Check-list of Recent Clausiliidae
Subspecies concept
Faunen Europas
Neniinae
Siciliaria Sicily
Dextral Clausiliidae
Clausiliidae Vietnam
Albinaria Crete
Papillifera bidens
Charpentieria itala
Cochlodina diversity
Systematics Clausilia
Alopia Dambovita
Alopia System
Revisoral remarks Phaedusinae
Systematics Baleinae
Plio-Pleistozän
Welter-Schultes 2012
Feher & al. 2013
Welter-Schultes 2010

Systematics of the genus Clausilia Draparnaud

Hartmut Nordsieck (VII.2013)

I. List of species taxa (with references):

The subspecies are given in alphabetical order.

Clausilia Draparnaud 1805

Clausilia (C.):

Diagnoses of species (Nordsieck 1990: 139-140, Nordsieck & Neubert 2002: 34-35), distribution (1990: 139-140, Nordsieck & Neubert 2002: 36);
list of taxa from S. W. Europe (1990: 164-166);
genital morphology of species (1990: 138-140).

C. rugosa (Draparnaud 1801) (1990: 145-152):
Distribution (: 145), subspecies (: 145-147).
r. andusiensis Coutagne 1886 (: 147), r. belonidea Bourguignat 1877 (: 148), r. lamalouensis Letourneux 1877 (: 148), r. magdalenica Salvañá 1887 (: 151), r. parvula A. Férussac 1807 (: 149-150), r. penchinati Bourguignat 1876 (: 150-151), r. pinii Westerlund 1878 (: 151-152), r. provincialis Coutagne 1886 (: 149), r. reboudii Dupuy 1850 (: 148-149), r. rugosa (: 147).

C. bidentata (Ström 1765) (1990: 157-160):
Distribution (: 157), subspecies (: 157).
b. abietina Dupuy 1849 (: 158), b. bidentata (: 157-158), b. crenulata Risso 1826 (: 159), b. moniziana Lowe 1852 (: 158-159).

C. cruciata Studer 1820 (1990: 160-162):
Distribution (: 160), subspecies (: 160).
c. amiatae E. Martens 1873 (: 161-162), c. bonellii E. Martens 1873 (: 161), c. pedemontana H. Nordsieck 1990 (: 160-161).

C. dubia Draparnaud 1805 (1990: 144-145, Nordsieck 2002: 38-42).
Distribution (1990: 145), taxa from S. W. Europe (1990: 144-145);
remaining taxa (2002: 38-39), main subspecies (2002: 39-40).
d. vindobonensis A. Schmidt 1856 sensu lato (: 40-41):
Coarsely ribbed forms (: 40-41, d. bucculenta Klemm 1960 (: 40), d. carpathica Brancsik 1888 [non Hazay] (: 40), d. „costata“ Brancsik 1888 (: 41), d. floningiana Westerlund 1888 (: 40), d. gratiosa Sajó 1968 (: 40-41), d. grimmeri L. Pfeiffer 1848 (: 40), d. ingenua Hudec & Brabenec 1963 (: 40), d. otvinensis M. Gallenstein 1895 (: 40), unnamed forms from Romania (: 40-41));
d. vindobonensis sensu stricto (: 40) (dwarf forms of d. vindobonensis (: 40, runensis Tschapeck 1883, steinbergensis Edlinger 2000), moldanubica Klemm 1960 = d. vindobonensis s. str. (: 40));
weakly ribbed forms (: 40, 42, d. gobanzi L. Pfeiffer 1868 (: 40), d. schlechtii A. Schmidt 1856 (: 42), d. speciosa A. Schmidt 1856 (: 40), unnamed forms from Slovakia (: 40)).
Intermediate forms (: 41-42):
d. alpicola Clessin 1878 (: 41), d. didyma F. J. Schmidt 1847 (: 42), d. tettelbachiana Rossmässler 1838 (: 42) (kaeufeli Klemm 1960 = d. tettelbachiana (: 42)), further intermediate forms (: 41-42).
d. dubia s. l. (: 41):
d. dubia s. str. (: 41) (fagotiana Bourguignat 1877 + suttoni Westerlund 1881 = d. dubia (: 41), pauperata Brancsik 1888 = d. dubia (: 41), gracilior Clessin 1888 [non Mousson] = d. dubia (: 41)), d. huettneri Klemm 1960 (: 41-42).

Clausilia (Strobeliella) H. Nordsieck 1977:

Diagnosis (Nordsieck 1966: 20-21);
distribution (1966: 23-24, Nordsieck 2013: 54), species (1966: 24, 2013: 52);
genital morphology of species (1966: 40-44).
brembina Strobel 1850: b. alanica H. Nordsieck 2013 (2013: 53), b. brembina (1966: 25, 27, 2013: 54), b. klemmi H. Nordsieck 1966 (1966: 30-31, 2013: 54);
exoptata A. Schmidt 1856 (1966: 29-30, 2013: 55);
umbrosa (Käufel 1928) (1966: 32, 2013: 55);
whateliana Charpentier 1850 (1966: 28-29, 2013: 55).

II. Revision of Clausilia cruciata and C. pumila:

The revision of C. cruciata (Nordsieck 1990) is incomplete, a revision of C. pumila is overdue. The shell morphology of about 50 samples of each species distributed over the whole range has been more thoroughly examined. The results are as follows:

1. Clausilia cruciata:
c. cruciata, whole range (except Italy): Whorls widely to more densely ribbed (rib number on 1 mm of the penultimate whorl = R1 5.7-8.6); outer cormer of clausilium plate not upbent. Several geographical forms (among others type form, cuspidata form) can be distinguished by their size and rib density.
In Italy the following subspecies occur:
c. amiatae Martens 1873, Monte Amiata: Like the following, but whorls more densely ribbed (R1 11.0-13.0);
c. bonellii Martens 1873, remaining Italy and Corsica, hardly different from the nominotypical subspecies: Whorls more widely to densely ribbed (R1 6.5-10); outer corner of clausilium plate more frequently upbent;
c. pedemontana H. Nordsieck 1990, Piedmont: Whorls densely ribbed (R1 9.0-14.0).
In cruciata n. subsp. from Vado di Sole, Gran Sasso d'Italia, the outer corner of clausilium plate is nearly always upbent.

2. Clausilia pumila:
Two main subspecies of C. pumila (Figs. 1-2) have been recognized. They have a peculiar mosaic distribution and can be regarded as species in statu nascendi:
p. pumila (Fig. 1) in the eastern and southern part of the range:
Superior lamella continuous with spiral lamella; outer corner of clausilium plate upbent.
p. sejuncta Mörch 1864 (Fig. 2) in Scandinavia, N. Germany, Russia and parts of the Baltic countries, Moravia and W. Slovakia, E. Austria, Slovenia:
Superior lamella separated from spiral lamella; outer corner of clausilium plate mostly not upbent.
The ribbing of both subspecies is the same (p. pumila R1 4.7-6.6, p. sejuncta 5.3-6.5).
C. p. sejuncta was regarded by O. Boettger (1878) and Westerlund (e. g., 1884) as independent species, by Likharev (1962) and Ložek (1964) as subspecies. Likharev (: 226-228, figs. 148-149) pointed to differences of both subspecies in genital morphology and to the remarkable distribution of p. sejuncta.
Some examined samples are intermediate between both main subspecies:
S. E. Bavaria and neighbouring Austria, W. Carinthia and Kanal valley, Styria, Lower Austria: Superior lamella continuous with spiral lamella; outer corner of clausilium plate like p. sejuncta.
Silesia: Superior lamella in part separated from spiral lamella; outer corner of clausilium plate like p. pumila.
A subspecies from Croatia, p. leptostoma A. Schmidt 1856 = p. sabljarii Brusina 1870, which is defined by its sculpture, is also intermediate between both main subspecies: Whorls widely or very widely ribbed (leptostoma form R1 4, sabljarii form R1 2.5); superior lamella separated from spiral lamella; outer corner of clausilium plate like p. pumila.


Figs. 1-2. Clausilia pumila.
Body whorl opened, contact of upper lamellae and outer corner of clausilium plate marked by arrows; phot. S. Hof. Shell size = H (mm).
1. C. p. pumila, A, Wien-Dornbach, ex SMF 323325, H = 10.1.
2. C. p. sejuncta, S, Västergötland, Fristad, Kröklings Hage, ex SMF 200079, H = 12.7. 

References

Boettger, O. (1878): Systematisches Verzeichniss der lebenden Arten der Landschnecken-Gattung Clausilia Drap. mit ausführlicher Angabe der geographischen Verbreitung der einzelnen Species. – Ber. Thätigkeit Offenbach. Ver. Naturk., 17/18: 18-101.

Likharev, I. M. (1962): Fauna SSSR, Molljuski, III, 4 (Clausiliidae). – 318 pp. Moskva-Leningrad (Akad. Nauk SSSR).

Lozek, V. (1964): Quartärmollusken der Tschechoslowakei. – Rozpravy Ústředniho ústavu geologického, 31: 374 pp., 32 pls.

Nordsieck, H. (1966): Zur Anatomie und Systematik der Clausilien, III. Clausilia whateliana und ihre Beziehungen zu den übrigen Clausilia-Arten, besonders zum Subgenus Neostyriaca. – Arch. Molluskenk., 95 (1/2): 19-47.

Nordsieck, H. (1990): Revision der Gattung Clausilia Draparnaud, besonders der Arten in SW-Europa (Das Clausilia rugosa-Problem) (Gastropoda: Stylommatophora: Clausiliidae). – Arch. Molluskenk., 119 (4/6): 133-179, 3 pls.

Nordsieck, H. (2002): The subspecies classification of Clausilia dubia Draparnaud (Gastropoda: Stylommatophora: Clausiliidae), a critical revision. – Mitt. dtsch. malakozool. Ges., 68: 37-44.

Nordsieck, H. (2013): Beschreibung einer neuen Clausilia-Unterart (Gastropoda, Pulmonata, Clausiliidae) von den Bergamasker Alpen (Italien), mit revisorischen Bemerkungen zur Untergattung Clausilia (Strobeliella). – Conchylia, 43 (1/4): 51-58.

Nordsieck, H. & Neubert, E. (2002): How to determine clausiliid species, I: Widespread species of Clausilia Draparnaud (Gastropoda: Stylommatophora: Clausiliidae). – Mitt. dtsch. malakozool. Ges., 68: 31-36.

Westerlund, C. A. (1884-1886): Fauna der in der palaearctischen Region (...) lebenden Binnenconchylien. IV. Gen. Balea Prid. & Clausilia Dr. – [1884]: VII+212+18 pp.; [1886]: +8 pp.; [1886]: +16 pp. Karlskrona.

Comment on Jaksch (2012): Subspecies of Clausilia dubia from Lower Austria

The paper of Jaksch deals with the subspecies classification of Clausilia dubia in Lower Austria. This classification is tested by shell morphology and a sequence of the mtDNA gene COI.
Criticism:
1. Morphology: Only shell morphology is used, genital morphology is not even mentioned. For shell morphology the characters of major taxonomic importance are not sufficiently considered, the development of the clausilium plate not at all, that of inferior lamella not thoroughly enough. The definition of subspecies and the so-called morpho-groups in which subspecies are assembled by the author is given mainly by external shell characters, which are of lower taxonomic value. As result in morpho-group I even members of both main subspecies of C. dubia (Nordsieck 2002) are included.
2. Taxonomy and nomenclature: The paper is based on the subspecies classification of Klemm (1960). The taxonomic and nomenclatural corrections, which were given since then (Bank & Gittenberger 2000, Nordsieck 2002), are not considered. C. d. dubia sensu Jaksch is C. d. vindobonensis, C. d. vindobonensis sensu Jaksch therefore the same. C. d. moldanubica is a synonym of C. d. vindobonensis. C. d. obsoleta sensu Jaksch is C. d. dubia. C. d. kaeufeli (from Schneeberg) is a synonym of C. d. tettelbachiana. The name C. d. gracilior cannot be used, because it is preoccupied. C. d. speciosa sensu Jaksch is not the same as C. d. speciosa A. Schmidt from Carinthia, C. d. runensis sensu Jaksch not the same as C. d. runensis Tschapeck from Styria.
3. COI analysis: As result of the COI analyses of Lower Austrian C. dubia five clades have come out. These clades do correspond neither to morphological nor to geographical units. This is not so astonishing, because by hybridization mtDNA genes can be forwarded from one morphological unit to another. Because several subspecies sensu Jaksch are or have been in the past sympatric or even syntopic, such hybridization often occurred. Therefore, all clades can include several of those subspecies. This is not discussed by the author.
4. Characters of Clausilia species: In the determination key of the Clausilia species of Lower Austria taxonomic important characters like development of subcolumellar lamella and clausilium plate are missing. The development of its subcolumellar lamella is the essential shell character by which C. dubia is distinguished from the other Clausilia species.

Jaksch, K. (2012): Phylogeographie und Unterartklassifikation von Clausilia dubia Draparnaud, 1805 im östlichen Österreich (Gastropoda: Pulmonata: Clausiliidae): 93 pp. Univ. Wien.

References

Bank, R. A. & Gittenberger, E. (2000): On the polytypic and problematic Clausilia dubia: notes on its nomenclature and systematics (Gastropoda, Pulmonata, Clausiliidae). – Basteria, 64: 15-27.

Klemm, W. (1960): Clausilia dubia Draparnaud und ihre Formen in Österreich. – Arch. Molluskenkunde, 89 (1/3): 81-109, pls. 10, 10a.

Nordsieck, H. (2002): The subspecies classification of Clausilia dubia Draparnaud (Gastropoda: Stylommatophora: Clausiliidae), a critical revision. – Mitt. dtsch. malakozool. Ges., 68: 37-44.


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