and Marchetti Lockwood, Hoopes, panion m w o e c b s e i e t e r Invasion Ecology f Second Edition Lockwood, Hoopes, and Marchetti INVASION This new edition of Invasion Ecology provides a comprehensive and updated introduction to all aspects of biological invasion by non-native species. Highlighting Invasion Ecology important research fi ndings associated with each stage of invasion, the book provides an overview of the invasion process from transportation patterns and causes of ECOLOGY establishment success to ecological impacts, invader management, and post-invasion evolution. The authors have produced new chapters on predicting and preventing invasion, managing and eradicating invasive species, and invasion dynamics in a changing climate. Second Edition Modern global trade and travel have led to unprecedented movement of non-native species by humans with unforeseen, interesting, and occasionally devastating consequences. Increasing Julie L. Lockwood, recognition of the problems associated with invasion has led to a rapid growth in research into the dynamics of non-native species and their adverse effects on native biota and human economies. Martha F. Hoopes, and This book provides a synthesis of this rapidly growing fi eld of research and is an essential text for undergraduate and graduate students in ecology and conservation management. Michael P. Marchetti Julie L. Lockwood is a Professor at Rutgers University. Her research interests include conservation biology, population biology, and biological invasions. Martha F. Hoopes is an Associate Professor at Mount Holyoke College. She has worked primarily with plants and insects on questions of spatial community dynamics and invasions. Michael P. Marchetti is the Fletcher Jones Professor of Ecology at St. Mary’s College of California. Second Edition He is an aquatic ecologist who primarily studies community and landscape aspects of freshwater invaders in the western U.S. panion m w There is a free companion website available for this book at o e c b s e i e t e r f www.wiley.com/go/invasionecology On the site you will fi nd: ■ Downloadable versions of the illustrations within the book ■ Downloadable versions of the tables within the book panion m w o e c b s e i e t e r f hb_9781444333640.indd 1 27/2/13 13:40:33 Invasion Ecology Invasion Ecology Second Edition Julie L. Lockwood Ecology, Evolution and Natural Resources Department Rutgers, The State University of New Jersey USA Martha F. Hoopes Department of Biological Sciences Mount Holyoke College USA Michael P. Marchetti Fletcher Jones Professor of Ecology Biology Department St Mary’s College of California USA A John Wiley & Sons, Ltd., Publication This edition first published 2013 © 2013 Julie L. Lockwood, Martha F. Hoopes, and Michael P. Marchetti First edition published 2007 © 2007 Julie L. Lockwood, Martha F. Hoopes, and Michael P. Marchetti Blackwell Publishing was acquired by John Wiley & Sons in February 2007. Blackwell’s publishing program has been merged with Wiley’s global Scientific, Technical and Medical business to form Wiley-Blackwell. Registered Office John Wiley & Sons, Ltd, The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK Editorial Offices 9600 Garsington Road, Oxford, OX4 2DQ, UK The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK 2121 State Avenue, Ames, Iowa 50014-8300, USA 111 River Street, Hoboken, NJ 07030-5774, USA For details of our global editorial offices, for customer services and for information about how to apply for permission to reuse the copyright material in this book please see our website at www.wiley.com/wiley-blackwell. The right of the authors to be identified as the authors of this work has been asserted in accordance with the UK Copyright, Designs and Patents Act 1988. All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, except as permitted by the UK Copyright, Designs and Patents Act 1988, without the prior permission of the publisher. Designation used by companies to distinguish their products are often claimed as trademarks. All brand names and product names used in this book are trade names, service names, trademarks or registered trademarks of their respective owners. The publisher is not associated with any product or vendor mentioned in this book. Limit of Liability/Disclaimer of Warranty: While the publisher and author(s) have used their best efforts in preparing this book, they make no representations or warranties with respect to the accuracy or completeness of the contents of this book and specifically disclaim any implied warranties of merchantability or fitness for a particular purpose. It is sold on the understanding that the publisher is not engaged in rendering professional services and neither the publisher nor the author shall be liable for damages arising herefrom. If professional advice or other expert assistance is required, the services of a competent professional should be sought. Library of Congress Cataloging-in-Publication Data Lockwood, Julie L. Invasion ecology / Julie Lockwood, Ecology, Evolution and Natural Resources Department Rutgers University, USA, Martha Hoopes, Biological Sciences Department,Mount Holyoke College, USA, Michael Marchetti, Fletcher Jones Professor of Biology, Biology Department, St Mary’s College of California, USA. – Second edition. pages cm Includes bibliographical references and index. ISBN 978-1-4443-3364-0 (hardback : alk. paper) – ISBN 978-1-4443-3365-7 (softback : alk. paper) – ISBN 978-1-118-57078-4 – ISBN 978-1-118-57080-7 – ISBN 978-1-118-57081-4 (emobi) – ISBN 978-1-118-57082-1 (epub) – ISBN 978-1-118-57083-8 (epdf) 1. Biological invasions. I. Hoopes, Martha F. II. Marchetti, Michael P. III. Title. QH353.L63 2013 577′.18–dc23 2013001797 A catalogue record for this book is available from the British Library. Wiley also publishes its books in a variety of electronic formats. Some content that appears in print may not be available in electronic books. Cover images: Front: Gorse (Ulex europaeus) at Lake Wakatipu just outside of Queenstown, New Zealand. Gorse is an extremely invasive weed in New Zealand, and covers approximately 700,000 hectares (1,700,000 acres) of the New Zealand landscape. Gorse is native to Western Europe and has become established in NZ, Australia, the US and Chile. Photo courtesy of Michael Marchetti. Back: Nutmeg Mannikin (Lonchura punctulata), a passerine bird native to tropical Asia that has been introduced to Puerto Rico, Australia and the US. The picture was taken in Puerto Rico. Photo courtesy of Alberto Lopez. Cover design by Nicki Averill Design & Illustration Set in 10.5/12pt Classical Garamond by SPi Publisher Services, Pondicherry, India 1 2013 Contents Preface ix 1 An Introduction to Invasion Ecology 1 What are invaders and why do we care about them? 2 A brief history of invasion ecology 5 The wicked terminological web we weave 7 The invasion process 13 Summary 18 2 Transport Vectors and Pathways 24 What’s the difference between a vector and a pathway? 25 Does human-mediated dispersal differ from natural dispersal? 26 Transport vectors 29 Which species are transported via what vector group? 41 Dynamics of transport pathways 44 Summary 48 3 Trends in Numbers of Invaders 50 Invasion rates through time 51 Geographic patterns in numbers of invaders 63 Summary 72 4 Propagules 74 What are propagules? 75 Donor region and propagule pressure 75 vi Contents Biological mechanisms 79 Empirical evidence 85 The hidden influence of propagule pressure 92 Summary 97 5 Disturbance 99 History and definition of disturbance 100 Disturbance facilitates invasion? 104 Restoration and disturbance 112 Agriculture and urbanization as disturbance 115 Biotic disturbance 118 Summary 127 6 Establishment Success: The Influence of Biotic Interactions 129 Conceptual issues 130 Resistance to invasion 131 Facilitation of establishment 146 Summary 155 7 Modeling the Geographical Spread of Invasive Species 157 What exactly is geographical spread? 158 Why do we want to model geographical spread? 162 The reaction–diffusion model 163 Long-distance dispersal 170 Directional dispersal 173 Stratified dispersal 176 Other forms of heterogeneity 182 Summary 187 8 Ecological Processes and the Spread of Non-native Species 189 Population growth 190 Dispersal 194 Biotic interactions 202 The role of heterogeneity 207 Lag times 210 Boom and bust 215 Summary 216 Contents vii 9 Ecological Impacts of Invasive Species 218 Genetic impacts 219 Individual impacts 222 Population impacts 228 Community impacts 233 Ecosystem impacts 240 Landscape, regional, and global impacts 242 Summary 244 10 Impact Synthesis 246 Perception and recognition of impact 247 Integrating perception with ecological determinants of impact 255 A theory of impact? 258 Finding common currencies 263 A cross-stage impact formula 273 Summary 275 11 Evolution of Invaders 277 Founding process 279 Losses and gains in genetic variability via transport mechanisms 279 Genetics and post-release success 288 Local adaptation and life-history evolution 291 Evolution of native species in response to non-natives 296 Summary 298 12 Predicting and Preventing Invasion 299 Explanation versus risk assessment 301 Inherent limitations to prediction 301 Risk analysis 303 Screening risky species 304 Screening risky transportation vectors 317 Summary 333 13 Eradication and Control of Invaders 335 Cause for optimism? 336 Rapid response 337 Lazarus effect 343 Long-term control 346 Sisyphus effect 350 Summary 354 viii Contents 14 Global Climate Change and Invasive Species 356 Global climate change 101 357 Non-native species and global climate change 364 Transport 365 Establishment 368 Spread 373 Impact 379 Human responses 387 Summary 391 References 393 Index 428 A colour plate section falls between pages 372 and 373 Preface It has been our experience that placing the reams of observations, examples, and research within the broad context of invasion ecology gets more difficult as the field grows each year.