Montenegrina subcristata (Pfeiffer, 1848)

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 : 100-105

Taxon classification Animalia Stylommatophora Clausiliidae

Montenegrina subcristata (Pfeiffer, 1848) Fig. 34 A–E

Clausilia subcristata Pfeiffer, 1848: 438. - Küster 1844-1862: 39-40, plate 4, figs 10-13. - Schmidt 1868: 70.

Clausilia cattaroensis (partim) - Walderdorff 1864: 509.

Clausilia cattaroensis var. minor Boettger, 1877b: 66.

Clausilia (Herilla) klecaki (sic!) Westerlund, 1881: 55.

Clausilia subcristata Clausilia (Delima) subcristata - Westerlund 1884: 54. - Clessin 1885: 182. - Wohlberedt 1907: 551-552. - Wohlberedt 1909: 675, plate 14, figs 158-161.

Clausilia (Delima) kleciaki - Westerlund 1884: 54.

Clausilia (Delima) wohlberedti Möllendorff, 1899: 169-170.

Clausilia (Delima) wohlberedti var. - Möllendorff 1899: 170.

Delima wohlberedti var. sublabiata v. Möll. - Wohlberedt 1901: 198, 206. (nomen nudum)

Clausilia (Delima) kleciaki var. brunnea Boettger 1907 in Wohlberedt 1907: 553. - Wohlberedt 1909: 676-677.

Clausilia (Delima) subcristata var. interior Boettger in Wohlberedt 1907: 552. - Wohlberedt 1909: 675, plate 14, fig. 162-163.

Clausilia (Delima) subcristata var. sublabiata Wohlberedt 1907: 553. - Wohlberedt 1909: 676.

Clausilia (Delima) subcristata f. minor - Wohlberedt 1907: 552.

Clausilia (Delima) subcristata wohlberedti - Wohlberedt 1907: 552-553. - Wohlberedt 1909: 675-676, plate 14, figs 164-169.

Delima (Delima) cattaroensis - Sturany and Wagner 1915: 73.

Delima (Delima) cattaroensis kleciaki - Sturany and Wagner 1915: 73.

Delima (Albanodelima) subcristata - Wagner 1924: 118. - Wagner 1925: 67, plate 2, fig. 23. (genital anatomy).

Delima (Albanodelima) kleciaki - Wagner 1924: 118. - Wagner 1925: 67, plate 3, fig. 26. (genital anatomy).

Delima (Albanodelima) subcristata wohlberedti - Wagner 1924: 119.

Delima (Albanodelima) subcristata interior - Wagner 1924: 119.

Clausilia laxa - Wagner 1924: 119.

Montenegrina subcristata - Nordsieck 1969: 259. (genital anatomy).

Montenegrina subcristata subcristata - Zilch 1981: 131, plate 13, fig. 19. - Nordsieck 2009: 73.

Montenegrina subcristata wohlberedti - Zilch 1981: 131, plate 13, figs 20-22. - Nordsieck 2009: 73.

Diagnosis.

Shell small to large, often ventricose, light corneous. All whorls smooth. Neck weakly inflexed, finely striate to moderately costate. Basal crest strong, peripheral crest well recognizable. Peristome attached, ovoid to angular, with somewhat swollen margin. Lamellae superior and spiralis do not or only barely overlap. In front view lamella inferior well emerged, medium-bent subcolumellaris mostly visible. Lunella lateral to ventrolateral, separate from the long to very long basalis. Subclaustralis is straight continuation of the lunella. Sulcalis strong. Long and strong anterior plica superior diverges from the principalis, remains separate from the lunella complex.

Dimensions

(in mm). Hs: 13.7-25.3 (lectotype brunnea 17.0, lectotype interior 25.3, holotype sublabiata 16.5, lectotype wohlberedti 22.8), Ws: 4.2-6.7 (lectotype brunnea 4.5, lectotype interior 6.7, holotype sublabiata 4.8, lectotype wohlberedti 5.8) (SMF 176323, SMF 176328, SMF 176308, SMF 176327, NHMW 110.430/MN/0138, NHMW 110.430/MN/0144, NHMW 110.430/MN/0145, HNMW 110430/MN/0119, NHMW 110430/MN/0157, HNHM 99489, NHMW 110.430/MN/0140, NHMW 38413, NHMW 110.430/MN/0143).

Type locality.

"in montibus Montenegrinis, 3000' supra mare" ( Pfeiffer 1848), more precisely by the collector ( Küster 1844-1862): "Grenzgebirge Montenegro’s gegen Cattaro" = Montenegro, E of the Bay of Kotor, ca. 900 m [ subcristata ]; “Albanien” [ minor ]; Niksič (due to subsequent lectotype designation) [ interior ]; "Dalmatia ad Cattaro" [ kleciaki ]; “Sabljach” = Montenegro, Žabljak Crnojevića (due to subsequent lectotype designation) [ wohlberedti ]; “Wirbasar” = Montenegro, Virpazar (in: Möllendorff 1899) [ sublabiata ]; "Skadar kastel" = Albania, Shkodër fortress (due to subsequent lectotype designation) [ brunnea ].

Type material.

Niksič, ex Boettger, ex. Wohlberedt, 1906, lectotype [ interior ] (SMF 176323), paralectotypes [ interior ] (SMF 176324/5); "zw. Rijeka und Komarno", ex Boettger, ex Wohlberedt 1906, paralectotype [ interior ] (SMF 176325); "Rijeka Quelle", ex Boettger, ex Wohlberedt 1906, paralectotype [ interior ] (SMF 176326); “Sabljack”, ex Möllendorff, ex Wohlberedt, lectotype [ wohlberedti ] (SMF 176328), paralectotypes [ wohlberedti ] (SMF 176329/11); "Festung Skutari", ex Boettger, ex Nägele, lectotype [ brunnea ] (SMF 176308), paralectotypes [ brunnea ] (SMF 176309/6); "Tarabos-Gebirge, Skutari", ex Boettger, ex Wohlberedt 1906, paralectotypes [ brunnea ] (SMF 176310/6); Mikulić, ex Boettger, ex Wohlberedt 1906, paralectotype [ brunnea ] (SMF 176311); "Dalmatien, Praesieka", ex Boettger, paralectotype [ brunnea ] (SMF 176312); “Virbazar”, ex Möllendorff, ex Wohlberedt, holotype [ sublabiata ] (SMF 176327).

Other material.

Montenegro, Zatrijebac, leg. Sturany, 24.iv.1905 (NHMW 43362); “Mužura Planina bei Dulcigno" (= Ulcinj), leg. Winneguth, 1904 (NHMW 41258); Rumija Mts, Đuravci, 340 m, 42.1677°N, 19.1959°E, leg. ZF, TK, DM, 16.vi.2012 (HNHM 99484); 12 km S of Virpazar, along the road to Petrovac, 570 m, 42.2188°N, 19.0246°E, leg. LD, ZF, JK, DM, 14.x.2008 (HNHM 99537); N of Tuđemili along the Bar to Virpazar road, 410 m, 42.1410°N, 19.1356°E, leg. MD, EH, KJ, HS, 7.vii.2015 (NHMW 110430/MN/0156); same locality, leg. ZE, ZF, 19.iv.2000 (HNHM 83481); near the Sutorman Pass, 760 m, 42.1568°N, 19.0996°E, leg. MD, EH, KJ, HS, 7.vii.2015 (NHMW 110430/MN/0157); Sutorman Pass, 42.1558°N, 19.1025°E, leg. TN, 1.vi.2014 (HNHM 99647); 9 km S of Virpazar, along the road to Bar, 440 m, 42.1912°N, 19.1081°E, leg. LD, ZF, JK, DM, 14.x.2008 (HNHM 99648); Virpazar, fortress, 42.244°N, 19.093°E, leg. ZE, ZF, 19.iv.2000 (HNHM 95414); Virpazar, 42.2460°N, 19.0917°E, leg. TD, ZE, ZF, 25.v.2015 (NHMW 110.430/MN/0122); 1 km E of Godinje, 40 m, 42.2205°N, 19.1241°E, leg. TD, ZE, ZF, 25.v.2015 (NHMW 110.430/MN/0123); 3.5 km SW of Virpazar, near Gluhi do, 42.2256°N, 19.0659°E, leg. TD, ZE, ZF, 25.v.2015 (NHMW 110.430/MN/0124); 0.5 km N of Virpazar, 42.2513°N, 19.0873°E, leg. TD, ZE, ZF, 25.v.2015 (NHMW 110.430/MN/0125); 1.3 km N of Virpazar, 42.2521°N, 19.0875°E, leg. TD, ZE, ZF, 25.v.2015 (NHMW 110.430/MN/0126); 3.5 km N of Virpazar, Vidikovac Observation Point, 150 m, 42.2547°N, 19.0896°E, leg. TD, ZE, ZF, 25.v.2015 (NHMW 110.430/MN/0127); Poseljani, 150 m, 42.3058°N, 19.0481°E, leg. TD, ZE, ZF, 25.v.2015 (NHMW 110.430/MN/0128); Začir, Pečina u Pećkom brdu, leg. LP, PS, AS, 20.vii.1972 (HNHM 36936); Komarno, near the Jabukov do Pečina, leg. ZE, ZF, 19.iv.2000 (HNHM 95418); Obodska Pećina, W of Rijeka Crnojevića, 150 m, 42.3523°N, 19.0047°E, leg. TD, ZE, ZF, 25.v.2015 (NHMW 110.430/MN/0129); 6.5 km NW of Rijeka Crnojevića, along the road to Cetinje, 310 m, 42.3651°N, 18.9924°E, leg. TD, ZE, ZF, 26.v.2015 (NHMW 110.430/MN/0130); "Rijeka Brücke”, leg. Bischoff, ex Klemm, viii.1934 (NHMW 21644); Ulići junction at the Podgorica to Cetinje road, leg. Kiss, LP, 9.vii.1985 (HNHM 43124); 1 km E of the Ulići junction, 470 m, 42.3815°N, 19.0084°E, leg. TD, ZE, ZF, 26.v.2015 (NHMW 110.430/MN/0131); Dobrska Župa junction, 250 m, 42.3865°N, 19.0571°E, leg. TD, ZE, ZF, 26.v.2015 (NHMW 110.430/MN/0132); 3 km from Barutan toward Rijeka Crnojevica, leg. LP, PS, AS, 20.vii.1972 (HNHM 36723); Cetinje, Donji Kraj District, 690 m, 42.4005°N, 18.9152°E, leg. TD, ZE, ZF, 26.v.2015 (NHMW 110.430/MN/0133); Mt. Lovćen, Njeguši, Popova Pečina, 42.4340°N, 18.8189°E, leg. LP, PS, AS, 23.vii.1972 (HNHM 36732, HNHM 37764); Njeguši, 910 m, 42.4267°N, 18.8314°E, leg. ZB, DP, 12.x.2003 (HNHM 94951); Njeguši, near the church, 880 m, 42.4316°N, 18.8126°E, leg. TD, ZE, ZF, 29.v.2015 (NHMW 110.430/MN/0149); Popova Pečina near Njeguši, leg. Sturany, 30.v.1903 (NHMW 38413); Nikšić District, W of Zagorak, 280 m, 42.6306°N, 19.0085°E, leg. ZB, DP, 7.x.2003 (HNHM 94911); Zeta Valley, ca. 2 km S of Nikšić along the road to Podgorica, 620 m, 42.7319°N, 18.9382°E, leg. TD, ZE, ZF, 30.v.2015 (NHMW 110.430/MN/0150); Podgorica District, Vitoja Spring, 42.3252°N, 19.3623°E, leg. AR, NR, PR, x.2013 (HNMW 110430/MN/0119); same locality, leg. MD, EH, KJ, HS, 8.vii.2015 (NHMW 110430/MN/0158); 2 km W of Božaj, 42.31°N, 19.37°E, leg. ZE, ZF, 30.vi.1996 (HNHM 91032); “Titograd” (= Podgorica), leg. Jakucs, 1966 (HNHM 95285); Podgorica, Mt. Ljubovic, eastern side, 42.4306°N, 19.2544°E, leg. MD, EH, KJ, HS, 8.vii.2015 (NHMW 110430/MN/0160); Šipčanik, 60 m, 42.3707°N, 19.3139°E, leg. TD, ZE, ZF, 25.v.2015 (NHMW 110.430/MN/0120); Žabljak Crnojevića, fortress, 50 m, 42.3172°N, 19.1567°E, leg. TD, ZE, ZF, 25.v.2015 (NHMW 110.430/MN/0121); Bar District, Dobra Voda, Tunel Ujtin, leg. Kiss, LP, 8.vii.1985 (HNHM 43121); 1 km S of Zaljevo, at Tunel Cafe, 90 m, 42.0572°N, 19.1278°E, leg. TD, ZE, ZF, 26.v.2015 (NHMW 110.430/MN/0138); 2 km S of Buljarica, 70 m, 42.1906°N, 18.9898°E, leg. TD, ZE, ZF, 28.v.2015 (NHMW 10.430/MN/0145); same locality, leg. ZE, ZF, 29.vi.1996 (HNHM 90880); Ulcinj District, Šasko Lake, 41.9768°N, 19.3387°E, leg. ZF, TK, DM, 16.vi.2012 (HNHM 99489); 1 km N of Gornja Kležna, 110 m, 42.0040°N, 19.2630°E, leg. TD, ZE, ZF, 27.v.2015 (NHMW 110.430/MN/0140); same locality, leg. PJ, TK, Magos, GP, 26.v.2013 (HNHM 99535); 5 km N of Zoganje, 80 m, 41.9794°N, 19.2591°E, leg. TD, ZE, ZF, 27.v.2015 (NHMW 110.430/MN/0139); 4 km S of Arbnež, 350 m, 42.0582°N, 19.3535°E, leg. TD, ZE, ZF, 28.v.2015 (NHMW 110.430/MN/0144); Albania, Selca (= Selcë), ex Fuchs (NHMW 111238); Maranaj north of Skutari, leg. Dabović (NHMW 27733); Shkodër District, Mt. Tarabosh, Vidhgar, 42.0572°N, 19.3962°E, leg. ZB, DP, 14.v.2013 (HNHM 99478); Vallas, 190 m, 42.0506°N, 19.4162°E, leg. TD, ZE, ZF, 27.v.2015 (NHMW 110.430/MN/0141); above Vallas, S of the Tarabosh Summit, 300 m, 42.0524°N, 19.4182°E, leg. TD, ZE, ZF, 27.v.2015 (NHMW 110.430/MN/0142); Shkodër, Rozafa Hill, S side, 42.0451°N, 19.4902°E, leg. TD, ZE, ZF, 28.v.2015 (NHMW 110.430/MN/0143); same locality, leg. ZB, ZF, CN, DP, 14.viii.2004 (HNHM 94509); same locality, leg. ZB, DP, 14.v.2013 (HNHM 99649); Zusi near Shkodër, 42.0386°N, 19.4811°E, leg. AR, NR, PR, v.2015 (NHMW 110.430/MN/0151); Lezhë District, Torovicë, 41.9000°N, 19.5130°E, leg. ZE, ZF, JK, DM, 21.x.2002 (HNHM 91612).

Distribution.

Western Montenegro and northwestern Albania (Fig. 35). This species has a wide range and by far the highest number of known localities within the genus. It occurs primarily in the region of the Skutari Lake, and is particularly frequent along its northern and western shores, as well as in the area between the lake and the Adriatic coast. To the northwest it reaches Cetinje and the Njeguš Plateau, and in the north it has isolated occurrences to the Zeta valley around Niksič. Its southernmost occurrence was found near Torovicë (in Albania), whereas the easternmost record is probably from the Mt. Maranaj near Shkodër. Wohlberedt’s (1907) records from farther east, namely the Prokletije Mts (Medun, Zatrijebac, Selcë, between Kolašin and Andrijevica) could not be confirmed by recent field studies ( Fehér et al. 2004, Murányi et al. 2011), therefore they cannot be accepted without reservation.

Remarks.

In the description the type locality of subcristata was only vaguely defined by Pfeiffer (1848). However, Küster’s (1844-1862) description provides important clues, mentioning that the material originated from the border area of Montenegro (with that of the Habsburg Monarchy), a site somewhere along the road from Kotor (to Cetinje) at an altitude of 3000 feet (ca. 900 m). Based on these data the type locality of the nominotypical subspecies can be in the vicinity of Njeguši, E of the Bay of Kotor.

Plate 4 of Küster’s monograph with figures of subcristata was published in 1847, whereas pages 39-40 with the description only in 1950. The plates contained no names, only accompanying sheets (wrappers) were issued along with the plates, so that subscribers and librarians would know which species were figured. These printed sheets were not meant to provide permanent scientific records. Due to the publication of Plate 4 and its wrapper, Pfeiffer could refer to subcristata in 1848, and attribute its authorship to Küster. However, ICZN Art. 8.1.1 specifies that for the availability of scientific names only the publication dates of text pages are relevant, and thus the wrappers are not acceptable as published work. Accordingly, Pfeiffer (1848) used the name first in proper form, therefore he should be considered the author ( Welter-Schultes 1999: 165).

In 1899 Möllendorff gave description of a variety from Virpazar, but without a name. Later Wohlberedt (1901) mentioned the name sublabiata Möllendorff, without providing a description or indication according to ICZN Art. 12 (nomen nudum). Later Wohlberedt (1909) published the name sublabiata together with Möllendorff’s (1899) description, but he also published these, in literal Serbian translation, two years earlier ( Wohlberedt 1907). Therefore, 1907 should be regarded as the year of publication and Wohlberedt as the author. This little-known Wohlberedt paper in Cyrillic script was probably overlooked or misunderstood by Zilch (1981), who referred to its sublabiata as nomen nudum.

In his 1907 and 1909 papers Wohlberedt clearly indicated that the descriptions of the new clausiliids are Boettger’s contributions, therefore the authorships of interior and brunnea are Boettger’s.

Westerlund (1881) described Clausilia (Herilla) klecaki , with this spelling. Later ( Westerlund 1884) he used this name as kleciaki , and subsequently all authors adopted, probably incorrectly, this spelling form. Based on the original description, kleciaki is identical with the typical subcristata . By contrast, Sturany and Wagner used the name kleciaki in a different way ( Sturany 1905, Sturany and Wagner 1915, Wagner 1924), for the form that was described by Boettger as brunnea (see also: Nordsieck 1972).

Intraspecific variability.

Nordsieck (2007, 2009) accepted only two valid subspecies, namely s. subcristata and s. wohlberedti . These differ primarily in the position of the lunella: subcristata having it ventrolaterally, whereas wohlberedti laterally. In this view, minor , interior and kleciaki are synonyms of the nominotypical form, whereas sublabiata and brunnea are synonyms of wohlberedti . The examined populations were highly variable in the size and shape of the shell, as well as in the position and shape of the lunella, allowing no clear delineation of two (or more) morphotypes. Moreover, considerable morphological heterogeneity could sometimes be observed within the same museum lots, or between lots from the same localities. Therefore, with the present knowledge, we find most appropriate classifying this diverse species without subspecific division.