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The Evolutionary Strategies that Shape Ecosystems Companion Website This book has a companion website: www.wiley.com/go/grime/evolutionarystrategies with Figures and Tables from the book for downloading The Evolutionary Strategies that Shape Ecosystems J. Philip Grime FRS Department of Animal and Plant Sciences University of Sheffi eld, UK Simon Pierce Department of Plant Production, University of Milan, Italy A John Wiley & Sons, Ltd., Publication This edition fi rst published 2012 © 2012 by John Wiley & Sons, Ltd. Wiley-Blackwell is an imprint of John Wiley & Sons, formed by the merger of Wiley’s global Scientifi c, Technical and Medical business with Blackwell Publishing. 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Designations used by companies to distinguish their products are often claimed as trademarks. All brand names and product names used in this book are trade names, service marks, trademarks or registered trademarks of their respective owners. The publisher is not associated with any product or vendor mentioned in this book. This publication is designed to provide accurate and authoritative information in regard to the subject matter covered. It is sold on the understanding that the publisher is not engaged in rendering professional services. If professional advice or other expert assistance is required, the services of a competent professional should be sought. Library of Congress Cataloging-in-Publication Data Grime, J. Philip (John Philip) The evolutionary strategies that shape ecosystems / J. Philip Grime, FRS Simon Pierce. p. cm. Includes index. ISBN 978-0-470-67481-9 (cloth) – ISBN 978-0-470-67482-6 (pbk.) 1. Biotic communities. 2. Evolution (Biology) 3. Adaptation (Biology) I. Pierce, Simon. II. Title. QH541.G75 2012 577.8'2–dc23 2011045522 A catalogue record for this book is available from the British Library. Wiley also publishes its books in a variety of electronic formats. Some content that appears in print may not be available in electronic book. Set in 10.5/12 pt Classical Garamond BT by Toppan Best-set Premedia Limited 1 2012 Nothing in biology makes sense except in the light of evolution (Dobzhansky, 1973). Nothing in evolutionary biology makes sense except in the light of ecology (Grant & Grant, 2008). Nothing in evolution or ecology makes sense except in the light of the other (Pelletier, Garant & Hendry, 2009). Contents Preface x Chapter Summaries xii Acknowledgements xviii Introduction 1 1 Evolution and Ecology: a Janus Perspective? 3 Evolutionary biology 3 Ecology 4 The emergence of a science of adaptive strategies 6 Summary 7 2 Primary Strategies: the Ideas 8 MacArthur’s ‘blurred vision’ 9 The mechanism of convergence; trade-offs 10 The theory of r- and K-selection 11 CSR Theory 12 Summary 23 3 Primary Adaptive Strategies in Plants 25 The search for adaptive strategies 26 Theoretical work 26 Measuring variation in plant traits: screening programmes 28 Screening of plant growth rates 29 The Integrated Screening Programme 29 viii Contents Further trait screening 34 The application of CSR theory 34 Virtual plant strategies 36 Summary 38 4 Primary Adaptive Strategies in Organisms Other Than Plants 40 The architecture of the tree of life 41 r, K and beyond K 42 Empirical evidence for three primary strategies in animals 43 The universal three-way trade-off 44 Mammalia (mammals) 46 Aves (avian therapods) 53 Squamata (snakes and lizards) (with notes on other extant reptile clades) 56 Amphibia (amphibians) 60 Osteichthyes (bony fi shes) 61 Chondrichthyes (cartilaginous fi shes) 65 Insecta (insects) 68 Aracnida (spiders, scorpions, mites and ticks) 72 Crustacea (crustaceans) 74 Echinodermata (sea urchins, starfi sh, crinoids, sea cucumbers) 75 Mollusca (snails, clams, squids) 77 Annelida (segmented worms) 79 Cnidaria (corals, sea anemones, jellyfi sh, hydras, sea pens) 81 Eumycota (fungi) (including notes on lichens) 83 Archaea 84 Proteobacteria 86 Firmicutes 87 Cyanobacteria 88 Viruses 90 Extinct groups 94 Universal adaptive strategy theory – the evolution of CSR and beyond K theories 99 First steps towards a universal methodology 100 Summary 103 5 From Adaptive Strategies to Communities 105 Plant communities 106 Productive disturbed communities 107 Productive undisturbed communities 108 Unproductive relatively undisturbed communities 111 Plant community composition 111 Contents ix The humped-back model 114 Origins 114 Formulation 115 Independent confi rmation and compatibility with new research 116 Species-pools, fi lters and community composition 121 Evidence for the action of twin fi lters 128 Additional mechanisms promoting diversity 132 Genetic diversity, intraspecifi c functional diversity and species diversity 132 Microbial communities 136 The effects of plant strategies on soil microbial communities 139 Facilitation in bacterial communities 141 Coexistence in marine surface waters 142 Novel techniques for investigating microbial adaptive strategies 142 Animal communities 144 Primary producers delimit animal diversity/productivity relationships 145 Twin fi lters and animal community assembly 150 Adaptive radiation and community assembly 154 Summary 160 6 From Strategies to Ecosystems 163 Back to Bayreuth 164 The Darwinian basis of ecosystem assembly 167 How do primary adaptive strategies drive ecosystem functioning? 168 The plant traits that drive ecosystems 169 The propagation of trait infl uences through food chains 176 Complicating factors 178 Ecosystem processes 180 Dominance and mass ratio effects 180 Fluxes and feedbacks between communities 181 Top-down control by herbivores 187 Top-down control by carnivores 189 The key role of eco-evolutionary dynamics 190 Summary 192 7 The Path from Evolution to Ecology 194 What has been learned? 194 What are the implications for conservation and management? 198 Research priorities for the next decade 199 References 202 Organism Index 235 Subject Index 241 Preface One of the greatest achievements of biology in the 20th century, carried forward with vigour into the 21st, has been to translate the theory of natural selection into a practical reductionist science. This development, complemented by the ‘molecular revolution ’ ensuing from the cracking of the genetic code, has immense potential for our understanding of the evolutionary relationships between organisms and the basis of variation within and between populations. For many ecologists these advances have provided opportunities to expand and refi ne investigations of the population genetics of individual species. However, this has to a large extent left unattended the urgent requirement for ecologists to bring order and predictive science to the study of ecosystems, many of which are suffering degradation and extinction in an over-exploited and rapidly chang- ing world. In established sciences, including physics and chemistry, the revela- tions that eventually led to general theories were surprisingly discrete and testable. We believe that ecology, despite its reputation as an ‘ambitious but ramshackle science ’ (Calder, 2003), is no exception to this pattern, and that it is now opportune to attempt integration of the disparate fi elds of animal, plant and microbial ecology within a single theoretical framework that keeps natural selection at the fore. To address this need, this short book endeavours to understand how evolu- tionary adaptations govern the manner in which organisms assemble into com- munities and process matter and energy within ecosystems. We fi rst chart the tentative steps by which some ecologists over the course of more than a century have attempted to establish a path from evolutionary theory to an understanding of variation in the structure, functioning and vulnerabilities of ecosystems. In marked contrast to the technical diffi culties of microbial ecology and the limited insights into general principles gained by animal ecology, the accessibility of plants and their dominance of terrestrial ecosystems have revealed generalities in the constraints to evolution. These, in light of recent research, are now Preface xi apparent for all organisms, from the largest whale to the smallest virus. We have distilled the essence of these constraints to evolution and ecology into a novel universal adaptive strategy theory, in which only a restricted set of viable approaches to survival, or adaptive strategies, are possible and in turn set limits to the performance of ecosystem functions. We explain, in terms of a twin-fi lter model, how it is possible to distinguish between mechanisms of natural selection that control the functioning of ecosystems and those that merely determine which