What Is Biodiversity?

What Is Biodiversity?

What Is Biodiversity? James Maclaurin and Kim Sterelny The University of Chicago Press chicago and london

james maclaurin is a senior lecturer in the Department of Philosophy at the University of Otago, New Zealand, and has also been a Marsden Post Doctoral Fellow at Victoria University. He is the author of numerous articles published in professional journals. kim sterelny divides his time between Victoria University of Wellington, where he is a professor of philosophy, and the Research School of Social Sciences and the Centre for Macroevolution and Macroecology at the Australian National University. He is the editor of the journal Biology and Philosophy, and his books include Evolution of Agency and Other Essays; Thought in a Hostile World; Dawkins vs. Gould; and, with Paul Griffiths, Sex and Death: An Introduction to Philosophy of Biology, which is published by the University of Chicago Press. The University of Chicago Press, Chicago 60637 The University of Chicago Press, Ltd., London 2008 by The University of Chicago All rights reserved. Published 2008 Printed in the United States of America 17 16 15 14 13 12 11 10 09 08 1 2 3 4 5 isbn-13: 978-0-226-50080-5 (cloth) isbn-13: 978-0-226-50081-2 (paper) isbn-10: 0-226-50080-2 (cloth) isbn-10: 0-226-50081-0 (paper) Library of Congress Cataloging-in-Publication Data Maclaurin, James. What is biodiversity? / James Maclaurin and Kim Sterelny. p. cm. Includes bibliographical references and index. isbn-13: 978-0-226-50080-5 (hardcover : alk. paper) isbn 10: 0-226-50080-2 (hardcover : alk. paper) isbn-13: 978-0-226-50081-2 (pbk. : alk. paper) isbn-10: 0-226-50081-0 (pbk. : alk. paper) 1. Biodiversity. I. Sterelny, Kim. II. Title. QH541.15.B56M325 2008 333.95 dc22 o The paper used in this publication meets the minimum requirements of the American National Standard for Information Sciences Permanence of Paper for Printed Library Materials, ansi z 39.48 1992.

For Kristen and George and Melanie and Kate

Contents Acknowledgments xi 1 Taxonomy Red in Tooth and Claw 1 1.1 Biodiversity and Biodiversity 1.2 Biodiversity and Biodiversities 1.3 History and Taxonomy 1.4 Diversity as Cause; Diversity as Effect 1.5 Prospectus: The Road Ahead 2 Species: A Modest Proposal 27 2.1 Introduction 2.2 Species, Species Concepts, and Speciation 2.3 The Effect of Speciation 2.4 Species and Biodiversity 3 Disparity and Diversity 42 3.1 The Cone of Increasing Controversy 3.2 How Disparate Was the Cambrian Fauna? 3.3 Fossils in a Molecular World 4 Morphology and Morphological Diversity 60 4.1 Introduction 4.2 Morphological Diversity 4.3 Biological Possibility Spaces

viii contents 4.4 The Power of Morphospaces 4.5 Here There Be No Dragons: The Limits of Theoretical Morphology 4.6 Morphological Biodiversity 5 Development and Diversity 84 5.1 Diversity, Disparity, Plasticity 5.2 The Variety of Developmental Resources 5.3 From Gene Regulation to Modularity 5.4 Modularity in Development and Evolution 5.5 Developmental Biodiversity 6 Explorations in Ecospace 106 6.1 Ecological Systems 6.2 Communities, Ecosystems, and Ecosystem Functions 6.3 Individualism and Community Regulation 6.4 The Emergent Property Hypothesis 6.5 Boundaries 6.6 The Space of Population Assemblages 7 Conservation Biology: The Measurement Problem 132 7.1 Introduction 7.2 Counting Taxa 7.3 Measuring Phylogenetic Diversity 7.4 Measuring Genetic Diversity 7.5 Biodiversity Surrogates 8 Conservation Biology: The Evaluation Problem 149 8.1 Value 8.2 Is Biodiversity Intrinsically Valuable? 8.3 Demand Value 8.4 The Option Value Option 8.5 Applying Option Value: Case 1, Phylogeny 8.6 Applying Option Value: Case 2, Bioprospecting 8.7 Applying Option Value: Case 3, Ecological Option Value 8.8 The Conservation Consequences of Option Value Models

9 Concluding Remarks 172 9.1 Introduction: The Temptations of a Unified Measure 9.2 The Variety of Diversities 9.3 Should We Conserve Species? Notes 179 References 186 Index 207 Contents ix

Acknowledgments This book is the product of a Marsden Fund grant. The full-time focus the grant made possible kick-started the project. Even so, it has been a long project, and we have received help from many quarters. Our special thanks go to Ben Jeffares and Russell Brown for dedicated research assistance during the early phase of the project. James Maclaurin would like to thank Alan Musgrave for being extraordinarily helpful and supportive and also the Department of Philosophy at the University of Otago, which continues to be a stimulating and enjoyable place to think. Kim Sterelny would like to thank his two academic homes, the philosophy programs at Victoria University of Wellington and the RSSS, ANU. Both continue to be collegial and supportive environments in which to do empirically oriented philosophy. Many people were happy to be used as sounding boards and provided useful feedback. These include Nick Agar, Peter Anstey, Jochen Brocks, David Braddon-Mitchell, Lindell Bromham, Brett Calcott, David Chalmers, Geoff Chambers, James Chase, Colin Cheyne, Mark Colyvan, Tim Dare, Kath Dickinson, Steve Downes, Heather Dyke, Patrick Forber, Peter Godfrey Smith, Todd Grantham, Paul Griffiths, Mike Hannah, Frank Jackson, Ben Jeffares, Richard Joyce, John Matthewson, George McGhee, Andrew Moore, Karen Neander, Daniel Nolan, Samir Okasha, Charles Pigden, Josh Parsons, Gerhard Schlosser, Nick Shea, and Daniel Stoljar. Secretarial support at Otago has been ably provided by Sally Holloway and Kate Anscombe. The index was prepared by Meg Cordes. We reserve special thanks for Christie Henry at the University of Chicago Press, who has been unflaggingly helpful, and for our excellent copy editor, Dawn Hall.

xii acknowledgments Kim thanks his partner Melanie for her unfailing support of his research efforts despite her own intense research commitments, and his daughter Kate for making life much easier through her cheerfulness and goodwill. James would like to thank his wife Kristen and son George for their love and inspiration, and he extends grateful thanks to his coauthor, a mentor and friend without whom this book would not have happened.