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Control of Pests and Weeds By Natural Enemies CONTROL OF PESTS AND WEEDS BY NATURAL ENEMIES AN INTRODUCTION TO BIOLOGICAL CONTROL Roy Van Driesche, Mark Hoddle, and Ted Center 9781405145718 1 pre qxd. Beneficial Species: Several beneficial insect species play an important role for garden health. The most important group of beneficial insects are pollinators, biological control agents, and soil decomposers. Pollinators are insects which pollinate plants. Insect pollinators include honey bees, beetles, flies, ants, moths, butterflies, bumble bees, solitary bees, and wasps. Butterflies and moths are important pollinators of flowering plants in wild ecosystems and managed systems such as gardens and parks. Biological control of pests is part of an integrated pest management (IPM) strategy. It is the reduction of pest populations by natural enemies and typically involves an active human role. In fact, all insect species are also suppressed by naturally occurring organisms and environmental factors, with no human input. The natural enemies of insect pests, also known as biological control agents, include predators, parasitoids, and pathogens
9781405145718_1_pre.qxd 1/25/08 10:21 AM Page iii CONTROL OF PESTS AND WEEDS BY NATURAL ENEMIES AN INTRODUCTION TO BIOLOGICAL CONTROL Roy Van Driesche, Mark Hoddle, and Ted Center 9781405145718_1_pre.qxd 1/25/08 10:21 AM Page ii 9781405145718_1_pre.qxd 1/25/08 10:21 AM Page i CONTROL OF PESTS AND WEEDS BY NATURAL ENEMIES 9781405145718_1_pre.qxd 1/25/08 10:21 AM Page ii 9781405145718_1_pre.qxd 1/25/08 10:21 AM Page iii CONTROL OF PESTS AND WEEDS BY NATURAL ENEMIES AN INTRODUCTION TO BIOLOGICAL CONTROL Roy Van Driesche, Mark Hoddle, and Ted Center 9781405145718_1_pre.qxd 1/25/08 10:21 AM Page iv © 2008 by Roy Van Driesche, Mark Hoddle, and Ted Center BLACKWELL PUBLISHING 350 Main Street, Malden, MA 02148-5020, USA 9600 Garsington Road, Oxford OX4 2DQ, UK 550 Swanston Street, Carlton, Victoria 3053, Australia The right of Roy Van Driesche, Mark Hoddle, and Ted Center to be identified as the authors of this work has been asserted in accordance with the UK Copyright, Designs, and Patents Act 1988 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, electronic, mechanical, photocopying, recording or otherwise, except as permitted by the UK Copyright, Designs, and Patents Act 1988, without the prior permission of the publisher Designations used by companies to distinguish their products are often claimed as trademarks All brand names and product names used in this book are trade names, service marks, trademarks, or registered trademarks of their respective owners The publisher is not associated with any product or vendor mentioned in this book This publication is designed to provide accurate and authoritative information in regard to the subject matter covered It is sold on the understanding that the publisher is not engaged in rendering professional services If professional advice or other expert assistance is required, the services of a competent professional should be sought First published 2008 by Blackwell Publishing Ltd 2008 Library of Congress Cataloging-in-Publication Data Van Driesche, Roy Control of pests and weeds by natural enemies : an introduction to biological control / Roy Van Driesche, Mark Hoddle, and Ted Center – 1st ed p cm Includes bibliographical references and index ISBN 978-1-4051-4571-8 (pbk : alk paper) Pests–Biological control Weeds–Biological control I Hoddle, Mark II Center, Ted D III Title SB975.V38 2008 632'.96–dc22 2007038529 ISBN: 978-1-4051-4571-8 (paperback) A catalogue record for this title is available from the British Library Set in 9/11pt Photina MT by Graphicraft Limited, Hong Kong Printed and bound in Singapore by C.O.S Printers Pte Ltd The publisher’s policy is to use permanent paper from mills that operate a sustainable forestry policy, and which has been manufactured from pulp processed using acid-free and elementary chlorine-free practices Furthermore, the publisher ensures that the text paper and cover board used have met acceptable environmental accreditation standards For further information on Blackwell Publishing, visit our website at www.blackwellpublishing.com 9781405145718_1_pre.qxd 1/25/08 10:21 AM Page v CONTENTS Preface ix PART SCOPE OF BIOLOGICAL CONTROL 1 INTRODUCTION TYPES OF BIOLOGICAL CONTROL, TARGETS, AND AGENTS What is biological control? Permanent control over large areas Temporary pest suppression in production areas Kinds of targets and kinds of agents PART KINDS OF NATURAL ENEMIES PARASITOID DIVERSITY AND ECOLOGY 11 What is a parasitoid? 11 Terms and processes 11 Some references to parasitoid families 13 Groups of parasitoids 13 Finding hosts 15 Host recognition and assessment 19 Defeating host defenses 22 Regulating host physiology 24 Patch-time allocation 25 PREDATOR DIVERSITY AND ECOLOGY 29 Non-insect predators 29 Major groups of predatory insects 31 Overview of predator biology 33 Predator foraging behavior 34 Predators and pest control 37 Effects of alternative foods on predator impact 40 Interference of generalist predators with classical biological control agents 41 Predator and prey defense strategies 43 WEED BIOCONTROL AGENT DIVERSITY AND ECOLOGY 45 The goal of weed biological control 45 Terms and processes 45 Herbivory and host finding 46 Herbivore guilds 47 Groups of herbivores and plant pathogens 47 ARTHROPOD PATHOGEN DIVERSITY AND ECOLOGY 56 Bacterial pathogens of arthropods 56 Viral pathogens of arthropods 58 Fungal pathogens of arthropods 59 Nematodes attacking arthropods 61 Generalized arthropod pathogen life cycle 62 Epidemiology: what leads to disease outbreaks? 64 PART INVASIONS: WHY BIOLOGICAL CONTROL IS NEEDED 67 THE INVASION CRISIS 69 Urgency of the invasion crisis 69 Case histories of four high-impact invaders 70 The extent of harmful impact by invaders 73 How invasive species get to new places? 75 Why some invasions succeed but others fail? 77 Invader ecology and impact 78 WAYS TO SUPPRESS INVASIVE SPECIES 80 Prevention: heading off new invasions through sound policy 80 Eradication based on early detection 83 9781405145718_1_pre.qxd 1/25/08 10:21 AM Page vi vi Contents Invaders that no harm 84 Control of invasive pests in natural areas 84 Factors affecting control in natural areas 86 Control of invasive species in crops 87 PART NATURAL ENEMY INTRODUCTIONS: THEORY AND PRACTICE 89 INTERACTION WEBS AS THE CONCEPTUAL FRAMEWORK FOR CLASSICAL BIOLOGICAL CONTROL 91 Terminology 91 Forces setting plant population density 93 Forces setting insect population density 94 Predictions about pests based on food webs 95 10 THE ROLE OF POPULATION ECOLOGY AND POPULATION MODELS IN BIOLOGICAL CONTROL, BY JOSEPH ELKINTON 97 Basic concepts 97 Population models 104 11 CLASSICAL BIOLOGICAL CONTROL 115 Introduction 115 Classical biological control 115 New-association biological control 133 Summary 136 12 WEED BIOLOGICAL CONTROL 137 Differences and similarities between weed and arthropod programs 137 Why plants become invasive 138 Selecting suitable targets for weed biological control 139 Conflicts of interest in weed biological control 139 Faunal inventories: finding potential weed biological control agents 139 Safety: “will those bugs eat my roses?” 141 Pre-release determination of efficacy 142 How many agents are necessary for weed control? 143 Release, establishment, and dispersal 144 Evaluation of impacts 145 Non-target impacts 146 When is a project successful? 146 Conclusions 147 PART TOOLS FOR CLASSICAL BIOLOGICAL CONTROL 149 13 FOREIGN EXPLORATION 151 Planning and conducting foreign exploration 151 Shipping natural enemies 154 Operating a quarantine laboratory 156 Managing insect colonies in quarantine 157 Developing petitions for release into the environment 158 14 CLIMATE MATCHING 160 Climate matching 160 Inductive modeling: predicting spread and incursion success 162 Deductive modeling: predicting spread and incursion success 164 Conclusions 165 15 MOLECULAR TOOLS, BY RICHARD STOUTHAMER 167 Types of molecular data 168 Important biological control issues that molecular techniques can address 177 Conclusions 179 PART SAFETY 181 16 NON-TARGET IMPACTS OF BIOLOGICAL CONTROL AGENTS 183 Biological control as an evolving technology 183 The amateur to early scientific period (1800–1920) 184 A developing science makes some mistakes (1920–70) 188 Broadening perspectives (1970–90) 192 Current practice and concerns 195 “Re-greening” biological control 198 17 PREDICTING NATURAL ENEMY HOST RANGES 199 Literature records 199 Surveys in the native range 201 Laboratory testing to estimate host ranges 201 Interpretation of tests 207 Examples of host-range estimation 209 Risk assessment 213 9781405145718_1_pre.qxd 1/25/08 10:21 AM Page vii Contents 18 AVOIDING INDIRECT NON-TARGET IMPACTS 215 Kinds of potential indirect effects 215 Can risk of indirect impacts be reduced by predicting natural enemy efficacy? 216 PART MEASURING NATURAL ENEMY IMPACTS ON PESTS 221 19 FIELD COLONIZATION OF NATURAL ENEMIES 223 Limitations from the agent or recipient community 223 Managing release sites 225 Quality of the release 225 Caging or other release methods 228 Persistence and confirmation 229 20 NATURAL ENEMY EVALUATION 230 Natural enemy surveys in crops 230 Pre-release surveys in the native range for classical biological control 231 Post-release surveys to detect establishment and spread of new agents 232 Post-release monitoring for non-target impacts 233 Measurement of impacts on the pest 233 Separating effects of a complex of natural enemies 248 Economic assessment of biological control 251 PART CONSERVING BIOLOGICAL CONTROL AGENTS IN CROPS 253 21 PROTECTING NATURAL ENEMIES FROM PESTICIDES 255 Problems with pesticides 255 Super pests and missing natural enemies 256 Dead wildlife and pesticide residues in food 258 Cases when pesticides are the best tool 259 How pesticides affect natural enemies 259 Seeking solutions: physiological selectivity 261 Pesticide-resistant natural enemies 262 Ecological selectivity: using non-selective pesticides with skill 263 Transgenic Bt crops: the ultimate ecologically selective pesticide 264 22 ENHANCING CROPS AS NATURAL ENEMY ENVIRONMENTS 266 Problem 1: unfavorable crop varieties 266 vii Solution 1: breeding natural enemy-friendly crops 268 Problem 2: crop fields physically damaging to natural enemies 269 Solution 2: cover crops, mulching, no-till farming, strip harvesting 269 Problem 3: inadequate nutritional sources 270 Solution 3: adding nutrition to crop environments 271 Problem 4: inadequate reproduction opportunities 272 Solution 4: creating opportunities for contact with alternative hosts or prey 273 Problem 5: inadequate sources of natural enemy colonists 273 Solution 5: crop-field connectivity, vegetation diversity, and refuges 274 Other practices that can affect natural enemies 276 Conclusions 278 PART BIOPESTICIDES 279 23 MICROBIAL PESTICIDES: ISSUES AND CONCEPTS 281 History of microbial insecticides 281 What makes a pathogen a likely biopesticide? 282 Overview of options for rearing pathogens 283 Agent quality: finding it, keeping it, improving it 284 Measuring the efficacy of microbial pesticides 285 Degree of market penetration and future outlook 286 24 USE OF ARTHROPOD PATHOGENS AS PESTICIDES 289 Bacteria as insecticides 289 Fungi as biopesticides 291 Viruses as insecticides 295 Nematodes for insect control 298 Safety of biopesticides 301 PART 10 AUGMENTATIVE BIOLOGICAL CONTROL 305 25 BIOLOGICAL CONTROL IN GREENHOUSES 307 Historical beginnings 307 When are greenhouses favorable for biological control? 308 9781405145718_7_plates.qxd 1/25/08 10:18 AM Page (c) (d) Plate 3.1 (Cont’d) (c) The aphelinid Aphytis melinus DeBach attacking California red scale, Aonidiella aurantii (Maskell) (d) The trichogrammatid Trichogramma pretiosum Riley ovipositing in the egg of Helicoverpa zea (Boddie) 9781405145718_7_plates.qxd 1/25/08 10:18 AM Page (e) Plate 3.1 (Cont’d) (e) The mymarid Gonatocerus ashmeadi Girault (f ) The braconid Aphidius colemani Viereck Photographs courtesy of (a, c–f ) Jack Kelly Clark, University of California IPM Photo Library and (b) William Roltsch, California Department of Agriculture (f) 9781405145718_7_plates.qxd 1/25/08 10:18 AM Page (a) (b) Plate 4.1 (a) Orius tristicolor (White) (Anthocoridae), a species used for augmentative control of thrips in greenhouse crops (b) A chrysopid larva, a group predaceous on aphids and other pests 9781405145718_7_plates.qxd 1/25/08 10:18 AM Page (c) (d) Plate 4.1 (Cont’d) (c) An adult vedalia beetle [Rodolia cardinalis (Mulsant)] next to its prey, the cottony cushion scale (Icerya purchasi Maskell) (d) The histerid beetle Teretrius nigrescens (Lewis), a predator of the larger grain borer, Prostephanus truncatus (Horn), a pest of stored corn on subsistence farms in Africa 9781405145718_7_plates.qxd 1/25/08 10:18 AM Page (e) (f) Plate 4.1 (Cont’d) (e) An adult syrphid fly (f) Syrphid larvae are predators of aphids Photographs courtesy of (a–c, e, f) Jack Kelly Clark, University of California IPM Photo Library and (d) Georg Goergen, IITA 9781405145718_7_plates.qxd 1/25/08 10:18 AM Page (a) (c) (b) Plate 5.1 (a) Galls of Trichilogaster acaciaelongifoliae (Froggatt) (Hymen.: Pteromalidae) on Acacia longifolia (Andrews); (b) adult gall wasp; (c) close-up of a gall; and (d) bisected gall showing larvae (d) 9781405145718_7_plates.qxd 1/25/08 10:18 AM Page (e) (f) Plate 5.1 (Cont’d) (e) The eriophyiid rust mite Floracarus perrepae Knihinicki and Boczek, an herbivore associated with Old World climbing fern, Lygodium microphyllum (Cav.) R Br.; and (f ) damage from F perrepae Photographs courtesy of (a–d) S Neser, PPRI and (e, f) John Goolsby, USDA-ARS 9781405145718_7_plates.qxd 1/25/08 10:18 AM Page (a) (b) Plate 7.1 (a) A stand of the toxic alga Caulerpa taxifolia (Vahl) C Agardh covering the bottom of the Mediterranean Sea (b) Brown tree snake (Boiga irregularis Fitzinger), an invasive predator that has decimated the forest birds of Guam 9781405145718_7_plates.qxd 1/25/08 10:18 AM Page 10 (c) (d) Plate 7.1 (Cont’d) (c) The Guam Micronesian kingfisher (Todirhamphus cinnamominus cinnamominus) is one of the native birds of Guam decimated by the brown tree snake (Boiga irregularis Fitzinger) (d) Close-up of an adult hemlock woolly adelgid (Adelges tsugae Annand) and eggs 9781405145718_7_plates.qxd 1/25/08 10:18 AM Page 11 (e) (f) Plate 7.1 (Cont’d) (e) Scymnus camptodromus Yu et Liu, a coccinellid predator from China of the hemlock woolly adelgid (A tsugae Annand) (f) Laricobius nigrinus Fender, a derodontid predator from British Columbia (Canada) of the hemlock woolly adelgid (A tsugae Annand) Photographs courtesy of (a) Alexandre Meinesz, University of Nice, (b) Christy Martin, CGAPS, Hawaii, USA, (c) W.D Kesler, (d) Mike Montgomery, www.Forestryimages.org, (e) Dr Guoyue Yu, and (f ) Rob Flowers 9781405145718_7_plates.qxd 1/25/08 10:18 AM Page 12 (a) (b) Plate 12.1 (a, b) Rubber vine (Cryptostegia grandifolia R Br.) is a severe pest of natural areas in tropical Queensland, Australia, smothering native vegetation (a); the rust Maravalia cryptostegiae (b), a pathogen imported from Madagascar, damages rubber vine heavily and is providing control (Vogler & Linday 2002) 9781405145718_7_plates.qxd 1/25/08 10:18 AM Page 13 (c) (d) Plate 12.1 (Cont’d) (c, d) Biological control of waterhyacinth [Eichhornia crassipes (Mart.) Solms-Laub] on Lake Victoria in Africa (Kisuma, Kenya) by introduced weevils provided dramatic control, changing solid mats in May 1999 (c) to open water by December of the same year (d) 9781405145718_7_plates.qxd 1/25/08 10:18 AM Page 14 (e) Plate 12.1 (Cont’d) (e, f) The chrysomelid beetle Diorhabda elongata Brulle deserticola Chen, has been introduced in the southwestern USA for the biological control of Tamarix spp., which are Eurasian shrubs widely invasive in riparian areas Here, (e) larvae are shown defoliating Tamarix shrubs 9781405145718_7_plates.qxd 1/25/08 10:18 AM Page 15 (f) Plate 12.1 (Cont’d) (e, f) The chrysomelid beetle Diarhabda elongata Brulle deserticola Chen, has been introduced in the southwestern USA for the biological control of Tamarix spp., which are Eurasian shrubs widely invasive in riparian areas (f) The extent of defoliation is visible in an elevated view of a release site Photographs courtesy of (a, b) Colin Wilson, (c, d) Mic Julien, CSIRO, and (e, f) Ray Caruthers, USDA-ARS 9781405145718_7_plates.qxd 1/25/08 10:18 AM Page 16 (a) (b) 0.00–0.22 0.22–0.60 0.60–1.25 1.25–2.36 2.36–4.25 4.25–7.48 7.48–12.99 12.99–22.40 22.40–38.46 38.46–47.87 1–2 3–5 6–8 9–11 12–17 18–21 22–24 25–27 N N 15 30 60 90 Miles 120 15 30 60 90 Miles 120 PREPARED BY KAC GIS FACILITY Plate 14.1 GIS mapping of the estimated life-table statistic, net reproductive rate (Ro) for the Homalodisca coagulata egg parasitoids (a) Gonatocerus ashmeadi Girault and (b) Gonatocerus triguttatus Girault in California Ro was determined in the laboratory for each parasitoid species across five experimental temperatures and the resulting relationship between Ro and temperature was determined and modeled in GIS using data from 260 weather stations in California The results are very striking: G ashmeadi can be expected to infiltrate most of California and reproduce annually, whereas G triguttatus may be severely restricted to localized regions in southern California Maps drawn by M Hoddle ... parasitoid families see Clausen (1 962; useful but dated), Askew (1 97 1), Waage and Greathead (1 98 6), Gauld and Bolton (1 98 8), Grissell and Schauff (1 99 0), Godfray (1 99 4), Hanson and Gauld (1 99 5), ... Clausen (1 96 2), Arnett (1 96 8), Hodek (1 97 3), Foelix (1 98 2), Gerson and Smiley (1 99 0), Hagen et al (1 99 9), and Triplehorn and Johnson (2 00 5) Predatory thrips (Thysanoptera) Most thrips are phytophagous,... Gerson and Smiley (1 99 0), New (1 99 2), Sabelis (1 99 2), Dixon (2 00 0), and Triplehorn and Johnson (2 00 5) Spiders Spiders (Aranae) (Figure 4. 1) are all predaceous (Foelix 198 2) Spiders often show habitat