Revision of the Argninae, 1. The species of Agardhiella from

eschweizerbartxxx sng-

Arch. Molluskenkunde | 137 | (1) | 75 – 103 | 14figs,1plate | FrankfurtamMain,27.06.2008

©E.Schweizerbart’scheVerlagsbuchhandlung(Nägeleu.Obermiller),2008,ISSN0003–9284

Revision of the Argninae, 1. The species of Agardhiella from the countries bordering the Adriatic Sea

(Gastropoda: Pulmonata: Pupillidae)

Peter Subai

Kurzfassung

DieAgardhiella-ArtenimWestteilihresVerbreitungsgebieteswerdenrevidiert.EswerdenzweineueArtenausAlbanienbeschrieben:Agardhiella extravagantan.sp.undAgardhiella zoltanorum n.sp.AlleArtenwerdendurchFotosundZeichnungenabgebildet,ihrebisherbekannteVerbrei-tungaufKartendargestellt.FürA. formosa wirdeinNeotypusfestgelegt.

Schlüsselwörter:Taxonomie, Anatomie,Agardhiella, neueArten,Verbreitung.

Abstract

The Agardhiellaspeciesinthewesternpartoftheirareaofdistributionarerevised.TwospeciesfromAlbaniaaredescribedasnew:Agardhiella extravagantan.sp.and Agardhiella zoltanorum n. sp.Allspeciesareillustratedbyphotosanddrawings.Theirhithertoknownareaofdistributionispresentedonseveralmaps.ForA. formosa,aneotypeisdesignated.

Key words: Taxonomy, anatomy,Agardhiella,newspecies,distributionpattern.

Introduction

During several collecting tripsofHungarianmala-cologiststoAlbania,anumberofyetunknownlandsnailspecieshavebeendiscovered.Thesecollectorsentrustedmewith thedeterminationof theirAgardhiella speci-menstobeusedforarevisionofthewholegroup.

Comparingthesespecimenstomaterialofthisgenuscollected fromMontenegro, twospecies fromAlbaniaturnedouttobenewtoscienceandaredescribedhere.AtthesametimearevisionoftheArgninaeisstartedtoincrease thescarceknowledgeof thissubfamily. In thefirstpartof therevision, theAgardhiella-speciesof thewesternpartofthedistributionareaofthegenuswillbe

treated,i.e.Italy,Austria,Slovenia,Croatia,BosniaandHercergovina,Montenegro,AlbaniaandnorthwesternGreece.AfewrecordsofA. truncatellafromSerbiaandSlavicMacedoniaarealsomentioned.

MaterialandMethods

Theshellsof thespeciesofAgardhiella aremainlyfound in thedebris ofwaterbodies,while records fromprimaryhabitats are extremely rare.This canbe ex-plainedbythecrypticwayoflifeoftheseanimals.Agar

Author’saddress:Peter Subai,Kronenberg143,D-52074Aachen;email:[email protected]


76 Subai,P.:RevisionoftheArgninae,1

dhiella species live in the soil interstitialprovidedbyrootsandsmallstonesintheimmediatevicinityofrocks.Theyclearlyprefercalcifictypesofsoil.Theinterstitialkeepshumidityoveraconsiderableperiodof timeandthusconstitutesanidealbiotopeforseveralsmall-sizedsnail species.Forexample, theexcavatedsoilbetweentherootsofasmalltreenearLoutra(=5kmNofAma-randos)inEpirus,northwesternGreeceyielded,besidesAgardhiella truncatella,theshellsofspeciesofOrcula,Sphyradium,Acanthinula,Punctum,Vitrea,Gyralina andDaudebardia!

Thepreferredsamplingmethodisasfollows:carefullystir thesoil inabucketfullofwater,andthetinyshellswhicharefilledwithairwillswimat thesurface.Theythencanbeeasilyskimmedbyafinesieve.This is theonlymethodwhichyieldsnumerousspecimensofactuallyrareAgardhiellaspecies.Besidesthat,Agardhiellaspeciescanbe found incavesandrockcavitieswhichprovideaccess to thesubterraneanhabitats.SomeAgardhiella-specieslikeforexampleA. daboviciandotherspeciesinRomaniahaveexclusivelybeenfoundincaves.

Often,thenumberofribsonthelastwhorlisusedtocharacterizespecies(e.g.speciesofAgardhiella,Renea,Spelaeodiscusandothers).However,ithasbeenshownthatinshellswithvaryingsizes,thenumberofribsvar-iesaswell.Forexample, a slender specimenhas lessribson the lastwhorl thanabroaderoneof the samepopulation.Therefore, thenumberof ribswithin1mm(onthelastwhorl)asusedbyPilSbry(1924)isaddedtocompletetheusualsetofcharacters.

ThespellingofAlbanian localities followsFehéretal.(2004).

Allmeasurementsinmillimetres.

Abbreviations:

CNHM CroatianNaturalHistoryMuseum,ZagrebE privatecollectionZ. Erőss,BudapestH privatecollectiona. hunyadi,BudapestHNHM HungarianNaturalHistoryMuseum,Buda-

pestMaa privatecollectionJ. M. W. MaaSSen,Duiven-

drechtMMMB MunkácsiMihályMuseum,BékéscsabaNMW NaturalHistoryMuseum,ViennaNMWE CollectionedlauerinNMWNMWK CollectionKleMMinNMWNMWR CollectionrušnoWinNMWNNM NationaalNatuurhistorischMuseum,LeidenS privatecollectionP. Subai,AachenSch privatecollectionh. Schütt,Düsseldorf-

BenrathSMF NaturmuseumSenckenberg,Frankfurtam

Main± moreorlessalk. specimenpreservedinalcoholD shelldiameterAD diameterofapertureH heightoftheshellAH heightoftheaperturejuv. juvenilespecimenn.det. UTM-CodecouldnotbedeterminedUTM UTM-CodefortheinventoryofEuropean

Invertebrates

Taxonomicaccount

Agardhiella truncatella (l. PFeiFFer1841)Plate1,Figs.1–4,textfigs1–3

1841 Pupa truncatella l. PFeiFFer, Symbolaeadhistoriamheliceorum,1: 46 [locus typicus: “...inmonteKarstpropeCastelnuovo, interTergestumetFiume” (par-tim);typematerial:lost].

1842 Pupa truncatella,– roSSMäSSler, IconographiederLand-&Süßwasser-Mollusken,(1)2(5):12,Tafel53,Fig.733(partim,notthevarieties).

1848 Pupa truncatella,–l. PFeiFFer, Monographiaheliceo-rumviventium,2:303(partim).

1850 Pupa truncatella,–KüSter, In:Martini & cheMnitz:SystematischesConchylienCabinet,I (15):34,Tafel4Fig.20–21(partim).

1886 Pupa (Sphyradium) truncatella, – Pollonera,MonografiadegliSphyradiumitaliani:16.

1887 Sphyradium truncatella, –cleSSin, DieMollusken-FaunaMitteleuropas,II.Teil,DieMolluskenfaunaÖs-terreich-UngarnsundderSchweiz:242,Fig.144.

1887 Pupa (Coryna) truncatella,–WeSterlund, FaunaderinderpaläarctischenRegionlebendenBinnenconchylien,3:88.

1890 Pupa (Coryna) truncatella, –Flach, Verhandlungenderphysikalisch-medizinischenGesellschaftzuWürz-burg,(2)24:52.

1915 Agardhia (Agardhia) truncatella, – Sturany & Wag-ner,SchriftenderKaiserlichenAkademiederWissen-schafteninWien,mathematische-naturwissenschaftli-cheKlasse,91:64,Tafel18,Fig.102.

1926 Agardhia (Agardhiella) truncatella,–PilSbry, ManualofConchology,27:162,Plate.19Fig.2–5.

?1939 Agardhia truncatella depressa PetrboK,–VěstnikČs.zoologickéspolečnostivpraze,6/7:344[locustypicus:„...intherockycleftsoftheKarstatČebič...“typema-terial:unknown].

?1959 Sphyradium truncatella,–Tomić, Akademija,27:32(partim).

1971 Agardhia (Agardhiella) truncatella, –alzona, AttidellaSocietàItalianadiScienceNaturaliedelMuseoCivicodiStoriaNaturalediMilano,111:70.


Subai,P.:RevisionoftheArgninae,1 77

1974 Argna (Agardhiella) truncatella,–KleMM, Denkschrif-tenderÖsterreichischenAkademiederWissenschaften,mathematisch-naturwissenschaftlicheKlasse,117:169,distributionmap42(p.168).

1974 Agardhiella truncatella,–gittenberger, Basteria,38:3,Fig.1(shellsculpture),Fig.3–4(genitalorgans).

1975 Agardhiella truncatella truncatella, – gittenberger, Zoolo-gischeMededelingen,48(24):286,Fig.6(genitalorgans).

1983 Argna (Agardhiella) truncatella,–Kerney, caMeron & Jungbluth, DieLandschneckenNord-undMitteleu-ropas:123,Fig.p.124,distributionmap110.

1985 Agardhiella truncatella, – zilch, Archiv fürMol-luskenkunde,116(1/3):129.

1985 Agardhiella truncatella skipetarica, –MaaSSen, De Kreukel,21 (1–2):4,Plate1,Fig.1–3,distributionmap[nonskipetarica a. J.Wagner1915].

1985 Agardhiella truncatella truncatella, –MaaSSen, De Kreukel,21(1–2):4,Plate1,Fig.5–8,distributionmap.

1999 Agardhiella truncatella skipetarica, – Subai, SchriftenzurMalakozoologie,13:45,Fig.3(shell),Fig.5(distri-butionmap),Plate.9,Fig.2(shell)(partim).

D i a g n o s i s :small,aperturesmall,withinfraparie-talisandangularis.Lower2/3ofthepalatalwallslightlybulge-likethickened.

D e s c r i p t i o n : Shell relatively small, cylindri-calwitha roundedapex;upper1.75teleoconchwhorlssmooth,slightlygranulated;afterthesewhorls,theper-pendicularribbingpatternbeginsandincreasesrapidly;on the lastwhorl thereare34–63(9–14/mm)ribs.Be-tweenthemainribsthereareirregularperpendicularribsonlyrecognizablewithhighmagnifications.

Whorls5.25–6.75; the lastwhorlup to51–63%ofthe totalheight;umbilicusnarrow, somewhatpointed.Aperturewithaninfraparietalisandangularis.Bothteethweaklydeveloped,stretchingstraight towards the inte-rioroftheshell.Aperturalrimu-shapedinfrontalview,interiorlyslightlyenlargedlikealipandwidenedexte-riorly;at thecolumellar side, it slightlybents towardstheumbilicus; in thecentreof thepalatal sideof theaperture, theaperturemaybesomewhat thickenedbut

Textfig.1.Agardhiella truncatella (l. PFeiFFer1841).Austria,Carinthia,KarawankenMts.,TscheppaGorge(=SofKlagenfurt),atlimestonerocks,UTMVM44,(H=3.81,D=1.62mm).



neverformsatooth(seeremarks);sometimes,thisareaismarkedbyformingaslightfurrow;abovethistheap-ertureisrecessed,andtheaperturalrimisnarrowwith-out lip.Above theangularis,ashortsinulus ispresent.Theperistomial rimsconnectedbyawell recognisablecalluscovering theupperpartof theapertureandcon-cealingtheribsofthelastwhorl.Insideview,theupperpartoftheaperturalrimisslightlyprotruding.

M e a s u r em e n t s (n=106):H=2.87–4;D=1.31–1.75;AH=1.06–1.37;AD=0.75–0.94.

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 (accordingto2specimenfromAlbania,SofGurriiZi;gittenberger 1974:3–6,Fig.3–4and1975:286,Fig.6):Theretractormuscleoftherighteye-stalkrunsbetweenpenisandva-gina;penissubdividedinaslenderdistalandaswollenproximalpartofequallength;theappendixinsertsinthemiddlebetweenthetwopartsofthepenis;itscentralpartisnarrow,while thedistaland theproximalpart reachdoubleof thewidthof thecentralpart.gittenberger foundthebranchesofthepenisretractormuscletobeofdifferent length,while in thespecimens fromAlbania,thebranchesareofequallength.Additionally,ingitten-berger’sspecimen,theappendixretractorinsertsshortlybehindtheinsertionoftheappendixtothepenis,whileintheAlbanianspecimens,itconnectsattheendofthedistal(wider)partoftheappendix.TheepiphallusoftheAustrianspecimen (gittenberger1974:Fig.3–4)andoftheSlovenianspecimen(gittenberger1975:Fig.6)reachesaboutthesizeofthepenis.Itconsistsofalong,narrowproximalpartandashortdistalpart(1/4–2/5ofits total length)whichmay reachdouble thediameterof theproximalpart. In theAlbanian specimens, theepiphallusonly reacheshalfof the lengthof theAus-

trian/Slovenianspecimens,and the transitionzonebe-tween thenarrowand thebroadpart is inconspicuous.Theoviductislongerthanthevagina.ThepedunculusofthebursacopulatrixoftheAustrian/SlovenianspecimenisremarkablylongerifcomparedtothatoftheAlbanianspecimens.Inthelatter,thevesicleofthebursacopula-trixislargeandbroadonashortstem.

D i f f e r e n t i a l d i a g n o s i s :Apart from thepa-rietalteeth,A. biarmatahasawelldevelopedlongpari-etallamella,andsomespecimensuptothreecolumellarteeth.Intheaperture,itsparietalwallisstrongerwithamoredeveloped indentation. Itsapertural rim isnotasstronglylippedasinA. truncatella.However,thecolum-ellarsideoftheapertureisstrongerinA. biarmatathaninA. truncatella.

TheshellofA. skipetarica ismoreslender,and theribbingpatternismoredense.Moreover,itsaperturehasnoteeth.InA. skipetarica, thepalatalwallof theaper-tureismoreprotrudedbutlessstrengthened.

T y p e s :lost.M a t e r i a l e x a m i n e d :Austria:EastTyrol,Tristach-

erLakenearLienz,UTMUM38, (exWiedeMayr)NMWE8487/1; Tristacher Lakewall, UTM UM 38, leg.KleMM 30.7.1948, NMWK 27534/2 (juv./damaged); leg. Seidl 26.5.1967,NMWK54976/1.

UpperAustria,Steinbacha.d.Steyr,UTMVP41,leg.gan-SlMayr,NMWK50429/2+1 (damaged);Küpfern,UTMVP70,leg.ganSlMayr,NMWK48783/4;Weyera.d.Enns,UTMVP70,NMWK38390/1;Weyera.d.Enns,Bichlbauerkogel,UTMVP70,leg.ganSlMayr,NMWE37576/1.

Salzburg,SalzburgCity,UTMUN59,NMW68443/2.Styria, Leopoldsteiner Lake, Hochschwab, UTM VN

86/96,leg.Franz11.10.1943,NMWK61106/1;PretterauSWslope, roadHieflau-Eisenerz,Mörchjassingau,UTMVN87,

Textfig.2a–b.Agardhiella truncatella (l. PFeiFFer1841),situsofthegenitalorgans.Albania,PeriferiElbasan,SofGurriiZi,13kmfromtheElbasanjunctionontheroadtoQafaeShtyllës,limestonerocks,900malt.,UTMDL26.—Abbreviationstothedrawingsofgenitalorgans:A=appendix,B=bursa,E=epiphallus,O=oviductus,P1=proximalpartofpenis,P2=distalpartofpenis,Pd=pedunculus,R=retractormuscle,V=vagina,Vd=vasdeferens.



leg.Franz,NMWE39147/1 (juv.); frontof theRadmerVal-ley,„Gesäuse“,UTMVN87, leg.Franz3.10.1946,NMWK61107/1(juv.);leg.edlauer,NMWK56096/5;Radlpass,UTMWM16,leg.FranK10.10.1972,NMWK66375/2;Grand,WoftheHohenRannach,UTMWN22,leg.boecKel1930,NMWE9455/2;Glaserer,UTMWN22,NMW30902/1;PfaffenkogelnearStübing,UTMWN22, leg.KreiSSl19.3.1970,NMWK63393/1;Hohenstein,peaknearStiftRein,UTMWN22,leg.KleMM9.1943,NMWK21255/1;HohensteinnearStiftRein,slope toHörgasgraben,UTMWN22, leg.KleMM 9.1943,NMWK21256/1;HörgasgrabennearStiftRein, rocksat theslope ofHohenStein,UTMWN22,NMWE36300/1+1(juv.);Mühlbachgraben,UTMWN22,NMW30903/2;Peg-gau,UTMWN22, leg.Franz 12.7.1946,NMWK40829/2(juv.); rockson the right banksofMur river nearPeggau,UTMWN22,leg.KleMM9.1943,NMWK21257/1;PeggauerwallnearPeggau,UTMWN22,leg.KleMM9.1943,NMWK21258/1+3 (juv.);Waterfall at ruinsofPeggau,UTMWN

22, leg.KleMM9.1943,NMWK21259/3+4 (juv./damaged);southwesternslopeTanneben towardsPeggau,UTMWN22,NMWE27379/10(partly damaged); (exPaget),HNHM/1;Peggau, gorge above Badlgalerie, UTMWN 23, NMWE36744/9;gorgebelowBadlgalerie,MurValley,UTMWN23,leg.KleMM9.1943,NMWK21253/2+1(juv.);Badelgalerie,UTMWN 23, NMW 30890/3+6 (juv./damaged); Badl-galerie,Hochtrötsch,UTMWN23,leg.FuchS,NMWR/1;(exFuchS,exKäuFel)NMW59768/5;BadlgrabennearPeggau,UTMWN23,NMWE27717/18+3(juv),30412/1,NMWK38372/3,SMF200505/4;Badlgraben,MurValley,UTMWN23, leg.KleMM 9.1943,NMWK21252/1;wallon the rightbankofMurriveroppositeofBadlgalerie,UTMWN23,leg.KleMM 9.1943, NMWK 21254/1; [Mountain] GschwendtnearFrohnleiten,UTMWN23,NMW30899/1;Bärenschütz-klammnearMixnitz,UTMWN24,NMWK38368/1; leg.Franz 18.6.1947,NMWE53754/2 (juv.);Eggenberg,UTMWN31, leg.FranK 31.7.1973,NMWK66377/1+1 (juv.);



debrisofTurcavenearSemriach,UTMWN32,leg.KreiSSl 2.11.1969,NMWK60825/2;Andritz source,UTMWN32,NMW30901/1+2(juv.);RadegundnearGraz,UTMWN32,Novistein, leg.Franz 17.4.1950,NMWE55159/1;KlammnearSt.Radegund,climbtoGöstinger-hut,UTMWN32,leg.Franz19.5.1959,NMWK46529/1(juv.);Raabklamm,UTMWN33/44,leg.Franz,NMWK57266/1;WeiznearGleisdorf,UTMWN42,NMWK38389/2;Weizklamm,UTMWN43,leg.Franz23.5.1948,NMWE54584/1;leg.Franz17.7.1959,NMWK46534/3;Weizklamm,Trögersteig,UTMWN43,leg.KoFler28.4.1942,NMWK56901/2+1(juv.);WeiznearGleisdorf,WeizklammbehindFeister,limewallN-side,UTMWN43,leg.Franz25.5.1948,NMWE54236/1.

Carinthia,UTMn.det.,SMF51763/1,51776/1,51777/3,110480/3, 247179/3, 247180/1 (+ 3L. cylindracea); leg.KoKeil 1862,NMW68/4+1 (juv.); lowerWolayerValley,UTMUM36, leg.ehrMann 1906, SMF51758/4; Soleck,UTMUM37, leg.KleMM 6.1932,NMWK1518/14 (partlydamaged);WildensenderValley,UTMUM37,(exehrMann) SMF51756/1;Kötschach-Mauthen,Valentinklamm(=Mauth-nerKlamm),UTMUM46,leg.KleMM6.1931,NMWK941/1;leg.KleMM7.1940,NMWK7156/6+2(juv.);leg.SchlicKuM 26.8.1941,SMF274293/13; leg. 10.1942, (coll.S.H. Jae-cKel)SMF200506/2+1;leg.neutebooM13.8.1952,NNM/1;leg.Schütt7.8.1958,Sch123/1; leg.nordSiecK2.10.1962,SMF205799/1;leg.gittenberger9.1964,NNM/12;leg.git-tenberger&riPKen 9.1964,NNM/4; leg.Seidl 24.7.1965,NMWK54977/2; leg.StuMMer 8.8.1978, S 9362/12; leg.MaaSSen9.1979,Maa/7;NorthernpartValentinklamm,UTMUM46, leg.KleMM 29.8.1948,NMWK27410/1,NMWK27441/5+4 (juv.); lowerValentinklamm,UTMUM46, leg.Seidl27.5.1987,NMWK54978/1+1(juv.);PlöckenpassatEd-erwirt,UTMUM46,leg.SchlicKuM9.8.1958,SMF274294/1;leg.gittenberger9.1964,NNM/1;Plöckenpass,Valentinbank,1050malt.,UTMUM46,leg.neutebooM14.7.1958,NNM/5;PlöckenpassbelowCellonkofel,UTMUM46,leg.ehrMann,SMF51760/2;Plöckenpass,at limestone rocks,1360malt.,UTMUM46, leg.Subai 13.7.2006,S20810/15+3 (juv.);gravelsEofPlöckenpass,1370malt.,UTMUM46,leg.git-tenberger9.1964,NNM/8;Kronhofgraben,UTMUM56,leg.höbbel9.6.1956,NMWK40973/1;Weissbriach,UTMUM67, leg.KleMM 27.7.1948,NMWK27356/4+2 (juv./dam-aged);Obervellach,Groppenstein,UTMUM76,leg.thorSon 30.6.1930,NMWK62890/1+2 (juv.); ruinsofOrtenburg,UTMUM88,(exgallenStein),NMW53816/3;nearlyofdamofWeissenbach,betweenZlanandWiederschwing(=5.5kmWofPaternion),700malt.,UTMUM97,leg.breure17.6.1986,NNM/7;Federaun,[Mountain]Dobratsch,UTMVM05, leg.KleMM3.6.1960,NMWK44652/1(damaged);DobratschnearOber-Federaun,750malt.,UTMVM05, leg.gittenberger 5.9.1969,NNM/3;KarawankenMountains,Mallestig,UTMVM15, leg.KleMM1.8.1960,NMWK44577/2 (juv.);Kan-zianiMountainnearVillach,UTMVM15,leg.KleMM7.1940,NMWK7193/4;Goritschach,UTMVM15(orVM65), leg.Strouhal23.8.1957,NMWK57628/1+3(juv./damaged);Os-siacherTauernnearVillach,VM16,leg.Kühnelt10.10.1942,NMWK60949/2;AnnenheimatOssiacherLake,UTMVM16, leg.Sturany 1893,NMWwithoutNr./1 (juv.);Maria-Elend,GrosserSuhagraben, 750malt.,UTMVM25, leg.neutebooM 12.8.1952,NNM/3;Karawanken,Rosenbach inRosentalValley,UTMVM25, leg.KleMM6.1934,NNM/5;leg.KarneKaMP8.1970,Maa/4;leg.Mildner7.4.1971,NMW79388/1;RosenbacherSchweiz, rocknarrows,UTMVM25,

leg.KleMM6.1934,NMWK2182/5,2183/8(damaged);alongthe road toBärentalValley, 790malt.,UTMVM34, leg.gittenberger 9.1964,NNM/1; road fromBärentalValleytoBauerAurig,UTMVM34, leg.gaSchott, (exKäuFel) NMWK16190/1+1(juv.);Sattnitz,UTMapp.VM35/45/55,NMWK 33071/5; (exFuchS, exKäuFel) NMW59769/4;(cleSSin, p. 242)NMW22079/1; (exgallenStein)NMW53815/23(±damaged);TscheppaGorge (SofKlagenfurt),atlimestonerocks,UTMVM44,leg.KleMM3.7.1960,NMWK44224/7+5(juv.);leg.SchuSter1975,NMWK69306/1(juv.);leg.Subai 24.8.1981,S8344/23+3 (juv.);Tschaukofall atLoiblPass,UTMVM44,leg.gittenberger9.1964,NNM/1;KotlaofFerlacherHorns,UTMVM44,leg.KleMM6.7.1960,NMWK44276/1+1 (juv.);Loiblpassheight,UTMVM44,leg.KleMM10.7.1960,NMWK44351/1+3(juv.);Loiblslope,UTMVM44,NMW18233/2+6 (juv.);Loiblpass road (SofKlagenfurt), 2kmN from restaurant „DeutscherPeter“,UTMVM44, leg.MaaSSen 9.1979,Maa/2+2 (juv.); samespot,50mSof restaurant„DeutscherPeter“,UTMVM44,leg.Subai24.8.1981,S8353/10+2 (juv.); samespot,500mSofrestaurant„DeutscherPeter“,1000malt.,UTMVM44,leg.MaaSSen9.1980,Maa/4 (juv.);KarawankenMountains,Rabenberg,UTMVM45,NMWK40942/1 (juv.);Roggen-bergnearMaiernigg,UTMVM46,leg.Strouhal13.7.1956,NMWK57621/1;KoschuttaatKoschutta-hut,UTMVM54,leg.Franz31.8.1960,NMWK46549/2+2(juv.);KarawankenMountains,western sideofKuhberg (Hochobir),UTMVM54, leg.KleMM8.8.1960,NMWK44742/3;Freibachgraben,westernsideHochobir,UTMVM54/55,leg.Fauer31.8.1965,NMWK52698/3+3(juv.);Gurnitz,gorgeSattnitz,UTMVM56, leg.KleMM 25.7.1960,NMWK44475/1+1 (damaged);leg.Mildner1.5.1971,NMW79387/1;Launsdorf, castleofHochosterwitz, 650m alt.,UTMVM57, leg.neutebooM 8.7.1952,NNM/3;Magdalensberg,UTMVM57,(exgallen-Stein),NMW53813/2;leg.FalKner9.1966,NMWK56509/4(damaged); ruinsofKraig,UTMVM58, (exgallenStein) NMW53818/3+1 (juv.);S-partofVellacherKocna,1150malt.,UTMVM63, leg.gittenberger29.7.1981,NNM/2;Ei-senkappel,UTMVM64,leg.KleMM6.1932,NMWKwithoutNr./5;Eisenkappel,Vellachbanks,530malt.,UTMVM64,leg.neutebooM9.8.1952,NNM/1;LobnikgrabennearEisenka-ppel,UTMVM64, leg.KleMM8.1941,NMWK7533/6+3(juv.);LobniknearEisenkappel,UTMVM64, leg.KleMM 6.1932,NMWK1395/1+8 (damaged);TrögernklammSWofEisenkappel,UTMVM64, leg.MaaSSen9.1979,Maa/1;BozicevaGoranearEisenkappel,UTMVM64/65,leg.KleMM 8.1941,NMWK7901/3;waterfallofWildenstein,UTMVM65, leg.KleMM 12.8.1960,NMWK44822/3+1 (juv.); leg.G.&P. Subai 22.8.1981,S 8388/9; leg.MaaSSen 5.1985,Maa/22+5(juv./damaged);Eberstein,UTMVM68,(exgal-lenStein),NMW53814/1;KupitzklammnearEisenkappel,UTMVM74,leg.KleMM6.1932,NMWK1396/1(damaged);leg.KleMM8.1941,NMWK7565/3+2(damaged),7844/3+2(juv.);leg.neutebooM9.8.1952,NNM/5;leg.MaaSSen5.1985,Maa/17+3 (juv.); leg.Schütt26.5.1985,Sch1157/2;8kmWSWofBleiburg,caveentranceDumpelnicanearGlobasnitz,UTMVM75,leg.gittenberger8.1981,NNM/2;ruinsofRab-enstein,UTMVM97,(exgallenStein),NMW53817/2.

Italy: Pieve di Cadore, UTM TM 94, leg. FentroP 12.8.1959,NNM/12;Auronzo,FiumeAnsiei,845malt.,UTMUM05, leg.neutebooM20.7.1968,NNM/1;SáurisdiSopra,UTMUM24, (exehrMann),SMF51755/2;ForniAvoltri,UTMUM26, leg.ehrMann 1911,SMF51751/1; between



CimaSappadaandForniAvoltri,UTMUM26,leg.ehrMann 1911,SMF51759/4;Rigolato-Comegliáns,DeganoValley,UTMUM35, leg.ehrMann1906,SMF51748/1; lowerFra-snettanearCollina,UTMUM36, leg.ehrMann1911,SMF247178/1;Rivo,S.PietroValley,UTMUM45,leg.ehrMann 1905,SMF51757/2;Malborghetto(=Malborgeth),UTMUM75,(exc. r. boettger)SMF110482/3;ValCanale(=CanalValley),UTMUM75/85/95,NMW24809/3+2 (damaged);(exreSSMann, coll.Kobelt) SMF51773/6; (exreSSMann 1876, coll. JetSchin)SMF110479/3;ValCanale,Lußnitz-graben,UTMUM75, leg.ehrMann 1904, SMF51794/3;StupizzaalNatisone (=Stupizza),UTMUM81, leg.nord-SiecK27.8.1965,SMF205800/1;upperSeisseraValley,UTMUM84, leg.ehrMann 1904,SMF51754/5;Bartolograben,UTMUM85, leg.ehrMann 1904,SMF51747/1;SeebachValleyatWeißenfels,UTMUM94,leg.ehrMann1904,SMF51750/1;SlizzaCanyonnearTarvisio (=Tarvis),UTMUM94,NMWE46027/1;CoccaudiSopra(=GoggaunearTarvis),UTMUM95, (exStuSSiner1888,coll.o. boettger),SMF51768/4;dolinePercidolnearOpicina (=NofTriest),UTMVL05,leg.1916,NMWE8489/1.

Slovenia (=Krain,Carniolia):UTMn. det.,HNHM/1;SMF51767/1,51770/6,51779/4,110477/9,200508/3,NMW61278/2+1(juv.)(veryslenderspecimens);(exSchMidt1862)NMW94/2;Žaga,Boka-waterfall,UTMUM82, leg.MaaS-Sen 9.1982,Maa/2 (damaged); left shoreNofŽaga,UTMUM82,NMWK48604/6+1 (juv.); leg.KäuFel,NMWK2799/1;Žaga,sourceatriverSoča,600malt.,UTMUM82,leg.MaaSSen7.1982,Maa/3; rockbottom1kmbelowŽaga,UTMUM82,NMWE10914/10;(exedlauer,exSchlicKuM),SMF274292/2;Srpenica (=2kmSofŽaga),UTMUM82,leg.MaaSSen9.1985,Maa/1;StaroSelonearRobič,UTMUM82,leg.MaaSSen9.1985,Maa/5;sourcePerilonearRobičnearKobarid,UTMUM82,leg.Schütt17.5.1975,Sch775/1;Vrh[=peak]RombonnearBovec,UTMUM83, leg.MaaSSen 9.1985,Maa/3;KaninMountainnearBovec,900malt.,UTMUM83,leg.MaaSSen7.1982,Maa/2;samespot,2000malt.,UTMUM83, leg.MaaSSen7.1982,Maa/11;2kmNofKo-barid,200malt.,UTMUM92, leg.MaaSSen9.1982,Maa/1(damaged);SočaatriverSoča(=Isonzo)(NofFlitsch),UTMUM93,(exKuščEr)NMW50580/3;Soča,alongsideofTrenta[=rockwall],UTMUM93, leg.MaaSSen 9.1985,Maa/1;MountainNanos,UTMVL26/27,(exgySSer,exullePitSch),SMF51762/4;5kmSof Idrija,UTMVL29, leg.MaaSSen 9.1979,Maa/1;3kmEofPostojna,650malt.,UTMVL46,leg.gittenberger9.1970,NNM/3; rocksnearlyofPlaninskacavenearPlanina,UTMVL47,leg.MaaSSen9.1979,Maa/2;Vrhnika,rocksatsideSourceofLjubljanicanearMirke,UTMVL48,NMWE16088/1+1 (juv.);Vrhnika-DolenjiLogatec,UTMVL48, leg.nordSiecK10.9.1961,SMF205797/1;do-linenearLaze(=NNEofPlanina),UTMVL48, leg.MaaS-Sen 9.1979,Maa/1;Vrhnika (=Oberlaibach), rocks on theleftbankofRetovjeSource,UTMVL49,NMWE15974/2;Vrhnika,at theMočilnikSource,UTMVL49, leg.edlauer,NMWK2798/1; leg.nordSiecK 12.9.1961,SMF205798/1;Vrhnika,rocksaboveMočilnikSource,UTMVL49,NMWE15949/20+1 (juv.),withoutNr./2+1 (juv.); entranceof Ib-linecavenearKompolje,UTMVL76, (exKäuFel)NMWK16195/1+2 (damaged); in „stopar teva skedeunza“ nearPonikve,UTMVL77,(exKäuFel)NMWK16191/1;entranceof„RaskaSkedeunta“nearPonikve,UTMVL77,(exKäuFel) NMWK16194/1(damaged);Kočevje(=NEofRijeka),UTMVL85,NMWK38376/3;„sestra trehbratov“nearKočevje,

UTMapp.VL85,(exKuščEr)NMWE48840/8+2(damaged);caveNEofKočevje,UTMVL95,NMWK38364/1;caveofZeljne(=NEofKočevje),UTMVL95,NMWE45136/1+1(damaged);(exKäuFel)NMWK16192/2+2(damaged);NofTolminka,bifurcationofTolminkaandZadlascica,250malt.,UTMVM01, leg.gittenberger9.1970,NNM/1;slapSavi-canearBohinj,UTMVM02, leg.MaaSSen7.1980,Maa/3;Vršičpasssouthernside,1350malt.,UTMVM04,leg.MaaS-Sen9.1982,Maa/6;samespot,1600malt.,UTMVM04,leg.MaaSSen6.1987,Maa/25+3 (juv./damaged);Stopnik (=NWof Idrija),UTMVM10, leg.MaaSSen 9.1979,Maa/1 (juv./damaged);lakeofBohinj,UTMVM12,(exKuščEr)NMWE48895/1;Bohinj (=Wochein), „Hot.Zlatorog-kantine“, slapSavica,UTMVM12, (exKäuFel)NMWK16188/2;Gozd-Martuljek,at limestonerocksnearlyofJermanStream,UTMVM14, leg.Subai 20.8.1987,S13578/3;western slopeofBabjiZob,1000malt.,UTMVM23, (exKäuFel)NMWK16199/1;summitofVelikoOsojnica,750m,UTMVM23,(exKäuFel)NMWK16197;Mlinca,UTMVM24, (exKuščEr) NMWE48807/1;caveMlinca,UTMVM24,(exStuSSiner,exO.boettger),SMF51766/4;„StuncupriSelcah“nearlyoftherailwaystationSava,UTMVM24,NMWwithoutNr./1;(exKuščEr)NMWE49635/1;„Zaškakukavadoline“Noftherail-waystationPlanina,UTMVM24,NMWE20179/8,NMWK48603/3;Bled, castleMountain,UTMVM33, leg.KleMM 6.1934,NMWK2181/1 (juv.);SavebanknearRibno,Bled,UTMVM33, (exKäuFel)NMWK20219/1+1 (damaged);„Hud.SavaBohinjska“ nearRibno, 420m,UTMVM33,(exKäuFel)NMWK16196/1+1 (damaged);VintgarklammnearBled (=Veldes),UTMVM33,NMWE15109/12; leg.MaaSSen 7.1979,Maa/2;KarawankenMountains,Southernslopeof theMountainStol,UTMapp.VM34, (exKäuFel) NMWK51688/4,38367/1;ZavršnicaValley, slopeofStol,UTMVM34,NMWK29758/2+1 (juv.),38367/1; slopeofStol,ZavršnicaValley,N-slopeofPecit,650-700m,UTMVM34,leg.KäuFel,NMWE44525/2;Loiblpass3kmSfromtheborderofAustria,UTMVM44,leg.MaaSSen5.1984,Maa/3;„Zarica“SaveGorgebelowKranj,UTMVM51,(exKuščEr) NMWE15996/3;Duplica,UTMVM61,(exKuščEr),NMWE48518/1; rockson the leftbankofBistricaNofStahovica,UTMVM62,NMWE10654/2+4 (juv.); 2kmNofGrad(=11.5kmENEofKranj),550malt.,UTMVM62,leg.git-tenberger24.7.1981,NNM/1; rockwallat the roadStahovi-ca-Bistricakoča,UTMapp.VM62/72,NMWE14271/2+3(juv.); Grintovca, Kankar Saddle, UTMVM 63, NMWK38370/2;Jezersko,20kmNEofKranj,950malt.,UTMVM63,leg.de Winter29.9.1980,NNM/4;neighbourhoodofJez-ersko,UTMVM63,leg.MaaSSen9.1980,Maa/22(partlydam-aged);Grintavec-slope inKankarSaddle,200mbelowZois-hut,UTMVM63,NMWE10632/2;KamniškaBelaValley13kmNofKamnik (=NWofVelikaPlanina), 600malt.,UTMVM63, leg.gittenberger23.7.1981,NNM/1;1.5kmWof„DomvKamniškiBistrici“,12kmNofKamnik,610malt.,UTMVM63, leg.gittenberger 23.6.1981,NNM/11;grottosofMoräutsch,UTMVM71, (exStuSSiner, coll.O.boettger)SMF51775/3;entranceofgrottosnearAich,UTMVM71,(exrobič1879,exJetSchin),SMF110481/3;NWVe-likaPlaninain„DolskiGraben“,875malt.,UTMVM72,leg.gittenberger9.1970,NNM/1;rocksabovetheeasternBistricaSource,UTMVM73,NMWE10686/2;between„FrisaufovDom“and„Logarskadolina“,1350malt.,UTMVM73,leg.gittenberger 26.8.1971,NNM/2; „Savinjal. solcav“,UTMVM73/74,NMWK38373/2; „Naplov savinjepriSolčavi“



(=Solčava),UTMVM74,(exKuščer)NMWE48675/4;en-tranceofLisičnacavenearLuče,UTMVM83, (exKuščer) NMWE49095/2.

Croatia: Istria,Režanci,cave,UTMVK18, leg.Kovačić7.8.1980,CNHM/1;Istria,Motovun,debrisoftheMirna,UTMVL02,leg.MaaSSen5.1991,Maa/3(juv.);Istria,UčkaMoun-tain(=Mte.Maggiore),rocksimsmalldolinasbelowrefuge,UTMVL31,leg.1927,NMWE21351/1;UčkaMountainnearPoklon,970malt.,UTMVL31,leg.MaaSSen9.1982,Maa/1(juv.);UčkaMountainwesternsidenearVeprinac,350malt.,UTMVL42, leg.MaaSSen 9.1982,Maa/1+1 (juv.); forestabove Ičići (=betweenPoljaneandVeprinac), at limestonerocks,UTMVL42, leg.Subai 4.9.1986,S13157/1 (dam-aged);3kmWofGornjeJelenje(=NEofRijeka),UTMVL72, leg.MaaSSen 9.1982,Maa/1 (juv.);Delnice,UTMVL82, leg.atteMS 8.10.1907,NMW46132/1;DinaraPlateaunearKijevoSofProtoztop,app.1000malt,UTMXJ07/17,NMWE16379/3+1(juv.),NMWK38371/3;Vrlika,UTMXJ16,leg.MaaSSen8.1986,Maa/1;castleruininVrlika,atlime-stonerocks,UTMXJ16,leg.Subai 30.8.1986,S12188/3+1(juv.);KrupaValley(=SofGračac),UTMWJ69/79,NMWE33884/1+1 (juv.); caveMramornica near Otočac, UTMWK16, (exKuScer),NMWE49905/1;Brušane (=betweenKarlobagandGospić),UTMWK22, leg.MaaSSen8.1986,Maa/1;southernsideofBrušane,debrisofBrušankaStream,UTMWK22, leg.Subai 3.9.1986,S13170/2;Lika-Region,Tomašićevakuća,betweenPerušić andLičkoLešće,UTMapp.WK24/25/34,leg.štaMol28.6.1981,CNHM/1+1(juv.),1 (juv.), 5+9 (juv./damaged);Plitvice lakes, southern sideofPlitvičkiLjeskovac,UTMWK46, leg.verMeulen1988,NNM/6;Plitvicelakes,UTMWK46/47,leg.verMeulen1988,NNM/6;Plitvice(=WofBihać),limestonerocksatalargewa-terfall,UTMWK47,leg. Pintér, Subai & Szigethy 6.8.1972,HNHM/1,S2706/1 (damaged);NationalParkPlitvice,nearentrance2,UTMWK47, leg.MaaSSen 9.1985,Maa/1+3(juv.);Plitvice,40maboveMilanovacLake,UTMWK47,leg.MaaSSen9.1975,Maa/5;Plitvice,„MilkaTrnina“cavenearGavanovacLake,UTMWK47,leg.MaaSSen9.1975,Maa/1(damaged);Devčićevac,PlitviceLakes,UTMWK47, leg.Sturany6.1895,NMW23490/1+2 (juv.);2kmNofJošan,650malt.,UTMWK53,leg.MaaSSen5.1983,Maa/10(partlydamaged);Čerovačkacave4kmSSEofGračac,UTMWK60, leg.MaaSSen9.1988,Maa/3+1(damaged);Presika(=NofVrbovsko)600malt.,UTMWL02, leg.MaaSSen9.1982,Maa/13;Lukovdol,UTMWL13, leg.hirc,CNHM682/1;(exhirc,exMaceK)CNHM683/2+1(juv.);Kordun-Region,MikašinovićinearGornjeDubrave,closeZalacaves,UTMWL21,leg.Jalžić13.3.1983,CNHM/1;southernCroatia,Kapela,UTMn.det.,(exreitter,coll.O.boettger)SMF51771/1.

Bosnia and Hercegovina: Karanovac, UTM BQ 85,leg.nordSiecK3.8.1966,SMF205796/1; Ivan [=mountainWSWofSarajevo],UTMBP54/55,64/65, leg.aPFelbecK,NMW32350/2;Preslica-wallsagainstBradina (=Sof Ivan),UTMBP54/64,NMWE15926/3 (juv./damaged),NMWK38375/1;VreloBosna (=Wof IlidžanearSarajevo),UTMBP85, leg.MaaSSen7.1980,Maa/4+4 (juv./damaged); leg.MaaSSen5.1984,Maa/2;Željeznica-debrisnear Ilidža,UTMBP85, leg.Winneguth,NMW62147/1+1 (juv.);Miljacka-debrisnearSarajevo,UTMapp.BP85/95, leg.Winneguth,NMWE20834/6;leg.KuščEr,NMWK33619/2;(exKuščEr) NMWE49638/2;ReljevonearSarajevo,UTMBP86,NMWK38369/2 (juv./damaged),38374/1; (exKuščEr,exWinnegut) NMWE49373/4(damaged,without teeth);2kmSofDobro

Polje,alongsideofDobropoljankaStream,UTMBP92, leg.MaaSSen5.1984,Maa/37+6(juv./damaged);3kmEofDobroPolje (=35KmSSEofSarajevoalongsideofDobropoljankaStream,800malt.,UTMBP92, leg.MaaSSen 17.9.1980,NNM/6+1(damaged);4.5kmNofTrnovo,atrocksalongastream,800malt.,UTMBP94,leg.MaaSSen5.1984,Maa/2(juv.);hillslope11kmNEofMiljevina(=countryroadSara-jevo-Foča),alongsideofRijekaBistrica,800malt.,UTMCP02, leg.de Winter 18.9.1980,NNM/1+4 (juv./damaged);15kmoutsideMiljevina (=SofDobroPolje), alongsideofDobropoljankaStream,UTMCP02, leg.MaaSSen 9.1980,Maa/37+6 (juv./damaged); JahorinaPale (=EofSarajevo),at limestonerocksbesideofMiljacka-river,900malt.,UTMCP05,leg.Subai 17.8.1987,S16842/1;PračaValley,2kmSofKukavice (=±25kmESEofVišegrad),475malt.,UTMCP34, leg.MaaSSen&de Winter 27.9.1980,NNM/8+8(juv./damaged);NofUstiprača imPračaValley,500malt.,UTMCP44,leg.MaaSSen9.1980,Maa/11;MountainStolac,borderofBosniaandSerbia,UTMCP66,leg.Penther1905,NMW41080/1;10kmSofVinac(=SofJajce),UTMXJ89,leg.MaaSSen9.1988,Maa/1;3.4kmoutsideČađavica (=be-tweenKljučandMrkonjićGrad),UTMXK52,leg.MaaSSen &de Winter16.9.1980,Maa/2+3(damaged),NNM/1(dam-aged);22.1kmfromJajcetowardsBanjaLuka(=SofjunctiontowardsMrkonjićGrad), at limestone rocks,UTMXK72,leg.Subai 3.8.1987,S16813/2+2 (juv.);VrbasbanksnearKrupanaVrbasu,UTMXK74,leg.MaaSSen5.1984,Maa/14;VrbasValleyatS-marginofBanjaLuka,deciduousforestwithlimestonerocks,UTMXK75,leg.Subai3.8.1987,S16902/1;rocksontheriverbanknearJajce,UTMXK81,leg.MaaSSen 9.1980,Maa/1;Jajce,nearthebridge,UTMXK81,leg.MaaS-Sen9.1988,Maa/1;Jablanica→RamaValley,UTMYJ14/23,leg.nordSiecK 12.8.1966,SMF205801/1;mountain slope10.8kmfromJerezaDubrovicatowardsJablanica[=betweenJablanicaandMostar],atlimestonerocks,UTMYJ21/22,leg.Subai 3.8.1987,S16939/20+29(damaged);[Mountain]Plasanear Jablanica,UTMYJ23,NMWK16189/1; leg.Penther 1900,NMW33565/1+2(juv.);MountainVlašićnearTravnik,1500malt.,UTMYK10,leg.MaaSSen5.1984,Maa/3;sourcebelowVreloKnijeginjac nearSarajevo,UTMn. det., leg.Schütt8.9.1959,Sch158/2.

Serbia:NoviPazar,gorgeatpath tomonasterySopoćani,UTMDN57, leg.Schütt1.6.1960,NMWK45832/2 (dam-aged),Sch192/1;Užice,UTMDP05, (exKuščEr)NMWE49123/1;ZlatiborMountains,1kmSofUžice,UTMDP05,leg.Erőss,Fehér&KovácS 1.4.2001, coll.Erőss 7075/1;2kmSofUžice (=Užice-NovaVaroš country road), app.470malt.,UTMDP05,leg.Erőss,Fehér&KovácS6.4.2001,hnhM 85673/1;VreloRaškenearUžice,UTMapp.DP05,(exKuščEr) NMWE 49416/2+1 (damaged); „tudiVreloRaške“,UTMapp.DP05, (exKuščer)NMWE49903/1+1(damaged),NMWK38365/1.

Kosovo,RugovskaGorge2.5kmWofKućište,1300malt.,UTMDN22,leg.MaaSSen9.1980,Maa/1(juv.);Rugov-skaGorge5kmWofPeć,UTMDN32,leg.MaaSSen6.1987,Maa/4.

Montenegro:1kmfrom thebifurcationoncountry roadtowardsMratinjenearPlužine,UTMCN29, leg.MaaSSen 9.1980,Maa/5;Durmitor [Mountains],UTMCN37/47, leg.Dabović1936,NMWE21620/2;RijekaBijelaSofŠavnik,600malt.,UTMCN45, leg.MaaSSen9.1980,Maa/2;4kmSofKruševice(=betweenNikšićandŠavnik),UTMCN45,leg.MaaSSen9.1980,Maa/2;alongthecountryroadbetween



NikšićandŠavnik,8KmSofMokroKale,1450malt.,UTMCN45, leg.de Winter 20.9.1980,NNM/1 (juv./damaged);CrnoLakenearŽabljak,UTMCN47,1450malt.,leg.MaaS-Sen9.1980,Maa/4;BistricainTaraValley,UTMCN76,leg.MaaSSen 7.1979,Maa/3+2 (damaged);BiogradskoLake,UTMCN85,1200malt.,leg.MaaSSen6.1987,Maa/1(juv.);nearMotelinSjenogošte(=NofKolašin),UTMCN85,950malt., leg.MaaSSen9.1980,Maa/2; leg.de Winter23.9.1980,NNM/2+1 (juv.);5kmSofZatonalong the roadIvangrad-VrapčePolje,650malt.,UTMDN05,leg.MaaSSen9.1980,Maa/11;betweenRibarićeandRožaje (=20kmfromthebi-furcationatTutin),limestonerocks,UTMDN34,leg.Pintér, Subai & Szigethy 19.7.1972,S2707/10+1 (damaged); IbarValley3kmNofRožaje (=SofNoviPazar), 1000malt.,UTMDN34,leg.MaaSSen9.1980,Maa/2.

Macedonia: rocks longsideof theDrim-river3kmNofLukovo,UTMDL68, leg.MaaSSen 5.1975,Maa/5;Ohrid,UTMDL85, (exKuščEr)NMWE49046/1+2 (juv./dam-aged);outsideofVratnica,UTMEM16,(exKuščEr),NMWE48921/1;sourceatRašće(=WofSkopje),UTMEM25,(exKuščEr)NMWE48171/1+2(juv./damaged).

Albania:PeriferiMalësia,amountainpass2kmNofRra-ps, along the road fromHai iHotit toVermosh, limestonerocks,UTMCM79, leg.Erőss &Fehér 30.6.1996, hnhM 90976/1;PeriferiShkodër,WofShllak(=18kmfromtheMesbridge),1020malt.,limestonerocks,UTMCM96,leg.Erőss,Fehér, hunyadi&Murányi 16.4.2006,HNHM/1;PeriferiElbasan,SofGurriiZi,13kmfromtheElbasanjunctionontheroadtoQafaeShtyllës,limestonerocks,900malt.,UTMDL26, leg.Erőss,Fehér, hunyadi&Murányi 10.4.2006.,HNHM/2(alk.)+2(juv.,alk.);PeriferiTiranë,6kmSofQafaeShtylles,along the roadfromTiranë toKlos,karstplateau,limestonerocks,1420malt.,UTMDL27,leg.Erőss,Fehér, KontSchán&Murányi 22.10.2002, hnhM 91699/1;PeriferiPogradec, Shpellë(=4kmSSEofBishnicë),ShkembiQytetit,limestoneandconglomeraterocks,1140malt.,UTMDL52,leg.Erőss,Fehér, KontSchán&Murányi1.7.2003,HNHM93070/1;Qukës-Shkumbin,over thequarry in the left sideof theLumi iShkumbinit, limestonerocks,500malt.,UTMDL54, leg.Erőss,Fehér, KontSchán&Murányi30.6.2003,HNHM93055/1.

Greece:Macedonia,GrámmosMountain,slopetothelakeon theE-sideofmountEpánoAréna(=NWofPefkófito),atlimestonerocksandunderstones,1900malt.,UTMDK96,leg.Subai 15.7.1990,S16383/2.

Ípiros,app.38kmSofIoannina(=51kmalongthecountryroad), rocksWofcountry roadnearpaleoliticcaves,150malt.,UTMDJ84, leg.e. & e. J. gittenberger 31.5.1991,NNM/1 (damaged);Loutrá (=5kmNofAmárantos) lime-stonewesternhillside,1270malt.,UTMDK74,leg.riedel & Subai 12.4.1988,S14906/65+17(juv./damaged);LoutránearAmárantos,atN-exposed limestonerocksand inwoodsapp.500mSEofLoutrá,1250malt.,UTMDK74, leg. Subai & SzeKereS 16.5.1997,S17057/34+6(damaged);Zagoria,Kipi,limestonerocksnearoldbridge,UTMDK81,leg.SattMann 21.8.1990,NMW102816/4; rocksbetweenMikraPeristeriandMegaPeristeri,UTMEJ09, leg.reiSchütz7/87,NMW85654/2+1(juv).

R e m a r k s :Pupa truncatellawasdescribedbyl. PFeiFFer (1841:46)without illustrations.Hisspecifica-tionofthelocustypicusis:„...monteKarstpropeCastelnuovo, interTergestumetFiume“.Castelnouvo is the

formernameofPodgrad.Thesmalltownislocatedbe-tweenTriest (=Tergestum)andRijeka (=Fiume).The“smallspecies(=/truncatella/)”mentionedinPFeiFFer’soriginaldescriptionisfoundthere.

roSSMäSSler(1842:12)recapitulatedPFeiFFer’slocal-itiesandaddedarecordbySchMidtfromnearLjubljana(=Laibach);additionally,he illustrated two“varieties”hereceivedfromFrivaldSzKy“fromTurkey”.Itisonlyhisfigure733thatshowsA. truncatella.

KüSter (1842: 34) referred to both,PFeiFFer androSSMäSSler, repeated their statements and addedasnew locality“lowerDalmatia (neuMeier)“andfiguredthespecimensonhisplate4Fig.20–25.Itisimpossibleto identify thespecies illustrated inFigs.20–23.SinceKüStercharacterizedtheapertureas“...withoutfoldingsand teeth…”and thedrawing showsadense ribbingpattern it is likely that thefirstpictureshows the laterdescribedA. skipetarica,which in fact lives in lowerDalmatia.Butit isalsopossiblethathehadthetrueA. truncatella beforehim,butwasnotable to recognizethetinyteeth.The“varieties”(figs.24–25)clearlyshowthatspecieswhichwaslaterdescribedbyPFeiFFer(1848:331)underthenamePupa parreyssii(avalidtaxonliv-inginBulgaria).However,PFeiFFer(1848:303)suppliedamodifieddescriptionofA. truncatella repeating theinaccurate informationabout the locus typicus.Addi-tionally,hedescribedavarietyβunder thenamePupa formosa(seethere).

ItwasnotpossibletoidentifythespeciesPetrboKhadathandwhennaminghis“Agardhia truncatella var. depressa”(1939:344,footnote).Hisdescriptionisnotsuf-ficientenoughtodifferentiate depressafrom“Agardhia truncatella biarmata for. minor”whichlivesatthesamespots.Verylikely,bothtaxadescribethesmall,toothlessformofA. truncatella fromnorthernBosniasincebothoriginatefromČebič,aplacewhichcouldnotbelocated(seealsobiarmata).

Va r i a b i l i t y :A. truncatellaismoderatelyvariableconcerningshell sizeandshape, thepatternof ribbingandtheformationoftheaperturalteeth.MaaSSen(1985:4)describedaninterestingformofA. truncatellalivinginnorthernBosnia(Karanovac,VrbasValleynearBanjaLuka and10kmSofVinac). It is smaller andmoreslenderthantheusualformofA. truncatella,withlack-ingaperturalteeth(seeFig.2).NearKrupanaVrbasuintheVrbasValley,thepopulationislargerandwider,butsill the teetharemissing.Finally, the formnearVreloBosna (=SWofSarajevo) is slenderandonlyslightlysmallerthantheusualformwithhardlyrecognizablebutexistingteethandaveryfaint,hardlyelevatedkeel-likeparietal ridge.Thespecimen from themost southwardhabitatinGreece(app.38kmSofIoannina,fig.3.)hasapoorlydevelopedpalatal toothandan inconspicuousparietallamella.Allothercharactersaresharedwiththeusualform.NearKipiattheTurkishbridgeinZagoria,largespecimenswere found (seefig.4.).Ata localitybetweenMikra-andMegaPeristeri, twospecimensof



A. truncatellawerefound.Oneofthemshowsacoarseandwidelyspacedmodeofribbingwithanindicationofaninfraparietalis,whilethesecondspecimensisdenselyribbedwithoutaninfraparietalis.

AllthesedifferencesrecordedhereareinterpretedasindividualvariationsincealltheshellshithertoexaminedsharetheremainingcharactersofA. truncatella.

D i s t r i b u t i o n :This species isknown fromnearUpperAustriasouthofSalzburgtoSteyrareathroughoutAustria, north-eastern Italy,westernSlovenia, south-westernCroatia,BosniaandHercegovina,Montenegro,thewesternborderofSerbiaandKosovo,westernMac-edonia,AlbaniatonorthwesternGreece.Itclearlyisaninlandspecieswhichdoesnot live in themountainsoftheAdriaticcoastsoutheastofKvarnerBaytoGreece.

Agardhiella biarmata (o. boettger1880)Plate1,Figs.5–6, textfigs.4–6

1880 Pupa (Sphyradium) truncatella var. biarmataO.boett-ger, BerichtdesOffenbacherVereinsfürNaturkunde,1880:109[locustypicus:“...beiRagusaundbeiPrid-worjegesiebt”; lectotypusdesign.zilch 1985:SMF4658].

1887 Pupa (Coryna) truncatella var. biarmata,–WeSter-lund, FaunaderinderpaläarctischenRegionlebendenBinnenconchylien,3:88.

1890 Pupa (Coryna) biarmata,–Flach, Verhandlungenderphysikalisch-medizinischenGesellschaftzuWürzburg,(2)24:53.

1906 Coryna biarmata spelaea Kobelt, IconographiederLand-&Süßwasser-Mollusken,(2)12(1/6):23,Plate.311Fig.1979[locustypicus:“...ineinerHöhlebeiZa-valainderHerzegowina,...”,syntypes:NMW,SMF].

1915 Agardhia (Agardhia) truncatella biarmata, – Sturany & Wagner,DenkschriftenderKaiserlichenAkademiederWissenschafteninWien,mathematische-naturwis-senschaftlicheKlasse,91:65,Plate.18Fig.104a–c.

1926 Agardhia (Agardhiella) truncatella biarmata,–PilSbry, ManualofConchology,27:164,Plate.19Fig.9–14.

1939 Agardhia truncatella biarmata for. minor PetrboK, – VěstnikČs.zoologickéspolečnostivpraze,6/7:344[locus typicus: “...in the rockycleftsof theKarst atČebič,...”;typematerial:unknown].

1975 Agardhiella formosa, –gittenberger, ZoologischeMededelingen,48 (24):284,Fig.1 (shells) [partim,non formosa l. PFeiFFer1848].

1985 Agardhiella biarmata, – zilch, Archiv für Mol-luskenkunde,116(1/3):128,Plate2,Fig.19.

1985 Agardhiella formosa,–MaaSSen, DeKreukel,21 (1–2):4,distributionmap[partim,nonformosa l. PFeiFFer 1848].

D i a g n o s i s :shellsmall,aperturesmallwithangu-laris,parietallamellaandinfraparietalis.Stronglydevel-opedpalataltooth,sometimeswithcolumellarteeth.

D e s c r i p t i o n : shell relatively small, cylindri-calwitha roundedapex.After ca.1.75–1.95smooth,slightlygranulatedwhorls, a rapidly increasing radialribsculpturebegins.On the lastwhorl thereare35–55

(9–14/mm) ribs.Between themain ribs therearefine,irregularradialribswhichcanonlyberecognizedunderhighmagnifications.

Whorls6–7; the lastwhorloccupiesmore than thehalfofthetotalheight.Theumbilicusisnarrowcylindri-cal.

Aperturesmall,withan infraparietalisandanangu-laris. Ingeneral,both teetharepoorlydevelopedandstretchstraighttowardstheinterioroftheshell;aparietallamella ispresentaswellwhichoftenonlyreaches theheightofthemarginalteeth,butitssizeincreasesintheinteriorof theshell. In theupperpartof thecolumella,therearesometimestwocolumellarteeth,andatthebot-tomabasicolumellaris.Thesecolumellarteethareoftenveryweakormissingatall.Infrontalview,theaperturalrimisu-shaped.Interiorly,itisstrengthenedbyaslightlipandflaredtowardstheoutside.Intheupperpartofthecolumellarsideitisslightlybenttowardstheumbilicus.Thecentreofthepalatalsideoftheapertureisimpressedbyastrongtooth-likecallus.Theupperpalatalapertureisslightlybulgedwithoutalip.Sometimes,averyshortsinulusisdevelopedabovetheangularis.Theperistomi-alrimsareconnectedbyawelldevelopedcalluswhichalsocoverstheribsabovetheaperture.Insideview,theupperpartoftheaperturalrimisslightlyprotruding.

M e a s u r e m e n t s (n=23): H=3.18–3.93;D=1.43–1.75;AH=1.19–1.37;AD=0.75–0.87.

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 (after2spec-imenofTrebinje,Petrina,Matulicacave):Theretractormuscleof the right eye-stalk runsbetweenpenis andvagina;proximalhalfof thepenis slender, almost aslongastheslightlyswollendistalhalfofthepenis.Theappendix insertscentrallybetween the twopenisparts;thecentralpartisnarrow,thebasalandtheterminalpartsreach two to three timesof itsdiameter; theappendixretractormuscle inserts at thewidenedproximalpartof theappendix.Theepiphallus reachesabouthalf thelengthofthepenis;inoneofthespecimensdissected,thedistalepiphalluswasconstrictedbehindaslightswell-ing,while in theother specimen theepiphalluspassedsmoothlytothevasdeferens.Theoviductislongerthanthebroadvagina.Thepedunculusofbursacopulatrixismoderatelylongandinsertsataslightlyswollenpointatthedistalpartofthevagina.

D i f f e r e n t i a l d i a g n o s i s : Its closest relative,A. truncatella, hasasomewhatwideraperturewithonlytwomarginalparietal teeth. Italways lacks thecolum-ellar teethand theparietal lamella. Itsparietalwall isslightlyenlargedbyabarrencallusandnot indentateasinA. biarmata.InA. truncatella, theaperturalrimismorestrengthenedbasally,butthecolumellarsideisnotasenlargedasinA. biarmata.

InA. formosa, theaperture is largerandwider; therib sculptureof its teleoconchwithcoarserandwiderribs.This species is similar in thedevelopmentof theapertural teeth,but theparietal lamellaalreadystartsattheparietalcallus.Thelowerpartoftheapertureismore



stronglyenlarged,butapalataltoothisalwaysmissing.InA. formosa,theparietalwallisonlyslightlyenlarged.Sofar,nopalatalteethhavebeenfound.

T y p e s :Croatia,Dubrovnik(=Dalmatien,Ragusa),UTMBN52/62,(coll.O.BOETTGERexE.REITTER)lectotypebiarmata(design.zilch):SMF4658/1;Pridvorje,UTMBN81,(coll.O.BOETTGERexE.REITTER)paratypesbiarmata:SMF4571/1+2(juv.);cavenearZavala,UTMYH44,leg.Pa-ganetti1903,syntypesbiarmata spelaea:NMW39555/4+1(juv.);exSturany1904,(coll.o. boettger)SMF51691/1.

A d d i t i o n a l m a t e r i a l e x a m i n e d :Bosnia and Hercegovina:smallcavenearBihovonearTrebinje,UTMBN72,(excoll.ehrMann),leg.abSolon25.7.1912,SMF51692/5;leg.deeleMan-reinhold21.6.1975,NNM/1(damaged);cavenearTrebinje,UTMBN83, leg.abSolon,NMW50236/3(juv.);„cave3e“nearTrebinje,UTMBN83,NMW68444/1;

(exKäuFel)NMWK16200/1;Trebinje,Petrina,Matulicacave,UTMapp.BN83,leg.Pavić8.9.2006,HNHM/3(alk.);VelikacavenearTrebinje,UTMBN83,(exKuščEr)NMWE48720/2;TrebišnjicaSource nearBileća,UTMBN84, leg.Schütt 8.9.1967,Sch427/2(damaged);ČepelicaSourcenearBileća,UTMBN84,leg.Schütt14.9.1959,Sch140b/2(damaged);BunanearMostar,UTMYH39,(exKuščEr)NMWE48923/1(damaged);Zavala,UTMYH44,leg.MaaSSen7.1982,Maa/2;Bjelušica,smallcavenearZavala(Popovopolje),UTMYH44,(exedlauer16438),Sch/1 (damaged); (exKuščEr)NMWE48721/5;Zavala,smallcavenearmonastery,UTMYH44,leg.edlauer,NMWK47250/5,2941/1;uppermostrockyfrontofOstrogNofZavala,UTMYH44,NMWE16417/1;districtTrebinje,Djurkovinacave15minutesNEof„Gensdame ricgrebci“,UTMn.det.,leg.reMy27.8.1936,NMWE48275/1.

Croatia: cave „špilja za Gromačkom vlakom“ nearGromačanearDubrovnik,UTMBN53,leg.Jalžić, 6.3.1998,

Textfig.4.Agardhiella biarmata (o. boettger1880).Croatia,IslandKorčula,mountainslope2.8kmfromČaratowardsKorčula-City,atlimestonerocks,UTMXH55,(H=3.75,D=1.68mm).



CNHM/1+3 (juv.); 1kmE ofOsojnik (=8kmN ofDu-brovnik),450malt.,UTMBN63,leg.gittenberger8.5.1986,NNM/1;Pridvorje,UTMBN81,(exO.boettger1904)NMW39286/1;Konavle Region, Sv. StjepanChapel nearDubaKonavoska,UTMBN82, leg. Jalžić,KlETEčKi&štaMol 23.11.2003,CNHM/1;atafountatsouthernsideofSlano,5malt.,UTMYH34, leg.gittenberger 6.5.1986,NNM/2+1(juv.).

PeninsulaPelješac, southeasternmountain slopeSv. Ili-janearOrebić,200–650malt.,UTMXH76, leg.MaaSSen 10.1978,Maa/1+1,S5240/1;Kučište,150malt.,UTMXH76, leg.MaaSSen5.1984,Maa/1;Zagorje-Gebirge,MountainČarovićN-side600mofZaradežeindirectionOrebić,atlime-stone rocks,370malt.,UTMYH04, leg.Subai 6.8.1987,S13593/2+6(juv./damaged);2kmSWofSton,UTMYH14,leg.gittenberger6.5.1986,NNM/1;nearPrapatna (=3kmSWofSton,17kmWNWofSlano),UTMYH14, leg.git-tenberger6.5.1986,NNM/2;mountainslopeatwesternsideofSparagovići(=Ponikve),atlimestonerocks,450malt.,UTMYH14,leg.Subai 6.8.1987,S13643/1+2(juv.).

IslandKorčula,mountainslope2.8kmofČaraindirectiontoKorčula-City,at limestonerocks,UTMXH55, leg.Subai 8.8.1987,S13632/3+3(juv.);4.3kmofPupnatindirectionofPupnatskaLuka,atlimestonerocks,170malt.,UTMXH65,leg.Subai 8.8.1987,S13596/2+1(juv.).

IslandMljet,grotto200mEofSobra,40malt.,UTMYH13,leg.Pintér & Szigethy 2.8.1972, hnhM/1, S 2705/1.

Montenegro:Northwestern sideofMokrine (=NNWofHercegNovi), at limestone rocks,520malt.,UTMBN90,leg.Subai19.9.2005,S20673/8+4 (juv./damaged);decidu-ousforest4.2kmNWofMokrine(=indirectionofTrebinje,NWofHercegNovi), at limestone rocks,770malt.,UTMBN91, leg.Subai13.10.2004,S20537/1+1 (juv.); [mount]SokolnearDupilovillage,UTMCM47,leg.Dabović,NMWE49398/2;1kmfromVilusetowardsofTrebinje,inaforestonmountSkorča,at limestonerocks,1000malt.,UTMCN03,leg.Subai 12.8.1987,S16903/4+1(juv.);Njeguši,UTMCN20,NMWK38384/1(large!);(exFuchS)NMWE35216/2+2(juv./damaged), 35217/1+2 (juv)+1 (formosa!); PopovacavenearNjeguši,UTMCN20,NMWKwithoutNr./1; leg.Dabović,NMWE18273/1,49341/1+1(juv,damaged);Buko-vicaRivernearŠavnik,UTMCN45,NMWK38360/3.

U n i d e n t i f i e d l o c a l i t y :cave„pećinaMaricaVuk-sanovica“,UTMn.det.,leg.Dabović,NMWE49224/1(dam-aged).

R em a r k s :Intheoriginaldescriptionaswellasinsubsequentpublicationsofothermalacologists,A. biarmatawasrecognisedasasubspeciesofA. truncatella. However, it isawellcharacterisedspecieswitha rela-tivelysmalldistributionarea.

PetrboK (1939)described thevariabilityof “Agardhia truncatella”includingthetwonewtaxa“Agardhia

Textfig.5a–b.Agardhiella biarmata (o. boettger1880)situsofthegenitalorgans.BosniaandHercegovina,Trebinje,Petrina,Matulicacave,UTMapp.BN83.—Abbreviationstothedrawingsofgenitalorgans:A=appendix,B=bursa,E=epiphallus,O=oviductus,P1=proximalpartofpenis,P2=distalpartofpenis,Pd=pedunculus,R=retractormuscle,V=vagina,Vd=vasdeferens.



truncatella var. depressa“ and „Agardhia truncatella biarmatafor. minor“(bothonpage344inthefootnotes).TheillustrationsofA. biarmata(2.and3.rowonpage343)withvariouspicturesofapertural teethcontradicthisstatementthat„biarmata for. minor“istoothless.Hissomewhatconfusingidentificationkey(p.340)doesalsonothelp.Untilnow,thetypelocality“Čebič”couldnotbe identified, althoughdetailedmapswereconsulted.Theonly similar-sounding locality name isČepić inNorthern Istria betweenBuje andBuzet,which is inthedistributionareaofA. truncatella.Moreover, thetypespecimensofbothtaxacouldnotbeentraced.Con-cluding,itiscurrentlyimpossibletodecidewhetherthespecimensPetrboKhadathandbelongedtothetoothlessformofA. truncatellafromnorthernBosniaorinfacttoA. biarmata.

In thepast,A. biarmatawasconfusedwith thege-ographicallyneighbouringA. formosa (gittenberger 1975:284,MaaSSen1985:4).Mostlikelythisisduetothesimilaritiesinthedevelopmentoftheaperturalteeth.NearMokrine,A. biarmata andA. formosahavebeenfound sympatricwithout any transitional forms, andalsofromtheNjegušibothspeciesareknown(NMWE35216adv.35217)without anyoverlap in characters.However, it has to bementioned that fromNjeguši(NMWK38384/1)thereisonespecimenofA. biarmata withanextremelylargeshell.ItapproachestheaveragesizeofA. formosaandhasaweakerpalatal tooth.Butinallothercharactersas theroundedwhorls, thesmallaperture,onlyslightlyenlargedaperturalrimatthebaseand thedevelopmentof theparietal lamella, thisspeci-menistypicalfor A. biarmata.

Similaras inA. truncatella,A. biarmata is found incavesandintheduffofrocks.Thisseparatesbothspe-ciesfromA. formosa,whichlivesexclusivelysubterra-neousandisonlyfoundinrockcavities.

D i s t r i b u t i o n :ThedistributionofA. biarmata reachestheIslandofKorčula,thepeninsulaPelješacandtheCroatiancoastsoutheastoftheNeretvaValley.Addi-tionally,ithasbeenfoundinthemountainsofthesouth-easternpartofBosniaandHercegovinaand inwesterncoastalpartsofMontenegro.

Agardhiella stenostoma (Flach1890)Plate1,Fig.10;textfigs.7,8.

1890 Pupa (Coryna) stenostoma Flach, Verhandlungenderphysikalisch-medizinischenGesellschaftzuWürzburg,(2)24:53[locustypicus:„Dalmatia“;holotypeSMF4597].

1926 Agardhia (Agardhiella) stenostoma,–PilSbry, ManualofConchology,27:162.

1958 Argna (Agardhiella) stenostoma, – zilch, Archiv fürMolluskenkunde,87(4/6):150,Plate11,Fig.1a,Plate12,Fig.1.

1975 Agardhiella stenostoma, – gittenberger, ZoologischeMededelingen,48(24):285,Fig.3.

1985 Agardhiella stenostoma, – zilch, Archiv fürMol-luskenkunde,116(1/3):129,Plate2,Fig.17.

1985 Agardhiella stenostoma,–MaaSSen, DeKreukel,21 (1–2):4,distributionmap.

D i a g n o s i s :mediumsized shells, aperturewithinfraparietalis,angularisandbentparietallamella;strongvertical columellar lamella reaching theapertural rimandstronglyimpressedlongitudinalpalataltooth.

Textfig.6.DistributionmapofA. biarmata.



D e s c r i p t i o n : Shell medium-sized, cylindri-calwitha roundedapex.Thefirst1.75whorlssmooth,slightlygranulated, followedbyradial ribsculptureontheprecedingwhorls; lastwhorlwith54–73 (13–14/mm)hardlyelevated ribs;between themainribs, therearefine,irregularradialribswhichcanonlybeseenun-derhighmagnification.

Whorls5.75–6.25; the lastwhorlup to52–58%ofthetotalheight;umbilicusnarrow,falciform,partlycov-eredbythelastwhorl.

Aperture longand slender,withweeklydevelopedinfraparietalisandangular teeth runningstraight to theinteriorof the shell;between themaparietal lamella,reachingtwotothreetimestheheightoftheteeth;itispointedinfrontandprojectsabovetheleveloftheaper-

ture;itiscurved,narrowedinthemiddlebutsomewhatbroaderatitsendinsidetheaperture.Deepintheaper-ture,anelongate,strong,verticalcolumellarisisvisible.Aperturalrimu-shapedinfrontalview;atthecolumellaborderedbyavertical ridge;flaredat thebaseand thepalatalside; lowerpartof theapertureconstrictedbyastrong tooth-likevertical ridge;above this theapertureis recessed, and the apertural rimnarrowed; in somespecimens,ashortsinuluscanberecognisedabovetheangularis.Theperistomialrimsconnectedbyawellrec-ognisablecallusconcealingtheribsofthelastwhorl.Insideview,theupperpartoftheaperturalrimisslightlyprotruding,andtheupperthirddiagonallytruncated.

M e a s u r e m e n t s (n=19): H=3.62–4.06;D=1.37–1.56;AH=1.25–1.37;AD=0.68–0.87.

Textfig.7.Agardhiella stenostoma (Flach1890).Croatia,graveyardofKomolacnearDubrovnik,atlimestonerockbeneaththechapel,UTMBN62,(H=3.93,D=1.43mm).



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 :unknown.D i f f e r e n t i a l d i a g n o s i s : In general, A.

zoltanorum ishigherandwider than A. stenostoma. Itsribsarehigherandmorewidelyspacedandithasanal-mostcompletelyopenumbilicus;aperturemorestrongerconstricted in thebasalpart.Moreover,A. zoltanorum hasawelldevelopedbasaltoothbeneaththecolumellarlamella.

InA. formosa,theshellisalsohigherandwiderandtheribsaremorewidelyspaced than inA. stenostoma. Itsapertureiswiderandnotconstricted,theparietalla-mellaislowerandlesscurved,bluntanddoesnotprojectovertheaperturallevel;thecolumellarisismissing,thepalataltoothissmalltoalmostreduced.

Ty p e s : “Dalmatia”,holotypeSMF4597(coll.o. boett-ger).

A d d i t i o n a l m a t e r i a l e x a m i n e d :Croatia: sur-roundingsof„campingSolitude“nearLapadnearDubrovnik,UTMBN52, leg.MaaSSen 9.1988,Maa/2+1 (damaged);leg.MaaSSen4.1989,Maa/1;graveyardofKomolacnearDu-brovnik,at limestone rockbeneath thechapel,UTMBN62,leg.Schütt 8.6.1978,Sch868/1 (“beengiven toZagreb”);leg.MaaSSen7.1982,Maa/1 (broken); leg.MaaSSen5.1984,Maa/3;leg.Subai25.9.2006,S20850/11+2(juv.).

IslandMljet,grotto200mEofSobra,40malt.,UTMYH13, leg.Pintér & Szigethy2.8.1972,HNHM/1(lowerhalf),72068/1+1(juv.); rockcleft1kmEofSobra,UTMYH13,leg.MaaSSen6.1987,Maa/2.

PeninsulaPelješac,PrapatnanearSton,UTMYH14, leg.MaaSSen5.1984,Maa/1(fraction).

Montenegro:Sutorinamouth(=SWofHercegNovi),de-bris,UTMBN90,NMWK16201/3+1(damaged),38386/1;

leg.KäuFel, NMWE 44507/2, NMWR/2; (exKleMM, exSchlicKuM5381),SMF274290/2.

D i s t r i b u t i o n :A. stenostoma is recorded fromsoutheasternDalmatia fromhabitats close to the seaalongthecoastbetweenthepeninsulaPelješactosouthofHercegNoviintheKotorBayinMontenegro.Itlivessubterraneous.

Agardhiella skipetarica (a. J. Wagner1915)Plate1,Fig.8;textfigs.3,9.

1915 Agardhia (Agardhia) truncatella skipetarica a. J. Wagner, in Sturany & Wagner, DenkschriftenderKaiserlichenAkademiederWissenschaften inWien,mathematische-naturwissenschaftlicheKlasse,91:65,Plate18,Fig.105[locitypici:„ZabljakinMontenegro;KiribrückenächstMesibeiSkutariinAlbanien“;lecto-typedesign.gittenberger1975NMW43328/1].

1926 Agardhia (Agardhiella) truncatella skipetarica, –PilSbry, ManualofConchology,27:166,Plate19,Fig.15.

1975 Agardhiella truncatella skipetarica, – gittenberger, ZoologischeMededelingen,48(24):287.

1988 Agardhiella truncatella skipetarica, – gittenberger,AnnalendesNaturhistorischenMuseums inWien,90 (B):425.

1999 Agardhiella truncatella skipetarica, – Subai, SchriftenzurMalakozoologie,13:45,Fig.5(distributionmap)[partim,onlyfromKérkiraIsland].

D i a g n o s i s :shellslender,denselyribbed,aperturewithoutteeth.



D e s c r i p t i o n : shell relatively small, cylindricalwith a rounded apex.The first 1.5–2whorls slightlygranulated, followedbyradial ribsculptureon thepre-cedingwhorls;lastwhorlwith54–68(13–16/mm)hard-lyelevated ribs;between themain ribs, therearefine,irregularradialribswhichcanonlybeseenunderhighmagnification.

Whorls5.5–6.5;thelastwhorlupto55–58%ofthetotalheight;umbilicusnarrow,elongatecylindrical.

Aperturewithout teeth; in frontalview, theapertur-al rim isu-shaped; interiorlywitha slight lip,outsideflared;palatalwall slightly impressed in themiddle;abovethisarea,theapertureisbulged,andtheapertural

rimisnarrowandsimple;asmallsinulusispresent.Theperistomial rimsareconnectedbyamoderatelydevel-opedcallus.Insideview,theupperpartoftheaperturalrimisslightlyprotruding.

M e a s u r e m e n t s (n=23): H=3.43–4.31;D=1.43–1.68;AH=1.31–1.43;AD=0.81–1.

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 :unknown.D i f f e r e n t i a l d i a g n o s i s :Shells ofA. trun

catellaaregenerallybroaderthanthoseofA. skipetarica.Ithasroundedwhorls,andtheribsaremorewidelyspacedandhigher. InA. truncatella, an infraparietalisandangularisarepresent;itshowsafurrowbetweenthecolumellarwall and infraparietalis stretching towards

Textfig.9.Agardhiella skipetarica (a. J. Wagner1915).Greece,KérkiraIsland(=Korfu),southernsideofroad1.5kmofPetáliaindirectionofPantokrátor-climax,inacavityinlimestonerocks,720malt.,UTMDK00,(H=3.75,D=1.56mm).



theumbilicus;theaperturalrimisclearlylesserflaredifcomparedtoA. skipetarica.

Ty p e s :Albania,KiribridgenearMes(=Mesi)close toShkodër (=Skutari),UTMCM86, leg.Sturany27.4.1905,lectotypeNMW43328/1.

A d d i t i o n a l m a t e r i a l e x a m i n e d :Croatia: LokrumIsland,UTMBN62,legKuzMić,CNHM681/5.

Montenegro:Njeguši,UTMCN20, leg.FuchS,NMWE35215/1+2 (damaged);Petrovac, hillSof the road,UTMCM37, leg.gittenberger 6.5.1975,NNM/1 (damaged); incaveObodskapećinanearRijekaCrnojevica,UTMCM39,leg.MaaSSen 6.1978,Maa/1 (juv.);Virpazar,UTMCM47,leg.dabović,NMWE48742/1 (damaged);caveJabukovdo(=7KmofVirpazaron theway toKomarno),UTMCM48,leg.gittenberger6.u.11.5.1974,NNM/1.

Albania:PeriferiShkodër, app.15.5kmupstream fromdamatKoman,Liquen iKomanit rightbank,debris,180malt.,UTMDM07, leg.Erőss,Fehér,hunyadi&Murányi 15.4.2006,HNHM/2(juv.+damaged).

Greece:Kérkira Island (=Korfu),500mEofTroumpéta(=Pandeleimónas)atN-exposedrockcavitiesat theroadsideand atS-exposed limestone rocks,UTMCJ99, leg.Subai 6.8.1995,S14256/2(damaged);Sfakerá, largersource,UTMCK90,leg.Schütt16.7.1983,Sch1078/1;atthebridge1kmWofMessongi (=16kmSofKérkira-City),UTMDJ07,leg.gittenberger 17.4.1979,NNM/2 (juv./damaged); app.1kmWofBenítses (=8kmSofKérkira-City),100-150malt.,UTMDJ07, leg.gittenberger16.4.1979,NNM/1(juv./damaged); rockcavityat roadsidewesternpartofPantokrá-tor-highplateau,700malt.,UTMDK00, leg.gittenberger 10.1978,NNM/1+2(damaged);southernsideofroad1.5kmofPetália indirectionofPantokrátor summit, inacavity inlimestonerocks,720malt.,UTMDK00,leg.Subai 5.8.1995,S14255/20+5(juv./damaged).

R e m a r k s :a. J. WagnerdescribedA. skipetarica asasubspeciesof A. truncatella.Afterexaminationofmorethan30specimens,thistaxonisconsideredtoformanindependentspecies.Itisclearlydistinguishablefromallneighbouringspeciesbyitsslendershell,thepatternofdenselyarrangedribs,anditsaperturedevoidofanyteeth.

Aftergittenberger (1975:288), theparalectotypefromŽabljakisintheNaturalHistoryMuseuminWar-saw.Ithasnotbeenexaminedbyme.

D i s t r i b u t i o n :ThehithertoknowndistributionareaofA. skipetarica is split. In thenorth, it is foundfromLokrumIslandnearDubrovnikinanarrowrangealongthesouthDalmatiancoastMontenegroandnorth-ernAlbania.A second area of distribution known isknownontheGreekislandKérkira.Thegapofrecordsisprobablyduetolimitedcollectionefforts.

Agardhiella formosa (l. PFeiFFer1848)Plate1,Fig.7;textfigs.8,10.

1848 Pupa truncatella β [Pupa formosa]L. PFeiFFer, Mono-graphiaheliceorumviventium,2:304 [locus typicus:„...inmonteKarst,propeCastelnuovointerTergestumetFiume! InCarniolia,Dalmatia,Turcia.“ restricted

bygittenberger(1975:284):HercegNovi(=Castel-nuovo);typematerial:lost].

1887 Pupa (Coryna) truncatella var. formosa, –WeSter-lund, FaunaderinderpaläarctischenRegionlebendenBinnenconchylien,3:88.

1915 Agardhia (Agardhia) truncatella formosa, – Sturany & Wagner, DenkschriftenderKaiserlichenAkademiederWissenschafteninWien,mathematische-naturwis-senschaftlicheKlasse,91:65,Plate.18Fig.103.

1926 Agardhia (Agardhiella) truncatella formosa,–PilSbry, ManualofConchology,27:164,Plate.19Fig.6–8.

D i a g n o s i s :Largeshells,aperture large,withan-gularis,parietallamellaandinfraparietalis;palataltoothmissingorreduced.

D e s c r i p t i o n :Shell relatively large, cylindricalwithroundedapexAfterca.1.75smooth,slightlygranu-latedwhorls,arapidlyincreasingradialribsculpturebe-gins.Onthelastwhorlthereare36–50(7–11/mm)welldeveloped ribs.Between themain ribs therearefine,irregularradialribswhichcanonlyberecognizedunderhighmagnifications.

Whorls6.5–7.25; the lastwhorloccupies45–56%ofthetotalheight.Theumbilicusisnarrowcylindrical,elongate.

Aperture largewith infraparietalisandangularis. Ingeneral, both teeth are poorlydeveloped and stretchstraight towards the interiorof theshell; aparietal la-mellaispresentaswellwhichisslightlyhigherthanthemarginalteeth;itisstraightandrunsdeepintotheshellwithoutchangesinheight.Infrontalview,aperturalrimalmostu-shaped,slightlystrengthenedbyaninteriorlip,moderatelyflared;slightlydepressedbetweentheuppercolumellar rimand the infraparietalis, formingasmallfurrowpointing towards theumbilicus;on thepalatalside, it is slightly impressedwithweak, teeth-likecal-losities;above that, theaperture isslightlybulged, rimnarrowwithoutlip,sometimeswithaveryshortsinulusabovetheangularis.Theperistomialrimsareconnectedbyawelldevelopedcalluswhichalsocovers the ribsabove theaperture. Insideview, theupperpartof theapertural rimisstraight,and theupper thirddiagonallytruncated.

M e a s u r em e n t s (n=13):H=4.3–5;D=1.75–2;AH=1.43–1.75;AD=1–1.12.

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 :unknown.D i f f e r e n t i a l d i a g n o s i s :A. biarmata iscon-

stantlysmallerandhasasmalleraperture.The ribsontheshell’ssurfacearemoredenselysetandweaker.TheaperturaldentitionissimilarifcomparedtoA. formosa,however,theparietallamellastartsatasomewhatlowerpointon theupperapertural rim.The lowerpartof theapertural rim isnot thatstronglyenlargedandshowsawelldevelopedpalatal tooth.Moreover, insomespeci-mensofA. biarmata additionalcolumellar teethcouldbeobserved.

T y p e s :original typematerial lost.Neotype (herewithdesignated):SMF329426.



A d d i t i o n a l m a t e r i a l e x a m i n e d :Montenegro: Kameno,UTMBN90,leg.Paganetti1902,NMW36742/1;StoSWofthemouthofSutorinaRivernearIgalo(=WofHer-cegNovi),inrockcreasesandcaveniches,UTMBN90,leg.gittenberger13.5.1974,NNM/1; leg.gittenberger5.1975,NNM/2+3(juv.);leg.MaaSSen7.1977,Maa/2+5(juv./dam-aged);N-exposedmountainsidesouthof thecrossroadSuto-rina-Njivice-Igalo(=WofHercegNovi),arockcavity,UTMBN90,leg.Subai11.10.2004,S20417/1;leg.Subai19.9.2005,S20231/10+18 (juv.+ fragments);Njeguši,UTMCN20,

NMW35217/1(+1+2(juv.)biarmata);caveObodskapećinanearRijekaCrnojevica,UTMCM39, leg.Schütt5.6.1978,Sch862/1(juv.);RumijaMountains,17kmfromTuđemilito-wardsVirpazar,atlimestonerocksnextthecountryroad,app.470malt.,UTMCM47,leg.Subai25.9.2005,S20347/1.

R em a r k s :PFeiFFer(1848:303–304)separatedava-riety“β”ofPupa truncatellaandnameditPupa formosa. Thelocalitieshementionedattheendofhistextcovers

Textfig.10.Agardhiella formosa (l. PFeiFFer1848).Montenegro,N-exposedmountainsidesouthof thecrossroadSutorina-Njivice-Igalo(=WofHercegNovi),arockcavity,UTMBN90,(H=4.56,D=1.87mm).



almostthewholedistributionareaofthegenusAgardhiella fromSlovenia (=Carniola) toBulgaria/Greece (asapartofTurkeyatthetimebeing).Regardingtheexplicitlynamedlocalities, thevariety“β”isonlyfoundinthevi-cinityofHercegNovi(=Castelnuovo).Notypematerialcouldbetracedinallrelevantmuseum’scollections,andthecollectionoflouiS PFeiFFerisdestroyed.AsPFeiFFer didnot illustrate the typespecimen,clarificationof thetaxonomicalconceptcanonlybeachievedbydesignat-inganeotype.InordertoavoidfuturemisinterpretationsandtofixtheuseofthenameA. formosa, thespecimenfiguredinthispaper(Plate1,Fig.7)isdesignatedasneo-type.ItoriginatesfromtheW-sideofHercegNovi(Igalo),the locus typicusof thisspecies.AfterPFeiFFer itwereSturany & Wagner (1915:65,Plate.18Fig.103)whoillustratedalargeAgardhiellafromKameno(=ca.2kmfromHercegNovi).TheirdescriptionandmeasurementsagreewellwithPFeiFFer’s P. formosa,andtheyrecordedthespecimens fromDubrovnik (=Ragusa)andHercegNovi(=Castelnuovo).TheirinformationwasrepeatedbyPilSbry(1926:164).gittenberger(1975:284)considered A. biarmataasynonymofA. formosaduetothesimilardevelopmentof theaperturaldentition.However,heal-readystressedthe“remarkablylargemeasurements”forA. formosaasmentionedbypreviousauthors(gittenberger 1975:285).Recentlycollectedspecimens(2004–2006)ofA. formosafromtheareaaroundHercegNoviprovedtheconceptthatA. formosaandA. biarmataaretwoseparatespecies.TheirrespectiveareasofdistributionoverlapintheareaofKotorBay. A. formosa isexclusively foundeastofthebay,whileA. biarmataisfoundinalargerareaat thewesternsideofSkadarLake(=SkutariLake) inwesterndirectiontotheNeretvaValley.Sympatricoccur-rencecouldberecordedfromNjegušiandMokrine.

Thespecieslivessubterraneous.Allrecentlycollectedspecimenswerefoundinrockcavitiesandrockcrevices.

D i s t r i b u t i o n :So far this specieshasonlybeenfoundinthemountainssurroundingKotorBayandatthecoastalstripeeastoftheKotorBay.

Agardhiella dabovici gittenberger 1975Plate1,Fig.9;textfigs.11,12.

1975 Agardhiella dabovici gittenberger, ZoologischeMed-edelingen,48(24):281,Fig.2(shell),Figs.4–5(geni-talorgans),Plate1Fig.1–2 (radula),Fig3–4 (shell)[locus typicus:“Velika jama imSüdhangdesBergesSokobeiDupilo,westlichvonVirpazar,in300–400mHöhe”;holotypeNNM55043].

1985 Agardhiella dabovici, – zilch, Archiv für Mol-luskenkunde,116(1/3):128.

1985 Agardhiella dabovici,–MaaSSen, DeKreukel,21(1–2):4,distributionmap.

D i a g n o s i s :mediumsizedshells,aperturewithin-fraparietalis,angularisandparietallamella;palatalwallof theperistomeshortwith strongly impressedpalatalfurrow;columellarisandbasaliswelldeveloped.

D e s c r i p t i o n : shell medium-sized, cylindricalwitharoundedapex.Afterca.1.75–2slightlygranulatedwhorls,theradialribsculpturebegins.Onthelastwhorlthereare37–47(7–11/mm)ribs.Betweenthemainribstherearefine,irregularradialribswhichcanonlyberec-ognizedunderhighmagnifications.

Whorls6.5–7.25; the lastwhorloccupies51–56%ofthetotalheight.Theumbilicusisnarrowcylindrical,elongate.

Aperturewithinfraparietalisandangularis,bothteethelongate towards the interiorof theshell;aparietal la-mellaispresentaswellwhichcontinuouslyincreasesinheightafter theparietalcallus followedbyasmallde-pression;then,itrisesagaininheighttoacertainmaxi-mumandalmostdisappearsdeep in the interiorof theshell;insideviewitisoftriangularshape.Deepinsidetheaperturethereisawelldeveloped,verticalcolumel-lariscovering theupperhalfof thecolumellawith itslowerendslightlyenlarged.Thesmallbasalisisshiftedtowardstheaperturalrim.Occasionally,thereareoneormore small, roundednodulesbetween thebasalis andthecolumellarrim.Infrontalview,aperturalrimalmostu-shapedand strengthenedbyan interior lip, slightlyflared;aperturalrimlightlydepressedbetweentheuppercolumellar rimand the infraparietalis, formingasmallfurrowpointing towards theumbilicus;palatalwalloftheapertureconsiderablyimpressedformingashortpa-latal callus coveringup to¼of the apertural height;above that, theaperture is slightlybulged, rimnarrowwithoutlip.Insideview,theouterpartoftheaperturalrimiscentrallyprojectingabovetheaperturallevel.

M e a s u r em e n t s (n=63):H=4.18–5.8;D=1.87–2.3;AH=1.43–1.8;AD=0.87–1.3.

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 :seegitten-berger (1975:282,Fig.4–5).

D i f f e r e n t i a l d i a g n o s i s :A. zoltanorumn.sp.differs fromA. daboviciby itshigherbutnarrowerap-erture. Itsparietal lamellaprotrudesabove theaperturelevel.Ininterioroftheshellitissubdividedbyafurrowandshowsanundulatingshape. Itspalatal fold isverylong,andoccupiesca.2/3oftheheightoftheaperture.Additionally,A. zoltanorum has smaller teethon thesomewhat enlarged columellar rimanda remarkablylongcolumellaris.However, thebasalis (ifpresent atall),islocateddirectlybeneaththecolumellaris.

T y p e s (juvenile respectivelydamagedspecimenof theparticulartypeseriesarealsomentioned.Intheoriginalpubli-cation,thesehavenotalwaysbeenconsidered.Thesespecimen(noparatypes)areaddedinbrackets):Montenegro:caveVili-napećinaNWofDonjaSeoca(=VilinapećinanearKaluderacSofVirpazar),UTMCM47,leg.Dabović,NMWE19778/47(paratypes) [+3juv. o. damaged], 49630/75 (paratypes)[+1];caveVelikajama(=Velajama,Sokopećina)atsouthernsideofmountSokol(=“Soko”)nearDupilo(=WofVirpazar),UTMCM48,NMWwithoutNr./40 (paratypes) [+5juv.o.damaged],NMWE19050/90 (paratypes) [+1juv. o. dam-aged];leg.Dabović,NMWE48680/11(paratypes),49631/9+3



(juv./damaged) (paratypes),49632/61(paratypes) [+5juv.o.damaged];leg.gittenberger11.5.1974,SMF239645/3(para-types);Jabukovdocave(=SofKomarno,NNWofVirpazar),UTMCM48,leg.Dabović,NMWE48396/2(paratypes);leg.gittenberger6.u.11.5.1974,NNM55045/11+3(juv.)(para-types); caveRidpećinanearPopratnica,UTMCM48, leg.Dabović,NMWE20793/1(paratype)[+1].

A d d i t i o n a l m a t e r i a l e x a m i n e d :Montene-gro:caveVilinapećinaNWofDonjaSeoca(=VilinapećinanearKaluderacSofVirpazar),UTMCM47,NMWK/4(with-outNr.)+27(withoutNr.,mostlydamaged), 38357/30; (exedlauer)NMWK51693/3; caveVelika jama (=Velajama,

Sokopećina)atsouthernsideofmountSokol(=“Soko”)nearDupilo (=WofVirpazar),UTMCM48,NMWK38355/20,38356/12,38358/6,38359/15,51691/32,51692/8; Jabukovdocave(=SofKomarno,NNWofVirpazar),UTMCM48,leg.MaaSSen&Schütt3.6.1978,Maa/3,Sch/1,S5239/1;leg.Subai23.9.2005,S20401/3+1(juv.).

R em a r k s :Although thespecimensfromKleMM’s seriesinthecollectionoftheNMWareobviouslyfromthesamehabitat,evenfromthesamecollector,theyarenottobeconsideredasparatypessincetheauthordidnot

Textfig.11.Agardhiella dabovici gittenberger 1975. Montenegro,Jabukovdocave(=SofKomarno,NNWofVirpazar),UTMCM48,(H=4.18,D=2mm).



seethembeforethedescription.Thecorrectnameofthemountainatthelocustypicusis“Sokol”andnot“Soko”.Also thecaveVelikapećinaonthe labels issometimesnamed„Sokolpećina”.

InthelotfromthecaveVilinapećinatherearesomere-markablylargespecimensofA. dabovici,buttheydonotdif-ferintheothershellcharacteristicsfromthehithertoknownspecimens. Sofar, A. daboviciisonlyknownfromcaves.

D i s t r i b u t i o n :EndemictocavesaroundVirpazarinsouthernMontenegroatthewesternsideofSkadarLake.

Agardhiella extravagantan.sp.Plate1,Fig.12,textfigs.12,13

D i a g n o s i s :mediumsizedbroadshells; aperturewith infraparietalisandangularis, a strongparietal la-

mellaandhorizontalcolumellaris;umbilicuswide, lastwhorlstronglyconstrictedatthebase.

D e s c r i p t i o n :shellmediumsized,broadlybiconi-calwitharoundedapex.Afterca.2slightlygranulatedwhorls,theradialribsculpturebegins.Onthelastwhorlthereare37–62(5–10/mm)ribs.Betweenthemainribstherearefine,irregularradialribswhichcanonlyberec-ognizedunderhighmagnifications.

Whorls6.5–7;thelastwhorloccupies53–55%ofthetotalheight.Theshellisstrongly,horizontallyconstrict-edatthebaseabouthalfawhorlawayfromtheaperture;umbilicuswide,roundedtoelongatecylindrical.

Aperturewitha strong infraparietalisandangulariswhichprojectstraighttotheinterioroftheshell.Thepa-rietalisiscurved,startsattheaperturalrimandstretchesslowly increasing inheight towards the interiorof the



shell.Thehorizontalcolumellaris is strongstarting inthecentreof thecolumellaandissimilarlyshapedliketheparietalis. Inside theaperture, they reachabout thesameheightandrunparalleltotheinterioroftheshell.In frontalview, theapertural rim isu-shaped, slightlystrengthenedinteriorlybyalipandflared;aperturalrimslightlydepressedbetween theupper columellar rimandtheinfraparietalis,formingasmallfurrowpointingtowards theumbilicus.Theperistomial rimsare con-nectedbyawelldevelopedcalluswhich is indentatewhereittouchestheribsofthelastwhorls.Themiddleofthepalatalwalloftheapertureisimpressed,acallusis

missing;abovethat,theapertureisslightlybulged,rimnarrowwithoutlip.Insideview,thecentralpartoftheaperturalrimstronglyprojectsabovetheaperturallevel.

M e a s u r e m e n t s (n =4) : H=3.62–4.56;D=2.18–2.37;AH=1.37–1.68;AD=1–1.06;measure-mentsof theholotype:H=4.56;D=2.37;AH=1.68;AD=1.06.

D i f f e r e n t i a l d i a g n o s i s :ThisspeciescanbeeasilydistinguishedfromallotherspeciesofAgardhiella byitsbroadbiconicalshellandthestronghorizontalcolu-mellaris(whichismoretypicalforspeciesofArgna).Simi-larsizedshellsofA. zoltanorumn.sp.arelessbroadcy-

Textfig.13.Agardhiella extravagantan.sp.Albania,PeriferiShkodër,app.18kmupstreamfromdamatKoman,aleftsideValleyofLiqueniKomanit,limestonerocks,debris,170malt.,UTMDM07,paratype(H=3.62,D=2.25mm).



lindricalandneverofbiconicalshape;itsapertureismorestronglyconstricted,anditsangularishardlyrecognizable;astronghorizontalcolumellarisismissing,butithasastrongverticalcolumellarisdeepintheaperture,anditscolumellarrimisstraight;itspalatalisprojectsabovetheaperturallev-el,islowerthaninA. extravagantaandsomewhatundulat-ing;itsumbilicusisnarrowerandthebaseofthelastwhorlisonlyconstrictedshortlybeforetheaperture.

L o c u s t y p i c u s :Albania,PeriferiShkodër,app.18kmupstreamfromembankmentdamatKoman,attheleftsideinthevalleyofLiqueniKomanit,limestonerocks,debris,170malt.,UTMDM07.Besidestheholo-typeallspecimensmentionedareparatypes.

T y p e s :fromthelocustypicus,leg.Erőss,Fehér,huny-adi&Murányi14.4.2006,holotypeHNHM96786,paratypeh/1;Albania:districtShkodër,app.17.5kmupstreamfromembankmentdamatKoman, rightbank, stream,debris andlimestonerocks,170malt.,UTMDM07, leg.Erőss,Fehér,hunyadi&Murányi 15.4.2006,e/1; app.1.5kmupstreamembankment fromdamatKoman,Liquen iKomanit rightbank,sidestream,debris,180malt.,UTMDM06,leg.Erőss,Fehér,hunyadi&Murányi14.4.2006,HNHM96787/1(juv.).

E t ym o l o g y :fromLatin“extravaganta”foritsun-usualshellcharacteristics.

D i s t r i b u t i o n :Onlyknown fromnorthwesternAlbaniaNEofKoman(=app.20-25kmEofShkodër)fromthedebrisoftheDrimRiver.

Agardhiella zoltanorumn.sp.Plate1,Fig.11;textfigs.12,14.

D i a g n o s i s :mediumsizedshell,aperturestronglycompressed,withinfraparietalis,astrongparietallamellaandstrongverticalcolumellaris;sometimeswithabasa-lisandrudimentsoftoothatthecolumellarrim;baseofthelastwhorlconstrictedshortlybeforetheaperture.

D e s c r i p t i o n :shellmediumsized,compact,cylin-dricalwitharoundedapex.After1.75toalmost2slight-lygranulatedwhorls,theradialribsculpturebegins.Onthe lastwhorl thereare43–66 (7–11/mm)moderatelyelevated ribs.Between themain ribs therearefine, ir-regular radial ribswhichcanonlyberecognizedunderhighmagnifications.

Whorls5.75–6.75; the lastwhorloccupies54–70%ofthetotalheight;umbilicusmoderatelywide,roundedtoelongatecylindrical.

Aperturewithaninfraparietalisandaparietalis,bothteetharestrongandstretchstraighttotheinterioroftheshell;parietalisprojectsabove theapertural level; it issubdivided in twopart by a small depressionbehindtheapertureand isundulatinganddecreasing inheighttowards its enddeep in the interiorof the shell;usu-ally,anangularis ispresent,but itoften fuseswith thestronglyelevatedaperturalcallus;insomespecimens,afewsmall,hardlydevelopedteethcanbefoundbetweentheinfraparietalisandtheparietalis.Deepinsidetheap-

erturethereisalong,stronglyelevatedverticalcolumel-lariswithasmallbasalisdirectlybeneath it. In frontalview,theaperturalrimisu-shaped,slightlystrengthenedinteriorlybya lipandflared;apertural rimslightlyde-pressedbetweentheuppercolumellarrimandtheinfra-parietalis, formingasmall furrowpointing towards theumbilicus;columellarrimoftenwithrudimentaryteeth.Theperistomialrimsareconnectedbyawelldevelopedcalluswhich is slightly indentate above the aperture.Palatalwalloftheaperturestronglyimpressed,formingalargepalatalfoldoccupyingabout2/3oftheaperturalheight;above that, theaperture is slightlybulged, rimnarrowwithoutlip.Insideview,theouteraperturalrimisstraight,andtheupperthirddiagonallytruncated.

M e a s u r em e n t s (n=21):H=3.5–4.68;D=1.7–2.3;AH=1.43–1.87;AD=0.68–1.1;measurementsoftheholotype:H=4.37;D=2.18;AH=1.62;AD=1.

D i f f e r e n t i a l d i a g n o s i s :SimilarlysizedshellsofA. reinhardtiandof A. stenostomaaremoreslenderifcomparedtoA. zoltanorumandhaveadenserribbingpattern. Ithasaclearlyvisibleangularis,but infrapari-etalis,parietalisandcolumellarisareweaker.InA. reinhardti,theteethonthecolumellarrimaremissing,whileinA. stenostomathebasalisismissing.

TheapertureofA. dabovici islesshighandlessim-pressedlaterally.Itsparietalisislowattheapertureanddoesnotprojectabove theapertural level; towards theinterioroftheshell,itshowsashorthumpanddecreasestheninheight.ThepalatalisofA. daboviciisshorterandlessshiftedtowardsthecentreoftheaperture.Itonlyoc-cupies1/3oftheaperturalheight.OccasionallyoccurringsmallteethatthecolumellaarenotplacedatthesideasinA. zoltanorum,butdeeperinsidetheshell.However,thebasalisisclosertotheaperturalrim.Finally,itscolu-mellaris ismuchshorter than that inA. zoltanorum, itsouterendisslightlyenlarged.

ForthedifferenceswithA. extravagantan.sp.,pleaserefertothisspecies.

L o c u s t y p i c u s :Albania:PeriferiDibrë,SofZall-Reç(=5kmSofthebridgeofriverDriniZi),limestonerocks,510malt.,UTMDM43.Besidestheholotypeallfollowingspecimenareconsideredasparatypes.

T y p e s :fromthelocustypicus,leg.DEli, Erőss, FEhér & Murányi9.10.2005,holotypeHNHM95900,paratypeHNHM95901/2.Albania:app.1.5kmupstreamfromdamatKoman,Liquen iKomanit rightbank, side stream,debris, 180malt.,UTMDM06,leg.Erőss,Fehér,hunyadi&Murányi14.4.2006,h/2+1(juv.);app.15.5kmupstreamfromdamatKoman,LiqueniKomanitrightbank,debris,180malt.,UTMDM07,leg.Erőss,Fehér,hunyadi&Murányi15.4.2006,E/1+1(juv./damaged), H/1+1 (juv./damaged),HNHM96790/1+3 (juv./damaged),MMMB/1;app.17.5kmupstreamfromdamatKoman,LiqueniKomanitrightbank,stream,debrisandlimestonerocks,170malt.,UTMDM07,leg.Erőss,Fehér,hunyadi&Murányi15.4.2006,HNHM96788/1;app.18kmupstreamfromdamatKoman,aleftsideValleyofLiqueniKomanit,limestonerocks,debris,170malt.,UTMDM07,leg.Erőss,Fehér,hunyadi&Murányi14.4.2006,HNHM96789/3+11(juv./damaged),e/3,S20799/1;Tropojë



county,overGurorë,3kmNoftheriverValbonëmouth,limestonerocks,andsecondaryhornbeambush,289malt.,UTMDM18,leg.DEli, Erőss, FEhér & murányi7.10.2005,e/1,HNHM96791/1+1(juv.);Kukëscounty,GurriiArrënit,EofArrën(=35kmSoftheShkodër-Kukësroad),limestonerocks,devastatedbeechforest,1598malt.,UTMDM44,leg.DEli, Erőss, FEhér & murányi 8.10.2005,HNHM96802/1,MMMB/1(juv.).

E t y m o l o g y :Thisnewspecies isdedicated to itsdiscoverer, theHungarianmalacologistsDr. zoltán ErőssandDr.zoltán Fehér.

D i s t r i b u t i o n :Thisspeciesisknownfromnorth-ernAlbanianorthofLezhë in thewestand theKorabMountainsattheeasternborderofthecountry.

Textfig.14.Agardhiella zoltanorumn.sp.Albania,Tropojëcounty,overGurorë,3kmNoftheriverValbonëmouth,limestonerocks,andsecondaryhornbeambush,289malt.,UTMDM18,paratype(H=4.62,D=2.1mm).



Discussion

TheremarkablylargedistributionareaofA. truncatellacanbeexplainedbyitsautecology.Itlivesinrockduffandrockdebrisandthuscaneasilybetransportedoverlargerareaswiththehelpofrainfallsandstrongwinds.AsimilarwayoflifeisknownforA. biarmata,butthisspeciesisalsorecordedfromcavesalongtheDalmatiancoast.Thereareotherspecieswhichlivesubterraneousandareexclusivelyfoundinafewcaves(e.g.A. dabovici)orinrockcrevicesandrockholes(e.g.A. formosa,A. skipetarica,A. stenostoma).Theirareaofdistributionis relativelysmallandrestricted toa fewneighbouringcavesystemsorneighbouringmountains.

Specimensfoundintheirprimaryhabitatslikeacaveor fromrockcrevicesonlyshow little individualvari-ation,whereasspecimensof thesamespeciesofAgardhiellacollectedfromriverdebrisshowalargevariationinsize,numberofribs,aperturalshapeetc.Thisobserva-tioncanbeexplainedbythefactthatariverdebriscon-tainsshellsoriginatingfromvariouspopulationswithin

itscatchmentarea,washed inby rain from thesoilorrockcrevices.This is contrastedbypopulations fromcavesystemswith restrictedgeneticexchange.Agoodexample isA. dabovici,whichoccurs incaveswestofVirpazarinthesmalltypicalform,whilespecimensorig-inating from thecaveVilinapećinanearDonjaSeocaareconstantlymuchlargealthoughtheyshareallothershellcharacteristicsofthespecies.ThesecavesareeastofVirpazarandbelongtoanothermountainridgewithaseparatecavesystem.

Acknowledgment

Mysincere thanksgo to thedirectorsandownersof thementionedcollectionsandtheirallowanceforloaningandex-aminationofthespecimensmentionedinthetext.IalsowanttothankDr. eiKe neubert(Badenweiler)forpreparingthepho-tographsandtheplatesandhishelpwiththetext.

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Manuscriptsubmitted:03.12.2007Revisedmanuscriptaccepted:17.03.2008



Plate



Plate1allphotos×15,phot.E.neubert

Figs1–4. Agardhiella truncatella (l. PFeiFFer1841). 1)Austria,Carinthia,KarawankenMts.,TscheppaGorge(=S

ofKlagenfurt),atlimestonerocks,UTMVM44(characteristicspecimen,SMF329425H=3.62,D=1.62mm).

2)BosniaandHercegovina,Karanovac,UTMBQ85(small,teethless,SMF205796/1,H=3.1,B=1.5mm)

3)Greece,Ípiros,app.38kmSofIoannina(=51kmalongthecountryroad),rocksWofcountryroadnearpaleoliticcaves,150malt.,UTMDJ84(faintlydeveloppedpalataltoothandslightlybeginningparietallamella,NNM/1,H=3.37,B=1.68mm)

4)Greece,Ípiros,Zagoria,Kipi,limestonerocksnearoldbridge,UTMDK81(largespecimen,NMW102816/4,H=3.8mm)

Figs5–6. Agardhiella biarmata (o. boettger1880). 5)lectotypeSMF4658,Croatia,Dubrovnik(=Dalmatien,Ragusa),

UTMBN52/62(H=3.74mm). 6)Coryna biarmata spelaea Kobelt1906,syntypeSMF51691,

BosniaandHercegovina,cavenearZavala,coll.O.boettgerexSturany1904(H=4.1,B=1.9mm)

Fig.7. Agardhiella formosa (l. PFeiFFer1848). Montenegro,N-exposedmountainsidesouthofthecrossroad

Sutorina-Njivice-Igalo(=WofHercegNovi),arockcavity,UTMBN90(NeotypeSMF329426,H=4.8,D=1.87mm).

Fig.8. Agardhiella skipetarica (a. J. Wagner1915). LectotypeNMW43328,Albania,Kiri-BridgenearMes(=Mesi)

nearShkodër(=Skutari),UTMCM86(H=4.4mm).

Fig.9. Agardhiella dabovici gittenberger 1975. ParatypeSMF239645,Montenegro,caveVelikajama(=Velajama,

Sokopećina)atsouthernsideofmountSokol(=„Soko“)nearDupilo(=WofVirpazar),UTMCM48(H=4.1,D=1.7mm).

Fig.10. Agardhiella stenostoma (Flach1890). HolotypeSMF4597,“Dalmatien”(H=4.1,D=1.4mm).

Fig.11. Agardhiella zoltanorumn.sp. HolotypeHNHM95900,Albania,PeriferiDibrë,SofZall-Reç

(=5kmSofthebridgeofriverDriniZi),limestonerocks,510malt.,UTMDM43(H=4.37,D=2.18mm).

Fig.12. Agardhiella extravagantan.sp. HolotypeHNHM96786,Albania,PeriferiShkodër,app.18km

upstreamfromembankmentdamatKoman,attheleftsideinthevalleyofLiqueniKomanit,limestonerocks,debris,170malt.,UTMDM07(H=4.56,D=2.37mm).


Plate 1


Documents

Revision of the Argninae, 1. The species of Agardhiella from