Corresponding author: Barna Páll-Gergely (
Academic editor: Martin Haase
The Balkan genus
Páll-Gergely B, Deli T, Erőss ZP, Reischütz PL, Reischütz A, Fehér Z (2018) Revision of the subterranean genus
The conchologically similar genus
So far,
The new samples were collected between 2010 and 2017 during 13 collecting trips. Sampling was done scratching out fine granulate material from the superficial fissures of rocks applying long and narrow hand rakes and separated the shells either by sieving (“scratch and sieve” method) or by flotating (“scratch and flotate” method).
Shell whorls (± 0.25) were counted according to Kerney and Cameron (
The following variables were measured: angle of aperture (AA); aperture height (AH); aperture width (AW); shell height (SH); shell width (SW).
The genus
Distribution of the genus
Number of ribs on the body whorl, shell size, and key traits of
(Sub)species | No. of ribs | Shell diameter (in mm) | Key traits |
---|---|---|---|
|
43–93 | 3.6–4.3 | matte protoconch, weak palatal and two weak basal teeth |
35–54 | 3.6–4.2 | widely-spaced ribs, glossy protoconch glossy, toothless aperture | |
|
57–112 | 1.9–3.4 | dense, low ribs, smooth protoconch, toothless aperture |
116 | 3.7 | very low and dense ribs, toothless aperture | |
|
41–52 | 2.8–3.5 | widely spaced, strong ribs, rounded, toothless aperture, finely granular protoconch |
42 | 2.2 | strong, very widely spaced ribs, glossy protoconch, toothless aperture | |
47–54 | 1.9 | strong, widely spaced ribs, glossy protoconch, toothless aperture | |
42–48 | 2.1–2.2 | widely spaced, strong ribs, glossy protoconch, strongly oblique, toothless aperture | |
|
43–76 | 2.6–3.0 | elevated spire, roughly sculptured protoconch, strong ribs, toothless aperture |
|
74–118 | 2.4–3.2 | low basal tooth; palatal part of peristome with strong incision |
64–91 | 2.9–3.5 | pointed basal tooth; palatal part of peristome with shallow incision | |
40–70 | 3.3–3.6 | spiral sculpture, thickened callus |
The reproductive anatomy of
As for shell characters,
The habitat was the only “trait” mentioned by
A large species with usually widely spaced, strong ribs, and no or weak apertural teeth.
The most similar species in terms of shell size and shape is
Kiri-Brücke nächst Mesi b. Skutari, Albanien (im Genist), leg. Sturany, 27.04.1905,
Vrelo Pronisicut (or Pronifkut), coll. Edlauer ex coll. Kuščer,
Protoconch matte; aperture with a rather weak palatal and two weak basal teeth.
Shell rarely flat, usually spire somewhat elevated; protoconch consists of 1.5–1.75 whorls, very finely granulated, rather matte, not glossy; teleoconch with strong, equidistant ribs that are supported by fine periostracal filaments in fresh shells; rib density variable (43–93 ribs on body whorl), usually widely spaced; between main ribs some fine wrinkles discernible; entire shell with 4.25–4.5 whorls; aperture semilunar or due to the straight basal part triangular; peristome expanded and slightly reflected, especially on the palatal, basal and umbilical areas; palatal tooth of variable strength, usually weak, although present in all adult shells, palatal region of peristome without outer incision; basal portion of peristome usually straight, slightly thickened, with two low denticles that are visible in all adult shells; umbilicus funnel-shaped, wide (although width depends on spire height).
See under
Some variability was found between populations in terms of rib density, spire height, and strength of apertural teeth.
This subspecies is distributed in northwestern Albania. Most of the known distribution records are from the Kir River Valley. One shell found in the fluvial debris of the Drin River above the Koman Dam extends the range farther eastwards (Figure
See under
Shells of
Distribution of
Albania, rocks along the road Hani i Hotit to Vermosh, 7.2–7.8 km north of the junction, 370 m a.s.l.,
Albania, Malesia district, Xhajë NE 0.5 km, 650 m a.s.l.,
Albania, rocks along the road Hani i Hotit to Vermosh, 7.2–7.8 km north of the junction, 370 m a.s.l.,
Protoconch glossy; aperture without teeth.
This new subspecies differs from the nominotypical subspecies in the following traits: aperture relatively larger, and its basal area not straight; apertural barriers (teeth) absent; protoconch smooth, glossy; ribs somewhat less dense (rib density on body whorl: 35–54).
This subspecies shows some variability in terms of shell size and rib density.
This new subspecies is named after its toothless aperture, which distinguishes it from the nominotypical subspecies.
This taxon is found in the northeastern part of the Lake Shkodër Basin (Figure
See under
Shells of
Jama Nadjama bei Gnezdu (Izitovice), Krain, leg. Kuščer, coll. Edlauer,
Vetajama bei Sokol, coll. Edlauer,
A small to medium sized species with dense, low ribs, smooth protoconch, and toothless aperture.
Shell nearly flat, but spire somewhat always elevated; protoconch consists of 1.25–1.75 whorls, rather glossy; teleoconch with fine, equidistant, dense ribs; rib density variable (57–112 ribs on body whorl); between main ribs some fine wrinkles discernible; entire shell with 3.25–3.75 whorls; aperture semilunar; peristome slightly thickened and expanded; aperture toothless; umbilicus regular funnel-shaped, relatively narrow (width depends on spire height).
See under
This is a widely distributed species with numerous known populations, most of them with unique character states of spire height, shell size, and rib density.
This species is distributed in the Rumija Mountain between the Shkodër Lake Basin and the Adriatic Sea. Northwards the range extends to the Cetinje area. According to the original labels, type material is of Slovenian origin, however, this species was never again found in Slovenia. It can be reasonably supposed that it is due to mislabelling and the ‘type locality’ is not the site where the type material actually came from (
Assessed as Least Concern (
Shells of
Montenegro, Hotel/Restaurant Izvor north of Sutomore, no GPS available, leg. A. Reischütz, N. Reischütz & P. L. Reischütz, May 2015,
Montenegro, Hotel/Restaurant Izvor north of Sutomore.
A large species with very low and dense ribs; aperture toothless.
Spire slightly elevated; protoconch consists of slightly more than 1.5 whorls, rather glossy; teleoconch with very fine, low, equidistant riblets (approx. 112 on the body whorl); between main ribs some fine wrinkles discernible; entire shell with 3.75 whorls; aperture semilunar, peristome; peristome expanded and slightly reflected on the basal and umbilical areas; aperture toothless; umbilicus regular funnel-shaped, relatively wide.
The only known adult shell is the holotype. Therefore, the morphological diversity within population is unknown.
This new species is named for its dense ribs, which distinguishes it from the most similar
This species is known from the type locality only (Figure
Distribution of
The number of known locations of this species is less than five (i.e. known from a single site) and AOO is smaller than 20 km2, but there is no reason to suppose that AOO, EOO, number of locations, number of subpopulations or the number of mature individuals are declining or extremely fluctuating. Therefore, it should be assessed as Near Threatened (NT).
Krain: Krimberg-Grotte, ex coll. Schmidt,
Velikajama, Soko Kod, Sela Dopilo (geographic position unknown), coll. Edlauer ex coll. Dabović,
A medium sized species with very widely spaced, strong ribs, rounded, toothless aperture, and finely granular protoconch.
Spire somewhat elevated; protoconch consists of 1.5–1.75 whorls, very finely granulated, rather matte, not glossy; teleoconch with strong ribs that are supported by fine periostracal filaments in fresh shells; ribbing less regular than in other congeneric species; ribs widely spaced (41–52 ribs on body whorl); between main ribs some fine wrinkles discernible; entire shell with 3.75–4.25 whorls; aperture toothless, semilunar/rounded; peristome slightly thickened, slightly reflected in direction of umbilicus; umbilicus regular funnel-shaped, relatively narrow.
The most similar taxon to
This species is distributed in the southeastern Alps (Central Slovenia) relatively far from the ranges of its congeneric taxa.
As there are several known locations and no reason to suppose that the habitat quality, habitat extent or population are deteriorating or extremely fluctuating, it was assessed as Least Concern (
Shells of
Distribution of
Montenegro, SE of Virpazar, 4.3 km (in a straight line) SSE of Ðuravci, near Besa/Bes near Krone i Besit, 330 m a.s.l.,
Montenegro, SE of Virpazar, 4.3 km (in a straight line) SSE of Ðuravci, near Besa/Bes near Krone i Besit, 330 m a.s.l.,
A small, nearly flat species with strong, widely spaced ribs, glossy protoconch and strongly oblique, toothless aperture.
Spire somewhat elevated; protoconch consists of 1.25–1.5 whorls, smooth, glossy; teleoconch with very strong (thick), equidistant, widely spaced ribs; rib density: 42–48 ribs on body whorl; between main ribs some fine wrinkles discernible; entire shell with 3.5–3.75 whorls; aperture semilunar, toothless, strongly oblique to shell axis; peristome slightly thickened and expanded in direction of umbilicus; umbilicus funnel-shaped, relatively narrow.
Specimens of the type sample show no notable conchological variability.
This new species is named after our colleague and friend, András Hunyadi, who is one of those who first collected this species.
This species is known from the type locality only (Figure
To our present knowledge this species is very rare (currently known from a single location) and thus AOO is smaller than 20 km2. However, there is no reason to suppose that AOO, EOO, number of locations, number of subpopulations or the number of mature individuals are declining or extremely fluctuating. Therefore, it might be assessed as Near Threatened (NT).
Shells of
Montenegro, S of Virpazar, 0.8 km (in a straight line) E of Limljani, above the village, 400 m a.s.l.,
Montenegro, Seoća S 1 km, along the Virpazar-Ostros road, 280 m a.s.l.,
Montenegro, S of Virpazar, 0.8 km (in a straight line) E of Limljani, above the village, 400 m a.s.l.,
A small, nearly flat species with strong, very widely spaced ribs, glossy protoconch and a toothless aperture.
Spire somewhat elevated; protoconch consists of ca 1.25–1.5 whorls (the holotype is corroded at the protoconch-teleoconch junction), rather smooth, moderately glossy; teleoconch with very strong, equidistant, widely spaced ribs (42 on the body whorl); between main ribs some fine wrinkles discernible; entire shell with 3.75 whorls; aperture semilunar, toothless; peristome slightly thickened and expanded; umbilicus funnel-shaped, relatively narrow.
The widely spaced ribs are similar to
See remarks.
This new species is named after its remarkably widely spaced ribs.
See under Remarks and Figure
The holotype of this species was found in a large sample of
To our present knowledge this species is very rare (currently known from two locations) and thus AOO is smaller than 20 km2. However, there is no reason to suppose that AOO, EOO, number of locations, number of subpopulations or the number of mature individuals are declining or extremely fluctuating. Therefore, it might be assessed as Near Threatened (NT).
Obodska pećina, Rij. Crnojevića, Mtg., September 1956,
Vodna jama (Lovćen), coll. Edlauer ex coll. Kuščer,
A medium sized species with elevated spire, roughly sculptured protoconch, strong ribs on the teleoconch, and toothless aperture.
Spire elevated, shell low conical; protoconch consists of 1.25–1.5 whorls, roughly granulated/”hammered”, matte, not glossy; teleoconch with strong, equidistant ribs that are supported by fine periostracal filaments in fresh shells; rib density variable (43–76 ribs on body whorl), usually widely spaced; between main ribs some fine wrinkles discernible; entire shell with 3.75–4 whorls; aperture semilunar, toothless; peristome slightly thickened and expanded, especially in direction of the umbilicus; umbilicus funnel-shaped, relatively narrow.
The most similar species is
This species is the most variable in terms of shell size and rib density.
This species is found northwest of the Shkodër Lake Basin, as well as in the Zeta River Valley between Podgorica and the Ostrog Monastery (Figure
As there are at least seven known locations and no reason to suppose that the habitat quality, habitat extent or population are deteriorating or extremely fluctuating, it was assessed as Least Concern (
Type sample of
A small to medium sized species with dense riblets, and strong parietal and basal teeth/thickenings.
This species differs from most other
As there are more than five known locations and no reason to suppose that the habitat quality, habitat extent or population are deteriorating or extremely fluctuating, it was assessed as Least Concern (
Obodska pećina, Rij. Crnojevića, Mtg., 1956 September,
Bioče, N Titograd, Mtg. 23.09.1978,
Basal tooth/thickening low, not pointed; palatal part of peristome with strong incision at the position of the palatal tooth, palatal region strongly “pushed” from outside.
Spire elevated; protoconch consists of 1.5 whorls, roughly granulated/”hammered”, matte, not glossy; teleoconch with strong but dense, equidistant ribs that are supported by fine periostracal filaments in fresh shells; rib density variable (74–118 ribs on body whorl); between main ribs some fine wrinkles discernible; entire shell with 3.75–4.5 whorls; aperture semilunar or triangular due to straight basal part; peristome thickened, slightly expanded and slightly reflected on the palatal, basal and umbilical areas; palatal tooth strong, pointed; palatal region of peristome with strong outer incision (i.e. the position of the palatal tooth is indicated with a groove on the outer side); basal portion of peristome straight, thickened; occasionally two small denticles visible, sometimes only the one situated closer to the palatal side is developed as a low denticle; umbilicus funnel-shaped, wide to relatively narrow (width depends on spire height).
See under
This subspecies is particularly variable in terms of shell size. The shape of the thickening of the basal part of the peristome is also slightly variable, although it never develops to a pointed tooth.
This taxon is found north and northwest of the Shkodër Lake Basin in Montenegro (Figure
The only sample in the
Formerly, Peuta Cave population was incorrectly referred to as
Albania, rocks north of Hajmel at the bridge, north of Lezhe, 40 m a.s.l.,
Albania, Shkodër District, Vau i Dejës, Mjedë, near the dam, 45 m a.s.l., limestone rocks,
Basal tooth pointed; palatal part of peristome with slight incision at the position of the palatal tooth, palatal region not or slightly “pushed” from outside.
This subspecies is variable in terms of shell size and spire height, i.e. one population has a nearly flat shell.
This new subspecies is named after its pointed (Latin:
This subspecies is known from the southeastern side of the Shkodër Lake Basin in Albania (Figure
Shells of
Albania, Malësia district, a mountain pass 2 km N of Rraps-Starjë, 700 m a.s.l.,
Albania, Malësia district, a mountain pass 2 km N of Rraps-Starjë, 700 m a.s.l.,
A medium sized to large species with elevated spire, strong, very widely spaced ribs, fine spiral lines consisting of series of minute tubercles, hammered protoconch, weak basal thickening, weak parietal tooth, and a thickened parietal callus.
Spire elevated; protoconch consists of ca. 1.25–1.75 whorls, roughly sculptured, “hammered”; teleoconch with very strong, equidistant, widely spaced ribs (40–70 on the body whorl; addition to the radial ribs very fine, dense spiral striation is visible on the entire teleoconch, consisting of minute tubercles; entire shell with 4–4.75 whorls; aperture subcircular/triangular; basal part with two low swellings (similar to those of
This new species can be distinguished from all congeners (especially the most similar
The degree of the thickness of the parietal callus shows some recognisable variability within the single known population, but this trait might be due to the age (degree of development) of the examined shells.
This new species is named after its resemblance to
Initially we considered placing this species to the genus
Formerly, this population was incorrectly referred to as
To our present knowledge this species is very rare (currently known from a single location) and thus AOO is smaller than 20 km2. However, there is no reason to suppose that AOO, EOO, number of locations, number of subpopulations or the number of mature individuals are declining or extremely fluctuating. Therefore, it might be assessed as Near Threatened (NT).
We examined all available
Although the majority of species is known from multiple populations of relatively large areas, three single-site endemic species (
Co-occurrences of
Locality | co-occurring species |
---|---|
Montenegro, SE of Virpazar, 4.3 km (in a straight line) SSE of Ðuravci, near Besa/Bes near Krone i Besit, 330 m a.s.l. | |
Yugoslavia: Pečina u Peckom Brdu cave above Začir | |
Obodska pećina, Rij. Crnojevića | |
Montenegro, S of Virpazar, 0.8 km (in a straight line) E of Limljani, above the village, 400 m a.s.l. |
We have collected all
Literature data are very scarce about MSS dwelling gastropods, and still, that little is dealing mostly with scree slopes, habitats where fragments of rocks are accumulating at the bottom of rocky walls, and are covered over time by an evolving soil (
Our sites represent a different sort of habitat type, similar to “bedrock MSS” according to the subdivision of
“Scratch and flotate” method to collect shells of subterranean species.
It is still disputed whether such a bedrock MSS habitat is characteristically different from caves or it can be considered just as the extension of the cave system, and therefore, its fauna is composed of troglobionts of wider ecological tolerance or there are certain specialized elements exclusive of MSS (see
We are grateful to Anita Eschner (