Journal of The Lepidopterists' Society Volume 55 2001 Number4 JournaloftheLepidopterists'Society 55(4),2001,129-139 REVIEWOFREBINEA RAZOWSKI AND ELIACHNA RAZOWSKI (TORTRICIDAE: EULIINI)— SISTER GROUPS ENDEMIC TO CHILE AND ARGENTINA John W. Brown SystematicEntomologyLaboratory,PlantSciencesInstitute,Agricultural ResearchService,U.S. DepartmentofAgriculture, c/oNationalMuseumofNaturalHistory,Washington,DC20560-0168,USA e-mail:[email protected] AND TsitsiY. McPherson c/oFacultyofNaturalSciencesorFacultyofArts,UniversityofGuyana,TurkeyenCampus, GreaterGeorgetown,GUYANA ABSTRACT. RebineaRazowskiandEliachnaRazowski,twoformerlymonotypicgeneraknownonlyfrommales,areredescribedbasedon largeseriesofspecimens(n =320)includingbothsexes.Aspresentlydefined,Rebineaismonotypic,withasinglevariablespecies,R. erebina (Butler, 1883),anditssynonym,ArotrophorabalsamodesMeyrick, 1931.Itispossiblethattwo(ormore)speciesareconcealedwithinthevari- ation,butwewereunabletoseparatethemusingtraditionalmorphologicalcharacters.Eliachnaisrepresentedbythreespecies:E.chileanaRa- zowski,1999,E.digitanaBrownandMcPherson,newspecies,andE.hemicordata BrownandMcPherson,newspecies.Bothgenera arerestrictedtosouth-centralChileandsouthwesternArgentina, rangingfromcoastallowlands (5m)tomiddleelevations (1200-1700m)at thesouthernendoftheAndes.Aphylogeneticanalysisofthefourspecies(plustwoout-groupspecies)providessupportforthesisterrelation- shipofRebineaandEliachnabasedonthefollowingsynapomorphies:(1)elongatelabialpalpi(length3-4timeshorizontaldiameterofdiecom- poundeye);(2)apairofstout,digitate,submedialprocessesonthedorsumofthetranstilla;(3)adeep,roundedexcavationnearthemid-venter ofthevalva;and(4)apairofsemicircular,lateralflangesfromtheposterioredgeofthesterigma. Additionalkeywords: newspecies,genitalia,phylogenetics,leafrollers. The tortricid fauna ofChile andAndean Argentina Rebinea and Eliachna based on new information, de- is distinct from that of the remainder of South scribe two new species ofEliachna, provide data on America, comprised primarily of endemic, or nearly the geographic distribution of the included species, endemic genera (e.g., Accuminulia Brown, Acman- and examine the phylogenetic relationship between thina Brown,Argentulia Brown, Chapoania Razowski, the two genera. Chileulia Powell,Eliachna Razowski,Haemateulia Ra- Materials and Methods zowski, Nesochoris Clarke, Proeulia Clarke, Rebinea Razowski, Recintonia Razowski, Varifula Razowski). Weexamined320pinnedadults ofRebinea andEli- Although the contributions ofRazowski (1995, 1999) achna deposited in the following institutions: Ameri- and Brown (1998, 2000a, b) have added substantially can MuseumofNatural History, NewYork, NewYork, to our knowledge of Euliini of this region, phyloge- U.S.A. (AMNH); The Natural History Museum, Lon- neticrelationshipsamonggeneraofthetribe areunre- don, England (BMNH); Mississippi Entomological solved and manyspecies remain undescribed. Priorto Museum, Mississippi State, Mississippi, U.S.A. this study, Rebinea and Eliachna were considered (MEM); Essig Museum ofEntomology, Universityof monotypic, known only from ahandful ofmales. The California, Berkeley, California, U.S.A. (UCB); Na- discovery ofadditional specimens ofthe two genera, tional Museum ofNatural History, Smithsonian Insti- includingbothsexes, reveals aclosephylogeneticrela- tution, Washington, D.C., U.S.A. (USNM); and Zoo- tionship between them and the presence oftwo new logical Museum, Copenhagen, Denmark (ZMC). species. The purposes ofthis paper are to redescribe Specimens were sorted by forewing pattern, geo- 130 Journalofthe Lepidopterists' Society graphic location, and sex. The resulting groups then Diagnosis. Adults ofRebinea are superficiallyand were examined for differences in male and female morphologically most similar to those of Eliachna genitalia, which have been shownto provide the most amongdescribed Euliinigenera. Thetwo share asim- reliable morphological features for distinguishing ilar forewing shape, size, andpattern; extremely elon- among related species ofTortricidae. Preparation of gate labial palpi (3-4 times horizontal diameterofthe genitaliaslides followedthe methodologysummarized compound eye); a pair of stout, digitate, submedial in Brown and Powell (1991). Because ofthe pheno- processes from the dorsum of the transtilla; a deep, typic similarity of, and variation within the treated rounded excavation near the mid-venter ofthe valva; species, we examined the genitalia of all male speci- and apair ofsemicircular, lateral flanges at the poste- mens. For undissected specimens, we used a fine rior edge ofthe sterigma. Superficially, males ofmany camel-hair brush to remove scales from the external Rebinea can be distinguished from Eliachna by their margin ofone valva, which provided enough detail to slightlylongerforewings andpalergroundcolor. Gen- convincingly assign all males to a species-level taxon. italia characters that distinguish Rebinea from Eli- Specimens were examined using aWild M3Z dissect- achna include its broad, straight aedeagus, with a ing microscope; slide mounted genitaliawere studied single, large, compound conutus; the absence of an usingthe dissectingmicroscope andaZeiss compound elongate, freeprocess atthe distalendofthesacculus; microscope. Illustrations ofgenitaliawere drawn with and an extremelyshort, broadductus bursae. the aid of a Ken-A-Vision microprojector (model Redescription. Head: Antennal cilia approximately 1.5 times X1000-1). Unless indicated otherwise, genitalia illus- width offlagellomere in male; cilia short, unmodified in female. Labialpalpusporrect,3-4timeshorizontaldiameterofeyeinboth trations areofasinglepreparation.Textdescriptionsof sexes (i.e., without dimorphism). Vertex with overhanging tuft of all characters are composite, based on all available scales. Proboscis present, presumably functional. Ocellus moder- specimens. Measurementsofforewingandlabialpalpi atelylarge.Chaetosemapresent.Thorax:Smoothscaled. Malewith- were made with an ocular micrometer mounted in a outforeleghairpencil. Forewing(Figs. 5,9): Length2.3-2.6times Wild M3Z dissecting microscope under low power width; lengthofDCca. 0.6timesforewinglength;widthofDCca. (xlO-16). Forewinglengthwas measured in astraight v0e.2intsismeepsarDaCtelbeengydoin;dCDuCA;,corhiogridnaataensd0.M6--s0.t7emalaobnsgenDt;CCluenPgtwhe;akal,l line from the base to the apex ofthe wing, including presentonlyatmargin. Noupraisedscaletufts;malewithoutcostal thefringe. Forewingwiddiwas measuredatdiewidest fold. Hindwing: Sc + Rand Rscloselyapproximateatbase; Rsand place perpendicular to the length measurement. Mjstalked;M andCuAjconnateorshort-stalked;CuPpresent;M- 3 Where available, aminimum of15 individuals ofeach stemabsent;tuftofhairlikescalesatbaseof1A+ 2Ainbothsexes. Abdomen:Smoothscaled;dorsalpitsabsent;nomodifiedcorethrog- sexwere measured. ColorsfollowRidgway(1912); ter- ynescalinginfemale. Malegenitalia(Fig. 10): Uncusshort,moder- minology for wing venation and genitalia structures ately stout, curved, strongly sclerotized; socius moderately short, follows Horak (1984). Abbreviations and symbols are hairy, drooping, slightly expanded distally; gnathos arms slender, as follows: DC = discal cell; ca. = circa (approxi- withadelicateterminalplate;transtillaatransversebandwithapair mately); n = number of individuals examined; x = ofstout, digitate, submedialprocessesondorsum, sometimeswith W tipsslightlyexpanded;valvabroadatbase,withdeep,roundedexca- mean; N, E, S, = compass points. vationnearmid-venter;sacculusrestrictedtobasalportionofvalva, Polymorphism or moderate variation in phenotype stronglysclerotized, endingin short, dorsal-projectinghook;pulvi- is uncommon to rare in most Euliini. However, many nus absent; vinculum well developed, strongly sclerotized; juxta euliine species in Chile andArgentina (e.g., Chileulia, large, stout arrowhead-shaped. Aedeagus large, straight, with a Proenlia, Haemateulia), including the two genera single large, compound, capitate comutus, consisting of several fusedfilaments;vesicafinelyspiculate. Femalegenitalia(Fig. 14): treatedhere, arevariablein forewingpattern andmac- Apophyses anterioresandposteriores moderate inlength, slender. ulation. Consequently, examination ofthe genitalia is Sterigmarelativelybroad,withnarrow,shallowlyU-shaped,sclero- the only reliable method for accurately identifying tizedbandandapairofsemicircularlateral flangesfromposterior species. Comparison with the illustrations provided is edge. Ductusbursaeextremelyshort, broad,withashortmembra- highlyrecommended. Formales, theprofileofasingle nousregionimmediatelyanteradofantrum; afrail,obovate,acces- sorybursafrom amoderatelylongductusoriginatingfrom dorsum valvais adequate; females mustbe dissected. ofcorpusbursae inposteriorone-third; corpus bursae moderately large, ovoid, with dense spiculae and alarge, irregularly rounded, Systematica sclerotizedplate,usuallyalonglateralwall. Distribution and biology. Rebinea is known from Rebinea Razowski, 1986 Chile andArgentinabetween about 30° and45°S lati- Rebinea Razowski, 1986:22; Powell et al., 1995:145; tude, ranging from the coastal lowlands (50 m) to the Razowski, 1999:84. middle elevations (1400 m) of the southern Andes Type species. Sericoris erebina Butler, 1883:72, by (Fig. 1). Collecting localities seem to have little in original designation. common in terms of habitat type, ranging from 12 1 Volume 55, Number 4 131 Nothofagus forest (e.g., Alto Tregualemu) to xeric ar- White to pale brown, usually with brownish-gray mottling. Ab- eas dominated by succulents and leguminous trees domen:Paleyellowbrowntodarkbrown.Genitalia(Fig. 10):Asde- scribedforgenus (drawnfrom USNM slide90383; 15preparations (e.g., Nague, Los Vilos) (see Davis 1986 for descrip- examined). Female. Head,thorax,andabdomen: Essentiallyasde- tions ofthe habitat at manyofthe collecting localities scribedformale. Forewing: Length 6.2-8.2 mm (x = 7.4; n = 15); listed below). Adults have been captured primarilyin groundcolorburnt umbertocinnamon,with indistinctpatches of October(n = 16), November(n = 118), andDecember orange-brown,orange-red,andcreamscales;avariablydefined,red- (n - 95), with afewspecimens recordedfromJanuary brown fasciafrom nearmid-costatodorsum, angledoutwardnear through March. The earlystages are unknown. mfirdodmlUeSofNwMinsgl.idGee8ni1t2a3l9i;a1(0Fipgr.ep14a)r:atAisondsesecxraimbiendedf)o.rgenus (drawn Remarks. Razowski (1986) included two species in Types. Holotype6(erebina),Chile, [Mountainsofthehacienda Rebinea: R. erebina (Butler) and R. balsamodes ofCauquenes, Butler1883] (BMNH). Lectotype(newdesignation) (Meyrick). Powell et al. (1995) synonymized the two 6 (balsamodes), Argentina, Territory Rio Negro, Lake Gutierrez, without explanation. Although the types of the two 3-14.XI.1926(F&M. Edwards, BMNH). Materialexamined.ARGENTINA:ChubutProvince:ElBolson, nominal taxaare fairlydistinctin forewingsize andfa- LagoPuelo,220m,26, 17.XI.1978, 16,29,21.XI.1978(MisionCi- des, they have extremelysimilar genitalia. The abun- entifica Danesa, ZMC), 1 9, 22.X.81, 3 9, 23.X.1981 (Nielsen & dance of material now available from Chile and Ar- Karsholt,ZMC);Esquel,LagoMenendez,ElSagrarioPuerto,600m, gentinasuggeststhatthetwoprobablyrepresentforms 16, 19,2-4.1.1982(Nielsen&Karsholt,ZMC). NeuquenProvince: of an extremely variable species. Alternatively, there LagoLacar,Pucara,750m,49, 10.XI.1978,26,29,25.XI.1978,56, may be more than one species concealed within this 29,1.XII.1978,36,79,2.XII.1978,56,39,3.XII.1978(MisionCien- tifica Danesa, ZMC); Lago Lacar, Pucara, 600 m, 21 6, 3 9, variation (see discussion below). 28-29.XI.1981, 16,19,26-27.XII.1981(Nielsen&Karsholt,ZMC); LagoLacar,5km E Hua-Hum,640m,26,39,6.XI.1981,26, 19, Rebinea erebina (Butler, 1883) 25.XI.1981, 1 9, 26-27.XII.1981 (Nielsen & Karsholt, ZMC); San (Figs. 1, 5, 9, 10, 14) MartindelosAndes,640m, 16, 14.X.1981, 6,19, 17-31.X.1981,4 6,16.X.1981,36,2.XI.1981,19,5-6.XI.1981,36,69,7-15.XI.1981, Sericoriserebina Butler, 1883:72. 19,26.XI.1981 (Nielsen&Karsholt,ZMC); San MartindelosAn- Arotrophora balsamodes Meyrick, 1931:381; Clarke, des,CerroChapelco, 1400-1600m,26,2-19.XII.1981 (Nielsen& Karsholt, ZMC). Rio Negro Province: San Carlos de Bariloche, 1963:8 [illustration ofadultandmale genitalia]. Colonia Suiza, 810 m, 1 9, 6.XI.1978, 1 o*, 15.XI.1978, 3 9, Rebinea balsamodes: Razowski, 1986:22. 28.XI.1978, 3 6, 29.XI.1978, 1 9, 4.XII.1978, 1 9, 5.XII.1978, 3 9, Rebinea erebina: Razowski, 1986:22 [illustration of 11.XII.1978,16,12.XII.1978,16,15.XII.1978, 19,9.1.1979(Mision male genitalia]; Powell et al., 1995:145; Razowski, CientificaDanesa,ZMC); SanCarlosde Bariloche, ColoniaSuiza, 1999:84. 800 m, 1 6, 26.X.1981, 1 6, 31.X.1981, 1 6, 11.XI.1981, 2 6,19, 12-20.XI.1981,46,21-22.XI.1981, 19,23.XI.1981,16,24.XI.1981, Diagnosis. Rebinea erebina resembles Eliachna 16,29,29^30.XI.1981,26,39,3.XII.1981, 6,19,5-6.XII.1981, species in general facies; most individuals (especially 6, 7.XII.1981, 1 6, 8.XII.1981, 1 9, 22.XII.1981, 1 9, 5-7.1.1982 (Nielsen & Karsholt, ZMC); San Carlos de Bariloche, Camino de females) cannot be separated by forewing length and Tronador,29,29.XI.1978 (Mision Cientifica Danesa, ZMC); Lago pattern. The malegenitaliaofR. erebina canbe distin- Nahuel Huapi, Puerto Blest, 770 m, 1 o", 27.XI.1978, 1 9, guished easily from those of Eliachna by its broad, 18.XII.1978, 16,30.XII.1978(MisionCientificaDanesa,ZMC),16, straight aedeagus with a large compound cornutus, 3.XII.1981 (Nielsen & Karsholt, ZMC). CHILE: Aconcagua and the absence ofthe elongate, free process of the Province: LosAndes,Curimon,700m, 16,28.111.1979(MisionCi- entificaDanesa,ZMC).ChiloeProvince:Puntra,ca.30airkmSAn- sacculus. In addition, the overall shape ofthe valva is cud,50m, 16, 19,21-22.XII.1981 (D. Davis, USNM). Coquimbo distinct, with a short hooklike process at the terminal Province: FrayJorge National Park, ca. 70 km WOvalle, 46, 29, end ofthe sacculus; the latter likely represents an au- 6-9.XI.1981 (D. &M. Davis, USNM); Nague, 11km N LosVilos, 1 tapomorphy for Rebinea. The female genitalia of R. 6, 4-5.XI.1981 (D. &. M. Davis, USNM); Coquimbo, 1 9, erebina are similar to those of Proeulia, Argentulia, 1.VII-19.X.1883, "Walker" (BMNH). Llanquihue Province: Casa andotherrelatedChilean-Argentineangenera,withan Pangue, 19(paralectotypeoibalsanwdes),4-10.XII.1926 (F. & M. Edwards, USNM); Peulla, 1 9 (paralectotype of balsamodes), extremelybroadductus bursaand an irregularlyscler- 12-13.XII.1926(F.&M. Edwards, BMNH). MauleProvince: Paso otized, highlyspiculate corpus bursa. Garcia,ca.23kmNWCauquenes,300m, 19,29-30.XI.1981(D.R. Redescription.Male. Head:Lowerfronspaletantocream;up- Davis, USNM); Rio Teno, ca. 40 km E Curico, 800 m, 1 6, 1 9, per frons gray brown. Labial palpus light brown. Thorax: Mostly 25-27.XI.1981 (D. R. Davis, USNM). Nuble Province: Alto brown.Forewing(Fig.5):Length6.6-8.8mm(x=7.9mm;n=25); Tregualemu,ca.20km SEChovellen,500m,59, 1-3.XII.1981 (D. groundcolorpalegraytoburntumber,withsmall,scattered, indis- R.Davis,USNM).OsornoProvince:P.N.Puyehue,Ag.Calientesto tinct patches oforange-brown, orange-red, and cream scales; fre- 3kmW,600m,26, 12-20.XII.1981 (D. R. Davis,USNM);Parque quentlywith faint,parallelbands indistalone-thirdofwingrepre- NacionalPuyehue,AguasCalientes,450m,6c*, 19,12.XI.1981,16, sented by series of slightly disjunct, darker scales; often with a 29, 13.XI.1981,66,59, 10.XII.1981,46, 19, 11.XII.1981, 16,69, variablydeveloped, diagonalfasciaofdarkerbrownfromnearmid- 12.XII.1981, 29, 13.XII.1981 (Nielsen& Karsholt,ZMC); Parque costato dorsum, angled outward near middle ofwing. Hindwing: Nacional Puyehue,Anticura,350m, 26,29, 17.XI.1981, 26, 29, 132 Journalofthe Lepidopterists' Society Figs. 1-4. GeographicdistributionofRebineaandEliachna. 1,R. erebina;2,E. digitana, newspecies;3,E. hemicordata,newspecies;4, E. chileana. Volume55, Number4 133 .^: | *^ _; _ ; „- • ^IP^ 7 5? g Figs.5-8. AdultsoiRebineaandEliachna. 5,fl. erebina;6,£. chileana; 7,£.hemicordata, newspecies;8,E. digitana, newspecies. 18.XI.1981,29, 19.XI.1981, 19, 15.XII.1981,39, 17.XII.1981, 19, bina). Although a majority of the specimens of this 18.XII.1981 (Nielsen & Karsholt, ZMC). Santiago Province: Los phenotypeare males, afewfemales approachthisgen- Maitenes, Colorado River, 33°22'S, 70°17'W, 1200-1400 m, 2 6, eralaspect. Attheotherextreme are specimenswith a 16X1954(L. Pena,USNM);Pilay,RioPueco,ca.45km SSantiago, 800m,16,23-24.XI.1981(D.R.Davis,USNM).ValdiviaProvince:20 shorter forewing length (6.6-7.7 mm), a darker (red km SValdivia, Rincon delaPiedra, 180m,29, 14.XI.1981, 16,39, brown to brown) forewing ground color, and a more 15.XI.1981 (Nielsen & Karsholt, ZMC). Valparaiso Province: Val- uniformly dark hindwing (similar to the type ofbal- paraiso, 1 6, 30.IX- 8.X.1883, "Walker 3074" (BMNH). Unknown samodes). Although this phenotype is typical of fe- Province:CentralAustral,26,1-III.1898(V.Izquerdo,USNM). males, some males approach this aspect (see Clarke Discussion. Rebinea erebina is either a single, 1963). Male genitalia are onlyslightlyvariable among highlyvariablespecies ortwo (ormore) extremelysim- all the specimens examined (n > 100), and the varia- ilarspeciesthatcannotbeseparatedreliablyusingtra- tion is concordant with neither differences in facies, ditional morphological characters. At one extreme are forewing length, nor geographic distribution. In some specimens with alarge forewing length (7.3-8.8 mm), male specimens, the distal end of the venter of the apale (grayto beige) forewing ground color, a poorly valva is somewhat pointed and slightlyreflexed, while definedforewingpattern, andapalehindwing, usually in others it is somewhat rounded. The width of the mottled with gray brown (similar to the type ofere- paired processes from the transtilla is somewhatvari- 134 Journalofthe Lepidopterists' Society able, as is the development ofthe hooklike process at middle,usuallywithdistalspineprojectingdorsoposteriorly;vesica the distal end of the sacculus. Apparent variation in withorwithoutminutespinules and/orsmallpatchoftinycornuti. Female genitalia (Figs. 15-16): Papillae anales somewhat slipper- the latter feature, however, is likelyan artifact ofslide shaped. Apophysesanteriores andposterioresslender, nearlyequal mountingofgenitalia. inlength. Sterigmausuallycrescent-shaped,withapairofrounded, Remarks. In his description ofArotrophora bal- sclerotizedlateralflangesatposterioredge. Ductusbursaemoder- samodes, Meyrick (1931) indicated that he had six ex- atelyshort;corpusbursaeovoid,withdensespicules,atleastinpos- amples from "Argentina, Territory Rio Negro, Lake teriorone-half. Gutierrez, November; S. Chile, Llanquihue Province, Distribution and biology. Eliachna apparentlyis Casa Pangue and Peulla, December." Three ofthese confined to south-central Chile and adjacent Ar- specimens areinthe BMNH, one ofwhichis afemale gentina, ranging from coastal lowlands (5 m) to mon- ofEliachna; one specimen is in USNM. Clarke (1963) tane areas (1700 m). Adults have been collected from identified the male from Argentina as "type" without OctobertoApril. Nothingisknownoftheearlystages. formally designating it as the lectotype. Because the Eliachna digitana Brown & McPherson, type series consists ofmore than one species, we for- new species mally designate a lectotype, and we select the speci- men labeled as such in the BMNH and identified as (Figs. 2,8, 11, 15) suchbyClarke (1963). This designationis necessaryto Diagnosis. Superficially, E. digitana is difficult to establish the concept of the species and promote distinguish from other species in the genus; the nomenclatural stability. forewing length is usually a little shorter and the ground color slightly more orange brown rather than gray brown. The male genitalia can be distinguished Eliachna Razowski, 1999 easily from its congeners by the subrectangular distal Eliachna Razowski 1999:87. portionofdievalva; theshort, straight, digitateprocess Type species. Eliachna chileana Razowski 1999:87, by at the termination of the sacculus; and the weakly monotypy. curvedaedeagus. Female genitalia, likewise, are easily Diagnosis. Eliachna is most similar to Rebinea in distinguished; the lateral pouches ofthe sterigma and forewing length, shape, pattern, andvenation (see di- the sclerotized, knoblike diverticulaofthe ductusbur- agnosis ofRebinea above for details), and most speci- sae are uniqueto this species. Description. Male. Head: Labialpalpusdarkbrown. Forewing mens are difficult to distinguished superficially from (Fig. 8): Length 5.9-7.0 mm (x = 6.5; n = 15); ground color and Rebinea. Genitalicdifferencesbetweenthetwogenera maculationsomewhatvariable;groundcolorusuallygoldgraytored areconspicuousandaredetailedaboveunderRebinea. brown,withfaint,darkbrownreticulations,infrequentlywithill-de- Redescription. Head: Antennal cilia approximately 1.5 times fined, darkerareain basalone-third; avariablydevelopedmedian width offlagellomere in male; cilia short, unmodified in female. fasciafromcostatodorsalmargin,browntoredbrown,angledout- Labialpalpuselongate,porrect,length3-4timeshorizontaldiame- ward near middle offorewing; apex frequentlywith darkerpatch. ter ofcompound eye, slightly longer in female. Vertex with over- Hindwing: Brownish gray, infrequently with faint mottling. Ab- hanging tuft ofscales. Proboscis present, presumably functional. domen: Goldbrowntodarkbrown. Genitalia: AsinFig. 11 (drawn Ocellus moderately large. Chaetosema present. Thorax: Smooth from USNM slide90484; 7preparationsexamined). Uncus,socius, scaled.Legsunmodified,malewithoutforeleghairpencil.Forewing gnathos, andtranstillaas describedforthegenus. Valvalong, sub- (Figs. 6-8): Lengthca. 2.4timeswidth;lengthofDCca. 0.6times rectangular,onlyslightlynarrowednearmiddleandslightlybroad- forewinglength;widthofDCca. 0.2timesDClength; CuA2origi- eneddistally;sacculusbroadbasally,withafree,mostlystraight,dig- nates 0.6-0.7 along DC length; all veins separate beyond DC; itate process distally. Juxta as described for the genus. Aedeagus chordaand M-stem absent; CuPweak,presentonlyatmargin. No weakly undulate; spine at termination ofcoecum strongly sclero- upraisedscaletufts;malewithoutcostalfold. Hindwing: Sc+ Rand tized,slightlydisjunctfrom coecum;vesicadenselypunctate, espe- Rscloselyapproximateatbase; RsandM stalkedca.one-thirddis- ciallyindistalthree-fourths;arowofminutecornutinearthebase. tance from DC to margin; M and CuA 1connate orshort-stalked; Female. Essentiallyasdescribedformale. Forewinglength6.5-7.6 CuPpresent; M-stemabsent;3tuftofhairtlikescalesatbaseof1A+ mm (x = 6.9; n = 15). Genitalia: Asin Fig. 1.5 (drawnfrom USNM 2Ainbothsexes.Abdomen: Dorsalpitsabsent;nomodifiedcoreth- slide 81228; 6 preparations examined). Sterigma subrectangular, rogynescalinginfemales. Malegenitalia(Figs. 11-13): Uncusslen- withapairofshallowlateralpouchesatanterioredge. Ductusbur- der, short, simple, strongly sclerotized; socius moderately short, sae short,withashort, stout, knoblike, sclerotizeddiverticulador- broad,haiiy, slightlyexpandeddistally;gnathoswithslenderlateral sally;corpusbursaeoblong,uniformlycoveredwithfinespinules. armsconnectedtoterminalplatebymembrane.Transtillaasimple Type. Holotype6,Chile, Nuble Province, 17.5km S Curanipe, band,highlysclerotizedlaterally,weakermedially,withapairofsub- near coastal stream, 50 m, 25.1.1979 (D. & M. Davis & B. Aker- medial, digitate processes, slightly rounded apically. Valva broad bergs,USNM). basally,withvariableexcavationnearmid-venter; sacculuswellde- Paratypes. ARGENTINA: Chubut Province: El Bolson, Lago fined,witiifree,elongate-digitateterminalprocessofvariableshape Puelo,220m, 16,18.XI.1978(MisionCientificaDanesa,ZMC),19, andlength;pulvinusabsent;juxtastronglysclerotized, stoutarrow- 13.X.1981 (Nielsen&Karsholt,ZMC);Esquel,550m, 16, 1.1.1982 head-shaped. Aedeagus somewhat elongate, variably curved near (Nielsen&Karsholt,ZMC); SierraColorada,800m, 16,29.1.1983 Volume 55, Number 4 135 (M.&P.Gentili,USNM).NeuquenProvince:SanMartindeIosAn- des,640m, 19, 13X1981, 1d, 7-15.XI.1981 (Nielsen&Karsholt, ZMC); LagoLucar,Pacara,650m, 1d, 10.XI.1978(MisionCienti- fica Danesa, ZMC), 1 d, 26-27.XII.1981 (Nielsen & Karsholt, ZMC). Rio Negro Province: Lago Gutierrez, 1 9 (paralectotype of balsmodes),3-14.XI.1926 (F. & M. Edwards, BMNH); San Carlos de Bariloche, Colonia Suiza, 810 m, 1 6, 2.XII.1981, 1 9, 20.XII.1981, 1d,23.XII.1981, 1 d,5-7.1.1982(Nielsen&Karsholt, ZMC), 1 9, 9.XI.1978, 1 d, 19.XI.1978, 1 d, 29.XI.1978, 1 9, 9.XII.1978, 1 9, 12.XII.1978, 1 9, 9.1.1979, 1 9, 10.1.1979, 1 9, 11.1.1979 (Mision Cienti'fica Danesa, ZMC). CHILE: Cautin Province: Fundo Neltume, 2 km N Villarrica, 200 m, 1 d, 1 9, 27.11.1979 (D. & M. Davis & B. Akerbergs, USNM); Fundo el Coigue,27kmNEVillarrica,500m,2d,28.11-3.III.1979(D.&M. Davis&B.Akerbergs,USNM). MallecoProvince:RioManzanares, 59,19.X.1979(Flint&Barria,USNM).NubleProvince:ForelCar- rizalillo, 250 m, 1 9, 30.I-5.II.1981 (L. E. Pena, USNM); Alto Tregualemu, 500 m, 1 9, 27-28.1.1981 (L. E. Pena, USNM); Alto Tregualemu,ca.20kmSEChovellen,500m,2d,26-27.1.1979(D. &. M. Davis & B. Akerbergs, UCB),49, 1-3.XII.1981 (D. Davis, USNM); 17.5km S Curanipe, nearcoastal stream, 50m,3d,29, 25.1.1979 (D. & M. Davis & B. Akerbergs, USNM); Pierdrade la Iglesia,8km N Cobquecura,5m,49,25.1.1979(D. &M. Davis& B. Akerbergs, USNM), 1 d, 4 9, 24.1.1979 (D. & M. Davis & B. Akerbergs,USNM);Cachapoal,CajondeLisboa,Alhue,800m,1d, 19-21.XII.1987(L. E. Pefia, USNM). Llanquihue Province: Llan- quihue, Petrohue, 1 d, 8.III.959, 1 d, 12.111.959 (J. F G. Clarke, USNM).OsomoProvince:ParqueNacionalPuheyue,Anticura,250 m, 1d, 17.XII.1981(Nielsen&Karsholt,ZMC). SantiagoProvince: Rinconada Maipu, 450 m, 35°31'S, 70°47'W, 1 d, 14.IV.1966 (W. Hichins & M. E. Irwin, UCB);Pilay, RioPeuco,ca.45kmS Santi- ago, 800 m, 1 d, 23-24.XI.198l' (D. Davis, USNM). Valdivia Province:Valdivia, 19,7.III.1960(E.Krahmer,ZMC);20kmSVal- divia,RincondePiedra, 180m, 1d,24.XI.1981(Nielsen&Karsholt, ZMC). Distribution and biology. Eliachna digitana oc- curs from Santiago Province, Chile, to Chubut Province, Argentina (Fig. 2), ranging from coastal Nothofagus forests (5 m) to arid uplands (1300 m) dominated by Fabaceae and Lauraceae. Capture records are from October (n = 3), November (n = 5), Figs. 9-10. Rebineaerebina.9,Wingvenation; 10, Malegeni- December (n = 11), January (n = 20), February (n = talia,aedeagusremoved,valvaespread. 4), March (n = 3), andApril (n = 1). Nothingis known ofthe earlystages. portionofthevalva; theelongate, curved, digitate, free Etymology. The species name refers to the digitate process ofthe sacculus is similartothatofE. chileana. processthatcomprisesthedistalportionofthesacculus. The female is unknown. Remarks. N. Obraztsov probably was the first to BroDwensctroipdtairoknb:roMwanl.e.FoHreeawdi:ngLa(bFiiagl.p7)a:lpLuesngddairk7.b8r-o8w.n0.mTmhor(axx:= recognize this species as distinct and undescribed; he 7.9;n = 5);groundcolorpaleorangecream,widitinyblackspecks labeled the specimen from Llanquihue, Peulla throughout; moderatelybroadmedianfascia,extendingfromcosta (USNM) with a manuscriptname. ca. 0.6-0.7 from base to apex, to dorsum ca. 0.7-0.8 from baseto tornus, angled outward near middle offorewing Hindwing: Pale Eliachna hemicordata Brown & McPherson, gray brown with variably developed darker mottling. Abdomen: new species Gold brown to dark brown. Genitalia: As in Fig. 13 (drawn from USNM slide 81222; 7 preparations examined). Uncus, socius, (Figs. 7, 13) gnathos,transtillaasdescribedforgenus.Valvalong, distalportion Diagnosis. Eliachna hemicordata has a slighter ovoid,widishorthooklikeprocessfromventerofapex;sacculuswith greater forewinglength and apalerground colorthan elongate,slightlyflattened,weaklycurvedfreeprocess.Juxtaasde- scribed for genus. Aedeagus curved dorsadjust beyond coecum, otherspecies inthegenus. Malescanbe distinguished with dorsoposteriorlyprojectingspineatdistalend;vesicawithout from other Eliachna by the somewhat cordate distal spicules. Female.Unknown. 136 Journalofthe Lepidopterists' Society Figs. 14-16. FemalegenitaliaofRebineaandEliachna. 14,Re- bineaerebina; 15,E.digitana,newspecies; 16,E. chileana. fromJanuary(n = 3), February(n = 2), andDecember (n = 3). Nothingis known ofthe earlystages. Etymology. The species name refers to the half- Figs. 11-13. MalegenitaliaofEliachna,aedeagusremoved,val- hearted shape ofthe distalportion ofthevalva. vae spread. 11, E. digitana, new species; 12, E. chileana, new species; 13,E. hemicordata. Eliachna chileana Razowski, 1999 (Figs. 6, 12, 16) Type. Holotype6,Argentina, Nuequen, Chapelco Lenga, 1700 Eliachna chileana Razowski, 1999:88 [male genitalia m,24.1.1984(M. &P. Gentili,USNM). illustrated]. Paratypes.ARGENTINA:NeuquenProvince:ChapelcoTechos, 1400 m, 1 6, 21.1.1982 (M. & P. Gentili, USNM); Lago Lacar, Diagnosis. The malegenitaliaofE. chileana canbe Trompul,1200m,1i,6.1.1983(M.&P.Gentili,USNM);SanMartin distinguished from those ofotherspecies inthe genus de losAndes, Tr. Kura, 1000 m, 1 6, 29.XII.1985 (M. & P. Gentili, bythegreatlyexpandeddistalportionofthevalva, ter- USNM).RioNegroProvince:LagoNahuelHuapi,PuertoBlest, 16, minatingin an attenuate, pointedtip. The female gen- 23.XII. 1978 (Mision Cientifica Danesa, ZMC). CHILE: Bio-Bio Province: Lago El Barco, Guallali, Sta. Barbara, 1200 m, 1 6, italia can be distinguish by the simple U-shaped 25-28.11.1981 (L. E. Pena, USNM). Cautin Province: [Parque Na- sterigma. cional] Conguillio, 1200 m, 1 6, 4-8.II.1988 (L. E. Pena, USNM). Redescription. Male. Head: Labialpalpuslightbrown. Thorax: UnknownProvince:V.Villarica, 16km SPucon, 16,20.XII.1982(R. Mostlybrown.Forewing(Fig.6): Length7.1-7.8mm(x=7.5mm;n Brown, MEM). =4);groundcolordullsilverygray,faintlyoverscaledwithredorange Distribution and biology. Eliachna hemicordata and copper orange; basal one-fourth usually with patch ofslightly is known from Neuquen andRio Negroprovinces,Ar- darker scales; variably defined, red-brown median fascia from near mid-costatodorsum,angledoutwardnearmiddleofforewing;termi- gentina, and Bio-Bio and Cautinprovinces, Chile, be- nal areawith irregular patches ofblack and orange-red scales, de- tween 1000 and 1400 m (Fig. 3). Capture records are creasingtowardapex. Hindwing: Paleolivebrown,withfaintbrown- Volume 55, Number4 137 17 Distribution and biology. Eliachna chileana is / £ / /^> known only from Malleco and Nuble provinces (Fig. / / /a /J> # / 4s)p.ecWieisthappalelarcsapttourbeesrbesettrwiceteend t1o3h0i0g-h1e6r00elemv,atitohniss V V v V V than its congeners. Capture records are from Decem- ^' ber (n = 1) andJanuary (n = 10). Nothingis known of the earlystages. 7--1 Explanation of Characters and 10--2 PhylogeneticAnalysis B- -1 8--3 A phylogenetic analysis was performed on the four species thatcompriseRebinea andEliachna, plus two 14--1 12--1 out-group species, Proeulia triquetra Obraztsov and 11--1 Haemateulia haematitis (Meyrick). The use ofProeu- | 180----12 lia and Haemateulia as out-groups is somewhat arbi- trary because sister group relationships within this 13- clade of Euliini previously have not been demon- 9- strated. However, the two genera share a variety of 5- features with Rebinea and Eliachna (e.g., forewing 4- 3- pattern, polymorphism, broad, short valvae, etc.), and 2--1 1--2 all appearto belongto acomplexofendemic Chilean- Argentinean genera that is taxonomically isolated 14--2 from other Euliinipresentin SouthAmerica. Thepri- 12--1 11--1 15--1 marypurposes ofthe analysiswereto confirm the sis- 10--1 8--1 ter relationship ofRebinea and Eliachna, and ensure 3--1 1--1 that the two in-group genera are monophyletic with respect to each other. The analysis was based on 15 morphologicalcharacters (11 binaryand4 multistate), including two ofthe head, one ofthe thorax, nine of the male genitalia, and three ofthe female genitalia. taxaF.iNg.um1b7e.rsHoynpotthheesleifstroeffeprhtyolcohgaernaecttiecrsr(e1l-a1t5i)o;nsnhuipmbaemrosnogntthhee Character state polaritywas determined through the rightrefertocharacterstates(seeTable 1). out-group method and using Horak's (1984) assess- ment ofcharacters ofTortricinae. The characterstate ish-grayreticulations. Abdomen: Paleyellowbrown to darkbrown. data were subjected to parsimony analysis using the Genitalia: AsinFig. 12 (drawnfrom USNM slide81223;5prepa- "mhennig*" command ofHennig86 (Lipscomb 1994). rationsexamined).Uncus,socius,andgnathosasdescribedforgenus. Characters usedin the analysis are listed anddiscussed Valvabroad basally, narrowedat middle, greatlyexpanded distally, withelongate,curved,beaklikeprocess directedventrally; sacculus brieflybelow;thecharactermatrixispresentinTable 1. withelongate,free,slightlyflattened,curved,digitateprocess.Juxtaas 1. Labial palpi: (0) upturned, ca. 1.5-2.0 as long as describedforgenus. Aedeagus stronglycurvednearmiddle;vesica horizontal diameter ofthe compound eye; (1) some- wEisstehntsieavlelryalasmidneusctreisbpeidnfeosramnadlea.fFeowreswhoirntg,:caLpeintgattehc6o.m7u-t7i.0.mFemma(lxe.= what porrect, ca. 2.0-3.0 as long as horizontal diame- 6.9;n=4). Genitalia:AsinFig. 16(drawnfromUSNMslides90493 ter ofthe compound eye; (2) porrect, 3-4 times the and90177;5preparationsexamined).SterigmaweaklyU-shaped,uni- horizontal diameterofthe compoundeye. Whileelon- formindiickness.Ductusbursaemoderatelyshort,widilargeductus gate labial palpi occur in several groups scattered seminalisoriginatingdorsallyaboutmidwaybetweenostiumandcor- throughout Euliini (e.g., Proeulia, Seticosta Razowski, pusbursae.Corpusbursaeasdescribedforthegenus. Anopinella Powell, etc.), none ofthese taxahave palpi la,TCyhpiel.iaHnolaorteay,peScJE,ChRielcei,ntNou,bl1e5P0r0ovmin,ce,15L.aXsIIT.r1a9n8c3as,(LS.haPnegfriia-, as longas those ofRebinea andEliachna. AMNH). 2. Maleantennalcilia: (0) conspicuous, elongate, ca. Additional specimens examined. CHILE: MallecoProvince: 1.0-1.5 times thewidth ofthe flagellomere; (1) incon- nr. Los Gringos Camp, Nahuelbuta Nat. Park, 1300 m, 3 <$, 2 9, spicuous, extremely short, ca. 0.5 times the width of As6i-kd1ee1r.bV1eo.rl1gc9sa8,n2CU(hSDi.lNiMRa)n.,;D1aLv6ia0ss0,TmUr,aSn1Nc6Ma,s),1.92-N12u1kb.l1m.e1EP9r7Ro9evc(iiDnn.cte&o:.,SMhnae.nagDrraihv-iilgsah,&wSaBW-. ithsehefdlasgoemlelowmhearte.bTyhietsvvaalriuaebiolfittyhiisncthhaeraocutte-rgrisoudpimtianx-a terfall, 1300m, 16,39, 17.1.1979(D. &. M. Davis&B.Akerbergs, (e.g., length of male antennal cilia varies among USNM). species ofProeulia). . 138 Journalofthe Lepidopterists' Society Table1. Charactermatrix("?"= missingdata). considered a synapomorphy for E. chileana and E. hemicordata. haematitis 00100 00001 11020 triquetra 11000 00100 00001 9. Juxta: (0) shield-shaped, unmodifed; (1) stout ar- erebina 20111 00110 00101 rowhead-shaped. The presence ofa stout, arrowhead- digitana 20111 10211 11111 shapedjuxtais notparticularlycompellingas asynapo- chileana 20111 01312 11111 morphy for Eliachna and Rebinea because other hemicordata 20111 00312 11??? genera ofChilean-Argentinean Euliini maypossess a similarlyshapedjuxta. The structure frequentlyis not 3. Male foreleg hairpencil: (0) present; (1) absent. included in illustrations of male genitalia or is dis- Thepresenceofamaleforeleghairpencilis assumedto tortedbyslide mounting. represent the plesiomorphic condition in Euliini 10. Aedeagus: (0)broad, straight, relativelylarge; (1) (Brown 1990). However, because the structure is evo- slightlymoreslender,weaklycurved; (2) conspicuously lutionarily labile, there is no evidence that its shared more slender, strongly curved. A broad, straight, absenceistrulyasynapomorphyforthetaxathatlackit. relativelylargeaedeagus, characteristicofProeulia and 4. Transtilla: (0) a simple bridge; (1) with a pair of Rebinea, is consideredtheplesiomorphic condition. A stout, digitate, submedial processes on dorsum. slightly more slender, weakly curved aedeagus is con- Althoughdigitatestructures arepresentonthedorsum sidered the first step in a transformation series lead- ofthe transtilla ofInape Razowski and Ortognathosia ing to a conspicuously more slender, strongly curved Razowski (see Razowski 1988 for illustrations), few aedeagus. other features ofthe male or female genitalia ofthe 11. Aedeagus: (0) without external projections; (1) lattertwo generaindicate aclose relationshipwithRe- with a small dorsoposteriorly projecting spine from binea and Eliachna. Hence it is suspected that the nearjunction ofthe coecum and the phallobase. The structures represent convergent development in In- latter character state appears to represent a synapo- ape, Ortognathosia, andRebinea + Eliachna. morphyforEliachna. 5. Valva: (0) venteruniform; (1)venterwithadeeply 12. Vesica: (0)withoneorfewlargecapitatecornuti; excavated portion near middle resulting in a broad (1)withnumeroustinynon-capitatecornuti. Thepres- basalportion, anarrow"neck"nearthe middle, andan enceofoneorfewlargecornuti, assumedtorepresent expanded distal portion. The distinctive shape ofthe the plesiomorphic condition, is typicalofProeulia; the valva is apparently unique to Rebinea and Eliachna, vesica of Rebinea, likewise, has a single, large, com- and is reminiscent of the valva of some Eucosmiini poundcornutus. (Olethreutinae). 13. Sterigma: (0) unmodified; (1) with a pair of 6. Valva: (0) distalone-thirdsomewhat club-shaped; semicircular sclertotized flanges located at the poste- (1) distal portion narrowed, somewhat elongate- rior edge. The latter character state appears to repre- rectangular. Thelattercharacterstate is consideredan sent asynapomorphyforRebinea andEliachna autapomorphyforE. digitana. 14. Ductus bursae: (0) extremely broad, nearly as 7. Valva: (0) distalone-thirdsomewhat club-shaped; broad as the corpus bursae; (1) slightly more narrow, (1) distal portion greatly expanded into an elongate, weakly differentiated from corpus bursae; (2) rela- curved, beaklike process directedventrally The latter tively narrow, clearly differentiated from corpus character state is considered an autapomorphy for E. bursae. In Proeulia, Argentulia, and related Chilean- chileana. Argentinean genera, the ductus bursae is extremely 8. Sacculus: (0) weak, lacking free terminal broad. This condition, considered the plesiomorphic process; (1) well-defined, with short, free, distal ter- state, alsoispresentinRebinea. The slightlymorenar- mination; (2) well-defined, with slender, digitate rowductus bursae ofEliachna is consideredasynapo- process; (3)well-defined, with long, slightlyflattened, morphyforthis genus, and the relativelywell defined curved process. A sacculus lacking a free terminal ductus bursae ofHaemateulia is considered the most process, such as that in the genitalia ofHaemateulia, advanced state. Becausethis charactervaries through- is consideredtheplesiomorphiccondition. Awell-de- out Euliini (sometimes evenwithin asingle genus), it fined sacculus with a short, free, distal termination, is aless compellingindicatorofrelationship. such as thatin the genitaliaofProeulia, is considered 15. Corpus bursae: (0) finely punctate; (1) densely derived; and the development ofthe free tip into an spiculate. A denselyspiculate corpus bursae is shared elongate, digitate process is consideredasynapomor- byProeulia, Rebinea, Eliachna, Argentulia, and addi- phyforEliachna. Its further developmentinto anex- tional related Chilean-Argentinean genera. The finely tremely long, slightly flattened, curved process is punctate corpus bursaofHaemateulia, similarto most