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Research Article
The land snail family Clausiliidae (Gastropoda, Pulmonata, Stylommatophora) in Georgia: overview, novel records and a new species
expand article infoLevan Mumladze, Jozef Grego§, Miklós Szekeres|
‡ Ilia State University, Tbilisi, Georgia
§ Unaffiliated, Horná Mičiná, Slovakia
| Institute of Plant Biology, Biological Research Centre HAS, Szeged, Hungary
Open Access

Abstract

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.

Key words

Caucasus, endemisms, land snails, new species, zoogeography

Introduction

The Caucasus ecoregion is a major biodiversity hotspot in western Eurasia (Myers et al. 2000; Zazanashvili et al. 2005). The origin of its rich flora and fauna with a multitude of endemic and relict species can be traced back to the forest refuges of the Colchic and Hyrcanian regions, which preserved their mild and humid climate throughout the entire Pleistocene period (Kikvidze and Oshawa 2001; Wielstra et al. 2013; Tarkhnishvili 2014). The geological and climatic fragmentation of formerly continuous distribution areas also contributed to the species richness of the Caucasus Isthmus. This is well exemplified by the remarkable diversity of land snails, an animal group with low mobility (Walther et al. 2016, 2018; Hausdorf et al. 2018; Neiber et al. 2018, 2022).

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 Pfeiffer (1846), Albert Mousson (1856, 1863, 1876a, 1876b), Oskar Boettger (1877, 1878a, 1878b, 1878c, 1879a, 1879b, 1880a, 1880b, 1881a, 1881b, 1883, 1886, 1887, 1888) and Otto Retowski (1889, 1899) provided the descriptions, and also some distribution records, of most of the Georgian clausiliids. By contrast, the tumultuous first half of the 20th century proved unfavourable for mollusc studies, yielding only few clausiliid-related papers (Rosen 1914; Kokochashvili 1940, 1941; Javelidze 1941). Thereafter a renewed interest in the region's land snails resulted in several further contributions (Likharev and Rammelmeier 1952; Likharev and Lezhava 1961; Likharev 1962; Lezhava 1962, 1964, 2004; Nordsieck 1975, 1976; Natsvlishvili 1967; Sysoev and Schileyko 2009; Pokryszko et al. 2011; Fehér et al 2014; Mumladze et al. 2017; Hausdorf et al. 2018; Mumlaze and Szekeres 2020; Neiber et al. 2021) that considerably increased our knowledge of the Georgian Clausiliidae fauna. As a result, the taxonomic positions and distribution areas of the species, as well as the fauna composition became quite well assessed. However, there are still relatively few well defined occurrence records of the clausiliid taxa, data that could help better identifying and protecting their preferred habitats.

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.

Materials and methods

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 Majoros and Németh (1997), Pokryszko et al. (2011), Fehér et al. (2014), Mumladze et al. (2017) and Neiber et al. (2021).

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 (FMNH), Florida Museum of Natural History, Gainesville (UF), Hungarian Natural History Museum, Budapest (HNHM), Ilia State University, Tbilisi (ISU), Muséum National d'Histoire Naturelle, Paris (MNHN), Naturhistorisches Museum Wien (NHMW), Naturmuseum und Forschungsinstitut Senckenberg, Frankfurt am Main (SMF), Zoological Museum Moscow State University (ZMMU), Zoological Museum Hamburg (ZMH), as well as of Jozef Grego, Banská Bystrica (JG) and Miklós Szekeres, Budapest (MS).

Results

Family Clausiliidae Gray, 1855

Subfamily Phaedusinae Wagner, 1922

Caspiophaedusa perlucens (Boettger, 1877)

Figures 1A, 2A

Clausilia perlucensBoettger 1877: p. 73

Type locality

"Kaukasus".

Distribution

Southeastern areas of the Greater and Lesser Caucasus, as well as the Talysh and Alborz Mountains in Iran.

Habitat

Moist forests; in the texture and under the bark of decaying trees.

Occurrence data

KakhetiN41.9837°, 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).

Dobatia goettingi (Brandt, 1961)

Figure 1B

Serrulina (Serrulina) goettingi — Brandt 1961: p. 18, plate 2 fig. 17

Type locality

Turkey, Bolu Province, vicinity of Lake Abant.

Distribution

Southeastern Bulgaria and northwestern Turkey to Samsun toward east.

Habitat

Humid broadleaf forests; in the trunks and roots of decaying trees.

Remark

This species does not occur in Georgia. An empty shell (ZMMU Lc-29847), collected from sea flotsam at the Black Sea coast of Adjara, was presented erroneously as Serrulinella senghanensis (Germain, 1933) (Egorov 2001; Sysoev and Schileyko 2009). It belongs to Dobatia goettingi, which has also been recorded from coastal flotsam of the Crimean Peninsula (Retowski 1887; Reischütz et al. 2016).

Pontophaedusa funiculum (Mousson, 1856)

Figures 1C, 2B

Clausilia funiculumMousson 1856: p. 396

Type locality

Boztepe (Turkey, suburb of Trabzon).

Distribution

Southeastern and eastern coastal areas of the Black Sea between Trabzon (Turkey) and Tuapse (Russia).

Habitat

Colchic broadleaf forests; in the texture and under the bark of decaying trees.

Occurrence data

AdjaraN41.7074°, 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 Mumladze & Szekeres, 2020

Figures 1D, 2C

Pontophaedusa gregoiMumladze and Szekeres 2020: p. 151, figs 1, 2

Type locality

Katskhi (Imereti Region).

Distribution

Near Katskhi and Zastafoni in central Imereti Region.

Habitat

Karstic broadleaf forests; primarily in subterranean rock crevices.

Occurrence data

ImeretiN42.2836°, E43.2113°; 500 m; leg. JG-LM-MS, 2021 (ISU, NHMUK 20220497, NHMW-MO-113738, JG, MS).

Figure 1. 

(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.

Figure 2. 

Occurrence records of Caspiophaedusa perlucens (A), Pontophaedusa funiculum (B) and Pontophaedusa gregoi (C). Black and white dots corrsepond to novel and published data, respectively.

Pravispira semilamellata (Mousson, 1863)

Figures 1E, 3A

Clausilia semilamellataMousson 1863: p. 396

Type locality

"Réduktaleh" (misspelled Redutkaleh, = Kulevi, Samegrelo-Zemo Svaneti Region).

Distribution

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).

Habitat

Forests up to around 2000 m; in the texture and under the bark of decaying trees.

Occurrence data

AdjaraN41.7959°, 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). ImeretiN42.5028°, 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). KakhetiN42.2368°, 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 KartliN41.7511°, E44.5094°; 1610 m; leg. LM, 2021 (ISU). Mtskheta-MtianetiN41.8484°, E44.6379°; 480 m; leg. LM, 2014 (ISU). Racha-Lechkhumi and Kvemo SvanetiN42.5681°, 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 SvanetiN43.0652°, 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-JavakhetiN41.8092°, E43.3112°; 970 m; leg. LM, 2012 (ISU) • N41.7912°, E43.4665°; 1240 m; leg. LM, 2012 (ISU).

Serrulina serrulata (Pfeiffer, 1847)

Figures 1F, 3B

Clausilia serrulataPfeiffer 1847: p. 71

Type locality

"Tauria" (= Crimea; erroneous!).

Distribution

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.

Habitat

Humid, primarily broadleaf forests up to around 2000 m; in the texture and under the bark of decaying trees.

Occurrence data

AdjaraN41.7477°, 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). ImeretiN42.3829°, E43.0122°; 950 m; leg. JG-LM, 2018 (JG, MS). KakhetiN42.2368°, 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-MtianetiN42.0974°, E44.8236°; 1420 m; leg. LM, 2010 (ISU). Racha-Lechkhumi and Kvemo SvanetiN42.3914°, E42.9823°; 1520 m; leg. JG-LM-MS, 2021 (ISU, JG, MS) • N42.3876°, E42.9748°; 1500 m; leg. LM, 2018 (ISU). Samegrelo-Zemo SvanetiN42.9109°, 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).

Serrulina sieversi (Pfeiffer, 1871)

Figure 1G

Clausilia sieversiPfeiffer 1871: p. 70

Type locality

Lankaran (Azerbaijan).

Distribution

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).

Habitat

Moist broadleaf forests up to 1500 m; in the texture and under the bark of decaying trees.

Remark

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 (ZMMU Lc-246280, leg. N. Bekman, 1961), it was not found during several recent field trips to this region. Thus the presence of S. sieversi in Georgia needs to be confirmed.

Serrulinella senghanensis (Germain, 1933)

Figure 1H

Serrulina senghanensisGermain 1933: p. 391

Type locality

"Siah Senghan" (locality in Iran's Gilan Province).

Distribution

This species is known only from a small area in the Alborz Mountains south of Lahijan (Iran, Gilan Province).

Habitat

Moist broadleaf forests in karst areas; in subterranean rock crevices.

Remark

Reports of this Iranian species from Georgia (Egorov 2001; Sysoev and Schileyko 2009) were based on a marine flotsam specimen of Dobatia goettingi. See remark at that species.

Subfamily Clausiliinae Gray, 1855

Acrotoma (Iliamneme) baryshnikovi Likharev & Schileyko, 2007

Figures 3C, 4A

Acrotoma (Iliamneme) baryshnikoviLikharev and Schileyko 2007: p. 65, figs 1, 2

Type locality

Akhsargina (South Ossetia).

Distribution

Mountain ridge between Kvedi (Racha-Lechkhumi and Kvemo Svaneti Region) and Kvaisi (South Ossetia).

Habitat

Karst areas; on and underneath limestone cliffs.

Occurrence data

Racha-Lechkhumi and Kvemo SvanetiN42.5561°, E43.5721°; 1510 m; leg. JG-MS, 2021 (JG, MS) • N42.5561°, E43.5691°; 1560 m; leg. JG-MS, 2021 (ISU, NHMUK 20220498, NHMW-MO-113739, JG, MS); EC-JG-LM-MO-MS, 2022 (ISU, JG, MS) • N42.5467°, E43.5309°; 1080 m; leg. LM, 2022 (ISU); JG-LM-MS, 2022 (ISU, JG, MS).

Figure 3. 

Occurrence records of Pravispira semilamellata (A), Serrulina serrulata (B) and Acrotoma (Iliamneme) baryshnikovi (C). Symbols are as in Figure 2.

Figure 4. 

(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 Hausdorf, Walther & Neiber, 2018

Figures 4B, 5A

Acrotoma (Iliamneme?) enguriensis — Haudsorf et al. 2018: p. 698, figs 4, 28, 44, 45, 52

Type locality

Mountain ridge between the Enguri and Magana Rivers, north of Jvari (Samegrelo-Zemo Svaneti Region).

Distribution

Known only from the type locality.

Habitat

Karst areas; on and underneath limestone cliffs.

Occurrence data

Samegrelo-Zemo SvanetiN42.7500°, E42.0489°; 470 m; leg. JG-LM, 2018 (ISU, NHMUK 20220499, NHMW-MO-113740 JG, MS) • N42.7410°, E42.0483°; 420 m; leg. JG-MS, 2021 (JG, MS).

Armenica (Armenica) unicristata (Boettger, 1877)

Figures 4C, 5B

Clausilia unicristataBoettger 1877: p. 75

Type locality

"Ekatherinenfeld" (= Bolnisi, Kvemo Kartli Region).

Distribution

Occurs sporadically in southern and southeastern Georgia, northern and southeastern Armenia, as well as northern and western Azerbaijan.

Habitat

Exposed metamorphic or limestone cliffs; around and inside rock crevices.

Occurrence data

KakhetiN41.4896°, E46.0976°; 750 m; leg. DM, 2019 (HNHM 104483) • N41.4888°, E46.0956°; 760 m; leg. JG-LM-MS, 2021 (ISU, JG, MS). Samtskhe-JavakhetiN41.7014°, E43.5131°; 2210 m; leg. JG-LM-MS, 2021 (ISU, JG, MS).

Remarks

The Armenica from Sakavre (Shida Kartli Region) (ISU B27-501), mentioned by Lezhava (1964) as A. gracillima (Retowski, 1889), belongs to this species. Another sample from Akhalsopeli (Kvemo Kartli Region), mentioned in the same paper as Armenica brunnea (Rossmässler, 1939), could not be located but is assumed to be also A. (A.) unicristata. In Lezhava's collection we found a further sample of this species from Gujareti (Samtskhe-Javakheti Region) (ISU 485), a yet unpublished locality.

Armenica (Astrogena) griseofusca (Mousson, 1876)

Figure 4D

Clausilia (Alinda) griseofuscaMousson 1876b: p. 145, plate 5 fig. 3

Type locality

Vicinity of Lake Tabatskuri (Samtskhe-Javakheti Region; possibly erroneous!).

Distribution

All verified localities of this species are in Trabzon and Rize Provinces of Turkey.

Habitat

Broadleaf or mixed forests between 1000 and 1500 m; on tree trunks and small, shaded cliffs.

Remark

The description and a probable syntype (SMF 144161) identify Armenica (Astrogena) griseofusca as the species currently known from northeastern Turkey. However, the type locality around Lake Tabatskuri is far from the distribution area in Turkey and lacks similar habitats. Therefore, and because this species has not been found there ever since, the locality given by Mousson seems to be erroneous.

Elia (Caucasica) ossetica (Mousson, 1863)

Figures 4E, 5C

Clausilia somchetica var. osseticaMousson 1863: p. 399

Type locality

"Koischet" (unidentified site, likely on the northern slopes of the Greater Caucasus near the border between Chechnya and Dagestan, in Russia).

Distribution

Central regions of the Greater Caucasus, as well as the Lesser Caucasus between Borjomi (Samtskhe-Javakheti Region) and Lake Sevan (Armenia).

Habitat

Moist mountain forests and rocky subalpine meadows up to 2500 m; among leaf litter, in rock crevices and under stones.

Occurrence data

Mtskheta-MtianetiN42.6979°, 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). KakhetiN42.2368°, E45.3386°; 900 m; leg. JG-LM-MS, 2021 (ISU, MS) • N42.2229°, E45.3020°; 810 m; leg. DM, 2019 (HNHM 104475) • N41.8759°, E45.3564°; 1080 m; leg. LM, 2010 (ISU). Shida KartliN41.8359°, E44.2441°; 1100 m; leg. LM, 2009 (ISU).

Elia (Caucasica) somchetica (Pfeiffer, 1846)

Figures 4F, 6A

Clausilia somcheticaPfeiffer 1846: p. 94

Type locality

"Somchetia" (= Somkheti).

Distribution

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).

Habitat

Moist mountain forests and rocky subalpine meadows up to 2500 m; among leaf litter, in rock crevices and under stones.

Occurrence data

Samtskhe-JavakhetiN41.6976°, 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).

Remarks

Somkheti, the type locality, is a loosely defined historic region roughly corresponding to the Kvemo Kartli Region and Lori Province in Armenia.

Figure 5. 

Occurrence records of Acrotoma (Iliamneme) enguriensis (A), Armenica (Armenica) unicristata (B) and Elia (Caucasica) ossetica (C). Symbols are as in Figure 2.

Figure 6. 

Occurrence records of Elia (Caucasica) somchetica (A), Elia (C.) tuschetica (B) and Elia (Megaleuxina) derasa (C). Symbols are as in Figure 2.

Elia (Caucasica) tuschetica (Likharev & Lezhava, 1961)

Figures 4G, 6B

Euxina (Euxina) tuscheticaLikharev and Lezhava 1961: p. 473, figs 1–3

Type locality

Kakheti Region, near the village Zemo Omalo, at 1900 m.

Distribution

Mountain Tusheti.

Habitat

Mountain forests and subalpine meadows between 1000 and 2000 m; among leaf litter, in rock crevices and under stones.

Occurrence data

KakhetiN42.2765°, E45.3524°; 1280 m; leg. JG-LM-MS, 2021 (ISU, JG, MS) • N42.2744°, E45.3517°; 1200 m; leg. DM, 2019 (HNHM 104477) • N42.2447°, E45.4987°; 1630 m; leg. DM, 2019 (HNHM 104481) • N42.2422°, E45.4946°; 1530 m; leg. DM, 2019 (HNHM 104480, MS) • N42.2170°, E45.4761°; 1220 m; leg. DM, 2019 (HNHM 104479, MS); LM, 2019 (ISU).

Elia (Megaleuxina) derasa (Mousson, 1863)

Figures 4H, 6C

Clausilia derasaMousson 1863: p. 400

Type localities

"Réduktaleh" (misspelled Redutkaleh, = Kulevi, Samegrelo-Zemo Svaneti Region) and Kutaisi (Imereti Region).

Distribution

Between the ranges of the Greater and Lesser Caucasus from the Black Sea coast to Tbilisi toward east.

Habitat

Cliffs in moist forests; in leaf litter and loose soil under cliff vegetation.

Occurrence data

AdjaraN41.7477°, 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). GuriaN41.8992°, E42.3707°; 1240 m; leg. LM, 2017 (ISU) • N41.8367°, E42.3314°; 2070 m; leg. LM, 2017 (ISU). ImeretiN42.3829°, 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 (HNHM 104376) • N41.8695°, E42.7946°; 1690 m; leg. EC-JG-MO-MS, 2022 (JG, MS) • N41.8554°, E42.7910°; 1920 m; leg. EC-JG-MO-MS, 2022 (JG, MS). KakhetiN41.8759°, E45.3564°; 1080 m; leg. LM, 2010 (ISU) • N41.8647°, 45.3355°; 1260 m; leg. LM, 2010 (ISU). Racha-Lechkhumi and Kvemo SvanetiN42.5561°, E43.5721°; 1510 m; leg. JG-MS, 2021 (JG, MS) • N42.5561°, E43.5691°; 1560 m; leg. JG-MS, 2021 (JG, MS); JG-LM-MS, 2022 (ISU, JG, MS) • N42.5116°, E43.3983°; 1680 m; leg. EC-JG-LM-MO-MS, 2022 (ISU, JG, MS) • N42.5069°, E43.4005°; 1650 m; leg. EC-JG-LM-MO-MS, 2022 (ISU, JG, MS) • N42.4949°, E43.3523°; 1910 m; leg. LM, 2011 (ISU) • N42.4807°, E43.4404°; 1810 m; leg. JG-LM, 2018 (ISU, JG) • N42.4557°, E43.3847°; 2010 m; leg. AB-BJ-LM, 2021 (ISU) • N42.3914°, E42.9823°; 1520 m; leg. JG-MS, 2021 (JG, MS) • N42.3901°, E42.9737°; 1520 m; leg. JG-LM, (ISU, JG). Samegrelo-Zemo SvanetiN43.1116°, E42.7441°; 1690 m; leg. LM, 2015 (ISU) • N43.0249°, E42.7682°; 2440 m; leg. LM, 2018 (ISU) • N42.7563°, E42.5114°; 2280 m; leg. LM, 2014 (ISU) • N42.7528°, E42.5093°; 2050 m; leg. LM, 2014 (ISU) • N42.7493°, E42.5071°; 1850 m; leg. LM, 2014 (ISU) • N42.7367°, E42.5015°; 1600 m; leg. LM, 2015 (ISU) • N42.6599°, E42.4330°; 950 m; leg. LM, 2010 (ISU) • N42.4772°, E42.4593°; 740 m; leg. LM, 2018 (ISU) • N42.3944°, E41.8382°; 450 m; leg. LM, 2010 (ISU); JG-MS, 2021 (JG, MS) • N42.3933°, E41.5617°; 10 m; leg. LM-TM, 2021 (ISU) • N42.3112°, E42.0660°; 100 m; leg. EC-JG-MO-MS, 2022 (JG, MS). Samtskhe-JavakhetiN41.8449°, E43.3748°; 920 m; leg. JG, 2017 (JG) • N41.8096°, E43.3121°; 2230 m; leg. LM, 2012 (ISU) • N41.7968°, E43.4599°; 1150 m; leg. LM, 2012 (ISU) • N41.7922°, E42.8394°; 1600 m; leg. JG-LM-MS, 2021 (ISU, JG, MS) • N41.7863°, E43.2381°; 980 m; leg. LM, 2014 (ISU) • N41.7400°, E43.1572°; 1000 m; leg. LM, 2014 (ISU) • N41.7318°, E43.1361°; 1000 m; leg. LM, 2014 (ISU) • N41.6833°, E42.6537°; 1200 m; leg. LM, 2011 (ISU) • N41.6366°, E42.5797°; 1690 m; leg. LM, 2008 (ISU). Shida KartliN42.1339°, E43.6396°; 860 m; leg. LM, 2010 (ISU) • N42.0678°, E43.7848°; 670 m; leg. LM, 2014 (ISU) • N41.8130°, E44.2300°; 890 m; leg. LM, 2012 (ISU) • N41.7919°, E44.2567°; 1100 m; leg. LM, 2009 (ISU).

Euxinastra (Euxinastra) hamata (Boettger, 1888)

Figures 7A, 8A

Clausilia (Euxinastra) hamataBoettger 1888: p. 152

Type locality

Batumi (Adjara Region).

Distribution

Mountainous areas along the Black Sea coast from Rize Province (Turkey) to the Adjara Region.

Habitat

Colchic deciduous forests; on decaying tree trunks and in leaf litter.

Occurrence data

AdjaraN41.7959°, 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).

Figure 7. 

(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..

Filosa filosa (Mousson, 1863)

Figures 7B, 8B

Clausilia filosaMousson 1863: p. 395

Type locality

"Chysirkaleh" (unidentified locality somewhere southwest of Batumi, Adjara Region).

Distribution

Areas along the Black Sea coast from Trabzon Province (Turkey) to Adjara.

Habitat

Colchic deciduous forests; on decaying tree trunks and in leaf litter.

Occurrence data (Figure 8B): AdjaraN41.7959°, E41.9452°; 210 m; leg. LM, 2018 (ISU) • N41.7730°, E41.9692°; 280 m; leg. LM, 2008 (ISU) • N41.7627°, E41.9779°; 340 m; leg. LM, 2010 (ISU) • N41.7358°, E42.0410°; 1220 m; leg. LM, 2008 (ISU) • N41.7353°, E42.0196°; 720 m; leg. LM, 2008 (ISU) • N41.7020°, E41.7211°; 70 m; leg. JG, 2017 (JG) • N41.6984°, E41.7178°; 90 m; leg. LM, 2017 (ISU) • N41.6890°, E41.7055°; 40 m; leg. OD-IS, 2009 (IS, MS) • N41.6797°, E41.8880°; 450 m; leg. LM, 2014 (ISU) • N41.6769°, E41.8590°; 350 m; leg. LM, 2017 (ISU) • N41.6755°, E41.7069°; 40 m; leg. JG, 2017 (JG) • N41.6738°, E41.8543°; 450 m; leg. LM, 2014 (ISU) • N41.6713°, E41.6900°; 30 m; leg. JG, 2017 (JG) • N41.6699°, E41.8511°; 700 m; leg. LM, 2014 (ISU) • N41.6617°, E41.8560°; 860 m; leg. LM, 2014 (ISU) • N41.6527°, E41.7625°; 550 m; leg. LM, 2017 (ISU) • N41.5660°, E41.6136°; 60 m; leg. LM, 2008 (ISU) • N41.5652°, E41.5871°; 310 m; leg. LM, 2019 (ISU) • N41.5476°, E41.7564°; 80 m; leg. LM, 2017 (ISU) • N41.5150°, E41.7556°; 120 m; leg. LM, 2008 (ISU) • N41.4993°, E41.8398°; 420 m; leg. LM, 2017 (ISU).

Inobseratella lindholmi (Lindholm, 1912)

Figures 7C, 8C

Clausilia lindholmiLindholm 1912: p. 202

Type locality

"Berge Salolet" (= Mount Sallet south of Artvin, Turkey).

Distribution

The Parmak and Karçal Mountains in Artvin Province (Turkey) and the Meshkheti Range in the Guria Region.

Habitat

Deciduous forests above 800 m; on and around live trees.

Occurrence data

GuriaN41.8914°, E42.3697°; 1490, m; leg. EC-JG-LM-MO-MS, 2022 (ISU, JG, MS)

Remarks

The occurrence of I. lindholmi around the above mentioned site in the Guria Region has already been reported by Pokryszko et al. (2011). However, another record in the same publication from the Adjara Region (KIN3: N41.7730°, E41.9692°, 250 m) needs correction because, upon our examination, the specimen on which it had been based proved to be Euxinastra hamata.

Inobseratella monticola (Neubert, 1992)

Figure 7D

Kazancia monticolaNeubert 1992: p. 70, plate 1 fig. 4

Type locality

Çatköy south of Çamlıhemşin (Turkey, Rize Province).

Distribution

This species has been known to occur in the Fırtına Valley southwest of Çamlıhemşin (Turkey, Rize Province). Recently Mumladze et al. (2017) reported two specimens of this species from the valley of the Tsablarastskali Stream near Sairme (Imereti Region).

Habitat

Forested rocky areas; on trees, in vegetation beneath cliffs and under stones.

Mentissoidea rupicola (Mortillet, 1854)

Figures 7E, 9A

Clausilia rupicolaMortillet 1854: p. 13, fig. 7

Type locality

Tortum (Erzurum Province, Turkey; likely erroneous!).

Distribution

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.

Habitat

Broadleaf and mixed forests; on trees and under the bark of decaying logs.

Occurrence data

AdjaraN41.7477°, E42.0996°; 1860 m; leg. LM, 2018 (ISU) • N41.7351°, E42.0930°; 1180 m; leg. LM, 2010 (ISU). ImeretiN42.5023°, 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). KakhetiN42.2765°, 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-MtianetiN42.3275°, E44.6368°; 1280 m; leg. DM, 2019 (HNHM 104470) • N42.3105°, E45.1229°; 1490 m; leg. LM, 2010 (ISU). Racha-Lechkhumi and Kvemo SvanetiN42.5968°, E43.2395°; 770 m; leg. AB-BJ-LM, 2021 (ISU) • N42.5681°, E43.4953°; 1060 m; leg. LM, 2014 (ISU) • N42.5586°, E43.5741°; 1390 m; leg. EC-JG-LM-MO-MS, 2022 (MS) • N42.5561°, E43.5721°; 1510 m; leg. JG-MS, 2021 (JG, MS) • N42.5116°, E43.3983°; 1680 m; leg. EC-JG-LM-MO-MS, 2022 (MS) • N42.4807°, E43.4404°; 1810 m; leg. JG-LM, 2018 (ISU, JG) • N42.4557°, E43.3847°; 2010 m; leg. AB-BJ-LM, 2021 (ISU) • N42.3914°, E42.9823°; 1520 m; leg. JG-MS, 2021 (JG, MS) • N42.3876°, E42.9748°; 1500 m; leg. LM, 2018 (ISU). Samegrelo-Zemo SvanetiN43.0311°, E42.7317°; 1850 m; leg. LM, 2018 (ISU) • N42.9109°, E42.0765°; 740 m; leg. LM, 2010 (ISU) • N42.7493°, E42.5071°; 1850 m; leg. LM, 2014 (ISU) • N42.7103°, E42.1522°; 640 m; leg. LM, 2010 (ISU) • N42.6599°, E42.4330°; 950 m; leg. LM, 2010 (ISU) • N42.4774°, E42.4587°; 700 m; leg. JG, 2019 (JG) • N42.3944°, E41.8382°; 450 m; leg. LM, 2010 (ISU); JG-MS, 2021 (JG, MS). Samtskhe-JavakhetiN41.7968°, E43.4599°; 1150 m; leg. LM, 2012 (ISU) • N41.7922°, E42.8394°; 1600 m; leg. JG-LM-MS, 2021 (ISU, JG, MS) • N41.6833°, E42.6537°; 1200 m; leg. LM, 2011 (ISU) • N41.6366°, E42.5797°; 1690 m; leg. LM, 2008 (ISU).

Mucronaria (Mucronaria) acuminata (Mousson, 1876)

Figures 7F, 9B

Clausilia (Mentissa) acuminataMousson 1876b: p. 144, plate 5 fig. 4

Type locality

Vicinity of Lake Tabatskuri (Samtskhe-Javakheti Region).

Distribution

The Lesser Caucasus near Bakuriani (Samtskhe-Javakheti Region).

Habitat

Rocky subalpine meadows; among vegetation and under stones.

Occurrence data

Samtskhe-JavakhetiN41.7014°, E43.5131°, 2210 m; leg. JG-LM-MS 2021 (ISU, JG, MS) • N41.6976°, E43.5188°, 2420 m; leg. OD-IS 2019 (IS, MS)

Mucronaria (Mucronaria) duboisi (Charpentier, 1852)

Figures 7G, 9C

Clausilia duboisiCharpentier 1852: p. 402

Type locality

"Tauria" (= Crimea; erroneous!).

Distribution

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).

Habitat

Forests and rocky subalpine meadows; on trees, under loose bark and stones.

Occurrence data

AdjaraN41.4825°, E41.8629°; 540 m; leg. LM, 2017 (ISU). ImeretiN42.3821°, E43.0178°; 1000 m; leg. DM, 2018 (HNHM 104386) • N42.3254°, E43.2678°; 630 m; leg. LM, 2021 (ISU) • N42.2945°, E42.7677°; 400 m; leg. JG, 2017 (JG) • N42.2873°, E43.21640°; 620 m; leg. LM, 2011 (ISU) • N42.2836°, E43.2113°; 500 m; leg. JG-LM-MS, 2021 (ISU) • N42.2828°, E42.7583°; 210 m; leg. JG, 2017 (JG) • N42.2771°, E42.7044°; 200 m; leg. JG, 2017 (JG) • N42.2722°, E42.8530°; 380 m; leg. EC-JG-LM-MO-MS, 2022 (ISU, JG, MS) • N42.2716°, E42.8531°; 350 m; leg. LM, 2016 (ISU) • N42.2620°, E42.9575°; 240 m; leg. JG, 2017 (JG) • N42.2443°, E43.2780°; 400 m; leg. AB-BJ-LM, 2021. N42.2357°, E43.3115°; 560 m; leg. EC-JG-LM-MO-MS, 2022 (MS) • N42.1476°, E42.8037°; 130 m; leg. LM, 2012 (ISU) • N42.0634°, E42.2694°; 70 m; leg. EC-JG-LM-MO-MS, 2022 (MS) • N42.0741°, E43.1626°; 260 m; leg. LM, 2014 (ISU) • N42.0558°, E43.1791°; 270 m; leg. LM, 2014 (ISU) • N42.0530°, E43.1676°; 350 m; leg. LM, 2014 (ISU) • N42.0522°, E43.1628°; 450 m; leg. LM, 2014 (ISU) • N42.0521°, E43.1822°; 320 m; leg. LM, 2014 (ISU) • N42.0290°, E42.8303°; 270 m; leg. DM, 2018 (HNHM 104382) • N41.9054°, E42.7442°; 910 m; leg. DM, 2018 (HNHM 104385) • N41.8832°, E42.7556°; 1270 m; leg. DM, 2018 (HNHM 104383) • N41.8695°, E42.7944°; 1680 m; leg. DM, 2018 (HNHM 104384); EC-JG-MO-MS, 2022 (JG, MS) • N41.8554°, E42.7910°; 1920 m; leg. EC-JG-MO-MS, 2022 (JG, MS). KakhetiN42.2532°, E45.3321°; 890 m; leg. LM, 2019 (ISU) • N42.2368°, 45.3386°; 900 m; leg. JG-LM-MS, 2021 (ISU, JG, MS) • N42.2229°, E45.3020°; 810 m; leg. DM, 2019 (HNHM 104476) • N42.1523°, E45.4171°; 580 m; leg. LM, 2019 (ISU) • N42.0765°, E45.8462°; 1400 m; leg. LM, 2010 (ISU) • N41.9837°, 45.8456°; 680 m; leg. LM, 2010 (ISU) • N41.9823°, 45.8540°; 840 m; leg. JG-LM-MS, 2021 (ISU, JG, MS) • N41.9016°, E45.4918°, 750 m; leg. LM, 2019 (ISU) • N41.8998°, E45.4874°; 780 m; leg. DM, 2019 (HNHM 104472) • N41.8556°, E46.3410°; 1730 m; leg. LM, 2010 (ISU) • N41.4935°, E46.10086°; 680 m; leg. LM, 2010 (ISU) • N42.03319°, E45.3774°; 440 m; leg. JG-LM-MS, 2021 (ISU, JG, MS). Kvemo KartliN41.6461°, E44.7057°; 1250 m; leg. LM, 2012 (ISU) • N41.3889°, E44.4419°; 880 m; leg. LM, 2013 (ISU) • N41.2675°, E44.5183°; 880 m; leg. LM, 2017 (ISU). Mtskheta-MtianetiN42.0424°, E44.7249°; 750 m; leg. LM, 2012 (ISU) • N42.0346°, E44.7458°; 640 m; leg. LM, 2012 (ISU) • N41.8625°, E44.6422°; 740 m; leg. LM, 2017 (ISU) • N41.8383°, E44.7160°; 520 m; leg. LM, 2017 (ISU) • N41.8290°, E44.6780°; 650 m; leg. LM, 2015 (ISU). Racha-Lechkhumi and Kvemo SvanetiN43.0652°, E42.4118°; 1250 m; leg. LM, 2010 (ISU) • N42.8389°, E42.8211°; 980 m; leg DM, 2018 (HNHM 104380) • N42.5735°, E43.4569°; 850 m; leg. JG-LM, 2018 (ISU, JG) • N42.5681°, E43.4953°; 1060 m; leg. LM, 2014 (ISU) • N42.5596°, E43.2297°; 680 m; leg. AB-LM, 2021 (ISU) • N42.5586°, E43.5741°; 1390 m; leg. EC-JG-LM-MO-MS, 2022 (ISU, JG, MS) • N42.5561°, E43.5721°; 1510 m; leg. JG-MS, 2021 (JG, MS) • N42.5561°, E43.5691°; 1560 m; leg. JG-MS 2021 (JG, MS), EC-JG-LM-MO-MS, 2022 (ISU, JG, MS) • N42.5466°, E43.5308°; 1070 m; leg. EC-JG-LM-MO-MS, 2022 (ISU, JG, MS) • N42.5340°, E43.1892°; 600 m; leg. JG-LM, 2018 (ISU, JG) • N42.5169°, E43.3116°; 1420 m; leg. LM, 2018 (ISU) • N42.5116°, E43.3983°; 1680 m; leg. EC-JG-LM-MO-MS, 2022 (ISU, JG, MS) • N42.5083°, E43.3493°; 1850 m; leg. LM, 2010 (ISU) • N42.5004°, E43.3959°; 1780 m; leg. EC-JG-LM-MO-MS, 2022 (ISU, MS) • N42.4985°, E43.3885°; 1780 m; leg. EC-JG-LM-MO-MS, 2022 (ISU, MS) • N42.4949°, E43.3523°; 1910 m; leg. LM, 2011 (ISU) • N42.4837°, E43.0505°; 1050 m; leg. JG, (JG). Samegrelo-Zemo SvanetiN43.1116°, E42.7441°; 1690 m; leg. LM, 2015 (ISU) • N42.7103°, E42.1522°; 640 m; leg. LM, 2010 (ISU) • N42.6599°, E42.4330°; 950 m; leg. LM, 2010 (ISU) • N42.6122°, E42.3827°; 470 m; leg. LM, 2014 (ISU) • N42.5969°, E42.3322°; 400 m; leg. LM, 2014 (ISU) • N42.4772°, E42.4593°; 740 m; leg. LM, 2018 (ISU) • N42.3944°, E41.8382°; 450 m; leg. LM, 2010 (ISU), JG-MS, 2021 (JG, MS) • N42.3933°, E41.5617°; 10 m; leg. LM-TM, 2021 (ISU) • N42.3585°, E42.1997°; 120 m; leg. LM, 2010 (ISU) • N42.3112°, E42.0660°; 100 m; leg. EC-JG-MO-MS, 2022 (JG, MS). Samtskhe-JavakhetiN41.9093°, E43.4839°; 790 m; leg. LM, 2014 (ISU) • N41.8957°, E43.5128°; 940 m; leg. LM, 2011 (ISU) • N41.8699°, E43.4185°; 790 m; leg. LM, 2011 (ISU) • N41.8680°, E43.4125°; 790 m; leg. LM, 2011 (ISU) • N41.8449°, E43.3748°; 920 m; leg. JG, 2017 (JG) • N41.8197°, E43.4470°; 10 m; leg. LM, 2012 (ISU) • N41.8096°, E43.3121°; 2230 m; leg. LM, 2012 (ISU) • N41.8092°, E43.3112°; 970 m; leg. LM, 2012 (ISU) • N41.8075°, E43.4696°; 1050 m; leg. LM, 2018 (ISU) • N41.7968°, E43.4599°; 1150 m; leg. LM, 2012 (ISU) • N41.7962°, E42.8445°; 1700 m; leg. JG-LM-MS, 2021 (ISU, JG, MS) • N41.7675°, E43.3199°; 1230 m; leg. LM, 2011 (ISU) • N41.7649°, E43.3284°; 1370 m; leg. LM, 2020 (ISU) • N41.7400°, E43.1572°; 1000 m; leg. LM, 2014 (ISU) • N41.7318°, E43.1361°; 1000 m; leg. LM, 2014 (ISU) • N41.5187°, E43.0397°; 1510 m; leg. LM, 2020 (ISU). Shida KartliN42.1339°, E43.6396°; 860 m; leg. LM, 2010 (ISU) • N42.1048°, E43.6454°; 750 m; leg. LM, 2010 (ISU) • N41.9858°, E44.1096°; 610 m; leg. LM, 2020 (ISU) • N41.9463°, E43.5094°; 740 m; leg. LM, 2014 (ISU) • N41.8490°, E44.1731°; 1320 m; leg. JG-LM-MS, 2021 (ISU, JG, MS) • N41.7919°, E44.2567°; 1100 m; leg. LM, 2010 (ISU).

Mucronaria (Mucronaria) kartvelicasp. n.

Figures 10, 11, 12A

Differential diagnosis

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.

Type locality

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 material

Type locality; 14.10.2021; leg. JG-MS, holotype (ISU TM-T003-H); paratypes (FMNH 392791/1, HNHM 105342/1, MNHN-IM-2014-7037/1, NHMUK 20220496/1, NHMW-MO-113737/1, SMF 368736/1, UF 580908/1, ZMH-MOL-0141473/1, JG/4, MS/3). Georgia, Racha-Lechkhumi and Kvemo Svaneti Region, at the Kvedi 2 Cave, 2 km ESE of Kvedi village, 42.5561°, E43.5691°; 1560 m; 14.10.2021; leg. JG-MS, paratypes (ISU TM-T003-P1/1, SMF 368737/1, JG/15, MS/9).

Further material

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).

Description

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.

Dimensions

(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).

Anatomy

Unknown.

Habitat

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 11A–B), accompanied by Acrotoma (Iliamneme) baryshnikovi, Elia (Megaleuxina) derasa, Mentissoidea rupicola and Mucronaria (M.) duboisi.

Etymology

The name of the new species refers to its occurrence in Sa-kartvelo (native historic name for Georgia).

Remark

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.

Mucronaria (Mucronaria) strauchi (Boettger, 1878)

Figures 7H, 12B

Clausilia strauchiBoettger 1878b: p. 301, plate 10 fig. 6

Type locality

"Thianetauer Wald" (= Mtianeti, Mtskheta-Mtianeti Region).

Distribution

Endemic to central Georgia.

Habitat

Loosely forested areas; among rocks and weeds.

Occurrence data

Kvemo KartliN41.8125°, E44.9673°; 950 m; leg. MN, 1967 (ISU). Mtskheta-MtianetiN41.8624°, 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 KartliN41.9463°, 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). TbilisiN41.6841°, E44.7923°; 550 m; leg. LM, 2012 (ISU).

Figure 8. 

Occurrence records of Euxinastra hamata (A), Filosa filosa (B) and Inobseratella lindholmi (C). Symbols are as in Figure 2.

Figure 9. 

Occurrence records of Mentissoidea rupicola (A), Mucronaria (Mucronaria) acuminata (B) and Mucronaria (M.) duboisi (C). Symbols are as in Figure 2.

Figure 10. 

Mucronaria (Mucronaria) kartvelica sp. n., holotype (ISU TM-T003-H). Scale bar: 3 mm.

Figure 11. 

The habitat (A) and a live individual (B) of Mucronaria (Mucronaria) kartvelica sp. n.

Figure 12. 

Occurrence records of Mucronaria (Mucronaria) kartvelica sp. n. (A), Mucronaria (M.) strauchi (B) and Mucronaria (Index) index (C). Symbols are as in Figure 2.

Mucronaria (Index) index (Mousson, 1863)

Figures 13A, 12C

Clausilia indexMousson 1863: p. 401

Type localities

"Réduktaleh" (misspelled Redutkaleh, = Kulevi, Samegrelo-Zemo Svaneti Region) and "Chysirkaleh" (unidentified locality southwest of Batumi, Adjara Region).

Distribution

Coastal areas of the Black Sea between Batumi (Adjara) and Sukhumi (Abkhazia), inland to Borjomi (Samtskhe-Javakheti).

Habitat

Colchic forests; on or under loose bark of trees, among leaf litter and cliff vegetation.

Occurrence data

AdjaraN41.6890°, 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). GuriaN41.9081°, E42.1522°; 400 m; leg. LM, 2012 (ISU) • N41.9080°, E42.1529°; 380 m; leg. LM, 2012 (ISU). ImeretiN42.2828°, 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 SvanetiN42.4807°, E43.4404°; 1810 m; leg. JG-LM, 2018 (ISU, JG). Samegrelo-Zemo SvanetiN42.3944°, 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-JavakhetiN41.7863°, E43.2381°; 980 m; leg. LM, 2007 (ISU).

Mucronaria (Index) pleuroptychia (Boettger, 1878)

Figures 13B, 14A

Clausilia pleuroptychiaBoettger 1878b: p. 291, plate 10 fig. 1

Type locality

"Syrien" (erroneous!).

Distribution

Southern Racha-Lechkhumi and northern Imereti Regions.

Habitat

Karst areas; in rock crevices and among roots of cliff vegetation.

Occurrence data

ImeretiN42.3838°, E43.0118°; 970 m; leg. JG-LM-MS, (ISU, MS) • N42.3821°, E43.0178°; 1000 m; leg. DM, 2018 (HNHM 104387) • N42.3254°, E43.2678°; 630 m; leg. 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) • N42.2836°, E43.2113°; 500 m; leg. JG-LM-MS, 2021 (ISU, JG, MS) • N42.2833°, E43.2111°; 500 m; leg. LM, 2013 (ISU) • N42.2828°, E42.7583°; 210 m; leg. JG, 2017 (JG, MS) • N42.2719°, E42.7337°; 150 m; leg. LM, 2019 (ISU) • N42.2716°, E42.8531°; 350 m; leg. LM, 2016 (ISU) • N42.2712°, E42.8576°; 420 m; leg. JG, 2017 (JG) • N42.2620°, E42.9575°; 240 m; leg. JG, 2017 (JG, MS) • N42.2598°, E42.9583°; 990 m; leg. JG-LM-MS, 2021 (ISU, JG, MS) • N42.2575°, E42.7102°; 150 m; leg. DM, 2018 (HNHM 104377). Racha-Lechkhumi and Kvemo SvanetiN42.6679°, E42.7730°; 600 m; leg. MM, 2010 (ISU) • N42.5340°, E43.1892°; 600 m; leg. JG-LM, 2018 (ISU, JG-LM) • N42.5004°, E43.3959°; 1780 m; leg. EC-JG-LM-MO-MS, 2022 (ISU, MS) • N42.4985°, E43.3885°; 1780 m; leg. EC-JG-LM-MO-MS, 2022 (ISU, MS) • N42.4557°, E43.3847°; 2010 m; leg. AB-BJ-LM, 2021 (ISU) • N42.3901°, E42.9737°; 1520 m; leg. JG-LM, 2018 (ISU, JG) • N42.3876°, E42.9748°; 1500 m; leg. LM, 2018 (ISU).

Quadriplicata dipolauchen (Boettger, 1881)

Figures 13C, 14B

Clausilia (Euxina) dipolauchenBoettger 1881a: p. 126

Type locality

Gordi (Imereti Region).

Distribution

Southeastern Samegrelo-Zemo Svaneti and northwestern Imereti Regions.

Habitat

Karst areas; in rock crevices and among roots of cliff vegetation.

Occurrence data

ImeretiN42.4792°, 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 SvanetiN42.7514°, 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).

Quadriplicata lederi (Boettger, 1878)

Figures 13D, 14C

Clausilia lederiBoettger 1878a: p. 123

Type locality

Suram mountain ridge (Imereti and Shida Kartli Regions).

Distribution

Between the ranges of the Greater and Lesser Caucasus from the Black Sea to around Telavi toward east.

Habitat

Forests and rocky subalpine meadows; under leaf litter and bark, among stones and cliff vegetation.

Occurrence data

AdjaraN41.6458°, 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). GuriaN41.9081°, E42.1522°; 400 m; leg. LM, 2012 (ISU). ImeretiN42.4568°, E42.5991°; 310 m; leg. DM, 2018 (HNHM 104379) • N42.4560°, E42.5969°; 290 m; leg. DM, 2018 (HNHM 104378) • N42.4528°, E43.3903°; 1870 m; leg. EC-JG-LM-MO-MS, 2022 (MS) • N42.3838°, E43.0118°; 970 m; leg. EC-JG-LM-MO-MS, 2022 (ISU, MS) • N42.3778°, E42.6022°; 170 m; leg. LM, 2018 (ISU) • N42.3095°, E42.6697°; 390 m; leg. JG, 2017 (JG, MS) • N42.2945°, E42.7690°; 410 m; leg. LM, 2021 (ISU) • N42.2771°, E42.7044°; 200 m; leg. JG, 2017 (JG, MS) • N42.2716°, E42.8531°; 350 m; leg. LM, 2016 (ISU) • N42.2443°, E43.2780°; 400 m; leg. AB-BJ-LM, 2021 (ISU) • N42.1476°, E42.8037°, 130 m; leg. LM, 2012 (ISU) • N42.0986°, E43.3846°; 440 m; leg. DM, 2018 (HNHM 104375) • N42.0741°, E43.1626°; 260 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). KakhetiN41.8759°, E45.3564°; 1080 m; leg. LM, 2010 (ISU) • N41.8647°, E45.3355°; 1260 m; leg. LM, 2010 (ISU). Kvemo KartliN41.6461°, E44.7057°; 1250 m; leg. LM, 2012 (ISU). Racha-Lechkhumi and Kvemo SvanetiN42.8402°, E42.8802°; 1070 m; leg. DM, 2018 (HNHM 104381) • N42.5116°, E43.3983°; 1680 m; leg. EC-JG-LM-MO-MS, 2022 (JG, MS) • N42.5542°, E43.5083°; 900 m; leg. LM, 2014 (ISU) • N42.5083°, E43.3493°; 1850 m; leg. LM, 2010 (ISU) • N42.5069°, E43.4005°; 1650 m; leg. EC-JG-LM-MO-MS, 2022 (ISU, JG, MS) • N42.5004°, E43.3959°; 1780 m; leg. EC-JG-LM-MO-MS, 2022 (ISU, MS) • N42.4985°, E43.3885°; 1780 m; leg. EC-JG-LM-MO-MS, 2022 (ISU, MS) • N42.4949°, E43.3523°; 1900 m; leg. LM, 2011 (ISU) • N42.4844°, E43.4295°; 1750 m; leg. LM, 2021 (ISU) • N42.3876°, E42.9748°; 1500 m; leg. LM, 2018 (ISU). Samegrelo-Zemo SvanetiN42.7413°, E42.0485°; 430 m; leg. LM, 2015 (ISU) • N42.7367°, E42.5015°; 1600 m; leg. LM, 2015 (ISU) • N42.6599°, E42.4330°; 950 m; leg. LM, 2010 (ISU) • N42.6262°, E42.4014°; 520 m; leg. LM, 2014 (ISU) • N42.3944°, E41.8382°; 450 m; leg. JG-MS, 2021 (JG, MS). Samtskhe-JavakhetiN41.8957°, E43.5128°; 940 m; leg. LM, 2011 (ISU) • N41.8540°, E43.2392°; 1670 m; leg. LM, 2011 (ISU) • N41.8096°, E43.3121°; 2230 m; leg. LM, 2012 (ISU) • N41.8092°, E43.3112°; 970 m; leg. LM, 2012 (ISU) • N41.7968°, E43.4599°; 1150 m; leg. LM, 2012 (ISU) • N41.7962°, E42.8445°; 1700 m; leg. JG-LM-MS, 2021 (ISU, JG, MS) • N41.7936°, E42.8520°; 1490 m; leg. LM, 2019 (ISU) • N41.7912°, E43.4665°; 1250 m; leg. LM, 2012 (ISU) • N41.7863°, E43.2381°; 980 m; leg. LM, 2014 (ISU) • N41.6651°, E42.8279°; 1080 m; leg. LM, 2017 (ISU). Shida KartliN42.1339°, E43.6396°; 860 m; leg. LM, 2010 (ISU).

Quadriplicata quadriplicata (Schmidt, 1868)

Figures 13E, 15A

Clausilia quadriplicataSchmidt 1868: p. 163

Type locality

"Radtscha, Rudtscha oder Ructscha" (= Racha, Racha-Lechkhumi and Kvemo Svaneti Region).

Distribution

Southern slopes of the Greater Caucasus from Racha to the Caspian Sea, as well as the Lesser Caucasus east of Stepanavan (Armenia).

Habitat

Deciduous forests; on trees, in leaf litter and under stones.

Occurrence data

KakhetiN42.2532°, 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, (HNHM 104482, MS). Mtskheta-MtianetiN42.3105°, E45.1229°; 1490 m; leg. LM, 2010 (ISU).

Quadriplicata subaggesta (Retowski, 1887)

(Figures 13F, 15B)

Clausilia (Euxina) subaggestaRetowski 1887: p. 37

Type locality

Black Sea coast between Sudak and Feodosia (Crimea; only marine flotsam!).

Distribution

Coastal areas of the Black Sea from Trabzon Province (Turkey) to Adjara. Inland to southernmost Imereti and northernmost Ardahan Province (Turkey).

Habitat

Broadleaf and mixed forests: on trees, under stones and among vegetation at shaded cliffs.

Occurrence data

AdjaraN41.8047°, 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). GuriaN41.9785°, 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). ImeretiN41.8837°, E42.7587°; 1420 m; leg. LM, 2013 (ISU).

Quadriplicata sp.

Figures 13G, 15C

Distribution

Upper valley of the Alazani River (Kakheti).

Habitat

Cliffs in deciduous forests above 1000 m; on cliffs and under stones.

Occurrence data

KakhetiN42.2765°, E45.3524°; 1280 m; leg. JG-LM-MS, 2021 (ISU, JG, MS) • N42.2744°, E45.3517°; 1200 m; leg. DM, 2019 (HNHM 104478).

Remark

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.

Scrobifera taurica (Pfeiffer, 1848)

Figures 13H, 16A

Clausilia tauricaPfeiffer 1848: p. 412

Type locality

"Tauria" (= Crimea; erroneous!).

Distribution

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).

Habitat

Forests and mountain meadows up to the subalpine zone; on tree trunks, under the bark of decaying trees and among stones.

Occurrence data

ImeretiN42.5060°, 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). KakhetiN42.2765°, 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 (HNHM 104474, MS) • N42.0765°, E45.8462°; 1400 m; leg. LM, 2010 (ISU) • N42.0539°, E45.5274°; 480 m; leg. LM, 2019 (ISU) • N41.9918°, E45.5779°; 350 m; leg. DM, 2019 (HNHM 104473) • N41.9837°, E45.8456°; 680 m; leg. LM, 2010 (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) • N41.9761°, E45.8413°; 630 m; leg. JG-LM-MS, 2021 (ISU, JG, MS) • N41.9179°, E45.4898°; 660 m; leg. DM, 2019 (HNHM 104471) • N41.9016°, E45.4918°; 750 m; leg. LM, 2019 (ISU) • N41.8759°, E45.3564°; 1080 m; leg. LM, 2010 (ISU) • N41.8741°, E46.3788°; 2350 m; leg. LM, 2013 (ISU) • N41.8647°, E45.3355°; 1260 m; leg. LM, 2010 (ISU) • N41.8556°, E46.3410°; 1730 m; leg. LM, 2010 (ISU) • N41.8490°, E46.3314°; 1300 m; leg. LM, 2010 (ISU) • N41.8114°, E45.1527°; 900 m; leg. JG-LM-MS, 2021 (MS) • N41.4935°, E46.1009°; 680 m; leg. LM, 2010 (ISU) • N41.4921°, E46.0994°; 710 m; leg. JG-LM-MS, 2021 (ISU, JG, MS). Kvemo KartliN41.3745°, E45.0667°; 270 m; leg. LM, 2012 (ISU) • N41.2675°, E44.5183°; 880 m; leg. LM, 2017 (ISU). Mtskheta-MtianetiN42.3275°, E44.6368°; 1280 m; leg. DM, 2019 (HNHM 104469, MS). Racha-Lechkhumi and Kvemo SvanetiN42.5735°, E43.4569°; 850 m; leg. JG-LM, 2018 (ISU, JG) • N42.5681°, E43.4953°; 1060 m; leg. LM, 2014 (ISU) • N42.5466°, E43.5308°; 1070 m; leg. EC-JG-LM-MO-MS, 2022 (ISU, MS) • N42.5169°, E43.3116°; 1430 m; leg. LM, 2018 (ISU) • N42.5116°, E43.3983°; 1680 m; leg. EC-JG-LM-MO-MS, 2022 (MS) • N42.4985°, E43.3885°; 1780 m; leg. EC-JG-LM-MO-MS, 2022 (MS) • N42.4844°, E43.4295°; 1750 m; leg. LM, 2018 (ISU), AB-GJ-LM, 2021 (ISU) • N42.4837°, E43.0505°; 1050 m; leg. JG, 2017 (JG, MS) • N42.4807°, E43.4404°; 1810 m; leg. JG-LM, 2018 (ISU, JG) • N42.4629°, E43.1380°; 860 m; leg. DM, 2018 (HNHM 104388) • N42.3901°, E42.9737°; 1520 m; leg. JG-LM, 2018 (ISU, JG) • N42.3876°, E42.9748°; 1500 m; leg. LM, 2018 (ISU). Samegrelo-Zemo SvanetiN43.0652°, E42.4118°; 1250 m; leg. LM, 2010 (ISU) • N43.0078°, E42.2917°; 830 m; leg. LM, 2015 (ISU) • N42.9109°, E42.0765°; 740 m; leg. LM, 2010 (ISU) • N42.7766°, E42.0540°; 720 m; leg. LM, 2015 (ISU) • N42.7367°, E42.5015°; 1600 m; leg. LM, 2015 (ISU) • N42.6988°, E42.4582°; 1050 m; leg. LM, 2014 (ISU) • N42.6599°, E42.4330°; 950 m; leg. LM, 2010 (ISU) • N42.6495°, E42.2051°; 320 m; leg. LM, 2018 (ISU) • N42.6262°, E42.4014°; 520 m; leg. LM, 2014 (ISU) • N42.6006°, E42.3466°; 450 m; leg. LM, 2015 (ISU) • N42.5969°, E42.3322°; 400 m; leg. LM, 2014 (ISU) • N42.4907°, E42.4231°; 340 m; leg. JG, 2017 (JG) • N42.4774°, E42.4587°; 700 m; leg. JG, 2019 (JG, MS) • N42.4772°, E42.4593°; 740 m; leg. LM, 2018 (ISU) • N42.3944°, E41.8382°; 450 m; leg. JG-MS, 2021 (JG, MS) • N42.3112°, E42.0660°; 100 m; leg. EC-JG-MO-MS, 2022 (MS) • N42.1765°, E41.9452°; 10 m; leg. LM, 2009 (ISU). Samtskhe-JavakhetiN41.8449°, E43.3748°; 920 m; leg. JG, 2017 (JG) • N41.8197°, E43.4470°; 1120 m; leg. LM, 2012 (ISU) • N41.8096°, E43.3121°; 950 m; leg. LM, 2012 (ISU) • N41.8092°, E43.3112°; 970 m; leg. LM, 2012 (ISU) • N41.7968°, E43.4599°; 1150 m; leg. LM, 2012 (ISU) • N41.7863°, E43.2381°; 980 m; leg. LM, 2014 (ISU) • N41.7400°, E43.1572°; 1000 m; leg. LM, 2014 (ISU) • N41.7318°, E43.1361°; 1000 m; leg. LM, 2014 (ISU) • N41.6833°, E42.6537°; 1200 m; leg. LM, 2011 (ISU). Shida KartliN41.8492°, E44.2949°; 1040 m; leg. LM, 2014 (ISU).

Strigileuxina reuleauxi (Boettger, 1887)

Figures 13I, 16B

Clausilia (Euxina) reuleauxiBoettger 1887: p. 55

Type locality

Batumi (Adjara Region).

Distribution

Coastal areas of the Black Sea from Artvin Province (Turkey) to the Guria Region, inland to southwestern Imereti Region.

Habitat

Humid broadleaf forests; on trees and under the bark of decaying logs or among ground vegetation.

Occurrence data

AdjaraN41.7377°, 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). GuriaN41.9080°, E42.1529°; 380 m; leg. LM, 2012 (ISU) • N41.8914°, E42.3697°; 1490 m; leg. EC-JG-LM-MO-MS, 2022 (JG, MS). ImeretiN41.9236°, E42.7494°; 790 m; leg. JG-LM-MS, 2021 (MS).

Figure 13. 

(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

Figure 14. 

Occurrence records of Mucronaria (Index) pleuroptychia sp. n. (A), Quadriplicata dipolauchen (B) and Quadriplicata lederi (C). Symbols are as in Figure 2.

Figure 15. 

Occurrence records of Quadriplicata quadriplicata (A), Quadriplicata subaggesta (B) and Quadriplicata sp. (C). Symbols are as in Figure 2.

Figure 16. 

Occurrence records of Scrobifera taurica sp. n. (A) and Strigileuxina reuleauxi (B). Symbols are as in Figure 2.

Discussion

Origin and establishment of the Georgian Clausiliidae fauna

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 (Nordsieck 2007). Of its seven extant subfamilies only two, the Phaedusinae and Clausiliinae occur in the Caucasus region (Uit de Weerd and Gittenberger 2013; Bouchet et al. 2017).

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 (Uit de Weerd and Gittenberger 2013), from where all Caucasian Phaedusinae may have originated. Based on fossil records, the Caucasus region seems to have played an important role in the local diversification of the Phaedusinae (Likharev 1962; Steklov 1966). However, from the end of the Miocene period the gradually cooling and drying climate of western Eurasia caused a loss of diversity and territorial regression of this subfamily, the species of which usually require permanently humid and frost free habitats. The resulting modern Phaedusinae fauna of the Caucasus comprises relatively widespread species adapted to mountain forest habitats and also ones confined to narrow subterranean refugia along the temperate southeastern coast of the Black Sea.

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 (Uit de Weerd and Gittenberger 2013; Hausdorf and Neiber 2022). The colonization of the Caucasus by the Clausiliinae could have begun soon after the emergence of the Lesser Caucasus as part of the Pontic-Anatolian mountain complex in the middle Miocene, around 15 mya (Popov 2004).

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 (Hausdorf et al. 2018). In this genus the species level diversification could have resulted from the orogenic rearrangement of limestone areas, climate related habitat fragmentation and the colonization of novel, high mountain habitats (Koch et al. 2016; Hausdorf et al. 2018). The species of Mucronaria Boettger, 1877, Micropontica Boettger, 1881 and Quadriplicata Boettger, 1878 also diversified locally. The oldest fossil of Quadriplicata is known from the northwestern part of the Greater Caucasus and dates back to the transition between the Miocene and Pliocene periods (Steklov 1966). But in general, fossil records from the Caucasus region, and especially its Transcaucasian part, are scarce and require further exploration. Nevertheless, the available phylogenetic and paleontological data suggest that the radiation of the Caucasian Clausiliinae was driven primarily by climate transitions and the resulting habitat fragmentation, rather than by orogeny-related speciation scenarios (Hausdorf et al. 2018).

Species inventory

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 1), indicating a potential for future increase in the resolution of species distributions.

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.

Table 1.

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

Distribution and habitat preferences

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 2). Of the 30 known species of the study area 23 are forest-dwellers, and of these only Caspiophaedusa perlucens and Quadriplicata quadriplicata are restricted to the forests of central and eastern Georgia. The rest of the forest species are either endemic to western Georgia or have their centre of distribution along the eastern coast of the Black Sea. Among the seven petrophilic species six are obligate limestone-dwellers and only Armenica unicrsitata shows tolerance for other types of rocks. Of the cliff-dwelling species five occur exclusively in northwestern Georgia, whereas only Quadriplicata sp. is restricted to the eastern part of the country. Two narrow endemics, Mucronaria acuminata and Ponrophaedusa gregoi, are found only in subalpine or subterranean environments, respectively. By contrast, the widespread species Mucronaria duboisi and Scrobifera taurica occur almost over the entire territory of Georgia with high tolerance for different habitats, including anthropogenic ones. An assessment of the distribution and habitat types of the Georgian clausiliids is given in Table 2.

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 17, the clausiliids of Georgia show widely varied patterns of altitudinal distribution (see also: Mumladze et al. 2017). Eight species are present from sea level to the altitude of 2500 m, six are middle to high altitude species, and four are found only in high altitude habitats above 1200 m. The rest of the species inhabit regions of low or middle elevations, below 1700 m. Of the eight narrow endemic species two occur below 800 m, five at altitudes not higher than 2000 m, and only one above 2000 m (Table 2).

Table 2

. Distribution patterns and habitat types of the Georgian clausiliids.

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
Figure 17. 

Elevational distribution ranges of the Georgian clausiliids.

Species conservation and gaps in our knowledge

The Caucasus ecoregion has a relatively long and complex geological history (Popov 2004; Adamia et al. 2011) which, coupled with paleoenvironmental fluctuations, has resulted in a high diversity of landscapes, flora, and fauna. It is among the 36 global biodiversity hotspots, recognized as such because of their high species diversity, high rate of endemism and the vulnerability of their ecosystems as compared to similar-sized regions elsewhere (Myers et al. 2000; Zazanashvili et al. 2005; Noss et al. 2015). In this context terrestrial molluscs are one of the most exemplary groups of animals. In the Caucasus region they show exceptional diversity with over 300 species. Their remarkable average rate of endemism is 70%, but in certain groups it can be above 90% (Sysoev and Schileyko 2009). This is the case with the family of the Clausiliidae, in which this rate is 97% for the Caucasus ecoregion, involving 93% of the Georgian species. This clearly highlights conservation importance of this land snail group. Considering that most of the clausiliid species are habitat-specific, requiring forest or rock environments, they can be considered vulnerable. Notable exceptions are Mucronaria duboisi, Scrobifera taurica and, to a lesser extent, Quadriplicata subaggesta, which are frequently encountered in anthropogenic environments, such as gardens or concrete structures.

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 Mumladze et al. (2014) has shown that the current extent of protected areas in Georgia, corresponding to around 10% of the country’s territory, can at best ensure the protection of 50% of the ranges where the endemic species occur.

The extent of occurrence (EOO) and area of occupancy (AOO) data of the endemic Georgian clausiliids are given in Table 2. These show that in the case of four of these species the AOO, the approximate size of the distribution area, is less than 20 km2. Considering that the distribution ranges of the endemic clausiliids are relatively well documented, all these small range-restricted species might be assessed as endangered, whereas the others, except for Mucronaria index and Quadriplicata lederi, as vulnerable according to criteria of the IUCN (IUCN 2012).

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 (Grego and Mumladze 2019) and no measures have been taken so far to protect this species. Other range restricted clausiliids of Georgia have similarly grim prospects. Therefore, we propose measures ensuring their survival, primarily by means of habitat protection.

Acknowledgements

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.).

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