The Evolutionary Strategies that Shape Ecosystems

THE EVOLUTIONARY STRATEGIES THAT SHAPE ECOSYSTEMS In 1837 a young Charles Darwin took his notebook, wrote “I think”, and then sketched a rudimentary, stick-like tree. Each branch of Darwin’s tree of life told a story of survival and adaptation – adaptation of animals and plants not just to the environment but also to life with other living things. However, more than 150 years since Darwin published his singular idea of natural selection, the science of ecology has yet to account for how contrasting evolutionary outcomes affect the ability of organisms to coexist in communities and to regulate ecosystem functioning. In this book Philip Grime and Simon Pierce explain how evidence from across the world is revealing that, beneath the wealth of apparently limitless and bewildering variation in detailed structure and functioning, the essential biology of all organisms is subject to the same set of basic interacting constraints on life-history and physiology. The inescapable resulting predicament during the evolution of every species is that, according to habitat, each must adopt a predictable compromise with regard to how they use the resources at their disposal in order to survive. The compromise involves the investment of resources in either the effort to acquire more resources, the tolerance of factors that reduce metabolic performance, or reproduction. This three-way trade-off is the irreducible core of the universal adaptive strategy theory which Grime and Pierce use to investigate how two environmental filters selecting, respectively, for convergence and divergence in organism function determine the identity of organisms in communities, and ultimately how different evolutionary strategies affect the functioning of ecosystems. This book refl ects an historic phase in which evolutionary processes are finally moving centre stage in the effort to unify ecological theory, and animal, plant and microbial ecology have begun to find a common theoretical framework. Companion website This book has a companion website www.wiley.com/go/grime/evolutionarystrategies with Figures and Tables from the book for downloading.

Table of 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 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 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