Animal Dispersal. Small mammals as a model. WILLIAM Z. LIDICKER, JR Museum of Vertebrate Zoology, University of California, Berkeley, USA

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1 Animal Dispersal

2 Animal Dispersal Small mammals as a model Edited by NILS CHR. STENSETH Department of Biology, University of Oslo, Norway and WILLIAM Z. LIDICKER, JR Museum of Vertebrate Zoology, University of California, Berkeley, USA SPRINGER-SCIENCE+BUSINESS MEDIA, B.V.

3 First edition Springer Science+Business Media Dordrecht Originally published by Chapman & Hali in 1992 Softcover reprint of the hardcover 1 st edition 1992 Typeset in 10112pt Sabon by Graphicraft Typesetters Ltd, Hong Kong Apart from any fair dealing for the purposes of research or private study, or criticism or review, as permitted under the UK Copyright Designs and Patents Act, 1988, this publication may not be reproduced, stored, or transmitted, in any form or by any means, without the priof permission in writing of the publishers, or in the case of reprographic reproduction only in accordance with the terms of the licences issued by the Copyright Licensing Agency in the UK, or in accordance with the terms of licences issued by the appropriate Reproduction Rights Organization outside the UK. Enquiries concerning reproduction outside the terms stated here should be sent to the publishers at the London address printed on this page. The publisher makes no representation, express or implied, with regard to the accuracy of the information contained in this book and cannot accept any legal responsibility or liability for any errors or omissions that may be made. A catalogue record for this book is available from the British Library Library of Congress Cataloging-in-PublicatiOl. data Animal dispersal: small mammals as a model/edited by Nils Chr. Stenseth and William Z. Lidicker, Jr. p. cm. Includes bibliographical references and index. ISBN ISBN (ebook) DOI / Animals-Dispersal. 2. Mammals-Dispersal. 1. Stenseth, Nils ehr. II. Lidicker, William Zander, QH543.3.A dc CIP

4 Contents Contributors Acknowledgements Preface Part One. Introductory chapters 1 The study of dispersal: a conceptual guide N.C. Stenseth and W.Z. Lidicker, Jr 1.1 Introduction 1.2 Terminology 1.3 Dispersal or residency 1.4 A matter of scale 1.5 Dispersal: a conceptual 'glue' 1.6 The book x xi Xll To disperse or not to disperse: who does it and why? 21 W.Z. Lidicker, Jr and N.C. Stenseth 2.1 Introduction Who are the dispersers? What motivates dispersal? A classification of dispersal Conclusions The genetic consequences of dispersal 37 N.H. Barton 3.1 Introduction Dispersal, environmental heterogeneity, and genetic variation 3.3 Gene flow and spatial differentiation 3.4 Inference from genetic patterns 3.5 Conclusions

5 VI Contents Part Two. The process of dispersal 4. Habitat heterogeneity and dispersal: environmental and genetic patchiness A. Cockburn 4.1 Introduction 4.2 The adaptive basis of dispersal 4.3 Are dispersers genetically distinct? 4.4 Conclusions Social Factors in immigration and emigration C.A. Brandt 5.1 Introduction 5.2 Aggression, dominance, and spacing 5.3 Lack of social bonding 5.4 Mate access and mate choice 5.5 Social consequences of dispersal 5.6 Conclusions Part Three. Patterns: cyclic versus stable populations Patterns of dispersal in non-cyclic populations of small rodents 147 ]. Gliwicz 6.1 Introduction Patterns of seasonal dispersal Why do rodents disperse early in the breeding season? Why do rodents disperse in the autumn? Spring-summer dispersal, spatial heterogeneity, and population cycles Conclusions The role of dispersal in cyclic rodent populations 160 C.]. Krebs 7.1 Introduction Immigration and emigration are always area-dependent Dispersal can affect a population directly or indirectly

6 Contents Vll Part Four. Applications Small mammal dispersal in pest management and conservation 181 L. Hansson 8.1 Introduction Population eradication Damage related to dispersal from permanent habitats Pest outbreaks Settling in human habitations Disease transmission Models in conservation theory Discussion and generalizations Conclusions Part Five. Looking backward and forward Presaturation and saturation dispersal 15 years later: some theoretical considerations 201 N.C. Stenseth and W.Z. Lidicker, Jr 9.1 Introduction The evolution of dispersal Population dynamic consequences of dispersal 215 9:4 Presaturation and saturation dispersal 15 years later Part Six. Invited commentaries Do large mammals disperse like small mammals? 229 A.R.E. Sinclair 10.1 Introduction Patches and demes Dispersal in increasing and stationary populations 231 loa Natal, breeding, and transfer dispersal Breeding systems and the functions of dispersal Demographic consequences of dispersal Conclusions The primate perspective on dispersal A.E. Pusey 11.1 Introduction 11.2 Patterns of dispersal in primates 11.3 Proximate causes of dispersal

7 Vlll Contents 11.4 Evolutionary explanations of dispersal 11.5 Species with female-biased dispersal 11.6 Conclusions A red grouse perspective on dispersal in small mammals 260 A. Watson 12.1 Introduction Red grouse and small mammals as study animals Dispersal or emigration? Models on emigration and immigration as direct causes of demographic change in red grouse and voles Emigration or immigration as a direct cause of demographic change 12.6 Different types of emigration and their demographic consequences 12.7 Emigration, immigration, cycles, and irregular fluctuations 12.8 Emigration as an indirect cause of declines 12.9 Final remarks An entomological perspective on animal dispersal G.K. Roderick and R.L. Caldwell 13.1 Indroduction 13.2 Separate paths 13.3 Insect spatial movements 13.4 Cross-contributions and future directions Part Seven. Appendices Appendix 1. Where do we stand methodologically about experimental design and methods of analysis in the study of dispersal? N.C. Stenseth and W.Z. Lidicker, Jr ALl Introduction A1.2 Techniques for obtaining data A1.3 Methods of analysis Al.4 Future directions

8 Contents lx Appendix 2. Alternative approaches to the study of small mammal dispersal: insights from radiotelemetry 319 W.J. McShea and D.M. Madison A2.1 Introduction 319 A2.2 Verification of dispersal 320 A2.3 Description of movements 320 A2.4 The location of dispersers 323 A2.5 Dispersal distance 324 A2.6 Dispersal frequency 325 A2.7 Future studies 326 A2.8 Conclusions Appendix 3. The use of radioisotopes in the study of dispersal: with a case study 333 N.C. Stenseth and W.Z. Lidicker, Jr A3.1 Introduction 333 A3.2 A brief survey of radioisotope use in population studies 337 A3.3 The Myllymaki-Hansson-Hoff study 340 A3.4 Conclusions Epilogue Index

9 Preface Xlll ally. Hence, we wanted to broaden the scope of the book considerably relative to the ITC symposium. We had two options: (1) to ask the authors of the small mammal chapters to broaden their reviews and include references to other groups of organisms, or (2) to ask a group of additional authors who worked on dispersal in other kinds of organisms to comment on the core chapters of the book from their perspectives. We chose the latter option. By so doing, we would, we believed, achieve two important objectives. We could both go into depth with small mammals, exploiting the various authors' specialities and backgrounds, and still enjoy and profit from the enlarged perspectives provided by authors with experiences far beyond small mammals. Moreover, by adopting the 'commentary strategy' we hoped to produce a more cohesive, comprehensive, and broadly based product than we might have otherwise expected. The result, the present book, has therefore developed into a treatise on animal dispersal which goes into depth in the area of small mammal biology, but claims a much wider scope. In writing this book we have had both the professional researcher and beginning graduate students in mind. For both we have provided a synthetic and broad review of the literature. We hope that readers whose experiences lie outside the small mammal area will find items of interest and profit in this enterprise as well. For the students particularly, we have provided in addition a guide to the conceptual framework of animal dispersal and a methodological guide (both with respect to experimental design and data analysis) for studying dispersal in small mammals. We hope that this book will be especially useful to anyone about to begin a project involving dispersal behaviour. For everyone we have provided some speculations on directions for the future. In science, it is not enough to know where we are and where we have been, we want also to consider where we go from here. We have purposely not intended this book to be viewed merely as a symposium proceedings. Even though all but two of the papers presented at the ITC symposium are included in the book, several additional chapters have been written specifically for the book. In addition, the symposium chapters have been extensively revised in order to give them greater depth of coverage and to better connect the parts. We have enjoyed working on this book, and have found the collaboration with the authors to be professionally very rewarding; we thank them for their co-operation. We hope that readers will find the book as useful as we have found it fulfilling to produce. Nils Chr. Stenseth William Z. Lidicker, Jr

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