J Appl Entomol. 2018;142:731–744. wileyonlinelibrary.com/journal/jen  | 731© 2018 Blackwell Verlag GmbH

Received:13January2018  |  Revised:5April2018  |  Accepted:3May2018DOI:10.1111/jen.12528

O R I G I N A L C O N T R I B U T I O N

Improved monitoring of oriental fruit moth (Lepidoptera: Tortricidae) with terpinyl acetate plus acetic acid membrane lures

Valentina Mujica1 | Michele Preti2 | Esteban Basoalto3 | Liliana Cichon4 |  Eduardo Fuentes-Contreras5 | Wilson Barros-Parada5,6 | Greg Krawczyk7 |  Marcelo Z. Nunes7 | Jim F. Walgenbach8 | Randy Hansen9 | Alan L. Knight10

1InstitutoNacionaldeInvestigaciónAgropecuaria,Montevideo,Uruguay2FacultyofScienceandTechnology,FreeUniversityofBozen-Bolzano,Bolzano,Italy3InstitutodeProducciónySanidadVegetal,FacultaddeCienciasAgrarias,UniversidadAustraldeChile,Valdivia,Chile4EstaciónExperimentalAgropecuaria,INTA—AltoValle,GeneralRoca,RioNegro,Argentina5MillenniumNucleusCentreinMolecularEcologyandEvolutionaryApplicationsintheAgroecosystems(CEM),FacultaddeCienciasAgrarias,UniversidaddeTalca,Talca,Chile6EscueladeAgronomía,PontificiaUniversidadCatólicadeValparaíso,Quillota,Chile7DepartmentofEntomology,FruitResearchandExtensionCenter,ThePennsylvaniaStateUniversity,Biglerville,Pennsylvania8MountainHorticulturalCropsResearch&ExtensionCenter,NCStateUniversity,MillsRiver,NorthCarolina9HansenAssociates,Placerville,California10USDA,AgriculturalResearchService,Wapato,Washington

CorrespondenceAlanL.Knight,USDA,AgriculturalResearchService,Wapato,WA.Email:alan.knight@ars.usda.gov

Funding informationFondodeDesarrolloCientíficoyTecnológico(FONDECYT),Grant/AwardNumber:3140285;MillenniumNucleusCentreinMolecularEcologyandEvolutionaryApplicationsinAgroecosystemsandInciativaCientificaMilenio,Grant/AwardNumber:NC120027

AbstractMaleandfemalemothcatchesofGrapholita molesta(Busck)intrapswereevaluatedinstoneandpomefruitorchardsuntreatedortreatedwithsexpheromonesformat-ingdisruptioninUruguay,Argentina,Chile,USA,andItalyfrom2015to2017.Trialsevaluatedvariousblendsloadedintoeithermembranecupluresorsepta.Membranelureswereloadedwithterpinylacetate(TA),aceticacid(AA)and(Z)-3-hexenylace-tatealoneor incombinations.Twosepta lureswere loadedwitheitherthethree-component sex pheromone blend for G. molesta alone or in combination withcodlemone(2-PH),thesexpheromoneofCydia pomonella(L).Athirdseptumlurein-cludedthecombinationsexpheromoneblendpluspearester,(E,Z)-2,4-ethyldecadi-enoate (2-PH/PE), and a fourth septumwas loadedwith only β-ocimene. Resultswereconsistentacrossgeographicalareasshowingthattheadditionofβ-ocimeneor(Z)-3-hexenylacetatedidnotincreasemothcatches.Theadditionofpearestertothesexpheromoneluremarginally increasedmothcatches.TheuseofTAandAAto-gethersignificantlyincreasedmothcatchescomparedwiththeuseofonlyoneofthetwocomponents.TrapswiththeTA/AAlureoutperformedtheAjartrapbaitedwithaliquidTAplussugarbait.TheemissionrateofAAwasnotasignificantfactoraffect-ingtheperformanceoftheTA/AAlure.TheadditionofTA/AAsignificantlyincreasedmothcatcheswhencombinedwiththe2-PHlure.TheTA/AAlurealsoallowedtraps

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1  | INTRODUC TION

The use of sex pheromones for themanagement of oriental fruitmoth,Grapholita molesta(Busck),hasbeenbroadlyadoptedthrough-outitsworldwidedistributioninbothstone(Prunusspp.)andpome(Malus domesticaBorkhausen,andPyrusspp.)fruitsandnowincludestheuseofhand-applieddispensers,aerosols,sprayablesandattractandkillformulations(DeLame&Gut,2006;Evenden&McLaughlin,2004;Il’ichev,Stelinski,Williams,&Gut,2006;Knight,Light,Pickel,Kovanci,&Molinari,2008;Kovanci,Schal,Walgenbach,&Kennedy,2005;Steliniski,Gut,Haas,McGhee,&Epstein,2007;Steliniskietal.,2005;Trimble,Pree,Barszez,&Carter,2004;Vickers,Rothschild,&Jones,1985).Akeycomponentinthemanagementofthisimportantpest,aswithothertortricidtreefruitpests,iseffectivemonitoringtoestablishbothaction thresholdsand to timesupplementary in-secticidetreatmentsundereitherconventionalorcertifiedorganicpractices (Charmillot & Vickers, 1991; Knight & Light, 2005a,b).Unfortunately,theuseofsexpheromonelurestomonitorthesea-sonal population density of maleG. molesta is problematic in sexpheromone-treated orchards, but the absence of moth catch canbeagoodindicatorofeffectivedisruptionofsexualcommunication(Rice & Kirsch, 1990; Vickers, 1990).Many growers, instead, relyonbuckettrapswithliquidbaits(terpinylacetateplusbrownsugar,TAS)tomonitorpopulations(Il’ichev,Williams,&Gut,2007;Kovanci&Walgenbach, 2005; Rice&Kirsch, 1990; Rothschild, Vickers, &Morton,1984).TAS-baited traps catchboth sexesandcan reflecttheproportionofmatedfemaleswithinapopulation(Vickers,1990),butarecumbersometouseandmaintainduetoevaporation,spill-age and required sorting of various nontarget species (Kovanci &Walgenbach,2005).Developmentofdryterpinylacetateluresthatcanbeusedinstandardtrapswithstickylinershasbeenreported,butthesetrapswerenotaseffectiveasbuckettrapsusingtheliquidTASbait(Knightetal.,2011).

DevelopmentofthenovelAjartrapswherethe liquidbaitwasplaced in a screen container inside the trap and moths were re-tainedonstandardstickylinerssurroundingthescreenofferedanimprovedmonitoring technique, and these traps achieved compa-rableperformancetoTAS-baitedbuckettraps(Cichonetal.,2013;Knight, Cichon, etal., 2014; Padilha, Arioli, Boff, Rosa, & Botton,2017). However, the nonselectivity (high catches of dipterans,

noctuids,andvespids)ofAjartrapswasoftenfoundtorequirefre-quentlinerchanges.Unfortunately,theadditionofscreenstoAjartraps,whilesignificantlyreducingthenontargetcatch,alsoreducedmothcatches in some studies (Knight,Cichon, etal., 2014).WhileAjartrapseliminatedthesortingofnontargetsintheliquidbait,therequiredperiodichandlingofthebaitstillmadethesetrapsrelativelycumbersometouse,andtheyhavenotbeencommercialized.

Alternatively,variousstudieshavefocusedonidentifyingplantvolatilesaspotentialattractantsforadultG. molesta. Individual and blends of various peach, pear and apple volatiles have been eval-uated in laboratory assays, with some blends being identified asattractive (Molinarietal.,2010;Najar-Rodriguez,Orschel,&Dorn,2013; Natale, Mattiacci, Hern, Pasqualini, & Dorn, 2003, 2004;Natale,Mattiacci,Pasqualini,&Dorn,2004;Piñero&Dorn,2007;Varela,Avilla,Anton,&Gemeno,2011).Fieldstudieshavealsore-ported that several volatile blends were attractive, including inAustralia (Il’ichev,Kugimya,Williams,&Takabayashi, 2009) and inChina (Lu, Huang, &Wang, 2012; Lu,Wang,Wang, Luo, &Qiao,2015; Lu etal., 2014). However, these blendswere not shown tobeattractive insubsequent field trialsconducted inChile (Barros-Parada, Ammagarahalli, Basoalto, Fuentes-Contreras, & Gemeno,2018).StudiesintheUnitedStatesfoundthattheadditionofeither(E)-β-farnesene,(E)-β-ocimeneorbutylhexanoateseptaluresintheAjartrapwiththeTASbaitincreasedtotalmothcatches,and(E)-β-ocimenealsoincreasedfemalemothcatchescomparedwiththeTASbaitalone(Knightetal.,2013).Theadditionofthegreenleafvolatile(Z)-3-hexen-1-ylacetateto(E)-β-ocimeneincombinationwitheither(E)-β-farneseneorbutylhexanoatedidnot increasemothcatches.Previously, the addition of (Z)-3-hexen-1-yl acetatewas shown toincreasethenumberofG. molestacaughtinacleartrapwithadryterpinyl acetate lure (Knight etal., 2011). Acetic acid is typicallypresentinsugarybaits(Utrio&Erikson,1977)andhasbeendirectlyusedwithethanolinliquidbaittrapsinChina(He,Qin,&Zhu,2009).However,theuseofaceticacidcolureswithplantvolatileshasnotpreviouslybeeneffectivewithG. molesta(Knight,Hilton,Basoalto,&Steliniski,2014).Studieswithtrapsbaitedwithacombinationofterpinylacetateandaceticacidhavenotbeenpreviouslyreported.

Septaluresformulatedwiththethree-componentsexpheromoneofG. molesta plus the major sex pheromone component ofCydia pomonella (L.), codlemone, (E,E)-8,10-dodecadien-1-ol, significantly

tocatchbothsexes.CatchofC. pomonellawiththe2-PHlurewascomparabletotheuseofcodlemone;however,mothcatchwassignificantlyreducedwiththe2-PH/PElure.Optimizationofthesecomplexlurescanlikelyfurtherimprovemanagers’abilitytomonitorG. molestaandhelptodevelopmultispeciestortricidluresforuseinindi-vidualtraps.

K E Y W O R D S

apple,Cydia pomonella,Grapholita molesta,matingdisruption,peach

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increasedmothcatches innondisruptedorchards (Allred,Croft,&Riedl, 1995). However, these lures were only intermediate whencomparedwithanumberofcommercialsexpheromoneluresinor-chards treatedwith sex pheromone (Knight, Basoalto,& Stelinski,2015).Also,thecombinationsexpheromonelurewassignificantlylessattractive toC. pomonella than standard sexpheromone luresforthisspecies,andthismaylimititsuseinpomeorchardsmanag-ingbothC. pomonella and G. molesta(Evenden&McLaughlin,2005;Il’ichevetal.,2007;Steliniski,Il’ichev,&Gut,2009;Steliniskietal.,2007).Theadditionofaceticacid,(E)-β-ocimene,orpearesterwiththis combination sex pheromone only marginally increased mothcatches,includingfemalemoths(Knight,Cichon,etal.,2014).

Despitethis largeandrecentbodyofresearchtodevelopnewattractants and trapdesigns formonitoringG. molesta, further re-finements are likely still needed. Furthermore, the potential ef-fectiveness of various attractants should be tested over a widegeographical area to consider intraspecific variation in sensorycommunication channels, as was recently performed with maleG. molestatoitssexpheromoneblend(Knight,Barros-Parada,etal.,2015).Geographicaldifferencesinhostrangeexpansionscouldalsoprovideselectivepressureformoths’responsetovariouschemicalsignals.Forexample,initialstudieswithpearesterinvariousstonefruits in California found that it was not attractive to G. molesta (Knight&Light,2004).Yet,studiesinItalyreportedthatantennaeinbothsexesareresponsivetopearesterand, inanolfactometerassay, primarily males were more attracted to pear ester than toappleorpearshootvolatiles,butnotincomparisonwithpeachshootvolatiles(Molinarietal.,2010).

Herein, we report studies from Uruguay, Chile, Argentina,Italyand intheeasternandwesternUnitedStatesevaluatingsev-eral long-lasting membrane lures used alone and in combinationwith septa lures for G. molesta in apple and stone fruit orchardsuntreated or treated with sex pheromones for mating disruption.Variouskairomonalcombinationsweretestedandincludedterpinylacetate,aceticacid,pearester,β-ocimeneandgreenleafvolatiles.Kairomones were tested alone and in combination with the sexpheromoneofG. molesta and C. pomonella.Bisexuallureswerede-velopedwhichcansignificantlyimprovemonitoringofG. molesta in sex pheromone-treated orchards. Further studies fine-tuning lureblendsforbisexualcatchofmultipletortricidspeciesiswarranted.

2  | MATERIAL S AND METHODS

2.1 | Traps, lures and field study protocol

Studies were conducted with delta-shaped traps and several com-mercial and experimental lures during 2015–2017. Traps includedthe standard orange or white Pherocon VI trap (28×20×20cm)usedwithpaperlinerscoatedwithapolybutaneadhesive(TrécéInc.,Adair,OK).ThecommercialsexpheromoneluresPheroconOFM-L2(93:6:1blendof(E)-8-dodecenylacetate,(Z)-8-dodecenylacetateand(Z)-8-dodecenol)andPheroconCML2loadedwithcodlemonewereprovidebyTrécé Inc.Avial lure loadedwith5.0mlofglacialacetic

acid(99.7%,Sigma-Aldrich,StLouis,MO)wasmadebydrillinga3.2-mmholeintothecapof8-mlnalgenevials(NalgeNuncInternational,Rochester,NY)andloadingeachvialwithtwosmallcottonballs.

Experimental lures including grey halobutyl septa, red rubberseptaandwhiteplasticmembraneluresloadedwithsexpheromonesorhostplantvolatilesaloneorcombinedwerepreparedbyTrécéInc.chemists and shipped to the various researchers. The three com-ponents of G. molesta pheromone (E)-8-dodecenyl acetate (98.6%purity), (Z)-8-dodecenylacetate (98.1%purity)and (Z)-8-dodecenol(98.3%purity)and thesexpheromoneofC. pomonella, codlemone,(E,E)-8,10-dodecadien-1-ol (97% purity), were purchased fromBedoukianResearch,Danbury,CT.Greyhalobutylseptalureswereprepared with a 93:6:1 ratio of this three-component pheromoneblendalone(OFM-PH)orwithcodlemone(2-PH).Athirdgreyseptumlurewasloadedwithpearester,(E,Z)-2,4-ethyldecadienoate(>92%purity); and a fourth septum lure combined the combination sexpheromonewithpearester(2-PH/PE).Aredrubberseptum(BO)wasloadedwitharacemicmixtureofβ-ocimene(>90%(E)-isomer,Sigma-Aldrich).Severalproprietaryplasticmembranecuplures(1.8cmdi-ameter)werealsopreparedbyTrécé Inc.These includedTRE1114loadedwithterpinylacetate(TA);TRE1237loadedwith(Z)-3-hexenylacetate(Z3);TRE3321loadedwithaceticacid(AA);TRE1370loadedwith a 1:1 ratio of terpinyl acetate and acetic acid (TA/AA); andTRE1367loadedwitha1:1:1ratioofterpinylacetate,aceticacidand(Z)-3-hexenyl acetate (TA/AA/Z3). Two additional acetic acid lureswith different membranes used to achieve higher emission ratesthanTRE3321werealsoevaluated:TRE1468 (AA2) and the larger(36cm2) redTRE1531 (AA3) lure.Theweight lossofallmembraneluresloadedwithterpinylacetateoraceticacid(N=6)wasmeasuredin the laboratoryat25°C.Lurespreviouslykeptat−15°Cwere re-movedfromtheirpackaging,airedfor24handweighed.Lureswerethenspaced15cmapartonpaper-linedtraysandreweighedweeklyfor4weeks.Datawereexpressedasmglossperday.

A general protocol for all field studies was adopted acrossthe various geographical regions and during different years.Treatmentswererandomizedineachorchardwithfivereplicates.Trapswereplacedinthemid-canopy,≥30mapartand>20mfromthebordersoforchards.Mothswereremoved,sexedandcountedweeklyorbiweekly.CountdatawerecollectedforbothG. molesta and C. pomonella. Traps were rotated among positions on eachdatetheywerechecked.Studieslastedfrom1weekto3months.Linerswerereplacedasneededandatleastevery4weeksinthelongerstudies.Capturesofnontargetsonlinerswererecordedinafewstudies.Lurereplacementschedulesvaried:noreplacementinshorterexperiments,betaocimeneseptareplacedevery2weeksandotherluresreplacedatleastevery4weeks.Lureswereplacedatthecentreoftheliner.Afewexceptionstothisgeneralprotocoloccurredindifferenttrialsandwillbenoted.

2.2 | Oregon 2015

This studywas repeatedover twoperiods inamixed-cultivarpeachorchardsituatednearMedford,OR(42°14′47.37″N,119°52′21.88″W).

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Treeswere>30yearsoldandplantedatadensityof600/ha.Theor-chardwasnottreatedwithsexpheromoneduring2015.Mothcatchesin10luretreatmentswerecomparedinthisstudy,includingthecombi-nationsexpheromonelure(2-PH)aloneandinbinaryandternarylurecombinations.Thebinary lure treatments included2-PHplusoneofthefollowinglures:thePEorBOsepta,ortheZ3,AAorTAmembranelures.Ternarylurecombinationsincludedthetwoseptalures2-PHandBO,andtheadditionofeitherthePEseptumoroneofthethreemem-branelures:Z3,AAorTA.Thestudywasconductedtwicefrom25Julyto8Augustandrepeatedfrom20Augustto3September.

2.3 | Uruguay 2015–2016

Astudywasconductedinpeach(34°44′8″S,56°17′11″W)situatednearMoizo,Uruguay. The orchardwas planted primarilywith thecultivar Pavia Canario, trees <20years old and planted at 1,000/ha.TheorchardwastreatedwithIsomateOFMdispensers(PacificBiocontrol) at 250/ha and was adjacent to commercial apple or-chards. This study compared five lures, includingOFM-PH, 2-PH,2-PH+BO, 2-PH+TA+AA and 2-PH+TA+AA+BO. The studywasconductedfrom15Decemberandterminatedon18January.

2.4 | Oregon 2016

ThesameMedfordorchardused in2015was treatedwithCidetrakOFM-Ldispensers(TrécéInc.)appliedat250/hainlateMay.Dispenserswereloadedwith250mga.i.ofthethree-componentblend.Thisstudywasrepeatedonthreesetsofdates,including12–26June,14–29Julyand29Julyto8August.Sixluretreatmentswerecomparedinthesestudies,2-PHaloneandinbinaryandternarylurecombinations.Thebinaryluretreatmentsincluded2-PHplustheBOseptum,2-PHplusamembranelureloadedwitheitherAAorTA,orthetwomembraneluresTA andAA.Two ternary lure combinations included the septawith2-PHandBOplusamembranelurewitheitherTAorTA/AA.

2.5 | South America 2015–2016

StudieswereconductedinArgentina,ChileandUruguaycomparingmothcatchintrapswitheightdifferentluretreatments.TreatmentsincludedtheindividualseptaluresOFM-PHand2-PH,andthemem-brane luresAA,andTA;binary lures including2-PHpluseitherAAorTAandAAplusTA;anda ternary lurecombining2-PH,AAandTA.TwosetsofstudieswereconductedinArgentina,includinginamixed-cultivar(FlamekistandArcticSnowandO-Henry)nectarine/peachblock(39°5′14.64″S,67°38′27.31″W)andinamixed-cultivar(Williams, Red Bartlett, and Abate Fetel) pear block (39°6′2.98″S;67°35′32.80″W). Both orchards were treated with Isomate M100(PacificBiocontrol,Vancouver,WA)at400dispensersperhaappliedinmid-October.Thesedispenserswereloadedwith232mga.i.ofa93:6:1blendof(E)-8-dodecenylacetate,(Z)-8-dodecenylacetateand(Z)-8-dodecenol.ThepearblockwasalsotreatedwithIsomateCPlus(PacificBiocontrol)at1,000dispensersperha.Thesedispenserswereloadedwith182mga.i.ofa60:34:6blendofcodlemone,1-dodecanol

and1-tetradecanol.Bothstudieswereinitiatedon7Februaryandter-minatedon23March.Trapswerecheckedandrotatedevery3–7daysduringthetrial.Inadditiontotheeightluretreatmentsestablishedintheothertwocountries,trapsbaitedwiththePheroconCML2greysepta(TrécéInc.)tomonitorC. pomonellawereincludedinthepearstudy.Lureswerepinnedtotheinsideroofofthetraps.

ThesamestudyconductedinArgentinawasrepeatedinapeachorchard(cv.Carson)inChile(35°33′29″S,71°33′44″W).Thepeachorchardwas treatedwithCidetrakOFM-L (Trécé Inc.) at 450dis-pensersperhaon15October2015.Thesedispenserswereloadedwith250mga.i. of a93:6:1blendof (E)-8-dodecenyl acetate, (Z)-8-dodecenyl acetate and (Z)-8-dodecenol. Mean tree height was4.5m,andtreeswereplantedatadensityof1,111/ha.Trapswereplacedinthefieldon20February2016andcheckedweeklyuntil29March2016.Thematingstatusoffemaleswasdeterminedfollowingdissectionofthebursa copulatrix.

ThestudyinChileandArgentinawasrepeatedinthesamepeachorchardtreatedwithsexpheromonedispensersnearMoizoprevi-ously described. Like the pear orchard inArgentina, an additionalluretreatmentwasincluded,thePheroconCML2lure(TrécéInc.)forC. pomonella.Thisstudywasinitiatedon1Februaryandtermi-natedon7March2016.

2.6 | Italy 2016

Three sets of trials were performed in peach orchards in Emilia-Romagna region of northern Italy (44°17′32.06″N–12°5′43.26″Eto44°28′45.56″N–12°5′5.40″E)withrecenthistoriesoffruitinjuryfromG. molestaundersexpheromonematingdisruption.The firsttrial includedfivetreatments:thesingle luresOFM-PH,2-PH,and2-PH/PE; and thebinary lures2-PH+BOand2-PH/PE+BO.Thesecondtrialincludedsixtreatments:thesingleluresOFM-PH,and2-PH,thebinarylures2-PH+TA/AAand2-PH/PE+TA/AAandtheternarylures2-PH+TA/AA+BOand2-PH/PE+TA/AA+BO.Thethirdtrial includedsixtreatments:thesingleluresOFM-PHand2-PH, the binary lures 2-PH+TA/AA/Z3 and 2-PH/PE+TA/AA/Z3and the ternary lures 2-PH+TA/AA/Z3+BO and 2-PH/PE+TA/AA/Z3+BO.Thethreetrialswereconductedconcurrentlyfrom6Julyto1Augustandthenrepeatedindifferentorchardsfrom1to29August.

Peach orchards in Italy were <10years old and planted at1,100–1,700treesperha.Allorchardsweretreatedwithstandardagronomic practices including the applications of insecticides formanagement ofG. molesta. Sex pheromones were placed in eachorchard (5–10ha) formating disruption ofG. molesta beginning inlateMarch.Theseincludedboththehandapplicationofdispensers(Isomate-MRosso, PacificBiocontrol) loadedwith 250.2g a.i. perdispenserandappliedat600dispensersperha,theuseofaerosoldevices(CheckMatePufferOFM-O,Suterra,Bend,OR)loadedwith48ga.i.perunitwithspraysappliedat4.8mga.i.every15minfor12hours/dayat3units/ha,andsprayapplications(CheckMateOFMF,Suterra)of22.2ga.i./haevery3–4weeksat45–50mlin1,000Lwaterperha.

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2.7 | North Carolina 2016

ThesamethreetrialsconductedinItalywererepeatedinamixedvarietyblockofappletreesat theMountainHorticulturalCropsResearchStation(MillsRiver,NC).Treespacingvariedfrom2.4to4.2mwithinrows,rowswereplanted6mapart,andtreeheightrangedfromapproximately2.0to3.5m.Eachtreatmentwasrep-licated six times. A replicate of lure treatmentswas placed in arowof treesateye level (about1.7m).Theorchardwas treateduniformly with a standard fungicide and insecticide programmeappliedatapproximatelytwo-wkintervalsduringtheseason.Thethreeexperimentswere runconsecutively from1 to19 July,29Julyto26Augustand26Augustto3October,respectively.Trapswerecheckedweeklyduringtheexperiments,andmothswerenotsexed.

2.8 | Uruguay 2016–2017

The same three trials conducted in Italy and North Carolinawere also repeated in Uruguay. Trial 1 was conducted from 25January to 10 February in a “Fuji” apple block (34°39′14.22″S,56°39′14.22″W).TheorchardwastreatedwithOFM/CMpuffers(Suterra)at2.5units/ha.Theseunitsreleased6.6mga.i.ofa3:4ratioofcodlemonetothethree-componentG. molestapheromoneperpuffevery15minfor12hours/day.Thesecondtrialwascon-ductedfrom26Januaryto16Februaryinpeaches(34°44′2.36″S,56°17′5.80″W)treatedwithCheckmateOFM(Suterra)at250dis-pensersperha.Thethirdstudywasconductedfrom26Januaryto16February inpeaches (34°42′49.85″S,56°05′19.85″W) treatedwith the paste formulation, SPLAT OFM (ISCA Technologies,Riverside,CA)at3.7kg/ha.

2.9 | Pennsylvania 2016

Alonger-termstudywasconductedfrom15Julyto11OctoberinappleblocksatthePennStateUniversityFruitResearchExtensionCenternearBiglerville,PA (39°55′49″N–77°14′49″W), toevaluateluresthat includedtheadditionofpearesterand/or (Z)-3-hexenylacetate.Studieswereconductedinorchardsbothuntreated(N=5)andtreated(N=5)withsexpheromones.Sixluretreatmentswerecompared: the individual lures2-PHand2-PH/PE; and thebinarylures 2-PH+TA/AA, 2-PH/PE+TA/AA, 2-PH+TA/AA/Z3 and2-PH/PE+TA/AA/Z3.Allappleblockswereplantedasmixedcul-tivars (Delicious, Golden Delicious, Gala, Fuji and Yorking) andwereunsprayedforG. molestaduringtheseason.Fourblockswere>15yearsold andplantedat400–550 treesperha.Theother sixblockswere<10yearsoldandplantedattreedensitiesfrom1,500to2,200/ha.OneblockwastreatedwithIsomateCM/OFMTTdis-pensers(PacificBiocontrol)at500/ha.Thesedispenserswereloadedwith268mgofcodlemoneand105mgofthethree-componentsexpheromoneofG. molesta.FourblocksweretreatedwithCheckMatePufferCM/OFM (Suterra) at5units/ha.Lureswereplacedon theliner,and linerswere replacedwhencumulativecatchesexceeded

ca. 50 moths. All lures were replaced once after 4weeks on 12August.Mothswerenotsexed.

2.10 | California and Washington 2017

MothcatchwascomparedinAjartraps(describedinCichonetal.,2013) and delta traps baited with the TA/AA membrane lure. Asingle peach orchard situated near Clarksburg, CA (38°25′14″N,121°31′38″W),was treatedwith IsomateCM/OFMTTdispensersat500/haon17April.Fivereplicatesofeachlure/trapcombinationwererandomizedintheorchardwithtrapsspaced30mapart.Trapswerefirstcheckedon24Aprilandweeklyforthreeadditionalweeks(15May).Mothswerenotsexedinthisstudy.

Two studies were conducted in Washington to evaluate themembraneTA/AA lure.The first studywas run from18May to1JuneandwassimilartothepreviousstudyinCaliforniacomparingmothcatchesinanAjartrapbaitedwithTASandindeltatrapsbaitedwiththeTA/AAmembranelure(N=10);butinaMoxee(46°30′5″N,120°10′16″W) apple orchard not treated with mating disruptiondispensers,andmothsweresexed.Thisorchard is situatednearadairyandthenumberofmuscidflieswasrecordedinbothtraps.Thesecondstudyinthisorchardcomparedmothcatchesintraps(N=5)with the TA/AA lure against traps with the TA lure alone and incombinationwiththreedifferentaceticacidlures:AA,AA1orAA2.Thesestudieswereestablishedintwoblocks(“Fuji”and“Delicious”)on30Juneand3Julyandbothendedon10July.

2.11 | Statistical analyses

Catchdataforeachofthe5–15replicatesofeachtreatmentweresummarized across each test’s time period and analyzed withStatistix9(AnalyticalSoftwareInc.,Tallahassee,FL).Asquare-roottransformationwasusedtonormalizecountdatapriortoanalysisofvariance.ThenormalityofdatawasinspectedwithaShapiro–Wilkstest.Ifthetransformeddatacouldnotbenormalized,theywereana-lyzedwithanonparametricKruskal–Wallistest(W)followedbyanANOVAofranksifthetestwassignificant,p < 0.05.Normalizeddatawere analyzed with either a completely randomized or completeblockANOVA.Ap-valueof0.05wasusedtoestablishsignificance,Tukey’stest.

3  | RESULTS

3.1 | Oregon 2015

Significant differences in total G. molesta moth catches occurredamong the 10 lure treatments compared in this untreated peachorchard (Table1). The two lure treatmentswith thehighestmeancatcheswere2-PH+BOand2-PH+BO+AA.However,trapswithneither lure treatment caught significantlymore totalmoths thantrapswithjustthe2-PHlure(1.5-foldincrease).Theadditionofasep-tumloadedwithpearestertotrapsbaitedwith2-PHor2-PH+BOsignificantly reduced total catch. The addition ofmembrane lures

736  |     MUJICA et Al.

witheitherZ3,TAorAAtotrapsbaitedwith2-PHdidnotincreasemothcatch.TheadditionofamembranelurewithZ3totrapsbaitedwith2-PH+BOreducedmothcatch.Therewasnosignificantdif-ferenceinmothcatchintrapsbaitedwitheither2-PHor2-PH+BOwheneitherTAorAAwasadded.LownumbersoffemaleG. molesta werecaughtinthisstudyandonlyinthefourluretreatmentsthatincludedeitherTAorAA(Table1).Femalemothcatchintrapswasnotsignificantlydifferentamongthesefourluretreatments.

3.2 | Uruguay 2015–2016

SignificantdifferencesinthetotalnumbersofG. molestawerefoundamong the five treatments, F4,20=53.96, p < 0.0001 (Figure1).AddingtheTA+AAlurestotrapswith2-PHincreasedtotalcatches41-fold. Adding TA+AA lures to traps baitedwith 2-PH+BO in-creasedmoth catch nearly 20-fold. Trapswith eitherOFM-PH or2-PHdidnotcatchanyfemaleG. molesta,andthetwotreatmentsincludingTA+AAlurescaughtsignificantlymorefemalemothsthantrapswith2-PH+BOlures,F2,12=33.16,p < 0.0001.Thepercent-ageoffemaleG. molestarangedfrom48%to64%amongthesethreeluretreatments.NoC. pomonellawerecaughtintrapsbaitedwiththeOFM-PHlure.MeantotalmothcatchesofC. pomonellarangedfrom3.6(2-PH+TA+AA+BO)to14.2(2-PH)moths,andtheproportionoffemalemothcatchwas>40%intrapswithallfourattractivelures.Cumulativecatchofnontargets(primarilydipterans)waslowintrapsacrossall lure treatments,means=11–17, including inblank traps,mean=15.2, and nontarget catch was not significantly different,F5,24=0.65,p = 0.66.

3.3 | Oregon 2016

Mean totalmoth catches varied 10-fold across the six lure treat-ments compared in this study (Table2). Moth catch was signifi-cantly greater in traps baited with 2-PH+TA/AA lures with or

withouttheadditionoftheBOseptumcomparedwith2-PHaloneand2-PH+BO.Thetwoluretreatmentscombining2-PHwiththeTAlureandwithorwithouttheBOseptumwereintermediate.NofemaleG. molestawerecaught intrapsbaitedonlywith2-PH,andcatcheswereonlyincidentalwhentheBOseptumwasadded.Thetwo lure treatments including 2-PH+TA/AA caught significantlymorefemalesthanthetwoluretreatmentswith2-PH+TA.Thead-ditionofBOdidnotaffectfemalemothcatchwhenusedwitheither2-PH+TAor2-PH+TA/AAlures.

3.4 | South America 2015–2016

Significant resultsoccurred in the total catchofG. molesta amongeight lure treatments tested concurrently in Uruguay, Chile andArgentina (Table3). The highest moth catch occurred in all threecountrieswithtrapsbaitedwith2-PH+TA+AAlures.However,thesignificantmeanseparationofluretreatmentsvariedamongcoun-tries.Forexample, theAA+TAcombinationperformedaswellas2-PH+TA+AA in Chile and Uruguay, but not in Argentina. Also,2-PH+TA+AA outperformed 2-PH+TA only in Chile, althoughmeancatchwasca. threefoldhigher inUruguay.Adding the2-PHlurewithTA+AAtotrapscomparedwithTA+AAonlysignificantlyincreasedmothcatch inArgentina.Totalmothcatchwas low (de-finedhere as<90%of catch in trapsbaitedwith2-PH+TA+AA)intrapsbaitedwiththeOFM-PHlureinUruguayandChileandlowwith 2-PH in Chile (Table3). Across the four studies, total mothcatchwas16-foldhigherintrapsbaitedwith2-PH+TA+AAthanintrapswithOFM-PHandnearlyfivefoldhigherthanintrapsbaitedwithonly2-PH. Incidental catches (≤0.6mothsper trap)occurredwith several lures, includingAA inall threecountries,OFM-PH inUruguayandChile,TAinUruguayand2-PHand2-PH+AAinChile.

Differences in the catch of female G. molesta only occurred among lure treatments in Chile (Table3). Here, traps baited with2-PH+TA+AA caught significantly greater numbers than traps

Lures Septa Membranes

Mean (SE) moth catcha

Total Females

2-PH – – – – – 9.1(1.5)ab 0.0(0.0)

2-PH BO – – – – 13.9(3.5)a 0.0(0.0)

2-PH – PE – – – 4.4(1.2)c 0.0(0.0)

2-PH – – Z3 – – 6.4(1.0)abc 0.0(0.0)

2-PH – – – AA – 9.2(3.1)abc 0.5(0.3)

2-PH – – – – TA 9.0(2.4)abc 1.2(0.3)

2-PH BO PE – – – 5.4(1.8)c 0.0(0.0)

2-PH BO – Z3 – – 5.5(1.5)bc 0.0(0.0)

2-PH BO – – AA – 14.0(4.0)a 0.6(0.2)

2-PH BO – – – TA 10.2(4.5)abc 1.8(0.8)

Stats F9,89=2.36,p < 0.05

W=4.90,p = 0.18

Note.Columnmeansfollowedbyadifferentletterweresignificantlydifferent,p < 0.05;Tukey’stest,p < 0.05.aTreatmentswithnofemalemothcatchwereexcludedfromtheanalysis.

TABLE  1 ComparisonoftotalandfemaleGrapholita molestaadultscaughtintrapsbaitedwiththesexpheromoneblendforG. molesta and Cydia pomonella (2-PH)eitheraloneorinbinaryandternarylurecombinationswithβ-ocimene(BO)orpearester(PE)loadedinseptaor(Z)-3-hexenyl-acetate(Z3),aceticacid(AA)orterpinylacetate(TA)loadedinmembranecupluresintwostudies(N=10)conductedinAugustandSeptemberinanuntreatedpeachorchard,Oregon,USA,2015

     |  737MUJICA et Al.

withTAor2-PH+TA.TrapswithTA+AAcaughtan intermediatenumberoffemales.Ingeneral,femaleswereonlycaughtatnumbers>1mothper trap in lure treatments includingTA.ThepercentageoffemalemothscaughtwithluretreatmentsincludingTAwashighin bothUruguay (34%–55%) andChile (64%–71%), and somewhatlower in Argentina (17%–26%). In Chile, the proportion of femalemoths (N=332) thatweremated in traps baitedwith terpinyl ac-etatewas37.7%;andinArgentina,matingwas17.4%inpearsand23.6%inpeaches.

In Argentina, low numbers of C. pomonella were caught (≤1.2mothpertrap)acrossalltreatmentsinbothorchards,andnomothswerecaughtintrapsbaitedwithOFM-PH,AAorintheblank.Mean(SE)catchofC. pomonellaintrapsbaitedwithcodlemoneinthepearorchardwas1.2 (0.4).Mean (SE) catchofC. pomonellawashigherinUruguay,andnomothswerecaught intrapswithOFM-PH,TA,AAorTA+AA.Meanmothcountsweresimilarintrapsbaitedwithcodlemone(10.0[2.2]),or2-PHalone(10.2[2.7])or2-Phincombi-nationwith eitherAA (13.2 [6.1]) or TA (9.2 [4.0]).However, totalcatchwas significantly lower in trapsbaitedwith2-PH+TA+AA,(7.0[4.8]),F4,20=3.54,p < 0.05.

ThecumulativemeancatchesofnontargetsintrapsinArgentinaover the6-weekstudywas19–39among lure treatments,andnosignificant differencewas found,F8,80=0.70,p = 0.69.Nontargetsincluded low numbers of bees, lacewings, noctuids and coccinel-lids;and96%of thetotalcatchweredipterans,primarilymuscids.Cumulative catch of nontarget dipterans in the peach orchard inUruguaywerelowandconsistentacrossalltreatmentswithmeansranging from 11 to 20 flies across lure treatments, F9,40=0.92,p = 0.51.Itisinterestingthatthemostabundantnontargetcaughtintrapswerethysanopterans,withcumulativemeancatchesrangingfrom38to102thripsacrossluretreatments;however,thesemeanswerenotsignificantlydifferent,F9,40=1.06,p = 0.41.

3.5 | Uruguay 2016–2017

NosignificantdifferenceswerefoundfortotalmothcatchinTrial1(Table4).Femalemothswereeithernotcaughtoronlyincidentallyin theOFM-PH lure treatment.Totalmothcountswere fairly lowinTrial2,especiallyintrapsbaitedwithOFM-PHand2-PH.Meantotalcatchwas>eightfoldhigherintrapswiththeTA/AAlureadded

F IGURE  1 ComparisonoffemaleandtotalmothcatchesofGrapholita molesta(a)andCydia pomonella(b)intrapsbaitedwithindividualandcombinationsofthesexpheromone(OFM-PH),thecombinationsexpheromone(2-PH),β-ocimene(BO),terpinylacetate(TA)andaceticacid(AA)lures,Uruguay2015–2016.NofemaleG. molestawerecaughtintrapsbaitedwiththetwopheromone-onlylures,andthesedatawerenotincludedintheanalysis,denoted“−”.Treatmenthistogramswithdifferentlowercaseanduppercaselettersweresignificantlydifferentinfemaleandtotalmothcatches,p < 0.05.CatchesofC. pomonelladidnotdifferamongluretreatments,p > 0.05

(a)

(b)

738  |     MUJICA et Al.

comparedwiththeuseof2-PHonly,andnosignificantdifferencewasfoundfortotalorfemalecatchamongthefourluretreatmentsincluding TA/AA. The proportion of females caught in these fourluretreatmentswashigh,0.62–0.86. Incomparison,mothcatcheswereveryhighinallluretreatmentsinTrial3andfewfemaleswerecaught. These data were collected from a peach orchard severalmonthsaftertheSPLATpastewasappliedandsuggestthatthefor-mulationwasprovidingscantdisruptionofthesetraps.

3.6 | Italy 2016

Catch of G. molesta in these commercial orchards was low withthefive luretreatmentstested inTrial1,andmeansdidnotdiffer(Table4).However,significantdifferenceswerefoundintotalmothcatchamongluretreatmentsinTrial2.Thefourtreatmentsinclud-ing2-PHor2-PH/PEplusTA/AAhadthehighestmothcatchesanddidnotdiffer.Allofthesetreatmentscaughtsignificantlymoretotalmoths than trapswitheitherOFM-PHor2-PH lures.Thehighesttotalcatchwas intrapswith2-PH/PE+TA/AA+BO,andthiswas30-foldhigherthantrapswithOFM-PH.Femalemothcatchesoc-curred in all traps baited with TA/AA but did not differ amongthesefourluretreatments.InTrial3,significantdifferencesinbothtotalandfemalemothscatcheswerefoundamongluretreatments(Table4). All of the lure treatments with TA/AA/Z3 except with2-PH+TA/AA/Z3+BOcaughtsignificantlymoretotalmothsthantraps with 2-PH. Traps with 2-PH/PE+TA/AA/Z3 caught nearlyfivefold more moths than traps with OFM-PH. No female mothswere caught in trapswith eitherOFM-PH or 2-PH. Femalemothcatch was significantly higher in traps with 2-PH/PE+TA/AA/Z3thanwith2-PH+TA/AA/Z3,andtheothertwoluretreatmentshadintermediatefemalecatches.

3.7 | North Carolina 2016

Significantdifferences in totalmothcatchoccurred inTrial1withtraps baited with 2-PH/PE+BO catching more than traps withOFM-PH.Theotherthreeluretreatmentscaughtintermediatenum-bers ofmoths. Similarly, in Trial 2, significant differences inmothcatchoccurredamongthefiveluretreatments.Mothcatchintrapsbaitedwith either 2-PH/PE+TA/AAwith orwithout the addition

TABLE  2 ComparisonoftotalandfemaleGrapholita molesta adultscaughtintrapsbaitedwithaseptumloadedwithacombinationsexpheromoneblendforG. molesta and Cydia pomonella(2-PH)eitheraloneorincombinationwithaβ-ocimene(BO)septumormembranecupluresloadedwithterpinylacetate(TA)orTAplusaceticacid(AA)inthreeserialstudies(N=15)conductedinapeachorchardtreatedwithsexpheromonedispensers,Oregon,USA,2016

Lures Septa Membranes

Mean (SE) moth catch

Total Females

2-PH – – 0.9(0.3)c 0.0(0.0)

2-PH BO – 4.1(1.1)bc 0.2(0.1)

2-PH – TA 4.5(1.4)abc 0.7(0.3)b

2-PH BO TA 5.3(1.3)ab 0.7(0.3)b

2-PH – TA/AA 9.7(1.5)a 3.3(1.2)a

2-PH BO TA/AA 9.2(1.7)a 3.3(0.8)a

Stats F5,82=6.97,p < 0.0001

W = 15.91,p < 0.001 F3,56=6.89,p < 0.001

Note.Columnmeansfollowedbyadifferentletterweresignificantlydif-ferent,p < 0.05.Totalmothcatchwasanalysedwitharandomizedcom-pleteblockANOVA,andfemalecatchwasanalysedwithaKruskal–WallistestfollowedbyanANOVAofranks,p < 0.05.

TABLE  3 ComparisonoftotalGrapholita molestaadultscaughtinunbaitedtraps(Blank)andtrapsbaitedwithaceticacid(AA),terpinylacetate(TA),sexpheromone(OFM-PH),acombinationsexpheromoneblendforG. molesta and Cydia pomonella(2-PH),andthiscombinationblendpluseitherAA,TAorAA+TAinrepeatedstudiesinUruguay,ChileandArgenina,2015–2016

Lures Septa Membranes

Mean (SE) moth catchesa

Uruguay Chile Argentina

Total Females Total Females Total Femalesb

OFM-PH – – 0.2(0.2) 0.0(0.0) 0.0(0.0) 0.0(0.0) 2.1(0.8)c 0.1(0.1)

2-PH – – 3.2(2.0)b 0.0(0.0) 0.2(0.2) 0.0(0.0) 6.2(1.2)abc 0.7(0.3)

– AA – 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.0(0.0) 0.2(0.2) 0.2(0.2)

– – TA 0.4(0.2) 0.0(0.0) 10.4(3.8)bc 6.6(2.8)b 2.4(0.8)bc 0.4(0.2)

– AA TA 4.6(1.9)ab 2.4(1.2) 21.0(5.5)ab 14.4(3.8)ab 3.5(1.2)bc 0.9(0.5)

2-PH AA – 2.0(0.7)b 0.0(0.0) 0.6(0.4) 0.4(0.4) 5.2(1.5)abc 0.3(0.2)

2-PH – TA 4.4(1.2)ab 2.4(1.1) 7.6(2.1)c 5.0(1.1)b 10.5(3.1)ab 2.0(1.1)

2-PH AA TA 12.8(3.3)a 4.4(1.0) 27.2(6.8)a 19.4(4.6)a 15.2(4.3)a 3.6(1.8)

Stats F4,20=4.57 p < 0.01

F2,12 = 1.28 p = 0.31

F3,16 = 3.91 p < 0.05

F3,16=4.15 p < 0.05

F6,62=5.58 p < 0.0001

W4=5.60 p = 0.23

Note.Columnmeansfollowedbyadifferentletterweresignificantlydifferent,Tukey’stest,p < 0.05.aTreatmentswithnomothcatchoronlyaninci-dentalcatch(means≤0.6mothpertrap)wereexcludedfromtheANOVAs.Blanktrapsplacedinthestudydidnotcatchanymoths.bFollowingthesignificantKruskal–Wallisnonparametrictest,anANOVAoftherankswasconducted,F43,76=4.00,p < 0.05,andthiswasfollowedbyameanssepara-tiontest,p < 0.05.

     |  739MUJICA et Al.

ofBOcaughtsignificantlymoremothsthantrapsbaitedwith2-PH.Moth catch in traps with the 2-PH+TA/AA with or without BOcaughtintermediatenumberofmoths.Mothcountswerefairlycon-sistentacrossthefiveluretreatmentstestedinTrial3.LownumbersofC. pomonellawerecaught in theseappleblocksduringall threetrials.ThemostC. pomonellawerecaughtduringTrial1andthefivetrapsbaitedwith2-PH/PEcaught fourcomparedwith13 in trapsbaitedwith2-PH.

3.8 | Pennsylvania 2016

AlargenumberofG. molestawerecaughtintrapsinthereplicatedappleorchardsnottreatedwithsexpheromones(Table5).Significantdifferences occurred among lure treatments in these blocks andtrapsbaitedwith2-PH/PE+TA/AAcaughtsignificantlymoremothsthan theother five lure treatments.Also, trapsbaitedwith2-PH/PE+TA/AA/Z3 caught significantly more moths than traps with

2-PH.Mothcatchinblocksundersexpheromoneswere98%lowerthanintheuntreatedblocks,andnosignificantdifferencewasfoundamongluretreatments.C. pomonellawascaughtatfairlyhighlevelsinallluretreatmentsinthenondisruptedappleblocks.Thehighestcatchwasintrapsbaitedwith2-PH.CatchesweresignificantlylowerwhenpearesterwasaddedtotheseptumlureandwheneitherTA/AAorTA/AA/Z3wasaddedtothetrap.TrapsincludingtheTA/AA/Z3lurehadthelowestcatches.ThecatchofC. pomonellainblockstreatedwithsexpheromonewasmuchlowerthanintheuntreatedblocks.Onlytrapsbaitedwith2-PHor2-PH/PEcaught>1.0moth.

3.9 | California and Washington 2017

Significant differences were found in moth catch between theAjaranddeltatrapbaitedwiththeTRE1370 lure inCaliforniaandWashington.InCalifornia,themean(SE)mothcatchoverthe4weektrialinthedisruptedpeachorchardaveraged7.0(2.8)and29.8(10.9)

TABLE  4 ComparisonoflureswithvariousblendsinaseriesofthreetrialsconductedinUruguay,ItalyandNorthCarolinatestingseptaluresloadedwiththesexpheromoneofGrapholita molesta(OFM-PH),thecombinationsexpheromoneofG. molesta and Cydia pomonella (2-PH),thecombinationsexpheromonepluspearester(2-PH/PE)and/orβ-ocimene(BO),andamembranecupluresloadedwithterpinylacetate(TA)andaceticacid(AA)together(TA/AA)andwith(Z)-3-hexenylacetate(TA/AA/Z3),2016

Trial No.Lures Septa Membranes

Mean (SE) moth catchesa

Uruguay Italy NC

Total Females Total Females Total

1 OFM-PH – – 6.6(1.6) 0.2(0.2) 1.4(0.6) 0.0(0.0) 1.7(1.3)b

2-PH – – 17.6(4.7) 0.0(0.0) 1.7(0.9) 0.4(0.3) 4.5(0.9)ab

2-PH/PE – – 18.0(5.9) 0.0(0.0) 4.6(1.6) 0.2(0.1) 3.3(0.8)ab

2-PH BO – 13.8(3.3) 0.0(0.0) 2.6(0.9) 0.3(0.2) 5.0(1.6)ab

2-PH/PE BO – 18.0(5.7) 0.0(0.0) 4.3(1.2) 0.6(0.3) 6.8(0.9)a

Stats F4,20 = 1.37 p = 0.28

– F4,44=2.53 p = 0.05

– F4,20 = 4.21 p < 0.05

2 OFMPH – – 0.2(0.2) 0.0(0.0) 0.9(0.4)b 0.0(0.0) –

2-PH – – 0.2(0.2) 0.0(0.0) 4.8(2.0)b 0.0(0.0) 7.7(1.9)b

2-PH – TA/AA 2.6(0.7)ab 1.6(0.9) 13.5(4.7)a 3.7(0.8) 12.2(3.8)ab

2-PH/PE – TA/AA 2.4(0.9)ab 1.6(0.6) 17.2(2.3)a 4.8(1.2) 14.2(3.0)a

2-PH BO TA/AA 2.8(1.2)a 2.4(1.3) 14.9(4.0)a 6.0(1.4) 11.3(4.2)ab

2-PH/PE BO TA/AA 1.6(0.4)ab 1.2(0.4) 27.4(7.1)a 9.4(2.1) 15.7(4.6)a

Stats F3,16 = 0.31 p = 0.82

F3,16=0.05 p = 0.99

F5,53 = 14.37 p < 0.0001

F3,35=2.56 p = 0.07

F4,20 = 4.02 p < 0.05

3 OFM-PH – – 20.0(7.8) 0.0(0.0) 4.0(1.4)c 0.0(0.0) –

2-PH – – 45.6(7.5) 0.0(0.0) 6.7(2.5)c 0.0(0.0) 9.7(3.6)

2-PH – TA/AA/Z3 33.6(5.1) 1.0(0.4) 13.5(5.1)ab 1.3(0.3)b 9.3(3.5)

2-PH/PE – TA/AA/Z3 31.6(5.4) 0.4(0.2) 19.5(3.9)a 3.7(0.9)a 7.8(4.4)

2-PH BO TA/AA/Z3 35.0(4.4) 0.2(0.2) 6.3(1.4)bc 1.9(0.5)ab 8.8(5.2)

2-PH/PE BO TA/AA/Z3 33.6(4.6) 0.6(0.4) 15.5(3.7)a 3.1(0.5)ab 9.7(4.4)

Stats F5,24 = 2.21 p = 0.09

– F5,53 = 14.07 p < 0.0001

F3,35=3.85 p < 0.05

F4,20 = 0.88 p = 0.49

Note.Columnmeansfollowedbyadifferentletterweresignificantlydifferent,p < 0.05.aDatawereexcludedfromtheanalysesifthemeanwas≤0.6mothspertrap.

740  |     MUJICA et Al.

intheAjaranddeltatrapbaitedwiththeTRE1370lure,respectively;andthisdifferencewassignificant.F1,8=30.94,p < 0.001. In com-parison,twotrapsbaitedwithsexpheromoneluresandplacedinthesameorchardbutadjacent(distancesof30–50m)toourstudybothcaughttwomothsduringthistimeperiod.

InWashington,significantdifferencesinmothcatchoccurredinthenondisrupted appleorchardbetween theAjar anddelta trapsbaited with the TRE1370 lure, F1,18=21.73, p < 0.001. Mean (SE)mothcatchovera2-weekperiodaveraged1.2(0.5)and6.0(1.1)inthetwotraptypes,respectively.Bothtrap-luretypescaught>40%femalemoths. Inaddition,themean(SE)catchofmuscidflieswassignificantlyhigherintheAjartraps(28.0[5.6])thanthedeltatraps(10.2[2.0]),F1,18=11.75,p < 0.01.

Significantdifferencesinfemaleandtotalmothcatchesoccurredamong traps baited with TA, TA/AA and TA plus three differentacetic acid lures (Table6).While the threemembrane luresvariedwidelyintheirreleaseofaceticacid,meanmothcatchesdidnotdif-feramongthesethreelureswhenusedincombinationwiththeTAlure.Also,mothcatcheswerenotdifferentbetweentheTA/AAlureandthebinaryuseofTAplusanyofthethreeaceticacidlures.

4  | DISCUSSION

Grapholita molestaisakeyworldwideinsectpestofstonefruitsandof pome fruits in some geographical areas (Rothschild & Vickers,1991).Theabilitytoeffectivelymonitortheadultstagesofthispestallows farmmanagers todevelopeffectiveprogrammesthatmini-mizetheuseofinsecticides(Charmillot&Vickers,1991).Workre-portedbyourresearchgroupoverthepast5yearshasdetailednewcombinationlures,includingsexpheromonesandhostplantandmi-crobialvolatilesthatcanmonitorbothsexesinorchardstreatedwithsexpheromones formatingdisruption (Cichonetal.,2013;Knightetal., 2013; Knight, Cichon, etal., 2014; Knight, Barros-Parada,etal.,2015).Oneprerequisiteincludedinourdevelopmentofnew,low-costmonitoringtoolshasbeenthemanager’sabilitytouseany

new lures in standard traps with removable sticky liners (Cichonetal.,2013).DevelopmentoftheAjartrapwasanintermediatestepthat allowed bisexual monitoring using sticky liners with catchescomparabletothebuckettrapsbutwithfewernontargetstosort,butwas still a cumbersomedesigndue to the replacementof theliquidbaits(Knightetal.,2013;Knight,Cichon,etal.,2014;Padilhaetal.,2017).OurnewresultsdemonstratethatG. molesta can now bemonitoredwithacombinationofvolatilesreleasedbyseptaandmembranelureswithinstandardstickytraps.Also,thenumbersofnontargetscaughtintrapsbaitedwiththeTA/AAluredidnotappeartobeaproblemacrossthevariousstudies.

Earlier studies demonstrated thatmoth catches ofG. molesta in trapsbaitedwith the combination sexpheromone lure (2-PH),whichincludescodlemone,wereonly intermediate incomparisonwith a number of commercial sex pheromone lure forG. molesta when used in disrupted orchards (Knight, Basoalto, etal., 2015).Here, among our various studies, the mean catch in traps wasconsistently higher (3–5×) in traps baitedwith the 2-PH lure vs.

TABLE  6 Comparisonoftheweightlossofmembraneluresovera2-weekperiodat25°CandcatchesofGrapholita molestaindeltatrapsbaitedwithterpinylacetate(TA)andaceticacid(AA)lures,N=10,Moxee,Washington,USA,2017

Lures

Active: Mean (SE) daily weight loss (mg)

Mean (SE) moth catch

Total Females

TA TA:12.6(0.2) 1.1(0.2)b 0.4(0.3)b

TA/AA TA+AA:13.1(0.3) 4.0(1.4)ab 2.0(0.9)ab

TA+AA AA:5.2(0.05) 6.9(2.8)ab 2.1(1.1)ab

TA+AA2 AA:27.8(0.5) 8.6(3.1)a 3.7(1.1)a

TA+AA3 AA:73.1(1.0) 10.3(3.3)a 4.2(1.4)a

ANOVA F4,44=4.33,p < 0.01

F4,44=3.86,p < 0.01

Note.Meanmoth catch in each column followed by a different letterwere significantly different in a randomized complete block. ANOVA,Tukey’sHSDtest,p < 0.05.

Lures Septa Membranes

Mean (SE) moth catcha

G. molesta C. pomonella

MD No MD MD No MD

2-PH – 1.0(0.6) 169.8(61.6)c 3.8(2.2) 105.8(25.6)a

2-PH/PE – 0.8(0.4) 246.4(89.0)bc 2.0(0.6) 43.6(11.6)b

2-PH TA/AA 4.8(2.9) 267.2(100.7)bc 0.4(0.4) 25.6(6.7)bc

2-PH/PE TA/AA 4.6(1.5) 448.4(151.0)a 1.0(0.3) 36.6(11.3)bc

2-PH TA/AA/Z3 1.6(0.7) 231.0(76.9)bc 0.4(0.4) 12.4(5.8)cd

2-PH/PE TA/AA/Z3 1.8(1.1) 282.8(104.6)b 0.0(0.0) 5.6(3.2)d

Stats W6=5.51 p = 0.35

F5,20=13.54 p < 0.0001

W3=5.33 p = 0.15

F5,20 = 20.74 p < 0.0001

Note.Columnmeansfollowedbyadifferentletterweresignificantlydifferent,p < 0.05.aDatawereexcludedfromtheanalysesifthemeanwas≤0.6mothspertrap.

TABLE  5 ComparisonoflureswithvariousblendstestingseptaluresloadedwiththecombinationsexpheromoneofGrapholita molesta and Cydia pomonella (2-PH),thecombinationsexpheromonepluspearester(2-PH/PE)andmembranecupluresloadedwithterpinylacetateandaceticacidtogether(TA/AA)orincombinationwith(Z)-3-hexenylacetate(TA/AA/Z3),N=5appleorchardstreatedwith(MD)andwithoutsexpheromones(noMD)forbothG. molesta and C. pomonella,Pennsylvania,USA,2016

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astandardsexpheromonelure,butthesemeanswereneversta-tisticallysignificant.Previously,wefoundthatasignificanttrade-offwithusingthecombinationsexpheromonelurewasthatmalecatchofC. pomonellawasreducedincomparisonwithcodlemone-baited traps (Knight,Cichon, etal., 2014;Knight,Basoalto, etal.,2015).Conversely, studies reported fromArgentinaandUruguayfound thatC. pomonella catchwassimilar in trapsbaitedwithei-thercodlemoneorthe2-PHlure.However,significantnegativeef-fectsonthecatchofC. pomonellaoccurredwhenotherlureswereaddedtoboostthecatchofG. molesta,forexampleTA+AAluresinUruguayandTA/AA,TA/AA/Z3orpearester inPennsylvania.Similarly, previouswork had reported that the addition of eitheraceticacidortheTASliquidbaittotrapsbaitedwiththe2-PHlurereducedcatchofC. pomonellaby50%(Knight,Cichon,etal.,2014).However,nopreviousstudyhastheuseofpearesterbeenshowntoreducethecatchofC. pomonella.

Initial field screening of target lepidopteran pest species inNorthAmericafoundthatpearesterwasnotattractiveforG. mo-lesta (Knight&Light,2004).Yet, laterwork inItalysuggestedthatbothsexesofG. molestacandetectpearesterandthatmalesinlab-oratorychoiceolfactometerbioassayswerebehaviourallyattractedtopear ester (Molinari etal., 2010).Here, the importanceof add-ing pear ester to the grey septa loadedwith the combination sexpheromone(2-PH/PE)wassomewhatinconclusive:reducedcatchinOregonin2015,amarginal increase(p = 0.05) inItaly in2016andclearlynoeffectinthe2016studiesinUruguay,NorthCarolinaandPennsylvania. However, combining the 2-PH/PE septum with theTA/AAmembraneluredidsignificantlyincreasecatchofG. molesta inPennsylvania, but thisdidnotoccur in theother trials.Overall,includingpearesterinthebinarylureforG. molestadoesnotseemtobecritical.However,ourfindingthatinclusionofpearesterintheseptalurereducescatchofC. pomonellaisproblematicforanyfuturecommercializationoftheselures,especiallyinorchardswherebothpestsarepresentandmanagerswanttomonitorbothinthesametrap.

A number of laboratory and field studies have evaluated vari-oushostplantvolatilesforG. molestaandachievedvariableresults.Forexample,eitherβ-ocimeneor(Z)-3-hexenylacetatealoneorinvarious HPV blends was found to increasemoth catches (Il’ichevetal.,2009;Knightetal.,2011;Nataleetal.,2003;Piñero&Dorn,2007),orprovidedvariableresults,suchasβ-ocimeneimprovedtheperformanceofTAS-baitedtraps(Knightetal.,2013);butpearesterplusβ-ocimenedidnotimprovethe2-PHlure(Knight,Cichon,etal.,2014).Similartothislaterstudy,wefoundthatneitherβ-ocimenenor(Z)-3-hexenylacetateaffectedmothcatcheswhenusedwitheither2-PHorTA/AA lures, and likelyneither volatilewill be developedfurtherthroughourtrials.

The addition of acetic acid lures to traps did not improve theperformanceof host plant volatile-based lures or 2-PH forG. mo-lestainpreviousstudies(Knight,Cichon,etal.,2014;Knight,Hilton,etal.,2014).However,aceticacidhadbeenusedeffectivelyinaliq-uidbaitwithsugarandethanolinChina(Heetal.,2009).Here,wefound inseveralstudiesthataceticaciddidnot improvethe2-PH

lure.Similarly,addingaTAluretotrapswiththe2-PHluredidnotincreasemothcatches. Incomparison,mothcatcheswith theTASliquidbaitweresignificantlyincreasedwiththeadditionofthe2-PHlure(Knight,Cichon,etal.,2014).Thesedisparateresultscouldlikelybeduetotheundefinedandtemporallysensitivevolatileprofilere-leasedbytheTASbaitfromopencontainerswhencomparedwiththeclosedTAmembranelure.

ThemosteffectivelureforG. molestainourtrialswasclearlythe2-PHseptumandTA/AAmembraneused intrapstogether.Mothcatcheswiththisbinarylurecomparedwiththe2-PHlurealoneincreasedfrom<twofoldto>100-fold,andtheproportionof females caught in traps baited with this binary lure rangedfrom0.24to0.71acrossthevariousstudies.Atpresent,wehavenohypothesistoexplainthishighdegreeofvariationintherel-ativeperformancesoftheselures,andfurtherstudiesmightad-dresslureefficacyindifferenthostcrops(stonefruits,appleandpear)orpossible geographicaldifferences in thesebehaviouralresponsesamongG. molestapopulations(Knight,Barros-Parada,etal.,2015).Clearly,theTA/AAmembranelurehasnotyetbeenoptimized.Forexample,therewasaclearpositivetrendinbothtotal and female moth catch as a function of emission rate ofacetic acid.Also, theweight lossdata suggest that the loadingof terpinyl acetate in the TA/AA luremay be too low and thebinaryemissionprofileofthislureshouldbecharacterizedovertimeaswell.

Onefocusofourrecentworktodevelopbisexualluresforkeytortricidpestsoftreefruitshasbeentotestwhethermultiplecom-ponents that attract different species that co-occur in individualorchards can be formulated and used in individual traps (Knight,El-Sayed,Judd,&Basoalto,2017;Knight,Hilton,etal.,2014).Thiswork is motivated by the rapid advances in smart technologies,such as remotemonitoring of traps through cameras (Kim, Jung,Kim,&Lee,2011;Wen,Guyer,&Li,2009).Todate,ourworkhasfocusedprimarilyonC. pomonella andseveral tortricid leafrollerspresentinthewesternUnitedStates(Basoaltoetal.,2017;Knight,Hilton,etal.,2014;Knightetal.,2017).However,due to the fre-quentco-occurrenceofG. molesta and C. pomonellaplusasuiteofleafrollerspeciesinorchards,itwill likelybeimportanttoexpandthesestudiestoinclude2-PHandTA/AAlures.Fortunately,aceticacidhasbeenfoundtoincreasemothcatchesofallspecieswhenused inspecificways, suchas incombinationwithpearester forC. pomonella (Knight,Hilton, etal., 2014), or in combinationwith2-phenylethanol forChoristoneura rosaceana Harris (Knight etal.,2017).Interestingly,itappearsthattheemissionrateofaceticacidis an important factor affecting moth catches; C. pomonella wasmoreattractivetolowrates(AAlure)andC. rosaceanaismoreat-tracted to higher rates (AA2 andAA3 lures) in these studies. Todate,wedonothaveenoughinformationontheinfluenceofaceticacidrateoncatchofG. molesta,butourpreliminarydatasuggestthatthisspeciesmayhaveabroaderresponsethantheothertwospecies.

Theremaybeseriousdrawbacksindevelopingalurethatcaneffectively catch several species in the same trap. For example,

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theinclusionof (E)-4,8-dimethyl-1,3,7-nonatrienetoenhancethecatch of C. pomonella was found to depress the catch of C. ro-saceanawhenusedwith2-phenylethanolandaceticacid (Knightetal., 2017). Our unexpected finding in the Pennsylvania trialwhere adding pear ester to a septum loadedwith the sex pher-omoneofboth speciesdepressed thecatchofC. pomonella sug-gests that a heuristic approach will not be effective to developmultispecies trapping systems. Unrealized factors such as mothbehavioursneartraps,includingtheroleofvisionorothersensorycuesthatmaycauserepellencyandareusedbyrelatedsympatricspeciestomaintainmatingisolationhavenotbeenaddressedwithkairomones.Manyfactorscanimpacttherelativeeffectivenessofkairomonelures,includinggeography,season,cropandcultivarasshown for pear ester withC. pomonella (Knight & Light, 2005c).Likely, the complexity of combining sex pheromones with hostplant, andmicrobial volatiles for a suiteof specieswouldbe toogreat to create one optimized lure. Instead, perhaps developingsuboptimal lureswithsome limitedattractivenessacrossall spe-ciesmightbe sufficient toestablishaction thresholdsand to re-ducemonitoringcosts.

ACKNOWLEDG EMENTS

Wewould all like to thankBill Lingren,Trécé Inc.,Adair,OK, forproviding the experimental septa and membrane lures used ex-tensively in these trials.Wealsoappreciate the fundingmadebyFondodeDesarrolloCientíficoyTecnológico(FONDECYT)Grant3140285 to WBP and partial funding from Millennium NucleusCentre in Molecular Ecology and Evolutionary Applications inAgroecosystems and Inciativa Cientifica Milenio NC 120027 toEFC.WealsothankSteveSchoofandLindsayStumpofNCStateUniversity,Raleigh,NC, for technical assistance in theNCstudy.In Uruguay, we would like to thank the staff of the experimen-tal stationWilson FerreiraAldunate of the InstitutoNacional deInvestigaciónAgropecuariaincludingGonzaloVázquezandGastónTejerawhoparticipatedinthefieldactivitiesandMarianaSilverawhohelpedwith insect counts in the laboratory. Finally, in Italy,wewouldliketothankDomenicoAbetiofAgrintesaandtheotherIPMtechnicianswhohave indicatedthepeachorchards inwhichto perform the studies.Wewould like to thank RodneyCooper,ARS,Wapato,WA;CesarGemeno,UniversityofLleida,Spain;andValentinoGiacomuzzi,FreeUniversityofBozen-Bolzano,Italy,fortheircarefulreviewsofourroughdraft.

AUTHOR CONTRIBUTION

ALK conceived research, analysed the data, conducted statisticalanalysesandwrotethemanuscript.Allauthorsconductedresearch.Allauthorsreadandapprovedthemanuscript.

ORCID

Alan L. Knight http://orcid.org/0000-0003-4496-9000

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How to cite this article:MujicaV,PretiM,BasoaltoE,etal.Improvedmonitoringoforientalfruitmoth(Lepidoptera:Tortricidae)withterpinylacetateplusaceticacidmembranelures.J Appl Entomol. 2018;142:731–744. https://doi.org/10.1111/jen.12528