Montenegrina Boettger, 1877

Feher, Zoltan & Szekeres, Miklos, 2016, Taxonomic revision of the rock-dwelling door snail genus Montenegrina Boettger, 1877 (Mollusca, Gastropoda, Clausiliidae), ZooKeys 599, pp. 1-137 : 6-8

Taxon classification Animalia Stylommatophora Clausiliidae

Genus Montenegrina Boettger, 1877

Montenegrina [ Clausilia , Sect. X. Delima , gruppe der cattaroensis ] Boettger 1877a: 37.

Clausilia (Heteroptycha) Westerlund, 1884: 5.

Montenegrina [ Clausilia (Delima) gruppe A.] - Westerlund 1884: 50.

Montenegrina [ Delima , group i.] - Kennard and Woodward 1923: 307.

Heteroptychia (sic!) - Kennard and Woodward 1923: 308.

Delima (Albanodelima) Wagner, 1924: 118. - Wagner 1925: 59.

Montenegrina [ Alopia (Delima) a.] - Lindholm 1924: 57.

Heteroptycha [ Alopia (Delima) b.] - Lindholm 1924: 57.

Delima (Montenegrina) - Zilch in Wenz 1960: 429-430.

Delima (Heteroptycha) - Zilch in Wenz 1960: 429-430.

Montenegrina (Heteroptycha) - Brandt 1961: 1.

Montenegrina (Montenegrina) s. str. - Brandt 1961: 3.

Montenegrina (Beieriella) Klemm, 1962: 242.

Montenegrina - Brandt 1962: 141. - Nordsieck 1969: 259. - Nordsieck 1972: 26. - Nordsieck 1974: 149. - Gittenberger 2002: 131. - Welter-Schultes 2012: 326.

Type species.

[ Montenegrina ]: cattaroensis -to our best knowledge, Kennard and Woodward (1923) were the first mentioning this as type species, but without pointing out subsequent designation; [ Heteroptycha ]: helvola -by monotypy, see Lindholm (1924); [ Albanodelima ]: umbilicata -subsequent designation by Nordsieck (2001: 20); [ Beieriella ]: irmengardis -by original designation.

Distribution.

Coastal regions of Montenegro south of the Bay of Kotor, Albania, western Macedonia, and northwestern Greece (Fig. 1.).

Ecology.

Montenegrina species inhabit rocky habitats, where they feed on the microflora and find hiding place in crevices or among and under boulders. All known populations are associated with limestone environments. The formation and shape of the habitable outcrops can vary considerably even within a species. Suitable habitats include small to large cliffs, gorges, rocky forests, rocky alpine grasslands, or even artificial stone walls along roads (Fig. 10).

Historical background.

The first few described taxa of Montenegrina were initially classified within the genus Clausilia Draparnaud, 1805 and, later on, in the subgenus Clausilia (Delima) Hartmann, 1842. In his Clausilia monograph Schmidt (1868) already distinguished Clausilia cattaroensis , Clausilia helvola , Clausilia laxa , and Clausilia subcristata as the group of Clausilia cattaroensis . The name Montenegrina was first used by Boettger (1877a) to designate the cattaroensis group within the subgenus Clausilia (Delima) . A formal description of Montenegrina was provided by Westerlund (1884), who included here all aforementioned taxa except Clausilia helvola . For this species, based on its special aperture structure, he established the subgenus Clausilia (Heteroptycha) , which was then elevated to genus rank by Kennard and Woodward (1923). Later Zilch (1960) considered Montenegrina and Heteroptycha as subgenera of Delima , whereas Albanodelima Wagner, 1924 as a synonym of Heteroptycha .

Based on genital morphology, Wagner (1924) was the first to unite all Montenegrina species known at that time within one subgenus, Delima (Albanodelima) Wagner, 1924. A description of this taxon, originally outlined only by a list of the involved species, was given in a subsequent publication of Wagner (1925). In this he defined three species groups within Delima (Albanodelima) . The umbilicata group included Delima (Albanodelima) umbilicata , Delima (Albanodelima) cattaroensis , Delima (Albanodelima) interior (Boettger, 1907), Delima (Albanodelima) kleciaki , Delima (Albanodelima) skipetarica ( Soós, 1924), Delima (Albanodelima) subcristata , Delima (Albanodelima) weigneri Poliński, 1924, and Delima (Albanodelima) wohlberedti ( Möllendorff, 1899). Wagner pointed out that he regarded part of Montenegrina (as used by Kennard and Woodward, 1923) synonymous with this species group. The attemsi group comprised Delima (Albanodelima) attemsi Wagner, 1914, Delima (Albanodelima) janinensis , Delima (Albanodelima) ochridensis Wagner, 1925, Delima (Albanodelima) perstriata Wagner, 1919, and Delima (Albanodelima) rugilabris . Finally, the group of helvola included Delima (Alpidelima) helvola and Delima (Albanodelima) apfelbecki (Sturany, 1907), with a note that Heteroptycha was considered a synonym of this group. In contrast to this concept, Zilch (1960) referred to Albanodelima as if it had been used by Wagner only for the helvola group, and thus would have been a synonym of Heteroptycha .

Montenegrina was first regarded a distinct genus by Brandt (1961). In the following year Klemm (1962) erected the subgenus Montenegrina (Beieriella) , which was later rejected by Nordsieck (1972), claiming that it had been based merely on distinctive characters of its type species, Montenegrina (Beieriella) irmengardis Klemm, 1962. In the same article Nordsieck also dismissed Heteroptycha as subgenus. Subsequent publications also do not use subgeneric division of Montenegrina ( Nordsieck 1969, 1972, 1974, 2009; Erőss et al. 1999, 2006; Welter-Schultes 2012), only species groups are distinguished in Nordsieck’s works.

Assessments of the genital morphology of Montenegrina were provided by Nordsieck (1969, 1972, 2009), based on examinations of Montenegrina cattaroensis , Montenegrina janinensis , Montenegrina rugilabris , Montenegrina irmengardis , Montenegrina stankovici ( Urbański 1960), Montenegrina subcristata and Montenegrina umbilicata samples (Nordsieck, 2009). Genitalia data were also given on Delima (Albanodelima) apfelbecki , Delima (Albanodelima) kleciaki and Delima (Albanodelima) subcristata by Wagner (1925), Clausilia (Delima) skipetarica by Soós (1924), Montenegrina (Beieriella) irmengardis by Klemm (1962), Delima (Montenegrina) perstriata ochridensis by Loosjes (1966), Montenegrina janinensis maasseni Gittenberger, 2002 by Uit de Weerd and Gittenberger (2004) and Montenegrina dofleini sinosi Páll-Gergely, 2010 in its description ( Páll-Gergely 2010). Data on the microarmature of the clausilium plate were published by Gittenberger (2002).

The most recent revision of the genus was provided by Nordsieck (2009). In this he distinguished eight species groups, namely those of Montenegrina cattaroensis , Montenegrina drimmeri Fehér & Szekeres, 2006, Montenegrina fuchsi Brandt, 1961, Montenegrina janinensis , Montenegrina laxa , Montenegrina perstriata , Montenegrina rugilabris , and Montenegrina skipetarica . By and large this concept was adopted in the Fauna Europaea checklist ( Bank 2013), which mentions 22 species of Montenegrina with 87 subspecies. In the present paper a revised system of Montenegrina is provided on the basis of the material available at the HNHM, NHMW and SMF collections, as well as that of newly discovered populations.