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Bulletin of the British Museum (Natural History) Entomology PDF

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Preview Bulletin of the British Museum (Natural History) Entomology

Bulletin Museum British (Natural History) Entomology Series VOLUME NUMBER DECEMBER 60 2 19 1991 The Bulletin ofthe BritishMuseum (NaturalHistory), instituted in 1949, isissued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and an Historicalseries. The Entomology Seriesisproduced undertheeditorshipofthe KeeperofEntomology: Dr L. A. Mound Publications Manager(Entomology): DrP. C. Barnard Papersin the Bulletin areprimarily the resultsofresearchcarried out on the uniqueandever-growingcollectionsofthe Museum, both by thescientificstaff and by specialistsfrom elsewherewhomakeuseofthe Museum'sresources. Many ofthepapersareworksofreference that will remain indispensablefor years tocome. A volumecontainsabout288 pages, made upoftwo numbers: published Spring and Autumn. Subscriptionsmay beplaced forone ormore oftheserieson an Annual basis. Individual numbersand backnumberscan bepurchased anda Bulletincatalogue, by series, isavailable. Orders andenquiriesshould besent to: Sales Department, Natural History Museum Publications, British Museum (Natural History), Cromwell Road, London SW7 5BD Telephone:071-938-9386 Fax- 071-938-9212 WorldListabbreviation: Bull. Br. Mus. nat. Hist. (Ent.) © British Museum (Natural History), 1991 ISBN 565 06042 2 Entomology Series ISSN 0524-6431 Vol60,No.2,pp.205-288 British Museum (Natural History) Cromwell Road London SW7 5BD Issued 19 December 1991 TypesetbyJ&LComposition,Filey,N.Yorkshire PrintedinGreatBritainbyHenryLingLtd,Dorchester,Dorset Bull. Br. Mus.nat. Hist. (Ent.)60(2):205-241 Issued19December1991 Sattleria: a European genus of brachypterous alpine moths (Lepidoptera: Gelechiidae) LINDA M. PITKIN & K. SATTLER DepartmentofEntomology, The NaturalHistory Museum, CromwellRoad, London SW75BD CONTENTS Synopsis 205 Introduction 205 MaterialandMethods 207 Abbreviations 208 Acknowledgements 209 ThesystematicpositionofSattleria 209 SattleriaPovolny 210 ChecklistofthespeciesofSattleria 214 KeytothespeciesofSattleria 215 References 240 Index 241 SYNOPSIS.ThegenusSattleriaPovolny(Gelechiinae,Gnorimoschemini),endemictothe alpine zone of European mountains, is revised. Eight species, including four newly described, are recognized as valid; one species is recalled from synonymy, one taxon reverts from subspecific to specific status, one subspecies is transferred to a different speciesandone synonym is newlyestablished. Separate keysfor males and females are provided,basedoncharactersofthegenitalia. Maleandfemalemothsandtheirgenitalia are describedand illustrated. the disposal of refuse add to the pressure on the INTRODUCTION environment. At the same time the agricultural sector's drive for greater profitability contributes through excessive application of fertilizers to In many parts ofEurope the alpine zone was the reduced biodiversity even on remote alpine pas- last natural ecosystem almost undisturbed by tures, whilst long-range industrial air pollution man's activities and, as long as the mountains appearstoberesponsiblefordramaticdie-backof were remote and inaccessible, their unique flora alpine forests with resultant soil erosion (see, for andfaunacouldbeconsideredrelativelysafefrom example, B\ab etal., 1987: 170-177; Diem, 1988: destruction bymasstourism,intensiveagriculture 2-8). or industrial processes. Increasing recreational Amongst the insects likely to be threatened by useofthemountainsbyalarge,evermoreaffluent such degradation of their habitats a number of population, encouraged by commercial interests Lepidoptera species are particularly vulnerable, eagertoexploit Europe'slasttouristicfrontier, is because their females are flightless and they are now causing widespread overdevelopment and often confined to small disjunct colonies. An simultaneousdestructionorfragmentationofsen- increasedtrendtowardsflightlessnesswithamore sitive natural habitats. Thus, numerous areas orlesspronouncedreductionofthewingshaslong previously inaccessible to a wider public have been known tooccurinthefemalesofLepidoptera been opened up totourism through the construc- inhabiting the alpine zone of high mountains in tion ofnewroads, hotels, holidaychalets, ski lifts many parts of the world. Such abortion of flight andpistes,cablecarsandotherfacilities,andwith and reduction ofthe flight organs as an apparent the growing number ofvisitors such problems as responsetothe ecologicalconditionsin an extreme 206 LINDAM. PITKIN&K.SATTLER environment is not confined to high altitude representing one variable species, but would not speciesbut is also observed in many Lepidoptera exclude the possibility that some of the more endemic to small remote oceanic islands and in extreme morphologically separable forms might species which are"active as adults during the cold be distinctspecies. Atthesametime hedismissed season; the whole phenomenon is extensively Gelechia pyrenaica Petry as falling within the reviewed by Sattler (1991). range of variation of dzieduszyckii, and sub- IntheEuropean AlpsexamplesofLepidoptera sequently synonymized it formally with the latter species with wing reduction in the female sex are (Povolny, 1967: 175), although Petry had been foundin atleastsevendifferentfamilies(Huemer emphatic that both were distinct. As Povolny's & Sattler, 1989: 257) but the total number of views had never been challenged it came as a species, which is in excess of20, is still uncertain surprise when, during a field trip by members BMNH becauseseveralunresolvedspecies-complexesare of the Microlepidoptera Section (G. S. involved. Onesuchcomplexisthegelechiidgenus Robinson, K. Sattler & K. R. C. Tuck) in 1981, SattleriaPovolny(Gelechiinae: Gnorimoschemini), Petry'sobservations in the central Pyreneeswere which is endemic to Europe and inhabits the fully confirmed. On the slopes of Pic du Midi de highermountainsfromthePyreneesinthewestto Bigorre twoSattleria species, clearly separable in theCarpathiansintheeastinanumberofdisjunct bothsexesexternally(Fig. 1)andbytheirgenitalia, populations. It is likely that, together with their werefoundtoflytogetherin the samebiotope. In habitats, many of its taxa are endangered; how- the light of these findings it was decided to re- ever, to obtain meaningful data for assessing the assessthe taxonomicstatusofallSattleriapopula- impactofadversefactorson the alpine ecosystem tions because it is not unreasonable to assume and its constituent parts, it is imperative first to that, iftwo undisputed species occur side by side understand the taxonomy of the organisms in- in the central Pyrenees, other morphologically volved. It is therefore a purpose ofthis paper to separable forms elsewhere may also represent elucidatethehithertounresolvedtaxonomyofthe distinct species. known Sattleria populations and summarize the Preliminary results of our studies were pre- limited available ecological data. sented at the Sixth Innsbrucker Lepidopterolo- WhenheproposedthegenusSattleria, Povolny gengesprach, held on 20-21 October 1984 at the (1965:490-492)wasambivalentaboutthesystem- TirolerLandesmuseumFerdinandeum,wherewe atic status ofthe included taxa. He regarded the producedevidencetosupportourviewthatSattleria genusasmonotypic,withS. dzieduszyckii(Nowicki) comprisesacomplexofseveralcloselyrelatedbut Fig. 1 Males (left) andfemales (right) oftwospeciesofSattleriafromthe Pyrenees: S. arcuata (top), S.pyrenaica (bottom). SATTLER1A AEUROPEANGENUSOFBRACHYPTEROUSALPINEMOTHS 207 : morphologically distinguishable species. A second- most of the material recorded in earlier publica- handreportonourpresentationpromptedPovolny tions, usually as Gelechia dzieduszyckiiNowicki, (1987) to pre-empt our planned publication by a wasre-assessed, includingseveral ofthe specimens hastily produced paper in which he reiterated his illustratedorotherwise itemizedby Povolny(1965; beliefindzieduszyckiiasasinglevariablespecies. 1967; 1983; 1987). We were unable to trace any In essence he recognized several geographical examplesfrom Bulgaria(Rebel, 1903: 329)whilst forms, variously referred to as 'Rassenkomplex', the only known specimens from Montenegro 'Rassenkreis', 'Populationsgruppe'orsubspecies, (Rebel, 1913: 330) and Albania (Rebel & Zerny, butemphasizedthat,forthetime being, theseare 1931; 146),whichbelongtoNM,Vienna,havenot to be taken as representing merely statistical yet been returned by Povolny and were not trends. At the same time he stated that it is examined. impossibletoassign individual mothsofunknown We studied 117 genitalia slides (84 males, 33 geographic origin unambiguously to any ofthose females), most of which were specifically pre- subspecific groups. A key factor in his argument pared for this work. The male genitalia were was the strong variation in size, coloration and 'unrolled' in accordance with the technique pre- genitalia structures he believed to have observed viously described (Pitkin, 1986). Originally in his specimens from the Pyrenees; however, developed for the gelechiid genus Mirificarma from his illustrated examples it is clear that he Gozmany, this method also proved ideal for misinterpreted as intraspecific variation the dif- Sattleria males; without sacrificing any critical ferences between the widespread 5. pyrenaica characterit allowsthetwo-dimensional displayof (Pctry) and two geographically more restricted all taxonomically important structures - a pre- and hithertoundescribedspecies. Thushiscolour requisite for successful photomicrography. The figures, intended to demonstrate a remarkable advantagesofthe unrollingtechnique forinterpret- range ofvariation in '5. dzieduszyckiipyrenaica ing complex genitalia structures are immediately , represent two males ofS. pyrenaica (Pctry) (figs obviouswhen ourillustrationsarc comparedwith 14, 15) from Picdu Midi and MtCanigou respect- those of Povolny (1987, figs 1-13), which are ively,flankedbyS. arcuatasp. n. (fig. 13)from Pic based on conventional preparations ofthe 'entire', du Midi and S. angustispina sp. n. (fig. 16) from strongly three-dimensional genital armature. Mt Canigou. It may have further misled Povolny As a result of a literature survey over 70 thatapparentlyhe hadtransposed thegenitaliaof referencesto nominal taxaofSattleriawere traced; his specimens from Mt Canigou; the genitalia in however, most of them arc faunistic records or text-fig. 12belongtothemothincolour-fig. 16and entries in catalogues and check-lists of little those in text-fig. 13 to the moth in colour-fig. 15, information value. Only a limited number of not the other way round as the accompanying papers contain worthwhile biological observa- label data would suggest. tions (e.g., Burmann, 1954; 1977) or discuss Due to lack of material from several under- taxonomic aspects (e.g. Povolny, 1965; 1967; collectedareasinthewestern Alpsandmountains 1987) and are recorded in the systematic part of inYugoslaviaandBulgariatherestillremainsome our work. unresolvedproblems. Nevertheless,we believeto In the past, collecting of Sattleria was mostly have shown in our study that Sattleria comprises sporadic and only one lepidopterist ever made a several clear, morphologically distinguishable special study (Burmann, 1954) in the course of species. which heaccumulatedappreciable seriesofspeci- mens. Many of the mountains where Sattleria populations occur were rarely visited by microle- pidopterists and minimal materialexistsin collec- MATERIAL AND METHODS tions; for example, only one male each is known fromKorab(Albania)andDurmitor(Montenegro) and very few specimens, all males, were ever Our study of Sattleria is based on over 400 collected in the Julijske Alpe (Slovenija) and specimens (6:9 ratio 6: 1), about 200 of which Abruzzo (central Italy). Even in the much better originated from various museums and private exploredAlpscollectingwaspreviouslyextremely collections whilst a similar number was collected difficult in the higher regions. Many suitable by us. All specimens recorded under 'Material habitatscouldonly be reached afterlongstrenuous BMNH examined' are in unlessstatedotherwise. hikesand, astheyusually lackedovernightshelter, Theprimarytypesofthreeofthesevenpreviously fieldworkwaslimitedtojustafewdaytimehours, described nominal t«xa were examined, together often further curtailed by the notoriously un- with topotypical specimensoftheotherfour, and predictable weather conditions. Thus, like many 208 LINDAM. PITKIN&K.SATTLER otherMicrolepidopteraofthealpinezone,Sattleria 1954: 349; Klimesch, 1961: 650). The standard specimens are comparatively rare in collections, technique of running a mercury vapour light in particularly the brachypterous females, and front ofa white sheet can therefore be employed Povolny"s claim to have examined 'ample series' butappearstoproduceonlymodestresults,dueto A ('reichliche Serien') or 'extensive material' the limited motility of the moths. night search ('ausgedehntes Material') from 'all European betweenaboutmidnightand01.00hrswiththeaid mountain ranges, where [Sattleria] occurs' ('aus of a Petromax paraffin lantern was found to be alien europaischen Gebirgen, wo sie vorkommt') much more fruitful. A lantern placed on the (Povolny, 1965:492; 1987:85)isclearlyexaggerated. ground during that time in a suitable habitat will In recenttimescollectingandfieldstudies have attractthemalesinthenearvicinity.Theapproach- becomeverymucheasier,because morelocalities ingmothscan then be nettedorcollecteddirectly arenowfreelyaccessible by roadorcable carand into glass tubes, and every ten minutes or so the there are more facilities forspending the night in lantern is moved to a new location. In spite of a or near suitable habitats. We carried out field specific search no females were located with that work in the central and eastern Pyrenees (K. S., method and few other Microlepidoptera were 1981), southern Carpathians (L. M. P. & K. S., attracted at that time. 1984) and Lechtaler Alpen (K. S., 1987) whilst On occasions when a night search proved im- unsuccessful searches for Sattleria were made in practical or it was intended to sample different theeasternAlps(Raxalpe;K.S., 1985)andsouth- spots simultaneously, a Common trap was eastern Alps (Alpi Giulie/Julijske Alpe: Canin, employed. This trap is equipped with a 6 watt Montasio, Mangrt; L. M. P. & K. S., 1987). actinic tube operated by a 12 volt car battery and Duringtheday, Sattleriamalesareoccasionally chargedwith tetrachloroethane asakillingagent. flushed out of their hiding places, particularly in It is specifically designed to collect the larger earlymorningandlateafternoon,andcanthenbe and small moths in separate compartments and netted. Disturbed males fly only short distances, thus prevent damage to the Microlepidoptera unlessthey arecarriedawayby strongwinds, and (Common, 1986). Only the males were caught in as they stay close to the ground they are some- this trap, and inspections during its operation times difficult to see against the background of indicated that they were not usually attracted rocks and vegetation. before midnight. In suitable habitats with sparse vegetation and Rearingfromtheegghasnotyetbeentriedand many fiat stones a search of the ground can be may be difficult if larvae overwinter twice as mostproductive. Stonesinthe immediatevicinity suggested by Burmann (1954: 350), but a search of plant cushions can be turned over to expose for larvae and pupae was found to be successful. specimens of both sexes, either sitting on the ground or on the underside ofsuch stones. As for other insects hidden in inaccessible substratessuch asdeepscreeordensevegetation, a beesmoker can be used successfully to drive ABBREVIATIONS Sattleria males and females out of hiding. This wmhetehnotdheissmpoarkteicluilnagrelrys eofnfetchteivgeroounndd.aSmmpokdeayiss BMNH British Museum (Natural History), London, usually produced from cartridges made ofcorru- BURM EBnugrlmaannd.n collection, Innsbruck, Austria. gatedcardboard; however, locally available com- ETH Eidgenossische Technische Hochschule, bustiblessuchasdryplantmaterialorsheepdung Zurich. Switzerland. can be substituted. During field work in the HUEM Huemercollection, Innsbruck, Austria. central Pyrenees on Pic du Midi de Bigorre a MCSN Museo Civico di Storia Naturale, Milan, locally common lichen proved a most effective, Italy. longlastingfuelondampdayswhenithadthefeel MINGA MuzeuldeIstorieNaturala'GrigoreAntipa', ofgood pipe tobacco, whilst on dry days it dried MM Bucharest, Rumania. oaudtvaqnuticakgleyo,usbetcooimmipnrgegbnriattteleoannedeunndsuoiftatbhlee.caIrtdi-s MNHN MMoursaevusmkeNaMtuiosneaulmd,'HBisrtnooi,reCzNaetcuhroesllloev,aPkairai.s, France. boardcartridgewithpotassium nitrate forgreater NM NaturhistorischesMuseum, Vienna, Austria. ease oflightinginwindyweather. Careshouldbe NMB NaturhistorischesMuseum, Basle,Switzerland. taken not to cause fires when emptying the TM Termeszettudomanyi Muzeum, Budapest, beesmoker; in dry habitats it may be safer to Hungary. restrict its use to damp days. WHIT Whitebreadcollection, Magden, Switzerland. BothsexesofSattleriarespondtolight(Burmann, ZI Zoological Institute, Leningrad. U.S.S.R. SATTLERIA AEUROPEANGENUSOFBRACHYPTEROUSALPINEMOTHS 209 \ AlthoughitisuniversallyacceptedthatSattleria ACKNOWLEDGEMENTS belongstotheGelechiinae, tribeGnorimoschemini, thishasnotyetbeendemonstratedbeyonddoubt. The main synapomorphy of the subfamily Gele- Wearemostgratefultothefollowingspecialistsforloan chiinae is the division of the male abdominal ofmaterialandforhelpful information: Dr K. Burmann, segment 8 along the pleural line, which enables Innsbruck; DrRGacdike, Deutsches Entomologisches tergite and sternite to operate as independent Institut, Ebcrswalde,Germany; DrP. Hucmer,Tiroler coversforthegenitalia.Thisarrangementseparates Landesmuscum Ferdinandeum, Innsbruck; Mr O. Karsholt, Zoologisk Museum, Copenhagen; Dr J. the Gelechiinae from other subfamilies in which Klimesch, Linz; the late Dr F. Kasy, and Dr M. Lodl, segment 8 forms a continuous ring into which the NM; DrC. Leonardi, MCSN; Dr G. Luquet, MNHN; genitalia are usually withdrawn. In the Sattleria DrA. Lvovsky, ZI; DrA. Popescu-Gorj, MINGA; Dr male the structure of segment 8 (Figs 48-50) J. Razowski, Polish Academy of Sciences, Krakow; clearly shows the gelechiine condition. Prof. W. Sauter, ETH; Frau Dr I. Schatz, Innsbruck; The Gelechiinae are currently divided into the Prof. Sir Richard Southwood FRS, Department of tribesGelechiini,TeleiodiniandGnorimoschemini. Zoology,OxfordUniversity;MrS.Whitebread,Magden. Whilstthereappearstobeagreementthatmostof We also thank our colleagues, for helpful advice and the 40 genera united in the Gnorimoschemini criticism during the preparation of this paper. Our form a monophyletic group, this has yet to be fpioerltdewdobrykRionyRaulmSaonciieaty(1s9c8i4e)ntaifnidcAeuxscthrainage(1p98r7o)gwraamsmseusp.- establishedanditisheresuggestedthatthethorn- The photographs were taken by the Photographic likesignumbursaeofthefemaleisasynapomorphy Unit, BMNH. of this tribe. Regrettably, in the Sattleria female the signum is secondarily lost and, until further synapomorphies arc found, its gnorimoschemine statuscan onlybe deduced fromoverallsimilarities THE SYSTEMATIC POSITION OF in conjunction with a specialized anellus character SATTLERIA (seebelow)whichSattleriashareswithasubgroup of undisputed Gnorimoschemini. No classification of the Gnorimoschemini has S. dzieduszyckii (Nowicki), the typc-specics of ever been proposed, but an attempt was made by Sattleria Povolny, has had a chequered history Povolny (1967) toexpress hisviewson the phylo- with regard to its generic position. Originally geneticrelationshipsofthegenerawithinthetribe described in 'Gelechia (AnacampsisHS.)\ where it in a diagram, although it was not in the form of wasplaced nexttoAcompsiamaculosella(Hcrrich- a cladogram. A recent idiosyncratic numerical Schaffer) (Gelechiidae: Dichomerinac), it was analysis of phylogcnctic relationships in Gnori- subsequently transferred to Doryphora Heine- moschemini by Povolny & Sustek (1988) is less mann (= Xystophora Wocke) and within that helpful because it does not provide clear results genusassociatedwithspecieswhicharcnowaccom- and merely demonstrates the unsuitability of the modated in Monochroa Hcinemann (Gelechiidae: chosentechniqueforresolvingsuch relationships. Aristoteliinae) (Wocke, 1871: 298). Rebel (1901: In Povolny'sdiagramof 1967theGnorimoschemini 145) placed dzieduszyckii in Gelechia (subgenus are divided intothe gnorimoschemoid, scrobipal- Gelechia) amongst a group ofspecieswhich were puloid and scrobipalpoid branches, the last of later separated as Teleiopsis Sattler, a position in which issubdivided into the scrobipalpoid, ephy- which it was also retained by Mcyrick (1925: 77). steridandcaryocolidgroupsofgenera.Theplace- Following a revision ofthe collective genus Gele- ment of Sattleria in the scrobipalpoid group next chia, dzieduszyckii was removed to the Gnori- toScrobipalpopsisPovolnyandScrobipalpaJanse moschema group (Gelechiinae: Gnorimoschemini) wasbasedprimarilyon similaritiesofthe Sattleria (Sattler, 1960: 69), where a separate genus, fore wing pattern with that of Scrobipalpopsis Sattleria Povolny, was eventually proposed for petasitis(Pfaffenzeller) (Povolny, 1965: 490, text- it. fig.; 1967: 174); however, the mere presence of Sattleriaisestablishedasmonophyleticbyseveral the three typical stigmata in both genera is autapomorphies: inthe malegenitaliathe presence irrelevant for establishing phylogenetic relation- of the usually forked posterior processes of the ships, because it is a groundplan character of at vinculum, andtheanellusintheshapeofapairof least the ditrysian Lepidoptera, and no syna- perforated disks; in the female genitalia the pre- pomorphies were found to confirm Povolny's sence of characteristic sternal pockets. A likely view. further autapomorphy is the reduction of the In contrast to Povolny it is here believed with wings in the female sex. Huemer(1988:445)thatSattleriaiscloselyrelated 210 LINDAM. PITKIN&K.SATTLER to the so-called caryocolid genera Agonochaetia SATTLERIA Povolny Povolny, Pogochaetia Staudinger, Tila Povolny, Lutilabria Povolny, Klimeschiopsis Povolny, Sattleria Povolny, 1965: 490. Type-species: Caryocolum Gregor & Povolny and Cosmardia GelechiadzieduszyckiiNowicki, 1864,byoriginal Povolny, which are characterized morphologically designation and monotypy. by a reduction ofthe gnathos hook and develop- 6 6.5-11.5 mm. Frons evenly convex. Ocellus ment ofsclerotized anellusstructures in the male . present but small. Proboscis well developed, genitalia andbiologically bythe exclusive utiliza- longer than labial palpus, squamose at base. tion of Caryophyllaceae as host-plants of the Maxillarypalpuswith foursegments, foldedover larvae (host-plants ofAgonochaetia and Lutilabria base of proboscis. Labial palpus recurved, seg- unknown). A hooked gnathos is probably a groundplan ment 3 acute, about as long as 2. Antenna two- thirdstothree-quarterslengthofforewing, scape characterofthe Gelechiidae andiswidespreadin withoutpecten. Forewing broadestin distal half, the Gnorimoschemini; its reduction in the caryo- length about 4.5-5 times greatest width; costa colid genera is here considered a synapomorphy straight to weakly concave, weak indication of of that group, and the well developed gnathos tornus. Coloration various shades of grey, with hook ofSattleria is a plesiomorphic character. In diffuse light transverse line extended between Agonochaetia (Fig. 65), Pogochaetia, Tila and distal fifth of costa and tornus, stigmata black: Caryocolum(Fig. 64)theanellusisequippedwith short plical dash sometimes extended to base, a pair of needle-like or peg-like sclerotizations, a structure unknown in other Gelechiidae. In discal often small but distinct, discocellular sagit- tate; usuallycomplete rowofmarginalspotsfrom the first three genera they are striking needles distal fifthofcostaaroundapex andalongtermen of considerable size, sometimes exceeding the totornus. Costaofforewing(Fig. 2)withvariably aedeagus in length (Povolny, 1965: figs 8, 10, developedweak pterostigmabetween Sc andR5. 11; 1974: figs 1-3; Sattler, 1968: fig. 11), whilst Discal cell about three-fifths to two-thirds length ilnikeCaarnydomcaolyubme,arifaptreersmeinnta,ltsheetya(aHreuesmheorrt,,1p9e88g:- opafrawlilnegl,tdoistceorcmeellnu;laRrlvefirnomoblmiiqdudel,e,moRr2e—oRr4l+es5s figs 102, 123). The anellus of Sattleria, a pair freefromcell,freeendsofR4andR5one-quarter ofsclerotized disks with perforated centres (Fig. to one-third length ofcommon stalk; Ml close to 66), an autapomorphy of this genus, is here or connate with R4+5; distance at base M1-M2 interpretedasanextremereductionofsuchstruc- usually greater than M2-M3; distance CuAl- ture and thus a synapomorphy with caryocolid CuA2 variable, halfto twice distance M3-CuAl. genera. Hindwing(Fig. 2) aboutfour-fifthslengthoffore Our view that Sattleria is closely related to wing, broad, costa almost straight, termen more caryocolid Gnorimoschemini is also supported or less straight, very weakly concave beneath bythe host-plants ofthe larvae. Although, based apex. Sc+Rl to distal third of costa, Rl distinct on observations by Burmann (1954), it was between RsandSc; RsandMl separate, distance assumed that Sattleria larvae were Saxifragaceae at base M1-M2 about twice M2-M3; M3 and feeders, there is now increasing evidence that CuAl connateoronshortstalk;CuA2fromabout they live primarily on Caryophyllaceae, a plant distal third of cell. 9. 5.0-8.5 mm. As 6 but family rarely utilized by Gelechiidae other than the caryocolid Gnorimoschemini. As far as can antenna little shorter than fore wing. Fore wing (Fig. 3) broadly lanceolate, costa distinctly con- be ascertained from published and unpublished vex, no tornus, fringe very short. Coloration and sources only five other gelechiid species can markings similar to those of 8 but discocellular beassociatedwithCaryophyllaceae: Eulamprotes stigma rarely sagittate, marginal spots always wilkella (L.) on Cerastium; Bryotrophafigulella absent. All veins present and tubular as in 8 but (Staudinger) and B. tachyptilella (Rebel) pos- muchclosertogether. Hindwing(Fig. 3) narrow, sibly feeding on Silene; Teleiodes myricariella lanceolate, only about one-third length of fore (Frey) normally feeding on Tamaricaceae but wing,veinsstronglyreduced,nottubular;frenulum once reared on Silene (Huemer, pers. comm.); triple. and Scrobipalpa salinella (Zeller) on Cheno- podiaceae but occasionally found feeding on Genitalia 6 (Figs 27, 48-50). Tergite and Spergularia. sternite8separate,tergitenarrowerthansternite, No study has been made to resolve the phylo- posteriorly rounded; sternite broad, posteriorly geny within the caryocolid genera and it is not rounded orwith V-shaped median emargination; possibleatthisstagetoidentifythesister-groupof coremata absent. Uncus narrowerthan tegumen, Sattleria. distallymore orless rounded. Gnathoswithlarge SATTLERIA AEUROPEANGENUSOFBRACHYPTEROUSALPINEMOTHS 211 ; Figs2, 3 WingvenationofSattleriamelaleucella. 2, 6. 3, $ (differentscale). spiculate culcitula and strong, sharply pointed than tegumen, slender, straight; coecum hardly hook. Anterior margin of tegumen with broad inflated, base sometimes splayed; ventral surface emargination,pedunculilong. Valvausuallyslen- sometimes with median projection; apical arm der, digitate; sacculus clearly separated, always with sclerotized hook, usually at right angles to shorter, shape variable. Posterior margin ofvin- longitudinal axis ofaedeagus; ductus ejaculatorius culum with pair of characteristic, frequently enters dorsally near base. forked, strongly sclerotized processes; saccus of moderate width, with more or less parallel Genitalia 9 (Fig. 28). Papillaanalislongerthan margins. Anelluswithpairofcentrallyperforated wide, unspecialised; apophysis posterior about sclerotized disks. Aedeagus as long as or longer three-quarters to four-fifths length of abdomen; 212 LINDAM PITKIN&K SATTLER . . telescopic membranous part of ovipositor appa- observedinthedensegrassofalpine pasturesand rentlywithoutdorsalinvaginationorsac. Segment amongstlowgrowingRhododendron. Thefreshly 8 dorsally membranous, sclerotizations confined emergedmoths,withtheirwingsstill undeveloped, to ventral and ventrolateral surface. Sternite 8 have been observed in the early hours of the withpairofcharacteristicpockets, sternalsurface morning (Burmann, 1954: 348). Later in the day mediallyoftenwithconspicuousirregularfoldsor theyusuallyhidedeeperin scree, rockcrevicesor longitudinal ridges. Apophysis anterior rod-like, vegetation. Intheearlymorningand atdusk, less about one-third length of apophysis posterior; frequently in the daytime, males are sometimes sclerotized area at base extended into ostium disturbed from their hiding places and fly short bursae, sometimes fused with posterior end of distancesbefore settlingagain in ashelteredspot. antrum; narrow, strongly sclerotized band from Theyarerelativelypoorfliersbutcanrunwelland base to sternal pockets. Long tubular sclerotized on rare occasions make shortjumps. The brachy- antrum,moreorlesslengthofapophysisanterior. pterous females are incapable of flying but can Membranous ductus bursae usually shorter than alsorun and,when disturbed,jumpshortdistances antrum, corpus bursae spherical or oval, without ofup to 50 mm. signum. According to our observations in the Pyrenees Ductusseminalis arisingdorsallyfrom anterior (S.pyrenaica,S. arcuata)andsouthernCarpathians endofsclerotizedantrum, short,withoutpseudo- (5. dzieduszyckii) the males have a period of bursaorbullaseminalis. Vestibulumwithpapilla. particular activity between about midnight and Canalis spiralis short, about length of canalis 01.00hrs. Itseemslogicaltoassumethatthisflight receptaculi;vesiculaofmoderatesize;spermatheca is in search of the females and that the latter (receptaculum seminis) with small lagena, utri- should at the same time move from their hiding culus about three to four times length of lagena, placesto more exposedsites in ordertoawaitthe glandula receptaculi very long, filiform. Accessory males; however, no females were noticed by us glands with short ductus sebaceus, its half nearest duringnocturnalsearches.Thecourtshipbehaviour junctionofsaccisebaceianinflatedreservoir;pair has never been observed, but a mating pair was ofnarrowtubularsaccisebaceiterminatinginlong found in the early afternoon under a stone, with filiformglandulaesebaceae. Oviductuscommunis the male halfeaten by a spider. Burmann (1954: inferiormergedwithanaltubeshortlybeforeanal 349) suspected that a female he had seen walking opening to form cloaca. [Description of ductus from oneplantto anotherin the afternoonwasin seminalis - cloaca based on an examination of a the process ofdepositing her eggs. single female ofS. melaleucella.] The flight period of Sattleria can vary greatly from one year to the next and probably also Remarks. There isconsiderable individual varia- between different localities, depending to a con- tion in some genital characters of Sattleria, siderable extent on local conditions such as alti- notablythesaccusandthebaseoftheaedeagusin tude, exposure to the elements, and the weather themale, andanunusuallyhighlevelofdeformity conditionsinagivenyear. Adultsexaminedbyus was observed in the relatively small number of werecollectedorbredbetweenlateJuneandmid- females examined. For example, two females of September;however,themainflightperiodappears melaleucella have a strongly reduced apophysis to be mid-July to mid-August, unless these dates anterior whilst the genitalia of one female of merely reflect the most populartime forentomo- aAngbuasrteilsypidniascaerrenisbtlreonfgelaytuarseylmemssetvreisctaiglia(lFisgi.g5n5u)m. lEoxgciecpatliovinsailtlsyt,ostohmeehriegahreerdEaudrulotpsedaindmnootunetmaeirngse. was found in two individuals of pyrenaica; its until late September-early October. presenceisanindicationthatthelossofthisorgan The ovum, which is probably laid singly or in in Sattleria is secondary but it is too strongly smallclustersonpartsofthehost-plant,hasnever reducedtodrawinferencesastoitsoriginalshape. been observed and the duration of the egg stage Biology. Our knowledge of the biology of andthenumberoflarvalinstarsareunknown. Itis Sattleria is still sketchy, the only significant pub- alsounknownwhetherthefirstinstarlarvamines, lishedobservationsbeingthosemadebyBurmann like the early instars of many other Gelechiidae (1954) in the Austrian Lechtaler Alpen. Sattleria includingsome Caryocolum species. Larvae have species are restricted to the alpine zone of the been found in July-August, living singly in a European mountains where they can be found loosely spun silken tube that is often extended between 1500 m and 3500 m, although they fromtheupperpartsofthe host-planttotheroots usuallyoccurupwardsof2000m. Typical habitats or nearby stones, where the larva seeks shelter are open slopes with coarse scree and sparse when disturbed. The frass is deposited in a small vegetation, but several species have also been heapoutsidethelarvaltube. Pupationtakesplace

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