Cochlostoma (Clessiniella) waldemari ( Wagner, 1897 )

Zallot, Enrico, Mattia, Willy De, Fehér, Zoltán & Gittenberger, Edmund, 2021, Cochlostoma revised: the subgenus Clessiniella Zallot et al., 2015 (Caenogastropoda, Cochlostomatidae), European Journal of Taxonomy 762 (1), pp. 49-95 : 87-89

Cochlostoma (Clessiniella) waldemari ( Wagner, 1897)

Figs 1 , 3D , 14E , 15E , 16E , 31–33 , 34 (top), 35 (top), 36 (bottom)

Pomatias (Auritus) waldemari A.J. Wagner, 1897: 616 , pl. 9, 90 (“Ogulin”).

Cochlostoma waldemari – Welter-Schultes 2012: 103.

Diagnosis

Spotless shell. Moderately strong, more or less widely spaced ribs. Columellar lobe gently curved backwards. External lobe of type “II” in almost all the populations. The body spermiduct is huge and twisted instead of straight as in all the other species of Clessiniella .

Type material

Syntypes CROATIA • ♀; Ogulin; A.J. Wagner leg.; MIZPAS-W-6979a ( Fig. 33 ) • 2 ♀♀, 3 ♂♂; same collection data as for preceding; MIZPAS-W-6979/5 .

Other material

CROATIA • Kapela pass , Ogulin; 45.07534° N, 15.20837° E; 2009; De Mattia leg.; EZ0909 • Klek Mt , close to the top; 45.25904° N, 15.14044° E; 2009; Zallot leg.; EZ0904 • Klek Mt , cliffs halfway ; 45.25438° N, 15.14561° E; 1996; Zallot leg.; EZ0905 • Mt Risnjak ; 45.43095° N, 14.6188° E; 1400 m a.s.l.; 2009; De Mattia leg.; EZ0845 • Mt Risnjak ; 45.39072° N, 14.61748° E; 1000 m a.s.l.; 2010; Zallot leg.; EZ0907 • Ročko Polje ; 45.3607° N, 14.09534° E; 2009; De Mattia leg.; WdM6697 • Rudnica ; 45.23402° N, 15.33827° E; 2009; De Mattia leg.; EZ0906 • Spilja Vrelo ; 45.31777° N, 14.71065° E; 2009; Zallot leg.; EZ0899 • Vitunj-Ogulin ; 45.29114° N, 15.14041° E; 2005; Zallot leg.; EZ0903 .

SLOVENIA • Benete , 45.76192°N, 14.56752°E;2009; De Mattia leg.; WdM6805 • Bresovica ; 45.95607°N, 14.43533° E; 2006; De Mattia leg.; EZ0897 • Bresovica pri Predgradu ; 45.55164° N, 15.0437° E; 2009; De Mattia leg.; WdM6612 • Divje Jezero ; 45.9827° N, 14.0277° E; 2009; De Mattia leg.; WdM6840 • Dol-E of Čepovan ; 46.06445° N, 13.81189° E; 2009; De Mattia leg.; WdM6837 • Jelenja Vas ; 45.5115° N, 15.05346° E; 2009; De Mattia leg.; WdM6804 • Krempa Kolpa ; 45.53417° N, 15.09528° E; 1996; Mildner leg.; EZ0902 • Marija Ceplje Kolpa ; 45.53444° N, 15.09389° E; 2006; Mildner leg.; EZ0901 • Mt Snežnik 1150; 45.56913° N, 14.321° E; Jun. 2010; De Mattia leg.; WdM7014 • Mt Snežnik 1620; 45.58845° N, 14.44209° E; Jun. 2010; De Mattia leg.; EZ0963 • Mt Snežnik car parking ; 45.58235° N, 14.43117° E; Jun. 2010; De Mattia leg.; WdM7006 • Polog-E of Čepovan ; 46.06387° N, 13.81275° E; 2009; De Mattia leg.; WdM6842 • SW Pescena Glava; 46.05156° N, 13.73844° E; 843 m a.s.l.; 2009; De Mattia leg.; WdM6829 • Tisovec ; 45.81809° N, 14.76859° E; 2009; De Mattia leg.; WdM6749 • same collection data as for preceding; WdM6750 • Vrsic Pass ; 46.44278° N, 13.76417° E; 1985; Zallot leg.; EZ0896 • Zagradec- Polyane ; 45.86028° N, 14.83417° E; Bodon leg.; EZ0900 .

Description

SHELL. The shell is more or less light corneous brown and spotless. The size of the aperture varies considerably among the populations; shells with a large aperture and a correspondingly large lip occur next to shells with a poorly developed lip and a relatively small aperture. The lip is gently curved backwards on the columellar side.

FEMALE GENITALIA. The female genitalia are hardly distinguishable from those of Co. (Cl.) tergestinum if not for the more relaxed loops.

MALE GENITALIA. There is a long, inflated penis. The penial spermiduct is slightly twisted and occupies only the central part of the penis. The body spermiduct is twisted and conspicuously protruding (see Figs 34–35 ).

Distribution

The species inhabits the north-western and south-eastern part of Slovenia and an adjacent area in Croatia. It is reported from Austria for the Karawanken and the Gailtaler Alpen ( Edlinger & Mildner 1979). Its range largely surrounds that of Co. (Cl.) tergestinum (see Fig. 1 ).

Habitat preference

The snails are found on rocky cliffs as well as on boulders in woody areas.

Remarks

To assign a given population to Co. (Cl.) waldemari rather than to Co. (Cl.) tergestinum on the basis of the morphology of the shell and the female genitalia alone could be challenging if not impossible (see Figs 34–35 ). Only the structure of the male genitalia allows a convincing identification of the species. The H3 analysis ( Fig. 7 ) confirms that these sibling species belong to separate clades within Clessiniella .

Boeckel (1939) considered Co. (Cl.) waldemari a subspecies of Co. (Cl.) tergestinum . This view was contradicted by Bole (1994). In some areas, like on Mount Snežnik in southern Slovenia, the two species live not far from each other and here they can be distinguished also conchologically while differing in the presence or absence of the spots on the teleoconch (with Co. (Cl.) waldemari having a spotless shell) and in the shape of the body whorl approaching the aperture, in lateral view ( Fig. 36 ).

Figures

Fig. 1. Map of the sampling localities. Blue dots = Cochlostoma (Clessiniella) villae (Strobel, 1851); red dots = Co. (Cl.) stelucarum sp. nov.; green dots = Co. (Cl.) tergestinum (Westerlund, 1878); yellow dots = Co. (Cl.) waldemari (A.J. Wagner, 1897); orange dots = Co. (Cl.) anomphale Boeckel, 1939.

Fig. 3. Examples of protoconchs: the red dots mark the end of the protoconchs. A. Cochlostoma (Clessiniella) anomphale Boeckel, 1939 (EZ0034). A1. Co. (Cl.) anomphale (EZ0034): 1 = end of the first smooth part of the protoconch; 2 = end of the protoconch. B. Co. (Cl.) tergestinum (Westerlund, 1878) (EZ0799). C. Co. (Cl.) villae (Strobel, 1851) (EZ0885). D. Co. (Cl.) waldemari (A.J. Wagner, 1897) (WdM6697). E. Co. (Cl.) stelucarum sp. nov. (EZ0104).

Fig. 14. The five species of Clessiniella Zallot et al., 2015. A–D. Picture of a female (top) and male (bottom) specimen from the type localities. A. Cochlostoma (Clessiniella) anomphale Boeckel, 1939. Vellach, Austria (EZ0991). B. Co. (Cl.) stelucarum sp. nov., Cornino, Italy (EZ0104). C. Co. (Cl.) tergestinum (Westerlund, 1878), Val Rosandra, Italy (EZ0840). D. Co. (Cl.) villae (Strobel, 1851), Mizzole, Italy (EZ0878). E. Specimens from a locality close to the type one: Co. (Cl.) waldemari (A.J.Wagner, 1897), Mt Klek, Croatia (EZ0905).

Fig. 15.Comparison of femalegenitalia.A. Cochlostoma (Clessiniella) anomphaleBoeckel, 1939.Vellach, Austria (EZ0991). B. Co. (Cl.) stelucarum sp. nov., Cornino, Italy (EZ0104). C. Co. (Cl.) tergestinum (Westerlund, 1878), Val Rosandra, Italy (EZ0840). D. Co. (Cl.) villae (Strobel, 1851), Mizzole, Italy (EZ0878). E. Co. (Cl.) waldemari (A.J.Wagner, 1897), Mt Klek, Croatia (EZ0905).

Fig. 16.Comparisonof male genitalia.A. Cochlostoma (Clessiniella)anomphaleBoeckel, 1939.Vellach, Austria (EZ0991). B. Co. (Cl.) stelucarum sp. nov., Cornino, Italy (EZ0104). C. Co. (Cl.) tergestinum (Westerlund, 1878), Val Rosandra, Italy (EZ0840). D. Co. (Cl.) villae (Strobel, 1851), Mizzole, Italy (EZ0878). E. Co. (Cl.) waldemari (A.J.Wagner, 1897), Tisovec, Croatia (WdM6750).

Fig. 31. Shell variability in Cochlostoma (Clessiniella) waldemari (A.J. Wagner, 1897). A. Polog-E of Čepovan, Slovenia (WdM6842). B. Vrsic Pass, Slovenia (EZ0896). C. Benete, Slovenia (WdM6805). D. Mt Klek, Croatia (EZ0904). E. Bresovica pri Predgradu, Slovenia (WdM6612).

Fig. 32. Variation of the female genitalia in (A.J. Wagner, 1897). A. Polog-E of Čepovan, Slovenia (WdM6842). B. Dol-E of Čepovan, Slovenia (WdM6837). C. Divje Jezero, Slovenia (WdM6840). D. Tisovec, Slovenia (WdM6749). E. Vrsic Pass, Slovenia (EZ0896). F. Benete, Slovenia (WdM6805). G. Vitunin spring, Croatia (EZ0903). H. Mt Klek, Croatia (EZ0905).

Fig. 33. Cochlostoma (Clessiniella) waldemari (A.J. Wagner, 1897), syntype, ♀, at the Muzeum i Instytut Zoologii PAN, Warsaw, Poland (MIZPAS-W-6979a).

Fig. 34. Example of two populations of species of Clessiniella Zallot et al., 2015 which are impossible to distinguish without analyzing the male genitalia. Female shells. Top: Cochlostoma (Clessiniella) waldemari (A.J. Wagner, 1897), Polog-E of Čepovan, Slovenia (WdM6842). Bottom: Co. (Cl.) tergestinum (Westerlund, 1878), Jakačici, Croatia (WdM6699).

Fig. 7. Statistical parsimony network based on Histone H3 sequences of species in the subgenus Clessiniella Zallot et al., 2015. Cochlostoma (Cochlostoma) septemspirale (Razoumowsky, 1789) was used as an outgroup. For sample IDs we refer to Table 5.

Fig. 36. Top: Cochlostoma (Clessiniella) tergestinum (Westerlund, 1878), Mt Snežnik 1161 m a.s.l., Slovenia (WdM7012). Bottom: Co. (Cl.) waldemari (A.J. Wagner, 1897), Mt Snežnik 1150 m a.s.l., Slovenia (WdM7014).