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1、942| J Appl Entomol.2019;143:942947.2019 Blackwell Verlag GmbH Received:6 March 2019|Revised:18 July 2019|Accepted:22 July 2019DOI:10.1111/jen.12682 O R I G I N A L CO NTR I B U TI O NMonitoring codling moth(Lepidoptera:Tortricidae)with a fourcomponent volatile blend compared to a sex pheromonebased
2、 blendAlan L.Knight1|Valentina Mujica2|Sebastian Larsson Herrera3|Marco Tasin31USDA,Agricultural Research Service,Wapato,WA,USA2INIA,Montevideo,Uruguay3Department of Plant Protection Biology,IPP,Swedish University of Agricultural Science,Alnarp,SwedenCorrespondenceSebastian Larsson Herrera,IPP,Depar
3、tment of Plant Protection Biology,Swedish University of Agricultural Science,PO Box 102,Sundsvgen 14,SLU,230 53 Alnarp,Sweden.Email:sebastian.larsson.herreraslu.seAbstractMonitoring adult codling moth,Cydia pomonella(L.),is a crucial component in im-plementing effective integrated management program
4、mes in apple,Malus domestica Borkhausen.Use of sex pheromone lures to track male populations has been the tra-ditional approach,but their use in orchards treated with sex pheromone for mating disruption(MD)has been problematic.Development of kairomone and kairomonepheromone combination lures has all
5、owed the catch of female moths and has ben-efited several aspects of codling moth management through improved spray timings and action thresholds.Recently,a new fourcomponent volatile blend(4K)comprised of pear ester,(E,Z)2,4ethyl decadienoate(PE),(E)11 4,8dimethyl1,3,7nonatriene,all isomers of pyra
6、noid linalool oxide and acetic acid(AA)has been characterized that has increased female moth catch threefold versus any previous blend.Field trapping studies were conducted to compare moth catches in traps baited with 4K versus the use of sex pheromone,(E,E)8,10dodecadien1ol(PH)in combination with P
7、E and AA.Trials were conducted in orchards left either untreated,or treated with PH or PH+PE.Traps baited with 4K and 4K+PH lures caught significantly more females than traps baited with PH+PE+AA lures.Traps baited with 4K+PH lures caught significantly more total moths than traps baited with PH+PE+A
8、A lures in all three orchards.Adding a PH lure to traps with the 4K lure did not affect female catch,but significantly increased male and total moth catches.These studies demonstrate that codling moth can be trapped effectively in apple under MD without the use of sex pheromone lures.The significant
9、 increase in female codling moth catch with the 4K lure suggests that efforts to improve spray timings and action threshold determina-tions as well as mass trapping might be enhanced with this new lure.K E Y W O R D S(E,E)8,10dodecadien1ol,(E,Z)2,4ethyl decadienoate(E)4,8dimethyl1,3,7nonatriene,acet
10、ic acid,Cydia pomonella(L.),pyranoid linalool oxideThis article has been contributed to by US Government employees and their work is in the public domain in the USA.|943KNIGHT eT al.1|INTRODUCTIONThe recent report that a fourcomponent blend(4K)consisting of pear ester;(E,Z)2,4ethyl decadienoate(PE);
11、(E)4,8dimethyl1,3,7nonatriene(DMNT);pyranoid linalool oxide(LOX);and acetic acid(AA)is an attractive bisexual lure for codling moth,Cydia pomonella(L.),offers a renewed promise for enhanced monitoring and mass trapping of this key pome fruit pest(Knight,Light,Judd,&Witzgall,2019).Previously,PE alone
12、(Light et al.,2001),PE+AA(Knight,2010;Landolt,Suckling,&Judd,2007),DMNT+AA(Knight,Light,&Trimble,2011),DMNT+AA+PE(Knight&Light,2012)and nbutyl sulphide+PE+AA(Landolt et al.,2014)were found to be effective bisexual lures for codling moth.However,the 4K lure was shown to be threefold more attractive t
13、han any of these blends in com-parative field trapping trials(Knight,Mujica,Herrera,&Tasin,2019).Various multifunctional lures(attractants plus pheromones)including the use of PE,AA and DMNT with the sex pheromone(E,E)8,10dodecadien1ol,(PH)of codling moth have been evalu-ated in orchards treated wit
14、h PH for mating disruption(MD).In gen-eral,the addition of PE has improved the PH lure and allows females to be trapped(Fernndez,Cichon,Garrido,RibesDasi,&Avilla,2010;Hri,Penzes,Josvai,Ladanyi,&Toth,2011;Joshi,Hull,Rajotte,Krawczyk,&Bohnenblust,2011;Knight,Hilton,&Light,2005).The addition of AA to t
15、raps with either PE+PH or DMNT+PH has cre-ated a more attractive blend for codling moth and increased female moth catches fivefold to 10fold(Knight&Light,2012;Knight,Light,&Chebny,2013).Effective monitoring with these blends has also been demonstrated in orchards treated with MD dispensers loaded wi
16、th both PH+PE(Knight,Basoalto,Katalin,&ElSayed,2015;Knight et al.,2013).The most effective combinational lure reported in these orchards has been PE+DMNT+AA+PH.However,be-cause the 4K lure with the addition of LOX caught nearly threefold more total and female moths than the PE+DMNT+AA lure,we believ
17、ed a more effective combinational lure could be developed(Knight,Light,et al.,2019).The standard lure used in Washington and maybe worldwide for monitoring codling moth in orchards treated with MD orchards is likely the PH+PE lure(Ioriatti&Lucchi,2016;Knight,Light,et al.,2019;Witzgall,Stelinski,Gut,
18、&Thomson,2008).Prior to the study reported here,the 4K lure has not been tested against the standard(PH+PE)lure or used in combination with PH or in or-chards treated with MD.While several host plant volatiles have been shown to synergize male tortricids attraction to sex phero-mone(Light et al.,199
19、3;SchmidtBsser,von Arx,&Guerin,2009;Varela,Avilla,Anton,&Gemeno,2011;Von Arx,SchmidtBusser,&Guerin,2012),a synergistic or additive interaction between kai-romones and sex pheromones does not always occur in moth at-traction(Reddy&Guerrero,2004).For example,the combinations of either phenylacetonitri
20、le or 2phenylethanol with AA are both attractive bisexual binary blends for at least two tortricids,the eyespotted budmoth Spilonota ocellana(Denis and Schiffermller)and threelined leafroller Pandemis limitata(Robinson),that when used in traps in combination with these species sex pheromones,signifi
21、cantly reduced male but not female catches(Judd,Knight,&ElSayed,2017a,2017b).However,among the four components of the 4K lure,only LOX has not previously been evaluated in combination with PH for cod-ling moth.LOX is a known floral volatile from many plant genera(Knudsen,Tollsten,&Bergstrm,1993),but
22、 is not commonly found in apple(Giacomuzzi et al.,2016)and has not been reported from other hosts of codling moth,that is quince,Cydonia oblonga Mill.,pear,Pyrus communis L.and walnut,Juglans regia L.LOX can be gen-erated by the oxidation of linalool through microbial biotransforma-tion(Demyttenaere
23、&Willemen,1998),and linalool has been shown to enhance male codling moth attraction to PH in laboratory studies(Schmera&Guerin,2012;Yang,Bengtsson,&Witzgall,2004).Thus,without further testing,it is unclear whether the 4K lure can be used effectively in combination with sex pheromone for monitor-ing
24、codling moth.Also,it should be clarified whether this multifunc-tional lure affects female moth catches,especially when considering the use of attract and kill technologies which can maximize female moth removal(ElSayed,Suckling,Wearing,&Byers,2006).Herein,we report the first evaluation of the 4K lu
25、re used alone and in combination with PH to trap codling moth in both untreated and MDtreated apple orchards.Lures were compared in orchards treated with either PH or PH+PE mating disruption dispensers.The 4K and 4K+PH lures were evaluated against a widely used commercial lure,Pherocon CMDA Combo lo
26、aded with PH+PE and paired in traps with the AA colure,Pherocon AA.2|MATERIALS AND METHODS2.1|Chemicals and lures(E)4,8dimethyl1,3,7nonatriene(DMNT;99%purity)was obtained from the Plant Research Institute(Wageningin,the Netherlands)and loaded into grey halobutyl septa(West Co).Prior to loading,septa
27、 were extracted three times with dichloromethane(99.9%purity)and airdried overnight prior to storage in glass jars at 15C.DMNT lures were prepared by adding 100 l of a dilution in dichlorometh-ane to load 10 mg per septum.Similar volumes of dichloromethane were added three times after this initial l
28、oading to enhance penetra-tion into the substrate.Several commercial lures were provided by Trc Inc.(Adair,OK),including a 3.2 cm2 plastic membrane cup lure loaded with 720 mg of acetic acid(Pherocon AA,TRE3321),and grey halobutyl septa loaded with 3.9 mg of pear ester,(E,Z)2,4ethyl decadienoate(Phe
29、rocon DA,TRE3460),3.5 mg of(E,E)8,10dodecadien1ol(Pherocon CM L2,TRE4111)and combinational lures loaded at these rates with both pear ester and sex pheromone(Pherocon CMDA Combo,TRE3461).Linalool oxide pyranoid(LOX,99.1%purity,57.8%Zisomer and 41.3%Eisomer with for each isomer roughly 5050 of S and
30、R optical isomers)was supplied by Nippon Terpene Chemicals Inc,LOX was diluted in mineral oil(SigmaAldrich,#18512)(10%w/w),and 10 mg a.i.was loaded into 1.5 ml microcentrifuge plastic tubes which contained a dental cotton wick to adsorb the solution.A 2.0 mm aperture was drilled into the 944|KNIGHT
31、eT al.cap of the tube just prior to field application.All prepared lures were stored at 15C until used in field studies.2.2|Field trialsA general protocol for field studies was adopted which included the use of orange deltashaped 28 20 20 cm traps(Pherocon VI,Trc Inc.)with removable liners coated wi
32、th hotmelt pressuresensitive adhesive(Alpha Scents).Traps were attached to poles and placed in the midcanopy at 3.0m height,30 m apart and 20 m from the borders of orchards.Lure treatments were randomized in each orchard with five replicates per orchard,and the experiment was repeated 24.Unbaited tr
33、aps were included as a treatment in all experiments.However,the mean catch with blank traps was al-ways 0.2 moths per trap and this treatment was not included in the analyses.Moths were removed,sexed and counted in the laboratory.Trials were conducted in three“Delicious”apple orchards(48 ha)located
34、south of Wapato,WA(4245N,12042W).Orchards were planted at 720 trees/ha with average tree canopies of 4.04.5 m.Orchards were not sprayed with pesticides during 2018.One orchard was untreated,and the other two orchards were treated with 80 dispensers(Trc Inc.)/ha loaded with either 900 mg of PH(Pheroc
35、on CM MesoA)or 900 mg PH+600 mg PE(Pherocon CMDA MesoA).Four repeated experiments were each run in the orchard with no MD and in the orchard with PH dispens-ers.Two repeated experiments were run in the orchard treated with PH+PE dispensers.Each experiment lasted from 1 to 3 nights to minimize liner
36、saturation and was conducted during August 2018.New traps were placed on different trees within orchards for each experiment.2.3|Statistical analysisThe numbers of females male and total moths caught in traps were analysed using the software R(v.3.5.3,R Core Team,2019).Treatments and experiment repe
37、tition were set as explanatory variables and for each trial analysed using generalized linear mod-els(GLMs).After comparing deviance as well as Akaikes informa-tion criteria(AIC)for fitted GLMs with different distributions,the trap data were found to be nonnormally distributed and overdis-persed in
38、all trials,fitting best with a negative binomial distribution.For each individual test,a family theta value was extracted from a zero distribution(package MASS,Venables&Ripley,2002).To sepa-rate if the treatments differed significantly within an orchard,first a chisquare test(package car,Fox&Weisber
39、g,2011)followed by post hoc analysis using Tukeys allpair comparisons was performed using package multcomp(Hothorn,Bretez,&Westfall,2008),with a significance level(alpha)set at.05.TABLE 1Male,female and total catches of codling moth in delta traps baited with sex pheromone(PH),pear ester(PE),and ace
40、tic acid(AA)compared with traps baited with the 4K lure set alone consisting of PE,(E)4,8dimethyl1,3,7nonatriene,linalool oxide,and AA or in combination with PH in separate apple orchards either untreated or treated with dispensers loaded with sex pheromone or sex pheromone/pear ester dispensers for
41、 mating disruption during 2018Orchard#,treatment,no.trialsaMean(SE)moth catch per trapLuresbMalesFemalesTotalOrchard 1,no MD,4 trialsPH/PE+AA5.7 1.2b4.8 0.7b10.5 1.5b4K8.0 1.8b23.4 3.6a31.4 5.1a4K+PH14.2 2.1a16.0 2.2a30.2 3.1aGLM neg bin Residual degrees of freedom=54X2=22.60,p .001X2=51.66,p .001X2
42、=45.73,p .001Orchard 2,PH MD,4 trialsPH/PE+AA15.1 2.7b3.7 0.7b18.8 3.1c4K16.0 2.0b16.4 1.9a32.4 3.6b4K+PH33.0 4.8a19.7 3.5a52.7 8.0aGLM neg bin Residual degrees of freedom=54X2=16.95,p .001X2=57.97,p .001X2=25.16,p .001Orchard 3,PH/PE MD,2 trialsPH/PE+AA11.7 2.4b7.0 1.1b18.7 2.7c4K13.3 2.6b19.2 1.9a
43、32.5 3.7b4K+PH36.8 8.7a21.0 5.5a57.8 12.9aGLM neg bin Residual degrees of freedom=26X2=24.47,p .001X2=19.54,p .001X2=26.20,p .001Note:Data were analysed using a generalized linear model(GLM)with trial as a fixed effect.Lure treatments were replicated five times in each trial.Column means followed by
44、 a different letter were significantly different in all pairwise comparison,p .05.aOrchards were either untreated or treated with 60 Cidetrak Meso(Trc,Inc.)dispensers/ha loaded with sex pheromone(PH MD)or sex phero-mone plus pear ester(PH/PE MD).bLures consisted of PE,DMNT and PH in grey halobutyl s
45、epta,LOX in a plastic vial with a 2mm hole and AA loaded into a closed membrane.|945KNIGHT eT al.3|RESULTSAnalyses of male,female and total moth catches in traps baited with PH/PE+AA,4K and 4K+PH lures had both similar and dissimilar results across studies conducted in untreated and MDtreated or-cha
46、rds(Table 1).For example,adding the PH lure to traps baited with the 4K lure significantly increased male catch in all three orchards.Also,traps baited with the 4K lure alone or in combination with a PH lure always caught significantly more moths than traps baited with the PH/PE+AA lure.Otherwise,th
47、e results were congruent in the different orchards.The 4K+PH lure grouped statistically with the 4K lure in terms of number of females caught and always caught more than the PH/PE+AA lure.The 4K lure did catch more males but never significantly so than the PH/PE+AA lure.As a lure for males,the 4K+PH
48、 lure outperformed and always caught signifi-cantly more males than the two other lures.4|DISCUSSIONThe relative effectiveness of the 4K lure suggests it might be used to improve several aspects of the management of codling moth(Knight,Light,et al.,2019).The potential of increasing female moth captu
49、res compared with previous lures could,after establishing the efficacy in spring,increase growers ability to establish female moth“Biofix”to predict the start of egg hatch and time larvicidal sprays more effectively(BarrosParada,Knight,&FuentesContreras,2015;Knight&Light,2005a).Enhanced catch of fem
50、ales could also benefit the adoption of action thresholds based on female moth catches(Knight&Light,2005b)and promote the use of precise sitespecific management tactics(Knight,Hawkins,McNamara,Borman,&Hilton,2011).A practical advantage of the new potent attrac-tive lure may be that pest managers can