Front Matter

Cambridge University Press 978-0-521-88318-4 - Speciation and Patterns of Diversity Edited by Roger K. Butlin, Jon R. Bridle and Dolph Schluter Frontmatter More information

Speciation and Patterns of Diversity

Bringing together the viewpoints of leading ecologists concerned with the processes that generate patterns of diversity, and evolutionary biologists who focus on mechanisms of speciation, this book opens up discussion in order to broaden understanding of how speciation affects patterns of biological diversity, especially the uneven distribution of diversity across time, space and taxa studied by macroecologists. The contributors discuss questions such as: Are species equivalent units, providing meaningful measures of diversity? To what extent do mechanisms of speciation affect the functional nature and distribution of species diversity? How can speciation rates be measured using molecular phylogenies or data from the fossil record? What are the factors that explain variation in rates? Written for graduate students and academic researchers, the book promotes a more complete understanding of the interaction between mechanisms and rates of speciation and these patterns in biological diversity.

R OGER B UTLIN is Professor of Evolutionary Biology, Animal and Plant Sciences, at the University of Sheffield. He has held a prestigious Royal Society Research Fellowship at the University of Cardiff and his work has been recognized by honorary fellowships at the Natural History Museum, Zoological Society of London, and Royal Belgian Institute of Natural Sciences.

J ON B RIDLE is Lecturer in Biology, in the School of Biological Sciences, University of Bristol. Since completing his PhD in Evolutionary Genetics in 1998, he has conducted research in quantitative genetics and evolutionary biology at University College London, University of Cardiff, Universidad Auto´ noma de Madrid, and the Institute of Zoology, London. He was awarded a fellowship at the Zoological Society of London in 2002.

D OLPH S CHLUTER is Professor and Canada Research Chair, Biodiversity Research Centre and Zoology Department, at the University of British Columbia. He is a former President of the Society for the Study of Evolution and recipient of the Sewall Wright Award from the American Society of Naturalists. He is a Fellow of the Royal Society of London and of Canada.

© Cambridge University Press www.cambridge.org Cambridge University Press 978-0-521-88318-4 - Speciation and Patterns of Diversity Edited by Roger K. Butlin, Jon R. Bridle and Dolph Schluter Frontmatter More information

Ecological Reviews series editor Hefin Jones Cardiff University, UK series editorial board Mark Bradford University of Georgia, USA Jim Brown University of New Mexico, USA David Burslem University of Aberdeen, UK Lindsay Haddon British Ecological Society, UK Sue Hartley University of Sussex, UK Richard Hobbs Murdoch University, Australia Mark Hunter University of Michigan, USA Heikki Setala University of Helsinki, Finland Louise Vet Wageningen Agricultural University, The Netherlands

Ecological Reviews will publish books at the cutting edge of modern ecology, providing a forum for volumes that discuss topics that are focal points of current activity and are likely to be of long-term importance to the progress of the field. The series will be an invaluable source of ideas and inspiration for ecologists at all levels from graduate students to more-established researchers and professionals. The series will be developed jointly by the British Ecological Society and Cambridge University Press and will encompass the Society’s Symposia as appropriate.

Biotic Interactions in the Tropics: Their Role in the Maintenance of Species Diversity Edited by David F. R. P. Burslem, Michelle A. Pinard and Sue E. Hartley Biological Diversity and Function in Soils Edited by Richard Bardgett, Michael Usher and David Hopkins

Island Colonization: The Origin and Development of Island Communities by Ian Thornton Edited by Tim New

Scaling Biodiversity Edited by David Storch, Pablo Margnet and James Brown

Body Size: The Structure and Function of Aquatic Ecosystems Edited by Alan G. Hildrew, David G. Raffaelli and Ronni Edmonds-Brown.

Speciation and Patterns of Diversity

Edited by

ROGER K. BUTLIN University of Sheffield

JON R. BRIDLE University of Bristol

DOLPH SCHLUTER University of British Columbia

cambridge university press Cambridge, New York, Melbourne, Madrid, Cape Town, Singapore, Sa˜o Paulo, Delhi Cambridge University Press The Edinburgh Building, Cambridge CB2 8RU, UK

Published in the United States of America by Cambridge University Press, New York

www.cambridge.org Information on this title: www.cambridge.org / 9780521883184

This publication is in copyright. Subject to statutory exception and to the provisions of relevant collective licensing agreements, no reproduction of any part may take place without the written permission of Cambridge University Press.

First published 2009

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 Speciation and patterns of diversity / edited by Roger Butlin, Jon Bridle and Dolph Schluter. p. cm. — (Ecological reviews) Includes bibliographical references. 1. Species diversity. 2. Biodiversity. I. Butlin, Roger, 1955– II. Bridle, Jon. III. Schluter, Dolph. IV. Series. QH541.15.S64S62 2008 577—dc22 2008025507

ISBN 978-0-521-88318-4 hardback ISBN 978-0-521-70963-7 paperback

Cambridge University Press has no responsibility for the persistence or accuracy of URLs for external or third-party internet websites referred to in this publication, and does not guarantee that any content on such websites is, or will remain, accurate or appropriate.

Contents

List of contributors page vii Preface ix Acknowledgements xi

1 Speciation and patterns of biodiversity Roger K. Butlin, Jon R. Bridle and Dolph Schluter 1 2 On the arbitrary identification of real species Jody Hey 15 3 The evolutionary nature of diversification in sexuals and asexuals Timothy G. Barraclough, Diego Fontaneto, Elisabeth A. Herniou and Claudia Ricci 29 4 The poverty of the protists Graham Bell 46 5 Theory, community assembly, diversity and evolution in the microbial world Thomas P. Curtis, Nigel C. Wallbridge and William T. Sloan 59 6 Limits to adaptation and patterns of biodiversity Jon R. Bridle, Jitka Polechova´ and Tim H. Vines 77 7 Dynamic patterns of adaptive radiation: evolution of mating preferences Sergey Gavrilets and Aaron Vose 102 8 Niche dimensionality and ecological speciation Patrik Nosil and Luke Harmon 127 9 Progressive levels of trait divergence along a ‘speciation transect’ in the Lake Victoria cichlid fish Pundamilia Ole Seehausen 155 10 Rapid speciation, hybridization and adaptive radiation in the Heliconius melpomene group James Mallet 177

vi CONTENTS

11 Investigating ecological speciation Daniel J. Funk 195 12 Biotic interactions and speciation in the tropics Douglas W. Schemske 219 13 Ecological influences on the temporal pattern of speciation Albert B. Phillimore and Trevor D. Price 240 14 Speciation, extinction and diversity Robert E. Ricklefs 257 15 Temporal patterns in diversification rates Andy Purvis, C. David L. Orme, Nicola H. Toomey and Paul N. Pearson 278 16 Speciation and extinction in the fossil record of North American mammals John Alroy 301

Index 324

The colour plate section is placed between pages 180 and 181

Contributors

JOHN ALROY SERGEY GAVRILETS The Paleobiology Database, University of Department of Ecology and Evolutionary California Biology and Department of Mathematics, University of Tennessee TIMOTHY G. BARRACLOUGH Division of Biology, Imperial College LUKE HARMON London Department of Biological Sciences, University of Idaho GRAHAM BELL Biology Department, McGill University ELISABETH A. HERNIOU Division of Biology, Imperial College London JON R. BRIDLE School of Biological Sciences, University of JODY HEY Bristol Department of Genetics, Rutgers, the State University of New Jersey ROGER K. BUTLIN Animal and Plant Sciences, University of JAMES MALLET Sheffield Galton Laboratory, Department of Biology, University College London THOMAS P. CURTIS School of Civil Engineering and PATRIK NOSIL Geosciences, University of Newcastle upon Zoology Department and Biodiversity Tyne Research Centre, University of British Columbia DIEGO FONTANETO Division of Biology, Imperial College C. DAVID L. ORME London Division of Biology, Imperial College London

DANIEL J. FUNK PAUL N. PEARSON Department of Biological Sciences, School of Earth, Ocean and Planetary Vanderbilt University Sciences, Cardiff University

viii LIST OF CONTRIBUTORS

ALBERT B. PHILLIMORE DOLPH SCHLUTER NERC Centre for Population Biology and Zoology Department and Biodiversity Division of Biology, Imperial College Research Centre, University of British London Columbia

JITKA POLECHOVA´ OLE SEEHAUSEN Biomathematics & Statistics Scotland Centre of Ecology, Evolution and Biogeochemistry, EAWAG TREVOR D. PRICE Department of Ecology and Evolution, WILLIAM T. SLOAN University of Chicago Department of Civil Engineering, University of Glasgow ANDY PURVIS Division of Biology, Imperial College NICOLA H. TOOMEY London Division of Biology, Imperial College London CLAUDIA RICCI Universita` di Milano, Dipartimento di TIM H. VINES Biologia Centre d’Ecologie Fonctionelle et Evolutive Montpellier ROBERT E. RICKLEFS Department of Biology, University of AARON VOSE Missouri – St. Louis Department of Ecology and Evolutionary Biology and Department of Computer DOUGLAS W. SCHEMSKE Science, University of Tennessee Department of Plant Biology and W. K. Kellogg Biological Station, Michigan State NIGEL C. WALLBRIDGE University Nomad Digital Limited

Preface

This volume is derived from the Annual Symposium of the British Ecological Society on ‘Speciation and Ecology’ which was held at the University of Sheffield, 28–30 March 2007. The idea for this Symposium arose during a previous meeting in the series, the 2002 ‘Macroecology: Concepts and Consequences’ meeting organized by Tim Blackburn and Kevin Gaston. The 2002 meeting concentrated on large-scale diversity patterns. Many speakers acknowledged the role of speciation in generating diversity and influencing patterns of diversity. Although there was some discussion of the factors that determine rates of speciation, it was striking how little contact there seemed to be between the discipline of macroecology and the large and active field of research into mechanisms of adaptive divergence and speciation. ‘Ecological speciation’ has been an area of research growth in recent years, asking how ecological drivers influence the speciation process. However, the opposite direc- tion of effect, how speciation processes impact on ecological patterns, has been studied less. Therefore, we proposed a meeting whose central objective was to foster dialogue between these two fields. The meeting had an unusual mix of participants but we hope that they managed to communicate effectively with one another! The chapters in this book reflect the range of topics discussed and we hope that they will help to continue the conversations that were started in Sheffield. In our introduction, we try to set the scene by considering mechanisms of speciation and their potential impacts on biodiversity, both in terms of species’ geographical distri- butions, and their interactions within ecological communities. In particular, speciation mechanisms can be divided into those that generate ecologically distinct species and those that do not, for example because reproductive iso- lation is driven by sexual conflict. These two classes of speciation mechanism may contribute to diversity in different ways, adding to either local diversity or beta diversity, for example, or affecting levels of within-species functional diversity. We also introduce the problems surrounding the estimation of speciation rates, an essential preliminary to understanding the factors that influence those rates.

x PREFACE

We have not subdivided the book because we are reluctant to separate parts of a spectrum. However, the chapters are ordered in a progression, similar to that used at the Symposium, which begins with the nature of species, how we count the units of biological diversity and some of the limits on diversity; considers mechanisms of speciation and adaptation; and then looks at rates of speciation both from the perspective of molecular phylogenetics and that of the fossil record. We hope that readers will see some logic in this arrangement and will find the book stimulating and enjoyable.

Acknowledgements

We are very grateful to the British Ecological Society for the opportunity to organize the Symposium on ‘Speciation and Ecology’ and especially to Hefin Jones, Hazel Norman and Richard English for their excellent support in the organizational stages and at the meeting. The Department of Animal and Plant Sciences and the Conference Office of the University of Sheffield also helped to make the meeting a success. As with any meeting, the interest and excitement of the event was dependent on the contributors: we thank those who gave oral presentations, those who brought posters and all participants for making the meeting go so well. During the preparation of this volume, our contributors have done a great job of providing stimulating chapters more or less on schedule, as well as providing very valuable assistance with the review process, and responding constructively to reviewers’ comments. We have had strong support both from the British Ecological Society, Hefin Jones and Lindsay Haddon, and from Cambridge University Press, Dominic Lewis and Alison Evans. Alan Crowden was also a great help in the early stages.