An Introduction to Biological Control
An Introduction to Biological Control Robert van den Bosch P. S. Messenger A. P. Gutierrez Division of Biological Control University of California, Berkeley Albany, California SPRINGER SCIENCE+BUSINESS MEDIA, LLC
Library of Congress Cataloging in Publication Data Van den Bosch, Robert. An introduction to biological control. Rev. ed. of: Biological control. 1973. Includes bibliographical references in index. 1. Pest control- Biological control. 2. Insect control- Biological control. I. Messenger, P. S. II. Gutierrez, A. P. III. Title. SB97S.V36 1982 632'.96 ISBN 978-1-4757-9164-8 ISBN 978-1-4757-9162-4 (ebook) DOI 10.1007/978-1-4757-9162-4 81-21125 AACR2 This limited facsimile edition has been issued for the purpose of keeping this title available to the scientific community. First Printing- February 1982 Second Printing-October 1985 This volume is a revised version of BIOLOGICAL CONTROL, by Robert van den Bosch and P.S. Messenger, published in 1973 by Intext Press, New York. The original text has been thoroughly updated for this edition. 1982 Springer Science+Business Media New York Originally published by Plenum Press, New York in 1982 Softcover reprint of the hardcover 1st edition 1982 All rights reserved No part of this book may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, microfilming, recording, or otherwise, without written permission from the Publisher
TO ROBERT VAN DEN BOSCH ( 1922-1978) For the legacy of good works in biological control, for aspiring to the highest of ideals in environmental protection, for tenacity in the face of adversity, for faithfulness to human principles beyond the reach of most men, and for the genuine love his bright spirit cast upon those fortunate enough to have known him. TOP. S. MESSENGER (1920-1976) For the steady leadership of the Division of Biological Control he provided during difficult times, for his many fine scientific works, for his tenacious struggle to enhance and expand the field of biological control, and lastly for the model he provided for students to emulate. A. P. Gutierrez
Preface This volume is a revision of Biological Control by R. van den Bosch and P. S. Messenger, originally published by Intext Publishers. In the revision, I have attempted to keep the original theme, and to update it with current research findings and new chapters or sections on insect pathology, microbial control of weeds and plant pathogens, population dynamics, integrated pest management, and economics. The book was written as an undergraduate text, and not as a complete review of the subject area. Various more comprehensive volumes have been written to serve as handbooks for the experts. This book is designed to provide a concise overview of the complex and valuable field of biological control and to show the relationships to the developing concepts of integrated pest management. Population regulation of pests by natural enemies is the major theme of the book, but other biological methods of pest control are also discussed. The chapter on population dynamics assumes a precalculus-level knowledge of mathematics. Author names of species are listed only once in the text, but all are listed in the Appendix. Any errors or omissions in this volume are my sole responsibility. A. P. Gutierrez Professor of Entomology Division of Biological Control University of California, Berkeley vii
Acknowledgments Very special thanks must be given to my colleagues, Professors C. B. Huffaker and L. E. Caltagirone, for the very thorough review they provided and for the many positive suggestions they gave. Dr. Lowell Etzel helped develop the sections on insect pathology, and for this I am most grateful. The tireless efforts of Mr. William Voigt require special mention, as he served in very many capacities, among them illustrator, reviewer, typist, and in general an invaluable assistant. Mrs. Joanne Fox and Ms. Barbara Gemmill remained cheerful while typing the many revisions. Messrs. J. K. Clark (University of California, Davis) and P. F. Daley and F. Skinner (University of California, Berkeley) provided many of the photographs of natural enemies reproduced here. Dr. George Poinar (University of California, Berkeley) provided photography and illustrations for the sections on insect pathology, while Dr. D. F. Waterhouse (Chief, Division of Entomology, CSIRO, Australia) provided the illustrations dealing with biological control of the European rabbit and dung in Australia. A. P. G. ix
Contents Chapter 1 The Nature and Scope of Biological Control 1 Definition of Biological Control.............................. 2 A Comparison of the Biological Control of Pest Insects and Weeds.............................................. 3 The Role of Pathogens in Biological Control................... 5 Summary... 7 References................................................. 7 Chapter 2 The Ecological Basis for Biological Control................. 9 Biological Control of Native Species.......................... 17 References................................................. 19 Chapter 3 The History and Development of Biological Control.......... 21 Ancient Origins............................................ 22 North American Beginnings.................................. 23 Twentieth Century Developments.............................. 28 Quarantine Considerations.................................... 29 Development of International Organizations..................... 34 References................................................. 36 xi
xii Chapter 4 Natural Enemies Contents 37 Predators................................................. 38 Parasites................................................. 46 Host Specificity............................................ 55 Conclusion............................................... 56 References................................................. 57 Chapter 5 Microbial Control of Insects, Weeds, and Plant Pathogens 59 The Definition and Scope of Microbial Control................. 59 History.................................................. 60 Uses and Advantages of Microbial Control Agents.............. 61 The Kinds of Microbial Agents............................... 61 Microbial Control of Weeds and Plant Pathogens................ 68 The Future of Microbial Control.............................. 73 References................................................. 73 Chapter 6 Procedures in Natural-Enemy Introduction 75 Identification of the Pest as an Exotic Species.................. 75 Detennination of the Native Habitat of the Exotic Pest........... 77 Importation Agencies in the United States...................... 79 Foreign Exploration for Natural Enemies....................... 81 Quarantine Reception........................................ 82 Mass Culture of Entomophagous Insects....................... 85 Colonization.............................................. 87 Evaluation of Natural Enemies............................... 90 References................................................. 94 Chapter 7 Life Table Analysis in Population Ecology 95 Age-Specific Life Tables..................................... 95 Time-Varying Life Tables.................................... 104 What Use Are Models?..................................... 113 Appendix................................................ 114 References................................................. 115
Contents xiii Chapter 8 Factors Limiting Success of Introduced Natural Enemies 117 Characteristics of the Colonized Environment................... 118 Poorly Adapted Natural-Enemy Species and Strains.............. 120 The Importance of Biotypes in Biological Control............... 121 The Mechanics of Natural-Enemy Introductions................. 128 Conclusion............................................... 129 References................................................. 129 Chapter 9 Analysis of Classical Biological Control Programs 131 The Walnut Aphid... 142 The Winter Moth... 144 The Oriental Fruit Fly...................................... 146 The Imported Cabbageworm................................. 148 The Eucalyptus Snout Beetle in South Africa................... 151 The Olive Scale in California................................ 153 St. Johnswort or Klamath Weed.............................. 158 References................................................. 162 Chapter 10 Naturally Occurring Biological Control and Integrated Control............................ 165 Effects of Insecticides....................................... 166 Ecological and Economic Backlash............................ 167 Integrated Control.......................................... 175 References................................................. 184 Chapter 11 Other Kinds of Pests and Other Biological Methods of Pest Control......................................... 187 Other Kinds of Pests..................... 187 Other Biological Methods of Pest Control...................... 192 Summary.................................................. 206 References................................................. 207
xiv Contents Chapter 12 Economics of Biological Controls.......................... 209 Importation of Biological Controls............................ 209 Research and Development of Imported Biological Controls....... 214 Inundative Release of Imported or Native Natural Enemies and Environmental Enhancement............................ 215 A Working Example of Mass Culture and Augmentation of Imported Parasites 216 References................................................. 218 Chapter 13 The Future of Biological Control... 219 Classical Biological Control.................................. 220 Naturally Occurring Biological Control... 221 Special Manipulation of Natural Enemies... 221 Pest Management and Biological Control....................... 222 Conclusion............................................... 223 References................................................. 223 Appendix: List of Species Cited in the Text................. 225 Glossary............................................... 23 5 Index... 239