IMMATURE STAGES OF METALMARK MOTHS FROM THE GENUS BRENTHIA CLEMENS (CHOREUTIDAE): MORPHOLOGY AND LIFE HISTORY NOTES

2008·5.4 MB·English

by  Jadranka Rota

Journal of
The
Lepidopterists’
. I
Volume 62 2008 Number 3
JournaloftheLepidopterists’Society
62(3),2008,121-129
IMMATURE STAGES OF METALMARK MOTHS FROM THE GENUS BRENTHIA CLEMENS
(CHOREUTIDAE): MORPHOLOGY AND LIFE HISTORY NOTES
JADRANKA ROTA
DepartmentofEcologyandEvolutionaryBiology,UniversityofConnecticut,Storrs,CT06269, USA;Currentaddress: Departmentof
Entomology,NationalMuseumofNatural History,SmithsonianInstitution,Washington, DC20013-7012;email: [email protected]
ABSTRACT. InthispapertheimmaturestagesofBrenthia monolt/chna Meyrick(Choreutidae: Brenthiinae),aswellastheirul-
trastructure,aredescribedandfigured. ThisisthefirstdescriptionofaNewWorldbrenthiine. In addition, noteson lifehistoryfor
four NewWorldspecies ofBrenthia Clemens areprovided, includingmention of theirhostplants andparasitoids. Host plant uti-
lization ofthegenus is discussed. Aclarification ofthe nomenclatureofthelongestsetaon the larval abdominalsegment9is pro-
posed. Earlierliteraturedisagreesonwhetherthisisalateral,subdorsal,ordorsalseta-myexaminationsuggestsitisthesubdorsal
seta 1.TherecordeddistributionofBrenthiapavonaceUa Clemensisquestioned,andareviseddistributionissuggested. Moreover,
an escape mechanism, employedbyallknownBrenthia larvae, isdiscussed. Finally, alistofmorphologicalandbehavioralsynapo-
moiphiesforthesubfamilyBrenthiinaeandthegenusBrenthiaisprovided.
Additional keywords: Microlepidoptera,chaetotaxy, ultrastructure,larvalescapebehavior,parasitoids, Braconidae
Brenthia Clemens is a cosmopolitan genus of mechanism (see below; Williams 1951; Diakonoff 1986;
metalmark moths (Choreutidae). With more than 80 Aiello and Solis 2003).
described species and likely an even greater number of Immature stages ofBrenthia have been described for
undescribed species (Rota2003), it is amongthe largest ahandfulofspecies (e.g.,Williams 1951; Arita 1987). In
microlepidoteran genera. Together with the genus this paper, the larval and pupal stages of Brenthia
Litobrenthia Diakonoff, Brenthia is classified in the monohjchna Meyrick are described and figured. This is
subfamily Brenthiinae (Heppnerand Duckworth 1981). the first detaileddescription ofthe immature stages ofa
The relationship of brenthiines to the other two New World member of the subfamily. In addition, life
choreutid subfamilies, Choreutinae and Millieriinae, is historynotes are provided on B. monohjchna, as well as
understudy(J. Rotain prep.). three of its congeners: B. hexaselena Meyrick, B.
Moths in the genus Brenthia are small - wing size pavonaceUa Clemens, and B. stimulans Meyrick.
rangesfrom 6to 14mm.With blueorvioletmetallicand
Materials and Methods
white markings on dark backgrounds, wings of most
species look veiy similar (Figs. 1-3), so much so that Larvae of B. monohjchna B. hexaselena and B.
, ,
genitalic dissections are often necessary to confirm stimulanswere collected from 2001 to 2004in atropical
identification. AdultsofBrentliia mimicjumpingspiders wet forest at La Selva Biological Station, a lowland
by holding their wings to the side and above the body, reserve on the Atlantic slope ofCosta Rica, Province of
and movingin rapid,jerkymotion (Robinsonetah 1994; Heredia. B. monohjchna pupae were reared from a
RotaandWagner2006). collection oflarvaeinAugust2004alsofrom LaSelva. B.
Life history information is available for only a few pavonaceUa larvaewerecollectedonSeptember8, 2002,
species, mostly from Asia (e.g., Arita 1987). Brentliia in Illinois, Coles County, Fox Ridge State Park, byTerry
larvae appear to be relatively specialized, one species L. Harrison. Rearingwas done in alaboratoryin plastic
usually feeding on a single genus or closely related bags or plastic vials filled with foliage and some soft
genera ofplants (Arita 1987; Rota 2003). The larvae of papertissue forcontrolof humidity. Periods of light and
known species are surface-feeding leaf skeletonizers darkcorrespondedto naturallightcycles.
(Arita 1987; Aiello and Solis 2003). All known larvae For preservation, larvae and pupae were placed into
seem to have a similar predator/parasitoid avoidance nearly boiling water for less than a minute and then
122 Journalofthe Lepidopterists’ Society
transferred to 75% ethanol (Zimmerman 1978). Dl and D2 in vertical alignment, and with extremely long SD1,
Specimensforviewinginthe SEM weredehydratedin a correspondinginlengthto4-5abdominalsegments. Prolegslongand
slender,subcylindrical(Fig.21);crochetsuniserialanduniordinal;A3-6
graded ethanol series (ending in 100%), transferred to withapproximately12crochetsarrangedinmesalpennelipse(Fig.22);
freshsolutionsofhexamethyldisilazanethreetimes,each A10with 16crochetsinsemicirclewithposteriorgap.
time for approximately 15 minutes, and then immersed Pupa. Averagelength6.25mm (n=3);with long,fork-tippedsetae
(Figs. 29, 32), and short, thickly arranged, caudally oriented dorsal
into fresh hexamethyldisilazane and allowed to air-diy. spinesonA4-8;A4withdorsalspinespoorlydeveloped(Fig.23),A5
Dried specimens were sputter coated with spines larger than on A4, but smaller than on A6 or A7 (Fig. 24),
gold/palladium. A6-A7 spines well developed (Figs. 25-27), A8 with field ofpoorly
A LEO/Zeiss DSM 982 Gemini Field Emission SEM dcuervveeldopseldesnpdienressp(iFniegs.,2t8)w.oCpraeimrsasotfehrooonk-At1i0ppweidthsettwaoeptahirreseoftislmiegshtalsy
was used. Photographs ofadults and larvae were taken longasspines,andtwopairsofverylongfork-tippedsetaesixtoseven
with a Nikon D100 and Dl. The pupal photographwas timesaslongashooks(Figs.29,30).
taken with a digital camera attached to a Leica Life history' notes. As in otherchoreutids, larvae of
microscope, connected to a computer with Brenthia leave diagnostic feeding damage: superficial
Automontage® software (Synoptics Ltd., Cambridge, skeletonizationofundersidesofleaves (Figs. 4-7).While
UK). Line art was prepared with a camera lucida. All earlier instars often feed in groups (Fig. 5), older larvae
imageswere editedin Adobe Photoshop CS©. are normally solitary (Figs. 6, 7). Over the feeding area
Measurements were made using an ocular larvae create a thin, loose web into which they
micrometer. Chaetotaxy nomenclature follows Hinton incorporate fecal pellets (Figs. 7, 8). Most likely, the
(1946) and Stehr (1987). Usage of other terms is as incorporation of fecal pellets is not an active process;
defined in the Torre-Bueno Glossary of Entomology rather, it happens passively as the caterpillars move
(Nichols 1989). Larval description is based on the last about the shelter and their feculae accumulate in the
instar. Voucher specimens are deposited in the webbing. Larvae ofall four Brenthia species that I have
University of Connecticut Entomological Collection observed chew a roughly circular “escape hatch” - a
(UCMS). wormhole - somewhere in their feeding shelter. When
Results resting, they sit with their head next to the hole. If
Hostplants. Larvaewerecollectedandrearedondie disturbed, larvae dash through the wormhole to the
following host plants: B. hexaselena larvae from other side ofthe leaf(Figs. 7, 9, 11). After a littlewhile,
Byttneria aculeata (Jaeq.) Jacq. (Sterculiaceae) (Fig. 4); they wriggle through the opening backwards to their
B, monolychna larvae from Calathea crotalifera S. original position. Larvae of some species (e.g., B.
Watson (Marantaceae), other Calathea spp., and from monolychna) construct “fecal stalactites” (Aiello and
Heliconia sp. (Heliconiaceae) (Figs. 5-10); B. stimulans Solis 2003) in theirfeedingarea (Fig. 9). Located at the
larvae from Cecropia insignis Liebm. (Cecropiaceae); mouth ofthe escape hole on the leafunderside, these
and B. pavonacella larvae from Desmodium glutinosum structuresappeartoserveaslandmarksthatfacilitatethe
(Muhl.) Wood (Fabaceae) and other Desmodium spp. quick escape ofa larva (see Aiello and Solis 2003). The
(Figs. 11-12). fecal stalactites also seem to serve to suspend the
Morphology ofthe immature stages ofBrenthia monolychna webbing above the lana. Brenthia cocoons are often
Meyrick, 1915. Lana.Palegreeninlife (earlyinstarsalmostwhite) spun on the leaf undersides or somewhere on the stem
a(Fbidgso.me5n-,7)a,ndtrwaintshlucleonntg,sewtiateh;oauvteraangye lpeantgttehrn8i.n5gmomn (tnh=o5r)a.xOannldy and are composed oftwo principal parts: an inner one,
primarysetaepresent.Head.Withdarkbrownspots(Fig.6),widestat which is white, fusiform, and composed of multiple
aboutlevelofPI;vertexonlyshallowlyconcave;hypognathous;setaeas layersofthicksilk: andathin, outermostsilkenlayerthat
ilnenFgitghs.of1h3e-a1d5;;AA33t,hrAeFe1t,imaensdasPlIonegxtasreAmle,lyanedloAn1gaatned,Aa2ppsruobaecqhuianl;g forms a sheet over the cocoon proper (Fig. 10). The
AF1 five to six times as long as AF2; S2 posteriad of stemma 1; whole cocoon of B. monolychna is less than 2 cm along
spinneretwell developed, more than twice as long as labial palpus, its longest axis. The pupais protruded from the cocoon
salrernadnegred(Fiingsa.se1m5i,ci1r6c)l;e,swtietmhmsatVeImmhaig2hlpyosrietdiuocneedd;sloitghhtelryosuttesmimdeatoaf at eclosion. For all four species, the development from
semicircle (Fig. 14); frontoclypeus extending halfway to epicranial the pupato the adultstagetookabouttendays.
notch; labrum notched, with toothed underside (Figs. 17, 18); Larvae ofthe three Brenthia species reared from La
mandiblewithwell-developedteeth;hypopharyngealspinesprominent
(Figs.19,20).Thorax. DlandD2subequal;T1withLIthreetimesas Selva are heavily parasitized by braconid wasps
longasL2andL3,SVbisetose;T2withLItwotimesaslongasL2and (Hymenoptera: Braconidae) (Table 1 and Fig. 10). In B.
threetimesaslongasL3;T2andT3withSVunisetose,SD1andSD2 monolychna the parasitism rate is close to 85%
subequal, L setae on separate pinacula. Abdomen. Setae very long,
especiallyposteriad(Fig.31);DlandD2subequal,LIlongerthanL2 (n, =51). In B. hexaselena (n, =5) the rate is about
and L3; Al-2 and A7-9 with SV group numbering 3:3 and 2:2:1, 20%, and in B. stimulans (% =4) about 50%, but note
respectively;AlwithLsetaeonseparatepinacula;A2widrSVsetaein small sample sizes. I have inrvoaet reared any parasitoids
triangle;A8withsetaeDlandD2oncommonmiddorsalpinaculum;
A9with Dl, SD1, LI,andL2oncommonsubdorsalpinaculum,with from B. pavonacella (n|iwae=25).
Volume 62, Number 3 123
Figs. 1-12. 1. B. hexaselena adult. 2. B. monolychna adult. 3. B. pavonacella adult. 4. B. hexaselena larvaon its hostplant. B
monolychna:5.Gregariousfeedingofyounglarvae;6. Lastinstarlarva; 7. Larvagoingthroughanescapehatch;8. Larvalwebbing;
9. Fecalstalactite; 10. Cocoons ofahealthylarva(center) andofaparasitizedlarva(upperright). B. pavonacella: 11. Larvagoing
through anescapehatch; 12. Lastinstarlarva.
124 Journalofthe Lepidopterists’ Society
Figs. 13-18. B. monolychnalarva: 13. Head, dorsal; 14. Head,lateral; 15. Head,frontal; 16. Head,ventral; 17. Labrum,dorsal;
18. Labrum,ventral.
Volume 62, Number 3 125
Figs. 19—24. B. monolychna larva: 19 and 20. Hypopharyngealspines; 21. Ventralviewofabdominalprolegs (arrowpoints to
anterior); 22. Crochets on an abdominal proleg (arrows pointing A: anterior and M: mesad). Pupa: 2.3. Dorsal spines on A4;
24. DorsalspinesonA5.
126 Journalofthe Lepidopterists’ Society
Figs. 25-30. B. monolychna pupa: 25. Dorsal spines on A6; 26. Close-upofdorsal spines on A6; 27. Dorsalspines on A7; 28.
PoorlydevelopedfieldofdorsalspinesonA8; 29. Dorsalviewofthecremaster;30.Close-upofthecremaster.
Volume 62, Number 3 127
Figs. 31-33. B. monolychna: 31. Larvalchaetotaxy; 32. Pupa,dorsal,lateral, andventralview. B pavonacella: 33.pupa.
—
. .
128 Journalofthe Lepidopterists’ Society
B. monolychna isanunusuallyabundant—species atLa Table 1. GeneraofbraconidparasitoidsfromB. monolychna
Selva Biological Station, at least as a larva there were andB. hexaselena
traces of larval feeding damage on almost every host
plant examined (n>300). The adult can he seen during B. hexaselena Microgastrinae
the day on vegetation in the vicinity of its host plant; DolichogenideaViereck
occasionallyadults also come to mercuryvaporlight. B. monolychna Microgastrinae
B. hexaselena is uncommon in collections from La DolichogenideaViereck
Selva, but it can be frequently encountered during the Agathidinae
day around stands of its hostplant. I have never seen it PlesiocoelusvanAchterberg
come to lights at night (number ofblacklighting nights Orgilinae
Orgilus Haliday
at La Selvaca. 100).
B. stimulans appears to feed on mature Cecropia into the floor oftheir feeding shelter (Aiello and Solis
insignis plants only- I examineddozens ofsaplings and 2003;Williams 1951; Diakonoff1986; thispaper).While
neverfoundanysignsoflarvaeortheirfeedingdamage. clearly this represents a predator/parasitoid escape
This species, most likely because its host plant is a tall mechanism, it is also evident that this behavior is not
tree, is rarelyencountered. especiallysuccessful foravoidingparasitism bybraconid
—
Brenthia pavonacella is the only member of the wasps as noted above, parasitism rates were as high as
genus foundin North Americanorth of Mexico (Hodges 85% in B. monolychna
et al. 1983). It is widely distributed in the eastern Most species of Brenthia appear to be specialists
United States, north to southern New York (Forbes they feed on a single genus of plants or closely related
1923). B. pavonacella is often common where found. groups ofplants. Yet, when taking a look at the whole
Like B. hexaselena and monolychna, it is principally genus, Brenthia species have been recorded from a
diurnal. ratherextraordinaryarrayofunrelatedplantfamiliesthat
Discussion includes both monocots and dicots: Boraginaceae
In larvae of Brenthia, the longest seta on A9 is (Williams 1951), Cecropiaceae (LaPierre pers. comm.;
considered under different names by different authors: this paper), Asteraceae, Dipterocarpaceae,
Arita (1987) refers to it as an additional seta ofthe L- Euphorbiaceae (Robinson et al. 2007), Fabaceae (Arita
group, whereas Heppner and Duckworth (1981) call it 1987; Aiello and Solis 2003), Heliconiaceae (this paper),
D2. Basedonacarefulexaminationof setalarrangement Malvaceae (Heppner 1985), Marantaceae (Aiello and
on all the abdominal segments and comparisons with Solis 2003), Moraceae (Robinson et al. 2007),
Hinton (1946) and Stehr (1987), I agree with Williams Sapindaceae (Heppner 1985), Sterculiaceae
(1951), who designatedthis setaas SDL (Ilespenheide pers. comm.; this paper), Tiliaceae
Forbes (1923) characterized the distribution of B. (Robinsonetal. 2007), and Urticaceae (Diakonoff1986;
pavonacella as extending from New York southward to Arita 1987). There are also unconfirmed records of
Brazil. Thisisalmostcertainlyincorrect. Throughoutthe rearings from ferns (specimens in Costa Rica’s INBio
NeotropicstherearemanyspeciesofBrenthiawithwing collection). A species-level phylogeny of the genus, at
patterns very similar to that of pavonacella. My this point unattainable, would create an opportunityfor
examination ofgenitalic characters and molecular data the study of the evolution of Brenthia’s remarkable
suggests that many superficially similar species often ability to exploit novel and, evidently unrelated host
prove to be distantly related (Rota unpublished data). plants.
Brenthia collections at INBio1 (for Costa Rica) suggest A review ofliterature describing immature stages of
thereareatleast 10speciesinthecountry, mostofwhich 12 ehoreutid genera (Brenthia from Arita (1987),
are undescribed, and none of which is assignable to Williams (1951), and this paper; Litobrenthia,
pavonacella Furtherevidencesupportingthis revisionof Anthophila, Choreutis, Prochoreutis, Saptha, and
the species' distribution is that B. pavonacella larvae are Tebenna from Arita (1987); pupae of Anthophila,
strictlylimitedto Desmodium in NorthAmerica, andno Choreutis, Prochoreutis, and Tebenna from Patocka
Neotropical Brenthia have been reared from plants in (1999); Asterivora from Dugdale (1979); Rhobonda and
tins genus. Zodia from Rota (2005); Caloreas from Keifer (1937);
All known Brenthia larvae exhibit a similar escape Toidyra from Wille (1937)), as well as my own
behaviorinvolvingescapingthroughwormholes chewed investigation (unpublished data for Choreutis,
Hcmerophila, Prochoreutis, Tebenna, and Tortyra
),
TNBio-Institute) Nacionalde Biodiversidad suggestthefollowingcharacterstates assynapomorphies
Volume 62, Number 3 129
for the subfamily Brenthiinae: 1) hypognathous head Diakonoff, A. 1986. Glyphipterygidae auctorum sensu into. Mi-
S(sDe1mipsretoagnoanthoAu9s (inSDC1horleeuntgitnhaes)i;mil2a)retxotrLemesleytaelonign Dugdscaprleocleie,epsid|p.orpeSt.veiro1au9s7Pl9ay.laaeAsasricngteniwcead,gtevoonlSe.irmi7.aceGtn.haiBmsreaauulnco,troKtrahruelmsrN(uLeheewp.id4Zoe3pa6tleparpna.:d
Choreutinae); 3) A9with D1 setapresent (D1 absent in Choreutidae),with descriptionofanewspecies. NewZealand J.
C(hcoormepulteitneae)c;iraclned 4i)ncrCohcohreetustiinnaaen).incLoimkpelweistee,cirtchlee Dugdo1af9l9Ze8o.,ol.JT.h6S:e.,46sN1m.a—lP4.l6e6Kr.riMsitcernosleenp,idGo.ptSe.rRao-bgriandseons,up&erMf.amiJ.liSecso,blpep..
immatures of choreutine genera share numerous 219-220. In N. P. Kristensen(ed.),Lepidoptera, mothsandbut-
synapomorhies (Rota unpublished data). Togetherwith terflies. Volume 1: Evolution, systematic^, and biogeography.
Handbook for zoology. Volume IV Arthropoda: Insecta, (35).
preliminary molecular data (Rota unpublished data), Lepidoptera,mothsandbutteflies.WalterdeGruyter, Berlin.491
these different sets of shared derived characters for pp.
Brenthiinae and Choreutinae strongly suggest that both Forbes, W. T. M., 1923. Lepidopteraol NewYork andneighboring
states.PartI.Primitiveforms,Microlepidoptera,Pyraloids,Bom-
subfamilies are monophyletic, contra Dugdale et al. byces. Memoir 68, Cornell UniversityAgricultural Experiment
(1998) and in agreementwith Heppner and Duckworth Station Ithaca, NY. 729pp.
(1981). HHeeppppnneerr.. J. BB..,1&985W..WDe.stDuIcndkiweosrBtrhen.th1i9a8.1.InCsleacstsaifMicuantciohn1o:f1t3h-e26s.u-
Characterstatesthatappeartobesynapomorphies for perfamJi.lySesioidea(Lepidoptera: Ditrysia). Smithson. Contrib.
thegenusBrenthia are 1) crochetsin amesalpennelipse Zool.314:1-144.
(semicircle in Litobrenthia), 2) two SVsetae on A8 (one Hintoofnl,epHi.doE.pt1e9r4o6u.sOlanrvtahe,ewhiotmholsoogmyeannodtensoomnentchleapthuyrleoogfetnhyeosfetthaee
SV seta in Litobrenthia and choreutine genera except Lepidoptera.Trans. R. Entomol. Soc. Loncl.97: 1-37.
Rhobonda), 3) larval escape behavior through a Hodges, R. W., T. Dominick, D. R. Davis, D. C. Ferguson, J. G.
previously made hatch, 4) two pairs ofcurved spines in F19r8a3n.cClheemcokntL,istEo.lGt.heMLuenpirdooep,te&raJo.fAA.meProiwcealnlo,rtehtoafl.Me(xeidsc.o)..
the pupal cremaster, and 5) long forked setae on the E.W.Classey, London.284pp.
pupa. Characters3),4), and5)werenot discussedin the Kf.ifer, H. If 1937. California Microlepidoptera XII. Bull. Dept.
onlypublished description ofLitobrenthia (Arita 1987), Agric.California,Sacramento26:334—338.
Nichols, S.W. 1989.TheTorre-BuenoGlossaryofEntomology.The
so at this point it is impossible to say whether these NewYork Ent. Soc. and Amer. Mus. Nat. Hist., NewYork. 840
characters, or a subset of them, will prove to be PP
synapomorphic for Brenthia or are in fact PatoCka, |. 1999. UberdiePuppendermittel-unciwesteuropiiischen
Choreutidae (Lepidoptera: Choreutoidea). Entomol. Ber. 59:
synapomorphies for Brenthiinae. More work is needed, 53-58.
especially in the Neotropics where Brenthia appears to Robinson, G. S., K. R. Tuck, & M. Shaffer. 1994. Afieldguideto
be highly diverse, before we can begin answeringthese thesmallermothsofSouth-EastAsia, Malaysian NatureSociety,
KualaLumpur.309pp.
andotherquestions aboutthis fascinatinggroup. RobiMns.onH,erGn.aSn.,deP.z.R.2A0c0k7e.ryH,OIS.TJ.SKi-tachdiantga,baGs.eWo.f tBheecchaolsotpnlia,nt&sLo.f
Acknowledgements the world's Lepidoptera. http://www.nhm.ac.uk/researeh-
aspeIctasmogfrattheifsulprotjoecDta,vibudtL.espWeaciganlelyrffioerldhwisorkhelapndwictohmmneunmtersouosn Rotac,tuiorJ.nat2(i0Co0no3/s.ptraCohjReoiccrtaes)u/:thTiosadtaxpeolna(onLtmesyp/,.idUAolcptctreeasrsast)erduocfStueLrpaetSoeefmlbIvemarmBa2it0o0luo7rg.eicSatlagSetsa,-
earlier drafts ofthis manuscript. For hints on potential host plants
andLife Histoiy. Master'sThesis, UniversityofConnecticut. 160
and/or direct help with collecting, thanks are due to Lee A. Dyer,
Humberto Garcia, Grant Gentry, Terry L. Harrison, Henry A. pp.
TptiHodrueeocsncvpktoieild(fnleiehBcedetdMiibNdotyneHhs,Je).aLmbToaereunascidchsRonJnMiooic.nmdaaAlL.apnPaao(risBaewssuriicrsatketnoa,)idndacLsMene.adwrfiRUoisrozena(oCyUbaPtnSUrat.NiiienMSnsieot)nz.geafJnon.SardmEhIeeatMslnhpaBian.wmkniWatoKhghnieeytvasficmicnoweeualsRsssd RRoottaa,,uCmthiJiJo..md,ri2e&g0cue0tn5Die.t.dhraeLaLeia.)rrR.WvhaajoAlgunbnamnoen.pndirdEn.apngut2Wpo0aasm0llp6oki.lded.eePrsrrcSaerondicpadp.rtteZAiodoomrandt.ms/oiaro9msf8iH.:tceh3rpe7yPp-:LNn4oem7eoSe.rttra(oOLlpemNipacEiardlkopcmlth:oeoetrr4hae5s:.-
rmpSraeconvijuieesenccwctreeir(pFPtpio.sru:onFvdRuiaondtbediediornnthgeglKfrp.oafrnuCtltohlDicwsEoeBlmpl-rm,o0ej0neJ7tcot2sh7nc0oaT2n.meaaLnnoidnnegaipNranalrtoiti,eorfnearvtleoarmlsG.i.etoohnNgearotAaifpLohntAiahScle WSitlehlpdreoa,,in:yFJ..,10EW.D..1u3b1179u913q8/77uj..oeEu,IlmmaIbmloaa.wrtapr.uoenrn7eeo5.4id0n0eps0pel0.cot0ss4.b5r.Koetnedsadlel/laHuHnitguPeurblaiesnhienlgPCeorum-y
Societygrants7331-02and7751-04)andfromPennerandDeCoursey sucontrol. DirecciondeAgrieultura, GanaderiayColonizacion,
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tocosma Meyrick (Lep., Glyphipterygidae). Bull. Entomol. Res.
Natural Histoiy).
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