About Island Press Island Press is the only nonprofit organization in the United States whose princi- pal purpose is the publication of books on environmental issues and natural resource management.We provide solutions-oriented information to professionals, public officials,business and community leaders,and concerned citizens who are shaping responses to environmental problems. In 2004,Island Press celebrates its twentieth anniversary as the leading provider of timely and practical books that take a multidisciplinary approach to critical envi- ronmental concerns.Our growing list of titles reflects our commitment to bring- ing the best of an expanding body of literature to the environmental community throughout North America and the world. Support for Island Press is provided by the Agua Fund,Brainerd Foundation, Geraldine R.Dodge Foundation,Doris Duke Charitable Foundation,Educational Foundation of America,The Ford Foundation,The George Gund Foundation, The William and Flora Hewlett Foundation,Henry Luce Foundation,The John D. and Catherine T. MacArthur Foundation,The Andrew W. Mellon Foundation, The Curtis and Edith Munson Foundation, National Environmental Trust,The New-Land Foundation,Oak Foundation,The Overbrook Foundation,The David and Lucile Packard Foundation,The Pew Charitable Trusts,The Rockefeller Foun- dation,The Winslow Foundation,and other generous donors. The opinions expressed in this book are those of the author(s) and do not nec- essarily reflect the views of these foundations. ALIEN SPECIES AND EVOLUTION ALIEN SPECIES AND EVOLUTION The Evolutionary Ecology of Exotic Plants, Animals, Microbes, and Interacting Native Species George W. Cox Island Press Washington • Covelo • London Copyright © Island Press All rights reserved under International and Pan-American Copyright Conventions.No part of this book may be reproduced in any form or by any means without permission in writ- ing from the publisher:Island Press,1718 Connecticut Avenue,NW,Suite 300,Washington, DC 20009. ISLAND PRESS is a trademark ofThe Center for Resource Economics. Library of Congress Cataloging-in-Publication data. Cox,George W.,1935– The evolutionary ecology of exotic plants,animals,microbes,and interacting native species / George W.Cox. p.cm. Includes bibliographical references and index. ISBN 1–55963–008–6 (cloth :alk.paper) — ISBN 1–55963–009–4 (pbk.:alk.paper) 1.Introduced organisms--Environmental aspects. 2.Introduced animal— Evolution. 3.Exotic plants—Evolution. I.Title. QH353.C69 2004 578.6'2—dc22 2003027299 British Cataloguing-in-Publication data available. Design by Brighid Willson Printed on recycled,acid-free paper Manufactured in the United States of America 10 9 8 7 6 5 4 3 2 1 Contents Preface ix Part I. Basic Concepts of Alien Invasion and Evolution 1 1. Alien Species and Accelerated Evolution 3 2. Adaptation of Alien Species for Dispersal and Establishment 17 3. Founder Effects and Exotic Variability 32 4. Introduction Sources,Cryptic Species,and Invasion Routes 47 Part II. Processes of Evolutionary Change and Adaptation 61 5. Hybridization and Evolution of Exotics 63 6. Hybridization and Transgenic Organisms 76 7. Invasion Resistance of Native Communities 89 8. Adaptation of Alien Species to New Habitats 105 Part III. Evolutionary Interaction of Aliens and Natives 119 9. Evolutionary Adaptation by Alien Herbivores 121 10. Evolutionary Adaptation by Alien Predators and Parasites 134 11. Adaptation of Alien Diseases to Hosts and Vectors 148 12. Adaptation of Plants to Alien Herbivores and Diseases 161 13. Adaptation of Native Herbivores to Alien Plants 174 vii viii Contents 14. Adaptation of Animals to Alien Predators,Parasites,and Disease Agents 188 15. Accumulation of Herbivores,Predators,and Parasites by Alien Species 202 Part IV. Global Evolutionary Consequences of Alien Invasions 217 16. Alien Species as Agents of Extirpation and Extinction 219 17. Evolutionary Ecology of Alien Biological Control Agents 232 18. Counteradaptation and Integration into the Biotic Community 245 19. Dispersing Aliens and Speciation 258 20. Permanently Altered Biotic Communities 272 Literature Cited 287 Glossary 345 Index 355 Preface The phenomena of biological invasions and rapid evolution have come together in the last decade to reveal a host of cases of rapid evolutionary change by alien species and the native species with which they interact. All groups of organisms—plants, animals, and microorganisms of all types—are involved.These examples of rapid evolution are of great inter- est to theoretical biologists, but they include many aspects of applied importance,as well.The potential for the spread of transgenes from plants of economic importance to wild and weed relatives is substantial.The same is true of transgenes introduced into populations of animals,espe- cially those of importance in aquaculture and biological control.The growing number of species introduced for biological control also means a growing potential for ecological and evolutionary shifts that enable these agents to attack nontarget hosts. More generally, the mixing of genomes of alien and native species through hybridization increases the potential for the evolution of new invasive forms. Despite the numerous,well-publicized cases of evolutionary shifts in human disease agents and the diseases and pests of agricultural plants and animals, the public at large remains largely unaware of the accelerated pace of evolution resulting from the massive introduction of species to new geographical regions.In particular,systems of governmental regula- tion of alien species tend to regard them as stable genetic entities.In edu- cation,the concept of evolution is treated as a controversial theory of the history of life on earth,rather than an on-going process of great relevance to human welfare. Recognition of the importance of rapid evolution involving alien species culminates more than a half century of interest in the biology of introduced organisms.In 1958,in perhaps one of the most forward-look- ing publications in ecology,Charles Elton characterized introductions of ix
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