Biological Control

Biological Control

Roy G. VAN DRIESCHE Department of Entomology, University of Massachusetts, Amherst, MA

THOMAS S. BELLOWS, JR. Department of Entomology, University of California, Riverside, CA

KLUWER ACADEMIC PUBUSHERS BOSTONIDORDRECHTILONDON

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Library of Congress Cataloging-in-Publication

Van Driesche, R.G. Biological control I Roy G. Van Driesche and Thomas S. Bellows, IT.

p. cm. Includes bibliographical references and index.

ISBN-13:978-1-4612-8490-1 DOT: I 0.1 007/978-1-4613-1157-7

e-ISBN-13978-1-4613-1157-7

I. Pests--Biological control. 2. Pollination by animals. 3. Pollination by insects. 4. Angiosperms--Evolution. 1. Bellows, T.S. n. Title SB975.V375 1996 95-31557 632'.96--dc20

Copyright © 1996 by Chapman & Hall

Sotlcover reprint ofthe hardcover I st edition 1996

Third Printing 2001 by Kluwer Academic Publishers

Cover photos courtesy of: 1.A. MacDonald, P. Kenmore, and J.K. Clark. Cover design: Robert Freese

This printing is a digital duplication of the original edition.

CIP

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, mechanical, photo-copying, recording, or otherwise, without the prior written permission of the publisher, K1uwer Academic Publishers, 101 Philip Drive, Assinippi Park, Norwell, Massachusetts 02061

Printed on acid-free paper.

CONTENTS

Prtiface , , , xi

SECTION I ORIGINS AND SCOPE OF BIOLOGICAL CONTROL 1

CHAPTER 1 PEST ORIGINS, PESTICIDES AND THE HISTORY OF BIOLOGICAL CONTROL. . . . . . . . . . . . . . . . . . . . . . . . . . 3

Human Needs and the Origins of Pests ,.,.,',.,',.".".,",.,"" .... "., 3 Problems with Pesticides ' , , ' , , , , . , , ..... , , , , , , , . , , . , , , , , , , , , , , . , . ' , . , , 4 Definition of Biological Pest Control , .. """"', .. ," .... "','" ... ,,,',.' 6 History of Biological Control """ .. "."",."" .. " .. """"",.""" 7 The Historical Record of Biological Control Efforts """"""', .. ,"',.,"" 19 Biological Control and Integrated Pest Management . , ' , , ' . , , . , , , , , , ..... ' , , . , 20

CHAPTER 2 KINDS OF BIOLOGICAL CONTROL TARGETS, AGENTS, AND METHODS ........................ 21

Targets of Biological Control """""""""""""""""""""'" 21 Kinds of Biological Control Agents """"""""""""""""""'" 23 Principal Biological Control Methods, , , ' , , , , ' . ' . , .. , , , . , , . , , ' . , , , , , . , , , . , , 31

SECTION /I A REVIEW OF THE ORGANISMS EMPLOYED AS AGENTS OF BIOLOGICAL CONTROL. . . . . . . . . . . . . . 35

CHAPTER 3 PARASITOIDS AND PREDATORS OF ARTHROPODS AND MOLLUSCS ................................ 37

Introduction .... , , , . , , .. ' , , , , , , . , ' , , .. , , .. , , , . , ' .. , , , , , , , , , . , , .. , , , , , , , , 37 Insects Parasitic on Arthropods and Snails """",."""."."""",.",. 37 Insects Predacious on Arthropods """""""""""""""""""'" 53 Arachnids Predacious on Arthropods ", .... ,.', ... ,.,',",'., .. ,',",.,.,' 60 Predacious or Competitor Snails , , , , , , . , , , . , , , .. , , ' , . , , . , , , , , , , , .. , , , ... , , . 64 Vertebrates (Mammals, Birds, Fish) ",.""" ... """",."""" ... ,.,... 65

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CHAPTER 4 PATHOGENS AND NEMATODES OF ARTHROPODS, AND PATHOGENS OF VERTEBRATES .............. .

Pathogens of Pest Arthropods .................................. . Pathogens of Pest Vertebrates ...................................... .

CHAPTER 5 HERBIVORES AND PATHOGENS USED FOR BIOLOGICAL WEED CONTROL ................... .

Introduction ................................. . Insects and Mites ...................................................... . Fungal Pathogens ....................................... . Nematodes ................................. . Vertebrates .................................... .

CHAPTER 6 BIOLOGICAL CONTROL AGENTS FOR PLANT PATHOGENS ................................... .

Introduction ........................................................... . Root Pathogens ........................................................ . Stem Pathogens ....................................................... . Leaf Pathogens ........................................................ . Flower and Fruit Pathogens ............................................. . Plant-Parasitic Nematodes ............................................... . Summary .......................................................... .

SECTION III METHODS FOR BIOLOGICAL CONTROL

66

66 77

78

78 79 89 91 91

93

93 93 94 95 97 98

101

IMPLEMENTATION. . . .. . .. ... .... . .. .. . . .. ...... 103

CHAPTER 7 NATURAL ENEMY CONSERVATION

Introduction ........................................................... . concepts ................................... . ............ . Use of Agricultural Chemicals .................................... . Management of Soil, Water, and Crop Residues ............................. . Crop Patterns ......................................................... . Manipulation of Non-Crop Vegetation .................................... . Providing Food or Shelter .............................................. . Safety ........................................................... .

CHAPTER 8 INTRODUCTION OF NEW NATURAL ENEMIES: PRINCIPLES .................................... .

Introduction ........................................................... . Case History Examples ......................................... . Outcomes of Biological Control Programs ......................... . Ecological Rationale for Natural Enemy Introduction ............... .

105

105 105 108 117 119 122 125 127

128

128 129 134 136

CONTENTS vii

Programs for Natural Enemy Introduction Costs, Social Benefits, and Safety ..

CHAPTER 9 INTRODUCTION OF NEW NATURAL ENEMIES:

139 155

METHODS ...................................... 158

Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158 The Collecting Trip. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158 Shipping Procedures .................................................... 162 Processing Material in Quarantine ......................................... 166 Safety ....................... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 170 Field Colonization Procedures ............................................ 172

CHAPTER 10 AUGMENTATION OF PARASITOIDS, PREDATORS, AND BENEFICIAL HERBIVORES. . . . . . . . . . . . . . . . . .. 178

Introduction. .......................................................... 178 Agent Selection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 178 Quality. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180 Cost ............................ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184 Storage and Shipping. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190 Application Methods ....................... . . . . . . . . . . . . . . . . . . . . . . . . . 191 Product Evaluation ............................ . . . . . . . . . . . . . . . . . . . . . . . . . . 193 Market Development and Continuity. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196 Natural Enemies in Current Use .... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197 Safety ........... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 198

CHAPTER 11 AUGMENTATION OF PATHOGENS AND NEMATODES ................................... 201

Introduction . 201 Agent Selection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 201 Production . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 204 Storage and Shipping. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 214 Formulation and Application ................ ............................ 215 Evaluation ............................................................. 219 Product Development ................................................... 225 Safety . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 228

CHAPTER 12 METHODS FOR BIOLOGICAL CONTROL OF PLANT PATHOGENS ................................... 235

Introduction . .. ........................................................ 235 Paradigms of Biological Control of Plant Pathogens .......................... 235 Habitat Characteristics ................................................... 236 Mechanisms of Biological Control of Plant Pathogens ........................ 238 Conservation ................ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239 Augmentation ........... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 244 Developing and Using Beneficial Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 255

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SECTION IV EVALUATION AND INTEGRATION

CHAPTER 13 NATURAL ENEMY MONITORING AND EVALUATION .................................. .

Introduction ........................................................... . Surveys to Document Natural Enemy-Pest Associations ...................... . Monitoring Progress of Introduction Projects ............................... . Sampling Natural Enemies for Pest Management ............................ . Evaluation of Population Effects of Natural Enemies ......................... . Simulation Models and the Evaluation of Natural Enemies ................... . Documentation, Economic Evaluation, and Planning ........................ . Planning .............................................................. .

CHAPTER 14 INTEGRATION OF BIOLOGICAL CONTROL INTO PEST MANAGEMENT SYSTEMS .................. .

Introduction ........................................................... . Pest Management Systems: Foundations, and Transitions .. , .................. . Combinations of Factors and Their Integration ....... , .................... ,. Integrating Natural Enemies into Pest Management Systems .................. .

SECTION V NATURAL ENEMY BIOLOGY ..................... .

CHAPTER 15 BIOLOGY OF ARTHROPOD PARASITOIDS AND PREDATORS ................................... .

Introduction ....................................... , .. , , ............... . Finding Habitats and Hosts .............................................. . Host Patch Exploitation and Abandonment Patterns . , ... , .. , ...... , . , ..... , .. Host Recognition and Assessment ... ,", ................................ ,. Overcoming Host or Prey Defenses ...................... , ........... , .... . Host Regulation and Physiological Interactions ....... , ..................... . Learning by Natural Enemies ............................. , .............. . Effects of Plant Features on Natural Enemies ............................... . Synchrony and Natural Enemy Races ............... , ...... , ....... , ...... .

CHAPTER 16 BIOLOGY AND DYNAMICS OF PATHOGENS ..... .

Introduction .............. , .................. , , .. , ....... , ............. . Basic Processes in Pathogen Biology ..................... , ............... , . Epizootiology of Arthropod Pathogens .................................... . Bacterial Pathogens of Arthropods ............ , ........... , ............... , Viral Pathogens of Arthropods ............... ' .......... , ........... , .... . Fungal Pathogens of Arthropods ..................... , ................... , Protozoan Pathogens of Arthropods ... , ........................... , ...... . Nematodes Pathogenic in Arthropods . , ................................... . Case Histories I. Two Viruses ..................... , ................... , ..

257

259

259 259 260 261 264 293 294 295

296

296 296 300 303

307

309

309 309 316 320 325 329 331 333 335

337

337 337 342 345 346 347 348 349 351

CONTENTS ix

Case Histories II. Case Histories III.

Two Bacteria ......................................... . Two Nematodes ...................................... .

CHAPTER 77 BIOLOGY OF WEED CONTROL AGENTS

Introduction ............................................... . Herbivorous Invertebrates ............................................... . Plant Pathogens ................... . ............... . Herbivorous Fish ................................................. .

CHAPTER 78 POPULATION REGULATION THEORY AND IMPLICATIONS FOR BIOLOGICAL CONTROL

Introduction ........................................................... . Single-Species Populations .............................................. . Interspecific Competition ............................................... . Host-Parasitoid Systems ................................................. . Host-Pathogen Systems ................................................. . Multispecies Systems ................................................... . Implications for Biological Control ....................................... . Biological Control Systems and Ecological Evaluations , ...................... .

SECTION VI ADDITIONAL TOPICS .......................... .

CHAPTER 79 THE ROLE OF GROWER EDUCATION IN BIOLOGICAL CONTROL ........................ .

Introduction ...................................... , ......... , .......... . Type of Information Needed by Growers ........................... , ..... , . Training Media ........ ,................. . ...... ' ..................... . Training of Extension Agents ...... , ...... , ...... , ... , ................... . Teaching Biological Control to Politicians ...... , ........................... . Communicating with Environmental Groups ............................... .

CHAPTER 20 GOVERNMENT POLICY AND BIOLOGICAL CONTROL .................................... .

Introduction .. , .......... . Pesticide Policies ........ ,. Quarantine and Eradication Policies , .......... , ......... , ................ . Agricultural Development Policies ........................................ . Pest Management Policies ............................. , ... , ............ , . Biological Control Policies ...... . . . . . . . . . . . . . . . . .............. . Use of Genetically-Modified Organisms .................... , .......... , ... .

CHAPTER 27 BIOLOGICAL CONTROL IN SUPPORT OF NATURE CONSERVATION ............................... .

Introduction ............................................... , ........... . Restorative Ecology through Biological Control ... , .................... , .... .

351 352

354

354 354 364 365

367

367 369 372 375 390 393 397 398

399

401

401 401 408 409 413 414

415

415 415 417 419 420 421 423

424

424 424

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Factors Promoting Adventive Species Invasions . . . . . . . . . . . . . . . . . . . . . . . . 426 Effects of Adventive Species ............................. . . . . . . . . . 428 Biological Control for Conservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 433 Suppressing Adventive Species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 441 Safety ................................................................. 442

CHAPTER 22 FUTURE PROSPECTS 444

REFERENCES 447

INDEX...................................................... 522

PREFACE

Nearly twenty years have passed since the publishing of a broadly-based texthook on hiolog­ical control (Huffaker and Messinger 1976). In the interim, other works on biological control have been either briefer treatments (DeBach and Rosen 1991), or collections of essays on selected topics (Waage and Greathead 1986; Mackauer and Ehler 1990). Our text has been written to fill what we believe is a need for a well-integrated, broadly-based text of appropriate length and degree of technical detail for teaching a one semester upper level course in hiological control. We have attempted to focus on principles and concepts, rather than on biological control of particular taxa or hiological control by particular kinds of natural enemies. Therefore, for example, the reader will find the material on biological control of weeds integrated with biological control of insects and mites into chapters on principles, techniques, and applications rather than presented separately. Only biological control of plant pathogens is addressed independently, an appraoch made necessary by the many special features and concepts in plant pathogen biological control.

In addition to essential material on such expected topics as natural enemy introduction, taxonomy of natural enemies, and the history of the discipline, we have sought to emphasize several areas we view as of special importance. In particular, we have emphasized natural enemy evaluation (Chapter 13) as a key feature of biological control studies. We have devel­oped new treatments on conservation of natural enemies (Chapter 7) and biological control of environmental pests (Chapter 21). We have also explored topics of a broader nature, such as the philosophy of integration of biological control in pest management systems (Chapter 14), the role of extension in teaching biological control to farmers (Chapter 19), and the effects of government policies on the degree to which biological control is employed (Chapter 20).

This text has been written to encourage training of a new generation of biological control scientists committed to hoth the understanding of biological control and its safe use to solve pest problems. Biological control, while enjoying a century of practical use, is a critically important means of addressing current pest problems in both crop protection and the preserva­tion of natural systems. It is our hope that this text will contribute to the training of scientists who achieve tomorrow's successes in biological control.

Preparation of this book benefitted in many ways from interactions with and contributions from colleagues. The hook's early development and outline was aided by discussions with J. Waage and P. Kenmore. Reviews of the book in part or whole, were contributed by F. Bigler, C. Campbell, R. Charudattan, W. Coli, J. Coulson, E. Delfosse, J. Elkinton, T. Fisher, H. Frank, L. Gilkeson, R. Goeden, D. Headrick, M. Hoddle, M. Hoffmann, D. Hogg, M.Johnson, H. Kaya, J. van Lenteren, N. Lepla, J. Lewis, C. McCoy, D. Meyerdirk, O. Minkenberg, R. Prokopy, M. Samways, M. Schauff, J. Sutton, B. Vinson, J. Waage, and M. Wilson. M. Hassell assisted in composing portions of Chapter 18. Assistance in checking the names and authors of species in

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the text was provided by C. Breidenbaugh, C. Lombardi, C. Meisenbacher, and B. Orr.]. Rewa prepared many revisions of the typescript. Our thanks also go to Greg Payne, at Chapman & Hall, who has been unwavering in his patience and support, and to those who encouraged our early interest in entomology and biological control, P. Ritcher,]. Lattin, E. Dickason, S. Knapp,]. Owens, W. Whitford, M. Hassell, and our colleagues at the University of Massachusetts and the University of California. We also extend a special thanks to our wives and families for their patience and support in this project.

R. G. Van Drlesche T. S. Bellows

Biological Control