Cambridge University Press
978-0-521-84942-5 - The Geometry of Evolution: Adaptive Landscapes and Theoretical Morphospaces - by George R. McGhee Jr.
Front Matter

The Geometry of Evolution


The metaphor of the adaptive landscape – that evolution via the process of natural selection can be visualized as a journey across adaptive hills and valleys, mountains and ravines – permeates both evolutionary biology and the philosophy of science. The focus of this book is to demonstrate to the reader that the adaptive landscape concept can be put into actual analytical practice through the usage of theoretical morphospaces – geometric spaces of both existent and nonexistent biological form – and to demonstrate the power of the adaptive landscape concept in understanding the process of evolution. The adaptive landscape concept further allows us to take a spatial approach to the concepts of natural selection, evolutionary constraint and evolutionary development. For that reason, this book relies heavily on spatial graphics to convey the concepts developed within these pages, and less so on formal mathematics.

GEORGE McGHEE is Professor of Paleobiology in the Department of Geological Sciences at Rutgers University, New Jersey, USA. He is a past Fellow of the Konrad Lorenz Institute for Evolution and Cognition Research in Vienna, Austria.





The Geometry of Evolution

Adaptive Landscapes and Theoretical Morphospaces

GEORGE R. McGHEE JR.
Rutgers University





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© Cambridge University Press 2007

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First published 2007

Printed in the United Kingdom at the University Press, Cambridge

A catalogue record for this publication is available from the British Library

Library of Congress Cataloguing-in-Publication Data

McGhee, George R.
The Geometry of evolution : adaptive landscapes and theoretical morphospaces / George R. McGhee, Jr.
p. cm.
Includes bibliographical references and index.
ISBN-13: 978-0-521-84942-5 (hardback)
ISBN-10: 0-521-84942-X (hardback)
1. Evolution (Biology)–Mathematical models. 2. Adaptation (Biology)–Mathematical models. 3. Morphology–Mathematical models. I. Title.

QH371.3.M37M34 2006
576.801′5118–dc22 2006026495


ISBN-13 978-0-521-84942-5 hardback
ISBN-10 0-521-84942-X hardback

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For Marae

A leannain m’òige, mo bhean.





Contents



Preface page xi
 
1 The concept of the adaptive landscape 1
What is an adaptive landscape? 1
Modelling evolution in adaptive landscapes 4
 
2 Modelling natural selection in adaptive landscapes 6
Visualizing natural selection 6
Modelling directional selection 7
Modelling stabilizing selection 10
Modelling disruptive selection 13
Modelling less-than-optimum evolution 16
Evolutionary topology of adaptive landscapes 19
Why has evolution not ceased? 22
Modelling environmental and ecological change 24
 
3 Modelling evolutionary phenomena in adaptive landscapes31
Evolution in geological time 31
Modelling convergent evolution 32
Modelling iterative evolution 36
Modelling speciation by cladogenesis 38
Modelling hyperdimensionality in adaptive landscapes 45
Are adaptive landscapes of heuristic value only? 51
 
4 The concept of the theoretical morphospace 57
What is a theoretical morphospace? 57
Procedural phases of theoretical morphospace analyses 60
Creating theoretical morphospaces: an example 63
 
5 Analysing the role of adaptive evolution in theoretical morphospaces 71
Functional analysis in theoretical morphospace 71
Adaptive significance of existing and nonexistent ammonoid form 72
Adaptive significance of existing and nonexistent brachiopod form 77
Adaptive significance of existing and nonexistent plant form 82
 
6 Analysing evolutionary phenomena in theoretical morphospaces 90
Analysing evolution in geological time 90
Analysing convergent evolution in theoretical morphospace 90
Analysing iterative evolution in theoretical morphospace 96
Analysing biodiversity crises in theoretical morphospace 100
 
7 Evolutionary constraint in theoretical morphospace 108
Potential causes of empty morphospace 108
Modelling geometric constraint 109
Modelling functional constraint 111
Modelling phylogenetic constraint 112
Modelling developmental constraint 114
Analysing evolutionary constraint in theoretical morphospaces 117
Mapping geometric constraint boundaries 118
Mapping functional constraint boundaries 124
Analysing intrinsic constraints 127
Modelling the evolution of intrinsic constraints 134
Evolutionary constraint and the phenomenon of convergent evolution 144
Is the concept of phylogenetic constraint of heuristic value only? 150
 
8 Evolutionary development in theoretical morphospace 152
The concept of developmental constraint 152
Transformation theory and theoretical morphospaces 155
Epigenetic landscapes and theoretical morphospaces 159
Analysing development in theoretical morphospace 163
Theoretical developmental morphospaces? 167
How does an organism come to be? 172
 
9 There is much to be done … 174
Adaptive landscapes and theoretical morphospaces 174
Current progress in theoretical morphospace analyses 175
What more is to be done? 177
The re-emergence of the science of morphology 180
 
References 185
Index 197




Preface



The concept of the adaptive landscape is the creation of the great American geneticist Sewall Wright who, along with the equally great British scientists R. A. Fisher and J. B. S. Haldane, crafted the Neo-Darwinian synthesis of evolutionary theory in the 1930s. The metaphor of the adaptive landscape, that evolution via the process of natural selection could be visualized as a journey across adaptive hills and valleys, mountains and ravines, permeated both evolutionary biology and the philosophy of science through the succeeding years of the twentieth century. Yet critics of the adaptive landscape concept have maintained that the concept is of heuristic value only; that is, it is fine for creating conceptual models, but that you cannot actually use the concept in analysing the evolution of actual animals or plants. That criticism became invalid in the year 1966 when the palaeontologist David M. Raup used computer simulations to model hypothetical life forms that have never existed in the evolution of life on Earth, and who subsequently created the concept of the theoretical morphospace.

The focus of this book is to demonstrate to the reader the power of the adaptive landscape concept in understanding the process of evolution, and to demonstrate that the adaptive landscape concept can be put into actual analytical practice through the usage of theoretical morphospaces. The adaptive landscape concept allows us to visualize the possible effects of natural selection through simple spatial relationships, rather than complicated modelling of changing environmental or ecological conditions. For that reason, this book relies heavily on spatial graphics to convey the concepts developed within these pages, and less so on formal mathematics.

I thank the Santa Fe Institute for the invitation to visit and work on computational methods in theoretical morphology in 2000, for it was at the Santa Fe Institute that the idea of writing this book came to me in conversations with Dave Raup. I thank the Konrad Lorenz Institute for Evolution and Cognition Research for the Fellowship that enabled me to work at the institute in 2005, for it was there that I developed many of the ideas presented in Chapters 7 and 8 of this book. Finally, I thank my wife, Marae, for her patient love.


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