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Connectivity Conservation

One of the biggest threats to the survival of many plant and animal species is the destruction or fragmentation of their natural habitats. The conservation of landscape connections, where animal, plants, and ecological processes can move freely from one habitat to another, is therefore an essential part of any new conservation or environmental protection plan. In practice, however, maintaining, creating, and protecting connectivity in our increasingly dissected world is a daunting challenge. This fascinating volume provides a synthesis on the current status and literature of connectivity conservation research and implementation. It shows the challenges involved in applying existing knowledge to real-world examples and highlights areas in need of further study. Containing contributions from leading scientists and practitioners, this topical and thought-provoking volume will be essential reading for graduate students, researchers, and practitioners working in conservation biology and natural resource management.

K EVIN C ROOKS is an assistant professor in the Department of Fish, Wildlife, and Conservation Biology and the Graduate Degree Program in Ecology at Colorado State University. His research investigates the effects of habitat fragmentation, urbanization, and landscape connectivity on the behavior, ecology, and conservation of wildlife.

M. SANJAYAN is a lead scientist for The Nature Conservancy. His current work focuses on ensuring connectivity in applied conservation projects worldwide and on understanding the role of ecosystem services in human well-being and conservation.

Conservation Biology Conservation biology is a flourishing field, but there is still enormous potential for making further use of the science that underpins it. This new series aims to present internationally significant contributions from leading researchers in particularly active areas of conservation biology. It will focus on topics where basic theory is strong and where there are pressing problems for practical conservation. The series will include both single-authored and edited volumes and will adopt a direct and accessible style targeted at interested undergraduates, postgraduates, researchers and university teachers. Books and chapters will be rounded, authoritative accounts of particular areas with the emphasis on review rather than original data papers. The series is the result of a collaboration between the Zoological Society of London and Cambridge University Press. The series ethos is that there are unexploited areas of basic science that can help define conservation biology and bring a radical new agenda to the solution of pressing conservation problems.

1. Conservation in a Changing World, edited by Georgina Mace, Andrew Balmford and Joshua Ginsberg 0 521 63270 6 (hardcover), 0 521 63445 8 (paperback) 2. Behaviour and Conservation, edited by Morris Gosling and William Sutherland 0 521 66230 3 (hardcover), 0 521 66539 6 (paperback) 3. Priorities for the Conservation of Mammalian Diversity, edited by Abigail Entwistle and Nigel Dunstone 0 521 77279 6 (hardcover), 0 521 77536 1 (paperback) 4. Genetics, Demography and Viability of Fragmented Populations, edited by Andrew G. Young and Geoffrey M. Clarke 0 521 78207 4 (hardcover), 0 521 79421 8 (paperback) 5. Carnivore Conservation, edited by John L. Gittleman, Stephan M. Funk, David Macdonald and Robert K. Wayne 0 521 66232 X (hardcover), 0 521 66537 X (paperback) 6. Conservation of Exploited Species, edited by John D. Reynolds, Georgina M. Mace, Kent H. Redford and John G. Robinson 0 521 78216 3 (hardcover), 0 521 78733 5 (paperback) 7. Conserving Bird Biodiversity, edited by Ken Norris and Deborah J. Pain 0 521 78340 2 (hardcover), 0 521 78949 4 (paperback) 8. Reproductive Science and Integrated Conservation, edited by William V. Holt, Amanda R. Pickard, John C. Rodger and David E. Wildt 0 521 81215 1 (hardcover), 0 521 01110 8 (paperback) 9. People and Wildlife: Conflict or Coexistence? edited by Rosie Woodroffe, Simon Thergood and Alan Rabinowitz 0 521 82505 9 (hardcover), 0 521 53203 5 (paperback) 10. Phylogeny and Conservation, edited by Andrew Purvis, John L. Gittleman and Thomas Brooks 0 521 82502 4 (hardcover), 0 521 53200 0 (paperback) 11. Large Herbivore Ecology, edited by Kjell Danell, Roger Bergstrom, Patrick Duncan and John Pastor 0 521 83005 2 (hardcover), 0 521 53687 1 (paperback) 12. Top Predators in Marine Ecosystems, edited by Ian Boyd, Sarah Wanless and C. J. Camphuysen 0 521 84773 7 (hardcover), 0 521 61256 X (paperback) 13. Coral Reef Conservation, edited by Isabelle Coˆte´ and John Reynolds 0 521 85536 5 (hardcover), 0 521 67145 0 (paperback)

Connectivity Conservation

Edited by

KEVIN R. CROOKS Colorado State University

AND

M. SANJAYAN The Nature Conservancy

CAMBRIDGE UNIVERSITY PRESS Cambridge, New York, Melbourne, Madrid, Cape Town, Singapore, Sa˜o Paulo Cambridge University Press The Edinburgh Building, Cambridge CB2 2RU, UK

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

www.cambridge.org Information on this title: www.cambridge.org/9780521857062

ß Cambridge University Press 2006

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 2006

Printed in the United Kingdom at the University Press, Cambridge

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

ISBN-13 978-0-521-85706-2 hardback ISBN-10 0-521-85706-6 hardback ISBN-13 978-0-521-67381-5 paperback ISBN-10 0-521-67381-X 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 ix Acknowledgements xv

1 Connectivity conservation: maintaining connections for nature 1 KEVIN R. CROOKS AND M. SANJAYAN

PART I Approaches to connectivity research 21

Introduction: Connectivity research—what are the issues? 23 JOHN A. WIENS

2 Landscape connectivity: a return to the basics 29 PHILIP D. TAYLOR,LENORE FAHRIG, AND KIMBERLY A. WITH

3 Connectivity and metapopulation dynamics in highly fragmented landscapes 44 ATTE MOILANEN AND ILKKA HANSKI

4 Genetics and landscape connectivity 72 RICHARD FRANKHAM

5 Connectivity at the landÀwater interface 97 DREW M. TALLEY,GARY R. HUXEL, AND MARCEL HOLYOAK

6 Inﬂuence of natural landscape fragmentation and resource availability on distribution and connectivity of gray wolves (Canis lupus) in the archipelago of coastal British Columbia, Canada 130 PAUL C. PAQUET,SHELLEY M. ALEXANDER,PATRICIA L. SWAN, AND CHRIS T. D ARIMONT

vi Contents

7 Migratory connectivity 157 PETER P. MARRA,D.RYAN NORRIS,SUSAN M. HAIG, MIKE WEBSTER, AND J. ANDREW ROYLE

8 Connectivity in marine ecosystems: the importance of larval and spore dispersal 184 CLAUDIO DIBACCO,LISA A. LEVIN, AND ENRIC SALA

9 Connectivity and wide-ranging species in the ocean 213 AUTUMN-LYNN HARRISON AND KAREN A. BJORNDAL

10 Hydrologic connectivity: a neglected dimension of conservation biology 233 CATHERINE PRINGLE

11 Connectivity and ecosystem services: crop pollination in agricultural landscapes 255 TAYLOR H. RICKETTS,NEAL M. WILLIAMS, AND MARGARET M. MAYFIELD

PART II Assessing connectivity 291

Introduction: Evaluating and quantifying the conservation dividends of connectivity 293 PETER KAREIVA

12 Quantifying connectivity: balancing metric performance with data requirements 297 WILLIAM F. FAGAN AND JUSTIN M. CALABRESE

13 Assessing connectivity in salmonid ﬁshes with DNA microsatellite markers 318 HELEN NEVILLE,JASON DUNHAM, AND MARY PEACOCK

14 Individual-based modeling as a tool for conserving connectivity 343 JEFF A. TRACEY

15 Linking connectivity to viability: insights from spatially explicit population models of large carnivores 369 CARLOS CARROLL

Contents vii

16 Impacts of corridors on populations and communities 390 NICK M. HADDAD AND JOSH J. TEWKSBURY

17 Exploring the functional connectivity of landscapes using landscape networks 416 DAVID M. THEOBALD

PART III Challenges and implementation of connectivity conservation 445 Introduction: Don’t fence me in 447 THOMAS LOVEJOY

18 Hyperconnectivity, invasive species, and the breakdown of barriers to dispersal 451 JEFFREY A. CROOKS AND ANDREW V. S UAREZ

19 Disease and connectivity 479 HAMISH MCCALLUM AND ANDY DOBSON

20 Maintaining and restoring connectivity in landscapes fragmented by roads 502 ANTHONY P. CLEVENGER AND JACK WIERZCHOWSKI

21 Where to draw the line: integrating feasibility into connectivity planning 536 SCOTT A. MORRISON AND MARK D. REYNOLDS

22 South Coast Missing Linkages: restoring connectivity to wildlands in the largest metropolitan area in the USA 555 PAUL BEIER,KRISTEEN L. PENROD,CLAUDIA LUKE, WAYNE D. SPENCER, AND CLINT CABAN˜ ERO

23 Incorporating connectivity into broad-scale conservation planning 587 REED F. NOSS AND KATHLEEN M. DALY

24 Escaping the minimalist trap: design and implementation of large-scale biodiversity corridors 620 JAMES SANDERSON,GUSTAVO A. B. DA FONSECA, CARLOS GALINDO-LEAL,KEITH ALGER,VICTOR HUGO INCHAUSTY KARL MORRISON, AND ANTHONY RYLANDS

viii Contents

25 The role of connectivity in Australian conservation 649 MICHAEL E. SOULE´ ,BRENDAN G. MACKEY,HARRY F. RECHER, JANN E. WILLIAMS,JOHN C. Z. WOINARSKI,DON DRISCOLL, WILLIAM C. DENNISON, AND MENNA E. JONES

26 The future of connectivity conservation 676 ANDREW F. BENNETT,KEVIN R. CROOKS, AND M. SANJAYAN

Index 695

List of Contributors

S HELLEY M. ALEXANDER C ARLOS C ARROLL Department of Geography, Klamath Center for Conservation University of Calgary, Calgary, Research, PO Box 104, Orleans, Alberta, T2N 1N4, Canada CA 95556, USA

K EITH A LGER A NTHONY P. CLEVENGER Center for Applied Biodiversity Western Transportation Institute, Science, Conservation International, Montana State University, 1919 M Street NW, Suite 600, PO Box 174250, Bozeman, Washington, DC 20036, USA MT 59717, USA

P AUL B EIER J EFFREY A. CROOKS School of Forestry, Tijuana River National Estuarine Northern Arizona University, Research Reserve, Imperial Beach, Flagstaff, AZ 86011, USA CA 91932, USA

A NDREW F. BENNETT K EVIN R. CROOKS School of Life and Environmental Department of Fish, Wildlife, and Sciences, Deakin University, Conservation Biology, Colorado 221 Burwood Highway, Burwood, State University, Vic 3125, Australia Fort Collins, CO 80523, USA

K AREN A. BJORNDAL K ATHLEEN M. DALY Archie Carr Center for Sea Turtle Wildlands Project, Research and Department of Zoology, PO Box 455, Richmond, University of Florida, Gainesville, VT 05477, USA FL 32611, USA C HRIS T. D ARIMONT C LINT C ABAN˜ ERO Department of Biology, South Coast Wildlands, PO Box 1102, University of Victoria, PO Box 3020, Idyllwild, CA 92549, USA Victoria, BC V8W 3N5, Canada

J USTIN M. CALABRESE W ILLIAM C. DENNISON Department of Biology, University of Maryland Center for University of Maryland, Environmental Science, PO Box 775, College Park, MD 20742, USA Cambridge, MD 21613, USA

x List of Contributors

C LAUDIO D IBACCO Macquarie University, Department of Earth and Ocean NSW 2109, Australia; Sciences, University of British Department of Ecology and Genetics, Columbia, 1461-6270 University of Aarhus, University Boulevard, Vancouver, DK-8000 Aarhus C, Denmark; British Columbia, V6T 1Z4, Canada Australian Museum, 6 College Street, A NDY D OBSON Sydney NSW 2010, Australia Department of Ecology and C ARLOS G ALINDO-LEAL Evolutionary Biology, Princeton Coordinador Programa Bosques University, Princeton, NJ 08544, USA Mexicanos, Fondo Mundial para la D ON D RISCOLL Naturaleza (World Wildlife School of Biological Sciences, FundÀMe´xico), Av. Me´xico 51, Flinders University, GPO Box 2100, Col. Hipo´dromo, Me´xico, Adelaide, SA 5001, Australia D.F. 06100, Me´xico

J ASON D UNHAM N ICK M. HADDAD US Forest Service, Rocky Mountain Department of Zoology, PO Box 7617, Research Station, Suite 401, 322 North Carolina State University, E. Front St, Boise, ID 83702, USA; Raleigh, NC 27695, USA USGS Forest and Rangeland S USAN M. HAIG Ecosystem Science Center, US Geological Survey Forest and Corvallis Research Group, Rangeland Ecosystem Science Center, 3200 SW Jefferson Way, 3200 SW Jefferson Way, Corvallis, Corvallis, OR 97331, USA OR 97331, USA W ILLIAM F. FAGAN I LKKA H ANSKI Department of Biology, Metapopulation Research Group, University of Maryland, Department of Biological and College Park, MD 20742, USA Environmental Sciences, PO Box 65 L ENORE F AHRIG (Biocenter 3), FI-00014 University of Department of Biology, Carleton Helsinki, Finland University, Ottawa, Ontario, A UTUMN-LYNN H ARRISON K1S 5B6, Canada Department of Ecology and G USTAVO A. B. DA F ONSECA Evolutionary Biology, University of Center for Applied Biodiversity California Santa Cruz, Long Marine Science, Conservation International, Laboratory, 100 Shaffer Road, 1919 M Street NW, Suite 600, Santa Cruz, CA 95060, USA Washington, DC 20036, USA M ARCEL H OLYOAK R ICHARD F RANKHAM Department of Environmental Key Centre for Biodiversity and Science and Policy, University of Bioresources, Department California, 1 Shields Avenue, Davis, of Biological Sciences, CA 95616, USA

List of Contributors xi

G ARY R. HUXEL The Australian National University, Department of Biology, Canberra, ACT 0200, Australia University of South Florida, P ETER P. MARRA 4202 East Fowler Avenue, Smithsonian Migratory Bird Center, SCA110, Tampa, National Zoological Park, FL 33620, USA 3001 Connecticut Avenue NW,

V ICTOR H UGO I NCHAUSTY Washington, DC 20008, USA Director, Areas Protegidas, M ARGRARET M. MAYFIELD Ministerio Desarrollo Agropecuario Department of Ecology, Evolution Rural y Medio Ambiente, and Marine Sciences, University Loayza Edif. Lara Bisch, of California Santa Barbara, La Paz, Bolivia Santa Barbara, CA 93103, USA

M ENNA E. JONES H AMISH M C C ALLUM School of Zoology, School of Integrative Biology, University of Tasmania, The University of Queensland, Private Bag 5, Hobart, Brisbane, Qld 4072, Australia Tas 7004, Australia A TTE M OILANEN P ETER K AREIVA Metapopulation Research Group, The Nature Conservancy, Seattle, Department of Biological and WA 98105, USA Environmental Sciences,

L ISA A. LEVIN PO Box 65 (Biocenter 3), FI-00014 Integrative Oceanography Division, University of Helsinki, Finland Scripps Institution of Oceanography, K ARL M ORRISON 9500 Gilman Drive, Center for Applied Biodiversity La Jolla, CA 92093, USA Science, Conservation International, T HOMAS L OVEJOY 1919 M Street NW, The H. John Heinz III Center for Suite 600, Washington, Science, Economics and the DC 20036, USA Environment, Washington, DC 20004, USA S COTT A. MORRISON The Nature Conservancy, C LAUDIA L UKE 201 Mission Street, 4th Floor, Bodega Marine Laboratory and San Francisco, CA 94105, USA Reserve, University of California, PO Box 247, Bodega Bay, H ELEN N EVILLE Department of Biology, University CA 94923, USA of Nevada, Reno, NV 89557, USA; B RENDAN G. MACKEY US Forest Service, Rocky Mountain School of Resources, Research Station, Suite 401, Environment and Society, 322 E. Front St, Faculty of Science, Boise, ID 83702, USA

xii List of Contributors

D. RYAN N ORRIS T AYLOR H. RICKETTS Department of Integrative Biology, Conservation Science Program, University of Guelph, Guelph, World Wildlife Fund, Ontario, Canada N1G 2W1 1250 24th Street NW, Washington, DC 20037, USA R EED F. NOSS Department of Biology, University J. ANDREW R OYLE of Central Florida, Orlando, US Fish and Wildlife Service, FL 32816, USA; Wildlands Project, Division of Migratory Bird PO Box 455, Richmond, Management, 11510 American Holly VT 05477, USA Drive, Laurel, MD 20707, USA

NTHONY YLANDS P AUL C. PAQUET A R Faculty of Environmental Design, Center for Applied Biodiversity University of Calgary, Calgary, Science, Conservation International, Alberta, T2N 1N4, Canada; 1919 M Street NW, Suite 600, Raincoast Conservation Society, Washington, PO Box 26, Bella Bella, DC 20036, USA British Columbia, E NRIC S ALA V0T 1B0, Canada Center for Marine Biodiversity and Conservation, Scripps M ARY P EACOCK Institution of Oceanography, Department of Biology, 9500 Gilman Drive, University of Nevada, La Jolla, CA 92093, USA Reno, NV 89557, USA J AMES S ANDERSON K RISTEEN L. PENROD Center for Applied Biodiversity South Coast Wildlands, Science, Conservation International, PO Box 1102, Idyllwild, 1919 M Street NW, CA 92549, USA Suite 600, Washington, DC 20036, USA C ATHERINE P RINGLE

Institute of Ecology, M. SANJAYAN University of Georgia, Athens, The Nature Conservancy, GA 30602, USA 4245 North Fairfax Drive, Arlington, VA 22203, USA H ARRY F. RECHER School of Natural Sciences, M ICHAEL E. SOULE´ Edith Cowan University, Joondalup, PO Box 1808, Paonia, WA 6027, Australia CO 81428, USA

M ARK D. REYNOLDS W AYNE D. SPENCER The Nature Conservancy, 4th Floor, Conservation Biology Institute, 201 Mission Street, San Francisco, 815 Madison Avenue, CA 94105, USA San Diego, CA 92116, USA

List of Contributors xiii

A NDREW V. S UAREZ J EFF A. TRACEY Department of Entomology and Graduate Degree Program in Ecology Department of Animal Biology, and Program for Interdisciplinary School of Integrative Biology, Mathematics, Ecology, and Statistics, University of Illinois at Colorado State University, UrbanaÀChampaign, Fort Collins, CO 80523, USA Urbana, IL 61801, USA M IKE W EBSTER

P ATRICIA L. SWAN School of Biological Sciences, Department of Geography, Washington State University, University of Calgary, Calgary, PO Box 644236, Pullman, Alberta, T2N 1N4, Canada WA 99164, USA

J OHN A. WIENS D REW M. TALLEY Department of Environmental The Nature Conservancy, Science and Policy, University Arlington, VA 22203, USA of California, 1 Shields Avenue, J ACK W IERZCHOWSKI Davis, CA 95616, USA; Geomar, PO Box 1843, San Francisco Bay National Grand Forks, British Columbia, Estuarine Research Reserve, V0H 1H0, Canada San Francisco State J ANN E.WILLIAMS University/Romberg Tiburon Center, Centre for Sustainable Regional 3152 Paradise Drive, Tiburon, Communities, LaTrobe University, CA 94920, USA PO Box 199, Bendigo, P HILIP D. TAYLOR Vic 3550, Australia Department of Biology, N EAL M. WILLIAMS Acadia University, Department of Biology, Wolfville, Nova Scotia, Bryn Mawr College, 101 N Merion B4P 2R6, Canada Avenue, Bryn Mawr,

J OSH J. TEWKSBURY PA 19010, USA

Department of Biology, K IMBERLY A. WITH PO Box 351800, Division of Biology, 24 Kincaid Hall, Kansas State University, University of Washington, Manhattan, KS 66506, USA Seattle, WA 98195, USA J OHN C. Z. WOINARSKI D AVID M. THEOBALD Biodiversity Section, Natural Resource Ecology Natural Systems, Laboratory and Department of Department of Infrastructure, Recreation and Tourism, Planning and Environment, Colorado State University, PO Box 496, Palmerston, Fort Collins, CO 80523, USA NT 0831, Australia

Acknowledgements

We first would like to thank all chapter authors for their excellent contributions to this volume. Their collective breadth and depth of knowledge, as well as their enthusiasm, diligence, and patience throughout the publication process, made this volume possible. We are particularly grateful to A. L. Harrison for her tireless work in helping organize, compile, and format the volume prior to submission. All chapters in this volume received at least two peer reviews and we sincerely thank the reviewers for their thoughtful comments, edits, and suggestions: B. Allan, L. Angeloni, S. Baruch-Mordo, C. Baxter, S. Bayard de Volo, P. Beier, L. Bernatchez, C. Carroll, T. Clevenger, H. Crockett, J. Crooks, K. Daly, C. DiBacco, B. Dickson, M. DiGiorgio, B. Fagan, J. Forester, H. Fox, M. Freeman, B. Grenfell, S. Griffin, A. Gonzalez, N. Haddad, A. L. Harrison, B. Hastings, P. Kareiva, M. Kavermann, J. Kelly, P. Kevan, J. Kintsch, R. Lavier, S. Magle, B. Mila, T. Mildenstein, M. Miller, S. Mills, A. Moilanen, S. Morrison, M. Neel, H. Neville, C. Pague, H. Possingham, J. Powell, T. Ricketts, E. Ruell, J. Sanderson, R. Scherer, R. Shaw, P. Snelgrove, D. Smith, D. Spiering, A. Suarez, D. Talley, S. Tartowski, P. Taylor, T. Tear, M. Tilton, J. Tracey, and M. Wonham. Special thanks also to the EY 592 graduate seminar at Colarado State University (led by K. Crooks), and a graduate seminar at Scripps Institution of Oceanography (led by L. Levin), for their helpful reviews of earlier drafts of this book. We thank Cambridge University Press for their assistance in publication of the volume, specifically A. Crowden for his continued interest and encouragement, A. Hodson for her meticulous copy-editing skills, and C. Georgy, D. Lewis, and D. Preston for their help with the contract, editing, production, and design stages. Additionally, we acknowledge the funding and support of The Nature Conservancy, The Society for Conservation Biology, University of WisconsinÀMadison, and Colorado State University during the compilation of the volume.

xvi Acknowledgements

We are also indebted to our advisor and teacher, Michael Soule´, for his guidance, wisdom, and inspiration during our careers. Finally, we extend our deepest appreciation to our families for their continued love and encouragement. K. Crooks would like to thank his wife and daughter, Lisa and Elena, his parents, Bob and Mary, and his brothers, Jeff and Matt. M. Sanjayan would like to thank his parents, Muttu and Ranji, and his sister, Vaithi. We dedicate this volume to you.