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Semiochemicals or ladybird the considered. in deterrents by or the as aggregation tainted developed of wine being impact prevent for presently the treatments are to methods lower remedial vineyards Other quality; to wine vineyards in and in production or trapping practices grape various buildings on cultivation using grapes; on e.g. on insecticides buildings, or inside houses of beetles use buildings of entering the aggregates invasive from this remove methods; beetles manage to adult to or prevent methods autumn Current mechanically in grape biodiversity. to a native techniques nuisance, to human include: threat a species become a has and it pest where wine Americas and the and Europe in invasive beetle ladybird, Abstract eateto ilg,Ui fEooyadEouin nvriyo rbug hmnd Muse du Chemin Fribourg, of University Evolution, and Ecology of Unit Biology, of Department Zindel UK R. 2LS, PE28 Huntingdon Wood, Monks Hydrology [email protected] and e-mail: Ecology for Centre NERC Roy E. H. Switzerland Delemont, 2800 Grillons, [email protected] des e-mail: Rue 1, Europe-Switzerland, CABI eateto eeis nvriyo abig,DwigSre,CmrdeC23EH, CB2 [email protected] Cambridge e-mail: Street, Downing Cambridge, UK of England, University Genetics, of Department Majerus N. E. M. [email protected] e-mail: Switzerland Fribourg, 1700 & amnaaxyridis Harmonia ) hspprrvestecretadptnilmtost oto h harlequin the control to methods potential and current the reviews paper This Æ .Zindel R. ee .Roy E. Helen Æ Pla)(oepea ocnlia) nAinpredatory Asian an Coccinellidae), (Coleoptera: (Pallas) eaeZindel Renate 1 Æ ihe .N Majerus N. E. 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Russia, the and of China dynamics population i.e. enemies the range, natural in that distribution importance suggest little observations its America, of of North are and part Europe largest In unknown. the largely in enemies the natural on of available dynamics information population little the is in There enemies mite. natural parasitic of a importance and nematodes parasitoids, tal. et in,priual nisrgo foii.Rsac nmraiyfcosi edconditions field in factors (Osawa mortality Japan on in Research out origin. carried of been region has its in particularly tions, ucpil odsso 10 of doses to susceptible .bassiana B. 10 ulta feton effect sublethal a rtdon trated 2007 tal. et insects. nraeteripc ntetre et l hs ehd ol edvlpdand developed be could methods these All pest. target to natural practices, the cultural controlling strategy, various on for control’’ through or employed e.g. impact biological inundative favoured, their be ‘‘conservation regular can increase site a through on present increased in a already enemies be establishment, Finally, permanent can releases. for enemies introduction inoculative biological natural of ‘‘augmentative an region programme, in the Second, control’’ control’’. to biological ‘‘classical pest called the often method of origin of region .septempunctata C. uto infiaty(o tal. et (Roy significantly duction ( Ctrl n Shapiro-Ilan and (Cottrell n Krischik and samcisciieaantvrosps net.Ssetblt fvrosladybirds various of Susceptibility insects. pest vergens to various Coccinellidae) against (Coleoptera: mycoinsecticide a as Mlat Ctrl n Shapiro-Ilan and (Cottrell (Mulsant) mrc n fteoewneighbtt of Shapiro-Ilan habitats and overwintering (Cottrell the of one America .axyridis H. nti vritrn oainadaecmol eoddifce with infected recorded commonly are and location overwintering this in 2003 9 aua nme eotd rtse against tested or reported, enemies Natural w yoraenfnihv enfudifcigladybirds: infecting found been have fungi Hypocrealean Two prsml spores .Ms eerho h ietitrcin ewe ug n ayid a concen- has ladybirds and fungi between interactions direct the on research Most ). 1996 2002 eosrtdi aoaoysuisthat studies laboratory in demonstrated ) .bassiana B. Gue , and ) 2000 2006 to neto.Frhrsuisuigvrosioae of isolates various using studies Further infection. ´ - 2000 rin-Me 1 .bassiana B. .bassiana B. aelmtdipc on impact limited have ), n aecnieal oeta sbooia oto gnso pest of agents control biological as potential considerable have and ) .I otat ubro tde aedmntae h eiineof resilience the demonstrated have studies of number a contrast, In ). eagu parcesetosum Serangium Rye al. et (Roy ´ 2007 eil Jmse al. et (James neville .axyridis H. eas fteptnilnntre mato hsfnu hnused when fungus this of impact non-target potential the of because hwdthat showed ) .axyridis H. 2003 2003 Ctrl n Shapiro-Ilan and (Cottrell Jmse al. et (James 7 2007 prsml spores .Aut ftentv ladybird native the of Adults ). ;whereas ); .bassiana B. 2007 n vnlwdss(10 doses low even and ), 2003 letntwt h aeagents. same the with not albeit , ). oemglamclt lengi maculata Coleomegilla 1992 .axyridis H. .bipunctata A. - and ) 1 1995 , .axyridis H. 1993 iad(orwk tal. et (Poprawski Sicard 7 u vndssof doses even but 1995 a endemonstrated: been has .axyridis H. rpoamsmontrouzieri Cryptolaemus .axyridis H. oteladShapiro-Ilan and Cottrell ; n ot oe Pr tal. et (Park Korea South and ) ), However, . .axyridis H. .bipunctata A. and 2003 r o.Ctrl n Shapiro-Ilan and Cottrell not. are sudrbr npcnorchards pecan in bark under is 5 opieptoes predators, pathogens, comprise .septempunctata C. prsml spores o tal. et Roy ; .bassiana B. .v-nigrum O. .axyridis H. seteeyrssatto resistant extremely is .bassiana B. .anisopliae M. .bassiana B. ibrae(Todorova Timberlake 1998 Rye al. et (Roy - 1 ipdmacon- Hippodamia eueegpro- egg reduce ) 2007 ), 2003 usn (Smith Mulsant per ohave to appears nfil condi- field in r lofound also are lav-nigrum Olla upr this support .bassiana B. .I North In ). o tal. et Roy ; r highly are 1996 exceeding (Ginsberg 2007 ,but ), ), http://doc.rero.ch biaeetprst htifcsmn ocnliswrdie(aeaadWeir and (Thaxter (Nalepa Coccinellidae) worldwide coccinellids many virescens Hesperomyces infects that ectoparasite obligate virescens omyces eetgie yifce eae sarsl ftedahadcnupino hi male their of consumption and death the of result al. a et as (Hurst females siblings infected by gained ( benefit equilibrium male-killer an the to of efficiency population transmission Spiroplasma the vertical through the naturally by spread dictated prevalence would male-killer The eggs. the depress Williams ob eeca nrdcn otpplto ie n osqetybt intraguild both consequently and size, population host reducing in beneficial be to aiernei aibe u a ec 0.Teml-ilrhsntbe eotdfrom reported been not of has populations male-killer into females The infected 80%. killer reach can axyridis but H. variable, is range native 1967 ec tal. et Welch ospseiro eae;asxa iopimta eet h ao transmission major the Beakes and reflects (Weir mating that during contact direct dimorphism via be sexual to thought the a is and which males mechanism of females; ventroposterior the of on reported dorsoposterior often is fungus This observation). personal and neatosbetween interactions grgtos a ugltal itiue ttemrisadfotage fteelytra the of angles front and margins the Beakes at and distributed A thalli (Weir contact body. fungal the direct of had with for part aggregations, accordance anterior reported the in was on winter pattern distributed notably similar behaviour, were of orientation thalli and the beginning fungal aggregation exhibit through contrast, not the did In ladybirds at theory. overwintering sion These 22–38% of winter. distribution of of elytral end characteristic the levels by 62% infection to increasing demonstrated they Carolina h otbd ypouto facrua prsoim hc tahsadpntae the penetrates and attaches which appressorium, Beakes circular and a (Weir of cuticle host production by body host the virescens H. role. of significant a transmission plays for also mechanism winter important in most aggregation the is virescens conspecifics H. with contact through u otepeec faml-iln atru ftegenus the of bacterium male-killing a of presence the to due investigation. further requires this but mating and foraging h oao hs edml gsaalbea odsuc oifce eaesibling Hurst female and infected (Majerus to advantage source fitness food significant a a as with available them 1997 eggs makes providing male male-killer the larvae, dead so, neonate these doing In of embryogenesis. soma in early the hosts male kills it consequence, 1998 upce ohv eaieefc on effect negative al. a et (Applebaum have to suspected nw eeeiu mat ntebel Nlp n Weir and (Nalepa beetle the on impacts deleterious known hr soefrhrfna nooahgno ocnliswrh fnote: of worthy coccinellids of entomopathogen fungal further one is There aoleilscaatrsial ontpntaeteisc uil,however, cuticle, insect the penetrate not do characteristically Laboulbeniales ti loo oeta oeAinppltosof populations Asian some that note of also is It plbu tal. et Applebaum ; 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L)(oepea ocnlia)(abrve al. et (Kamburov Coccinellidae) (Coleoptera: (L.) 1967 2005 Spiroplasma .axyridis H. .bipustulatus C. 8 Wlhe al. et (Welch .Hayifcin ol loipd flight, impede also could infections Heavy ). .Nlp n er( Weir and Nalepa ). hc,we olce rmoverwintering from collected when which, 2007 shpteie ytesxa transmis- sexual the by hypothesised as .axyridis H. ndti.I edsre nNorth in survey field a In detail. in hlcrsstigma Chilocorus ocueta iettransmission direct that conclude ) nErp n ot mrc would America North and Europe in Lws Clotr:Coccinellidae) (Coleoptera: (Lewis) nppltosof populations in 2007 hog nrae mortality increased through , 2001 Spiroplasma .However, ). r togyfml biased, female strongly are ), .axyridis H. 2007 Sy (Coleoptera: (Say) aesuidthe studied have ) .axyridis H. Mjrse al. et (Majerus .virescens H. [ .9frthis for 0.99 (Garce Hesper- 2007 ´ 1996 and s nits in is ). ; http://doc.rero.ch hscladceia eecsaantpeainta otldbrs(e ele al. et Pell (see ladybirds most than predation against defences al. chemical et and physical (Pell coccinellids other by upon preyed h ulta fet fdsaerqiefrhrrsac n ol rvd ouinto solution a provide could out. and ruled research be further not strategy require should novel strategy disease a controlling control of complex of effects a development sublethal of the The component a Nevertheless, as insects. pathogens other utilising and coccinellids target aostak oisakli-ihdfnescein n t psmtccolouration. aposematic its and secretions defence ( alkaloid-rich Kuznedsov and its Nechayev Nevertheless, to thanks dators ladybirds, other As Predators control. to key the associated be parasites) could other these and of (nematodes One protists, ‘‘hitchhikers’’ (fungi, it. other pathogens with and of diversity viruses) the and assess bacteria thoroughly to distribution non-native and rcie smale As practice. .bassiana B. eae sulkl ola ordcdfriiyi eae.Mroe,as Moreover, females. in fertility reduced to lead to unlikely is females nteautpplto.Fnly sfml ayid a osdrbymr hnmales than more considerably eat ladybirds female reduction a as to not Finally, embryo- and during population. immatures, (Hodek of progeny adult cannibalism male in the of reduction a in proportion to significant lead simply a may of genesis loss cannibalistic, highly are rdtrwl vrpa infiatrl ntentrlcnrlof control natural the in role significant a play ever will predator o eiw.I sas esatce yat n pdr hnohrlddrs(uce tal. et (Dutcher ladydirds other than spiders and ants by attacked less 1999 also is It review). al. for et Clercq (De bugs predatory ocnlis including coccinellids, on feed canus .axyridis H. optto n rdto moe by imposed predation and competition utbebooia oto gnsfor agents conducted. control been has biological action a suitable an of such use of uncertainties, consequences these the to of Due analysis females. full reduce of to favour attempt an in in ratio male-killer sex population the biasing aenvrbe eotda en infiatsuc fmraiy h ahndfly tachinid The mortality. of source al. significant et (Nalepa a Carolina North being as triangulifera reported Strongygaster been on never found were have species parasitoid two Only Parasitoids Americas. the ewe ie n rmya oya u,o vrg,ol –%o h ete were beetles the of 2–4% only average, ( on Aliniazee but, and year Katsoyannos to Oregon, year In from parasitized. and sites between Aliniazee urn eerhidctsta,udrcretkolde ahgn r nieyt be to unlikely are pathogens knowledge, current under that, indicates research Current auaadKimura and Yasuda ; mln(iioms iia)and Picidae) (Piciformes: Gmelin 1996 .axyridis H. 1998 .axyridis H. akefiayadseict towards specificity and efficacy lack ) nRsi.Atog unto ( Kuznetsov Although Russia. in ,telvlo nrgidcmeiinmyatal eicesda eutof result a as increased be actually may competition intraguild of level the ), .Alwr erdfo vritrn ete.Prsts a eyvariable very was Parasitism beetles. overwintering from reared were All ). .axyridis H. .axyridis H. nms vritrn grgtos ecnldsta rdto of predation that concludes he aggregations, overwintering mass in Furthermore, . .axyridis H. 2001 Le)(itr:Tciia)wsfudi dl ete in beetles adult in found was Tachinidae) (Diptera: (Loew) 1996 a aewt ayfmls isi h e ai towards ratio sex the in bias a females, many with mate can scniee ob agl rtce rmgnrls pre- generalist from protected largely be to considered is .I svr nieyta n netbaeo vertebrate or invertebrate any that unlikely very is It ). ybrsi eeal ae gsadlra a lobe also may larvae and Eggs rare. generally is birds by , .axyridis H. aeaadKidd and Nalepa ; 2003 .axyridis H. .axyridis H. .axyridis H. ,atog,i general, in although, ), it europaea Sitta 1973 .axyridis H. 9 1997 ouain hudntb netknutla until undertaken be not should populations eto ih pce fbrsfeigon feeding birds of species eight mention ) 2007 hudb cendtruhu t native its throughout screened be should eot httetobr species bird two the that reports ) oee,ti a o etecs in case the be not may this However, . h ug xmnds a (mainly far so examined fungi The . .axyridis H. 2002 aeadMajerus and Ware ; 1998 nisitouto ag n these and range introduction its in .(asrfre:Stia)will Sittidae) (Passeriformes: L. n rgn(asyno and (Katsoyannos Oregon and ) bevdprsts ae of rates parasitism observed ) n ol mato non- on impact could and .axyridis H. .axyridis H. .axyridis H. 2007 nErp or Europe in a better has n by and ) larvae 2007 Picus , http://doc.rero.ch srcre rmmn ayid ftesbaiyCcielnebtsosstrong shows but attacks mainly Coccinellinae it UK subfamily the in the example, of For species. punctata ladybirds large some many for preferences from recorded is ot mrc ti ototnrcre from recorded often most al. et is Obrycki it America South quadripunctata efrac of performance .–54 ydseto n .%b rearing. by 4.8% and dissection by 6.7–15.4% aaiimwti h empea eitr,Ototr n eiotr Ree and of (Reeves reports Lepidoptera scattered and are Orthoptera most there is Hemiptera, and It Coleoptera Dermaptera, O polyphagous. adult the very of and within parasitoid America parasitism a North as in distributed reported widely frequently is that fly parasitic aaiieaut,atog aveadppecnas eatce,i hc aethe case which in attacked, be al. also et (Obrycki can beetle usually pupae adult Females the and Americas. in the continues larvae development and although Asia Europe, adults, in parasitise ladybirds of endoparasitoid common dnie as identified species, ubro eaoye rdcdb h aaiodin parasitoid the by produced teratocytes of number el etmucaabrucki septempunctata nella nAsia, In ilje al. et Firlej cainlyosre on observed occasionally oebri oe Pr tal. et (Park Korea in November luctuosa hra ayCcielde u o yet not but Coccinellidae, questionable many is whereas parasitoid this Europe, of identity in true because, The oviposition. and by development parasitism egg vented rearing, laboratory In 21.1%. tal. et ucsflprsts shge nAi oprdt uoe(Liu Europe to compared Asia in higher is parasitism successful 1969 hi mat ilg n clg r cre h ahndfly tachinid The scarce. are ecology and biology impact, their oerssatto resistant more to takdbtn aaioddvlpdi hs dls ilj( Firlej adults. these in developed parasitoid no but attacked inellae in .septempunctata C. ( Fre tal. et (Firlej agnlhs for host marginal eaewssatc h w ot edl,btlra eefudi 0 fteattacked the of 90% in found were larvae but readily, hosts maculata two C. the attack wasps female in avebigal ocmlt hi eeomn in development their complete to able being larvae aaiimrtsb iscino edcletdbels ayn rm9 o2% u the But 24%. to 9% in from higher varying much beetles, was field-collected rate of emergence dissection by rates parasitism 2007 0 Hara .axyridis H. .septempunctata C. h rcndwasp braconid The te aaiod of parasitoids Other .axyridis H. ake al. et Park ; aesonta hl h takrtsof rates attack the while that shown have ) 1996 a ucsflol hnteldbr avewr aaiie,weesaut were adults whereas parasitised, were larvae ladybird the when only successful was 2004 eiaseparata Medina , .coccinellae D. Mie)(itr:Tciia)wsfudi dl ete olce nOctober– in collected beetles adult in found was Tachinidae) (Diptera: (Meigen) ,sgetn ihrta sa ouain of populations Asian that either suggesting ), ocnlaundecimpunctata Coccinella 2005 .luctuosa M. ). n nol %o the of 7% only in and 2005 hnErpa ouain,o htivsv ouain of populations invasive that or populations, European than 1985 n h aieladybird native the and otpia Clotr:Ccielde (Majerus Coccinellidae) (Coleoptera: Pontoppidan .coccinellae D. ,Erp Kym n Majerus and (Koyama Europe ), .coccinellae D. .coccinellae. D. n,i h K(oaaadMajerus and (Koyama UK the in and, ) 1996 r iia,scesu eeomn in development successful similar, are et ih n Costa and Filho Berti ; .luctuosa M. ilj( Firlej . ioapscoccinellae Dinocampus .axyridis H. .I uoeadNrhAmerica, North and Europe In ). sls omnyrae from reared commonly less is Tcosi n Herting and (Tschorsnig .axyridis H. Mie)(itr:Tciia) hc a oeie wrongly sometimes was which Tachinidae), (Diptera: (Meigen) usn Clotr:Ccielde (Maeta Coccinellidae) (Coleoptera: Mulsant 2006 . in sseict dl hyoeiso h genus the of chrysomelids adult to specific is ognor n eme ( Heimpel and Hoogendoorn 1996 bevdta,in that, observed ) .axyridis H. r nw nyfo hi rao rgn u eal on details but origin, of area their from only known are .axyridis H. .maculata C. nNrhAeia(ognor n Heimpel and (Hoogendoorn America North in .Prsts ae utae ewe .%and 0.7% between fluctuated rates Parasitism ). .(oepea ocnlia)and Coccinellidae) (Coleoptera: L. .maculata C. .axyridis H. 10 1995 .luctuosa M. iia bevtoswr aei Canada in made were observations Similar . Shak Hmnpea rcnde sa is Braconidae) (Hymenoptera: (Schrank) a ea es atydet h lower the to due partly least at be may ognor n Heimpel and Hoogendoorn ; hnin than 1994 oemglamaculata Coleomegilla toggse triangulifera Strongygaster 2007 .axyridis H. r aaiie yaclosely-related a by parasitized are , .coccinellae D. .axyridis H. .Ageaiu ua aaiod the parasitoid, pupal gregarious A ). nteUA hyfudsimilar found They USA. the in .axyridis H. .axyridis H. n atAi (Liu Asia East and ) .axyridis H. .coccinellae D. hree h iesa n pre- and span life the shortened 1985 .axyridis H. .axyridis H. 2006 2007 2002 oprdt utbehosts. suitable to compared ). ugse httepoor the that suggested ) ioapscoccinellae Dinocampus .Kym n Majerus and Koyama ). aaiimby parasitism , 1950 nlbrtr assays, laboratory In . hnfo,e.g. from, than oprdparasitism compared ) adyayparasitoid any hardly , Medina 1997 on Maeta ; smc oe than lower much is scniee sa as considered is r etradapted better are 1969 .I ot and North In ). .axyridis H. .axyridis H. DGe)(e.g. (DeGeer) 1950 (= .However, ). 2002 1969 .septem- C. sasmall a is Harmonia Degeeria .cocc- D. Haltica Maeta ; Cocci- .I is It ). Park ; 2002 and are ) ; , http://doc.rero.ch eaoe(ae ecie as described (later nematode hrdfly phorid aa (Disney Japan oorkolde eaoe aenvrbe eoddfo idppltosof populations wild from recorded (Iperti populations been ladybird never infest to have nematodes axyridis H. knowledge, our To cmt Dpea hrde,prstz eea ocnli pce (Hodek species coccinellid Kearns of several and range parasitize host Phoridae), The (Diptera: pupa. Schmitz Europe, host in single a but, from unknown emerging adults 1–8 Korea, Tlnhd:Alnoeaia)i priadvnWeeee( and particularly Waerebeke ladybirds, Coccinellidae) several van (Coleoptera: of and adults of Iperti intestine in Allantonematidae) (Tylenchida: n ake l ( al. et Park and from reported is Phoridae) .coccinellae D. axyridis H. of control neto a o ehl u togyrdcdegmtrto (Iperti maturation egg reduced strongly but lethal, not was infection in rates Parasitism h aiernehsnvrbe netgtdfrprsts.Teprsti ope of seven complex least parasitoid at example, from The For parasitism. known species. of three part for are largest than investigated parasitoids the more whereas been comprises Korea, often never and ladybirds Japan has indigenous in studies range of native couple of are complex a the that parasitoid on It the based genera Coccinellidae. on is to family knowledge range the belong our native within that they least however, but noted, at investigated be species, par- must be polyphagous the rather to comprise of remains to specificity Asia known in The only programmes. found control asitoids biological augmentative or conservation Hodek ouain n eeeyafc hi mata pi rcci rdtr (Iperti predators coccid or aphid as impact their affect severely and populations 2007 r omnlra aaiod fCcielde(lunte n Klausnitzer and (Klausnitzer Coccinellidae of parasitoids larval common are Homalotylus iod ftegenera the of sitoids 1997 oyhgu tfml rsbfml ee KasizradKlausnitzer rather and often to specific but (Klausnitzer be sub-families, level may coccinellid sub-family Europe, to or be par- may or ladybird family exception Most Coccinellidae to America. at to specific or totally polyphagous Europe specific is into are that introduction to for found remains asitoids pre-requirement be It Siberia. a may and be parasitoid China would as a such whether surveyed, seen been rarely be or never have which regions (Hodek Clotr:Ccielde nRsi (Noyes Russia in Coccinellidae) (Coleoptera: Nematodes rmti uvy tapasta h oeta fuigprstisfrtebiological the for parasitoids using of potential the that appears it survey, this From .Lra aaiimb nytdeo uohdemykl pt 09%o ladybird of 90–95% to up kill may Eulophidae or Encyrtidae by parasitism Larval ). .Lra aaiimhsnvrbe eoddon recorded been never has parasitism Larval ). 1973 1973 hlcoohr philaxyridis Phalacrotophora aaii eaoe ftefmle latnmtdeadMriia r able are Mermitidae and Allantonematidae families the of nematodes Parasitic . .axyridis H. nErp n ot mrc r fmria motnead ntecs of case the in and, importance marginal of are America North and Europe in 1989 .Tu,i ol ewrhhl netgtn aaiimin parasitism investigating worthwhile be would it Thus, ). ,atog thsbe eetyrpre from reported recently been has it although ), Hmnpea nytde and Encyrtidae) (Hymenoptera: o elaatdto adapted well not , 1997 1996 ). oaoyu platynaspidis Homalotylus .Acneei species, congeneric A ). .quatuordecimpunctata P. Aprostocetus iea cite ) nisae fitouto slmtd h aaiod on on found parasitoids The limited. is introduction of area its in .fasciata P. .axyridis H. ltnsi luteorubra Platynaspis .septempunctata C. Phalacrotophora aaiyecu coccinellinae Parasitylenchus , eoi conglobata Oenopia Oomyzus .axyridis. H. 1964 n nte species, another and ua nteRsinFrEs yKzesv( Kuznetsov by East Far Russian the in pupae iny(itr:Poia)wsdsrbdfrom described was Phoridae) (Diptera: Disney .Freape pri( Iperti example, For ). 11 and oe Hmnpea nytde,wih in which, Encyrtidae), (Hymenoptera: Hofer hlcoohr fasciata Phalacrotophora Pachyneuron 2007 ece pt 6 neryatm.The autumn. early in 46% to up reached hs hr sltl rsetfrteruein use their for prospect little is there Thus, p aaiiig0467 fteppein pupae the of 0.4–6.7% parasitizing sp. Tetrastichus nErp KasizradKlausnitzer and (Klausnitzer Europe in ). rplaquatuordecimpunctata Propylea Gee Clotr:Coccinellidae) (Coleoptera: (Goeze) .axyridis H. L)(oepea Coccinellidae) (Coleoptera: (L.) hlcoohr berolinensis Phalacrotophora Hmnpea Pteromalidae) (Hymenoptera: cmu subvillosus Scymnus Hmnpea Eulophidae), (Hymenoptera: pri&VnWaerebeke Van & Iperti lhuhsvrlpara- several although , 1968 1964 1964 .axyridis H. )prstzn the parasitizing )) on parasitic a found ) (Falle .philaxyridis P. .axyridis H. .Commercially ). 1973 .axyridis H. ´ 1997 )(Diptera: n) 1997 Majerus ; sthis as , (Goeze) Noyes ; .An ). 1997 1964 nits in (L.) in is ) ; . http://doc.rero.ch et nwhich in tests moe on imposes axyridis H. eetse on tested were and Heterorhabditidae) ol erqie eoeitgaigti ilgclcnrlaetit management into agent control biological this assessment risk integrating appropriate an before mite, the required of range be host wide would the given However, there. play vial nooahgncnmtds( nematodes entomopathogenic available eotta ie a ucsflytase ewe pce uigitrpcfi hybrid interspecific during species between that transfer proven is successfully al. it If can et matings. (Pascoe mites Europe in that field report the in recorded been has loocr nNrhAeia(cailadMorrill and (McDaniel America North propagation. in further infested occurs for artificially populations also field naturally, to occur inoculated be not could does ladybirds European from bu aual,fritrseichbi aig between matings hybrid inter-specific for naturally, about .axyridis H. quadripunctata whether monia is known mite not the and once currently prevalence transmission high is to horizontal What leading mite, both established. the of facilitate has generations to the mul- down reproducing, generations) transmission before dormancy over-lapping for requirement tivoltinism, no promiscuity, (high characteristics ocnlia)and Coccinellidae) S.Te eosrtdta while that demonstrated They USA. s sabooia oto gn of agent control biological a as use h eaiu n clg of ecology and behaviour the vr9%o oeppltosaeifce ylt umr(ebre tal. et (Webberley voltinism has (Webberley and coccinellids, promiscuity summer European to al. four late correlated et on positively by mite, is the prevalence infected of that indicated levels are prevalence on populations of prevalence analysis some highest Comparative shows of but 90% coccinellids, of over variety female a males on on found found in been have particularly mite al. mortality, the et of (Hurst overwintering effects zero increasing to eventually hosts, and eggs al. adults, female et mite (Webberley with covered and is elytra a male the Within al. of adults. et surface into (Webberley ventral metamorphosing larvae the haemolymph, of on much feed weeks, and few elytra host the of surface al. et podamiae eulytasitdetprst fautccield nErp MDne n Morrill and (McDaniel Europe in coccinellids adult 1969 of ectoparasite transmitted sexually mite The two than mite Parasitic resistant more both were they nematodes to ( susceptible species are native USA) in species u otehg eaieefcsthat effects negative high the to Due ebre tal. et Webberley ; 2006 1995 asbtenml n eaehssdrn ouain nete ieto (Hurst direction either in copulation, during hosts female and male between pass ocplpshippodamiae Coccipolipus ). .Oc nanwhs,lra ie me hi otprsit h ventral the into mouthparts their embed mites larval host, new a on Once ). .bipunctata A. 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promising cultural based most methods the new negative environment, their of the prevent and to chemical development of avoided health use be human the preferably on Besides should damage. impact but fruit efficient and developed are houses be which in to insecticides, aggregations continue will control methods or control prevent Suitable to effects. non-target avoid to specific is n eerht aehsbe eylmtd utemr,a otohrldbrsare ladybirds other most strate- requires as against management Furthermore, beetle, methods new limited. control predatory base very insects, other to been beneficial has which any date considered on or to experience ladybird, research previous predatory and no gies, a is time there first Thus, several for control. the challenge, is a remains It ladybird harlequin reasons. the against methods control Developing Conclusions ratio. sex 1:1 a with population a as in such STD, than an male-killer, a of presence ilee ii h mato outbreak methods these of of any impact that the unlikely very limit is it ever vineyards, will or orchards buildings, in problems oemtn atesads oeopruiyt ohcnrc n ops ntedisease the on pass to and contract have will both scarcity, to their opportunity to more due (Majerus so males, and because partners biased female mating are more ratios sex which in lations as as considered such promiscuous, be may continents these on congeneric occur other that although considered, ecologies be alternatives. similar not with should Africa) mites America, (South reported nyi hr r la in htntv ayid,o te piohgu net,are insects, aphidophagous other or ladybirds, native that signs clear are there if only can- a control addition, to In to challenging. introduced the specific be be be of may should to part bad agent needs control turned a biological agent that and didate agent control authorities control biological public, biological a another the that convincing community Thus scientific practices. control of invasion biological the because partly ouaindniisi utial a steipraino aua nme rmthe from enemies natural of importation the invasive is However, lower origin. eventually way may of sustainable that region a method in only the densities control, population provides and invader new the rgams nitouto ocniet rmwhich from continents to introduction An programmes. .virescens H. oee,teeaemn fetv oe ytm sn eiceia ehd for for strategies methods developing in semiochemical informative using be to systems prove model could controlling which effective insects, many pest controlling are there However, gis h iko on ohn rters eae otaiinlcnrlmaue.I the In measures. weighted control be traditional to to bio- related has classical risk agent all the exotic in or of As an nothing doing negligible. case introducing of not of risk but risk the low against the is programmes, Asia control in logical parasitoid or pathogen specific nesantv uoeno mrcnntrleey(e.g. enemy natural American or European native a Unless htas novsteml-ilrof male-killer the involves also that hl oto ehd rsnl sdo ne eeomn a vnulyslethe solve eventually may development under or used presently methods control While h fcc fsxal rnmte iess(T)sc as such (STD) diseases transmitted sexually of efficacy The .axyridis H. 2003 .axyridis H. scnrligaet for agents controlling as .Tu,i ouainof population a in Thus, ). h nrdcino ilgclcnrlaetsol econsidered be should agent control biological a of introduction the , .axyridis H. . .axyridis H. .axyridis H. h aeo pedo nSDwl egetri popu- in greater be will STD an of spread of rate the , .axyridis H. .axyridis H. sadfcl agtfrcasclbooia control, biological classical for target difficult a is .axyridis H. .hippodamiae C. 13 .axyridis H. s nisl,ms rbbytersl fbad of result the probably most itself, in is, nseisi hc ohsxsaehighly are sexes both which in species In . ih eipoe na P strategy IPM an in improved be might .axyridis H. ouain nntv biodiversity. native on populations .axyridis H. hti eaebae u othe to due biased female is that .hippodamiae C. ol ec ihrprevalence higher reach would , .hippodamiae C. .axyridis H. h hneo niga finding of chance The . ilhv ob highly be to have will .hippodamiae C. ssillimited. still is , dpsisl to itself adapts ) a o been not has and http://doc.rero.ch obas irsadape (Koch apples and citrus biodiversity. introduction, soybeans, native for for dramatic considered where become be of regions identification could invaded the the that to allow in not analysis, agents also situation do would the table control which should studies life biological and These through range introduction. classical native e.g. of potential the regions out, in the carried densities in be population occur limit should factors introduction which of understand areas the and seset fmraiyfcosafcigppltosof populations affecting factors mortality of assessments diesT rnur ,Me 20)TemlioordAinladybird Asian multicoloured The (2003) D Maes E, Branquart T, Adriaens References (GOCE-CT-2003-506675). ALARM project FP6 EU the and (F232-0377) Environment the for Office Acknowledgements the of present, impact exact At the on range. data distribution quantitative and their reliable more of axyiridis gather H. part to is significant research for a priority over threatened seriously diesT a atnyGmzG asD(07 nainhsoy aia rfrne n hnlg of phenology and preferences habitat history, Invasion (2007) D Maes G, Gomez y Martin San T, Adriaens plbu ,KrR esnU amrU(91 tde ntesme eln of decline summer the on Studies (1971) U Tadmor U, Gerson R, Kfir S, Applebaum ( ladybeetle Asian Multicolored and alert. foraging on pest feces Viticulture heterospecific (2005) and Anonymous conspecific of Effect (2003) Y Kajita H, Yasuda BK, Agarwala eeso pi icnrlad hs raeacnito neetbtenteagricultural the between interest of conflict a create conservationists. thus, and and, sector biocontrol aphid of levels aiciJ aanM,Febr ,Cee ,Rcmn 20)Mliooe sa ayid( ladybird Asian Multicolored (2004) D Richmond B, Cheves J, Freeborn ME, Dabaan J, Baniecki ol rvd rawd eutosin reductions wide area provide would et ih ,CsaV (1995) VA Costa E, Filho Berti rw M,Aran ,Bto ,Cpe ,GlaaeaA Ha A, Goldarazena J, Cuppen H, Bathon T, Adriaens PMJ, Brown otelT,SaioIa I(03 ucpiiiyo aieada xtcld ete(Coleoptera: beetle lady exotic an and native a of Susceptibility (2003) Annu DI management. pest Shapiro-Ilan integrated in TE, strategies push-pull Cottrell of use The (2007) JA Pickett ZR, Khan KR, Day SM, F, Cook Lieutier In: landscape. olfactory complex a in beetles bark of ecology Chemical (2004) JA Byers un A conl J epin -,Lga ,DxnAG(98 aercgiini h two-spot the in recognition Mate (1998) AFG Dixon G, Lognay J-L, Hemptinne JJ, McDonnell DA, Burns ial,i utb oe ht nNrhAeia hr r aiu giutrlcontexts agricultural various are there America, North in that, noted be must it Finally, h naieladybird 133:201–287 Zool J invasive Belg Belgium? in the predators aphid native for threat a Coccinellidae), : (Coleoptera vpsto ftopeaoyldbrs oeo ea usi rdto viac.JCe cl29: Ecol 2007 Chem Aug J avoidance. University. predation State Missoury in cues fecal of role ladybirds: 357–376 predatory two of oviposition s10526-007-9137-6 axyridis stulatus ,LoasAM aeu E,Nde ,Pdre ,Rbtc ,RyH,TrosV Zakharov V, Ternois HE, Roy W, Rabitsch J, Pedersen (2007) O, DB Nedved Roy MEN, IA, Majerus AJM, Loomans I, ia) aaiodof parasitoid a rinae), ladybeetle.htm iCnrl(hsise.doi: issue). (this BioControl 70:80 (Piracicaba) ocnlia)to Coccinellidae) 52:375–400 Entomol Rev Dordrecht Publishers, Academic Kluwer synthesis. Gre A, Battisti 44:1163–1171 ayidbeetle, ladybird .axyridis H. .Ws ignaUiest xeso Service. extension University Virginia West ). nctu rvsi sal noohg 16:433–444 Entomophaga Israel. in groves citrus in ntentv an nteivddrgos ntematm,comparative meantime, the In regions. invaded the in fauna native the on ´ ie 0Ag2007 Aug 20 Cited . or -,EasH es akadwo oigiscsi iigtesi uoe a Europe, in trees living in insects boring wood and Bark (eds) HF Evans J-C, goire acKnsadRnt idlaespotdb rn rmteSisFederal Swiss the from grant a by supported are Zindel Renate and Kenis Marc shgl auda ilgclcnrlaet atclryi corn, in particularly agent, control biological a as valued highly is euei bassiana Beauveria dlabipunctata Adalia amnaaxyridis Harmonia oemglamaculata Coleomegilla amnaaxyridis Harmonia http://www.mtngrv.missouristate.edu/Publications/PestAlert1.htm eiiu coccinellae Perilitus 10.1007/s10526-007-9132-y 2003 oeo hmcladbhvorlce.JIsc Physiol Insect J cues. behavioural and chemical of role : netb ahl84:137–144 Pathol Invertebr J . nErp:sra n itiuino o-aiecoccinellid. non-native a of distribution and spread Europe: in .axyridis H. evzadOmkar and Pervez ; 14 nBlim iCnrl(hsise.doi: issue). (this BioControl Belgium. in Shak 82 Hmnpea rcnde Eupho- Braconidae, (Hymenoptera, 1802) (Schrank, DGe)(oepea ocnlia) e Agric Rev Coccinellidae). (Coleoptera, (DeGeer) ouain ih lorsl nreduced in result also might populations http://www.wvu.edu/ ¨ gT ei ,KasizrBM Kova BEM, Klausnitzer M, Kenis T, gg .axyridis H. 2006 amnaaxyridis Harmonia .Acnrlmto that method control A ). amnaaxyridis Harmonia ntentv range native the in * agexten/ipm/insects/ hlcrsbipu- Chilocorus ngrapes. on ) Harmonia ie 20 Cited . 10.1007/ Pallas ´ r ˇ http://doc.rero.ch priG a arbk 16)Dsrpin ilgee motneduenuel espe nouvelle d’une importance et biologie Basses-Alpes. Description, les (1968) et D Alpes-Maritimes Waerebeke les van dans G, aphidiphages Iperti coccinelles des lady parasites Asian multicoloured Les the (1964) Managing G (2006) KB Iperti McClure DS, Fickle MA, Ellis RN, Williams DJ, Inclan eCec ,PeesI egueG hsO(03 neatosbetween Interactions (2003) O Thas G, Vergauwe I, Peeters P, Clercq De uce D se M uce J(99 neatosi noooy pis piohg n nsin ants and aphidophaga parasitoı aphids, du entomology: Interaction in (2006) Interactions A (1999) Firlej MJ Dutcher PM, Estes JD, Dutcher of Autodissemination (2003) beetle FE ladybird Vega Asian wide-spread PF, a parasitises Dowd that (Diptera) Phoridae of species new A (1997) RHL Disney avnT,Brns C ucio D(07 nmrtv n ioilsqeta apigpasfor plans sampling Garce multicolored sequential binomial the and to Enumerative spinosad (2007) and WD the Hutchison indoxocarb of EC, of Burkness risk Toxicity TL, the (2006d) Galvan reducing WD Midwest: Hutchison the RL, in multi- Koch grapes the TL, Wine of Galvan (2006c) infestations WD on Hutchison injury EC, berry Burkness of TL, Influence Galvan (2006b) WD Hutchison EC, the Burkness of TL, management for Galvan insecticides selected and of corn Efficacy sweet (2006a) in WD insecticides Hutchison used EC, commonly Burkness of TL, Toxicity Galvan (2005b) WD Hutchison RL, Koch TL, development Galvan survival, on indoxacarb and spinosad of Effects (2005a) WD Hutchison RL, Koch TL, Galvan isegH,Lbu A ee ,Ziu 20)Ptnilnnagtefcsof effects nontarget Potential (2002) E Zhioua K, Heyer RA, Lebrun HS, Ginsberg us D,Sap G rofil H akrL,MjrsTO ahrvI,MjrsMN(1995) MEN Majerus IA, Zakharov TMO, Majerus LE, ( Walker beetle AH, lady Broomfield Asian RG, multicolored Sharpe ladybird the GDD, of native Hurst treatment a and and Behavior introduced (2004) J an Kovach between MF, interactions Huelsman Indirect (2002) Academic Kluwer GE coccinellidae. 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South for implications Hemisphere: Western the in cinellidae) pce fccieldfo aa n rti.Booto ti su) doi: issue). (this BioControl Britain. and Japan from coccinellid of species cinellidae noo es61:207–208 News Entomol coccinellae by 62:1456–1458 Am Soc Entomol Ann Podopolipidae). (Acarina: Dakota 19)Eteevraini h rvlneo neie aekligmcoraim ewe three between microorganisms male-killing inherited of ladybird prevalence the the of populations in variation Extreme (1998) noohg 42:13–20 Entomophaga 103:1–7 Entomol J Eur 30:68–70 10:15–24 Hist Nat Entomol J Br Braconidae). noptera: ahl94:196–203 Pathol virescens Service. beetle, lady Asian axyridis Harmonia amnaaxyridis Harmonia n fet ntepaahdadanntre ocnli nafla icnrlSiTcnl5:425–438 Technol Sci Biocontrol alfalfa. in HSE-1030-01. coccinellid Entomology non-target a and aphid pea the on effects and eils,afnu soitdwt rmtr otlt of mortality premature with associated fungus a beniales), gi e eoo 17:131–134 Rehovot Res Agric mrc.CnEtml130:905–906 Entomol of Can parasitoid America. a as nidae) fteJpns beetle Japanese the of toggse triangulifera Strongygaster iCnrl(hsise.doi: issue). (this BioControl . http://www.wvu.edu/ Acmcts aoleils:rl fmtn ttsadageainbhvo.JInvertebr J behavior. aggregation and status mating of role Laboulbeniales): (Ascomycetes: Shak,aprst fautCcieia nNrhCia(yeotr,Braconidae). (Hymenoptera, China North in Coccinelidae adult of parasite a (Schrank), Shak.Kny 37:147–1166 Kontyu (Schrank). amnaaxyridis Harmonia nBian nen 29:196–208 Antenna Britain. in , Pla)(oepea ocnlia) nBian clEtml31:207–215 Entomol Ecol Britain. in Coccinellidae), (Coleoptera: (Pallas) oilajaponica Popillia amnaaxyridis Harmonia amnaaxyridis Harmonia http://www.ohioline.osu.edu/hse-fact/1030.html * Dpea ahnde nNrhCrln.JEtmlSi37:124–127 Sci Entomol J Carolina. North in Tachinidae) (Diptera: agexten/ipm/insects/beetle.pdf amnaaxyridis Harmonia 10.1007/s10526-007-9121-1 Clotr:Ccielde.Ws ignaUiest Extension University Virginia West Coccinellidae). (Coleoptera: Clotr:Saaaia) icnrlSiTcnl9:151–158 Technol Sci Biocontrol Scarabaeidae). (Coleoptera: ¨ e.WsapWseshfe,Magdeburg Wissenschaften, Westarp fer. Tetrapolipus Pla)(oepea ocnlia)i etr North western in Coccinellidae) (Coleoptera: (Pallas) 16 Clotr:Ccielde.Hrdt 81:683–669 Heredity Coccinellidae). (Coleoptera: toggse triangulifera Strongygaster amnaaxyridis Harmonia Clotr:Ccielde by Coccinellidae) (Coleoptera: amnaaxyridis Harmonia ioapscoccinellae Dinocampus hlcrsbipustulatus Chilocorus eprmcsvirescens Hesperomyces from ie 0Ags 2007 August 20 Cited . ° ipdmaconvergens Hippodamia 0 ihmn ul,Slough publ., Richemond 10. euei bassiana Beauveria ioapscoccinellae Dinocampus eiwo t ilg,ue in uses biology, its of review a : 10.1007/s10526-007-9138-5 Pla)(oepea Coc- (Coleoptera: (Pallas) ie 0Ags 2007 August 20 Cited . Le)(itr:Tachi- (Diptera: (Loew) .i sal salJ Israel Israel. in L. Shak (Hyme- (Schrank) hxe (Laboul- Thaxter t persistence its : Hesperomyces eiiu coc- Perilitus rmSouth from Harmonia Perilitus n two and http://doc.rero.ch aeaC,KneyG,BoneC(05 oeo iulcnrs nteaihigbhvo of behavior alighting the in contrast visual of Role (2005) Coccinellidae): C (Coleoptera: Brownie beetle GG, lady Kennedy Asian CA, multicolored Nalepa The (2000) DI Hopkins KA, Kidd CA, Nalepa aeaC,Kd A hsrmK 19)Booyof Biology (1996) KR Ahlstrom KA, Kidd CA, Nalepa ikrn J i Y enlsA oesG isnR(06 h vlaino eeiltetet for treatments remedial of evaluation The (2006) R Riesen G, Soleas A, Reynolds of JY, influence The Lin (2005) GJ, I Brindle Pickering R, Riesen G, Soleas A, environment— Reynolds complex JY, a Lin in ladybird, Foraging GJ, Asian (2005) Pickering JA multicoloured Pickett MA, of Birkett application R, Glinwood control V, Ninkovic biological J, and Pettersson Ecology (2006) A Omkar involving predation A, Intraguild Pervez (2007) MEN Majerus RL, Ware HE, Roy J, Baverstock JK, Pell aceS rw M aeu E 20)Hbi aigbetween mating Hybrid (2007) MEN Majerus PM, Brown S, Pascoe o E aesokJ elJ 20)Mnpltn eaiu:asrtg o etcnrl n ks S, Ekesi In: control? pest for strategy a behaviour: entomo- Manipulating endgames: (2008) and JK interactions Pell Bizarre J, (2006) Baverstock JK HE, Pell A, Roy Hajek E Eilenberg D, Steinkraus repels HE, Deet Roy multi- (2004) the JC of Davis behavior JR, the Aldrich modifying EW, for Riddick Potential (2000) JC Davis A, Milo De JR, Aldrich EW, fungus Riddick parasitic of distribution and density Occurrence, (2005) PW Schaefer EW, Asian Riddick multicolored the of pheromones and attractants repellents, for Search (2004) JR Aldrich EW, Riddick a N ec L(04 ayu yesniiiy eoto aeadrve fltrtr.Allergy literature. of review and case a of O report WK, hypersensitivity: Reeves Ladybug (2004) HL Pence JN, of College Ray Kentucky, of University structures. of invasion beetle lady Asian (2005) L on Townsend fungi MF, Potter entomopathogenic of Influence (1998) PE Parker JC, Legaspi TJ, Poprawski ikrn J e ,Sla J(07 eemnto fteciia tgso rcsigadtlrnelimits tolerance and processing of stages critical the of Determination (2007) GJ Soleas K, Ker GJ, Pickering ehyvV,KzesvV 17)Ainpeaino ocnlisi rmrergo.Entomofauna museum. history region. Natural Primorie database. in chalcidoidea Universal coccinellids (2007) on J predation Noyes Avian (1973) VN Kuznetsov VA, Nechayev sw 19)Alf al fteldbr beetle ladybird the of table life A (1992) N Osawa (1985) CA Tauber MJ, Tauber JJ, Obrycki sw 19)Pplto edsuiso h piohgu ayidbeetle ladybird aphidophagous the of studies field Population (1993) N Osawa akH akY,Hn K h Y(96 aaiod fteahdpaosladybeetles, aphidophagous the of Parasitoids (1996) SY Cho OK, Hong YC, Park H, Park axyridis 35:150–157 Sci Entomol J sites. aggregation to orientation 89:681–685 Am Soc Entomol Ann aggregations. winter ieafce by affected 70:S128–S135 wine Sci Food J 102:365–370 aging. Entomol and J composition Eur wine ladybird. on spot seven the of behaviour searching support semiochemicals axyridis Harmonia 10.1007/s10526-007-9125-x axyridis Pla)(oepea ocnlia)i hnho ra,Kra oenJEtml26:143–147 Entomol J punctata Korean Korea. areas, Chuncheon in Coccinellidae) (Coleoptera: (Pallas) 761() otPO,Tiadu-9 2,Krl,India Signpost Kerala, Research 023, management. Trivandrum-695 pest P.O., biological in Fort fungi (2), entomopathogenic 37/661 of Use 51:331–357 (eds) Entomol NK Maniania Rev Annu hosts. arthropod their and fungi 39:373–386 pathogenic Sci Ann Entomol J products. bioassays. natural laboratory in plant-derived adults lidae) with Coccinellidae) (Coleoptera: 93:1314–1321 Am beetle Soc Entomol lady Asian colored 98:615–624 Am Soc Entomol Ann dae). environ- urban and virescens agriculture in beetles 50:167–168 lady Entomol Asian Am symposium—multicolored ments. Instant beetle. lady beetle, 136:661–662 cantharid a of asitoid shaPo 25:133–136 Proc Asthma ef416.asp Agriculture.ENTFACT-416. of Department Agriculture, 27:785–795 Entomol Environ setosum for oesooDleoVsoa Vladivostok Vostoka, Dalnego Sovetskogo n eeomn nrlto ohs-tg takd n noo o m78:852–854 Am Soc Entomol Ann attacked. host-stage to relation in development and curation/projects/chalcidoids/ eaint h pi bnac.JnJEtml60:575–579 Entomol J Jpn abundance. aphid the to relation pea ocnlia) ietbe n e atraayi.RsPplEo 35:335–348 Ecol Popul Res analysis. factor key and tables life Coccinellidae): optera: amnaaxyridis Harmonia eiwo urn nweg n uueprpcie.Booto ti su) doi: issue). (this BioControl perspectives. future and knowledge current of review a : ulAa noo o 66:156–159 Soc Entomol Amat Bull . Clotr:Ccielde toewneigsts nio noo 34:425–431 Entomol Environ sites. overwintering at Coccinellidae) (Coleoptera: Clotr:Ccielde,a motn rdtro htfle Hmpea Aleyrodidae). (Homoptera: whiteflies of predator important an Coccinellidae), (Coleoptera: ie 0Ag2007 Aug 20 Cited . Lbubnae:Lbubnaee nmlioordAinbel Clotr:Coccinelli- (Coleoptera: beetle Asian multicoloured on Laboulbeniaceae) (Laboulbeniales: 0 aaJ 20)Frtrpr of report First (2004) JE Hara amnaaxyridis Harmonia eiw icnrlSiTcnl16:111–128 Technol Sci Biocontrol review. a : o ldbgtit nwn.Vts46:85–90 Vitis wine. in taint’ ‘ladybug for huigahspennsylvanicus Chauliognathus ie 0Ags 2007 August 20 Cited . n odSiTcnl41:77–86 Technol Sci Food J Int . eiiu coccinellae Perilitus toggse triangulifera Strongygaster amnaaxyridis Harmonia 17 amnaaxyridis Harmonia amnaaxyridis Harmonia http://www.ca.uky.edu/entomology/entfacts/ Hmnpea rcnde:parasitization Braconidae): (Hymenoptera: Clotr:Cnhrde.CnEntomol Can Cantharidae). (Coleoptera: als(oepea ocnlia)in Coccinellidae) (Coleoptera, Pallas amnaaxyridis Harmonia http://www.nhm.ac.uk/research- Dpea ahnde sapar- a as Tachinidae) (Diptera: pla)(oepea Coccinel- (Coleoptera: (pallas) Clotr:Ccielde in Coccinellidae) (Coleoptera: amnaaxyridis Harmonia amnaaxyridis Harmonia amnaaxyridis Harmonia eagu parce- Serangium and Hesperomyces dlabi- Adalia Harmonia Harmonia (Cole- http://doc.rero.ch oooaS,CdreD oeJC(00 ahgnct of Pathogenicity (2000) J-C Cote D, Coderre SI, Todorova hxe 19)Splmnaynt nNrhAeia aoleica.Po mAa rsSci two of Arts effects Acad the of Am Evaluation Proc (1996) D Laboulbeniaceae. Coderre American J-C, Cote North SI, on Todorova note (Coleoptera: Supplementary species (1891) coccinellid R four on Thaxter pesticides biorational of Effects entomo- to (2000) Coccinellidae) VA (Coleoptera: Krischik beetles SF, lady Smith of Susceptibility (2005) TE Cottrell DI, Shapiro-Ilan o E rw M,RteyP aeR,MjrsMN(07 neatosbtentefna pathogen fungal the between Interactions (2007) MEN Majerus RL, Ware P, Rothery PMJ, Brown HE, Roy ehge J ae ,HuknS onyG rni ,Hurg 20)Eetohsooia and Electrophysiological (2007) E Haubruge F, Francis UK G, Press, Lognay University Cambridge S, York control. Heuskin New vector Hall, Q, and & Fagel Pest Chapman (2004) FJ, control. MW Verheggen Biological Service (1996) HF, Jr Emden TS Van Bellows RG, Driesche Bestimmung- Van Mitteleuropas: Tachinidae) (Diptera: Raupenfliegen Die (1994) B Herting H-P, Tschorsnig aeR,MjrsMN(07 nrgidpeaino mauesae fBiihadJpns coccinellids Japanese and British of stages immature of predation Intraguild (2007) MEN Majerus RL, Ware auaH iuaT(01 neseicitrcin natitohcatrpdsse:efcso pdron spider a multi- of the effects system: of arthropod control tri-trophic a for in evaluations interactions Interspecific Chemical (2001) T (2006) Kimura DJ H, Yasuda Inclan KB, McClure of prevalence DS, the Fickle in develop- variation RN, clinal the Williams Short-range on (2001) GDD studies Hurst microscope KM, Webberley electron JJ, scanning Sloggett and VL, Welch light- Correlative (1996) GW Beakes A, Weir ebre M ise ,Sogt J aeu E,HrtGD(06 pta aito nteincidence the in variation Spatial (2006) GDD Hurst MEN, Majerus JJ, Sloggett M, Tinsley Majerus KM, J, Webberley Buzcko V, Isham JJ, Sloggett JHGVD, in Schulenburg selection RW, sexual Husband mediated GDD, parasite Hurst of KM, Lack Webberley (2002) MEN Majerus J, Buzcko GDD, Hurst KM, Webberley a) plEtml120:159–163 Entomol of Appl development J dae). the on strains Vuillemin decemlineata ocnlia)hvn oeta sbooia oto gnsi neircps cnEtml93: Entomol Econ J 25:261–270 interiorscapes. in agents control biological as 732–736 potential having Coccinellidae) 89:150–156 Pathol Invertebr J nematodes. pathogenic and euei bassiana Beauveria eairlrsosso h utclrdAinld beetle, lady Asian multicolored the of responses behavioral 506:1–170 (Biol) O und A Verbreitung zur Angaben und stabellen lengi maculate Coleomegilla yteivsv ladybird invasive the by doi: press). (in Ecol Chem J semiochemicals. h uvvlo aveo he rdtr ayid nrlto oahd.EtmlEpAp 98:17–25 Appl Exp Entomol aphids. to relation in ladybirds predatory three of 31:C11 larvae of Tests survival Manag the Arthropod 2005. grapes, on beetle lady 26:547–550 Asian Entomol coloured Ecol urbanisation. with associated fungus transmitted sexually a of morphology mental E 20)Hs erdcinadasxal rnmte ies:cue n osqecsof consequences beetle ladybird and the causes of parasites disease: transmitted sexually transmitted of sexually a and reproduction hippodamiae Coccipolipus Host (2004) MEN 63:131–141 Behav Anim system. insect-STD an dlabipunctata Adalia Clotr:Chrysomelidae), (Coleoptera: n he pce fcoccinellid: of species three and iCnrl(hsise.doi: issue). (this BioControl . eseoye virescens Hersperomyces amnaaxyridis. Harmonia itiuino ocnli ete.JAi cl73:1195–1200 Ecol Anim J beetles. coccinellid on distribution Clotr:Ccielde.Pyortcin81:15–22 Phytoprotection Coccinellidae). (Coleoptera: oemglamclt lengi maculate Coleomegilla yu persicae Myzus ¨ ooi e izle re.SutatrBirNtr Ser Naturk Beitr Stuttgarter Arten. Einzelnen der kologie iCnrl(hsise.doi: issue). (this BioControl 10.1007/s10886-007-9370-6 18 yooi 88:677–693 Mycologia . 10.1007/s10526-007-9122-0 euei bassiana Beauveria amnaayii,Cciel septempunctata Coccinella axyridis, Harmonia dlabipunctata Adalia amnaaxyridis Harmonia Hmpea piia)adterpredator their and Aphididae) (Homoptera: euei bassiana Beauveria u noo 103:793–797 Entomol J Eur . ibrae(o,Coccinelli- (Col, Timberlake sltstoward isolates 10.1007/s10526-007-9135-8 als osesquiterpene to Pallas, Leptinotarsa (Balsamo)