Cascading trophic interactions Cambridge Studies in Ecology presents balanced, comprehensive, up-to-date, and critical reviews of selected topics within ecology, both botanical and zoological. The Series is aimed at advanced final-year undergraduates, graduate students, researchers, and university teachers, as well as ecologists in industry and government research. It encompasses a wide range of approaches and spatial, temporal, and taxonomic scales in ecology, including quantitative, theoretical, population, community, ecosystem, historical, experimental, behavioral and evolutionary studies. The emphasis throughout is on ecology related to the real world of plants and animals in the field rather than on purely theoretical abstractions and mathematical models. Some books in the Series attempt to challenge existing ecological paradigms and present new concepts, empirical or theoretical models, and testable hypotheses. Others attempt to explore new approaches and present syntheses on topics of considerable importance ecologically which cut across the conventional but artifi- cial boundaries within the science of ecology. CAMBRIDGE STUDIES IN ECOLOGY Series Editors H. J. B. Birks Botanical Institute, University of Bergen, Norway, and Environmental Change Research Centre, University College London, UK J. A. Wiens Department of Biology, Colorado State University, USA Advisory Board P. Adam University of New South Wales, Australia R. T. Paine University of Washington, Seattle, USA R. B. Root Cornell University, USA F. I. Woodward University of Sheffield, UK ALSO IN THE SERIES H. G. Gauch, Jr Multivariate Analysis in Community Ecology R. H. Peters Hie Ecological Implications of Body Size C. S. Reynolds The Ecology of Freshwater Phytoplankton K. A. Kershaw Physiological Ecology of Lichens R. P. Mclntosh Tlie Background of Ecology: Concepts and Tlieory A. J. Beattie Tlie Evolutionary Ecology of Ant-Plant Mutualisms F. I. Woodward Climate and Plant Distribution J. J. Burdon Diseases and Plant Population Biology J. I. Sprent Tlie Ecology of the Nitrogen Cycle N. G. Hairston, Sr Community Ecology and Salamander Guilds H. Stolp Microbial Ecology: Organisms, Habitats and Activities R. N. Owen-Smith Megaherbivores: Tlie Influence of Large Body Size on Ecology J. A. Wiens Tlie Ecology of Bird Communities N. G. Hairston, Sr Ecological Experiments R. Hengeveld Dynamic Biogeography C. Little Tlie Terrestrial Invasion: An Ecophysiological Approach to the Origins of Land Animals P. Adam Saltmarsh Ecology M. F. Allen Tlie Ecology of Mycorrhizae D. J. Von Willert et al. Life Strategies of Succulents in Deserts J. A. Matthews Tlie Ecology of Recently-deglaciated Terrain E. A. Johnson Fire and Vegetation Dynamics D. H. Wise Spiders in Ecological Webs J. S. Findley Bats: A Community Perspective G. P. Malanson Riparian Landscapes S. R. Carpenter & Tlie Trophic Cascade in Lakes J. F. Kitchell (Eds.) R. J. Whelan Tlie Ecology of Fire R. C. Mac Nally Ecological Versatility and Community Ecology The trophic cascade in lakes edited by STEPHEN R. CARPENTER and JAMES F. KITCHELL CAMBRIDGE UNIVERSITY PRESS Published by the Press Syndicate of the University of Cambridge The Pitt Building, Trumpington Street, Cambridge CB2 1RP 40 West 20th Street, New York, NY 10011-4211, USA 10 Stamford Road, Oakleigh, Melbourne 3166, Australia © Cambridge University Press 1993 First published 1993 First paperback edition 1996 Printed in Great Britain at the University Press, Cambridge A catalogue record for this book is available from the British Library Library of Congress cataloguing in publication data The trophic cascade in lakes/edited by Stephen R. Carpenter and James F. Kitchell. p. cm. — (Cambridge studies in ecology) Includes bibliographical references and index. ISBN 0-521-43145-X 1. Lake ecology. 2. Biological productivity. 3. Food chains (Ecology) I. Carpenter, Stephen R. II. Kitchell, James F. III. Series. QH541.5.L3T76 1993 574.5'26322-dc20 92-36737 CIP ISBN 0 521 43145 X hardback ISBN 0 521 56684 3 paperback SE To our mentor Arthur Hasler, pioneer ecosystem experimenter; and in memory of David Benkowski, whose good humor and skill live on among the legends of the northern lakes Contents Contributors page xi Preface xiii 1 Cascading trophic interactions 1 J. F. Kitchell and S. R. Carpenter 2 Experimental lakes, manipulations and measurements 15 S.R. Carpenter andj. F. Kitchell 3 Statistical analysis of the ecosystem experiments 26 S. R. Carpenter 4 The fish populations 43 J. R. Hodgson, X. He and]. F. Kitchell 5 Fish behavioral and community responses to manipulation 69 X. He, R. Wright and]. F. Kitchell 6 Roles offish predation: piscivory and planktivory 85 X. He,}. F. Kitchell, J. R. Hodgson, R. Wright, P. A. Soranno, D. M. Lodge, P. A. Cochran, D. Benkowski and N. Bouwes 1 Dynamics of the phantom midge: implications for zooplankton 103 P. A. Soranno, S. R. Carpenter and S. M. Moegenburg 8 Zooplankton community dynamics 116 P. A. Soranno, S. R. Carpenter and M. M. Elser 9 Effects of predators and food supply on diel vertical migration oiDaphnia 153 M. L. Dini, P. A. Soranno, M. D. Scheuerell and S. R. Carpenter x • Contents 10 Zooplankton biomass and body size 172 P. A. Soranno, S. R. Carpenter and X. He 11 Phytoplankton community dynamics 189 S. R. Carpenter, J. A. Morrice,J.J. Elser, A. L. St. Amand and N. A. MacKay 12 Metalimnetic phytoplankton dynamics 210 A. L. St. Amand and S. R. Carpenter 13 Primary production and its interactions with nutrients and light transmission 225 S. R. Carpenter, J. A. Morrice, P. A. Soranno, J.J. Elser, N. A. MacKay and A. L. St. Amand 14 Heterotrophic microbial processes 252 M. L. Pace 15 Annual fossil records of food-web manipulation 278 P. R. Leavitt, P. R. Sanford, S. R. Carpenter, J. F. Kitchell and D. Benkowski 16 Simulation models of the trophic cascade: predictions and evaluations 310 S. R. Carpenter and}. F. Kitchell 17 Synthesis and new directions 332 J. F. Kitchell and S. R. Carpenter References 351 Index 381 Contributors David Benkowski (Deceased) Nicolaas Bouwes, Department of Fisheries and Wildlife, Utah State Univer- sity, Logan, UT84322, U.S.A. Stephen R. Carpenter, Center for Limnology, University of Wisconsin, Madison, WI53706, U.S.A. Philip A. Cochran, Division of Natural Sciences, St. Norbert College, DePere, WI54115, U.S.A. Michael L. Dini, Department of Biological Sciences, Texas Tech University, Box 43131, Lubbock, TX 79409, U.S.A. James J. Elser, Department of Zoology, Arizona State University, Tempe, AZ 85287, U.S.A. Monica M. Elser, 1843 East Laguna Drive, Tempe, AZ 85282, U.S.A. XiHe, Center for Limnology, University of Wisconsin, Madison, WI 53706, U.S.A. James R. Hodgson, Department of Biology, St. Norbert College, De Pere, WI54115, U.S.A. James F. Kitchell, Center for Limnology, University of Wisconsin, Madison, WI 53706, U.S.A. Peter R. Leavitt, Zoology Department, Biological Sciences Building, Univer- sity of Alberta, Edmonton, Alberta, Canada K6G 2E9 David M. Lodge, Department of Biological Sciences, University of Notre Dame, Notre Dame, IN46556, U.S.A. Neil A. MacKay, Zoology Department, Arizona State University, Tempe, AZ 85287, U.S.A. xii • Contributors Susan M. Moegenburg, Department of Zoology, 233 Bartram, University of Florida, Gainesville, FL 32611, U.S.A. John A. Morrice, Department of Biology, University of New Mexico, Albuquerque, NM 87131, U.S.A. Michael L. Pace, Institute for Ecosystems Studies, Cary Arboretum, Box AB, Millbrook, NY 12545, U.S.A. Patricia R. Sanford, Center for Limnology, University of Wisconsin, Madison, WI53706, U.S.A. Mark D. Scheuerell, Cornell Biological Field Station, R.D. #1, Bridgeport, NY 13030, U.S.A. Patricia A. Soranno, Center for Limnology, University of Wisconsin, Madison, W153706, U.S.A. Ann L. St. Amand, Phycotech, Inc., P.O. Box 218, Baroda, MI 49101, U.S.A. Russell Wright, Center for Limnology, University of Wisconsin, Madison, WI53706, U.S.A.