Research Article |
Corresponding author: Armen Seropian ( armen.seropiani@iliauni.edu.ge ) Academic editor: Nils Hein
© 2024 Armen Seropian, Natalia Bulbulashvili, Giorgi Makharadze, Andrei Baznikin.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Seropian A, Bulbulashvili N, Makharadze G, Baznikin A (2024) New and interesting spiders (Arachnida, Araneae) from Vashlovani National Park and Chachuna Managed Reserve (Georgia). Caucasiana 3: 183-213. https://doi.org/10.3897/caucasiana.3.e132501
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Spiders collected individually between 19–23 May 2024 during a trip to Vashlovani National Park and Chachuna Managed Reserve were determined, resulting in 24 species, of which two species are described as new: Synema inexpectata sp. nov. (Thomisidae) and Prodidomus trihelicoides sp. nov. (Prodidomidae). Ten species - Acanthinozodium parysatis Zamani & Marusik, 2021 (Zodariidae), Berlandina nabozhenkoi Ponomarev & Tsvetkov, 2006, Cryptodrassus helvolus (O. Pickard-Cambridge, 1872), Synaphosus palearcticus Ovtsharenko, Levy & Platnick, 1994, Synaphosus turanicus Ovtsharenko, Levy & Platnick, 1994, Talanites involutus (O. Pickard-Cambridge, 1885), Turkozelotes attavirus Chatzaki, 2019 (all Gnaphosidae), Evarcha armeniaca Logunov, 1999 (Salticidae), Lycosa soboutii Shafaie, Nadolny & Mirshamsi, 2022 (Lycosidae), Rhysodromus rikhteri (Logunov & Huseynov, 2008) (Philodromidae), and Sardinidion blackwalli (O. Pickard-Cambridge, 1871) (Theridiidae) are recorded in Georgia for the first time, of which six species and three genera are new to the Caucasus region. The male of L. soboutii is described for the first time. Diagnostic drawings of preserved specimens and first photos of several preserved and alive species are provided along with the detailed collecting data.
Arthropoda, biodiversity, new records, new species, South Caucasus, taxonomy
Nestled in the extreme southeast of Georgia, the Vashlovani National Park (hereafter Vashlovani NP) presents a remarkable example of geographical and biological diversity. Covering approximately 35,000 hectares, it features a mosaic of landforms, including expansive steppes, rugged semideserts, imposing badlands, sinuous river valleys, and sporadic pockets of vegetation, such as dry open woodlands of mastic (Pistacia mutica) and juniper species (Juniperus spp.) (
This habitat supports a rich array of wildlife, harboring endangered species such as the Caucasian leopard (Panthera pardus tulliana), Eurasian lynx (Lynx lynx), and goitered gazelle (Gazella subgutturosa). The biodiversity of Vashlovani NP underscores its pivotal role in conservation initiatives and ecological research. As a microcosm of natural systems, it offers a unique opportunity for scientific inquiry and conservation efforts, providing a valuable context to explore the intricate interplay between geological formations, climatic conditions, and biological diversity. The same refers to the Chachuna Managed Reserve (hereafter Chachuna MR) bordering Vashlovani NP from the south-west.
Despite all these, until recently, the history of spider research in the specified territories was abysmal and included only three species: Eresus kollari Rossi, 1846, Orthobula charitonovi (Mikhailov, 1986), and Neon rayi (Simon, 1875) (
Most samples were collected individually during a four-day trip to Vashlovani NP and Chachuna MR. Additional material was collected within short-term individual trips to Gori and Kumisi Lake. Sampling details are given below. The elevations and GPS coordinates (given in WGS84) were obtained via Garmin GPS MAP 64s. Collected specimens were then preserved in 96% ethanol and stored in a freezer at -22˚C at the scientific collections of Ilia State University (Georgia, Tbilisi).
Identification was made using literature sources on Caucasian spiders (see list in
ALE – anterior lateral eyes;
AME – anterior median eyes;
PLE – posterior lateral eyes;
PME – posterior median eyes;
RTA – retrolateral tibial apophysis;
Tr – trochanter;
Fe – femur;
Pa – patella;
Ti – tibia;
Mt – metatarsus;
Ta – tarsus;
ISU – Ilia State University, Tbilisi;
mun – municipality.
In total, 53 spiders, including 19 males, 33 females, and 1 juvenile, collected during the sampling period were examined, comprising 24 species from 22 genera and 12 families. The list of all the recorded taxa is arranged alphabetically in the following section. Species marked with an asterisk (*) are reported from Georgia for the first time, while those marked with a double asterisk (**) represent the new records in the Caucasus region.
Brigittea
is a small genus of five valid species distributed in the Western Palaearctic (with B. civica (Lucas, 1848) introduced to South Africa), most of which were previously classified under Dictyna Sundevall, 1833 (
Brigittea innocens:
GEORGIA • 1♂, 1♀; Kakheti, Dedoplistskaro mun., Chachuna MR, Dalis Mta Reservoir; N41.2804°, E45.8968°; semidesert, Tamarix sp.; leg. Seropian A; 19 May 2024; CaBOL-IDs 1037662, 1038175.
This species is distributed from the eastern Mediterranean to eastern Kazakhstan (
Brigittea latens:
GEORGIA • 1♂; Kakheti, Dedoplistskaro mun., Chachuna MR, Dalis Mta Reservoir; N41.2804°, E45.8968°; semidesert, Tamarix sp.; leg. Seropian A; 19 May 2024; CaBOL-ID 1038114.
This species is distributed from the Iberian Peninsula east to eastern Kazakhstan and north to Steinkjer (Norway) (
Stegodyphus
is the second-largest genus of velvet spiders, with 20 valid species mainly diversified in Africa (
Stegodyphus lineatus:
GEORGIA • 2♂♂; Kakheti, Dedoplistskaro mun., Chachuna MR, Osiaskheoba; N41.2314°, E46.1473°; semidesert, on vegetation; leg. Seropian A; 19 May 2024; CaBOL-IDs 1038105, 1038146.
A recent record of this species in Georgia was based on a couple of males from Mijniskure (Vashlovani NP) (
Berlandina
is a relatively large genus in the subfamily Gnaphosinae with 41 valid species distributed in the Middle East, Caucasus, Mediterranean, East Asia, and West and East Africa (
Berlandina nabozhenkoi
GEORGIA • 2♀♀; Kakheti, Dedoplistskaro mun., Chachuna MR, Dalis Mta Reservoir; N41.2804°, E45.8968°; semidesert, under rocks; leg. Seropian A; 19 May 2024; CaBOL-IDs 1037707, 1037708.
This species is distributed from Turkey east to Iran and north to the Astrakhan Oblast of Russia (
Cryptodrassus
is a small genus of small spiders with 11 accepted species distributed in the Palaearctic and India (
Zelotes helvolus:
Zelotes helvolus:
GEORGIA • 1♂; Kakheti, Dedoplistskaro mun., Chachuna MR, Dalis Mta Reservoir; N41.2804°, E45.8968°; semidesert, under rocks; leg. Seropian A; 19 May 2024; CaBOL-ID 1037661.
This species is distributed from Cyprus north to Stavropol Kray of Russia, east to Kazakhstan, and south to Iraq and Iran (
Marinarozelotes
is a genus of small to medium-sized (2–12 mm) Zelotinae spiders with 23 accepted species distributed in the Palaearctic (
Marinarozelotes jaxartensis:
GEORGIA • 4♀♀, 1♂; Kakheti, Dedoplistskaro mun., Chachuna MR; Osiaskheoba; N41.2314°, E46.1473°; semidesert, under rocks and soil crevices; leg. Bulbulashvili N, Makharadze G; 19 May 2024; CaBOL-IDs 1037660, 1037394, 1037395, 1037870, 1037396. • 2♀♀; Kakheti, Vashlovani NP; Mijniskure; N41.1245°, E46.6456°; semidesert, under rocks; leg. Makharadze G; 22 May 2024; CaBOL-IDs 1038113, 1038156.
This species is distributed from North Africa to the Caucasus (Azerbaijan, Dagestan), Iran, Russia (Europe), and Central Asia. Introduced to Hawaii, USA, Mexico, South Africa, India, China, and Japan. (
Nomisia
is a relatively large genus with 39 currently recognized species mainly distributed in temperate and subtropical regions of the Palaearctic (
Nomisia ripariensis:
GEORGIA • 1♀; Kakheti, Dedoplistskaro mun., Chachuna MR, Dalis Mta Reservoir; N41.2804°, E45.8968°; semidesert, under rocks; leg. Makharadze G; 19 May 2024; CaBOL-ID 1037388.
This species is distributed from the eastern Mediterranean north to Dagestan and east to Iran (
Synaphosus
is a relatively large genus of small spiders, with 36 accepted species mainly distributed in the Palaearctic (
Synaphosus palearcticus:
Synaphosus palearcticus:
GEORGIA • 1♂, 1♀; Kakheti, Dedoplistskaro mun., Chachuna MR, Dalis Mta Reservoir; N41.2804°, E45.8968°; semidesert, under rocks; leg. Seropian A; 19 May 2024; CaBOL-IDs 1037705, 1038162. • 1♀, 1♂, 1juv.; Shida Kartli, Gori, Kvernaki ridge; N41.9926°, E44.1330°; heathland, under rocks; leg. Seropian A; 31 Mar. 2024; CaBOL-IDs 1037287, 1037285, 1037680.
This species is distributed from Crete to Kazakhstan (
Synaphosus turanicus:
Synaphosus turanicus:
GEORGIA • 1♀; Kakheti, Vashlovani NP; Mijniskure; N41.1245°, E46.6456°; semidesert, under rocks; leg. Makharadze G; 22 May 2024; CaBOL-ID 1037406.
Talanites
is a relatively small genus with 19 recognized species mainly distributed in temperate regions of the Holarctic (
Talanites fagei:
Talanites involutus:
GEORGIA • 2♂♂(1sub.); Kakheti, Dedoplistskaro mun., Chachuna MR, Dalis Mta Reservoir; N41.2804°, E45.8968°; semidesert, under rocks; leg. Seropian A; 19 May 2024; CaBOL-IDs 1037856, 1037857.
This species is known from Astrakhanskaya Oblast of Russia, Caucasus (Azerbaijan, Dagestan), Pakistan or India, and China (
Turkozelotes microb Kovblyuk & Seyyar, 2009.
Turkozelotes
is a small genus of small-sized ground spiders with six accepted species distributed in the West Palaearctic and most diversified in the Mediterranean region (
Turkozelotes attavirus: Chatzaki and Van Keer 2019: 451, figs 50–51, 56–57 (♂♀).
Since, in our opinion, the female description by Chatzaki and Van Keer (2019) lacks several notable features, we decided to describe the females from Georgia.
GEORGIA • 2♀♀; Kakheti, Vashlovani NP; Mijniskure; N41.1245°, E46.6456°; semidesert, under rocks; leg. A. Seropian; 22 May 2024; CaBOL-IDs 1038314, 1038315.
Female. Habitus as in Fig.
Epigyne and endogyne as in Figs
Two females were individually collected under rocks at the base of hills in the semidesert of Mijniskure, Vashlovani NP (Figs
Previously known from the type locality (Rhodes) only (
Several spider species have been known to mimic the velvet ants, from which at least two gnaphosids, namely Zelotes albobivittatus (Strand, 1906) and Titus lugens O. Pickard-Cambridge, 1901, are known to be mimics (although the first one is nomen dubium) (
Spiders of the genus Lycosa are relatively large and fossorial, with 220 recognized species distributed worldwide. The highest species diversity is found in Asia and South America (
Lycosa soboutii:
Lycosa soboutii:
GEORGIA • 3♂♂, 4♀♀; Kakheti, Dedoplistskaro mun., Chachuna MR, Osiaskheoba; N41.2314°, E46.1473°; semidesert, at night with flashlight; leg. Makharadze G, Seropian A; 19 May 2024; CaBOL-IDs 1038296, 1038298, 1038073, 1038297, 1038299, 1038074, 1038075. • 1♀; Kakheti, Vashlovani NP, Datviskhevi; N41.2382°, E46.3637°; semidesert, lured from burrow; leg. Bulbulashvili N; 20 May 2024; CaBOL-ID 1038300. • 1♀; Kakheti, Mijniskure; N41.1245°, E46.6456°; semidesert, at night with flashlight; leg. Makharadze G; 22 May 2024; CaBOL-ID 1038062. The material is deposited in the scientific collections of Ilia State University, Georgia, Tbilisi.
Diagnosis. In having extra processes of the tegular apophysis, the males of L. soboutii are most similar to those of L. elymaisa Zamani et Nadolny, 2022 and L. aragogi Nadolny et Zamani, 2017 from Iran, from which it can be easily distinguished by the less curved tegulum (Tg) [vs strongly curved in L. elymaisa (
Male. Total length 13.87. Carapace 7.49 long, 5.41 wide. Carapace light reddish brown, covered with dense off-white setae and medially radiating darker bands (Fig.
Female. See
Iran (
Comments: At the time of capture, all specimens were in the subadult stages and then captive-reared during May.
Lycosa soboutii (21: male, left palp, ventral view; 22: ditto, retrolateral view; 23: female, epigyne, ventral view; 24: endogyne, dorsal). Abbreviations: Em – embolus, Se – synembolus, Ta – tegular apophysis, Tg – tegulum, To – ventral outgrowth of tegular apophysis, Tp – distal process of tegular apophysis, Tr – tegular ridge. Scale bars: 0.5 mm.
Genus Oecobius Lucas, 1846
Oecobius maculatus:
GEORGIA • 1♂, 3♀♀; Kakheti, Dedoplistskaro mun., Chachuna MR, Dalis Mta Reservoir; N41.2804°, E45.8968°; semidesert, under rocks; leg. Makharadze G, Seropian A; 19 May 2024; CaBOL-IDs 1038125, 1038126, 1038127, 1038128.
This species is distributed from the Mediterranean to Azerbaijan, and has been introduced to the USA and Mexico (
Rhysodromus
is a relatively large genus with 27 valid species generally distributed in the Palaearctic, the majority of which were previously classified under Philodromus Walckenaer, 1826 (
Philodromus rikhteri
Rhysodromus rikhteri:
GEORGIA • 1♀; Kakheti, Dedoplistskaro mun., Chachuna MR; Osiaskheoba; N41.2314°, E46.1473°; semidesert, Tamarix sp.; leg. Seropian A; 20 May 2024; CaBOL-ID 1037865. • 1♀; Kvemo Kartli, Gardabani mun., Kumisi Lake; N41.5768°, E44.8172°; steppe, Tamarix sp.; leg. Seropian A; 22 Apr. 2024; CaBOL-ID 1037864.
This species is known solely by females and was recorded in Armenia and Azerbaijan (
Rhysodromus rikhteri , female (25: endogyne, dorsal view). Prodidomus trihelicoies sp. nov., holotype, ♀ (26: general habitus of live holotype female from Osiaskheoba; 27: epigyne, ventral view; 28: endogyne, dorsal view). Abbreviations: B – basal hood, Ic – insemination coils, Id – insemination duct, Lh – lateral hood, S – septum, Re – receptacula. Scale bars: 0.1 mm.
Prodidomus
is the largest genus in this gnaphosoid family, accounting for 53 valid species, originally distributed in Palaearctic, Afrotopic, Indomalaya, and Australasia (P. rufus Hentz, 1847, introduced to the Americas) (
Holotype. ♀ (CaBOL-ID 1037720); Georgia, Kakheti, Dedoplistskaro mun., Chachuna MR, Osiaskheoba; N41.2314°, E46.1473°; semidesert, in soil cracks; 20 May 2024; leg. Bulbulashvili N. Paratype. ♀ (CaBOL-ID 1037833); Georgia, Kakheti, Vashlovani NP, Mijniskure; N41.1245°, E46.6456°; semidesert, under rocks; 22 May 2024; leg. Makharadze G. Both specimens are deposited in the scientific collections of ISU.
The new species resembles P. redikorzevi Spassky, 1940 (
Female (holotype/paratype). Total length 3.46/4.44. Carapace ovoid, 1.46/1.50 long, 1.18/1.23 wide. Sternum 1.16/1.11 long, 0.82/0.85 wide. Carapace, sternum, labium, chelicerae, and maxillae orange, without any pattern, covered with light yellow setae. Abdomen light yellow-orange without any pattern, dorsally covered with light yellow setae and dark brown setae posteriorly. Ocular area triangular. Chelicerae without teeth. Legs the same color as carapace, without annulations. Eyes: AME 0.12/0.13, ALE 0.11/0.12, PME 0.14/0.15, PLE 0.13/0.14. Leg measurements: I 4.67/4.76 (1.16/1.18+0.93/0.94+0.98/1.01+0.84/0.85+0.76/0.78), II 3.89/4.02 (0.96/0.98+0.74/0.77+0.84/0.86+0.69/0.73+0.66/0.68), III 3.41/3.53 (0.88/0.91+0.62/0.65+0.71/0.73+0.61/0.63+0.59/0.61), IV 4.25/4.33 (0.85/0.87+0.79/0.81+1.28/1.30+0.71/0.72+0.62/0.63). Epigyne and endogyne as in Figs
Male. Unknown.
The specific epithet is an adjective referring to the three coils of insemination ducts.
Two females were collected in soil cracks and under rocks in semideserts of Chachuna MR and Vashlovani NP (Figs
Known from two localities in Georgia only.
In his paper,
Aelurillus
is a large genus with 75 valid species mainly distributed in the Palaearctic, with only ten species being recorded from outside its limits (
Aelurillus concolor:
GEORGIAi • 1 ♂; Kakheti, Dedoplistksaro mun., Vashlovani NP; Mijniskure; N41.1245°, E46.6456°; semidesert; leg. Makharadze G; 22 May 2024; CaBOL-ID 1037400.
This species is distributed from Greece to Central Asia (
Chalcoscirtus
is a large genus of small-sized jumping spiders with 46 valid species with a Holarctic distribution (
Chalcoscirtus tanasevichi:
Chalcoscirtus tanasevichi:
GEORGIA • 1 ♂; Kakheti, Dedoplistskaro mun., Chachuna MR, Dalis Mta Reservoir; N41.2804°, E45.8968°; semidesert; leg. Seropian A; 19 May 2024; CaBOL-ID 1037713.
This species is distributed from Turkey to Central Asia (
Evarcha
is a large genus with 94 valid species mainly distributed in the Holarctic, Oriental, and Afrotopical regions (for diagnosis, see
Evarcha armeniaca Logunov, 1999b: 301, figs 1–4 (♂♀).
GEORGIA • 3♂♂, 3♀♀; Kakheti, Dedoplistksaro mun., Vashlovani NP; Mijniskure; N41.1245°, E46.6456°; semidesert; leg. Bulbulashvili N, Makharadze G; 22 May 2024; CaBOL-IDs 1037663, 1037397, 1037398, 1037664, 1038129, 1038115.
This species was described from Armenia and Azerbaijan (Logunov 1999) and then recorded in Turkey (
A monotypic genus with a single species previously classified under Theridion Walckenaer, 1805, and distributed in the Western Palaearctic (
Sardinidion perplexum:
GEORGIA • 1♂; Kakheti, Dedoplistksaro mun., Vashlovani NP; Datviskhevi; N41.2361°, E46.3709°; semidesert; leg. Seropian A; 20 May 2024; CaBOL-ID 1038179.
This species is distributed from the Eastern Mediterranean to eastern European Russia (
Sardinidion blackwalli , male (41: left palp, retrolateral view). Synema inexpectata sp. nov., holotype, male (42: general habitus of holotype male from Osiaskheoba; 43: left palp, ventral view; 44: ditto, retrolateral view; 45: retrolateral tibial apophysis, dorsal view). Abbreviations: Et – embolic tip, RTA – retrolateral tibial apophysis. Scale bars: 0.2 mm (41, 43–45); 1 mm (42).
GEORGIA • 1♀; Kakheti, Dedoplistskaro mun., Chachuna MR, Dalis Mta Reservoir; N41.2804°, E45.8968°; semidesert; leg. Seropian A; 19 May 2024; CaBOL-ID 1038158.
Species with Holarctic distribution (
Coscinida
is a relatively small genus with 18 valid species mainly distributed in Africa, East, and South Asia (
Coscinida tibialis:
GEORGIA • 1♀; Kakheti, Dedoplistksaro mun., Vashlovani NP; Datviskhevi; N41.2361°, E46.3709°; semidesert; leg. Seropian A; 20 May 2024; CaBOL-ID 1038179.
This is the southeasternmost record in Georgia and the first one in the Kakheti region.
Aranea rotundata Walckenaer, 1802
Synema
is a large genus of brightly colored crab spiders with 126 valid species distributed on all continents except Australia (
Holotype. ♂ (CaBOL-ID 1038144): Georgia: Kakheti, Dedoplistskaro mun., Chachuna MR, Osiaskheoba; N41.2314°, E46.1473°; semidesert, vegetation; 20 May 2024; leg. Makharadze G; Type specimen is deposited in the scientific collections of Ilia State University, Georgia, Tbilisi.
Diagnosis. The new species resembles S. utotchkini Marusik & Logunov, 1995 (
Male (holotype). Total length 3.82. Carapace 2.03 long, 2.08 wide. Abdomen 1.92 long, 1.79 wide. Cheliceral length 0.75. Eye measurements: AME 0.11, ALE 0.16, PME 0.08, PLE 0.09. Carapace dark-brown with lighter median part. Abdomen dorsally bright red (coloration dull in ethanol) with extensive black pattern (Fig.
Female unknown.
The specific epithet is an adjective and refers to the unexpected discovery of a new species of this genus in Georgia.
A single male of the new species was individually collected from vegetation in semidesert (Fig.
Known from the type locality only.
During the fieldwork conducted by the CaBOL team in July 2023 on the territory of Chachuna MR, several new and interesting spiders were collected (
Titanoeca
is the largest genus within the family, comprising 31 valid species mainly distributed in the Palaearctic, with only seven species being recorded from outside its limits. Seven species, namely T. caspia Ponomarev, 2020, T. caucasica Dunin, 1985, T. nivalis Simon, 1874, T. schineri L. Koch, 1872, T. tristis L. Koch, 1872, T. ukrainica Guryanova, 1992, and T. veteranica Herman, 1879, have been recorded in the Caucasus (
Titanoeca caucasica:
Titanoeca caucasica:
GEORGIA • 2♀♀, 3♂♂; Kakheti, Dedoplistksaro mun., Vashlovani NP; Datviskhevi; N41.2361°, E46.3709°; semidesert; leg. Bulbulashvili N, Makharadze G; 20 May 2024; CaBOL-IDs 1037698, 1037700, 1037695, 1037699, 1037401.
This species is distributed in the Caucasus, Turkey, and Iran (
Acanthinozodium
currently comprises 24 valid species, distributed in Iran, Central Turkmenistan, the Arabsphere, West Africa, and Cameroon. None of the species have been recorded in the Caucasus (
Acanthinozodium parysatis
GEORGIA • 1♂; Kakheti, Dedoplistskaro mun., Vashlovani NP; Mijniskure; N41.1245°, E46.6456°; semidesert, under rocks; leg. Seropian A; 22 May 2024; CaBOL-ID 1038313.
This species was previously known from the Ardabil and Qazvin Provinces of Iran only (
Zodarion
is the largest genus within the family, comprising 171 valid species mainly distributed in the Palaearctic region. Eight species, namely Z. abantense Wunderlich, 1980, Z. caucasicum Dunin & Nenilin, 1987, Z. italicum (Canestrini, 1868), Z. morosum Denis, 1935, Z. petrobium Dunin & Zacharjan, 1991, Z. rhodiense Caporiacco, 1948, Z. rubidum Simon, 1914, and Z. thoni Nosek, 1905, have been recorded in the Caucasus (
Zodarion nigrifemur:
Zodarion nigrifemur:
Zodarion rhodiense:
GEORGIA • 1♂; Kakheti, Dedoplistksaro mun., Vashlovani NP; Mijniskure; N41.1245°, E46.6456°; semidesert; leg. Makharadze G; 22 May 2024; CaBOL-ID 1037399.
This species was recorded in Greece, Turkey, and recently from several localities in Georgia (
We are grateful to Nils Hein (subject editor) and an anonymous reviewer for their valuable comments, which helped to improve the quality of our manuscript. Our team is indebted to the Agency of Protected Areas for the extension of the collection permit #655-0-2-202103182033..
The authors have declared that no competing interests exist.
No ethical statement was reported.
The study was partly founded by the Federal Ministry of Education and Research under grant number 01DK20014A.
Conceptualization: AS. Material collection: AS, NB, GM. Methodology: AS. Material sorting and identification: AS GM. Data Curation: AS. Writing and drawing - original draft: AS. All authors read and approved the final draft of the manuscript.
Armen Seropian https://orcid.org/0000-0003-3777-9954
Natalia Bulbulashvili https://orcid.org/0000-0002-6802-1209
Giorgi Makharadze https://orcid.org/0009-0008-4443-5432
Andrei Baznikin https://orcid.org/0009-0001-9149-1735
All of the data that support the findings of this study are available in the main text or Supplementary Information.