ulanowicz 8 Size-based analyses of aquatic food webs 86 Simon Jennings vii viii CONTENTS 9 Food-web theory in marine ecosystems 98 Jason S. Link, William T. Stockhausen, and Elizabeth T. Methratta PART III Stability and diversity in food webs 115 10 Modeling food-web dynamics: complexity–stability implications 117 Jennifer A. Dunne, Ulrich Brose, Richard J. Williams, and Neo D. Martinez 11 Is biodiversity maintained by food-web complexity?—the adaptive food-web hypothesis 130 Michio Kondoh 12 Climate forcing, food web structure, and community dynamics in pelagic marine ecosystems 143 L. Ciannelli, D. Ø. Hjermann, P. Lehodey, G. Ottersen, J. T. Duffy-Anderson,