STERILE INSECT TECHNIQUE Sterile Insect Technique Principles and Practice in Area-Wide Integrated Pest Management Edited by V. A. DYCK J. HENDRICHS and A.S. ROBINSON Joint FAO/IAEA Programme Vienna, Austria A C.I.P. Catalogue record for this book is available from the Library of Congress. ISBN-10 1-4020-4050-4 (HB) ISBN-13 978-1-4020-4050-4 (HB) ISBN-10 1-4020-4051-2 ( e-book) ISBN-13 978-1-4020-4051-1 (e-book) Published by Springer, P.O. Box 17, 3300 AADordrecht, The Netherlands. www.springeronline.com Printed on acid-free paper Photo Credits: A.S. Robinson and M.J.B. Vreysen provided some of the photos used on the front and back covers. All Rights Reserved © 2005 IAEA All IAEA scientific and technical publications are protected by the terms of the Universal Copyright Convention on Intellectual Property as adopted in 1952 (Berne) and as revised in 1972 (Paris). The copyright has since been extended by the World Intellectual Property Organization (Geneva) to include electronic and virtual intellectual property. Permission to use whole or parts of texts contained in IAEA publications in printed or electronic form must be obtained and is usually subject to royalty agreements. Proposals for non- commercial reproductions and translations are welcomed and considered on a case-by-case basis. Inquiries should be addressed to the Publishing Section, IAEA, Wagramer Strasse 5, A-1400 Vienna, Austria. Printed in the Netherlands. PREFACE It is a challenge to bring together all relevant information about the sterile insect technique (SIT) and its application in area-wide integrated pest management (AW- IPM) programmes; this book is the first attempt to do this in a thematic way. Since SIT practitioners tend to operate in the context of only one insect pest species, it was also a challenge for authors to develop and write their chapters from generic and global points of view, stressing the principles of the technology, and including examples from a range of pest species. We appreciate the understanding shown by the authors in accepting our many suggestions to emphasize the principles of the technology and to minimize details of field programmes. We also thank them for their patience with the prolonged editing process of the book. We are especially grateful to the authors for writing the chapters without financial compensation. Authors who are retired, or worked on their own time, deserve special commendation. Each chapter was peer-reviewed, and we thank the reviewers for helping to make the book accurate, complete, up-to-date, and generic in content. The need for this book has been evident for many years, and now that it has finally been published, it is expected to serve the scientific community for many years to come. We are pleased to have been able to participate in its development. The Editors March 2005 DISCLAIMER This publication has been prepared from the original material submitted by the authors, and then edited and revised by the editors according to style guidelines acceptable to the publisher. The views expressed do not necessarily reflect those of the Food and Agriculture Organization of the United Nations (FAO), the International Atomic Energy Agency (IAEA), and the governments of the Member States. The use of particular designations of countries or territories does not imply any judgement by the publisher, the FAO or the IAEA as to the legal status of such countries or territories, of their authorities and institutions or of the delimitation of their boundaries. The mention of names of specific companies or products (whether or not indicated as registered) does not imply any intention to infringe proprietary rights, nor should it be construed as an endorsement or recommendation on the part of the FAO or IAEA. The authors are responsible for having obtained the necessary permission for the reproduction, translation or use of material from sources already protected by copyrights. FOREWORD For several major insect pests, the environment-friendly sterile insect technique (SIT) is being applied as a component of area-wide integrated pest management (AW-IPM) programmes. This technology, using radiation to sterilize insects, was first developed in the USA, and is currently applied on six continents. For four decades it has been a major subject for research and development in the Joint FAO/IAEA Programme on Nuclear Techniques in Food and Agriculture, involving both research and the transfer of this technology to Member States so that they can benefit from improved plant, animal and human health, cleaner environments, increased production of plants and animals in agricultural systems, and accelerated economic development. The socio-economic impacts of AW-IPM programmes that integrate the SIT have confirmed the usefulness of this technology. Numerous publications related to the integration of the SIT in pest management programmes, arising from research, coordinated research projects, field projects, symposia, meetings, and training activities have already provided much information to researchers, pest-control practitioners, programme managers, plant protection and animal health officers, and policy makers. However, by bringing together and presenting in a generic fashion the principles, practice, and global application of the SIT, this book will be a major reference source for all current and future users of the technology. The book will also serve as a textbook for academic courses on integrated pest management. Fifty subject experts from 19 countries contributed to the chapters, which were all peer reviewed before final editing. vii INTRODUCTORY REMARKS As evidenced by the successful area-wide insect pest control programmes described in this book, the sterile insect technique (SIT), a component of these programmes, has come of age. The technology has expanded rapidly — additional target species, new rearing techniques, studies on genetics and insect behaviour, and especially integration into operational area-wide integrated pest management (AW-IPM) programmes. The SIT has matured to the point where a critical overview of its principles and practice will greatly facilitate further research, development, and application in the field. The SIT was among the first biological insect control methods designed for area- wide application. While the SIT gained its reputation in insect eradication programmes, it is essential that the scientific community now recognizes its potential as a part of IPM strategies for the area-wide suppression, containment, prevention and, where advisable, eradication of pests. th Insect control methods in the first 70 years of the 20 century were based largely on chemical insecticides; this was especially so after the Second World War with the introduction of synthetic insecticides. The concept of IPM became popular after 1970, and a more selective use of insecticides was emphasized. Attempts to significantly reduce insecticide applications have only gradually become more prominent. Biological control of pest insects, together with the breeding of insect- tolerant or resistant plants, is probably now receiving the major emphasis in IPM programmes. According to an international standard under the International Plant Protection Convention (IPPC), the SIT is now officially considered as one type of biological control, and it is ideally suited for incorporation into AW-IPM programmes. The scientific underpinning of SIT programmes has broadened as new areas of science have developed, e.g. insect mass production and quality, geographic information systems and data management systems, genetics and molecular biology, insect behaviour, aerial release of sterile insects, and modelling of AW-IPM. The practical success of a programme incorporating the SIT requires a holistic and multidisciplinary approach, and effective management, since in the last analysis programmes must produce substantial economic benefits. This is clearly evident in the major successes using the SIT against screwworms, fruit flies, and moths. In spite of documented successes, many colleagues in the scientific community are partially or inadequately informed on the application and importance of this powerful addition to the biological weapons that can be used against insect pests that are economically important or a threat to human health. The credibility and impact of the technology needs to be described in an objective, comprehensive, and balanced fashion, and in an accessible format. New insect pest problems, new restrictive legislation, as well as older problems such as insecticide resistance and minimum residue levels, require new solutions. There is a real need, and an x increasing demand, for information on the SIT so that its potential for addressing some of these problems can be assessed. The chapters have been written by well-known experts on the SIT and other technologies that are integrated into IPM systems. A “first” in its field and worldwide in scope, this book will provide an in-depth resource for the whole range of documented scientific information about the SIT. The target audience of the book is the scientific community worldwide. It will assist animal health and plant protection practitioners, as well as students, teachers, and researchers, in understanding and applying the SIT. It is anticipated that the book will have a considerable impact on the science and practice of pest control systems. Research workers new to this field have difficulty accessing the literature — it tends to be widely scattered in multiple publications (some with very limited distribution), in conference proceedings, and in unpublished programme reports. To further the science and application of the SIT, the accumulated knowledge and experience needs to be integrated and synthesized from a generic standpoint. The consolidation of comprehensive information into one volume, with references to the large amount of previous work, is long overdue. Such a consolidation will facilitate the application of the SIT to those pest problems for which it is appropriate. It will also lay the groundwork for future applications. The present book is uniquely designed to fill this gap. The strengths and weaknesses, and successes and failures, of the SIT have rarely been evaluated openly and fairly from a scientific perspective. This is just the beginning. This book will help develop further the use of the SIT for pest suppression, and where advisable, eradication. It will be a gold mine for graduate students who want to learn about the history, accomplishments, problems, and promises of the SIT. As an “autocidal” biological control method, it fits into present-day concerns regarding human health and the environment. There is great potential for significant advances that will make the SIT more effective and economically viable, such as commercializing the different components, developing genetic sexing strains that permit the release of only males, treating sterile insects hormonally and semiochemically to increase their quality and competitiveness, releasing insects from improved aerial systems, and using modern biotechnology. It is an honour to have been asked to write these introductory remarks. The developments in this technology are exciting, and I will always remain a part of them. Maurice Fried Director, Research Support Program National Academy of Sciences 500 Fifth Street, NW Washington, DC 20001, USA TABLE OF CONTENTS PART I. INTRODUCTION Chapter 1.1. History of the Sterile Insect Technique..................................................................3 W. Klassen and C. F. Curtis PART II. PRINCIPLES OF THE STERILE INSECT TECHNIQUE Chapter 2.1. Area-Wide Integrated Pest Management and the Sterile Insect Technique.....39 W. Klassen Chapter 2.2. Biological Basis of the Sterile Insect Technique..................................................69 D. R. Lance and D. O. McInnis Chapter 2.3. Genetic Basis of the Sterile Insect Technique......................................................95 A. S. Robinson Chapter 2.4. Inherited Sterility in Insects................................................................................115 J. E. Carpenter, S. Bloem and F. Marec Chapter 2.5. Mathematical Models for the Use of Sterile Insects..........................................147 H. J. Barclay PART III. TECHNICAL COMPONENTS OF THE STERILE INSECT TECHNIQUE Chapter 3.1. Role of Population and Behavioural Ecology in the Sterile Insect Technique..............................................................................................................177 Y. Itô and K. Yamamura xii Chapter 3.2. Mass-Rearing for Sterile Insect Release.............................................................209 A. G. Parker Chapter 3.3. Sterilizing Insects with Ionizing Radiation.........................................................233 A. Bakri, K. Mehta and D. R. Lance Chapter 3.4. Sterile Insect Quality............................................................................................269 C. O. Calkins and A. G. Parker Chapter 3.5. Sterile Insect Supply, Emergence, and Release..................................................297 R. V. Dowell, J. Worley and P. J. Gomes Chapter 3.6. Monitoring Sterile and Wild Insects in Area-Wide Integrated Pest Management Programmes...................................................................................325 M. J. B. Vreysen Chapter 3.7. Procedures for Declaring Pest Free Status.........................................................363 H. J. Barclay, J. W. Hargrove, A. Clift and A. Meats PART IV. SUPPORTIVE TECHNOLOGIES TO IMPROVE THE STERILE INSECT TECHNIQUE Chapter 4.1. Role of Population Genetics in the Sterile Insect Technique............................389 E. S. Krafsur Chapter 4.2. Population Suppression in Support of the Sterile Insect Technique................407 R. L. Mangan Chapter 4.3. Genetic Sexing Strains in Mediterranean Fruit Fly, an Example for Other Species Amenable to Large-Scale Rearing for the Sterile Insect Technique..............................................................................................................427 G. Franz Chapter 4.4. Use of Geographic Information Systems and Spatial Analysis in Area-Wide Integrated Pest Management Programmes that Integrate the Sterile Insect Technique.................................................................................453 J. St. H. Cox and M. J. B. Vreysen