An Introduction to Arthropod Pest Control J.R.M. Thacker Dept of Biological Sciences University of Paisley, UK The Pitt Building, Trumpington Street, Cambridge, United Kingdom The Edinburgh Building, Cambridge CB2 2RU, UK 40 West 20th Street, New York, NY 10011-4211, USA 477 Williamstown Road, Port Melbourne, VIC 3207, Australia Ruiz de Alarcón 13, 28014 Madrid, Spain Dock House, The Waterfront, Cape Town 8001, South Africa http://www.cambridge.org © J.R.M. Thacker 2002 This book is in copyright. Subject to statutory exception and to the provisions of relevant collective licensing agreements, no reproduction of any part may take place without the written permission of Cambridge University Press. First published 2002 Printed in the United Kingdom at the University Press, Cambridge Typeface Monotype Garamond 11/13pt System QuarkXPress™ [] A catalogue record for this book is available from the British Library Library of Congress Cataloguing in Publication data Thacker, J. R. M. (Jonathan Richard MacDougall), 1962– An introduction to arthropod pest control / J.R.M. Thacker. p. cm. Includes bibliographical references and index. ISBN 0 521 56106 X (hbk.) – ISBN 0 521 56787 4 (pbk.) 1. Arthropod pests – Control. I. Title. SB931.T52 2002 632Ј.7–dc21 2002023436 ISBN 0 521 56106 X hardback ISBN 0 521 56787 4 paperback Contents Preface page xiii Acknowledgements xv 1 A brief history of arthropod pest control 1 Introduction 1 Neolithic agricultural development and pest control 4 Early attempts at pest control 5 Pest control after the sixteenth century 7 The development of modern chemical pesticides 11 The development of integrated pest management 16 Alternatives to chemical pest control 22 High-tech pest control for the twenty-first century 24 Summary 25 Further reading 26 2 Botanical insecticides 28 Introduction 28 Sabadilla 30 Nicotine 32 Quassia 34 Unsaturated isobutylamides 35 Ryanodine 37 Naphthoquinones 38 Rotenone 39 Sweet flag 40 Marigolds 41 Pyrethrum 41 Azadirachtin 43 Essential oils 44 Botanical insecticides developed for use in the west 44 Botanical insecticides in developing countries 47 viii Contents Summary 48 Further reading 49 3 Modern synthetic insecticides 50 Introduction 50 Early synthetic insecticides 50 Modern synthetic insecticides 52 The synthetic insecticide market 52 Global crop losses and insecticide use 55 Organochlorine insecticides 57 Organophophate insecticides 62 Pyrethroid insecticides 67 Carbamate insecticides 71 Neonicotinoids 74 Other insecticidal chemical groups 76 Summary 76 Further reading 79 4Formulation, application and the direct and indirect side-effects of insecticides 81 Introduction 81 Formulation of insecticides 83 Categories of formulation 83 Application of insecticides 88 Direct and indirect side-effects of insecticides 94 Resistance to insecticides 94 Resurgent pests, secondary pests and nontarget mortality 100 Ecosystem side-effects associated with insecticide use 102 Human problems associated with exposure to insecticides 104 Legislation and pesticides 107 Summary 108 Selected further reading 110 5 Biological control agents 112 Introduction 112 Predatory invertebrates 114 Parasitoids 116 Predators 120 Methods for using biocontrol agents 121 Classical biological control 122 Augmentation/inoculation techniques 128 Conserving natural enemies and habitat manipulation 129 Vertebrate biocontrol agents 130 Worldwide results from biocontrol programmes 131 Summary 132 Further reading 133 6 Microbial pest control 135 Introduction 135 Bacteria 137 Contents ix Fungi 143 Nematodes 147 Protozoa 150 Viruses 152 Summary 157 Further reading 157 7 Pheromones and pest control 159 Introduction 159 Types of pheromone 160 Chemistry of pheromones 164 Formulation of pheromones 165 Pheromone traps 167 Using pheromones 169 Monitoring 169 Mass trapping 171 Mating disruption 173 Lure and kill 174 Other applications of pheromones 175 Development of pheromones for pest control 176 Allelochemicals 177 Summary 178 Further reading 178 8 Insect growth regulators 180 Introduction 180 Arthropod growth and development 180 Juvenile hormone mimics 181 Moulting hormone mimics 185 Chitin synthesis inhibitors 186 Summary 188 Further reading 188 9 Genetic manipulation of pest species 190 Introduction 190 Methods of genetic manipulation 191 Sterile insect release technique 193 Mass rearing 194 Sterilisation 195 Insect release 196 Mating 196 Control of the American screwworm fly 197 Control of the screwworm in Libya 200 Control of the Mediterranean fruit fly 201 Control of the tsetse fly 202 Other species controlled using SIRT 203 Insect orders not controlled by SIRT 203 Summary 203 Further reading 205 x Contents 10 Host-plant resistance 207 Introduction 207 Crop plant development 209 Secondary plant substances 210 Types of resistance 212 Germplasm collections 216 Techniques for producing resistant plants 218 Commercialised plants with resistance to pests 221 Potential problems with host-plant resistance 224 Summary 227 Further reading 227 11 Cultural techniques and organic farming 229 Introduction 229 Soil cultivation 231 Clean cultivation 232 Manuring/fertilising 232 Irrigation 233 Crop rotations 233 Trap crops 234 Planting date manipulation 235 Harvest date manipulation 235 Planting density 235 Abiotic environmental manipulation 236 Intercropping 236 Boosting natural enemy numbers 239 Organic farming 239 Sustainable farming 242 Summary 243 Further reading 244 12 Integrated pest management (IPM) 246 Introduction 246 Defining IPM 247 Steps involved in establishing an IPM programme 257 Cotton IPM 259 Cotton IPM programmes worldwide 261 Apple IPM 262 Rice IPM 264 Vegetable IPM 265 Greenhouse IPM 266 The impacts of biotechnology on IPM programmes 268 Future developments with IPM 269 Summary 272 Further reading 273 13 Biotechnology and pest control 275 Introduction 275 Species for manipulation 276 Biotechnological methods 277 Contents xi Transgenic plants 281 Transgenic predators 286 Transgenic pests 287 Advantages associated with biotechnology for pest control 288 Disadvantages associated with biotechnology for pest control 290 Predictions for future developments 295 Summary 300 Further reading 301 Glossary and commentary on text 302 Arthropod pest index 333 Beneficial species index 335 Subject index 337 Plate sections between pages 144 and 145, and pages 240 to 241. 1 A brief history of arthropod pest control Introduction Two of the most important challenges facing humanity in the twenty-first century comprise food production and disease control. These are challenges that are associated with the increasing global human population (Figure 1.1) and with the control of arthropod pests, the subject of this book. The impor- tance of these challenges cannot be overemphasised. Arthropod pests are responsible for global pre- and postharvest crop losses of approximately 20–50% of potential production and for transmitting a number of the world’s most important diseases (Table 1.1). For example, it has been estimated that the protozoan organism causing malaria infects approximately 500 million people worldwide – almost 10% of the people on earth. It is undoubtedly the case that humanity’s problems with arthropod pests are not new and they certainly predate the development of agriculture approx- imately 10000–16000 years ago. Arthropods first appear in the fossil record over 500 million years ago during the Cambrian 1 period at the start of the Palaeozoic. The oldest insect fossils to have been found so far are dated to the Devonian, a period that began 400 million years ago. Modern insects begin to appear in the fossil record about 280 million years ago. However, the time scale over which different insect orders are detected in the fossil record 1 Geological time is split into four major phases that are known as the Precambrian, Palaeozoic, Mesozoic and the Cenozoic. Each of these major phases is then further subdivided, e.g. the Cambrian is the first phase or subdivision of the Palaeozoic. See glossary for further details. 7 6 5 4 3 2 World Population, Billions 1 0 -12000 -10000 -8000 -6000 -4000 -2000 0 2000 4000 Time (years BC) Fig. 1.1. Growth in the global human population since 10000 BC. Introduction 3 Table 1.1. The world’s most important arthropod-borne diseases Disease Vector Agent Cases Deaths Malaria Mosquito Protozoa 300–500 m 1.5–2.7 m (Plasmodium) Filariasis Mosquito Filarial 120 m NAa worms Dengue Mosquito Virus 50 m 0.5 m Onchocerciasis Blackfly Filarial 18 m NAb worms Chagas Triatomine Protozoa 16–18 m 20000– bugs (Trypanosoma) 50000 Leishmaniasis Sand flies Protozoa 12 m NAa (Leishmania) Trypanosomiasis Tsetse flies Protozoa 300000– NAa (Trypanosoma) 500000 Yellow fever Mosquito Virus 200000 30000 Notes: a Accurate estimates not available. b Onchocerciasis or ‘river blindness’ causes chronic rather than acute illness. The WHO estimates that, of the 18 m people infected, 6.5 m suffer from severe dermatitis and 270000 are blind. Source: Data collected from the World Health Organization (WHO) World Health Report 1998. stretches up until 75 million years ago, during the Cretaceous period at the end of the Mesozoic. For example, silverfish (order Thysanura) are detected in the fossil record from the Devonian while caterpillars (order Lepidoptera) are not detected until the Cretaceous. By contrast, Homo sapiens has been around for about a 100000 years. It should therefore come as no great surprise to know that so far we have not been able to conquer the problems that arthropod pests pose. These animals have had far more time