Nematodes as Biocontrol Agents This page intentionally left blank Nematodes as Biocontrol Agents Edited by Parwinder S. Grewal Department of Entomology Ohio State University, Wooster, Ohio USA Ralf-Udo Ehlers Department of Biotechnology and BiologicalControl Institute forPhytopathology Christian-Albrechts-University Kiel,Raisdorf Germany David I. Shapiro-Ilan United States Department ofAgriculture Agriculture ResearchService Southeastern Fruitand Tree Nut Research Laboratory,Byron,Georgia USA CABI Publishing CABIPublishingisadivisionofCABInternational CABIPublishing CABIPublishing CABInternational 875MassachusettsAvenue Wallingford 7thFloor OxfordshireOX108DE Cambridge,MA02139 UK USA Tel:þ44(0)1491832111 Tel:þ16173954056 Fax:þ44(0)1491833508 Fax:þ16173546875 E-mail:[email protected] E-mail:[email protected] Website:www.cabi-publishing.org (cid:1)CABInternational2005.Allrightsreserved.Nopartofthispublication maybereproducedinanyformorbyanymeans,electronically,mech- anically,byphotocopying,recordingorotherwise,withouttheprior permissionofthecopyrightowners. A catalogue record for this book is available from the British Library, London,UK. LibraryofCongressCataloging-in-PublicationData Nematodes as biocontrol agents / edited by Parwinder S. Grewal, Ralf- UdoEhlers,DavidI.Shapiro-Ilan. p.cm. Includesbibliographicalreferencesandindex. ISBN0-85199-017-7(alk.paper) 1. Nematoda as biological pest control agents. I. Grewal, Parwinder S. II.Ehlers,Ralf-Udo.III.Shaprio-Ilan,DavidI. SB976.N46N462005 632’.96–dc22 2004030022 ISBN0851990177 TypesetbySPIPublisherServices,Pondicherry,India PrintedandboundintheUKbyBiddlesLtd.,King’sLynn ThisvolumeisdedicatedtoDrHarryK.Kayainrecognitionofhisnumerouscontributions toinsectpathology.Hisearlycontributionstotheecologyofentomopathogenicnematodes provided a solid foundation for the development of application strategies for use of nematodes in insect biocontrol. Dr Kaya has co-edited four books on entomopathogenic nematodes and insect pathology/biological control and co-authored a book on insect pathology.Hehaspublishedmorethan230researchpapersontheecologyandapplication of insect nematodes and other pathogens and is one of the co-founding editors of the journal Biological Control. His outstanding leadership and scholarly accomplishments, spanningnearlythreedecades,haveplayedakeyroleinexpandingresearchandapplica- tionininsectnematologytolaboratoriesandindustriesaroundtheworld. This page intentionally left blank Contents Contributors xi Preface xv GlossaryofTerms xvii PARTI. NEMATODEMORPHOLOGYANDTAXONOMY 1 1. MorphologyandSystematicsofNematodesUsedinBiocontrol 3 S.P.StockandD.J.Hunt PARTII. ENTOMOPATHOGENICNEMATODES 45 2. BiologyandBehaviour 47 C.T.Griffin,N.E.BoemareandE.E.Lewis 3. MassProduction 65 R.-U.EhlersandD.I.Shapiro-Ilan 4. FormulationandQuality 79 P.S.GrewalandA.Peters 5. ApplicationTechnology 91 D.J.Wright,A.Peters,S.SchroerandJ.P.Fife 6. ForumonSafetyandRegulation 107 R.-U.Ehlers 7. Lawn,TurfgrassandPastureApplications 115 P.S.Grewal,A.M.Koppenho¨ferandH.Y.Choo 8. GlasshouseApplications 147 M.Tomalak,S.PiggottandG.B.Jagdale 9. NurseryandTreeApplications 167 R.W.H.M.VanTolandM.J.Raupp vii viii Contents 10. MushroomApplications 191 S.Jess,H.SchweizerandM.Kilpatrick 11. OrchardApplications 215 D.I.Shapiro-Ilan,L.W.Duncan,L.A.LaceyandR.Han 12. SoftFruitApplications 231 R.S.Cowles,S.Polavarapu,R.N.Williams,A.ThiesandR.-U.Ehlers 13. VegetableandTuberCropApplications 255 G.Be´lair,D.J.WrightandG.Curto 14. Cereal,Fibre,OilseedandMedicinalCropApplications 265 H.E.Cabanillas,R.J.WrightandR.V.Vyas 15. ForestryApplications 281 P.Torr,M.J.WilsonandS.Heritage 16. ApplicationsfortheControlofPestsofHumansandAnimals 295 I.Glazer,M.SamishandF.G.delPino 17. ApplicationsforSocialInsectControl 317 D.H.Gouge 18. ASystemsApproachtoConservationofNematodes 331 M.E.BarbercheckandC.W.Hoy 19. InteractionswithPlant-parasiticNematodes 349 E.E.LewisandP.S.Grewal 20. CompatibilityandInteractionswithAgrochemicalsandOther BiocontrolAgents 363 A.M.Koppenho¨ferandP.S.Grewal PARTIII. ENTOMOPHILICNEMATODES 383 21. ApplicationofBeddingiasiricidicolaforSirexWoodwaspControl 385 R.A.BeddingandE.T.Iede 22. TheEntomophilicThripinema 401 J.FunderburkandK.S.Latsha 23. MermithidNematodes 411 E.G.Platzer,B.A.MullensandM.M.Shamseldean PARTIV. SLUG-PARASITICNEMATODES 419 24. Biology,ProductionandFormulationofSlug-parasiticNematodes 421 M.J.WilsonandP.S.Grewal 25. ApplicationofSlug-parasiticNematodes 431 A.EsterandM.J.Wilson PARTV. PREDATORYNEMATODES 445 26. PotentialofPredatoryNematodestoControlPlant-parasiticNematodes 447 A.L.BilgramiandC.Brey Contents ix PARTVI. FUNGAL-FEEDINGNEMATODES 465 27. PotentialofFungal-feedingNematodesfortheControlofSoil-borne PlantPathogens 467 N.Ishibashi PARTVII. CONCLUSIONSANDFUTUREDIRECTIONS 477 28. CriticalIssuesandResearchNeedsforExpandingtheUseof NematodesinBiocontrol 479 P.S.Grewal,R.-U.EhlersandD.I.Shapiro-Ilan Index 491 This page intentionally left blank