The Biology of Lakes and Ponds BIOLOGY OF HABITATS Series editors: M. Crawley, C. Little, T. R. E. Southwood, and S. Ulfstrand The intention is to publish attractive texts giving an integrated overview of the design, physiology, ecology, and behaviour of the organisms in given habitats. Each book will provide information about the habitat and the types of organisms present, on practical aspects of working within the habitats and the sorts of studies which are possible, and will include a discussion of biodiversity and conservation needs. The series is intended for naturalists, students studying biological or environmental sciences, those beginning independent research, and biologists embarking on research in a new habitat. The Biology of Rocky Shores Colin Little and J. A. Kitching The Biology of Polar Habitats G. E. Fogg The Biology of Lakes and Ponds Christer Brönmark and Lars-Anders Hansson The Biology of Streams and Rivers Paul S. Giller and Björn Malmqvist The Biology of Mangroves Peter J. Hogarth The Biology of Soft Shores and Estuaries Colin Little The Biology of the Deep Ocean Peter Herring The Biology of Lakes and Ponds, second edition Christer Brönmark and Lars-Anders Hansson The Biology of Soil Richard Pardgeti The Biology of Lakes and Ponds Christer Brönmark Department of Ecology University of Lund, Sweden and Lars-Anders Hansson Department of Ecology University of Lund, Sweden 1 3 Great Clarendon Street, Oxford OX2 6DP Oxford University Press is a department of the University of Oxford. It furthers the University’s objective of excellence in research, scholarship, and education by publishing worldwide in Oxford New York Auckland Cape Town Dar es Salaam Hong Kong Karachi Kuala Lumpur Madrid Melbourne Mexico City Nairobi New Delhi Shanghai Taipei Toronto With offices in Argentina Austria Brazil Chile Czech Republic France Greece Guatemala Hungary Italy Japan Poland Portugal Singapore South Korea Switzerland Thailand Turkey Ukraine Vietnam Oxford is a registered trade mark of Oxford University Press in the UK and in certain other countries Published in the United States by Oxford University Press Inc., New York © Christer Brönmark and Lars-Anders Hansson, 1998, 2005 The moral rights of the authors have been asserted Database right Oxford University Press (maker) First editon published 1998 Reprinted 1999, 2000, 2001, twice 2002, 2003, 2004 Second edition published 2005 Reprinted 2005 (with corrections) 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, 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 the same condition on any acquirer A catalogue record for this title is available from the British Library Library of Congress Cataloging-in-Publication Data Brönmark, Christer. The biology of lakes and ponds / Christer Brönmark, Lars-Anders Hansson.—2nd ed. p. cm. ISBN 0–19–851612–6 (alk. paper)—ISBN 0–19–851613–4 (alk. paper) 1. Limnology. 2. Lake ecology. 3. Pond ecology. I. Hansson, Lars-Anders. II. Title. QH96.B724 2005 577.63—dc22 2004025884 Typeset by Newgen Imaging Systems (P) Ltd., Chennai, India Printed in Great Britain on acid-free paper by Biddles, King’s Lynn. ISBN 0 19 851612 6 (Hbk) 978 0 19 851612 5 ISBN 0 19 851613 4 (Pbk) 978 0 19 851613 2 1098765432 Foreword Lakes and ponds cover only a very small portion of the earth’s surface, yet their study has played a disproportionately large role in the development of ecological theory. From broad ecosystem-level concepts such as trophic structure and trophic dynamics (Elton 1927, Lindeman 1942, Hutchinson 1948), the trophic cascade (Zaret and Paine 1973, Carpenter et al. 1985), and ecological stoichiometry (Tilman 1982, Sterner and Elser 2002), to the impacts of competition and predation in community structure (Hutchinson 1961, Hrbáek 1961, Brooks and Dodson 1965), to key features of species adaptations such as induced morphological and behavioral defenses (Gilbert 1967, Krueger and Dodson 1981, Dodson 1988), fundamental discoveries have flowed from the study of these relatively small lenses of freshwater resting on our planet’s crust. One reason for this is that lakes and ponds are superb natural laboratories for understanding how the ecological world functions. Like plants and animals on ocean islands, the populations and communities of organisms in lakes are more isolated from exchange with nearby habitats than are their terrestrial counterparts. The microbes, algae, plants and animals that live submersed beneath the water’s surface are nearly all restricted to their liquid environment, unable to cross the land that surrounds them. Thus confined, their interac- tions are often intense, easily discerned, and complex in fascinating ways. The diversity of genetic and species assemblages to be found among lakes and ponds, even within close proximity to each other, has provided exceptional opportunities for dissecting the forces that produce variation in nature. The thought that lakes and ponds are isolated from the surrounding landscape goes back at least to Forbes’ classic 1887 paper “The lake as a microcosm” (reprinted 1925). This independence, of course, should not be overstated (Likens 1984, Wetzel 1990): for example, the population genetic structure of even obligately aquatic organisms suggests occasional exchange of individuals, and there are certainly animal taxa that regularly move from lake to lake or between water and land. In addition, there are substantial subsidies of lake primary and secondary production by organic and inor- ganic chemicals flowing from the watershed (Pace et al. 2004). Indeed, the sediments of lakes store the historical record of a wide variety of ecological and evolutionary events that have taken place not only inside the confines of the lake proper (Kilham et al. 1986, Hairston et al. 1999), but also within its watershed and beyond (Davis ans Shaw 2001). Yet, as Forbes (1887) vi FOREWORD noted so long ago: “One finds in a single body of water a far more complete and independent equilibrium of organic life and activity than on any equal body of land.” The idea of a tight interdependence among the organisms resident in lakes that he described has remained both a significant paradigm and a useful foil for evolutionary biologists, ecologists and lim- nologists studying the adaptations, abundance, species composition and trophic interactions of lake ecosystems. A second reason underlying the central role of lakes in the formation of basic ecological understanding lies in the nature of water itself: it is a solvent for organic and inorganic molecules, a trap for heat, a filter for light, and a dense and viscous medium in which fluid movements large and small play a criti- cal role. As a result of this tight coupling between the physical, chemical and biological world in lakes, limnologists were among the first to take a whole- ecosystem approach to ecology. When August Thienemann (1925) tied lake basin shape, nutrient and oxygen concentrations, and organism abundance together in a single conceptual framework, he summarized the accumulated knowledge of a field that was already some four decades old (Forbes 1887, Forel 1892). This linkage between geological, chemical, physical and biologi- cal processes remains a hallmark of limnology. Christer Brönmark and Lars-Anders Hansson are ideal scientists to summa- rize the diversity of interactions among freshwater organisms and the aquatic context in which these links take place. Their research has been among the most illuminating in showing both the important and fascinating ways by which organisms from microscopic primary producers to large predatory fish detect and respond to each other, and how these interactions play out in the broader community and ecosystem context. The first edition of their book received well-deserved praise as a readable text for an undergraduate course and a useful summary of the ecological aspects of limnology for graduate students and instructors (Knight 2000, Kilham 2000). In this second edition, Brönmark and Hansson expand their coverage of population and commu- nity processes, link these to whole-ecosystem dynamics, and address a com- ment of the first edition by strengthening their coverage of the physical and chemical context in which the intra- and interspecific interactions that they expertly summarize take place. Each of these additions enhances the value of this book as an introduction first to the ecology of lakes, and second to lakes as model systems for studying general ecological processes. In addition, Brönmark and Hansson have expanded their final chapter on the challenges that human activities, intentional and unintended, have on lake and pond ecosystems. It is the basic ecological and limnological knowledge captured so well by the book you have in your hands that makes possible informed deci- sions about how to manage lakes and ponds. Nelson Hairston Jr. Department of Ecology and Evolutionary Biology, Cornell University Literature cited Brooks, J. L. and Dodson, S. I. (1965). Predation, body size, and composition of plankton. Science, 150, 28–35. Carpenter, S. R., Kitchell, J. F., and Hodgson, J. R. (1985). Cascading trophic inter- actions and lake productivity: Fish predation and herbivory can regulate lake ecosystems. Bioscience, 35, 634–9. Davis, M. B. and R. G. Shaw. 2001. Range shifts and adaptive responses to Quarternary climate change. Science, 292, 673–9. Dodson, S. I. (1988) The ecological role of chemical stimuli for zooplankton: preda- tor-avoidance behavior in Daphnia. Limnology and Oceanography, 33, 1431–9. Elton, C. (1927). Animal ecology. MacMillan Co., New York Forbes, S. (1925). The lake as a microcosm. Bulletin of the Illinois Natural History Survey, 15, 537–50.