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

eschweizerbartxxx sng-

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:[email protected]. Zoltán Fehér, HungarianNaturalHistoryMuseum,Barossutca13,H-1088Budapest,email:[email protected].


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.


Subai, P. & Fehér, Z.:RevisionoftheAriantinae,3.Superba nov.gen. 207

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



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.



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.



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



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.



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



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.



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



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.



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



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



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.



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



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.


Plate 1



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.


Plate 2

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

Documents

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