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Arch. Molluskenkunde | 135 | (2) | 205–223 | 8figs,2pls | FrankfurtamMain,18.12.2006

©E.Schweizerbart’scheVerlagsbuchhandlung(Nägeleu.Obermiller),2006,ISSN0003-9284

Revision of the Ariantinae, 3. Superba n. gen., with the description of three new species

(Gastropoda: Pulmonata: Helicidae)

Péter Subai & Zoltán Fehér

Abstract

Inthepresentwork,thegenusSuperban.gen.,whichbelongstothesubfamilyAriantinaeoftheHelicidae, isdescribed.Threenewtaxa,S. skipetarica asketa n.ssp.,S. grisean.sp.andS. kulmakanan.sp.areconfinedtothenewgenus.Recordsforalltaxaarelistedandmapped.ThenewgenusiscomparedtootherrelatedgeneraofthesubfamilyAriantinae,suchasCattania, Jose-phinella, andLiburnica.

Keywords:Ariantinae,Superba, taxonomy,anatomy,distribution,newgenus,newspecies,Albania,Greece.

Kurzfassung

IndieserArbeitwirddieGattungSuperban.gen. innerhalbderAriantinaebeschrieben. IndieserGattungwerdendreiTaxaneubeschrieben:S. skipetarica asketa n.ssp.,S. grisean. sp.und S. kulmakanan.sp.NebenBeschreibungenundVerbreitungderArtenwerdendieUntersc-heidungsmerkmalegegenüberdenbenachbartenGattungenderAriantinae,Cattania, Josephinella und Liburnicadiskutiert.

Schlüsselwörter:Ariantinae,Superba,Taxonomie,Anatomie,Verbreitung,neueGattung,neueArten,Albanien,Griechenland.

Introduction

DuringAugust 2004andAugust 2006, theTomorMountains insouthernAlbaniawerevisitedbyexpedi-tionsconducted in the frameworkof theScientificRe-searchProgrammeon theBalkanareaonbehalfof theHungarianNaturalHistoryMuseum(Fehéretal2004),inordertocollectplantandanimalspecimens1.Amongothers,livingspecimensofthreehithertounknowntaxa

of theAriantinaecouldbe found.The investigationofthegenitalorgans revealed that theyconstituteagroupof ratherclosely relatedspecieswhichhas tobesepa-ratedfromothergroupsofAriantinaeongenericlevel.

Sinceseveralyears,theseniorauthorofthispublica-tion investigates the taxonomyof thecentral-andeastEuropeanAriantinae.Asa result it is foreseeable that

1ThisresearchwaspartlysupportedbytheHungarianNationalR&DProgramme,title:TheoriginandgenesisofthefaunaoftheCarpathianBasin:diversity,biogreographicalhotspotsandnatureconservationsignificance;contractno:3B023-04.

Authors’addresses:Péter Subai,Kronenberg143,D-52074,Aachen,email:subai@bio1.rwth-aachen.de.Dr. Zoltán Fehér, HungarianNaturalHistoryMuseum,Barossutca13,H-1088Budapest,email:feher@nhmus.hu.

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206 Subai, P. & Fehér, Z.:RevisionoftheAriantinae,3.Superba nov.gen.

mostofthegroupshithertorankedassubgeneraofHeli-cigona A. FéruSSac1821have tobe raised togenericlevel.Thesubdivisionparticularlyfocusesonthecharac-terstatesofthepenialpapillaandtheatrialstimulator,asmallpadoftissuewhichisusuallylocatedintheatriallumenandwhichturnedouttoprobablybecharacteris-ticongenericlevel.Thisorganwasforexamplecalled“crest-likestructure(cls)”byGiuStietal.(1997).

Asafirststep,thenewgenusSuperbaisdescribedforthesmallgroupofspeciesof theAriantinae inhabitingSouthAlbaniaand theneighbouringareasofnorthernGreece.

Abbreviations

F privatecollectionW.Fauer(nowintheZoologicalMuseumoftheUniversityofHamburg)

H privatecollectionA.hunyadi,BudapestHNHM HungarianNaturalHistoryMuseum,BudapestIZPAN InstytutZoologiczny,PolskaAkademiaNauk,

WarszawaNEUB privatecollectione. neubert,BadenweilerNMW NaturalHistoryMuseum,Vienna

NNM NationaalNatuurhistorischMuseum,LeidenREI privatecollectionP.L.reiSchütZ,HornS privatecollectionP. Subai,AachenSMF NaturmuseumSenckenberg,Frankfurta.M.STU privatecollectionStummer(nowinthe

InstituteofPalaeontologyoftheUniversityofVienna)

ZMUA ZoologicalMuseumoftheUniversityofAthens

Additionalabbreviations

± moreorlessAD diameterofapertureAH heightoftheaperturealk. specimenpreservedinalcoholAST atrialstimulatorD shelldiameterH heightoftheshelljuv. juvenilespecimenUTM UTM-CodefortheinventoryoftheEuropean

Invertebrates

Allmeasurementsaregiveninmm.

Systematicaccount

Superban.gen.

Ty p e s p e c i e s :Helicigona skipetarica Subai 1995.

D i a g n o s i s : Shells small tomedium sized, lowconicaltoalmostflat.Thecolourpatternbasicallycon-sistsofthreedeepbrownspirals(insometaxareducedtoalmost lacking).Theshell surfacegenerallywithouthairs,usuallysmoothandshining.Insomespecies, theradialstripesformthickribs.Theapertureisroundedtoobliquelyoval, theapertural fringesareclose together.Theumbilicus is alwaysopen, funnel-shaped,moder-atelytoverywide.

Thebodyisgreytogreyish-brown.Mantlewithgrey,irregularlyfusingspotsofpigment.Thesecondaryureteriscompletelyopen.Thegenitalorganswithsimpleorsubdividedglandulae(insomespecimens,bothtypesofsubdivisionmayoccur).Thestemofthebursacopulatrixisveryshort.Diverticulumusuallyaslongasvesicularstem+vesicle.

ThemainpartoftheASTissituatedcentrallyintheatriumandformsanelevatedtriangularpadoftissue.Itsometimesshowsashortpilasterpointing towards the

peniallumen,andrarelyashortsecondpilasterpointingtowards thevaginal lumen.Thepenialpapilla is shortandconical,with adeep lateral furrow, its surface isstructuredbyfineobliquefolds.Theporeofthepapillaissituatedbasicallyinthelateralperforation.

Thedartisshortandclumsy,itsstemisthin.dieKro-neundderSpitzenteilsinddoppeltsobreitwiejene.Thetipofthedartreaches40%oftheoveralllengthofthedart2.

D i f f e r e n t i a l d i a g n o s i s :SuperbadiffersfromthegenusCattania bruSina1904byisgenerallysmallerandlessconicalshellsandthemuchlowerwhorls.TheASTofSuperba ismuchsmallerandflatter.ThepenialpapillaofCattania iscontractedin themiddle(notcon-tractedinSuperba).Theporeofthepenialpapillaissitu-atedonthetipofthepapillainCattania,whileinthedeeplateral furrow inSuperba.Thenewgenusdiffers fromLiburnica Kobelt1904andJosephinella F.haaS1934byitshairlesssurfaceof theshelland the low,almostflatwhorls.Additionally,shellsofspeciesofLiburnicadis-playacharacteristicwhitecallusordenticleatthebasisoftheaperture.InspeciesofJosephinella,thecolouroftheshellisgreenishtobrownish,whileinSuperbaitiswhit-

2 ThebothspeciesfromGreecehavedartswhicharelonger,moreslenderandwhichhavealsorelativelysmallerandmoreslenderapicalparts.

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ishtogreyish;theshellshavealwaysasingledarkspiralin themediumposition,while inSuperba, threecolourspiralsarepresent.InspeciesofJosephinella,theASTislargewithtwozigzaglikeprocesses,butinSuperba,theASTisflatterwithreducedprocesses.

D i s t r i b u t i o n p a t t e r n s :SpeciesofCattania inhabit anareanorthandeastof theareaofSuperba;towards north it reaches theBanat in SWRomania,andeastwardsandsouthwardstotheBlackSeaandtheAegeanSea.LiburnicaisfoundalongtheDalmatianandAlbaniancoastandreachesNWGreece.SpeciesofJose-phinellaliveinanareafromCentralAlbaniatoWestandCentralGreeceandreachthesouthofthePeloponnesus.SeveralspeciesofJosephinellalivesympatricwithspe-ciesofSuperba.

E t ym o l o g y : fromLatin“superbus, -a, -um=ex-cellent“forthewonderfulappearanceoftheshells;gen-derfemale.

Superba grisean.sp.Plate1,fig.1;textfigs1,7–8

Ty p e s a n d l o c u s t y p i c u s :HolotypeHNHM95542.ParatypesHNHM95468/4+1 (preserved),SMF328953/1,S20211/1 (partlydamaged);Albania,PeriferiSkrapar,QafaeDëvris,NEofRadeshalong theÇorovodë-Zaloshnjë road,eastsideofthegorge,limestonerocks,1150malt,UTMDK38, 8.8.2004, leg.Fehér.—Additional paratypes:HNHM95978/14+7(alk.)+3(juv.,alk),H/21,22.8.2006,leg.Fehér, hunyadi, huSZár & murányi.

A d d i t i o n a l m a t e r i a l (fromthetypelocality,butnotypematerial):HNHM95469/17(juv.+fragments),leg.Fehér, 8.8.2004;HNHM95979/8,H/8(juv.+fragments),22.8.2006,leg.Fehér, hunyadi, huSZár & murányi.

D i a g n o s i s :Shellmediumsized,basicshellcolourgrey,withextraordinarilynarrowspiralbands, ribletsblunt,diverticulumshorterthanthestemofbursacopu-latrix,flagellumlong.

D e s c r i p t i o n :Theshellofmediumsize,depressedto slightly conical with 4½–5regularly increasingwhorls; shell colourgrey to slightlybrownish, in theupper thirdof thewhorlswith twonarrowbrownspi-rals (width0.5–0.8mm); the thirdspiralalwayson theperipheryoftheshellandofthesamewidthastheothertwo;usually,theinitial2–2.5whorlsarebrownish.

Theprotoconchof1.5–1.75whorls,whicharesmoothinthebeginning;after1.5whorls,theprotoconchsurfaceissculpturedbyfineradiallyarrangedstripesconsistingofminutegranules.Teleoconchsculpturedby low, ir-regularlyarranged riblets,whichbecome increasinglycoarserandmorebluntonthelastwhorlsoftheshell.Onthelastwhorl,minutespiralthreadscanbefound,whichsometimesmaybearrangedinshortobliqueareas(highmagnificationrequired).

The lastwhorl reaches1.7–2thesizeof thepenulti-matewhorl,obliquelydescendingbefore theaperture.Sutureofmediumdepthandslightlyundulating.Umbili-

cusofmediumwidth,perspective;itincreasesregularlyfromtheinitialwhorl,andincreasesrapidlyinsizeinthelastwhorlreachingthedoublewidthofthepenultimatewhorl(ca.3.4–4.8mm).Aperturespherical tooval, theinsertionswithadistanceof2.8–4.2mmandconnectedbyathinbutwellvisiblecallus;aperturalrimsharp,lat-erallyandbasallysomewhatextended(0.6–1mm).

S h e l l m e a s u r em e n t s (n=7):H:10.1–11.3;D:22–23.3;AH:9.2–10.7;AD:11.1–12.

Theanimal’shead (after sixadult andone juvenilespecimen)andthedorsumaregreyishtobrownish;lateralflanksandtailusuallybrighter;footwhitishwithagreyfringe;mantlewithweakandirregularlyplacedspotsofpigment;thereisnoregularpatternofspots.Thesecond-aryureter reachesa lengthof20–27mm, iscompletelyopen;at itsendit issubdividedintwoseparateductsof1.5–2and3–4mmlength,respectively.Themandible ismediumtochestnutbrownandhas5–6narrowridges.

C h a r a c t e r s o f g e n i t a l o r g a n s :femalepart:glandulaeelongate,somewhatnarrowatthetip,reachingfive tosix times the lengthof thedartsac;usually, theglandulaesplitat40%of theirabsolute length,but intwospecimens,acombinationofasubdividedandasim-pleglandulacouldbeobserved.Onespecimenhad twosimpleglandulae.Stemofbursacopulatrixshort,reach-ing20–25%ofthevesicularstem;diverticulumreaches2/3ofvesicularstem+vesicle.Vaginaaslongasorevenshorterasthedartsac(whichisquiteshortifcomparedtotheotherspecies)withseveralbroadtransversepilasters.Dartshort,somewhatclumsy,2.5–3mmlong,withanar-rowstem,slightlycurved;crownfunnel-shaped.

TheASTstartsatthegenitalporewithathickpilaster,bendsperpendicularlyinthemiddleofthegenitalatriumforminga lowpyramidalpadwithflattenedflanksandconnectstoasmallpilasterpointingtowardsthepeniallumen. Insomespecimens, thispilaster formsabroadlongitudinalfold.

Malepart:flagellumreachesalmosttwicethelengthofpenis+epiphallus; it is slightlycoiledorundulatedinitsbasalpart;peniallumenwithseveralbroadandflatlongitudinalpilastersreachinginthedistalpartofthepe-nis;inthesurroundingofthepenialpapillawithnarrowtransversepilasters;penialpapilla± longwithslightlyoblique transverse foldson its surface; theporeof thepapilla insideadeep longitudinal furrowwhich issitu-atedlaterallyonthepapillaandreachesfromthebaseofthepapillatoalmostthetip.

E t ym o l o g y : fromLatin“griseus, -a, -um=grey”describingthegreyishbasiccolouroftheshell.

D i f f e r e n t i a l d i a g n o s i s : the shell ofS. kul-makanan.sp.displaysacreamybasiccolourwithmuchbroaderspiralbands,itswhorlsareflatter;thediverticu-luminS. kulmakana isrelativelylongerifcomparedtoS. grisea,andithasalwaysdeeplysubdividedglandulae.InS. s. skipetarica Subai 1995,theshellisusuallysmaller,thebasiccolourisbrighterwithmoreintensivelydeeply

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browncolouredbands,thesurfaceisgranulated(smoothinS. grisea), and theumbilicus increases regularly insize.Theglandulae aregrey, and thediverticulum isrelativelylonger.InS. reischuetzi (Subai 1990),theshellisusuallysmallerandmoredepressed,thebasiccolouriswhitewithbroadandbrightbrownish spiralbands.TheradialribletsinS. reischuetzi aremuchfiner,andtheaperturalrimislessdilated;thediverticuluminS. reis-chuetzi hasthesamelengthasvesicularstem+vesicle.

D i s t r i b u t i o n :S. grisealivesinsouthernAlbania,inthesouthernpartoftheKulmakMts.Thisspeciesisonlyknownfromthelocustypicus.

Superba kulmakana n.sp.Plate1,fig.3;textfigs2,7–8

T y p e s a n d l o c u s t y p i c u s : Holotype HNHM95543.Paratypes:HNHM95470/45+8(alk.),SMF328954/1,S20210/2;Albania,PeriferiSkrapar,PeriferiSkrapar, smallcanyon1KmNWofMajaeKulmakut,on limestone rocks,2070malt.,UTMDK39,9.8.2004,leg.Fehér.

A d d i t i o n a l m a t e r i a l : from the locus typicus,HNHM95471/15(juv.+fragments,notypes).

D i a g n o s i s :Shellofmediumsize,spirelowconi-cal,teleoconchwiththreespiralbandsofchestnutbrown

Textfig.1.Superba grisean.sp.—a)wholebodyextractedfromtheshell;b)situsofthegenitalorgans;c)dart;d)lumenofthedistalgenitalorgansshowingAST,penialpapillaandvagina.Albania,PeriferiSkrapar,QafaeDëvris,NEofRadeshalongtheÇorovodë-Zaloshnjëroad.

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colour; surfaceof teleoconchsmooth, sculpturedwithfinegrowthlines,umbilicuswide;glandulaelong,deep-lysubdivided;diverticulumlongerthanthestemofbursacopulatrix.

D e s c r i p t i o n :The shell isdepressed to slightlydepressed conicalwith4.5–5.25regularly increasingwhorls,apexsometimesmammillate.Thebasiccolouriscreamwiththreespiralbandsofchestnutbrowncolour.Theirwidthisca.0.5–1mmandusuallyofthesamesize.In some specimens, the third spiral below the shell’speripherycan reachawidthofup to2.5mmandfadeout towards theumbilicus.Theupper2–2.5whorlsareusuallyyellowishbrown.

Theprotoconchwhorlsconsistof1.5whorlswhichare sculpturedwith finegranules.The surfaceof theteleoconch is shining,granulatedand sculpturedbyafewlowradialstripes.Onthelowerwhorls,thissculp-turepatternbecomes somewhatmore coarse.On thelastwhorl, a fewcloselyspacedspiral furrowscanbeobserved(highmagnification required).The lastwhorlreaches 1.8–2the size of the penultimatewhorl andbendsslightly tostrongly towards theaperture.Sutureofmediumdepth,slightlyindentate;umbilicusdeepandfunnel-shaped, regularly increasing inwidth,butdou-blingitssizeinthelastwhorl.Here,itreachesadiameterof4–5.5mm.Aperturespherical tooval, the insertions

Textfig.2.Superba kulmakanan.sp.—a)wholebodyextractedfromtheshell;b)situsofthegenitalorgans;c)dart;d)lumenofthedistalgenitalorgansshowingAST,penialpapillaandvagina.Albania,PeriferiSkrapar,MaliiKulmakës,1kmNWofMajaeKulmakut.

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withadistanceof1.5–3.1mmandconnectedbyathin,almostinvisiblecallus;aperturalrimsharp,laterallyandbasallyregularlyextendedreaching0.8–1.25mm,some-whatenlargedtowardstheumbilicus.

S h e l l m e a s u r em e n t s (n=48):H:9.1–14.5;D:20.3–27.3;AH:8.5–12;AD:10.2–14.7.

Theanimal’sbody (after8specimens from1kmNofMaja eKulmakut) isuniformlygreyish tobrown-ishgreyish,thetailsometimesbrighter,thesoleoffootcreamwithdarker fringes.Themantledisplaysapat-ternof irregularlyarranged,blurreddarkspots.On theupperwhorls, these spots areonly fewbut theirden-sityincreasesonthelowerwhorl.Thesecondaryureterreachesa lengthof17–25mm, iscompletelyopenandsplitsterminallyintwocanalsof1.5and2.5mmlength,respectively.Themandibleismediumtochestnutbrownandhas4–5ridges.

C h a r a c t e r s o f g e n i t a l o r g a n s :femalepart:glandulaeelongate,somewhatnarrowatthetip,reachingfivetoseventimes the lengthof thedartsac; theglan-dulaesplitalreadyat30–40%of theirabsolute length.Stemofbursacopulatrixshort,reaching20–25%ofthevesicular stem; thediverticulum isas longasoronlyslightly shorter than thecombined lengthofvesicularstem+vesicle.Thevaginareachesthesamelengthofthedartsacorisslightlyshorterandmaybenarrowedbyatransversefoldinsomespecimens.Dartshort,somewhatclumsy,2.7–3mmlong,slightlycurved;crownbroadlyfunnel-shaped,narrowinthemiddle.

TheASTwitha±broadandflatstempartandstrong-lyupraised in thecentreof theatrium.ThepenialarmoftheASTpointstowardsthepenialpapillaandreachesapproximatelyhalfof thepenial lumen; itbecomesflatatitsend.Thevaginalarmisusuallyreduced,inafewspecimens,suchapilasterentersthevaginauntilhalfofitslength.

Malepart:Thepenial lumen is filledwithnarrowpilastersarrangedinparalleltothepenialbranchoftheAST.Thepenialpapilla is short conicalwith slightlysunkentransversefolds;thetipisbluntlyrounded.Later-ally, thereisafurrowreachingfromthebasetothetipof thepapilla; thepore issituated in this furrowat thebaseof thepapilla.Theflagellumreaches1.3–1.5thesizeofpenis+epiphallus; ithasoneor two loops in itsbasalpart.

E t y m o l o g y : the name kulmakana refers to theKulmakMts.,wherethisnewspeciesisliving.

D i f f e r e n t i a l d i a g n o s i s :Superba kulmakana differs from S. griseaby itsbasicshellcolour (greyishinS. grisea, cream inS. kulmakana) and thebroaderandmoredistinct spiralbands.The teleoconchsurfaceisgranulated inS. kulmakanabut smoothandcoveredwithawhitishsurfacelayerinS. grisea.Thediverticu-luminS. grisea is relativelyshorter ifcompared toS. kulmakana.InS. s. skipetarica,thelowerspiralisoftenmuchbroader.Theumbilicusofthisspeciesisrelatively

narrower, theapertural insertionsarenot soclose, theapertural rim is lessdilated, itsglandulaearegreyishpigmentedandusuallynot subdividedoronly theup-perquarteroftheirlength.TheshellofS. reischuetziissmallerandmoredepressed,itsspiralsarelightbrown,onlytheprotoconchwhorlsaregranulated,anditsteleo-conchsurfaceissmoothbesidessomefineradialstripes.Theaperturalinsertionsareconsiderablydistanttoeachothers,theaperturalrimislessdilated,theglandulaearerelativelyshorterandcylindricaluptothetip,thedartismuchmoreslender.

D i s t r i b u t i o n :Until now,S. kulmakana isonlyknownfromitslocustypicus.

Superba reischuetzi (Subai 1990)Plate1,fig.2;textfigs3,7–8

1990 Chilostoma reischuetzi Subai,–AnnalendesNaturhis-torischenMuseums inWien,91 (B):244,Taf.1Fig.1–3 (shell),Abb.1 (genitalorgans) [Locus typicus:“Griechenland,Epirus,Felsenb.d.türkischenBrückeüberdenAóos-FlußbeiKónitsa(UTMDK73)”,Holo-type:NMW84363/1].

1990 Chilostoma reischuetzi,–reiSchütZ & Sattmann,An-nalendesNaturhistorischenMuseumsinWien,91(B):258.

T y p e s a n d l o c u s t y p i c u s : Holotype NMW84363/1.Paratypes:NMW84364/2,REI/3,STU/1,Greece,Ípiros, rocksat theTurkishbridgecrossing theAóos riveratKónitsa,UTMDK73,12.7.1987,leg.reiSchütZ & Stummer;SMF328991/1,S10726/5+2(juv.)+2(alk),NNM/1,ditto,12.4.1988,leg.riedel & Subai.

A d d i t i o n a l m a t e r i a l :ZMUA20002/3+1 (juv.),Greece, Ípiros, canyon of theAóos river, UTM DK 73,29.5.1981, leg.mylonaS;ditto,11.8.1972, leg.FauerF/4+1(juv.)(rightsideofthecanyon)and2+1(juv.)(leftsideofthecanyon);F/2+2(juv.),S11670/4+8(juv.),samelocality,600mfromtheTurkishbridge,onlimestonerocks,580malt.,UTMDK73,26.9.1989,leg.Fauer & Subai;NMW/102938/1+1(juv.),samelocality,0.5–1.5kmabovetheTurkishbridgeatKónitsa,UTMDK73,25.6.1988, leg.Sattmann;ZMUA20085/2+3 (juv.),canyonof theAóos river,MoniStómiou,UTMDK73,18.5.1982, leg.mylonaS;ZMUA20182/3,ca-nyonof theAóos river,Monastíri,UTMDK73,19.5.1982,leg.mylonaS;N/1 (fragment), rocks at theTurkishbridgecrossing theAóos riveratKónitsa,UTMDK73,18.7.1990,leg.neubert;S19162/1(juv.),mountainslope1kmfromtheroadcrossingKónitsa-Kozáni indirection toAmárantos,onlimestonerocks,UTMDK73,12.4.1988,leg.riedel & Subai.

D i a g n o s i s :Shellsmall,depressed,teleoconchwiththreespiralbandsoflighttomediumbrowncolour;sur-faceofteleoconchsmooth,umbilicusmoderatelywide;penialpapillashort,dartslenderandlong.

D e s c r i p t i o n :Theshellisdepressedconicaltoal-mostdiscoidalwith3.75–5regularlyincreasingwhorls.Thebasicshellcolouriswhitishwiththreespiralbandsoflighttomediumbrowncolour.Usually,theuppermostspiralisweak,narrowandfaint,whilethemediumspiralissharpwithawidthofca.0.8–1mm.Thethirdspiralis

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thebroadest,alwayssituatedbelowtheperipheryoftheshellandupto2mmwidth.Theupper2–2.5whorlsaresometimesyellowishbrown.

Theprotoconchwhorls consist of1.5–1.75whorlswhicharesculpturedwithminutegranules.Thesurfaceoftheteleoconchissmooth,itssculptureconsistsofir-regularly spaced radial riblets.The lastwhorl reaches1.7–1.9thesizeofthepenultimatewhorlandbendsob-liquely towards theaperture.Sutureofmediumdepth,slightly indentate;umbilicus funnel-shaped, regularlyincreasing inwidth. In the lastwhorl, itsdiameter in-creases1.5timesifcomparedtothepenultimatewhorl.Itreachesadiameterof3.5–5.5mm.Aperturetransver-sallyoval,theinsertionswithadistanceof3.5–5.5mm

andconnectedbyathinalmostinvisiblecallus;aperturalrimsharp,upperpartflattenedextended; laterallyandbasallyenlarged reaching0.8–1.25mm,somewhaten-largedtowardstheumbilicus.

S h e l l m e a s u r e m e n t s (n=29):H: 7–10;D:16–22.2;AH:7.5–9.5;AD:8.5–11.5.

Theanimal’sheadandbody(after2specimensfromthe type locality)aregrey, the tail somewhatbrighter;soleofthefootcream.Themantledisplaysapatternofirregularlyarranged,blurreddarkspotsontheentiresur-face.Ontheupperwhorls,thesespotsareonlyfewbuttheirdensityincreasesonthelowerwhorl.Thesecond-aryureterreachesalengthof15.5–17mm,iscompletelyopenandsplitsterminallyintwocanalsof1and2mm

Textfig.3.Superba reischuetzi (Subai 1990). — a)wholebodyextractedfromtheshell;b)situsofthegenitalorgans;c)dart;d)lumenofthedistalgenitalorgansshowingAST,penialpapillaandvagina.Greece,Ípiros,rocksattheTurkishbridgecrossingtheAóosriveratKónitsa.

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length,respectively.Themandibleismediumbrownandhas4ridges.

C h a r a c t e r s o f g e n i t a l o r g a n s :femalepart:glandulaeofmoderate length, reaching three to fourtimesthelengthofthedartsac;inbothspecimensinves-tigated,oneglandulawasundivided,theotheronesplitat30–40%oftheirabsolutelength.Stemofbursacopu-latrixshort,reaching15–20%ofthevesicularstem;thediverticulumisaslongasthecombinedlengthofvesicu-larstem+vesicle.Thevaginareachesthesamelengthofthedartsac;internallywithseveralnarrowlongitudinalpilasters.Dartslender,3.1–3.3mmlong,slightlycurved;crownbroadlyfunnel-shaped,narrowinthemiddle.

TheASTwitha±broadandflatstempartandform-ing a strongly upraised triangular or linguiformpadin thecentreof theatrium.Thepenialarmof theASTreachesthepenialpapilla(orisevenlonger);itbecomesflatatitsend.Thevaginalarmisshortpointingtowardstheboundarybetweenatriumandvagina.

Malepart:Thepenial lumen is filledwithnarrowpilastersarrangedinparalleltothepenialbranchoftheAST.Therearenarrowtransversefoldsintheproximalpartofthepenisaroundthepapilla.Thepenialpapillaisshortconicalwithslightlysunkentransversefolds.Lat-erally,thereisafurrowreachingfromthebasealmosttothetipofthepapilla;theporeissituatedinthisfurrowatthebaseofthepapilla.Theflagellumreachesthesizeofpenis+epiphallus;ithasuptofourloopsinitsbasalpart.

D i f f e r e n t i a l d i a g n o s i s :TheshellofS. viko-sensis ismoredepressedwith less developed colourspirals; the surfaceof its teleoconch is sculpturedbystronger radial riblets; theuppersideof theaperture ismoredepressedandtheumbilicusiswiderifcomparedtoS. reischuetzi.Therearealmostnodifferencesintheanatomyofthegenitalorgansexcepttheundividedglan-dulaeandthelargesizeoftheASTinS. vikosensis. TheshellofS. kulmakana is largerand lessdepressedwithdistinctcolourspirals,and its teleoconch isgranulated;theaperturalinsertionsarecloser,andtheaperturalrimismoredilated; theglandulaeare longer,moredeeplysubdivided and thediverticulum is relatively longer;thedartisbroader.TheshellofS. griseaislargerwithagreybasicshellcolour,theradialribletsaremorebluntbroader; thediverticulumisconspicuouslyshorter.TheshellofS. s. skipetaricaislessdepressedandhaschest-nutbrownspiralsdiffering in size;on the teleoconch,theradialribletsarestronger(formingalmostribs);theglandulaeandthediverticulumarerelativelylonger,theASTisflatterwithonlyasingle(thepenial)arm;thedartisbroaderwithalargertip.

D i s t r i b u t i o n :Asfarasknown,thisspecieslivesrestrictedto thevalleyof theAóosriver inÍpiros(NWGreece).Quite recently, itwasalsocollected from thevalleyof theSarantáporos river.Thisnew localityex-tends the rangeof this speciesabout5km innortherndirection.

Superba skipetarica skipetarica (Subai 1995)Plate2,figs4–5;textfigs4,7–8

1995 Helicigona skipetaricus Subai,–AnnalendesNaturhis-torischenMuseumsinWien,97(B):88,Abb.2(shell),Abb. 13 (genital organs),Abb. 14 (dart),Abb. 22(distribution) [Locus typicus: “Süd-Albanien, To-morgebirge, Kar N Çuka Partizan, E-Hang, Kalk-felsen,1850malt.,UTMDL20”,Holotype:NMW87770/1].

1996 Helicigona skipetaricus,–Welter-SchulteS,SchriftenzurMalakozoologie,9:22,29.

1996 Helicigona skipetaricus,–dhora & Welter-Schul-teS,SchriftenzurMalakozoologie,9:164(distributionmap),166.

1996 Helicigona skipetaricus,–dhora & Welter-SchulteS,SchriftenzurMalakozoologie,9:209.

2002 Helicigona skipetaricus,–dhora,Studimembimo-lusqeteShqipërisë:135,193.

T y p e s a n d l o c u s t y p i c u s : Holotype NMW87770/1.Paratypes:NMW87771/11+2(juv.),NMW87772/9(alk.),S14076/2;KarNÇukaPartizan,easternslope, lime-stone rocks,1850malt.,UTMDL20,16.8.1992, leg.Satt-mann & WeiSSenSteiner;NMW87768/8,ditto,2000–2100malt.,UTMDL20,16.8.1992, leg.Sattmann & WeiSSenStei-ner;NMW87769/6+11(juv.),87775/4+1(juv.alk.),KarNÇukaPartizan,limestonegravel,1730malt.,UTMDL20,15–16.8.1992, leg.Sattmann & WeiSSenSteiner;NMW87752/1,NslopeofÇukaPartizan,beechforest,1330malt.,UTMDL20,15.8.1992,leg.Sattmann & WeiSSenSteiner.

A d d i t i o n a l m a t e r i a l (populationswith transitionalcharacterstates toS. s. asketa):HNHM95456/21+4(juv.),S20214/2,Albania,PeriferiBerat,Mali iTomorrit,6KmNofMaja eTomorrit, on limestone rocks along themoutainridge,2240malt.,10.8.2004,leg.Fehér;HNHM95457/98+14(juv.),SMF328955/1,S20215/2,ditto,onlimestonerocksbelow thewesternslopeof themountain ridge,2060malt.,11.8.2004,leg.Fehér.

D i a g n o s i s :Shell small tomedium sized, basicshell colourwhitish,with threechestnutbrownspiralbands;surfaceoftheteleoconchwithfineradialstripesonly;umbilicusofmoderatewidth;genitalorganswithgreypigment.

D e s c r i p t i o n :Theshell isdepressedconicaltoal-mostflatwith4.5–5regularlyincreasingwhorls;basicshellcolourwhitish,with threechestnutbrownspiralbands;usually,theuppertwospiralsaredistinctandreachawidthofca.0.5–1mm(infewspecimens,themiddlespiralmayfade); the lowerspiral isawideas theothers,butoftenincreasesinwidthto3–9mmandfadestowardstheumbili-cus.Theupper2–2.5whorlsareusuallyyellowishbrown.

Theprotoconchwhorlsaresculpturedwithveryfinegranules,surfacedull.Thesurfaceof the teleoconch isshining,finelygranulatedandsculpturedbya few lowradial stripes (whichmaybestronger insomepopula-tions).On the lastwhorl, a fewclosely spaced spiralfurrowscanbeobserved(highmagnificationrequired).Sometimes, the apex ismammillate.The lastwhorlreaches1.6–2.1the sizeof thepenultimatewhorl and

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bendsslightly towards theaperture.Sutureofmediumdepth, slightly indentate; umbilicusdeepand funnel-shaped, reachingadiameterof3.1–4.4mm.Aperturespherical tooval, the insertionswithadistanceof2.6–4.7mmandconnectedbyathin,almostinvisiblecallus;apertural rimsharp, regularlyextended reaching0.6–1mm,inconspicuouslyenlargedtowardstheumbilicus.

S h e l l m e a s u r em e n t s (n=165):H:7.9–11;D:17–23.7;AH:7–10.3;AD:8.5–12.

Theanimal’sbody(after5specimensfromKarNÇukaPartizan,eastern slope) isuniformlybrown tobrown-ishgreyish; thesoleof footcreamwithbrightgreyishfringes.Usually, themantlewithoutpigments,but inafewspecimensthereisapatternofgreypigmentaroundthelungvenation.Thesecondaryureterreachesalength

of22–25mm,iscompletelyopenandsplitsterminallyintwocanalsof1.5and2.5–3mmlength,respectively.Themandibleisopaquetoreddishbrownandhas4–5ridges.

C h a r a c t e r s o f g e n i t a l o r g a n s : Internally,manypartsofthegenitalorganswithgreypigment.

Femalepart:glandulaeelongate,greyish, somewhatnarrowat the tip, reachingfour times the lengthof thedartsac;usually,theglandulaearenotdivided,butinafewspecimens therearesubdividedglandulae (splitat25–30%oftheirabsolutelength).Stemofbursacopu-latrixshort,reaching20–25%ofthevesicularstem;thediverticulumisaslongasoronlyslightlylongerthanthecombinedlengthofvesicularstem+vesicle.Thevaginais short, reaches30–50 %of the lengthof thedart sacandmaybenarrowedbyatransversefolds.Dartshort,

Textfig.4.Superba skipetarica skipetarica (Subai 1995). — a)wholebodyextractedfromtheshell;b)situsofthegenitalorgans;c)dart;d)lumenofthedistalgenitalorgansshowingAST,penialpapillaandvagina.Albania,TomorMts.,KarNÇukaPartizan,easternslope.

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somewhat clumsy,2.3–2.7mm long, slightly curved;crownslightlyfunnel-shaped,narrowinthemiddle.

TheASTwithabroadandflatstempart,stronglyup-raisedandinaslightlyobliquepositioninthecentreoftheatrium.ThepenialarmoftheASTlongandreachesapproximatelyhalfof thepenial lumen; itbecomesflatatitsend.

Malepart:Thepenialpapillaiselongateconicalwithslightlysunkentransversefolds.Laterally,thereisafur-rowreachingfromthebasetothetipofthepapilla;theporeissituatedinthisfurrowatthebaseofthepapilla.Theflagellumreaches1.5thesizeofpenis+epiphallusandisslightlyloopedinitsbasalpart.

D i f f e r e n t i a l d i a g n o s i s : fordifferences to S. s. asketa n.ssp.refertotherespectiveparagraphunderthistaxon.S. skipetarica differsfromallotherspeciesofSuperbabythepresenceofpigmentationof thegenitalorgans(greyinS. s. skipetaricaandgreytodarkgreyinS. s. asketa).

TheshellofS. reischuetzi ismoredepressedandhasflatwhorlswithamorefaintpatternofyellowish-brown-ishspiralbands, thelowestbandissituatedmoreclosetotheperipheryoftheshellandnarrower,andtheradialstripesaremuchweaker; thegranulationof theproto-conchiscoarser;theglandulaearecylindrical(notnar-rowedatthetip),theASTislargerwithavaginalarm;thedartismoreslenderwithasmallertip.TheshellofS. kulmakanaislargeronaveragewithbandsofalmostthe samewidth; theumbilicus is relatively larger,andtheapertural insertionsarecloserand theapertural rimislarger;theglandulaearedeeplysubdivided.TheshellofS. grisea isusuallylarger,greyish,andthebandsaremuchweaker;theshellsurfaceissmooth,withanwhit-ish surface layerandblunt riblets; thediverticulum ismuchshorter.TheshellofS. vikosensis isalmostdiscoi-dalwith lowwhorlsanda relatively largerumbilicus;theaperture ismore transverselydilated,and theaper-turalrimnarrow;thebandsofS. vikosensis areweaker,and the radialstripes lessdeveloped; theglandulaearerelativelyshorterandcylindrical,theASTisbroaderandsituatedatthegenitalopening;thedartismoreslender.

D i s t r i b u t i o n :S. s. skipetarica lives insouthernAlbania in thenorthernpartof theTomorMts,aroundtheÇukaPartizanpeak.Itoccursinanaltitudinalrangeof1330–2240m.

R e m a r k s : In contrast toprevious idea aspoint-edout in theoriginaldescription,S.skipetarica isnotcloselyrelatedtoS. reischuetzi fromGreece.Thenomi-notypicalsubspeciesdisplayssomeremarkablevariationofshellmorphology.NorthandeastoftheÇukaPartizanpeak,analmostsmoothandmorenarrowlyumbilicateformcanbefound,wherethelowerbandisquitebroad,fadingintheperiomphalicareaandofteninterruptedbyradialstripesofbasicshellcolour.Recently,thisspecieswasfoundsouthof theÇukaPartizanpeak, too,wheretheshell’sumbilicus iswider, the lowerbandnarrower

andmoredistinct.On the teleoconch, the radialstripesarestronger,a regularpatternofnarrowribletscanbeobservedand tracesof a spiral threadscanbe found.ThispopulationisheretakenasbeingtransitionaltoS. s. asketa.

Superba skipetarica asketan.ssp.Plate2,fig.6;textfigs5,7–8

T y p e s a n d l o c u s t y p i c u s : Holotype HNHM95544.Paratypes:HNHM95466/5;Albania,PeriferiBerat,MaliiTomorrit,4.5KmNvonMajaeTomorrit,onlimestonerocksbelow the ridge,westernslope,2060malt.,UTMDL20,leg.Fehér,10.8.2004.—Additionalparatypes:coniferousforest2KmNofMajaeTomorrit,onlimestonerocksbelowtheridge,easternslope,2050malt.,UTMDL20,10.8.2004,leg.Fehér,HNHM95464/6;HNHM95462/8+2(alk.),SMF328956/1,S20213/1,1kmNofMajaeTomorrit,on lime-stonerocksbelowtheridge,easternslope,2230malt.,UTMDK29, leg.Fehér,10.8.2004;HNHM95460/10+2(alk.),S20212/1,700mNofMajaeTomorrit,onlimestonerocksbe-lowtheridge,2375malt.,UTMDK29,leg.Fehér,10.8.2004;hnhm 95982/34, h/34,ditto, leg.Fehér, hunyadi, huSZár & murányi; HNHM95458/2,2KmSofMaja eTomorrit,onlimestonerocks,2100malt.,UTMDK29,10.8.2004,leg.Fehér.

A d d i t i o n a l m a t e r i a l from the abovementionedlocalities(notconsideredastypespecimens):HNHM95467/2(juv);HNHM95465/2 (juv);HNHM95463/15 (juv.+ frag-ments);HNHM95461/8 (juv.+ fragments);HNHM95459/1(juv.),HNHM95983/3(juv.).

D i a g n o s i s : Shell small to medium sized, de-pressedconical,greyishbrown;spiralbands indistinct,surfaceof teleoconch sculpturedwith strong ribsandspiralthreads.

D e s c r i p t i o n :The shell is depressed conical toalmostflatwith4–5regularly increasingwhorls,whit-ish togreyishwith threechestnutbrownspiralbands.Theupper twobandsarenarrow reachingawidthofca. 0.5–1mmeach, the lowerband is situatedbelowtheperipheryandcanreachupto2mmwidth.Usually,theinitial2whorlsareyellowish-brownish.Theapexissometimesmammillate.

Theprotoconch consists of 1.5–1.75finelygranu-latedwhorls. Initially, thesurfaceof the teleoconch isfinelygranulated anddisplays low ribletswhichbe-camestrongerwith the increaseof thewhorls.On thelastwhorls,theseribsarestrongandaccompaniedbyasculptureoffinespiralthreads.

Thelastwhorlreaches1.6–2.1thesizeofthepenul-timatewhorl,weaklytostronglydescendingbeforetheaperture.Sutureofmediumdepthandslightlyundulat-ing.Umbilicusdeep,perspective; it increases regularlyfrom the initialwhorl,and increases rapidly insize inthe lastwhorlalmost reaching thedoublewidthof thepenultimatewhorl(ca.2.6–4.8mm).Aperturesphericaltooval,theinsertionswithadistanceof2.3–3.9mmand

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connectedbyathinbutcallus;aperturalrimsharp,later-allyandbasallysomewhatextended(0.8–1.25mm).

S h e l l m e a s u r em e n t s (n=35):H:6.5–10.3;D:14.4–24;AH:6.5–11;AD:7.5–12.8.

Theanimal’sbody(after4specimensfrom“700mN”and“1KmNofMajaeTomorrit”)isuniformlybrown,greytodarkgreywithabrownishhue;soleofthefootcream towhitishwithabrownish fringe;mantlewithweakand irregularlyplaced spotsofpigment,whichbecomelessnumerousintheupperwhorls.Thesecond-aryureterreachesalengthof10.5–20mm,attheenditissubdivided in twoseparateductsof1and1.5–2mmlength,respectively.Themandibleismediumbrownandhas3ridges.

C h a r a c t e r s o f g e n i t a l o r g a n s : Internally,manypartsofthegenitalorganswithgreyish-brownishpigment.

Femalepart:glandulaeelongate, somewhatnarrowatthetip,withbrownishpigmentation,reachingfivetosix times the lengthof thedart sac;usually, theglan-dulaeare simple,but ina fewspecimens, subdividedglandulaesplitat50%oftheirabsolutelength.Stemofbursacopulatrixshort,reaching20–25%ofthevesicu-larstem;diverticulumas longasorup to25%longerthanvesicularstem+vesicle.Vaginaaslongasoronlyhalfthelengthofthedartsac.Distally,thevaginallumenisnarrowedbya±broadandflat transverse fold.Dartshort,somewhatclumsy,1.9–2.35mmlong,withanar-rowstem,slightlycurved;crownfunnel-shaped.

TheASTasinS. s. skipetarica.Malepart:Distalpeniallumenlacksanyfoldsorpi-

lasters,but therearefine transverse foldon thepenialwallclose thepenialpapilla.ThepapillaandflagellumasinS. s. skipetarica,butflagellumslightlylonger.

Textfig.5.Superba skipetarica asketan.ssp.—a)wholebodyextractedfromtheshell;b)situsofthegenitalorgans;c)dart;d)lumenofthedistalgenitalorgansshowingAST,penialpapillaandvagina.a)–b)from:Albania,PeriferiBerat,MaliiTomorrit,700mNofMajaeTomorrit,onlimestonerocksbelowtheridge,2375malt.c)–d)from:PeriferiBerat,1kmNofMajaeTomor-rit,onlimestonerocksbelowtheridge,easternslope,2230malt.

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Etymology:Wewouldliketodedicatethisnewsub-speciestothreeyoungHungarianbotanists:Zoltán ba-rina, cSaba németh anddániel PiFKó,whoaccompa-niedthesecondauthoronhiscollectingtrip in2004tosouthernAlbania.Thename“asketa” isderived fromtheirnickname„theascetics“.

D i f f e r e n t i a l d i a g n o s i s :Superba s. asketa differs from thenominotypical subspeciesby the lessdistinctspiralcolourbands,andthepresenceofribsandwidelyspacedspiralthreadsontheteleoconchwhorls.Ifpresentinotherspecies (S. s. skipetarica,S.kulmakana orsometimes inS. grisea), thesespiral threadsareal-waysverynarrow,seldomdeepandrestrictedtothelow-erwhorl.TheshellofS. vikosensis ismoredepressed,itssurfacelacksanyspiralthreads,andtheradialribletsarelessdistinct;itsapertureismoretransverselydilated,andtheumbilicuswider.Themorphologyofthegenitalorgansissimilar,butitdiffersintheformofitsmorecy-lindricalglandulae,thelargerASTandthemoreslenderdart.

D i s t r i b u t i o n :Superba s. asketa ishithertoonlyrecordedfromtheTomorpeakinthesouthernpartoftheTomorMts. inavertical rangebetween2050–2375maltitude.

Superba vikosensis (Subai 1990)Plate2,fig.7;textfigs6,7–8

1990 Chilostoma vikosensis Subai,–AnnalendesNaturhis-torischenMuseums inWien,91 (B):245,Taf.1Fig.4–6 (shell),Abb.2 (genitalorgans) [Locus typicus:“Griechenland,Epirus,Vikos-Schlucht50–100mvomKlosterAg.ParaskevibeiMonodendri(UTMDK71)”,Holotype:NMW84365/1].

1990 Chilostoma vikosensis,–reiSchütZ & Sattmann, An-nalendesNaturhistorischenMuseumsinWien,91(B):,258,Abb.13(genitalorgans).

T y p e s a n d l o c u s t y p i c u s : Holotype NMW84365/1.Paratypes:NMW84366/2,REI/2,STU/5;Greece,Ípiros,VíkosGorge,50–100mfromthemonasteryAgíaPara-skevíatMonodéndri,UTMDK71,15.7.1987,leg.reiSchütZ & Stummer.—Additional paratypes: IZPAN/18,NNM/2,SMF328992/2,S10727/22+27 (juv., fragments+1alk.),150–200mfromthemonasteryAgíaParaskevíatMonodéndri,UTMDK71,11.4.1988, leg.riedel & Subai; STU/4,VíkosGorgeatthemonasteryÁgiosProfítisIlías(NofMonodéndri),UTMDK71,15.7.1987,leg.reiSchütZ & Stummer.

A d d i t i o n a l m a t e r i a l :F/5(+1alk.),S11671/12+4(juv.),Greece,Ípiros,VíkosGorge,50–100mfromthemona-steryAgíaParaskevíatMonodéndri,UTMDK71,27.9.1989,leg.Fauer & Subai;HNHM91015/7,ditto,10.8.1996, leg.Erőss & FEhér;NEUB/4+6 (juv.),VíkosGorge, vantagepointatMonodéndri,UTMDK71,19.7.1990, leg.neubert;S11772/3+6 (juv.),7.2KmSofKepéssovo, indirection toAsfákaat theTurkishbridge,on limestonerocks,700malt.,UTMDK81,23.5.1991,leg.Subai.

D i a g n o s i s :Shellsmall,almostdiscoidal,stronglydepressedwhorls; three indistinct spiralcolourbands;

surfaceoftheteleoconchwithstrongradialstripes;um-bilicuswide,aperturetransverselydepressed;glandulaeshort,dartslender.

D e s c r i p t i o n :Theshellisdepressedconicaltoal-mostdiscoidalwith4–4.5regularlyincreasing,bluntlykeeledwhorls.Thebasicshellcolouriswhitishwiththreenarrowspiralbandsoflighttomediumbrowncolour.Thetwoupperspiralsarenarrowandreachawidthofca.0.5mm.Thethirdspiralisalwayssituatedbelowtheperipheryoftheshell.Theupper2–2.5whorlsaresometimesyellowishbrown.Theprotoconchwhorls consist of1.5–1.75whorls

whicharesculpturedwithminutegranules.Thesurfaceof the teleoconch is shinywithveryfinegranulesonthefirsttwowhorls,itssculptureconsistsofirregularlyspacedradialriblets.Thelastwhorlreaches1.6–2.4thesizeofthepenultimatewhorlandbendsslightlytowardstheaperture.Sutureofmediumdepth,slightlyindentate;umbilicus funnel-shaped, regularly increasing inwidth.Inthelastwhorl,itsdiameterincreases1.5timesifcom-paredtothepenultimatewhorl.Itreachesadiameterof3.4–4.2mm.Aperturetransversallydepressed,theinser-tionswithadistanceof2.8–4.1mmandconnectedbyathinalmost invisiblecallus;apertural rimsharp,upperpartflattened; laterally andbasally enlarged reaching0.5–0.6mm,somewhatenlargedtowardstheumbilicus.

S h e l l m e a s u r e m e n t s (n=37):H: 6–7.5;D:16–20.3;AH:5.8–8;AD:6.8–10.1.

Theanimal’sheadandbody(afterasinglespecimenfromtheVíkosGorge,50–100mvomKlosterAgíaPar-askevíatMonodéndriandafterreiSchütZ & Sattmann (1990:Abb.13))aregreytobrownish,thetailandflankssomewhatbrighter; themantlewithapatternofirregu-larlyarranged,blurred light-grey spots.On theupperwhorls, these spots areonly fewbut theirdensity in-creasesonthelowerwhorl.Thesecondaryureterreachesalengthof16mm,iscompletelyopenandsplitstermi-nally in twocanalsof1and2mmlength, respectively.Themandibleismediumbrownandhas4ridges.

C h a r a c t e r s o f g e n i t a l o r g a n s :femalepart:glandulaesimple, reaching four times the lengthof thedartsac;stemofbursacopulatrixshort,reaching20%ofthevesicularstem;thediverticulumisaslongasthecom-binedlengthofvesicularstem+vesicle.Thevaginareach-esthesamelengthofthedartsac;distally,itisnarrowedbyalargetransversefold,whiletheinteriorissculpturedbylongitudinalpilasters;dartslender,elongate.

TheASTissituatedclosetothegenitalopening.Itformsastronglyupraisedtransversepadandreachesitsmaxi-mumheightinthecentreoftheatrium.ThepenialarmoftheASTreachesthepenialpapilla(orisevenlonger); itbecomesflatatitsend.Thevaginalarmisshortpointingtowardstheboundarybetweenatriumandvagina.

Malepart:Thepeniallumenisfilledwithbroadpilas-ters.Therearenarrowtransverse folds in theproximalpartofthepenisaroundthepapilla.Thepenialpapillaiselongateconical.Laterally,thereisadeepfurrowreach-ingfromthebasealmosttothetipofthepapilla;thepore

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issituatedinthisfurrowatthebaseofthepapilla.Theflagellumreaches1.3–1.5thesizeofpenis+epiphallus;ithasuptofourloopsinitsbasalpart.

D i f f e r e n t i a l d i a g n o s i s :TheshellofS. reis-chuetzi (which isgeographicallyclose) ismoreconicalwithmoredistinct spiralbands; thesurfaceof its tele-oconchissculpturedbyfinerradialstripes; theapertureismorespherical,andtheumbilicusnarrower;glandulasometimessubdivided,ASTwithtwoarms.TheshellofS.

s. asketa ismoreconicalwith,sculpturedstrongwhiterib-letsandsunkenspiralthreads;itsapertureismorespheri-cal, theaperturalrimmoredilated,andtheumbilicus isnarrower;theglandulaearerelativelylongerandacuteatthetip,theASTsmaller,andthedartmoreclumsy.

D i s t r i b u t i o n :Allhithertoknownrecordsarefo-cusedon a small area in theuppermost reachof theVíkosgorgeinnorthernÍpiros.

R em a r k s :Thestempartofthesingledartislost;forthisreason,onlythetipofthedartcouldbeillustrated.

Textfig.6.Superba vikosensis (Subai 1990). — a)wholebodyextractedfromtheshell;b)situsofthegenitalorgans;c)dart;d)lumenofthedistalgenitalorgansshowingAST,penialpapillaandvagina.Greece,Ípiros,VíkosGorge,50–100mfromthemon-asteryAgíaParaskevíatMonodéndri.

Discussion

ThespeciesofSuperban.gen.canbesubdivided intwogroups.ThefirstcontainsthespeciesfromsouthernAlbania,S. s. skipetarica,S. s. asketa,S. griseaandS. kul-

makana.Theyare found in theneighbouringmountainrangesoftheTomorandtheKulmakandinhabitthealpineranges.TheonlyexceptionisS. griseawhichwasfound

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218 Subai, P. & Fehér, Z.:RevisionoftheAriantinae,3.Superba nov.gen.

Textfig.7.DistributionofspeciesofSuperba.

inagorgeat1150maltitude(textfig.8).Onaverage,theirshellsarelargerandlessdepressedthanintheirrelativesfromGreece.ThedartsoftheAlbanianspeciesareshorterwithalongertip,andtheASTislow.

Incontrast,thetwospeciesfromGreece,S. reischuet-zi und S. vikosensis,turnedouttoliveinloweraltitudes(althoughthetypelocalityofS. vikosensisisat1100maltitude)(textfig7).Onaverage,theirshellsaresmaller,moredelicateanddepressed if compared to theAlba-nianspecies.TheirASTismoreelevated,andthedartislongerandmoreslender.

ItshouldbestressedthatthedistributionareasofallspeciesofSuperban.sp.areremarkablysmall.Often,it

islimitedtoarangewithadiameterof100mtoafewkilometresandcoverseachamountainsummitorridgeorasmallvalley.Incomparison,thedistributionrangesofspeciesofthemorecloselyrelatedgenusJosephinella usuallycover50kmormore,andspeciesofthegeneraCattaniaandLiburnicainhabitevenlargerareas.

Acknowledgements

WearegratefultoDr h. Schütt(Düsseldorf)forlinguisticrevisionoftheGermantext,Dr.E.neubert(Badenweiler)fortranslation,thephotosandthearrangementofthefiguresusedinthiswork.

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Subai, P. & Fehér, Z.:RevisionoftheAriantinae,3.Superba nov.gen. 219

Textfig.8.DistributionoftheAlbanianspeciesofSuperba in theTomorandKulmakësMountains.

References

dhora, dh.(2002):StudimembimolusqeteShqipërisë[Stud-ieson themolluscsofAlbania]:1–209. „Camaj -Pipa“,Shkodër.

dhora, dh. & Welter-SchulteS, F. W. (1996):Listof spe-ciesandatlasof thenonmarinemolluscsofAlba-nia.—SchriftenzurMalakozoologieausdemHausderNatur-Cismar,9:90–197;Cismar.

—&— (1996):List of taxaof non-marinemolluscs, de-scribed fromAlbanian localities.—Schriften zurMalakozoologieausdemHausderNatur-Cismar,9: 198–223;Cismar.

FEhér, Z., Erőss, Z., Kontschán, J. & Murányi, D. (2004):Collecting sites of the zoological expeditions oftheHungarianNaturalHistoryMuseumtoAlbania(1992–2003).Foliahistorico-naturaliaMuseiMat-raensis28:67–82;Gyöngyös.

GiuSti, F, manGanelli, G, & Schembri,P.J.(1995):Thenon-marinemolluscsof theMaltese Islands.—607pp;Torino.

reiSchütZ, P. l. & Sattmann, h. (1990):BeiträgezurMol-luskenfaunadesEpirus,II.—AnnalendesNaturhis-torischenMuseums inWien,91 (B):253–272.Taf.1–4;Wien.

Subai, P. (1990):BeiträgezurMolluskenfaunadesEpirus, I.ZweineueChilostoma-ArtenausNord-Epirus(Gas-tropoda:Ariantinae).—AnnalendesNaturhistor-ischenMuseumsinWien,91 (B):243–247;Wien.

— — — (1995):BeitragzurKenntnisalbanischerundgrie-chischerAriantinae (Gastropoda:Helicidae).—An-nalendesNaturhistorischenMuseums inWien,97 (B):75–94;Wien.

Welter-SchulteS,F.W.(1996):Non-marinemolluscsrecentlycollected inAlbania.—SchriftenzurMalakozoolo-gieausdemHausderNatur-Cismar,9:21–31;Cis-mar.

Manuscriptsubmitted:06.02.2006 Revisedmanuscriptaccepted:31.10.2006

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220 Subai, P. & Fehér, Z.:RevisionoftheAriantinae,3.Superba nov.gen.

Plate1Allfigures×3,phot.E.neubert

Fig.1. Superba grisean.sp. HolotypeHNHM95542,Albania,PeriferiSkrapar,QafaeDëvris,

NEofRadeshalongtheÇorovodë-Zaloshnjeroad,limestonerocks,1150malt.,D=22.7mm.

Fig.2. Superba reischuetzi (Subai 1990). ParatypeSMF328991;Greece,Epirus,rocksattheTurkishbridge

crossingtheAóosriveratKónitsa,D=16.9mm.

Fig.3. Superba kulmakana n.sp. HolotypeHNHM95543,Albania,PeriferiSkrapar,smallcanyon

1KmNWofMajaeKulmakut,onlimestonerocks,2070malt.,D=22.25mm.

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Plate 1

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222 Subai, P. & Fehér, Z.:RevisionoftheAriantinae,3.Superba nov.gen.

Plate2Allfigures×3,phot.E.neubert

Fig.4–5 Superba skipetarica skipetarica (Subai 1995). 4)Paratypeexcoll.Subai,Albania,KarNÇukaPartizan,eastern

slope,limestonerocks,1850malt.,D=18.95mm. 5)SMF328955,transitionalformofS. s. skipetaricatoS. s. asketa

n.ssp.;PeriferiBerat,MaliiTomorrit,6KmNofMajaeTomorrit,onlimestonerocksbelowthewesternslopeofthemountainridge,2060malt.,D=20.1mm.

Fig.6. Superba skipetarica asketan.ssp. HolotypeHNHM95544;Albania,PeriferiBerat,MaliiTomorrit,

4.5KmNvonMajaeTomorrit,onlimestonerocksbelowtheridge,westernslope,2060m,D=20.1mm.

Fig.7. Superba vikosensis (Subai 1990). ParatypeSmF 328992, Greece,Ípiros,VíkosGorge,50–100mfrom

themonasteryAgíaParaskevíatMonodéndri,D=18.75mm.

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Plate 2

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