[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.

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

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).

Figure 10. 

Typical rocky habitats preferred by Montenegrina species. A Portitsa Gorge of the Venetikos River (type locality of M. hiltrudae maasseni) B rocky ridge of the Ostrovica Mts north of the Faqekuq Summit (type locality of M. sturanyana ostrovicensis ssp. n.) C valley of the Drin upstream of the Koman Dam (near the type locality of M. prokletiana kovacsorum ssp. n.) D roadside cliff near Zaljevo (M. subcristata) E ruins of the medieval fortress at Haj-Nehaj (M. cattaroensis umbilicata) F rocky forest in the valley of the Tropojë Stream (type locality of M. sporadica tropojana ssp. n.) G cliffs along the shore of Lake Prespa near the Sv. Marena church (M. dofleini prespaensis) H rocky grassland in the alpine region of the Galičica Mts south of the Bugarska Summit (M. dofleini dofleini).

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 C. cattaroensis, C. helvola, C. laxa, and C. subcristata as the group of C. cattaroensis. The name Montenegrina was first used by Boettger (1877a) to designate the cattaroensis group within the subgenus C. (Delima). A formal description of Montenegrina was provided by Westerlund (1884), who included here all aforementioned taxa except C. helvola. For this species, based on its special aperture structure, he established the subgenus C. (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 D. (Albanodelima). The umbilicata group included D. (A.) umbilicata, D. (A.) cattaroensis, D. (A.) interior (Boettger, 1907), D. (A.) kleciaki, D. (A.) skipetarica (Soós, 1924), D. (A.) subcristata, D. (A.) weigneri Poliński, 1924, and D. (A.) 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 D. (A.) attemsi Wagner, 1914, D. (A.) janinensis, D. (A.) ochridensis Wagner, 1925, D. (A.) perstriata Wagner, 1919, and D. (A.) rugilabris. Finally, the group of helvola included D. (A.) helvola and D. (A.) 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 M. (Beieriella), which was later rejected by Nordsieck (1972), claiming that it had been based merely on distinctive characters of its type species, M. (B.) 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 M. cattaroensis, M. janinensis, M. rugilabris, M. irmengardis, M. stankovici (Urbański 1960), M. subcristata and M. umbilicata samples (Nordsieck, 2009). Genitalia data were also given on Delima (Albanodelima) apfelbecki, D. (A.) kleciaki and D. (A.) 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 M. cattaroensis, M. drimmeri Fehér & Szekeres, 2006, M. fuchsi Brandt, 1961, M. janinensis, M. laxa, M. perstriata, M. rugilabris, and M. 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.

Shell small, light corneous. All whorls smooth, glossy. Neck inflexed, densely striate. Basal and peripheral crests well visible. Peristome detached, ovoid, with simple margin. Lamellae superior and spiralis do not overlap. In front view lamella inferior moderately emerged, weakly-bent subcolumellaris not visible. Lunella dorsal-dorsolateral. Basalis mostly absent, rarely residual and separate from the lunella. Subclaustralis very short, sulcalis well developed. Anterior plica superior absent. Clausilium plate almost entirely visible through the aperture.

(in mm). H_s: 12.6–16.0, W_s: 3.4–4.2 (syntypes NHMW 41166).

Figure 11. 

A Montenegrina apfelbecki (Sturany, 1907), syntype, NHMW 41166 B M. attemsi attemsi (Wagner, 1914), syntype, SMF 167038 C M. attemsi jakupicensis Fauer, 1993, holotype, SMF 309239 D M. cattaroensis cattaroensis (Rossmässler, 1835), lectotype, SMF 156967 E M. cattaroensis antivaricostata nom. n., lectotype of costata (Boettger, 1907), SMF 176320 F M. cattaroensis umbilicata (Boettger, 1879), lectotype, SMF 176318 G M. chiasma Nordsieck, 1972, paratype, NHMW 75000/-E 32343 H M. dofleini dofleini (Wagner, 1928), Tomoros, original series, ZSM 20150453 I M. dofleini fagorum Nordsieck, 1974, holotype, SMF 227680 J M. dofleini dofleini (Wagner, 1928), paratype of kaiseri, SMF 201563 K M. dofleini kastoriae Nordsieck, 1972, holotype, 221042 L M. dofleini pinteri Nordsieck, 1974, holotype, SMF 227684 M M. dofleini prespaenis Nordsieck, 1988, NHMW 84019 N M. dofleini sinosi Páll-Gergely, 2010, holotype, HNHM 97145 O M. dofleini wagneri Szekeres, 2006, holotype, NHMW 103375. Scale bar: 5 mm.

“Mal i Shêit bei Oroshi, Merdita, in einer Höhe von zirka 1500 m” = Albania, Mt. Shënt, ca. 1500 m.

Type locality, leg. Buljubašić, 1905, syntypes (NHMW 41166/pl., NHMW-E 38290/2, NHMW-K 9368/3, SMF 32771/2, SMF 201619); same data, ex Oberwimmer, syntypes (NHMW 65490/2), same data, ex Käufel, syntypes (NHMW 70409/3).

Mount Shënt (Mali i Shëntit) in Northern Albania (Fig. 12).

Figure 12. 

Distribution of Montenegrina apfelbecki. M. attemsi and M. chiasma. M. apfelbecki (star); M. chiasma (circle); M. attemsi attemsi (triangle); M. attemsi jakupicensis (square).

Of this taxon only the type series are known. The material was collected during the 1905 expedition organized by Rudolf Sturany, then curator of the Mollusc Collection at NHMW. Sturany was accompanied by Viktor Apfelbeck, curator of the entomological collection at the National Museum of Sarajevo, and Latif Buljubašić, Apfelbeck’s assistant. After a few days of joint work in the Oroshi area Buljubašić was left alone for two months to explore the surrounding Shënt, Munellë and Koritnik mountains (Sturany 1905). During that period Buljubašić acquired a large and diverse invertebrate material, but M. apfelbecki was found only at one site. This suggests that the species was highly local, although the discovered population could have been quite large. In 2014 a thorough search by the first author over the plateau of the Shënt Mountain, along its western slope and northern side toward the Mount Zebë, failed to re-discover the type locality of this species (Angyal et al. 2015). Apparently it is not frequent in the Shënt Mountain, and may only have a very narrow range.

Shell small, slender, light yellowish-corneous. All whorls smooth. Peristome attached, ovoid to angular, with somewhat swollen margin. Lamellae superior and spiralis overlap. In front view lamella inferior hidden or weakly emerged, subcolumellaris broadly-bent. Lunella dorsolateral to lateral, diffuse, not connected to the basalis. Subclaustralis absent or residual, sulcalis present. Anterior plica superior not connected to the lunella complex. Clausilium plate only barely visible through the aperture. Differs from similar-looking M. janinensis by the smaller, more elongate, entirely smooth shell, as well as the less emerged lamella inferior.

Neck striate to striate-costate. Basal crest strong, peripheral crest weak. In front view lamella inferior weakly emerged, broadly-bent subcolumellaris mostly visible. Basalis short, subclaustralis weak. Anterior plica superior mostly present.

(in mm). H_s: 12.0–14.8, W_s: 3.0–3.5 (Nordsieck 1974).

“Treska bei Üsküb” = Macedonia, Matka Gorge near Skopje.

Type locality, leg. Apfelbeck, ex Oberwimmer [10525], syntypes (NHMW 65491/3); same locality, ex Wagner, ex Käufel, syntypes (NHMW-E 9407/3); same locality, ex Käufel, syntypes (NHMW-K 9366/4); same locality, ex Wagner, ex Fuchs, ex Rušnov, ? syntypes (NHMW 70410/3); same locality, leg. Apfelbeck, ex Wagner, ex Brandt, syntype (SMF 167038/1); same locality, ex Käufel, ex Jaeckel, syntype (SMF 201603).

Macedonia, gorge of the Treska near Skopje, leg. Penther, 11.v.1918 (NHMW 52313); Skopje District, Matka Gorge near the Sv. Andreja Church, 300 m, 41.9610°N, 21.2943°E, leg. ZE, ZF, AH, 10.iv.2004. (HNHM 94429); same locality, leg, TD, 2008. (HNHM 99519).

The subspecies is known only from the valley of the Treska River (Matka Gorge) in northwestern Macedonia (Fig. 12).

Although the description was formally published in Sturany and Wagner (1915), a reprint of the paper had already been printed and circulated in 1914, and the authorship of the taxon has been unequivocally attributed to Wagner.

Neck smooth, basal and peripheral crests weak. In front view lamella inferior mostly hidden, broadly-bent subcolumellaris also not visible. Basalis mostly absent, subclaustralis absent or residual. Anterior plica superior mostly absent.

(in mm). H_s: 12.4–16.9 (holotype 14.2), W_s: 2.7–3.4 (holotype 2.9).

Macedonia, Jakupica Mts, Nežilovo, 6.8 km to Dom Čeplez, ca. 1170 m.

Type locality, leg. PS, 12.vii.1990, holotype (SMF 309239), paratypes (NMBE 535051/6, ZMH 91629/3); Nežilovo, 6.4 km towards Dom Čeplez, ca. 1120 m, leg. PS, 12.vii.1990, paratypes (SMF 309240/1, NMBE 535050/4, ZMH 91628/1, ZHM 91630/1).

Macedonia, Nežilovo, quarry ca. 10 km on the road to Dom Čeplez, 1230 m, 41.6810°N, 21.4303°E, leg. ZE, ZF, AH, 4.iv.2004. (HNHM 95358); Babuna Spring NW of Nežilovo, 1280 m, 41.6903°N, 21.4162°E, leg. DM, TK, 3.x.2013 (HNHM 99520).

Southern part of the Jakupica Mts in Macedonia. Known only from a few sites, within a narrow range northwest of Nežilovo village (Fig. 12).

Shell medium to large, light corneous. Whorls smooth or indistinctly costate. Neck weakly inflexed, basal and peripheral crests weak to well recognizable. Peristome attached, ovoid to angular, with somewhat swollen margin. Lamellae superior and spiralis mostly overlap. In front view lamella inferior well emerged, medium-bent subcolumellaris mostly visible. Lunella dorsal to dorsolateral, separate from the strong basalis. Subclaustralis well developed, sulcalis present. Anterior plica superior long, separate from the lunella complex. Clausilium plate partly visible through the aperture.

Shell mostly large, elongate. All whorls smooth. Neck finely costate. Basal and peripheral crests well recognizable. Lamellae superior and spiralis overlap. Lunella dorsolateral. Plica principalis often connected to the superior. Sulcalis well developed.

(in mm). H_s: 16.8–26.5 (lectotype 26.0); W_s: 4.3–5.9 (lectotype 5.8) (Kotor NHMW 110430/MN/0068, SMF 156967, SMF 221221). According to Westerlund (1884) the height and width of the smallest specimens (forma parvula) can be 15 mm and 3 mm, respectively.

“Cattaro” = Montenegro, Kotor.

“Dalmatien, Cattaro”, ex Rossmässler, lectotype (SMF 156967), paralectotype (SMF 5021/3, SMF 176307/5 SMF 221221/9)

Montenegro, Kotor, above the town, leg. Kiss, LP, 9.vii.1985 (HNHM 43127); Kotor, bus terminal, 42.4197°N, 18.7718°E, leg. ZE, ZF, 29.vi.1996 (HNHM 91037); Kotor, Škurda Gorge and N slope of the fortress hill, 80 m, 42.4269°N, 18.7756°E, leg. TD, ZE, ZF (HNHM 99601, NHMW 110430/MN/0068); Bay of Kotor, N of Dobrota, gorge of the Ljuta, 20 m, 42.4862°N, 18.7670°E, leg. LD, ZF, JK, DM, 8.x.2008 (HNHM 99599); Mt. Lovćen, gorge ca. 8 km from Kotor on the Kotor to Njeguši road, 400 m, 42.4024°N, 18.7750°E, leg. LD, ZF, JK, DM, 8.x.2008 (HNHM 99600); same locality, leg. ZE, ZF, 21.iv.2000 (HNHM 95412); same locality, leg. TD, ZE, ZF (HNHM 99602, NHMW 110430/MN/0069); 9 km from Kotor on the Kotor to Njeguši road, 450 m, 42.4095°N, 18.7787°E, leg. TD, ZE, ZF (HNHM 99603, NHMW 110430/MN/0070); “Risano über Bocche di Cattaro” (= Risan, above the Bay of Kotor) ex Käufel, ex Rušnov (NHMW Kau-70412); Bay of Kotor, Orahovac, 30 m, 42.4894°N, 18.7641°E, leg. TD, Domokos, Nacsa, Páll-Gergely, 5.iv.2006 (MMM-B01329).

This taxon lives at the foot and slopes of the mountains surrounding the Bay of Kotor between Risan and Kotor. We found it along the serpentine road from Kotor to the Mt. Lovćen up to 450 m. It is not syntopic with M. subcristata, which occurs somewhat farther on the Njeguši Plateau, above 800–900 m (Fig. 13).

Figure 13. 

Distribution of Montenegrina cattaroensis. M. cattaroensis cattaroensis (square); M. cattaroensis antivaricostata (empty circle with dot); M. cattaroensis umbilicata (circle).

The monograph of Küster (1844-1862) was published in parts over several years. Page 40, where the name gracilior was first mentioned, dates to 1850 (Welter-Schultes 1999).

The distribution record from Herceg-Novi (“Castelnuovo”) by Schmidt (1868) is probably erroneous, as during recent field trips no Montenegrina could be found in that area. Gluhi do, mentioned by Wohlberedt (1901), is a locality of M. subcristata.

Shell medium, tumid. All whorls and the neck with widely-spaced and indistinct wrinkle-ribs. Basal and peripheral crests well recognizable. Lamellae superior and spiralis mostly overlap. Lunella dorsal-dorsolateral. Weak sulcalis usually present.

(in mm). H_s: 14.4–23.1, W_s: 4.5–6.0 (topotypes HNHM 83482).

“Am Saumpfade zwischen Antivari und Mikulić” = Montenegro, along the path between Stari Bar and Mikulić [costata (Boettger, 1907)].

Type locality, ex Boettger, ex Wohlberedt, 1906, lectotype (SMF 176320), paralectotype (SMF 176321) [costata (Boettger, 1907)].

Montenegro, Rumija Mts, 3 km above Stari Bar, on the footpath to Veliki Mikulići, 340 m, 42.0973°N, 19.1458°E, leg. LD, ZF, JK, DM, 14.x.2008 (HNHM 99612); same locality, leg. ZE, ZF, 19.iv.2000 (HNHM 83482); same locality, leg. MD, EH, KJ, HS, 7.vii.2015 (NHMW 110430/MN/0155).

The new replacement name refers to the historical name of Stari Bar (Antivari).

Western side of the Rumija Mts in Montenegro. This taxon was found only at the type locality, parapatrically with M. c. umbilicata. (There is a very narrow transitional contact zone between the two taxa). M. c. umbilicata is wider distributed in the area, and can be found along a large section of the footpath from Stari Bar to Veliki Mikulići (Fig. 13).

Although the description of Clausila umbilicata costata was formally published in Wohlberedt (1907), the authorship is unequivocally attributed to Boettger. This name is a primary junior homonym (ICZN Art. 53.3) of Clausilia costata Pfeiffer, 1828 (currently Cochlodina costata), therefore, it must be rejected and replaced by a new substitute name (ICZN Art. 60.3, 72.7).

Shell medium to large, tumid. All whorls smooth. Neck finely costate. Basal and peripheral crests weak. Lamellae superior and spiralis mostly overlap. Lunella dorsal-dorsolateral. Subclaustralis fused to lunella as its almost straight continuation. Plica principalis often connected to the superior. Weak sulcalis usually present.

(in mm). H_s: 18.5–25.3 (lectotype 19.4), W_s: 4.8–6.6 (lectotype 5.4) (lectotype SMF 176318, paralectotypes SMF 176319, topotypes HNHM 99604, Petrovac NHMW 110430/MN/0073, Haj Nehaj NHMW 110430/MN/0075).

“Antivari” = Montenegro, Stari Bar.

Type locality, ex Boettger, ex Benoit, lectotype (SMF 176318), paralectotype (SMF 176319/5).

Montenegro, Stari Bar, fortress, 130 m, 42.0939°N, 19.1342°E, leg. LD, ZF, JK, DM, 14.x.2008 (HNHM 99604); Rumija Mts, 2 km above Stari Bar, along the footpath to Veliki Mikulići, 320 m, 42.0978°N, 19.1447°E, leg. LD, ZF, JK, DM, 14.x.2008 (HNHM 99606); 1 km above Stari Bar, both sides of the gorge, 220 m, 42.0996°N, 19.1426°E, leg. LD, ZF, JK, DM, 14.x.2008 (HNHM 99605, HNHM 99607); Mt. Menka near Zaljevo, leg. Papp, 6.iv.1972 (HNHM 31109); Rijeka Reževići, along the Budva to Petrovac road, 100 m, 42.2270°N, 18.9117°E, leg. TD, ZE, ZF, 26.v.2015 (HNHM 99608, NHMW 110430/MN/0072); S of Novoselje, 3.5 km from Petrovac toward Virpazar, 400 m, 42.2178°N, 18.9545°E, leg. TD, ZE, ZF, 26.v.2015 (HNHM 99609, NHMW 110430/MN/0073); Čanj, valley of the Suvi Potok, road to the beach, 40 m, 42.1649°N, 19.0069°E, leg. TD, ZE, ZF, 26.v.2015 (HNHM 99610, NHMW 110430/MN/0074); Haj Nehaj (near Sutomore), quarry, 140 m, 42.1530°N, 19.0237°E, leg. TD, ZE, ZF, 26.v.2015 (HNHM 99611, NHMW 110430/MN/0075).

This taxon is distributed sporadically along the southern Montenegrin coast between Budva and Zaljevo (Fig. 13).

Earlier M. c. umbilicata used to be classified as a distinct species, but its geographical vicinity and very similar shell characters indicate close relationship with M. cattaroensis.

Wohlberedt (1909) mentions a more southern occurrence, in the Mužura Mts near Ulcinj. This record, however, corresponds to M. subcristata (leg. Winneguth, 1904, NHMW 41258/2).

Shell medium, light violet-brown with whitish tint. Whorls flattened, smooth, only weakly inflexed neck costate. Basal and peripheral crests weak. Peristome broadly attached, ovoid, with somewhat swollen margin. Lamellae superior and spiralis do not overlap. In front view lamella inferior moderately emerged, medium-bent subcolumellaris hidden. Lunella dorsolateral, fused to the short basalis. Subclaustralis also short, sulcalis residual. Anterior plica superior mostly absent, if present short and separate from the lunella complex. Clausilium plate almost entirely visible through the aperture.

(in mm). H_s: 16.1–18.0 (holotype 18.0), W_s: 4.2–4.6 (holotype 4.5).

Albania, Tiranë District, N slope of Mt. Dajti, 1600 m.

Type locality, ex Edlauer, paratypes (NHMW 75000/-E 16652/4); Mal i Dajtit, N-slope, 100 m beneath the summit, ex Edlauer, paratypes (NHMW 75000/-E 32343/4). The holotype (NHMW 76600) could not be found in the NHMW collection.

This taxon is known from the summit region of the Mt. Dajti (1613 m), east of Tiranë (Fig. 12).

The type material is from the collection of Edlauer and, according to the label information, it has been determined as weigneri by Käufel. Therefore, it could have been collected in the 1930s, possibly by Fuchs or Bischoff. Although the type locality seems to be limited to the summit region of the Mt. Dajti, we were unable to find this species there.

Shell small to medium, light corneous. Neck variably inflexed, basal and peripheral crests weak. Peristome attached, its margin somewhat swollen. Lamellae superior and spiralis separate to overlapping. In front view lamella inferior visible, medium-bent subcolumellaris variably exposed. Lunella dorsal to dorsolateral. Basalis and subclaustralis weak to absent. If present, basalis not connected to the lunella. Anterior plica superior, if present, close and parallel to plica principalis, often connected to the lunella complex. Clausilium plate partly or entirely visible through the aperture.

Shell medium, occasionally with whitish tint. Lower whorls smooth, upper ones very densely striate-costate. Neck weakly inflexed, densely striate-costate. Peristome rounded to ovoid. Lamellae superior and spiralis mostly overlap. In front view lamella inferior more or less well emerged, subcolumellaris mostly visible. Lunella dorsolateral, separate from the short basalis. Subclaustralis and sulcalis absent. Anterior plica superior mostly connected to the lunella complex.

(in mm). H_s: 16.6–21.1, W_s: 4.2–5.5 (Nordsieck 1974).

“Tomoros, 1500 m” = Macedonia, Galičica Mts, Tomoros Summit [dofleini]; Macedonia, Galičica Mts, W of Peštani, ca. 1800 m [kaiseri].

“Tomoros, 1500 m”, leg. Doflein, 24.vii.1918, original series [dofleini], syntype status doubtful (ZSM 20150453/2); Peštani, E of the Tomoros Summit, 1800 m, leg. Kaiser, viii.1958, holotype [kaiseri] (ZMH 2926, ex SMF 163942), paratypes [kaiseri] (SMF 163943/4, SMF 201563, ZHM 2278/8, NMBE 9827/2, NHMW-K 44014/2).

Macedonia, Ohrid District, Galičica Mts, ca. 4 km N of the junction at the Sveti Naum to Carina road, 1590 m, 40.9934°N, 20.8569°E, leg. ZF, EH, KJ, HS, 16.x.2014 (NHMW 110430/MN/0038); ca. 600 m E of the Bugarska Chuka, 1650 m, 41.0026°N, 20.8534°E, leg. ZF, EH, KJ, HS, 16.x.2014 (NHMW 110430/MN/0039); ca. 500 m E of the Bugarska Chuka, 1680 m, 41.0019°N, 20.8523°E, leg. ZF, EH, KJ, HS, 16.x.2014 (NHMW 110430/MN/0040); ca. 250 m E of the Bugarska Chuka, 1750 m, 41.0033°N, 20.8488°E, leg. ZF, EH, KJ, HS, 16.x.2014 (NHMW 110430/MN/0041); Bugarska Chuka, E side, 1800 m, 41.0037°N, 20.8470°E, leg. ZF, EH, KJ, HS, 16.x.2014 (NHMW 110430/MN/0042); ca. 300 m W of the Bugarska Chuka, 1740 m, 41.0046°N, 20.8432°E, leg. ZF, EH, KJ, HS, 16.x.2014 (NHMW 110430/MN/0043); ca. 200 m N of the Bugarska Chuka, 1800 m, 41.0051°N, 20.8462°E, leg. ZF, EH, KJ, HS, 16.x.2014 (NHMW 110430/MN/0044); N of the Magaro Summit, 1820 m, 40.9444°N, 20.8270°E, leg. ZE, ZF, JG, 29.vi.2015 (HNHM 99567, NHMW 110430/MN/0046); N of the Magaro Summit, 1870 m, 40.9439°N, 20.8283°E, leg. ZF, 29.vi.2015 (HNHM 99568, NHMW 110430/MN/0047); N of the Magaro Summit, 1900 m, 40.9429°N, 20.8252°E, leg. ZF, 29.vi.2015 (HNHM 99569, NHMW 110430/MN/0048); SE slope of the Galičica Mts, rocky forest along the Carina to Sveti Naum road, 1000–1200 m, leg. LP, PS, AS, 14.vii.1972 (HNHM 36717, SMF 227682); Elšani, above the Elešec Camp, 760 m, 41.0345°N, 20.8105°E, leg. ZF, KJ, HS, 17.x.2014 (NHMW 110430/MN/0045); same locality, leg. LP, PS, AS, 15.vii.1972 (HNHM 36737, HNHM 54405, HNHM 67012, SMF 22768); same locality, leg. ZE, ZF, AH, 7.iv.2004 (HNHM 94433).

This taxon is frequent in the alpine region of the Galičica Mountain. We have found it on the eastern slope of the Bugarska Summit above 1600 m, and on the northern slope of the Magaro Summit above 1800 m. Although sporadically there are occurrences at lower altitudes (e.g. in Elšani at 750 m), on the western slopes of the Galičica, below 1600 m, it is replaced by M. d. fagorum Nordsieck, 1974 (Fig. 14).

Figure 14. 

Distribution of Montenegrina dofleini. M. dofleini dofleini (square); M. dofleini fagorum (inverted empty triangle); M. dofleini kastoriae (star); M. dofleini pinteri (triagle); M. dofleini prespaensis (circle); M. dofleini sinosi (empty circle with dot); M. dofleini wagneri (diamond).

According to Wagner (see: Hesse 1928), the material he studied consisted of six shells, only one of them in good condition, which had been dead-collected from the Tomoros, 1500 m, on July 24, 1918. In the ZSM collection there are only two shells, one of these relatively well preserved (Fig. 11H). These are apparently original specimens collected by Doflein. Although no material could be found in the MIZPAS collection (Dominika Mierzwa, personal communication), there is no evidence that the ZSM specimens are part of the material that had been studied by Wagner. Therefore, the syntype status of this sample is doubtful.

Although not synonymizing formally, Nordsieck (2009) has already questioned the subspecific status of kaiseri. Two arguments seem to support Nordsieck’s opinion that kaiseri could be regarded as junior synonym of dofleini. First, according to the label information, the type series of kaiseri were collected in the vicinity of the Tomoros Summit, which is the type locality of dofleini. Second, the paratypes of kaiseri show high variation in the traits defined by Brandt as distinguishing features.

In the original description the type locality of kaiseri was erroneously defined as ‘W of Peštani’ (Brandt 1961), because the Galičica range is east of this village.

The type series of kaiseri was collected by Kaiser, the curator of the ZMH Mollusc Collection at that time, and the entire lot was originally housed in the ZMH. As indicated in the description, Brandt (1961) intended depositing the holotype in the SMF. As this would have been done without authorization, the type was returned to the ZMH (Zilch 1981, Bernhard Hausdorf, personal communication).

Shell small, tumid, with whitish tint. All whorls smooth and glossy. Neck inflexed, behind the aperture finely striate. Peristome rounded to ovoid. Lamellae superior and spiralis mostly do not overlap. In front view lamella inferior moderately emerged, subcolumellaris not or only slightly barely visible. Lunella dorsal-dorsolateral, basalis and subclaustralis absent. Sulcalis mostly absent or residual. Anterior plica superior absent or weak, not connected to the lunella complex.

(in mm). H_s: 11.8–14.4 (holotype 13.5), W_s: 3.4–4.0 (holotype 3.7).

Macedonia, Galičica Mts, above Trpejca, 10 km toward Carina along the road crossing the Galičica Mts, 1400 m.

Type locality, leg. Nordsieck, 26.viii.1971, holotype (SMF 227680), paratypes (SMF 227681, NHMW-K 65049/10, SMNS-N 5483), same locality, leg. Fauer, 17.viii.1971, paratypes (SMNS-N 5571, NMBE 535049/3), same locality, leg. LP, PS, AS, 14.vii.1972, paratypes (NMBE 535047/8, HNHM 11915/15, HNHM 67026/2, HNHM 36721/8).

Macedonia, near the type locality, Korita Spring, 40.9674°N, 20.8149°E, leg. ZF, EH, KJ, HS, 17.x.2014 (NHMW 110430/MN/0053); same locality, leg. ZE, ZF, AH, 6.iv.2004 (HNHM 94244); Galičica Mts, 12 km from the Sveti Naum to Ohrid road toward Carina, 1540 m, 40.9627°N, 20.8171°E, leg. A. Fehér, T. Fehér, ZF, LT (NHMW 110430/MN/0052); same locality, leg. ZE, ZF, AH, 6.iv.2004 (HNHM 94450); 14 km from the Sveti Naum to Ohrid road toward Carina, Baba observation point, 1600 m, 40.9541°N, 20.8126°E, leg. ZE, ZF, JG, 29.vi.2015 (NHMW 110430/MN/0054); same locality, leg. ZE, ZF, AH, 6.iv.2004 (HNHM 94451).

This taxon inhabits the western slopes of the Galičica Mountain. It was found along the road crossing the mountain from ca. 1300 m to the mountain pass at 1600 m. Nordsieck (1974) mentions this taxon from the Mt. Thatë (southern extension of the Galičica), however those records correspond to another taxon, M. d. wagneri Szekeres, 2006. Our data suggest that the ranges of fagorum and wagneri are disjunct, as we found typical dofleini populations between them in the Magaro Summit area (NHMW 110430/MN/0046–48).

Shell medium, elongate, whitish-corneous. Lower whorls smooth to indistinctly ribbed, upper ones very densely striate to costate. Neck weakly inflexed, behind the aperture striate to costate. Peristome ovoid to angular. Lamellae superior and spiralis overlap. In front view lamella inferior moderately emerged, subcolumellaris not or barely visible. Lunella dorsal to dorsolateral, weaker toward the basis. Basalis rarely recognizable, never connected to the lunella. Subclaustralis and sulcalis absent. Anterior plica superior mostly does not reach the lunella complex.

(in mm). H_s: 14.4–20.4 (holotype 19.0), W_s: 3.8–4.6 (holotype 4.5).

Greece, Western Macedonia, Kastoria, S side of the peninsula.

Type locality, leg. H. Nordsieck, 24.viii.1971, holotype (SMF 221042), paratypes (SMF 221043, NHMW-K 65048/10, SMNS-N 5479).

Greece, Western Macedonia, Kastoria, N side of the peninsula, leg. H. Nordsieck, 24.viii.1971 (SMNS-N 5477, NHMW 77938/5); same locality, behind Yacht Club, 640 m, 40.5233°N, 21.2747°E, leg. ZE, ZF, JG, 28.vi.2013 (HNHM 99570); same locality, leg. ZF, EH, KJ, HS, 17.x.2014 (NHMW 110430/MN/0050); Florina District, Krystallopigi, above the cemetery, 1120 m, 40.6357°N, 21.0903°E, leg. ZE, ZF, JG, 29.vi.2013 (HNHM 99571); same locality, leg. ZF, EH, KJ, HS, 17.x.2014 (NHMW 110430/MN/0049).

Western part of the Verno Mts in Western Macedonia Province, Greece. Originally this taxon was known only from Kastoria, but recently it has also been found at Krystallopigi, ca. 20 km northwest of the type locality (Fig. 14).

Shell medium, elongate, whitish-corneous. Lower and upper whorls with dense, sharp ribs. Neck not or barely inflexed, ribs behind the aperture denser and finer. Basal and peripheral crests very weak. Peristome rounded to somewhat angular. Lamellae superior and spiralis long overlap. In front view lamella inferior moderately emerged, subcolumellaris mostly not visible. Lunella dorsal-dorsolateral, separate from the short basalis. Subclaustralis and sulcalis absent. Anterior plica superior mostly connected to the lunella complex.

(in mm). H_s: 14.8–18.9 (holotype 16.9), W_s: 4.1–5.0 (holotype 4.4).

Macedonia, Trpejca, north of boat harbor.

Type locality, leg. LP, PS, AS, 16.vii.1972, holotype (SMF 227684), paratypes (SMF 227685, SMF 265700/4, SMNS-N 6146, NMBE 535039/118, ZMH 91605/3, HNHM 11960/272, HNHM 37763/207, HNHM 37762/3, HNHM 36708/3, HNHM 52687/10, HNHM 67011/6).

Macedonia, Ohrid District, N of Trpejca, shore of the Lake Ohrid, 700 m, 40.9656°N, 20.7858°E, leg. ZF, 9.viii.2014 (NHMW 110430/MN/0051); same locality, leg. ZE, ZF, AH, 6.iv.2004 (HNHM 94438); Sv. Zaum, shore of Lake Ohrid, 40.949°N, 20.774°E (DED).

This taxon is known only from the vicinity of Tpejca at the shore of the Lake Ohrid (Fig. 14). At the type locality it occurs in the immediate vicinity of M. stankovici. The two taxa can often be found on the same cliffs, with M. stankovici preferring the lowest regions directly above the water surface, and M. d. pinteri inhabiting higher parts.

See considerations on niche segregation at M. stankovici.

Shell medium, elongate. Lower whorls smooth to indistinctly wrinkled, upper ones almost smooth to bluntly costate. Neck very weakly inflexed, behind the aperture striate to finely costate. Peristome attached, ovoid. Lamella superior weak, with spiralis does not overlap. In front view lamella inferior moderately emerged, subcolumellaris mostly not visible. Lunella dorsal-dorsolateral, weaker toward the basis. Basalis and subclaustralis mostly absent, residual sulcalis occasionally recognizable. Anterior plica superior absent or, if present, occasionally connected to the lunella complex.

(in mm). H_s: 13.9–19.6 (holotype 16.7), W_s: 3.6–4.5 (holotype 3.9).

Greece, Western Macedonia, Psarades, opposite to the village.

Type locality, leg. HS, 1986, holotype (NHMW 84019), paratypes, (NHMW 84020/46); Psarades, peninsula NW of the village, leg. HS, 1985, paratypes (NHMW 84021/6, SMNS-N 9519).

Greece, Psarades, N of the village, 860 m, 40.8315°N, 21.0288°E, leg. PS, 10.v.1995 (HNHM 12040); Psarades, E side of the bay, 870 m, 40.8316°N, 21.0287°E, leg. ZE, ZF, JG, 28.vi.2013 (HNHM 99573); same locality, leg. ZF, EH, KJ, HS, 18.x.2014 (NHMW 110430/MN/0056); SW of Psarades, Panagia Eleousa cave temple, 850 m, 40.8087°N, 20.9979°E, leg. ZF, EH, KJ, HS, 18.x.2014 (NHMW 110430/MN/0057); Albania, Korçë District, Kallamas, 850 m, 40.8924°N, 20.9394°E, leg. ZF, EH, KJ, HS, 17.x.2014 (NHMW 110430/MN/0055); same locality, leg. ZE, ZF, JG, 26.vi.2015 (HNHM 99575); Sveta Marena cave temple ca. 4 km E of Kallamas, 850 m, 40.8872°N, 20.9729°E, leg. ZE, ZF, JG, 29.6.2015 (NHMW 110430/MN/0134); 2 km SE of Kallamas, 850 m, 40.8809°N, 20.9590°E, leg. ZE, ZF, JG, 29.6.2015 (NHMW 110430/MN/0135).

Lake Prespa area, with sporadical occurrences are along the western and southern shores. Occasinally sympatric, but not syntopic, with M. hiltrudae sattmanni Nordsieck, 1988 and M. h. desaretica ssp. n. (Fig. 14).

From the confluence of the Great and Small Prespa Lakes Nordsieck (1988) mentions a further lot of paratypes (NHMW 84022/3) which, in fact, belongs to M. d. sinosi (see there).

Shell medium, elongate. Lower and upper whorls strongly wrinkled or ribbed. Neck very weakly inflexed, behind the aperture densely striate-costate. Peristome ovoid. Lamella superior weak, does not overlap spiralis. In front view lamella inferior moderately emerged, subcolumellaris mostly hidden. Lunella dorsal-dorsolateral, weaker toward the basis. Basalis and subclaustralis mostly absent, residual sulcalis occasionally recognizable. Anterior plica superior absent or, if present, occasionally connected to the lunella complex.

(in mm). H_s: 13.9-16.3 (holotype 15.0), W_s: 3.4-3.7 (holotype 3.5 mm).

Greece, Western Macedonia, southern shore of the Prespa Lake, Agios Achillios, near the junction to Psarades, 850 m, 40.8105°N, 21.0702°E.

Type locality, leg. Páll-Gergely, 25.v.2009, holotype (HNHM 97145), paratypes (HNHM 97146/3, SMF 333885/3, NMBE 28607/2, NHMW 107878/2); at the confluence of the Great and Small Prespa Lakes, leg. HS, 21.v.1986, paratypes [prespaensis] (NHMW 84022/3).

Type locality, leg. ZE, ZF, JG, 28.vi.2013 (HNHM 99580); same locality, leg. ZF, EH, KJ, HS, 18.x.2014 (NHMW 110430/MN/0058).

The subspecies is known only from its type locality, which is only at 4 km distance from the nearest occurrence of M. d. prespaensis. Potentially suitable intermediate localities have not been searched, therefore it is not clear whether there are transitional populations between the two taxa (Fig. 14).

Morphologically M. d. sinosi might be an extremely ribbed morph of the closely related M. d. prespaensis. Its distinct subspecific status remains questionable until the finer distribution pattern and population genetic relationship of the two taxa becomes better known.

Shell small, with whitish tint. All whorls smooth and glossy. Neck very weakly inflexed, behind the aperture finely striate. Basal and peripheral crests very weak. Peristome ovoid. Lamellae superior and spiralis do not overlap. In front view lamella inferior barely emerged. Deep subcolumellaris visible only in obliquely through the aperture. Lunella dorsolateral. Basalis mostly, subclaustralis always absent, sulcalis strong. Anterior plica superior usually absent or, if present, residual and connected to the lunella complex.

(in mm). W_s: 13.2–14.9 (holotype 14.9), W_s: 3.4–4.3 (holotype 4.2).

Albania, Mt. Thatë.

“Mal i That”, leg. Fuchs, holotype (NHMW 103375), paratypes (NHMW 47961/2).

Albania, Korçë District, Mt. Thatë, E of Podgorije, ca. 1.3 km E of the Bregu i Stanit Summit, 1830 m, 40.8255°N, 20.8528°E, leg. ZB, CN, DP, 20.v.2007 (HNHM 99576); ca. 4.3 km NW of Liqenas, ca. 700 m south of a summit, 1910 m, 40.8183°N, 20.8728°E, leg. ZB, DP, 22.v.2007 (HNHM 99577); ca. 4.3 km NW of Liqenas, 400 m ESE of a summit, 1780 m, 40.8220°N, 20.8789°E, leg. ZB, CN, DP, 22.v.2007 (HNHM 99578); along the path from Goricë to the ridge, 40.8485°N, 20.8624°E, leg. ZB, CN, Schmotzer, 11.viii.2008 (HNHM 99579); near the ‘Tower of the war’, alpine rocky grassland, 1750 m, 40.9122°N, 20.8498°E, leg. ID, 25.v.2013 (DED); same locality, 1810–1830 m, 40.9136°N, 20.8419°E, leg. ID, 24.ix.2013 (DED); Plaja e Pusit Summit, 2200 m, 40.8799°N, 20.8419°E, leg. ID, 25.vi.2013 (DED).

Alpine region (above 1700 m) of the Mt. Thatë in southern Albania. Dedov found this taxon syntopically with M. hiltrudae fusca Fehér & Szekeres, 2006 in the ‘Tower of the war’ area, near the Albanian–Macedonian border (Fig. 14).

Due to misinterpretation of the label information, the original description erroneously indicated the type locality as “Maja e Madhe (1796 m), WNW of Peshkopi”.

In Nordsieck’s view (Nordsieck 2009) M. d. wagneri is synonymous with M. d. fagorum. However, in our opinion their distinct subspecies status is well supported by stable differences in shell morphology.

Shell small, tumid. Lower whorls very pale, almost colorless. All whorls costate, ribs stronger, sharper and wider spaced toward the apex. Neck deep inflexed, densely costate. Basal and peripheral crests strong. Peristome attached, rounded to somewhat angular, with wide, simple margin. Lamellae superior and spiralis mostly do not overlap. In front view lamella inferior barely emerged, medium-bent subcolumellaris hidden, often not at all visible through the aperture. Plica principalis fused to the superior. Lunella dorsolateral, connected to the basalis. Subclaustralis shorter than the basalis, sulcalis residual or absent. Anterior plica superior separate from the lunella complex. Clausilium plate only barely visible through the aperture.

(in mm). H_s: 11.1–15.4 (holotype 12.6), W_s: 3.8–4.7 (holotype 4.5).

Albania, Dibrë District, Lunarë, at the bridge of the Lumi i Murrës (22 km W of Fushë-Muhurr along the Peshkopi to Burrel road) 730 m, 41.6256°N, 20.2498°E.

Type locality, leg. ZE, ZF, JK, DM, 26.vi.2003, holotype (HNHM 94829), paratypes (HNHM 94830/18, NHMW 103271, RMNH 100312, SMF 328075, NMBE 534905).

Type locality, leg. ZF, DM, ZU, 18.v.2010 (HNHM 99636).

Southern part of the Lura-Dejë mountain group in northern Albania. Known only from the type locality (Fig. 16).

Nordsieck (2009) placed M. drimmeri into the same species-group with M. fuchsi.

Figure 15. 

A Montenegrina haringae sp. n., holotype, NHMW 111219 B M. fuchsi fuchsi, Brandt, 1961, holotype, SMF 163939 C M. fuchsi klemmi Brandt, 1962, holotype, SMF 167014 D M. fuchsi muranyii Fehér & Szekeres, 2006, holotype, HNHM 94846 E M. fuchi pallida Fauer, 1993, holotype, SMF 309235 F Montenegrina grammica grammica, Nordsieck, 1988, holotype, NHMW 84026 G M. grammica erosszoltani ssp. n., holotype, HNHM 99512 H M. grammica improvisa ssp. n., holotype, HNHM 99518 I M. helvola helvola (Küster, 1860), Mt. Krujë, HNHM 86052 J M. helvola carinata Erőss & Szekeres, 1996, holotype, HNHM 70836 K M. helvola magna Fehér & Szekeres, 2006, holotype, HNHM 94848 L M. helvola ornata Erőss & Szekeres, 1996, holotype, HNHM 70838 M M. helvola pageti Brandt, 1962, holotype, SMF 167016 N M. drimmeri Fehér & Szekeres, 2006, holotype, HNHM 94829. Scale bar: 5 mm.

Shell very small to small, light corneous, whorls smooth to indistinctly wrinkled-costate. Neck deep inflexed, behind the aperture costate. Basal crest strong, peripheral crest well visible to very strong. Peristome with simple margin, narrowly attached to detached and projected. In front view lamella inferior moderately emerged, broadly-bent subcolumellaris not or only barely visible. Lunella dorsal to dorsolateral, subclaustralis short. Anterior plica superior not connected to the lunella complex. Clausilium plate partly visible through the aperture.

Shell surface indistinctly wrinkled, neck densely costate. Peripheral crest very strong. Peristome detached, strongly projected, rounded to pear-shaped. Lamella subcolumellaris visible through the aperture. Plica principalis often fused to the superior. Lunella dorsal, sulcalis well developed.

(in mm). H_s: 13.6–13.7, W: 2.9–3.0 (width of the penultimate whorl).

“An der Straße von Tepelene nach Korzyra” = Albania, valley of the Vjosë River between Kelcyrë and Tepelenë.

Type locality, ex Käufel, ex Zilch, holotype (SMF 163939); Type locality, leg. Fuchs, 1936, paratypes (NHMW-K 4491/3, NMBE 9825/2, ZMH 113581/3).

Albania, right bank of the Vjosë at the bridge along the Tepelenë to Kelcyrë road, leg. Fuchs, 1936 (NHMW-K 47960, NHMW-K 32317, NHMW-K 48293, NHMW 21443, ZSM 20150454); “Shendeli bei Tepelenë, 1000 m”, leg. Fuchs (NHMW-K 3238, NHMW-K 48292); Tepelenë District, 7 km W of Këlcyre, near the bridge at the Peshtan junction, 170 m, 40.2961°N, 20.1076°E, leg. ZE, ZF, 6.vii.1996 (HNHM 86048); same locality, leg. ZE, ZF, KK, 13.iv.2001 (HNHM 85949), same locality, leg. PJ, TK, DM, GP, 14.x.2013 (HNHM 99563); same locality, leg. DA, ZE, ZF, JG, 29.vi.2014 (HNHM 99333, NHMW 110430/MN/0036).

Southern part of the Mt. Shëndel in southern Albania.

Most of the museum lots of this taxon are from the valley of the Vjosë River, west of Kelcyrë, at the foot of the Mt. Shëndel. All of these are probably from the same locality, near the bridge to Peshtan. However, the record by Fuchs (“Shendeli bei Tepelene, 1000 m”) indicates a presumably wider distribution in the Shëndel, north of the type locality (Fig. 16).

Figure 16. 

Distribution of Montenegrina drimmeri and M. fuchsi. M. drimmeri (star); M. fuchsi fuchsi (square); M. fuchsi klemmi (triangle); M. fuchsi muranyii (empty circle with dot); M. fuchsi pallida (circle).

Franz Käufel apparently intended to describe this as a new species, and started to distribute material as types. The HNMW 21443 lot contains a handwritten label from Käufel with the name fuchsi, and the indication “Paratypen”. But Käufel’s manuscript has never been published and this taxon was formally described only by Brandt in 1961. Therefore, only the lots designated in Brandt’s (1961) description are considered type material.

Shell surface smooth to indistinctly wrinkled, neck irregularly costate. Peripheral crest strong. Peristome narrowly attached or detached, rounded. Lamella subcolumellaris barely visible through the aperture. Lunella dorsal-dorsolateral, sulcalis well developed.

(in mm). H_s: 10.4–14.0, W_s: 2.6–2.9.

“Suhe, Tal d. Lum i Suhes ö. Argyrokastro” = Albania, Gjirokastër District, Suhë, valley of Lumi i Suhës.

Type locality, leg. Fuchs, 1931, holotype (SMF 167014), paratypes (SMF 167015/2, NHMW-E 32491/23, NMBE 9826/4, ZMH 113579/2).

Type locality, ex Klemm (SMF 201621/2) [although mentioned as paratypes by Zilch (1981), the original description does not mention this lot]; same locality, without date (NHMW-K 48294/7) [although indicated on the label as type, the original description does not mention it]; Gjirokastër District, between Suhë and Poliçan, leg. Fuchs, 1936. (NHMW-E 32441, NHMW-K 48295); 3 km NE of Suhë, along the Libohovë to Sheper road, 430 m, 40.0882°N, 20.2716°E, leg. ZF, JK, DM, 12.x.2004 (HNHM 95049); same locality, leg. DA, ZE, ZF, JG, 27.vi.2014 (HNHM 99272, NHMW 110430/MN/0037); 5 km NE of Suhë, 420 m, 40.0915°N, 20.2922°E, leg. ZF, JK, DM, 12.x.2004 (HNHM 95050).

Lunxhëri Mts in southern Albania. Known only from the gorge of the Suhë Creek crossing the mountain east to west between Suhë and Poliçan (Fig. 16).

Shell very small, light corneous. All whorls smooth or very finely, indistinctly wrinkled. Neck weakly inflexed, densely wrinkled-costate. Basal and peripheral crests well recognizable. Peristome rounded to ovoid, with simple margin. Lamellae superior and spiralis do not overlap. In front view lamella inferior moderately emerged, subcolumellaris not visible. Lunella dorsolateral. Basalis absent or weak and separate from the lunella. Subclaustralis absent, sulcalis weak to residual. Anterior plica superior mostly present, not connected to the lunella complex. Clausilium plate not or only barely visible through the aperture.

(in mm). H_s: 8.9–11.9 (holotype 10.9), W_s: 2.8–3.4 (holotype 3.4).

Albania, Berat District, Tomorr Mts, Kalaja e Tomorrit, 1100 m, 40.7025°N, 20.1093°E.

Type locality, leg. Harmos, DM, 26.v.2004, holotype (HNHM 94846), paratypes (NHMUK 20050219, HNC 63183, HNHM 94847/39, NHMW 103277, RMNH 100307, SMF 328081).

Tomorr Mts in southern Albania. This taxon is known only from the type locality, which is a western extension of the Tomorr (Fig. 16).

Shell surface indistinctly wrinkled-costate, stronger at the upper whorls. Neck with sharp ribs and well visible recognizable peripheral crest. Peristome detached, projected, ovoid. Lamella subcolumellaris visible through the aperture. Lunella dorsal, sulcalis weak to residual.

(in mm). H_s: 10.7–15.5 (holotype 12.5), W_s: 2.8–4.1 (holotype 3.4).

Greece, Epirus, Pogonisko junction on the Konitsa to Molivdokepastos road, 500 m.

Type locality, leg. Fauer, PS, 26.ix.1989, holotype (SMF 309235), paratypes (SMF 309236/2, NMBE 534897/115, SMNS-N 9792, NHMW 103730/1); same locality, leg. PS, 16.vii.1990, paratypes (NMBE 534896/95).

Type locality, 40.0468°N, 20.5648°E, leg. ZE, ZF, JG, 26.vi.2013 (HNHM 99566).

Southern part of the Dousko (Nemërçkë) Mountain, in Epirus, northwestern Greece. Known only from the type locality (Fig. 16).

Shell very small to small, neck inflexed, with strong peripheral crest. Peristome attached, with swollen margin. Lamellae superior and spiralis do not or only occasionally overlap. In front view lamella inferior barely to moderately emerged, medium-bent subcolumellaris mostly not visible. Lunella broad, dorsolateral to lateral. Basalis absent or short, if present fused to or separate from lunella. Subclaustralis and sulcalis residual or absent. Anterior plica superior absent or weak and separate from the lunella complex. Differs from M. janinensis by the smaller, more conical and stronger sculptured shell, as well as the deeper inflexed neck and broadly-bent lamella subcolumellaris.

Shell light corneous. Lower whorls smooth to indistinctly costate-weinkled, upper ones with strong ribs. Neck smooth to weakly wrinkled, with weak basal crest. Peristome rounded. Lamellae superior and spiralis do not overlap. In front view lamella inferior barely emerged, subcolumellaris mostly not visible. Lunella dorsolateral. Basalis absent or residual and fused to the lunella. Anterior plica superior mostly absent, rarely weak and separate from the lunella complex.

(in mm). H_s: 9.8–10.3 (holotype 9.8), W_s: 2.9–3.4 (holotype 3.1).

Greece, Gramos Mts, mountain ridge NNW of the Epano Arena, ca. 2000 m.

Type locality, leg. HS, 3.vii.1986, holotype (NHMW 84026), paratypes (NHMW 84027/8).

Greece, Gramos Mts, W of Epano Arena summit, 2000 m, 40.3090°N, 20.8981°E, leg. ZE, ZF, JG, 27.vi.2013 (HNHM 99628).

Gramos Mts in northwestern Greece. Known only form the northern side of the Epano Arena. In that area it lives syntopically with the more frequent and wider distributed M. skipetarica pindica Nordsieck, 1988 (Fig. 17).

Figure 17. 

Distribution of Montenegrina grammica and M. haringae sp. n. M. grammica erosszoltani ssp. n. (square); M. grammica grammica (diamond); M. grammica improvisa ssp. n. (triangle); M. haringae sp. n. (circle).

Very small subspecies with dorsolateral lunella, missing basalis and non-connected anterior plica superior.

The very small, somewhat conical, greyish-corneous shell consists of 7½ to 9 whorls. The entire surface is strongly costate, with denser ribs behind the aperture and toward the apex. The deep neck inflection extends to the lateral side. The basal and peripheral crests are well developed. The ovoid to circular aperture has broadly attached, simple margin. The lamellae superior starts at the same depth as that of the end of the spiralis, occasionally with a slight overlap. The lamella inferior and medium-bent subcolumellaris are only rarely and slightly visible in front view. The broad lunella is dorsolateral. The basalis and subclaustralis are missing, the sulcalis is present. The weak anterior part of the plica superior initiates far from the lunella complex. Only the parietal edge of the clausilium plate is visible through the aperture.

(in mm). Holotype H_s: 8.9, W_s: 3.0, H_a: 2.4, W_a: 2.1; paratypes (HNHM 99513, n = 12): H_s: 8.5–11.4 (mean 9.5, S.D. 0.77), W_s: 2.6–3.3 (mean 2.9, S.D. 0.21), H_a: 2.3–2.9, W_a: 1.9–2.4.

The new subspecies differs from M. g. grammica by its strongly costate whorls and the presence of the anterior plica superior, whereas from M. g. improvisa by its smaller size, costate surface, deeper ending lamella subcolumellaris, and missing basalis.

Albania, Mat District, W of the Shkëmb i Skanderbeut (1 km from the Peshkopi to Burrel road toward Fushë-Lurë), 1160 m, 41.6461°N, 20.1807°E.

Type locality, leg. ZE, ZF, AH, DM, 13.iv.2006, holotype (HNHM 99512), paratypes (HNHM 99513/52, ER/52, HU/52, SZ/5); same locality, leg. ZE, ZF, 11.x.2005, paratypes (HNHM 99514/138+5aj, MMM-B01327/138, ER/138, SZ/10); same locality, leg. ZF, DM, ZU, 18.v.2010, paratypes (HNHM 99515/3a+75, NHMW 111233/10); on the Peshkopi to Burrel road, 1 km W of the Fushë-Lurë junction, 1090 m, 41.6451°N, 20.1823°E, leg. ZE, ZF, AH, DM, 13.iv.2006, paratypes (HNHM 99516/27, ER/27, HU/27, SZ/5); Mali i Dejës, limestone hill of 1700 m height above the headwaters of the Lumi i Varoshit, 1540 m, 41.6746°N, 20.2112°E, leg. ZF, DM, ZU, 18.v.2010, paratypes (HNHM 99517/3+1a, SZ/2).

The new subspecies is named after our friend and colleague Zoltán Erőss, who accompanied the first author on several Balkan field trips, including the one during which this taxon was discovered.

Southern part of the Lura-Dejë mountain group, within the catchment area of the Varosh Stream (Fig. 17).

At one of the localities it was found syntopically with M. skipetarica puskasi ssp. n.

Small subspecies with almost smooth shell, dorsolateral to lateral lunella, short, non-connected basalis, and missing anterior plica superior.

The medium-size, light brownish-corneous shell consists of 9½ to 11 whorls. The surface is smooth at the lower whorls and becomes weakly, indistinctly costate toward the apex. The neck is irregularly and finely wrikled-costate, and behind the aperture densely striate. The basal and peripheral crests are well developed. The peristome is attached, ovoid to somewhat angular, with slightly swollen margin. The lamellae superior and spiralis occasionally overlap. In front view the straight descending end of the lamella inferior is visible, the subcolumellaris is hidden. The lunella is broad, dorsolateral to lateral, separate from the short basalis. The subclaustralis is absent, the sulcalis is residual or also missing. The clausilium plate cannot be viewed through the aperture.

(in mm). Holotype H_s: 14.4, W_s: 3.8, H_a: 3.6, W_a: 2.4; paratypes (HNHM 99019, n = 12): H_s: 12.8–14.4 (mean 13.7, S.D. 0.50), W_s: 3.3–4.2 (mean 3.6, S.D. 0.26), H_a: 3.0–3.6, W_a: 2.0–2.6.

The new taxon can be distinguished from M. g. grammica and M. g. erosszoltani by its larger and smoother shell, deep-ending lamella subcolumellaris, deeper lunella, and the presence of a short basalis.

Albania, Mat District, Gropa Mts, 3.5 km W of Gurri i Bardhë, N slope of the Maja e Bastarit, 1160 m, 41.4361°N, 20.0485°E.

Type locality, leg. ZF, TN, EM, 15.iv.2014, holotype (HNHM 99518), paratypes (HNHM 99019/29+18a+8aj, NHMW 111234/5, SZ/3).

The name improvisa refers to the unexpected, accidental discovery of the new subspecies.

Gropa Mts in Central Albania. Known only from the type locality (Fig. 17).

Small to medium-size species with broadly attached peristome, broadly-bent lamella subcolumellaris, dorsal lunella, and weak, separate basalis.

The small to medium-size, tumid, light corneous shell consists of 8 to 9½ whorls. The entire surface is opaque and smooth, except for the inflexed neck that is weakly costate behind the aperture. The basal and peripheral crests are weak. The aperture is ovoid, somewhat angular, broadly attached, with simple peristome margin. The long lamella superior only barely overlaps with the spiralis. In front view the lamella inferior is moderately emerged, the end of the broadly-bent subcolumellaris is usually visible. The dorsal lunella is separate from the weak, lump-like basalis. The subclaustralis is absent, the sulcalis is residual or also absent. The anterior part of the plica superior is absent or weak, non-parallel to the principalis, separate from the lunella complex. The narrow clausilium plate is partly visible through the aperture, its proximal half is obscured by the deep-emerged lamella inferior.

(in mm). Holotype H_s: 14.8, W_s: 4.3, H_a: 3.6, W_a: 2.9; paratypes (NHMW 111220, n =12): H_s: 12.5–17.4 (mean 14.7, S.D. 1.44), W_s: 3.7–4.8 (mean 4.4, S.D. 0.26), H_a: 3.3–4.1, W_a: 2.9–3.7 mm.

Compared to species of the surrounding coastal regions, the new taxon can be distinguished from both M. cattaroensis and M. subcristata by its smaller size, less deep lunella, missing subclaustralis and weak or absent anterior plica superior. From all other species of the genus differs by the combination of a dorsal lunella and the absence of subclaustralis.

Albania, Shkodër District, Mt. Renc, dry gorge S of Baks-Rjollë, (SW of the Maja e Zezë), 30 m, 41.8577°N, 19.4923°E.

Type locality, leg. TD, ZE, ZF, 27.v.2015, holotype (NHMW 111219), paratypes (NHMW 111220/65, HNHM 99483/61, MMM-B01324/61, SZ/5); above Baks-Rjollë, S slope of the Maja e Zezë, 100 m, 41.8569°N, 19.5044°E, leg. ZB, DP, GP, 4.v.2014, paratypes (HNHM 99061/4, SZ/1).

The new species is named after Elisabeth Haring, head of the Central Research Laboratories at NHMW, who provided support and inspiration to this study.

Mt. Renc in northwestern Albania (Fig. 17).

Shell small to large, light corneous. Whorls smooth to indistinctly costate. Neck deep inflexed, costate. Basal and peripheral crests strong. Peristome detached, projected, rounded to pear-shaped, with simple margin. Lamellae superior and spiralis long overlap. In front view lamella inferior high-ending, hidden to moderately emerged. Broadly-bent subcolumellaris retracted, not visible through the aperture. Lunella dorsolateral to ventrolateral. Basalis absent to long and fused to the lunella as its almost straight continuation. Subclaustralis absent or short, perpendicular to the lunella. Sulcalis absent or residual. Anterior plica mostly present, separate from the lunella complex. Clausilium plate not or only barely visible through the aperture.

Shell small to medium. Lower whorls smooth, upper ones indistinctly wrinkled-costate. Neck wrinkled-costate. Peristome rounded to pear-shaped. In front view lamella inferior very high, barely emerged. Lunella dorsolateral. Basalis absent or residual and fused to the lunella. Subclaustralis weak and short, sulcalis residual. Anterior plica superior short, occasionally absent. Clausilium plate barely visible through the aperture.

(in mm). H_s: 12.5–19 (neotype: 18.5), W_s: 3.5–4.6 (neotype: 4.5) (SMF 94110a, HNHM 86051).

Albania, Krujë, Skanderbeg Fortress (due to the present neotype designation). In the original description the locality was given as “Dalmatien”. According to Wagner (1924), the original material was collected in Slano (southern Dalmatia) from marine flotsam.

Albania, Mali i Krujës, inside the Skanderbeg Castle, ex K. L. Pfeiffer, ex Fuchs, 1941, neotype (SMF 94110a) (Zilch in Wenz 1960: fig. 1527.).

Same as that of the neotype (NHMW-E, NHMW-E 33201, NHMW-E 32547, NHMW-K 50602, NHMW-K 48291 NHMW-K 54388); Krujë, castle, 600 m, 41.5107°N, 19.7947°E, leg. ZE, ZF, KK, 10.iv.2001 (HNHM 86054); Mali i Krujës, S slope at 1200 m, “zum Derwischgrab”, leg. Fuchs (NHMW-E 16727/6); same locality at 1000 m, leg. Fuchs (NHMW-E 16501/3); Mali i Krujës, above Krujë, ca. 1000 m, 41.5222°N, 19.8055°E, leg. ZF, Kónya, 14.ix.1994 (HNHM 86051); same locality, leg. ZE, ZF, KK, 10.iv.2001 (HNHM 86052); Krujë, road to the Qafa e Shtamës, 41.5220°N, 19.7939°E, leg. AR, NR, PR, ix.2013 (NHMW 10.430/MN/0076); Mali i Krujës above Noi, along the Burrel to Krujë road, 700 m, 41.5229°N, 19.8164°E, leg. ZE, ZF, JK, DM, 26.x.2002 (HNHM 91606).

Mt. Krujë in central Albania. Known from the fortress of Krujë and some other sites on the Mt. Krujë, above and east of the town (Fig. 18).

Figure 18. 

Distribution of Montenegrina helvola. M. helvola carinata (square); M. helvola helvola (diamond); M. helvola magna (inverted triangle); M. helvola ornata (triangle); M. helvola pageti (circle).

The first volume of the Systematisches Conchylien-Cabinet (Küster 1844–1862) was published in parts. Plate 19, with a figure of M. helvola but without description, was printed in 1853. The description (at p. 176), however, was published only in 1860 (Welter-Schultes 1999).

Without specific type locality it was not possible to ascertain where Küster’s material originated from. Zilch (1960) depicted a specimen from Krujë, which was subsequently used as reference for the taxon without any nomenclatural relevance. The original type material could not be found in the public collections that currently hold remnants of Küster’s collection. Therefore, we can reasonably assume that it has been lost. In order to clarify and fix the taxonomic status and the type locality of helvola it is necessary to designate a neotype, in accordance with ICZN Art. 75.3. To preserve nomenclatural stability, we think it is justified to designate as neotype the specimen that has been used as reference for the taxon since Zilch (1960).

Wagner (1924) mentioned helvola as a species of Alpidelima. But in the same paper he listed it under Albanodelima, suggesting Alpidelima could have been merely a typographic error.

Shell small to large, slender. Lower whorls smooth to finely, indistinctly costate, upper ones wrinkled-costate. Neck strongly costate. Peripheral crest particularly strong, extends to the entire last whorl. Peristome pear-shaped. In front view lamella inferior hidden or only barely emerged. Lunella ventrolateral, fused to the long basalis. Subclaustralis short, sulcalis absent or residual. Anterior plica superior long. Clausilium plate not visible through the aperture.

(in mm). H_s: 15.4–21.2 (holotype 18.9), W_s: 3.4–4.2 (holotype 3.7).

Albania, Mallakastër District, 4 km SSE of Greshicë, gorge of the Përroi i Povlit (in the description erroneously given as Konakut Valley), 150 m, 40.5356°N, 19.7964°E.

Type locality, leg. ZE, 17.viii.1993, holotype (HNHM 70836), paratypes (HNHM 70837/2, NHMW 100077/2, SMF 312506/2, NMBE 534907/1, NMBE 22595/1).

Type locality, leg. ZF, JK, DM, 11.x.2004 (HNHM 95051); Byllis, 40.5447°N, 19.7389°E, leg. AR, NR, PR, ix.2003 (NHMW 110430/MN/0078); Poçem, bank of the Vjosë, 50 m, 40.4926°N, 19.7261°E, leg. ZB, DP, GP, 11.v.2014 (HNHM 99618).

Mallakastër Hills in southern Albania, in the lower valley of the Vjosë River. Known localities are within 5 km from each other (Fig. 18).

Nordsieck (2009) mentions this taxon from Mollas, which record presumably refers to M. h. ornata Erőss & Szekeres, 1999.

Shell medium to large, ventricose. Lower whorls smooth, upper ones smooth to indistinctly wrinkled-costate. Neck wrinkled-costate. Peristome pear-shaped to somewhat angular. In front view lamella inferior barely emerged. Lunella lateral, fused to the short basalis. Subclaustralis and sulcalis residual. Anterior plica superior long, rarely connected to the lunella complex. Clausilium plate not visible through the aperture.

(in mm). H_s: 14.5–20.6 (holotype 18.7), W_s: 3.7–5.0 (4.2).

Albania, Mat District, Ura e Vashës, in the gorge of the Lumi i Matit, at its confluence with the Përroi i Gurri i Bardhit, 350 m, 41.4677°N, 20.1048°E.

Type locality, leg. ZF, JK, DM, 9.x.2004, holotype (HNHM 94848), paratypes (HNHM 94849/26, NHMW 103278, SMF 328082).

Type locality, leg. ZF, TN, EM, 15.iv.2014 (HNHM 99010); between Fshat and Shkallë, N slope of the Mt. Kjuteti, 450 m, 41.4794°N, 20.0814°E, leg. ZB, DP, B. Pintér, 27.v.2007 (HNHM 99619); 6 km S of Gurri i Bardhë along the Klos to Elbasan road, gorge of the Lumi i Guisës, 1030 m, 41.4306°N, 20.0920°E, leg. LD, ZE, ZF, AH, DM, 30.vi.2007 (HNHM 99622); Librazhd District, Orenjë, 720 m, 41.2802°N, 20.2100°E, leg. ZF, EM, 13.iv.2014 (HNHM 98962).

Gropa-Bizë-Martanesh mountain group in central Albania. The most distant sites are ca. 25 km from each other (Fig. 18).

Shell small to medium. All whorls indistinctly wrinkled-costate. Neck wrinkled-costate. Peristome pear-shaped. In front view lamella inferior barely emerged. Lunella lateral-ventrolateral, fused to the short basalis. Subclaustralis residual, sulcalis residual or absent. Anterior plica superior long. Clausilium plate not visible through the aperture.

(in mm). H_s: 13.2–16.9 (holotype 15.7), W_s: 3.0–3.9 (holotype 3.3).

Albania, Elbasan District, Petresh (= 5 km SSE of Graçen), along the Tiranë to Elbasan road, S of the village, 440 m, 41.1030°N, 20.0064°E.

Type locality, leg. ZE, 19.viii.1993, holotype (HNHM 70838), paratypes (HNHM 70839/2, NHMW 100079/2, SMF 312507/2, NMBE 534906/1, NMBE 22594/1).

Type locality, leg. ZF, TK, DM, 22.vi.2012 (HNHM 99617); “Elbasan Paß”, leg. Jaeckel (NHMW-K 57868/1, NHMW-K 47293/1); 1 km SE of Shkamë (near Mollas), W side of the Mt. Sulovë, 480 m, 40.9280°N, 20.0305°E, leg. ZE, ZF, JK, DM, 29.vi.2003 (HNHM 94904).

Krabba and Sulovë Mts in central Albania. The type locality is in the southern part of the Krraba Mountain, and presumably the site “Elbassan-paß, 800 m” given by Jaeckel and Schmidt (1961) can be in the vicinity. We found this subspecies in the Mt. Sulovë, at ca. 20 km from the type locality, and isolated from it by a wide, non-rocky flysch zone. Some other locality records in the literature from the Mt. Sulovë (Dhora and Welter-Schultes 1999, Nordsieck 2009) presumably also correspond to this taxon (Fig. 18).

Shell small to medium. Lower whorls smooth, upper ones indistinctly wrinkled-costate. Neck densely costate. Basal crest strong, peripheral crest very strong. Peristome pear-shaped. In front view lamella inferior more or moderately emerged. Lunella lateral, fused to the short basalis. Subclaustralis and sulcalis residual or absent. Anterior plica superior long. Clausilium plate not visible through the aperture.

(in mm). H_s: 13.2–16.9, W_s: 3.4–4.2.

“Berat (Perat) in S-Albanien, am Abhang des Festungsberges” = Albania, Berat district, Berat, slope of the castle hill.

“Perat, Festungberg”, ex Zilch, leg. Fuchs, 1936, holotype (SMF 167016), paratypes (SMF 167017/2, SMF 201625/2, NHMW-E 16552/20, NHMW-K 48296/5, ZMH 85772/3); “Berat, Festungsberg”, leg. Fuchs, ex Brandt, ? paratypes (type status not indicated on the label) (ZSM 20150455/3).

Berat, SW side of the castle hill, 70 m, 40.7041°N, 19.9479°E, leg. DA, ZE, ZF, JG, 27.vi.2014 (HNHM 99616, NHMW 110430/MN/0077).

Known only from the castle hill of Berat (type locality), which is one of the westernmost foothills of the Tomorr Mts (Fig. 18).

Shell small to large, light corneous to light brown. Lower whorls smooth to costate, sculpture stronger toward the apex. Neck variably inflexed. Basal crest weak to strong, peripheral crest weak to not recognizable. Peristome circular to ovoid, with simple to swollen margin. Lamellae superior and spiralis with no to long overlap. In front view lamella inferior more or less well emerged, broadly-bent subcolumellaris variably exposed. Lunella dorsal to dorsolateral. Basalis absent to strong, fused to or separate from the lunella. Subclaustralis absent to weak, sulcalis rudimentary to strong. Anterior plica superior present, connected to or separate from the lunella complex. Clausilium plate partly visible through the aperture.

Shell small to medium, tumid, light corneous. Lower and upper whorls distinctly costate, ribs sharper behind the aperture. Neck weakly inflexed. Basal crest strong, peripheral one not recognizable. Peristome detached, circular to ovoid, with swollen, reflexed margin. Lamellae superior and spiralis mostly overlap. In front view lamella inferior well emerged, medium-bent subcolumellaris mostly visible. Lunella dorsal-dorsolateral, weakens toward the basis. Basalis absent or weak and separate from the lunella. Subclaustralis very short, sulcalis present. Anterior plica superior occasionally connected to the lunella complex.

(in mm). H_s: 13.9–17.4 (holotype 15.9), W_s: 4.0–5.0 (holotype 4.3).

Figure 19. 

A Montenegrina hiltrudae hiltrudae Nordsieck, 1972, holotype, SMF 221046 B M. hiltrudae costulata Erőss & Szekeres, 2006, holotype, HNHM 94893 C M. hiltrudae dennisi Gittenberger, 2002, paratype, SMF 331782 D M. hiltrudae densicostulata Nordsieck, 1974, holotype, SMF 227686 E M. hiltrudae desaretica ssp. n., holotype, NHMW 111250 F M. hiltrudae fusca Fehér & Szekeres, 2006, holotype, HNHM 94895 G M. hiltrudae maasseni Gittenberger, 2002, paratype, SMF 331784 H M. hiltrudae protruda Gittenberger, 2002, paratype, SMF 331780 I M. hiltrudae robusta Nordsieck, 2009, SMF 328778 J M. hiltrudae sattmanni Nordsieck, 1988, holotype, NHMW 84028 K M. hiltrudae selcensis ssp. n., holotype, NHMW 111216 L M. minuscula Erőss & Szekeres, 2006, holotype, HNHM 94831. Scale bar: 5 mm.

Greece, Western Macedonia, near Vogatsikon, on the road to Germas (= Yermas).

Type locality, leg. H. Nordsieck, 23.viii.1971, holotype (SMF 221046), paratypes (SMF 221047, SMNS-N 5476, NHMW 77935/5, NHMW-K-65052/11).

Southern part of the Verno Mts in Western Macedonia (Greece). Known only from the type locality (Fig. 20).

Figure 20. 

Distribution of Montenegrina hiltrudae. M. hiltrudae costulata (empty circle with dot); M. hiltrudae dennisi (triangle); M. hiltrudae densicostulata (triangle pointing right); M. hiltrudae desaretica ssp. n. (empty circle); M. hiltrudae fusca (square); M. hiltrudae hiltrudae (triangle pointing left); M. hiltrudae maasseni (inverted triangle); M. hiltrudae protruda (empty triangle); M. hiltrudae robusta (diamond); M. hiltrudae sattmanni (circle); M. hiltrudae selcensis ssp. n. (star).

Shell small to medium, conical, whitish-corneous. Surface distinctly costate, ribs sharper on the upper whorls, denser at the neck. Neck very weakly inflexed. Basal crest strong, peripheral crest not recognizable. Peristome detached, ovoid, with somewhat swollen margin. Lamellae superior and spiralis long overlap. In front view lamella inferior more or less well emerged, broadly-bent subcolumellaris mostly not visible. Lunella dorsal, short and broad, mostly separate from the strong basalis. Subclaustralis short, sulcalis strong. Anterior plica superior mostly connected to the lunella complex.

(in mm). H_s: 14.1–20.4 (holotype 16.0), W_s: 3.7–5.3 (holotype 4.8).

Albania, Korçë District, Qafa e Zvezdës, 4 km from Zvezdë along the road to the Prespa Lake, 1030 m, 40.7330°N, 20.8729°E.

Type locality, leg. ZE, ZF, JK, DM, 2.vii.2003, holotype (HNHM 94893), paratypes (NHMUK 20050231, HNC 63195, HNHM 94894/77, NHMW 103293, RMNH 100323, SMF 328097).

Type locality, leg. ZE, ZF, JG, 29.vi.2013 (HNHM 99590); transitional forms to M. h. sattmanni: Korçë District, Mt. Thatë, ca. 2.7 km NNW of Zvezdë, ca. 500 m SW of the Zvezdë Summit, 1720 m, 40.7565°N, 20.8494°E, leg. ZB, CN, DP, 25.v.2007 (HNHM 99589); Zaroshka, 870 m, 40.7668°N, 20.9277°E, leg. ID, 27.v.2013 (DED).

Southern part of the Mt. Thatë, in the vicinity of the Zvezdë Pass that connents the Prespa and Korçë Basins (Fig. 20).

Shell medium, light corneous. Lower whorls smooth, upper ones moderately wrinkled-costate. Neck weakly inflexed, finely costate. Basal and peripheral crests weak. Peristome attached, ovoid, with somewhat swollen margin. Lamellae superior and spiralis mostly overlap. In front view lamella inferior well emerged, broadly-bent subcolumellaris usually not visible. Lunella dorsal, short and broad. Basalis absent or residual, separate from the lunella. Subclaustralis absent, sulcalis well developed. Anterior plica superior weak, not connected to the lunella complex.

(in mm). H_s: 15.7–21.2 (holotype 18.7), W_s: 4.3–5.2 mm.

Greece, Western Macedonia, 1.5 km SE of Zakas, 2.5 km along the road to Spileo, N slope, 900 m.

Type locality, leg. Gittenberger, Uit de Weerd, holotype (RMNH 94939), paratypes (RMNH 94940/50); same locality, leg. Gittenberger, Maassen, 23.v.2001, paratypes (RMNH 82021/30, SMF 331782/3, NMBE 534890/3); Spileo, west side, 3.25 km S of Zakas, SSW slope, 930 m, leg. Gittenberger, Maassen, 24.v.2001, paratypes (RMNH 82027/32, SMF 331783/3, NMBE 534889/3).

Greece, 0.5 km N of Spileo, toward Zakas, 930 m, 40.0080°N, 21.2847°E, leg. ZE, ZF, JG, 22.vi.2013 (HNHM 99591); 3 km E of Zakas, 900 m, 40.0422°N, 21.2823°E, leg. ZE, AH, 15.vii.2004 (HNHM 95393); 0.5 km W of Spileo, toward Portitsa Farangi, 940 m, 40.0077°N, 21.2805°E, leg. ZE, ZF, JG, 22.vi.2013 (HNHM 99592).

Western Macedonia (Greece). All known localities are within a range of a few km on the Mt. Orliakas, reaching to the Venetikos River in the the north (Fig. 20).

Originally M. h. dennisi was described as a separate species (Gittenberger 2002). However, its shell features (and also those of the closely related M. h. protruda Gittenberger, 2002) fit well in the varability range of M. hiltrudae subspecies.

Shell small to medium, tumid, light corneous. Lower whorls striate to densely costate, upper ones with stronger ribs. Neck weakly inflexed. Basal crest strong, peripheral crest not recognizable. Peristome detached, circular to ovoid, with swollen, reflexed margin. Lamellae superior and spiralis mostly overlap. In front view lamella inferior well emerged, medium-bent subcolumellaris usually visible. Lunella dorsal-dorsolateral, weakens toward the basis. Basalis absent or short, separate from the lunella. Subclaustralis very short, sulcalis well developed. Anterior plica superior occasionally connected to the lunella complex.

(in mm). H_s: 15.8–22.4 (holotype 21.9), W_s: 4.3–6.1 (holotype 6.1).

Greece, Western Macedonia, Srianovon, along the Neapolis to Vogatsikon road.

Type locality, leg. H. Nordsieck, 23.viii.1971, holotype (SMF 227686), paratypes (SMF 227687, SMNS-N 5475, NMBE 534967/2, NHMW-K-65051/10).

Greece, Kastoria District, above a quarry along the Vogatsiko to Driovouno road, 680 m, 40.3782°N, 21.4160°E, leg. ZE, ZF, JG, 28.vi.2013 (HNHM 99584); 1 km W of Germas, dry gorge, 840–880 m, 40.4430°N, 21.4111°E, leg. ZE, ZF, JG, 28.vi.2013 (HNHM 99582, HNHM 99583).

Southern part of the Verno Mountain in Western Macedonia (Greece). Found at some sites around Vogatsiko, southeast of the Kastoria Lake (Fig. 20).

Originally Nordsieck (1972) classified M. h. densicostulata with the nominotypical subspecies, but only two years later he described it as a distinct taxon (Nordsieck 1974).

Medium-size subspecies with costate shell, attached peristome, and non-overlapping lamellae superior and spiralis.

The light greyish-corneous shell of 8½ to 10½ whorls is tumid, with relatively large aperture. All whorls are more or less costate, more distinctly toward the apex. The neck is not or only very weakly inflexed, its ribs become sharper and near the aperture. The basal crest is weak, the peripheral one is not recognizable. The aperture is almost circular, its broadly attached margin is somewhat swollen and deflexed. The lamella superior is low but long, it does not overlap with the spiralis. In front view the lamella inferior is moderately emerged, the end of the broadly-bent subcolumellaris is mostly visible. The short and broad lunella is dorsal-dorsolateral, it is connected to the often weak basalis. The subclaustralis is absent, the sulcalis is rudimentary. The anterior part of the plica superior is weak, not connected to the lunella complex. The clausilium plate is partly visible through the aperture.

(in mm). Holotype H_s: 17.6, W_s: 5.5, H_a: 4.5, W_a: 4.1; paratypes H_s: 15.0–18.9 (mean 16.9, S.D. 1.33), W_s: 4.4–5.9 (mean 5.3, S.D. 0.39), H_a: 3.8–5.2, W_a: 3.3–4.3.

The new subspecies resembles the nearby occurring M. h. sattmanni, but can be distinguished from it by the costate shell, broadly attached peristome, non-overlapping lamellae superior and spiralis, as well as the reduced subclaustralis, sulcalis and anterior plica superior. From all other M. hiltrudae subspecies differs by the combination of an attached peristome and costate lower whorls.

Albania, Korçë District, cave temple ca. 2 km S of Glloboçeni, at the Prespa Lake, 850 m, 40.8422°N, 20.9616°E.

Type locality, leg. ZE, ZF, JG, 30.vi.2015, holotype (NHMW 111250), paratypes (NHMW 111251/20+5a+11aj, HNHM 99690/25, GR/25, SZ/5); same locality, leg ID, 23.ix.2013, paratypes (DED/17).

Albania, Korçë District, 1 km NE of Liqenas, Sveti Atanas i Veliki Antoni Church, 850 m, 40.7919°N, 20.9152°E, leg. ZE, ZF, JG, 28.vi.2013 (HNHM 99587); same locality, leg. ZE, ZF, JK, DM, 2.vii.2003 (HNHM 94901); Liqenas, 860 m, 40.7891°N, 20.9072°E, leg. ZE, ZF, JK, DM, 2.vii.2003 (HNHM 94898); Ishull i Vogël, E of Liqenas, 860 m, 40.7915°N, 20.9324°E, leg. ZF, LT, 17.viii.2007 (HNHM 99572).

The new taxon is named after Desaretia, the ancient Roman name of the Ohrid–Prespa region.

Prespa Lake area. Apart from the type locality, it was also found in the vicinity of Liqenas (Fig. 20).

Formerly the Liqenas population was mentioned by Erőss et al. (2006) as differing from the typical M. h. sattmanni by the stronger apical sculpture. However, more recent morphological and distribution data suggest that the populations around Liqenas belong to M. h. desaretica, but differ from its typical form by the weaker sculpture and the occasionally connected anterior plica superior.

Shell medium, light brown, paler along the suture. Lower whorls smooth, upper ones indistinctly striate-costate. Neck weakly inflexed, densely striate behind the aperture. Basal crest weak, peripheral crest not recognizable. Peristome attached, ovoid, with somewhat swollen margin. Lamellae superior and spiralis overlap. In front view lamella inferior well emerged, broadly-bent subcolumellaris not visible. Lunella dorsal-dorsolateral, short and broad, mostly fused to the basalis. Subclaustralis short, sulcalis strong. Anterior plica superior connected to the lunella complex, forward reaches as far as the plica principalis.

(in mm). H_s: 18.3–22.4 (holotype 22.4), W_s: 4.5–5.3 (holotype 5.3).

Albania, Korçë District, Korit e Bregas, 6 km S of Podgorje along the Pogradec to Zvezdë road, 900 m, 40.7688°N, 20.8275°E.

Type locality, leg. ZE, ZF, JK, DM, 2.vii.2003, holotype (HNHM 94895), paratypes (HNHM 94896/9, NHMW 103294, RMNH 100324, SMF 328098).

Type locality, leg. ZE, ZF, JG, 29.vi.2013 (HNHM 99596); Veliterna, leg. Megyeri, 7.vii.1960 (HNHM 99597); Mt. Thatë, near the ‘Tower of the war’, 1830 m, 40.9136°N, 20.8419°E, leg. ID, 24.ix.2013 (DED).

Mt. Thatë in southern Albania. The type locality is at the western foot of the mountain, and this taxon was also found in the alpine region at the ‘Tower of the war’, near the Albanian–Macedonian border (Dedov, unpublished data). At this locality it occurs syntopically with M. dofleini wagneri (Fig. 20).

Shell small to medium, light corneous. Lower whorls smooth, upper ones indistinctly wrinkled-costate. Neck inflexed, costate. Basal and peripheral crests weak. Peristome attached, ovoid, with simple margin. Lamellae superior and spiralis mostly overlap. In front view lamella inferior more or less well emerged, broadly-bent subcolumellaris occasionally visible. Lunella dorsal, short and broad. Basalis residual, separate from the lunella. Subclaustralis absent, sulcalis well developed. Anterior plica superior mostly not connected to the lunella complex.

(in mm). H_s: 14.5–18.6 (holotype 16.0), W_s: 3.6–4.1.

Greece, Western Macedonia, Portitsa bridge, 4 km SE of Zakas, 625 m.

Type locality, leg. Gittenberger, Maassen, 24.v.2001, holotype (RMNH 94949), paratypes (RMNH 82024/28, NMBE 535056/2, SMF 331784/2).

Greece, Portitsa Farangi, near Spileo, 39.9965°N, 21.2855°E, leg. ZE, ZF, JG, 22.vi.2013 (HNHM 99595); same locality, leg. ZE, AH, 15.vii.2004 (HNHM 99594).

Western Macedonia (Greece). Known only from its type locality, the gorge of the Venetikos River. M. h. dennisi occurs only a few few km farther north, at the slope of the Mt. Orliakas, bordered by the Venetikos Valley (Fig. 20).

Montenegrina h. maasseni was described by Gittenberger (2002) as a subspecies of M. janinenis, but its shell morphology suggests a close relationship to M. h. dennisi (see: Erőss et al. 2006).

Shell medium, light to brownish-corneous. Lower whorls smooth, upper ones finely striate. Neck very weakly inflexed, finely costate. Basal and peripheral crests weak. Peristome detached, ovoid, with somewhat swollen margin. Lamellae superior and spiralis mostly overlap. In front view lamella inferior well emerged, broadly-bent subcolumellaris not visible. Lunella dorsal, short and broad, not fused to the basalis. Subclaustralis not recognizable. Sulcalis well developed. Anterior plica superior weak, not connected to the lunella complex. Lunella dorsal-dorsolateral to dorsolateral, short and broad, mostly fused to the basalis. Subclaustralis weak or absent, sulcalis well developed. Anterior plica superior not connected to the lunella complex.

(in mm). H_s: 16.0–20.6 (holotype 19.0), W_s: 4.8–5.4 mm.

Greece, Western Macedonia, 2.3 km before Aetia along the Aetia to Anavrita road, along the path to the Nymphoon Cave, W slope, 1000 m.

Type locality, leg. Gittenberger, Uit de Weerd, holotype (RMNH 94942), paratypes (RMNH 94941/21); same locality, leg. Gittenberger, Maassen, 23.v.2001, paratypes (RMNH 82016/19, SMF 331780/3, NMBE 534891/3); hill 1.75 km E of Aetia, E slope, 940 m, paratypes (RMNH 82017/7).

Type locality, 990 m, 40.0741°N, 21.2017°E, leg. ZE, ZF, JG, 22.vi.2013 (HNHM 99593).

Western Macedonia (Greece). Known only from the type locality (Fig. 20).

Shell medium to large, brownish-corneous. Very weak and indistinct ribs of the lower whorls become stronger and better defined toward the apex. Weakly inflexed neck has more robust ribs. Basal and peripheral crests weak. Peristome ovoid, mostly attached, with swollen, reflexed margin. Lamellae superior and spiralis overlap. In front view lamella inferior more or less well emerged, broadly-bent subcolumellaris visible. Lunella short and broad, dorsal-dorsolateral to dorsolateral, mostly fused to the basalis. Subclaustralis residual or absent, sulcalis well developed. Anterior plica superior long, often connected to the lunella complex.

(in mm). H_s: 18.0–25.1 (holotype 21.4), W_s: 4.55–5.8 (holotype 5.2).

Greece, Western Macedonia, 4 km from Gavros toward Kotas along the Kastoria to Florina road, 850 m.

Type locality, leg. Hemmen, 13.iv.1995, holotype (SMF 328778), paratypes (SMNS-N 10260/5).

Same as types (NHMW 110430/MN/0117); type locality, 40.6532°N, 21.1789°E, leg. ZF, EH, KJ, HS, 18.x.2014 (NHMW 110430/MN/0118).

Western part of the Verno Mts in Western Macedonia (Greece). Known only from the type locality, located between Kastoria and the Prespa Lakes (Fig. 20).

Some specimens of the original series (e.g. those of NHMW 110430/MN/0117) were not designated as types.

Shell medium, light corneous. Lower whorls smooth, upper ones finely striate to indistinctly costate. Neck not or very weakly inflexed, densely striate. Basal crest very weak, peripheral crest not recognizable. Peristome attached, ovoid, with somewhat swollen margin. Lamellae superior and spiralis overlap. In front view lamella inferior well emerged, broadly-bent subcolumellaris mostly visible. Lunella dorsal-dorsolateral, short and broad, separate from the basalis. Subclaustralis short, sulcalis strong. Anterior plica superior long, mostly connected to the lunella complex.

(in mm). H_s: 17.3–21.4 (holotype 20.1), W_s: 4.6–5.5 (holotype 5.3).

Greece, Western Macedonia, Mikrolimni, near the Biological Station.

Type locality, leg. HS, 24.v.1986, holotype (NHMW 84028), paratypes, (NHMW 84029/7, SMNS-N 9520/2).

Type locality, 40.7429°N, 21.1102°E, leg. ZE, ZF, JG, 28.vi.2013 (HNHM 99585); 500 m N of the Agios Achillios junction (Psarades to Vrondero road), 870 m, 40.7998°N, 21.0715°E, leg. ZF, EH, KJ, HS, 18.x.2014 (NHMW 110430/MN/0065); 3 km SE of Vrondero, Petros Kokkalis Cave, 1000 m, 40.7192°N, 21.0341°E, leg. ZF, EH, KJ, HS, 18.x.2014 (NHMW 110430/MN/0066); Albania, Korçë District, Tren, Shpella e Trenit, bank of the Small Prespa Lake, 860 m, 40.6723°N, 20.9871°E, leg. ZE, ZF, JG, 28.vi.2013 (HNHM 99586); Mt. Thatë, ca. 1.7 km N of Zvezdë, 2.1 km southeast of the Zvezdë Summit, 1180 m, 40.7419°N, 20.8635°E, leg. ZB, CN, DP, 25.v.2007 (HNHM 99588); Zagradets, 870 m, 40.6767°N, 21.0039°E, leg. ID, 24.vi.2013 (DED); Cerie, 1130 m, 40.7523°N, 20.9434°E, leg. ID, 24.vi.2013 (DED); Rakitsko, 1100 m, 40.7245°N, 20.9768°E, leg. ID, 26.v.2013 (DED); Glloboçeni, S side, boat harbor, 850 m, 40.8584°N, 20.9474°E, leg. ZE, ZF, JG, 30.vi.2013 (NHMW 110430/MN/0067); N of Glloboçeni, 860 m, 40.8580°N, 20.9426°E, leg. ZF, EH, KJ, HS, 17.x.2014 (NHMW 110430/MN/0064);

The area of the Prespa Lakes (Fig. 20).

Originally M. h. sattmanni was described as a distinct species (Nordsieck 1988), but its shell characteristics and distribution range indicate its belonging to M. hiltrudae.

The paratype lot NHMW 84030/4 could not be found in the NHMW collection.

Medium-size subspecies with large, very narrowly attached peristome and in front view well visible lamella subcolumellaris.

The medium-size, light brownish-corneous shell of 9 to 10½ whorls is tumid, with brad basis. The surface is smooth at the lower whorls but becomes indistinctly and densely striate-costate toward the apex. The neck is only very weakly inflexed, behind the peristome it is finely and densely striate. The basal crest is weak, the peripheral crest is not recognizable. The aperture is very large, with the peristome its height reaches one quarter of that of the entire shell. The light whitish-brown peristome is wide, narrowly attached, its margin is swollen, deflexed. The lamellae superior and spiralis overlap. The lamella inferior is strongly emerged, its inner part bends close to the spiralis. The broadly-bent lamella subcolumellaris is visible in front view. The lunella is dorsal-dorsolateral, short and broad, mostly separate from the basalis. The subclaustralis short, the sulcalis is well developed. The anterior part of the plica superior is long, ends in a small lump behind the aperture. Its inner end occasionally extends to the lunella complex. The clausilium plate is entirely visible through the aperture.

(in mm). Holotype H_s: 19.9, W_s: 6.2, H_a: 5.5, W_a: 4.4; paratypes H_s: 16.5–21.4 (mean 19.4, S.D. 1.40), W_s: 5.1–6.2 (mean 5.7, S.D. 0.39), H_a: 4.6–5.8, W_a: 4.0–4.9 (NHMW 111217, NHMW 111218, HNHM 99481, HNHM 99482, n = 12).

The new subspecies can be distinguished from M. h. dennisi and M. h. maasseni by its much finer neck sculpture, from M. h. fusca by its more conical shell and stronger lamella inferior, from M. h. sattmanni by the finer apical and neck sculpture, weaker basalis and more dorsal lunella, whereas from the rest of M. hiltrudae subspecies by the attached peristome.

Albania, Korçë District, NE of Strelcë, in the limestone gorge of the Lumi i Verbës at the foot of the Shkëmb i Selcës, 990 m, 40.7480°N, 20.5219°E.

Type locality, leg. ZF, JG, 30.vi.2014, holotype (NHMW 111216), paratypes (NHMW 111218/1+2a, HNHM 99482/3, GR/2); between Strelcë and Selcë, 1090 m, 40.7462°N, 20.5123°E, leg. DA, ZE, 30.vi.2014, paratypes (NHMW 111217/19+2a, HNHM 99481/20, GR/20, SZ/5).

The new taxon is named after the Selcë Cliff (Shkëmb i Selcës), its type locality.

Southern part of the Mokër Mts. Two nearby localities are known in the vicinity of Strelcë. At the type locality this taxon is found syntopically with M. laxa errans Erőss & Szekeres, 2006 (Fig. 20).

Shell small, light corneous. Lower whorls smooth, upper ones with fine, widely-spaced, indistinct ribs. Neck inflexed, wrinkled-costate. Basal and peripheral crests well visible. Peristome attached, rounded to somewhat angular, with slightly to strongly swollen, reflexed margin. Lamellae superior and spiralis overlap. In front view lamella inferior moderately emerged, broadly-bent subcolumellaris visible. Lunella dorsolateral, mostly separate from the weak basalis. Subclaustralis residual, sulcalis well developed. Anterior plica superior not connected to the lunella complex. Clausilium plate partly visible through the aperture.

(in mm). H_s: 12.2–17.1, W_s: 3.2–4.2 (Perama, HNHM 99560).

Figure 21. 

A Montenegrina laxa laxa (Küster, 1861), Tërkuza Valley HNHM 99033 B M. laxa dedovi Nordsieck, 2009, holotype, SMF 332465 C M. laxa delii ssp. n., holotype, NHMW 111237 D M. laxa disjuncta Fehér & Szekeres, 2006, holotype, HNHM 94871 E M. laxa errans Erőss & Szekeres, 2006, holotype, HNHM 94874 F M. laxa iba Nordsieck, 1972, holotype, SMF 201629a G M. laxa kontschani Erőss & Szekeres, 2006, holotype, HNHM 94878 H M. laxa lakmosensis Nordsieck, 2009, paratype, SMF 332354 I M. laxa miraka Nordsieck, 1996, Mirakë, HNHM 99630 J M. janinensis (Mousson, 1859), syntype, SMF 176316 K M. janinensis (Mousson, 1859), holotype of crassilabris, SMF 309237 L M. lillae sp. n., holotype, HNHM 99496. Scale bar: 5 mm.

“environs de Ianina” = Greece, Epirus, Ioannina [janinensis]; Greece, Epirus, Kria near Perama, above the southeastern cemetery, 600 m [crassilabris].

“Janina, Epirus”, ex Schaefli, ex Mousson, syntypes [janinensis] (SMF 176316/6); same locality, ex Boettger, syntypes [janinensis] (SMF 176317/3); same locality, ex Shuttleworth, syntypes [janinensis] (NMBE 21041/6); Kria near Perama, leg. Fauer, 10.viii.1971, holotype [crassilabris] (SMF 309237), paratypes [crassilabris] (SMF 309238/3, HNHM 39968/3, SMNS-N 9791, ZMH 91624/pl.); same locality, leg. PS, Fauer, 27.ix.1989, paratypes [crassilabris] (NMBE 535019, NMBE 22589/3, NHMW 103731, ZMH 91623/pl.); Kria, above the church, leg. Hausdorf, 31.v.1985, paratypes [crassilabris] (ZMH 91622/pl.); Kria, above the southeastern cemetery, 600 m, leg. Fauer, 22.iv.1988, paratypes [crassilabris] (ZMH 91627/pl.); 1.2 km from Kria toward Kranoula, ex Fauer, 10.viii.1971, paratypes [crassilabris] (ZMH 91625/6); Perama, exit of the stalactite cave, leg. PS, 29.vii.1975 paratypes [crassilabris] (HNHM 67025/4, NMBE 535017/60, ZMH 91626/9).

Greece, Epirus, Amfithea–Spothi road N of Lake Pamvotis, 2 km E of Ligkiades junction, 640 m, 39.6802°N, 20.9052°E, leg. ZE, ZF, JG, 24.vi.2013 (HNHM 99559); Perama, near entrance of the cave, 500 m, 39.6947°N, 20.8463°E, leg. ZE, ZF, JG, 24.vi.2013 (HNHM 99560); Mitsikeli Mts, S of Dikórifo, 1070 m, 39.7828°N, 20.8041°E, leg. PS, 23.v.1991 (NMBE); Kria, dry gorge above the cemetery, 660 m, 39.7245°N, 20.8429°E, leg. ZE, ZF, JG, 24.vi.2013 (HNHM 99562); Perama, near the exit of the stalactite cave, 520 m, 39.6971°N, 20.8433°E, leg. ZE, ZF, JG, 24.vi.2013 (HNHM 99561).

Southwestern part and foothills of the Mitsikeli Mts in northwestern Greece. Most of the known occurrences are along the northern shore of the Pamvotis Lake (between Kria and Spothi), but a record from Dikórifo indicates that the species may be wider distributed (Fig. 22).

Figure 22. 

Distribution of Montenegrina janinensis; M. lillae sp. n. and M. minuscula. M. janinensis (circle); M. lillae sp. n. (square); M. minuscula (triangle).

Although not synonymized formally, Nordsieck (2009) already questioned the distinct taxonomic status of crassilabris, assuming that it might only be a local form of M. janinensis with thickened peristome. This morphotype is known from Kria and Perama, but its range is not clearly distinct from that of the typical janinensis. Moreover, on the Goritsa Hill at Perama they live in parapatry: at the entrance of the stalactite cave typical janinensis, whereas at the exit (only a few hundred meters farther) the form with the thickened peristome (Fauer 1993, and also our personal observation).

Based mainly on their small shell size, originally several non-related Montenegrina taxa were described as subspecies of M. janinensis. These include M. soosi Erőss & Szekeres, 2006, and some subspecies currently classified with M. attemsi, M. dofleini, M. grammica, M. hiltrudae, M. sporadica Nordsieck, 1974, and M. tomorosi Brandt, 1961.

Shell small to large, yellowish- to brownish-corneous. Lower whorls smooth, upper ones smooth to finely wrinkled-costate. Neck variably inflexed, finely striate to wrinkled-costate. Basal and peripheral crests recognizable. Peristome attached, ovoid to angular, with simple to swollen and reflexed margin. Lamellae superior and spiralis distant to long overlapping. In front view lamella inferior mostly well emerged. Weakly-bent subcolumellaris and the clausilium plate not or only barely visible through the aperture. Lunella short, broad and diffuse, dorsolateral to ventrolateral. Basalis is fused to the lunella as its almost straight continuation. Lunella complex also fused to the plica principalis. Subclaustralis short, sulcalis mostly residual or absent. Anterior plica superior usually long, distant and diverging from the principalis, in most subspecies not connected to the lunella complex.

Shell medium to large, yellowish-corneous. All whorls smooth. Neck weakly inflexed, wrinkled-costate. Basal and peripheral crests weak. Peristome ovoid to somewhat angular, with simple margin. Lamellae superior and spiralis mostly do not overlap. Subcolumellar lamella barely, clausilium plate not visible through the aperture. Lunella lateral. Subclaustralis shorter than the basalis, sulcalis residual or absent. Anterior plica superior long, almost parallel to the plica principalis, not connected to the lunella complex.

(in mm). H_s: 16.3–21.8, W_s: 4.6–5.9 mm (Nordsieck 1972).

“wahrscheinlich aus der Gegend von Castel nuovo” = Montenegro, probably from the area of Herceg-Novi [laxa]; “monts Mali Dajtit à l’Est de la ville de Tirana, près de la route de Tuffina à Safa Muzizes” = Albania, Mt. Dajti east of Tiranë, route from Tufinë to the Qafa e Murrizës [weigneri]; “la vallée de la rivière Terküza” = Albania, N of Tiranë, gorge of the Lumi i Tërkuzës [weigneri].

Syntypes of weigneri are thought to be in the MIZPAS (not seen).

Albania, Tirana, “Römerbrücke” (= Roman bridge), leg. Fuchs (SMF 94109a [Nordsieck 1972: fig. 40], NHMW-E 32318, NHMW-K 54402, NHMW-K 50578, SMF 94109); “Mal i Krujë” (likely erroneous locality record), leg. Fuchs (NHMW-E 32479, NHMW-K 50575); Tiranë District, NE of Herraj, gorge of the Lumi i Tërkuzës near the dam of the Lumi i Bovillës, 330 m, 41.4446°N, 19.8661°E, leg. ZF, TN, EM, 16.iv.2014 (HNHM 99032); gorge of the Lumi i Tërkuzës 0.5 km beneath the dam, 240 m, 41.4412°N, 19.8636°E, leg. ZF, TN, EM, 16.iv.2014 (HNHM 99033); gorge of the Lumi i Tiranës, 5 km NE of Ferraj, 290 m, 41.3948°N, 19.9000°E, leg. ZF, TN, EM, 16.iv.2014 (HNHM 99035); Shkalla e Tiranit, gorge of the Lumi i Tiranës, 400 m, 41.4022°N, 19.9012°E, leg. ZE, ZF, KK, 10.iv.2001 (HNHM 86044); between Ferraj and Shtish (NE of Tiranë), at the bridge of the Lumi i Tiranës, 200 m, 41.3798°N, 19.8592°E, leg. ZE, ZF, KK, 10.iv.2001 (HNHM 86043).

Dajti-Kruja mountain range in central Albania. Known from the gorges of the Tiranë and Terkuzë Rivers, north of Tiranë (Fig. 23).

Figure 23. 

Distribution of Montenegrina laxa. M. laxa dedovi (empty circle with dot); M. laxa delii ssp. n. (empty circle); M. laxa disjuncta (diamond); M. laxa errans (circle); M. laxa iba (empty triangle); M. laxa kontschani (inverted triangle); M. laxa lakmosensis (star); M. laxa laxa (triangle); M. laxa miraka (square).

The illustration with the name laxa was published in 1860, but a description of the species only in 1861 (both in Küster 1844–1862). Therefore the latter is considered the year of description (Welter-Schultes 1999).

Montenegrina species do not occur in the area of Herceg-Novi, which was given by Küster (1844–1862) as the type locality. This record is either incorrect or based on marine flotsam (Nordsieck 1972). Without type material and reliable type locality this taxon could be identified using the precise description of Schmidt (1868), which was based on the type series. In the view of Wagner (1924), laxa is a synonym of wohlberedti and not identical with weigneri. By contrast, Nordsieck (1972) claims that the Terküza Valley population fits well Schmidt’s description, therefore he considers laxa a senior synonym of weigneri. The latter view has been widely accepted in the past 40 years. In order to preserve nomenclatural stability, we stick to this practice.

Shell small to medium, light corneous. All whorls smooth. Neck weakly inflexed, finely and densely striate. Basal and peripheral crests weak. Peristome, ovoid to somewhat angular, with simple margin. Lamellae superior and spiralis do not overlap. Subcolumellar lamella not or barely, clausilium plate not at all visible through the aperture. Lunella lateral. Subclaustralis short to residual, sulcalis very weak or absent. Anterior plica superior distant and diverging from the plica principalis, not connected to the lunella complex.

(in mm). H_s: 13.6–20.3 (holotype 16.1), W_s: 3.7–4.4 (holotype 4.15) (Nordsieck 2009, Dedov and Neubert 2009).

Macedonia, Jablanica Mts, Vartop region, after the Čuma Summit, 1868 m.

Type locality, leg. ID, 15.vii.2006, holotype (SMF 332465), paratypes (SMF 331838/4).

Macedonia, Jablanica Mts, up to Gorna Belitsa, alpine limestone meadow, ca. 1600–1700 m, 41.22°N, 20.54°E, leg. ID, Minkov, 10.vii.2009 (DED-580); near the Kokal Summit, 1800–1928 m, 41.203°N, 20.532°E, leg. ID, 15.vii.2009 (DED-560); Tchumata area (up to the Krustets area), alpine limestone meadows, 1980 m, leg. ID, 12.vii.2009 (DED-582).

This taxon is known from the alpine region of the Jablanica Mts (Fig. 23).

Originally M. l. dedovi was described as a separate species (Nordsieck 2008), but its shell structure (and particularly its palatal plicae) positions it among the subspecies of M. laxa.

Large subspecies with tumid shell, long overlapping lamellae superior and spiralis.

The large, tumid, brownish-corneous shell consists of 8½ to 10½ whorls. The surface is smooth at the lower whorls, whereas toward the apex it becomes indistinctly stiate-costate. The neck is also indistinctly costate with riblets that become crowded behind the aperture. The basal and peripheral crests are well recognizable. The light brownish, wide ovoid peristome is broadly attached, with simple or only slightly swollen margin. The lamellae superior and spiralis long overlap. The lamella inferior is well emerged. The weakly-bent subcolumellaris is retracted, its end is barely visible in slanted view through the aperture. The lateral lunella is short and broad, with diffuse outline. It is fused, through the plica superior, to the plica principalis, and downward in an almost straight line also to the long basalis. In most cases the subclaustralis is short and weak, the reduced sulcalis is fused to the lunella complex. The anterior plica superior of variable strength starts diverging from, then becomes parallel to, the plica principalis. It is not connected to the lunella complex. The clausilium plate is not visible through the aperture.

(in mm). Holotype H_s: 16.7, W_s: 4.9, H_a: 4.4, W_a: 3.7; paratypes (NHMW 111235, n = 12): H_s: 16.1–20.7 (mean 18.0, S.D. 1.31), W_s: 4.9–6.0 (mean 5.3, S.D. 0.39), H_a: 4.4–5.5, W_a: 3.6–4.6.

Differs from nearest occurring M. l. laxa and M. laxa disjuncta Fehér & Szekeres, 2006 by its larger, stronger sculptured shell and weaker, shorter anterior plica superior, whereas from all other M. laxa subspecies by the combination of a large, tumid shell and long overlapping lamellae superior and spiralis.

Albania, Kukës District, S of Draj-Reç, entrance of the Vilë Gorge, 430 m, 41.8830°N, 20.3370°E.

Type locality, leg. ZF, JG, 2.vii.2005, holotype (NHMW 111237), paratypes (NHMW 111235/58+5a+4aj, HNHM 99645/63, GR/63); Draj-Reç (11 km S of the Përroi i Bushtricës mouth), 550 m, 41.8908°N, 20.3370°E, leg. TD, ZE, ZF, DM, 9.x.2005, paratypes (HNHM 99646/15, MMM-B01328/15, ER/15, SZ/4).

The new taxon is named after Tamás Deli, malacologist (MMM-B), who participated in several Balkan field trips, including the one that led to the discovery of this subspecies.

Western foothills of the Korab Mts in northeastern Albania. Known from two nearby sites in the valley of the Black Drin (Drin i Zi), around Draj-Reç. This is the northernmost subspecies of this polytypic species (Fig. 23).