ACADEMIC PRESS RAPID MANUSCRIPT REPRODUCTION Proceedings of the Symposium on Biological Control in Agricultural Integrated Pest Management Systems held at the Citrus Research and Education Center, University of Florida, at Lake Alfred, June 4-6, 1984. Biological Control in Agricultural IPM Systems Edited by Marjorie A. Hoy Department of Entomological Sciences University of California Berkeley, California and Donald C. Herzog Department of Entomology and Nematology University of Florida Gainesville, Florida and Agricultural Research and Education Center Quincy, Florida 1985 ACADEMIC PRESS, INC. (Harcourt Brace Jovanovich, Publishers) Orlando San Diego New York London Toronto Montreal Sydney Tokyo COPYRIGHT © 1985, BY ACADEMIC PRESS, INC. ALL RIGHTS RESERVED. NO PART OF THIS PUBLICATION MAY BE REPRODUCED OR TRANSMITTED IN ANY FORM OR BY ANY MEANS, ELECTRONIC OR MECHANICAL, INCLUDING PHOTOCOPY, RECORDING, OR ANY INFORMATION STORAGE AND RETRIEVAL SYSTEM, WITHOUT PERMISSION IN WRITING FROM THE PUBLISHER. ACADEMIC PRESS, INC. Orlando, Florida 32887 United Kingdom Edition published by ACADEMIC PRESS INC. (LONDON) LTD. 24-28 Oval Road, London NW1 7DX LIBRARY OF CONGRESS CATALOGING-IN-PUBLICATION DATA Main entry under title: Biological control in agricultural IPM systems. Proceedings of a conference held at the Citrus Research and Education Center of the University of Florida at Lake Alfred on June 4-6, 1984. Includes index. 1. Agricultural pests—Biological control—Congresses. 2. Agricultural pests—Biological control —United States—Congresses. 3. Agricultural pests—Integrated control —Congresses. 4. Agricultural pests— Integrated control —United States—Congresses. I. Hoy, Marjorie A. II. Herzog, D. C. (Donald C.) SB933.3.B548 1985 632'.96 85-9157 ISBN 0-12-357030-1 PRINTED IN THE UNITED STATES OF AMERICA 85 86 87 88 987654321 Contributors and Conference Participants Numbers in parentheses indicate the pages on which the authors' contributions begin. George S. Abawi (433), Department of Plant Pathology, New York State Agricultural Experiment Station, Cornell University, Geneva, New York 14456 *Perry L. Adkisson (41), Deputy Chancellor for Agriculture, Texas A&M University System, College Station, Texas 77843 * William W. Allen (572), Department of Entomological Sciences, University of California, Berkeley, California 94720 *Lloyd A. Andres (388), Biological Control of Weeds Laboratory, USDA/ARS, Albany, California 94607 *Ralph T. Baker (25), Department of Plant Pathology, Colorado State University, Ft. Collins, Colorado 80973 *Carl S. Barfield (88, 101, 121), Department of Entomology and Nematology, University of Florida, Gainesville, Florida 32611 Steven K. Beckendorf (167), Department of Molecular Biology, University of California, Berkeley, California 94720 Elizabeth H. Beers (103), Department of Entomology, Pennsylvania State University, University Park, Pennsylvania 16802 *Elizabeth A. Bernays (373), Division of Biological Control, University of California, Albany, California 94706 *Drion G. Boucias (263, 280), Department of Entomology and Nematology, University of Florida, Gainesville, Florida 32611 *Leopoldo E. Caltagirone (189), Division of Biological Control, University of California, Berkeley, California 94720 *James R. Cate (537), Department of Entomology, Texas A&M University, College Station, Texas 77843 *Raghaven Charudattan (347), Department of Plant Pathology, University of Florida, Gainesville, Florida 32611 *Those names with asterisks were conference participants; others contributed to various presentations at the conference and chapters to this proceedings. ix X Contributors Brian A. Croft (123), Department of Entomology, Oregon State University, CorvalliSy Oregon 97331 *Louis A. Falcon (229), Department of Entomological Sciences, University of California, Berkeley, California 94720 *Peter Faulkner (263), Department of Microbiology, Queen's University, Kingston, Ontario K7L 3N6, Canada *Donald L. Flaherty (501), Agricultural Building, County Civic Center, Visalia, California 93277 *Raymond E. Frisbie (41), Integrated Pest Management Coordinator, Texas A&M University, College Station, Texas 77843 * Joseph E. Funderburk(67), Department of Entomology and Nematology, University of Florida, North Florida Research and Education Center, Quincy, Florida 32351 *Daniel Gonzalez (50), Division of Biological Control, University of California, Riverside, California 92521 Joseph G. Hancock (415), Department of Plant Pathology, University of California, Berkeley, California 94720 * Joseph C. Headley (53), Department of Agricultural Economics, University of Missouri, Columbia, Missouri 65201 *Donald C. Herzog (3, 67, 557), Department of Entomology and Nematology, University of Florida, North Florida Research and Education Center, Quincy, Florida 32351 Harvey C. Hoch (433), Department of Plant Pathology, New York State Agricultural Experiment Station, Cornell University, Geneva, New York 14456 Keith R. Hopper (201), Southern Field Crops, Insect Management Research Laboratory, USDAIARS, Stoneville, Missouri 38776 *Marjorie A. Hoy (3,151,167), Department of Entomological Sciences, University of California, Berkeley, California 94720 *Carl B. Huffaker (13), Division of Biological Control, University of California, Berkeley, California 94720 *Larry A. Hull (103), Department of Entomology, Pennsylvania State University, Fruit Research Laboratory, Biglerville, Pennsylvania 17307 * Carlo M. Ignoffo (243), Biological Control of Insects Research Laboratory, USDAIARS, Columbia, Missouri 65201 *Harry K. Kaya (283), Division of Nematology, University of California, Davis, California 94616 Hiroshi Kido (501), Department of Entomology, University of California, Davis, California 95616 *Edgar G. King (201), Southern Field Crops Insect Management Research Laboratory, USDAIARS, Stoneville, Missouri 38776 *W. Joe Lewis (89), Insect Biology and Population Management Research Laboratory, USDAIARS, Tifton, Georgia 31793 *StevenE. Lindow(395), Department of Plant Pathology, University of California, Berkeley, California 94720 Preface χι * James J. Marois (461), Department of Plant Pathology, University of California, Davis, California 95616 *S. Bruce Martin (433), Department of Plant Pathology, North Carolina State University, Raleigh, North Carolina 27650 *Clayton W. McCoy (481), Department of Entomology and Nematology, University of Florida, Citrus Research and Education Center, Lake Alfred, Florida 32850 *Christopher A. Mullin (123), Department of Entomology, Pennsylvania State University, University Park, Pennsylvania 16802 *Carolyn Napoli (455), Department of Plant Pathology, University of California, Berkeley, California 94720 Donald A. Nordlund (89), Insect Biology and Population Management Research Laboratory, USDAIARS, Tif ton, Georgia 31793 *Robert J. O'Neil1 (323), Department of Entomology and Nematology, University of Florida, Gainesville, Florida 32611 Janine E. Powell (201), Southern Field Crops insect Management Research Laboratory, USDAIARS, Stoneville, Missouri 38776 *Reece I. Sailer (166, 187, 200, 226), Department of Entomology and Nematology, University of Florida, Gainesville, Florida 32611 *Milton N. Schroth (415), Department of Plant Pathology, University of California, Berkeley, California 94720 Merle Shepard2 (557), Department of Entomology and Zoology, Clemson University, Clemson, South Carolina 29631 *Grover C. Smart (302, 478), Department of Entomology and Nematology, University of Florida, Gainesville, Florida 32611 Brian J. Staskawicz (455), Department of Plant Pathology, University of California, Berkeley, California 94720 Vernon M. Stern (501), Department of Entomology, University of California, Riverside, California 92521 * Jerry L. Stimac (323), Department of Entomology and Nematology, University of Florida, Gainesville, Florida 32611 *Ronald E. Stinner (62, 150, 322, 344), Department of Entomology, North Carolina State University, Raleigh, North Carolina 27650 *Maurice J. Tauber (3), Department of Entomology, Cornell University, Ithaca, Ne\ York 14853 * Seymour D. Van Gundy (467), Department of Nematology, University of California, Riverside, California 92507 *L. Theodore Wilson (303, 501), Department of Entomology, University of California, Davis, California 95616 * Kenneth V. Yeargan (521), Department of Entomology, University of Kentucky, Lexington, Kentucky 40546 'Present address: Department of Entomology, Purdue University, West Lafayette, Indiana 47907. 2Present address: Department of Entomology, International Rice Research Institute, Manila, Philippines. Preface This volume on "Biological Control in Agricultural IPM Systems" evolved, at least in part, as an outgrowth of a trip to the Peoples' Republic of China in 1982. As members of a ten-person National Academy of Sciences Team on biological control of arthropod pests, we were able to contrast the use of biological control in Chinese agriculture and in the United States, and we subsequently asked the question, How can biological control be used more extensively and effectively in U.S. agriculture? Further impetus to organization of the conference from which this volume arose came during a National Interdisciplinary Biological Control Conference held in Las Vegas, Nevada, in February, 1983. The large number of participants attending that con- ference suggested that interest in biological control was expanding. Originally, we outlined "Biological Control in Agricultural IPM Systems" as a one-half day symposium for presentation at the 1983 or 1984 annual meeting of the Entomological Society of America. However, our half-day symposium quickly grew into a three-day conference and we realized that a separate work conference was needed to cover the subject adequately, and that participation of and input was needed from entomologists, plant pathologists, weed scientists, nematologists, toxicolo- gists, and economists. Thus the conference was conceived and held on June 4-6, 1984, at the Citrus Research and Education Center of the University of Florida at Lake Alfred. Participants in the conference worked diligently to review critically the status of biological control in their areas of expertise and as an implementable component in agricultural IPM systems. We are not aware o fany comparable review of the subject matter in print. While there are several important reference volumes on both biologi- cal control and on IPM, none have focused specifically on the role and status of biological control within the text of agricultural IPM systems. Because of the complexity of the subject matter, we deemed it desirable to limit our discussions to agriculture in the United States and to exclude animal, rangeland, urban, and glasshouse IPM systems. Many students in recent years have been taught that biological control and plant resistance are cornerstones of IPM. Despite the supposed, and often-quoted status of biological control as a basic subdiscipline underlying IPM, research to incorporate biological control into agricultural cropping management systems is still relatively uncommon, and funding for such effort is scarce. It is our perception that IPM xiii XIV Preface program managers have often tended to support projects that focused on research designed to time pesticide applications effectively, to model plant/pest interactions, to identify and/or refine economic injury levels, and to predict pest populations. These subjects are certainly important and deserve the attention that they have received. However, funds for biological control—whether by importation, augmen- tation, or conservation—probably constitute 20% or less of the funding allocated to IPM programs. Biological control research has therefore coped with extraordinarily limited resources, often not keeping up with inflation. This perception raised in our minds a series of questions relative to biological con- trol programs and resources (sources and adequacy of funding) for such programs. The following questions were articulated in the introduction to the conference and were subsequently discussed, both overtly and covertly, by participants during the three days: • Is biological control currently an integral part of agricultural IPM programs in U.S. agriculture? Are the costs, risks, and benefits associated with and accruing to biological control sufficient to justify the statement that it is, along with plant resis- tance, a foundation upon which IPM can rest? • If biological control is not fully implemented and/or utilized in U. S. agricul- ture, should more research and especially more funding be directed toward such im- plementation and utilization? • What factors limit the expanded use of biological control in agriculture? Is it limited only because of a lack of necessary funding? Or, are new attitudes, approaches, or technologies needed to fully exploit the potential of biological con- trol? For example, can genetic engineering technologies be useful tools to improve biological control of arthropod pests, weeds, plant pathogens, or nematodes? • Integrated pest management is described as an interdisciplinary approach to the solution of agricultural problems. Is such cooperation only painfully, and too rarely, achieved? If so, are interdisciplinary meetings such as this conference fruitful? Or, are our individual disciplines so filled with incomprehensible jargon, highly detailed research needs, and divergent interests that our efforts to communicate are a waste of time? Can we exchange ideas, concepts, and perspectives in order to provide a new focus in our approach to biological control in agricultural IPM systems? During our conference, we attempted to scrutinize very carefully the current status of biological control in our agricultural IPM systems in a broad sense. We also provided perspectives and perceptions of some of the more specific research needed to enhance the use of biological control of arthropod pests, weeds, plant pathogens, and nematodes in agricultural IPM systems. We hope that in doing so we have fostered an expanded horizon for biological control. We recognize that new beginnings do not develop overnight, but we do hope that our conference and this proceedings volume will be a constructive contribution to the development of a new horizon for biological control. That there have been two large national meetings in the last two years on biological control suggests that we may be at the threshold of a new era relative to biological control. Integrated pest management has undergone evolutionary changes during the Preface xv past 20 years. Many important advances have been made in managing pests. The important role of biological control organisms in regulating pests, especially insects and mites, of agricultural crops was not fully recognized until they were decimated by applications of pesticides harmful to the beneficial species, but not to the pest. Overt utilization of biological control agents has only just begun; much yet remains to be done to make biological control of arthropod pests, weeds, plant pathogens, and nematodes reliable and accepted components in agricultural IPM programs. Reliance on pesticides as the major, in some cases sole, suppression tactic in IPM is seen as having an increasingly limited future. Thus, while pesticides will continue to be, for the forseeable future, important in IPM programs, problems surrounding their con- tinued use are expanding and intensifying. Pesticide resistance among pests is a major concern. Increased restrictions on pesticide use and uncertainty about the discovery and development of replacement chemicals gives great cause for concern. Biological control is highly compatible with quarantines, plant resistance, and cultural controls. The appropriate and full deployment of biological control in agricultural IPM systems remains to be achieved. We hope that this volume will assist in this endeavor. Finally, we must thank many people, not least of whom are the conference par- ticipants, who unselfishly supported the aims and goals of the conference. Crucial financial support was provided by the University of Florida, Institute of Food and Agricultural Sciences; the University of California; and the National Science Founda- tion Office of Interdisciplinary Research, which provided funds to "foster a compre- hensive, interdisciplinary view of research needed to advance biological control as a strategy in integrated pest management." The editors express special appreciation to the following employees of the North Florida Research and Education Center, Uni- versity of Florida, who worked tirelessly, spending many long hours preparing the various versions of this volume: Ms. J.M. Waiden, Ms. K.M. Lurding, and Ms. J. V. Smith. Without their assistance and cooperation, completion of this project would have been extremely difficult. Marjorie A. Hoy Donald C. Herzog BIOLOGICAL CONTROL IN AGRICULTURAL IPM SYSTEMS: A BRIEF OVERVIEW OF THE CURREHT STATUS ATO FUTURE PROSPECTS N.J. Tauber, M.A. Hoy and D-C. Herzog Our Conference was qoncerned with biological control in agricul- tural IPM systems, a subject of considerable current interest because of a perceived urgency to develop and adopt safe and efficient methods for managing agricultural pests. Problems associated with pest suppression (including environmental pollution, deleterious effects of pesticides on non-target organisms, pesticide resistance, resurgence of target pests, secondary pest outbreaks, escalating costs of developing, producing and applying pesticides) all affect the vitality and profitability of agriculture and the well-being of our society. General challenges of the Conference were to: 1) provide up-to-date reviews of past work on biological control of arthro- pods, weeds, nematodes and plant pathogens in the context of agriculture, 2) identify the status of biological control in cur- rent IPM programs, and 3) identify research required to accomp- lish greater integration of biological control into agricultural IPM systems. In many, if not most cases, biological control by itself does not provide economically acceptable pest suppression in agricul- tural cropping systems. Therefore, biological control must be developed and implemented as a component of IPM. However, if it is to be an integral part of IPM (along with plant resistance, cultural methods and pesticidal controls) biological control must be nurtured to become a strong, vital entity. This is especially true, as stressed in many of the following chapters, if biologi- cal control is to be a vital component of agricultural IPM sys- tems. NATIONAL ISSUES The papers presented and subsequent discussions at the Conference addressed crucial biological control and integrated pest manage- ment issues and issued specific recommendations. These are pre- sented in individual papers and discussion summaries; they will not be repeated here in detail. In this summary, emphasis will BIOLOGICAL CONTROL Copyright © 1985 by Academic Press, Inc. IN AGRICULTURAL IPM SYSTEMS 3 H rigAhts of reproduction in any form reserved. ISBN 0-12-357030-1