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Biocontrol Science and Technology Publication details, including instructions for authors and subscription information: http://www.informaworld.com/smpp/title~content=t713409232 Predation by Halmus chalybeus (Coleoptera: Coccinellidae) on Ceroplastes destructor and C. sinensis (Hemiptera: Coccidae: Ceroplastinae) Infesting Citrus in Northland, New Zealand P. L. Lo; R. B. Chapman

Online Publication Date: 01 February 2001 To cite this Article: Lo, P. L. and Chapman, R. B. (2001) 'Predation by Halmus chalybeus (Coleoptera: Coccinellidae) on Ceroplastes destructor and C. sinensis (Hemiptera: Coccidae: Ceroplastinae) Infesting Citrus in Northland, New Zealand', Biocontrol Science and Technology, 11:1, 57 - 66 To link to this article: DOI: 10.1080/09583150020029745 URL: http://dx.doi.org/10.1080/09583150020029745

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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: 10:27 11 September 2007 icnrlSineadTcnlg 20)1,57±66 11, (2001) Technology and Science Biocontrol rdto by Predation a cl) u o hr instar third not but scale), both wax of instars second and fecoigctu rnhsi as on htlra n adult and larval technique that exclusion found an bags, using in experiments branches Field citrus Zealand. enclosing New Northland, of in orchards citrus on hl aveae9.7 ate larvae while uvvdps h eodisa tg where stage chalybeus instar H. second the past survived pce eenttse u r aey fee,atce by attacked ever, if rarely, are but tested not were species dlscnue naeae15.6 average on consumed Adults euto fsaepopulations. destructor C. scale of reduction Keywords: n ayidppltosand populations ladybird and ilgclcontrol biological aeokNrh e eln.Tel: Priv Centre, Zealand. Research Bay New Hawke’s important HortResearch, North, address: Present Havelock became Lo. L. P. They to: ago. Correspondence decades several Australia ( and citrus of Zealand pests New into imported C eolse destructor Ceroplastes INTRODUCTION SN05-17(rn)IS 3007 (online)/01/010057-10 1360-0478 (print)/ISSN 0958-3157 ISSN O:1.0005350027 5 10.1080/0958315002002974 DOI: e Zealand; New rdto by Predation . 1 h otclueadFo eerhIsiueo e eln,Whangarei, Zealand, New of Institute Research Food and Horticulture The sinensis ocde eolsia)IfsigCtu nNorthland, in Citrus Infesting Ceroplastinae) Coccidae: on ( eevdfrpbiain1Nvme 99 eie aucitacpe etme 2000 September 8 accepted manuscript revised 1999; November 1 publication for Received eolse destructor Ceroplastes amschalybeus Halmus e uri Cieewxsae Hmpea ocde eeaccidentally were Coccidae) (Hemiptera: scale) wax (Chinese Guercio Del amschalybeus Halmus and nsettled on Citrus , 2 .sinensis C. ..Bx8,Lnon atruy e Zealand New Canterbury, Lincoln, 84, Box P.O. ol ln n clgclSine iiin icl University, Lincoln Division, Sciences Ecological and Plant Soil, citrus .destructor C. amschalybeus Halmus p. nbt onre eas h oedwte xrt evsa a as serves excrete they honeydew the because countries both in spp.) Nwta)(ht a cl,sf a cl nNwZaad and Zealand) New in scale wax soft scale, wax (white (Newstead) , W e Zealand New .L LO L. P. s n eodisa clswsetmtdisd agdbranches. bagged inside estimated was scales instar second and rst eolse destructor Ceroplastes hnisatvt sntdsutdb pesticides. by disrupted not is activity its when , W .chalybeus H. + selleldbr)o w pce fwxsaewsstudied was scale wax of species two on ladybird) (steelblue l xeiet hwdta e clso ihrspecies either of scales few that showed experiments eld .destructor C. eolse destructor Ceroplastes e a.Teefeigrtsacutdfrteexperimental the for accounted rates feeding These day. per 468789;Fax: 8196; 877 6 64 .destructor C. 1 e Zealand New and .B CHAPMAN B. R. .chalybeus H. and hr instar Third . saueu aua nm o h oto of control the for enemy natural useful a is n 13.3 and wiewxsae and scale) wax (white Clotr:Coccinellidae) (Coleoptera: + .sinensis C. 468746;Emi:[email protected] E-mail: 4761; 877 6 64 € .chalybeus H. , a ueos h edn aeof rate feeding The numerous. was 01Tyo rni Ltd Francis & Taylor 2001 .sinensis C. eolse sinensis Ceroplastes .sinensis C. .chalybeus H. 2 (Hemiptera: h apigo scale of sampling The . e a e ladybird, per day per .sinensis C. n dlso both of adults and rydon preyed predation , t a 1401, Bag ate (Chinese ) W rst , Downloaded By: [NEICON Consortium] At: 10:27 11 September 2007 ncieaut a efudsetrn nocad.Ti omnyi o rediapause true controlling in a facultative a of not as clusters is described is small dormancy and This 1995). winter rise temperatures orchards. (Flynn, when During once oligopause in summer immediately 2000). resume sheltering early (Lo, can found in activity population since be abundant active the are can most of Adults adults are 1995). majority inactive larvae (Flynn, the while Zealand comprise Northland, New in they northern in year year the per throughout generations two least at H 90 ln,19) eeomn rmegt dl okama f3 asa constant a at 1995). days (Flynn, laboratory 37 the of armoured in mean and weeks a 22 took (Coccidae) for adult lived soft Gordon, to females & egg two both Drea from and 1983; Gellatley, 25ëC, Development includes & 1995). Beattie prey Flynn, 1967; (Valentine, 1990; its mites and group; scales, ) this (Diaspididae among s polyphagou L,20) tcmrsd9%o h ayid nvsa assessments. visual in ladybirds the of but 97% Zealand, comprised New It 2000). Northland, (Lo, in orchards citrus forestieri oto civd Hely achieved. control 90 in,18)and 1981) Milne, 1970; ouain of populations eti,pr.cm. 99.Fv fteeseiswr motdit e eln as Zealand New into imported were species established, (Cameron these became aphids and of species mealybugs insects, Five scale other 1999). for predators comm., pers. Beattie, opee ylt aur.Lv eodisasaefudutlArl hr ntr and instars Third comparison, April. In respectively. until March, found are and February instars from second Live develop January. adults late by completed ilgclcnrlaet fsaeiscsfroe etr.I utai,sxseisof species six Australia, In century. a over for on insects preying October. scale recorded been to of have ladybird through respectively. agents onwards, occur June control instars and biological March second from live found and are adults March, and and instars February Third in mainly hatch C nombl rwesta ipreo olae n ete Both settle. body and female’s leaves which the 1999). to under size comm., from on scale pers. hatch average disperse Beattie, eggs the that C. The on crawlers A. 1995). depends G. mobile (Lo, and 1995; into orchards (Lo, fewer between considerably eggs greatly is 5000 varied over fecundity lay average can The species both and gns oadCamn(98 eotdo h aaiod n uglptoesthat pathogens fungal and parasitoids the on control biological reported as (1998) act Chapman could attack that and present Lo already agents. species utilising preferably programme, these claims. of their None support ladybirds. of to data elimination presented the to authors applications insecticide following populations ntr eoestln emnnl.Nihrseisifsstefuto irstes unlike trees, citrus of fruit the infests species Neither scales. permanently. armoured settling before instars ac ± otsbfr hs of those before months 2±3 hatch tgswr unrbeto vulnerable were stages H 92.Aut cu n12ya l tm,atog bu 0 fadult of 10% about although stems, old (Lo Northland year in 1±2 leaves citrus and on on photosynthesis developed occur reduces and Adults fruit 1972). the to 1993). injury Brink, cosmetic 1986; (Brun, causes yield that fruit layer black ( a fungi with mould sooty for substrate 58 . . . utainsuishv rdcdcnitn ttmnso h motneo ladybirds of importance the on statements con¯icting produced have studies Australian H ayid Clotr:Ccielde aebe sdscesul rudtewrdas world the around successfully used been have Coccinellidae) (Coleoptera: Ladybirds h e eln irsidsr sdvlpn nitgae etaddsaemanagement disease and pest integrated an developing is industry citrus Zealand New The Both eae fbt pce aetreisasadassieautsae(iles 97 Pollet, 1967; (Cilliers, stage adult sessile a and instars three have species both of Females destructor chalybeus chalybeus . chalybeus C C . Mlat and (Mulsant) . destructor destructor ac anydrn eebr(al umr n rwe eteeti largely is settlement crawler and summer) (early December during mainly hatch sapeao fteetopss h betvswr odtriewihscale which determine to were objectives The pests. two these of predator a as a nipratpeao of predator important an was C . C eog otetieCiooiiadi aiet utai.I srelatively is It Australia. to native is and Chilocorini tribe the to belongs destructor . destructor and and C tal et rpoamsmontrouzieri Cryptolaemus C H . C . sinensis . and . sinensis amschalybeus Halmus oee,te loatiue eugnein resurgence a attributed also they However, . chalybeus 18)sae htpeaosrdcd u i o control, not did but reduced, predators that stated (1982) . sinensis C Capnodium . .L O&R .CHAPMAN B. R. & LO L. P. sinensis Ple,17)mv rmlae oyugwo sthird as wood young to leaves from move 1972) (Pollet, C nNrhad hssuyeaie h motneof importance the examined study This Northland. in . r nvliei e eln,but Zealand, New in univoltine are sinensis n t oeta au santrlenemy. natural a as value potential its and C . mt 17)adSobl 17)rpre that reported (1972) Snowball and (1970) Smith . destructor p..Svr nettoso ug oe plants cover fungi of infestations Severe spp.). tal et C (Lo . Bidvl selleldbr), ladybird) (steelblue (Boisduval) destructor ,19) erdcini parthenogenetic is Reproduction 1996). ., H tal et Sih 90 nwal 92 .A C. A. G. 1972; Snowball, 1970; (Smith, usn.Altreldbrsocrin occur ladybirds three All Mulsant. . chalybeus ,19) nNrhad h gsof eggs the Northland, In 1996). ., ihu niaigtelvlof level the indicating without , tal et a h oiatspecies dominant the was C . ,18) he fthese of Three 1987). ., destructor C H . C . destructor . chalybeus C sinensis . C (Snowball, Rhyzobius destructor . sinensis eggs eggs has Downloaded By: [NEICON Consortium] At: 10:27 11 September 2007 ouainsof Sampling Population s Population Ladybird and Scale METHODSMATERIALS AND est.Ec wgwsasmdt aeteae faclne ihadaee esrdat measured diameter the a calculate with to cylinder outer wood a the of of on area area m the the have 1±2 by to of mid-point. divided assumed its was height was immature branch a for twig per leaves from Each 50 scales density. collected and of scales, randomly 1991. Kerikeri, number adult February near were The for 1 radius canopy. leaves tree) and km and January per 2 16 (1 Branches a between branches scales. within 20 twice orchards or comprised nine once sample on ladybirds. scales Each size and for in scales sampled ha immature were scales, 1 Northland, adult about of to densities up the blocks Twelve compare to orchards citrus same aoaoyexperiment Laboratory 0900± between days Predation ®ne Ladybird on disturbed made not were orchard was per examined foliage counts were three block The or per h. two recorded. trees 1900 of 20 were total Approximately A adults orchard. time. the each and through pupae slowly walking larvae, while all which during aur 91t o®mta ohsae eepeaoso a cls igetneo( tangelo Single scales. wax of predators were stages both reticulata that con®rm to 1991 January edn aeof rate feeding ntr eevleal opeainby predation at to vulnerable kept were were instars on later. h lids experiments. 24 with recounted Field of dishes were replicates The scales seven treatment. the were adult and There daylight the dish. natural One of each food. under nine to without 20ëC and introduced overnight been treatment, was had held larval ones ladybird and the dead ®eld adult obviously the or any from instar after collected fourth counted were were ladybirds scales The The removed. dishes. Petri in placed were lsdbg,oewtotldbrsadoewt ± adult 1±2 with one and ladybirds without one bags, closed ottasomd nEprmn ,tasomto ftedt a nbet tblz the of stabilize predation to 1: unable Experiment was square- data was analysed. the scales were of of data number transformation rank the either end 2, tested so necessary and were was variances, where Experiment start means and ladybirds In the of 1996), transformed. at (Zar, homogeneity and test root counted The ANOVA. bags ratio was were by variance scales voile treatments analysed the of were and by terylene Data number trees, experiment. The long separate each branch. on cm of each located to 60 were assigned replicates by randomly The wide excluded. or cm introduced in 40 occur dominated. with can species covered species one scale site trial both each Although at trees. orchard, tangelo same on the conducted were experiments All nteocadwsasse v ie uigteeprmn y1-i iulsearches. settled visual of 10-min predation by 2: experiment Experiment the during population ladybird times The emergence. later. ®ve crawler weeks 10 assessed 4 of end was counted had start distal was orchard the leaves the treatments before the the with on 1990, two in end scales December other proximal settled 11 of the on the number at The commenced and branches trial to replicates The attached open. ®ve were left had bags open treatments The bag replicates. closed The bags. xeietwscnutdwith conducted was experiment ouainsof Population s o aheprmn,bace ihsmlrdniiso h eie cl tg were stage scale desired the of densities similar with branches experiment, each For Blanco C H 3 . . H destructor C chalybeus eiso he xeiet a odce odtriewihscale which determine to conducted was experiments three of series A . . paradisi chalybeus avladadult and Larval . RDTO FWXSAE YLADYBIRDS BY SCALES WAX OF PREDATION .destructor C. and n xmndisptnilfrcnrligsaepopulations. scale controlling for potential its examined and af)lae netdwt rtadscn instar second and ®rst with infested leaves Macf.) eeasse ycnutn 0mnsace e block, per searches 10-min conducting by assessed were C W H . s n eodinstar second and rst . destructor chalybeus H crawlers. . chalybeus H . rm1Fbur o2 ac 91 tan at 1991, March 27 to February 1 from chalybeus eesmlda h aetm n nthe in and time same the at sampled were h ortetet opie w with two comprised treatments four The w ute xeiet esrdthe measured experiments further Two . .destructor C. eetse ntelbrtr in laboratory the in tested were H . chalybeus and pnbg n no and bags open , .snni.This sinensis. C. C . destructor Citrus 59 Downloaded By: [NEICON Consortium] At: 10:27 11 September 2007 rnhswr one n1 pi 91 h rnhswr hnbge n eo w or two zero, and bagged then were branches The 1991. April adult 11 eight instar on two third counted every were of branches counted predation were 3: stages Experiment immature ladybird damaged All or count. searches. dead visual initial and 10-min scales the by adult before weeks All removed unbagged. were left scales or excluded ladybirds with ahadters oe oro h ashdautldbrsadtohdtidinstar third had two and ladybirds adult using had repeated was bags experiment The the later. days of 7 Four recounted were none. scales rest The larvae. the and each 91 2bace ih200±500 with branches 12 1991, dl ayid e a.Tediycnupino clswsetmtdfo h di eight the or from zero estimated was and scales scales of 1300±1800 consumption had daily The branches bag. These per 1992. ladybirds April adult March±3 28 between xeiet4 edn aeof rate feeding 4: Experiment reassessed. rhr ihhg ubr fldbrs twsrpae ewe pi n 4My1991 May 24 and April 5 between had repeated which was orchard It ladybirds. another of at numbers high with orchard 60 ewe h oso clso rnhswt n ihu ladybirds. e without 5: and Experiment with branches on scales of loss the between netdwt adult with infested f2. cls(E .)adarneo ±4sae.A vrg f05sae disappeared scales 0.5 of mean of average a predation An 1: were experiment scales. adults Field 2±64 nine ladybirds. of the without range for dishes a ®gures control and respective four 7.9) the The from (SEM h. scales 24 27.0 during of scales, 2±138 of range experiment Laboratory leaf. per scales instar about second and than Predation and ®rst Ladybird ®rst more six of was with than scales that Blocks fewer instar averaged 1(b)). and search second (Figure 10-min scales and ladybirds per ®rst adult ladybirds of of 30 of numbers density density with the However, correlated the 1(a)). inversely (Figure between scales relationship instar clear second no was Sampling There Population Ladybird and Scale and ®rst hatched. settled had and eggs RESULTS scales all when breeding later of Six weeks numbers 12 scales/bag). branches The counted to 18.0 densities. were assigned (mean scale randomly scales were low instar density ) ladybirds second and adult `low’ scales 8 high a of or both 4 had density 2, of `high’ half (0, treatment a and each having of as adults/bag) replicates categorized breeding were 35.7 branches (mean the Half production. crawler w ie Tbe2.I h rhr with orchard the In 2). (Table sites two a bu . ie oeautsae hnohrtetet Tbe1.Fu ek later, weeks Four di those 1). no than was (Table branches There treatments 4). these 3, on other 2, settled (treatments than scales ladybirds more to scales exposed times adult 12 approximately more were there times however, 1.5 about had hseprmn a odce wc,wt itntydi distinctly with twice, conducted was experiment This il xeiet2 rdto fsettled of predation ladybirds 2: of experiment number Field high count). a 2.5)/10-min was (SEM There 52.8 (mean ones. block whereas protected the branches, in on exposed present on observed instar subsequently third to were survived they scales No treatments. exposed on scales H . chalybeus V c fadult of ect C . sinensis h ee aveaea vrg f5. cls(E 31,wt a with 23.1), (SEM scales 54.0 of average an ate larvae seven The . eeitoue.Fu ek ae h ubr fsae were scales of numbers the later weeks Four introduced. were .chalybeus H. .chalybeus H. eecvrdwt ol aso ac 92a h tr of start the at 1992 March 3 on bags voile with covered were .destructor C. C C . . .L O&R .CHAPMAN B. R. & LO L. P. sinensis destructor W .dsrco.Tenme fsae nselected on scales of number The destructor. C. s n eodinstar second and rst C n e ayid.Bace eeete bagged either were Branches ladybirds. few and . nsaeppltos. populations scale on on destructor crawlers. eebge,wt afrciigtoladybirds two receiving half with bagged, were W s n eodisa scales. instar second and rst V n bnatldbrs(en70.4 (mean ladybirds abundant and h nlsdbace tetet1) (treatment branches enclosed The V rn uvvlo clsbtenthe between scales of survival erent rnei h ubr fimmature of numbers the in erence .destructor C. iial ie branches sized Similarly and n7March 7 On .sinensis. C. C . V sinensis erence Downloaded By: [NEICON Consortium] At: 10:27 11 September 2007 ih9%o hs rtce ybg.I otat hr ayid eesac ma 3.7 (mean scarce compared were ladybirds branches of where exposed proportion contrast, on similar In survived a bags. scales min), by the protected 0.3)/10 those half (SEM on than 90% fewer with min), 11.0)/10 (SEM IUE1 orltosbten()ma este faut n rtadscn instar second and ®rst and adult, of densities mean (a) between Correlations 1. FIGURE il xeiet3 rdto ftidinstar third of predation H 3: experiment Field branches. il xeiet4 edn aeof rate instar feeding with second disappeared, 4: trial. had experiment the scales survived Field the ladybirds of the 5% of about All later weeks present. Four ladybirds count. without initial and the in scales of . chalybeus b endniyo rtadscn instar second and ®rst of density mean (b) eesmlda h aetm nctu rhrsi otln ewe 6Jnayand January 16 between Northland in orchards citrus on time same 1991. February the 1 at sampled were H rydo hr ntrsae Tbe3.Altretetet a iia numbers similar had treatments three All 3). (Table scales instar third on preyed C . . chalybeus destructor bevddrn 0mnsace.Autadimtr clsadladybirds and scales immature and Adult searches. 10-min during observed RDTO FWXSAE YLADYBIRDS BY SCALES WAX OF PREDATION and C . sinensis .cayesTecag ntenme f®s and ®rst of number the in change The chalybeus H. nbace ihadwtotldbrsi shown is ladybirds without and with branches on C .dsrco.Teewsn vdnethat evidence no was There destructor. C. . sinensis C . destructor uvvdo xoe n protected and exposed on survived n ennme falsae of stages all of number mean and C . destructor and , 61 Downloaded By: [NEICON Consortium] At: 10:27 11 September 2007 h hnei ubro rtadscn ntrsaeo oto rnhs(.)was (1.0) 4) branches (Table control scales/day/ladybird on 15.6 ladybirds on of was adult rate for 31.2 scale a estimated of give instar were ®gure to rates ladybirds This second of feeding (32.2). number Similar ladybirds and the with by ®rst disappeared divided that then of number the number from in subtracted change the ayid 1. . 0. . 5.6 4.5 5.4 6.2 5.8 8.2 106.0 100.5 Decrease % 114.0 6.3 SEM 6.0 6.5 Final 112.3 104.2 120.5 adult of SEM rate feeding The 6 4. Table in 6 6 Initial ANOVA bags of Ladybirds No. 8 Ladybirds 2 Ladybirds 0 Treatment 0.1 E 3. TABLE 0.5 0.6 1.6 0.5 0.4 0.9 4.7 1.0 4.9 3.6 6.9 A. 3.6 2.2 25.6 26.2 25.0 10 E 6.4 8.2 2. 10 TABLE 22.3 36.5 4 3, Replicates 2, vs 1 Trt. 5 5 ANOVA bag No 4. open Bag ladybirds 3. 1±2 closed, Bag ladybirds 0 2. closed, Bag 1. Treatment E 1. TABLE 62 B. il xeiet5 e 5: experiment Field ladybirds. of mortality ayidsgicnl eue ubr fstldsae tbt este fautscales adult of densities both at scales settled of numbers reduced signi®cantly ladybird r.2v ,404n . ns 0.1 ns 0.4 4 vs 3 4 Trt. 3, vs 2 Trt. ramn o fbg E ia E Decrease % SEM Final SEM bags of No. Treatment C obg8417. 3 3621.6 53.6 16.2 334 35.5 74.8 347 431 51.7 8 421 8 ANOVA bag No ladybirds 0 closed, Bag C obg6534. 8 . 51.3 9.0 10.4 285 51.4 41.0 451 593 61.1 6 505 6 data) (rank ANOVA bag No ladybirds 0 closed, Bag . . sinensis destructor ae n8May 8 on later 1Dcme 90we aswr u n n ubr fstld®s ntr eeasse on assessed were instars ®rst settled of numbers and on, put 1991 were January bags 8 when 1990 December 11 eecutdo 1Arl19,when 1991, April 11 on counted were 4My19.Tetoocad a epciemaso 0ad4ldbrs1-i count ladybirds/10-min 4 and and April 70 5 between of B means trial respective and had 1991, orchards March two 27 The and 1992. February May 1 24 between conducted was A Trial V V V c fadult of ect of numbers low or high of ect of ect H . V chalybeus c of ect H . chalybeus .chalybeus H. nstlmn of settlement on F 5 .L O&R .CHAPMAN B. R. & LO L. P. nsria ftidinstar third of survival on . ns 1.7 H . F Initial H F chalybeus enn.adults no. Mean 5 ennme fscales/branch of number Mean s24ns 2.4 ns 0 . 10cm /100 5 P F P F n chalybeus . ns 3.1 ns 0 H ennme fscales/branch of number Mean nsaeppltos. populations scale on . C chalybeus 2 . odMa o rtinstars/leaf ®rst no. Mean wood destructor on < SEM nsria f®s n eodisa a scales. wax instar second and ®rst of survival on .561.1 0.05 C . eeitoue otebg,ad4weeks 4 and bags, the to introduced were destructor F Adult . C 5 . C . ns 1.0 F F destructor . 5 5 sinensis 6.2 . ns 0.1 C n a acltda follows: as calculated was . destructor l he este of densities three All niecoe as Scales bags. closed inside cls hr a no was There scales. P < Ï 0.05 x eecutdon counted were < SEM 0.001 . Downloaded By: [NEICON Consortium] At: 10:27 11 September 2007 hc eue clst iia ees l ayid eeaiea h n ftetil There trial. the of end the at alive were ladybirds All di levels. a similar was to scales reduced which B. and di A. mn ramns13n 42.3 0.2 27.2 ns 0.6 743 1.3 28.4 e greater a 2.7 had 821 treatment ladybird eight The 5). 40.0 (Table 1.9 6 Final 18.6 Ladybirds 0 treatments 1. Among SEM 6 ANOVA Initial Ladybirds 0 1. bags High Treatment Low density Scale E 5. TABLE hssuydmntae that demonstrated study This DISCUSSION approximately consume would ladybirds scales two (26 experiment, the scales throughout 2300 day per scales 13 of aey fee,pe for prey ever, if rarely, ete rtadscn ntr ftetosaeseis lhuhpeaino hr instar third of predation Although species. scale two the of instars C second and ®rst settled r s2 ,4 3, 2, vs 1 Trt neato r sDniy14n . ns 1.2 ns 1.4 92.2 Density vs Trt Interaction densities scale Between immature ramn asIiilSMFnlSMn.dyFeigrate Feeding no./day SEM Final SEM Initial bags Treatment of scales/day/ladybird) of (number rates Feeding 4. TABLE ra n cl pce o of No. species scale and Trial ee enfeigo dlso ihrseis is n eodinstar second and First species. either of adults on feeding seen never C . . V C ayid 14011017. 103.0 1178.0 121.0 1164.0 2 Ladybirds 0 dl ayid 5505. 8. 129.7 881.9 53.2 1505.0 8 ladybirds Adult 8 avlldbrs24409. 6. 48.0 261.0 98.0 404.0 2 ladybirds Larval 2 C ayid 4. 9026236.5 236.2 39.0 243.5 6 Ladybirds 0 dl ayid 1. 8515077.2 185.0 48.5 410.5 4 ladybirds Adult 2 sinensis sinensis rnecudb cone o yteetmtdfeigrt.Asmn osatrate constant a Assuming rate. feeding estimated the by for accounted be could erence . . C sinensis destructor . sinensis V rneo pt bu 0 clsstldo rnhswt r28ldbrs This ladybirds. 2±8 or 0 with branches on settled scales 800 about to up of erence H este fautsae.Tenme fimtr clswsasse 2wesltro 9May 29 on later weeks 12 assessed was scales immature of number The scales. adult of densities rnho ac 92 when 1992, March 3 on branch cls ra on A Trial scales. a odce ewe 8Mrhad3Arl1992 April 3 and March 28 between conducted was n dl clso ihrseiswsnteprmnal etd hs tgsare stages these tested, experimentally not was species either of scales adult and ert rihwxta nyprilycvr h oy hra hr instars third whereas body, the covers partially only that wax dryish a secrete V . c fadult of ect chalybeus nNwZaad ohautadlra ayid rydo rwesand crawlers on preyed ladybirds larval and adult Both Zealand. New in .8Ldbrs63. . 3350.2 0.8 3.5 0.6 9.8 0.1 10.7 23 0.5 0.4 192 0.9 202 2.9 3.9 3 3.6 4.1 27 2.5 30 30.7 34.5 37.5 1.4 0.7 6 2.3 6 19.3 6 16.0 Ladybirds 8 4. 18.0 Ladybirds 4 3. 6 Ladybirds 2 2. 6 6 Ladybirds 8 4. Ladybirds 4 3. Ladybirds 2 2. 3 eeosre edn ntidinstar third on feeding observed were H 7days). 87 H . RDTO FWXSAE YLADYBIRDS BY SCALES WAX OF PREDATION C . chalybeus . chalybeus destructor H . chalybeus o of No. nstlmn of settlement on ennme fscales/branch of number Mean uigmn or fsuy(o 94,autand adult 1994), (Lo, study of hours many During . a odce ewe ±4Mrh19.TilBon B Trial 1991. March 7±14 between conducted was H . chalybeus enn.bedn enn.lv immature live no. Mean breeding no. Mean sa motn aua nm of enemy natural important an is . s122.3 ns 2.4 clsbac scales/branch scales/branch P F P F C eeitoue,frbace ihlwadhigh and low with branches for introduced, were . sinensis < H .0 . ns 4.6 0.001 . chalybeus V ubro reigsae e bagged per scales breeding of Number . c hntoo orladybirds, four or two than ect C . sinensis n®s n eodisa wax instar second and ®rst on hnein Change 2 2 2 0. 13.3 103.9 nyoc n were and once only + 2 049.7 20.4 2215.6 32.2 C . Ð 2.3 . Ð 1.0 . Ï destructor x C . destructor C < < . SEM sinensis 0.001 0.001 and 63 Downloaded By: [NEICON Consortium] At: 10:27 11 September 2007 rdc di a produce 64 nadults. in hce n opeelyro a.Nvrhls,teltrsae fwxsae r not are scales wax of stages of later larvae The the ladybirds. Nevertheless, by wax. predation of to invulnerable layer necessarily complete and thicker on for found ntr hc a iia otenmesi h oto ramn.Peainby third Predation to treatment. survive control would the branch in per numbers scales the 400±800 to mortality similar of was level which instar, this At 1994). (Lo, instar ae ntr of instars later atrfrwxsae.Lgcly hr hudb oiiecreainbtentenumber the between correlation positive a be should there adult Logically, of scales. wax for factor hrfr rdc netmtd2 0 n 500eg e rnh nteasneof absence the In branch. per eggs by 000 woul produced 45 experiment eggs this and the in of adults 000 23 98.3% of estimated densities estimated an `high’ and predation, an `low’ The produce 1995). (Lo, therefore eggs 1268 be to ol edt d ute .±.%mraiyt euetenmeso clso branches on scales 5. of Experiment numbers the in reduce observed to those mortality to 1.3±1.6% ladybirds further with a add to need would ewe h este fadult of densities the between cl tgsaepeet rdto by Predation present. are stages scale some and for time su leaves than in the more distributed on evenly remain occur. not factors more did environmental been this data), by have and the d unpublishe killed time would were Lo, scales factors losses environmental L. Furthermore, then If (P. space. population. factor, branches the mortality on exposed unevenly main other acted disappeared than mortality scales that more branches Also, showing leaves. exposed the fragments on wax abundant. some left of were were remains that from numerous ladybirds ladybirds scales the by was where instar feeding predation with second likely branches of consistent and were importance exposed ®rst the scales on of on few evidence survived predation found Further where scales (1994) orchards Lo few two which investigated, the at was In levels maturity. to to species survive both to of pattern. scales observed instar the second cause to unlikely therefore Mendel were 1981; and manner Milne, environmental to density-independent vulnerable 1980; suggested highly ladybirds are (Hill, of crawlers abundance scale the conditions Although and predation. was scales correlation inverse factor of this The generation that population. next each the on of unequally settlement acted between factor mortality a that suggested oml nEprmn ,tefcniyo naeaeszdadult than branches sized searching average to time an less con®ned of spend were to fecundity likely ladybirds the were 5, the they Experiment because Consequently prey. In high of normal. density arti®cially high been a have with may rate the philippiae tasmlrtm fya as year of time similar a at h ucsflbooia oto fteepss(o 94.Fwsae fete species either of scales Few 1994). numerous. (Lo, was ladybird pests this where these stage of second-instar the control past survived biological successful the and fteetoseis di A species. two these of ral oteetn htte r o cre(.Hry .Wlet,pr.cm. 1999). are by comm., fungicides scales pers. copper wax Willetts, about and of brought J. insecticides predation been Harty, have to e Organophosphat may (A. programmes. disruptive status highly spray pest scarce in in now change changes are this by that they evidence circumstantial that is extent There the to greatly a nrdcdagainst introduced was w fteitoue pce fecri ap Mle 91 Sands 1981; (Milne, wasps encyrtid of species introduced the of two ouainsmln rvddidrc vdneta ayid eea motn mortality important an were ladybirds that evidence indirect provided sampling Population ietbesuisof studies table Life h siae edn aeo adult of rate feeding estimated The that showed experiments ®eld The ic hssuywscnutd h bnaceo a clsi otln a declined has Northland in scales wax of abundanc e the conducted, was study this Since C . sinensis C ivsr Hmnpea peiia),wihi h nyprsti of parasitoid only the is which , Aphelinidae) (Hymenoptera: Silvestri H . Ceroplastes destructor . V Y chalybeus rn yeo a htosue h hl oslsraeadepnsgreatly expands and surface dorsal whole the obscures that wax of type erent eolse rubens Ceroplastes nNwZaad(o&Camn 1998). Chapman, & (Lo Zealand New in in oacutfrtedsperneo clsi xeiet5 However, 5. Experiment in scales of disappearance the for account to cient C n h ubro rwespoue,a adn n ai (1984) Ravid and Yardeni as produced, crawlers of number the and X C a rsmbydtre rmatcigteeltrsae ytheir by stages later these attacking from deterred presumably was . oridensis . destructor V C destructor rn iuto xssi utai,weeasieo parasitoids of suite a where Australia, in exists situation erent . destructor C . osoki sal nta,telc farelationship a of lack the Instead, Israel. in Comstock akl Rcad,1981). (Richards, Maskell .L O&R .CHAPMAN B. R. & LO L. P. destructor and h eln fteps hr a trbtdlreyto largely attributed was there pest the of decline The . H H oislra r otaudn hntevulnerable the when abundant most are larvae its so , . H C chalybeus . . . chalybeus chalybeus sinensis n hi ete rgn mns orchards amongst progeny settled their and tal et H . opeet h ciiyof activity the complements chalybeus hwdthat showed ,18) hs atr c nalargely a in act factors these 1984), ., ol eueppltoso rtand ®rst of populations reduce could on C E C . . philippiae . sinensis sinensis L ln,19;L,1994), Lo, 1992; Blank, & (Lo H . C R aldt ec h third the reach to failed chalybeus . . tal et tak h ae stages later the attacks nEprmn ,was 4, Experiment in sinensis forestieri H ,1986). ., . chalybeus a estimated was a rca to crucial was C Euxanthellus H ryo the on prey . . destructor chalybeus breeds d Downloaded By: [NEICON Consortium] At: 10:27 11 September 2007 B B David REFERENCES Charles, John manuscript. by the New Comments improved the referees funding. and anonymous Technology and provided Clapperton and Council Kay Science Steven, Planning Research David by for Citrus of analysed Foundation were Planning Zealand Zealand data sites. The trial New Blank. supplied The Roddy also with Saville. who discussions Hibbert- Willetts from Miles John bene®ted especially and experiments collected, Stang were Murray samples Rhodes, orchards Lyn Foy, whose from growers the thank We ACKNOWLEDGEMENTS insects. u h s fteepout a elndi eetyas optbepsiiespray pesticide compatible A years. recent in e declined the maximize has to products needed these is of programme use the but C B D C H L F H L L L L L M M P ollet, rink, eattie, run, o, lynn, o, o, o o, o, illiers, ameron, rea, ely, ill, endel, ilne, ..(00 pce n bnac fldbrs(oepea ocnlia)o irsocad in orchards citrus on Coccinellidae) (Coleoptera: ladybirds of abundance and Species (2000) P.L. , P P P P giutr,NwSuhWls Australia. 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G & D L . . ., enman, oansug ot fia 8Jn± uy19,p 130. p. 1993, July June±1 28 Africa, South Johannesburg, , . 19)Terl fprstisadetmptoei ug nmraiyo third- of mortality in fungi entomopathogenic and parasitoids of role The (1998) osen, R & noooia eor fteDprmn fArcluevl 3 Technical 13. vol. Agriculture of Department the of Memoirs Entomological ora fteAsrla noooia Society Entomological Australian the of Journal 19)Ccielde in Coccinellidae, (1990) .sinensis C. V ellatley, RDTO FWXSAE YLADYBIRDS BY SCALES WAX OF PREDATION alentine, V osen, J D c fpsiie npeaino otwxsaeb h te-leldbr,in ladybird, steel-blue the by scale wax soft of predation on pesticides of ect . G (C 24, D . d) leir mtra,p.19±40. pp. Amsterdam, Elsevier, Ed.). , e eln Entomologist Zealand New 49, . . avalloro, R 18)Mt et fctu.Afc 2A.,6p.Dprmn of Department pp. 6 H2.AE.3, Agfact citrus. of pests Mite (1983) 315±321. D Hmpea ocde nctu.PDtei,LnonUniversity, Lincoln thesis, PhD citrus. on Coccidae) (Hemiptera: ) . J 939±957. . 19)Peooyadrltv bnac of abundance relative and Phenology (1996) . G E and 18)Pplto yaiso h eierna lc scale, black Mediterranean the of dynamics Population (1984) . V . W (1982) c npooytei ae fcitrus. of rates photosynthetic on ect . .sinensis C. craseychellarum Icerya V T & R cieesof ectiveness eolse destructor Ceroplastes . netPsso ri n eealsi e ot Wales South New in Vegetables and Fruit of Pests Insect D & homas, roe cl net,TerBooy aua Enemies Natural Biology, Their Insects, Scale Armored 8, 573±582. i Hmpea ocde eolsia)o irsin citrus on Ceroplastinae) Coccidae: (Hemiptera: M eolse ceriferus Ceroplastes W artino, Ceroplastes . 18, P H . elntn e eln,1±3August 11±13 Zealand, New Wellington, , ora fteEtmlgclSceyof Society Entomological the of Journal Mraoia:Hmpea nAldabra on Homoptera) (Margarodidae: (1987) 63±69. . chalybeus rboeaimandersoni Cribrolecanium E . d) ..Blea Rotterdam, Balkema, A.A. Eds). , n hns a cl ( scale wax Chinese and ) acri destructor Gascardia netbae motdit New into imported Invertebrates pce Hm ociee and Coccoideae) (Hem. Species 20, amschalybeus Halmus 167±170. Fbiis and (Fabricius) sapeao fscale of predator a as e eln ora of Journal Zealand New rceig fte9th the of Proceedings eolse destructor Ceroplastes eolse destructor Ceroplastes tKenthurst, at ) e Zealand New (Hemiptera: (Boisduval) Ceroplastes .sinensis C. Integrated from 65 ) ) . Downloaded By: [NEICON Consortium] At: 10:27 11 September 2007 R S 66 S S Z S Y V ands, mith, nowball, nowball, ar, alentine, ichards, ardeni, hi ilg n au o cl netcontrol. insect scale for value and biology their Hmpea ocie:Ccia) h hss ignaPltcncIsiueadSaeUniversity. State and Institute Polytechnic Virginia thesis, PhD Coccidae). Coccoidea: (Homoptera: sinensis oto of control oia Society logical Australia, fScience of fcalr n h orsodn netto eeomn nactu grove. citrus a in development infestation corresponding the and crawlers of Hmpea ocie:Ccia) nesenAustralia. Entomology eastern of in Coccidae), Coccoidea: (Homoptera: J . H D D . . . A P (1996) 17)Wiewxsaeadiscontrol. its and scale wax White (1970) A . . E A G G . R & e uri nVrii nldn eecito of redescription a including Virginia in Guercio Del M . ., W . . ora fteAsrla noooia Society Entomological Australian the of Journal 10, J J acri destructor Gascardia . L . . . (1981) isaitclAnalysis Biostatistical avid, ukins, 16)Als ftehsso noohgu net fNwZealand. New of insects entomophagous of hosts the of list A (1967) (1970) 17)Sau fntrleeiso ht a scale, wax white of enemies natural of Status (1972) abra utai,2±0Ags 92 .212. p. 1972, August 22±30 Australia, Canberra, , 1100±1209. 25, 51±59. M hzbu ventralis Rhyzobius R . eolse sinensis Ceroplastes . 18)Foiawxsae( scale wax Florida (1984) G . S & nowball, Nwta)(eitr:Coccidae). (Hemiptera: (Newstead) rnieHl,NwJersey. New Prentice-Hall, . .L O&R .CHAPMAN B. R. & LO L. P. Eiho)and (Erichson) G e uri Hmpea ocde,awxsaenwto new scale wax a Coccidae), (Homoptera: Guercio Del . J . 18)Aet nrdcdit utai o h biological the for Australia into introduced Agents (1986) uesadArclua Journal Agricultural Queensland ultno noooia Research Entomological of Bulletin Ceroplastes .forestieri R. rceig fteXVItrainlCongress International XIV the of Proceedings 9, X 57±66. oridensis Mlat Clotr:Coccinellidae), (Coleoptera: (Mulsant) Ceroplastes ora fteAsrla Entomo- Australian the of Journal osok oioigb capture by monitoring Comstock) acri destructor Gascardia Alon-Hanotea 96, 704±708. X e eln Journal Zealand New 71, oridensis 33±46. 39, Comstock 121±123. (Newst.)