Monograph |
Corresponding author: Zoltán Fehér ( feher@nhmus.hu ) Academic editor: Thierry Backeljau
© 2016 Zoltán Fehér, Miklós Szekeres.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Fehér Z, Szekeres M (2016) Taxonomic revision of the rock-dwelling door snail genus Montenegrina Boettger, 1877 (Mollusca, Gastropoda, Clausiliidae). ZooKeys 599: 1-137. https://doi.org/10.3897/zookeys.599.8168
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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: M. haringae sp. n., M. lillae sp. n., M. prokletiana sp. n., M. sturanyana sp. n., M. grammica erosszoltani ssp. n., M. grammica improvisa ssp. n., M. hiltrudae desaretica ssp. n., M. hiltrudae selcensis ssp. n., M. laxa delii ssp. n., M. nana barinai ssp. n., M. prokletiana kovacsorum ssp. n., M. rugilabris golikutensis ssp. n., M. rugilabris gregoi ssp. n., M. skipetarica danyii ssp. n., M. skipetarica gurelurensis ssp. n., M. skipetarica pifkoi ssp. n., M. skipetarica puskasi ssp. n., M. sporadica tropojana ssp. n., M. sturanyana gropana ssp. n., M. sturanyana ostrovicensis ssp. n., and M. tomorosi hunyadii ssp. n. A neotype is designated for M. helvola (Küster, 1860), and M. 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.
Balkan, Albania, Greece, Montenegro, Macedonia, new taxa, distribution data, types, polytypic species, limestone habitat
Montenegrina is an obligate rock-dwelling door snail genus, associated with habitats of limestone outcrops. According to
The geographic range of Montenegrina includes the coastal regions of Montenegro south of the Bay of Kotor, Albania, western Macedonia, and northwestern Greece (Fig.
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 (
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 (
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:
After the turn of the century, Rudolf Sturany and Otto Wohlberedt began exploring the Ottoman-ruled northern Albanian area (
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 (
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.
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:
brana dam (Macedonian)
chuka summit (Macedonian)
çuka summit (Albanian)
farangi canyon (Greek)
gora mountain (Serbian)
izvor spring (Serbian)
lumi river (Albanian)
maja summit (Albanian)
mali mountain (Albanian)
pečina cave (Serbian)
përroi stream, creek (Albanian)
planina mountain (Serbian)
potok stream (Serbian)
qafa mountain pass (Albanian)
shpella cave (Albanian)
shkëmb cliff (Albanian)
ura bridge (Albanian)
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:
HNC
MIZPAS
MMM
Munkácsy Mihály
DED private collection of Ivailo Dedov, Sofia
ER private collection of Zoltán Erőss, Budapest
GR private collection of Jozef Grego, Banská Bystrica
HU private collection of András Hunyadi, Budapest
SZ private collection of Miklós Szekeres, Szeged
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.
The names of the frequently mentioned collectors are abbreviated as follows: AH – András Hunyadi, AR – Alexander Reischütz, AS – Anna Szigethy, CN – Csaba Németh, DA – Dorottya Angyal, DM – Dávid Murányi, DP – Dániel Pifkó, EH – Elisabeth Haring, EM – Edvárd Mizsei, GP – Gellért Puskás, HS – Helmut Sattmann, ID – Ivailo Dedov, JG – Jozef Grego, JK – Jenő Kontschán, KJ – Katharina Jaksch, KK – Kornél Kovács, LD – László Dányi, LP – László Pintér, LT – Lilla Tamás, MD – Michael Duda, NR – Nicole Reischütz, PJ – Péter Juhász, PR – Peter L. Reischütz, PS – Péter Subai, TD – Tamás Deli, TK – Tibor Kovács, TH – Tamás Huszár, TN – Tamás Németh, ZB – Zoltán Barina, ZE – Zoltán Erőss, ZF – Zoltán Fehér, ZU – Zsolt Ujvári.
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.
The terminology of distinctive shell structures is used according to the detailed descriptions provided by
Shell structures of the last whorl. lin: lamella inferior, lsc: lamella subcolumellaris, lsu: lamella superior, lun: lunella, ni: neck inflection, nr: neck ribs, pba: plica basalis, ppr: plica principalis, psa: anterior plica superior, psc: plica subclaustralis, psl: plica sulcalis, psu: plica superior.
Montenegrina
[Clausilia, Sect. X. Delima, gruppe der cattaroensis]
Clausilia (Heteroptycha) Westerlund, 1884: 5.
Montenegrina
[Clausilia (Delima) gruppe A.] –
Montenegrina
[Delima, group i.] –
Heteroptychia
(sic!) –
Delima (Albanodelima)
Wagner, 1924: 118. –
Montenegrina
[Alopia (Delima) a.] –
Heteroptycha
[Alopia (Delima) b.] –
Delima (Montenegrina) – Zilch in Wenz 1960: 429–430.
Delima (Heteroptycha) – Zilch in Wenz 1960: 429–430.
Montenegrina (Heteroptycha)
–
Montenegrina (Montenegrina)
s. str. –
Montenegrina (Beieriella) Klemm, 1962: 242.
Montenegrina
–
[Montenegrina]: cattaroensis—to our best knowledge,
Coastal regions of Montenegro south of the Bay of Kotor, Albania, western Macedonia, and northwestern Greece (Fig.
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.
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
Based on genital morphology,
Montenegrina was first regarded a distinct genus by
Assessments of the genital morphology of Montenegrina were provided by
The most recent revision of the genus was provided by
Clausilia
apfelbecki
Sturany, 1907: 233. –
Delima (Delima) apfelbecki
–
Delima (Albanodelima) apfelbecki
–
Montenegrina
apfelbecki
–
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). Hs: 12.6–16.0, Ws: 3.4–4.2 (syntypes
A Montenegrina apfelbecki (Sturany, 1907), syntype,
“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 (
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
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.
Delima (Delima) attemsi
Wagner, 1914 in
Delima (Albanodelima) attemsi
–
Montenegrina
janinensis
attemsi
–
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). Hs: 12.0–14.8, Ws: 3.0–3.5 (
“Treska bei Üsküb” = Macedonia, Matka Gorge near Skopje.
Type locality, leg. Apfelbeck, ex Oberwimmer [10525], syntypes (
Macedonia, gorge of the Treska near Skopje, leg. Penther, 11.v.1918 (
The subspecies is known only from the valley of the Treska River (Matka Gorge) in northwestern Macedonia (Fig.
Although the description was formally published in
Montenegrina
janinensis
jakupicensis
Fauer, 1993: 56–57, plate 1, fig. 8. –
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). Hs: 12.4–16.9 (holotype 14.2), Ws: 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 (
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. (
Southern part of the Jakupica Mts in Macedonia. Known only from a few sites, within a narrow range northwest of Nežilovo village (Fig.
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.
Clausilia
cattaroensis
Rossmässler, 1835: Heft 2, pp. 8–9, plate 7, fig. 100. – Küster 1844–1862: 40–41, plate 4, figs 14–17. –
Clausilia cattaroensis var. gracilior Küster, 1850 in Küster 1844–1862: 40.
Clausilia
cattaroënsis
(partim) –
Clausilia (Delima) cattaroensis
–
Clausilia (Delima) cattaroensis f. parvula Westerlund, 1884: 54.
Clausilia (Delima) catarvensis
(sic!) –
Clausilia (Delima) lævigata
Mhlf. –
Clausilia (Delima) lesinacensis
Parr. –
Clausilia (Delima) cattaroensis var. gracilior
–
Delima (Delima) cattaroensis
–
Delima (Albanodelima) cattaroensis
–
Delima (Montenegrina) cattaroensis – Zilch in Wenz 1960: 429–430, fig. 1526.
Montenegrina
cattaroensis
–
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). Hs: 16.8–26.5 (lectotype 26.0); Ws: 4.3–5.9 (lectotype 5.8) (Kotor
“Cattaro” = Montenegro, Kotor.
“Dalmatien, Cattaro”, ex Rossmässler, lectotype (
Montenegro, Kotor, above the town, leg. Kiss, LP, 9.vii.1985 (
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.
The monograph of
The distribution record from Herceg-Novi (“Castelnuovo”) by
non Clausilia costata Pfeiffer, 1828: 42, plate 7, figs 17–18.
Clausilia (Delima) umbilicata var. costata
Boettger, 1907 in
Delima (Albanodelima) umbilicata costata
–
Montenegrina
umbilicata
costata
–
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). Hs: 14.4–23.1, Ws: 4.5–6.0 (topotypes
“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 (
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 (
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.
Although the description of Clausila umbilicata costata was formally published in
Clausilia umbilicata Boettger, 1879: 102, plate 2, fig. 3.
Clausilia (Delima) umbilicata
–
Delima (Albanodelima) umbilicata
–
Montenegrina
umbilicata
–
Montenegrina
umbilicata
umbilicata
–
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). Hs: 18.5–25.3 (lectotype 19.4), Ws: 4.8–6.6 (lectotype 5.4) (lectotype
“Antivari” = Montenegro, Stari Bar.
Type locality, ex Boettger, ex Benoit, lectotype (
Montenegro, Stari Bar, fortress, 130 m, 42.0939°N, 19.1342°E, leg. LD, ZF, JK, DM, 14.x.2008 (
This taxon is distributed sporadically along the southern Montenegrin coast between Budva and Zaljevo (Fig.
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.
Montenegrina laxa chiasma Nordsieck, 1972: 30, plate 5, fig. 42.
Montenegrina
chiasma
–
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). Hs: 16.1–18.0 (holotype 18.0), Ws: 4.2–4.6 (holotype 4.5).
Albania, Tiranë District, N slope of Mt. Dajti, 1600 m.
Type locality, ex Edlauer, paratypes (
This taxon is known from the summit region of the Mt. Dajti (1613 m), east of Tiranë (Fig.
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.
Delima (Montenegrina) dofleini
Wagner, 1928 in
Montenegrina
dofleini
dofleini
–
Montenegrina (Montenegrina) kaiseri Brandt, 1961: 4–5, plate 1, fig. 3.
Montenegrina
dofleini
kaiseri
–
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). Hs: 16.6–21.1, Ws: 4.2–5.5 (
“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 (
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 (
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.
According to Wagner (see:
Although not synonymizing formally,
In the original description the type locality of kaiseri was erroneously defined as ‘W of Peštani’ (
The type series of kaiseri was collected by Kaiser, the curator of the
Montenegrina
janinensis
fagorum
Nordsieck, 1974: 152, plate 5, fig. 30. –
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). Hs: 11.8–14.4 (holotype 13.5), Ws: 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 (
Macedonia, near the type locality, Korita Spring, 40.9674°N, 20.8149°E, leg. ZF, EH, KJ, HS, 17.x.2014 (
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.
Montenegrina kastoriae Nordsieck, 1972: 34, plate 4, fig. 33.
Montenegrina
dofleini
kastoriae
–
Montenegrina
kastoriae
kastoriae
–
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). Hs: 14.4–20.4 (holotype 19.0), Ws: 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 (
Greece, Western Macedonia, Kastoria, N side of the peninsula, leg. H. Nordsieck, 24.viii.1971 (
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.
Montenegrina janinensis pinteri Nordsieck, 1974: 154, plate 6, fig. 33.
Montenegrina
dofleini
pinteri
–
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). Hs: 14.8–18.9 (holotype 16.9), Ws: 4.1–5.0 (holotype 4.4).
Macedonia, Trpejca, north of boat harbor.
Type locality, leg. LP, PS, AS, 16.vii.1972, holotype (
Macedonia, Ohrid District, N of Trpejca, shore of the Lake Ohrid, 700 m, 40.9656°N, 20.7858°E, leg. ZF, 9.viii.2014 (
This taxon is known only from the vicinity of Tpejca at the shore of the Lake Ohrid (Fig.
See considerations on niche segregation at M. stankovici.
Montenegrina dofleini prespaensis Nordsieck, 1988: 199–200, fig. 4.
Montenegrina
dofleini
ssp. –
Montenegrina
kastoriae
prespaensis
–
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). Hs: 13.9–19.6 (holotype 16.7), Ws: 3.6–4.5 (holotype 3.9).
Greece, Western Macedonia, Psarades, opposite to the village.
Type locality, leg. HS, 1986, holotype (
Greece, Psarades, N of the village, 860 m, 40.8315°N, 21.0288°E, leg. PS, 10.v.1995 (
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.
From the confluence of the Great and Small Prespa Lakes
Montenegrina dofleini sinosi Páll-Gergely, 2010: 150–152, figs 1–2. (genital anatomy, clausilium plate).
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). Hs: 13.9-16.3 (holotype 15.0), Ws: 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 (
Type locality, leg. ZE, ZF, JG, 28.vi.2013 (
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.
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.
Montenegrina
apfelbecki
wagneri
Szekeres, 2006 in
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). Ws: 13.2–14.9 (holotype 14.9), Ws: 3.4–4.3 (holotype 4.2).
Albania, Mt. Thatë.
“Mal i That”, leg. Fuchs, holotype (
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 (
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.
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 (
Montenegrina
drimmeri
Fehér & Szekeres, 2006 in
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). Hs: 11.1–15.4 (holotype 12.6), Ws: 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 (
Type locality, leg. ZF, DM, ZU, 18.v.2010 (
Southern part of the Lura-Dejë mountain group in northern Albania. Known only from the type locality (Fig.
A Montenegrina haringae sp. n., holotype,
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.
Montenegrina (Heteroptycha) fuchsi Brandt, 1961: 2–3, plate 1, fig. 1.
Montenegrina
fuchsi
fuchsi
–
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). Hs: 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 (
Albania, right bank of the Vjosë at the bridge along the Tepelenë to Kelcyrë road, leg. Fuchs, 1936 (
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.
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
Montenegrina (Heteroptycha) fuchsi klemmi Brandt, 1962: 143, plate 5, fig. 13.
Montenegrina
fuchsi
klemmi
–
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). Hs: 10.4–14.0, Ws: 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 (
Type locality, ex Klemm (
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.
Montenegrina
fuchsi
muranyii
Fehér & Szekeres, 2006 in
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). Hs: 8.9–11.9 (holotype 10.9), Ws: 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 (
Tomorr Mts in southern Albania. This taxon is known only from the type locality, which is a western extension of the Tomorr (Fig.
Montenegrina
fuchsi
pallida
Fauer, 1993: 54–55, plate 1, fig. 6. –
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). Hs: 10.7–15.5 (holotype 12.5), Ws: 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 (
Type locality, 40.0468°N, 20.5648°E, leg. ZE, ZF, JG, 26.vi.2013 (
Southern part of the Dousko (Nemërçkë) Mountain, in Epirus, northwestern Greece. Known only from the type locality (Fig.
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.
Montenegrina
janinensis
grammica
Nordsieck, 1988: 198–199, fig. 2. –
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). Hs: 9.8–10.3 (holotype 9.8), Ws: 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 (
Greece, Gramos Mts, W of Epano Arena summit, 2000 m, 40.3090°N, 20.8981°E, leg. ZE, ZF, JG, 27.vi.2013 (
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.
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 Hs: 8.9, Ws: 3.0, Ha: 2.4, Wa: 2.1; paratypes (
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 (
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.
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 Hs: 14.4, Ws: 3.8, Ha: 3.6, Wa: 2.4; paratypes (
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 (
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.
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 Hs: 14.8, Ws: 4.3, Ha: 3.6, Wa: 2.9; paratypes (
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 (
The new species is named after Elisabeth Haring, head of the Central Research Laboratories at
Mt. Renc in northwestern Albania (Fig.
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.
Clausilia
helvola
Küster, 1860 –
Clausilia (Heteroptycha) helvola
–
Delima (Alpidelima) helvola
–
Delima (Heteroptycha) helvola – Zilch in Wenz 1960: 429–430, fig. 1527.
Montenegrina
helvola
helvola
–
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). Hs: 12.5–19 (neotype: 18.5), Ws: 3.5–4.6 (neotype: 4.5) (
Albania, Krujë, Skanderbeg Fortress (due to the present neotype designation). In the original description the locality was given as “Dalmatien”. According to
Albania, Mali i Krujës, inside the Skanderbeg Castle, ex K. L. Pfeiffer, ex Fuchs, 1941, neotype (
Same as that of the neotype (
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.
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 (
Without specific type locality it was not possible to ascertain where Küster’s material originated from.
Montenegrina
helvola
carinata
Erőss & Szekeres, 1999 in
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). Hs: 15.4–21.2 (holotype 18.9), Ws: 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 (
Type locality, leg. ZF, JK, DM, 11.x.2004 (
Mallakastër Hills in southern Albania, in the lower valley of the Vjosë River. Known localities are within 5 km from each other (Fig.
Montenegrina
helvola
magna
Fehér & Szekeres, 2006 in
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). Hs: 14.5–20.6 (holotype 18.7), Ws: 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 (
Type locality, leg. ZF, TN, EM, 15.iv.2014 (
Gropa-Bizë-Martanesh mountain group in central Albania. The most distant sites are ca. 25 km from each other (Fig.
Montenegrina
helvola
ornata
Erőss & Szekeres, 1999 in
Montenegrina
n. sp. –
Montenegrina
helvola
carinata
–
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). Hs: 13.2–16.9 (holotype 15.7), Ws: 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 (
Type locality, leg. ZF, TK, DM, 22.vi.2012 (
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
Montenegrina (Heteroptycha) pageti Brandt, 1962: 143–144, plate 5, fig. 14.
Montenegrina
helvola
pageti
–
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). Hs: 13.2–16.9, Ws: 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 (
Berat, SW side of the castle hill, 70 m, 40.7041°N, 19.9479°E, leg. DA, ZE, ZF, JG, 27.vi.2014 (
Known only from the castle hill of Berat (type locality), which is one of the westernmost foothills of the Tomorr Mts (Fig.
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.
Montenegrina hiltrudae Nordsieck, 1972: 33–34, plate 4, fig. 32.
Montenegrina
hiltrudae
hiltrudae
–
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). Hs: 13.9–17.4 (holotype 15.9), Ws: 4.0–5.0 (holotype 4.3).
A Montenegrina hiltrudae hiltrudae Nordsieck, 1972, holotype,
Greece, Western Macedonia, near Vogatsikon, on the road to Germas (= Yermas).
Type locality, leg. H. Nordsieck, 23.viii.1971, holotype (
Southern part of the Verno Mts in Western Macedonia (Greece). Known only from the type locality (Fig.
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).
Montenegrina
sattmanni
costulata
Erőss & Szekeres, 2006 in
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). Hs: 14.1–20.4 (holotype 16.0), Ws: 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 (
Type locality, leg. ZE, ZF, JG, 29.vi.2013 (
Southern part of the Mt. Thatë, in the vicinity of the Zvezdë Pass that connents the Prespa and Korçë Basins (Fig.
Montenegrina
dennisi
dennisi
Gittenberger, 2002: 134, figs 7–8. –
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). Hs: 15.7–21.2 (holotype 18.7), Ws: 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 (
Greece, 0.5 km N of Spileo, toward Zakas, 930 m, 40.0080°N, 21.2847°E, leg. ZE, ZF, JG, 22.vi.2013 (
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.
Originally M. h. dennisi was described as a separate species (
Montenegrina
hiltrudae
densicostulata
Nordsieck, 1974: 155, plate 6, fig. 37. –
Montenegrina
hiltrudae
(partim) –
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). Hs: 15.8–22.4 (holotype 21.9), Ws: 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 (
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 (
Southern part of the Verno Mountain in Western Macedonia (Greece). Found at some sites around Vogatsiko, southeast of the Kastoria Lake (Fig.
Originally
Montenegrina
sattmanni
sattmanni
–
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 Hs: 17.6, Ws: 5.5, Ha: 4.5, Wa: 4.1; paratypes Hs: 15.0–18.9 (mean 16.9, S.D. 1.33), Ws: 4.4–5.9 (mean 5.3, S.D. 0.39), Ha: 3.8–5.2, Wa: 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 (
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 (
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.
Formerly the Liqenas population was mentioned by
Montenegrina
sattmanni
fusca
Fehér & Szekeres, 2006 in
Montenegrina
skipetarica
fusca
–
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). Hs: 18.3–22.4 (holotype 22.4), Ws: 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 (
Type locality, leg. ZE, ZF, JG, 29.vi.2013 (
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.
Montenegrina
janinensis
maasseni
Gittenberger, 2002: 131–134, figs 5–6. –
Montenegrina
janinensis
maasii
(sic! typographic error) –
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). Hs: 14.5–18.6 (holotype 16.0), Ws: 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 (
Greece, Portitsa Farangi, near Spileo, 39.9965°N, 21.2855°E, leg. ZE, ZF, JG, 22.vi.2013 (
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.
Montenegrina h. maasseni was described by
Montenegrina
dennisi
protruda
Gittenberger, 2002: 135, figs 9–10, 16 (clausilium plate), 17–19 (clausilium plate microarmature). –
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). Hs: 16.0–20.6 (holotype 19.0), Ws: 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 (
Type locality, 990 m, 40.0741°N, 21.2017°E, leg. ZE, ZF, JG, 22.vi.2013 (
Western Macedonia (Greece). Known only from the type locality (Fig.
Montenegrina skipetarica robusta Nordsieck, 2009: 77–78, plate 2, fig. 8.
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). Hs: 18.0–25.1 (holotype 21.4), Ws: 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 (
Same as types (
Western part of the Verno Mts in Western Macedonia (Greece). Known only from the type locality, located between Kastoria and the Prespa Lakes (Fig.
Some specimens of the original series (e.g. those of
Montenegrina sattmanni Nordsieck, 1988: 200–201, fig. 5.
Montenegrina
sattmanni
sattmanni
–
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). Hs: 17.3–21.4 (holotype 20.1), Ws: 4.6–5.5 (holotype 5.3).
Greece, Western Macedonia, Mikrolimni, near the Biological Station.
Type locality, leg. HS, 24.v.1986, holotype (
Type locality, 40.7429°N, 21.1102°E, leg. ZE, ZF, JG, 28.vi.2013 (
The area of the Prespa Lakes (Fig.
Originally M. h. sattmanni was described as a distinct species (
The paratype lot
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 Hs: 19.9, Ws: 6.2, Ha: 5.5, Wa: 4.4; paratypes Hs: 16.5–21.4 (mean 19.4, S.D. 1.40), Ws: 5.1–6.2 (mean 5.7, S.D. 0.39), Ha: 4.6–5.8, Wa: 4.0–4.9 (
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 (
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.
Clausilia janinensis Mousson, 1859: 276.
Clausilia (Delima) janinensis
–
Delima (Albanodelima) janinensis
–
Montenegrina
janinensis
janinensis
–
Montenegrina
janinensis
crassilabris
Fauer, 1993: 55–56, plate 1, fig. 7. –
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). Hs: 12.2–17.1, Ws: 3.2–4.2 (Perama,
A Montenegrina laxa laxa (Küster, 1861), Tërkuza Valley
“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] (
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 (
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.
Although not synonymized formally,
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.
Clausilia
laxa
Küster, 1861 in Küster 1844–1862: 276, plate 31, figs 14–16. –
Delima (Albanodelima) weigneri
Poliński, 1924: 143–145, plate 4, figs 7–8. –
Clausilia (Delima) laxa
–
Montenegrina
laxa
laxa
–
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). Hs: 16.3–21.8, Ws: 4.6–5.9 mm (
“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 (
Dajti-Kruja mountain range in central Albania. Known from the gorges of the Tiranë and Terkuzë Rivers, north of Tiranë (Fig.
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 (
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 (
Montenegrina
dedovi
dedovi
Nordsieck, 2009: 75–77, plate 2, fig. 4. –
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). Hs: 13.6–20.3 (holotype 16.1), Ws: 3.7–4.4 (holotype 4.15) (
Macedonia, Jablanica Mts, Vartop region, after the Čuma Summit, 1868 m.
Type locality, leg. ID, 15.vii.2006, holotype (
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.
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 Hs: 16.7, Ws: 4.9, Ha: 4.4, Wa: 3.7; paratypes (
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 (
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.
Montenegrina
laxa
disjuncta
Fehér & Szekeres, 2006 in
Shell medium, light corneous. All whorls smooth. Neck inflexed, finely striate-costate. Basal and peripheral crests well recognizable. Peristome angular, with broad, somewhat swollen margin. Lamellae superior and spiralis do not overlap. Subcolumellaris barely, clausilium plate not visible through the aperture. Lunella lateral to ventrolateral, basalis strong. Subclaustralis short, sulcalis residual or absent. Anterior plica superior long, starts diverging from, then turns parallel to the plica principalis. It is occasionally connected to the lunella complex.
(in mm). Hs: 16.1–21.3 (holotype 17.7), Ws: 3.6–4.5 (holotype 4.1).
Albania, Dibrë District, 4 km E of Selishtë, along the Peshkopi to Burrel road, 13 km W of the bridge of Fushë-Muhurr, 760 m, 41.6308°N, 20.2983°E.
Type locality, leg. ZE, ZF, JK, DM, 26.vi.2003, holotype (
Type locality, leg. ZF, TN, EM, 14.iv.2014 (
This taxon is known from the vicinity of Selishtë (west of Peshkopi in Central Albania) (Fig.
Montenegrina
laxa
errans
Erőss & Szekeres, 2006 in
Shell slender, medium to large, yellowish-corneous. Lower whorls smooth, upper ones finely striate. Neck inflexed, striate. Basal and peripheral crests weak. Peristome angular, with somewhat swollen margin. Lamellae superior and spiralis do not overlap. Subcolumellaris and clausilium plate not visible through the aperture. Lunella lateral to ventrolateral, basalis strong. Subclaustralis short, sulcalis residual or absent. Anterior plica superior long, starts diverging from, then turns parallel to the plica principalis. It is not connected to the lunella complex.
(in mm). Hs: 16.0–19.9 (holotype 19.9), Ws: 3.6–4.3 (holotype 4.0).
Albania, Pogradec District, Çervenakë, 6 km from the Lin to Pogradec road toward the TV transmission tower, 1150 m, 40.9444°N, 20.6109°E.
Type locality, leg. ZE, ZF, JK, DM, 2.vii.2003, holotype (
Type locality, leg. ZE, ZF, JG, 29.vi.2013 (
This is the southernmost M. laxa subspecies, found sporadically around the Mokër Mts in southeastern Albania (Fig.
Montenegrina
laxa
iba
Nordsieck, 1972: 30, plate 5, fig. 41. –
Shell medium, yellowish-corneous. Lower whorls smooth, upper ones finely striate. Neck weakly inflexed, densely costate. Basal and peripheral crests well recognizable. Peristome ovoid to somewhat angular, with simple margin. Lamellae superior and spiralis mostly do not overlap. Subcolumellaris and clausilium plate barely visible through the aperture. Lunella dorsolateral to lateral, basalis short. Subclaustralis shorter than the basalis, sulcalis residual or absent. Anterior plica superior weak, not connected to the lunella complex, rarely absent.
(in mm). Hs: 12.3–17.8 (holotype 16.2), Ws: 3.4–4.2 (holotype 3.7).
“Arzen-Durchbruch bei Ibë” = Albania, Tiranë District, gorge of the Erzen near Ibë.
Type locality (“beim Krabapaß”), leg. Fuchs, holotype (
Original series but not type (
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.
Montenegrina
laxa
kontschani
Erőss & Szekeres, 2006 in
Shell medium to large, yellowish-corneous. All whorls smooth. Neck inflexed, striate. Basal and peripheral crests well recognizable. Peristome rounded to somewhat angular, with swollen, reflexed margin. Lamellae superior and spiralis overlap. Subcolumellar lamella not or barely, clausilium plate not at all visible through the aperture. Lunella lateral. Subclaustralis shorter than the basalis, sulcalis present. Anterior plica superior very long, starts diverging from, then turns parallel or even converging to, the plica principalis. It is mostly separate from the lunella complex.
(in mm). Hs: 14.8–19.3 (holotype 19.0), Ws: 3.7–4.8 mm (holotype 4.1).
Albania, Tiranë District, 6 km S of the Qafa e Shtyllës, 7.7 km S of the Elbasan junction along the Tiranë to Klos road, 1420 m, 41.3512°N, 20.0546°E.
Type locality, leg. ZE, ZF, JK, DM, 22.x.2002, holotype (
Albania, Tiranë District, Gropa Mts, ca. 3 km E of the Shën Mëri junction along the Tiranë to Bizë road, 1400 m, 41.3512°N, 20.0502°E, leg. ZF, TK, DM, 20.vi.2012 (
Southwestern part of the Gropa-Bizë-Martanesh mountain group in Central Albania (Fig.
Montenegrina
janinensis
–
Montenegrina dedovi lakmosensis Nordsieck, 2009: 77, plate 2, fig. 5.
Shell small, tumid, yellowish-corneous. All whorls smooth. Neck weakly inflexed, densely striate. Basal and peripheral crests weak. Peristome ovoid to somewhat angular, with slightly swollen margin. Lamella superior weak and short, distant from the spiralis. In front view lamella inferior moderately emerged. Subcolumellaris and clausilium plate not or barely visible through the aperture. Lunella dorsolateral to lateral, basalis short. Subclaustralis shorter than the basalis, sulcalis residual or absent. Anterior plica superior mostly absent, if present separate from the lunella complex.
(in mm). Hs: 12.8–15.0 (holotype 14.1), Ws: 3.8–4.1 (holotype 4.0).
Greece, Lakmos Mts, Epirus–Thessalia border, near Metsovon, S of the Peristeri Summit, ca. 2000 m.
Type locality, leg. HS, 2.vii.1988, holotype (
Original series but not type (
Lakmos (= Peristeri) Mts in northwestern Greece. Known only from the type locality (Fig.
Originally M. l. lakmosensis was described by Nordsieck (2008) as a subspecies of dedovi, which is now also classified with M. laxa (see above).
Montenegrina
laxa
miraka
Nordsieck, 1996: 8–9, plate 2, fig. 2. –
Shell medium to large, yellowish-corneous. All whorls smooth. Neck inflexed, densely striate-costate. Basal and peripheral crests well recognizable. Peristome ovoid to somewhat angular, with simple to weakly swollen margin. Lamellae superior and spiralis long overlap. In front view lamella inferior mostly well emerged. Subcolumellaris and clausilium plate not visible through the aperture. Lunella ventrolateral-lateral to ventrolateral, basalis long. Subclaustralis shorter than the basalis, sulcalis residual or absent. Anterior plica superior very long, starts diverging from, then turns parallel or even converging to the principalis. It is not connected to the lunella complex.
(in mm). Hs: 16.6–22.5 (holotype 22.1), Ws: 4.0–5.4 (holotype 5.1).
Albania, Librazhd District, Mirakë, 5 km W of Librazhd = 1 km NW Ura e Kamarës, S slope, 300 m.
Type locality, leg. Welter-Schultes, 17.ix.1995, holotype (HNC 40580), paratypes (HNC 40577,
Albania, Mirakë, Ura e Kamarës, 190 m, 41.1634°N, 20.2301°E, leg. ZF, TK, DM, 22.vi.2012 (
Çermenikë Mountain in central Albania. Known from a few localities around this mountain within a circle of 10–15 km radius around Librazhd (Fig.
Large, elongate species with wide, broade-lipped aperture, lateral to ventrolateral lunella and long, fused basalis.
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.
(in mm). Holotype Hs: 23.0, Ws: 5.0, Ha: 5.6, Wa: 4.7; paratypes (
Differs from all other species of the genus by the combination of a broadly attached peristome with dissolved upper margin, weak lamella superior, and deep subcolumellaris. From M. subcristata, the only other species of the genus with lateral to ventrolateral lunella, it is distinguished by the basalis that is fused to the lunella complex.
Albania, Shkodër District, Koman, ferry harbor near the dam, 180 m, 42.1087°N, 19.8264°E.
Type locality, leg. ZE, ZF, AH, DM, 14.iv.2006, holotype (
Albania, Koman Lake, right bank, ca. 12 km upstream of the Koman Dam, 170 m, 42.1872°N, 19.8696°E, leg. ZE, ZF, AH, DM, 15.iv.2006 (
This species is named after Lilla Tamás, the wife of the first author.
Lower part of the Drin Valley. Known from the Koman Dam and 12 km upstream of that (Fig.
Montenegrina
janinensis
–
Montenegrina
minuscula
Erőss & Szekeres, 2006 in
Shell very small, light corneous. All whorls smooth. Neck finely striate, deep inflexed, with strong basal and peripheral crests. Peristome detached, projected, pear-shaped, with simple margin. Lamellae superior and spiralis overlap. In front view lamella inferior hidden or barely emerged. Subcolumellaris and clausilium plate barely visible through the aperture. Lunella dorsolateral-lateral, separate from the short to rudimentary basalis. Subclaustralis absent, sulcalis residual or absent. Anterior plica superior only exceptionally present, either fused to or separate from the lunella complex.
(in mm). Hs: 9.1–11.5 (holotype 9.9), Ws: 2.3–2.9 (holotype 2.7).
Albania. Mat District, 3 km W of the Qafa e Murrës, gorge of the Lumi i Varoshit near the Shkëmb i Skanderbeut, 970 m, 41.6465°N, 20.1898°E.
Type locality, leg. ZE, ZF, JK, DM, 26.vi.2003, holotype (
Type locality, leg. TD, ZE, ZF, DM, 11.x.2005 (
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” (
In the gorge of the Varosh Stream this species occurs syntopically with M. s. skipetarica.
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.
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.
Montenegrina
perstriata
nana
Fehér & Szekeres, 2006 in
Shell small to medium, light corneous. All whorls smooth. Neck weakly inflexed, smooth to very finely striate. Basal and peripheral crests well recognizable. Lamellae superior and spiralis mostly overlap. Lamella subcolumellaris not or only barely visible through the aperture. Lunella lateral, fused to the short basalis. Subclaustralis residual or absent, sulcalis present. Anterior plica superior absent or weak, parallel to the principalis, separate from the lunella complex. Clausilium plate not visible through the aperture.
(in mm). Hs: 14.3–16.1 (holotype 15.1), Ws: 3.3–3.7 (holotype 3.7).
A. Montenegrina nana nana Fehér & Szekeres, 2006, holotype,
Albania, Librazhd District, 3 km NE of Lunik, on the Librazhd–Peshkopi road, 1050 m, 41.2966°N, 20.3741°E.
Type locality, leg. ZE, ZF, JK, DM, 24.ix.2002, holotype (
Type locality, leg. ZF, TN, EM, 13 Apr. 2014 (
Gollobordë Plateau in northeastern Albania (Fig.
Distribution of Montenegrina nana and M. perstriata. A M. nana gracilis and M. nana barinai ssp. n. (circle); M. nana nana (diamond); M. perstriata diminuta (square); M. perstriata mavrovoensis (empty circle with dot); M. perstriata occidentalis (empty triangle); M. perstriata perstriata (empty inverted triangle); M. perstriata plenostoma (inverted triangle); M. perstriata subcristatula (triangle) B M. perstriata callistoma (empty triangle); M. perstriata drimica (circle); M. perstriata ochridensis (square); M. perstriata radikae (empty circle); M. perstriata tenebrosa (star); M. perstriata steffeki is not shown.
Small subspecies with strongly costate shell, widely separate lamellae inferior and spiralis, missing anterior plica superior.
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.
(in mm). Holotype Hs: 13.2, Ws: 3.6, Ha: 3.5, Wa: 2.9; paratypes (
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.
Albania, Bulqizë District, Valikardhë, S slope of the Maja e Temlishit, 770 m, 41.514°N, 20.316°E.
Type locality, leg. ZF, TN, EM, 15.iv.2014, holotype (
Type locality, 880 m, 41.5138°N, 20.3142°E, leg. ZB, DP, 29.v.2008, (
The new taxon is named after Zoltán Barina, botanist (
Known only from its type locality at the southern slope of the Mt. Temlishi in northern Albania. (Fig.
The type locality of the new subspecies is sepatated only by a few hundred meters from that of M. nana gracilis Erőss & Szekeres, 2006. M. n. barinai ssp. n. occurs syntopically with M. perstriata ochridensis.
Montenegrina
perstriata
gracilis
Erőss & Szekeres, 2006 in
Shell small to medium, light to dark corneous. Lower whorls smooth, upper ones indistinctly costate. Inflexed neck with sharp, irregular ribs. Basal crest strong, peripheral crest well recognizable. Lamellae superior and spiralis mostly do not overlap. Lamella subcolumellaris not visible through the aperture. Lunella dorsolateral-lateral to lateral, mostly fused to the short basalis. Subclaustralis and sulcalis well developed. Anterior plica superior absent or weak, parallel to the principalis, separate from the lunella complex. Clausilium plate not visible through the aperture.
(in mm). Hs: 14.2–20.1 mm (holotype 16.5), Ws: 3.5– 4.7 mm (holotype 3.9).
Albania, Bulqizë District, 10 km E of Bulqizë, bank of Lumi i Zalli i Qytetit, 620 m, 41.5096°N, 20.3149°E.
Type locality, leg. ZE, ZF, JK, DM, 25.x.2002, holotype (
Type locality, leg. LD, ZE, ZF, AH, DM, 30.vi.2007 (
Mt. Temlishi in northern Albania. Known only from the type locality at the foot of this mountain (Fig.
Erroneously this taxon was also mentioned from the vicinity of the Tre Çesmë Spring near Zerqan (
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.
Montenegrina
apfelbecki
okolensis
Szekeres, 2006 in
Montenegrina
janinensis
caesia
(partim) –
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.
(in mm). Hs: 13.9–15.6 (holotype 13.9), Ws: 3.4–4.4 mm (holotype 3.4) (see type and other material).
“Theth valley” = Albania, Prokletije Mts, one of the mountains surrounding the Theth Valley (probably Mt. Jezercë).
“Theth valley”, ex Nádai, ex Holzinger, collected by an unknown mountaineer, vii.1996, holotype (
Albania, Malesia District, above the N side of the Qafa e Valbonës, along the footpath between Rragam and Theth, 1850 m, 42.4068°N, 19.8122°E, leg. TD, ZE, ZF, DM, 06.x.2005 (
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” (
Montenegrina
apfelbecki
caesia
Fehér & Szekeres, 2006 in
Montenegrina
janinensis
caesia
(partim) –
Shell small, tumid. In front view lamella inferior hidden. Lunella dorsal-dorsolateral. Sulcalis residual. Anterior plica superior absent. Clausilium plate almost entirely visible through the aperture.
(in mm). Hs: 12.2–16.3 (holotype 14.3), Ws: 3.9–4.8 (holotype 4.5 mm).
Albania, Prokletije Mts, N side of the Qafa e Tërthorës, 11 km from Bogë toward Theth, 1800 m, 42.392°N, 19.730°E.
Type locality, leg. ZE, ZF, JK, DM, 20.x.2002, holotype (
Central part of the Prokletije Mts in northern Albania. Known only from the type locality (Fig.
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.).
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, medium-bent 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.
Delima laxa perstriata Wagner, 1919: 71–72.
Delima (Albanodelima) perstriata
–
Montenegrina
perstriata
perstriata
–
Shell medium, tumid, dark corneous. Lower whorls smooth, flattened, nearly of the same width. Upper whorls very finely, indistinctly wrinkled-costate. Neck densely costate. Peristome with simple, somewhat reflexed margin. Lamellae superior and spiralis overlap. In front view lamella subcolumellaris not visible. Lunella dorsolateral, not connected to the basalis. Subclaustralis short, sulcalis present. Anterior plica superior absent or weak, separate from the lunella complex. Clausilium plate barely visible through the aperture.
(in mm). Hs: 21.2, Ws: 5.5 (holotype according to
A Montenegrina perstriata perstriata (Wagner, 1919), Galičnik,
Macedonia, Mavrovo District, Galičnik.
According to
Macedonia, 1 km E of Galičnik, 1440 m, 41.5942°N, 20.6716°E, leg. ZF, EH, KJ, HS, 14.x.2014 (
Western part of the Bistra Mts in the vicinity of Galičnik (Fig.
Montenegrina
perstriata
callistoma
Fehér & Szekeres, 2006 in
Montenegrina
perstriata
crassa
–
Montenegrina
perstriata
“crassa” –
Shell medium, light corneous. Lower whorls smooth, upper ones very finely and densely striate. Neck densely striate. Peristome rounded to angular, with strongly swollen, reflexed margin. Lamellae superior and spiralis overlap to long overlap. Subcolumellaris not or barely, clausilium plate not at all visible through the aperture. Lunella lateral, separate from the basalis. Sulcalis residual to weak. Anterior plica superior separate from the lunella complex.
(in mm). Hs: 14.3–18.1 (holotype 16.0), Ws: 3.2–4.3 (holotype 4.2).
Albania, Bulqizë District, Klenjë, near a cave along the Librazhd to Peshkopi road, 1220 m, 41.3646°N, 20.4685°E.
Type locality, leg. ZE, ZF, JK, DM, 24.x.2002, holotype (
Type locality, leg. ZF, TN, EM, 13.iv.2014 (
Gollobordë and Çermenikë Plateaus in eastern central Albania (Fig.
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
diminuta
Fehér & Szekeres, 1999 in
Shell medium, slender, light corneous. Lower whorls flattened, nearly of the same width. All whorls indistinctly wrinkled-costate, more prominently toward the apex. Neck densely costate. Peristome with simple margin. Lamellae superior and spiralis mostly overlap. In front view lamella subcolumellaris not visible. Lunella lateral to ventrolateral, not connected to the basalis. Subclaustralis residual, sulcalis present. Anterior plica superior absent or weak, not connected to the lunella complex. Clausilium plate not visible through the aperture.
(in mm). Hs: 14.0–18.5 (holotype 17.4), Ws: 3.0–3.8 (holotype 3.7).
Macedonia, Galičica Mts, 8 km NW of the Bukovo Pass (= 1 km E of the Zavoj junction) on the Ohrid to Resen road, 950 m, 41.1978°N, 20.9173°E.
Type locality, leg. ZE, ZF, 11.viii.1996, holotype (
Type locality, leg. ZE, ZF, AH, 7.iv.2004 (
Northern extensions of the Galičica Mts (Fig.
Montenegrina
perstriata
drimica
Nordsieck, 1972: 32, plate 4, fig. 38. –
Montenegrina
perstriata
crassa
Erőss & Szekeres, 1999 in
Shell small to large, tumid, light to dark corneous. Lower whorls smooth, flattened, nearly of the same width. Upper whorls finely striate-costate. Neck densely striate-costate. Peristome with weakly to very strongly swollen, reflexed margin. Lamellae superior and spiralis overlap. In front view lamella subcolumellaris often visible. Lunella lateral, separate from the weak basalis. Subclaustralis short, sulcalis present. Anterior plica superior absent or weak, separate from the lunella complex. Clausilium plate not visible through the aperture.
(in mm). Hs: 14.0–21.5 (holotype 16.5), Ws: 3.8–5.0 (holotype 4.3).
Macedonia, valley of the Crni Drin, Lukovo near Struga [drimica]; Macedonia, valley of the Crni Drin, left side, 2 km N of Lukovo and 1 km S of the road junction to Modrić, 41.3654°N, 20.6048°E [crassa].
Lukovo near Struga, leg. H. Nordsieck, 27.viii.1971, holotype [drimica] (
Macedonia, Debar District, S of Debar, at the dam of the Crni Drin, 590 m, 41.4948°N, 20.5052°E, leg. ZF, EH, KJ, HS, 14.x.2014 (
Upper valley of the Crni Drin between the Globočica Lake and Debar (Fig.
This taxon shows considerable intra- and inter-population heterogeneity in terms of shell size and peristome thickness.
Montenegrina perstriata mavrovoensis Nordsieck, 2009: 78, plate 3, fig. 11.
Shell medium, elongate, light corneous. Lower whorls nearly of the same width, almost smooth to finely, indistinctly costate. Upper whorls indistinctly wrinkled-costate. Neck costate. Peristome with simple margin. Lamellae superior and spiralis mostly overlap. Lamella subcolumellaris deep, even in oblique view barely visible. Lunella dorsolateral to lateral, not connected to the weak basalis. Subclaustralis residual, sulcalis present. Anterior plica superior absent or weak and separate from the lunella complex. Clausilium plate mostly not visible through the aperture.
(in mm). Hs: 15.4–18.4 (holotype 17.7), Ws: 3.9–4.5 (holotype 4.3).
Macedonia, Bistra Mts, Mavrovo 2 km towards Galičnik.
Type locality, leg. H. Nordsieck, 4.viii.1974, holotype (
Bistra Mts in western Macedonia. Known only from the type locality (Fig.
Montenegrina
dofleini
occidentalis
Nordsieck, 1977: 85, plate 4, fig. 15. –
Shell small to medium, elongate, light corneous. Lower whorls flattened, nearly of the same width. Surface costate, ribs sharper and wider spaced over the upper whorls and the neck. Peristome rounded to angular, with somewhat swollen margin. Lamellae superior and spiralis long overlap. In front view lamella subcolumellaris not visible. Lunella dorsolateral to lateral, separate from the weak basalis. Subclaustralis residual, sulcalis strong. Anterior plica superior mostly absent, if present separate from the lunella complex. Clausilium plate occasionally visible through the aperture.
(in mm). Hs: 12.8–17.8 (holotype 16.7), Ws: 3.6–4.4 (holotype 4.1).
Macedonia, Kališta near Struga.
Type locality, leg. Nordsieck, 5.viii.1974, holotype (
Macedonia, Kališta, orthodox cemetery, 710 m, 41.1493°N, 20.6493°E, leg. ZE, ZF, AH, 7.iv.2004 (
Western shore of the Ohrid Lake. Known only from two localities (Fig.
Delima (Albanodelima) perstriata ochridensis
–
Delima (Delima) perstriata ochridensis Wagner, 1925: 62, plate 14, figs 98a–c.
Delima (Montenegrina) perstriata ochridensis
–
Montenegrina
perstriata
ochridensis
–
Montenegrina
ochridensis
–
Shell medium, light corneous. Lower whorls smooth to very densely striate, upper ones densely costate. Neck densely costate. Peristome angular, with swollen, reflexed margin. Lamellae superior and spiralis do not overlap. Subcolumellaris not or only barely, clausilium plate not at all visible through the aperture. Lunella lateral, fused to the basalis. Sulcalis absent or residual. Anterior plica superior separate from the lunella complex.
(in mm). Hs: 16.8–20.7, Ws: 4.4–5.1 (
Macedonia, Ohrid.
Syntypes are thought to be in the MIZPAS (not seen).
Macedonia, Ohrid, Izvor Studeničiśta, 700 m, 41.1030°N, 20.8143°E, leg. ZE, ZF, AH, 7.iv.2004 (
Originally this taxon was known only from the eastern coastline of the Ohrid Lake, between Ohrid and Gradište (see:
First ochridensis was regarded by
In the original description the subgenus name Delima is likely a typographic error (
Montenegrina
dofleini
plenostoma
Fehér & Szekeres, 2006 in
Shell medium to large, slender, light corneous. Lower whorls flattened, nearly of the same width. Surface costate, ribs shaper and denser over the upper whorls and the neck. Peristome rounded to angular, with strongly swollen, reflexed margin. Lamellae superior and spiralis overlap. In front view lamella subcolumellaris mostly visible. Lunella dorsolateral, separate from the basalis. Subclaustralis short, sulcalis strong. Anterior plica superior well developed, often connected to the lunella complex. Clausilium plate occasionally visible through the aperture.
(in mm). Hs: 14.8–18.9 (holotype 17.6), Ws: 3.7–4.4 (4.0).
Albania, Pogradec District, 2 km E of the Qafa e Thanës, along the Librazhd to Pogradec road, 860 m, 41.0632°N, 20.6253°E.
Type locality, leg. ZE, ZF, JK, DM, 23.x.2002, holotype (
Type locality, leg. ZF, TN, EM, 12.iv.2014 (
West of the Ohrid Lake (see
Montenegrina
perstriata
radikae
Nordsieck, 1972: 32, plate 4, fig. 37. –
Shell medium to large, tumid, light corneous. All whorls smooth, lower ones nearly of the same width. Neck wrinkled-costate. Basal crest stronger than the peripheral. Peristome with swollen margin. Lamella superior weak, distant from the spiralis. In front view lamella subcolumellaris not or only barely visible. Lunella lateral, fused to the short basalis. Subclaustralis and sulcalis residual. Anterior plica superior absent or short and mostly fused to the lunella complex. Clausilium plate not visible through the aperture.
(in mm). Hs: 17.8–23.1 (holotype 19.7), Ws: 4.6–5.5 (holotype 4.9).
Macedonia, Radika gorge near Debar, at the bridge.
Type locality, leg. H. Nordsieck, 29.viii.1971, holotype (
Macedonia, Mavrovo District, Radika gorge, bridge 1 km N of the Sence junction along the Debar to Gostivar road, 880 m, 41.6994°N, 20.6478°E, leg. ZF, EH, KJ, HS, 14.x.2014 (
Radika Valley in western Macedonia. Known to occur within a small range. All known occurrences are in the immediate vicinity of the type locality. A few kilometers farther upstream in the Radika Valley this taxon is replaced by M. p. subcristatula Nordsieck, 1977 (Fig.
Montenegrina
perstriata
steffeki
Erőss & Szekeres, 2006 in
Montenegrina
dofleini
steffeki
–
Shell medium to large, tumid and somewhat conical, light corneous. Lower whorls almost smooth to variably wrinkled, upper ones indistinctly wrinkled-costate. Neck strongly, irregularly costate. Relatively large peristome with simple margin. Lamellae superior and spiralis overlap. In front view lamella subcolumellaris occasionally visible. Lunella dorsal-dorsolateral to dorsolateral, mostly separate from the basalis. Subclaustralis short, sulcalis strong. Anterior plica superior weak, not connected to the lunella complex. Clausilium plate partly visible through the aperture.
(in mm). Hs: 14.9–21.7 (holotype 17.7), Ws: 3.9–5.4 (holotype 4.9).
Macedonia, valley of the Crni Drin, gorge above and W of Jablanica, about 25 km NNW of Struga.
Type locality, leg. Šteffek, 28.vii.1985, holotype (
Upper valley of the Crni Drin Valley in western Macedonia. The subspecies was collected only once at the type locality. Due to the vague locality data, during later attempts the site could not be found.
In
Montenegrina
perstriata
perstriata
–
Montenegrina
perstriata
subcristatula
Nordsieck, 1977: 84, plate 4, fig. 14. –
Shell medium, slender, light corneous. Lower whorls smooth, flattened, nearly of the same width. Upper whorls indistinctly wrinkled-costate. Neck strongly wrinkled-costate. Peristome with simple margin. Lamellae superior and spiralis overlap. In front view lamella subcolumellaris often visible. Lunella lateral, mostly connected to the basalis. Subclaustralis as long as the basalis, sulcalis present. Anterior plica superior absent or very weak and separate from the lunella complex.
(in mm). Hs: 15.6–18.6 (holotype 18.6), Ws: 3.8–4.2 (holotype 4.2) (
Macedonia, gorge of the Radika near Debar, 1.5 km upstream of the bridge.
Type locality, leg. Nordsieck, 28.viii.1971, holotype (
Macedonia, Gostivar District, upper valley of the Radika, 6 km from the Debar to Gostivar road toward Ničpur, 1100 m, 41.7655°N, 20.6666°E, leg. ZF, EH, KJ, HS, 14.x.2014 (
Valley of the Radika in western Macedonia. This taxon is known to occur within a few km section of the valley. Downstream it is replaced by M. p. radikae (Fig.
In the description,
Montenegrina
perstriata
–
Montenegrina perstriata tenebrosa Nordsieck, 2009: 78–79, plate 3, fig. 14.
Shell medium to large, dark corneous. Lower whorls smooth, upper ones very finely and densely striate. Neck densely striate. Peristome rounded to angular, with strongly swollen, reflexed margin. Lamellae superior and spiralis mostly do not overlap. Subcolumellaris mostly not, clausilium plate not visible through the aperture. Lunella dosolateral-lateral, fused to the basalis. Sulcalis well developed. Anterior plica superior frequently connected to the lunella complex.
(in mm). Hs: 18.3–21.4 (holotype 20.9), Ws: 4.75–5.6 (holotype 5.3).
Albania, Mali i Gurit, Qafa e Zylit, 1800 m.
Type locality, leg. Dhora, 16.ix.1996, holotype (
Albania, Bulqizë District, Maja a Zyllit (= Maja e Shullanit), 6 km from the Librazhd to Peshkopi road toward Sebisht, 1360 m, 41.3611°N, 20.3967°E, leg. ZF, 13.iv.2014 (
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.
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.
Very small, slender subspecies with strongly inflexed neck, weak basalis, but long subclaustralis.
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.
(in mm). Holotype Hs: 10.8, Ws: 2.7, Ha: 2.6, Wa: 2.1; paratypes (
The new subspecies differs from M. p. kovacsorum ssp. n. by its smaller size, non-marginal lamellae inferior and subcolumellaris, as well as its well developed subclaustralis.
Albania, Tropojë District, Dragobi (14 km N of Bajram Curri), gorge of the Përroi i Thatë, 540 m, 42.4364°N, 19.9846°E.
Type locality, leg. TD, ZE, ZF, DM, 6.x.2005, holotype (
The new taxon is named after the Prokletije Mts, the area where this species is distributed.
Northern Albania, Valbona Valley in the southern part of the Prokletije Mts. Known only from the type locality (Fig.
Small, very slender subspecies with strongly inflexed neck and emerged, almost marginal, swollen end of the lamella subcolumellaris.
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.
(in mm). Holotype Hs: 15.5, Ws: 3.5, Ha: 3.5, Wa: 2.7; paratypes (
Differs from M. p. prokletiana ssp. n. by the larger size, marginally ending lamellae inferior and subcolumellaris, and the absence of a subclaustralis.
Albania, Shkodër District, Toplanë, Drin Valley 20.5 km upstream of the Koman Dam, limestone gorge and a cave on the right bank of Koman Lake, 180 m, 42.2339°N, 19.8741°E.
Type locality, leg. ZF, TK, DM, 18.vi.2012, holotype (
Albania, Drin Valley ca. 17.5 km upstream of the Koman Dam, right bank, 42.2258°N, 19.8924°E, leg. ZE, ZF, AH, DM, 15.iv.2006 (
The new taxon is named after Tibor Kovács, entomologist (
Lower Drin Valley in northern Albania. Known to occur upstream of the Koman Dam up to the mouth of the Valbonë (Fig.
The specimens near the mouth of the Valbonë are usually smaller (Hs: 9.0–12.0 mm) with fewer (8–9) whorls. Some flotsam specimens have strongly wrinkled surface, but otherwise very similar shell structure.
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.
Clausilia rugilabris Mousson, 1859: 275–276.
Clausilia (Delima) rugilabris
–
Delima (Albanodelima) rugilabris
–
Montenegrina
rugilabris
–
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.
(in mm). Hs: 16.5–20.4, Ws: 5.1–6.3 (near Ligkiades junction
A Montenegrina rugilabris rugilabris (Mousson, 1859), syntype,
“Ianina” = Greece, Epirus, Ioannina.
“Janina”, ex Mousson, ex Schaefli, syntypes (
Greece, Epirus, Mitsikeli Mts, N of Ligkiades, 1220 m, 39.7019°N, 20.8946°E, leg. JK, DM, Szederjesi, ZU, 4.v.2011 (
Southwestern part and foothills of the Mitsikeli Mts, north of the Lake Pamvotis. Its range overlaps with that of M. janinensis (Fig.
Distribution of Montenegrina rugilabris. M. rugilabris edmundi (empty circle with dot); M. rugilabris golikutensis ssp. n. (triangle); M. rugilabris gregoi ssp. n. (empty triangle); M. rugilabris irmengardis (square); M. rugilabris lambdaformis (star); M. rugilabris rugilabris (circle); M. rugilabris welterschultesi (inverted triangle).
Montenegrina
irmengardis
edmundi
Szekeres, 2006 in
Montenegrina
skipetarica
edmundi
–
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.
(in mm). Hs: 20.2–23.9 (holotype 22.2), Ws: 4.8–6.1 (holotype 5.3).
Greece, Epirus, 6 km from Lia toward Lista, NE of Filiates, 460 m, 39.7358°N, 20.4537°E.
Type locality, leg. PS, Szekeres, 16.v.1997, holotype (
Type locality, 39.7358°N, 20.4537°E, leg. ZE, ZF, JG, 25.vi.2013 (
Southeastern part of the Mt. Mourgana in Epirus, northwestern Greece. In addition to the type locality a site is also known near Tsamantas (Fig.
In a recent assessment Nordsieck (2008) classified M. r. edmundi (as well as the M. rugilabris subspecies irmengardis, lambdaformis Reischütz & Sattmann, 1990 and welterschultesi Fehér & Szekeres, 1999) with M. skipetarica.
Medium-size, light to darker purplish-brown subspecies with smooth whorls, not overlapping lamellae superior and spiralis, in front view not visible lamella subcolumellaris, and mostly separate anterior plica superior and lunella complex.
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.
(in mm). Holotype Hs: 17.6, Ws: 4.7, Ha: 4.3, Wa: 3.5; paratypes Hs: 16.2–20.1 (mean 18.3, S.D. 1.17), Ws: 4.6–5.8 (mean 5.0, S.D. 0.33), Ha: 4.3–5.3, Wa: 3.3–4.2 (
Differs from M. r. irmengardis and M. r. lambdaformis Reischütz & Sattmann, 1990 by the non-overlapping lamellae superior and spiralis, deeper lunella, and the separate basalis, whereas from all other M. rugilabris subspecies by its purplish-brown shell surface.
Albania, Tepelenë District, Kendrevicë Mts, N slope of the Maja e Golikut, 1350 m, 40.2758°N, 20.0947°E.
Type locality, leg. DA, ZF, JG, 28.vi.2014, holotype (
The new taxon is named after the Mt. Goliku (Mali i Golikut), its type locality.
Kendrevicë Mts in southern Albania. Known only from the type locality and its vicinity (Fig.
Medium-size, tumid subspecies with costate apical whorls, long overlapping lamellae superior and spiralis, and in front view visible lamella subcolumellaris.
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.
(in mm). Holotype Hs: 19.7, Ws: 5.7, Ha: 4.9, Wa: 4.0; paratypes Hs: 15.3–19.9 (mean 17.8, S.D. 1.34), Ws: 4.5–5.7 (mean 5.1, S.D. 0.35), Ha: 3.9–4.9, Wa: 3.5–4.1 (
Distinguishable from all other M. rugilabris subspecies by its costate apical whorls.
Greece, Epirus, Ioannina District, 3 km NE of Prosilio, along the road to Syrrako, 1130 m, 39.5768°N, 21.0969°E.
Type locality, leg. ZE, ZF, JG, 23.vi.2013, holotype (
Greece, Epirus, 1 km W of Kipina Monastery, 620 m, 39.5668°N, 21.1231°E, leg. ZE, ZF, JG, 23.vi.2013 (
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.
Lakmos Mts in northwestern Greece. Known from two nearby localities in the western part of the Lakmos (Fig.
The specimens from the vicinity of the Kipina Monastery have weaker sculpture than those of the type locality.
Montenegrina (Beieriella) irmengardis Klemm, 1962: 242–244, plate 3, figs 2 (genital anatomy), 8 (shell).
Montenegrina
irmengardis
irmengardis
–
Montenegrina
skipetarica
irmengardis
–
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.
(in mm). Hs: 20.1–26.2, Ws: 5.2–6.5 (
Greece, Epirus, “Platanusa” (= Platanoussa), 750–800 m.
Type locality, leg. Beier, 3–10.vi.1933, holotype (
Type locality, 600 m, 39.4279°N, 21.0106°E, leg. ZE, ZF, JG, 23.vi.2013 (
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.
Montenegrina irmengardis lambdaformis Reischütz & Sattmann, 1990: 260, plate 3, fig. 7.
Montenegrina
skipetarica
lambdaformis
–
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.
(in mm). Hs: 17–20 (holotype 19.5), Ws: 4.0–5.0 (holotype 4.5 mm).
Greece, Epirus, Pateron Monastery, W of Zitsa.
Type locality, leg. PR, vii.1987, holotype (
Greece, Epirus, E of Lithino, ca. 1 km N of the Pateron Monastery, 360 m, 39.7720°N, 20.6180°E, leg. ZE, ZF, JG, 25.vi.2013 (
Epirus in northwestern Greece. Known only from the vicinity of the Pateron Monastery (Fig.
Montenegrina
irmengardis
welterschultesi
Fehér & Szekeres, 1999 in
Montenegrina
skipetarica
welterschultesi
–
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.
(in mm). Hs: 16.0–22.7 (holotype 22.1), Ws: 4.0–5.0 (holotype 4.8).
Albania, Përmet District, Petran, at the confluence of the Vjosë and Lengaricë Rivers, 300 m, 40.2094°N, 20.4126°E.
Type locality, leg. ZE, ZF, 7.vii.1996, holotype (
Type locality, leg. ZE, ZF, KK, 13.iv.2001 (
Nemerçkë Mts in southern Albania.
Shell small to large, light corneous to violet-brown. Neck variably inflexed. Peristome mostly attached. In front view lamella inferior moderately to well emerged. Subcolumellaris broadly-bent. Lunella short and broad, dorsal-dorsolateral to dorsolateral. Lunella complex with the basalis can form a lambda-like structure. Plica superior fused to the principalis. Basalis and subclaustralis of comparable length. Sulcalis weak or residual. Anterior plica superior absent to strong. Clausilium plate well visible through the aperture. Distinguishable from M. rugilabris by the longer subclaustralis and weak sulcalis.
Clausilia (Delima) skipetarica Soós, 1924: 181–183, figs 2 (shell), 3 (genital anatomy).
Delima (Albanodelima) skipetarica
–
Montenegrina
perstriata
skipetarica
–
Shell large, tumid, reddish-brown with whitish suture. Lower whorls smooth, upper ones finely costate. Neck weakly inflexed, wrinkled-costate. Basal crest well recognizable, peripheral crest weak. Peristome attached, ovoid to somewhat quadrangular, with simple margin. Lamellae superior and spiralis overlap. In front view lamella inferior well emerged, subcolumellaris mostly not visible. Lunella dorsal-dorsolateral, usually connected to the short basalis. Subclaustralis short, sulcalis residual. Anterior plica superior absent or week and separate from the lunella complex.
(in mm). Hs: 18.5–23.4, Ws: 5.6–6.8 (topotypes
A Montenegrina skipetarica puskasi ssp. n., holotype,
“Ura i Lopez” = Albania, Kukës District, 2.5 km N of Bushtricë along the Kukës to Peshkopi road, Ura e Lapavës at the gorge of the Përroi i Vaut të Çajës, 600 m.
The original series, collected by Csiki, were destroyed in 1956 by a fire at the
Type locality, 41.8949°N, 20.4169°E, leg. ZE, ZF, JK, DM, 25.vi.2003 (
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.
Distribution of Montenegrina skipetarica. A M. skipetarica ersekensis (triangle); M. skipetarica flava (star); M. skipetarica konitsae (circle); M. skipetarica nobilis (inverted triangle); M. skipetarica pindica (empty triangle); M. skipetarica rugosa (diamond); M. skipetarica thysi (empty circle with dot); M. skipetarica voidomatis (square) B M. skipetarica csikii (square); M. skipetarica danyii ssp. n. (empty triangle); M. skipetarica gurelurensis ssp. n. (empty circle with dot); M. skipetarica pifkoi ssp. n. (triangle); M. skipetarica puskasi ssp. n. (circle); M. skipetarica remota (star); M. skipetarica skipetarica (empty circle).
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 (
Montenegrina