Taxonomic revision of the rock-dwelling door snail genus Montenegrina Boettger, 1877 (Mollusca, Gastropoda, Clausiliidae)

Abstract The genus Montenegrina is revised on the basis of material available at the Hungarian Natural History Museum (Budapest), Naturhistorisches Museum Wien (Vienna), and the Naturmuseum Senckenberg (Frankfurt am Main), as well as newly discovered populations. The following new taxa are described: Montenegrina haringae sp. n., Montenegrina lillae sp. n., Montenegrina prokletiana sp. n., Montenegrina sturanyana sp. n., Montenegrina grammica erosszoltani ssp. n., Montenegrina grammica improvisa ssp. n., Montenegrina hiltrudae desaretica ssp. n., Montenegrina hiltrudae selcensis ssp. n., Montenegrina laxa delii ssp. n., Montenegrina nana barinai ssp. n., Montenegrina prokletiana kovacsorum ssp. n., Montenegrina rugilabris golikutensis ssp. n., Montenegrina rugilabris gregoi ssp. n., Montenegrina skipetarica danyii ssp. n., Montenegrina skipetarica gurelurensis ssp. n., Montenegrina skipetarica pifkoi ssp. n., Montenegrina skipetarica puskasi ssp. n., Montenegrina sporadica tropojana ssp. n., Montenegrina sturanyana gropana ssp. n., Montenegrina sturanyana ostrovicensis ssp. n., and Montenegrina tomorosi hunyadii ssp. n. A neotype is designated for Montenegrina helvola (Küster, 1860), and Montenegrina cattaroensis antivaricostata nom. n. was introduced to replace the junior homonym Clausilia umbilicata costata Boettger, 1907 (non Pfeiffer, 1928). Of each taxon types or specimens from the type localities are figured, and distribution maps are provided.


Introduction
Montenegrina is an obligate rock-dwelling door snail genus, associated with habitats of limestone outcrops. According to Nordsieck (2007), the genus belongs to the subfamily Alopiinae and, within that, the tribe Montenegrinini, together with Protoherilla Wagner, 1921. A family-wide molecular phylogenetic reconstruction confirmed its belonging to the Alopiinae, and found Alopia Adams & Adams, 1855 and Herilla Adams & Adams, 1855 the closest related genera (Uit de Weerd and Gittenberger 2013).
The geographic range of Montenegrina includes the coastal regions of Montenegro south of the Bay of Kotor, Albania, western Macedonia, and northwestern Greece (Fig. 1). In geographical terms it extends from the southernmost parts of the Dinaric Mountains to the northern part of the Pindos Mountains, and is separated from the eastern Balkans (Moesia) by the Vardar Basin and the Kosovo Plain. Biogeographically this area can be classified as part of the South Adriatic-Ionian province (Griffiths et al. 2004: 262).
The region is rich in limestone, which occurs in discrete karstic patches that are separated by intervening non-limestone (e.g. Quaternary fluvial deposits, flysch, volcanic and metamorphic) regions. For obligate limestone-dwelling gastropods with limited capacity of active dispersal these karst outcrops function as 'islands' that offer home to isolated populations (Gittenberger 2007). Nevertheless, in some rare events, local forms can be subject to jump-dispersal, allowing colonization at remote locations (Uit de Weerd et al. 2005). Allopatric distribution, in combination with the restricted dispersal, provides favourable conditions for non-adaptive speciation (e.g. via founder effect, genetic drift), and can give rise to radiation into multiple, morphologically diverse and geographically separated forms (Gittenberger 1991).
Unambiguous application of species and subspecies concepts under these circumstances can be quite challenging. In Balkan species of the rock-dwelling alopiinid door snails related morphotype groups often show higher divergence between than within their isolated populations and, when groups of populations show morphological diversity, their features tend to follow discernible geographical partitioning. In the taxonomical practice such species have been regarded polytypic, in the sense of Mayr's subspecies concept (Mallet 2007). In Montenegrina the number of traditionally recognized subspecies is high, representing about three-quarters of the described taxa in the genus (Nordsieck 2007). Occasionally the unambiguous delimitation of species can also be problematic. As different morphs only rarely co-occur or meet in a contact zone, reproductive and competitive relationships under natural conditions cannot be ascertained. Consequently, in some cases it can be difficult to judge objectively which forms can or cannot be regarded conspecific (Gittenberger 2007). During the 19th century the range of Montenegrina, except for the Montenegrin coast, belonged to the reclusive Ottoman Empire and, as a result, remained mostly inaccessible to zoological exploration. Therefore, the first taxa of the genus became known from the Adriatic coast of Montenegro, part of which then belonged to the Austrian Empire. The description of the first species, Clausilia cattaroensis Rossmässler, 1835, was followed by those of C. subcristata Pfeiffer, 1848, C. cattaroensis var. gracilior Küster, 1850, C. helvola Küster, 1860, C. laxa Küster, 1861 (the latter two likely from marine flotsam, see : Nordsieck 1972), C. umbilicata Boettger, 1879, and C. (Herilla) klecaki Westerlund, 1881. Exceptions were C. janinensis Mousson, 1859 and C. rugilabris Mousson, 1859 from the Turkish-occupied region, which Mousson (1859) described from material collected by the Schaefli Expedition to the Ioannina region.
After the turn of the century, Rudolf Sturany and Otto Wohlberedt began exploring the Ottoman-ruled northern Albanian area (Sturany 1905, Wohlberedt 1907. During the 1st World War, wide-scoped Austrian, Hungarian and German expeditions were organized to the northern territories of present-day Albania, eastern Montenegro, Kosovo and western Macedonia, which were then occupied by the armies of the Central Powers (Doflein 1921, Kümmerle 1926, Teleki and Csiki 1922-1940. The molluscs collected on these expeditions were studied by Wagner (1919), Soós (1924) and Hesse (1928). There were only few follow-up malacological studies (e.g. those of Anton Fuchs) between the two World Wars. Molluscs were typically collected as by-products by experts focusing on other groups (e.g. Bischoff, Dabović, Weigner and Winkler). Since the 1960s and 1970s the exploration of Greek and former Yugoslavian areas became intense due to the contributions of Wolfgang Fauer, Hartmut Nordsieck, László Pintér, Péter Subai and, somewhat later, Ivailo Dedov, Peter Reischütz, and Helmut Sattmann. By contrast, Albania was inaccessible for naturalists until the last decade of the 20th century, when finally a substantial set of malacological data from Albania could be published Welter-Schultes 1996, Welter-Schultes 1996).
As Albania and the neighbouring regions still remained the least explored parts of Europe, it became one of the focus areas in the research of the Hungarian Natural History Museum (HNHM). From the late 1990s the HNHM organized nearly a hundred zoological and botanical field expeditions to this area (Barina et al. 2014, Erőss and Fehér 2009, Murányi 2007, Murányi et al. 2011, which sampled at thousands of localities that have never been surveyed by naturalists before. These field trips resulted in significant accessions of the Mollusc Collection, providing several undescribed taxa, as well as new distribution records for little-known species of Montenegrina.
This in itself could have justified the need for a taxonomic revision of this genus. But there are also other reasons why Montenegrina became the focus of our interest. The size of its range and the number of known populations deemed large enough, but still accessible to almost comprehensive sampling. Furthermore, its patchy distribution makes this genus an attractive system for molecular phylogenetic, phylogeographic and evolutionary studies that can help better understanding the general mechanisms of speciation and spatial distribution of rock-dwelling gastropods. Such studies, however, require a thorough, morphology-based revision of the taxa.

Methods
In the text common geographic names and terms are given according to their current English usage. However, in the sections listing the available material these names and their spelling often follow those of the literature records or original labels. These, as well as historic or obsolete names, are given between quotation marks. To allow unambiguous identification of locality names geographical objects are often defined by locally used terms. These are as follows: In some of the countries different names are used for transboundary objects (e.g. Crni Drin/Drin i Zi/Black Drin, Vjosë/Aoós, Skadar /Shkodër/Skutari, Nemerçkë/ Dousko, etc.). In such cases these are mentioned in the form used in the country where the actual site belongs.
The studied samples are in the following collections: Private collections are referred to only when they include type material of the newly described taxa. Numbers of the type specimens are given separately for the dry and alcohol-preserved specimens. Among the latter 'a' and 'aj' denote adult and recognizable juvenile individuals, respectively.

HNC
The  Georeferenced distribution records used in this paper are deposited at the Global Biodiversity Information Facility (GBIF), http://ipt.pensoft.net/resource?r=montenegrina Dimensions and whorl numbers are given as shown in Fig. 2. For most of the taxa size data are taken from the literature, preferentially from the original descriptions (unless indicated otherwise). For the new taxa the measurements of the holotypes and 10-12 randomly selected paratypes are given. These were taken from front view images by ImageJ software (Schneider et al. 2012) using landmarks placed on the pictures. The dimensions of the shells are given by size ranges, as well as means and standard deviation values, whereas those of the apertures only by size ranges. In the diagnoses shell height is given simply as large (>20 mm), medium (15-20 mm), small (10-15 mm), or very small (<10 mm).

Ecology.
Montenegrina species inhabit rocky habitats, where they feed on the microflora and find hiding place in crevices or among and under boulders. All known populations are associated with limestone environments. The formation and shape of the habitable outcrops can vary considerably even within a species. Suitable habitats include small to large cliffs, gorges, rocky forests, rocky alpine grasslands, or even artificial stone walls along roads (Fig. 10).
Historical background. The first few described taxa of Montenegrina were initially classified within the genus Clausilia Draparnaud, 1805 and, later on, in the subgenus Clausilia (Delima) Hartmann, 1842. In his Clausilia monograph Schmidt (1868) already distinguished 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.  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.
Remarks. 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.

Fig. 11C
Montenegrina janinensis jakupicensis Fauer, 1993: 56-57 Distribution. Southern part of the Jakupica Mts in Macedonia. Known only from a few sites, within a narrow range northwest of Nežilovo village (Fig. 12). (Rossmässler, 1835) Diagnosis. 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, mediumbent subcolumellaris mostly visible. Lunella dorsal to dorsolateral, separate from the  Distribution. 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).
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.
Remarks. 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). Distribution. This taxon is distributed sporadically along the southern Montenegrin coast between Budva and Zaljevo (Fig. 13).

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

Montenegrina dofleini dofleini
Distribution. 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).
Remarks. 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 MIZ-PAS 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.

Montenegrina dofleini pinteri
Distribution. 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.
Remarks. See considerations on niche segregation at M. stankovici.
Distribution. 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).
Remarks. 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. Fig. 15N Montenegrina drimmeri Fehér & Szekeres, 2006in Erőss et al. 2006 fig. 1 Distribution. Southern part of the Lura-Dejë mountain group in northern Albania. Known only from the type locality (Fig. 16).

Montenegrina drimmeri Fehér & Szekeres, 2006
Remarks. Nordsieck (2009) placed M. drimmeri into the same species-group with M. fuchsi. Brandt, 1961 Diagnosis. 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. Brandt, 1961  Diagnosis. 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.
Type locality. "An der Straße von Tepelene nach Korzyra" = Albania, valley of the Vjosë River between Kelcyrë and Tepelenë.  Brandt, 1961, holotype, SMF 163939 C M. fuchsi klemmi Brandt, 1962 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).
Remarks. 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. Brandt, 1962    Distribution. 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).

Fig. 15E
Montenegrina fuchsi pallida Fauer, 1993: 54-55 Distribution. Southern part of the Dousko (Nemërçkë) Mountain, in Epirus, northwestern Greece. Known only from the type locality (Fig. 16). Nordsieck, 1988 Diagnosis. 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.
Description. 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 mediumbent 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.
Type locality. Albania Etymology.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.
Distribution. Southern part of the Lura-Dejë mountain group, within the catchment area of the Varosh Stream (Fig. 17).
Remarks. At one of the localities it was found syntopically with M. skipetarica puskasi ssp. n.
Description. 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.
Description. 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.
Type locality. 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.
Type material. Albania Distribution. 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).
Remarks. The first volume of the Systematisches Conchylien-Cabinet (Küster 1844(Küster -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.

Diagnosis.
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.
Dimensions ( Distribution. Mallakastër Hills in southern Albania, in the lower valley of the Vjosë River. Known localities are within 5 km from each other (Fig. 18).
Dimensions ( Distribution. 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 Welter-Schultes 1999, Nordsieck 2009) presumably also correspond to this taxon (Fig. 18). Diagnosis. 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.

Montenegrina hiltrudae hiltrudae
Distribution. Southern part of the Verno Mts in Western Macedonia (Greece). Known only from the type locality (Fig. 20). Distribution. 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).
Description. 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.
Etymology.The new taxon is named after Desaretia, the ancient Roman name of the Ohrid-Prespa region.
Distribution. Prespa Lake area. Apart from the type locality, it was also found in the vicinity of Liqenas (Fig. 20). 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. Distribution. 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). Gittenberger, 2002 Fig. 19G Montenegrina janinensis maasseni Gittenberger, 2002: 131-134 (Fig. 20).

Diagnosis.
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.
Dimensions ( Distribution. 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).

Montenegrina hiltrudae sattmanni
The paratype lot NHMW 84030/4 could not be found in the NHMW collection.
Montenegrina hiltrudae selcensis ssp. n. http: Diagnosis. Medium-size subspecies with large, very narrowly attached peristome and in front view well visible lamella subcolumellaris.
Description. 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.
Dimensions ( Distribution. 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).
Remarks. 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).

Montenegrina laxa laxa
Remarks. The illustration with the name laxa was published in 1860, but a description of the species only in 1861 (both in Küster 1844Küster -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 (1844Küster ( -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.  22°N, 20.54°E, leg. ID, Minkov, 10.vii.2009 (DED-580); near the Kokal Summit, 1800-1928m, 41.203°N, 20.532°E, leg. ID, 15.vii.2009; Tchumata area (up to the Krustets area), alpine limestone meadows, 1980 m, leg. ID, 12.vii.2009 (DED-582).

Montenegrina laxa dedovi
Distribution. This taxon is known from the alpine region of the Jablanica Mts (Fig. 23).
Remarks. 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.
Description. 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.
Dimensions ( 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. Type locality. Albania Etymology.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.
Distribution. 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).

Montenegrina laxa errans Erőss & Szekeres, 2006 Fig. 21E
Montenegrina laxa errans Erőss & Szekeres, 2006in Erőss et al. 2006 fig. 20 Distribution. This is the southernmost M. laxa subspecies, found sporadically around the Mokër Mts in southeastern Albania (Fig. 23). Distribution. Central Albania. For a long time this taxon has been known only from the type locality, but a recently discovered population more than 30 km farther south indicates that it may have a wider, sporadic distribution (Fig. 23).

Montenegrina laxa lakmosensis
Remarks. Originally M. l. lakmosensis was described by Nordsieck (2008) as a subspecies of dedovi, which is now also classified with M. laxa (see above).
Dimensions ( Distribution. Çermenikë Mountain in central Albania. Known from a few localities around this mountain within a circle of 10-15 km radius around Librazhd (Fig. 23).
Description. The large, elongate shell of 10½ to 12 whorls is light brown. The surface of the whorls is opaque, very finely wrikled, almost smooth. The weakly inflexed neck has strong whitish ribs that become dense toward the aperture. The basal crest is strong, the peripheral one is weak. The peristome is large, angular, broadly attached. It has wide, whitish, somewhat swollen margin, which is missing at the upper columellar side. The lamella superior is weak, it does not overlap with the spiralis. The terminal part of the lamella inferior is well emerged, descends seeply and ends low at the columellar side of the peristome. The weakly-bent lamella subcolumellaris is retracted, its end is not visible through the aperture. The plica superior often becomes fused to the principalis. The broad, lateral to ventrolateral lunella is connected to the long basalis. The subclaustralis is short, diffuse, the sulcalis is well developed. The anterior part of the plica superior is long, often separate from the lunella complex. The clausilium plate is not visible through the aperture.
Dimensions ( Etymology.This species is named after Lilla Tamás, the wife of the first author. Distribution. Lower part of the Drin Valley. Known from the Koman Dam and 12 km upstream of that (Fig. 22). Distribution. Lura-Dejë mountain group in northern Albania. Known only from two sites: the type locality at the southern foothills of the Mt. Lura in the gorge of the Varosh Stream, and the locality in the valley of the Urakë River, at the western foot of the Dejë. Although the elevation seems different, the record "Kurbnesh, 400 m" (Dhora and Welter-Schultes 1999) most probably refers to the latter occurrence (Fig. 22).

Montenegrina nana Fehér & Szekeres, 2006
Diagnosis. Shell small to medium, smooth to strongly wrinkled-costate. Neck weakly inflexed to inflexed, basal and peripheral crests well recognizable. In front view lamella inferior moderately emerged, medium-bent subcolumellaris not or only barely visible. Lunella dorsolateral to lateral, mostly fused to the basalis. Subclaustralis and sulcalis present. Anterior plica superior absent or weak. Clausilium plate not or barely visible through the aperture.
Remarks. M. nana, which was formerly classified as a subspecies of M. perstriata, differs from this species by its smaller size, oblique peristome, and deeper inflexed neck. One of the subspecies, M. n. barinai ssp. n., occurs syntopically and does not hybridize with M. perstriata ochridensis, indicating that they are distinct species.  Distribution. Gollobordë Plateau in northeastern Albania (Fig. 25A).
Description. The small shell of 9½ to 10½ whorls is light brownish-corneous. All whorls are strongly costate. The sharp whitish ribs become crowded behind the aperture. The neck is inflexed, the basal and peripheral crests are well recognizable. The peristome is ovoid, attached, its wide, light brown margin is slightly deflexed. The weak lamella superior ends far from the outer end of the spiralis. The weakly emerged, straight descending end of the lamella inferior is barely visible in front view. The deep-ending, medium-bent lamella subcolumellaris cannot be viewed through the aperture. The broad lunella is dorsolateral. It is fused, via the superior, to the plica principalis, and at its basal end to the short or residual basalis. The subclaustralis is weak, the sulcalis is present. The anterior part of the plica superior is missing. The parietal edge of the clausilium plate is often visible through the aperture. Dimensions (in mm). Holotype Hs: 13.2, Ws: 3.6, Ha: 3.5, Wa: 2.9; paratypes (HNHM 99008, n = 12): Hs: 13.0-15.8 (mean 13.8, S.D. 0.95), Ws: 3.4-4.6 (mean 3.9, S.D. 0.34), Ha: 3.1-3.9, Wa: 2.3-3.3. Differential diagnosis. Distinguishable from the two other subspecies of M. nana by its smaller, strongly costate shell, wide separate lamellae superior and spiralis, dorsolateral lunella, and the absence of the anterior plica superior.
Type locality. Albania Etymology.The new taxon is named after Zoltán Barina, botanist (HNHM), who collected invaluable zoological material during his field trips in Albania.
Distribution. Known only from its type locality at the southern slope of the Mt. Temlishi in northern Albania. (Fig. 25). Remarks

Distribution.
Mt. Temlishi in northern Albania. Known only from the type locality at the foot of this mountain (Fig. 25A).
Remarks. Erroneously this taxon was also mentioned from the vicinity of the Tre Çesmë Spring near Zerqan (Erőss et al. 2006), a locality of M. perstriata ochridensis (see also there).

Montenegrina okolensis Szekeres, 2006
Diagnosis. Shell small to medium, elongate, light to dark corneous. All whorls smooth. Neck inflexed, almost smooth, basal and peripheral crests weak. Peristome attached, ovoid to somewhat angular, with somewhat swollen margin. Lamella superior absent to residual and distant from the spiralis. In front view lamella inferior hidden to weakly emerged. Medium-bent subcolumellaris only obliquely visible. Lunella dorsal-dorsolateral to dorsolateral. Basalis and subclaustralis absent. Sulcalis present or only residual. Anterior plica superior mostly absent. Clausilium plate visible through the aperture. Distinguishable from M. apfelbecki by the attached peristome and reduced lamellae. Diagnosis. Shell small to medium, elongate. In front view lamella inferior weakly emerged. Lunella dorsolateral. Sulcalis present. Anterior plica superior mostly absent. Clausilium plate partly visible through the aperture.

Montenegrina okolensis okolensis Szekeres, 2006
Dimensions ( Distribution. In the central part of the Prokletije Mts in northern Albania. The description of this subspecies was based on a single shell without precise locality and collector information. Accordingly, type locality was defined only as "Maja e Jezerces region" (Erőss et al. 2006). Recently the subspecies has been re-discovered near the Valbona Pass (Qafa e Valbonës), south of the Jezercë Summit. Notably, along the Theth to Rragam footpath that crosses the main mountain ridge, M. o. okolensis could be found only around its highest point, near to and above the pass, between 1760 and 1850 m (see also : Fehér and Erőss 2009) (Fig. 26).
Remarks. Nordsieck (2009) considered okolensis a synonym of caesia Fehér & Szekeres, 2006. New okolensis samples confirm the stability of its morphological differences from caesia, namely the elongate shell, deeper lunella and better developed sulcalis. Therefore we maintain it as a distinct subspecies. Distribution. Central part of the Prokletije Mts in northern Albania. Known only from the type locality (Fig. 26).

Montenegrina okolensis caesia
Remarks. Originally M. o. okolensis and M. o. caesia were described as subspecies M. apfelbecki, with which they share similar plicae. However, their reduced clausiliar apparatus and isolated occurrence supports their classification within a separate species, which seems more closely related to other north Albanian representatives of the genus (e.g. M. prokletiana sp. n.).

Montenegrina perstriata (Wagner, 1919)
Diagnosis. Shell medium to large, light to dark corneous. Lower whorls in some subspecies flattened and nearly of the same width. Whorls smooth to indistinctly costate. Neck weakly inflexed, basal and peripheral crests well recognizable. Peristome attached, angular, with simple to strongly swollen margin. Lamellae superior and spiralis distant to overlapping. In front view lamella inferior moderately emerged, mediumbent subcolumellaris mostly not visible. Lunella dorsolateral to ventrolateral, mostly not connected to the basalis. Subclaustralis often short, sulcalis present. Plica superior frequently fused to the principalis. Anterior plica superior absent or weak. Clausilium plate not or only barely visible through the aperture.  Remarks. Wagner (1925) clarified that the unintentional assignement of this taxon to laxa in the original description was due to an editorial error during publication.

Remarks.
Shell sizes show unusually large variability between the different localities. They are smallest at the type locality, but at other sites can be as large as those of the closely related M. p. tenebrosa Nordsieck, 2009.

Montenegrina perstriata ochridensis
Distribution. Originally this taxon was known only from the eastern coastline of the Ohrid Lake, between Ohrid and Gradište (see : Brandt 1962;Loosjes 1966).
Recently it was also found in Central Albania, east of Bulqizë (Fig. 25). On the Mt. Temlishi it lives syntopically with M. nana barinai ssp. n.
Remarks. First ochridensis was regarded by Nordsieck (1972) a subspecies of M. perstriata but later, pointing out the differences in shell morphology, he classified it as a distinct species (Nordsieck 1988). More recently he again mentioned this taxon as M. perstriata ochridensis (Nordsieck 2009).
In the original description the subgenus name Delima is likely a typographic error (Wagner 1925: 70), because earlier and in the plate caption of the same paper he used Albanodelima (Wagner 1924(Wagner , 1925.
Type locality. Macedonia, gorge of the Radika near Debar, 1.5 km upstream of the bridge.
Remarks. In the description, Nordsieck (1972) mentions material from from three sites, without specifying which of these are designated as paratypes. Sample SMF 247211 is labelled as paratypes and  lists it accordingly. But SMNS-N 6933, the other part of the same lot, is without indication of type status.  3611°N, 20.3967°E, leg. ZF, 13.iv.2014 (HNHM 98975).

Montenegrina perstriata tenebrosa
Distribution. Gollobordë Plateau in northeastern Albania. The precise location of the type locality is unknown, but the material from the road to Sebisht is thought to be from its vicinity (Fig. 25).
Remarks. Nordsieck (2009) noted that M. p. tenebrosa is close to the 'crassa' mentioned by Fehér and Erőss (2009) from central Albania. Based on a large set of recently obtained samlpes we are of the opinion that the form mentined as crassa (Fehér and Erőss 2009) belongs to M. p. callistoma. We regard M. p. tenebrosa an extremely large member of this heterogeneous group, but maintain its status as a separate subspecies.

Montenegrina prokletiana sp. n.
http://zoobank.org/3223E4A9-37AB-4ECD-B4F2-FDA88CC135B9 Diagnosis. Shell very small to small, elongate, light corneous. Whorls flat, smooth to indistinctly wrinkled, lower whorls nearly of same width. Neck costate, its strong inflection extends to the lateral side. Basal crest weak, peripheral crest strong. Peristome attached, ovoid to somewhat angular, with more or less swollen margin. Lamellae superior and spiralis overlap. In front view lamella inferior moderately, medium-bent subcolumellaris strongly emerged. Lunella dosal-dorsolateral to dorsolateral, separate from the basalis. Subclaustralis of variable strength, sulcalis present. Anterior plica superior mostly absent. Clausilium plate not or only barely visible through the aperture. Differs fom all other species of the genus by its very prominent, horizontally ending lamella subcolumellaris.
Description. The light brown shell is very small and slender, its 8½ to 10½ whorls are separated by a deep suture. The lower three whorls are almost of the same width. Except the finely costate neck the entire shell surface is smooth. The strong inflection of the neck extends to the ventral side. The basal crest is weak, the peripheral one is strong. The light brown peristome is inflexed at the parietal side. The swollen and deflexed peristome margin is absent at the upper columellar side. The strong, projected lamella superior barely overlaps with the spiralis. The lamella inferior is weakly emerged, it turns horizontal before ending. In front view the medium-bent subcolumellaris is visible. Neither the inferior, nor the subcolumellaris reach the peristome margin. The dorsolateral lunella is not connected to the residual basalis. The subclaustralis is long and well developed, the sulcalis is present. The anterior plica superior is missing. The clausilium plate is not or only barely visible through the aperture.
Description. The small, very slender, light brown shell consists of 8½ to 10½ whorls, which are separated by a deep suture. The lower three whorls are almost of the same width. The surface is smooth over the entire shell except the neck, which is strongly wrinkled-costate from the dorsal side. The neck inflection is very strong and extends to the ventral side. The basal crest is weak, the peripheral one is strong. The light brown peristome is inflexed at the parietal side. The peristome margin is narrow, swollen and deflexed, but it is entirely absent over the upper columellar side. The strong lamella superior emerges from the plane of the peristome. Inward it does not overlap the spiralis. The lamella inferior is emerged, its end abruptly turns toward the aperture. Both the inferior and the medium-bent subcolumellaris terminate in thick, horizontal plicae at the peristome margin. The dorsal-dorsolateral to dorsolateral lunella is separate from the basalis. The subclaustralis is absent, the sulcalis is present. The anterior part of the plica superior is mostly missing, rarely a weak, separately standing residue of it is visible. The clausilium plate is not or only barely visible through the aperture.
Distribution. Lower Drin Valley in northern Albania. Known to occur upstream of the Koman Dam up to the mouth of the Valbonë (Fig. 26). Along this section of the river we found Montenegrina shells in fluvial flotsam, which likely belong to this subspecies, but their poor condition does not allow unambiguous identification.
Remarks. The specimens near the mouth of the Valbonë are usually smaller (H s : 9.0-12.0 mm) with fewer (8-9) whorls. Some flotsam specimens have strongly wrinkled surface, but otherwise very similar shell structure.

Montenegrina rugilabris (Mousson, 1859)
Diagnosis. Shell medium to large, yellowish-to purplish-brown. Lower whorls mostly smooth, upper ones smooth to indistinctly costate. Neck not or weakly inflexed. Basal crest well recognizable, peripheral crest weak. Peristome rounded to ovoid, with wide, swollen margin. Lamellae superior and spiralis mostly overlap. In front view lamella inferior variably emerged, broadly-bent subcolumellaris usually visible. Lunella dorsal to dorsal-dorsolateral, fused to or separate from the basalis. Lunella complex with the basalis can form a lambda-like structure. Subclaustralis always shorter than the basalis, sulcalis strong. Anterior plica superior fused to or separate from the lunella complex. Clausilium plate well visible through the aperture. Distinguishable from the similar M. skipetarica by the shorter subclaustralis and well developed sulcalis. Diagnosis. Shell medium, tumid, yellowish-brown. Lower whorls smooth, upper ones very finely, indistinctly costate. Neck not to weakly inflexed, finely and densely costate. Basal crest well recognizable, peripheral crest weak. Peristome attached, with strongly swollen, reflexed margin. Lamellae superior and spiralis overlap. In front view lamella inferior moderately emerged, broadly-bent subcolumellaris visible. Lunella dorsal, separate from the short basalis. Subclaustralis absent or residual. Anterior plica superior often connected to the lunella complex. Clausilium plate almost entirely visible through the aperture.
Diagnosis. Shell large, purplish-brown with strong whitish overlay. Lower whorls are opaque and densely striate, upper ones increasingly glossy with diffuse striae. Neck not or barely inflexed, basal and peripheral crests very weak. Peristome attached, with somewhat swollen margin. Lamellae superior and spiralis overlap. In front view lamella inferior well emerged, subcolumellaris mostly hidden. Lunella dorsal-dorsolateral, fused to the long basalis. Subclaustralis short. Anterior plica superior mostly separate from the lunella complex.
Dimensions ( PS, 17.vii.1990 (MNBE) Distribution. Southeastern part of the Mt. Mourgana in Epirus, northwestern Greece. In addition to the type locality a site is also known near Tsamantas (Fig. 29).
Description. The medium-size, dark to lighter purplish-brown shell consists of 9 to 10 whorls, which are separated by a whitish suture. The entire surface is smooth, except that of the weakly inflexed, very finely and densely costate neck. The basal and peripheral crests are weak. The light brown, ovoid to somewhat quadrangular, narrowly attached peristome has slightly swollen margin. The lamella superior initiates at the end of the spiralis, without overlap. The lamella inferior is moderately emerged, its end part descends straight. The broadly bent subcolumellaris is not visible in front view. The dorsal to dorsolateral lunella is short and wide, fused to the short subclaustralis but not to the much longer basalis. The sulcalis is strong. The anterior par of the plica superior is long, often connected to the lunella complex. The clausilium plate is partly visible through the aperture.  Reischütz & Sattmann, 1990, holotype, NHMW 84367 F M. rugilabris irmengardis Klemm, 1962 (Urbański, 1960), Sveti Naum, HNHM 90879 J M. sporadica sporadica Nordsieck, 1974 Etymology.The new taxon is named after the Mt. Goliku (Mali i Golikut), its type locality.
Distribution. Kendrevicë Mts in southern Albania. Known only from the type locality and its vicinity (Fig. 29).
Description. The medium-size, tumid, light corneous shell consists of 9½ to 11½ whorls. The surface of the lower whorls varies between almost smooth to indistinctly costate with widely spaced, wrinkle-like ribs. The apex is and the weakly inflexed neck are strongly costate. The basal and peripheral crests are weak. The wide-rimmed peristome is almost circular, its upper margin is narrowly attached. There is a long overlap between the lamellae superior and spiralis. The moderately emerged lamella inferior and the broadly-bent subcolumellaris are both well visible in front view. The dorsal lunella is fused to the basalis, which is much longer than the subclaustralis. The sulcalis is strong. The anterior part of the plica superior is not connected to the lunella complex. Part of the clausilium plate is visible through the aperture.
Differential diagnosis. Distinguishable from all other M. rugilabris subspecies by its costate apical whorls.  Etymology.The new taxon is named after Jozef Grego, malacologist, and also to his son Maroš Grego, who accompanied the first author on several Balkan field trips, including the one during which when this subspecies was discovered.
Distribution. Lakmos Mts in northwestern Greece. Known from two nearby localities in the western part of the Lakmos (Fig. 29).
Remarks. The specimens from the vicinity of the Kipina Monastery have weaker sculpture than those of the type locality. Klemm, 1962 Fig. 28F Montenegrina (Beieriella) irmengardis Klemm, 1962: 242-244 Diagnosis. Shell large, dark purplish-brown with whitish suture. Lower whorls smooth, upper ones smooth to finely wrinkled-costate. Neck not to slightly inflexed, finely and densely striate. Basal crest well recognizable, peripheral crest weak. Peristome attached, with somewhat swollen margin. Lamellae superior and spiralis overlap. In front view lamella inferior well emerged, subcolumellaris mostly visible. Lunella dorsal-dorsolateral, fused to the basalis. Subclaustralis short. Anterior plica superior mostly separate from the lunella complex.

Montenegrina rugilabris irmengardis
Dimensions ( Distribution. Arachtos Valley west of the Lakmos and Tzoumerka mountains in northwestern Greece. In addition to the type locality and its surroundings, this taxon is also known from another site ca. 17 km N of the type locality (Fig. 29).
Diagnosis. Shell medium to large, dark purplish-brown with whitish suture. All whorls smooth, lower ones separated by whitish suture. Neck not to slightly inflexed, densely striate. Basal crest well recognizable, peripheral crest weak. Peristome mostly detached, with somewhat swollen margin. Lamellae superior and spiralis overlap. In front view lamella inferior well emerged, subcolumellaris mostly visible. Lunella dorsal-dorsolateral, fused to the basalis. Subclaustralis short. Anterior plica superior separate from the lunella complex.
Dimensions ( Distribution. Epirus in northwestern Greece. Known only from the vicinity of the Pateron Monastery (Fig. 29).
Diagnosis. Shell medium to large, purplish-brown with whitish suture. All whorls smooth, lower ones separated by whitish suture. Neck barely inflexed, densely striate. Basal and peripheral crests weak. Peristome attached, with somewhat swollen margin. Lamellae superior and spiralis long overlap. In front view lamella inferior moderately emerged, subcolumellaris mostly hidden visible. Lunella dorsal-dorsolateral, fused to the basalis. Subclaustralis short. Anterior plica superior separate from the lunella complex.

Montenegrina skipetarica skipetarica
Distribution. Lura-Dejë mountain group and the Korab Mts in northern Albania. Most of the known occurrences are in the southern part of the Lura-Dejë mountain group. The type locality, however, is 30-40 km northeast of this area, and an isolated occurrence is in the northwestern part of the Korab Mts (Fig. 31B).
Remarks. Soós' paper (1924) describing M. p. skipetarica is a chapter of a larger study assessing the results of Csiki's zoological explorations in northern Albania. Whereas the complete volume was published only in 1940, the chapter written by Soós had already been printed and circulated in 1924. Therefore, in the case of skipetarica this earlier date is to be regarded as the year of description (Erőss et al. 2006).
Description. The small, slender, light brownish-corneous shell consists of 10 to 11½ whorls. The lower whorls are smooth, the upper ones are increasingly costate toward the tip. The weakly inflexed neck is densely costate. The basal crest is weak, the peripheral one is well visible. The ovoid to somewhat quadrangular peristome is broadly attached, its margin is somewhat swollen. The lamella superior is long, its frontal half is of even height. Inward it overlaps with the lamella spiralis. The moderately emerged lamella inferior ends with a straight descent. The broadly-bent lamella subcolumellaris is mostly visible in front view. The inner end of the plica principalis is often fused to the superior. The dorsolateral lunella is short and broad, it is mostly connected to the plica basalis. The basalis and subclaustralis are of the same length. The sulcalis is week. The anterior part of the plica superior is weak, separate from the lunella complex, occasionally absent. The clausilium plate is only barely visible through the aperture.
Dimensions ( Etymology.The new taxon is named after László Dányi, entomologist and pedozoologist (HNHM), who was the companion to the first author on several Balkan field trips, including the one during which this subspecies was collected.
Distribution. Western extensions of the Gramos Mts in southern Albania (Fig.  31A). Known from a few sites of the limestone mountain between Barmash and Gërmenj along the Leskovik to Ersekë main road (see also : Welter-Schultes 1996).
Description. The shell is medium-to large-sized, light brownish-corneous, somewhat tumid, of its 9½ to 12½ whorls the lower three are almost of the same width. The surface is strongly costate, with denser ribs at the weakly inflexed neck. The basal crest is weak, the peripheral one is well visible. The light brown, ovoid peristome has simple, detached (rarely very narrowly attached) margin. The lamellae superior and spiralis overlap. The lamella inferior is emerged, well visible in front view. The broadly-bent lamella subcolumellaris is not visible in frontal, only in slanted view at the aperture. The lunella is dorsolateral, broad, mostly fused to the basalis that is as long as the subclaustralis. The sulcalis is week. The anterior part of the plica superior is long, ends in a lump behind the aperture, mostly does not reach the lunella complex. Part of the clausilium plate is visible through the aperture.
Etymology.The new taxon is named after the village Gurë-Lurë, as the type locality is along the footpath leading there.
Distribution. Southern part of the Lura Mountain in northern Albania. This taxon is known only from the Setë Gorge, where it was found at a short section of the footpath between Cidhnë and Gurë-Lurë. Towards Cidhnë, at ca. 1.5 km from the hydroelectric station, there is a narrow contact zone between this taxon and M. skipetarica puskasi, and to the east of that zone only M. skipetarica puskasi could be found (Fig. 31B). Nordsieck, 1972 Fig. 30B Montenegrina irmengardis konitsae Nordsieck, 1972: 35, plate 4, fig. 35. -Zilch 1981 129, plate 15, fig. 41. Montenegrina skipetarica konitsae -Nordsieck 2009: 73. Diagnosis. Shell large, tumid, light reddish-brown with whitish suture. All whorls smooth. Neck weakly inflexed, finely and densely costate. Basal crest well recognizable, peripheral crest weak. Peristome attached, ovoid, with simple margin. Lamellae superior and spiralis overlap. In front view lamella inferior well emerged, subcolumellaris mostly visible. Lunella dorsolateral, fused to the short basalis. Subclaustralis is short, sulcalis weak. Anterior plica superior short and weak, mostly separate from the lunella complex.
Distribution. Tymfi Mts in Epirus, northwestern Greece. Known from a few localities around Konitsa, from the northern and western sides of the Tymfi Mts (catchment area of the Aoos River), and from the gorge of the Sarantaporos River, ca. 8 km northwest of Konitsa (Fig. 31A).
Description. The small, light brownish-corneous shell consists of 9 to 11½ whorls. The lower whorls are smooth, the upper ones are indistinctly wrinkled-costate. The weakly inflexed neck is strongly, irregularly wrinkled. The basal crest is weak, the peripheral one is well recognizable. The ovoid to somewhat quadrangular peristome is broadly attached, its narrow margin is somewhat swollen. The lamella superior initiates at the depth where the spiralis ends, without overlap. The moderately emerged terminal part of the lamella inferior descends in a straight line. In front view the broadly-bent lamella subcolumellaris is mostly well visible. The inner end of the plica principalis is often fused to the superior. The dorsolateral lunella is short and broad, it is connected to the similarly short plica basalis. The basalis and subclaustralis are of the same length. The sulcalis is week. The anterior part of the plica superior is separate from the lunella complex. The clausilium plate is partly visible through the aperture.
Etymology.The new taxon is named after Dániel Pifkó, botanist (HNHM), who collected invaluable zoological material during his field trips in Albania.
Distribution. Mt. Dejë in northern Albania. This taxon was found at the western slope along the climbing path from Macukull village to the Dejë Summit, between 1250 and 2000 m (Fig. 31B).
Distribution. Gramos Mts in northwestern Greece. Known from the Epano Arena area where it is common. Sympatric, and in some sites syntopic, with the much rarer M. grammica grammica (Fig. 31A).
Description. The shell is medium-to large-sized, light brownish-corneous, somewhat tumid, of its 9 1 / 2 to 11 whorls the lower three are almost almost of the same width. The surface is smooth at the lower whorls but becomes indistinctly costate over the short apex. The weakly inflexed neck has dense, sharp ribs behind the peristome. The basal crest is weak, the peripheral one is well visible. The light brown, ovoid peristome has simple, narrowly attached (rarely somewhat detached) margin. The lamellae superior and spiralis overlap. The lamella inferior is emerged, well visible in front view. The broadly bent lamella subcolumellaris is not visible in frontal, only in slanted view at the aperture. The lunella is dorsolateral, broad, mostly fused to the basalis that is as long as the subclaustralis. The sulcalis is week. The anterior part of the plica superior is long, ends in a lump behind the aperture and often also reaches the lunella complex. Part of the clausilium plate is visible through the aperture.
Dimensions ( Etymology.The new taxon is named after Gellért Puskás, entomologist (HNHM), who collected invaluable mollusc material during his Balkan field trips.
Distribution. Southern part of the Lura-Dejë mountain group in northern Albania. This taxon was found at some nearby localities along the Setë Stream (Fig. 31B).
Remarks. This taxon shows high inter-population variability of the shell sculpture, ranging from indistinctly wrinkled to sharply ribbed.
Diagnosis. Shell large, tumid, light reddish-brown with whitish tint. Whorls with strong, often whitish ribs, which become less distinct toward the apex. Neck weakly inflexed, densely costate behind the aperture. Basal and peripheral crests are recognizable. Peristome attached, ovoid to angular, with simple margin. Lamellae superior and spiralis mostly overlap. In front view lamella inferior moderately emerged, subcolumellaris not or barely visible. Lunella dorsal-dorsolateral, fused to the strong basalis. Subclaustralis also strong, sulcalis weak, anterior plica superior long, often connected to the lunella complex.
Dimensions ( Distribution. Southern part of the Tymfi Mts in Epirus, northwestern Greece, within the catchment area of the Voidomatis River (Fig. 31A).
Remarks. As its shell does not reveal a clear affiliation to any other taxa, M. soosi is tentatively regarded as a distinct species. In the Lapavë Gorge it occurs syntopically with the nominotypical form of M. skipetarica. Nordsieck, 1974 Diagnosis. Shell small, light corneous. Lower whorls smooth, upper ones smooth to indistinctly wrinkled-costate. Neck inflexed, striate. Basal and peripheral crests weak. Peristome attached, ovoid to somewhat angular, with slightly swollen margin. In front view lamella inferior moderately emerged, medium-bent subcolumellaris visible or hidden. Lunella dorsolateral to dorsolateral-lateral. Basalis absent or weak and not fused to the lunella, subclaustralis absent. Anterior plica superior mostly missing, if present separate from the lunella complex. The clausilium plate is partly visible through the aperture. Differs from M. janinensis by the stronger sculpture, weaker-bent lamella subcolumellaris, and the lack of anterior plica superior.
Description. The small, elongate, light brownish-corneous shell has 9½ to 11 whorls. The lower three whorls are nearly of the same width. The entire surface of the shell is smooth, except the finely striate-costate neck. The neck is weakly inflexed, the basal and peripheral crests are also wealky developed. The light brownish, simple peristome is ovoid to somewhat angular, broadly attached, its upper columellar margin is missing. The lamellae superior and spiralis do not overlap. The moderately emerged terminal part of the lamella inferior descends in a straight line. In front view the medium-bent lamella subcolumellaris is not visible. The plica principalis is often fused to the superior. The broad lunella is dorsolateral-lateral to lateral. The plicae basalis and subclaustralis are absent. The sulcalis is strong. The anterior part of the plica superior is missing or weak, separate from the lunella complex. The clausilium plate is barely visible through the aperture.
Dimensions ( Etymology.The new subspecies is named after the Tropojë Stream, in the gorge of which the type locality is located. Distribution. Mt. Shkëlzen in the eastern part of the Prokletije in northeastern Albania. Known only from the limestone gorge of the Tropojë Stream (Fig. 32).
Remarks. Inhabits limestone cliffs along the shore of the Ohrid Lake, always very close to the water surface. This is the only known Montenegrina taxon that prefers such type of habitat. Occasionally it occurs together with, or in the vicinity of, M. dofleini pinteri or M. perstriata ochridensis, but in a strict sense is not syntopic with these. Apparent spatial segregation can be observed on a fine scale: its congeners prefer cliff regions farther from the water surface. This kind of niche partitioning is unique within the genus.

Montenegrina sturanyana sp. n.
http://zoobank.org/28674C8F-6124-45D9-B2B4-4ECFE74CF88B Diagnosis. Shell small, glossy, light corneous. All whorls smooth. Neck weakly inflexed, striate to striate-costate. Basal crest weak to well developed, peripheral crest absent to recognizable. Peristome attached, ovoid to somewhat angular, its margin wide and swollen. In front view lamella inferior is weakly emerged, medium-bent subcolumellaris hidden or only barely visible. Lunella broad, dorsolateral to dorsolaterallateral. Plica superior often fused to the principalis and can reach farther forward than its contact with the lunella. Basalis absent to short and separate from the lunella. Sub-claustralis absent to residual, sulcalis of variable strength. Anterior plica superior absent or weak and not fused to the lunella complex. The clausilium plate is not or only barely visible through the aperture. Differs from M. tomorosi Brandt, 1961 by its slender stature, stronger basal crest, deeper lamella subcolumellaris and weaker subclaustralis, and from all other Montenegrina species by the combination of the dorsolateral to lateral lunella and strongly reduced basal plicae.
Description. The small, light corneous shell is comprised of 9½ to 10 whorls, of which the last three are of nearly the same width. The weakly inflexed neck is finely striate, otherwise the entire shell surface is smooth. The basal crest is weak, the peripheral one is not recognizable. The ovoid peristome is attached, its margin is wide and swollen. The lamellae superior and spiralis do not overlap. In front view the straight descending end of the lamella inferior is moderately emerged, the medium-bent subcolumellaris is not visible. The lunella is broad, dorsolateral-lateral. The plica superior occasionally reaches farther forward than its fusion with the lunella. Of the lower plicae the basalis is absent, the subclaustralis is residual, the sulcalis is well developed. The anterior plica superior is missing. The clausilium plate is not visible through the aperture.
Dimensions ( Etymology.The new species is dedicated to Rudolf Sturany (1867-1935, Austrian zoologist, former curator of the Mollusc Collection at NHMW, in appreciation of his pioneering contribution to the research of the Balkan fauna. Distribution. Western side of the Jablanica Mts in central eastern Albania. Known from two nearby localities (Fig. 33).
Description. The small, slender, light corneous shell consists of 9½ to 11½ whorls. The entire surface is smooth, except for the weakly inflexed neck that is finely striate-costate. The basal crest is well developed, the peripheral one is recognizable. The peristome is somewhat angular, attached, its margin is wide and swollen. The lamellae superior and spiralis often overlap. In front view the straight descending lamella inferior is moderately emerged. The retracted, medium-bent lamella subcolumellaris is only barely visible through the aperture. The broad, dorsolateral-lateral to lateral lunella is not fused to the short basalis. The plica superior can reach forward somewhat farther than its attachment to the lunella. The subclaustralis and sulcalis are residual. The anterior plica superior is absent, or weak, parallel to the principalis, separate from the lunella complex. The clausilium plate cannot be viewed through the aperture.
Dimensions ( Etymology.The new taxon is named after the Gropa Mts in Central Albania. Distribution. Gropa-Bizë-Martanesh mountain group in Central Albania. Known from two nearby sites around the village Bizë in Central Albania (Fig. 33).
Description. The small, light corneous shell is comprised of 8½ to 10 whorls, of which the last three are of comparable width. Except for the finely striate-costate neck the entire shell is smooth. The neck is weakly inflexed, the basal and peripheral crests are recognizable. The ovoid to somewhat angular peristome is attached, its margin is wide and swollen. The lamellae superior and spiralis often overlap. In front view the straight descending lamella inferior is weakly emerged. Through the aperture the medium-bent subcolumellaris can only be seen at an angle. The broad lunella is dorsolateral. The plica superior can reach forward farther than its fusion with the lunella. The basalis is absent, the subclaustralis is also absent or residual. The sulcalis is well developed. The anterior part of the plica superior is missing or weak, parallel to the principalis, separate from the lunella complex. The parietal edge of the clausilium plate is visible through the aperture.
Type locality. Albania

Remarks.
In the description the type locality of subcristata was only vaguely defined by Pfeiffer (1848). However, Küster's (1844Küster's ( -1862 description provides important clues, mentioning that the material originated from the border area of Montenegro (with that of the Habsburg Monarchy), a site somewhere along the road from Kotor (to Cetinje) at an altitude of 3000 feet (ca. 900 m). Based on these data the type locality of the nominotypical subspecies can be in the vicinity of Njeguši, E of the Bay of Kotor.
Plate 4 of Küster's monograph with figures of subcristata was published in 1847, whereas pages 39-40 with the description only in 1950. The plates contained no names, only accompanying sheets (wrappers) were issued along with the plates, so that subscribers and librarians would know which species were figured. These printed sheets were not meant to provide permanent scientific records. Due to the publication of Plate 4 and its wrapper, Pfeiffer could refer to subcristata in 1848, and attribute its authorship to Küster. However, ICZN Art. 8.1.1 specifies that for the availability of scientific names only the publication dates of text pages are relevant, and thus the wrappers are not acceptable as published work. Accordingly, Pfeiffer (1848) used the name first in proper form, therefore he should be considered the author (Welter-Schultes 1999: 165).
In 1899 Möllendorff gave description of a variety from Virpazar, but without a name. Later Wohlberedt (1901) mentioned the name sublabiata Möllendorff, without providing a description or indication according to ICZN Art. 12 (nomen nudum). Later Wohlberedt (1909) published the name sublabiata together with Möllendorff's (1899) description, but he also published these, in literal Serbian translation, two years earlier (Wohlberedt 1907). Therefore, 1907 should be regarded as the year of publication and Wohlberedt as the author. This little-known Wohlberedt paper in Cyrillic script was probably overlooked or misunderstood by , who referred to its sublabiata as nomen nudum.
In his 1907 and 1909 papers Wohlberedt clearly indicated that the descriptions of the new clausiliids are Boettger's contributions, therefore the authorships of interior and brunnea are Boettger's. Westerlund (1881) described Clausilia (H.) klecaki, with this spelling. Later (Westerlund 1884) he used this name as kleciaki, and subsequently all authors adopted, probably incorrectly, this spelling form. Based on the original description, kleciaki is identical with the typical subcristata. By contrast, Sturany and Wagner used the name kleciaki in a different way (Sturany 1905, Sturany and Wagner 1915, Wagner 1924, for the form that was described by Boettger as brunnea (see also : Nordsieck 1972).
Intraspecific variability. Nordsieck (2007Nordsieck ( , 2009) accepted only two valid subspecies, namely s. subcristata and s. wohlberedti. These differ primarily in the position of the lunella: subcristata having it ventrolaterally, whereas wohlberedti laterally. In this view, minor, interior and kleciaki are synonyms of the nominotypical form, whereas sublabiata and brunnea are synonyms of wohlberedti. The examined populations were highly variable in the size and shape of the shell, as well as in the position and shape of the lunella, allowing no clear delineation of two (or more) morphotypes. Moreover, considerable morphological heterogeneity could sometimes be observed within the same museum lots, or between lots from the same localities. Therefore, with the present knowledge, we find most appropriate classifying this diverse species without subspecific division.

Montenegrina tomorosi coerulescens
Description. The small, light corneous shell consists of 8½ to 9½ whorls. The surface is smooth, only at the neck is wrinkled-costate. The neck is inflexed, the basal and peripheral crests are well developed. The ovoid to somewhat angular peristome is attached, its margin is slightly swollen. The lamellae superior and spiralis usually do not overlap. In front view the lamella inferior is moderately emerged, the weakly-bent subcolumellaris is mostly not visible. The broad, dorsolateral lunella is fused, via the plica superior, to the plica principalis. The basalis is weak, not connected to the lunella. The subclaustralis is absent, the sulcalis is well developed. The anterior plica superior of variable strength is separate from the lunella complex. The clausilium plate is partly visible through the aperture.
Distribution. Gorge of the Portaikos River at the western edge of the Thessaly Basin. Known only from the type locality (Fig. 36).

Discussion
The widely accepted species and subspecies concepts of Ernst Mayr provide a good theoretical background for taxonomists (Haffer 1997, Mallet 2007, Winker 2010. In theory, the lack of hybridization between different morphs under natural conditions indicates that they represent distinct species. When a morphologically variable species comprises parapatric populations with geographically partitioned morphs, and there are abrupt transitions between well discernible morphotypes with only narrow hybrid zones, then the species is polytypic. Their subspecific morphs with the mentioned characteristics represent biologically meaningful units that are reasonable to distinguish as subspecies (Welter-Schultes 2010).
However, the practical application of species and subspecies categories in the classification of allopatrically distributed groups is hampered by considerable difficulties (Gittenberger 2007). In Montenegrina, there are only a few known cases when different morphs co-occur or meet in a contact zone (Table 1). In a rare exception, the syntopic occurrence of M. perstriata ochridensis and M. nana barinai ssp. n. showed that M. perstriata, in the sense of Nordsieck (2009), consists of more than one species.
In the absence of natural contact there is no way of directly verifying reproductive isolation and competitive interactions. In such cases there are only indirect ways of inferring taxonomic differences. Namely, by comparing the degree of differences between related sympatric species, between intergrading subspecies of widespread species, or between hybridizing populations of related species (Mayr and Ashlock 1991: 104-105, Coyne and Orr 2004: 45, Haffer 1997. This makes the expert decision, whether allopatric morphs represent distinct subspecies within a polytypic species or only morphologically divergent populations of a monotypic taxon, inevitably subjective. A further complication is that the actual classification of a group can also depend on the extent of its exploration. In poorly explored groups the complexity and variability range of the populations/morphs is only superficially known. Newly discovered populations often represent unknown morphotypes that are usually described under new names. New taxa are established as long as the new morphotypes seem clearly steep cliffs on rocky grassland On exposed rock surface and among stones at the foot of the cliffs The former one seems to be common and widespread in the whole summit region whereas the latter one was found to be within a narrow range rock wall by the footpath On exposed rock surface along a transect of the footpath between Cidhnë and Gurrë-Lurë, morphologically transitional specimens were found only within a narrow zone (<100 m) distinguishable and geographically well separated. However, more detailed studies can reveal continuous spatial gradients of several traits or entangling of the distribution areas, and thus can necessitate the merging of former taxa. At a certain exploration level a new type of classification may become necessary, one that involves substantially fewer taxa than the earlier approach. Such was the case, for instance, with Cretan Albinaria, where a novel classification concept resulted in massive synonymization of the formerly accepted subspecies (Welter-Schultes 2010). Though Welter-Schultes (2012: 217) suggests the extension of the same 'lumper' approach to other rock-dwelling clausiliid genera, in particular Alopia, Bulgarica Boettger, 1877 and Montenegrina, the current taxonomy of rock-dwelling Balkan alopiinids still follows he traditional 'splitter' way (Nordsieck 2007, Bank 2013. Apparently, there is no single model that could be fully suitable for the classification of all these genera. For instance, in contrast to the Cretan Albinaria, in the case of Charpentieria itala (Martens, 1824) genetic relationships strongly supported the suitability of the polytypic species concept (Scheel and Hausdorf 2012).
Whereas taxonomic decisions unavoidably contain arbitrary elements, the nature of intra-generic heterogeneity provides guidance as to which of the approaches is to be used. The lumper approach is well applicable for species that have numerous populations within a geographically uninterrupted range, and shows high inter-, as well as intra-population diversity. By contrast, the splitter approach (or "traditional way", as termed by Welter-Schultes 2010) is best suited for species that have spatially well separated, and by stable character state combinations unambiguously distinguishable populations, with high inter-but low intra-population variability.
Despite a substantial increase of occurrence data from Albania and the neighbouring regions, most of the known Montenegrina taxa still seem to show distinct, spatially structured distribution. That is, they occur sporadically, within narrow, geographically clustered ranges, which are well isolated from each other. These conditions justify their classification according to the polytypic species concept. However, M. subcristata, a species having a large number of populations within a geographically uninterrupted range, and showing high inter-, as well as intra-population diversity, seems to be best classified without subspecific division. Therefore, at least at the current level of knowledge, we find the traditional method to be the most suitable way of classification, in which all taxa are recognized that are reliably distinguishable from the others. Accordingly, the present classification of Montenegrina contains 29 species and 106 subspecies, making it the second most speciose alopiinid genus after Albinaria.