Accepted Article

are incorporated. strategies withmutuallybene Successful programmes aremost likelygenerated when preventative andtherapeutic and innovative approaches emphasized, highligh controlhost plant and resistance as areasof in integrated tactics pestmanage main control alternative strategiesis therefore, an ofincreasingissue urgency. This paperreviews the strategies, that fortactics areless pushing crops worldwide. RecentEUlegislation hascauseddramatic a shift in management pest Western floweris one (WFT) of themost Abstract Keywords: thrips; article as doi: 10.10 differencesbetween version lead to this may been through the copyediting, typesetting, pagination andproofreading process, which This article hasbeen accepted for publication andundergone full peer review but has not Sanae Mouden * Institute of biology, Leiden University, P.O. ofbiology,Leiden University, Institute Integrated pest management Integrated pestmanagement Correspondence to:SanaeMouden,Research * Frankliniella occidentalis , KryssFacunSarmiento, Peter G.L. 02/ps.4531 This articleis protectedbycopyright. Allrightsreserved. [email protected] ficial, cost-effective andenvironmentallysoundfoundations resistance, -omic techniques in westernflower thrips: ; integratedpest management; biological control; Box 9505, 2300 Box 2300 RA, 9505, The Netherlands. E-mail:

major progress. Knowledge aregaps identified reliable on chemicals. The development of development chemicals. The on reliable and the Versionof Record. Please cite this economically importantpest insects manyof ment (IPM) of WFT with focus on biological focusonbiological (IPM) ofWFTwith ment group Plantand EcologyPhytochemistry, group ting the advances in -omics technologies. Klinkhamer andKirstenA. Leiss

past, present and future past, presentandfuture

This articleis protectedbycopyright. Allrightsreserved. Accepted Article high dispersal capability cause a high pest pressure. pest ahigh cause capability dispersal high towards beneficialnon-target organism resistance to majorinsecticide groups, residue problems on crops, marketable toxicity frequent use ofinsecticides. This overuse of important. by of whichTomato virus transmission, Spotted flowers on leaves, oviposition plant feeding and protection products (EC N° 1107/2009). Ten protection ye products (ECTen N°1107/2009). 2009/ (Directive pesticides use of sustainable and implementationofalternativecontrolme worldwide destabilisation of IPM programs fo management alternativesmanagement toth dueimportant pests damageto theirvast many field and greenhouse crops. WFT developedonemost economically into ofthe horticultural worldwide. pest Thiscosmopol Western flowerthrips(WFT), 1. Introduction and competitive ago AsMorsand reviewedtenyears and competitive species. Hoddle resulted in a resurgence of WFT and associat for protectedcrops.Howeverdevelop mainly with the judicioususe IPM prog of pesticides. mechanical measures, host plant resistance, bi complementary tactics are necessary, incl Therefore, in the framework ofintegratedpest (IPM)programmesmanagement multiple 2,3 Theirsmallaffinity size, forenclosed spaces, high reproductive potential and Frankliniella occidentalis Frankliniella e pesticide-dominated methods. s and contamination of theenvironment. of s and contamination pesticides has led to the ledtodevelopment has pesticides WFT of itan and polyphagous invaderitan andpolyphagous is abundant in ed viruses while depletingitsnatural enemies ed viruses r many crops. To emphasize the developmentTo emphasizethe r manycrops. uding monitoring,uding cultural, physical and rammes forcontrol of WFT have started to 128/EC) as well as on regulation of plant plant of on regulation as well as 128/EC) ological control,andsemiochemicals along Wilt Virus (TSWV) is economically the most most the is economically Wilt Virus(TSWV) asures, theEU issuednew legislation on , continued injudicious of pesticidesuse potential andconcurrent lack ofviable and fruits whileis indirectdamage caused ars after Mors and Hoddle, we aim at at weaim Morsand Hoddle, after ars (Pergande), forms a key agri- and forms (Pergande), akeyagri- 4 Control of WFT mainly relied on relied WFT Control of mainly 1 Direct damage results from from results damage Direct 1 , this ledtoa 5-7

This articleis protectedbycopyright. Allrightsreserved. Accepted Article leafminers. and aphids, for monitoring used be also can traps sticky yellow whereby WFT semiochemicals as lures which sign semiochemicals aslureswhich pepper, eggplant, cucumberpepper, eggplant, andstrawberry. only a feweconomic thresholds for damage WFT have been established such as in tomato, assessment of the economicof WFThas impact intervention andthedevelopment ofeconom andcurrentIn ordertoeffectivelymanage 2. 1 Monitoring WFT2. control tactics IPMapproachesbased havealready been extensively reviewed elsewhere. are discussed. WFT biology and ecology,f identified and newinnovative approaches with control andplant host resistance asof areas reviewing thecurrent knowledge about WFTcontro developed as potentialdecision toolsfor IPMprogrammes. samplings, models forpredictions of WFTpopul in crops with high threat of virus transmis use of sticky traps. Monitoring isbased on regular visualscouting economic to thresholdsareassessed deci Monitoring information onthedevelopmentof

The use of monitoringtools has been expanded bytheaddition of

10 Compared to yellow sticky traps,bluetraps have shown to catchmore ificantly increase thrips catches. ificantly increasethrips undamental tothedevelopment knowledge- of sion, a near zero tolerance forWFTprevails. sion, anearzerotolerance 8,9 majorprogress. Resulting knowledge are gaps de on the employment of control tactics. ofcontrol de ontheemployment of WFT adults on flowersandfruitsor of WFTadults on the However, in high-value ornamentalcrops or emphasisonemergingtechniques the -omics ation growth and spread ofTSWV havebeen only recently begundevelop. Therefore, to anticipate future pest outbreaks,anticipateearly futurepest WFT populations levels relative to the ic thresholds is critical. However, the However, critical. is ic thresholds l in relation to IPM, stressing biological biological stressing l in torelation IPM, 12

11 1,4,7 Based on WFT WFT on Based

7 6

This articleis protectedbycopyright. Allrightsreserved. Accepted Article colonizing throughcolonizing adults interruptionof did not prevent greenhouse invasion bythrips. greenhouse invasion did notprevent combination pressure force ventila ofapositive tomatoes, butnot in another. phenylalanine and femaleWFTabundance wa pupation in pupation in theplant canopy. incidence inwasincidence protectedtomato20% debris forms the first line of WFT defence. WFT lineof first the debrisforms Sanitarypracticessuch pests. of management Since ancient time,farmershave been relyingon cultural orphysical practicesforthe Cultural, mechanicaland physical control of WFT 2.2 it canbe controlled more easily. damage in avegetative crop. damage chrysanthemum population numbers in ornamentals. also effects WFT abundance. Increasedlevels adults. Irrigation, creatinga less favorable enviro promotedWFT larval development in different vegetables. WFT immigration intoprotected crops but requires optimization of ventilation. did not lead to increased thrips damage. increasing marketable yield. potato orbaby corncompromised to beanyieldbutreduced damage thebeanpods 20 In contrast, high relative humidity favo 28 18,24 18,24 21 Fertilization increases plan increases Fertilization 26 High rates ofdevelopment rates phosphorusfavoredthrips High but Flowering lowered chrysanthemumsastrap plants WFT 22 25 Similarly, high levels of aromatic amino acids ofaromatic acids amino levels high Similarly, Trap crops draw WFTaway Trapcrops fromthe crop decreased by greenhouse window screens. nment forthrips, decreased numbersWFT of 13,14 13,14 orientationandhost-finding behavior. 27 as removing weeds, old plantmaterial and tion system with insect provetion systemwithinsect screensthough IntercroppingFrench beans with sunflower, red WFT larval development and stimulated developmentandstimulated red WFTlarval Screening greenhouse openings prevented of nitrogen fertilization increasedWFT of nitrogenfertilization s observed in onestudyon field-grown 17 UV-reflective WFT mulch repelled UV-reflective 23 t development and growth but, Apositive correlation between 15

where WFT WFT 16 18,19 A

This articleis protectedbycopyright. Allrightsreserved. Accepted Article numbers ofWFT. trichomesapplicationto methyljasmonatehigher glandular type VI inresponse trapped negatively correlated with the amount of epicuticularwaxongladiolus leaves. cuticles, waxy and/oras epidermal structures including trichomes.was WFTdamage The surface of ahost plant can serve as aphysi structures defense Morphological 2.3.1 pests. protection against nature ofplant defensive traits plays a critical chemical traits that influence the amount damage of caused by pests. Understanding the webofcomplex a large involves resistance to reduceinsectattack andled tohost this plant resistance.Thehost plant of study of de variety have evolveda evolution, plants and for haveco-existed Plants more insects 2.3 Host plant resistance damage. we amino acids ofconcentrations aromatic concentrations theycanbe involved also inWFT resistance.Among different crops,low asnutritionalmetabolites, chemicals, are gene Plantchemical defense can arisefrom both resistance plant host Chemical 2.3.2 by chemical host plant composition. area. feeding damage and morphologicaltraits assuch 31,32 23 Instead, thelatterInstead, provided clear indications thatresistance was mainly influenced Nevertheless, theseuniversal compounds doprovideuniquenessnot any and are 30 However, other studies did not ob not did studies other However, interactions, mediated by morphological and interactions, mediatedby fense mechanisms,fense constitutive andinducible, role indesigning with cropvarieties enhanced primary andsecondary primary metabolites. Primary than 350 million years. In the course of thecourse In years. million 350 than cal barrier through morphological traits such re correlated withreduced WFT feeding rally beneficial for thrips. However, at low hairiness, leaf age, dry hairiness, weight leaf and serve anycorrelation between WFT 29 Induction of of Induction This articleis protectedbycopyright. Allrightsreserved. Accepted Article alanine in carrot. alanine in feroluylquinic acidinchrysanthemum, acylsugars inthewildplant species glucoside kaempferol practical applicationpest managementso for ofthese interference potential kininogen domainCdid cystatin inhibitWFT. not 3 and overexpressingstefinor equistatin,were A equistatin, when incorporated into artifi an anti-elastaseprotease inhibitor, noticeably delayed WFT damage. forinsect ofan growth aminoacids availability Plantprotease inhibitors(PIs) arenaturally Transgenic 2.3.3 plants varieties, isobutylamide was suggested tobe investigated chemical host plant resistanceto a study WFT.In different on chrysanthemum focuses role ofmetabolitesin on the secondary not likelytobeeffective in resistance on th protease inhibitors in potato signific functions in human health prevention. antioxidant attentionfor their considerable receive negative onWFT, butalso effect did nota show clear effecton WFTfecundity. Transgenicover-expressing chrysanthemums, mult in subsequent susceptible compounds plants, forconstitutive Developing aneco-metabolomiccomp approach in-vitro 31,33,35,36 31,33,35,36 bioassays. Interestingly, some of these meInterestingly, some ofthese multidomain with basiccellfunctions proteins hindered has a 34 Identifiedcompoundsincluded jaconineand jacobine, antly improvedresistancethrips. to

occurring plant defense compounds reducing the reducing compounds defense plant occurring cial reduced diets, WFT ovipositionrates. deterrent to thrips whileoverexpressionof thrips to deterrent d development. Transgenic development. alfalfa, d expressing associated with WFThost plant resistance. eir own. Therefore, the majority of studies WFT resistance were identified and validated validated and identified were resistance WFT plant defense. Up to now few studies havestudies to nowfew Up plantdefense. aring metabolomic profiles of resistant and ofresistant profiles metabolomic aring intomatoandsinapic and acid, luteolin, far. Targeting virus resistance, transgenic transgenic resistance, virus Targeting far. 39 icystatin, a potato inhibitor,icystatin, a proteinase Cysteine potatoPI transgenic Cysteine plants Jacobaea vulgaris, tabolites did not onlyshow a 40 Expression of multi-domain multi-domain Expression of 37 Purified cystatin and chlorogenic-and 41 However, the However, β 33 38 -

This articleis protectedbycopyright. Allrightsreserved. Accepted Article 35S::prosystemin transgene, constitutively activatingJAdefense. Arabidopsis The JA-responsive genes thrips. against defense in role important an plays JA-pathway The and pathogens. insects wellas naturally asethylene (ET) trigger occu signalingcommunicating The planthormones pathways. salicylic- (SA) and jasmonic acid(JA) attack, presumably Inducedcosts. minimize to toconstitutive defenses,plants In addition Induced 2.3.4 resistance modified thecommercializationlimit ofgenetically crops in Europe. the scientificcommunity.However, highly re strategies, is recognized as a promising approach for thrips and tospovirus management by uninfected ones. infection, enhancedthe performanceofWF defense andSA-regulated inresponse systems interactionbetween toTSWV JA- the infection in tomato with acibenzolar-S-methyl a func (ASM), WFT larvae. tomato expressing G tomato expressing tomato mutant tomato mutant resulted in a decreased preference, performance and abundance of WFT. response ofaplant,thereby enhancing resist plants to thrips. 42 Arabidopsis The use of transgenic plants, alternated or simultaneously used with additionalofwith transgenicor simultaneouslyuse plants,alternatedused The 45 def-1 Treatmentswith exogenouselicit N glycoprotein, interfered with TSWV acquisition and transmissionand by acquisition withTSWV interfered glycoprotein, , deficient in JA, in tothe mutantexpressing, deficientcomparison a inJA, induced SA-regulated gene expression.SA-regulated gene induced VSP2 43 WFTreached maximal in reproductiveperformance the and and PDF1.2 rring chemical responses responses chemical rring were stronglyof stimulated were upon exposure use inducible defenses as a response to pest pest to response asa defenses inducible use strictive political and regulatory frameworks frameworks politicalandregulatorystrictive defenses defenses are regulated bya networkofcross- ance to thrips. Application of JA in tomato of JAin tomato Application tothrips. ance tionalSA incidence, analogof reducedTSWV T preferringinfectedTSWV plantsover ors activate the natural activate defensiveors 43 44 Theresulting antagonistic In contrast to WFT, TSWV protecting plantsfrom protecting 46 Treatmentof This articleis protectedbycopyright. Allrightsreserved. Accepted Article females showed WFTlarvae. ahigherpropensity kill toattack and survival while both predators showed similar efficacy showedwinter. similar in predators while both survival swirskii WFTbiocontrol agent is also influenced by degenerans seeking to find a superior WFTpredator. Species such as controlDue toinadequate achievements anumberof other mites havebeen studied, but didnotinfluence WFTpopulationdensities. N Neoseiulus efficacy. predatory Thefirst mites used forWFTcontrol were been recordedas predators of WFT and various havespecies beenassessed for their mites ( The principal predatorsassociated Predatory mites 2.4.1 WFT. against agents used biocontrol available the mostcommonlycommercially 1summarizes nematodes. Table predators and parasitoids and microbials beingsubdivided in enthomopathogenic fungi and are known toattack WFT,which canbe separated in two groups: macrobials including approaches to sustaintheir abundance andefficiency. Alarge number of natural enemies Biological control uses the augmentativereleas Biological 2.4. control andinterest particularly might in beof valueconjunction withother IPMapproaches. . cucumeris provided higher thrips higher control provided than (formerly (formerly and , A. swirskii A. montdorensis spp.) and pirate bugs ( bugs andpirate spp.) Amblyseius) cucumerisAmblyseius) proved be to a better WFTpredator thaninsweet pepper proved to beeffective predators of WFT. Orius host plant specieswhereby increased trichome N. cucumeris which primarily feed upon first instar larvae. instar first upon feed whichprimarily 47 e ofe natural enemiesasas conservation well with WFT biological control with WFTbiological are phytoseiid spp.). Several species of Several spp.). Induced resistance is recently gainingmore insummer, dueto abetter likely 51 A. limonicus, A.swirskii, A. Efficiencyof 50 Amblyseius barkeri Inchrysanthemum 48,49 Amblyseius A. swirskii Compared to

since have as a a as and A.

This articleis protectedbycopyright. Allrightsreserved. Accepted Article swirskii densities hinder mite performance. thrips control isnotnegatively affected by the presence of Although the presence of canleadto of thrips at ovipositionsites,pres predators were observed to preferentially ovipos Success of to beingan efficient predator ofWFT, short day conditionsin autumn as theyenter diapause. insidious Observations from fieldand glasshouse experiments insweet pepper demonstrated that of Severalthrips. speciesand of andlarvae awideadults range of insect species such as aphids, whiteflies, mitesspider Orius Predatory 2.4.2 bugs been investigated. Supplyingbeen investigated. pollenimproved of performance food sourcemites. for Therefore worldwide. cont forbiological used main predator the successful inyearroundcont all biological production. chrysanthemum as did the addition chrysanthemum asdid of decapsulated brineshrimp( cysts O. laevigatus , commonly known aspirate bugs, are known to begeneralist predators, preying on isa better predator onwhitefly inthe presence ofthrips. suppressed WFTtoalmostextinction,but Orius 49 49 In addition to controlofWFT, In addition Since its commercial introductionin2005 Sinceitscommercial in ornamentals depends in on the complexityof flower structure. hasbeen WFTresistancewound in tomato shown toinduce through Orius umably toprotect their offspring. addition food to ofsupplemental 52 have been tested to evaluate their use against WFT. Thrips can also consume A. swirskii rol of WFT in vegetables andornamentals. rol ofWFTinvegetables A. swirskii rol ofWFTin vegetables andornamentals a short-term escape of from thrips predation, is easilyreared which allowseconomic mass it at sites without or thrips, to kill more failed to control WFT properly under 57 also provides control of whiteflies. providescontrolof also In contrast, In contrast, A. swirskii

A. swirskii A. swirskii whitefly, while in contrast in contrast whitefly, while 55,56 A. swirskii 53 Thrips are not the best not thebest Thripsare has, therefore, has, become

A. swirskii O. laevigatus Artemia in control in of WFT in eggs and female 59 has recently recently has Oviposition Oviposition sp.). has been been has 54 Next Next 59,59 O. A.

This articleis protectedbycopyright. Allrightsreserved. Accepted Article expensive to mass rear. for their potential to control WFT. To date, Parasitoids 2.4.4 response. ornamentals. been repeatedly showntosuccessfully contro coriaria miles investigated of which one-third of their life as pupae inthe soil. Di Most research on WFT biocontrolfocused on Soil-dwelling 2.4.3 predators results. Heterorhabditis various nematodespecies and in strains the nematode genera agents WFT biocontrol used as Entomopathogens Entomopathogens 2.4.5 control.developmentWFTwasp time hindersefficient in resulting larvae, intofirst-instar oviposit treatment wasinsufficientfor control ofWFT. flower and buds foliar non-leth was terminals, )

and 65,66 ) , Ceranisus menes are commercially biocontrol producedas agents against WFTpupae. 60 Gaeolaelaps aculeifer Although akeyenemy in ofWFT, natural biocontrol However, foliar application However,of foliar 67,68 against soil-inhabiting WFT pupae pr pupae WFT against soil-inhabiting Treatmentwith Macrocheles robustulus, Stratiolaelaps scimitus 59

and C. americensis asthe wellasbeetlerove Thripinema 64 Underlaboratory conditions, these parasitic wasps S. feltiae death of thepre-pupal However, stage. deathof slow , are the only two parasitoid wasps investigated fferent soil-dwelling predatory mites havebeen 67 adult However,and larval stages. WFT spend

l WFT adults and larvae in vegetables and nematodes, infecting within WFT residing al andcaused sterility WFT. of female This consist of nematodes and fungi. The use ofThe use consist ofnematodesandfungi. , in the presence of a wettingagent, has oduced low andinconsistent control Dalotia coriaria Orius (formerly spp. are spp. relatively Steinernema (formerly 61-63 61-63 Hypoaspis Atheta and This articleis protectedbycopyright. Allrightsreserved. Accepted Article populationsand floral in greenhouse vegetable crops. WFTpreference ordevelopment havebeen observed. causing disease symptoms, has beenexploredfor formulation. augmentative biological control agentsremain anisopliae applications ofdifferentfu addition, larvae have thicker cuticles, which maydelaypenetration offungus. larval and pupal stages possibly because moltin nutrients for growth reproduction.In and gene Entomopathogenic fungal conidiainfect thrips by penetrating their cuticle to obtain alone. on WFT greenhousein crops butthe effect was not greaterthan using one predator chrysanthemum. Formulations targeting the soil stage efficacy has been inconsistent likely due to buttheir applications forfoliar now available are fungi ofentomopathoghenic formulations predation. led tonegative inte in laboratory trials andtiming compatibility oftreatments. Application of entomopathogens are used asalternative or requires back-uptreatments.This careful Combinatorialof natural treatmentsenemieswi Combinatorial ofbiological use control 2.4.6 bassiana 58,77 showed sublethaleffects on the progeny of treated WFT adults. Incontrast,a combination of and 76 Simultaneous useof predatory mites andpirate bugsdidhavea negative effect 73 Recently,theuse ofendophytic developing fungi, within plant tissues without Lecanicillium lecanii Lecanicillium 72 major A constraint to the useof entomophatogenic fungi as ngal strains belonging to belonging strains ngal (formerly ractions on WFT control through intra-guild intra-guild through control on WFT ractions O. laevigatus have shown promising results in potted in potted results promising have shown varying ambienthumidity and temperature. difficulties in mass difficulties production, storageand ral, adult thrips are more susceptible than than aremoresusceptible thrips adult ral, th different arthropodsarthropods with or g avoidscontactg withfungalinoculum. In Verticillium WFT control. So far no negative effectsSo farnonegative on WFT control. 74,75 74,75 A. swirskii A. swirskii 69,70 Beauveria bassiana Beauveria

and Besides the direct effects, ) significantly reduced thrips Macrolophus pygmaeus together withtogether , N. cucumeris Metarhizium Metarhizium 71 Several Several

Foliar Foliar , a B.

This articleis protectedbycopyright. Allrightsreserved. Accepted Article control in green beancomparedcontrolreleases. ingreen to individual ornamentals beyond thatcaused by foliagepredators alone. aculeifer feltiae Heterorhabditis fine-tuning suitingthecropinquestion. forpr of WFT are control agents biocontrol in sweet pepper. predator achieved tocontrolaphids, andgeneralist thrips controlaphidsenhanced ofboth Likewise, laboratorycombinationsof Likewise, treatment of different mites orpiratebugs WFT all st life targeting agents biocontrol overlap duringasinglecrop production He cycle. Thrips generally complete their life cycle with mites with the nematode mites with dissemination seemed to be hindered by mite grooming. not inhibitenhanc did nor predatorymites fungi improve control,thrips while combinations of combinations of treatments. M. brunneum increased mortality increased ofWFTpupaeingreenbean. , D. coriaria 82 Concurrent use of the soil dwelling mite the dwelling ofsoil use Concurrent nematodes with the mite foliar-dwelling O. laevigatus 78 and and Combinations of the entomopathogenic of fungus Combinations or the nematodeS. the or B. bassiana S. feltiae with the respective nematode achieved higher control ofWFTcomparedtosingle different soildwellingpredators with achieved WFTcontrolgood incyclamen,while as foliage predators with thesoilpredators predators foliage as omising butmanagementrequire careful and ages havebeen investigated. Simultaneous these predatorswith entomopathogenic the in twoweeks causing several generations to feltiae nce, combinations of foliar and soil-dwelling andsoil-dwelling combinationsof foliar nce, e the controle of WFT, because fungal did not reduce thrips numbers in 81 83 Combinations of different predatory predatory ofdifferent Combinations 69,79 H. aculeifer It is apparent that combinations of N. cucumeris

s failed to controlthrips. 80 In contrast,the use of with the provided superior B. bassiana S. feltiae

nematode didnot with with G. S. 59

This articleis protectedbycopyright. Allrightsreserved. Accepted Article with poor taste. Methyl isonicotinate, the acti the isonicotinate, Methyl with poortaste. a plant defensestrategy against WFTas a flor methylbutanoate. Two keypheromones in maleWFT wereidenti Pheromones2.5.1 mo for lures as used are Semiochemicals communication betweenarthropods alleloch while semiochemicals functioning assignalcompounds.Pheromones for intraspecific serve inAn importantfocus applied control the pest manipulation is of adult insectbehavior using 2.5 Behavioral control chrysanthemum plants, they were deterred by linaloolglycosides. theyweredeterred chrysanthemum plants, WFT wereattractedbypure and asesquiterpene attracted adult female pyridines phenylpropanoids, monoterpenes, benzenoids, including scents, volatile Various forfeedin hosts usedtolocateplant Volatiles Allelochemicals 2.5.2 suggesting its function rates, landing decrease and oviposition reduce rates, take-off and movement to increase producedalarm as are pheromones in anal larvalSynthetic droplets. equivalents causedWFT areinusecommercially.(Biobest), Decylanddodecyl respectively, 10-and acetate, 12-AC The syntheticanalogues,Thrifemale WFT. male specific cuticular hydrocarbonwas to suggested inhibitmating. Systems), is anattractantSystems), and femaleWFT forbothas well andismale asotherthripsspecies 84 84 The latter is a sexual aggregation pheromone attracting both maleand pheromone attracting aggregation asexual is The latter as anpheromone. alarm linalool emittedby aswell linalool engineered as nitoring aswell control purposes. F. occidentalis plineAMS(Syngenta Bioline)andThriPher ve ingredient of Lurem- 85,86 al antagonist, balancingfragrance attractive g and oviposition can can beapplied andoviposition as g lures. fied: (R)-lavandulylacetate and neryl(S)-2- emicals mediate plant-insectinteractions. mediate emicals More recently,7-methyltricosane, WFT a in a dose-dependent way. dose-dependent in a 89 The lattermayrepresent 57

TR (Koppert Biological TR (KoppertBiological 88 While While This articleis protectedbycopyright. Allrightsreserved. Accepted Article attracting thrips attracting toparticular tr predator insecticide, insecticide, increased larvalmortality ofWFT. insecticides including insecticides pyrethroids, neonicit volatile (S)-verbenone, was describedfrompine pollen. strategies. semiochemicals hold promise great forthrips mass trapping “lureand as well kill’ as feeding andreproduction byutilizingdiffer conjunctionwith sticky cardtraps. Adultthri Currently the three commercially availableWFT semiochemicals are mainly used as lures in previous previous reviews to which we refer to for further detail. Managementof on thripshasrelied application as ofinsecticides has been described in at European level,are listedin Table 2. application of insecticides. Commonly usedinse almost wherean particularly forornamentals, Chemical control is among one of the most frequently used methods to suppress WFT, 2.6 Chemical control used tolocate hostplants. employs LUREM-T for autodisseminationentompathogenic the fungusemploys LUREM-T of carvacrol exhibited both a feeding as well resulted in adose-dependent habituation. jasmone deterred WFTlarvae from feeding althoughand settling repeated exposure andcis- ofmethyl-jasmonate finding. can beutilizedfor ofhost Applications disruption O. laevigatus 96,97 Combination of dodecyl acetate with maldison, an organo-phosphorous increased theabundanceof 90 Recently, a new potential active ingredient forthripslures, Recently, anewpotential aps provided withfungal inoculum. as a oviposition deterrent oviposition effect toWFT. as a inoids, organophospates and carbamates kills 92,93 92,93 zero damage tolerance encourages intensive 98 ps constantlyps exploretheirhost rangefor cticides formanagement of thrips,approved ent cues including vo ent cues Use of LUREM-T together ofLUREM-T withtheUse WFT The monoterpenoid phenols thymol and

the latter. 91 Volatiles with repellent activities 4,7 99 The use ofbroad spectrum use The The ‘lure and infect’ strategy 100 latiles. Therefore, latiles. M. anisopliae 94,95 94,95

by This articleis protectedbycopyright. Allrightsreserved. Accepted Article controlWFT. of compatible with natural enemies and, currently, provides the most effective chemical greenhouse conditions.greenhouse programs including entomopathogenic organisms successfullycontrolled WFT under schemes need to be complemented with other compatible control approaches. control compatible other with complemented tobe need schemes applications should be accurate and precise econ if beused only insecticides should does not necessarily focuson rotation schemes coupledamong differentclassesinsecticides. withrotations of resistance insecticide as revi most chemical classes as hasbeen extensively revised elsewhere. including spinosad, have led to the development of WFT resistance to active ingredients of lufenuron. However, frequent applications computational systemstool providepowerful a The mechanisms. emergencedefense ofin advances omic-based as techniques well as systems-based Suchapproaches. integrativeapproaches enable acomprehensive viewof hasbe metabolites, to methods, fromgenomes preventionpest andmanagement insects.of Pest management programs are constantly se Future ofWFT directions 3. native outcompeting thrips species and naturalenemies disrupting WFT management. 7,101 predatorymites, bugs, and othercompeting thrips species.

Spinosad, anaturalreduced-risk insecticidede Spinosad, 4 New, narrow-spectrum includepyridalyl insecticides,forWFTcontrol and 103 103 Various insecticideshavebeen shown to becompatible withWFT ewed in other publications ewed inotherpublications comprises resistance monitoring control: technologies ‘Omics’ omic damage threshold are reached whereby threshold whereby damage are reached omic of broad and narrow sp narrow and of broad arching forinnovative approaches advancing The development ofnon-targeted analytical while conservingnatural enemies. Rotation reducingoverall insecticideTherefore, use. to drive innovationin cropprotection. todrive en amajor driver forthe adaptation of rived from anactinomycete bacteria is 103,104 103,104 5-6 However, development of 5-6,102 Management of WFT of Management WFT ectrum insecticides, 5 Rotation Rotation 1,4-

This articleis protectedbycopyright. Allrightsreserved. Accepted Article as viruses. as alternative control based strategies onthemolecu toel possibilities newpowerful open will ancestors, therefore, provide a promising sour promising therefore,providea ancestors, reducedgreatly phenotypic and diversitygenetic While domestication of wild plantsthrough se genomics Plant 3.1. ourtechnologies byimprovingknowle resistant crop varieties, generates valuable Understanding plant-insect interactions, geneticvariations among insect populations and peruvianum known toconferresistan control.Genes TSWV for polyploid ornamentals. Atpresent, successful of double-stranded RNA into adult thrips. affecting the larval developmentofthrips. quantitative traitlocusmapped (QTL), to chro resistanceinsect intobreedinglines. traits Identifying sets ofgenesor metabolites as observed in variousvegetables andor the presenceBesides, ofconsiderablevariatio collection of assembled an annotated sequences. an annotated ofassembled collection a recentlygenomic project hasonlyand proteomic developed forWFT including tools as world-wide theireconomicimportance Despite genomics Insect 3.2. 111 ), Sw-7( An RNAinterferencetool has beendevelo L. chilense ) and Tsw ( Tsw ) and dgecomplex of resistance traits. C. chinense 112 106 namentals, canbe exploitedaswell. information that provide new opportunities and opportunities providenew information that Targeting theATP synthase vacuolar Targeting subunit-B This approach,however, beless might suitable biomarkers enables the introduction of novel theintroductionofnovel enables biomarkers n in to resistance WFT between accessions,as Inahighlyresistant a pepper accession, ucidate function thrips anddevelop gene ce forbreedingofWFT resistance traits. ce against TSWVisolates ce against include: Sw-5( mosome 6, confers resistance to WFT by lection improved yield and palatability, it it lection improvedyieldand palatability, breeding of breeding resistant iscultivars limited to 109,110 crop pests, the ‘i5k’ (5000 insect genome) insect the‘i5k’(5000 croppests, lar interaction of thrips with plants as well as lar interactionofthripswithplants leading to loss of insect resistance. Wild of insectresistance.Wild toleading loss ). 107,108 The availability of the thrips genome of thethrips genome availability The ped using microinjection for delivery

32,35,36,105 32,35 L.

This articleis protectedbycopyright. Allrightsreserved. Accepted Article yet developed. proteins reacting toviral infection havebeen leafminer basis for a multi-resistance breeding program. compounds conferring resistance to different he resistancetoand thrips opensopportunities analysesmetabolomics provide powerful oppo integrated network underlyingdefense mechan suppressing or inducingsuppressing ordefense plant responses. the development datawillfacilitate sequence inhibition responses. of plantdefense susceptible. approaches using genotypes with contrasting leve untargetedfashion. Sofar,metabolomicsha Metabolomics has a potential great to detect a wide range in of ancompounds unbiased or Metabolomics 3.3. WFT larvae. of mortality high in Alternatively, down-regulatingmediated RNAi, an symbiontgene, essential resulted tubulin resultedgene WFT inincreased mortality and reduced fecundity of surviving females. infected WFT lead to theputative annota involved in thrips resistance are resistance enhancedinvolved inthrips upon UV-Bexposure. plants thereby, enhancing WFT control. Rutin and chlorogenic acid, two phenolic compounds environmental factorsmayincrease concentratio secondary metabolites involvedWFT resistance in could beused to develop new protection

( Liriomyza 34 Addressing the metabolome, the however,allowsAddressing investigatingthecomplexand 109 Sequencing the salivary gland transcriptome of TSWV-infected and non- and TSWV-infected of transcriptome gland salivary the Sequencing

trifolii ) has been described in chrysanthemum. in described been has ) 113 For transmission of TSWV a suit ofTSWV of Fortransmission WFT candidate 110 availabilityThe ofWFT and genome transcriptome tion of genes involvedtion inof genes detoxification identified approachbut noRNAi for disruption An overlap of resistance toWFTandcelery for ‘metabolitebreeding’. Identification of rtunities identifying metabolic markers for metabolicmarkers identifying rtunities s mainly beenrestricteds tocomparative rbivores, i.e. cross-resistance,rbivores, i.e. a could form of approaches identifying thripseffectors of approaches isms. Combinedwithgenetic approaches,

ns ofresistance related within metabolites ls ofresistance, classified as resistant or 114 105 105 Inaddition,plant Manipulation of

and This articleis protectedbycopyright. Allrightsreserved. Accepted Article for identification of resistance sources is vital for resistance breeding programmes. throughput screening (HTS) system for thrips resistance. Screeningnumbers large of plants Employinggenomic aswell metabolomi as High-throughput 3.4. screening into endophytes increasing resistance to WFT have not been successful so far. successful notbeen have toWFT resistance increasing into endophytes video tracking of WFT behaviour. WFT of tracking video Recently, ahigh-throughput phenotypingmethod has beendescribed using automated induced resistance to Coloniza emission. chemistryorvolatile plant resistance. Assembly ofmicrobial communities microbials offeraNext plants, hugesource to organisms. provide the new minimizing of actionswithimprovedeffects modeon selectivity, non-target agents whichenhance or activate the plan potential to be exploredplant for defense to WFT. syringae explored. Similarly,the effect need tobebacterium of thrips, still the directedmicrobialcommunities defense ground onplantabove against insects, suchas bacteria opens the way for breeding of plants maximizing bacterial benefits. bacterial maximizing of plants breeding way the for opens bacteria Geneticvariationgeneral. to inresponse thecapa Rhizobacteria are known to play an importantro against WFT deriving from plants or microbials are absent. Development of stable thrips thrips stable of Development absent. are microbials or fromplants deriving WFT against lacking.Simila isstill thrips damage assessing producing the JA analogue coronatine and thus triggering herbivore defense has a likely due arepellent effectof volatiles. 118 However, a reproducible high-throughput method rly, HTS systems testingfor active metabolites ts’ ownmechanisms defense orwhich may tion of onion seedlings by fungal endophytes endophytes byfungal seedlings onion of tion may influence performanceofthrips throughmay influence cs techniques however, requires ahigh- requires however, techniques cs city of plants in reactingcity ofplantstothesebeneficial le inplant growth,le nutritionandhealth in of metabolites of for insect tobeused 116

115

The effect of soil soil of The effect Investigations Investigations Pseudomonas 74,75 117 117

This articleis protectedbycopyright. Allrightsreserved. Accepted Article of different inonepests cropping scarce.is system vertical Besides, into anthese ‘IPM toolbox’. respectivemethods assingle-solutionapproaches little evidence that IPM hasbeen largely adopted. Many studies seek to develop their management. However, despite the various benefits expected from IPM, there seems to be fromAs 2014, farmers in the EU are obliged to implement the principles of integrated pest 4Conclusions derived cell lines,beyond primary cellcultures, has been unsuccessful until now. development. ofHTS receptorstoformthebasis orolfactory gustatory opportunity toidentify unprecedented an provides sequence genome thrips the of availability the However, becomes increasinglybecomes important.Some breede WF incontrolof research progress Significant research. by transdisciplinary partners withcommercial operation should but, researchlevels multidisciplinary effectiveness andprogress, strategies should determine theindividual contributions and,he inapproaches sustainablemultiple pest management, while retainingthe capacity to scientists in different groups develop protoc activities that cannot be pinned down into a methodologicalsingle blueprint. How can Clearly, researchintotheintegration ofme fundamental to achieve successful control ofWF task. remains a In complex knowledge-based T has been made. Host plant resistance to WFT plant resistancetoWFT beenmade.Host T has integration of control atIPM measureslooking be driven through applied outcomes inco- ols and tests that allowthe combination of thods involves cooperative, jointly planned cooperative,jointly involves thods nce, modify improveand these? For optimal notatinter-and only beintegrated rs already have varieties with different withdifferent already varieties rs have to pest toproblems ratherthanintegratingpest T, yet, it presents significant challenges. significant challenges. itpresents yet, T, tegrating different control tactics is tactics differentcontrol tegrating 7 Developingand implementing IPM 119

This articleis protectedbycopyright. Allrightsreserved. Accepted Article of thrips. of thrips. remains theneed for approach transdisciplinary basic question remains of how one gets consis different production systems.Controllingpests order toachievesuccessfulcontrol, strategies it becomes even more complex. In crop protecti well asonecological, environmental andeconom on as compatibility programs withemphasis IPM in thecontextof ones shouldbeviewed enhancing resistance. However, not only new st allow development improvedthat of cultivars developmentof novelbiological products, including microorganismsand will metabolites, protection strategies against thrips. Molecula important contributionsfor biologic developing Nevertheless, muchremains tobelearnedabout plant-insect interactionsto makefurther of thrips.With the‘omics’ revolution, wehave there are manyexciting(bio)-technologicadva semiochemicals, not only formonitoringbut also controlchallenginparticular, makebiological crops. Neverthelessprotected shortcrop in not asstraightforward.Recently,more emphasi su crops certain for however, ratings, resistance cycles and lowcycles thresholds for ornamentals in g. Another promising approach isthe use of r modifications, genetic engineering andthe the tools athand tofully this grasp potential. thatnces will undoubtedly boostthecontrol should be tailored to fit the requirements of requirements should betailoredtofitthe ally, environmental friendly,sustainable crop is not a trivial issue, andneverbeen.The issue, has not trivial is a ch as polyploid ornamentals this approach ornamentals this ch aspolyploid is s hasbeen put controlon ofWFT biological es integrating different practices for control are able to respond to WFTattackby are ablerespond to ic consequences. Looking atdifferentcrops tent long-term control.Most importantly rategies need to be explored but existing needtoexisting bebut rategies explored on, as in life, one size does not fit all. In not does fitall. onesize in on, aslife, forthripscontrol. Looking tothefuture, This articleis protectedbycopyright. Allrightsreserved. Accepted Article 35. 13553. This information. review is part of a project funded by Technology FoundationSTW,project is publication in this trade names use of The Acknowledgements solely for the purpose of providing specific providing of for thepurpose solely This articleis protectedbycopyright. Allrightsreserved. Accepted Article References References 5. 4. 3. 2. 1. 6. 7. 8.

thrips, Technol ornamental crops grown in greenhouses: have we reachedan impasse? Western FlowerThrips. consequence for theacquisition andinoculat Exp Entomol Appl Genetic variation inchrysanthemum forresistance to 89 Bielza P, Insecticide resistance manageme P, Insecticideresistance Bielza Cloyd RA, ( PetersMaris PC,JoostenNN,DandThrips GoldbachRW, inpepperresistance and its De JagerCM,ButôtRPT, KlinkhamerPGL,DeJongTJ,WolffKandvander MeijdenE, Morse JG mechanisms, andmanagementmechanisms, strategies. andReitzLei Z SR,WesternGao Y, flower thripsresistanceto insecticides: detection, occidentalis Demirozer O,Tyler-Julian K,Funderburk J,LepplaNand ReitzSR, vegetables in Florida. Coll M, Shakya S, Shouster I, Nenner Y a Y Nenner I, Shouster S, Shakya M, Coll markets. occidentalis Frankliniella ( 2006) Frankliniella occidentalis

3 Entomol ExpAppl . : 1–9 (2009).

and and (Pergande) integrated pestmana integrated (Pergande) Hoddle MS

77: 277-287 (1995). 277-287 (1995). Pest Manag Sci Phytopathology

, 122: management in strawberry: consideration oftarget consideration in strawberry: management of thrips Invasion biology . 59-67 (2007). 59-67 Pest Manag Sci

68 Frankliniella occidentalis : 1537–1545 (2012). (2012). : 1537–1545 93: nd Steinberg S, Decision makingtoolsfor Pest Manag Sci nt strategies againsnt strategies 96-101 (2003). (2003). 96-101 ion of Tomato Spotted Wilt Virus bythe ion ofTomatoSpottedWilt

64 :1131–1138 (2008). (2008). :1131–1138 gement programs forfruiting .

Annu Rev Entomol Rev Annu

68 Frankliniella occidentalis. : 1111–1121 (2012). : 1111–1121 t the western flower ) management on management ) Frankliniella Frankliniella 51 : 67 Pest Pest – This articleis protectedbycopyright. Allrightsreserved. Accepted Article 11. 10. 9. 12. 13. 15. 14.

protected crops. attractant for monitoring FickenAbdullah ZS,GreenfieldKJ,TaylorP, M, andButt Bnew JW,Wood TM,A incidence-mean models. sampling ofwestern flower thrips infe Ugine TA, SandersonJP, WraightShippSP, L,WangKand Nyrop JP,Binomial 30: occidentalis Wang Kand ShippSimulationmodel JL, forpopulationdynamics of influenced by thevirus. Virusconsidering spreaddynamics, Ogada PA, Moualeu DP,PoehlingH-M,PredictiveSpotted models forTomato Wilt Northfield TD, Paini DR,Funderburk JE and Reitz SR, AnnualCycles of 217-222 ( 2008). ( 217-222 reproductivehosts: Predicting poss spp. (Thysanoptera: Thripidae) thrips abundance on North Floridauncultivated protected crops while minimizingin ofventilation Ben-YakirTeitel M,and M, Optimizing D,TannyJ,ChenMBarak Entomol Am of oviposition Feedingand H-M, Poehling and Subramanian Meyhöfer S, JO, Nyasani R, Frankliniella occidentalis Frankliniella 1073-1081 (2001). (2001). 1073-1081 101: 769-778 (2008). 769-778 (2008). 137: (Thysanoptera: Thripidae) on greenhouse cucumber. 204-213 (2013). 204-213 (2013). SpringerPlus PLoS One Environ Entomol for crops and weeds in Kenyan French bean French fields. Kenyan forcropsand weedsin westernflower thrips

4 : 89 (2015). (2015). : 89

11: Frankliniella occidentalis ible sourcespestoutbreaks.of e0154533 (2016). e0154533 vasion by whitefly and thrips. thrips. and by whitefly vasion 40: sting floweringgreenhouse crops using 381-390 (2011). (2011). 381-390 , Frankliniella occidentalis , Frankliniella

specific life processes as as processes life specific Ann Entomol Soc Ann EntomolSoc Environ Entomol Environ Entomol Acta Hort Frankliniella Franklinella J Appl 797: 797: in

This articleis protectedbycopyright. Allrightsreserved. Accepted Article 18. 16. 23. 20. 19. 22. 17. 21.

lycopersicum L. (Pergande) (Thysanoptera:AustJ Thripidae). Entomol thrips, flower ofwestern behaviour dropping and pepper management tactics for Exp Appl determines resistance to Reitz SR Sánchez D,Screenporosity and exclusio GuitiérrezMartinez TinocoC,AquinoEscamirosa GA, Bolanosand Martinez T Mollema C and Cole RA, Low aromatic amino Hortic Sci growth and susceptibility ofchrysanthemum to westernflower thrips. Schuch UK, RdakSchuch UK, RAandBehtkeCultivar, JA, Entomol western flower thrips, thrips, flower western Kigathi RandPoehling films and nets H-M,UV-absorbing affected thedispersal of western flower thrips on roses. roses. on thrips flower western against tactic management fertilization: A Reducing KM, Heinz A and A,Chau Chow 49: pressure forced ventilation to prevent invasion by insect pests. positive-andT, Preliminarygreenhouse N Saitoof evaluation K, Ohishi Sugiyama Steiner MY, Spohr LJand RelatiGoodwin S, 553-559 (2014). (2014). 553-559 .

, J Econ Entomol 78 Yearby EL Yearby 136: 123: : 325–333 (1996). (1996). : 325–333 761-771 (2012). 761-771 (2012). 727-733 (1998). (1998). 727-733 ). Southwest Entomol , Funderburk JE Funderburk

96 Frankliniella occidentalis Frankliniella Frankliniella Frankliniella occidentalis : 1201 J Appl Entomol J Appl – 1214 (2003) 1214 , Stavisky J Stavisky

in thrips (Thysanoptera:Thripidae)field-grown 39: n of pests in greenhousen ofpests tomatoes( 625-634 (2014). (2014). 625-634 fertilizer and irrigation effect vegetative effectvegetative and irrigation fertilizer ve humidity controls pupation success . ,

Momol MT Momol 136

acid concentrationsinproteins leaf (Thysanoptera: Thripidae). Thripidae). (Thysanoptera: : 520-529 (2012). : 520-529 (2012). in four vegetable crops. infourvegetable 50: 179-186 (2011). 179-186 and Frankliniella occidentalis occidentalis Frankliniella Olson SM Olson Appl EntomolZool ,

Integrated Integrated J AmSoc Solanum Solanum Entomol J Appl This articleis protectedbycopyright. Allrightsreserved. Accepted Article 27. 25. 24. 26. 28. 31. 30. 29.

63 occidentalis insecticide-treated and non-treated trapplants for ofthe management Buitenhuis R,Shipp JL,Jandricic S, Murphy of GandShort M,Effectiveness management. impatiens. westernon flower thrips population leveland quality ofsusceptibleand resistant esculentum of populations and status Chen Y, Story R and Samuel-Foo M, Effects ofnitrogenand fertilization phosphorous R andSamuel-FooM,Effects Chen Y,Story StaviskyBrodbeckBV, J,Funderburk Andersen PC JE, andOlsonSM,Flower nitrogen Cook SM, Khan ZR and Pickett JA, The use of push-pull strategies in integrated pestof push-pullstrategies use andPickett JA,The SM,Cook ZR Khan 142: thrips species composition and population abundanceinKenya. Nyasani Jo, Meyhöfer R, Subramanian S and PoehlingH-M, Effectofintercrops on Chem Ecol metabolomics of thrips (Frankliniella occi (Frankliniella ofthrips metabolomics RandKlinkhamerAbdel-Farid IB,Verpoorte PG,NMR KA, Choi YH, Leiss Chem Ecol increased densities of glandular trichomes ontomato, esculentum.Lycopersicon HooverBoughton K,andFelton AJ, Methyl GW, jasmonate application induces Plenum Press,NewT. Yo breeding, in Thrips Biology and Management, ed. Skinnerby Parker BL, MandLewis Zeier P and Wright MG, Thripsresistance : 910 236-246 (2012). (2012). 236-246 – 917 (2007). (2007). 917

35 31 Adv Crop Sci Tech Sci Crop Adv . (Thysanoptera:Thripidae) ornamentals. ingreenhouse Entomol ExpAppl : 219–229 (2009). (2009). : 219–229 : 2211–2216 (2005). (2005). : 2211–2216 Annu Rev Entomol rk, USA. pp. 411–416 (1995). (1995). rk, USA.pp.411–416

2 :145 (2014). :145 (2014). 99 Frankliniella occidentalis

:165–172 (2001). :165–172 (2001). 52 : 375 –400 dentalis) resistance in Senecio resistance hybrids.dentalis) in Gladiolus spp.: potential for IPM and forIPMand potential in spp.: Gladiolus (2006). (2006). feeding on Entomol Exp ApplEntomol Exp Pest ManagSci Frankliniella Frankliniella Lycopersicon J J

This articleis protectedbycopyright. Allrightsreserved. Accepted Article 32. 33. 34. 35. 37. 36. 38. 39.

tomatoes. PG, Metabolomics analysis of host plant resistance tothrips inwild andcultivated Romero-GonzalezMirnezhad M, Leiss RR, Frankliniella occidentalis Frankliniella for anisobutylamideassociated with host-plant resistance towestern flower thrips, R,Tsao Broadbent MarvinCH, AB, FriesenM,Allen WRand McGarvey BDEvidence resistance. metabolomics to identify secondary plantcompounds involved in host plant KA, Choi Leiss YH, Verpoorte R and Klinkhamer PG, An overview of NMR-based biofortified and wild carrots. wild and biofortified plantresistance metabolomic studyofhost eco-Steenis PG, An R,VerpoorteRandKlinkhamer KA,CristoforiG, van Leiss and expressionof aninsect inhibitorproteinase inalfalfa JC, WasmannCC,C,Dunn RL, HJ Thomas and Echt Bohnert McCoy TJ,Introduction (2009). 1567–1575 chlorogenic acidasresistancea factor forthrips inchrysanthemum. KA,Maltese F,ChoiYH, VerpoorteLeiss R and Klinkhamer PG,Identification of Cell Rep Cell Physiol oviposition rate ofthewesternflower thrips, rate oviposition Udayakumar M and Jongsma MA, Effects of cysteine protease inhibitors on Annadana S, PetersJ,Gruden K,SchipperBeekwilder A,Outchkourov NS, MJ, Campus GK, Expression ofCampus GK,Expression potatomultic WJ, and MA Jongsma Kogel Udayakumar M, Visser PB,de Annadana S,KuiperG,

48

14 : 701–706 (2002). (2002). : 701–706 Phytochem Anal Phytochem : 31–36 (1994). : 31–36 (1994). Phytochem Rev , in chrysanthemum. 1

21 0 Phytochemistry : 205–216 (2011). : 205–216(2011). : 110–117 (2009). (2009). : 110–117 ystatin inflorets of chrysanthemum and KA, ChoiKA, YH,Verpoorte Rand Klinkhamer

93 to westernflower thripsincultivated, J Chem Ecol : 63–70 (2013). : 63–70 (2013). Frankliniella occidentalis Frankliniella

31 Medicago sativa : 103–110 (2005). : 103–110 Plant Physiol Plant Physiol . L. J Insect J Insect Plant Plant 50 : This articleis protectedbycopyright. Allrightsreserved. Accepted Article 40. 41. 45. 43. 42. 44.

212 (2012). 212 (2012). Biotechnol J Frankliniella occidentalisFrankliniella MA, Specific cysteine protease inhibitors OutchkourovSchuurman-de NS,de KogelWJ, and Jongsma Abrahamson M Bruin A, Hortic assessment of resistance to western flower thrips, Engineered multidomain cysteine protease Outchkourov GL,WJ, Wiegers AbrahamsonMand Jongsma NS,deKogel M, 448 (2004). interactions among vector insect, thrips andatospovirus. M, Antagonisticplant defense system regulatedby phytohormones assists H,TomitakaAbe Y,Shimoda Seo S,Saku T, (2002). western flowerthrips ( Microbe Interact Microbe transcriptome changesof Arabidopsis during pathogen and insectattack. LoonLC,Dicke van MétrauxJP, Buchala AJ, De Vos Van OostenVR, M,Poecke Pelt RM, Van JA, PozoMJ,Mueller Van glycoprotein. viral German andTL Whitfield AE,Disruption of Montero-Astúa M, Rotenberg D,Leach-Kieffa the systemic wound response of tomato. Li L, Li C, Lee GI and Howe GA, Distinct GIand HoweGA, Li L,C,Lee

572 : 121–129 (2002). (2002). : 121–129

2 : 449–458 (2004). :449–458

18 Mol Plant-MicrobeInteract : 923–937 (2005). (2005). : 923–937 (Pergande), in transgenic potato. Frankliniella occidentalis roles for jasmonateroles synthesisand actionin act as deterrents of western flowerdeterrentsthrips,western act as of rai T, Kugimiya TsudaS S, and Kobayashi vector transmission by aplant-expressed M and PieterseCM,Signal signatureand M and Proc NatlAcad Sci USA 27 ber A,Schneweis BA, ParkS, ParkJK, inhibitorsyield resistance against : 296–304 (2014). : 296–304 (2014). ) in greenhouse trials. trials. ingreenhouse ) Frankliniella occidentalis Plant Biotechnol J Biotechnol Plant Plant CellPhysiol 99 : 6416–6421 : 6416–6421 Mol Plant-

53

2 : 439– : 204– . Plant Acta This articleis protectedbycopyright. Allrightsreserved. Accepted Article 48. 47. 46. 49. 50. 51. 52. 53.

Acaralogia limonicus Knapp M, van Houten HoggerbruggeY, H and Bolckmans K, tobacco. in flue-cured Tospovirus and onofacibenzolar-S-methyl imidacloprid suppression tomato wilt spotted Pappu HR, CsinosAS,McPherson RM, Jones DC andStephenson MG, Effect of plants. tomato in yield and herbivory Thaler JS, 1999.Induced resistance in agricultural crops: effects ofjasmonic acid on greenhouse production systems: afloriculture perspective. Buitenhuis R,Murphy G,ShippL andScott DupreeC, 28 cucumeris flowerwestern thripson sweet pepper using the predatory mites H an Ostilie ML,Hoogerbrugge YM, Houten Van 464 (2015). greenhouse ornamental crops. influence the efficacyofpredatorymites used Scott R, and Buitenhuis L, Shipp LC, Hewitt the behavior and performance of Buitenhuis R, ShippScott- Dupree L,C, Br Acarol predators De Almeida AAandJansenA, Juvenile preyinduce antipredator behavior in : 283–286 (2005). (2005). : 283–286 62: (: ) as a biocontrol agent: review and new findings. reviewandnewfindings. agent: (Acari: Phytoseiidae)asabiocontrol , Exp ApplAcarol 171-180 (2014). (2014). 171-180

Iphiseius degenerans 53: 102-202 (2013). (2013). 102-202 59: 275-282 (2013). (2013). 275-282 Exp Appl Acarol , Crop Prot A. andersoni Amblyseius swirskii Environ Entomol Entomol Environ ommit A and Lee W, Host onommit planteffects A andLee W, 19 Dupree C, Seasonal climatic variations climaticvariations C,Seasonal Dupree forcontrolofwesternflower in thrips : 349–354 (2000). (2000). : 349–354 65: and d Bolckmans controlK, Biological of 435-450 (2015). 435-450 (2015). A. swirskii (Acari: Phytoseiidae) 28 : 30-37 (1999). (1999). : 30-37 Exp ApplAcarol Amblyseius swirskii Amblyseius . IOBC/WPRS Bulletin IOBC/WPRS Amblydromalus Amblydromalus Amblyseius Exp Appl 65: 65: 451- in

This articleis protectedbycopyright. Allrightsreserved. Accepted Article 54. 61. 55. 59. 58. 56. 60. 57.

pepper? effective western flower thrips, Anthocoridae) infield(Thysanoptera:by Thripidae) peppers greenhouse crops. (Kraatz) (Coleoptera:Staphylinidae)as foe? food for Clercq P,Supplemental Vangansbeke D,Nguyen DT,Audenaert J, Verhoeven R, GobinB, TirryLandDe Carney VA,Diamond JC,MurphyGD Entmol temporarily escapefrom predation in the presence ofasecond prey species GJ, Holstein- MaanenR, van Van Messelink agents against canopy- and soil-dwelling stage of of stage and soil-dwelling canopy- against agents Pozzebon and A,Boaria DusoC,Singlean A cyclamen. manyWeintraub S,How PG, PivoniaS andSteinberg Messelink VanMaanen GJ, R,VanSteenpaal 44: thrips and whiteflies by ashared predator: two pests are better than one De Puysseleyr V, Höfte M and De Clercq P, Ovipositing V, HöfteMandDeClercqP, Ovipositing De Puysseleyr Funderburk J, Stavisky J and Olsen S,Predationof Olsen FunderburkJ, StaviskyJand response. tomato resistance against 372-379 (2008). (2008). 372-379 Pest Manag Sci 37: Crop Prot Crop Arthrop Plant-Inter BioControl 1443-1448 (2012). (2012). 1443-1448 .

Environ Entomol Environ

30: IOBC/WPRS Bull IOBC/WPRS Bull 72: 60: 1443-1448 (2011) 1443-1448 466-473 (2016). 466-473 (2016). 341-350 (2015). (2015). 341-350 Frankliniella occidentalisFrankliniella 5: 71-80 (2011). 71-80 (2011).

29 Amblyseius swirskii swirskii Amblyseius Frankliniella occidentalis : 25: 376 and MarshallD,Thepotentialof 37-40 (2002). (2002). 37-40 – . 382 (2000 382

a biological control agent for use in control agentforuse biological a d combined releases of biological control ofbiological releases d combined Saj MW R,andJanssenSabelis A, Prey SEF and Janssen A, Biological control of SEF andcontrolof JanssenBiological A, ). feeding by inducing thewound in the control of thrips: friend or in thecontrolof thrips:friend

Orius laevigatus Orius insidiosus insidiosus Orius Frankliniella occidentalis Frankliniella Orius laevigatus Orius Frankliniella occidentalis Frankliniella management in sweet are needed for Atheta coriaria (Hemiptera: increase Biol Cont Biol Ecol in

This articleis protectedbycopyright. Allrightsreserved. Accepted Article 66. 68. 67. 62. 63. 65. 64.

(2008). predatory mite Messelink G and van Holstein-Saj R,Improving MesselinkHolstein-Saj soil-dwelling thripsG andvan controlbythe Wu S, Gao Y, Xu X, Wang E, Wang Y and Lei Z, Evaluation of WangYandLei XuX, WangE, Y, Wu S,Gao Biocontrol Sci Techn Techn Sci Biocontrol and 29: occidentalis entompathogenic nematodes for control ofwestern flower thrips ( EbssaL, Borgemeister C,PoehlingH-M,Effectiveness ofdifferent species/strainsof conditions. occidentalis nematodes on different soil-dwellingwestern flower of stages thrips, Ebssa L, Borgemeister C, Berndt O and Poehling H-M, Impact of entomopathogenic 83 (2006). ofthrips. parasitoids for hymenopterous AJ,Exploration Loomans feltiae Buitenhuis Rand JL, EfficacyShipp ofentomopathogenic nematode (Thysanoptera: Thripidae) developm 155 (2006). occidentalis Frankliniella Thripinema nicklewoodi 98: Arthurs SandHeinzArthurs KM,Evaluationof thenematodes 145-154 (2004). (2004). 145-154 1480-1485 (2005). (2005). 1480-1485 Neoseilus barkeri (Rhabditida: Steinernematidae) as influenced by

Biocontrol Sci Technol Sci Biocontrol ) at various concentrations,host densities and temperatures. (Thysanoptera : Thripidae),(Thysanoptera : inthe Macrochelesrobustulus 10: for biological control ofthrips on greenhouse cucumbers.

1110-1121 (2014). (2014). 1110-1121 as biological control agents of western flowerwestern thrips of control agents biological as infesting chrysanthemum.

11 : 515–525 (2001). (2001). : 515–525 ental stage andhost plant stage. ental stage (Berlese).

laboratory and under semi-fieldlaboratoryand under Biocontrol Sci Technol Biocontrol IOBC/WPRS Bull IOBC/WPRSBull Frankliniella occidentalis occidentalis Frankliniella Steinernema feltiae Stratiolaelaos scimitus B Insectol B Insectol J EconEntomol Frankliniella Frankliniella Steinernema Steinernema 32: Frankliniella Frankliniella Biol Control Biol

16 135-138 135-138 59 : 141– : 69– and and

This articleis protectedbycopyright. Allrightsreserved. Accepted Article 70. 69. 71. 73. 72. 74. 75.

Ugine TA, Wraight SP andSanderson JP, Ef western flower thrips, thrips, flower western of on efficacy parameters Techn occidentalis with Vuilleman Jacobson RJ, Chandler D, Fenlon J and Russel KM.Compatibility of (Balsamo) crops. Zhang T, Reitz SR, Wang H and Lei Z, Sublethal effects of effects Z,Sublethal T, ReitzZhang SR, Lei H Wang and (Thysanoptera: Thripidae). New opportunities forthe integration ofmi F,Gonzalez Tkaczuk Dinu C,MM,Fiedler Z, Vidal S, Zchori- Fein EandMesselink GJ, fungi. entomopathogenic occidentalis M, Gouli S,FrankSkinner CE, ParkerBLandKim JS, Managementof of tableparameters onlife Hypocreales) (Ascomycota: systems in greenhouse crops. (Thysanoptera: Thripidae). mycorrhizal populations influence fungi oftheherbivore Demirozer O, Tyler-Julian Kand Fundeburk J,Association ofpepper with arbusucular Inter do not affect host accept mycorrhizal fungus Koschier EH, Khaosaad TandVierheilig H,

2:

11: Biocontrol Sci Techn Sci Techn Biocontrol 11-15 (2007). 11-15 (2007). 391-400 (2001) 391-400 Pergande(Thysanoptera: Thripidae) oncucumber plants (Thysanoptera: Thripidae) with granular formulations of Amblyseiolus cucumeris Glomusmosseae Frankliniella occidentalis . anceand development of Biol Control the entomopathogenic fungus the entomopathogenic fungus

J EconEntomol J Entomol Sci J Entomol 17:

J Pest J Pest Sci 193-219 (2007). 193-219 (2007). and enhanced phosphorous levels in cucumber cucumber in levels phosphorous and enhanced 63:

89:

49: 246-252 (2012). 246-252 (2012). 108: (Phytoseiidae) tocontrol (Phytoseiidae) 95-311 (2016). (2016). 95-311 156-165 (2004). 156-165 fects of manipulating sprayapplication ofmanipulating fects croorganisms in Root colonization by the arbuscular bythearbuscular Root colonization 975-985 (2015). 975-985 (2015). , infesting greenhouse impatiens Frankliniella occidentalis Frankliniella Beauvaria bassiana Frankliniella occidentalis Frankliniella Frankliniella occidentalis Frankliniella

Beauvaria bassiana Biocontrol Sci Sci Biocontrol Frankliniella Frankliniella Frankliniella Frankliniella . against J Plant

This articleis protectedbycopyright. Allrightsreserved. Accepted Article 77. 79. 76. 82. 81. 78. 80.

cucumeris chrysanthemums and roses. thrips Manners AG, DembowskiBR Healey and MA, Control barkeri entomopathogenic fungus Chow A, Chau A and Heinz KM,Compatibilityof AandHeinz Chow A,Chau Wu S, Gao Y, SmaggheG, Xu X and LeiZ, Interactions andfitness preference of between prey: Dupreeextraguild C,Intra-guildvspredator ShippL andScott- Buitenhuis R, the Frankliniella occidentalis Frankliniella and agents theirefficacy incontrolling soil BrownbridgeM,Compatibility and Saito T Frankliniella occidentalis Frankliniella entomophatogenic nematodes tocontrol andwesternflower predatorymites thrips Ebssa L, Borgemeister C,Poehling Control predation. intraguild in involved predatorybugs ofgeneralist species with two Messelink GJandJanssen A,Increased cont Thripidae) on roses. on roses. Thripidae) control of for biological Anthocoridae) (Athias-Henriot) (Acari : Phytoseiidae) and : Phytoseiidae) (Acari (Athias-Henriot) Frankliniella occidentalis and biological controlshared prey/host and oftheir 98: 79: (Oudemans) Phytoseiidae) (Acari: (Oudemans) 43-51 (2016). 1-7 (2014). 1-7 (2014). Biol Control . .

Biol Control Biol Control Biol Control Biol Control Beauveria bassiana Aust J Entomol Amblyseius swirskii (Pergande) (Thysantoptera; Thripdae), in gerberas, 53 : 188–196 (2010). : 188–196 92: 39: 92-100 (2016). 92-100 (2016). 66-76 (2006). 66-76 (2006). Frankliniella occidentalis -dwelling stages stages westernflower-dwelling of thrips

52: ofsoil-dwelling predatorsand microbial Bull EntomolRes Bull rol of thrips and aphids in greenhouses in greenhouses aphids and of thrips rol H-M, Simultaneous application of 246-258 (2013). (2013). 246-258 andthepredatorymite Biological control of western flower flower western of control Biological Amblyseius (Athias-Henriot) and and (Athias-Henriot) Orius insidiosus Orius Frankliniella occidentalis Frankliniella ()

100: 167-173 (2010). 167-173 (2010). (Thysanoptera: (Hemiptera: Neoseiulus Neoseiulus Neoseiulus Neoseiulus swirskii . Biol Biol Biol Biol This articleis protectedbycopyright. Allrightsreserved. Accepted Article 83. 84. 87. 85. 86. 88. 89.

Hypoaspis aculeifer Hypoaspis Combined release ofentomopathogenic nematodes and thepredatory mite H-M, andPoehling R-U Ehlers Borgemeister O, Premachandra WTSD, C,Berndt Frankliniella occidentalis Frankliniella pheromone inthewestern flower thrips Hamilton JG, Hall DR and Kirk WDJ, Identification of amale-produced aggregation contact pheromone in the Western flowerthrips WD, A male-predominant cuticular hydrocarbon, 7-methyltricosane,used as is a Olaniran OA, Sudhakar DrijfhoutFP, AV, Du (2003). (2003). 282 (2002). on landingand take-off by adult flower western thrips. R an JG, Jacobson KM,Hamilton MacDonald 1369–1379 (2005). (2005). 1369–1379 Ecol the Western flower thrips MacDonald KM,Hamilton JG, Jacobson RandKi Koschier EH, Kogel WJand de compounds to westernflowerthrips Mediated Linalool MetabolismtheForm and andWerc Leiss K Miesch M,SchneiderB, HeinrichA, C,Gingliner J-F, L, Allouche JE, L,Bassard R,Miesch B,Junker Boachon 2655 (2000). (2000). 2655 39 : 559–568 (2013). (2013). : 559–568

to control soil-dwelling stages of western flower thrips flower ofthrips western to controlsoil-dwelling stages . BioControl BioControl Frankliniella occidentalis Visser JH, Assessing 48: Frankliniella occidentalis Frankliniella 529-541 (2003). (2003). 529-541 k-Reichhart D, CYP76C1 (Cytochrome P450)- (Cytochrome k-Reichhart D,CYP76C1 Vincent B, WahyuniBeran DSC,C, Paetz F, Frankliniella occidentalis Höfer R, Caillieaudeaux R, Seidel DE, Lesot Lesot DE, Seidel R, Höfer R,Caillieaudeaux dKirk WD, Effects ofalarmpheromone blon IA, Hall DR, Hamilton JG and Kirk, JGand DR,Hamilton IA,Hall blon ation of Volatile and Soluble Linalool Linalool and Soluble ation ofVolatile rk WD, ofAnalysis of anal drop -lets Frankliniellaoccidentalis the attractiveness of volatile plant . J Chem Ecol JChemEcol Entomol ExpAppl . J Chem Ecol . J ChemEcol 29 : 2385–2389

103 26 . : 2643– J Chem : 279–

31 :

This articleis protectedbycopyright. Allrightsreserved. Accepted Article 90. 91. 92. 94. 93. 95. 96.

Plant Cell Plant Oxides inArabidopsis Flowers:A Strategy management. NZ Plant Prot NZ management. Recent developmentswith methylisonicotin DA,DavidsonMM,PerryNB, NielsenTeulon MC, anddeKogelWJ van TolRW Evaluation of new volatile compounds as compounds volatile new of Evaluation ButlerRC HallC, Kean AM, J, Poulton DA,Teulon C, Castañé Nielsen MC, El-Sayed AM, Davidson MM,Gardner-Gee R, thrips inNewZealand and Spain.NZPlantProt Egger B and Koschier Egger EH, Behavioural responses of Pergande larvae toand methylPergande larvae jasmonate cis-jasmone. on Peneder S and KoschierEH,Toxicandbeha (2014). (2014). deterrent plant compounds andtheir blends. B B,Spangl and KoschierEH,Egger Habituation in carvacrol as oviposition deterrents for as oviposition deterrents carvacrol E, Allsopp Prinsloo GJ, Smart LEand Dewh One economicallywesternviable? Management flowerof the strawberry. thripsin Sampson CandKirkWD,Can blossoms. Arthropodblossoms. PlantInteract F. occidentalis 8 : e80787 (2013). (2013). : e80787

27 : 2972–2990 (2015). (2015). : 2972–2990 larvae. larvae. J Plant Dis Dis Protect J Plant 64 : 287 (2011). : 287 (2011). mass trappingreducemass thripsdamageandis it 8 : 421–427 (2014). (2014). : 421–427 Frankliniella occidentalis Frankliniella , Sansom CE, Suckling DMand Perry NB, lures for western flower thripsandonion forwestern lures

118 for Defense against forDefense Antagonists. Floral ate, a usedinsemiochemical thripspest vioural effects of carvacrolandthymol irst SY, Methyl salicylate, thymol and SY, Methylsalicylate,thymoland irst 67 : 26–30 (2011). 26–30 : : 175–183 (2014). (2014). : 175–183 Entomol Exp Appl J PestSci Frankliniella occidentalis Frankliniella Frankliniella occidentalisFrankliniella (Pergande) onplum

87 : 53–59 (2014).

151 : 231–238 : 231–238 PLoS PLoS to

This articleis protectedbycopyright. Allrightsreserved. Accepted Article 103. 102. 99. 97. 100. 98. 101.

predators?. Can semiochemicals attract both wester Entomol (Thysanoptera: Thripidae) adultpopulations under greenhouse conditions. forentomopathogenic suppression organisms of Western Flower Thrips occidentalis (2002). (2002). roses. Davidson MM,Nielsen MC, Butler RC, C, Castañé Alomar O,Riudavets J, TeulonDA, potential application of trapping inmass their semiochemicalsoffor Evaluation and RahmanT, S, CousinsDA Broughton Kivett JM, Cloyd RA and Bello NM, Insecticide Rotationwith Insecticide Programs Bello NM, and Kivett JM,CloydRA Jensen SE, Insecticide resistance inthewestern flower thrips autoinoculation system for thrips management. entomopathogenic fungicompatibility toenhancetheir an andinfectivity in separation of semiochemical SandManianiaNK,Spatial Lurem-TRand Ekesi H, Plessis du S, Niassy WJ, Kogel GL,de Wiegers RW, Tol S, van Subramanian K, Mfuti occidentalis western flowerthrips, improvementpossible controlofthe the the of Cook DF, Dadour IR andBailey WJ, Addition occidentalis invasiveness the limits resistance Funderburk J,FrantzG, Mellinger C, Ty Crop Prot

108 . Pergande Pergande (Thysanoptera: Thripidae). ) in Florida. Entomol Exp Appl Entomol Exp : 1936–1946 (2015). (2015). : 1936–1946 Integ Pest Mang Rev Pest Integ Mang

67 : 130–135 (2015). (2015). : 130–135 Insect Sci Insect

155 23:

: 54-63 (2015). : 54-63 (2015). 5: 131-146 (2000). 131-146 (2000). of thewestern flowerthrips ( 175-182 (2016). of alarm pheromone to insecticides and pheromone ofalarm toinsecticides n flowertheirthrips and anthocorid kler-Julian K and Srivastava M, Biotic M, and Srivastava K kler-Julian Frankliniella occidentalis occidentalis Frankliniella Pest Manag Sci Int JPest Manage 72 : 131–139 (2015). :(2015). 131–139

(Pergande) in 48 Frankliniella Frankliniella Frankliniella Frankliniella Frankliniella Frankliniella : 287–290 : 287–290 J Econ This articleis protectedbycopyright. Allrightsreserved. Accepted Article 104. 105. 108. 107. 106. 111. 110. 109.

97 occidentalis and competitorofenemies thrips insecticides against Srivistava M,Funderburk J,Demirozer O, flower thrips,celery leafminer, and two-spotted mite. spider western to chrysanthemum of Cross-resistance KA, PGandLeiss SP, Klinkhamer Kos wilt tospovirus in Boiteux LSandDeAvilla AC,Inheritance of (2011). wiltspotted virus (Bunyaviridae:Tospovirus) in tomato. Riley DG,Joseph SV, Kelly SandSc WT, Olson (2015). 1956 Voorrips RE,QTLmapping of thrips resistance in pepper. RG and G,PelgromPurwito A,Visser Steenhuis-Broers K, Maharijaya A,VosmanB, (2014). 198–208 tospovirus. tospovirus. of transmission Thrips AE, Whitfield and DJ Schneweis AL, Jacobson D, Rotenberg (2014). of the salivary gland transcriptome gland of the of salivary JohnsonStafford-Banks CA,RotenbergD, WhitfieldAE BR, andAnalysis Ullman DE, (2012). AE, Proteomicanalysis of AE, Badillo-Vargas IE, Rotenberg D, Schneweis DJ, Hiromasa Y, Tomich JM and Whitfield proteins in response proteins inresponse to : 337–348 (2014). (2014). : 337–348 Curr Opin Vir Curr Opin (Pergande) (Thysanoptera: Thripidae) in fruiting vegetables. vegetables. fruiting in Thripidae) (Thysanoptera: (Pergande) Capsicum chinense 15: Tomato spotted wilt virus wilt Tomato spotted 80-89 (2015). 80-89 (2015). Frankliniella occidentalis Frankliniella ‘PI 159236’. ‘PI159236’. Frankliniella occidentalis Frankliniella

Olson S andReitzSR,Impactson natural a resistance aresistance specific totomatospotted ott ott J, Host plant resistanceto Tomato Euphitica infection. and differentially expressed expressed differentially and HortScience Theor AppGen

75 : 139–142 (1994). (1994). :139–142 JVirol Entomol Exp Appl . PloS One One PloS

46 86: 86: : 1626–1633 : 1626–1633 Frankliniella Frankliniella 128 Fla Entomol 8739-8809 8739-8809 9:

: 1945– e94447 e94447 151 :

This articleis protectedbycopyright. Allrightsreserved. Accepted Article 113. 115. 112. 114. 116. 118. 117. 119.

Soc. B PJ, Symbiont-mediated and Dyson RNA inBodger OG interference insects. Whitten MM, FaceyPD,DelSolR,Fernández-Martínez LT, Evans MC, MitchelJJ, 46 (2015). plants. Behavioral responses of Muvea AM, Meyhöfer R, Maniania NK,Po tools forthe westernflowertools thrips, Badillo-Vargas IE,RotenbergD,Schnewei phenolics and antiherbivoredefense. independent pathways mediatespecific effect Demkura PV,Abdala G, Baldwin ITandBallaré CL,Jasmonate-dependent and- in edible brassicas. as DC3000a virulencepv. tomato functions factor and defense influences pathways BenderElizabeth SV and CL, Thephytotoxincoronatine from Automated video tracking of thrips behavi MP,Kloth Thoen GL, KJ, Wiegers Krips OE 2015). insect interactions: picturing more resistant crops. Goggin FL,Lorence andA ToppCN, Applyinghigh-throughput phenotyping to plant– wilt virus in primary cell cultures virus wilt Nagata T, Storms MM, Goldbach R and Pe parallel two-choice multiple setups. (1997). (1997).

283 J PestSci : 20160042 (2016). (2016). : 20160042

88 :555–562 (2015). (2015). :555–562 Mol Plant Pathol Mol Plant Thrips tabaci derived from twothrips species.

Frankliniella occidentalis Plant Methods 8 Plant Physiol Plant Physiol : 83-92 (2007). Lindeman to endophyte-inoculated onion ss DJ and Whitfield RNAAE, interference ehling H-M,EkesiS and Subramanian S, , Noldus LP,DickeM and JongsmaMA, ters D, Muliplication oftomato spotted or toassess resistancehost-plant in s of solar ultraviolet B radiation on leaf on Bradiation solarultraviolet of s , 152: 12 Curr Opin Insect Sci Insect Curr Opin : 1 (2016). 1084–1095 (2010). 1084–1095 . J Insect Physiol J Insect Pseudomonas syringae Virus Res

9

49: 49: : 69–76 (

Proc. R. Proc. R. 76 59-66 59-66 : 36– ( Table 1 www.herts.ac.uk This article isprotected bycopyright. Allrights reserved. Accepted Article entomopathogen Parasitoids Predator

Soildweller Crop-dwellers . Biological control agentsof s Fungi Rove beetle Mites Minute bugs WFTstageaf (foliar) Mites agent Typeof Classification Nematodes Parasitic wasp ). Lecanicillium lecaniiLecanicillium Atheta coriaria Hypoaspis aculeifer Macrocheles robustulus laevigatus Orius insidious Orius Amblyseius cucumeris Orius armatus Orius majusculus Orius albidipennis limonicus Amblydromalus montdorensis Amblyseius Amblyseius andersoni Amblyseius swirskii Amblyseius californicus Amblyseius degenerans Amblyseius barkeri Isaria fumosorosea fumosorosea Isaria Beauveria bassiana Metarhizium anisopliae Ceranisus americensis Hypoaspis miles Steinernema feltiae Ceranisus menes F. occidentalis . Information. retrieved from ‘Bio-pesticide Database’of University of Hertfordshire 1st and 2nd instar larvae 2005 Europe Europe 2005 larvae 2nd instar 1st and Adults most susceptible 2012 Europe Europe 2012 most Adults susceptible Adults most susceptible 2012 Europe and America Europe Netherlands 2012 2012 most susceptible Adults most Adults susceptible Pupae, pre-pupae and Larvae and adults 1900s Worldwide Worldwide America North- 1900s 1900s and adults Larvae and adults Larvae Larvae and adults 1993 EU and US EUandUS Australia Europe 2008/2009 1993 1991 and adults Larvae and adults Larvae and adults Larvae Parasitizes larvae 1996 Netherlands Netherlands 1996 Parasitizes larvae Parasitizes larvae 1996 Netherlands Netherlands 1996 Parasitizes larvae 1st instar larvae 1995 Worldwide Worldwide 1995 larvae 1st instar 1st instar larvae 1981 Worldwide Worldwide 1981 larvae 1st instar Larvae 2010-2011 Netherlands 2010-2011 Larvae 2010-2011 Larvae Netherlands 2007 Larvae Netherlands 1985 Larvae Europe 1993 Larvae Worldwide Larvae 2012 2012 Larvae Netherlands Pupae 2008 Europe 2008 Pupae 2002 Canada Pupae 1995 Europe Pupae Pupae 1994 Pupae 1994 Europe larvae 2005 Worldwide 2005 larvae etd is s Commercially available Firstuse fected Crops Target Tradename Typeofcompound Table 2

This article isprotected bycopyright. Allrights reserved. Accepted Article Synthetic origin Natural origin . Overviewofnatural syntheticand compounds used agains

Broad chemical spectrum Selective chemicals ueuo Mth htnboytei Rose, cutflowers rose, Capsicum,cutflowers, lettuce, Nicotinic acetylcholine receptor Conserve Chitinbiosynthesis Match Spinosad Lufenuron Pyridalyl Nocturn Protein Azadirachtin NeemAzal Spruzit/Raptol Pyrethrins Ecdysone Rose Synthesis Sodium recept Ch prttaa Mvno ctlCAcroyae Chrysanthemum Capsicum,Chrysanthemum, rose, Chrysanthemum, rose, cutflowers Acetyl CoA carboxylase Movento Sodium channel Sodium channel Decis EC Spirotetramat Nicotinicacetylcholine re Sumicidin Deltamethrin Actara Esfenvaleraat Methiocarb Mesurol Thiametoxam Acetylcholinesterase (Avermectin, Milbemycin) Abamectin Vertimec Glutamate-gated chloride channel Capsicum, Chrysanthemum, rose, t thrips t commercial based 2015. on sprayadvicecards annel Lettuce, cutflowers, strawberry or Rose, chrysanthemum, cutflowers Chrysanthemum, rose, cutflowers etr Chrysanthemum, rose, cutflowers ceptor cutflowers, lettuce, strawberry lettuce, cutflowers, cucumber, strawberry strawberry cutflowers, lettuce, cucumber,