The Evolutionary Strategies that Shape Ecosystems
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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 reflects 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.

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Philip Grime is a Professor Emeritus at the University of Sheffield where he currently maintains long-term experiments at the Buxton Climate Change Impacts Laboratory in North Derbyshire. As a pioneer of experimental approaches to communities and ecosystems Professor Grime is an elected member of the Dutch and British Royal Societies and was the inaugural recipient in 2011 of the Alexander von Humboldt Medal awarded by the International Association for Vegetation Science.

Simon Pierce is a researcher and lecturer at the University of Milan, Italy, and at the time of writing taught plant physiological ecology at the University of Insubria, Varese, Italy. His research encompasses plant community ecology and ecophysiology, and the reproductive biology, cultivation and conservation of terrestrial orchids. During his career he has lived and worked in the Republic of Panama, as an Andrew W. Mellon research fellow at the Smithsonian Tropical Research Institute, for the University of Cambridge, UK. He holds a doctorate from the University of Durham, UK, and a degree from the University of Wales, Bangor.


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


“In summary, The Evolutionary Strategies that Shape Ecosystemsis well-written and stimulating, and encourages its readers to think about how all the pieces of ecology might fit together, from the scale of an individual organism to entire ecosystems. It would make a valuable addition to the library of any scientist interested in ecological and evolutionary strategies.”  (Austral Ecology, 1 October 2013)

“Certainly I have found this a useful way to think about conservation Management.”  (British Ecological Society, 1 April 2013)

“The case studies range from microbes to animals, and even palaeontology is included in the mix, making the book a very comprehensive resource for those interested in eco-evolutionary dynamics.”  (Teaching Biology, 20 December 2012)

“I recommend this book to people interested in evolutionary and ecological strategies in ecosystems, to those who think about universal patterns in organism life history tactics and also to those who love the challenge of linking ecology and evolution.”  (Basic and Applied Ecology, 1 November 2012)

“A significant contribution to the field and a must read for ecologists.  Summing Up:  Highly recommended.  Upper-division undergraduates and above.”  (Choice, 1 October 2012)