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Online Publication Date: 01 June 2003 To cite this Article: Omkar and Pervez, A. (2003) 'Ecology and Biocontrol Potential of a Scale-Predator, ', Biocontrol Science and Technology, 13:4, 379 - 390 To link to this article: DOI: 10.1080/0958315031000104505 URL: http://dx.doi.org/10.1080/0958315031000104505

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The publisher shall not be liable for any loss, actions, claims, proceedings, demand or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material. © Taylor and Francis 2007 Downloaded By: [NEICON Consortium] At: 10:31 11 September 2007 icnrlSineadTcnlg Jn 03,Vl 3 o ,379 4, No. 13, Vol. 2003), (June Technology and Science Biocontrol O:10.1080/0958315031000104505 DOI: (online)/03/4379-12 1360-0478 (print)/ISSN 0958-3157 ISSN [email protected] E-mail: Omkar. to: Correspondence effecti an is It nigritus Chilocorus REVIEW Se asraigadrlae sre de is and releases growth and global habits, rearing its predation mass on and information range scattered prey The foraging, crops. distribution, horticultural other and sugarcane, infesting asi h nomto,sc s nlecso t g,reproducti age, its of influences as, such information, the in reproducti gaps and ecological gi its been whereas has emphasis much where drawn, inferences and ra;o hs,tefre eemr ucsfli eltn h etppltos(eBach, prevalent (De populations pest of pest species in the the depleting released , especially in parasitoids, been successful of families number have more A were predators 1964). former include potential. and the these, damage parasitoids which of as, tremendous areas; Coccoidea, have such which enemies, (Superfamily etc.), Natural Eriococcidae, Pseudococcidae, phytophagous of scale Coccidae, enemies natural have the like biodiversity of manipulation and pests, subsequent environment and on introduction pesticides the to of led implications the of awareness Increasing INTRODUCTION in further require which diets Keywords: artificial and guild the in role its strategies, si,hv lydamjrrl ntemngmn fsae:i particular, in scales: of management the in role (Maskell), major a played have Ishii, bifasciata Comperiella ayidRsac aoaoy eateto olg,Uni Zoology, of Department Laboratory, Research Ladybird v rlatmt ha attempts eral clg n icnrlPtnilo Scale-Predator, a of Potential Biocontrol and Ecology hlcrsnigritus Chilocorus oiilacitrina Aonidiella v ooit nati colonist, e v Fbiis sacmo rdtro cl net,epcal Diaspididae. especially insects, scale of predator common a is (Fabricius) enmd ntercn att xli tfrtebooto fdiaspids of biocontrol the for it exploit to past recent the in made been e ( Recei Howard, v v v rnee.Teei naymtyi h ieauea literature the in asymmetry an is There needed. er nt apig asraigadfedrlae ftepredator, the of releases field and rearing mass sampling, to en d3 coe 01 cetd2Dcme 2002 December 2 accepted 2001; October 31 ed OMKAR v v ocnlia,Dapdde cl net,diaspids insects, scale Diaspididae, , , (Coquillett), ee.Teitrrtto fteeprcldt a enmade been has data empirical the of interpretation The iewed. oIdaadmyesl sals nps pre pest in establish easily may and India to e hlcrsnigritus Chilocorus uko-207 India Lucknow-226007, arlpi dalmani Habroleptis v set r orydatwt.Teeaeanme of number a are There with. dealt poorly are aspects e AND asei oleae Saissetia PERVEZ A Wswo)and (Westwood) Aphytis Á / 390 Brad and (Bernard) and # v ,adapti e, v Metaphycus ) lpet reproduction, elopment, 03Tyo rni Ltd Francis & Taylor 2003 v riyo Lucknow, of ersity pyu timberlakei Aphycus v estigation. oiilaaurantii Aonidiella v Asterolecanium n sur and e v v ln regions. alent ial,since ailable, ln with along v i v al Downloaded By: [NEICON Consortium] At: 10:31 11 September 2007 v 380 lcnu miliaris of 1991a; olecanium adults Samways, with the & region, infested Hattingh this 1986a,b; orchards In Samways, 2000). 1964; Samways, Bach, & (De Magagula Africa South of orchards melas certain of ecology The species. pest target of biocontrol in utility ladybeetles, their and ladybeetles ocnlaseptempunctata Coccinella 20)peetda nltclacuto prey of account analytical 1996; ladybeetles, Dixon Honek, an British and Coccinellidae, of been & of presented history ecology have Hodek natural the reviewed (2000) books 1994; the comprehensively described (1996) (Majerus, Three Honek (1994) decade and 2003). Majerus Hodek last these, Pervez, Of the & 2000). pests, during Dixon, Omkar phytophagous ladybeetles 2000; other on with (Dixon, published along etc. 2000). scales Dixon, psyllids, 1964; the aleyrodids, time Bach, of generation (De enemies low species natural the pest target and the specificity and host bioagent efficiency, the searching between high ratio their on dependent aecaatrsi akpthso eod hr n eet onnhsget,which segments, ninth to seventh and far third two second, these on However, patches species dark support. (1981). this corroborative characteristic further in Chazeau have receive instars they by larval until seriously five supported taken of be report was cannot reports a which dispersed also (Ponsonby is instars 1953), larval there to four (Fitzgerald, Interestingly, undergo substrata The to 1996). 1953). thrice the Basavanna, moult Copland, Channa usually on & & and (Puttarudriah (singly) out ends hatch eggs their instars of first spindle-shaped early one by yellow attached bright are they lays which ladybeetle female The History 1989). Life (Samways, winters dry it cool that and revealed summers studies humid Biogeographical and 1989). (Samways, hot rainfall) prefers summer with (tropical II nigritus Chilocorus CHARACTERISTICS GENERAL been have data empirical the from been inferences required. have wherever and suggestions predator, drawn, Interpretations the research. some to of further Owing for aspects management. made important pest certain in on utilization information its ambiguous for beneficial be establishment & may and Hattingh that guidelines utilization 1964; Bach, successful (De revealing density Evidence prey of the 2000). in Dixon, changes 1991a; the Samways, to response quick and efficiency nigritus tlzda iaeti aybooto rgamsadcasclyitoue nscale- and in parasitoids The introduced 1998). classically 1984, Samways, successfully and 1964; been nigritus programmes Bach, has C. (De biocontrol again It many and 2003). time Pervez, in insects, regions & many prevalent bioagent of Omkar a predator 1995; active as Bind, an & utilized and (Omkar subcontinent scales Indian the the to especially native is It 1998). ladybeetles. (Booth, of list the to Bach, added (De yields crop Earlier Samways the 2003). 1976; increasing (Leeper, Pervez, thereby species & and Omkar infestations heavy 1964; removing of capable ariolosum eti rdtr,i atclr ayete Clotr:Ccielde r lopotential also are Coccinellidae) (Coleoptera: ladybeetles particular, in predators, Certain h rys elwlra ee ihhi-oee pn rcse ntedra surface dorsal the on processes spiny hair-covered with beset larvae yellow greyish The mns rdcosldbels pce of species ladybeetles, predaceous Amongst .nigritus C. Booth, n te ocdpaosbelshsbe trbtdt hi ihsearching high their to attributed been has coccidophagous other and r onl epnil o elnn h ouainof population the declining for responsible jointly are Rteug D ah 94.Tesceso ilgclcnrlaet slargely is agents control biological of success The 1964). Bach, (De (Ratzeburg) v iz hlcrssubindicus Chilocorus nteps rmtdu ogte ctee nomto n oformulate to and information scattered gather to us prompted past the in . hioee sexmaculata Cheilomenes Bidvl Smas 94 atnh&Smas 91) ucs of Success 1991b). Samways, & Hattingh 1984; (Samways, (Boisduval) rgntsfo h ninsbrgo,casfe sCiai Zonobiome Climatic as classified sub-region, Indian the from originates .aurantii A. ines(ma evz 02 a enrcnl reviewed. recently been has 2002) Pervez, & (Omkar Linnaeus hlcrsnigritus Chilocorus tal et ot and Booth MA .PERVEZ A. & OMKAR n in abopat abuigdiaspid, harbouring plants bamboo giant and ,19)bttrenwOina species, Oriental new three but 1999) ., Chilocorus Fbiis Aawl aua 00 and 2000) Yasuda, & (Agarwala (Fabricius) hlcrsgracilior Chilocorus Chilocorus Á Fbiis soeo h retlspecies Oriental the of one is (Fabricius) / a nw ocnan6 predaceous 62 contain to known was rdtrrltosiso predaceous of relationships predator .nigritus C. r motn scale-predators, important are A . ot aercnl been recently have Booth aurantii htl ewe citrus between shuttle v iz pis adelgids, aphids, . netn citrus infesting , v iz . Chilocorus Aster- C. Downloaded By: [NEICON Consortium] At: 10:31 11 September 2007 h niiul of individuals The pesticides hazardous FORAGING to tolerant highly and they fields conditions the in stressful since unclear, the 1994). is to (Schoeman, captivity in in reason adapted when The females males females. better older older than the are the of faster of mortality scarce mortality increased increased become 6 1984). this the 1984). for and females to Tate, Tate, 2 attributed that & be Africa, between (Samways conclusion & may age 1:0.42 interesting which captivity, South of of (Samways an (male:female) adults to of sex-ratio respectively older led a the This revealed prey-sites 1:1.71, of insectary sampling an two and the in observations, months at 1:2.07 field the adults was to sex- young Contrary Tshaneni The longevity. high the and relatively its between Hazyview was ratio (male:female) female-biased high ratio this for reason The 1991). erhte eetdbc oetniesac Hsel&Suhod 98 atr&Dixon, & Carter 1978; Southwood, & (Hassell search intensive extensive the prey 1984). to during in encountered back culminating not reverted were search they prey intensive search If and to 1995). adults Samways, while extensive Both & distances, from 1995). (Hattingh Samways, patterns short consumption & foraging over (Hattingh their cues cues tactile altered olfactory on larvae and dependent visual more were both prey latter through the high prey of locating searching sites of at capable their concentrate aid to to them margins, and enable photo- predators and similar the show veins density. also of species leaf responses prey as such the Since geotaxes, such 1995). Copland, plants, & (Ponsonby of behaviour features prominent use with infested varied orchard (male:female) sex-ratio the the The in in edge 1984). Tate, 1:3.55 the & at to (Samways scalloped absent 1:1.44 females weakly are from in is patches rounded sternite similar is eighth however, but The dark; 1984). males are Tate, sternite & the ones, seventh (Samways older females and the in fourth particularly the males, of In plates). areas (abdominal lateral sternite their in on prominent present patches is dimorphism Sexual Sex-Ratio and Dimorphism Sexual pronotum. the between of areas tips orange antero-lateral with elytra the shiny on highly Bind, from and & and black Omkar ranging eyes all 1984; sizes the Samways, by 1953; with of characterized Basavanna, are shape Channa concavity They & semi-spherical 1995). (Puttarudriah the almost mm their 5.50 inside ladybeetles to adult to 3.00 congregate The owing 1953). recognizable instars Basavanna, easily Channa fourth & are (Puttarudriah Mature pupate to appearance. leaves crumpled banded a impart oeatatv hntehrzna ns h lnsta eerglryecutrdb the e.g by oblong, encountered or regularly were elliptic were ovate, plants that i.e., the plants leaf-structure, of The similar lines ones. have Vertical predator horizontal 1995). the Samways, than & attractive (Hattingh more trees individual with plant, horizon host more of spent presence and further the walked in cyanophylli hosts, prey them Abgrallaspis in their activity for was to their attracted searching response enhance enable highly and time directional mechanisms habitats were host strong These females favoured odour. the The A in prey searching. volatiles. active with remain plant supplemented to host predator was the odour of plant in presence increased when the Orthokinesis exhibited 1995). in klinokinesis Copland, searching increased beetles & and the (Ponsonby starved prey, turning) speed) of (forward with intensity orthokinesis encounter and decreased an (frequency by After modified responses. was olfactory activity on based location habitat h oaigpten naut n avewr naibydfeet stefre were former the as different, invariably were larvae and adults in patterns foraging The hlcrsnigritus Chilocorus hlcrsnigritus Chilocorus .nigritus C. CLG N ICNRLOF BIOCONTROL AND ECOLOGY a iulyatatdt rmnnl ihute etrs uhas, such features, silhouetted prominently to attracted visually was Sgoe)oor Pnob oln,1995). Copland, & (Ponsonby odours (Signoret) a irrhclpe-oainmcaim hc novshost involves which mechanism, prey-location hierarchical a has r oiieypooatcadngtvl etci n edto tend and geotactic negatively and phototactic positively are .nigritus C. HLCRSNIGRITUS CHILOCORUS h w ee a eietfe ythe by identified be can sexes two The . siitsnerii oau melongena Solanum Bouche . irs uv,mango, guava, citrus, , (Erichsen ´ inesand Linnaeus tal et 381 v iz. ., Downloaded By: [NEICON Consortium] At: 10:31 11 September 2007 ni Mn rsnmoty 98.Bt aveadaut edo h spiralling the on feed adults and larvae Both 1998). Krishnamoorthy, whitefly, & (Mani India uaa,Ida(Murlidharan India Gujarat, on exclusively development their complete h hr one adbe epteldbel ntecmlt eoa ftesaefrom scale of larvae the for of behaviour removal Feeding complete 1988). Wilson, the & in (Samways ladybeetle surface the fruit help the mandibles pointed sharp The Behaviour Feeding of predator incidental 1988). an Ghosh, considered was aphids, It on 1995). Bind, feeding reported were nerii adults Aphis The 1948). erysimi (Sharga, Lipaphis India Kanpur, from recorded nigritus Chilocorus Range to Prey order in used PREDATION are only However, prey Samways, agents. alternative & reservoirs. biocontrol growing to (Hattingh preferred the by orchards orchards for from scales less migration affected reservoirs of minimize harbouring valuable the management plantations as of the serve for periphery peripheral may exploited the which being in 1991b), is plants predator host the alternative of aspect of This success the for reason ( prey essential of plants, absence bamboo the in Africa, of South species In nerii other 1989). Hattingh, in 1986; as Henen, prey-recognition, & facilitates probably it as visual 1991b), important an 1995). also Samways, is & shape (Hattingh leaf that prey-location suggests for which leaves, cue plant fig and coffee banyan, 382 mns h w ee,hwvr i o aysgiiaty(osny&Cpad 2000). Copland, & (Ponsonby significantly vary compared not when (Ponsonby did instars, respectively however, larval 170, sexes, the was and by two instars instar, 127 consumption the larval 107, prey second 104, in amongst male instar, was differences dehydration fourth instars The first and female 2000). rapid by Copland, third include and & their second, 160 (which first, and to 135 killed by 96, females) leads scales 122, & adult or of scales Ponsonby and behind number 1988; males the cuticle total Wilson, puparial completely of the & The instars leaving (Samways piercing/puncturing 1990). three contents, eaten (Foldi, The insect body other sucked entire 2000). the the and the out Copland, whilst cover was sucked Rarely behind, scale and eaten. cuticle formed cover partially the newly scale leaving the the juices, through removed body chewed the instars out First scale, 2000). cyanophyllum Copland, using investigated eielsalataniae Hemiberlesia rsnmoty 99.Tepeao skont eantrleeyo ubrof number a of enemy natural a be to known is , infesting predator scale-insects The 1999). Krishnamoorthy, sualcsi pentagona Pseudaulacaspis a,19) h rdtrwsietfe edn ntescales, the on feeding identified was predator The 1994). man, diaspids, on preys din Linnaeus Green hlcrsnigritus Chilocorus Samways, & (Hattingh adults in pattern foraging area-restricted enhanced intake Prey tal., et ,tepeao htlst lentv rysts(e.g., sites prey alternative to shuttles predator the ), isnastellifera Vinsonia , luoiu dispersus Aleurodicus suoocscitriculus Pseudococcus , 91.I savrcosfee nrdscale, red on feeder voracious a is It 1991). oe eFnclmefo h uko ein fUtrPaeh(ma & (Omkar Pradesh Uttar of regions Lucknow the from Fonscolombe de Boyer edoaau giganteus Dendrocalamus (Kaltenbach), ed navreyo netpss t piohgu ai a been has habit aphidophagous Its pests. insect of variety a on feeds hyopau aonidum Chrysomphalus ry nsf re scale, green soft on preys Nwta)and (Newstead) .nigritus C. agoiTzet and Targioni-Tozzetti (Westwood) tal et usl(ai&Kihaorh,19) h avefe and feed larvae The 1999). Krishnamoorthy, & (Mani Russel v Bre z siitsdestructor Aspidiotus iz. ,19;Mrihrn 1993). Murlidharan, 1992; ., Green v MA .PERVEZ A. & OMKAR crn brassicae icoryne nbooto fsae Htig awy,1991b). Samways, & (Hattingh scales of biocontrol in heioocsmarlatti Phoenicococcus ocsacutissimus Coccus , Palmicultor , brlapscyanophylli Abgrallaspis Munro (Linnaeus) .dispersus A. Coccus ) uaapstubercularis Aulacaspis hc c srsror n r major a are and reservoirs as act which , Linnaeus, .aurantii A. p and sp. v pi cracci Aphis iridis (Signoret), eioahsbeckii Lepidosaphes , seoeaimmiliaris Asterolecanium ntelbrtr Mn & (Mani laboratory the in Gen netn cdlm in lime acid infesting (Green) Pseudaulacaspis yu persicae Myzus Green altrablanchardi Parlatoria okrl ndt lnsin plants date on Cockerell Smas 99,adalso and 1989), (Samways, v eolse actiniformis Ceroplastes spe Pnob & (Ponsonby prey as ora ocshesperidum Coccus , Chilocorus oh(graa& (Agarwala Koch esed(Schoe- Newstead .nigritus C. Sle)and (Sulzer) p (Jalalud- sp. (Newman), Aspidiotus (Podoler ngiant on Hirst was , Downloaded By: [NEICON Consortium] At: 10:31 11 September 2007 a)(awy isn 98,wietelra preferred larvae the while 1988), Wilson, & (Samways day) Smas&Wlo,18) htsae niaiiyfrfrtisa avet ryo second on prey as to such larvae insects, instar scale first cyanophylli for other Abgrallaspis inability of an nymphs stated that instar 1988), Wilson, & instar of nymphs (Samways first instar the second that on observed prey (2000) easily Copland and Ponsonby 1992). miliaris Samways, Asterolecanium aurantii, ie(20 nine rysbttto eprrl eue h ins,ie,imtr eeomn n adult and development immature i.e., fitness, the reduces in Development reproduction temporarily on Conditions substitution and Abiotic Prey and male Biotic adult of of Influence longevity The respectively. 1994). days, (Murlidharan, 76.1 and respectively 50.2 days, was 6.6 female and 2.5 for scale, on fed when hr n orhlra ntr of instars larval fourth and third of development The 1989). on Singh, & of (Jalali predator potential scale biotic the madagassa Sticholotis on data Empirical scale the the on DEVELOPMENT increased AND enemies GROWTH perhaps natural two parasitization these that of impact suggests combined This the population. reducing 1988). for Wilson, likelihood & (Samways utblt over suitability nigritus hl dlscnue 60.7 consumed adults while h rdtr oeta n rycnupinb h aveadaut of adults and larvae the by consumption prey and potential predatory The Preference and Consumption Prey re a tde yJll ig 18) h avepee n29.7 on preyed larvae The (1989). Singh & Jalali by studied was Green otlt Pnob oln,19) h hoeia oe hehl o uvvlo first of survival for threshold of lower instar theoretical first The of 1996). development Copland, 20 and & of survival (Ponsonby Temperatures the mortality regions. in for sub-tropical helped suitable and has not which tropical predator, were the the of in strategy adaptive establishment of an its species is different tolerance Temperature of 1. tolerance Table in Thermal 1994). Samways, for & patches Larvae (Hattingh prey 1992). the Samways, select adults & as (Hattingh fields, the serious diet in in types of emphasizing prey change them. two maintenance, risk to the adult exposed to the generally on not larvae a than were reduces had of development substitution sensitivity also larval Prey greater the 1992). prey the Samways, on & unfamiliar effect (Hattingh prominent progeny an of more of fitness the presence to repercussion in oviposition reduced Singh, consumption & lifespan (Jalali 1989). total Singh, a (34.8) & with instars (Jalali diaspids/day third diaspids 20.9 average the 1317.1 an of by on followed consumed (55.6), Adults 1989). diaspids of number maximum preferred larvae perniciosus hlsaonidum phalus hlcrsnigritus Chilocorus preferred adults The hlcrsnigritus Chilocorus .perniciosus Q. Á hnpoie ihscn ntr of instars second with provided when / 5dyod isi species, diaspid 25-day-old) (Comstock), siitsnerii Aspidiotus .nigritus C. siitsdestructor Aspidiotus (Linnaeus), ncmaio oteohrto h uaino h g tg,frt second, first, stage, egg the of duration The two. other the to comparison in .blanchardi P. CLG N ICNRLOF BIOCONTROL AND ECOLOGY a eoeo h pce ottlrn oteetee ftemperature of extremes the to tolerant most species the of one be may Wie nnn isissoe httefre a h otpotent most the was former the that showed diaspids nine on (Weise) edl e on fed readily oiilaaurantii Aonidiella .Ti eeldfeiiiyi oaigpten ffrtisa of instar first of patterns foraging in flexibility revealed This ). siitsdsrco,H aaie oiilaaurantii Aonidiella lataniae, H. destructor, Aspidiotus Htig awy,19) tmyb osdrdaatv,as adaptive, considered be may It 1992). Samways, & (Hattingh Á / oioyiu albus Aonidomytilus 14dapd/a.Teaut preferred adults The diaspids/day. 71.4 Mrihrn 94.Tepeua n ua eid lasted periods pupal and prepupal The 1994). (Murlidharan, Pnob oln,2000). Copland, & (Ponsonby .nigritus C. and brlapscyanophylli Abgrallaspis Jll ig,18) h orhisasconsumed instars fourth The 1989). Singh, & (Jalali v z eaapsglomerata iz. hyopau aonidum Chrysomphalus oiilaaurantii Aonidiella 2. clsdy to scales/day) (21.6 .ngiu,Cioou bijugus Chilocorus nigritus, C. a .,35 .,38ad38dy,respectively, days, 3.8 and 3.8 3.8, 3.5, 6.1, was HLCRSNIGRITUS CHILOCORUS siitsnerii Aspidiotus brlapscyanophylli Abgrallaspis Cceel and (Cockerell) .nigritus C. siitsnerii Aspidiotus aaiie by parasitized otayt nerirreport earlier an to contrary , siitsnerii Aspidiotus mrhlgclysmlrto similar (morphologically (Green), Lnau)(atnh& (Hattingh (Linnaeus) .nigritus C. eioahscornutus Lepidosaphes a ats 3. days) (32.3 fastest was .perniciosus Q. Chilocorus Á / 80diaspids/day, 38.0 Quadraspidiotus niaigits indicating , over Mlat and (Mulsant) .nigritus C. .nigritus C. 8 .bifasciata C. n below and C 1. scalea/ (11.4 u ohigh to due , Aonidiella Chrysom- sshown is while , 383 can on C. Downloaded By: [NEICON Consortium] At: 10:31 11 September 2007 n . g epciey totmmcntn eprtr n . gs 10 n . gat mg 8.4 and 71.0% eggs, 2.2 (3 and temperatures on temperature constant exposure fluctuating optimum of at respectively, days mg, 40 8.8 after and weight adult and viability eodadtidisa uain,wr lorpre yPnob n oln (1996). Copland than and larger Ponsonby relatively by were 30 reported which also instars, were fourth durations, and instar third first and of second durations larval similar much osmto ytepeao rm39 o88 cls h edn aewshgeta a at highest was rate feeding The scales. cyclic 8.83 or to 26 3.94 fluctuating of from temperature at predator constant the 1998). spermathecae by Copland, & consumption in (Ponsonby adults of mortality ability reproductive sperm the pause reducing a thus and/or induces temperatures, probably exposure spermatogenesis Cold 1994). Samways, in & (Hattingh ones fluctuating than h r-aigpro of period pre-mating The REPRODUCTION oln,20) h ylctmeaue(14 temperature cyclic The 2000). Copland, fdfeetisaswsntafce Pnob oln,19) h riaia eidat period preimaginal The 1996). Copland, (14 & duration temperatures (Ponsonby relative affected the cyclic not but temperature, was in instars increase different 30 the of and with increased 28 thus, 26, rate 24, developmental 22, 20, of temperatures 16.6 was instars simoni Chilocorus infernalis Chilocorus distigma Chilocorus nigritus Chilocorus Species of survival Percent 1. TABLE 384 siiae oditk aidfo .9 gdya 13 at mg/day 0.097 from varied intake food assimilated hc a ihrta htosre t26 at observed that than higher was which pdrwb rayohrsbtaacmoe floefbe Smas&Mp,18)and 1983) Mapp, 1953). & Basavanna, (Samways scales Channa fibres dead loose & of of (Puttarudriah shields leaves composed the substrata scale-infested in other on eggs any four or to Dorge webs two eggs spider 1970; constituting the Mostly, Ahmad, groups 1970). small 1953; (Ahmad, mating in (Fitzgerald, first or the singly after laid days 10 are oviposited females and min 28 eudt agn ewe 0ad30eg Tbe2.Arcn td lutae htoptimum that illustrated study recent A 2). (Table eggs 370 and 90 between ranging fecundity ifrnei aiu ilgclatiue of when (33 attributes days) (24.0 biological slower various was in It difference days). (18.7 species When former the 2000). nigritus fed Copland, first when & faster was (Ponsonby diaspids, rate constant two development on at reared complete that were to larvae than sexes of higher mg the 16.24 significantly of both was mean A temperature mg/day). cycling (1.98 temperatures the at intake However, 8 of rate feeding The ewe 0ad30 and 20 Between h erdcieatiue of attributes reproductive The n hratrdcie Htig awy,19) h al vpsto,percent oviposition, daily The 1994). Samways, & (Hattingh declined thereafter and C Á / 5)(osny&Cpad 1998). Copland, & (Ponsonby 75%) avewr is e ntelte ry(atnh&Smas 91) osignificant No 1991b). Samways, & (Hattingh prey latter the on fed first were larvae atr atnh&Smas 1994) Samways, & Hattingh (after, Lnau)109 59 0 0 0 100 100 95 95 95 100 (Linnaeus) iad109 410108 0 88 100 100 94 93 100 Sicard 8 Fbiis 0 8109 59 0 95 95 95 100 88 100 (Fabricius) .Tedvlpetpro a 45 79 80 40 86ad2. asat days 25.2 and 28.6 34.0, 48.0, 57.9, 74.5, was period development The C. Ku)1010101010100 100 100 100 100 100 100 (Klug) usn 51010109 40 84 95 100 100 100 85 Mulsant Lnau)8 4101010100 100 100 100 100 94 85 (Linnaeus) 8 .nigritus C. ,a nraei osattmeaue nrae h aeo scale of rate the increased temperatures constant in increase an C, Á / 30 .nigritus C. Chilocorus 8 8 Á )ws4. as(osny&Cpad 96.Unexpectedly, 1996). Copland, & (Ponsonby days 47.0 was C) ,wt eae en oevrcosta ae Pnob & (Ponsonby males than voracious more being females with C, / 38 8 ) hwn htcntn eprtrsaemr suitable more are temperatures constant that showing C), nrae ihices ntmeaue ecigapa at peak a reaching temperature, in increase with increased .nigritus C. v MA .PERVEZ A. & OMKAR p tvroscntn eprtrs( temperatures constant various at sp. z Asterolecanium iz. a . asa 24 at days 5.4 was 8 tal et ,rsetvl Pnob oln,19) The 1996). Copland, & (Ponsonby respectively C, .nigritus C. Á ,17) eae looioi ntesl of silk the on oviposit also Females 1972). ., / 30 aebe eotdfrsvrlhsswith hosts several for reported been have 8 Pnob oln,20) Mean 2000). Copland, & (Ponsonby C 8 )icesdtevrct nbt sexes, both in voracity the increased C) .cyanophylli A. 1017243138413 8 a on tdfeetlvl fR.H. of levels different at found was p and sp. siitsnerii Aspidiotus Amd 90.Mtn atdfor lasted Mating 1970). (Ahmad, C 8 o142m/a t30 at mg/day 1.432 to C .nerii A. eprtr ( Temperature a eurdfrlra of larvae for required was n agdbten1 n 21) and 14 between ranged hi developmental their , a . gs 74.0% eggs, 4.3 was 8 C) .nigritus C. 8 C. C. brlapscyanophylli Abgrallaspis blanchardi Parlatoria nerii Aspidiotus beckii Lepidosaphes albus Aonidomytilus aonidum Chrysomphalus aurantii Aonidiella lataniae Hemiberlesia tubercularis Aulacaspis destructor Aspidiotus perniciosus Quadraspidiotus glomerata Melanaspis tegalensis Aulacaspis ryseisPeoioiinpro dy)Fcniy(gs vpsto eid(as vpsto aeReference rate Oviposition (days) period Oviposition (eggs) Fecundity (days) period Pre-oviposition glomeratus Aspidiotus orientalis Aonidiella species Prey ladybeetle, female of response Reproductive 2. TABLE Dt o available. not *Data Nwta)1. . ha (1970) Ahmad 3.1 * * 15.4 (Newstead) .Lbe 1 Labdev G.

Downloaded By: [NEICON Consortium] At: 10:31 11 September 2007 40**46Htig awy (1994) Samways & Hattingh (1989) Singh & Jalali (1989) Singh & Jalali (1989) Singh & Jalali (1989) 4.6 Singh & Jalali (1989) Singh & Jalali (1989) * Singh & Jalali (1989) * Singh & Jalali (1989) * Singh & Jalali (1989) * Singh & Jalali * (1977) Pope & Greathead * * * * * * * * * * * * * * * * 71.0 24 57.0 72.0 81.0 79.0 87.0 93.0 86.0 14.0 80.0 13.0 370 16.0 17.0 16.0 15.0 17.0 12.0 12.0 12.0 23.0 311059 osny&Cpad(1998) Copland (1994) & Murlidharan Ponsonby 5.92 * 180 * 1361 151.2 * 5.0 Á / 3241 .nigritus C. Á / Dorge * 3 tal et

(1972) .

385 CLG N ICNRLOF BIOCONTROL AND ECOLOGY HLCRSNIGRITUS CHILOCORUS Downloaded By: [NEICON Consortium] At: 10:31 11 September 2007 AL .Itouto of Introduction 3. TABLE gnaFie ilas&Geted(1973) Greathead & Williams (1987) Sekeroglu & Uygun (1981) Failed Chazeau (1978) (1984) (1991) Samways Lobel Chazeau Kinawy De (1988) Established Rossler & (1978) Argov Gutierrez (1981) Chazeau Established Established Established Established Established (1977) Pope & Greathead Established Established (1980) (1972) Blackmore Davis & Hill (1971) (1989) Failed Greathead Samways (1989) Samways ReferenceUganda Established Swaziland/Zimbabwe Islands Soloman Established Established Established Established Turkey Established Islands Society Samoa American Reunion Oman Caledonia Status Madagascar/New Kenya/Tanzania Java Israel Guam/Hawaii Ghana/Brazil Seychelles Agalega introduction of Place eesddrn January during released ae Htig awy,19) hsbhvori knt hs fprstis ste do they as feeding parasitoids, of affect those not to akin do is densities behaviour This predator 1990). varying Samways, of & destruction interference; (Hattingh mass mutual rates in results minimal prey-site the with at scales, predators of congregation The 1990). Samways, 95frtebooto of biocontrol the for 1985 e efadfu ete e rne(atnh&Smas 1990). Samways, & beetles (Hattingh two orange reached per density predator beetles the four scale, and red leaf by per encrusted completely were plants host ifrn osattmeaue ewe 0ad28 and 20 26 between of temperature temperatures constant 1998). different Copland, & (Ponsonby fecundity cyanophylli, enhances quality prey and temperature 386 t eesshv seilybe iepedadefciei ein fSuhAfrica, South conditions. climatic of inappropriate (Samways to regions 3) attributed (Table in be Africa can South effective attempts and unfruitful and Central few of widespread The regions 1999). climatic been the especially in particularly have releases Its nigritus Chilocorus RELEASES AND REARING CONTROL, BIOLOGICAL the inhibited & (Hattingh prey-substitution days several and for that oviposition prey-alteration to the similar delaying The was 1992). females, Samways, eggs/day) the workers. (5.92 of In rate activity 1998). other reproductive Copland, reproductive mean & rate by (Ponsonby the reproductive days fecundity, mean 50 recorded high and peak this 30 The with between of days. days, age spite 238 reproductive 180 of a of at 1998). period period attained Copland, oviposition reproductive was & prolonged an (Ponsonby a having date in to ladybeetle record resulted one highest also the fecundity is maximum which The eggs, 1890 laid female One h salsmn of establishment the of Infestation tree. per larvae fpssprla rawsbten0ad4.,weesps ubrrne rm148to 174.8 from ranged number pest whereas 49.7, of and Number 0 number release. median between the the was release, before area after 285.0 years leaf Two population. per pest pests the of in decline subsequent a and h ucs of success The niiul of Individuals spe,ehne eudt ihasih aito nnme feg adat laid eggs of number in variation slight a with fecundity enhanced prey, as .nigritus C. 8 .nigritus C. scmaaieymr utbe iligahg eudt f16 eggs. 1361 of fecundity high a yielding suitable, more comparatively is C a enclnzdi ubro onre ocnrlsaeinfestations. scale control to countries of number a in colonized been has .nigritus C. Á .nigritus C. / pi 95 hc eutdi ai ul-po rdtrnumbers predator of build-up rapid a in resulted which 1985, April siitsdestructor Aspidiotus xotdfo ni eeitoue fetvl nOa during Oman in effectively introduced were India from exported oiilaaurantii Aonidiella a edet o nrseii optto Htig & (Hattingh competition intraspecific low to due be may ntergo,teps rbe snwnts ct.We the When acute. so not now is problem pest the region, the in tdfeetplaces different at MA .PERVEZ A. & OMKAR .nigritus C. sacmo rbe nSuhArc.After Africa. South in problem common a is Knw,19) oa f63aut were adults 683 of total A 1991). (Kinawy, vnulyrahd38.2 reached eventually 8 Pnob oln,19) A 1998). Copland, & (Ponsonby C tal. et (1974) Á / 4. dlsand adults 144.5 Abgrallaspis tal et ., Downloaded By: [NEICON Consortium] At: 10:31 11 September 2007 cl 1. n 35scales/cm 23.5 and (15.8 scale fdapd Aoyos 95.Honey 1995). (Anonymous, diaspids of awy 19) reedidatfca it ae nbod of broods on based of diets which rearing artificial diets, mass Freeze-dried artificial Certain (1993). the 1995). Samways for (Anonymous, honey used g be 400 can and sugar g 200 powder, agar the to leads which maturation, predator for required is scales. density of control prey effective high Thus, 2000). was cyanophylli adults Abgrallaspis and instars of larval individuals late of in 500 of requirement resulted prey release populations total the scale Therefore, The red There mortality. advised. severe 1988). instar when at Wilson, larval starve introductions & early larvae egg (Samways high hatched where females reports the adult field of of several mostly predator’s most were composed the since advisable, is of not population introduction preferred is scale The mandibles more area the operations. are pointed scale-infested biocontrol adults the in sharp the in larvae their rate, young eggs feeding using and faster eggs surface their the fruit to than Owing the 1988). from Wilson, foraging scales & (Samways the female in adult cessation the remove temporary no is there conspecifics. and encountering crowding after increased pattern with disperse not h rdtrcnb utpido eea isis uhas, 1995). such Anonymous, diaspids, several 1993; on Sadakathulla, multiplied 1986; be Tate, can & predator Samways The 1983; Mapp, & (Samways ern a lob oeb sn honey a using by done be also can rearing of production glomerata its Thus, survive. not would prey of offspring infestation their density medium which with the sites on allows the (Erichsen to that fruits at compared recommended mechanism rotten as are built-in levels releases a such augmentative scale possibly avoid high is with to There infested infestation. adults fruits scale rotten medium avoid to with tendency those a has It 1991). epl germanica Vespula ern of rearing ucsfletbihet cainly h orhisa aveaeatce ythe by attacked are larvae instar fourth the Occasionally, parasitoid, establishment. successful enaeteps ak ieadaanb htln nt lentv ot.Teei very a is There hosts. alternative to of on may parasitism shuttling and fauna by of local again the frequency and suggested of member time Evidence low permanent ranks biocontrol. a pest classical becomes need it the in the area, reinvade utilization highlights an further in regions established and prevalent once rearing pest that scale different large in its establishment for successful Its past. the nigritus Chilocorus CONCLUSION 1953). Basavanna, Channa & (Puttarudriah of brood on based aiiae t uvvl ins n rdcin nyasnl parasitoid, single a Only production. and fitness survival, its facilitates enrpre,s a takn it. attacking far so reported, been eeal goe scmae oohrpeaeu ayete.Soai aaaeavailable are data Sporadic ladybeetles. is predaceous it other predators, other to and compared parasitoids as as ignored such bioagents, generally other amongst also but Coccinellidae hlcrsnigritus Chilocorus completely to larvae mature and adults the of ability the is success its for reason Another h ehiuso asraigo hssaepeao aebe ecie ndetail in described been have predator scale this of rearing mass of techniques The hlcrsnigritus Chilocorus Though tal. et Chilocorus Aoyos 95.Sdktul 19)dsrbdamto o h mass the for method a described (1993) Sadakathulla 1995). (Anonymous, .nigritus C. Homalotylus .nigritus C. 1991). , brlapscyanophylli Abgrallaspis CLG N ICNRLOF BIOCONTROL AND ECOLOGY a rvdisefcc sa motn icnrlaeto cl-net in scale-insects of agent biocontrol important an as efficacy its proved has .germanica V. Fbiis n ua of pupae and (Fabricius) a oeefceti eoigmdu n medium and medium removing in efficient more was srrl takdb ahgn rprstisadti ute isits aids further this and parasitoids or pathogens by attacked rarely is pce eeas eomne (Henderson recommended also were species soeo h otscesu iaet,ntol ntefamily the in only not bioagents, successful most the of one is r eurdfrtefrtlra tg ln Pnob Copland, & (Ponsonby alone stage larval first the for required are p,bttepretg fprsts suulyvr o nnature in low very usually is parasitism of percentage the but sp., yraigi on it rearing by 2 hna eyhg ns( ones high very at than ) a h otsuccessful. most the was .nigritus C. .nigritus C. .nigritus C. Á / gri rprdb dig1lwtrt 0gagar g 10 to water l 1 adding by prepared is agar Á / grde sa lentv oreo odinstead food of source alternative an as diet agar rmr.O hs,mr hn10frtisasof instars first 100 than more these, Of more. or .destructor, A. ppyspost Epiphyas ncnrs oteohrldbels which ladybeetles, other the to contrast in HLCRSNIGRITUS CHILOCORUS aeas ensgetdb atnhand Hattingh by suggested been also have o h eeomn fimtr tgsis stages immature of development the for  oou eti ni.Prolific India. in pest coconut a / 00scales/cm 60.0 v .arni,H lataniae H. aurantii, A. ittana psmellifera Apis Wle)frlaboratory for (Walker) tal. et Homalotylus 2 (Erichsen ) Á 92.Tediet The 1992). , / ihdniisof densities high inesor Linnaeus and p has sp. tal. et 387 M. Á , / Downloaded By: [NEICON Consortium] At: 10:31 11 September 2007 C A C REFERENCES assistance. of financial Council for and Pradesh suggestions Uttar valuable Technology, their certain and for providing Science referees and manuscript anonymous the two reading Pietermaritzburg, suggestions; critically Natal, valuable literature, of providing University for Samways, Africa Michael South Professor to thankful are Authors valuable ACKNOWLEDGEMENTS provide may reduce studies help and and failures. predator These the also intraspecific occasional the of of for performance. is utilization including possibilities factors diets result-oriented reproductive the abiotic and effective guild, of specific its cost more optimization a for The enhance and information lacking. Suitable in to are explored. predator predator needed be its scale this the and to of attributes, yet of augmentation reproductive is and role interactions biological heterospecific The its strategies. surrounding ambiguities adaptive the on needed D D E F C A B D D A F D A the to limited contribution mostly little is are there fact, studies on In biology. Indian India its subcontinents. from on information Indian brief with the records from distributional occurrence its on 388 A OLDI, ITZGERALD, RICHSEN, OOTH, HAZEAU, HAZEAU, ARTER, GARWALA, GARWALA, RGOV, NONYMOUS HMAD, AVIS, ORGE, IXON, E E hr tl eanmn asi h ieaueon literature the in gaps many remain still There B L oeta gnsfrbooia control. biological for agents potential hrsynshriWieand Weise horni Pharoscymnus glomeratus Phytoparasitica bibliography. nvriyPes abig,p 257. p. Cambridge, Press, University Clotr)ta taksaeiscs(o. Coccoidea). (Hom., scale-insects attack that (Coleoptera) Biologique destructor search. hi ilg,NtrlEeisadControl and Enemies Natural Biology, Their noooia Society Entomological Caledonie. Francais ultnd uemNtoa ’itreNtrle3 ei No. Serie 3e Naturalle d’Historie National Museum du Bulletin ultno noooia Research Entomological of Bulletin fAphidology of hioee sexmaculata Cheilomenes F)Clotr:Ccielde fadoe cl otresource. host scale doomed a of Coccinellidae) (F.)(Coleoptera: aglr (India) Bangalore 493 ACH OBEL I C A . Y R Á R . S 19)Mutn n cl oe omto,in formation, cover scale and Moulting (1990) .(d)(1964) (Ed.) P. , J M / . . 498. F . . . .(1978) A. , . J K G &R J . . C 17)Suisi etPksa ntebooyo n iiui pce n w ocnli species coccinellid two and species nitidulid one of biology the on Pakistan West in Studies (1970) . 17)Rcn nrdcin o ilgclcnrli aaiXVII. Hawaii in control biological for introductions Recent (1972) G ,E . C B noooi xeietlse Applicata et Experimentalis Entomologia . . B ,D fc el ehrh cetfiu tTcnqeOteMr eted oma Nouvelle Noumea, de Centre Outre-Mer, Technique et Scientifique Recherche la de Office . . . ,S enemies. natural of production for Technology (1995) . . 18)L ut ilgqecnr acnhnletasaet ucocotier du transparente conchinelle la contre biologique lutte La (1981) D . K 19)Arve fteseisresembling species the of review A (1998) K TIENNE, (2000) &D OSSLER, . 44 . Sgoe)axNuelsHbie Hmpea Diaspididae). (Homoptera, Hebrides Nouvelles aux (Signoret) .Labdev. G. V . AMWAYS, ALAYA, &G .nigritus C. . &Y rpclPs Management Pest Tropical ,11 15)Rve ftebooia oto fcciso oou am nteSeychelles. the in palms coconut on coccids of control biological the of Review (1953) 14 IXON, 16 HOSH, ,1 netPeao-Pe yais ayidBelsadBooia Control Biological and Beetles Ladybird Dynamics, Prey Predator- Insect Á ASUDA, otiuinal oniac eletmfuedicdneeooiu nPolynesie en economique d’incidence l’entomofaune de connaisance la a Contribution p 221. pp. , 303 , / J 22. . Y Á V &F / ilgclCnrlo netPssadWeeds and Pests Insect of Control Biological . 20. A . M P 18)Itouto fbnfiiliscsit salfrcnrlo netpests. insect of control for Israel into insects beneficial of Introduction (1988) 21 ora fSineadTcnlg (B) Technology and Science of Journal A Á . . F / . URSH, osbybcuesaeifsaini o ao rbe there. problem major a not is infestation scale because possibly , 316. J 187 , . Fb. Clotr:Ccielde ihawrdiecekito preys. of checklist worldwide a with Coccinellidae) (Coleoptera: (Fabr.) . &P K . H G . &H . . 18)Pe eod fahdpaosCcieldei ni.Arve and review A India. in Coccinellidae aphidophagous of records Prey (1988) 18)Frgn eaiu fccieldlra:drto n intensive and duration larvae: coccinellid of behaviour Foraging (1984) 20)Cmeiieaiiyo ayidpeaoso pis eiwof review A aphids: of predators ladybird of ability Competitive (2000) Á RADHAN, / H 190. ATTINGH, . hlcrsnigritus Chilocorus 17)LsCcnlia eIied aRuin(net.Coleoptera). (Insecta. Reunion la de I’ile de Cocinellidae Les (1974) 44 MA .PERVEZ A. & OMKAR 405 , ultno noooia Research Entomological of Bulletin 34 A Á . ,1 G V / 413. (R . . Á OSEN, 19)Aodneb h ladybird the by Avoidance (1991) / 17)Suiso w rdtr ocnli beetles, coccinellid predatory two on Studies (1972) 14. 36 ol rpPs,Vlm A rordSaeInsects: Scale Armoured 4A. Volume Pest, Crop World 133 , D hlcrsnigrita Chilocorus . a. edn nsgraescales, sugarcane on feeding Fab., .nigritus C. d) leir mtra,p.217 pp. Amsterdam, Elsevier, Ed.). , Á / 136. ultno noooia Research Entomological of Bulletin rjc ietrt fBooia Control Biological of Directorate Project 10 210 138 , hpa al odn p 844. pp. London, Hall, & Chapman . ora fApidEntomology Applied of Journal ,1 iehsoytat.Rsac is Research traits. history life Á Á / 297. / 141. 88 Clotr:Coccinellidae): (Coleoptera: 361 , rceig fHawaiian of Proceedings air RT,Serie ORSTM, Cahiers Á / 367. hlcrsnigritus Chilocorus Temnaspidotus Cambridge . Á Aspidiotus / 60 230. 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American in Chilocorus Coccinellidae). 289 Msel Hmpea isiia) nSuhr fia vdnefo aoaoyadfil trials. field and laboratory from evidence Africa: Southern Entomology in African scale, Diaspididae), red citrus (Homoptera: of enemy (Maskell) natural non-target a Coccinellidae), (Coleoptera: (Fabricius) 22 ocdpaosbooto agents biocontrol coccidophagous 19)Cci ope fccnti ai Nadu. Tamil in coconut of complex Coccid (1991) 64. p. 1989, July 13 Clotr:Ccielde ntelbrtr eaiet fetvns ntefil sbooto agents. biocontrol as field the in effectiveness to Entomology relative Economic laboratory of Journal the in Coccinellidae) (Coleoptera: ecito fanwspecies. new a of description omnelhIsiueo ilgclControl, Slough. Biological Royal, of Institute Commonwealth fteSe the of 118 aegoe fKcch(Gujarat). Kachchh of groves date altrablanchardi Parlatoria ,M.&K M HAMBERS, M 279 , S . I Á Á J G M . . . . / / K 119. 326. M R M &K . .(99 oaigo he predatory three of Foraging (1989) V. , &H . .(98 ilgclcnrlo h oou ipd( hispid coconut the of control Biological (1978) J. , . V V . V M &B V . V Á ..(91 eiwo ilgclcnrli h tipa region. Ethiopian the in control biological of review A (1971) D.J. , V V . P &S 17)Arve fteHwia Coccinellidae. Hawaiian the of review A (1976) . . E . / v . . C 306. . . . 62 S D R . &S &S &S 81 nhEtmlgclCnrs noooia oit fSuhr fia Pietermaritzburg Africa. Southern of Society Entomological Congress Entomological enth Chilocorus . RISHNAMOORTHY, &S &S . ONEK, N . &S 19)Booia oto fteccntsaeinsect, scale coconut the of control Biological (1991) &S M &S N . . LACKMORE, ladybirds. 720 , RISHNAMOORTHY J C C C 169 , INGH, . . R C . AMWAYS, AMWAYS, . . AMWAYS, p 210 pp. , . (1994) . ,T ,H M OUTHWOOD, &P M . . AMWAYS, &S AMWAYS, Oecologia M J AMWAYS, AMWAYS, ultno noooia Research Entomological of Bulletin . ,D . . Á HIRUMOORTHY, altrablanchardii Parlatoria . Á CLG N ICNRLOF BIOCONTROL AND ECOLOGY 19)Booyadfeigptnilo lc ete( black of potential feeding and Biology (1994) A 19)Saeiscso aepl ( palm date of insects Scale (1993) ,S ILL, / 721. / OPE, 174. S AMWAYS, . IXON, . Chilocorus Ladybirds P (1996) 8 ODAGAR, ot aicCmiso,Nue,NwCaledonia New Noumea, Commission, Pacific South p.(oepea Coccinellidae). (Coleoptera: spp. ,anwhs fbakbel ( beetle black of host new a ), . e eln Entomologist Zealand New ,47 Á M M M hlcrsnigritus Chilocorus 18)Boi oeta ftreccieldpeaoso aiu isiehosts. diaspine various on predators coccinellid three of potential Biotic (1989) M / 212. R P M M . . . . J J . G 45 M J M . A J Á . . D . . . M . / T . 19)Vsa n latr oaino itps rypths n individual and patches, prey biotopes, of location olfactory and Visual (1995) of establishment field for method effective most the of Determination (1991a) J 56. J clg fCoccinellidae of Ecology . 360 , 19)Asneo nrseicitreec uigfeigb h predatory the by feeding during interference intraspecific of Absence (1990) . . . F . . J . J R &W .(98 upeso ftesf re scale green soft the of Suppression (1998) A. , A M . ultno noooia Research Entomological of Bulletin . . . 3 19)Eauto fatfiildesadtoseiso aua ryas prey natural of species two and diets artificial of Evaluation (1993) some of and biology the on Studies (1977) G 19)Pyilgcladbhvorlcaatrsisof characteristics behavioural and Physiological (1994) aprClis odn .367. p. London, Collins, Harper . 18)Itrcin ewe nsadtecoccid, the and ants between Interactions (1980) . ,20 . 19)Pe hieadsbttto in substitution and choice Prey (1992) 19b ocdsic npe yedrn editoutosof introductions field during type prey in switch forced A (1991b) E N . spp. . J (1999) . . Á &H . S N 87 17)Frgn taeiso insects. of strategies Foraging (1978) / IGLEY, 365. . Chilocorus 20)Efcso netgot euaoson regulators growth insect of Effects (2000) Á ,M ln rtcinBulletin Protection Plant . Chilocorus / ,R ,31 23. noooi xeietlse Applicata et Experimentalis Entomologia ODEK, ). Aulacaspis OHANASUNDARAM, hlcrsnigritus Chilocorus AMDEVPUTRA, Á v ninJunlo giutrlSciences Agricultural of Journal Indian / 38. rnetlipiain n hut,Bnaoe ni,15 India, Bangalore, thrusts, and implications ironmental . P noooi xeietlse Applicata et Experimentalis Entomologia . J . I Chilocorus . d..Aaei ulsig h au,p.143 pp. Hague, The Publishing, Academic Eds.). , 19)Fez-re rica it o he pce of species three for diets artificial Freeze-dried (1992) Clotr:Coccinellidae). (Coleoptera: pce Clotr:Ccielde:i ta important an it is Coccinellidae): (Coleoptera: species o ,Cmowat giutrlBra,Farnham Bureau, Agricultural Commonwealth 5, No. Beha 15 HLCRSNIGRITUS CHILOCORUS p.(eitr:Dapdde nEs fia ihthe with Africa, East in Diaspididae) (: spp. rceig fteFrtNtoa ypsu nPest on Symposium National First the of Proceedings 82 hei dactylifera Phoenix ninCcntJournal Coconut Indian lwrAaei ulses odeh,Boston, Dordrecht, Publishers, Academic Kluwer . ,83 hlcrsnigritus Chilocorus v 375 , oradIpc fAphidophaga of Impact and iour clgclEntomology Ecological M rceig fHwia noooia Society Entomological Hawaiian of Proceedings Á p.(oepea ocnlia) in Coccinellidae), (Coleoptera: spp. rpclPs Management Pest Tropical / rnip longissima Brontispa 87. . Á V nsialn whitefly. spiralling on M / 334. . 45 &P . ,C siitsdestructor Aspidiotus ,31 67 HINNIAH, ATEL, Á 259 , / 33. hlcrsnigritus Chilocorus n hi aua nme nthe in enemies natural their and ) nulRe Annual ). Á . P / Chilocorus ninJunlo Agricultural of Journal Indian 270. . K 22 C . ultno Entomological of Bulletin n uvyo ri flies fruit of survey a and ) . 69 17. , 15 19)Dt amscale palm Date (1992) ehia ultno the of Bulletin Technical &C 64 Coccus ,13 craseychellarum Icerya 385 , v netEn Insect e fEntomology of iew 270 , HINNASWAMI, hlcrsnigritus Chilocorus 37 Á 75 in (Homoptera: Sign. oiilaaurantii Aonidiella / p.(Coleoptera: spp. 20. Á 387 , ,87 / 390. Chilocorus Chilocorus Á ,apeao on predator a ), v / 271. iridis (P v Á Á Proceedings OLGA ironment / 98. / 389. (Green) ´ Á R, / 148. ,10 K 389 spp. spp. Á . 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J J &H 14)Booia oto fsm eitru netpsso rp nIndia. in crops of pests insect hemipterous some of control Biological (1948) J . J . hlcrsnigritus Chilocorus . . J . . J . . hlcrsnigritus Chilocorus . . . . . ,O . 345 , . &T 18a pta n eprlpplto atrsof patterns population temporal and Spatial (1986a) . 18)Booyadeooi au ftesaepredator scale the of value economic and Biology (1984) &T &C &G (1998) &C 18)Ciaedarm n ilgclcnrl neapefo h rorpyo the of areography the from example an control: biological and diagrams Climate (1989) &W &M EKEROGLU 19)Ms ern n eesso h fetv cl predator scale effective the of releases and rearing Mass (1994) &C 6 &C M 15 S SBORN, ora fAd of Journal ENEN, .(02 clg fahdpaosld beetle, lady aphidophagous of Ecology (2002) A. , 1,101 (1), 607 ATE, . uky .Etmlj ogei idrlr,13 Bildirileri, Kongresi; Entomoloji I. Turkiye Á . .(03 ocnlisi netPs aaeet in Management, Pest Insect in Coccinellids (2003) A. , R OPLAND, ATE, ,87 Á / OPLAND, &C 19)Tcnqeo aspouto ftepeaoycoccinellid, predatory the of production mass of Technique (1993) REATHEAD, 562. / . OPLAND, 351. ILSON, OPLAND, B APP, ,4 . hlcrsnigritus Chilocorus B Á HANNA B 19)Rcrso pisntrleeiscmlxo ta rds.I.The IV. Pradesh. Uttar of complex enemies aphids-natural of Records (1995) / J Á . 95. A . R / . 5. Á J A 18)Frgn eaiu ftoseiso h genus the of species two of behaviour Foraging (1986) .(97 oeta s fipre aua nme o ilgclcnrlin control biological for enemies natural imported of use Potential (1987) E. , . . 14 . / 109. ,H 18)Ms-ern ftesaepredator, scale the of Mass-rearing (1986) . S M 18)Anwmto o h asitouto of mass-introduction the for method new A (1983) M ,11 18)Sxn of Sexing (1984) . J M v . M hlcrsnigritus Chilocorus ASTINGS, . v J rTois nSbrpeeGewasse Subtropiese en Tropiese ir . B J . ne Zoology anced D 18)Apcso h edn eaiu of behaviour feeding the of Aspects (1988) W . . ora fBiogeography of Journal . J ASAVANNA, W Á Fbiis 78 Isca oepea Coccinellidae). Coleoptera, (Insecta, 1798) (Fabricius, ihanwmto fetmtn odintake. food estimating of method new a with J (S siitsdestructor Aspidiotus . . . / W J . 23. W . . 20)Mxmmfeigptnilo aveadaut ftesaeinsect scale the of adults and larvae of potential feeding Maximum (2000) RIVASTAVA . 19)Efc ftmeaueo eeomn n mauesria in survival immature and development on temperature of Effect (1996) 17)Tesgraesaeinsect scale sugarcane The (1973) Phelsuma . 19)Evrnetliflecso eudt,egvaiiyadegg and viability egg fecundity, on influences Environmental (1998) 19)Ofcoyrsossb h cl netpredator insect scale the by responses Olfactory (1995) 630 H irs&SbrpclFutJournal Fruit Subtropical & Citrus ultno noooia Research Entomological of Bulletin MA .PERVEZ A. & OMKAR . Fbiis Clotr:Ccielde nCuieIln sixty Island Cousine on Coccinellidae) (Coleoptera: (Fabricius) ,9 5 &H ,91 G hlcrsnigritus Chilocorus icnrlSineadTechnology and Science Biocontrol Á .. d) nentoa okDsrbtn o,Lucknow. Co., Distributing Book International Ed.). R.P., , 75 / hlcrsnigritus Chilocorus . 14. P 6 16 ATTINGH, . ,87 Á 379 , 15)Bnfiilccield fMsr I. Mysore of coccinellids Beneficial (1953) ,67 / 105. F)(oepea Coccinellidae). (Coleoptera: (F.) ora fAphidology of Journal Á Pans / 88. Á Á / / 71. 393. Sign. 26 V 19 . 795 , 353 , ninCcntJournal Coconut Indian 19)Goa lmt hneadacrc of accuracy and change climate Global (1999) . Á BioControl Á / 6Ei 97 g U Ege 1987, Ekim 16 F)(Coccinellidae). (F.) / Á oiilaaurantii Aonidiella ora fApidEntomology Applied of Journal 812. 262 / 367. ocnlaseptempunctata Coccinella hlcrsnigritus Chilocorus 15. , uaapstegalensis Aulacaspis hlcrsnigritus Chilocorus 16 598 ipsiie n Bioagents and Biopesticides 76, 43 176 , hlcrsnigritus Chilocorus ,39 ,4 BioControl 671 5 Chilocorus ,83 Á Á / Á 6. / hlcrsnigritus Chilocorus 201. / Á icnrlSineand Science Biocontrol 52. / 23 Msel (Hemiptera: (Maskell) 683. Á ¨ / irs&Subtropical & Citrus 93. hlcrsnigritus Chilocorus ni ,12 urn Science Current hlcrsnigritus Chilocorus (F.)(Coccinellidae). ora fBiogeo- of Journal oiilaaurantii Aonidiella v riei Borno ersitesi, ninJunlof Journal Indian Á 45 comparative a : (F.)(Coccinelli- Zn. n its and (Zhnt.) / 13. 295 , (Coleoptera: Chilocorus 106 (F.)(Col., Á / 310. 177 , (F.) 17 v in a, Á . , /