{"id":650,"date":"2022-02-12T18:56:50","date_gmt":"2022-02-12T17:56:50","guid":{"rendered":"https:\/\/hnords.de\/wordpress\/?page_id=650"},"modified":"2022-04-05T23:31:55","modified_gmt":"2022-04-05T21:31:55","slug":"clausilia-dubia","status":"publish","type":"page","link":"https:\/\/hnords.de\/wordpress\/clausilia-dubia\/","title":{"rendered":"Clausilia dubia"},"content":{"rendered":"

Clausilia dubia<\/em><\/strong> \u2013 a revision based on informative shell characters<\/strong><\/p>\n

Hartmut Nordsieck (III. 2022)<\/p>\n

I. Introduction<\/h4>\n

The subspecies division of C<\/em>. dubia<\/em> is difficult, because across the range of the species many forms of different rank are present. The previous revisions of the species (Klemm 1960, Nordsieck 1990, Bank & Gittenberger 2000, Jaksch 2012, Mason et al. 2020) are incomplete, because they are restricted to certain parts of the range. The only complete revision (Nordsieck 2002) is a survey on the different forms of the species with recognition of two major subspecies, d<\/em>. vindobonensis<\/em> A. Schmidt 1856 sensu lato and C<\/em>. d<\/em>. dubia<\/em> sensu lato, but without a detailed subdivision into further subspecies.
\nIn the papers of Klemm, Jaksch and Mason et al. a wrong nomenclature is used (see Nordsieck 1990; 145, footnote 16, Bank & Gittenberger 2000: 16-17, 20): C<\/em>. d<\/em>. dubia<\/em> sensu Klemm = C<\/em>. d<\/em>. vindobonensis<\/em>, C<\/em>. d<\/em>. obsoleta<\/em> sensu Klemm = C<\/em>. d<\/em>. dubia<\/em>. It is difficult to understand why this has been continued by Jaksch and Mason et al.
\nIn the same papers from several localities more than one subspecies are given. This is not acceptable.
\nIn Mason et al. no qualitative shell characters are used (though examined according to the authors).
\nInformative shell characters within the subgenus Clausilia<\/em> (C<\/em>.) have been discussed by Neubert & Nordsieck (2002). For this revision those shell characters, which within C<\/em>. dubia <\/em>have revealed to be informative were in the focus of attention (see Figs. 1-2):
\n1. development of inferior lamella;
\n2. formation of clausilium plate;
\n3. sculpture.
\nShell measurements are no informative shell characters of Clausilia<\/em> species. K\u00e4ufel (1929, see also Klemm 1960) united forms of C<\/em>. dubia<\/em> with more slender shell, C<\/em>. rugosa<\/em> and C<\/em>. bidentata<\/em> to one species, without recognizing the differences of the species in the Pyrenees (C<\/em>. rugosa<\/em>, C<\/em>. bidentata<\/em>, see Nordsieck 1990) or in central Europe (C<\/em>. dubia<\/em>, C<\/em>. bidentata<\/em>, see Klemm under C<\/em>. d<\/em>. moldanubica<\/em>). Therefore it is not astonishing that also the differentiation of subspecies of C<\/em>. dubia<\/em> by shell measurements by Jaksch and Mason et al. failed.
\nMore than fifty intraspecific taxa of C<\/em>. dubia<\/em> have been described. In former papers (Nordsieck 1990, 2002) I registered most of these taxa (not considering several taxa, mainly from Westerlund, which refer to infrasubspecific forms of C<\/em>. dubia<\/em> from central and northern Europe). Type or original material of the majority of the registered taxa (35) could be studied.
\nFor this revision samples of C<\/em>. dubia<\/em> from 115 localities across the whole range have been examined more thoroughly (character states quantified, rib density measured). The rib density is given as rib number per 1 mm of the penultimate whorl = R1<\/sub>.
\nIn the meantime (IV. 2022) the examination has been extended to samples from 160 localities.<\/p>\n

\"M\u00fcndung<\/a>
\nFigs. 1-2. Clausilia dubia<\/em>, characters of CA.
\nBody whorl partly removed. Shell height = H (mm).
\n1. C<\/em>. dubia<\/em> vindobonensis<\/em>, A, Lower Austria, Helenental near Baden; H 12.25.
\n2. C<\/em>. d<\/em>. dubia<\/em>, D, Hesse, K\u00f6nigstein \/ Taunus; H 12.7.
\nAbbreviations: cp = clausilium plate; il = inferior lamella; ilp = folds of inferior lamella;\u00a0 inp = fold of interlamellar; lu = lunella; pr = principal plica; sc = subcolumellar lamella; sl = superior lamella; sp = spiral lamella.
\nOuter lobe of clausilium plate marked by an arrow.<\/h5>\n

II. Systematic part<\/h4>\n

Subspecies:
\nMajor subspecies: d<\/em>. vindobonensis<\/em> sensu lato, d<\/em>. dubia<\/em> sensu lato.<\/p>\n

d<\/em>. vindobonensis<\/em> sensu lato
\nInferior lamella mostly high (more s-like ascending); terminal (outer) lobe of clausilium plate mostly upbent and pointed (see Fig. 1).
\nEastern and southeastern part of the range.
\nIncluding
\n1. d<\/em>. vindobonensis<\/em>, containing:
\na. normal forms
\n(infrasubspecific: trencsiniensis<\/em> Brancsik 1888, manina<\/em> Brancsik 1888, podolica<\/em> Bakowski 1880, transilvanica<\/em> A. Schmidt 1856 [non A. Schmidt 1853], moldanubica<\/em> Klemm 1960):
\nDensely ribbed (R1<\/sub> 6.6-9.8); inferior lamella and clausilium plate like major subspecies.
\nEastern countries, in the west reaching R\u00fcgen and Passau, Germany, and in the south Carinthia. traced up to East and South Tyrol (Matrei, Villn\u00f6ss valley).
\nNotes: C<\/em>. d<\/em>. moldanubica<\/em>\u00a0 is a densely ribbed form of d<\/em>. vindobonensis<\/em>. Some forms from Lower Austria (e. g., Bad Fischau, R1 <\/sub>11) are transitional to d<\/em>. huettneri<\/em>.
\nb. dwarf forms
\n(infrasubspecific: runensis<\/em> Tschapeck 1883, steinbergensis<\/em> Edlinger 2000):
\nSmall; more densely ribbed (R1<\/sub> 13).
\nE. Austria.
\nNote: The runensis<\/em> form is from Styria (near Rein). The other forms affiliated to C<\/em>. d<\/em>. runensis<\/em> by Klemm (1960) are small forms of C<\/em>. d<\/em>. vindobonensis <\/em>(e. g., the form from T\u00fcrkensturz, Lower Austria).
\n2. Weakly sculptured subspecies:
\nd<\/em>. gobanzi<\/em> L. Pfeiffer 1868,\u00a0 d<\/em>. speciosa<\/em> A. Schmidt 1856.
\nVery densely ribbed or rib-striated (R1<\/sub> 10-14.5); inferior lamella and clausilium plate like d<\/em>. vindobonensis<\/em>.
\nS. E. Carinthia (d<\/em>. speciosa<\/em>), E. Styria (d<\/em>. gobanzi<\/em>).
\nNote: C<\/em>. d<\/em>. speciosa<\/em> is restricted to its type locality in Carinthia (Rabenstein ruin). The forms from Lower Austria determined by Klemm (1960) as d<\/em>. speciosa<\/em> are C<\/em>. d<\/em>. dubia<\/em> (Traisen valley) or C<\/em>. d<\/em>. huettneri<\/em> or C<\/em>. d<\/em>. schlechtii<\/em>, respectively (Schwarza valley).
\n3. Coarsely sculptured subspecies:
\nd<\/em>. bucculenta<\/em> Klemm 1960, d<\/em>. carpathica<\/em> Brancsik 1888 [non Hazay 1885], d<\/em>. \u201ecostata<\/em>\u201c Brancsik 1888 [name not available, because infrasubspecific], d<\/em>. floningiana<\/em> Tschapeck 1886, d<\/em>. grimmeri<\/em> L. Pfeiffer 1848, d<\/em>.\u00a0 grimmeri<\/em> sensu Bielz 1867 = gratiosa<\/em> Saj\u00f3 1968, d<\/em>. ingenua<\/em> Hudec & Brabenec 1963, d<\/em>. otvinensis<\/em> M. Gallenstein 1895.
\nCoarsely ribbed (R1<\/sub> 3.2-6.9); inferior lamella and clausilium plate in most subspecies like d<\/em>. vindobonensis<\/em>.
\nE. and S. Austria, Slovakia, Romania.
\nNote: C<\/em>. d<\/em>. floningiana<\/em> from N. Styria and C<\/em>. d<\/em>. „costata<\/em>\u201d from W. Slovakia exhibit a clausilium plate like d<\/em>. dubia<\/em> (subspecies transitional to d<\/em>. dubia<\/em>).<\/p>\n

d<\/em>. dubia<\/em> sensu lato
\nInferior lamella mostly moderately high (more steeply ascending); terminal (outer) lobe of clausilium plate not upbent, forming an outer corner (right or obtuse angle) (see Fig. 2).
\nWestern part of the range.
\nIncluding
\n1. Forms transitional to d<\/em>. vindobonensis<\/em>
\n(infrasubspecific: alpicola<\/em> Clessin 1878, didyma<\/em> F. J. Schmidt 1847, dydima<\/em> M. Gallenstein 1852 [non F. J. Schmidt 1847] = ovirensis<\/em> A. Schmidt 1856, sordellii<\/em> Adami 1876, langobardica<\/em> Pini 1883, ? reticulata<\/em> Pini 1883):
\nRibbing like d<\/em>. dubia<\/em>; inferior lamella high or moderately high; terminal lobe of clausilium plate like d<\/em>. dubia<\/em>, but more frequently somewhat upbent and pointed.
\nNorthern Carpathians, southern Alps (Karawanken to Dolomites, South Tyrol, Lombardy).
\nNote: Bank & Gittenberger (2000) classify most dubia<\/em> samples from N. E. Italy with d<\/em>. dubia,<\/em> some from the Dolomites between Bozen and Belluno as subspecies of its own d<\/em>. alpicola.<\/em> These samples shall be more densely ribbed than d<\/em>. dubia<\/em> (R1<\/sub> 11), but the sample from Schlern mountain (type locality), which I examined, exhibits a rib density like other samples from that region (R1<\/sub> 8.5).
\n2. d<\/em>. dubia<\/em>, containing:
\na. normal forms
\n(infrasubspecific: gracilis<\/em> C. Pfeiffer 1821, roscida<\/em> Studer 1820 = obsoleta<\/em> A. Schmidt 1856, fagotiana<\/em> Bourguignat 1877 = geretica<\/em> Bourguignat 1877 (name fagotiana<\/em> chosen by the first revisor, K\u00e4ufel 1929), suttoni<\/em> Westerlund 1881 = cravenensis<\/em> Taylor 1895, pauperata<\/em> Brancsik 1888).
\nDensely ribbed (R1<\/sub> 7.6-12.4); inferior lamella and clausilium plate like major subspecies.
\nWestern countries from N. Spain (Catalonia) and England (Pennine mountains) to the boundary against d<\/em>. vindobonensis<\/em> in central Europe, northern Carpathians, and eastern and southern Alps.
\nb. form transitional to the weakly sculptured subspecies
\n(infrasubspecific: gracilior<\/em> Clessin 1888 [non Mousson 1863]):
\nLess densely ribbed than d<\/em>. huettneri<\/em> (R1<\/sub> 10-12).
\nS. E. Lower Austria, N. E. Styria.
\n3. Weakly sculptured subspecies:
\nd<\/em>. huettneri<\/em> Klemm 1960, d<\/em>. schlechtii<\/em> A. Schmidt 1856, d<\/em>. tettelbachiana<\/em> Rossm\u00e4ssler 1838.
\nVery densely ribbed (d<\/em>. huettneri<\/em> R1<\/sub> 12-14.5) or indistinctly rib-striated (d<\/em>. schlechtii<\/em>, d<\/em>. tettelbachiana<\/em> R1<\/sub> \u226515); inferior lamella mostly high or moderately high; clausilium plate like d<\/em>. dubia<\/em>, in one form of d<\/em>. schlechtii<\/em> terminal lobe upbent and pointed.
\nS. Lower Austria and N. Styria.
\nNotes: The examination of the original sample of C<\/em>. d<\/em>. var. tettelbachianae affinis A. Schmidt 1856 = schmidtiana<\/em> Clessin 1888 had the result that this form from \u201cStyria\u201d is corresponding with d<\/em>. huettneri.<\/em> The latter name, however, is not replaced, because the exact locality of C<\/em>. d<\/em>. var. schmidtiana<\/em> is unknown.
\nC<\/em>. d<\/em>. schlechtii<\/em> has the same sculpture as d<\/em>. tettelbachiana<\/em>, but is larger and therefore more slender. It contains two different forms. In the form from the type locality Stixenstein the terminal lobe of clausilium plate is more or less upbent and pointed (form transitional to d<\/em>. vindobonensis<\/em>). In the form from the lower part of H\u00f6llental (between Hirschwang and Singerin) it is not upbent (like in d<\/em>. huettneri<\/em>). A. Schmidt (1856: 36) gave for C<\/em>. d<\/em>. var. schlechtii<\/em> a clausilium plate, which is described as \u201c… flache Rinne, … vorn abgerundet\u201d, i. e., like that of the H\u00f6llental form. Therefore, both forms are provisionally not separated as subspecies.
\nC<\/em>. d<\/em>. speciosa<\/em> sensu Klemm from H\u00f6llental and the Rax area belongs to C<\/em>. d<\/em>. huettneri<\/em> or C<\/em>. d<\/em>. schlechtii<\/em>, depending on the sculpture.
\nThe characters of C<\/em>. d<\/em>. kaeufeli <\/em>Klemm 1960 from Hochschwab mountain, N. E. Styria, which according to Klemm shall differentiate it from d<\/em>. tettelbachiana<\/em> (shell shape, sculpture, aperture form, formation of clausilium plate) are useless, because they are also found in that subspecies. E. g., the tettelbachiana<\/em> form from Schauerkogel, Schneealpe (see Pl. 2, Fig. 13), is more distinctly rib-striated (a character differentiating d<\/em>. kaeufeli<\/em>) than specimens of other samples of the subspecies. Therefore, kaeufeli<\/em> is regarded as a synonym of that subspecies This is confirmed by the fact that Klemm could pick kaeufeli<\/em> specimens out of tettelbachiana<\/em> samples from Schneealpe, Schneeberg and Rax mountains. On the material from the other localities farther westwards given by Klemm (1960) as kaeufeli<\/em> cannot be judged, because they could not be examined.<\/p>\n

Comment on the material used by Mason et al. 2020:
\nMaterial collected in more than 30 localities of Lower Austria has been determined as two dubia<\/em> subspecies, in some cases even as three.
\nThe sample lists are commented as follows:
\nd<\/em>. dubia<\/em> = d<\/em>. vindobonensis<\/em>.
\nLists of d<\/em>. huettneri<\/em>, d<\/em>. schlechtii<\/em> and d<\/em>. gracilior<\/em> (= form of d<\/em>. dubia<\/em>) each include also the other two named subspecies.
\nd. kaeufeli<\/em> = d<\/em>. tettelbachiana<\/em>.
\nd<\/em>. obsoleta<\/em> includes d<\/em>. vindobonensis<\/em> and gracilior<\/em> form of d<\/em>. dubia<\/em>.
\nd<\/em>. runensis<\/em> = small forms of d<\/em>. vindobonensis<\/em>.
\nd<\/em>. speciosa<\/em> = d<\/em>. schlechtii.
\n<\/em>The final remarks of Mason et al. on their subspecies (: 49) are given as follows: \u201e … C<\/em>. d<\/em>. dubia<\/em>, C<\/em>. d<\/em>. moldanubica <\/em>and C<\/em>. d<\/em>. vindobonensis<\/em> are very similar and their ranges overlap widely. Likewise, the … ranges of the quite similar C<\/em>. d<\/em>. kaeufeli<\/em> and C<\/em>. d<\/em>. tettelbachiana<\/em> overlap to some extent and they occurred frequently at the same locality. Thus, uniting them into one subspecies might be considered.\u201c
\nThese subspecies sensu Mason et al. belong to one and the same subspecies (C<\/em>. d<\/em>. vindobonensis<\/em> or C<\/em>. d<\/em>. tettelbachiana<\/em>, respectively).
\n\u201eThe same is true for C<\/em>. d<\/em>. huettneri<\/em>, C<\/em>. d<\/em>. schlechtii<\/em> and C<\/em>. d<\/em>. speciosa<\/em> …\u201c.
\nC<\/em>. d<\/em>. speciosa<\/em> sensu Mason et al. belongs to C<\/em>. d<\/em>. schlechtii<\/em>. C<\/em>. d<\/em>. huettneri<\/em> and C<\/em>. d<\/em>. schlechtii<\/em> sensu Mason et al., if collected at the same place, are confused.
\nI have tried to improve the COI gene tree of Mason et al. (: fig. 3) by correcting the determinations, based on my knowledge of the species from the respective localities. The result was as follows: Now as before, the subspecies are randomly distributed among the clades, and the geographic regions are not revealed. But several (14) subclades with haplotypes of specimens coming from more than one collected sample include only one (or two) subspecies, that is four subclades from the north and east only d<\/em>. vindobonensis<\/em>, five from the Schneeberg-Rax region only d<\/em>. huettneri <\/em>+ d<\/em>. schlechtii<\/em> and five from Schneeberg mountain only d.<\/em> tettelbachiana<\/em>. This result shows at least a partial separation of these subspecies and besides the close relationship of d<\/em>. huettneri <\/em>and d<\/em>. schlechtii<\/em>.<\/p>\n

Material
\nThe material given as follows is mainly deposited in the collection of the Forschungsinstitut Senckenberg (SMF).<\/p>\n

d<\/em>. vindobonensis<\/em><\/p>\n

(normal forms):<\/p>\n

A, Liechtenstein ruin near M\u00f6dling (topotypes vindobonensis<\/em>, SMF 31899):
\nR1<\/sub> mean (7) 9.0.<\/p>\n

A, Wien (Dornbacher Park) (SMF 163024):
\nR1<\/sub> mean (7) 7.8.<\/p>\n

A, Glei\u00dfenfeld near Seebenstein (street towards Sollgraben) (SMF 356627):
\nR1<\/sub> mean (5) 7.8.<\/p>\n

A, Aggstein ruin \/ Wachau (paratypes moldanubica<\/em>, SMF 163041):
\nR1<\/sub> mean (12) 9.8.<\/p>\n

A, Hangender Stein near Bad Fischau (SMF 86547):
\nR1<\/sub>:mean (10) 10.95.<\/p>\n

A, Hermannsh\u00f6hle near Kirchberg am Wechsel (SMF 356613):
\nClausilium plate in part not upbent.
\nR1<\/sub> mean (8) 10.25<\/p>\n

A, G\u00f6sting ruin near Graz (SMF 31940):
\nR1<\/sub> mean (9) 8.0.<\/p>\n

A, Glanegg ruin near St. Veit (SMF 130331):
\nClausilium plate in part not upbent.
\nR1<\/sub> mean (12) 7.4.<\/p>\n

A, Wei\u00dfe Wand near Launsdorf (SMF 31906):
\nClausilium plate in part not upbent.
\nR1<\/sub> mean (12) 8.9.<\/p>\n

A, Prossegg-Klamm near Matrei (SMF 182981):
\nR1<\/sub> mean (10) 8.1.<\/p>\n

I, lower Villn\u00f6ss valley near Klausen (SMF 130164):
\nClausilium plate in part not upbent.
\nR1<\/sub> mean (12) 6.8.<\/p>\n

D, Passau (Oberhaus) (SMF 264 818):
\nR1<\/sub> mean (12) 8.9.<\/p>\n

D, Granitz between Binz and Sellin \/ R\u00fcgen (SMF 129822):
\nR1<\/sub> mean (4) 7.5.<\/p>\n

RUS, Warnicken = Lesnoje \/ Samland (SMF 86673):
\nR1<\/sub> mean (10) 8.1.<\/p>\n

LT, Girstupis gorge near Kaunas (SMF 64734):
\nR1<\/sub> mean (12) 7.7.<\/p>\n

SK, Tren\u010dianske Teplice (Mount Klecpacs) (trencsiniensis<\/em>, SMF 31275):
\nR1<\/sub> mean (12) 9.65.<\/p>\n

SK, Manin gorge near Tren\u010din (manina<\/em>, SMF 31276):
\nR1<\/sub> mean (9) 9.1.<\/p>\n

SK, Igl\u00f3 = Spi\u0161sk\u00e1 Nov\u00e1 Ves (SMF 130377):
\nClausilium plate in part not upbent.
\nR1<\/sub> mean (6) 7.7.<\/p>\n

SK, Sz\u00e1dell\u00f6 = Zadiel gorge near Tur\u0148a (SMF 87189):
\nRibbing weakened.
\nR1<\/sub> mean (4) 9.1.<\/p>\n

PL, Panie\u0144skie Ska\u0142y near Krakau = Krak\u00f3w (SMF 130313):
\nR1<\/sub> mean (12) 7.9.<\/p>\n

UA, Czastowska skala near Lemberg = Lviv (podolica<\/em>, SMF 199966):
\nR1<\/sub> mean (10) 6.8.<\/p>\n

HU, Budapest, Fehetefaj (SMF 264754):
\nR1<\/sub> mean (8) 7.0.<\/p>\n

RO, Hunedoara (Castel) (transilvanica<\/em>, SMF 323455):
\nR1<\/sub> mean (10) 7.2.<\/p>\n

RO, Poiana Galdei near Teiu\u015f (towards \u00centregalde) (SMF 323469):
\nR1<\/sub> mean (10) 7.1.<\/p>\n

RO, Cheile Runcului near Ocoli\u015f (SMF 327352):
\nR1<\/sub> mean (10) 6.9.<\/p>\n

RO, Valea Rud\u0103reasa near Ciungetu 6 km above branch towards mountain (SMF 353664):
\nR1<\/sub> mean (10) 9.55.<\/p>\n

RO, between Moroeni and Z\u0103noaga \/ Bucegi mountains, 7.1 km from Moroeni (SMF 353615):
\nR1<\/sub> mean (10) 6.6.<\/p>\n

RO, Valea \u015eipoaia \/ Piatra Mare mountains (upper part, ca. 1300 m a. s. l.) (SMF 324514):
\nClausilium plate in part not upbent.
\nR1<\/sub> mean (8) 7.5.<\/p>\n

BIH, Trebesin near Sarajevo (1500-1600 m a. s. l.) (SMF 199987):
\nR1<\/sub> mean (8) 9.4.<\/p>\n

RKS, Stra\u017ea near Ka\u010danik (on street Tetovo-Doganovi\u0107) (coll. Nordsieck 5516):
\nR1<\/sub> mean (4) 9.5.<\/p>\n

MK, Ljuboten \/ \u0160ar mountains (1850-2000 m a. s. l.) (SMF 31284):
\nR1<\/sub>: 11.<\/p>\n

(dwarf forms):<\/p>\n

A, Steinberg near Grillenberg-Steinhof (paratypes steinbergensis<\/em>, SMF 323346):
\nR1<\/sub>: 12, 14.
\n(SMF 356658):
\nClausilium plate in part not upbent.
\nR1<\/sub> mean (10) 13.15.<\/p>\n

A, Matterleitner Hoher Stein near Rein (syntypes runensis<\/em>, SMF 86540):
\nR1<\/sub> mean (5) 13.2.<\/p>\n

d<\/em>. speciosa<\/em>:<\/p>\n

A, Rabenstein ruin \/ Lavant valley (SMF 163026):
\nRibbing weakened.
\nR1<\/sub> mean (6) 10.3.<\/p>\n

d<\/em>. gobanzi<\/em>:<\/p>\n

A, Badlgraben near Peggau (f. magna<\/em>, SMF 86574, 163025):
\nNearly smooth.
\nR1<\/sub> mean (18) 10.3.<\/p>\n

A, lower B\u00e4rensch\u00fctz valley near Mixnitz (SMF 86602):
\nLike preceding.
\nR1<\/sub> mean (8) 14.4.<\/p>\n

d<\/em>. otvinensis<\/em>:<\/p>\n

A, Otwinskogel near Launsdorf (SMF 163033):
\nR1<\/sub> mean (7) 5.2.<\/p>\n

d<\/em>. grimmeri<\/em>:<\/p>\n

A, Tollgraben near Graz (SMF 163034):
\nTransitional to d<\/em>. vindobonensis<\/em>.
\nR1<\/sub> mean (4) 6.9.<\/p>\n

A, Jungfernsprung near Deutsch-Feistritz (SMF 31909):
\nR1<\/sub> mean (10) 5.0.<\/p>\n

A, right Mur bank near Peggau (SMF 199947):
\nR1<\/sub> mean (8) 5.2.<\/p>\n

d<\/em>. floningiana<\/em>:<\/p>\n

A, Floningalpe near Kapfenberg (SMF 264778):
\nClausilium plate not upbent.
\nR1<\/sub> mean (12) 6.1.<\/p>\n

d<\/em>. bucculenta<\/em>:<\/p>\n

A, Innerschildgraben near Scheiblingkirchen (paratypes bucculenta<\/em>, SMF 163038):
\nR1<\/sub> mean (12) 4.2.<\/p>\n

d<\/em>. carpathica<\/em>:<\/p>\n

SK, Mount Kraszina near Tren\u010din (SMF 86643):
\nR1<\/sub> mean (6) 4.9.<\/p>\n

SK, Rajec-Teplice near Tren\u010din (f. minima<\/em>, SMF 86592):
\nR1<\/sub> mean (6) 6.6.<\/p>\n

d<\/em>. \u201ecostata<\/em>\u201c:<\/p>\n

SK, Szuly\u00f3 = S\u00fal’ovsk\u00e9 sk\u00e1ly near \u017dilina (f. costata<\/em>, SMF 86594):
\nClausilium plate not upbent.
\nR1<\/sub> mean (10) 5.2.<\/p>\n

d<\/em>. ingenua<\/em>:<\/p>\n

SK, Spi\u0161ska Kotlina near Drevenik \/ Spi\u0161sk\u00e9 Podhradie (SMF 199954):
\nR1<\/sub> mean (4) 3.2.<\/p>\n

d<\/em>. grimmeri<\/em> sensu Bielz = gratiosa<\/em>:<\/p>\n

RO, Cheile Turzii near Turda (paratypes gratiosa<\/em>, SMF 191281):
\nR1<\/sub>: 6\u2154, 5, 5\u00bc.<\/p>\n

RO, Mada near Geoagiu (SMF 323459):
\nR1<\/sub> mean (10) 4.9.<\/p>\n

RO, Pe\u015ftera Bolii near Petro\u015fani (SMF 86619):
\nR1<\/sub> mean (12) 5.0.<\/p>\n

d<\/em>. dubia<\/em><\/p>\n

(intermediary forms):<\/p>\n

DK, M\u00f6ens Klint (SMF 264829):
\nR1<\/sub>: 8\u00bc, 8, 8.<\/p>\n

SLO, Alpe Ursula = Ur\u0161lja gora near Slovenj Gradec (SMF 31929):
\nR1<\/sub> mean (10) 9.2.<\/p>\n

SLO, Alpe Saplata = Zaplata \/ Savinja Alps (didyma<\/em>, SMF 130589):
\nR1<\/sub> mean (8) 10.6.<\/p>\n

A, Wildensteiner Graben \/ Obir mountain (SMF 62582):
\nR1<\/sub> mean (7) 9.1.<\/p>\n

A, Hochobir \/ Obir mountain (dydima<\/em>, SMF 130060):
\nR1<\/sub> mean (8) 9.9.<\/p>\n

SLO, Suha valley near H\u00f6flein = Preddvor (SMF 130057):
\nR1<\/sub> mean (10) 10.2.<\/p>\n

A, Wurzen pass 4 km above branch of Wurzen street (SMF 312696):
\nR1<\/sub> mean (10) 9.55.<\/p>\n

SLO, Smrekova draga \/ Trnovski gozd (SMF 129793):
\nR1<\/sub>: 7, 8, 12.<\/p>\n

I, Raibl = Cave del Predil (SMF 130372):
\nR1<\/sub>: 11, 8, 8\u00bd.<\/p>\n

I, Luscharigraben = Lussari near Tarvisio (SMF 31253):
\nR1<\/sub> mean (5) 9.0.<\/p>\n

I, Timau-Pramosio (~ 1100-1200 m a. s. l.) (SMF 31991):
\nR1<\/sub> mean (10) 8.6.<\/p>\n

I, Prags valley (SMF 264795):
\nR1<\/sub> mean (10) 8.1.<\/p>\n

I, Schlern (2565 m a. s. l.) (alpicola<\/em>, SMF 130163):
\nR1<\/sub> mean (5) 8.25.<\/p>\n

I, Borno \/ Valcamonica (syntypes? sordellii<\/em>, SMF 130240):
\nR1<\/sub>: 12, 10\u00bd, 12.<\/p>\n

I, Esino \/ Lecco (SMF 130247):
\nR1<\/sub> mean (10) 10.5.<\/p>\n

I, Porto Ceresio \/ Varese (syntypes? langobardica<\/em>, SMF 130252):
\nR1 <\/sub>mean (5) 13.7.<\/p>\n

(normal forms):<\/p>\n

F (lectotype, coll. Draparnaud, Naturhistorisches Museum Wien, Bank & Gittenberger 2000: fig. 5):
\nR1<\/sub> ~ 12.<\/p>\n

S, Garpenberg \/ Dalarna (cemetery) (SMF 323906):
\nR1<\/sub> mean (4) 8.6.<\/p>\n

D, Xanten (eastern city wall) (SMF 264798):
\nR1<\/sub> mean (10) 11.7.<\/p>\n

D, N\u00fcrburg ruin \/ Eifel (SMF 264823):
\nR1<\/sub> mean (10) 9.4.<\/p>\n

D, K\u00f6nigstein ruin \/ Taunus (SMF 330654):
\nR1<\/sub> mean (10) 9.5.<\/p>\n

F, Odilienburg = Sainte-Odile \/ Vosges (SMF 335914):
\nR1 <\/sub>mean (12) 8.5.<\/p>\n

D, M\u00fchlheim \/ Donau (Wulfbach spring) (SMF 335850):
\nR1<\/sub> mean (12) 7.7.<\/p>\n

D, Jena (SMF 87278):
\nR1<\/sub> mean (10) 8.3.<\/p>\n

D, Berchtesgaden (St. Bartholom\u00e4, way to Eiskapelle) (SMF 130016):
\nR1<\/sub> mean (12) 8.5.<\/p>\n

CZ, Spitzstein near Saubsdorf = Supikovice (SMF 130288):
\nR1<\/sub> mean (12) 9.3.<\/p>\n

PL, Mount Czantor near Ustro\u0144 (SMF 87182):
\nR1<\/sub> mean (10) 8.9.<\/p>\n

SK, Vr\u00e1tna valley \/ Fatra mountains (pauperata<\/em>, SMF 130440):
\nR1<\/sub> mean (4) 10.1.<\/p>\n

A, upper Traisen valley between Freiland and Gsteinach (SMF 31901):
\nR1<\/sub> mean (10) 8.95.<\/p>\n

A, Kapfenberg (SMF 87225):
\nR1<\/sub> mean (12) 10.3.<\/p>\n

A, Hallstatt \/ Salzkammergut (SMF 86654):
\nR1<\/sub> mean (12) 7.6.<\/p>\n

CH, Via Mala (obsoleta<\/em>, SMF 130212):
\nR1<\/sub> mean (10) 9.4.<\/p>\n

CH, L’Abbaye (towards Les Bioux, 1004 m a. s. l.) (SMF 342559):
\nR1<\/sub> mean (12) 10.4.<\/p>\n

F, Grande Chartreuse (gallica<\/em>, dupuyana<\/em>, coll. Bourguignat):
\ngallica<\/em>: R1<\/sub> mean (10) 10.3, dupuyana<\/em>: R1<\/sub> mean (9) 10.1.<\/p>\n

F, Bar\u00e8ges \/ Hautes-Pyr\u00e9n\u00e9es (fagotiana<\/em>, SMF 129798):
\nR1<\/sub> mean (5) 10.4.<\/p>\n

F, Bar\u00e8ges \/ Hautes-Pyr\u00e9n\u00e9es (gallica<\/em>, coll. Bourguignat):
\nR1<\/sub> mean (17) 11.2.<\/p>\n

E, Valle de Ar\u00e1n \/ Lleida (SMF 317363):
\nR1<\/sub> mean (5) 12.4.<\/p>\n

UK, England, Cark \/ Lancashire (suttoni,<\/em> SMF 86672):
\nR1<\/sub> mean (4) 9.9.<\/p>\n

UK, England, Malham \/ Yorkshire (SMF 129726):
\nR1<\/sub>: 9\u00bc, 9, 8\u2153.<\/p>\n

d<\/em>. huettneri<\/em>:<\/p>\n

A, Springlessteig \/ Hohe Wand (paratypes huettneri<\/em>, SMF 264850):
\nRibbing weakened.
\nR1<\/sub> mean (9) 13.0.<\/p>\n

A, Gutenstein (SMF 86565):
\nLike preceding.
\nR1<\/sub> mean (9) 14.2.<\/p>\n

A, Eng at Schneeberg (SMF 31939):
\nLike preceding.
\nR1<\/sub> mean (13) 12.7.<\/p>\n

A, Hinterm Faden near Puchberg am Schneeberg (towards Puchberg) (SMF 356615):
\nLike preceding.
\nR1<\/sub> mean (6) 13.6.<\/p>\n

A, H\u00f6llental between Schwarzau im Gebirge and Singerin (SMF 31900):
\nLike preceding.
\nR1<\/sub> mean (11) 12.4.<\/p>\n

A, Na\u00dfwald near Schwarzau im Gebirge (N\u00e4gelegraben) (SMF 31941):
\nLike preceding.
\nR1<\/sub> mean (12) 13.0.<\/p>\n

A, \u201eStyria\u201c (types schmidtiana<\/em>, SMF 130052):
\nR1<\/sub> mean (8) 13.0.<\/p>\n

(forms transitional to d.<\/em> dubia<\/em>):<\/p>\n

A, Miesenbach near Puchberg am Schneeberg (street towards Lanzing) (SMF 356626):
\nR1<\/sub> mean (10) 10.45.<\/p>\n

A, Schottwien \/ Semmering (gracilior<\/em>, SMF 163031):
\nR1<\/sub> mean (7) 10.4.<\/p>\n

A, Klamm ruin near Schottwien (SMF 199944):
\nR1<\/sub> mean (8) 11.1.<\/p>\n

A, Semmering, between station and kurhaus 0.8 km behind branch towards Breitenstein (SMF 356636):
\nR1<\/sub> mean (8) 11.75.<\/p>\n

A, Semmering (pass, near to restaurant Erzherzog Johann, 1000 m a. s. l.) (SMF 86554, 130477):
\nR1<\/sub> mean (12) 11.3.<\/p>\n

d<\/em>. schlechtii<\/em>:<\/p>\n

A, Stixenstein near Ternitz (Schlo\u00dfpark) (SMF 163030):
\nClausilium plate like d<\/em>. vindobonensis<\/em>.
\nNearly smooth.
\nR1<\/sub> 15-20.<\/p>\n

A, Stixenstein near Ternitz (towards Puchberg am Schneeberg) (SMF 356614):
\nLike preceding.
\nR1<\/sub> mean (10) 16.45.<\/p>\n

A, Weichbach valley near Kaiserbrunn (behind entrance) (SMF 356653):
\nClausilium plate like d<\/em>. huettneri<\/em>.
\nNearly smooth.
\nR1<\/sub> mean (10) 15.5.<\/p>\n

d<\/em>. tettelbachiana<\/em>:<\/p>\n

A, Schneeberg, between Waxriegel and Kaiserstein (SMF 31934, 163032):
\nNearly smooth.
\nR1<\/sub>: 18-22.<\/p>\n

A, Rax, Jakobskogel (SMF 264861):
\nLike preceding.
\nR1<\/sub>: 15-19.<\/p>\n

A, Rax (summit) (SMF 265663):
\nLike preceding.
\nR1<\/sub>: 16-21.<\/p>\n

A, Schneealpe (1900 m a. s. l.) (SMF 199982):
\nLike preceding.
\nR1<\/sub>: 15-19.<\/p>\n

A, Kleinschwab \/ Hochschwab (2100 m a. s. l.) (types kaeufeli<\/em>, SMF 163042, 163043):
\nVery densely rib-striated.
\nR1<\/sub>: 13-15, holotype 16.<\/p>\n

Appendix: C<\/em>. dubia<\/em> in Austria, E. Styria:<\/p>\n

In E. Styria C<\/em>. dubia<\/em> exhibits a striking diversity (Figs. 1-14); this region on the edge of the East Alps is thus for the species a centre of endemism. In the region six subspecies could be traced, with two additional geographic forms: C<\/em>. d<\/em>. vindobonensis<\/em> (Figs. 1-2, with dwarf = runensis<\/em> form, Fig. 3), C<\/em>. d<\/em>. gobanzi<\/em> (Figs. 4-6), C<\/em>. d<\/em>. grimmeri<\/em> (Figs. 7-8), C<\/em>. d<\/em>. floningiana<\/em> (Fig. 9), C<\/em>. d<\/em>. dubia<\/em> (Figs. 10-11, with gracilior<\/em> form, Fig. 12) and C<\/em>. d<\/em>. tettelbachiana<\/em> (Figs. 13-14).
\nAccording to the examined material their distributional ranges are as follows:
\nC<\/em>. d<\/em>. vindobonensis <\/em>is distributed in the mountainous region on both sides of the lower Mur valley. Some samples, e. g. from Stra\u00dfengel and Wildon ruins near Graz (Fig. 1), are because of their clausilium plate transitional to d<\/em>. dubia<\/em>. The runensis<\/em> form was found on a mountain near Rein.
\nC<\/em>. d<\/em>. gobanzi<\/em> is distributed mainly on the left side of the lower Mur valley from Mixnitz to Peggau. In contrast to the information given by Klemm in the B\u00e4rensch\u00fctz valley near Mixnitz only C<\/em>. d<\/em>. gobanzi <\/em>(Fig. 6) could be traced.
\nC<\/em>. d<\/em>. grimmer<\/em>i has been found in the lower Mur valley on both sides of the Mur, C<\/em>. d<\/em>. floningiana<\/em> in and near to the upper Mur valley from Leoben to Kapfenberg. The grimmeri<\/em> form from Tollgraben, Graz (Fig. 7), is in sculpture transitional to C<\/em>. d<\/em>. vindobonensis<\/em>. There is also a sample of floningiana<\/em> collected by A. J. Wagner in Judenburg.
\nC<\/em>. d<\/em>. dubia<\/em> is distributed in the west, in the east up to the lower Mur valley (Frohnleiten) and the M\u00fcrz valley. The forms from Salza and M\u00fcrz valleys are in sculpture similar to the gracilior<\/em> form of C<\/em>. d<\/em>. dubia<\/em>, which has been traced on Semmering pass.
\nThe examined samples of C<\/em>. d<\/em>. tettelbachiana<\/em> (including typical C<\/em>. d<\/em>. kaeufeli<\/em> sensu Klemm) come from higher altitudes of Hochschwab and Schneealpe mountains (1600-2200 m a. s. l.). The forms differ in the strength of their rib-striation (Figs. 13-14).
\nAccording to my examination C<\/em>. d<\/em>. huettner<\/em>i and C<\/em>. d<\/em>. schlechtii<\/em> could not been traced in Styria with certainty. C<\/em>. d<\/em>. var. schmidtiana<\/em> from \u201eStyria\u201c = C<\/em>. d<\/em>. huettneri <\/em>(see Material) has no exact locality. C<\/em>. d<\/em>. huettneri<\/em> sensu Klemm from Styria, as far as examined, refers to C<\/em>. d<\/em>. dubia<\/em> (M\u00fcrz valley) or C<\/em>. d<\/em>. vindobonensis<\/em>, respectively (lower Mur valley). C<\/em>. d<\/em>. schlechtii<\/em> sensu Klemm from Styria (Schneealpe) belongs to C<\/em>. d<\/em>. tettelbachiana<\/em>.
\nIn no locality of the region more than one subspecies could be found. Several localities given by Klemm for the different subspecies are according to my examination based on wrong determinations.<\/p>\n

 <\/p>\n

\"\"<\/a><\/p>\n

\"\"<\/a><\/h4>\n
Plate 1
\nFigs. 1-6: Diversity of Clausilia dubia<\/em> in\u00a0 A, E. Styria.
\nShell frontal, body whorl dorsal and views into aperture; H = shell height (mm).; phot. S. Hof.
\nFig. 1: C<\/em>. dubia vindobonensis<\/em>, Wildon castle near Graz, ex SMF 130061, H 15.3.
\nFig. 2: C<\/em>. d<\/em>. vindobonensis<\/em>, G\u00f6sting ruin near Graz, ex SMF 31940, H 10.0.
\nFig. 3: C<\/em>. d<\/em>. vindobonensis<\/em> (dwarf form), syntype of runensis<\/em>, Matterleitner Hoher Stein near Rein, ex SMF 86540, H 7.8.
\nFig. 4: C<\/em>. d<\/em>. gobanzi<\/em>, Peggau ruin near Peggau, ex SMF 264763, H 13.5.
\nFig. 5: C<\/em>. d<\/em>. gobanzi<\/em>, Badlgraben near Peggau, ex SMF 86574, H 16.45.
\nFig. 6: C<\/em>. d<\/em>. gobanzi<\/em>, lower B\u00e4rensch\u00fctz valley near Mixnitz, ex SMF 86602), H 11.2.
\nPlate 2
\nFigs. 7-14. Continued.
\nFig. 7: C<\/em>. d<\/em>. grimmeri<\/em>, Tollgraben near Graz, ex SMF 163034, H 10.6.
\nFig. 8: C<\/em>. d<\/em>. grimmeri<\/em>, Jungfernsprung near Deutsch-Feistritz, ex SMF 31909, H 10.7.
\nFig. 9: C<\/em>. d<\/em>. floningiana<\/em>, Floning mountain near Kapfenberg, ex SMF 264778, H 10.8.
\nFig. 10: C<\/em>. d<\/em>. dubia<\/em>, Kapfenberg, ex SMF 87225, H 13.3.
\nFig. 11: C<\/em>. d<\/em>. dubia<\/em>, Eppenstein ruin near Judenburg, ex SMF 86578, H 11.4.
\nFig. 12: C<\/em>. d.<\/em> dubia<\/em> (gracilior<\/em> form), Semmering (pass, near restaurant Erzherzog Johann, 1000 m a. s. l.), ex SMF 86554, H 12.9.
\nFig. 13: C<\/em>. d<\/em>. tettelbachiana<\/em>, Schneealpe, Schauerkogel (1750-1788 m a. s. l.), ex SMF 31916), H 10.5.
\nFig. 14: C<\/em>. d<\/em>. tettelbachiana<\/em>, Hochschwab (1625-2278 m a. s. l.), ex SMF 130459), H 8.45.<\/h5>\n

III. Fossil record<\/h4>\n

C.<\/em> dubia<\/em> is frequent in Quaternary faunas. Only specimens from Early Pleistocene and early Middle Pleistocene deposits of Lower Austria (and Poland) and central \/ southern Germany (2.6 \u2013 0.4 Ma ago) have been examined.
\nSince the first appearance of the species (Early Pleistocene, 2.6 Ma ago) the major subspecies of C<\/em>. dubia<\/em> can be traced. The layers of Lower Austria yielded C<\/em>. d<\/em>. vindobonensis<\/em>, those of central \/ southern Germany C<\/em>. d<\/em>. dubia. <\/em>
\nThe samples are ordered according to age.
\nOnly the SMF numbers are given. The rest of the material (except of the samples from Kielniki 3A and Wei\u00dfenburg 7) is deposited in the Naturhistorisches Museum Wien.<\/p>\n

Early Pleistocene, A, Stranzendorf D, SMF 344014:
\nR1<\/sub>: 9, 8\u00bd.
\n= cf. d<\/em>. vindobonensis<\/em>.<\/p>\n

Only from the following samples the clausilium plate of specimens could be examined:<\/p>\n

Early Pleistocene, A, Stranzendorf I:
\nR1<\/sub>: 9.
\n= d<\/em>. vindobonensis<\/em>.<\/p>\n

Early Pleistocene, A, Krems-Schie\u00dfst\u00e4tte 7\/2:
\nR1<\/sub> mean (13) 9.9.
\nEarly Pleistocene, PL, Kielniki 3A:
\nR1<\/sub> mean 8.4.
\nEarly Pleistocene, A, Radlbrunn, SMF 341500:
\nR1<\/sub> mean (12) 8.4.
\n= d<\/em>. vindobonensis<\/em> (transitional to d<\/em>. dubia<\/em>).<\/p>\n

Early Pleistocene, A, Deutsch-Altenburg 37, SMF 344015:
\nR1<\/sub> mean (8) 9.4.
\nEarly Pleistocene, A, Deutsch-Altenburg 4B, SMF 344016:
\nR1<\/sub> mean (16) 8.3.
\n= d<\/em>. vindobonensis<\/em>.<\/p>\n

Middle Pleistocene, D, Wei\u00dfenburg 7:
\nR1<\/sub>: 10, 9, 9.
\n= d<\/em>. dubia<\/em>.<\/p>\n

Middle Pleistocene, A, Hundsheim:
\nR1<\/sub>: 6; 11, 8.
\n= d<\/em>. vindobonensis<\/em>.<\/p>\n

Middle Pleistocene, D, Wiesbaden (Mosbach III), SMF 334855:
\nR1<\/sub> mean (10) 7.8.
\nMiddle Pleistocene, D, Wiesbaden (Hessler), SMF 258296:
\nR1<\/sub> mean (12) 8.35.
\n= d<\/em>. dubia<\/em>.<\/p>\n

References<\/strong><\/p>\n

Bank, R. A. & Gittenberger, E. (2000): On the polytypic and problematic Clausilia dubia<\/em>: notes on its nomenclature and systematics (Gastropoda, Pulmonata, Clausiliidae). \u2013 Basteria, 64<\/strong>: 15-27.<\/p>\n

Bielz, E. A. (1867): Fauna der Land- und S\u00fcsswasser-Mollusken Siebenb\u00fcrgens. 2. Aufl. \u2013 VIII + 216 pp. Hermannstadt.<\/p>\n

Jaksch, K. (2012): Phylogeographie und Unterartklassifikation von Clausilia<\/em> dubia<\/em> Draparnaud, 1805 im \u00f6stlichen \u00d6sterreich (Gastropoda: Pulmonata: Clausiliidae): 93 pp. Diplomarbeit, Vienna University.<\/p>\n

K\u00e4ufel, F. (1929): Beitrag zur Kenntnis der spanischen Clausiliiden, gesammelt von Dr. F. Haas in den Jahren 1914-1917. \u2013 Senckenbergiana, 11<\/strong>: 13-21.<\/p>\n

Klemm, W. (1960): Clausilia dubia<\/em> Draparnaud und ihre Formen in \u00d6sterreich. \u2013 Archiv f\u00fcr Molluskenkunde, 89<\/strong> (1\/3): 81-109, pls. 10, 10a.<\/p>\n

Mason, K., Sattmann, H., Eschner, A., Duda, M., Harl, J., Kruckenhauser, L., Leeb, C. & Haring, E. (2020): Variation in mitochondrial sequences and shell morphology of Clausilia dubia <\/em>Draparnaud, 1805 (Gastropoda: Clausiliidae) in eastern Austria. \u2013 Arianta, 8<\/strong>: 38-57.<\/p>\n

Nordsieck, H. (1990): Revision der Gattung Clausilia<\/em> Draparnaud, besonders der Arten in SW-Europa (Das Clausilia rugosa<\/em>-Problem) (Gastropoda: Stylommatophora: Clausiliidae). \u2013 Archiv f\u00fcr Molluskenkunde, 119<\/strong> (4\/6): 133-179, 3 pls.<\/p>\n

Nordsieck, H. (2002): The subspecies classification of Clausilia dubia <\/em>Draparnaud (Gastropoda: Stylommatophora: Clausiliidae), a critical revision. \u2013 Mitteilungen der deutschen malakozoologischen Gesellschaft, 68<\/strong>: 37-44.<\/p>\n

Nordsieck, H. & Neubert, E. (2002): How to determine clausiliid species, I: Widespread species of Clausilia<\/em> Draparnaud (Gastropoda: Stylommatophora: Clausiliidae). \u2013 Mitteilungen der deutschen malakozoologischen Gesellschaft,\u00a0 68<\/strong>: 31-36.<\/p>\n

Schmidt, A. (1856): Die kritischen Gruppen der europ\u00e4ischen Clausilien. I. \u2013 VI + 63 pp., 11 pls. Leipzig.<\/p>\n

 <\/p>\n

\"\"<\/a><\/p>\n","protected":false},"excerpt":{"rendered":"

Clausilia dubia \u2013 a revision based on informative shell characters Hartmut Nordsieck (III. 2022) I. Introduction The subspecies division of C. dubia is difficult, because across the range of the species many forms of different rank are present. The previous revisions of the species (Klemm 1960, Nordsieck 1990, Bank & Gittenberger 2000, Jaksch 2012, Mason et al. 2020) are incomplete, because they are restricted to certain parts of the range. The only complete revision (Nordsieck 2002) is a survey on…<\/p>\n

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