Research Article |
Corresponding author: Miklós Szekeres ( szekeres.miklos@gmail.com ) Academic editor: Barna Páll-Gergely
© 2023 Levan Mumladze, Jozef Grego, 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:
Mumladze L, Grego J, Szekeres M (2023) The land snail family Clausiliidae (Gastropoda, Pulmonata, Stylommatophora) in Georgia: overview, novel records and a new species. Caucasiana 2: 29-61. https://doi.org/10.3897/caucasiana.2.e101013
|
The Caucasus ecoregion is one of the most important biodiversity hotspots in western Eurasia. Georgia is situated in the middle of this area, between the ranges of the Greater and Lesser Caucasus, a position providing exceptionally balanced environmental conditions for its flora and fauna. Animal groups of low mobility, such as land snails, are important indicators of the past and present geographic and climatic events shaping these unique ecosystems. We give an overview of the land snail family Clausiliidae in Georgia that includes 30 species, among which eight are endemic to this country. Their distribution areas are reassessed in the light of novel georeferenced occurrence data resulting from recent field work. Aspects of the origin, diversity, habitat requirements and conservation status of the Georgian Clausiliidae fauna are discussed. Additionally, Mucronaria (Mucronaria) kartvelica sp. n. is described from the Racha-Lechkhumi and Kvemo Svaneti Region.
Caucasus, endemisms, land snails, new species, zoogeography
The Caucasus ecoregion is a major biodiversity hotspot in western Eurasia (
The family of the Clausiliidae is one of the most species-rich and best studied groups of land snails in the Caucasus region. Remarkably, 89 of the 92 clausiliid species occurring here are endemic to this ecoregion. A particularly high diversity can be seen in Georgia, which is home to more than 30% of the region's clausiliid species.
In Georgia the second half of the 19th century brought a golden age of natural history research. Extensive field trips of the devoted naturalist explorers Frédéric Dubois de Montpéreux, Gustav Radde, Hans Leder, Alexander Schläfli, Carl Reuleaux and Alexander Brandt made available a trove of unknown species, which were then forwarded to specialists for identification. The resulting pioneering publications of Ludwig
This paper offers an overview of the clausiliids of Georgia without the territories of Abkhazia and South Ossetia (i.e., parts of the Racha-Lechkhumi, Shida Kartli and Mtskheta-Mtianeti Regions). New georeferenced locality data, stemming mainly from the authors' field trips during the past two decades, as well as considerations on the historic and zoogeographic aspects of the distribution areas, give an updated assessment of this unique fauna. Additionally, we provide a taxonomic description of a recently discovered species of the genus Mucronaria Boettger, 1877.
The occurrence data listed in the Results are based on georeferenced samples that were collected during the past two decades. For comparison, the maps showing the occurrence points also include well defined locality data that have been provided in the publications of
The new occurrence data are given by the administrative regions of Georgia and in a north to south order. The geographic coordinates and altitude are followed by the abbreviated name(s) of the collector(s), the collection year and, in brackets, the abbreviations identifying the collection(s) where the voucher material is housed. In the maps black dots denote the occurrence points reported in this paper, whereas white dots correspond to those that are derived from the aforementioned five publications. Estimation of extent of occupancy (EOO) was based on the minimum convex polygon delimiting the entire distribution range of a species. Area of occupancy (AOO) was calculated as the sum of 2 x 2 km grid cells containing georeferenced occurrence of a species in Georgia. For areas exceeding 10,000 km2 the infinity symbol is used. EOO and AOO data are given only for species that are endemic to Georgia or near-endemics with distribution ranges extending only slightly beyond the borders of the country.
The listed samples were collected by Ani Bikashvili (AB), Elizaveta Chertoprud (EC), Olga Danilova (OD), Jozef Grego (JG), Bella Japoshvili (BJ), Levan Mumladze (LM), Tornike Mumladze (TM), Dávid Murányi (DM), Mari Murtskhvaladze (MM), Mate Natsvlishvili (MN), Mário Olšavský (MO), Igor Solodovnikov (IS) and Miklós Szekeres (MS). The collections mentioned in the text are those of the Field Museum of Natural History, Chicago (
Subfamily Phaedusinae Wagner, 1922
Clausilia perlucens
—
"Kaukasus".
Southeastern areas of the Greater and Lesser Caucasus, as well as the Talysh and Alborz Mountains in Iran.
Moist forests; in the texture and under the bark of decaying trees.
Kakheti • N41.9837°, E45.8456°; 680 m; leg. LM, 2010 (ISU) • N41.9823°, E45.8540°; 840 m; leg. JG-LM-MS, 2021 (MS) • N41.9823°, E45.8540°; 840 m; leg. LM, 2010 (ISU) • N41.9761°, E45.8413°; 630 m; leg. JG-LM-MS, 2021 (MS) • N41.8490°, E46.3314°; 1300 m; leg. LM, 2010 (ISU) • N41.4935°, E46.1009°; 680 m; leg. LM, 2010 (ISU) • N41.4921°, E46.0994°; 710 m; leg. JG-LM-MS, 2021 (ISU, JG, MS).
Serrulina (Serrulina) goettingi — Brandt 1961: p. 18, plate 2 fig. 17
Turkey, Bolu Province, vicinity of Lake Abant.
Southeastern Bulgaria and northwestern Turkey to Samsun toward east.
Humid broadleaf forests; in the trunks and roots of decaying trees.
This species does not occur in Georgia. An empty shell (
Clausilia funiculum
—
Boztepe (Turkey, suburb of Trabzon).
Southeastern and eastern coastal areas of the Black Sea between Trabzon (Turkey) and Tuapse (Russia).
Colchic broadleaf forests; in the texture and under the bark of decaying trees.
Adjara • N41.7074°, E41.7759°; 40 m; leg. LM, 2016 (ISU) • N41.7020°, E41.7211°; 70 m; leg. JG, 2017 (JG) • N41.6713°, E41.6900°; 30 m; leg. JG, 2017 (JG, MS) • N41.5652°, E41.5871°; 310 m; leg. LM-TM, 2019 (ISU).
Pontophaedusa gregoi
—
Katskhi (Imereti Region).
Near Katskhi and Zastafoni in central Imereti Region.
Karstic broadleaf forests; primarily in subterranean rock crevices.
Imereti • N42.2836°, E43.2113°; 500 m; leg. JG-LM-MS, 2021 (ISU, NHMUK 20220497,
(A) Caspiophaedusa perlucens, Kakheti, Artsivi Gorge northwest of Dedoplistskaro, N41.4921°, E46.0994°. (B) Dobatia goettingi, Turkey, 9 km northeast of Lake Abant, N40.6569°, E31.3892°. (C) Pontophaedusa funiculum, Adjara, Makhinjauri northeast of Batumi N41.6713°, E41.6900°. (D) Pontophaedusa gregoi, Imereti, between Chiatura and Katskhi, N42.2836°, E43.2113°. (E) Pravispira semilamellata, Samegrelo-Zemo Svaneti, northwest of Khobi, N42.3944°, E41.8382°. (F) Serrulina serrulata, Kakheti, valley of the Alazani River near Birkiani, N42.2368°, E45.3386°. (G) Serrulina sieversi, Iran, Gilan Province, valley of the Shim Rud south of Siyahkhal, N37.0239°, E49.8769°. (H) Serrulinella senghanensis, Iran, Gilan Province, valley of the Shim Rud at Lunak, N37.0089°, E49.8642°. Scale bar: 5 mm.
Clausilia semilamellata
—
"Réduktaleh" (misspelled Redutkaleh, = Kulevi, Samegrelo-Zemo Svaneti Region).
Mountains along the Black Sea coast between Rize (Turkey) and Tuapse (Russia), the Lesser Caucasus and the southern slopes of the Greater Caucasus west of Tbilisi. Isolated occurrences near Vladikavkaz (Russia), Gebele (Azerbaijan), and in the Talysh Mountains (Iran).
Forests up to around 2000 m; in the texture and under the bark of decaying trees.
Adjara • N41.7959°, E41.9452°; 210 m; leg. LM, 2018 (ISU) • N41.7377°, E41.9839°; 440 m; leg. LM, 2010 (ISU) • N41.6458°, E42.4853°; 1630 m; leg. LM, 2017 (ISU). Imereti • N42.5028°, E42.5579°; 860 m; leg. JG, 2017 (JG) • N42.5023°, E42.5595°; 880 m; leg. JG-LM, 2018 (JG) • N42.2443°, E43.2780°; 410 m; leg. AB-BJ-LM, 2021 (ISU) • N42.1612°, E43.3577°; 830 m; leg. AB-BJ-LM, 2021 (ISU) • N42.1499°, E43.3025°; 540 m; leg. AB-BJ-LM, 2021 (ISU) • N42.1476°, E42.8037°; 130 m; leg. LM, 2012 (ISU). Kakheti • N42.2368°, E45.3386°; 900 m; leg. JG-LM-MS, 2021 (ISU, JG, MS) • N41.8759, E45.3564; 1080 m; leg. LM, 2010 (ISU) • N41.8647°, 45.3355°; 1260 m; leg. LM, 2010 (ISU). Kvemo Kartli • N41.7511°, E44.5094°; 1610 m; leg. LM, 2021 (ISU). Mtskheta-Mtianeti • N41.8484°, E44.6379°; 480 m; leg. LM, 2014 (ISU). Racha-Lechkhumi and Kvemo Svaneti • N42.5681°, E43.4953°; 1060 m; leg. LM, 2014 (ISU) • N42.5561°, E43.5691°; 1560 m; leg. EC-JG-LM-MO-MS, 2022 (ISU) • N42.5466°, E43.5308°; 1070 m; leg. EC-JG-LM-MO-MS, 2022 (MS) • N42.4030°, E42.9697°; 1500 m; leg. JG, 2021 (JG) • N42.3914°, E42.9823°; 1520 m; leg. JG-MS, 2021 (JG, MS). Samegrelo-Zemo Svaneti • N43.0652°, E42.4113°; 1250 m; leg. LM, 2010 (ISU) • N42.9109°, E42.0765°; 740 m; leg. LM, 2010 (ISU) • N42.7528°, E42.5093°; 2050 m; leg. LM, 2014 (ISU) • N42.7367°, E42.5015°; 1600 m; leg. LM, 2015 (ISU) • N42.4774°, E42.4587°; 700 m; leg. JG, 2019 (JG, MS) • N42.3944°, E41.8382°; 450 m; leg. JG-MS, 2021 (JG, MS). Samtskhe-Javakheti • N41.8092°, E43.3112°; 970 m; leg. LM, 2012 (ISU) • N41.7912°, E43.4665°; 1240 m; leg. LM, 2012 (ISU).
Clausilia serrulata
—
"Tauria" (= Crimea; erroneous!).
Northern Anatolia, as well as the Greater and Lesser Caucasus ranges to Zaqatala (Azerbaijan) toward east. Isolated occurrences in the Northern Caucasus near Vladikavkaz, the Ukrainian Carpathians, eastern regions of Romania and Bulgaria, as well as in Hatay Province in southern Turkey.
Humid, primarily broadleaf forests up to around 2000 m; in the texture and under the bark of decaying trees.
Adjara • N41.7477°, E42.0996°; 1860 m; leg. LM, 2018 (ISU) • N41.7351°, E42.0930°; 1180 m; leg. LM, 2010 (ISU) • N41.6527°, E41.7625°; 550 m; leg. LM, 2017 (ISU). Imereti • N42.3829°, E43.0122°; 950 m; leg. JG-LM, 2018 (JG, MS). Kakheti • N42.2368°, E45.3386°; 900 m; leg. JG-LM-MS, 2021 (ISU, JG, MS) • N42.0539°, E45.5274°; 480 m; leg. LM, 2019 (ISU) • N41.9823°, E45.8540°; 840 m; leg. JG-LM-MS, 2021 (ISU, JG, MS) • N41.9823°, E45.8540°; 840 m; leg. LM, 2010 (ISU) • N41.8759°, E45.3564°; 1080 m; leg. LM, 2010 (ISU) • N41.8556°, E46.3410°; 1730 m; leg. LM, 2010 (ISU). Mtskheta-Mtianeti • N42.0974°, E44.8236°; 1420 m; leg. LM, 2010 (ISU). Racha-Lechkhumi and Kvemo Svaneti • N42.3914°, E42.9823°; 1520 m; leg. JG-LM-MS, 2021 (ISU, JG, MS) • N42.3876°, E42.9748°; 1500 m; leg. LM, 2018 (ISU). Samegrelo-Zemo Svaneti • N42.9109°, E42.0765°; 740 m; leg. LM, 2010 (ISU) • N42.6599°, E42.4330°; 950 m; leg. LM, 2010 (ISU) • N42.3944°, E41.8382°; 450 m; leg. LM, 2010 (ISU), JG-MS, 2021 (JG, MS).
Clausilia sieversi
—
Lankaran (Azerbaijan).
The Talysh Mountains in Azerbaijan and Iran, as well as the Alborz Mountains in Iran to eastern Mazandaran Province. Isolated along the southern slopes of the Greater Caucasus around Zaqatala (Azerbaijan).
Moist broadleaf forests up to 1500 m; in the texture and under the bark of decaying trees.
According to its description, the subspecies S. sieversi occidentalis Likharev, 1962 occurs between Lagodekhi (Kakheti Region) and Zaqatala in Azerbaijan. Though in the Zoological Museum of Moscow State University there is a sample of this form with locality given as the Lagodekhi Nature Reserve (
Serrulina senghanensis
—
"Siah Senghan" (locality in Iran's Gilan Province).
This species is known only from a small area in the Alborz Mountains south of Lahijan (Iran, Gilan Province).
Moist broadleaf forests in karst areas; in subterranean rock crevices.
Reports of this Iranian species from Georgia (
Acrotoma (Iliamneme) baryshnikovi
—
Akhsargina (South Ossetia).
Mountain ridge between Kvedi (Racha-Lechkhumi and Kvemo Svaneti Region) and Kvaisi (South Ossetia).
Karst areas; on and underneath limestone cliffs.
Racha-Lechkhumi and Kvemo Svaneti • N42.5561°, E43.5721°; 1510 m; leg. JG-MS, 2021 (JG, MS) • N42.5561°, E43.5691°; 1560 m; leg. JG-MS, 2021 (ISU, NHMUK 20220498,
(A) Acrotoma (Iliamneme) baryshnikovi, Racha-Lechkhumi and Kvemo Svaneti, Kvedi 1 Cave southeast of Kvedi, N42.5561°, E43.5721°. (B) A. (I.) enguriensis, Samegrelo-Zemo Svaneti, 6 km N of Jvari, N42.7500°, E42.0489°. (C) Armenica (Armenica) unicristata, Kakheti, Artsivi Gorge northwest of Dedoplistskaro, N41.4888°, E46.0956°. (D) A. (Astrogena) griseofusca, Turkey, Trabzon Province, Sümela south of Trabzon, N40.6894°, E39.6589°. (E) Elia (Caucasica) ossetica, Kakheti, Batsara Nature Reserve, N42.2229°, E45.3020°. (F) E. (C.) somchetica, Samtskhe-Javakheti, Didveli south of Bakuriani, N41.6976°, E43.5188°. (G) E. (C.) tuschetica, Kakheti, southwest of the Abano Pass, N42.2422°, E45.4946°. (H) E. (Megaleuxina) derasa, Samtskhe-Javakheti, east of the Goderdzi Pass, N41.6603°, E42.6042°. Scale bar: 5 mm.
Acrotoma (Iliamneme?) enguriensis — Haudsorf et al. 2018: p. 698, figs 4, 28, 44, 45, 52
Mountain ridge between the Enguri and Magana Rivers, north of Jvari (Samegrelo-Zemo Svaneti Region).
Known only from the type locality.
Karst areas; on and underneath limestone cliffs.
Samegrelo-Zemo Svaneti • N42.7500°, E42.0489°; 470 m; leg. JG-LM, 2018 (ISU, NHMUK 20220499,
Clausilia unicristata
—
"Ekatherinenfeld" (= Bolnisi, Kvemo Kartli Region).
Occurs sporadically in southern and southeastern Georgia, northern and southeastern Armenia, as well as northern and western Azerbaijan.
Exposed metamorphic or limestone cliffs; around and inside rock crevices.
Kakheti • N41.4896°, E46.0976°; 750 m; leg. DM, 2019 (
The Armenica from Sakavre (Shida Kartli Region) (ISU B27-501), mentioned by
Clausilia (Alinda) griseofusca
—
Vicinity of Lake Tabatskuri (Samtskhe-Javakheti Region; possibly erroneous!).
All verified localities of this species are in Trabzon and Rize Provinces of Turkey.
Broadleaf or mixed forests between 1000 and 1500 m; on tree trunks and small, shaded cliffs.
The description and a probable syntype (
Clausilia somchetica var. ossetica
—
"Koischet" (unidentified site, likely on the northern slopes of the Greater Caucasus near the border between Chechnya and Dagestan, in Russia).
Central regions of the Greater Caucasus, as well as the Lesser Caucasus between Borjomi (Samtskhe-Javakheti Region) and Lake Sevan (Armenia).
Moist mountain forests and rocky subalpine meadows up to 2500 m; among leaf litter, in rock crevices and under stones.
Mtskheta-Mtianeti • N42.6979°, E44.6188°; 1910 m; leg. LM, 2017 (ISU) • N42.5377°, E44.4927°; 2280 m; leg. LM, 2017 (ISU) • N42.4858°, E44.4788°; 2440 m; leg. OD-IS, 2009 (IS, MS) • N42.4148°, 44.5500°; 1690 m; leg. LM, 2010 (ISU) • N42.3105°, E45.1229°; 1490 m; leg. LM, 2010 (ISU). Kakheti • N42.2368°, E45.3386°; 900 m; leg. JG-LM-MS, 2021 (ISU, MS) • N42.2229°, E45.3020°; 810 m; leg. DM, 2019 (
Clausilia somchetica
—
"Somchetia" (= Somkheti).
Widely distributed along the ranges of the Greater and Lesser Caucasus. Isolated occurrences in the Oshten-Fisht mountain complex and at Stavropol (both in Russia), as well as in Kars Province (Turkey).
Moist mountain forests and rocky subalpine meadows up to 2500 m; among leaf litter, in rock crevices and under stones.
Samtskhe-Javakheti • N41.6976°, E43.5188°; 2420 m; leg. OD-IS, 2009 (IS, MS) • N41.7014°, E43.5131°; 2210 m; leg. JG-LM-MS, (ISU, MS) • N41.5260°, E43.0310°; 1320 m; leg. LM, 2020 (ISU) • N41.5187°, E43.0397°; 1510 m; leg. LM, 2020 (ISU) • N41.3995°, E43.6196°; 2050 m; leg. LM, 2017 (ISU).
Somkheti, the type locality, is a loosely defined historic region roughly corresponding to the Kvemo Kartli Region and Lori Province in Armenia.
Euxina (Euxina) tuschetica —
Kakheti Region, near the village Zemo Omalo, at 1900 m.
Mountain Tusheti.
Mountain forests and subalpine meadows between 1000 and 2000 m; among leaf litter, in rock crevices and under stones.
Kakheti • N42.2765°, E45.3524°; 1280 m; leg. JG-LM-MS, 2021 (ISU, JG, MS) • N42.2744°, E45.3517°; 1200 m; leg. DM, 2019 (
Clausilia derasa
—
"Réduktaleh" (misspelled Redutkaleh, = Kulevi, Samegrelo-Zemo Svaneti Region) and Kutaisi (Imereti Region).
Between the ranges of the Greater and Lesser Caucasus from the Black Sea coast to Tbilisi toward east.
Cliffs in moist forests; in leaf litter and loose soil under cliff vegetation.
Adjara • N41.7477°, E42.0996°; 1860 m; leg. LM, 2018 (ISU) • N41.7435°, E42.0823°; 1290 m; leg. LM, 2018 (ISU) • N41.7351°, E42.0930°; 1180 m; leg. LM, 2010 (ISU) • N41.6835°, E41.8889°; 550 m; leg. LM, 2014 (ISU). Guria • N41.8992°, E42.3707°; 1240 m; leg. LM, 2017 (ISU) • N41.8367°, E42.3314°; 2070 m; leg. LM, 2017 (ISU). Imereti • N42.3829°, E43.0122°; 950 m; leg. JG-LM, 2018 (ISU, JG) • N42.3778°, E42.6022°; 170 m; leg. LM, 2018 (ISU) • N42.3254°, E43.2678°; 630 m; leg. LM, 2021 (ISU) • N42.3095°, E42.6697°; 390 m; leg. JG, 2017 (JG) • N42.2998°, E42.9622°; 530 m; leg. AB-LM, 2021 (ISU) • N42.2945°, E42.7677°; 400 m; leg. JG, 2017 (JG); LM, 2021 (ISU) • N42.2890°, E43.2898°; 390 m; leg. LM-TM, 2021 (ISU) • N42.2873°, E43.2164°; 620 m; leg. LM, 2011 (ISU) • N42.2828°, E42.7583°; 210 m; leg. JG, 2017 (JG, MS) • N42.2771°, E42.7044°; 200 m; leg. JG, 2017 (JG, MS) • N42.2716°, E42.8531°; 350 m; leg. LM, 2016 (ISU) • N42.2620°, E42.9575°; 240 m; leg. LM, 2017 (ISU) • N42.2575°, E42.7102°; 150 m; leg. DM, 2018 (
Clausilia (Euxinastra) hamata
—
Batumi (Adjara Region).
Mountainous areas along the Black Sea coast from Rize Province (Turkey) to the Adjara Region.
Colchic deciduous forests; on decaying tree trunks and in leaf litter.
Adjara • N41.7959°, E41.9452°; 210 m; leg. LM, 2018 (ISU) • N41.7351°, E42.0930°; 1180 m; leg. LM, 2010 (ISU) • N41.6617°, E41.8560°; 860 m; leg. LM, 2014 (ISU).
(A) Euxinastra hamata, Adjara, valley of the Tchorokhi River south of Mirveti, N41.5165°, E41.7170°. (B) Filosa filosa, Adjara, 2 km northeast of Korolistavi, N41.6502°, E41.7647°. (C) Inobseratella lindholmi, Guria, between Ckhakoura and Bakhmaro, N41.8914°, E42.3697°. (D) I. monticola, Turkey, Rize Province, Fırtına Valley S of Çamlıhemşin, N40.9600°, E40.9622°. (E) Mentissoidea rupicola, Samtskhe-Javakheti, south of Didveli, N41.7197°, E43.4949°. (F) Mucronaria (Mucornaria) acuminata, Samtskhe-Javakheti, north of the Tskhratskaro Pass, N41.7014°, E43.5131°. (G) M. (M.) duboisi, Imereti, near the Motsameta Monastery, N42.2828°, E42.7583°. (H) M. (M.) strauchi, Kvemo Kartli, NW of Sakavre, N41.8424°, E44.1628°. Scale bar: 5 mm..
Clausilia filosa
—
"Chysirkaleh" (unidentified locality somewhere southwest of Batumi, Adjara Region).
Areas along the Black Sea coast from Trabzon Province (Turkey) to Adjara.
Colchic deciduous forests; on decaying tree trunks and in leaf litter.
Occurrence data (Figure
Clausilia lindholmi
—
"Berge Salolet" (= Mount Sallet south of Artvin, Turkey).
The Parmak and Karçal Mountains in Artvin Province (Turkey) and the Meshkheti Range in the Guria Region.
Deciduous forests above 800 m; on and around live trees.
Guria • N41.8914°, E42.3697°; 1490, m; leg. EC-JG-LM-MO-MS, 2022 (ISU, JG, MS)
The occurrence of I. lindholmi around the above mentioned site in the Guria Region has already been reported by
Kazancia monticola
—
Çatköy south of Çamlıhemşin (Turkey, Rize Province).
This species has been known to occur in the Fırtına Valley southwest of Çamlıhemşin (Turkey, Rize Province). Recently
Forested rocky areas; on trees, in vegetation beneath cliffs and under stones.
Clausilia rupicola
—
Tortum (Erzurum Province, Turkey; likely erroneous!).
Coastal areas of the Black Sea from Artvin Province (Turkey) to Gelendzhik (Russia), the Western Caucasus, southern slopes of the Greater Caucasus to Lagodekhi (Kakheti Region) and northern slopes of the Lesser Caucasus to the Murovdag (Azerbaijan) toward east. Isolated occurrences in Northern Ossetia and Chechnya.
Broadleaf and mixed forests; on trees and under the bark of decaying logs.
Adjara • N41.7477°, E42.0996°; 1860 m; leg. LM, 2018 (ISU) • N41.7351°, E42.0930°; 1180 m; leg. LM, 2010 (ISU). Imereti • N42.5023°, E42.5595°; 880 m; leg. JG-LM, 2018 (ISU, JG) • N42.3829°, E43.0122°; 950 m; leg. JG-LM, 2018 (ISU, JG) • N41.9236°, E42.7494°; 790 m; leg. JG-LM-MS, 2021 (ISU, JG, MS). Kakheti • N42.2765°, E45.3524°; 1280 m; leg. JG-LM-MS, 2021 (ISU, JG, MS) • N42.1523°, 45.4171°; 580 m; leg. AB-BJ, 2019 (ISU) • N41.9823°, 45.8540°; 840 m; leg. JG-LM-MS, 2021 (ISU, JG, MS) • N41.8647°, 45.3355°; 1260 m; leg. LM, 2010 (ISU) • N41.8556°, 46.3410°; 1730 m; leg. LM, 2010 (ISU) • N41.8490°, 46.3314°; 1300 m; leg. LM, 2010 (ISU). Kvemo Kartli • N41.7511°, 44.5094°; 1610 m; leg. LM, 2021 (ISU) • N41.6461°, E44.7057°; 1250 m; leg. LM, 2012 (ISU) • N41.3889°, E44.4419°; 880 m; leg. LM, 2013 (ISU). Mtskheta-Mtianeti • N42.3275°, E44.6368°; 1280 m; leg. DM, 2019 (
Clausilia (Mentissa) acuminata
—
Vicinity of Lake Tabatskuri (Samtskhe-Javakheti Region).
The Lesser Caucasus near Bakuriani (Samtskhe-Javakheti Region).
Rocky subalpine meadows; among vegetation and under stones.
Samtskhe-Javakheti • N41.7014°, E43.5131°, 2210 m; leg. JG-LM-MS 2021 (ISU, JG, MS) • N41.6976°, E43.5188°, 2420 m; leg. OD-IS 2019 (IS, MS)
Clausilia duboisi
—
"Tauria" (= Crimea; erroneous!).
Coastal areas of the Black Sea from Sinop (Turkey) to Sochi (Russia), the Western Caucasus, mainly the southern slopes of the Greater and mainly the northern slopes of the Lesser Caucasus. Isolated occurrence range in the Alborz Mountains in Mazandaran Province (Iran).
Forests and rocky subalpine meadows; on trees, under loose bark and stones.
Adjara • N41.4825°, E41.8629°; 540 m; leg. LM, 2017 (ISU). Imereti • N42.3821°, E43.0178°; 1000 m; leg. DM, 2018 (
The reduced, diffuse palatal plicae and the less elongate shell shape distinguish the new species from members of the subgenus Mucronaria (Index) Boettger, 1877 and position it in the subgenus Mucronaria (Mucronaria) Boettger, 1877. Mucronaria (M.) kartvelica sp. n. differs from all other species of its subgenus by the slender and strongly costate shell. By contrast, Mucronaria (M.) acuminata has ventricose shell with glossy, densely striate surface and very weak lamella superior, Mucronaria (M.) duboisi has strong, to the peristome margin extended lamella inferior and long, well outlined palatal plicae, whereas Mucronaria (M.) strauchi (Boettger, 1878) is less elongate, more densely costate, and its ribs are more widely spaced on the neck than on the rest of the last whorl.
Georgia, Racha-Lechkhumi and Kvemo Svaneti Region, at the Kvedi 1 Cave 2.2 km ESE of Kvedi village, N42.5561°, E43.5721°; 1510 m.
Type locality; 14.10.2021; leg. JG-MS, holotype (ISU TM-T003-H); paratypes (
Georgia, Racha-Lechkhumi and Kvemo Svaneti Region, at the Kvedi 2 Cave 2 km ESE of Kvedi village, 42.5561°, E43.5691°; 1560 m; leg. EC-JG-LM-MO-MS 12.09.2022 (ISU, JG, MS).
The brownish-corneous, slender spindle-shaped shell consists of 12.3 to 13.5 whorls. The apical part is elongate, its tip is pointed. The teleochonch whorls have strong and sharp ribs (25 to 28 on the last whorl), which become bulkier but not more widely spaced behind the aperture. The last whorl becomes narrower toward the base, which has a prominent basal crest. The relatively small aperture is oval to somewhat diamond-shaped, its wide peristome is slightly projected. The thin, weakly emerged lamella superior reaches the margin of the peristome. Inward it converges slightly toward the lamella inferior and terminates well before reaching the lamella spiralis, which starts ventrally and ends dorsally. The lamella inferior ends high and deep, its terminal part is only barely visible in apertural view of the shell. Ending even deeper, the lamella subcolumellaris cannot be viewed through the aperture. The plica principalis spans one third of the last whorl starting from the lateral side. The well discernable, dorsolaterally positioned upper plica runs close and parallel to it. Farther toward the base there are two or three lump-like reduced plicae, and a stronger and larger but also diffuse, callus-like basal plica. The broad clausilium plate widens before ending abruptly in a short, pointed tip. In some of the specimens its parietal margin is visible in oblique view through the aperture.
(in mm): Shell height: 12.4–13.9 (holotype 12.9), spire width: 2.8–3.1 (holotype 2.9), aperture height: 2.4–2.6 (holotype 2.5), aperture width: 1.9–2.1 (holotype 2.0).
Unknown.
Live individuals of Mucronaria (M.) kartvelica sp. n. were found on shaded limestone cliffs and among weeds (primarily Hedera helix) growing on those cliffs (Figure
The name of the new species refers to its occurrence in Sa-kartvelo (native historic name for Georgia).
Whereas Mucronaria (M.) duboisi is one of the widest distributed clausiliids in the Caucasus region, reaching as far as Turkey's Sinop Province toward west and northern Iran toward east, all the other species of the subgenus Mucronaria (Mucronaria) are endemic to Georgia.
Clausilia strauchi
—
"Thianetauer Wald" (= Mtianeti, Mtskheta-Mtianeti Region).
Endemic to central Georgia.
Loosely forested areas; among rocks and weeds.
Kvemo Kartli • N41.8125°, E44.9673°; 950 m; leg. MN, 1967 (ISU). Mtskheta-Mtianeti • N41.8624°, E44.6422°; 740 m; leg. LM, 2017 (ISU) • N41.8484°, E44.6379°; 480 m; leg. LM, 2014 (ISU) • N41.8383°, E44.7160°; 520 m; leg. LM, 2017 (ISU). Shida Kartli • N41.9463°, E43.5094°; 740 m; leg. LM, 2014 (ISU) • N41.8492°, E44.2949°; 1040 m; leg. LM, 2010 (ISU) • N41.8424°, E44.1628°; 1370 m; leg. JG-LM-MS, 2021 (MS). Tbilisi • N41.6841°, E44.7923°; 550 m; leg. LM, 2012 (ISU).
Clausilia index
—
"Réduktaleh" (misspelled Redutkaleh, = Kulevi, Samegrelo-Zemo Svaneti Region) and "Chysirkaleh" (unidentified locality southwest of Batumi, Adjara Region).
Coastal areas of the Black Sea between Batumi (Adjara) and Sukhumi (Abkhazia), inland to Borjomi (Samtskhe-Javakheti).
Colchic forests; on or under loose bark of trees, among leaf litter and cliff vegetation.
Adjara • N41.6890°, E41.7055°; 40 m; leg. OD-IS, 2009 (IS, MS) • N41.5663°, E41.5736°; 20 m; leg. LM, 2019 (ISU) • N41.5645°, E41.5703°; 10 m; leg. LM 2020 (ISU). Guria • N41.9081°, E42.1522°; 400 m; leg. LM, 2012 (ISU) • N41.9080°, E42.1529°; 380 m; leg. LM, 2012 (ISU). Imereti • N42.2828°, E42.7583°; 210 m; leg. JG, 2017 (JG, MS) • N42.2620°, E42.9575°; 240 m; leg. JG, 2017 (JG, MS) • N42.0587°, E42.2730°; 90 m; leg. LM 2014 (ISU) • N41.9236°, E42.7494°; 790 m; leg. JG-LM-MS, 2021 (MS). Racha-Lechkhumi and Kvemo Svaneti • N42.4807°, E43.4404°; 1810 m; leg. JG-LM, 2018 (ISU, JG). Samegrelo-Zemo Svaneti • N42.3944°, E41.8382°; 450 m; leg. JG-MS, 2021 (JG, MS) • N42.3112°, E42.0660°; 100 m; leg. EC-JG-MO-MS, 2022 (JG, MS) • N42.2113°, E41.7020°; 10 m; leg. LM, 2007 (ISU) • N42.1765°, E41.9452°; 10 m; leg. LM 2007 (ISU). Samtskhe-Javakheti • N41.7863°, E43.2381°; 980 m; leg. LM, 2007 (ISU).
Clausilia pleuroptychia
—
"Syrien" (erroneous!).
Southern Racha-Lechkhumi and northern Imereti Regions.
Karst areas; in rock crevices and among roots of cliff vegetation.
Imereti • N42.3838°, E43.0118°; 970 m; leg. JG-LM-MS, (ISU, MS) • N42.3821°, E43.0178°; 1000 m; leg. DM, 2018 (
Clausilia (Euxina) dipolauchen
—
Gordi (Imereti Region).
Southeastern Samegrelo-Zemo Svaneti and northwestern Imereti Regions.
Karst areas; in rock crevices and among roots of cliff vegetation.
Imereti • N42.4792°, E42.5440°; 710 m; leg. JG, 2017 (JG, MS) • N42.4558°, E42.5291°; 640 m; leg. LM, 2021 (ISU) • N42.3750°, E42.5966°; 170 m; leg. JG-MS, 2021 (JG, MS) • N42.3295°, E42.6172°; 120 m; leg. LM, 2014 (ISU) • N42.3095°, E42.6697°; 390 m; leg. JG, 2017 (JG, MS). Samegrelo-Zemo Svaneti • N42.7514°, E42.0472°; 470 m; leg. JG-LM, 2018 (ISU, JG, MS) • N42.7410°, E42.0483°; 420 m; leg. JG-LM, 2018 (ISU, JG) • N42.7295°, E42.0923°; 680 m; leg. JG-MS, 2021 (JG, MS) • N42.6262°, E42.4014°; 520 m; leg. LM, 2014 (ISU) • N42.5969°, E42.3322°; 400 m; leg. LM, 2014 (ISU) • N42.4907°, E42.4231°; 340 m; leg. JG, 2017 (JG, MS) • N42.4791°, E42.3903°; 350 m; leg. JG-MS, 2021 (JG, MS) • N42.3832°, E42.2540°; 130 m; leg. LM, 2010 (ISU) • N42.3585°, E42.1997°; 120 m; leg. LM, 2010 (ISU).
Clausilia lederi
—
Suram mountain ridge (Imereti and Shida Kartli Regions).
Between the ranges of the Greater and Lesser Caucasus from the Black Sea to around Telavi toward east.
Forests and rocky subalpine meadows; under leaf litter and bark, among stones and cliff vegetation.
Adjara • N41.6458°, E42.4853°; 1630 m; leg. LM, 2017 (ISU) • N41.5663°, E41.5736°; 20 m; leg. LM, 2019 (ISU) • N41.5652°, E41.5871°; 310 m; leg. LM-TM, 2019 (ISU). Guria • N41.9081°, E42.1522°; 400 m; leg. LM, 2012 (ISU). Imereti • N42.4568°, E42.5991°; 310 m; leg. DM, 2018 (
Clausilia quadriplicata
—
"Radtscha, Rudtscha oder Ructscha" (= Racha, Racha-Lechkhumi and Kvemo Svaneti Region).
Southern slopes of the Greater Caucasus from Racha to the Caspian Sea, as well as the Lesser Caucasus east of Stepanavan (Armenia).
Deciduous forests; on trees, in leaf litter and under stones.
Kakheti • N42.2532°, E45.3321°; 890 m; leg. LM, 2019 (ISU) • N41.8611°, E46.3397°; 1830 m; leg. LM, 2013 (ISU) • N41.8572°, E46.3103°; 800 m; leg. LM, 2013 (ISU) • N41.8559°, E46.3115°; 830 m; leg. LM, 2013 (ISU) • N41.8556°, E46.3410°; 1730 m; leg. LM, 2010 (ISU) • N41.8537°, E46.3181°; 1020 m; leg. LM, 2013 (ISU) • N41.8490°, E46.3314°; 1300 m; leg. LM, 2010 (ISU) • N41.4935°, E46.1009°; 680 m; leg. LM, 2010 (ISU) • N41.4921°, E46.0994°; 710 m; leg, JG-LM-MS, (ISU, JG, MS) • N41.4896°, E46.0976°; 750 m; leg:, DM, (
Clausilia (Euxina) subaggesta
—
Black Sea coast between Sudak and Feodosia (Crimea; only marine flotsam!).
Coastal areas of the Black Sea from Trabzon Province (Turkey) to Adjara. Inland to southernmost Imereti and northernmost Ardahan Province (Turkey).
Broadleaf and mixed forests: on trees, under stones and among vegetation at shaded cliffs.
Adjara • N41.8047°, E41.8843°; 90 m; leg. LM, 2018 (ISU) • N41.7959°, E41.9452°; 210 m; leg. LM, 2018 (ISU) • N41.7627°, E41.9779°; 340 m; leg. LM, 2010 (ISU) • N41.7433°, E42.0843°; 1250 m; leg. LM, 2018 (ISU) • N41.7377°, E41.9839°; 450 m; leg. LM, 2010 (ISU) • N41.7353°, E42.0196°; 720 m; leg. LM, 2008 (ISU) • N41.7351°, E42.0930°; 1180 m; leg. LM, 2010 (ISU) • N41.7269°, E42.0683°; 930 m; leg. LM, 2018 (ISU) • N41.7020°, E41.7211°; 70 m; leg. JG, 2017 (JG) • N41.6984°, E41.7178°; 90 m; leg. LM, 2017 (ISU) • N41.7074°, E41.7759°; 40 m; leg. LM, 2016 (ISU) • N41.6890°, E41.7055°; 40 m; leg. OD-IS, 2009 (IS, MS) • N41.6835°, E41.8889°; 550 m; leg. LM, 2014 (ISU) • N41.6806°, E41.8601°; 250 m; leg. LM, 2014 (ISU) • N41.6769°, E41.8590°; 340 m; leg. LM, 2017 (ISU) • N41.6755°, E41.7069°; 40 m; leg. LM, 2017 (ISU) • N41.6738°, E41.8543°; 450 m; leg. LM, 2014 (ISU) • N41.6713°, E41.6900°; 30 m; leg. JG, 2017 (JG) • N41.6617°, E41.8560°; 860 m; leg. LM, 2014 (ISU) • N41.6527°, E41.7625°; 550 m; leg. LM, 2017 (ISU) • N41.6315°, E42.4139°; 1120 m; leg. LM, 2017 (ISU). Guria • N41.9785°, E42.0714°; 130 m; leg. LM, 2018 (ISU) • N41.9081°, E42.1522°; 400 m; leg. LM, 2012 (ISU) • N41.8914°, E42.3697°; 1490 m; leg. EC-JG-LM-MO-MS, 2022 (MS). Imereti • N41.8837°, E42.7587°; 1420 m; leg. LM, 2013 (ISU).
Upper valley of the Alazani River (Kakheti).
Cliffs in deciduous forests above 1000 m; on cliffs and under stones.
Kakheti • N42.2765°, E45.3524°; 1280 m; leg. JG-LM-MS, 2021 (ISU, JG, MS) • N42.2744°, E45.3517°; 1200 m; leg. DM, 2019 (
This Quadriplicata represents a new species, which was discovered earlier by Bernhard Hausdorf's research team (Hamburg) at a location farther northwest in Kakheti. Its description is expected to be published soon by members of the Hamburg group.
Clausilia taurica
—
"Tauria" (= Crimea; erroneous!).
Southern slopes of the Greater Caucasus from the Black to the Caspian Sea, northeastern part of Artvin Province (Turkey) and northern slopes of the Lesser Caucasus to Mount Kapaz (Azerbaijan) toward southeast. Isolated occurrences in Northern Ossetia and near Pyatigorsk (Russia), as well as in the Zangezur Mountains (Armenia).
Forests and mountain meadows up to the subalpine zone; on tree trunks, under the bark of decaying trees and among stones.
Imereti • N42.5060°, E42.5581°, 880 m; leg. JG-MS, 2021 (JG, MS) • N42.5028°, E42.5579°; 860 m; leg. JG-MS, 2017 (JG) • N42.5023°, E42.5595°; 880 m; leg. JG-LM, 2018 (ISU, JG) • N42.3829°, E43.0122°; 950 m; leg. JG-LM, 2018 (ISU, JG) • N42.3778°, E42.6022°; 170 m; leg. LM, 2018 (ISU) • N42.3750°, E42.5966°; 170 m; leg. JG-MS, 2021 (JG, MS) • N42.2945°, E42.7690°; 410 m; leg. JG, 2017 (JG), LM, 2021 (ISU) • N42.2828°, E42.7583°; 210 m; leg. JG, 2017 (JG) • N42.2716°, E42.8531°; 350 m; leg. LM 2016 (ISU) • N42.1612°, E43.3577°; 830 m; leg. AB-BJ-LM, 2021 (ISU) • N42.1523°, E45.4171°; 580 m; leg. AB-BJ-LM, 2021 (ISU) • N42.0530°, E43.1676°; 350 m; leg. LM, 2014 (ISU) • N42.0529°, E43.1724°; 270 m; leg. LM, 2014 (ISU) • N42.0522°, E43.1628°; 450 m; leg. LM, 2014 (ISU) • N42.0521°, E43.1822°; 310 m; leg. LM 2014, (ISU) • N41.9236°, E42.7494°; 790 m; leg. JG-LM-MS, 2021 (ISU, JG, MS). Kakheti • N42.2765°, E45.3524°; 1280 m; leg. JG-LM-MS, 2021 (ISU, JG, MS) • N42.2532°, E45.3321°; 890 m; leg. LM, 2019 (ISU) • N42.2368°, E45.3386°; 900 m; leg. JG-LM-MS, 2021 (ISU, JG, MS) • N42.2229°, E45.3020°; 810 m; leg. DM, 2019 (
Clausilia (Euxina) reuleauxi
—
Batumi (Adjara Region).
Coastal areas of the Black Sea from Artvin Province (Turkey) to the Guria Region, inland to southwestern Imereti Region.
Humid broadleaf forests; on trees and under the bark of decaying logs or among ground vegetation.
Adjara • N41.7377°, E41.9839°; 450 m; leg. LM, 2010 (ISU) • N41.7351°, E42.0930°; 1180 m; leg. LM, 2010 (ISU) • N41.7020°, E41.7211°; 70 m; leg. JG, 2017 (JG, MS) • N41.6984°, E41.7178°; 90 m; leg. LM, 2017 (ISU) • N41.6797°, E41.8880°; 420 m; leg. LM, 2014 (ISU). Guria • N41.9080°, E42.1529°; 380 m; leg. LM, 2012 (ISU) • N41.8914°, E42.3697°; 1490 m; leg. EC-JG-LM-MO-MS, 2022 (JG, MS). Imereti • N41.9236°, E42.7494°; 790 m; leg. JG-LM-MS, 2021 (MS).
(A) Mucronaria (Index) index, Adjara, near Mtsvane Kontskhi, N41.6890°, E41.7055°. (B) Mucronaria (Index) pleuroptychia, Imereti, near the Motsameta Monastery, N42.2828°, E42.7583°. (C) Quadriplicata dipolauchen, Imereti, Satsiskvilo, N42.4792°, E42.5440°. (D) Quadriplicata lederi, Samtskhe-Javakheti, at the Mtsvane Monastery, N41.8036°, E43.3180°. (E) Quadriplicata quadriplicata, Kakheti, Artsivi Gorge northwest of Dedoplistskaro, N41.4921°, E46.0994°. (F) Quadriplicata subaggesta, Adjara, Makhinjauri northeast of Batumi, N41.6713°, E41.6900°. (G) Quadriplicata sp., Kakheti, Khadori Gorge, N42.2765°, E45.3524°. (H) Scrobifera taurica, Kakheti, valley of the Alazani River near Birkiani, N42.2368°, E45.3386°. (I) Strigileuxina reuleauxi, Imereti, north of Sairme, N41.9236°, E42.7494°. Scale bar: 5 mm
The family Clausiliidae is one of the most diverse groups of terrestrial gastropods with around 1300 species occurring in large parts of Eurasia, Africa and tropical America (
The Phaedusinae constitute an ancient clade that diverged from a common ancestor in southwestern Europe about 50 million years ago (mya). Among its genera occurring in Georgia Pontophaedusa Lindholm, 1924 represents the most basal lineage that is thought to have diverged around 30 mya in the Anatolian region (
Most of the clausiliids in the Caucasus region belong to the Clausiliinae subfamily. Compared to the Phaedusinae, this subfamily of western Eurasia is much more diverse and includes a number of species-rich genera. It has been shown that the subgeneric radiation of the Clausiliinae species is relatively recent, dating to the Pliocene or the second half of the Miocene period (
During the late Miocene and the Pliocene the Greater Caucasus played an important role in the diversification within the Clausiliinae genera. One of the best examples is that of Acrotoma Boettger, 1881, comprising 13 known species, which are all endemic to the western half of the Greater Caucasus (
In the Caucasus region the clausiliids represent one of the best studied families of invertebrates. Despite the recent discovery of Pontophaedusa gregoi and Mucronaria kartvelica sp. n., the diversity, distribution and habitat preferences of the Georgian species are already well understood. Nevertheless, the geographic coverage of species inventories is still far from complete. For some of the clausiliids this information is still partial, leaving room for refining our knowledge on their distribution. For instance, the regions of Imereti and Kakheti are the best studied ones with 95 and 58 georeferenced sampling sites (comprising our data and published locations), representing 13 species in both regions. By contrast, in the cases of the Guria, Shida Kartli and Kvemo Kartli Regions the numbers of the sampling sites are only 12, 13 and 6, which correspond to 11, 6 and 6 species, respectively. Generally, the Pearson’s correlation coefficient between the number of georeferenced sampling sites and species density among the regions is 0.82 (Table
The distribution areas of the common species Scrobifera taurica or Mucronaria duboisi are well documented, whereas those of some others (i.e., Pontophaedusa gregoi or Inobseratella lindholmi) are based merely on a few occurrence records. Such disparities reflect differences in the actual distribution and abundance of the species, but can also be exaggerated by unequal efficiencies of the sampling. For instance, collecting subterranean Pontophaedusa gregoi requires sampling from limestone crevices. Furthermore, its discontinuous, patchy habitat makes finding this species even harder. While perhaps it has a wider distribution area in the karst belt of western Georgia, discovering further localities could be difficult and may take some time. Another example is that of Inobseratella lindholmi, which occurs in virgin forests of beech (Fagus orientalis). The steep slopes of these mountain habitats and the dense undergrowth, dominated by Rhododendron ponticum and Rubus caucasicus, make their accessing rather difficult. For some species biased sampling may explain the absence of historical records and point out the limitations of further information gains. Even so, less studied regions and species should be subjected to further, more targeted research.
Summary statistics of samplings and species densities in the studied regions of Georgia.
Region | Territory (km2) | Sampling sites | km2 per sample | Species density |
Adjara | 2800 | 50 | 58 | 12 |
Guria | 2400 | 12 | 200 | 11 |
Imereti | 6100 | 95 | 64 | 13 |
Kakheti | 11300 | 58 | 198 | 13 |
Kvemo Kartli (including Tbilisi) | 6600 | 6 | 1100 | 6 |
Mtskheta-Mtianeti | 7000 | 23 | 304 | 9 |
Racha-Leckhumi and Kvemo Svaneti | 5100 | 33 | 155 | 11 |
Samegrelo-Zemo Svaneti | 7400 | 38 | 200 | 9 |
Samtskhe-Javakheti | 6300 | 38 | 168 | 10 |
Shida Kartli | 5600 | 13 | 438 | 6 |
The occurrence data of the Georgian clausiliids reveal that the diversity of the species is highest in the western regions of the country and gradually decreases toward east (Table
Of the 30 clausiliid species occurring in our study area 12 (40%) are endemic to Georgia. Only one of these, Mucronaria acuminata, is restricted to the Lesser Caucasus, whereas nine, Acrotoma baryshnikovi, Acrotoma enguriensis, Mucronaria kartvelica sp. n., Mucronaria pleuroptychia, Mucronaria strauchi, Pontophaedusa gregoi, Quadriplicata dipolauchen, Elia tuschetica, and Quadriplicata sp., are confined to localities in the Greater Caucasus. Except for the last three these occur in the karst zone of western Georgia. Further two of the Georgian endemics, Mucronaria index and Quadriplicata lederi, are present in areas of both the Greater and Lesser Caucasus. Among the non-endemic species seven have their centres of distribution in the Pontic region, one in the Hyrcanian area, and further 10 are relatively widespread along the Caucasus ranges. In the uniquely local Clausiliidae fauna of Georgia 28 species are endemic to the Caucasus ecoregion and only the distribution areas of Mucronaria duboisi and Serrulina serrulata extend beyond its borders.
As shown in Figure
Species | Biogeographic element | Distribution* | Habitat preference | Altitude range [m] | Georeferenced records | EOO | AOO [km2] ** |
Acrotoma baryshnikovi | Caucasian | GE | Limestone cliffs | 1100–1600 | 3 | 10 | <15 |
Acrotoma enguriensis | Caucasian | GE | Limestone cliffs | 400–500 | 2 | <5 | <5 |
Armenica unicristata | Caucasian | RE | Cliffs | 700–2300 | 3 | – |
Caspiophaedusa perlucens | Hyrcanian | ERE | Forest (decaying wood) | 600–1800 | 15 | – |
Elia derasa | Caucasian | CRE | Forest (various microhabitats) | 0–2500 | 80 | – |
Elia ossetica | Caucasian | CRE | Forest, rocky meadow | 800–2500 | 16 | – |
Elia somchetica | Caucasian | RE | Forest, rocky meadow | 1300–2500 | 8 | – |
Elia tuschetica | Caucasian | GE | Forest, rocky meadow | 1200–1900 | 6 | <2000 | <200 |
Euxinastra hamata | Pontic | WRE | Forest (various microhabitats) | 0–1200 | 6 | – |
Filosa filosa | Pontic | WRE | Forest (various microhabitats) | 0–1300 | 33 | – |
Inobseratella lindholmi | Pontic | WRE | Forest (tree trunks) | 700–1500 | 2 | – |
Inobseratella monticola | Pontic | WRE | Forest (various microhabitats) | 700–800 | 1 | – |
Mentissoidea rupicola | Caucasian | RE | Forest (various microhabitats) | 0–2200 | 61 | – |
Mucronaria acuminata | Caucasian | GE | Subalpine rocky meadow | 2200–2500 | 2 | <5 | <5 |
Mucronaria duboisi | Caucaso-Anatolian | W | Forest, rocky meadow | 0–2300 | 131 | – |
Mucronaria index | Caucasian | GE | Forest (various microhabitats) | 0–2200 | 27 | ∞ | ∞ |
Mucronaria kartvelica sp. n. | Caucasian | GE | Limestone cliffs | 1500–1600 | 2 | <5 | <5 |
Mucronaria pleuroptychia | Caucasian | GE | Limestone cliffs | 0–2100 | 27 | 2500 | <1000 |
Mucronaria strauchi | Caucasian | GE | Forest, rocky meadow | 400–1400 | 8 | ∞ | <1000 |
Pontophaedusa funiculum | Pontic | WRE | Forest (decaying wood) | 0–400 | 6 | – |
Pontophaedusa gregoi | Caucasian | GE | Subterranean | 200–500 | 3 | <100 | <15 |
Pravispira semilamellata | Caucasian | RE | Forest (decaying wood) | 100–2300 | 48 | – |
Quadriplicata dipolauchen | Caucasian | GE | Limestone cliffs | 100–800 | 14 | <1500 | <1000 |
Quadriplicata lederi | Caucasian | GE | Forest, rocky meadow | 0–2300 | 61 | ∞ | ∞ |
Quadriplicata quadriplicata | Caucasian | ERE | Forest (various microhabitats) | 600–2300 | 18 | ∞ | ∞ |
Quadriplicata subaggesta | Pontic | WRE | Forest (various microhabitats) | 0–1500 | 36 | – |
Quadriplicata sp. | Caucasian | GE | Limestone cliffs | 1200–1300 | 2 | <5 | <5 |
Scrobifera taurica | Caucasian | RE | Forest, rocky meadow | 0–2400 | 124 | – |
Serrulina serrulata | Caucaso-Anatolian | W | Forest (decaying wood) | 0–1900 | 29 | – |
Strigileuxina reuleauxi | Pontic | WRE | Forest (various microhabitats) | 0–1500 | 18 | – |
The Caucasus ecoregion has a relatively long and complex geological history (Popov 2004;
The conservation status has not been assessed for any of the Georgian (or Caucasian) clausiliids and, accordingly, none of these species is protected. Presently conservation is provided only by protected areas, which are supposed to shelter their habitats. However, a study by
The extent of occurrence (EOO) and area of occupancy (AOO) data of the endemic Georgian clausiliids are given in Table
For the protection of natural environments in Georgia the main concern is habitat alteration, which is an ever-increasing problem. One example is the vulnerability of the subterranean limestone crevices that provide habitat for Pontophaedusa gregoi. The entire distribution area of this species is within human settlements with unregulated heavy disturbances, such as grazing, logging, limestone mining and polluting. Due to their limited accessibility the populations of this species are difficult to monitor, thus the only feasible way of protecting them could be the protection of their distribution ranges. Another good example is that of Acrotoma enguriensis, which also has a very narrow distribution range along the main road to Svaneti, a popular tourist area in the Caucasus. Its habitat is under increasing pressure from human disturbance (
The authors are thankful to Robert Cameron, Bernhard Hausdorf, Dávid Murányi, Marco T. Neiber, Beata Pokryszko, Igor Solodovnikov, Dimitry Palatov, Barna Páll-Gergely and Frank Walther for providing material, locality information or technical support to this study, to Valeri Barbakadze, Elizaveta Chertoprud, Gogita Chitaia, Joerg Dreybrodt, Janiko Janashia, Mário Olšavský, Igor Pichkhaia, Conny Straub, Rainer Straub and Tamar Tolordava for their valuable help during the field work, as well as two reviewers for their useful comments. This project was supported by the European Speleological Federation (grant ESP 2021-06 "Biodiversity of Georgian Caves and Karst Areas 2021" to J.G. and M.S.).