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AREA-WIDE CONTROL OF INSECT PESTS

AREA-WIDE CONTROL OF INSECT PESTS · A C.I.P. Catalogue record for this book is available from the Library of Congress. P.O. Box 17, 3300 AA Dordrecht, The Netherlands

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Page 1: AREA-WIDE CONTROL OF INSECT PESTS · A C.I.P. Catalogue record for this book is available from the Library of Congress. P.O. Box 17, 3300 AA Dordrecht, The Netherlands

AREA-WIDE CONTROL OF INSECT PESTS

Page 2: AREA-WIDE CONTROL OF INSECT PESTS · A C.I.P. Catalogue record for this book is available from the Library of Congress. P.O. Box 17, 3300 AA Dordrecht, The Netherlands

Edited by

Joint FAO/IAEA Programme of Nuclear Techniques in Food and Agriculture,

M.J.B. VreysenA.S. RobinsonJ. Hendrichs

From Research to Field ImplementationArea-Wide Control of Insect Pests

Vienna, Austria

Page 3: AREA-WIDE CONTROL OF INSECT PESTS · A C.I.P. Catalogue record for this book is available from the Library of Congress. P.O. Box 17, 3300 AA Dordrecht, The Netherlands

A C.I.P. Catalogue record for this book is available from the Library of Congress.

P.O. Box 17, 3300 AA Dordrecht, The Netherlands.

www.springer.com

Printed on acid-free paper

ISBN 978-1-4020-6058-8 (HB)

ISBN 978-1-4020-6059-5 (e-book)

moth), Scott Bauer (USDA Agricultural Research Service, -

vil), Hendrik Hofmeyr (false codling moth), and the Locust Group (FAO) (desert locust).

Photo’s on the back cover were provided by Patrick Robert (tsetse fly), Ignacio Baez (cactuswww.forestryimages.org

Mediterranean fruit fly, melon fly), Jack Dykinga (USDA Agricultural Research Service,www.forestryimages.org - Mexican fruit fly), Susan Ellis (www.forestryimages.org - Asiantiger mosquito), Alton N. Sparks (University of Georgia, www.forestryimages.org - boll wee-

All Rights Reserved© 2007 IAEAAll IAEA scientific and technical publications are protected by the terms of the UniversalCopyright Convention on Intellectual Property as adopted in 1952 (Berne) and as revised in1972 (Paris). The copyright has since been extended by the World Intellectual PropertyOrganization (Geneva) to include electronic and virtual intellectual property. Permission to usewhole or parts of texts contained in the IAEA publications in printed or electronic form mustbe obtained and is usually subject to royalty agreements. Proposals for non-commercial repro-ductions and translations are welcomed and considered on a case-by-case basis. Inquiriesshould be addressed to the Publishing Section, IAEA, Wagramer Strasse 5, A-1400 Vienna,

Photo’s on the front cover were provided by Hendrik Hofmeyr, Frantisek Marec, Ilan Mizrahi,Marc Vreysen, Petr Pavlicek and SECNA (FAO).

Austria.

Published by Springer,

Photo credits:

Page 4: AREA-WIDE CONTROL OF INSECT PESTS · A C.I.P. Catalogue record for this book is available from the Library of Congress. P.O. Box 17, 3300 AA Dordrecht, The Netherlands

The world population is still growing at analarming rate, requiring ever increasing pro-ductivity and less waste in agriculture to copewith the increasing demands to satisfy foodsecurity for all humans. Alleviation of povertyis in many countries hampered by a myriad ofinsect pests that cause enormous economiclosses to agricultural commodities, both at thepre- and postharvest stages. Initially, most ofthese insect pests were controlled to a varyingdegree by the use of broad-spectrum insecti-cides. However, the indiscriminate use of thesechemicals as a control tactic is no longer sus-tainable in view of increased development ofresistance, pollution of soils and surface water,residues in food and the environment, repre-senting risks to human health and biodiversity,etc. As a consequence, demands have beenvoiced at least since “Silent Spring” in 1962for control tactics and approaches that are notonly efficient, but also sustainable and friend-lier to the environment.

Integrated pest management (IPM) hasbeen accepted since the 1960’s and 70s as aviable pest management strategy that aims atintegrating control tactics to maintain damagelevels below a certain economic thresholdlevel whilst also protecting the environment bythriving to limit the use of pesticides. ClassicalIPM is however a localized approach, with theobjective of protecting crops or livestock thatis largely under the control of each farmer,with little collaboration or any coordinatingstructure. Control is exercised only in the areasof economic interest, often resulting in themain or residual pest population pocketsremaining in the surrounding areas that haveno economic value. These constitute perma-nent sources from where the commercial areasunder control are re-invaded.

A quite different, more efficient and sus-tainable approach is the integration of controltactics against an entire pest population, i.e.area-wide integrated pest management (AW-

IPM) or total population management. TheAW-IPM is a coordinated, sustainable and pre-ventive approach that targets pest populationsin all areas, including non-commercial urban

The coordination required among farmersand all other stakeholders for an area-wideapproach, makes AW-IPM programmes com-plex, management intensive, requiring long-term commitment and funding. Although theyresult in more sustainable control of insectpests, there is by no means a guarantee for suc-cess. This new textbook on area-wide controlof insect pests collates a series of selectedpapers that attempts to address various funda-mental components of AW-IPM, e.g. theimportance of relevant problem-solvingresearch, the need for essential baseline data,the significance of adequate tools for appropri-ate control strategies, and the value of pilot tri-als, etc. Of special interest are the numerouspapers on pilot and operational programmesthat pay special attention to practical problemsencountered during programme implementa-tion.

The book is a compilation of 66 papers thatare authored by experts from more than 30countries. Each paper was peer-reviewed by atleast one, in most case two or more independ-ent, outside experts and edited for the Englishlanguage by Dr James Dargie, former Directorof the Joint FAO/IAEAProgramme of NuclearTechniques in Food and Agriculture. We boththank the many reviewers and Jim whosemeticulous work and suggestions improvedmany of the papers. In addition, the editorssubjected each paper to an in-depth technicalquality control process. As a result, we trustthat the technical quality of the papers is opti-mal, the information provided accurate, up-to-date and of a high international standard. Thisprocess of peer-review, editing and formattinghas taken considerable time and we appreciatethe patience of the authors.

Preface

The EditorsJune 2007

v

settings, non-cultivated and wild host areas.

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DISCLAIMER

This publication has been prepared from the original material as submitted by the authors. Theviews expressed do not necessarily reflect those of the IAEA, the governments of the nomi-nating Member States or the nominating organizations. The use of particular designations ofcountries or territories does not imply any judgement by the publisher, the IAEA, as to thelegal status of such countries or territories, of their authorities and institutions or of the delim-itation of their boundaries.The mention of names of specific companies or products (whether or not indicated as regis-tered) does not imply any intention to infringe proprietary rights, nor should it be construed asan endorsement or recommendation on the part of the IAEA.The authors are responsible for having obtained the necessary permission for the IAEA to

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reproduce, translate or use material from sources already protected by copyrights.

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Since area-wide integrated pest management(AW-IPM) programmes almost always aresocial enterprises each with a diverse set ofstakeholders reliant on advanced technology,they tend to be complex to implement, espe-cially in terms of management. Therefore, theappearance of this book, “Area-wide controlof insect pests: from research to field imple-mentation” is most timely, and it will beinvaluable for informing concerned scientists,leaders of private firms, commodity organiza-tions and public agencies, bankers, legislators,students and those interested in placing man-agement of major insect pest problems on asustainable and environmentally acceptablefooting.Although the need to develop and imple-

ment more effective strategies of combatingpests and pathogens has always been dire, theurgency of this challenge has increasedsharply for two reasons. The first reason is therapid increase in the world population, whichmore than doubled from roughly 2.5 billion to6 billion people in the second half of the 20thcentury; and the second reason is that therapid globalization of travel and trade in agri-cultural and other products has dramaticallyincreased the spread of pests, pathogens andother invasive harmful organisms.Many of the economically most damaging

pests are invasive alien species that haveescaped the constraints, which keep their pop-ulations in check in their regions of origin. InNorth America roughly one-half of the majorpests originated abroad, and this seems also tobe true in other continents. Major exotic pestsand pathogens – many adapted for wide dis-persal and high rates of reproduction – arebecoming established with increasing fre-quencies on all continents and on many eco-logically sensitive islands. Therefore, to facil-itate the expansion of international agricultur-al trade while minimizing the further spreadof some major pests, commodities of whichthey are hosts are increasingly produced forexport in pest free areas or in areas of low pestprevalence that obtained their favourable phy-

tosanitary status through AW-IPM approach-es.Currently, for the most part, the control of

many highly mobile and very destructiveinsect pests is still carried out by individualproducers who rely heavily on the use ofbroad-spectrum insecticides. Although othercontrol technologies are often incorporatedinto the producer’s IPM system, these tech-nologies, too, are usually applied by produc-ers independently of other producers, andwithout due consideration of surrounding hostand non-host areas. Such an uncoordinatedfarm-by-farm IPM pattern provides opportu-nities for the pest population to build up and toestablish damaging infestations.Consequently, on most farms insect pest

populations increase to damaging levels eachyear, and the farmer is forced to apply fast-acting insecticides as a rescue treatment. Thisdefeats the primary goal of the IPM system,which is to take maximum advantage of natu-rally occurring biological control agents.Similarly in combating pests and pathogens ofconcern to human and animal well-being, lessthan thorough treatment of the entire popula-tion fails to provide durable relief. Thus thekey concept of the AW-IPM strategy is toaddress the whole pest population includingall places of refuge or foci of infestation fromwhich recruits could come to re-establishdamaging densities of the pest population inareas of concern.The area-wide approach is not new, but

originated several thousand years ago. In theRoman Empire it was recognized that someservices carried out area-wide were more effi-cient and cheaper than when left to the actionof individual citizens. As such, garbage wasdiligently removed from some cities, cleanwater was brought from distant sources andpublic baths were provided. The suddenappearance in 1347 of Black Death, a bacteri-al disease transmitted by the flea, Xenopsyllacheopis (Rothschild), led to the invention ofquarantine to contain the epidemic and tostamp it out. Beginning in the late 1920s,

Introductory Remarks

vii

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when catastrophic locust plagues were wide-spread in Africa and southwest Asia, conti-nent-wide campaigns have been organized toprotect against highly devastating locustspecies; and these campaigns, now led byFAO’s Locust Group, employ sophisticatedtechnologies. During the past one-half centurythe area-wide application of the sterile insecttechnique in combination with other technolo-gies against an array of major insect pests hasserved to focus the attention of scientists andadministrators on ways of applying the area-wide approach in the combat against manyother pests and diseases.The principles ofAW-IPM are addressed in

this book’s introductory chapter. The chapterargues that each fundamental component ofclassical IPM, be it a cultural, biological orchemical control tactic, applied against anentire insect population (total population man-agement) will lead in most cases to more sus-tainable pest control as compared to a local-ized farm-by-farm approach. Some funda-mental management and strategic challengesof AW-IPM programmes are likewiseaddressed, including the make or break envi-ronmental and economic issues.Successful AW-IPM programmes require

basic research and preparatory activitiesincluding methods development, feasibilitystudies, pilot trials and a regulatory frame-work. These aspects are dealt with in subse-quent sections. Section 2 covers and illumi-nates several important basic research areasincluding genetics, transgenesis, genetic sex-ing, cryobiology, physiology, insect sym-bionts and mating behaviour strategies.Ecological heterogeneity at within field,

within farm, and broader spatial scales pro-foundly affects the population dynamics ofpests and their natural enemies and otheraspects of their ecology. Methods of systemsanalysis, mathematical modelling and a num-ber of geo-spatial technologies (geographicinformation systems and global positioningsystem) have been adapted to cope with thespatio-temporal complexity in AW-IPM pro-grammes and have contributed greatly toincreased effectiveness and efficiency of pro-

gramme activities. These and other methodsdevelopment tools are described in section 3.Feasibility studies addressing economic,

social and technical considerations arerequired prior to any major and costly fieldprogramme. A science-based analysis of theseconsiderations will enable a judgment to bemade as to whether the various control tacticscan be applied on an area-wide basis andwhether the envisioned control strategy is themost appropriate for the particular pest situa-tion. Section 4 provides examples of how suchelucidating studies have been conducted fordifferent pest situations.AW-IPM programmes are dependent on

the synergistic collaboration of many stake-holders. They require the entry onto privateproperties. They can affect the movement ofgoods, and they can also impact or inconven-ience the non-farming community. Thus AW-IPM requires that a sensitive and effectiveregulatory framework be developed by therelevant national and international regulatoryagencies. Commercialization of part or evenentire AW-IPM programmes, a complex andsometimes contentious issue, holds the prom-ise of properly capitalizing such programmes,introducing efficiencies and tackling pestproblems that government cannot afford toaddress. These regulatory and privatizationissues are discussed in section 5.Pilot field programmes are often carried

out following a feasibility study with afavourable outcome. Such programmes areneeded to evaluate and fine-tune various con-trol tactics and field methodologies toincrease their effectiveness and efficiency.Pilot programmes can vary in size and scopeas is described in the chapters in section 6.Section 7 describes operational AW-IPM

programmes against key pests such as the bollweevil, several lepidopteran pests, the bonttick, termites, mosquitoes, fruit flies, etc.Several of the chapters emphasize the techni-cal and managerial difficulties encounteredduring the implementation of eradication, sup-pression, containment or preventive AW-IPMprogrammes and attempt to extract importantlessons.

W. KLASSENviii

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As a concluding chapter (section 8) a criti-cal review is provided of AW-IPM pro-grammes in terms of their successes and fail-ures, and key factors are identified whichmust be addressed in order to improve thechance of success.The chapters in this text book originate

from papers and selected posters presented atthe 2nd FAO/IAEA International conferenceon area-wide control of insect pests. To com-plete the book, several invited chapters havebeen included. This book is an invaluablecompendium of reports on operational AW-IPM programmes. It will help to further devel-op the theory, technology and practice of suchprogrammes. Graduate students will learn

much about the history, accomplishments,problems and the great potential of the area-wide strategy. Entrepreneurs and policy-mak-ers will gain in-depth perspective on aspectsof commercialization.I am honored greatly to have been asked to

write these introductory comments in this textbook devoted to the area-wide management ofinsect pests. The prodigious progress in AW-IPM made in recent decades confirms that thearea-wide strategy has a far greater potentialthan any other approach to achieve sustain-able management of many major insect pests.Truly we are now at the beginning of an era ofdecidedly improved and sustainable insectpest management.

INTRODUCTORY REMARKS ix

Waldemar KlassenProfessor and Program DirectorTropical Research and Education Center,University of FloridaHomestead, Florida 33031, USA

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TABLE OF CONTENTSSECTION 1: SETTING THE SCENE

Area-Wide Integrated Pest Management (AW-IPM): Principles, Practice andProspects....................................................................................................................................3J. HENDRICHS, P. KENMORE, A. S. ROBINSON and M. J. B. VREYSEN

Area-Wide Pest Management: Environmental, Economic and Food Issues......................35D. PIMENTEL

SECTION 2: BASIC RESEARCH

Engineering Insects for the Sterile Insect Technique............................................................51L. S. ALPHEY

The hobo, Hermes and Herves Transposable Elements of Insects.......................................61P. W. ATKINSON, D. A. O'BROCHTA and N. L. CRAIG

Improving the Ecological Safety of Transgenic Insects for Field Release: New Vectors forStability and Genomic Targeting...........................................................................................73A. M. HANDLER, G. J. ZIMOWSKA and C. HORN

M. F. SCHETELIG, C. HORN, A. M. HANDLER and E. A. WIMMER

New Sexing Strains forMediterranean Fruit Fly Ceratitis capitata: Transforming Femalesinto Males.................................................................................................................................95G. SACCONE, A. PANE, A. DE SIMONE, M. SALVEMINI, A. MILANO, L. ANNUNZIATA,

Developing Transgenic Sexing Strains for the Release of Non-Transgenic Sterile MaleCodling Moths Cydia pomonella ..........................................................................................103F. MAREC, L. G. NEVEN and I. FUKOVA

Sex Chromatin Body as a Cytogenetic Marker of W Chromosome Aberrations in Cydiapomonella Females.................................................................................................................113H. MAKEE and N. TAFESH

Potential Use of a Conditional Lethal Transgenic Pink Bollworm Pectinophora gossypiella

Wolbachia-Induced Cytoplasmic Incompatibility to Control Insect Pests?.....................125K. BOURTZIS

Symbiosis-Based Technological Advances to Improve Tsetse Glossina spp. SITApplication.............................................................................................................................137S. AKSOY and B. L. WEISS

Colony Maintenance and Mass-Rearing: Using Cold Storage Technology for Extendingthe Shelf-Life of Insects.........................................................................................................149R. A. LEOPOLD

xi

itata............................................................................................................................................85

in Area-Wide Eradication or Suppression Programmes...................................................119

T. A. MILLER and R. T. STATENG. S. SIMMONS, L. ALPHEY, T. VASQUEZ, N. I. MORRISON, M. J. EPTON, E. MILLER,

U. MAURO and L. C. POLITO

Development of an Embryonic Lethality System in Mediterranean Fruit Fly Ceratitis cap-

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Improving the Efficacy of the Sterile Insect Technique for Fruit Flies by Incorporation ofHormone and Dietary Supplements into Adult Holding Protocols...................................163P. E. A. TEAL, Y. GOMEZ-SIMUTA, B. D. DUEBEN, T. C. HOLLER and S. OLSON

Unfaithful Mediterranean Fruit Fly Ceratitis capitata Females: Impact on the SIT?......175M. BONIZZONI, L. M. GOMULSKI, S. BERTIN, F. SCOLARI, C. R. GUGLIELMINO,

Assessing Genetic Variation in New World Screwworm Cochliomyia hominivorax

T. T. TORRES, M. L. LYRA, P. FRESIA and A. M. L. AZEREDO-ESPIN

Emerging Mosquito-Borne Flaviviruses in Central Europe: Usutu Virus and Novel West

N. NOWOTNY, T. BAKONYI, Z. HUBALEK, H. WEISSENBÖCK, J. KOLODZIEJEK,

SECTION 3: MODELLING AND METHODS DEVELOPMENT

The Role of Geographic Information Systems and Spatial Analysis in Area-Wide VectorControl Programmes.............................................................................................................199J. ST. H. COX

Optimizing Strategies for Eradication of Discrete-Generation Lepidopteran Pests UsingInherited Sterility..................................................................................................................211J. M. KEAN, A. E. A. STEPHENS, S. L. WEE and D. M. SUCKLING

ADiffusion Model for Glossina palpalis gambiensis in Burkina Faso...............................221J. BOUYER, A. SIBERT, M. DESQUESNES, D. CUISANCE and S. DE LA ROCQUE

Current Advances in the Use of Cryogenics and Aerial Navigation Technologies forSterile Insect Delivery Systems.............................................................................................229G. TWEEN and P. RENDÓN

Area-Wide IPM for Commercial Wheat Storage...............................................................239P. W. FLINN, D. W. HAGSTRUM, C. R. REED and T. W. PHILLIPS

Development, Validation and Use of a Simulation Model to Deliver National Predictionsof Ovine Cutaneous Myiasis Risk in the British Isles.........................................................247R. WALL and K. E. PITTS

Problems with the Management of the Golden Apple Snail Pomacea canaliculata: anImportant Exotic Pest of Rice in Asia..................................................................................257R. C. JOSHI

Mass-Rearing and Field Performance of Irradiated Carob Moth Ectomyelois ceratoniaein Tunisia................................................................................................................................265J. MEDIOUNI and M. H. DHOUIBI

Autodissemination of Semiochemicals and Pesticides: a New Concept Compatible withthe Sterile Insect Technique..................................................................................................275P. HOWSE, C. ARMSWORTH and I. BAXTER

SECTION 4: FEASIBILITY STUDIES

Strategies to Control the Desert Locust Schistocerca gregaria..........................................285A. VAN HUIS

TABLE OF CONTENTSxii

B. YUVAL, G. GASPERI and A. R. MALACRIDA

H. LUSSY and B. SEIDEL

Nile Viruses ............................................................................................................................193

Populations from Uruguay...................................................................................................183

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TheMountain Pine BeetleDendroctonus ponderosae inWestern NorthAmerica: Potentialfor Area-Wide Integrated Management..............................................................................297A. L. CARROLL

A Strategy for an Area-Wide Control Campaign with an SIT Component to Establish aTsetse- (Glossina austeni and Glossina brevipalpis) Free SouthAfrica..............................309K. KAPPMEIER GREEN, F. T. POTGIETER and M. J. B. VREYSEN

Area-Wide Control of Tsetse and Trypanosomosis: Ethiopian Experience in the SouthernRift Valley...............................................................................................................................325

Don't Let Cacto Blast Us: Development of a Bi-National Plan to Stop the Spread of theCactus Moth Cactoblastis cactorum in North America......................................................337K. BLOEM, S. BLOEM, J. CARPENTER, S. HIGHT, J. FLOYD and H. ZIMMERMANN

Preventive Programme Against the Cactus Moth Cactoblastis cactorum in Mexico......345J. HERNÁNDEZ, H. SÁNCHEZ, A. BELLO and G. GONZÁLEZ

Area-Wide Control Tactics for the False Codling Moth Thaumatotibia leucotreta in SouthAfrica: a Potential Invasive Species.....................................................................................351J. CARPENTER, S. BLOEM and H. HOFMEYR

SIT for the Malaria Vector Anopheles arabiensis in Northern State, Sudan: an HistoricalReview of the Field Site.........................................................................................................361C. A. MALCOLM, D. A. WELSBY and B. B. EL SAYED

Integrated Management of Rice Stem Borers in the Yangtze Delta, China......................373Z-R. ZHU, J. CHENG, W. ZUO, X-W. LIN, Y-R. GUO, Y-P. JIANG, X-W. WU, K. TENG,

Management of Cotton Insect Pests in Tajikistan..............................................................383S. M. MUKHITDINOV

Insecticidal Wound Treatment of Livestock on Isla de la Juventud, Cuba: an EfficientSuppression Method of New World Screwworm Cochliomyia hominivorax Prior to theRelease of Sterile Insects ......................................................................................................393R. GARCIA, L. MENDEZ, E. SERRANO, T. GIL MORALES and M.J.B. VREYSEN

SECTION 5: COMMERCIALIZATION AND REGULATION

Area-Wide Integrated Pest Management Programmes and Agricultural Trade:Challenges and Opportunities for Regulatory Plant Protection.......................................407C. DEVORSHAK

Systems Approaches as Phytosanitary Measures: Techniques and Case Studies............417E. V. PODLECKIS

High Dose Applications.........................................................................................................425P. A. FOLLETT

Tools for the Trade: the International Business of the SIT...............................................435M. M. QUINLAN and A. LARCHER-CARVALHO

Privatizing the SIT: a Conflict Between Business and Technology?..................................449B. N. BARNES

TABLE OF CONTENTS xiii

G. WOLDEYES, K. BEKELE and U. FELDMANNT. ALEMU, B. KAPITANO, S. MEKONNEN, G. ABOSET, M. KIFLOM, B. BANCHA,

B-P. ZHAI, J. LUO, X-H. JIANG and Z-H. TANG

Postharvest Phytosanitary Radiation Treatments: Less-Than-Probit 9, Generic Dose, and

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Private Sector Investment in Mediterranean Fruit Fly Mass-Production and SITOperations – The “Sheep” of the Private Sector Among the “Wolves” of the PublicGood?.....................................................................................................................................457Y. BASSI, S. STEINBERG and J. P. CAYOL

SECTION 6: PILOT PROGRAMMES

Assessment of the Sterile Insect Technique to Manage Red Palm Weevil Rhynchophorusferrugineus in Coconut..........................................................................................................475R. KRISHNAKUMAR and P. MAHESWARI

Area-Wide Suppression of Invasive FireAnt Solenopsis spp. Populations.......................487R. K. VANDER MEER, R. M. PEREIRA, S. D. PORTER, S. M. VALLES and D. H. OI

ACultural Method for the Area-Wide Control of Tarnished Plant Bug Lygus lineolaris inCotton.....................................................................................................................................497C. A. ABEL, G. L. SNODGRASS and J. GORE

Use of the Sterile Insect Technique Against Aedes albopictus in Italy: First Results of aPilot Trial................................................................................................................................505R. BELLINI, M. CALVITTI, A. MEDICI, M. CARRIERI, G. CELLI and S. MAINI

Area-Wide Integrated Control of Oriental Fruit Fly Bactrocera dorsalis and Guava FruitFly Bactrocera correcta in Thailand.....................................................................................517W. ORANKANOK, S. CHINVINIJKUL, S. THANAPHUM, P. SITILOB and W. R. ENKERLIN

Establishment of a Mediterranean Fruit Fly Ceratitis capitata, Fruit Fly Parasitoids andCodling Moth Cydia pomonella Rearing Facility in North-Eastern Brazil ......................527A. MALAVASI, A. S. NASCIMENTO, B. J. PARANHOS, M. L. Z. COSTA and J. M. M.WALDER

Pilot Mediterranean Fruit Fly Ceratitis capitata Rearing Facility in Tunisia: Constraintsand Prospects.........................................................................................................................535M. M'SAAD GUERFALI, A. RAIES, H. BEN SALAH, F. LOUSSAIEF and C. CÁCERES

SECTION 7: OPERATIONALAW-IPM PROGRAMMES

Progress of Boll Weevil Anthonomus grandis Eradication in the United States of America,2005.........................................................................................................................................547O. EL-LISSY and B. GREFENSTETTE

Regional Management Strategy for Cotton Bollworm Helicoverpa armigera inChina......................................................................................................................................559K. M. WU

Integrated Systems for Control of the Pink Bollworm Pectinophora gossypiella inCotton.....................................................................................................................................567T. J. HENNEBERRY

Pulling Out the Evil by the Root: the Codling Moth Cydia pomonella EradicationProgramme in Brazil............................................................................................................ 581A. KOVALESKI and J. MUMFORD

Suppression of the Codling Moth Cydia pomonella in British Columbia, Canada Using anArea-Wide IntegratedApproach with an SIT Component................................................591S. BLOEM, A. McCLUSKEY, R. FUGGER, S. ARTHUR, S. WOOD and J. CARPENTER

TABLE OF CONTENTSxiv

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Eradication of the Australian Painted Apple Moth Teia anartoides in New Zealand:Trapping, Inherited Sterility, andMale Competitiveness..................................................603D. M. SUCKLING, A. M. BARRINGTON, A. CHHAGAN, A. E. A. STEPHENS, G. M.BURNIP, J. G. CHARLES and S. L. WEE

Area-Wide Management of the Formosan Subterranean Termite Coptotermes formosanusin New Orleans’ French Quarter..........................................................................................617A. R. LAX, F. S. GUILLOT and D. R. RING

A Multi-Institutional Approach to Create Fruit Fly-Low Prevalence and Fly-Free Areasin Central America................................................................................................................627J. REYES, X. CARRO, J. HERNANDEZ, W. MÉNDEZ, C. CAMPO, H. ESQUIVEL, E. SAL-GADO and W. ENKERLIN

The Fruit Fly Exclusion Programme in Chile.....................................................................641J. GONZALEZ and P. TRONCOSO

Expansion of the National Fruit Fly Control Programme in Argentina...........................653D. GUILLÉN and R. SÁNCHEZ

The Augmentative Biological Control Component in the Mexican National CampaignAgainst Anastrepha spp. Fruit Flies.....................................................................................661P. MONTOYA, J. CANCINO, M. ZENIL, G. SANTIAGO and J. M. GUTIERREZ

The Hawaii Area-Wide Fruit Fly Pest Management Programme: Influence ofPartnerships and a Good Education Programme...............................................................671R. F. L. MAU, E. B. JANG and R. I. VARGAS

Area-Wide Management of Fruit Flies in Australia...........................................................685A. J. JESSUP, B. DOMINIAK, B. WOODS, C. P. F. DE LIMA, A. TOMKINS and C. J. SMALL-RIDGE

Five Years of Mosquito Control in Northern Greece..........................................................699N. PIAKIS, G. IATROU, S. MOURELATOS and S. GEWEHR

The Carribean Amblyomma variegatum Eradication Programme: Success or Failure?.................................................................................................................................................709R. G. PEGRAM, A. J. WILSMORE, C. LOCKHART, R. E. PACER and C. S. EDDI

SECTION 8: LESSONS LEARNED

Lessons from Area-Wide Integrated Pest Management (AW-IPM) Programmes with anSIT Component: an FAO/IAEAPerspective.......................................................................723M. J. B. VREYSEN, J. GERARDO-ABAYA, and J. P. CAYOL

Author Index..........................................................................................................................745

Subject Index.........................................................................................................................749

TABLE OF CONTENTS xv

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Section 1

Setting the Scene

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Area-Wide Integrated Pest Management(AW-IPM): Principles, Practice and

Prospects

J. HENDRICHS1, P. KENMORE2, A.S. ROBINSON3

and M.J.B. VREYSEN1

1Joint FAO/IAEA Programme of Nuclear Techniques in Food andAgriculture, Insect Pest Control Sub-Programme, IAEA,

Wagramerstrasse 5, A-1400 Vienna, Austria2Plant Production and Protection Division, Food and AgricultureOrganization of the United Nations, Viale delle Terme di Caracalla

00100 Rome, Italy3Joint FAO/IAEA Programme of Nuclear Techniques in Food andAgriculture, FAO/IAEA Agriculture and Biotechnology Laboratory,

Seibersdorf A-2444, Austria

ABSTRACT Integrated pest management (IPM) has remained the dominant paradigm of pest controlfor the last 50 years. IPM has been endorsed by essentially all the multilateral environmental agreementsthat have transformed the global policy framework of natural resource management, agriculture, and trade.The integration of a number of different control tactics into IPM systems can be done in ways that greatlyfacilitate the achievement of the goals either of field-by-field pest management, or of area-wide (AW) pestmanagement, which is the management of the total pest population within a delimited area. For severaldecades IPM and AW pest control have been seen as competing paradigms with different objectives andapproaches. Yet, the two “schools” have gradually converged, and it is now generally acknowledged thatthe synthesis, AW-IPM, neither targets only eradication, nor relies only on single control tactics, and thatmany successful AW programmes combine a centrally managed top-down approach with a strong grass-roots bottom-up approach, and that some are managed in a fully bottom-up manner. AW-IPM is increas-ingly accepted especially for mobile pests where management at a larger scale is more effective and prefer-able to the uncoordinated field-by-field approach. For some livestock pests, vectors of human diseases, andpests of crops with a high economic value and low pest tolerance, there are compelling economic incen-tives for participating in AW control. Nevertheless issues of free riders, public participation and financingof public goods, all play a significant role in AW-IPM implementation. These social and managerial issueshave, in several cases, severely hampered the positive outcome of AW programmes; and this emphasisesthe need for attention not only to ecological, environmental, and economic aspects, but also to the socialand management dimensions. Because globalization of trade and tourism are accompanied by the increasedmovement of invasive alien pest species, AW programmes against major agricultural pests are often beingconducted in urban and suburban areas. Especially in such circumstances, factors likely to shift attitudesfrom apathy to outrage, need to be identified in the programme planning stage and mitigated. This paperreviews the evolution and implementation of the AW-IPM concept and documents its process of develop-ment from basic research, through methods development, feasibility studies, commercialization and regu-lation, to pilot studies and operational programmes.

KEYWORDS area-wide IPM, field-by-field IPM, suppression, eradication, feasibility studies, pilotprogrammes, operational programmes, commercialization, regulation, public good, free rider, public par-ticipation

3

© 2007 IAEA., 3–33.M.J.B. Vreysen, A.S. Robinson and J. Hendrichs (eds.), Area-Wide Control of Insect Pests

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1. IntroductionIf major advances are to be made in coping withmost of the major arthropod pest problems, thenthe tactics and strategies for managing suchinsects, ticks and mites must change. They mustchange from the current, limited scale, reactive,broad-spectrum measures to preventive meas-ures that are target-pest specific and rigidlyapplied on an area-wide basis (Knipling 1992).

Around 850 million people remain malnour-ished (FAO 2006) in spite of significantprogress over the last four decades towardsfood security in several regions of the world,and numerous positive developments in thearea of food and agriculture. In 1996, theWorld Food Summit held in Rome, Italyaddressed this persistent crisis by launchingthe very ambitious goal, later incorporatedinto the Millennium Development Goals, ofhalving the number of hungry people by 2015(FAO 1996). However, the Food andAgriculture Organization of the UnitedNations (FAO) has analysed the world foodinsecurity situation and indicated that, unfor-tunately, progress is insufficient to meet theSummit’s target (FAO 2006). Furthermore,the world population is expanding by ca 75-80million people each year, and most likely willrise to about 9000 million by the year 2050,and thus require food production levels atleast 50% higher than those in 2000(Alexandratos 1999). At the same time agri-cultural research and extension budgets andaid to agriculture are shrinking, naturalresources are degrading, and the growth of theworld’s agricultural production is continuingto slow down (Bautista and Valdés 1993,Braun et al. 1993, Alexandratos 1999).

This continued rapid growth of the worldpopulation, which is causing the biggest surgein demand for food in history, includingdemands for significantly more animal pro-teins (Delgado et al. 1999) and biofuels,stands in stark contrast to the shrinking percapita land area available for agriculture. Thisdiscrepancy cannot be addressed by horizon-tal agricultural expansion, i.e. by an increasein cultivated surface area, but requires devel-opment and promotion of more intensive

cropping and livestock systems on existingfarmland (Borlaug 1997). Access to new tech-nologies and the knowledge to use or adaptthem locally will be vital, coupled with lesswastage and improved penetration intonational and global markets. As expansion ofarable land will only play a minor role in someregions and attaining substantial increases incrop yields will become increasingly difficultin others, the focus will have to shift towardsa more efficient use of agricultural resources(Trewavas 2001). There could hardly be a lessefficient use of resources than to invest land,water, fertilizer, seeds, labour, and energy toproduce agricultural commodities, only tohave the investment partially or totallydestroyed by insects or other pests. Preharvestlosses in developing countries are estimated atmore than one third of attainable crop produc-tion, while postharvest losses add at leastanother 10-20%. Insects, followed bypathogens and weeds, cause the largest por-tion of these losses (FAO 1975, Oerke et al.1995, Yudelman et al. 1998, Thomas 1999).

Insects have proven to be among the mostformidable adversaries of mankind. Sincethey appeared on the scene some 390 millionyears ago, they have diversified into severalmillion species that have adapted to almost allavailable ecosystems. This large diversity hasallowed them to effectively compete withmankind since the introduction of agricultureover the last ten millennia. The increase inagricultural production of the past decades hasnot been accompanied by a comparable reduc-tion in overall losses inflicted by insect pests:on the contrary, agricultural intensificationhas increased both yields and vulnerability topests (Yudelman et al. 1998). Furthermore ourmobile society is redistributing species aroundthe globe at an unprecedented pace with majorconsequences for agriculture and ecosystems(FAO 2001). Undoubtedly, insects will contin-ue to challenge mankind, and their resource-fulness is forcing a review of established waysof dealing with these pests and stimulating thedevelopment of new innovative control tactics(Klassen 2005). Investing in improved pestmanagement should therefore be an integral

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component of national strategies to raise pro-ductivity and to assure future global foodsecurity.

The sudden availability of very effectiveand persistent synthetic organic insecticidesimmediately after the Second World Warmarked the onset of chemically-based warfareagainst most insect pests. Before that, insectpests had to be kept at bay by making use ofvarious natural control factors. The impor-tance of the new synthetic chemicals to con-trol vectors of major diseases and their contri-butions to the green revolution cannot bequestioned (Oerke et al. 1995). Chemical con-trol has offered an effective and economicalway to deal with a multitude of arthropodproblems, to quell outbreaks, to decimate vec-tors of parasitic and infectious diseases ofhumans and livestock, to suppress noxiousinsect developing in dung on rangelandswhere domestic animals are produced, to sup-press mites in honey bee hives, and to controltermites and numerous household pests inurban ecosystems, etc. Pesticides have offeredthe pesticide user the freedom and flexibilityto control pests on his property at any timewithout regard for the opinions and actions ofhis neighbours. Without chemical pest con-trol, global agricultural productivity wouldhave been less, food prices much higher andthe available food of lower quality (Knipling1979). The ready accessibility of these “off-the-shelf”, relatively cheap and often subsi-dized chemicals was undoubtedly one of themain reasons the uncoordinated control of keyinsect pests on a field-by-field basis becameso widely and firmly established during thelast 60 years.

The drawbacks of the widespread use ofthese broad-spectrum insecticides were, how-ever, recognized early (Carson 1962). Theypollute soils and water, are a hazard to manynon-target and beneficial insects, lead to out-breaks of secondary pests, cause acute andchronic poisoning of farmers, accumulate andbiomagnify in the food chain and representserious concerns to human health (Repettoand Baliga 1996). The general public’sincreased awareness and demand for more

environment-friendly pest control tactics, theswift development of pesticide resistance, andthe need for ever new, more complex andmuch costlier products, gave rise to the con-cept of integrated pest control (IPC) (Stern etal. 1959, FAO 1966), later called integratedpest management (IPM) (Bottrell 1979). Inthe 1960-70s, stimulated by this need forreduced and more selective use ofinsecticides, IPM became gradually acceptedas a viable and sustainable pest managementstrategy that incorporates some of the tradi-tional practices of the pre-insecticide era(such as field sanitation, biological controland use of pest and disease resistant livestockbreeds and crop varieties) together with moreselective synthetic organic pesticides to main-tain pest population levels below an economicthreshold. IPC-IPM has remained the domi-nant paradigm of pest control for the last 50years (Kogan 1998).

2. Integrated Pest Management(IPM): a Reaction Against the

Abuse of Insecticides

IPM offers a strategic approach to solving pestproblems in an ecosystem context whileguarding human health and the environment(Brader 1979). It has been endorsed by essen-tially all the multilateral environmental agree-ments that, since the United NationsConference on Environment andDevelopment in Rio de Janeiro in 1992, havetransformed the global policy framework ofnatural resource management, agriculture, andtrade. More than half the world’s human pop-ulation lives in countries that are guided bynational IPM policies, and that account formost of the world’s staple crop production.IPM is increasingly practiced in agroecosys-tems featuring perennial tree crops, annualfield crops, crops, in protected cultivation,ornamental crops, rangelands, intensive pas-tures, roadways, recreational parks, forests,dairies, barnyards, and urban ecosystems.Small-scale family farms in tropical and tem-perate zones as well as multinational corpo-rate food producing and processing firms

AREA-WIDE INTEGRATED PEST MANAGEMENT 5

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apply IPM. IPM systems range over a contin-uum, from those still dependent to a consider-able degree on the use of pesticides all theway to those called biointensive that rarelyrequire chemical treatments (Vandeman et al.1994). Along this continuum, reliance on pes-ticide treatment-oriented interventionsdecreases and reliance on biological and cul-tural practices increases and requires anincreasing number of available methods inmanaging pests (Benbrook et al. 1996).

2.1. FAO’s Code of Conduct on theDistribution and Use of Pesticides

The International Code of Conduct on theDistribution and Use of Pesticides (revisedfrom the original 1985 version) was adoptedby the 123rd session of the FAO Council inNovember 2002. The Code embodies a mod-ern approach, leading to sound managementof pesticides with a focus on risk reduction,protection of human and environmentalhealth, and support for sustainable agricultur-al development by selectively using pesticidesas a component of various IPM strategies(FAO 2003a). Thus, the Code designed stan-dards of conduct to promote IPM, includingthe integrated management of public healthvectors.

The Code defines IPM as:

…the careful consideration of all available pestcontrol techniques and subsequent integrationof appropriate measures that discourage thedevelopment of pest populations and keep pesti-cides and other interventions to levels that areeconomically justified and reduce or minimizerisks to human health and the environment. IPMemphasizes the growth of a healthy crop with theleast possible disruption in agroecosystems andencourages natural pest control mechanisms(FAO 2003a).

The Code has been approved and adoptedby 185 FAO member governments, endorsedby non-governmental organizations and bythe pesticide industry. Therefore the definitionof IPM in the Code carries a degree of acces-sible authority that many other published def-initions of IPM do not (Bajwa and Kogan

1996). In its article on pesticide management,the Code calls for support to alternatives toconventional pesticides:

Governments, with the support of relevant inter-national and regional organizations, shouldencourage and promote research on, and thedevelopment of, alternatives posing fewer risks:biological control agents and techniques, non-chemical pesticides and pesticides that are, asfar as possible or desirable, target-specific, thatdegrade into innocuous constituent parts ormetabolites after use and are of low risk tohumans and the environment.

The Code reflects evolving responses tochanging conditions with emphasis on pro-tecting the integrity of agroecosystems,encouraging natural pest control mechanisms,and reducing risks to human health and theenvironment. In the articles of the Code onpesticide management, the wider range ofstakeholders, and the emphasis on promotingincreased participation of farmers, women’sgroups, and others reflect recent experienceswith successful IPM (van den Berg 2004).

2.2. Selected Successes of IPM

IPM has had a varied history, and simply mix-ing different management tactics does not con-stitute IPM (Ehler and Bottrell 2000).Successful IPM is “knowledge intensive” andhas never been successful without basicresearch on ecosystems, particularly to compre-hend the food webs or communities of speciesthrough which energy flows in agroecosystems(Barfield and Swisher 1994, Wood 2002).Understanding why a pest becomes a pest(Lewis et al. 1997) comes from a fundamentalunderstanding of the pest’s life history and theecology of crop-pest-natural enemy interac-tions, and rarely from a revolutionarily newcontrol tactic (Kogan 1998, Thomas 1999).

The rice brown planthopper Nilaparvatalugens (Stål) was the key insect pest of theAsian green revolution in rice. While singletactics of insecticides and vertical host-plantresistance largely dominated research and pestcontrol application, a series of studies in themid 1970s elucidated the mechanism of out-

J. HENDRICHS ET AL.6

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breaks of secondary pests due to the overuse ofsubsidized insecticides. This was shownthrough analyses of multi-species rice fieldagroecosystems that concentrated on the ricecrop, the planthoppers, their predators, and par-asitoids. The results, made available by FAO,were then successfully applied in the mid 1980sthrough national IPM policies and programmesin the Philippines, Indonesia, India, Vietnam,China, and other major rice-producing coun-tries (DeBach and Rosen 1991, Kogan 1998,Bartlett 2005). As a result of these policies,over USD 250 million per year in governmen-tal insecticide subsidies were eliminated.

Subsequently Settle et al. (1996) showedhow the rice field’s aquatic food web, driven bydecomposition of rice roots, rice straw, andother organic matter from previous seasons,through dozens of aquatic arthropods, producedsufficient numbers of predators, to protect therice crop from the seedling stage to maturity.Rice field dwelling arthropod species that donot feed on rice still serve as important foodsources in building up populations of naturalenemies of rice pests. The species richness ofnatural freshwater ecosystems (streams, ponds,rivers, and lakes) permits irrigated rice agroe-cosystems to draw from their larger naturalspecies pools to quickly fill in the essentialguild structure of the cultivated aquatic riceagroecosystem, buffering it from immigrantpest populations. In addition, Ives and Settle(1997) challenged the conventional wisdom onsynchronous rice planting, showing that in thepresence of natural enemies asynchronous riceplanting results in the lowest overall pest densi-ties, since early arriving generalist predatorsdecimate incipient infestations of pests andsuppress their populations to a greater extentthan is accomplished by killing large numberslater.

3. Field-by-Field and Area-Wide Pest Management

Approaches

3.1. Field-by-Field Pest Management

Insect pest control measures can be applied

either field-by-field (Fig. 1) or on an AW basis(Fig. 2), the latter addressing the total pestpopulation within a delimited area. The sim-plest and most widely used strategy has beenfield-by-field management, which addressesonly small fractions of a pest population atany given time. It allows individual crop andlivestock producers, households, and busi-nesses, to control pests independently, withoutinvesting effort in coordination, without hav-ing to obtain the consent or collaboration ofother stakeholders and most importantly with-out taking into account the pest individualsthat frequently migrate into the treated areafrom infestations in the untreated surround-ings. Insects, themselves, are mobile but, canalso be transported passively with wind, onanimal hosts, or in infested commodities trad-ed locally or internationally. This mobilityseverely compromises the effectiveness ofuncoordinated farm-by-farm, orchard-by-orchard, or herd-by-herd control efforts, andresults in the frequent need for curative ortherapeutic back-up measures (Lewis et al.1997) and the eventual overreliance on them.However, field-by-field pest control does notdemand long-term commitment to an organ-ized effort and its funding requirements, butrelies on remedial interventions triggeredwhen a pest population reaches a certainthreshold. Field-by-field pest control is, there-fore, largely a reactive approach to protecthumans, animals, crops, forests, houses,wooden structures, etc., rather than a preven-tive pest population management approach(Pedgley 1993, Abeku 2007).

As a result of the complexity of manyagroecosystems, as well as the site-specificnature of a majority of pest problems, pre-determined thresholds often become opera-tionally intractable and in some pest situa-tions, the threshold is zero tolerance (Ehlerand Bottrell 2000). Thus a field-by-field IPMapproach is often insufficient, particularlywhen pests are quite mobile. Furthermore, thecost of generating the large amount of ecolog-ical information needed to develop and imple-ment functional IPM systems for such localsituations cannot be afforded by most devel-

AREA-WIDE INTEGRATED PEST MANAGEMENT 7

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J. HENDRICHS ET AL.8

Figure 1. Graphic display of field-by-field IPM, where the pest is suppressed below an eco-nomic threshold level in areas of commercial interest, but often not in abandoned crops, alter-nate hosts, backyard hosts or on wild hosts. As a result, significant untreated refugia of the pestremain from which recruits re-establish damaging densities of the pest population.

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AREA-WIDE INTEGRATED PEST MANAGEMENT 9

Figure 2. Graphic display of AW-IPM, where the pest is suppressed below an economic thresh-old level in all areas, including abandoned crops, alternate hosts, backyard hosts or on wildhosts. As a result, no significant untreated refugia of the pest remain from which recruits canre-establish damaging densities of the pest population.

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oping countries (Morse and Buhler 1997).

3.2. The Principle of Total PopulationManagement

Knipling (1960, 1972) used simple populationmodels to show that small fractions of aninsect pest population left uncontrolled canrapidly nullify the benefits of strongly sup-pressing the main pest population in a largearea. One of these models compares thedynamics of an hypothetical insect pest popu-lation that has a fivefold natural rate ofincrease per year in an area, and where eachyear 99 percent of the population is destroyedon 90% of the host resources (but no control isconducted on the remaining 10%) with an areawhere only 90% of the population isdestroyed on 100% of the total host resources(Table 1). The model shows that because nocontrol was exercised on 10% of the hostresources, 100 times more pests were pro-duced in the first scenario (where 99% killwas achieved each generation in the largetreated fraction) than in the second scenariowhere no refugia were left but the total popu-lation was subjected to only 90% kill in eachgeneration.

From this Knipling (1972, 1979) deduced

the basic principle of total population control:Uniform suppressive pressure applied againstthe total population of the pest over a period ofgenerations will achieve greater suppressionthan a higher level of control on most, but notall, of the population each generation.

These refugia are permanent and prolificsources of immigrants and therefore representa constant economic threat, requiring applica-tions in cultivated areas of significantamounts of insecticides year after year, evenin situations where major pests are managedunder an IPM approach. This failure toaddress the total pest population in an areaoften compromises the basic goal of IPM, i.e.the reduction of insecticide use. In contrast thecentral paradigm of AW control, variouslyalso called landscape, large-scale, preventiveor total population management (Pedgley1993, Ekbom et al. 2000, Carrière et al. 2001,Smith et al. 2006), recognizes the need toaddress the existence of all foci/refugia fromwhich recruits can invade suppressed orcleared areas (Byers and Castle 2005, Klassen2005).

Total population management is alsorequired to reduce the probability of the devel-opment of insecticide resistance, or of theemergence of strains of a pest capable of over-

J. HENDRICHS ET AL.10

Generation

Number of insects in an area with

99 percentage control on 90 percentage of the targetpopulation

90 percentage control on 100 per-centage of the target population

treated area (90%) untreated area (10%) treated area (100%)

1 900 000 90001 100 000 1000 000 100 00012 45 000 450 500 000 500 000 50 0003 2250 22 2 500 000 250 000 25 0004 110 12 500 000 125 000

Total insects in 4th generation: 12 500 110 125 000

Table 1. Relative number of insects developing each generation in two hypothetical populationmanagement systems. Each of the two populations has an initial population size of one millioninsects and a fivefold increase in reproducing insects is assumed with each generation (afterKnipling 1979).

1number of insects surviving treatment

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coming host resistance. In particular, AW pestmanagement strategies, guided by the effec-tive use of geographic information systems(GIS) technology, can be applied to achieveeffective resistance management (Carrière etal. 2001, Sexson and Wyman 2005, Wu, thisvolume). Nevertheless, there is a potentialdanger in AW population suppression, sincecontinuous and thorough suppression appliedon an AW basis will select genes in the pest’sgene pool that enable the pest to overcome thesurvival threatening control agent. For exam-ple, in the absence of refugia, as part of aresistance management programme, repeatedAW applications of an insecticide will selectfor resistance to that insecticide, repeated AWuse of baits will select for avoidance behav-iour, and repeated AW planting of the sameresistant crop variety will select genes forovercoming the crop’s host resistance.

3.3. Origins of Area-Wide (AW)Approaches

The AW approach is not new but evolved cen-turies ago. Reduced cost, increased effective-ness and greater physical protection were gener-ally the underlying reasons for the developmentof AW services. In earlier times, essential itemssuch as the water supply, fuel, lighting, andsewage disposal were provided on an individualbasis and each extended family satisfied theseneeds independently of its neighbours. This wasan uncoordinated approach with the result thatthese individual services often proved to be notonly unreliable, but also inefficient and couldonly be obtained at great individual expense.The desire for more effective and efficientdelivery of these essential services transformedthem from “individual” to “area-wide” and ledeventually to the creation of public or privatecompanies that supplied clean water and elec-tricity, and that took responsibility for the col-lection of the garbage and disposal of sewage.Likewise, originally each head of the clan tookresponsibility for the physical protection of hiskin. Soon feudal societies developed in whichsome individuals and later institutions special-ized to provide protection for much larger

groups. Eventually, police forces, state andnational armies emerged that were able to dealwith larger-scale military threats. The AWapproach for these services made them moreeffective and less costly because the providerscould use technically more advanced methodsthat were not available to the individual(Lindquist 2000). Today, AW services (e.g. mailservice, retailing, ambulance, fire protection,public health, high-speed transport, telephone,internet services, etc.) are much more wide-spread than ever before.

3.4. Area-Wide Approaches Applied toInsect Pest Control

Applied to insect pests, AW approaches havetheir ancient roots in coping with vector-bornediseases and locust plagues (Klassen 2000,2005). In the 14th century, the systematic useof quarantines in some European city-statescontained bubonic plague transmitted by theoriental rat flea Xenopsylla cheopsis(Rothschild) and this approach was graduallyadopted throughout Europe. The late 19th cen-tury included AW approaches such as thedevelopment of classical biological controland the use of pest-resistant plants (for exam-ple the grafting of all European grapes onphylloxera-resistant American rootstocks inthe 1870s).

A campaign to eradicate the invasive gypsymoth Lymantria dispar (L.) from 1890 to1901 in Massachusetts mainly using a mod-estly efficient phytotoxin, was quite success-ful initially but had to be abandoned after pub-lic opposition to the spraying. Since then, theobjective of this AW campaign has reverted tolimiting or retarding the spread of this pest(Sharov et al. 2002).

In 1906, a massive effort in the southernUSA was initiated to eradicate two Boophiluscattle tick species that transmit cattle tickfever (Klassen 1989). A strategy of starvingthe ticks by making most pastures cattle free,combined with an arsenic-dipping programmeand restricting cattle movement to tick-freecounties was rigorously implemented for 37years. In 1943, the ticks had been eliminated

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from the USA at a total cost that was equiva-lent to the yearly losses before the programmewas initiated. To date, the southern USA hasremained free of these ticks, as the result of aneffective quarantine programme.

In 1911 the Government of Portugal decid-ed to eradicate the tsetse fly Glossina palpalispalpalis Robineau-Desvoidy and trypanoso-mosis from the Island of Principe off the westcoast of equatorial Africa. The suppressivesystem consisted of sticky black cloths wornby workers, treatment of people with the try-panocidal arsenical, atoxyl, clearing of vege-tation, corralling of domestic livestock anderadication of feral pigs. The campaign wasconcluded in 1914 (McKelvey 1973): it wasthe first successful tsetse eradication pro-gramme.

After the World Health Assembly urgedthe World Health Organization (WHO) toorganize the eradication of malaria in 1955,enormous progress was made in the following15 years, and malaria was declared eradicatedin 37 countries (Spielman et al. 1993). Socialpressure to devolve more control to the locallevel and the banning of DDT resulted in thedisintegration of the programme. To date,more than 400 million malaria cases arereported annually worldwide and more thanone million people, mainly children in Africa,succumb to the disease every year (Marshall2000).

In the 1950s and 1960s, a significant AWcampaign was implemented against the intro-duced Khapra beetle Trogoderma granariumEverts which had become established in thesouth-western USA and northern Mexico. Amajor quarantine and treatment effort, whichinvolved the fumigation of all warehouses,seed storage facilities, ships and other trans-port facilities, was initiated cooperatively byall states in these regions. Funding for this AWprogramme was shared between the federaland state governments and the private sector.By 1966 all known populations of the pest hadbeen eradicated. Another introduction of thispest was eliminated between 1980 and 1983in the north-eastern USA (Klassen 1989).

In 1973, the exotic cassava mealybug

Phenacoccus manihoti Matile-Ferero wasobserved attacking cassava around Kinshasa(Democratic Republic of Congo, formerlyZaire) and Brazzaville (Republic of Congo)and in subsequent years it dispersed through-out sub-Saharan Africa, causing starvation ofmore than 200 million people(Neuenschwander et al. 1988). TheInternational Center for Tropical Agriculture,Cali, Colombia, identified a suitable para-sitoid Epidinocarsis lopezi DeSantis inParaguay, which was brought to Africa by theInternational Institute for Tropical Agriculturein Ibadan, Nigeria. The parasitoid was suc-cessfully mass-reared and released in 38African countries. This is an outstandingaccomplishment in classical biologicalcontrol, and it continues to keep the mealybugat bay.

Cereal aphids have become a major threatto production of sorghum, wheat and barley incentral-western North America following theemergence of virulent biotypes of the greenbug Schizaphis graminum (Rondani) duringthe 1960s, the establishment of the Russianwheat aphid Diuraphis noxia (Kurdjumov) in1986, and the subsequent emergence of newbiotypes of the latter species. These pests havebeen causing annual losses in grain produc-tion in excess of USD 250 million, and havecaused large and unsustainable increases inthe use of insecticides on these low profit mar-gin crops. Therefore, a very large classicalbiological control programme involving theintroduction of parasitoids from Eurasia wasinititated (Brewer and Elliott 2004). However,an analysis of the wheat agroecosystem hasrevealed the presence and economic impact ofmany species of indigenous natural enemies,and it has been difficult to demonstrate anycontribution from imported exotic species. Asin the above mentioned problem of the brownplanthopper on rice, understanding the com-munity structure of the agroecosystem, initial-ly intended as a side product of the classicalbiological control campaign, turned out to bethe crucial foundation of the AW-IPM pro-gramme. Many other examples of successfulor unsuccessful AW programmes to suppress

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or eradicate insect pest populations are listedin Klassen (1989, 2005).

3.5. Public Participation, Public Good,and Free Riders

As discussed above and by Vreysen et al. (thisvolume), AW programmes are not always suc-cessful. Social concern over methods, toomany free riders, or insufficient complianceby all stakeholders have in several casesseverely hampered success (Klassen 2000),emphasizing the need for attention not only toecological, environmental, and economic, butalso to social and managerial dimensions.

3.5.1. Public ParticipationPublic participation or “ownership” by thegeneral public of AW programmes is crucialfor their success. Yet often such support isweak, and special public information effortsare needed to convince the stakeholders of thewider benefits. Mumford (2000) summarizesthis aspect well:

The main area in which problems lie with area-wide pest control appears to be in the mecha-nisms for public participation. Reports overmany years cite technical, economic and envi-ronmental success with the concept, but there isstill indifference, reluctance and antagonism.

... these attitudes may be the result of a lack ofopportunity for involvement and ownerships ofprogrammes, which may be seen as beingimposed from above/outside, managed by tech-nocrats or otherwise not arising from or meetingthe needs of the people directly concerned. Noneof these issues should be insurmountable, but itis worth noting that area-wide pest managementis an activity in which social participation andattention to the “reasonable person” is asimportant as technical proficiency.

Outrage factors pose a great threat to AWprogrammes. Once a large segment of thepublic has become outraged, it is almostimpossible to lead them back to an attitude oftrust and support (Sandman 1987). People insignificant numbers have expressed outrageagainst AW programmes conducted for thebenefit of agriculture or public health in urbansettings, and such instances are likely to

increase because of the recent surge in fre-quency of establishment of invasive alienpests near international airports, seaports andelsewhere in metropolitan areas. Protesters inurban communities fearful of mandatory pes-ticide applications attempted to halt the cam-paigns to eradicate the Mediterranean fruit flyCeratitis capitata (Wiedemann) in the LosAngeles Basin by the State of California andthe United States Department of Agriculture(USDA) during the 1980s (Lorraine andChambers 1989). During the recent campaignto eradicate citrus canker from Florida (1995-2006), protesters in urban communities insouth-eastern Florida, outraged by the entry ofinspectors and workers into backyards ofhomes without search warrants and thedestruction of apparently healthy citrus trees,delayed programme implementation from2001 to 2004 by challenging its legality incourt. In 2004 three hurricanes spread thecanker bacterium from infected trees leftuntouched during the three-year litigation sowidely that the programme was judged to benot feasible, and it was terminated in 2006after the expenditure of ca USD 875 million(Bouffard 2006). In India a WHO programmeto eradicate Aedes aegypti (L.), the vector ofyellow fever and dengue, was terminated dur-ing the 1970s because journalists incited irra-tional fear of the sterilized male mosquitoes(Nature 1975). Clearly potential outrage fac-tors need to be identified during the planningstages of AW programmes, and a well fundedpublic information programme must belaunched at the outset of each programme.Public information specialists must be trainedto be up front, open and honest about allissues which may be perceived negativelyespecially by urban stakeholders.

3.5.2. Public GoodPublic health, a clean environment, ecosystemservices, roads, public education, a whole-some food supply, etc., have all been labelledas public goods (Johnson 2005). The applica-tion of IPM in agriculture and human healthgives rise to positive externalities for the com-mon or public good, resulting in benefits such

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as reduced risks to humans and the environ-ment. In contrast a negative externality ariseswhen a farmer, through injudicious pesticideuse, pollutes the environment and harms hisneighbours. In the case of mobile pests, dam-age is a function of the total pest population inthe entire area. Pest control by any farmerresults, therefore, in pest suppression and lessdamage for the neighbours as well, althoughonly the one farmer incurs the cost of the con-trol. This spill-over benefit for the other farm-ers in the vicinity is a positive externality,which unfortunately encourages free ridersand is often not conducive for a collaborativeprogramme against the pest within a commu-nity or at larger scales (Yu and Leung 2006).

3.5.3. Free RidersFree riders are individuals choosing to benefitfrom a public good or a positive externalitywithout contributing to the costs of producingthe benefits (Johnson 2005), and they repre-sent a problem for IPM implementationagainst mobile pests. Farmer associations or“community IPM”, effectively promoted byFAO through IPM farmer field schools indeveloping countries (van den Berg 2004),can help address this problem. Actually com-munity IPM is more than pest managementand offers an entry point to improve the farm-ing system as a whole, developing theenhanced management skills necessary forsustainable environment-friendly agriculturaland rural development (Dilts 2001).Nevertheless, the cultural and socio-economicbackground of stakeholders significantlyaffects the collaboration rate in community-based projects (Smith et al. 2006, Stonehouseet al. 2007).

Non-collaborators represent a major weak-ness when operating programmes at a largerscale (for example regional public health vac-cination campaigns, or global rinderpest orpolio eradication). A few free riders or“refuseniks” can negate many positiveimpacts of AW programmes. Since externali-ties may affect a large number of stakeholders,obtaining full participation and dealing withsceptical, negligent, or even antagonistic third

parties is an unavoidable and difficult chal-lenge. In addition, merely because a good issaid to be public, such as an unpolluted or dis-ease-free environment, does not automaticallyimply that all people value it equally.Nevertheless, often a public good cannot beprovided, or provided to an adequate extent,without strong support for a mechanism ofcollective action.

3.5.4. Subsidies, Government Provision, andLegal AuthorityState subsidies, or complete government pro-vision, and/or regulation or even legal author-ity for enforcement by authorities can at leastpartially overcome this type of market failure.All of these solutions represent an “involun-tary” provision through taxation to partially orfully fund the public good. With benefits atleast as large for society as for growers andother IPM practitioners who carry the finan-cial burden of implementation, a strong casehas been made for financial incentives as astimulus to entice more participation andexpansion of IPM adoption (Brewer et al.2004). Since benefits to society of AW inter-ventions tend to be unusually large, the alloca-tion of public funds to many of such pro-grammes in public health and agricultureappears to be strongly justified, and indeed,many AW pest control programmes are par-tially funded by the state (Bassi et al., this vol-ume). The establishment and enforcement oflegal rules by authority (Klassen 2000), forexample, the removal of mosquito breedingsites, the implementation of quarantines, orthe removal of wild hosts in the surroundingsof crops (Kovaleski and Mumford, this vol-ume) is greatly facilitated by establishing such“official” AW pest management activities.

3.6. Linking Area-Wide Approaches withIPM (AW-IPM)

The concept of AW pest control, developed byUSDA under the direction of E. F. Knipling,has been closely identified with the pro-gramme to eradicate the New World screw-worm Cochliomyia hominivorax (Coquerel),

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which started in Florida in 1957 and reachedPanama in 2001 (Klassen and Curtis 2005).During these same decades many academicinstitutions developed and promoted the IPMparadigm. For several decades IPM and AWpest control were seen as competing para-digms with different objectives and approach-es (Perkins 1982), and each competing againstthe other with its own core organizationalbase, i.e., state universities (IPM) versus fed-eral research and regulatory agencies (AW).

3.6.1. Suppression versus EradicationWhile under IPM suppression, pests can betolerated at certain levels as part of healthyagroecosystems in which all components havea functioning role, IPM practitioners per-ceived AW pest control as targeting only erad-ication, an end point to which some had strongphilosophical objections. Practitioners of AWcontrol were accused of only wanting to erad-icate pest populations based entirely on theinitial promise of synthetic insecticides andthat they were unwilling to manage and other-wise learn to live with pests. However in real-ity, Knipling (1966, 1969, 1972) had early onincluded suppression in AW control of keypests, although the possibility of eradicationof selected populations of major pest species,if practical and economically and environ-mentally advantageous, was preferable(Kogan 1998). For example, since about 1960Knipling worked toward the eradication ofboll weevil Anthonomus grandis Boheman inthe USA, but in this he was effectivelyopposed for decades by IPM practitioners.Eradication of this key invasive pest on cot-ton, which was finally initiated in the 1980sand is now nearing completion (El-Lissy andGrefenstette, this volume), was foreseen tosignificantly and permanently reduce insecti-cide use and provide major environmental andeconomic benefits. It was also foreseen thatremoval of the pest would cause much cottonproduction to shift from the central part of thecotton belt to the south-eastern USA, where inthe absence of this pest cotton production isthe most profitable. Currently none of the sev-eral ongoing AW control programmes are

intended to eradicate the target plant pest orpest complex; instead they are designed tomanage selected pests across an expansivegeographic landscape (Pedgley 1993, Coop etal. 2000, Hendrichs et al. 2005, Sexson andWyman 2005, Carrière et al. 2006, Abeku2007).

In recent years organizations concernedwith the preservation of biodiversity and con-servation of natural ecosystems have becomealarmed that their investments in conservationare at serious risk of being undone by invasivealien species (Sklad et al. 2003). Moreover,conservation scientists have found eradicationto be essential for the restoration of certainnatural ecosystems badly damaged by inva-sive species, and attitudes toward eradicationare being revisited (Myers et al. 2000, Cloutand Veitch 2002, Simberloff 2002, PérezSandi Cuen and Zimmermann 2005).

3.6.2. Area-Wide IntegrationWhile the integration of compatible controltactics was seen as the cornerstone of the IPMconcept, adherents of the IPM school did notperceive such integration to be integral to AWpest control systems, even though most AWsuppression or eradication campaignsemployed simultaneously several control tac-tics to achieve their goal. However, the inte-gration of all available weapons is a funda-mental requirement for the success of bothfield-by-field and AW-IPM. For example theNew World screwworm eradication pro-gramme relied not only on the release of ster-ile insects, but integrated their use into a sys-tem including quarantines to prevent spreadand reinvasion, closely scheduled examina-tion of all livestock, collection and identifica-tion of specimens from wounds, diligent treat-ment of all wounds, navels and other sites ofoviposition of the parasite, scheduling of cul-tural practices related to livestock manage-ment such as branding, castration and dehorn-ing only when the parasite was least prevalent,and extensive public information activities toobtain transparency and secure the collabora-tion of all stakeholders. Clearly, the NewWorld screwworm eradication programme

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targeted both the adult and immature stages ofthe parasite. Indeed Knipling (1979) proposedintegrating control tactics that address imma-ture stages (for example natural enemies) withthose that target the adult stage (sterile insects,mating disruption), or those that are veryeffective against high population densities(use of bio-pesticides and baits) with thosethat are effective against low population den-sities (mating disruption systems, sterileinsects, parasitoids, etc.).

3.6.3. Top-Down ApproachWhile for IPM a bottom-up approach at thefarmer and community level was the opera-tional mode, AW control was seen as needinga top-down approach, centrally managed byan organization, and with a mandatory com-ponent to insure full participation of stake-holders within a region. Even though AW pro-grammes are usually, but not always, central-ly managed, they cannot afford to ignore theconcerns of farmers and communities. IndeedAW programmes cannot succeed unless theysecure the active enthusiastic participation ofall stakeholders, especially that of farmers andrural communities to achieve their goals. Andwhere such a programme involves urban com-munities, it is even more important to assurethat the urban people understand the impor-tance of the programme and are willing to tol-erate and contribute to its costs.

While not all AW-IPM programmesinclude mandatory components to insurestakeholder participation, a regulatory frame-work undoubtedly facilitates programmeeffectiveness. Some regulations establishedby regulatory authorities with stakeholderinput are critical to success of certain AW pro-grammes. These may include mandatory cropdestruction by a certain date to prevent over-wintering of the pest, mandatory seeding orplanting dates to provide for suicidal emer-gence of the pest, access to private propertyby inspectors, etc. Since such regulations maybe difficult to enforce, it is important that atthe outset a referendum is held to determine ifat least two-thirds of the stakeholders wouldwillingly comply. On the other hand special

regulations are not needed to assure the suc-cess of many AW programmes, and in suchinstances it would be most unwise to imposethem.

3.6.4. What is Area-Wide?The term area-wide, coined for total popula-tion management is widely entrenched, eventhough it can be misleading (and has been atleast partially responsible for misunderstand-ing the concept), since the concept deals pri-marily with a total population in a delimitedarea, the influence of migration/dispersal onits dynamics, and its ecological relationshipswithin its ecosystem. Including the distribu-tion of the pest population in space and avail-ability in time appears to be the logical nextstep in the ongoing evolution of IPM fromoriginally managing a single pest in a field, tomultiple pests in a field, to a larger scale thatincludes multiple fields and multiple crops.Actually large geographic areas are not a pre-requisite for the AW approach, becauseaddressing pest populations within a closedgreenhouse, for example, also involves man-aging them at the population level, where theirtemporal dynamics and spatial distribution areknown (Casey et al. 2007).

3.6.5. Merging of IPM and AW ControlOver time, as described above, the twoschools have gradually converged and the dif-ferences between them have turned out to beless critical than originally perceived(Coppedge 1994, Kogan 1994, 1995, Parry1995, Kogan 1998, Coop et al. 2000, Tan2000, Faust 2001, Yu and Leung 2006). AW-IPM is a very broad and flexible concept andis increasingly accepted for those situations ofmobile pests where management at a largerscale is advantageous to maximize the AW,not necessarily local, efficacy of managementtactics (Cronin et al. 1999). At the same timegrower education, for example through theFAO-organized IPM farmer field schools, isalso leading to some concerted action to scaleup the IPM movement at the community level(Dilts 2001, Pontius et al. 2002). Rice farmersare gradually moving from field-by-field to

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AW-IPM implementation by coordinating atthe village, subdistrict, and in some caseseven district levels (Matteson 2000).

The interaction of AW application of tac-tics and coordination of activities amongstakeholders is also not straightforward.While synchronous AW application of controlmeasures is generally more efficient to pre-clude pest population refugia (Byers andCastle 2005), in other situations AW spatialasynchrony is deliberately adopted over broadgeographic regions. For example, spatialasynchrony of control measures has a greatereffective suppressive effect on the total pestpopulation in the rice agroecosystem, becauseit favours earlier migration of natural enemiesinto the rice fields (Ives and Settle 1997).

3.6.6. Situations Advantageous for Area-WideControlEconomics undoubtedly plays a major role inthe initial grower decision to participate inAW-IPM (Sexson and Wyman 2005), anddeteriorating market conditions may cause thegrower to neglect or even abandon a crop in afield or an orchard. Farmers who cultivatecrops with a high economic value and lowpest tolerance risk suffer greater losses thanfarmers who cultivate crops with a low eco-nomic value and high pest tolerance (Yu andLeung 2006). In the latter situation there arefewer incentives for farmers to cooperatethrough an AW approach, whereas in the firstcase the economic advantages of participationare much greater (Stonehouse et al. 2007).This is particularly so for crops such as veg-etables and fruit, or for some livestock orhuman diseases, where the acceptable thresh-olds are so low that the presence of even a fewpest or vector individuals often triggers theneed for remedial applications.

Using a mathematical model, Yu andLeung (2006) derived several favourable andunfavourable conditions for implementingAW-IPM. In their view, AW-IPM is more like-ly to succeed where the number of farmers issmall, and the cultivated crops are similar(low farm heterogeneity). The stability of thecooperation among the farmers is enhanced

by short detection times and high discountrates. The model likewise demonstrates that aone-off suppression of the pest under the lead-ership of a third party facilitates the coopera-tion of heterogeneous groups of farmers inAW-IPM.

4. Area-Wide Control of InsectPests: from Research to Field

Implementation

Operational AW-IPM programmes are com-plex, long-term and proceed from basicresearch, through methods development, fea-sibility studies, commercialization and regula-tion, to field pilot studies, which could even-tually culminate into an operational pro-gramme. The various components of thisdevelopment process are briefly documentedand discussed.

4.1. Basic Research

AW-IPM is a science-based activity and,hence, relies on knowledge generated in basicscience to provide new tools and technologies.The link between basic science and AW-IPMprogrammes is often seen as tenuous, as muchtime and applied research is required to turn abasic invention into a product for use in a fieldprogramme. One example of this process hasbeen the basic genetic research conductedover ten years that culminated in the develop-ment of genetic sexing strains for theMediterranean fruit fly. Genetic sexing hasrevolutionized the use of the sterile insecttechnique against this pest. Unfortunately thistechnology cannot be transferred directly toother species, although it could be created inmany pest species with somewhat less effortthan was required in the Mediterranean fruitfly. Moreover, genetic transformation ofinsect pests may lead to the development ofgeneric sexing systems. Many of the proposeduses of genetic transformation aim to replaceor complement existing technologies, and,therefore, do not represent completely newconcepts. As such, genetic sexing, insectmarking, and sterilization are all proposed as

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targets for genetic transformation. In a fewpest species all these goals have beenachieved at the laboratory scale, but the largerchallenge of scaling up and validating thesesystems at meaningful levels for AW-IPMprogrammes remains to be done.

Genetic transformation was first developedto study gene function in Drosophilamelanogaster Meigen and then roughly tenyears of effort were required to simulate thisaccomplishment in the first major pestspecies, the Mediterranean fruit fly. Now, it ispossible, through the development of bettervector systems and transformation markers, togenetically engineer most pest species, includ-ing several dipteran and lepidopteran pests(Atkinson, this volume). Lepidoptera have adifferent sex determination system thanDiptera and this difference could be exploitedto produce a genetic sexing strain with non-transgenic males for release (Marec et al., thisvolume). The identification of specific chro-mosome markers (Makee and Tafesh, this vol-ume) will aid this approach.

Each AW-IPM programme requires a regu-latory framework tailored to its specific needs.However none of the currently existing regu-latory frameworks provides for the deploy-ment of any genetically transformed product.This issue was discussed in Rome at a meet-ing on risk assessment, which provided someguidance on deploying transgenic arthropods(IAEA 2006). Currently, the North AmericanPlant Protection Organization (NAPPO) isdeveloping a regional standard (RSPM) tofacilitate the importation and confined releaseof transgenic arthropods (NAPPO 2007).Strategies to facilitate the acceptance ofdeploying transgenic insects in AW-IPM pro-grammes are being developed (Handler et al.,this volume).

Characterization of pest populations to betargeted by an AW-IPM programme is anessential first step in mounting an AW-IPMprogramme. To this end advances in DNAanalysis are providing increasingly sophisti-cated tools to assess the genetic variability ofdifferent field populations of a pest speciesincluding the degree of isolation between

them (Torres et al., this volume). However,these studies do not provide information onthe presence or absence of behavioural matingbarriers between various field populations orbetween released sterile insects and targetpopulations. The existence of behavioural bar-riers to mating must be determined throughthe conduct of mating compatibility studies;and these should be conducted under semi-natural conditions (Cayol et al. 2002, Vera etal. 2006).

The quality of performance in the field ofinsects released in AW-IPM programmes is ofparamount importance. Therefore, great dili-gence is exercised in the mass-rearing andhandling of predators and parasitoids in aug-mentative biocontrol programmes and of ster-ile males released in SIT programmes. Thus itis important to note that dietary, hormonal,and semiochemical treatment of sterile malesprior to release (Teal et al., this volume) cangreatly increase their effectiveness. Similarconclusions were reached with respect topredators reared for release in augmentativebiocontrol programmes (Thompson 1999).

Likewise, new techniques that allow thestorage of insects for varying lengths of timeand at various stages could be very beneficialto AW-IPM programmes. Maintenance of cer-tain strains of the pest insect that have partic-ular characteristics is labour intensive, expen-sive and tends to expand as new strains aredeveloped. Strains also need to be maintainedover long periods of time even though theymay only be used for limited periods. In addi-tion, storage of insects can alleviate potentialhazards of the rearing process such as straindeterioration, disease outbreaks, labourunrest, or mechanical failure. Cryopreser-vation is considered to be a possible solution,and recent advances have made this technolo-gy available for some pest insects such astransgenic New World screwworm (Leopold,this volume).

Population suppression in AW-IPM pro-grammes can be achieved by reducing thereproductive potential of the target populationusing either natural or artificially inducedsterility. Four decades ago, cytoplasmic

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