98 urn:lsid:arphahub.com:pub:3686FA2D-1003-5130-A1A9-6F0A4E63DF4D urn:lsid:zoobank.org:pub:A3B9B45C-7F69-42DF-85B6-B3189A5B43F2 Caucasiana Caucasiana 2667-9809 Pensoft Publishers 10.3897/caucasiana.4.e169350 169350 Checklist Invertebrata Species Inventories Cenozoic Checklist of Invertebrates from the Racha and Khvamli Karst Massifs, Sakartvelo (Georgia) Margalitadze Ana ana.margalitadze.1@iliauni.edu.ge https://orcid.org/0009-0004-4805-7783 1 Kiria Eleonora https://orcid.org/0009-0007-1711-2163 2 Gogshelidze Mariam https://orcid.org/0009-0000-3614-213X 1 Shavadze Lado https://orcid.org/0000-0002-7643-2382 1 Modebadze Naia https://orcid.org/0009-0009-9213-5466 1 Barjadze Shalva https://orcid.org/0000-0001-8992-4987 1 Institute of Zoology, Ilia State University, Tbilisi, Georgia Institute of Zoology, Ilia State University Tbilisi Georgia Institute of zoology, Ilia State University, Tbilisi, Georgia Institute of zoology, Ilia State University Tbilisi Georgia

Corresponding author: Ana Margalitadze (ana.margalitadze.1@iliauni.edu.ge)

Academic editor: Levan Mumladze

 2025 10 12 2025 4 203 229 8DA6C573-CC13-50DF-9DD7-BE71EF32581A 17908393 67243229-C5F7-4F67-A7EE-EB5C5DFCC37B 20 08 2025 03 12 2025 This is an open access article distributed under the terms of the CC0 Public Domain Dedication. https://zoobank.org/67243229-C5F7-4F67-A7EE-EB5C5DFCC37B

The Caucasus region is recognized as a global biodiversity hotspot, hosting a remarkable diversity. Among its unique ecosystems, karst caves in western Georgia provide habitat for highly specialized invertebrate fauna. Despite the ecological significance of these cave systems, biospeleological research in the region remains limited. Based on recent studies, an updated and expanded checklist of invertebrates from the Racha and Khvamli karst massifs is provided. We report 102 species, representing 25 orders, 57 families, and 78 genera, including eight newly recorded taxa for the caves of Racha and Khvamli karst Massifs: Globochthonius satapliaensis, Harpolithobius perplexus, H. spinipes, Lithobius viriatus, Chaetoleptophyllum flexum, Leucogeorgia gioi, Porrhomma convexum and Colchidoniscus kutaissianus. Of these 102 species, 20 are troglobionts, 11 are stygobionts, 45 are troglophiles, two are stygophiles, 13 are trogloxenes, six are accidental records and for five species ecological status is not defined.

 South Caucasus endemism subterranean fauna Diversity of Cave-Dwelling Millipedes (Arthropoda, Myriapoda, Diplopoda) of the Racha-Khvamli Karst Massifs (Grant No. MR-23-140) and by the Mohamed bin Zayed Species Conservation Fund as part of the grant: Conservation of the cave-dwelling invertebrates, cave habitats and education of local communities on the cave conservation in Racha-Lechkhumi karst massifs (242532420). Introduction

The first documented information on the cave invertebrate fauna of the Racha karst massif was provided by Birstein & Lopaschov (1940) in the paper, “Investigation of the cave fauna in the USSR in 1935–1939”. Until the 21st century, most data on cave invertebrates from the Racha massif came from Tsakhi (= Gogolati) Cave (Borutzkii 1950; Birstein 1950; Golovatch 1990), with additional records from a few other caves, such as Melia (= Nikortsminda) Cave (Golovatch 1985), Sakishore Cave, and Tsivtskala I Cave (Kobakhidze 1958). For several decades thereafter, the karst massif was relatively poorly studied, until the late 2010s, when Georgian biospeleologists began collecting new data. Barjadze et al. (2015a, 2015b) published an annotated list of the invertebrates in the caves of the Racha region (Western Georgia), reporting 73 species from 11 caves.

The Khvamli karst massif, located close to the Racha massif (both belonging to the Racha-Lechkhumi and Kvemo Svaneti region), has only recently been investigated biologically. According to the 2009 cadastre, 16 caves were documented speleologically in the Khvamli massif (Tatashidze et al. 2009), although additional caves have been discovered since then, and the number has increased to 20. While seven species were previously recorded from four caves (Barjadze et al. 2019), this article presents 11 species from 5 caves.

Here, we present an updated checklist of the cave-dwelling invertebrates of the Racha and Khvamli karst massifs, including new records and updated distributions of previously reported species.

 Material and methods

This study was conducted by reviewing the existing literature published between 1940 and 2025. Additional data were collected from caves between 2018 and 2024 using hand-collection methods. Specimens were stored in the collections of the Institute of Zoology, Ilia State University, Tbilisi, Georgia (ISUIZ) in pure ethanol.

Collected material was identified by taxonomic specialists. Specimens were sorted and identified using a UNITRON Z650HR Stereo Microscope and an Accu-Scope Exc-350 Light Microscope. A map was created using QGIS version 3.40.6. Graphs were prepared in Microsoft Excel. Ecological groups were assigned based on previous literature (Turbanov et al. 2016a, 2016b; Barjadze et al. 2019; original descriptions for all existing species and personal comunications).

The photos used in the article were provded by Dragan Antić captured with Olympus Stylus Tough TG-6 digital camera.

Study Area

The study area comprises the Racha and Khvamli karst massifs (RKM and KKM respectively), located in the South Caucasus, in north-western Georgia (Tsikarishvili et al. 2010) (Fig. 1; Table 1).

The Racha karst massif lies within the Oni and Ambrolauri municipalities and includes up to 45 documented caves and chasms. The Khvamli karst massif is situated between the Tskhenistskali and Rioni river valleys, within the Tskaltubo and Tsageri municipalities. A total of 16 speleologically studied caves have been recorded there, along with a bedrock formation (Tatashidze et al. 2009).

List of studied caves in Racha and Khvamli massifs with geographic data and recorded species numbers.

Cave Geographic Coordinates Elevation (m a.s.l.) Number of species References
Khvamli Karst massif
Lagernaia I N42.545479, E42.686656 742 1 Antić and Makarov 2016
Tvishi N42.525686, E42.793161 468 2 Tatashidze et al. 2009
Tvishi V N42.528530, E42.794096 398 2 –
Verdzistava I N42.529234, E42.792486 493 1 Tatashidze et al. 2009
Verdzistava II N42.528723, E42.793847 461 5 Tatashidze et al. 2009
Racha Karst massif
Dolabistavi N42.451083, E43.177639 1202 10 Tatashidze et al. 2009
Ekis (=Eckis) N42.540000, E42.890000 414 1 Antić and Reip 2020
Gorgoleti N42.517570, E42.916524 599 2 Grego et al. 2020
Mghvime Gzis Piras N42.379620, E43.040900 1198 3 Tsikarishvili and Bolashvili 2013
Kakhidze N42.520050, E42.912390 585 7 Tatashidze et al. 2009
Kidobana N42.434014, E43.145800 1166 17 Tatashidze et al. 2009
Kvedi I N42.556100, E43.572100 1561 2 Mumladze et al. 2023
Kvedi II N42.556100, E43.569100 1623 4 Mumladze et al. 2023
Melia N42.450980, E43.073470 1136 16 Tatashidze et al. 2009
Muradi N42.395955, E42.978848 1501 1 Asanidze et al. 2017
Racha 2001 N42.398900, E42.989060 1427 1 Tatashidze et al. 2009
RAW020 N42.422690, E43.050090 1182 1 Ravanne 2022
RAW024 N42.397470, E42.993660 1393 1 Ravanne 2022
RAW025 N42.397110, E42.995770 1400 2 Ravanne 2022
Sakavria N42.404300, E43.002900 1474 3 Ravanne 2022
Sakinule N42.463440, E43.067410 1193 8 Tatashidze et al. 2009
Sakishore N42.442342, E43.159486 1165 14 Tatashidze et al. 2009
Shareula I N42.470000, E43.067778 1141 4 Tatashidze et al. 2009
Shareula II N42.471057, E43.063830 1128 1 –
Shkmeri N42.489233, E43.401644 1665 1 Chertoprud et al. 2023
Tsakhi N42.522167, E42.912583 676 20 Tatashidze et al. 2009
Tsivtskala I N42.470948, E43.064330 1166 3 Tatashidze et al. 2009
Tsivtskala II N42.471667, E43.065000 1130 3 Tsikarishvili and Bolashvili 2013
Tskhrajvari I N42.388253, E42.984672 1438 4 Tatashidze et al. 2009
Tskhrajvari II N42.387569, E42.983540 1497 1 Tatashidze et al. 2009
Usholta N42.499270, E43.386690 1817 7 Tatashidze et al. 2009
10.3897/caucasiana.4.e169350.figure1 8E250F8F-B2EC-5C31-811B-56533C02C7E8

Species richness in caves of Racha and Khvamli karst massifs: Lagernaia I (1); Verdzistava I (2); Tvishi (3); Verdzistava II (4); Tvishi V (5); Ekis (6); Kakhidze (7); Tsakhi (8); Gorgoleti (9); Muradi (10); Tskhrajvari II (11); Tskhrajvari I (12); Racha 2001 (13) RAW024 (14); RAW025 (15);Sakavria (16); Mghvime Gzis Piras (17); RAW020 (18); Shareula II (19); Tsivtskala I (20); Tsivtskala II (21); Sakinule (22); Shareula I (23); Melia (24); Kidobana (25); Sakishore (26); Dolabistavi (27); Usholta (28); Shkmeri (29); Kvedi II (30); Kvedi I (31).

https://binary.pensoft.net/fig/1488302 Results

The species list provided here is taxonomically arranged. For each species, the distribution within the study area (specific caves) and within Georgia (Karst massifs) is provided. The boundaries of Karst massifs follows Tatashidze et al. (2009). Ecological groups are delineated based on taxon-specific literature and our own observations.

Phylum <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="phylum">Annelida</tp:taxon-name-part></tp:taxon-name> Lamarck, 1809

Class Clitellata Michaelsen, 1919

Order Arhynchobdellida Blanchard, 1894

Family Erpobdellidae Moore, 1908

Genus Dina Blanchard, 1892

1. <italic>Dina kobakhidzei</italic> Grosser, Maghradze & Barjadze, 2025

Distribution in the study area. Tskhrajvari I Cave (Grosser et al. 2025); endmeic to RKM (Fig. 2).

Ecological group. Stygobiont (Grosser et al. 2025).

10.3897/caucasiana.4.e169350.figures2-7 9A457A47-3FED-5CA2-A989-D1132D033751

2: Live specimen of Dina kobakhidzei Grosser, Maghradze & Barjadze, 2025 from Tskhrajvari I Cave; 3: Live specimen of Inotrechus usholtaensis Maghradze, Lohaj & Faille, 2025 from Usholta Cave; 4: Live specimen of Acanthophorella didi Antić, Margalitadze & Šević, 2025 from Sakinule Cave; 5: Live specimen of Neobisium verae (Lapschoff, 1940) from Sakinule Cave; 6: Live specimen of Plutomurus weinerae Barjadze, Kováč & Parimuchová, 2024 from Usholta Cave; 7: Live specimen of Xiphocaridinella lechkhumensis Marin & Barjadze, 2022 from Verdzistava II Cave (Photo credit: Dragan Antić).

https://binary.pensoft.net/fig/1488303 2. <italic>Dina ratschaensis</italic> Kobakhidze, 1958

Distribution in the study area. Sakishore and Tsivtskala I caves (Grosser et al. 2023); endmeic to RKM.

Ecological group. Stygobiont (Grosser et al. 2023).

 Phylum <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="phylum">Mollusca</tp:taxon-name-part></tp:taxon-name> Linnaeus, 1758

Class Gastropoda Cuvier, 1795

Order Littorinimorpha Golikov & Starobogatov, 1975

Family Hydrobiidae Stimpson, 1865

Genus Hausdorfenia Grego & Mumladze, 2020

3. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hausdorfenia">Hausdorfenia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pseudohauffenia">pseudohauffenia</tp:taxon-name-part></tp:taxon-name></italic> Grego & Mumladze, 2020

Distribution in the study area. Dolabistavi, Kidobana, and Sakishore caves (Grego et al. 2020); endmeic to RKM.

Ecological group. Stygobiont (Grego et al. 2020).

 4. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hausdorfenia">Hausdorfenia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="shareula">shareula</tp:taxon-name-part></tp:taxon-name></italic> Grego & Mumladze, 2020

Distribution in the study area. Shareula I and Tsivtskala II caves (Grego et al. 2020); endmeic to RKM.

Ecological group. Stygobiont (Grego et al. 2020).

Genus Imeretiopsis Grego & Mumladze, 2020

5. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Imeretiopsis">Imeretiopsis</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gorgoleti">gorgoleti</tp:taxon-name-part></tp:taxon-name></italic> Grego & Mumladze 2020

Distribution in the study area. Gorgoleti Cave (Grego et al. 2020); endmeic to RKM.

Ecological group. Stygobiont (Grego et al. 2020).

Genus Sitnikovia Chertoprud, Palatov & Vinarski, 2020

6. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Sitnikovia">Sitnikovia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="ratschuli">ratschuli</tp:taxon-name-part></tp:taxon-name></italic> Chertoprud, Palatov, Vinarski, 2020

Distribution in the study area. Gorgoleti and Sakishore caves (Chertoprud et al. 2020); endmeic to RKM.

Ecological group. Stygobiont (Chertoprud et al. 2020).

Genus Tschernomorica Vinarski & Palatov, 2019

7. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tschernomorica">Tschernomorica</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="kopidophora">kopidophora</tp:taxon-name-part></tp:taxon-name></italic> Chertoprud, Grego & Mumladze, 2023

Distribution in the study area. Shkmeri Cave (Chertoprud et al. 2023).

Distribution in the Georgian Karst massifs. Okriba, Racha, Zemo Imereti plateau (Barjadze et al. 2019).

Ecological group. Stygophile (Chertoprud et al. 2023).

Order Stylommatophora Schmidt, 1855

Family Clausiliidae J. E. Gray, 1855

Genus Acrotoma O. Boettger, 1881

8. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Acrotoma">Acrotoma</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="baryshnikovi">baryshnikovi</tp:taxon-name-part></tp:taxon-name></italic> I. M. Likharev & Schileyko, 2007

Distribution in the study area. Kvedi I Cave (Mumladze et al. 2023).

Distribution in the Georgian Karst massifs. Kudaro, Racha (Barjadze et al. 2019).

Ecological group. Probably a troglophile (Turbanov et al. 2016a).

Genus Mucronaria O. Boettger, 1877

9. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mucronaria">Mucronaria</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="kartvelica">kartvelica</tp:taxon-name-part></tp:taxon-name></italic> Mumladze, Grego & Szekeres, 2023

Distribution in the study area. Kvedi I and Kvedi II caves (Mumladze et al. 2023); endmeic to RKM.

Ecological group. Stygophile (Mumladze et al. 2023).

Family Oxychilidae P. Hesse, 1927 (1879)

Genus Oxychilus Fitzinger, 1833

10. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Oxychilus">Oxychilus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="mingrelicus">mingrelicus</tp:taxon-name-part></tp:taxon-name></italic> koutaisanus (Mousson, 10863)

Distribution in the study area. Kakhidze Cave (Barjadze et al. 2015a, 2015b).

Distribution in the Georgian Karst massifs. Okriba, Racha, Sataplia-Tskaltubo, Zemo Imereti plateau (Barjadze et al. 2015a, 2015b).

Ecological group. Trogloxene (L. Mumladze personal communication).

Genus Schistophallus A. J. Wagner, 1914

11. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Schistophallus">Schistophallus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="sucinacius">sucinacius</tp:taxon-name-part></tp:taxon-name></italic> (O. Boettger, 1883)

Distribution in the study area. Sakishore Cave (Barjadze et al. 2019).

Ecological group. Trogloxene (L. Mumladze personal communication).

 Phylum <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="phylum">Arthropoda</tp:taxon-name-part></tp:taxon-name> Gravenhorst, 1843

Class Arachnida Cuvier, 1812

Order Araneae Clerck, 1757

Family Amaurobiidae Thorell, 1869

Genus Amaurobius C.L. Koch, 1837

12. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Amaurobius">Amaurobius</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="fenestralis">fenestralis</tp:taxon-name-part></tp:taxon-name></italic> (Ström, 1768)

Distribution in the study area. Tsakhi Cave (Barjadze et al. 2015a, 2015b).

Ecological group. Trogloxene. Species of the genus Amaurobius often occur at the cave entrances (Mamolla et al. 2022).

Family Dysderidae C. L. Koch, 1837

Genus Dysdera Latreille, 1804

13. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dysdera">Dysdera</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="anatoliae">anatoliae</tp:taxon-name-part></tp:taxon-name></italic> Deeleman-Reinhold, 1988

Distribution in the study area. Dolabistavi Cave (Deltshev et al. 2023).

Ecological group. Trogloxene (Deltshev et al. 2023).

Family Linyphiidae Blackwall, 1859

Genus Porrhomma Simon, 1884

14. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Porrhomma">Porrhomma</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="convexum">convexum</tp:taxon-name-part></tp:taxon-name></italic> (Westring, 1851)

Material examined. GEORGIA • 1♂; western Georgia, Ambrolauri municipality, RAW025 Cave; 12 May 2025; leg. Sh Barjadze, E Kiria, N Modebadze, L Shavadze.

Distribution in the study area. RAW025 Cave.

Distribution in the Georgian Karst massifs. Racha, Sataplia-Tskaltubo (Deltshev et al. 2023).

Ecological group. Trogloxene (Barjadze et al. 2019).

Family Lycosidae Sundevall, 1833

Genus Pardosa Koch, 1847

15. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pardosa">Pardosa</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hortensis">hortensis</tp:taxon-name-part></tp:taxon-name></italic> (Thorell, 1872)

Distribution in the study area. Kakhidze Cave (Barjadze et al. 2015a, 2015b).

Ecological group. Accidental record (A. Seropian personal communication).

 16. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pardosa">Pardosa</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="monticola">monticola</tp:taxon-name-part></tp:taxon-name></italic> (Clerck, 1757)

Distribution in the study area. Tsakhi Cave (Barjadze et al. 2015a, 2015b).

Ecological group. Accidental record (A. Seropian personal communication).

Family Nesticidae Simon, 1894

Genus Aituaria Esyunin & Efimik, 1998

17. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Aituaria">Aituaria</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="borutzkyi">borutzkyi</tp:taxon-name-part></tp:taxon-name></italic> (Reimoser, 1930)

Material examined. GEORGIA • 1♂, 1♀; western Georgia, Ambrolauri municipality, RAW024 Cave; 12 Jun 2025; leg. Sh Barjadze, E Kiria, N Modebadze, L Shavadze. • 2 Juv; western Georgia, Ambrolauri municipality, Melia Cave; 14 Jun. 2025; leg. E Kiria, N Modebadze. • 1♀, 1 Juv.; western Georgia, Ambrolauri municipality, RAW025 Cave; 12 May 2025; leg. Sh Barjadze, E Kiria, N Modebadze, L Shavadze. • 1 Juv.; western Georgia, Ambrolauri municipality, Tskhrajvari II Cave; 25 Aug 2024; leg. E Maghradze. • 1 Juv.; western Georgia, Ambrolauri municipality, RAW020 Cave; 14 Jun 2025; leg. N Modebadze.

Distribution in the study area. Dolabistavi, Melia, RAW020, RAW024, RAW025, Sakishore, Tsahki, and Tskhrajvari II caves (Barjadze et al. 2019).

Distribution in the Georgian Karst massifs. Askhi, Odishi plain, Okriba, Racha, Sataplia-Tskaltubo, Zemo Imereti plateau (Barjadze et al. 2019).

Ecological group. Troglophile (Nentwig et al. 2025).

Family Pholcidae C. L. Koch, 1850

Genus Holocnemus Simon, 1875

18. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Holocnemus">Holocnemus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pluchei">pluchei</tp:taxon-name-part></tp:taxon-name></italic> (Scopoli, 1763)

Distribution in the study area. Dolabistavi Cave (Deltshev et al. 2023) .

Ecological group. Trogloxene (Barjadze et al. 2019).

Family Pisauridae Simon, 1890

Genus Pisaura Simon, 1885

19. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pisaura">Pisaura</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="mirabilis">mirabilis</tp:taxon-name-part></tp:taxon-name></italic> (Clerck, 1757)

Distribution in the study area. Tsakhi Cave (Barjadze et al. 2015a, 2015b).

Distribution in the Georgian Karst massifs. Odishi plain, Racha. (Barjadze et al. 2015a, 2015b).

Ecological group. Accidental record (A. Seropian personal communication).

Family Theridiidae Sundevall, 1833

Genus Phylloneta Archer, 1950

20. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phylloneta">Phylloneta</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="impressa">impressa</tp:taxon-name-part></tp:taxon-name></italic> (L. Koch, 1881)

Distribution in the study area. Tsakhi Cave (Barjadze et al. 2019).

Ecological group. Accidental record (A. Seropian personal communication).

Genus Theridion Walckenaer, 1805

21. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Theridion">Theridion</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="melanurum">melanurum</tp:taxon-name-part></tp:taxon-name></italic> Hahn, 1831

Distribution in the study area. Tsakhi Cave (Barjadze et al. 2019).

Ecological group. Accidental record (A. Seropian personal communication).

Order Mesostigmata G. Canestrini 1891

Family Veigaiaidae Oudemans, 1939

Genus Veigaia Oudemans, 1905

22. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Veigaia">Veigaia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="planicola">planicola</tp:taxon-name-part></tp:taxon-name></italic> (Berlese, 1892)

Distribution in the study area. Melia Cave (Barjadze et al. 2015a, 2015b).

Ecological group. Not defined.

Order Opiliones Sundevall, 1833

Family Nemastomatidae Simon, 1872

Genus Nemaspela Šilhavý, 1966

23. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Nemaspela">Nemaspela</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="femorecurvata">femorecurvata</tp:taxon-name-part></tp:taxon-name></italic> Martens, 2006

Distribution in the study area. Sakishore Cave (Barjadze et al. 2015a, 2015b).

Distribution in the Georgian Karst massifs. Gumista-Psirtskha, Racha (Modebadze et al. 2023).

Ecological group. Troglobiont (Modebadze et al. 2023).

Family Phalangiidae Latreille, 1802

Genus Rilaena Šilhavý, 1965

24. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Rilaena">Rilaena</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="zakatalica">zakatalica</tp:taxon-name-part></tp:taxon-name></italic> Snegovaya & Chemeris, 2005

Distribution in the study area. Usholta Cave (Barjadze et al. 2019).

Ecological group. Accidental record (own observation).

Order Sarcoptiformes Reuter, 1909

Family Achipteriidae Thor, 1929

Genus Parachipteria Hammen, 1952

25. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Parachipteria">Parachipteria</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="fanzagoi">fanzagoi</tp:taxon-name-part></tp:taxon-name></italic> (Jacot, 1929)

Distribution in the study area. Kidobana Cave (Barjadze et al. 2015a, 2015b).

Ecological group. Troglophile (Turbanov et al. 2016b).

Family Autognetidae Grandjean, 1960

Genus Autogneta Hull, 1916

26. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Autogneta">Autogneta</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="longilamellata">longilamellata</tp:taxon-name-part></tp:taxon-name></italic> (Michael, 1885)

Distribution in the study area. Kidobana Cave (Barjadze et al. 2015a, 2015b).

Ecological group. Troglophile (Turbanov et al. 2016b).

Genus Conchogneta Grandjean, 1963

27. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Conchogneta">Conchogneta</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="dalecarlica">dalecarlica</tp:taxon-name-part></tp:taxon-name></italic> (Forsslund, 1947)

Distribution in the study area. Tsakhi Cave (Barjadze et al. 2015a, 2015b).

Distribution in the Georgian Karst massifs. Okriba, Racha, Zemo Imereti plateau (Barjadze et al. 2015a, 2015b).

Ecological group. Troglophile (Turbanov et al. 2016b).

Family Ceratozetidae Jacot, 1925

Genus Ceratozetes Berlese, 1908

28. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ceratozetes">Ceratozetes</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gracilis">gracilis</tp:taxon-name-part></tp:taxon-name></italic> (Michael, 1884)

Distribution in the study area. Kidobana Cave (Barjadze et al. 2015a, 2015b).

Ecological group. Troglophile (Turbanov et al. 2016b).

Genus Sphaerozetes Berlese, 1885

29. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Sphaerozetes">Sphaerozetes</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="tricuspidatus">tricuspidatus</tp:taxon-name-part></tp:taxon-name></italic> Willmann, 1923

Distribution in the study area. Kidobana Cave (Barjadze et al. 2015a, 2015b).

Distribution in the Georgian Karst massifs. Askhi, Gaucha, Okriba, Racha, Zemo Imereti plateau (Barjadze et al. 2015a, 2015b).

Ecological group. Troglophile (Turbanov et al. 2016b).

Family Chamobatidae Grandjean, 1954

Genus Chamobates Hull, 1916

30. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chamobates">Chamobates</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cuspidatus">cuspidatus</tp:taxon-name-part></tp:taxon-name></italic> (Michael, 1884)

Distribution in the study area. Kidobana and Tsakhi caves (Barjadze et al. 2015a, 2015b).

Ecological group. Troglophile (Turbanov et al. 2016b).

 31. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chamobates">Chamobates</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="voigtsi">voigtsi</tp:taxon-name-part></tp:taxon-name></italic> (Oudemans, 1902)

Distribution in the study area. Kidobana, Tsakhi and Shareula I caves (Barjadze et al. 2015a, 2015b).

Distribution in the Georgian Karst massifs. Askhi, Okriba, Racha, Zemo Imereti massifs (Barjadze et al. 2015a, 2015b).

Ecological group. Troglophile (Turbanov et al. 2016b).

Family Damaeidae Berlese, 1896

Genus Epidamaeus Koch, 1835

32. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epidamaeus">Epidamaeus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pinguis">pinguis</tp:taxon-name-part></tp:taxon-name></italic> (Kulijev, 1967)

Distribution in the study area. Kidobana and Shareula I caves (Barjadze et al. 2015a, 2015b).

Ecological group. Troglophile (Turbanov et al. 2016b).

Genus Metabelba Grandjean, 1936

33. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metabelba">Metabelba</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="flagelliseta">flagelliseta</tp:taxon-name-part></tp:taxon-name></italic> Bulanova-Zachvatkina, 1965

Distribution in the study area. Kidobana Cave (Murvanidze 2014).

Ecological group. Troglophile (Turbanov et al. 2016b).

Family Galumnidae Jacot, 1925

Genus Pilogalumna Grandjean, 1956

34. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pilogalumna">Pilogalumna</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="tenuiclava">tenuiclava</tp:taxon-name-part></tp:taxon-name></italic> (Berlese, 1914)

Distribution in the study area. Kidobana Cave (Barjadze et al. 2015a, 2015b).

Ecological group. Troglophile (Turbanov et al. 2016b).

Family Liacaridae Sellnick, 1928

Genus Dorycranosus Woolley, 1969

35. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dorycranosus">Dorycranosus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="splendens">splendens</tp:taxon-name-part></tp:taxon-name></italic> (Coggi, 1898)

Distribution in the study area. Sakinule Cave (Barjadze et al. 2015a, 2015b).

Ecological group. Troglophile (Turbanov et al. 2016b).

Family Liebstadiidae Balogh & P. Balogh, 1984

Genus Liebstadia Oudemans, 1906

36. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Liebstadia">Liebstadia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="similis">similis</tp:taxon-name-part></tp:taxon-name></italic> (Michael, 1888)

Distribution in the study area. Melia Cave (Barjadze et al. 2015a).

Ecological group. Troglophile (Turbanov et al. 2016b).

Family Microzetidae Grandjean, 1936

Genus Miracarus Kunst, 1959

37. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Miracarus">Miracarus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hurkai">hurkai</tp:taxon-name-part></tp:taxon-name></italic> Kunst, 1959

Distribution in the study area. Tsivtskala I Cave (Barjadze et al. 2019).

Ecological group. Troglophile (Turbanov et al. 2016b).

Family Mycobatidae Grandjean, 1954

Genus Minunthozetes Hull, 1916

38. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minunthozetes">Minunthozetes</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pseudofusiger">pseudofusiger</tp:taxon-name-part></tp:taxon-name></italic> (Schweizer, 1922)

Distribution in the study area. Kidobana Cave (Barjadze et al. 2015a, 2015b).

Distribution in the Georgian Karst massifs. Gaucha, Okriba, Racha, Zemo Imereti plateau (Barjadze et al. 2015a, 2015b).

Ecological group. Troglophile (Turbanov et al. 2016b).

Family Oppiidae Grandjean, 1951

Genus Graptoppia Balogh, 1983

39. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Graptoppia">Graptoppia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="foveolata">foveolata</tp:taxon-name-part></tp:taxon-name></italic> (Paoli, 1908)

Distribution in the study area. Melia Cave (Barjadze et al. 2015a).

Distribution in the Georgian Karst massifs. Racha, Sataplia-Tskaltubo (Barjadze et al. 2015a).

Ecological group. Troglophile (Turbanov et al. 2016b).

Family Oribatulidae Thor, 1929

Genus Phauloppia Berlese, 1908

40. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phauloppia">Phauloppia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lucorum">lucorum</tp:taxon-name-part></tp:taxon-name></italic> (C.L.Koch, 1841)

Distribution in the study area. Sakinule Cave (Barjadze et al. 2015a, 2015b).

Distribution in the Georgian Karst massifs. Okriba, Racha (Barjadze et al. 2015a, 2015b).

Ecological group. Troglophile (Turbanov et al. 2016b).

Genus Zygoribatula Berlese, 1896

41. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Zygoribatula">Zygoribatula</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="frisiae">frisiae</tp:taxon-name-part></tp:taxon-name></italic> (Oudemans, 1900)

Distribution in the study area. Kidobana and Sakinule Caves (Murvanidze 2014).

Ecological group. Troglophile (Turbanov et al. 2016b).

Family Phenopelopides Petrunkevitch, 1955

Genus Eupelops Ewing, 1917

42. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Eupelops">Eupelops</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="acromios">acromios</tp:taxon-name-part></tp:taxon-name></italic> (Hermann, 1804)

Distribution in the study area. Kidobana Cave (Barjadze et al. 2015a, 2015b).

Distribution in the Georgian Karst massifs. Okriba, Racha, Zemo Imereti plateau (Barjadze et al. 2015a, 2015b).

Ecological group. Troglophile (Turbanov et al. 2016b).

 43. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Eupelops">Eupelops</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="torulosus">torulosus</tp:taxon-name-part></tp:taxon-name></italic> (Koch, 1839)

Distribution in the study area. Kakhidze Cave (Barjadze et al. 2015a, 2015b).

Ecological group. Troglophile (Turbanov et al. 2016b).

Family Zetorchestidae Michael, 1898

Genus Zetorchestes Berlese, 1888

44. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Zetorchestes">Zetorchestes</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="falzonii">falzonii</tp:taxon-name-part></tp:taxon-name></italic> Coggi, 1898

Distribution in the study area. Kakhidze Cave (Barjadze et al. 2015a, 2015b).

Ecological group. Troglophile (Turbanov et al. 2016b).

Family Tectocepheidae Grandjean, 1954

Genus Tectocepheus Berlese, 1896

45. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tectocepheus">Tectocepheus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="velatus">velatus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="velatus">velatus</tp:taxon-name-part></tp:taxon-name></italic> (Michael, 1880)

Distribution in the study area. Kakhidze and Tsivtskala I caves (Murvanidze 2014).

Distribution in the Georgian Karst massifs. Askhi, Okriba, Racha, Zemo Imereti plateau (Murvanidze 2014).

Ecological group. Troglophile (Turbanov et al. 2016b).

Family Zetomotrichidae Grandjean, 1954

Genus Ghilarovus Krivolutsky, 1966

46. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ghilarovus">Ghilarovus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="kvavadzei">kvavadzei</tp:taxon-name-part></tp:taxon-name></italic> Murvanidze, 2014

Distribution in the study area. Tsakhi Cave (Murvanidze 2014).

Distribution in the Georgian Karst massifs. Okriba, Racha (Murvanidze 2014).

Ecological group. Troglophile (Turbanov et al. 2016b).

Order Pseudoscorpionida de Geer, 1778

Family Chthoniidae Daday, 1889

Genus Globochthonius Finnegan, 1932

47. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Globochthonius">Globochthonius</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="satapliaensis">satapliaensis</tp:taxon-name-part></tp:taxon-name></italic> (Schawaller & Dashdamirov, 1988)

Material examined. GEORGIA • 1♂; western Georgia, Tsageri municipality, Verdzistava II cave; 23 Jul 2022; leg. E Kiria, L Shavadze. • 2♂; western Georgia, Ambrolauri municipality, Tsakhi Cave; 7 Apr 2024; leg. A Margalitadze, M Gogshelidze.

Distribution in the Study Area. Tsakhi and Verdzistava II caves.

Distribution in the Georgian Karst massifs. Khvamli, Racha, Sataplia-Tskaltubo (Zaragoza et al. 2021).

Ecological group. Troglobiont (Zaragoza et al. 2021).

Family Neobisiidae Chamberlin, 1930

Genus Neobisium Chamberlin, 1930

48. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Neobisium">Neobisium</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="sakishorense">sakishorense</tp:taxon-name-part></tp:taxon-name></italic> Novák & Gogshelidze, 2025

Distribution in the study area. Sakishore Cave (Novák and Gogshelidze 2025); endmeic to RKM.

Ecological group. Troglobiont (Novák and Gogshelidze 2025).

 49. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Neobisium">Neobisium</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="verae">verae</tp:taxon-name-part></tp:taxon-name></italic> (Lapschoff, 1940)

Distribution in the study area. Melia and Tsakhi caves (Barjadze et al. 2015a, 2015b).

Distribution in the Georgian Karst massifs. Racha (Fig. 5), Sataplia-Tskaltubo (Barjadze et al. 2015a, 2015b).

Ecological group. Troglobiont (Turbanov et al. 2016b).

Class Collembola Lubbock 1870

Order Entomobryomorpha Börner 1913

Family Entomobryidae Folsom, 1937

Genus Pseudosinella Schäffer, 1897

50. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pseudosinella">Pseudosinella</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="alba">alba</tp:taxon-name-part></tp:taxon-name></italic> (Packard, 1873)

Distribution in the study area. Kakhidze Cave (Barjadze et al. 2015a, 2015b).

Ecological group. Trogloxene (own observation).

 51. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pseudosinella">Pseudosinella</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="octopunctata">octopunctata</tp:taxon-name-part></tp:taxon-name></italic> Börner, 1901

Distribution in the study area. Kakhidze Cave (Barjadze et al. 2015a, 2015b).

Ecological group. Trogloxene (own observation).

Family Isotomidae Schäffer, 1896

Genus Folsomia Willem, 1902

52. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Folsomia">Folsomia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="candida">candida</tp:taxon-name-part></tp:taxon-name></italic> Willem, 1902

Distribution in the study area. Melia Cave (Barjadze et al. 2015a, 2015b).

Distribution in the Georgian Karst massifs. Chaama, Gumista-Psirtskha, Odishi plain, Sataplia-Tskaltubo, Unknown karst massif in Samegrelo, Racha, Zemo Imereti plateau (Barjadze et al. 2015a, 2015b).

Ecological group. Troglophile (Turbanov et al. 2016b).

 53. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Folsomia">Folsomia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="inoculata">inoculata</tp:taxon-name-part></tp:taxon-name></italic> J. Stach, 1946

Distribution in the study area. Melia Cave (Barjadze et al. 2015a, 2015b).

Ecological group. Troglophile (Turbanov et al. 2016b).

 54. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Folsomia">Folsomia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="spinosa">spinosa</tp:taxon-name-part></tp:taxon-name></italic> Kseneman, 1936

Distribution in the study area. Melia Cave (Barjadze et al. 2015a, 2015b).

Ecological group. Troglophile (Turbanov et al. 2016b).

Genus Lepidocyrtus C. Bourlet, 1839

55. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lepidocyrtus">Lepidocyrtus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lignorum">lignorum</tp:taxon-name-part></tp:taxon-name></italic> (Fabricius, 1793)

Distribution in the study area. Kidobana Cave (Barjadze et al. 2015a, 2015b).

Distribution in the Georgian Karst massifs. Askhi, Okriba, Odishi plain, Racha (Barjadze et al. 2015a, 2015b).

Ecological group. Trogloxene (own observation).

Family Neanuridae Börner, 1901

Genus Neanura A.D. MacGillivray, 1893

56. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Neanura">Neanura</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="muscorum">muscorum</tp:taxon-name-part></tp:taxon-name></italic> (Templeton, 1835)

Distribution in the study area. Melia Cave (Barjadze et al. 2015a, 2015b).

Ecological group. Troglophile (Turbanov et al. 2016b).

Family Tomoceridae Schäffer, 1896

Genus Tomocerus H. Nicolet, 1842

57. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tomocerus">Tomocerus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="vulgaris">vulgaris</tp:taxon-name-part></tp:taxon-name></italic> (Tullberg, 1871)

Distribution in the study area. Kidobana Cave (Barjadze et al. 2015a, 2015b).

Distribution in the Georgian Karst massifs. Odishi plain, Racha, Zemo Imereti plateu (Barjadze et al. 2015a, 2015b).

Ecological group. Possibly a troglophile (Turbanov et al. 2016b).

Genus Plutomurus R.Yosii, 1956

58. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Plutomurus">Plutomurus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="birsteini">birsteini</tp:taxon-name-part></tp:taxon-name></italic> Djanashvili & Barjadze, 2011

Material examined. GEORGIA • 2♂; western Georgia, Ambrolauri municipality, Nikortsminda Sakinule Cave; 1 May 2023, leg. E Maghradze. • 1♂; western Georgia, Oni municipality, Kvedi II Cave; 24 May 2023, leg. M Gogshelidze. • 2♀; western Georgia, Ambrolauri municipality, Mghvime Gzis Piras (a cave by the roadside near the Shaori Reservoir); 22 Jun 2023; leg. L Shavadze. • 2♂; western Georgia, Tkibuli municipality, Tskhrajvari I Cave; 8 Oct 2023; leg. M Gogshelidze.

Distribution in the study area. Mghvime Gzis Piras, Kvedi II, Melia, Sakinule, Sakishore, Shareula II, Tsakhi, Tskhrajvari I and Tvishi caves (Barjadze et al. 2015a, 2015b; present study).

Distribution in the Georgian Karst massifs. Okriba, Khvamli, Racha, Sataplia-Tskaltubo, Zemo Imereti plateau (Barjadze et al. 2015a, 2015b; present study).

Ecological group. Troglophile (own observation).

 59. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Plutomurus">Plutomurus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="weinerae">weinerae</tp:taxon-name-part></tp:taxon-name></italic> Barjadze, Kováč & Parimuchová, 2024

Distribution in the study area. Usholta Cave (Barjadze et al. 2024) (Fig. 6); endemic to RKM.

Ecological group. Troglophile (Barjadze et al. 2019).

Order Poduromorpha Börner 1913

Family Hypogastruridae Börner 1906

Genus Ceratophysella Börner, 1932

60. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ceratophysella">Ceratophysella</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cavicola">cavicola</tp:taxon-name-part></tp:taxon-name></italic> (Börner, 1901)

Distribution in the study area. Melia Cave (Barjadze et al. 2015a, 2015b).

Distribution in the Georgian Karst massifs. Chaama, Racha (Barjadze et al. 2015a, 2015b).

Ecological group. Troglophile with some troglomorphic features (Turbanov et al. 2016 b).

Order Symphypleona Börner, 1901

Family Arrhopalitidae Stach, 1956

Genus Pygmarrhopalites Vargovitsh, 2009

61. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pygmarrhopalites">Pygmarrhopalites</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="principalis">principalis</tp:taxon-name-part></tp:taxon-name></italic> (Stach, 1945)

Distribution in the study area. Melia and Tsakhi Caves (Barjadze and Djanashvili 2008).

Distribution in the Georgian Karst massifs. Okriba, Racha, Sataplia-Tskaltubo, Zemo Imereti plateau (Barjadze et al. 2015a).

Ecological group. Troglophile (Turbanov et al. 2016b).

Family Onychiuridae Lubbock, 1867

Genus Argonychiurus Bagnall, 1949

62. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Argonychiurus">Argonychiurus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="multiocellatus">multiocellatus</tp:taxon-name-part></tp:taxon-name></italic> Djanashvili, Barjadze, Jordana & Burkhardt, 2014

Distribution in the study area. Tvishi Cave (Djanashvili et al. 2014); endmeic to KKM.

Ecological group. Troglobiont (Djanashvili et al. 2014).

Class Chilopoda Latreille, 1816

Order Lithobiomorpha Pocock, 1895

Family Lithobiidae Pocock, 1895

Genus Harpolithobius Verhoeff, 1904

63. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Harpolithobius">Harpolithobius</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="perplexus">perplexus</tp:taxon-name-part></tp:taxon-name></italic> Zalesskaja, 1972

Material examined. GEORGIA • 2♀; western Georgia, Ambrolauri municipality,Sakavria Cave; 26 Aug 2024; leg. E Maghradze.

Distribution in the study area.Sakavria Cave.

Distribution in the Georgian Karst massifs. Racha (Kiria et al. 2023).

Ecological group. Troglophile (Turbanov et al. 2016b).

 64. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Harpolithobius">Harpolithobius</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="spinipes">spinipes</tp:taxon-name-part></tp:taxon-name></italic> Folkmanová, 1958

Material examined. GEORGIA • 1♂; western Georgia, Ambrolauri municipality, Gzispiras Cave; 3 May 2023; leg. M Gogshelidze, L Shavadze.

Distribution in the study area. Mghvime Gzis Piras.

Ecological group. Not defined.

Genus Lithobius Leach, 1814

65. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lithobius">Lithobius</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="coloratus">coloratus</tp:taxon-name-part></tp:taxon-name></italic> Sseliwanoff, 1881

Distribution in the study area. Tsivtskala II Cave (Kiria et al. 2023).

Ecological group. Troglophile (Turbanov et al. 2016b).

 66. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lithobius">Lithobius</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="erythrocephalus">erythrocephalus</tp:taxon-name-part></tp:taxon-name></italic> Koch, 1847

Distribution in the study area. Tsivtskala II Cave (Kiria et al. 2023).

Ecological group. Troglophile (Turbanov et al. 2016b).

 67. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lithobius">Lithobius</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="reconditus">reconditus</tp:taxon-name-part></tp:taxon-name></italic> Zalesskaja, 1972

Distribution in the study area. Dolabistavi and Sakishore Cave (Kiria et al. 2023).

Distribution in the Georgian Karst massifs. Gumista-Psirtskha, Sataplia-Tskaltubo, Racha, Zemo Imereti plateau (Kiria et al. 2023).

Ecological group. Troglophile (Turbanov et al. 2016b).

 68. <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lithobius">Lithobius</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="viriatus">viriatus</tp:taxon-name-part></tp:taxon-name> Sseliwanoff, 1880

Material examined. GEORGIA • 1♀; western Georgia, Oni Municipality, Kvedi II Cave; 2 May 2023; leg. M Gogshelidze, L Shavadze.

Distribution in the study area. Kvedi II Cave.

Ecological group. Not defined.

Order Scutigeromorpha Pocock, 1895

Family Scutigeridae Leach, 1814

Genus Scutigera Lamarck, 1801

69. Scutigera coleoptrata (Linnaeus, 1758)

Material examined. GEORGIA • 1♀; western Georgia, Tsageri Municipality, Tvishi V Cave; 8 April 2024; leg. M Gogshelidze, L Shavadze, A Margalitadze.

Distribution in the study area. Tvishi V Cave.

Distribution in the Georgian Karst massifs. Arabika (Kiria et al. 2023), Khvamli.

Ecological group. Troglophile (Turbanov et al. 2016b).

Order Chordeumatida Pocock, 1894

Family Anthroleucosomatidae Verhoeff, 1899

Genus Acanthophorella Antić & Makarov, 2016

70. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Acanthophorella">Acanthophorella</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="barjadzei">barjadzei</tp:taxon-name-part></tp:taxon-name></italic> Antić & Makarov, 2016

Distribution in the study area. Dolabistavi Cave (Antić & Makarov 2016; Antić et al. 2025); endmeic to RKM.

Ecological group. Troglobiont (Antić & Makarov 2016).

 71. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Acanthophorella">Acanthophorella</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="didi">didi</tp:taxon-name-part></tp:taxon-name></italic> Antić, Margalitadze & Šević, 2025

Material examined. GEORGIA • many; western Georgia, Oni Municipality,Sakavria Cave; 26 Aug 2024; leg. L Mumladze, E Maghradze, Sh Barjadze. 3 Nov 2024; leg: N Modebadze, A. Margalitadze. • 3♂, 3♀ ; western Georgia, Ambrolauri municipality, Racha 2001; 26 Aug 2024; leg. L Mumladze, E Maghradze, Sh Barjadze. • 2♂, 4♀; western Georgia, Ambrolauri municipality, Tskradjvari II (kveda) Cave; 25 Aug 2024; leg. E Maghradze, Sh Barjadze.

Distribution in the study area. Muradi, Nikortsminda, Sakinule (Fig. 4), Tskhrajvari I (all from Antić et al. 2025), Racha 2001,Sakavria, and Tskrajvari II Caves; endemic to RKM.

Ecological group. Troglobiont (Antić et al. 2025).

 72. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Acanthophorella">Acanthophorella</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gaumarjos">gaumarjos</tp:taxon-name-part></tp:taxon-name></italic> Antić, Margalitadze & Šević, 2025

Material examined. GEORGIA • 2♂, 1♀; western Georgia, Ambrolauri municipality,Sakavria Cave; 26 Aug 2024 leg. L Mumladze, E Maghradze, Sh Barjadze.

Distribution in the study area. Mghvime Gzis Piras,Sakavria (Antić et al. 2025); endmeic to RKM.

Ecological group. Troglobiont (Antić et al. 2025).

 73. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Acanthophorella">Acanthophorella</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="valerii">valerii</tp:taxon-name-part></tp:taxon-name></italic> Antić, in <xref ref-type="bibr" rid="B3">Antić et al., 2023</xref>

Distribution in the study area. Usholta Cave (Antić et al. 2023); endemic to RKM.

Ecological group. Troglobiont (Antić et al. 2023).

Genus Georgiosoma Antić & Makarov, 2016

74. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Georgiosoma">Georgiosoma</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="bicornutum">bicornutum</tp:taxon-name-part></tp:taxon-name></italic> Antić & Makarov, 2016

Distribution in the study area. Lagernaia I Cave (Antić and Makarov 2016); endemic to KKM.

Ecological group. Troglobiont (Antić and Makarov 2016).

Genus Ratcheuma Golovatch, 1985

75. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ratcheuma">Ratcheuma</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="excorne">excorne</tp:taxon-name-part></tp:taxon-name></italic> Golovatch, 1985

Distribution in the study area. Melia Cave (Golovatch 1985); endmeic to RKM.

Ecological group. Troglobiont (Golovatch 1985).

Order Glomerida Brandt, 1833

Family Glomeridae Leach, 1816

Genus Trachysphaera Heller, 1858

76. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Trachysphaera">Trachysphaera</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minuta">minuta</tp:taxon-name-part></tp:taxon-name></italic> Golovatch, 1976

Distribution in the study area. Shareula I Cave (Golovatch 1990).

Ecological group. Troglophile (Turbanov et al. 2016b).

 77. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Trachysphaera">Trachysphaera</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="orientalis">orientalis</tp:taxon-name-part></tp:taxon-name></italic> Golovatch, 1976

Distribution in the study area. Tsakhi Cave (Golovatch, 1990).

Distribution in the Georgian Karst massifs. Racha, Zemo Imereti plateau (Golovatch, 1990; Barjadze et al. 2015a, 2015b).

Ecological group. Troglophile (Turbanov et al. 2016b).

Order Julida Brandt, J. F. (1833)

Family Julidae Leach, W. E. (1814)

Genus Chaetoleptophyllum Verhoeff, 1898

78. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetoleptophyllum">Chaetoleptophyllum</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="flexum">flexum</tp:taxon-name-part></tp:taxon-name></italic> Golovatch, 1979

Material examined. GEORGIA • 4♂, 1♀; western Georgia, Ambrolauri Municipality, Kidobana Cave; 3 Nov 2018; leg. G Nebieridze.

Ecological group. Trogloxene (Evsyukov et al. 2020).

Genus Cylindroiulus Verhoeff, 1894

79. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Cylindroiulus">Cylindroiulus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="schestoperovi">schestoperovi</tp:taxon-name-part></tp:taxon-name></italic> Lohmander, 1932

Distribution in the study area. Sakinule Cave (Read 1992).

Ecological group. Trogloxene (Read 1992).

Genus Leucogeorgia Verhoeff, 1930

80. <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucogeorgia">Leucogeorgia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="uncertainty-rank">aff.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lobata">lobata</tp:taxon-name-part></tp:taxon-name> Antić & Reip, 2020

Material examined. GEORGIA • 1♂, 4♀, 4 juv.; western Georgia, Tsageri Municipality, Verdzistava II Cave; 4 Nov 2021; leg: E Magradze, Sh Barjadze.

Distribution in the study area. Ekis (Antić and Reip 2020) and Verdzistava II Caves; endemic to RKM, KKM.

Ecological group. Troglobiont (Antić and Reip 2020).

 81. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucogeorgia">Leucogeorgia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gioi">gioi</tp:taxon-name-part></tp:taxon-name></italic> Antić & Reip, 2020

Material examined. GEORGIA • 1♂, 1♀, 1 juv.; western Georgia, Ambrolauri municipality, Melia Cave; 1 Mar 2024; leg. M Gogshelidze, E Kiria, N Modebadze, L Shavadze and A Margalitadze.

Distribution in the study area. Melia Cave.

Distribution in the Georgian Karst massifs. Zemo Imereti plateau (Antić and Reip 2020) and Racha.

Ecological group. Troglobiont (Antić and Reip 2020).

 82. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucogeorgia">Leucogeorgia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="golovatchi">golovatchi</tp:taxon-name-part></tp:taxon-name></italic> Antić & Reip, 2020

Distribution in the study area. Tsakhi Cave (Antić and Reip 2020); endemic to RKM.

Ecological group. Troglobiont (Antić and Reip 2020).

 83. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucogeorgia">Leucogeorgia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="longipes">longipes</tp:taxon-name-part></tp:taxon-name></italic> Verhoeff, 1930

Distribution in the study area. Dolabistavi, Sakishore and Tskhrajvari I caves (Antić and Reip 2020; Antic et al. 2025).

Distribution in the Georgian Karst massifs. Okriba, Racha, Zemo Imereti plateau (Antić and Reip 2020; Antić et al. 2025).

Ecological group. Troglobiont (Antić and Reip 2020).

Order Polydesmida Pocock, 1887

Family Polydesmidae Leach, 1816

Genus Brachydesmus Heller, 1858

84. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Brachydesmus">Brachydesmus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="kalischewskyi">kalischewskyi</tp:taxon-name-part></tp:taxon-name></italic> Lignau, 1915

Material Examined. GEORGIA • 1♂, 4♀; western Georgia, Ambrolauri municipality, Kidobana Cave; 3 Nov 2018; leg. G Nebieridze. • 1♂; western Georgia, Tsageri municipality, Tvishi V Cave; 17 May 2018; leg. G Nebieridze.

Distribution in the study area. Kidobana, Melia, Sakinule and Tvishi V caves (Golovatch et al. 2016; present study).

Distribution in the Georgian Karst massifs. Kudaro, Racha (Golovatch et al. 2016; Turbanov et al. 2016b) and Khvamli.

Ecological group. Trogloxene (Golovatch et al. 2016).

Genus Polydesmus Latreille, 1802

85. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Polydesmus">Polydesmus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="abchasius">abchasius</tp:taxon-name-part></tp:taxon-name></italic> Attems, 1898

Distribution in the study area. Sakinule Cave (Golovatch et al. 2016).

Distribution in the Georgian Karst massifs. Gumista-Psirtskha, Racha (Golovatch et al. 2016).

Ecological group. Trogloxene (Golovatch et al. 2016).

Class Insecta Linnaeus, 1758

Order Hymenoptera Linnaeus, 1758

Family Formicidae Latreille, 1809

Genus Lasius Fabricius, 1804

86. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lasius">Lasius</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="fuliginosus">fuliginosus</tp:taxon-name-part></tp:taxon-name></italic> Latreille, 1798

Distribution in the study area. Usholta Cave (Barjadze et al. 2019).

Ecological group. Not defined.

Order Lepidoptera Linnaeus, 1758

Family Geometridae Leach, 1815

Genus Triphosa Stephens, 1829

87. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Triphosa">Triphosa</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="dubitata">dubitata</tp:taxon-name-part></tp:taxon-name></italic> (Linnaeus, 1758)

Distribution in the study area. Dolabistavi, Sakishore and Usholta caves (Barjadze et al. 2015a, 2015b).

Ecological group. Troglophile (Turbanov et al. 2016b).

Family Noctuidae Latreille, 1809

Genus Scoliopteryx Germar, 1810

88. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Scoliopteryx">Scoliopteryx</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="libatrix">libatrix</tp:taxon-name-part></tp:taxon-name></italic> (Linnaeus, 1758)

Distribution in the study area. Dolabistavi and Sakishore caves (Barjadze et al. 2015a, 2015b).

Ecological group. Troglophile (Turbanov et al. 2016b).

Order Coleoptera Linnaeus, 1758

Family Carabidae Latreille, 1802

Genus Duvalius Delarouzée, 1859

89. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Duvalius">Duvalius</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gregoi">gregoi</tp:taxon-name-part></tp:taxon-name></italic> Maghradze, Barjadze, Lohaj & Faille, 2025

Distribution in the study area. Kvedi II Cave (Maghradze et al. 2025), endemic to RKM.

Ecological group. Troglobiont (Maghradze et al. 2025).

Genus Inotrechus Dolzhanski & Ljovushkin, 1989

90. <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Inotrechus">Inotrechus</tp:taxon-name-part></tp:taxon-name> injaevae Dolzhanskij et Ljovuschkin, 1989

Distribution in the study area. Sakishore Cave (Barjadze et al. 2015a, 2015b); endmeic to RKM.

Ecological group.Troglobiont (Barjadze et al. 2019).

 91. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Inotrechus">Inotrechus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="usholtaensis">usholtaensis</tp:taxon-name-part></tp:taxon-name></italic> Maghradze, Lohaj & Faille, 2025

Distribution in the study area. Usholta Cave (Maghradze et al. 2025); endemic to RKM (Fig. 3).

Ecological group. Troglobiont (Maghradze et al. 2025).

Genus Laemostenus Bonelli, 1810

92. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Laemostenus">Laemostenus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="ljovushkini">ljovushkini</tp:taxon-name-part></tp:taxon-name></italic> Vereschagina, 1985

Distribution in the study area. Usholta Cave (Barjadze et al. 2019).

Distribution in the Georgian Karst massifs. Okriba, Racha, Sataplia-Tskaltubo (Barjadze et al. 2019).

Ecological group. Not defined.

Genus Troglocimmerites Ljovuschkin, 1970

93. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Troglocimmerites">Troglocimmerites</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="djanaschvilii">djanaschvilii</tp:taxon-name-part></tp:taxon-name></italic> Ljovuschkin, 1970

Distribution in the study area. Melia Cave (Barjadze et al. 2019).

Ecological group. Troglobiont (Barjadze et al. 2019).

Family Staphylinidae Latreille, 1802

Genus Bryaxis Kugelann, 1794

94. <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Bryaxis">Bryaxis</tp:taxon-name-part></tp:taxon-name><italic>hypocritus</italic> Plavilshchikov, 1940

Distribution in the study area. Tsakhi Cave (Barjadze et al. 2015a, 2015b).

Ecological group. Probably Troglophile (Turbanov et al. 2016).

Order Orthoptera Latreille, 1793

Family Rhaphidophoridae Walker, 1869

Genus Dolichopoda Bolívar, 1880

95. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dolichopoda">Dolichopoda</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="euxina">euxina</tp:taxon-name-part></tp:taxon-name></italic> Semenov, 1901

Distribution in the study area. Tsakhi (Barjadze et al. 2015a, 2015b).

Distribution in the Georgian Karst massifs. Arabika, Bach-Otkhara, Chaama, Duripshi plateau, Fanavi, Gumishta-Psirtskha, Okriba, Racha, Sataphlia-Tskaltubo, Tsebelda, Unknown Cave in Gudauta Municipality (Barjadze et al. 2015a, 2015b).

Ecological group. Troglophile (Turbanov et al. 2016b).

Class Malacostraca Latreille, 1802

Order Amphipoda Latreille, 1816

Family Niphargidae Bousfield, 1977

Genus Niphargus Schiödte, 1849

96. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Niphargus">Niphargus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="rachalechkhumensis">rachalechkhumensis</tp:taxon-name-part></tp:taxon-name></italic> Marin, Barjadze, Maghradze & Palatov, 2023

Distribution in the study area. Dolabistavi, Sakishore, and Verdzistava II caves (Marin et al. 2023); endemic to RKM, KKM.

Ecological group. Stygobiont (Marin et al. 2023).

 97. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Niphargus">Niphargus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="tvishiensis">tvishiensis</tp:taxon-name-part></tp:taxon-name></italic> Marin, Barjadze, Maghradze & Palatov, 2023

Distribution in the study area. Verdzistava I Cave (Marin et al. 2023); endemic to KKM.

Ecological group. Stygobiont (Marin et al. 2023).

Order Decapoda Latreille, 1802

Family Atyidae De Haan, 1849

Genus Xiphocaridinella Sadowsky, 1930

98. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Xiphocaridinella">Xiphocaridinella</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lechkhumensis">lechkhumensis</tp:taxon-name-part></tp:taxon-name></italic> Marin & Barjadze, 2022

Distribution in the study area. Verdzistava II Cave (Marin and Barjadze 2022) (Fig. 7); endmeic to KKM.

Ecological group. Stygobiont (Marin and Barjadze 2022).

Order Isopoda Latreille, 1816

Family Ligiidae Leach W.E. 1814

Genus Caucasoligidium Borutzky, 1950

99. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Caucasoligidium">Caucasoligidium</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cavernicola">cavernicola</tp:taxon-name-part></tp:taxon-name></italic> Borutzky, 1950

Distribution in the study area. Tsakhi Cave (Barjadze et al. 2015a, 2015b).

Distribution in the Georgian Karst massifs. Arabika, Chaama, Gumishta-Psirtskha, Tsebelda, Racha, Unknown karst massif in Gudauta Municipality (Barjadze et al. 2015a, 2015b).

Ecological group. Troglophile (Turbanov et al. 2016a).

Family Trichoniscidae G. O. Sars, 1899

Genus Colchidoniscus Borutzky, 1974

100. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Colchidoniscus">Colchidoniscus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="kutaissianus">kutaissianus</tp:taxon-name-part></tp:taxon-name></italic> Borutzky, 1974

Material examined. GEORGIA • 4♂, 2♀; western Georgia, Tsageri municipality, Verdzistava II Cave; 4 Nov 2021; leg. Sh Barjadze, M Gogshelidze, N Modebadze, E Maghradze.

Distribution in the study area. Verdzistava II Cave.

Distribution in the Georgian Karst massifs. Askhi, Sataplia-Tskaltubo (Barjadze et al. 2015b) and Khvamli.

Ecological group. Troglobiont (Turbanov et al. 2016a).

Class Copepoda H. Milne-Edwards, 1840

Order Harpacticoida G. O. Sars, 1903

Family Canthocamptidae G. O. Sars, 1906

Genus Bryocamptus Chappuis, 1929

101. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Bryocamptus">Bryocamptus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="birsteini">birsteini</tp:taxon-name-part></tp:taxon-name></italic> Borutzky, 1940

Distribution in the study area. Tsakhi Cave (Barjadze et al. 2015a, 2015b); endemic to RKM.

Ecological group. Stygobiont (Turbanov et. al 2016a).

 102. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Bryocamptus">Bryocamptus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="innominatus">innominatus</tp:taxon-name-part></tp:taxon-name></italic> Borutzky, 1940

Distribution in the study area. Tsakhi Cave (Barjadze et al. 2015a, 2015b).

Distribution in the Georgian Karst massifs. Fanavi, Racha, Sataplia-Tskaltubo (Barjadze et al. 2015a, 2015b).

Ecological group. Stygobiont (Barjadze et al. 2019).

Additional material examined. There are 12 species that differ from other species observed in the study area, that are not listed above: Caucasogeyeria sp.1 from Verdzistava I and Verdzistava II caves; Nemaspela sp. 1 from Melia and Sakinule caves and Nemaspela sp. 2 from Sakavria Cave; Plutomurus sp. 1 from Tvishi II Cave, Plutomurus sp. 2 from Verdzistava II Cave and Plutomurus sp. 3 from caves of Khvamli I, Khvamli II, and Khvamli III; Caucasoligidium sp. 1 from Khvamli III; Caucasoligidium sp. 2 from Khvamli I, Khvamli II andSakavria caves; Ligidium sp. 6 from Tekenteri Chasm; Ligidium sp. 7 from Kvedi II Cave; Trachelipus sp. 1 from Chkondidi Cave; Trichoniscus sp. 1 from Melia Cave.

 Discussion

Here we presented data derived from 30 caves, some of which (Sakavria, Mghvime Gzis Piras, etc.) are not included in the latest cadastre of Georgian caves (Tatashidze et al. 2009) because they were discovered later. Among the five studied caves in the Khvamli massif, the highest species number (five) was recorded in Verdzistava II Cave. In contrast, 25 caves have been surveyed in the Racha massif, where Tsakhi Cave is populated with the highest number of species — 20 in total (Table 1).

According to current data, 102 species have been recorded in the Racha and Khvamli karst caves (instead of previously known 94 species). These species belong to 10 classes, 25 orders, 57 families, and 78 genera. Out of 95 and 11 species recorded in caves of the Racha and Khvamli karst massifs, respectively, only five species are shared, including two local endemics: Niphargus rachalechkhumensis and Leucogeorgia aff. lobata. In total, 20 local endemics are recorded.

Among the studied caves, no invertebrates were reported fromSakavria Cave prior to our study. Thus, the record of two millipede species, Acanthophorella gaumarjos and A. didi (an example of syntopy), Harpolithobius perplexus, and several other unidentified taxa are the first invertebrate records forSakavria Cave.

The discovery of a few taxa resulted in a significant increase of known distribution ranges for a number of species previously recorded in other areas. For example, Globochthonius satapliaensis, known from the caves of the Sataplia-Tskaltubo karst massif, is now recorded in both studied massifs. Harpolithobius perplexus was earlier reported from unnamed caves near Sokhumi, but it has now been found in theSakavria Cave in the Racha karst massif, which represents another significant range extension. It is worth to note that two species, Harpolithobius spinipes and Lithobius viriatus, have been observed in caves for the first time.

In total, 31 troglobiotic and stygobiotic species have been identified, which represents approximately 30% of the total number of species. In addition, we recorded 45 troglophiles, two stygobionts, 13 trogloxenes, six accidental records, and five species without ecological status due to insufficient material of these taxa. Troglobionts are represented predominantly by nine millipede species, and stygobionts by four mollusc species. Two species occur in both regions: the troglobiont Globochthonius satapliaensis and the stygobiont Niphargus rachalechkhumensis. Many caves in the studied massifs still remain biologically unexplored, which hinders an in-depth understanding of the species distribution and community composition in the subterranean environments of the Racha and Khvamli karst massifs. Moreover, several recorded taxa differ from all other known species from the nearby areas. So, the number of taxa is expected to increase provided further research, with the potential to describe a number species new to science.

 Acknowledgements

We thank Dragan Antić for identifying Diplopoda specimens and providing species photos; Levan Mumladze for his valuable insights on mollusks and guidance; Armen Seropian for spider identification; Tea Arabuli for information and literature on Acari and speleologist Valeri Barbakadze for his support during field expeditions. We are also grateful to all the collectors.

 Additional information Conflict of interest

The authors have declared that no competing interests exist.

 Ethical statement

No ethical statement was reported.

 Funding

This work was supported by the SRNSF (Grant No. MR-23-140 and Grant ID: FR-23-9905), by the Mohamed bin Zayed Species Conservation Fund (project ID: 242532420) and Institute of Zoology of Ilia State University.

 Author contributions

AM wrote the original draft of the manuscript and created the map. EK, MG, LS, NM, and SB edited the draft and identified the specimens of Chilopoda, Pseudoscorpiones, Isopoda, Opiliones, and Collembola, respectively.

 Author ORCIDs

Ana Margalitadze https://orcid.org/0009-0004-4805-7783

Eleonora Kiria https://orcid.org/0009-0007-1711-2163

Mariam Gogshelidze https://orcid.org/0009-0000-3614-213X

Lado Shavadze https://orcid.org/0000-0002-7643-2382

Naia Modebadze https://orcid.org/0009-0009-9213-5466

Shalva Barjadze https://orcid.org/0000-0001-8992-4987

 Data availability

All of the data that support the findings of this study are available in the main text or Supplementary Information.

 References AntićDŽMakarovSE (2016) The Caucasus as a major hotspot of biodiversity: Evidence from the millipede family Anthroleucosomatidae (Diplopoda, Chordeumatida).Zootaxa4211: 001–205. http://doi.org/10.11646/zootaxa.4211.1.1 AntićDŽReipHS (2020) The millipede genus Leucogeorgia Verhoeff, 1930 in the Caucasus, with descriptions of eleven new species, erection of a new monotypic genus and notes on the tribe Leucogeorgiini (Diplopoda: Julida: Julidae).European Journal of Taxonomy713: 1–106. https://doi.org/10.5852/ejt.2020.713 AntićDŠevićMBarjadzeSMakarovS (2023) Review of the genus Acanthophorella Antić & Makarov, 2016 (Diplopoda, Chordeumatida, Anthroleucosomatidae), with descriptions of three new species from the Caucasus.European Journal of Taxonomy908: 39–76. https://doi.org/10.5852/ejt.2023.908.2337 AntićDMargalitadzeAŠevićM (2025) Four new cavernicolous species of the genus Acanthophorella Antić & Makarov, 2016 (Diplopoda, Chordeumatida, Anthroleucosomatidae) from Georgia, Caucasus.Zootaxa5609(1): 41–69. https://doi.org/10.11646/zootaxa.5609.1.3 AsanidzeLChikhradzeNLezhavaZTsikarishviliKPolkJSLominadzeGBolashviliN (2017) Complex speleogenetic processes and mineral deposition in the Caucasus region of Georgia.Journal of Environmental Biology38: 1107–1113. BarjadzeSDjanashviliR (2008) Checklist of the springtails (Collembola) of Georgia.Caucasian Entomological Bulletin4(2): 187–193. BarjadzeSMurvanidzeMArabuliTMumladzeLPkhakadzeVDjanashviliRSalakaiaM (2015a) Annotated list of invertebrates of the Georgian karst caves. Georgian Academic Book, Tbilisi pp.119. BarjadzeSMurvanidzeMArabuliTPkhakadzeVMumladzeLChunashviliTGratiashviliNSalakaiaM (2015b) Annotated List of the Invertebrates in the Caves of Racha Region (Western Georgia).Proceedings of the Institute of Zoology24: 46–56. BarjadzeSArabuliTMumladzeLMaghradzeEAsanidzeZShavadzeLGogshelidzeM (2019) Cave Biodiversity of Transcaucasia, Open Access Database. Institute of Zoology at Ilia State University. https://cbg.iliauni.edu.ge (Accessed: 07/01/2025) BarjadzeSMayvanMMParimuchováAMaghradzeEKováčĽ (2024) Two new species of Plutomurus Yosii (Collembola: Tomoceridae) from Georgia and Iran.Zootaxa5463(4): 559–572. https://doi.org/10.11646/zootaxa.5463.4.6 BirsteinIA (1950) Cave fauna of Western Trans-Caucasia. Zoologicheskii zhurnal 29(5): 354–366. (in Russian) BirsteinJALopashovGV (1940) Investigations of the cave fauna in the USSR in 1935–1939. Bulletin Moskowskogo Obshhestva Ispytatelej Prirody, Biology 49(3/4): 29–38. BorutzkyEW (1950) Biospeologica Sovietica 12. Ground Isopods from the caves of the Bulletin of the Georgian SSR Academy of Sciences 21(5): 591–592. (in Russian) ChertoprudEMPalatovDMVinarskiMV (2020) Revealing the stygobiont and crenobiont Mollusca biodiversity hotspot in Caucasus: Part II. Sitnikovia gen. nov., a new genus of stygobiont microsnails (Gastropoda: Hydrobiidae) from Georgia.Zoosystematica Rossica29(2): 258–266. https://doi.org/10.31610/zsr/2020.29.2.258 ChertoprudEGregoJMumladzeLHofmanSPalatovDOsikowskiAFalniowskiA (2023) Revealing the Stygobiotic and Crenobiotic Molluscan Diversity in the Caucasus: Part IV—Crenobiotic Belgrandiellinae Radoman, 1983 (Mollusca, Hydrobiidae) from Georgia.Diversity15(3): 450. https://doi.org/10.3390/d15030450 DeltshevCBarjadzeSMaghradzeE (2023) On the cave spiders of Georgia with description of a new blind Centromerus (Arachnida: Araneae).Historia Naturalis Bulgarica45(11): 283–292. https://doi.org/10.48027/hnb.45.113 DjanashviliRBarjadzeSJordanaRBurkhardtU (2014) Redefinition of the genus Argonychiurus Bagnall, 1949 Collembola, Onychiuridae) with description of a new species from Georgia. Zootaxa. 3835. 381–391. https://doi.org/10.11646/zootaxa.3835.3.8 EvsyukovAGolovatchSReipHVandenspiegelD (2020) The millipede tribe Leptoiulini in the Caucasus, with notes on its generic classification (Diplopoda: Julida: Julidae).Zootaxa4778(2): 237–280. https://doi.org/10.11646/zootaxa.4778.2.2 GolovatchSI (1985) Two new genera of cave-dwelling millipedes (Diplopoda) with remarks on the millipede fauna of Western Caucasian caves. International Journal of Speleology 14(1/4): 39–50. GolovatchSI (1990) Diplopoda of the Caucasus. 3. Trachysphaeridae, with contributions to the fauna of Turkey. Senckenbergiana Biologica 70(4/6): 331–358. GolovatchSIEvsyukovAPReipH (2016) The millipede family Polydesmidae in the Caucasus (Diplopoda: Polydesmida).Zootaxa4085(1): 1–51. https://doi.org/10.11646/zootaxa.4085.1.1 GregoJMumladzeLFalniowskiAOsikowskiARysiewskaAPalatovDMHofmanS (2020) Revealing the stygobiotic and crenobiotic molluscan biodiversity hotspot in Caucasus: Part I. The phylogeny of stygobiotic Sadlerianinae Szarowska, 2006 (Mollusca, Gastropoda, Hydrobiidae) from Georgia with descriptions of five new genera and twenty-one new species.ZooKeys955: 1–77. http://doi.org/10.3897/zookeys.955.51983 GrosserCBarjadzeSMaghradzeEShavadzeLPešićVFailleA (2023) On the taxonomic status of Dina ratschaensis Kobakhidze, 1958 with a description of two new species–Dina imeretiensis sp. nov. and D. samegreloensis sp. nov. (Annelida, Hirudinida: Erpobdellidae).European journal of taxonomy891: 110–127. https://doi.org/10.5852/ejt.2023.891.2275 GrosserCMaghradzeEBarjadzeS (2025) Dina kobakhidzei sp. nov. (Hirudinea: Erpobdellidae): a new Caucasian cave leech from Georgia.Zoology in the Middle East75(1): 1–9. https://doi.org/10.1080/09397140.2025.2459501 KiriaEBarjadzeSTufIH (2023) Checklist of Georgian centipedes (Myriapoda: Chilopoda).Caucasiana2: 177–188. https://doi.org/10.3897/caucasiana.2.e108535 KobakhidzeDN (1958) New subspecies of the cave dwelling leech from Georgian SSR.Bulletin of the Georgian SSR Academy of Sciences 21(5): 591–592. (in Russian) NentwigWBosmansRGloorDHänggiAKropfC (2025) Spiders of Europe. Version: 12.2022, Available from: https://www.araneae.nmbe.ch (Accessed: 17/11//2025) MaghradzeEBarjadzeSLohajRFailleA (2025) Two new Trechini (Coleoptera: Carabidae) from Georgian caves, Caucasus.Zootaxa5609(1): 083–096. https://doi.org/10.11646/zootaxa.5609.1.5 MammolaSPavlekMHuberBABallarinFTolveMČupićIHesselbergTLunghiEMouronSGraco-RozaCCardosoP (2022) A trait database and updated checklist for European subterranean spiders.Scientific Data9(236): 1–13. https://doi.org/10.1038/s41597-022-01316-3 MarinINBarjadzeS (2022) A new species of stygobiotic atyid shrimps of the genus Xiphocaridinella (Crustacea: Decapoda: Atyidae) from the Racha-Lechkhumi and Kvemo Svaneti, with a new record of X. kumistavi from the Imereti, Western Georgia.Invertebrate Zoology19(1): 24–34. https://doi.org/10.15298/invertzool.19.1.04 MarinIBarjadzeSMaghradzeEPalatovD (2023) Diversity, taxonomy and phylogenetic relationships of the “Niphargus borutzkyi” ingroup (Crustacea: Amphipoda: Niphargidae) in Western Georgia, SW Caucasus.Zootaxa5352(4): 477–500. https://doi.org/10.11646/zootaxa.5352.4.2 ModebadzeNMartensJSnegovayaNBarjadzeS (2023) An annotated list of the Georgian harvestmen (Arachnida, Opiliones).Caucasiana2: 211–230. https://doi.org/10.3897/caucasiana.2.e106544 MumladzeLGregoJSzekeresM (2023) The land snail family Clausiliidae (Gastropoda, Pulmonata, Stylommatophora) in Georgia: overview, novel records and a new species.Caucasiana2: 29–61. https://doi.org/10.3897/caucasiana.2.e101013 MurvanidzeM (2014) Oribatid mites of Georgian (Caucasus) caves including the description of a new species of Ghilarovus Krivolutsky, 1966.International Journal of Acarology40(6): 463–472. https://doi.org/10.1080/01647954.2014.950604 NovákJGogshelidzeM (2025) Three new cavernicolous Neobisium species from Caucasian caves in Georgia (Arachnida: Pseudoscorpiones).Zootaxa5646(4): 575–585. https://doi.org/10.11646/zootaxa.5646.4.6 Ravanne (2022) Rapport de l’expédition spéléologique Racha 2022 (Géorgie). Fédération française de spéléologie. https://depots.ffspeleo.fr/uploads/100-2019_rapport_Racha_2019_-_v6.pdf (Accessed: 08/12/2025) ReadHJ (1992) The genus Cylindroiulus Verhoeff 1894 in the faunas of the Caucasus, Turkey and Iran (Myriapoda: Diplopoda: Julidae).Senckenbergiana biologica72(4–6): 373–433. TatashidzeZKTsikarishviliKDJishkarianiJM (2009) The Cadastre of the Karst Caves of Georgia.Petiti Publishing House, Tbilisi, 670 pp. (in Georgian) TurbanovISPalatovDMGolovatchSI (2016a) The state of the art of biospeleology in Russia and other countries of the former Soviet Union: a review of the cave (endogean) invertebrate fauna. 1. Introduction—Crustacea.Entomological Review96: 926–963https://doi.org/10.1134/S0013873816070162 TurbanovISPalatovDMGolovatchSI (2016b) The state of the art of biospeleology in Russia and other countries of the former Soviet Union: a review of the cave (endogean) invertebrate fauna. 2. Arachnida—Acknowledgments.Entomological Review96(9): 1297–1333. https://doi.org/10.1134/S0013873816090116 TsikarishviliKBarjadzeSKvavadzeEBolashviliNDjanashviliRMartkoplishviliI (2010) Speleology of Georgia: aspects of its current situation and perspectives.Cave and Karst Science,37(3): 73–78. TsikarishviliKDBolashviliN. (2013) The karst caves of Georgia: a short reference book.TSU press, Tbilisi, 157 pp. (in Georgian) ZaragozaJNovákJGardiniGMaghradzeEBarjadzeS (2021) Thetaxonomic status of the Caucasian cave-dwelling pseudoscorpion Chthonius satapliaensis (Arachnida: Pseudoscorpiones).Zoology in the Middle East67(4): 356–364. https://doi.org/10.1080/09397140.2021.1965072