BioControl 43: 325–338, 1998.© 1998Kluwer Academic Publishers. Printed in the Netherlands.

The life history and host range ofEctaga garcia, abiological control agent for Lantana camaraandL. montevidensisin Australia

M.D. DAY1, B.W. WILLSON2 and K.J. LATIMER1

1Alan Fletcher Research Station, P.O. Box 36, Sherwood, Qld, 4075, Australia;2present address: 33 Ben St, West Chermside, Qld, 4032, Australia

Received 3 April 1998; accepted in revised form 22 December 1998

Abstract. The life cycle and host specificity ofEctaga garciawere investigated. Mothsemerge in the morning and are inactive during the day. Eggs are laid and larvae feed onthe undersurface of leaves ofLantana camaraandL. montevidensis. Larvae spin protectivecocoons from which they feed and in which they pupate. Development from egg to adulttakes approximately 48 days. Forty-five plant species were tested to determine host speci-ficity. Females laid eggs only onL. camaraandL. montevidensis. In no-choice trials, neonatelarvae fed but failed to complete development on nine test species.E. garciawas subsequentlyapproved for release in Australia.

Key words: creeping lantana, host specificity, leaf-roller, Lepidoptera, Oecophoridae,phytophagous, varietal preference

Introduction

Lantana camaraL. andL. montevidensis(Sprengel) Briquet (Verbenaceae)are native to tropical America.L. camarais a serious weed of Mexican orCaribbean origin (Stirton, 1977) affecting cropping lands and forest areas in47 countries, including Australia (Parson and Cuthbertson, 1992). It was firstrecorded in Australia in 1841 and is now found throughout the coastal andsubcoastal areas of eastern Australia, from northern Queensland to south-ern New South Wales (Swarbrick et al., 1995).L. camaraeasily hybridisesand was cultivated for several centuries in Europe prior to its introductionto Australia and other countries. Several hundred varieties now exist world-wide (Parson and Cuthbertson, 1992) with 29 present in Australia (Smithand Smith, 1982).L. montevidensisis native to southern Brazil and northernArgentina. It has been introduced to many tropical and subtropical countriesas an ornamental, however, it is only considered a weed in Australia, where it

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is found along coastal and subcoastal areas of Queensland and northern NewSouth Wales (Holm et al., 1977; Swarbrick et al., 1995).

In Australia, bothLantanaspp. have seriously affected agricultural areasreducing productivity, whileL. camara has also invaded many nationalparks. In national parks,L. camarais quick to colonise gaps where it maybecome the dominant understorey species (Parsons and Cuthbertson, 1992).In pastoral areasL. camarareduces available grazing lands, affects mustering,and some varieties are poisonous to cattle and sheep. Costs ofL. camaraare estimated to exceed Au $7.7 million annually, including stock deaths ofup to 1500 head per year (Culvenor, 1985; Parson and Cuthbertson, 1992).L. montevidensiscolonises stony ridges, roadside embankments and otherdisturbed areas, spreading to surrounding pastures. The plant is a seriousproblem in grazing areas of central Queensland, outcompeting favourablespecies to form monocultures, and rendering pastures unproductive (Parsonsand Cuthbertson, 1992).

The control ofLantanaspp. is difficult, as many infestations are too largeto economically use conventional means such as herbicides. The widespreadburning or slashing ofL. camarais not possible in conservation areas whilemany other areas are inaccessible. The use of chemicals is not economicallyfeasible on large infestations although a number of herbicides are effec-tive when treating regrowth after slashing. Isolated plants can be removedmechanically before infestations become too large (Parsons and Cuthbertson,1992).L. montevidensiscan be controlled by repeated herbicide applicationsand sowing with perennial pasture species, however the cost per unit ofproduction is often uneconomically high (Parsons and Cuthbertson, 1992).Burning is not a viable option as there is insufficient fuel to maintain a fireand the root system may not be killed.

While biological control ofL. camarahas been attempted in Australiasince 1914, no agents have been released to controlL. montevidensis(Julien,1992). In recent field surveys in southern Brazil, a number of agents werefound to attack several species of lantana (Garcia, unpubl. data). The larvaeof one agent, the leaf rolling mothPyramidobelasp., was found to heavilydamage leaves ofL. camara, L. fucataLindley andL. tiliaefolia Chamisso,causing stunted growth and a reduction in flowering on a seasonal basis.The insect was imported into Australia asPyramidobelasp. and was laterdescribed asEctaga garciaBecker (Lepidoptera: Oecophoridae). Winder andHarley (1983) reported finding anEctagasp. attacking severalLantanaspp.during their surveys but it has not been determined if this is the same species.It is hoped thatE. garciamay establish and assist in the control ofL. camarain areas where other agents have failed and that it may aid the control of

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L. montevidensis. This paper reports on the biology and host specificity ofE. garcia.

Materials and methods

Biology and rearing

Sixty-five larvae ofE. garcia were collected from leaves ofL. fucata inCuritiba, Brazil and imported into an insect quarantine unit (27◦C day and22 ◦C night with natural lighting) at the Alan Fletcher Research Station(AFRS), Sherwood, Queensland, Australia in June 1991. From this ship-ment, a colony was maintained to conduct biology and host specificity tests.A further shipment of 90 larvae and 81 pupae collected fromL. fucata inCuritiba, Brazil was imported in May 1997 to facilitate field releases after theoriginal colony died out in January 1994.

Imported larvae were transferred to clear, plastic containers (265 mm×185 mm× 85 mm) lined with dry paper towel. Fresh leaves ofL. montevi-densisand of the five common varieties (pink, pink-edged-red, white, red andorange) ofL. camarafound in south-east Queensland were provided. Thesevarieties were determined using the key by Smith and Smith (1982). Thedifferent lantana varieties were provided so larvae could feed on preferredhosts if they were selective. The leaves were changed daily and pupaewere collected and held in petri dishes lined with dry filter paper for adultemergence.

Female moths could be distinguished by their abdomens which end in atube, while in males, the end of the abdomen is obliquely truncated. Approx-imately six male–female pairs of newly emerged adults were placed in eachof eight organza-screened cages (900 mm× 550 mm× 870 mm) with onepotted plant ofL. montevidensisand one potted plant of each of the fivecommon varieties ofL. camaraas mentioned above. A choice in lantanavarieties determined whether adults had preferred hosts for oviposition andon which varietiesE. garcia could be reared and released. Such tests areimportant as earlier field studies have shown that some previous agents intro-duced to control lantana have displayed differences in their preference andperformance on some lantana varieties (Haseler, 1966; Taylor, 1989).

The initial number of larvae present on each variety of lantana after allthe adults had died was recorded. Counting eggs though preferable, was toodifficult without damaging the plants. The cages were left for five weeks,by which time the developing larvae were nearing maturity. The plants werethen harvested and the larvae placed in plastic containers with leaves of thesame variety from which they had been removed. Clean containers and fresh

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foliage were provided three times per week. Pupae were collected and thenumber that developed on each variety was recorded and the mean percentof larvae that completed development was calculated. Pupae were sexed andkept separate according to their host plant.

Five male–female pairs of newly emerged adults were placed in smallcages (400 mm× 400 mm× 900 mm) containing one potted plant ofthe species or variety of lantana on which they had developed, in order todetermine whether a particular species or variety of lantana could sustainpopulations ofE. garcia. Five replicates of each variety or species were setup and the plants were left for five weeks before being harvested. Larvae andpupae were collected and treated as above.

Adult survival and fecundity were determined by placing up to 13 newlyemerged females, singly with 2–3 males, in a small gauze cage for 24 hoursto initiate mating. Cages contained cut tips of lantana of the variety on whichboth the males and female had been reared. Fresh flowers were provided as afood source. The adults were then transferred to cylindrical plastic containers(150 mm diameter× 250 mm high) for ease of handling. Gauze cages wereused initially as plastic containers were not conducive to mating. Fresh tips insmall sealed vases were provided daily, when adult survival and the numberof eggs present were recorded.

Host specificity tests

The host range ofE. garciawas determined by testing 45 species representing14 families, including six species from the family Verbenaceae (Table 1). Thetest list was compiled by staff at AFRS using the centrifugal phylogeneticmethod devised by Wapshere (1975) in consultation with 21 Governmentagencies throughout Australia. This method places an emphasis on testingmore plants closely related to the target weed and testing fewer plants thatare less related. The family Verbenaceae is not well represented in Australiabut the closely related Lamiaceae has many representatives. Hence the appar-ent disproportionate number of species tested from these families. The plantswere propagated and grown in a heated glasshouse to produce foliage rang-ing from young tips to more mature leaves. Testing consisted of choiceoviposition trials and no-choice larval feeding trials.

Choice oviposition trialsForty unsexed adults were placed in organza-screened cages with up to eighttest plant species andL. camaraas a control. After 14 days, the plants wereremoved and examined for eggs or the presence of larvae. Plants with eggsor larvae were held for subsequent larval development. Each species wastested three times, using different adults on each occasion. The number of

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Table 1. Plants tested to determine host-specificity ofEctaga garcia

Family Genus/species Common name

Avicenniaceae Avicennia marina(Forsskaol) Vierh. var grey mangrove

australasica(Walpers) J. Everett

Bignoniaceae Jacaranda mimosifoliaD. Don jacaranda

Pandorea pandorana(Andrews) Steenis wonga vine

Boraginaceae Cordia dichromataG. Forster

Ehretia acuminataR. Brown silky ash

ConvolvulaceaeEvolvulus alsinoides(L.) L. tropical speedwell

Ipomoea batatus(L.) Lamarck sweet potato

I. plebeiaR. Brown bellvine

Polymeria calycinaR. Brown polymeria

Lamiaceae Ajuga australisR. Brown Australian bugle

Callicarpa pedunculataR. Brown velvet leaf

Chloanthes parvifioraWalpers

Clerodendrum cunninghamiiBentham

Clerodendrum tomentosum(Ventenat) R. Brown

Faradaya splendidaF. Mueller

Gmelina leichhardtiiF. Mueller white beech

Lavandula dentataL. French lavender

Mentha viridisL. mint

Plectranthus argentatusS.T. Blake

Premna lignum-vitae(Cunningham ex Shauer) lignum-vitae

Pieper

Prosthanthera ovalfoliaR. Brown mintbush

Salvia coccineaJussieu ex J. Murray red salvia

Scutellaria formosaN.E. Brown

Spartothamnela juncea(Cunningham ex Walpers)

Briquet

Teucrium argutumR. Brown native germander

Vitex trifolia L.

Westringia fruticosa(Willdenow) Druce coastal rosemary

Mimosaceae Acacia concurrensPedley black wattle

Archidendron lucyiF. Mueller

Myoporaceae Myoporum parvifoliumR. Brown dwarf native myrtle

Myrtaceae Eucalyptus tessellarisF. Mueller Moreton Bay ash

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Table 1. Continued

Family Genus/species Common name

Pedaliaceae Sesamum indicumL. sesame

Proteaceae Macadamia integrifoliaMaiden & Betche Queensland nut

Rutaceae Flindersia australisR. Brown crows ash

ScrophulariaceaeMazus pumilioR. Brown

Solanaceae Duboisia myoporoidesR. Brown corkwood

Lycopersicon esculentumMiller tomato

Solanum nemophilumF. Mueller

Solanum tuberosumL. potato

Verbenaceae Duranta repensL. duranta

Lantana camaraL. lantana

Lantana montevidensis(Sprengel) Briquet creeping lantana

Phyla nodiflora(L.) Greene carpet weed

Stachytarpheta urticifolia(Salisbury) Sims dark blue snakeweed

Verbena bonariensisL. purpletop

plant species and the combination of plants in each trial varied as test plantsbecame available. Replicating trials with the same plant combinations whileimportant statistically, was not viewed as being imperative in determiningspecificity of the agent. Mixing the combinations of plants in each trial, mayovercome the possibility that some plant species may mask the preference ofother species by the agent. In addition, the synchronisation of plant specieswas difficult as annual species needed to be tested before they senesce, whilethe timing of using perennial species was less critical. Larvae that completeddevelopment on any test species, were returned as adults to fresh plants of thesame species to determine whether the plant species could sustain successivegenerations ofE. garcia. Moths emerging fromL. camarawere returned tofresh plants as a control.

No-choice larval feeding trialsTen neonate larvae were placed on cut tips of each test plant species. Cuttips were placed in small jars of water with a cotton wool seal around theopening to prevent larvae from drowning. The jar was placed in a 600 mlclear plastic container and sealed with a gauze lid. Each species was testedthree times, withL. camaraused as the control. After seven days, the cut

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tips were examined and the number of larvae surviving and the degree ofdamage on each tip were recorded. Species with partial larval developmentwere held longer to determine if larvae could complete development. Mothsthat developed on test species were placed on plants of the same species asin theChoice oviposition trialsto determine whether successive generationscould be maintained.

Results

Biology

Adults are mottled brown-grey and about 10 mm long. A full description isgiven in Becker (1994). The adults feed on nectar and will drink water or adilute solution of honey. They usually oviposit on the under side of leavesadjacent to veins. Mated females live for up to 16 days (mean = 10.6, S.E.= 0.8, n = 35) and can lay up to 150 eggs (mean = 79.8, S.E = 8.0, n = 35),with peak egg laying occurring on days three and four. Males live for up to14 days (mean = 8.0, S.E. = 0.8, n = 35).

Eggs are white and shaped like a truncated cone measuring 0.35 mmdiameter at the base and 0.18 mm across the top, with a length of 0.35 mm.In the laboratory, larvae emerge after approximately eight days (mean = 8.1,S.E. = 0.1, n = 10) and feed on the lower surface of leaves ofL. camaraandL. montevidensis. As the larvae mature, they spin silk cocoons, causing theleaves to fold and form a protective covering. Larvae move out of the cocoonsto feed and may form new cocoons if they move to feed on new leaves. Thelarvae feed for 23 to 36 days (mean = 29.8, S.E. = 0.2, n = 500), have fiveinstars and pupate in the cocoons within the folds. Adults emerge after tendays (mean = 10.0, S.E. = 0.2, n = 30) and mate on the first night. Ovipositionusually begins on the second night.

Oviposition occurred and larvae developed onL. montevidensisand allvarieties of L. camara tested. In choice trials, there were greater initialnumbers of larvae on the red (mean = 109.2 larvae/plant, S.E. = 20.3), pink(mean = 91.8 larvae/plant, S.E. = 19.2) and orange (mean = 93.7, S.E. = 17.2)L. camaravarieties than the pink-edged-red or whiteL. camaravarietiesand L. montevidensis, however, this difference was not significant (F5,24 =1.992,p = 0.116) (Table 2). There was no significant difference in the meanpercent of larvae that completed development to pupae on each of the vari-eties (F5,24 = 0.235,p = 0.943). The lowest percent development occurred onthe pink variety (mean = 49.0%, S.E. = 13.4) while the highest was on thepink-edged-red variety (mean = 70.0%, S.E. = 10.0) (Table 2).

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Table 2. Mean percent larval development, adult survival and fecundity ofE. garciareared onL. montevidensisand each variety ofL. camara

Mean no. of Mean no. of Mean % larvae No. females Mean female Mean male Total eggs

initial larvae pupae formed that completed paired survival survival laid/female

present (S.E.) (S.E.) d’ment (S.E.) (days) (S.E.) (days) (S.E.) (S.E.)

L. montevidensis 66.3 (9.2) 36.7 (10.8) 55.3 (14.3) 11 7.8 (0.8) 5.7 (2.2) 53.3 (21.1)

L. camara

Orange 93.7 (17.2) 65.2 (14.9) 69.6 (9.2) 8 9.5 (1.3) 6.0 (1.2) 69.0 (16.2)

PER 42.5 (8.6) 29.8 (4.9) 70.0 (10.0) 8 12.8 (1.9) 9.8 (1.0) 112.3 (26.8)

Pink 91.8 (19.2) 45.0 (13.4) 49.0 (14.0) 13 11.1 (0.8) 9.8 (1.2) 73.5 (12.6)

Red 109.2 (20.3) 70.8 (18.5) 64.8 (10.1) 10 10.1 (1.2) 7.6 (0.9) 73.9 (10.3)

White 64.2 (16.6) 38.7 (9.2) 60.3 (11.1) 10 12.5 (1.8) 9.3 (1.4) 100.7 (31.8)

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First generation females reared on the white and pink-edged-red varietiesof L. camara tended to have greater survival (mean 12.5 and 12.8 days,respectively) and lay more eggs (mean 100.7 and 112.3 eggs, respectively)than those reared onL. montevidensisand the other varieties ofL. camara(Table 2). These differences though, were not significant (survival: F5,24 =1.243,p = 0.32; fecundity: F5,24 = 1.017,p = 0.429).

Host specificity testing

Choice oviposition trialsFemales did not lay eggs on any non-target test plant in any replicate,however, over 400 eggs were laid on each plant ofL. camaraor L. montev-idensisin each of the 18 trials set up. Between 250 and 300 adults emergedfrom each of the control plants.

No-choice larval feeding trailsLarvae completed development only onL. camara, where a total of 13 mothsdeveloped from a total of 30 neonates placed on each of three cut tip trials(Table 3). Larval feeding was recorded on nine test species. Where larvae fedon test plants, mainly exploratory or minimal feeding was observed and mostlarvae died within seven days. Five larvae however, survived for 14 days anda further three larvae survived for 27 days onVerbena bonariensisL. causinglittle damage to some leaves (Table 3).

Discussion

In host specificity trials, females did not oviposit and neonate larvae didnot complete development on any non-target test plant, while adults werereared from bothL. camaraandL. montevidensis, suggesting thatE. garciais specific toLantanaspp. Most larvae that fed on test plants died withinnine days and caused little damage to the plants. EvenV. bonariensis, onwhich some larvae survived for 27 days, is unlikely to be at risk fromE.garcia. Females did not oviposit on this or on any other test species, so it isimprobable thatE. garcia larvae would encounter these plants in the field.Consequently,E. garciawas approved for field release in 1993.

The use of choice trials to determine whether non-target species areacceptable for oviposition during host specificity testing were supported byCullen (1989) and Harley and Forno (1992). They advocate that choice trialsrelate more closely to the natural environment and many plants can be quicklyscreened. However, recent developments in host specificity testing suggest

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Table 3. Plant species on whichE. garciafed as neonate larvae during choice larval feeding trials

Plant species No. of Mean no. No. of Mean no. Mean no. Mean no. Damage caused by larvae

Replicates larvae trials with larvae after larvae after adults feeding

feeding 7 days (S.E.) 9 days (S.E.) emerged

(out of 3)

L. camara 3 10 3 4.7 (0.3) 4.7 (0.3) 4.3 (0.3) extensive damage to all leaves

C. pedunculata 3 10 1 0.3 (0.3) 0.3b (0.3) minimal feeding on a few leaves

I. batatus 3 10 2 0.0 0.0 exploratory feeding (<2 mm dia)

P. pandorana 3 10 2 0.0 0.0 exploratory feeding (<2 mm dia)

P. nodiflora 3 10 1 0.3a (0.3) 0.0 minimal feeding on a few leaves

S. indicum 3 10 3 2.3 (0.3) 0.7b (0.3) minimal feeding on a few leaves

S. urticfolia 3 10 2 0.0 0.0 exploratory feeding (<2 mm dia)

T, argutum 3 10 2 0.3 (0.3) 0.0 minimal feeding on a few leaves

V. bonariensis 3 10 3 2.7 (0.3) 1.7c (0.3) little feeding to a few leaves

W. fruticosa 3 10 1 0.0 0.0 exploratory feeding (<2 mm dia)

alarva died after 9 days.blarvae died after 10 days.clarvae died after 27 days.

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that such trials risk returning false negatives in potential host plant accep-tance. Marohasy (1998) suggested that adults placed in cages with both thetarget species and non-target species, will tend to only lay eggs on the targetspecies. By placing adults in cages without the natural host, the adults havethe option to lay on less preferred species, which may not be laid on if thetarget or preferred species is present as well. Perhaps a more appropriatemethod to determine host range or host acceptance in oviposition tests, wouldbe to use a combination of choice and no-choice trials.

There has been a similar debate on the testing of neonates and their abilityto develop on non-target species. Placing neonate larvae directly onto plantsby-passes the normal host finding behaviour of females. Consequently, larvaeencounter plants in laboratory tests which they would not normally encounterin the field, often resulting in a higher degree of initial feeding and returningfalse positives in host acceptance (Cullen, 1989). However, this method hasbeen used to determine host specificity in other moths. Larvae ofCarmentamimosaEichlin & Passoa andNeurostrota gunniellaBusck were unable tocomplete development after initial bouts of feeding on non-target species,when eggs or neonates were placed on plants (Forno et al., 1991; Davis et al.,1991).

E. garcia is one of a number of recent introductions to Australia in anattempt to controlLantanaspp. (Palmer et al., 1996). The introduction ofother agents is sought as the biological control of a perennial weed is seldomachieved by a single species. The key to a reduction in host plant densities liesin the combination of the actions of different agents (Hoffman and Moran,1998). Though twenty-seven species have now been released in Austialia and35 have been released worldwide, lantana is still a problem in many countries.In Australia, only 17 species have established, of which just four are causingsignificant damage. Similar trends in the low establishment rate of agents onlantana have been observed in South Africa and Fiji (Julien, 1992). Winderand Harley (1982) suggested that the low establishment rates maybe a resultof some agents being incompatible with some of the lantana varieties presentand the failing of other agents to adapt climatically.

Some of the agents that have established in Australia have shown a prefer-ence for certain varieties of lantana, which may influence their establishment,abundance and distribution in the field. Haseler (1966) reported thatNeogaleasunia(Druce) prefers the pink and white forms of lantana, whileSalbia haem-orrhoidalis Guenée prefers varieties of red lantana, and Taylor (1989) foundthat Teleonemia scrupulosaDistant performed poorly on pink lantana vari-eties. Preference and performance tests in the laboratory indicateE. garciareadily acceptsL. montevidensisand all of the varieties ofL. camaratested.

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Another factor proposed as influencing the establishment of biocontrolagents on lantana is climate. There are numerous examples of biocontrolagents that have been easily reared in the laboratory, but have failed toestablish when released in the field. Other species have only established inlocalised areas (Neser and Cilliers, 1989; Taylor, 1989). Neser and Cilliers(1989) proposed that climatic matches between the country of origin andareas of infestation should be conducted prior to field release to aid estab-lishment. AsE. garcia is a leaf feeder, field releases should be restricted toareas of high rainfall, where lantana would have healthy foliage all year. Indrier areas, lantana can periodically lose its leaves, thus preventing defoli-ating agents from maintaining sufficiently high populations to exert control.Climatic matches with its geographic range in its country of origin (southernBrazil) suggest thatE. garcia would be suitable for release in most coastalparts of Queensland and New South Wales.

The potential ofE. garciaas an agent for lantana is yet to be determined,however, the field release ofE. garciashould complement the action of flowerand seed feeding agents already established as they utilise different parts ofthe plant. It is acknowledged that the establishment or abundance ofE. garciain some areas, maybe influenced by other leaf feeding agents onL. camarasuch as the widespread hispine beetles,Uroplata girardi Pic andOctotomascabripennisGuérin-Méneville. These beetles can cause extensive damage tothe leaves of lantana through the presence of mines and so possibly limit thenumber of oviposition and larval feeding sites forE. garcia.

Iwao and Rausher (1997) however, suggested that the presence of oneherbivore does not necessarily affect the level of damage by another herbi-vore. There will always be refuge areas or new growth not attacked by thebeetles, which could support oviposition or larval development of the moth.The introduction of another agent in areas where lantana is not adequatelycontrolled may lead to greater pressure on the weed resulting in stunted plantsand reduced flowering (Winder and Harley, 1982; Harris, 1984). In the nativerange of lantana there are numerous insect species of the same guild attackinglantana, and it is this combined effect that assists in suppressing plant density.Also, L. camarain Hawaii has been partially controlled by three species ofleaf-feeding Lepidoptera and the tingid,T. scrupulosa(Harris, 1973).

E. garciawas re-imported in 1997 and is currently being mass reared andreleased in Queensland and northern NSW. Low numbers of the agent werereleased at a few sites in 1993 prior to the laboratory colony dying out andthe project being suspended, but it did not establish.E. garcia will also bereleased onL. montevidensis, on which only one other agent is currently beingtested (Day et al., unpublished data). Regular monitoring is being conductedto evaluate its establishment.

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Acknowledgements

We wish to thank Cesar Garcia for field collecting and conducting prelimi-nary studies on the agent in Brazil, Justin Robazza for supplying the plantsand colleagues at AFRS for their helpful comments on the manuscript. Thisproject was funded by the Rural Lands Protection Board of Queensland.

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