Quality Control and Production of Biological Control Agents Theory and Testing Procedures i Quality Control and Production of Biological Control Agents Theory and Testing Procedures Edited by J.C. van Lenteren Laboratory of Entomology Wageningen University Wageningen The Netherlands CABIPublishing CABIPublishingis a division of CABInternational CABI Publishing CABI Publishing CAB International 44 Brattle Street Wallingford 4th Floor Oxon OX10 8DE Cambridge, MA02138 UK USA Tel: +44 (0)1491 832111 Tel: +1 617 395 4056 Fax: +44 (0)1491 833508 Fax: +1 617 354 6875 E-mail: [email protected] E-mail: [email protected] Web site: www.cabi-publishing.org © CAB International 2003. All rights reserved. No part of this publication may be reproduced in any form or by any means, electronically, mechani- cally, by photocopying, recording or otherwise, without the prior permis- sion of the copyright owners. A catalogue record for this book is available from the British Library, London, UK. Library of Congress Cataloging-in-Publication Data Quality control and production of biological control agents : theory and testing procedures / edited by J.C. van Lenteren. p. cm. Includes bibliographical references (p. ). ISBN 0-85199-688-4 1. Biological pest control agents. 2. Biological pest control agents industry--Quality control. I. Lenteren, J.C. van. SB975 .Q35 2003 632’.96--dc21 2002151406 ISBN 0 85199 688 4 Typeset by ColumnsDesign Ltd, Reading, Berkshire Printed and bound in the UK by Biddles Ltd, Guildford and King’s Lynn Contents Contributors vii Preface ix Acknowledgements xi PARTI. QUALITYCONTROLFORNATURALENEMIES 1. Need for Quality Control of Mass-produced Biological Control Agents 1 J.C. van Lenteren 2. Aspects of Total Quality Control for the Production of Natural Enemies 19 N.C. Leppla PARTII. VARIABILITYINFORAGINGBEHAVIOUROFNATURALENEMIES 3. AVariable-response Model for Parasitoid Foraging Behaviour 25 L.E.M. Vet, W.J. Lewis, D.R. Papaj and J.C. van Lenteren 4. Variations in Natural-enemy Foraging Behaviour: Essential Element of a Sound Biological-control Theory 41 W.J. Lewis, L.E.M. Vet, J.H. Tumlinson, J.C. van Lenteren and D.R. Papaj 5. The Parasitoids’Need for Sweets: Sugars in Mass Rearing and Biological Control 59 F.L. Wäckers PARTIII. COPINGWITHVARIATIONINFORAGINGBEHAVIOUR 6. Managing Captive Populations for Release: a Population-genetic Perspective 73 L. Nunney 7. Adaptive Recovery after Fitness Reduction: the Role of Population Size 89 R.F. Hoekstra 8. The Use of Unisexual Wasps in Biological Control 93 R. Stouthamer 9. Comparison of Artificially vs. Naturally Reared Natural Enemies and Their Potential for Use in Biological Control 115 S. Grenier and P. De Clercq 10. Pathogens of Mass-produced Natural Enemies and Pollinators 133 S. Bjørnson and C. Schütte v vi Contents SECTIONIV. MASS-PRODUCEDNATURALENEMIES 11. Commercial Availability of Biological Control Agents 167 J.C. van Lenteren 12. Mass Production, Storage, Shipment and Release of Natural Enemies 181 J.C. van Lenterenand M.G. Tommasini 13. Regulation of Import and Release of Mass-produced Natural Enemies: a Risk-assessment Approach 191 J.C. van Lenteren, D. Babendreier, F. Bigler, G. Burgio, H.M.T. Hokkanen, S. Kuske, A.J.M. Loomans, I. Menzler-Hokkanen, P.C.J. van Rijn, M.B. Thomas and M.G. Tommasini SECTIONV. QUALITYCONTROLTESTINGOFNATURALENEMIES 14. Quality Assurance in North America: Merging Customer and Producer Needs 205 C.S. Glenister, A. Hale and A. Luczynski 15. State of Affairs and Future Directions of Product Quality Assurance in Europe 215 K.J.F. Bolckmans 16. The Relationship between Results from Laboratory Product-control Tests and Large-cage Tests Where Dispersal of Natural Enemies is Possible: a Case- study with Phytoseiulus persimilis 225 S. Steinberg and H. Cain 17. Quality of Augmentative Biological Control Agents: a Historical Perspective and Lessons Learned from Evaluating Trichogramma 231 R.F. Luck and L.D. Forster SECTIONVI. QUALITYCONTROLTESTS 18. Towards the Standardization of Quality Control of Fungal and Viral Biocontrol Agents 247 N.E. Jenkinsand D. Grzywacz 19. Guidelines for Quality Control of Commercially Produced Natural Enemies 265 J.C. van Lenteren, A. Hale, J.N. Klapwijk, J. van Schelt and S. Steinberg 20. Basic Statistical Methods for Quality-control Workers 305 E. Wajnberg Index 315 Contributors D. Babendreier, Swiss Federal Research Station for Agroecology and Agriculture, Reckenholzstrasse 191, CH-8046 Zurich, Switzerland. F. Bigler, Swiss Federal Research Station for Agroecology and Agriculture, Reckenholzstrasse 191, CH-8046 Zurich, Switzerland. S. Bjørnson,Department of Biology, Saint Mary’s University, 923 Robie Street, Halifax, Nova Scotia, Canada B3H 3C3. K.J.F. Bolckmans, Koppert Biological Systems, PO Box 155, 2650AD Berkel and Rodenrijs, The Netherlands. e-mail [email protected] G. Burgio,Department of Agroenvironmental Sciences and Technologies (DISTA), University of Bologna, via F. Re 6, 40126 Bologna, Italy. H. Cain,Bio-Bee Biological Systems Sde Eliyahu, Bet Shean Valley, 10810 Israel. P. De Clercq, Laboratory of Agrozoology, Department of Crop Protection, Faculty of Agricultural and Applied Biological Sciences, Ghent University, Coupure Links 653, B-9000 Ghent, Belgium. L.D. Forster,Department of Entomology, University of California, Riverside, CA92521, USA. C.S. Glenister, IPM Laboratories, Inc., 980 Main Street, Locke, NY 13092-0300, USA. e-mail [email protected] S. Grenier, UMR INRA/INSA de Lyon, Biologie Fonctionnelle, Insectes et Intéractions, Institut National des Sciences Appliquées, Bât. Pasteur, 20 av. A. Einstein, 69621 Villeurbanne Cedex, France. e-mail [email protected] D. Grzywacz,CABI Bioscience, Silwood Park, Ascot, Berkshire SL5 7TA, UK. A. Hale,The Bug Factory, 1636 East Island Highway, Nanoose Bay, British Columbia, Canada V9P9A5. R.F. Hoekstra, Laboratory of Genetics, Wageningen University, Arboretumlaan 4, 6703BD Wageningen, The Netherlands. e-mail [email protected] H.M.T. Hokkanen,Department of Applied Biology, University of Helsinki, Finland. N.E. Jenkins, CABI Bioscience, Silwood Park, Ascot, Berkshire SL5 7TA, UK. e-mail n.jenk- [email protected] J.N. Klapwijk,Koppert Biological Systems, PO Box 155, 2650AD Berkel and Rodenrijs, The Netherlands. S. Kuske, Swiss Federal Research Station for Agroecology and Agriculture, Reckenholzstrasse 191, CH-8046 Zurich, Switzerland. vii viii Contributors N.C. Leppla, Department of Entomology and Nematology, University of Florida, Natural Area Drive, PO Box 110630, Gainesville, FL32611-0603, USA. e-mail [email protected]fl.edu W.J. Lewis, Insect Biology and Population Management Research Laboratory, USDA-ARS, PO Box 748, Tifton, GA31793, USA. e-mail [email protected] A.J.M. Loomans,Laboratory of Entomology, Wageningen University, PO Box 8031, 6700EH Wageningen, The Netherlands. R.F. Luck,Department of Entomology, University of California, Riverside, CA92521, USA. e- mail [email protected] A. Luczynski,Biobugs Consulting Ltd, 16279 30B Ave., Surrey, British Columbia, Canada V4P 2X7. I. Menzler-Hokkanen, Department of Applied Biology, University of Helsinki, Finland. L. Nunney, Department of Biology, University of California, Riverside, CA 92521, USA. e-mail [email protected] D.R. Papaj,Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, AZ 85721, USA. C. Schütte, Laboratory of Entomology, Wageningen Agricultural University, PO Box 8031, 6700EH Wageningen, The Netherlands. S. Steinberg, Bio-Bee Biological Systems Sde Eliyahu, Bet Shean Valley, 10810 Israel. e-mail [email protected] R. Stouthamer, Laboratory of Entomology, Wageningen University, PO Box 8031, 6700EH Wageningen, The Netherlands. Present address: Department of Entomology, University of California at Riverside, Riverside, CA92521, USA. e-mail [email protected] M.B. Thomas,CABI Bioscience, Silwood Park, Ascot, Berkshire SL5 7TA, UK. M.G. Tommasini,CRPV (Centro Ricerche Produzioni Vegetali), Via Vicinale Monticino 1969, 47020-Diegaro di Cesena (FC), Italy. e-mail [email protected] J.H. Tumlinson, Insect Biology and Population Management Research Laboratory, USDA- ARS, PO Box 14565, Gainesville, FL32604, USA. J.C. van Lenteren,Laboratory of Entomology, Wageningen University, PO Box 8031, 6700EH Wageningen, The Netherlands. e-mail [email protected] P.C.J. van Rijn,CABI Bioscience, Silwood Park, Ascot, Berkshire SL5 7TA, UK. J. van Schelt, Koppert Biological Systrems, PO Box 155, 2650AD Berkel and Rodenrijs, The Netherlands. L.E.M. Vet, Laboratory of Entomology, Wageningen University, PO Box 8031, 6700EH Wageningen, The Netherlands; and Netherlands Institure of Ecology, POBox 40, 6666 ZG Heteren, The Netherlands. e-mail [email protected] F.L. Wäckers, Laboratory of Entomology, Wageningen University, PO Box 8031, 6700 EH Wageningen, The Netherlands; and Netherlands Institute of Ecology, PO Box 40, 6666ZG Heteren, The Netherlands. e-mail [email protected] E. Wajnberg,INRA, 37 Blvd du Cap, 06600 Antibes, France. e-mail [email protected] Preface The use of biological control agents is increasing worldwide and there are now many companies mass producing and selling such organisms. However, there is a great need for quality control in the production and use of these natural enemies, because deterioration of mass-reared biological control agents leads to failures in pest management. The area of quality control is rather new for biological control workers. Therefore, the first book on this topic specifically for biological control agents contains several chapters with background information, before discussing the quality control guidelines that have recently been developed. The first section of the book is devoted to emergence of quality control for natural enemies. In Chapter 1 the need for quality control for mass-produced biological control agents is discussed. In Chapter 2 the aspects of total quality control for the production of nat- ural enemies are described. The second section of the book – the basis of variability in foraging behaviour of natural enemies – comprises chapters dealing with background information on sources of variation in behaviour that are regularly encountered, but not understood and often misinterpreted in mass rearing. In Chapters 3 and 4, factors are analysed that induce the variability in searching behaviour of natural enemies, and technologies are described that illustrate how to manage this variation. Searching behaviour is influenced by the insect’s genetic constitution, its physiological state and its experience. Chapter 5 presents an overview of the information on the topic of food ecology of natural enemies, and illustrates that a certain physiological state is needed before a natural enemy is able to search for hosts. These chapters make it clear that insight into behavioural variability in the foraging behaviour of natural enemies is a pre- requisite for proper mass rearing and efficient application of natural enemies in pest manage- ment. The third section focuses on how to cope with this variation. In Chapter 6 a population genetic perspective is given on how to manage captive populations. Examples of adaptation to captive rearing and of the trade-off with field performance are presented. Chapter 7 discusses the effects of a transfer of natural enemies from the field to a mass production facility, such as reduction of fitness and enhancing the possibility of fixation of deleterious mutations in the population by genetic drift. Ways to prevent these negative effects are presented. In Chapter 8 the possibilities and advantages of unisexual reproduction for biological control are discussed. Some evidence is found for two advantages of unisexual reproduction: (i) unisexuals are ix