S
TORED
-P
RODUCT AND
Q
UARANTINE
E
NTOMOLOGY
Laboratory Rearing of Lesser Appleworm (Lepidoptera: Tortricidae)
K. D. MANTEY, H. R. MOFFITT,
AND
L. G. NEVEN
Yakima Agricultural Research Laboratory, 5230 Konnowac Pass Road, Wapato, WA 98951
J. Econ. Entomol. 93(3): 1021Ð1024 (2000)
ABSTRACT
The lesser appleworm,
Cydia prunivora
(Walsh), was reared successfully in thelaboratory. Larvae of various instars were collected in the Þeld from hawthorn fruit,
Crataegus
spp.Initially, immature apples served as the food source for the larvae in the laboratory. Rearing wasconducted in a greenhouse and later in combination with a controlled environment room at 25
C,50Ð60% RH, and a photoperiod of 18:6 (L:D) h. Under these conditions, a generation required
30d. Fifty-six adult lesser appleworm moths emerged from the original Þeld collected hawthorn fruits.After a decline in the number of the F
1
generation to 39 moths, the colony on mature apples,increased to in excess of 10,000 moths by the Þfth generation with a mean survival rate to adult of 68.0%.WhenproductiononimmatureappleswascomparedwiththatonfourartiÞcialdiets,themostpromising of the artiÞcial diets was the lima bean-based diet currently used to rear the oriental fruitmoth,
Cydia molesta
(Busck), with a mean survival rate of 46.4%. The other bean-based diets testedwerenotassatisfactory.Pearfoliagewasthepreferredovipositionsubstrateofthosetested,includingappleandhawthornfoliage.Noeggsweredepositedonplainwaxedpaperorglassmicroscopeslides;however,largenumbersofeggsweredepositedonwaxedpapertreatedwithawaterextractofpearfoliage and immature apples.
KEY WORDS
Cydia prunivora,
lesser appleworm, rearing, hawthorn,
Crataegus,
diets
L
ESSER APPLEWORM
,
Cydia prunivora
(Walsh), is widelydistributed in North America and is probably a speciesnativetoeasternNorthAmerica.Thelesserapplewormwas described by Walsh (1868) as infesting plums asearly as 1867 in Illinois and was Þrst reported to attackapples in British Columbia, Canada, in 1895 (Fletcher1898).Lesserapplewormhasnotbeenaprobleminmostcommercial orchards. Because the lesser applewormÕslifehistoryandhabitsareverysimilartothoseofcodlingmoth, the chemical control measures used to controlcodling moth also control lesser appleworm (Faurot1912, Quaintance and Siegler 1922). Turmel and Fisher(1978) stressed the importance of not altering controlprogramsforinsectsonapplesinNewHampshirewith-out considering the possible impact on lesser apple-worm. With the increasing use of species speciÞc bio-rationaltechniquesforcontrolofcodlingmoth(Howell1992) and the subsequent decrease in the use of pesti-cides, consideration of the impact of these techniquesupon lesser appleworm becomes even more important.Thelarvaeofthisspecieshavebeenreportedtoattackthe fruit of the family Rosaceae (Chapman and Lienk1971, Brunner and Howitt 1981). The principal generaattacked are
Crataegus
(hawthorn) (Quaintance 1908,Wellhouse 1922),
Prunus
(prune, plum, cherry, peach,andapricot)(Brown1953),
Pyrus
(apple,pear,andcrabapple) (Fletcher 1898, Wellhouse 1920),
Rosa
(rosehips) (Mackie 1942),
Photinia
(toyon) (Keifer 1933),and
Amelanchier
(shadbush or serviceberry) (Heinrich1926).Lesserapplewormlarvaehavealsobeenreportedon galls of the oak (Fagaceae) and elm (Ulmaceae)(Fletcher 1898). The larvae are internal feeders similarto codling moth,
Cydia pomonella
(L.), oriental fruitmoth,
Cydia molesta
(Busck), and cherry fruitworm,
Cydiapackardi
(Zeller).Thelarvaeofallfourspeciesaresimilar in appearance. However, the codling moth larvalacksananalcomb,astructurethatispresentintheotherthree species (Garman 1918). Codling moth larvae arelargerthantheothersandfeeddirectlytothecoreofthefruit. Oriental fruit moth larvae are larger than lesserappleworm and cherry fruitworm larvae. Oriental fruitmothlarvaedonotfeedontheseeds.Lesserapplewormand cherry fruitworm larvae are very close to the samesize and the feeding habits are similar (Mackay 1959)(ChapmanandLienk1971).Larvaeofbothspeciesfeedbelowthesurfaceoftheskinofthefruitcausingsurfaceblotches.Lesser appleworm had remained relatively obscureuntil the Japanese government made it a quarantineissue for U.S.-produced apples in 1991 (H.R.M., un-published data). Lesser appleworm is not reported tooccur in Japan. Because lesser appleworm had beenreported as a pest of stone and pome fruits in thePaciÞc Northwest in the Milton-Freewater area of Oregon (Brown and Jones 1953), the export of applestoJapanwasdelayed.ThisstudywasinitiatedtolocateintheÞeldandrearinthelaboratoryenoughnumbersof lesser appleworm to determine if the two compo-nent treatment, fumigation with methyl bromide fol-lowedbycoldstorage,(MofÞtt1988)devisedtomeetquarantine security requirements for codling mothwouldalsomeetrequirementsforlesserappleworminapples destined for export to Japan.
Materials and MethodsRearing on Thinning Apples.
Larvae of various in-stars were collected initially in the Þeld from fruit of the hawthorn,
Crataegus
spp. Hawthorn fruits werebrought to the laboratory and placed in a plastic con-tainer(20.0cmlongby14.0cmwideby10.0cmdeep)with a cheesecloth cover. The container was held in agreenhouseat25
C,50Ð60%RH,andaphotoperiodof 18:6 (L:D) h for emergence. A greenhouse was se-lected to provide sunlight for the developing larvaeandthesubsequentmatingoftheemergingadults.An18-h day was selected to prevent the larvae from en-tering diapause (Brown 1991). The emerging adultswere collected and the sex of each adult was deter-mined. The adults were then placed in a plastic con-tainer, identical to the original container, with haw-thorn and apple fruits placed in the bottom forpossible oviposition stimulation and ovipositionalsites. The adults from this generation and those of subsequent generations were placed in mating andoviposition cages. The mating and oviposition cageswere 30.0-cm cubes. The cage was constructed of awooden frame with a wooden bottom. The frame wascovered on three sides and the top with nylon or-gandy. The fourth side of the cage had an armholeopening for access to the inside of the cage. Tenthinning apples were placed on the bottom of eachcageasapossibleovipositionstimulant,anovipositionsite and/or food source for any larvae that mighthatch. A 250-ml Erlenmeyer ßask of water with acottonwick(15.0cmdentalroll)servedasasourceof moisturefortheadultmoths.Adultfemalemothsfromthe Þrst few generations were dissected for the pres-ence of a spermatophore in the bursa copulatrix todetermine if mating had occurred.
Oviposition Substrates.
The search for an accept-able oviposition substrate included foliage of apple,pear and hawthorn, waxed paper, and glass micro-scope slides. Each of these possible oviposition sub-strates was presented to the female moths in the mat-ing and oviposition cages.Eachofthethreetypesoffoliagewascollectedfromthe Þeld and presented with the leaves attached. A250-ml Erlenmeyer ßask was Þlled with water, theshoots were wrapped with a 2.5 by 15.0-cm piece of rolled cotton to prevent the adults from drowning inthe water, and placed in the ßask. A 7.0-cm piece of plastic drinking straw was inserted through the rolledcotton into the ßask to allow water to be added to theßaskdaily.Threeoftheßaskswithshootsandoneßaskwiththemoisturewickwereplacedineachcage.Also,to supply pear foliage during the winter, pear shootswere collected from the Þeld in the fall. The methodof pear shoot collection and preservation was de-scribed by Fye (1981).The waxed paper was presented plain and with anextract that was developed to enhance oviposition onthepaper.Theextractconsistedof250mlwater,eightthinning apples, and 16 pear leaves. The ingredientswere blended for 1 min, forming a slurry. The slurrywas Þltered through qualitative Þlter paper, VWRgrade 613. A cotton ball was dipped into the Þlteredsolution and the liquid was spread across a 40.0 by30.0-cm piece of waxed paper. The waxed paper wasallowed to dry at room temperature. It was then cutintofour10.0by7.5-cmpieces.Thewaxedpaper,boththeplainandtheextractenhanced,werecrinkledandsuspendedalongthesidesofthecage.Theglassslideswere suspended from the top of the cage using tapeand wire. Glass slides were also laid in the bottom of the cage.A maximum of 500 adults was added to each cage.The conditions in the greenhouse were 25
C, 50Ð60%RH,andaphotoperiodof18:6(L:D)h.After5d,eachovipositionsubstratewiththeeggswasremovedfromthe cage and placed in a clear plastic container (35.0cm long by 27.0 cm wide by 9.0 cm deep). A sectionwas removed from the plastic lid of the container andorgandywasgluedovertheholeforventilation.Thin-ning apples, measuring
3.0 cm in diameter, wereplacedinthecontainertoserveasthefoodsourceforthe maturing larvae. The containers were transferredto a holding room at 25
C, 50% RH, and photoperiodof16:8(L:D)h.CorrugatedÞberboardstrips,(33.0cmlong by 1.3 cm wide) ßute size 125BU (Tharco, Au-burn, WA), were placed on the bottom and along thesides of the container. The ßutes of the strips servedas cocooning sites for mature larvae. The Þberboardstripscontainingthelarvaeweretransferredtoacleanplastic container of the same dimensions, where theadults emerged and were collected. The adults werecollected by using a battery operated aspirator.
RearingonArtificialDiets.
FourartiÞcialdietsusedfor the rearing of other insects were evaluated forpossible use as a food source for lesser applewormlarvae. The diets tested were as follows: the codlingmothdiet(CMdiet)(Howell1970),theorientalfruitmoth diet (OFM diet) (Yokoyama et al. 1987), theMcNeildietwithpintobeans(McNeilw/pinto)(Sho-rey and Hale 1965), and the McNeil diet with limabeans (McNeil w/lima). The ingredients for the dietsarelistedinTable1.Thinningappleswereusedasthestandard for comparison. The diets were mixed andpoured into individual 30.0-ml plastic cups. Fifteenmillilitersofthedietoronethinningapplewasplacedintoeachcup.Therewere25cupsforeachtypeofdietand each diet was replicated Þve times.A single lesser appleworm egg was placed on eachindividualcupofdietoreachthinningapple.Theeggswere collected by placing pear foliage in an oviposi-tion cage with 500 adult moths. The moths were al-lowed to mate and oviposit for a 48-h period. Foliagewith eggs was removed from the cage and single eggswere isolated by cutting a piece of leaf with a singleeggonitandpiercingtheleafwitha2.0-cmlongpieceofwire.Theotherendofthewirewasinsertedintothediet or thinning apple, suspending the egg over thediet or apple to diminish the possibility of contami-nation through direct contact. After 9 d, the foliagewith the egg was removed from the diet and the eggwas evaluated for hatch. The larvae were allowed todevelopandthenumberandsexoftheemergingadultmothswererecorded.Egghatch,survival,andrearing1022 J
OURNAL OF
E
CONOMIC
E
NTOMOLOGY
Vol. 93, no. 3
weight data were subjected to analysis of variance(ANOVA) and the Tukey mean separation (SAS In-stitute 1989). Percentages were transformed by
sin
1
before analysis.
Results and Discussion
Fifty-six adult lesser appleworm moths emergedfrom the original Þeld-collected hawthorn fruits. Theratioofmaletofemalemothswas1.5:1.Thefrequencyofmatingamongthenativefemalesthatemergedwas42.9%. The F
1
generation declined in number andproduced only 39 moths. The male to female ratio of this generation was 0.8:1 with 94.1% of the femalesmating. The total numbers of moths for the F
2
and F
3
generations were 228 and 697, respectively. Each suc-cessive generation continued to increase in numbersuntil space in the rearing facility became the limitingfactor (Fig. 1). The typical size of each generationmaintained is currently at 10Ð15,000 moths.A gold color variant was observed starting with theF
5
generation (Fig. 1). The color variant has the samemarkings as the normal lesser appleworm adult, butthe colors are muted resulting in a moth that has amorepaleappearance.Atotaloffourgoldadultmoths(two males and two females) was collected. Thesefour moths were held in a separate cage to form asubcolony of homozygous gold moths. This gold col-onywasnotasproliÞcasthewildcolony.Agoldcolorvarianthasalsobeenreportedforcodlingmoth(Huttand White 1975). The results of their laboratory andÞeldstudiesindicatedsimilarresults,thatgoldcodlingmoth population was not as vigorous as the wild pop-ulation.Pear foliage was observed to be the preferred ovi-position substrate. The female moths laid eggs singlyon the upper and lower surfaces of the leaves as wellasthestems,withmostoftheeggsbeingfoundontheunderside of the leaf. Female moths did not lay anyeggs on the plain, waxed paper or the glass slides.However, waxed paper treated with the extract resi-due was found to be an acceptable substrate for ovi-position. When the waxed paper with the extract res-idue and the pear foliage was monitored for a year inthe mass rearing facility, the treated waxed paperaveraged 448 eggs with a range of 28Ð4039 eggs perovipositioncage.Theaveragehatchwas75%.Thepearfoliage averaged 2,152 eggs per oviposition cage witha range of 262Ð3,248. The average hatch was 86.6%.Sunlight played a signiÞcant role in selection of ovi-position sites. Pear foliage and waxed paper stripstreatedwithextractthatwereorientedtowardthesunhad signiÞcantly more eggs than those not oriented
Table2. Comparisonoffourartificialdietsandthinningapplesas a food source for lesser appleworm larvae
Food source Mean % hatch Mean % survivalThinning apples 96.0a 68.0aOFM diet 94.4a 46.4bCM diet 96.8a 27.9bMcNeil w/pinto 94.4a 38.5bMcNeil w/lima 94.4a 27.7bFormeanpercenthatch:
F
0.20,df
4,
P
NS.Formeanpercentsurvival
F
6.77; df
4, 20;
P
0.01. Means followed by the sameletterwithinacolumnarenotsigniÞcantlydifferent(ANOVA,Tukeymean separation test).
Table3. Weightsoflesserapplewormadultsrearedonthinningapples compared to weights of adults reared on OFM diet
Food source Sex Mean wt Range STDThinning apple
&
4.6a 2.5Ð6.3 0.7Thinning apple
(
4.3b 3.0Ð5.7 0.6OFM diet
&
3.6c 1.4Ð5.6 0.9OFM diet
(
3.8c 1.6Ð5.5 0.9For mean weight:
F
34.95; df
3, 396;
P
0.01. Means followedbythesamesmallletterwithinacolumnarenotsigniÞcantlydifferent(ANOVA, Tukey mean separation test).
Table 1. List of ingredients and quantities for each artificialdiet tested as a possible food source for lesser appleworm larvae
IngredientOFMdietMcNeilw/PintoCMdietMcNeilw/LimaAgar 10.0 g 5.0 g 18.0 g 5.0 gL-ascorbic acid 6.0 g 0.8 g 12.6 g 0.8 gBrewerÕs yeast 60.0 g 8.0 g 8.0 gFructose 9.0 gMPHB 0.6 g 0.5 g 3.0 g 0.5 gSoy protein 15.0 gSorbic acid 1.5 g 0.25 g 2.7 g 0.25 gVitamin mixture 11.5 g 1.0 g 33.0 g 1.0 gWheat germ 30.0 g 180.0 gWater 900.0 ml 290.0 ml 3600.0 ml 290.0 mlLima beans 300.0 g 53.25 gFormaldehyde 0.75 ml 8.1 ml 0.75 mlPinto beans 53.25 gSoybean meal 495.0 gWheat starch 81.0 gMineral salts 5.4 gBenlate 0.54 gPropionic acid 8.1 mlPropylene glycol 30.0 mlAuremycin 8.0 gSucrose 87.0 g
Fig. 1.
Propagation of laboratory-reared wild and goldpopulations of lesser appleworm.
June 2000 M
ANTEY ET AL
.: L
ABORATORY
R
EARING OF
L
ESSER
A
PPLEWORM
1023
of 00
USDA: Neven 2000 42%20Mantey%20LAW%20rearing
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