98urn:lsid:arphahub.com:pub:3686FA2D-1003-5130-A1A9-6F0A4E63DF4Durn:lsid:zoobank.org:pub:A3B9B45C-7F69-42DF-85B6-B3189A5B43F2CaucasianaCaucasiana2667-9809Pensoft Publishers10.3897/caucasiana.5.e189214189214ChecklistAmphibiaReptiliaBiodiversity & ConservationBiogeographySystematicsCenozoicNeogeneAn Annotated Checklist of the Amphibians and Reptiles of Georgia with DNA Barcoding Data and Taxonomic RemarksTarkhnishviliDaviddavid_tarkhnishvili@iliauni.edu.gehttps://orcid.org/0000-0003-1479-98801IankoshviliGiorgihttps://orcid.org/0000-0002-0429-14881SeropianArmenhttps://orcid.org/0000-0003-3777-995412KachlishviliNinohttps://orcid.org/0000-0002-5632-89592ToduaMariamihttps://orcid.org/0000-0003-1870-30301Caucasus Leibniz Biodiversity Research Center, Ilia State University, Tbilisi, GeorgiaCaucasus Leibniz Biodiversity Research Center, Ilia State UniversityTbilisiGeorgiahttps://ror.org/051qn8h41Institute of Zoology, Ilia State University, Tbilisi, GeorgiaInstitute of Zoology, Ilia State UniversityTbilisiGeorgiahttps://ror.org/051qn8h41
Corresponding author: David Tarkkhnishvili (david_tarkhnishvili@iliauni.edu.ge)
Academic editor: Levan Mumladze
20260807202655913890335C41-0356-5786-A015-CC4242A532E9212913636111B0E9-6FDD-4FA7-BE9E-8ACEC972C8D61902202615052026David Tarkhnishvili, Giorgi Iankoshvili, Armen Seropian, Nino Kachlishvili, Mariami ToduaThis 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.https://zoobank.org/6111B0E9-6FDD-4FA7-BE9E-8ACEC972C8D6
Despite the publication of numerous papers, spanning the period from the 19th century to recent times, that have been devoted to the faunistics and taxonomy of the herpetological fauna of the Caucasus Biodiversity Hotspot, the actual species diversity of the region still remains unclear. Some formally described taxa lack comprehensive validation based on integrative phylogenetic and biogeographic evidence, whereas other nominal species have been questioned or synonymized in subsequent studies. In this paper, we present an annotated checklist of 12 amphibian and 56 reptile species recorded from Georgia, accompanied by taxonomic remarks based on current molecular, morphological, and biogeographic knowledge. Most taxa are represented by DNA barcodes (COI) generated within the Caucasus Barcoding of Life (CaBOL) project and deposited in institutional collections and international databases. We explicitly distinguish between biologically confirmed species, effectively isolated from their closest relatives, and candidate species whose status requires further validation; particularly in cases where contact zones and potential clinal variation remain insufficiently studied. The checklist is supplemented with specimen photographs and generalised distribution maps. Overall, we aim to contribute to a more transparent and critically evaluated taxonomic framework for the herpetofauna of Georgia and the broader Caucasus region.
AmphibiaReptiliaspecies listtaxonomyShota Rustaveli National Science Foundation501100004801https://ror.org/00xc87681http://doi.org/10.13039/501100004801Citation
Tarkhnishvili D, Iankoshvili G, Seropian A, Kachlishvili, N, Todua M (2026) An Annotated Checklist of the Amphibians and Reptiles of Georgia with DNA Barcoding Data and Taxonomic Remarks. Caucasiana 5: 59–138. https://doi.org/10.3897/caucasiana.5.e189214
Introduction
The herpetofauna of the Caucasus, and specifically Georgia, has been explored since the early 19th century initially by naturalists such as Eichwald (1831, 1841), Ménétriés (1832), and Hohenacker (1837). The first treatise devoted to the subject was Nikolskii’s (1913) “The Amphibians and Reptiles of the Caucasus”. Later, Terentyev and Chernov (1940) and Bannikov et al. (1977) included extensive information on amphibians and reptiles from Georgia in their books, with further significant contributions to the subject area from both Darevsky (1967), and Muskhelishvili’s (1970). In more recent years we have seen the publication of “The Amphibians of the Caucasus” (Tarkhnishvili and Gokhelashvili 1999) and “Amphibians of the Former Soviet Union” (Kuzmin 2012). Speybroeck et al. (2020) published a list of amphibians and reptiles of Europe, which included most of the species found in Georgia. Tuniyev and Ananjeva (2024) also published an annotated species list of the amphibians and reptiles of the Caucasus ecoregion, with short accounts of formally described 18 amphibian and 110 reptile species. Many shorter individual publications, including conference abstracts, have reported new individual findings, for instance, Tarkhnishvili et al. (2002) attempted to explore declines in amphibian and reptilian populations using data published since Nikolskii (1913). Finally, in recent years, a significant amount of additional information has become available through both recently published papers (e.g., Mebert et al. 2025a, b) and unpublished findings posted on social media (see Iankoshvili and Tarkhnishvili 2021). Some of these have been published on the website https://biodiversity.iliauni.edu.ge/en, where digital images of live or dead animals are commonly provided as evidence of a new finding; this information is awaiting summary and publication.
There are some discrepancies in the nomenclature used to describe the amphibians and reptiles of Georgia. The most commonly used nomenclature for reptiles is from the Reptile Database, a source listing species worldwide (Uetz et al. 2025), and the most widely used source of amphibian names is Amphibian Species of the World (Frost 2026). The nomenclatures used by IUCN (www.redlist.org) and the ITIS Catalog of Life (Bánki et al. 2026) are largely consistent with these two sources. However, some research papers have suggested changing the nomenclature for certain species, but these suggestions are not always been adopted by the previously mentioned sites or in global nomenclatural sources. The recent species list of Tuniyev and Ananjeva (2024) is the most exhaustive account of the herpetological fauna of the Caucasus; however, the validity of some nominal species included in their list is questioned by other researchers. All of which highlights that there can be inconsistencies in names given to herpetofauna in the country.
Currently, the most commonly used definition of a ‘species’, is the Integrative Species Concept (ISC), introduced by De Queiroz (2007). This concept considers that species are taxonomic units with “separate evolutionary pathways”, which can be validated through the presence of differences at multiple independent loci (and which does not exclude some hybridization among different species). However, in most cases, taxonomists currently describing new species do not have sufficient information to strictly validate these criteria, and clearly, they were not available to taxonomists of the previous generations. An alternative approach is validating species status by the presence of at least one phenotypically distinct population in the same geographic space, or by narrow contact zones among the ranges of closely related phenotypes.
Hillis (2019) underlined that the use of molecular genetic data may lead to premature taxonomic solutions when contact zones between different populations remain insufficiently studied. Recent studies, some affecting Georgain taxa, argue that the increasing reliance on mitochondrial markers in herpetology has, in some groups, contributed to taxonomic inflation, whereby geographically distant populations exhibiting moderate genetic divergence are elevated to species rank without sufficient evaluation of contact zones or reproductive isolation (Freitas et al. 2020; Dufresnes et al. 2024a; Tarkhnishvili 2024). Mitochondrial differentiation, even when accompanied by phenotypic differences, does not necessarily indicate irreversible evolutionary divergence but often reflects intraspecific geographic variation. Wüster (2025) published a valuable review paper in which he offered a methodology for validating the status of candidate species and disqualifying taxonomic solutions. He distinguishes between (i) a critical approach, aiming to disqualify the taxonomic status in case of firm evidence that there are no species-level differences between the candidate taxa, and (ii) a confirmatory approach that suggests the absence of sufficient support for a taxonomic status of a species, but does not aim to reject it until either the hypothesis (valid vs non-valid species) is supported. He found that in most recent species descriptions, both molecular and morphological data were used to confirm rather than to be critical, in both cases of the presence or absence of contact areas between the candidate species. The essence of the critical approach is testing whether the coincidental distribution of phenotypic or genomic vs mitochondrial DNA markers shows a gradual change between the type localities or a stepped cline. Only in the latter case is the status of a candidate species confirmed; if data from the contact area are absent, the validity of a species cannot be supported or rejected.
Some formally described species of amphibians and reptiles from Georgia do not have a status validated by the presence of coinciding differences at multiple alleles among related taxa, which would validate their status under either the traditional Biological Species (Mayr 1949) or Integrative Species (DeQueiroz 2007; Wüster 2025) concept. For sister and other closely related species, we do not know whether stepped clines of molecular and/or morphological traits are present in their contact zones, or whether there is just geographic variation in phenotypic traits and allele frequencies (e.g., Dubey et al. 2015). If a species description is based on acceptable criteria but the presence of stepped clines or full spatial isolation has never been validated, in a species account, we indicate that this is a “candidate species”, regardless of its current formally accepted taxonomic status.
There are also some formally described species whose status has not been confirmed and have rejected by later studies. (e.g., Schmidtler et al. 2002; Kindler et al. 2013; Mebert et al. 2025b). In the latter case, we do not include these nominal taxa in the checklist of Georgian amphibians and reptiles as separate species and downscale their status to subspecies or form comb. nov. Hence, we indicate, along with formal taxonomic status, the following details that affect the future status of a species: (1) Whether a species is sympatric with its closest relative and maintains the integrity of its species-specific characters (biologically confirmed species); (2) Whether a species is parapatric with its closest relatives or occupies distinct habitat types, with no overlap or a narrow contact zone with stepped clines of interspecific divergent characters across this area, indicating a limited gene flow (local hybridization) between these species whereas the species retain diagnostic phenotypic characters outside the contact zone throughout most of their ranges (biologically confirmed species); (3) Whether a species is allopatric with its closest evolutionary relatives, but the estimated split time is exceptionally long (~ 10 Mya or more, inferred from the molecular clock analysis), and there is a clear phenotypic hiatus between the species and its closest relatives (biologically confirmed species); (4) A species has a set of diagnostic phenotypic and genetic characters, but the area of contact with its closest relative is not sampled/studied, hence it is unclear whether the difference between the species and its closest relative can be reduced to within-species clinal variation (Candidate species); (5) A species is allopatric with its closest relatives and has some diagnostic phenotypic and/or genotypic characters. However, the time of split is modest, hence in case of secondary contact we cannot exclude merger of the evolutionary lineages (candidate species); (6) A species is diagnosed by some phenotypic characters, but genetic differentiation is minute (candidate species); (7) It is clearly suggested in some publications that the candidate species does not have fixed differences with its closest relatives and is not separated genetically (rejected taxon).
We underline that the genetic distance is not used here as a single piece of evidence for a separate status, until it is very high (conventionally, ~9-10% of mtDNA difference or more for less conservative genes such as Cytochrome b or Cytochrome Oxidase I) (Mebert 2008; Tarroso et al. 2014; Mebert et al. 2016; Stümpel et al. 2016). A distance of ~4-6% in mtDNA may indicate separate species status (Freitas et al. 2020; Kindler et al. 2017); however, it does not exclude lineage merger across the contact zone.
Simultaneously, little genetic difference does not exclude separate species status if there are big phenotypic or ecological differences that trigger prezygotic isolation of the lineages. The purpose of this contribution is not to propose new taxonomic acts, but to critically synthesize existing evidence and provide a transparent evaluation of species status within a regional framework. All taxonomic decisions presented here are explicitly linked to published data and are intended to facilitate future integrative revisions.
Materials and methodsGeneral approach
For each species listed below, if applicable, the following sections are included: (1) scientific and vernacular names in English and Georgian; (2) synonymy, restricted to names historically applied to Georgian populations or widely used in the literature; (3) nomenclatural sources followed, including the first publication using the currently accepted name and major modern taxonomic databases; (4) infraspecific variation, listing subspecies occurring in Georgia when applicable. Subspecific names considered invalid are discussed in the taxonomic remark; (5) taxonomic remark, summarizing phylogenetic affinities, historical taxonomic treatments, and the biological validation status (see Introduction); (6) barcoded individuals, including CaBOL identification numbers, BOLD BINs, and closest matches in BOLD Systems and/or NCBI databases. Supplementary Material 1 contains information about location, CaBOL (Caucasus Barcode of Life project – https://ggbc.eu/) identification number, and BOLD BIN for each study specimen.
Assignment of biological validation status (biologically confirmed species, candidate species, rejected taxon) was based on a qualitative assessment of available phylogenetic, population genetic, morphological, and biogeographic evidence from the published literature. Particular emphasis was placed on (i) evidence of sympatry or parapatry with full or partial reproductive isolation, (ii) presence or absence of stepped clines across contact zones, (iii) depth of divergence inferred from multilocus datasets, and (iv) consistency between mitochondrial and nuclear markers when available. No fixed genetic distance threshold was used as a standalone criterion.
Images of barcoded specimens
The images of preserved specimens were taken with a Canon EOS 90D camera equipped with a Canon EF 60 mm f/2.8 Macro USM lens and a Canon Macro Twin Lite MT-26EX-RT. Digital images were then prepared using Zerene Stacker image stacking software and Adobe Photoshop CS6 (version 13.0). Photographs of live specimens were taken with a Canon EOS 550D camera and a Canon EF 100 mm f/2.8 Macro USM lens.
Barcoding
DNA isolation from tissue samples was processed by using Quick-DNA miniprep Plus Kit (Zymo Research). Partial sequences of cytochrome oxidase subunit I (COI) were amplified by polymerase chain reaction (PCR) using the primer pair LCOI490-JJ and HCO2198-JJ (Astrin and Stüben, 2008). Thermal conditions included denaturation at 95 °C for 1 min, followed by the first cycle set (15 cycles): 94 °C for 30 s, annealing at 55 °C for 1 min (–1 °C per cycle), and extension at 72 °C for 1:30 min. Second cycle set (25 cycles): 94 °C for 35 s, 45 °C for 1 min, 72 °C for 1:30 min, followed by 1 cycle at 72 °C for 3 min and final extension step at 72 °C for 5 min. Sequencing of the unpurified PCR products in both directions was conducted at the Beijing Genomics Institute (Hong Kong, CN) by using the amplification primers. Extracted DNA was deposited in the scientific collections of Ilia State University, Tbilisi, Georgia, and aliquots will be deposited at LIB Biobank at Museum Koenig, Bonn, Germany. At the same time, the sequences (length 500-658 bp) have been submitted to Barcode of Life Data System (BOLD) databases. The newly obtained DNA barcodes of COI sequences were compared against the Barcode of Life Data System (BOLD; www.boldsystems.org. Barcode Index Number (BIN) (Ratnasingham and Hebert, 2013) for the sequenced taxa and their nearest neighbor in BOLD Systems BIN are also given. Sequence analysis for COI was performed using Geneious Prime 2022.1.1 (http://www.geneious.com).
Maps
Distribution maps were prepared in QGIS (QGIS Development Team 2023). Species ranges were approximated based on verified occurrence records, habitat suitability inferred from MODIS land cover classes (Missions 2020) and known climatic constraints, elevation range and annual precipitation (Hijmans et al., 2005). Maps are intended to illustrate general distribution patterns/expected distribution rather than precise range boundaries.
https://binary.pensoft.net/fig/1711428Records in Georgia.
Waga (1876), Nikolskii (1913), Bannikov et al. (1977), Tartarashvili and Bakradze (1989), Tarkhnishvili and Gokhelashvili (1999), Tarkhnishvili et al. (2000, 2008).
Infraspecific variation.
Mtirala reduced-pigmentation population (M.c.djanaschvilii Tartarashvili and Bakradze, 1989); “M. sp. 1” lineage (Borjomi Gorge/Meskheti) per Tarkhnishvili et al. (2000) – treated here as a lineage label, not a formal taxon.
Taxonomic remark.
Biologically confirmed species. Separated from the closest relative, Chioglossalusitanica, by a broad morphological and genetic hiatus, the distribution of both species is disjunct, and the time of split is estimated as 40-70 million years (Myr) (Kumar et al. 2017). Eocene relict, endemic of the western part of the Lesser Caucasus.
One undescribed but genetically isolated evolutionary lineage, perhaps a cryptic species, is present in Borjomi Gorge (Central Georgia) and the northern slopes of the Meskheti range: M. sp. 1 (Tarkhnishvili et al. 2000). The barcoded individuals belong to M. sp. 1 rather than to the “typical” M.caucasica. We do not follow the recent treatment of Mertensielladjanashvilii, as a separate species: the boundary between the nominal form and M. sp. 1 is not yet clear, and M.c.djanaschvilii does not genetically differ from the other populations of the nominal M.caucasica (Tarkhnishvili et al. 2000). This form is treated as M.djanaschvilii Tartarashvili and Bakradze, 1989 (status elevated by Raffaëlli 2022; followed by Tuniyev and Ananjeva 2024) (not accepted here). Frost (2026) considers it as a subspecies. The taxonomy of M.caucasica requires detailed genetic studies across its range before a formal revision.
Barcoded individuals.
CaBOL-IDs 1027447 (Tashiskari), 1027448 (Bakhvistskali River) (BOLD: AAB0814; similarity 98.78%). The closest homologous sequences: M.caucasica (M. sp. 1) (BOLD: AAB0814; Georgia, Akhaldaba; similarity 100–98.7%). The second-closest taxon is M.caucasica RuHF with a private status (Turkey, Ordu environs; similarity 88.48%).
AnimaliaCaudataSalamandridaeF464692B-EC29-54AE-A60E-1FE2CAAAE003Lissotritonlantzi(Wolterstorff, 1914)Figs 2, 121 (Engl. Caucasian Smooth Newt; Geo. კავკასიური ჩვეულებრივი ტრიტონი, kavkasiuri chveulebrivi tritoni)Tritonvulgaris subsp. typica forma lantzi Wolterstorff, 1914Molgevulgarislantzi: Nikolskii (1918)Triturusvulgarislantzi: Bannikov et al. (1977: 22); Tarkhnishvili and Gokhelashvili (1999: 55), et auct.Lissotritonvulgarislantzi: Speybroeck et al. (2020); Raffaëlli (2022)Taxonomic sources followed.
Dubois and Raffaëlli (2009), Kuzmin (2012), Bánki et al. (2026), Raffaëlli (2022), Tuniyev and Ananjeva (2024), Frost (2026).
Records in Georgia.
Lantz (1912), Nikolskii (1913), Bannikov et al. (1977), Tarkhnishvili and Gokhelashvili (1999).
Taxonomic remark.
Candidate species. L.lantzi is a geographically isolated evolutionary lineage within the L.vulgaris species complex. Pabijan et al. (2015) suggest that L.lantzi is the oldest isolated lineage within the complex, evolving in isolation from its closest relatives since late Miocene. However, the isolation time (~6.4 Mya; Arntzen et al. 2015) may not be sufficient for the development of postzygotic isolation. This form is listed as a separate species in Frost (2026) but not in Raffaëlli (2022).
AnimaliaCaudataSalamandridaeBE29A2C3-9628-5A37-A3A0-FA13E630F740Ommatotritonophryticus(Berthold, 1846)Figs 3, 121 (Engl. Northern Banded Newt; Geo. კავკასიური ტრიტონი, kavkasiuri tritoni)Tritonophryticus Berthold, 1846Molgevittata: Nikolskii (1913: 234)Triturusvittatusophryticus: Bannikov et al. (1977: 27); Tarkhnishvili and Gokhelashvili (1999: 40), et auct.Ommatotritonophryticusophryticus: Raffaëlli (2022)Taxonomic sources followed.
Litvinchuk et al. (2005), Kuzmin (2012), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Frost (2026).
Records in Georgia.
Lantz (1912), Nikolskii (1913), Bannikov et al. (1977), Tarkhnishvili and Gokhelashvili (1999).
Infraspecific variation.
O.o.ophryticus Berthold, 1846.
Taxonomic remark.
Biologically confirmed species. Separated from the closest relative (O.vittatus) in early Miocene or late Oligocene, no introgression was recorded based on the genomic data analysis (van Riemsdijk et al. 2022; Kalaentzis et al. 2025).
A member of species complex O.vittatus, which contains three allopatric species. Formerly was referred to as a geographic subspecies of O.vittatus. Mitochondrial Cytochrome b differences (p-distance) between O.ophryticus and O.nesterovi from NW Anatolia (NCBI accession numbers KX682130 and DQ821264) exceed 10%. The range of O.ophryticus is fully isolated from that of O.vittatus and O.nesterovi. Found throughout the entire mountain forest belt of Georgia.
AnimaliaCaudataSalamandridaeDDFA436E-10D8-53F5-9C26-C72162363241Trituruskarelinii(Strauch, 1870)(Figs 4, 121) (Engl: Eastern Crested Newt; Geo. კარელინის სავარცხლიანი ტრიტონი, karelinis savartskhliani tritoni)Tritonkarelinii Strauch, 1870Molgecristata var. karelinii: Nikolskii (1913: 232)Trituruscristatuskarelinii: Bannikov et al. (1977: 24); Tarkhnishvili and Gokhelashvili (1999: 66), et auct.Taxonomic sources followed.
Bucci-Innocenti et al. (1983), Kuzmin (2012), Bánki et al. (2026), Speybroeck et al. (2020), Raffaëlli (2022), Tuniyev and Ananjeva (2024), Frost (2026).
Records in Georgia.
Nikolskii (1913), Bannikov et al. (1977), Tarkhnishvili and Gokhelashvili (1999).
Taxonomic remark.
Biologically confirmed species. Trituruskarelinii builds a monophyletic group together with two allopatric Triturus species. One of these, T.ivanbureschi Arntzen & Wielstra, 2013, is separated by a narrow hybrid zone from other crested newts of genus Triturus (Arntzen et al. 2014).
A member of T.cristatus species complex, with seven allopatric and parapatric species. Formerly was referred to as a subspecies of T.cristatus. Wielstra et al. (2019) suggest that the north-western Anatolian Eastern crested newts belong to two different species, T.anatolicus and T.ivanbureschi. It is not completely clear whether there is a geographic gap or narrow hybrid zone between T.anatolicus and T.karelinii. Mitochondrial Cytochrome b differences (p-distance) between T.karelinii from NE Türkiye and T.ivanbureschi from the Balkan Peninsula (NCBI accession # DQ821247 and DQ821249) is 1.3%, which does not exceed the differences commonly recorded for conspecific populations. However, this does not challenge the taxonomic status of T.karelinii, which is the first species of Triturus described for Anatolia. Found throughout Western Georgia and the forest belt of Eastern Georgia.
https://binary.pensoft.net/fig/1711429Records in Georgia.
Nikolskii (1913), Bannikov et al. (1977), Tarkhnishvili and Gokhelashvili (1999).
Infraspecific variation.
P.s.boettgeri C. M. Cooke & Pilsbry, 1920 — subspecies concept follows Dufresnes et al. (2019); Georgian populations tentatively assigned based on regional affiliation.
Taxonomic remark.
Biologically confirmed species. Parapatrically distributed with the closest relative, P.fuscus (Laurenti, 1768), without signs of hybridization or common presence of intermediate phenotypes or genotypes (Veith et al. 2006). The genomic data of Dufresnes et al. (2019) suggest that the spadefoots from the Balkans traditionally attributed to P.syriacus may belong to a separate species, P.balcanicus Karaman, 1928. MtDNA (COI) differences between the spadefoots from Georgia and the Balkans exceed 10%. Found in Eastern Georgia.
Family: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Pelodytidae">Pelodytidae</tp:taxon-name-part></tp:taxon-name>AnimaliaAnuraPelodytidaeF5BF7DC7-EFFD-5AE9-B3F2-957C65364F95PelodytescaucasicusBoulenger, 1896Figs 6, 122 (Engl. Caucasian Parsley Frog, Caucasian Mud-Diver; Geo. კავკასიური ჯვარულა, კავკასიური ჯვრიანა, kavkasiuri jvriana)Taxonomic sources followed.
Boulenger (1896), Kuzmin (2012), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Frost (2026).
Records in Georgia.
Nikolskii (1913), Bannikov et al. (1977), Tarkhnishvili and Gokhelashvili (1999).
Taxonomic remark.
Biologically confirmed species. Relict species, diverged from its closest relatives since the Eocene, separated from the P.punctatus species group by broad morphological and genetic hiatus (Veith et al. 2006). Endemic of the western part of the Caucasus ecoregion.
Barcoded individuals.
CaBOL-ID 1027450 (not uploaded to BOLD; 558 bp) (Gombori). The Closest homologous sequence: P.caucasicus (NCBI MK014015; Türkiye; similarity 96.90%).
Family: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Bufonidae">Bufonidae</tp:taxon-name-part></tp:taxon-name>AnimaliaAnuraBufonidaeEE9D47C9-E04B-55E2-A258-CA6A0B6D0169Bufotesviridis(Laurenti, 1768)Figs 7, 122 (Engl. Green Toad, Geo. მწვანე გომბეშო, mtsvane gombesho)Bufoviridis Laurenti, 1768Pseudepidaleaviridis: Frost et al. (2006)Bufotesvariabilis: Bánki et al. (2026), et auct., partim.Taxonomic sources followed.
Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Frost (2026).
Taxonomic remark.
Biologically confirmed species. This is the nominal species of the B.viridis complex. The other candidate species (some with unconfirmed species status) from this complex do not exist in Georgia.
The subspecies B.v.sitibundus (green toads from the Caucasus, West Asia, and West Kazakhstan) treated as species (B.sitibundus) by Dufresnes et al. (2019). However, Frost (2026) retain the name B.viridis. Mitochondrial Cytochrome b differences (p-distance) between nominal B.viridis from Denmark and Türkiye (the latter potentially can be attributed to the subspecies B.v.sitibundus) (NCBI accession # OQ130164 and AB159263) exceed 5%, but there is no evidence of either geographic or genetic isolation of green toads from the Caucasus and Europe.
Candidate species. Sinsch et al. (2008) suggest that there is no firm evidence of separate species status for B.verrucosissimus. Mitochondrial DNA (Cytochrome b) difference between B.verrucosissimus from Georgia and nominal B.bufo from Slovenia (NCBI Accession # ON564369 and MW350662) is less than 2%. No evidence for sharp geographic border between nominal B.bufo and B.verrucosissimus in Anatolia. Moreover, recent study of Gür and Kalaycı (2025) suggests presence of toads with mitochondrial haplogroups associated with B.bufo and B. verrucosissimus in a limited geographic area of NE Türkiye. The separate species status for B.verrucosissimus still requires validation, although we list it here as a separate species following the majority of authors.
Candidate species. The separate species status of H.orientalis has been primarily supported by mitochondrial phylogenetic differentiation (Stöck et al. 2008). However, the contact zone between H.orientalis and the nominal H.arborea sensu stricto has not been sufficiently investigated using multilocus genomic data. In the absence of detailed analysis of clinal variation and potential introgression across this boundary, the strength of reproductive isolation remains incompletely demonstrated. Therefore, although H.orientalis is listed here as a distinct species following most recent taxonomic treatments, its status requires further integrative validation.
Biologically confirmed species. This species represents a well-supported evolutionary lineage within the H.arborea complex. Its species-level status is supported by multilocus genetic analyses, consistent differences in mating calls, and morphological traits (Kaya and Simmons 1999; Kaya et al. 2009; Litvinchuk et al. 2006; Stöck et al. 2008). Although H.savignyi and H.orientalis are parapatric in Eastern Georgia, and the extent of gene flow across their contact zone is not fully resolved, possible limited introgression does not preclude species status when lineages remain phenotypically and genetically distinct across their ranges. Divergence time estimates between H.savignyi and related taxa exceed 11 Mya (Kumar et al. 2017), further supporting long-term evolutionary independence.
Biologically confirmed species. Ishchenko (1978) and Tarkhnishvili et al. (1999, 2001) showed that some morphological and genetic traits have geographically distinct distribution. The latter authors accept the presence of two parapatric subspecies of this species in Georgia, R.m.macrocnemis and R.m.camerani. However, the distribution of the two subspecies does not coincide with that described by Veith et al. (2003). The taxonomic status of R.m.camerani deserves further studies on the population genomic level.
AnimaliaAnuraRanidaeD92DE3DF-15B6-59F7-9B4F-53F5497FA804Pelophylaxridibundus(Pallas, 1771)Figs 11, 122 (Engl. Lake Frog, Marsh Frog. Geo. ტბორის ბაყაყი, tboris bak’ak’i)Ranaridibunda Pallas, 1771Ranaesculenta var. ridibunda: Nikolsky (1913: 200)Taxonomic sources followed.
Kuzmin (2012), Bánki et al. (2026), Tuniyev and Ananjeva (2024), Frost (2026).
Taxonomic remark.
Biologically confirmed species. Often treated at species rank as P.bedriagae (Camerano, 1882) in parts of the literature (e.g., Bülbül et al. 2011). Recent syntheses/databases have also treated bedriagae as a subspecies within P.ridibundus (e.g., ASW running log notes “Pelophylaxbedriagae→Pelophylaxridibundusbedriagae”). It was shown that the COI sequences of individuals from Georgia are closer to P.bedriagae sensu lato from the Levant than to P.ridibundus from Europe, differing from the latter by over 5%. However, because both haplogroups are found in the type area of the species, Tarkhnishvili et al. (2024) suggest that P.bedriagae is a junior synonym of P.ridibundus, or a geographic subspecies of the latter.
Biologically confirmed species. Van der Kuyl et al. (2005) found that only two subspecies are clearly distinguishable (graeca and ibera), which is consistent with treating local Caucasus names like nikolskii within ibera. However, we maintain the subspecific name nikolskii considering phenotypic differences and distinct range of the northeastern Caucasian population.
Genetic distance from the closest species, T.kleinmanni, suggests 12–15 Mya of isolation (Kumar et al. 2017). The subspecies T.g.nikolskii is described from Western Georgia and is considered rare. T.g.ibera is widespread in Eastern Georgia. However, no data supporting or rejecting the genetic distinction between the two subspecies is available.
Family: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Emydidae">Emydidae</tp:taxon-name-part></tp:taxon-name>AnimaliaColeopteraEmydidaeA7606F26-45F7-53C4-8BC3-0AD09F5AB018Emysorbicularis(Linnaeus, 1758)Figs 13, 125 (Engl. European Marsh Turtle; Geo. ჭაობის კუ, chaobis ku)Testudoorbicularis Linnaeus, 1758Taxonomic sources followed.
Radde (1886), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Muskhelishvili (1970), Bannikov et al. (1977).
Infraspecific variation.
E.o.orbicularis (Linnaeus, 1758) – W Georgia; E.o.persica Eichwald, 1831 – E Georgia.
Taxonomic remark.
Biologically confirmed species. Subspecies E.o.colchica (Fritz, 1994) and E.o.iberica (Eichwald, 1831) (Eastern Georgia) were described for the southern Caucasus; the former was then synonymized with E.o.orbicularis and the latter with E.o.persica (Fritz et al. 1998; Bánki et al. 2026; Uetz et al. 2025).
Conant and Collins 1998, Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Reshetnikov et al, 2023.
Infraspecific variation.
T.s.elegans Wied, 1839.
Taxonomic remark.
Biologically confirmed species. The species unintentionally introduced during the last decade: a common pet species. Currently, reproducing populations exist in vicinities of Tbilisi and Batumi.
McDowell (1964), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Muskhelishvili (1970), Bannikov et al. (1977).
Infraspecific variation.
M.c.caspica (Gmelin, 1774).
Taxonomic remark.
Biologically confirmed species. The name has a priority related to the closely related species of turtles. Distinct species found from the eastern Mediterranean area to Western and Central Asia. Taxonomic status of the western subspecies of this polytypic species, M.c.rivulata, was elevated to a separate species. Found in SE Georgia.
Biologically confirmed species. Large settlements of Eastern Georgia, Kutaisi and Sokhumi in Western Georgia. The natural range covers the extreme south-east of the country.
Family: AngiudaeAnimaliaSquamataAnguidae3295019E-80F9-54AE-BB2B-AF3F689084F2Anguiscolchica(Nordmann, 1840)Figs 17–18, 131 (Engl. Colchis Slow Worm; Geo: კოლხური ბოხმეჭა, ძლოკვი, bokhmecha).Otophiseryx var. colchica Nordmann, 1840Anguisfragilis var. colchica: Nikolskii (1913: 40)Anguisfragilisfragilis: Muskhelishvili (1970: 39)Anguisfragiliscolchicus: Bannikov et al. (1977: 142); Dunayev and Orlova (2017), et auct.Taxonomic sources followed.
Gvoždík et al. (2010), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Muskhelishvili (1970), Bannikov et al. (1977); Gvoždík et al. (2010).
Taxonomic remark.
Candidate species. The species belongs to a widespread western Eurasian complex A.fragilis. Gvoždík et al. (2010) elevated the Caucasian form of slow worm (A.f.colchica) to a species level. This species is likely allopatric with A.graeca and A.fragilis. Mitochondrial DNA (Cytochrome b) difference between A.colchica and its closest relatives, A.graeca and A.fragilis (NCBI Accession # MK107684, OP493517, and MN122840) varies between 6.5-8%.
AnimaliaSquamataAnguidaeC80EA791-28E9-5123-8B23-046E0639B272Pseudopusapodus(Pallas, 1775)Figs 19–20, 126 (Engl. European Glass Lizard, Scheltopusik; Geo: გველხოკერა, ხოკერა, gvelkhokera)Lacertaapoda Pallas, 1775Ophisaurusapus: Nikolskii (1913: 36)Ophisaurusapodus: Terentyev and Chernov (1940: 111); Muskhelishvili (1970: 31); Bannikov et al. (1977: 140), et auct.Taxonomic sources followed.
Mertens and Müller (1928), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Muskhelishvili (1970), Bannikov et al. (1977).
Infraspecific variation.
P.a.apodus Pallas, 1775.
Taxonomic remark.
Biologically confirmed species. A distinct species from a monospecific genus, with a broad western Eurasian distribution. Found only in Eastern Georgia.
Barcoded individuals.
CaBOL-1027419 (BOLD:AEO0661; Kesalo). Closest homologous sequences: P.apodus with a private status (Tajikistan; similarity 100%); P.apodus (NCBI NC_071252; Georgia; similarity 100%). Next closest taxon: A.graeca Bedriaga, 1881 (NCBI NC_048977; Greece; similarity 88.30%).
Family: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Agamidae">Agamidae</tp:taxon-name-part></tp:taxon-name>AnimaliaSquamataAgamidae3FAB9F5B-F582-50B8-A700-CD4E81C00050Paralaudakiacaucasia(Eichwald, 1831)Figs 21–22Stelliocaucasius Eichwald, 1831Stelliocaucasicus: Ananjeva et al. (1988)Agamacaucasica: Nikolskii (1913: 25), Muskhelishvili (1970: 19), Bannikov et al. (1977: 111) et auct.Laudakiacaucasia: Ananjeva et al. (2006); Speybroeck et al. (2020)Taxonomic sources followed.
Baig et al. (2012), Bánki et al. (2026), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Muskhelishvili (1970), Bannikov et al. (1977).
Taxonomic remark.
Biologically confirmed species. Differs from the closest species, P.microlepis (Blanford, 1874), by a broad genetic and phenotypic hiatus (estimated divergence time 9-40 Mya; Kumar et al. 2017). A distinct species with a broad western Asian distribution. Rocky areas throughout the eastern and southern Georgia, locally in the Western Georgia.
Boulenger (1891), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Muskhelishvili (1970), Bannikov et al. (1977).
Infraspecific variation.
E.s.princeps (Eichwald, 1839) – Caucasus/Iran form. Sometimes treated at species rank as Novoeumecesprinceps (Eichwald, 1839) (e.g., Griffith et al. 2000).
Taxonomic remark.
Biologically confirmed species. Distinct species found in West Asia and North Africa. The subspecies princeps is found in the Caucasus and Iran. Found in the extreme south-east of Georgia.
Fitzinger (1823), Bánki et al. (2026), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Taxonomic remark.
Biologically confirmed species. Distinct species with a broad range in Western and Central Asia. Bakradze and Darevsky (1973) reported finding this species from Vashlovani Nature Reserve (Eastern Georgia). No further confirmation is available.
Kessler (1872), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Muskhelishvili (1970), Bannikov et al. (1977).
Infraspecific variation.
E.v.caucasia Lantz, 1928.
Taxonomic remark.
Biologically confirmed species. This is a distinct species with a broad range in the Caucasus and Central Asia. Fairly common in the south-east of Georgia. Simultaneously, judging from substantial differences in the structure of COI gene, E.v.caucasica may be a species separate from E.velox. Infraspecific taxonomy requires detailed population-level study.
Barcoded individuals.
CaBOL-IDs 1010909, 1010910 (BOLD:AAJ1050; Pantishara Canyon; similarity 99.01%). Closest homologous sequences: E.velox ssp. 2RuHF (BOLD:ACE8498; W Kazakhstan; similarity 93.49%); E.v.caucasia with a private status (Azerbaijan, Salyan; similarity 99–99.34%); E.velox (NCBI HQ733909.1; China, Shengdu, Sychuan; similarity 91.6%).
Ménétries (1832), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Muskhelishvili (1970), Bannikov et al. (1977); Tarkhnishvili et al. (2022).
Infraspecific variation.
O.e.elegans Ménétriés, 1832.
Taxonomic remark.
Biologically confirmed species. A distinct species with a broad western Asian range. Sporadically found in SE Georgia. Sequences of this species deposited in NCBI GenBank are highly divergent, which can lead to the revision of the taxonomy of O.elegans sensu lato.
Barcoded individuals.
CaBOL-IDs 1010913, 1027423 (BOLD:AAK5540; Taribana; similarity 99.48%). Closest homologous sequences: O.elegans (BOLD:AAK5540; Azerbaijan; similarity 99.82–100%); O.elegans (MN356545; Azerbaijan, Zarat; similarity 99.85%); O.elegans (NCBI MN536543; Türkiye, Akseki) has only 88.11% similarity with the barcoded specimen.
Linnaeus (1758), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Muskhelishvili (1970), Bannikov et al. (1977), Yablokov (ed.) (1976), Tarkhnishvili et al. (2002).
Infraspecific variation.
L.a.ioriensis Peters & Muskhelishvili, 1968 (Eastern Georgia); L.a.grusinica Peters, 1960 (Western Georgia); L.a.brevicaudata Peters, 1958 (southern and central Georgia).
Taxonomic remark.
Biologically confirmed species. A distinct species with a broad western Eurasian range. Everywhere in Western Georgia, at the altitude above 600–800 m in Eastern Georgia.
Barcoded individuals.
CaBOL-IDs 1010922 (BOLD:AAK8295; Tskruti), 1027422 (BOLD:AFT4708; Tianeti), similarity 95.06%. Closest homologous sequences: CaBOL-ID 1010922 – L.a.brevicaudata with a private status (Armenia; similarity 99.47%), 1027422 – L.a.exigua with a private status (Ukraine; similarity 95.87%).
AnimaliaSquamataLacertidaeDC3A4E3D-B1F4-5EE3-AFC5-EFCAE7631FC3LacertastrigataEichwald, 1831Lacertastrigata Eichwald, 1831Lacertaviridis var. strigata: Boulenger (1881)Taxonomic sources followed.
Eichwald (1831), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Muskhelishvili (1970), Bannikov et al. (1977), Tarkhnishvili et al. (2002).
Taxonomic remark.
Biologically confirmed species. A distinct species found throughout West Asia and the Caucasus, separated from its closest relatives > 10 Mya (Kumar et al. 2017). Found in semiarid habitats throughout the country.
Barcoded individuals.
CaBOL-ID 1027421 (BOLD:AFT6170; Chankaani). The closest homologous sequence: L.strigata (NCBI OM267848; Georgia, Tianeti; similarity 100%). Next closest taxa: L.agilisexigua with a private status (Ukraine; similarity 88.62%); L.a.boemica (NCBI OM267791; unknown location; similarity 90.52%).
Suchow (1948), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Muskhelishvili (1970), Bannikov et al. (1977), Tarkhnishvili et al. (2002).
Infraspecific variation.
L.v.media Lantz & Cyrén, 1920.
Taxonomic remark.
Candidate species. The separate status is indirectly supported by phylogenetic differences from the closest relatives and parapatric distribution in western Anatolia; however, a clear contact zone between this species and the species of L.trilineata group remains unexplored (Ahmadzadeh et al. 2013). Belongs to a monophyletic L.trilineata species complex. Found in Central and Eastern Anatolia, Iran, and the Caucasus. Widespread in semiarid habitats throughout Eastern Georgia and sporadically in the northwestern Georgia. Mitochondrial DNA (Cytochrome b) difference between L.media and its closest relative, L.trilineata (NCBI Accession # MZ06558 and MK330106) reaches 6.5%.
Barcoded individuals.
CaBOL-IDs 1010923 (Pantishara Gorge), 1010924 (Tmogvi) (BOLD:AAJ3156; similarity 100%). The closest homologous sequences: L.trilineatamedia with a private status (Türkiye ; similarity 99.69%); L.media (NCBI OM267813; Gelendzhik, Russia; similarity 99.68%). Next closest taxon: L.trilineata (NCBI MN015090; Milos, Greece; similarity 92.68%).
AnimaliaSquamataLacertidaeB11B03CA-DCC8-588E-B1EC-F63195B276B3Phoenicolacertalaevis(Gray, 1838)Lacertalaevis Gray, 1838Archaeolacertalaevis: Sindaco et al. (2000).Taxonomic sources followed.
Arnold et al. (2007), Bánki et al. (2026), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Tarkhnishvili et al. (2017a).
Taxonomic remark.
Biologically validates species status. Found at a single location in Georgia, genetically close to populations from southern Türkiye. Unintentionally introduced species.
Arribas et al. (2022a), Bánki et al. (2026), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Darevsky (1967), Bannikov et al. (1977), Murphy et al. (2000), Tarkhnishvili and Iankoshvili (2023).
Taxonomic remark.
Candidate species. Formerly considered to be a geographic subspecies of D.parvula. The range extends over SW Georgia. Mitochondrial DNA (Cytochrome b) difference between D.ajarica from Georgia and D.parvula from NE Turkey (NCBI Accession # U88609 and MK092688) reaches 3.7%. The transition zone between species of the complex D.parvula is not studied on the population level, hence we cannot reject free gene flow across the transition zones given current knowledge.
Barcoded individuals.
CaBOL-IDs 1027440, 1007137, 1007138, 1007139 (Shuakhevi). The closest homologous sequences: D.ajarica (NCBI MN015094; deposited as D.parvula, Ajaristskali, Georgia; similarity 99.21%). Next closest taxon: D.rudis (NCBI NC_046011; unknown location; similarity 89.87%).
AnimaliaSquamataLacertidaeC4F5B8AE-7672-5BBD-91A3-1B2EF1466277Darevskiaalpina(Darevsky, 1967)Lacertacaucasicaalpina Darevsky, 1967Lacertaalpina Darevsky, 1967: Murphy et al. (2000), et auct.Taxonomic sources followed.
Arribas (1999), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Taxonomic remark.
May be a hybrid species between species of the complexes “caucasica” and “saxicola” (Murphy 2000; Tarkhnishvili 2012).
Arribas (1999), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Darevsky (1967), Bannikov et al. (1977), Murphy et al. (2000), Tarkhnishvili and Iankoshvili (2023).
Taxonomic remark.
Conventional species status. Parthenogenetic form, supposedly a hybrid between males of D.valentini and females of D.mixta (Murphy et al. 2000; Yanchukov et al. 2022), or a backcross between D.valentini and another parthenogen (Tarkhnishvili et al. 2017b, 2020), with a small range in southern Georgia, and no remarkable differences between populations.
Barcoded individuals.
CaBOL-ID 1027438 (BOLD:AAE7437; Akhalkalaki). The closest homologous sequences: D.armeniaca (BOLD:AAE7437; NE Armenia, NE and E Türkiye; similarity 100%); D.armeniaca (NCBI NC_046006; Saghamo Lake; similarity 100%), D.mixta (NC_046008; Baniskhevi; similarity 99.85%).
AnimaliaSquamataLacertidae2A2D4D93-D437-513D-9C0D-7AED535685DBDarevskiabrauneri(Méhely, 1909)Lacertasaxicola var. brauneri Méhely, 1909.Lacertasaxicolabrauneri: Terentyev and Chernov (1940: 98); Darevsky (1967: 43).Taxonomic sources followed.
Arribas (1999), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Darevsky (1967), Tarkhnishvili et al. (2016).
Candidate species. Member of the Darevskia (saxicola) complex; historically treated as a subspecies in broad treatments (e.g., Bannikov et al. 1977). Although currently species status of D.brauneri is widely accepted, it is not clear to what extent there are stepped genetic and phenotypic clines separating it from D.saxicola or D.szczerbaki. Tarkhnishvili et al. (2017b) showed that mitochondrial haplogroups from some populations of nominal D.brauneri from Georgia are closer to D.saxicola from the North Caucasus than to formally conspecific populations. Distribution restricted to the western and central Greater Caucasus.
Tuniyev et al. (2023) described a population near Ertso Lake (Georgia) as D.arribasi. The diagnostic scalation and morphometric characters reported in that description fall within variation documented for nominal D.brauneri from the western Caucasus (Darevsky 1967) and for D.brauneri populations from the Enguri and Tekhuri basins in Western Georgia (Tarkhnishvili et al. 2016). Based on BLAST comparisons of available mtDNA/mitogenome data (e.g., D.arribasi OQ414972 versus D.brauneri NC_050005/NC_046012), the divergence is low (≈2% scale), and – until clear evidence for genetic isolation is demonstrated – we treat the Ertso population as subspecies D.b.arribasi (not a separate species). Subspecies D.b.myusserica occurs locally in the north-western Georgia. More generally, contact zones and isolation among several nominal taxa in the D.saxicola complex remain insufficiently tested.
AnimaliaSquamataLacertidae72E9DA08-8AD5-58E2-9113-47C994A7AA15Darevskiacaucasica(Méhely, 1909)Lacertacaucasica Méhely, 1909Lacertamuralis var. caucasica: Boulenger (1913)Lacertasaxicolacaucasica: Terentyev and Chernov (1940: 97)Lacertacaucasica: Nikolsky (1913: 82), Darevsky (1967: 111), et auct.Taxonomic sources followed.
Arribas (1999), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Darevsky (1967), Muskhelishvili (1970); Bannikov et al. (1977); Doronin et al. (2021), Tarkhnishvili and Iankoshvili (2023).
Taxonomic remark.
A recent phylogeographic study by Doronin et al. (2021) suggests strong genetic differences between D.caucasica from Georgia and the populations from the northern Caucasus. A recent genomic study (Tarkhnishvili et al. 2025) supports the presence of a few genetically distant units within D.caucasica.
Barcoded individuals.
CaBOL-IDs 1027425 (BOLD:AGJ4669; Zakagori Fortress), 1027426 (BOLD:AAJ4512; Mount Tbatana), 1027427 (BOLD:AGJ4670; Lebarde), similarity 91.59–95.29%. The closest homologous sequences: D.caucasica (NCBI NC_049999; Truso, Georgia; similarity 95.29–100%). The next closest taxa: D.mixta (NCBI NC_046008; Baniskhevi; similarity 94.98–96.05%).
Arribas (1999), Bánki et al. (2026), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Darevsky and Vedmederja (1977), Darevsky and Tuniyev (1997), Bischoff (2007), Tarkhnishvili and Iankoshvili (2023).
Records in Georgia.
Darevsky and Vedmederja (1977), Darevsky and Tuniyev (1997), Bischoff (2007), Tarkhnishvili and Iankoshvili (2023).
Infraspecific variation.
D.c.dryada (Darevsky & Tuniyev, 1997) comb.nov. — although we do not find sufficient evidence for recognising this form as a species, considering some phenotypic differences from typical D.clarkorum, we suggest treating it as a geographic subspecies.
Taxonomic remark.
Biologically confirmed species. A phenotypically and genetically distinct species geographically isolated from its closest relatives, D.mixta and D.caucasica (Tarkhnishvili et al. 2025). Nominal D.dryada is suggested to be recognised as a species, based on the phenotypic analysis (Arribas et al. 2021), although Schmidtler et al. (2002) treat it as an ecological race. Genetic differences based on the mitochondrial Cytochrome b gene between nominal D.clarkorum from Türkiye and nominal D. “dryada” from SW Georgia (Tarkhnishvili et al. 2025) and nominal D.clarkorum from NW Türkiye vary between 99.76 and 96.22%. For these reasons, we decide to treat the form as geographic subspecies of D.clarkorum.
Barcoded individuals.
CaBOL-IDs 1027431 (BOLD:AGJ4668), 1007143 (BOLD:AGJ4668) (Charnali riv. Valley). The closest homologous sequences: D.clarkorumdryada (NCBI NC_050002; deposited as D.clarkorum, Charnali; similarity 98.74–98.9%). Next closest taxon: D.armeniaca (BOLD:AAE7437; Armenia; similarity 95.74–95.81).
AnimaliaSquamataLacertidaeB950A2FD-0623-53EB-9C30-BD132F9AB664Darevskiadaghestanica(Darevsky, 1967)Lacertasaxicoladaghestanica Darevsky, 1967Lacertacaucasica: Nikolskii (1913: 82), partimLacertacaucasicadaghestanica: Bannikov et al. (1977: 214), et auct.Lacertadaghestanica: Murphy et al. (2000), et auct.Taxonomic sources followed.
Arribas (1999), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Darevsky (1967), Muskhelishvili (1970), Bannikov et al. (1977), Doronin et al. (2021), Tarkhnishvili and Iankoshvili (2023).
Infraspecific variation.
Not applicable, although deep genetic differences among the populations are shown (Doronin et al. 2021).
Taxonomic remark.
Commonly considered as a sister species of D.caucasica or even its subspecies (Darevsky 1967), however, this is a basal lineage equidistant from D.caucasica, D.mixta, and D.clarkorum (Murtskhvaladze et al. 2020).
Arribas (1999), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Darevsky (1967), Muskhelishvili (1970), Bannikov et al. (1977), Murphy et al. (2000), Tarkhnishvili et al. (2010).
Taxonomic remark.
A parthenogenetic hybrid form. The parental species D.mixta and D.portschinskii (Murphy et al. 2000; Tarkhnishvili et al. 2017, 2020; Yanchukov et al. 2022). The mitochondrial genome is shared with D.mixta and parthenogenetic D.armeniaca (Murphy et al. 2000).
Barcoded individuals.
CaBOL-ID 1027435 (BOLD:AAE7437; Kojori). The closest homologous sequences: D.dahli (NCBI NC_046007; Kojori, Georgia; similarity 99.85%); D.armeniaca (BOLD:AAE7437; Armenia; similarity 99.85%). Both D.dahli and D.armeniaca are hybrid parthenogens, and D.mixta is their matrilineal ancestor, hence very high similarity of mtDNA among these species is not surprising.
AnimaliaSquamataLacertidaeEA34C86D-6876-5692-A9B6-A0347D8A548FDarevskiaderjugini(Nikolskii, 1898)Lacertaderjugini Nikolskii, 1898Lacertaderjugini: Darevsky (1967: 8); Muskhelishvili (1970: 79); Bannikov et al. (1977: 207); Murphy et al. (2000), et auct.Taxonomic sources followed.
Arribas (1999), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Muskhelishvili (1970), Bannikov et al. (1977), Tarkhnishvili and Iankoshvili (2023).
Biologically confirmed species. Sympatric with other Darevskia, remaining distinct phenotypically and genetically. Although phenotypic diversity is high, genetic diversity within this species is modest (our data). This is one of two ground-dwelling species of the genus Darevskia from Georgia. Found throughout Western Georgia and the forest belt of Eastern Georgia.
Arribas (1999), Bánki et al. (2026), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Darevsky (1967), Bannikov et al. (1977), Tarkhnishvili and Iankoshvili (2023).
Taxonomic remark.
Biologically validates species status. In spite of little genetic (mtDNA) differences from D.caucasica, forms a narrow contact zone with this species in the Greater Caucasus. This species was previously thought to be a hybrid between D.derjugini and D.saxicola. The vernacular name “Ajarian lizard” should be changed to “Georgian lizard”, because the species is endemic to Western Georgia, but is absent from the Ajara region.
Biologically confirmed species. This species is closely related to D.rudis, D.obscura, and D.valentini, but is substantially smaller and forms a narrow contact zone with all three species. Hybrid zone with D.obscura present in the Tana River valley (Tarkhnishvili et al. 2013). Endemic of the central Lesser Caucasus.
Barcoded specimens: CaBOL-ID 1027436 (BOLD:AGJ4671; Kojori). The closest homologous sequences: D.portschinskii (BOLD:AAJ4522; Armenia; similarity 97.38%). Next closest taxon: D.obscura (NCBI NC_046011; deposited as D.rudis. Akhaldaba, Georgia; similarity 94.07%).
Arribas (1999), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Darevsky (1967), Muskhelishvili (1970), Tarkhnishvili and Iankoshvili (2023).
Infraspecific variation.
D.p.praticola (Eversmann, 1834) – Eastern and central Georgia.
Taxonomic remark.
Biologically confirmed species. A widespread, distinct species with a fragmented range; Georgian populations referred here to D.praticola represent the eastern/nominotypical lineage. Pontic populations from NW Georgia are treated separately below as D.pontica (candidate species), reflecting ongoing disagreement in the literature and databases about rank.
AnimaliaSquamataLacertidae7BF14684-913C-51A6-9B12-D5A857E7334DDarevskiapontica(Lantz & Cyrén, 1918)Lacertapraticolapontica Lantz & Cyrén, 1918Lacertapraticola: Nikolskii (1913: 55), Terentyev and Chernov (1940: 96), Darevsky (1967), et auct., partim.Darevskiapraticolapontica: Bannikov et al. (1977: 209), et auct.Taxonomic sources followed.
Arribas (1999), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Darevsky (1967), Muskhelishvili (1970), Tarkhnishvili and Iankoshvili (2023).
Taxonomic remark.
Candidate species. Pontic populations are phenotypically diagnosable in some characters, but published discussions also note very small genetic differentiation and lack of demonstrated effective genetic isolation from D.praticola sensu lato; rank remains inconsistently treated (species vs subspecies) across modern sources and databases.
Gabelaia et al. (2018), Arribas et al. (2022b), Bánki et al. (2026), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Darevsky (1967), Muskhelishvili (1970), Tarkhnishvili and Iankoshvili (2023).
Infraspecific variation.
D.o.obscura (Lantz & Cyrén, 1936) – upper flow of the Mtkvari (Kura) River from the westernmost part of Trialeti Mountains, eastern part of Meskheti Mountains, and mountains surrounding upper currents of Mtkavi (Kura) River and its tributaties; D.o.macromaculata (Darevsky, 1967) – western part of Javakheti Plateau in Georgia; D.o.bischoffi (Böhme & Budak, 1977)– SW Georgia/adjacent NE Turkey; D.o.chechenica (Eiselt & Darevsky, 1991) comb. nov. — Eastern Greater Caucasus (east of Enguri valley); D.o.svanetica (Darevsky & Eiselt, 1980) comb. nov. — western Greater Caucasus (e.g., upper Enguri basin).
Taxonomic remark.
Candidate species. D.obscura was elevated from within the D.rudis complex mainly based on a distinct mitochondrial lineage separating populations of the Caucasus/ Georgia from D.rudis found in the northern and central Anatolia (Candan et al. 2021; Arribas et al. 2022b). Following this logic, and based on the mitochondrial phylogeny (Tarkhnishvili et al. 2013; Candan et al. 2021) all populations of D.rudis s.l. from Georgia, including subspecies chechenica and svanetica, should be attributed to D.obscura and not to D.rudis s. str. Remarkably, this taxonomy does not rely on the original vewpoint of Lantz and Cyrén (1936), who described the lizards from Borjomi Gorge as a new subspecies of the rock lizard, Lacertasaxicola, based on their phenotypic differences and not associating them with D.rudis from other parts of Georgia. Darevsky (1967) placed this form into a broader polytypic D.rudis. Tarkhnishvili et al. (2013) showed monophyletic mitochondrial origin of D.o.obscura + D.o.macromaculata related to other subspecies of D.rudis s.l. from Georgia, and Gabelaia et al. (2018) suggested to elevate its status to a species, based on its phenotypic differences from other subspecies of D.rudis s.l. The current species status of polytypic D.obscura is, however, based solely on the differences of mitochondrial DNA of the lizards from the Caucasus from the mitochondrial DNA of D.rudis s.str. from Anatolia, without reference to morphology or nuclear genome. There is no evidence for (or against) gene flow across a contact zone between D.obscura and D.rudis. Consequently, we suggest candidate species status of this form, which may require revision in the future.
Barcoded specimens: CaBOL-1010930 (BOLD:AAF7228; Akhalkalaki, Saaghdgomo Mt.). The closest homologous sequences: D. “rudis” (BOLD:AAF7228; unknown location; similarity 100%); D.obscura (NCBI NC_046011; Borjomi Gorge, deposited as D.rudis; similarity 99.7%).
AnimaliaSquamataLacertidae91353700-0F1F-5142-B0BF-5EF76F347540Darevskiaraddei(Boettger, 1892)Lacertamuralis var. raddei Boettger, 1892Lacertacaucasica var. tenuis: Nikolskii (1913: 86), partimLacertasaxicolaraddei Darevsky (1967: 83)Lacertaraddei Bannikov et al. (1977: 217), et auct.Taxonomic sources followed.
Arribas (1999), Bánki et al. (2026), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Darevsky (1967), Tarkhnishvili and Iankoshvili (2023).
Infraspecific variation.
D.r.nairensis (Boettger, 1892) comb. rev.
Taxonomic remark.
Biologically confirmed species. Genetic analyses (Grechko et al. 2007) do not support clear species-level separation between D.raddei and D.nairensis. Mitochondrial DNA data indicate that the Georgian populations are genetically closer to D.r.nairensis than to nominal D.r.raddei from Armenia (Tarkhnishvili et al. 2020). Given the lack of demonstrated reproductive isolation and limited divergence, we retain nairensis at the subspecific rank. In Georgia, the species is restricted to the upper reaches of the Mtkvari (Kura) River, near the border with Türkiye.
AnimaliaSquamataLacertidaeF0820129-BDBC-5B46-8698-BDF916102F3BDarevskiavalentini(Boettger, 1892)Lacertamuralis var. valentini Boettger, 1892Lacertasaxicolavalentini: Méhely (1909); Nikolskii (1913: 77); Darevsky (1967: 93), et auct.Lacertavalentini: Engelmann et al. (1993); Murphy et al. (2000), et auct.Taxonomic sources followed.
Arribas (1999), Bánki et al. (2026), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Darevsky (1967), Bannikov et al. (1977), Tarkhnishvili and Iankoshvili (2023).
Infraspecific variation.
D.v.valentini — nominotypical subspecies.
Taxonomic remark.
Biologically confirmed species. This species is closely related with D.rudis, D.obscura and D.portschinskii, but is phenotypically distinct and forms a narrow contact zone with D.obscura and D.portschinskii. There is a broad gene introgression between this species, D.portschinskii, and D.obscura across their contact zones (Tarkhnishvili et al. 2013).
Stejneger (1907), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Muskhelishvili (1970), Bannikov et al. (1977), Iankoshvili and Tarkhnishvili (2021).
Infraspecific variation.
N.n.scutata (Pallas, 1771).
Taxonomic remark.
Biologically confirmed species. Distinct species, separated from its closest relative, N.helvetica, with a narrow contact zone (Kindler et al. 2017). All grass snakes from Georgia, including striped morph described as N.n.persa (Pallas, 1814) belong to the mitochondrial lineage of subspecies N.n.scutata, which is also present in Eastern Europe, Türkiye, and Central Asia (Kindler et al. 2013). Nominal species Natrix “megalocephala” (Orlov and Tuniyev, 1987) was synonymised with N.n.scutata (Kindler et al. 2013), based on the lack of genetic divergence between this form and ring snakes from the same geographic area. Specimens that have diagnostic characters of nominal N. “megalocephala” are occasionally found in populations of typical N.n.scutata; we suggest treat it as a morph N.n.scutata f. megalocephala comb. nov. (regular font is used as suggested in Mebert et al. 2025c).
Merrem (1820), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Muskhelishvili (1970), Bannikov et al. (1977), Iankoshvili and Tarkhnishvili (2021).
Taxonomic remark.
Biologically confirmed species. Genetically, dice snakes from eastern and southern Georgia belong to the “Caspian lineage” closer to the Central Asian than to the Anatolian populations (Jablonski et al. 2024). Genetic affinities of the snakes from the west of the country remain unknown.
Barcoded individuals.
CaBOL-ID 1027402 (BOLD:AEO3649; Akhalkalaki). The closest sequences in BOLD Systems: N.natrix with a private status (Unknown location; similarity 99.08%). Second-best match: N.tessellata with a private status (Tajikistan; similarity 98.92%). NCBI: N.tessellata (NCBI JN871607; Ukraine; similarity 99.84%). Next closest taxon: N.helvetica (NCBI MN993168; Austria; similarity 89.06%).
Biologically confirmed species. Smooth snakes from the Caucasus belong to the “eastern clade”, genetically closest to the snakes from Central Europe (Sztencel-Jablonka et al. 2015), but non-named to the present time.
AnimaliaSquamataColubridae153F63FB-C353-55BB-AFAC-7C2DBF8E79D2ElapheurarticaJablonski, Kukushkin, Avcı, Bunyatova, Ilgaz, Tuniyev, & Jandzik, 2019Elapheurartica Jablonski et al., 2019Coluberquatuorlineatus: Nikolskii (1913: 146), partim.Elaphequatuorlineatasauromates: Terentyev and Chernov (1940: 146), Muskhelishvili (1970: 163), Bannikov et al. (1977: 287), et auct. partim.Elaphesauromates: Lenk et al. (2001), et auct. partim.Taxonomic sources followed.
Jablonski et al. (2019), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Muskhelishvili (1970), Bannikov et al. (1977), Iankoshvili and Tarkhnishvili (2021).
Taxonomic remark.
Candidate species. Formerly considered as subspecies E.quatuorlineatasauromates, after elevation of the status of this subspecies, between 2001 and 2019 referred to as E.sauromates; elevated to a separate species as a result of genetic study of Jablonski et al. (2019). The two species have coinciding differences in mitochondrial and nuclear genes. Mitochondrial DNA (Cytochrome b) difference between E.urartica and its closest relatives, E.druzei and E.sauromates (NCBI Accession # NC073569, NC073568, and MK070315) varies between 4.3–5.6%. Yet, the transition zone of 600 km width between populations of E.urartica and E.sauromates in the north-western Caspian area was never studied explicitly, including genetic markers or phenotypic characters.
AnimaliaSquamataColubridaeD24B7A98-81D4-5D30-BC41-4A3B36B80AC2Zamenislongissimus(Laurenti, 1768)Natrixlongissima Laurenti, 1768Coluberlongissimus: Nikolskii (1913: 154)Elaphelongissima: Terentyev and Chernov (1940: 144), et auct.Elaphelongissimalongissima: Muskhelishvili (1970: 163).Taxonomic sources followed.
Utiger et al. (2002), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Muskhelishvili (1970), Bannikov et al. (1977), Iankoshvili and Tarkhnishvili (2021).
Taxonomic remark.
Biologically confirmed species. Published phylogeographic work recognizes multiple mitochondrial clades within the species; one (“Asian clade”) is native to the eastern Black Sea area (Musilová et al. 2010).
AnimaliaSquamataColubridaeC2CF8336-C3C3-574D-A521-0009881EC31AZamenishohenackeri(Strauch, 1873)Coluberhohenackeri Strauch, 1873Elaphehohenackeri: Terentyev and Chernov (1940: 143); Muskhelishvili (1970: 159); Bannikov et al. (1977: 280), et auct.Taxonomic sources followed.
Utiger et al. (2002), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Muskhelishvili (1970), Bannikov et al. (1977), Iankoshvili and Tarkhnishvili (2021).
Infraspecific variation.
Z.h.hohenackeri (Strauch, 1873) — subspecies occurring in Georgia (the Reptile Database also lists Z.h.lyciensisHofmann et al. 2018 and Z.h.aurica (Werner 1898) for parts of the wider range).
Taxonomic remark.
Biologically confirmed species. Genetically close to other populations from the Caucasus and eastern Türkiye.
Nagy et al. (2004), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Muskhelishvili (1970), Bannikov et al. (1977), Iankoshvili and Tarkhnishvili (2021).
Taxonomic remark.
Candidate species. This snake belongs to the eastern species group of a large superspecies D.jugularis (Nagy et al. 2010). Formerly was considered to be a geographic subspecies of D.jugularis. The species is not isolated geographically from the nominal form of D.jugularis. Mitochondrial DNA (Cytochrome b) difference between D.schmidti and its closest relative, D.caspius (NCBI Accession # AY486923 and PP378902) reaches 6.5% and p-distance based on mitochondrial cytochrome b structure exceeds 10%, and reaches 6–7% between D.schmidti and D.caspius. Yet, populations of the latter two species may have a contact zone in the western Caspian area, and population-level studies are needed to validate their separate species status.
AnimaliaSquamataColubridae290D3D4B-3C97-522A-A0E2-7FAFC19A2770Dolichophiscaspius(Gmelin, 1789)Colubercaspius Gmelin, 1789Zamenisgemonensis var. caspius: Nikolskii (1913: 136)Coluberjugulariscaspius: Terentyev and Chernov (1940: 136), et auct.Taxonomic sources followed.
Nagy et al. (2004), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Bannikov et al. (1977).
Taxonomic remark.
Candidate species. This snake belongs to the eastern species group of a large superspecies D.jugularis (Nagy et al. 2010). Formerly considered to be a geographic subspecies of D.jugularis. However, it is isolated geographically from the nominal form of D.jugularis. Reported mitochondrial differentiation from the geographically distant population of D.schmidti (cytochrome b) is ~6–7%. Yet, populations of these taxa may approach or contact in the western Caspian area, and population-level studies are needed to validate their separate species status.
AnimaliaSquamataColubridaeDE289DAC-273C-51F7-9C1D-FC4CAD516709Hemorrhoisravergieri(Ménétries, 1832)Coluberravergieri Ménétries, 1832Zamenisravergieri: Nikolskii (1913: 142)Coluberravergieri: Terentyev and Chernov (1940: 141), et auct.Coluberravergieriravergieri: Muskhelishvili (1970: 153)Taxonomic sources followed.
Nagy et al. (2004), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Muskhelishvili (1970), Bannikov et al. (1977), Iankoshvili and Tarkhnishvili (2021).
Taxonomic remark.
Biologically confirmed species. The species with a broad distribution in the Caucasus and Central Asia.
Barcoded individuals.
CaBOL-ID 1007153, 1027408 (BOLD:AGI4826; Tmogvi; similarity 100%). Closest homologous sequences: H.ravergieri with a private status (Tajikistan; similarity 96.9%); H.ravergieri (NCBI OQ417080; NW China; similarity 96.54%). Next closest taxon: H.hippocrepis (Linnaeus, 1758) (NCBI OZ203535; W Mediterranean; similarity 88.28%).
AnimaliaSquamataColubridae02688091-9666-504E-8489-28992E16333EPlatycepsnajadum(Eichwald, 1831)Tyrianajadum Eichwald, 1831Zamenisdahlii: Nikolskii (1913: 139)Colubernajadum: Terentyev and Chernov (1940: 137); Muskhelishvili (1970: 147); Bannikov et al. (1977: 262), et auct.Taxonomic sources followed.
Schätti and Monsch (2004), Bánki et al. (2026), Speybroeck et al. (2020), Tuniyev and Ananjeva (2024), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Muskhelishvili (1970), Bannikov et al. (1977), Iankoshvili and Tarkhnishvili (2021). Original description name: Tyrianajadum (Eichwald, 1831).
Biologically confirmed species. Belongs to the most basal lineage of dwarf snakes (Pyron et al. 2013). Species with a broad Eastern Mediterranean range. Eastern and central Georgia.
Candidate species. Belongs to a widespread Mediterranean species group M.monspessulanus. Mitochondrial DNA (Cytochrome b) difference between M.insignitus and its M.monspessulanus (NCBI Accession # AY612033 and AY188029) is below 5% (and 8.4% when Cyt b and 16S combined cit. in Carranza, Arnold and Pleguezuelos 2006). The transition area between M.monspessulanus and M.insignitus in the northeastern Morocco never studied on population level, and at least one hybrid has been reported (Jimenez-Cazalla 2012). Potentially, free gene flow across the transition zone cannot be excluded on a priori basis.
Barcoded individuals.
CaBOL-ID 1027416 (BOLD:AGH9517; Chachuna Managed Reserve). There were no sequences of properly identified M.insignitus in the BOLD Systems. The closest sequences: Malpolon sp. (BOLD:AEO4390; Algeria; similarity 94.82%). Malpolon sp. (NCBI ON943657; Algeria; similarity 94.87%).
Family: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Viperidae">Viperidae</tp:taxon-name-part></tp:taxon-name>AnimaliaSquamataViperidae716854A3-DB20-5137-B285-A3C8F847D839Macroviperalebetinus(Linnaeus, 1758)Coluberlebetinus Linnaeus, 1758Viperalebetina: Nikolskii (1913: 184), et auct.Viperalebetinaobtusa: Terentyev and Chernov (1940: 163); Muskhelishvili (1970: 200); Bannikov et al. (1977: 328), et auct.Daboialebetina: Engelmann et al. (1993)Macroviperalebetina: Tuniyev et al. (2019); Tuniyev and Ananjeva (2024)Taxonomic sources followed.
Fretey (2019), Bánki et al. (2026), Speybroeck et al. (2020), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Muskhelishvili (1970), Bannikov et al. (1977), Iankoshvili and Tarkhnishvili (2021).
Infraspecific variation.
M.l.obtusa (Dwigubsky, 1832).
Taxonomic remark.
Biologically confirmed species. Oviparous. A venomous snake, and the most medically significant viper in Georgia. In recent decades, the range has expanded from extreme south-eastern Georgia to Alazani Valley and Tbilisi City in the north and northwest.
Barcoded individuals.
CaBOL-IDs 1027396 (Chachuna Managed Reserve), 1027397 (Tbilisi Botanical Garden) (BOLD:AGH9530), similarity 99.39%. The closest homologous sequences: M.lebetina (sic!) with a private status (Tajikistan; similarity 97.25%); M.schweizeri (NCBI NC_044966.1; Milos, Cyclades Archipelago, Greece; similarity 96.50%).
AnimaliaSquamataViperidae7D0B8199-7E22-570A-9746-8F390BCF51FDViperarenardi(Christoph, 1861)Peliasrenardi Christoph, 1861Viperaursinirenardi: Terentyev & Chernov (1940: 160); Muskhelishvili (1970: 190); Bannikov et al. (1977: 322)Viperaursiniieriwanensis: Joger (1984); Nilson et al. (1988); Engelmann et al. (1993)Vipera (Pelias) eriwanensis: Nilson et al. (1999)Peliaseriwanensis: Tuniyev (2016)Peliasshemakhensis: Tuniyev et al. (2013)Peliasshemakhensiskakhetiensis: Tuniyev et al. (2018)Taxonomic sources followed.
Mebert et al. (2025b)
Records in Georgia.
Nikolskii (1913), Muskhelishvili (1970), Bannikov et al. (1977), Iankoshvili and Tarkhnishvili (2021), Mebert et al. (2025b).
Infraspecific variation.
V.r.eriwanensis (Reuss, 1933).
Taxonomic remark.
Candidate species. The species belongs to a widespread western Eurasian complex Viperaursinii. The nominal subspecies is found in the Javakheti region (southern Georgia); the subspecies V.e.shemakhensis occurs in grasslands of Eastern Georgia, except the Javakheti Plateau. Some authors describe/recognize subspecies kakhetiensis for Georgian populations, but in this checklist, we treat Georgian “kakhetiensis”-type material within V.e.shemakhensis for stability/consistency. Zinenko et al. (2016) found very little genetic differentiation between the nominal V.renardi, V.shemakhensis, and V.lotievi, the same applies to V.eriwanensis; following the most of species’ lists, we keep the name V.eriwanensis for all steppe vipers from Georgia, although suggest that separate species status for V.eriwanensis has insufficient evidence. Further population-level genomic studies might lead to synonymization of this species with V.renardi. Viperae.shemakhensis is mentioned as a separate species in Tuniyev and Ananjeva (2023), but Uetz et al. (2025) consider it as a subspecies.
Nikolskii (1909b), Bánki et al. (2026), Speybroeck et al. (2020), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Muskhelishvili (1970), Bannikov et al. (1977), Iankoshvili and Tarkhnishvili (2021), Mebert et al. (2025b).
Taxonomic remark.
Biologically confirmed species. A easily identified viper that is clearly distinct from other sympatric/parapatric vipers in the Caucasus (incl. V.r. where ranges meet). However, within the kaznakovi complex (incl. dinniki, darevskii, and “tuniyevi”), species boundaries and contact-zone gene flow remain insufficiently tested with nuclear/genomic data; therefore, we treat the latter taxa as candidate species pending integrative assessment. Viperapontica (Billing et al. 1990) was proven to be a hybrid between V.kaznakovi and V.ammodytestranscaucasiana and not a separate species (Dufresnes et al. 2024b).
Barcoded individuals.
CaBOL-IDs 1027394 (Gomaro), 1027395 (Kveda Kinchkha) (BOLD:ADX0147), similarity 99.85%. The closest homologous sequences: V.kaznakovi with a private status (Artvin, Türkiye; similarity 99.85–100%). The next closest taxa: V.dinniki (NCBI KC122720; Krasnodar, Russia; similarity 98.94%) and V.eriwanensis (NCBI KC122723; Kars, Türkiye; similarity 95.74%).
Vedmederja et at. (1986), Bánki et al. (2026), Speybroeck et al. (2020), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Muskhelishvili (1970), Bannikov et al. (1977), Iankoshvili and Tarkhnishvili (2021).
Infraspecific variation.
V.d.dinniki (Nikolskii 1913); V.d.tuniyevi (Ananjeva et al. 2021) comb. et stat. nov.; PeliastuniyeviAnanjeva et al. 2021 – original combination.
Taxonomic remark.
Candidate species. Belongs to the diverse species complex of V.kaznakovi. There is no clear evidence of genetic isolation between V.dinniki and V.kaznakovi, which may be an ecological or geographic subspecies of the latter. The genetic distance (p-distance based on the Cytochrome b structure) between this form and nominal V.dinniki varies between 0.009 and 0.017, with no evidence of fixed genetic differences. V.d.tuniyevi nests within V.dinniki lineages (Ananjeva et al. 2021), However, we suggest keeping subspecies status for this form considering distinct geograpic and environmental differences between V.d.dinniki and V.d.tuniyevi (the former is found above the timberline of the Greater Caucasus, the latter – in forest and mountain habitats of central Georgia).
Vedmederja et al. (1986), Bánki et al. (2026), Uetz et al. (2025).
Records in Georgia.
Nikolskii (1913), Muskhelishvili (1970), Bannikov et al. (1977), Iankoshvili and Tarkhnishvili (2021), Mebert et al. (2025a).
Infraspecific variation.
V.d.darevskii Vedmederja, Orlov and Tuniyev, 1986 (nominotypical).
Taxonomic remark.
Candidate species. Belongs to the V.kaznakovi complex. There is no clear evidence of genetic isolation between V.darevskii and V.kaznakovi; the form may represent an ecological/geographic derivative. This is a highland-adapted taxon, in Georgia it is restricted to the Lesser Caucasus ranges and areas to the south with an elevation greater than 2000m. Its range extends elsewhere into northern Armenia and west to the Black Sea Mountains in Türkiye. Mitochondrial DNA is shared with neighbouring populations of V.kaznakovi; mtDNA identity may reflect introgression rather than species-level isolation. However, phylogenomic data show Black Sea coastal V.kaznakovi being sister (closest relationship) to high mountain V.darevskii in Artvin province, Türkiye, even closer than morpho-ecological equivalent Sako's Viper V.d.sakoi from centro-western Black Sea mountains (Dufresnes et al. 2024b; Mebert et al. 2025a).
Barcoded individuals.
CaBOL-IDs 1010920 (Didi Abuli environs), 1027400 (Tskordza), 1027401 (Didi Abuli Mt.), 1007142 (Didi Abuli Mt.) (BOLD:ADX0147), similarity 99.47–100%. There were no sequences of properly identified V.darevskii in the BOLD Systems. The closest sequence is V.kaznakovi with a private status (Artvin, Türkiye; similarity 99.81–100%). NCBI: V.dinniki (KC122720; Krasnodar, Russia; similarity 99.09%). Next closest taxon: V.eriwanensis (KC122723; Kars, Türkiye; similarity 95.9%).
AnimaliaSquamataViperidae04C9DE6B-A927-5191-A623-EFE4E9530C6FViperaammodytes(Linnaeus, 1758)Coluberammodytes Linnaeus, 1758Viperaammodytes: Nikolskii (1913: 181); Terentyev and Chernov (1940: 162); Muskhelishvili (1970: 196); Bannikov et al. (1977: 326), et auct., partim.Viperatranscaucasiana: Nilson et al. (1999), et auct.Viperameridionalistranscaucasiana: Thanou et al. (2023); Dufresnes et al. (2024b)Taxonomic sources followed.
Boulenger (1913); Terentyev and Chernov (1940); Muskhelishvili (1970); Bannikov et al. (1977).
Records in Georgia.
Nikolskii (1913), Muskhelishvili (1970), Bannikov et al. (1977), Iankoshvili and Tarkhnishvili (2021).
Infraspecific variation.
V.a.transcaucasiana Boulenger, 1913.
Taxonomic remark.
Biologically confirmed species. Either a polytypic species or species complex. Recent genomic study suggested a closer relation of the vipers from the Southern Caucasus (V.a.transcaucasiana) to the southern Balkan lineage V.meridionalis, than to “typical” V.ammodytes (Thanou et al. 2023). However, the p-distance between mitochondrial cytochrome b gene of V.m.transcaucasiana and nominal V.a.ammodytes from Balkans is less than 5%. We prefer to follow a conservative approach and maintain the subspecific status for V.ammodytes from Georgia.
Barcoded individuals.
CaBOL-IDs 1010927 (Gomaro), 1007150 (Tashiskari) (BOLD:AFE9729; similarity 99.82%). The closest homologous sequences: V.transcaucasiana with a private status (Türkiye and Russia; similarity 99.48–100%); V.ammodytes (KU986335; unknown location; similarity 96.79%) Second-closest taxon: V.ammodytes (BOLD:ACH1831; uknown location; similarity 96.01–96.72%); V.eriwanensis (NCBI KC122723; Kars, Türkiye; similarity 89.13%).
Anecdotal reports
There are five species that have been reportedly found in Georgia but some authors, rejected their presence in the country. These species are:
1. Ranadalmatina (Fitzinger, 1839). Nikolskii (1913) suggests the presence of this species for three locations in Western Georgia. We assume that these individuals could be potentially be confused with R.macrocnemis: specimens of this species with very long legs and small metatarsal tubercle are common in the lowland populations from Western Georgia (Tarkhnishvili et al. 2001). No further findings of this species from Georgia or the Caucasus were ever recorded.
2. Cheloniamydas (Linnaeus, 1758). Tuniyev and Ananyeva (2024) mentioned two findings of this turtle on the coast of NW Georgia. There are few other records from Georgia’s Black Sea coast (anecdotic information), and no evidence of reproduction of this species in Georgia is available.
3. Eremiasstrauchi (Kessler, 1878). The species is found in southern Azerbaijan, and Balavadze (1959 as cited by Muskhelishvili 1970) reported finding of three specimens of E.strauchi in the extreme southeast of Georgia. However, Muskhelishvili (1970) suggested that the individuals stored in the collection were rock lizards (Darevskia), hence it is highly likely that this report is based on a misidentification.
4. Darevskiachlorogaster (Boulenger, 1908). Nikolskii (1913) indicated the presence of this species (under the synonym Lacertaboettgeri) near Lagodekhi (Eastern Georgia). Erroneous labeling of the sample is the most likely reason. There is a broad geographic gap between Lagodekhi and the actual range of this species in NE Azerbaijan.
5. Elaphesitula (Linnaeus, 1758). Terentyev and Chernov (1940) mentioned finding of this species near Tbilisi, without further reference. Subsequently, there have been no indications of this species’ presence in Georgia. Potentially, this note could result from misidentification with Zamenishohenackeri or Elaphedione.
Species found near Georgia’s border and that could potentially occur in the country:
1. Viperarenardilotievi (Nilson et al. 1995) is present on the northern slopes of the Greater Caucasus Mountains and may occur at mid elevations on slopes adjacent to the Tergi (Terek) River and its tributaries in Georgia near the border with Russia. Speybroeck et al. (2020) synonymized this form with V.renardi, whereas Mebert et al. (2025b) revealed syntopy and admixture (hybrids or intergrades) between the eriwanensis and lotievi lineages of V.renardi in the eastern Greater Caucasus. The putative finds of V.lotievi on the southern slopes of the Georgian section of the Greater Caucasus by Bekoshvili and Doronin (2015) require confirmation, as the site is within the Tusheti National Park an area well visited by hikers and herpetologists, with no confirmed observations of V.lotievi, whereas V.dinniki is found in the area. In addition, Bekoshvili and Doronin (2015) published a second case in the same article that requires confirmation, they report Dolichophiscaspius phenotypes from within a well-known D.schmidti region, the nearest indigenous D.caspius are known from a location several hundred kilometres away. Having both Dolichophis taxa syntopic would pose a severe problem due to the ecological competition between them.
2. Viperaberusbarani (Bohme & Joger, 1983) has erroneously been placed by Baran et al. (2001) into the area of the two known, barani-like specimens of Vipera “pontica” (actual hybrids between V.a.transcaucasiana and V.kaznakovi (Freitas et al. 2020; Dufresnes et al. 2024b)) in Artvin province near the south-western border of Georgia, (pers. comm. K. Mebert). Its eastern known limit is within Rize province, and it does not occur in Artvin province to the east (Gül, 2020).
Discussion
There are a total of 12 formally described amphibian species and 56 formally described reptiles recorded for Georgia (excepting the five erroneously mentioned species – see the list above). Two species (unintentionally introduced Phoenicolacertalaevis and captive-released Trachemysscripta) are introduced species, from distant geographic regions (Tarkhnishvili et al. 2017b; Reshetnikov et al. 2023). Tenuidactyluscaspius, native to extreme south-eastern Georgia, forms a set of exclave populations in Georgian cities, including those on the Black Sea coast (Prondzynska 2025).
Our analysis suggests that six nominal species (Mertensielladjanaschvilii, Darevskiaarribasi, D.dryada, Viperatranscaucasiana, V.eriwanensis, V.tuniyevi) had minute genetic differences from their closest relatives with no evidence of full or partial genetic isolation; consequently, we suggest downscaling their status to subspecies until more thorough genetic data are available. The status of Natrix “megalocephala” was rejected by previous authors. The other 15 species, some of which have been accepted for a long period (30 years or more) have been assigned unconfirmed species status, according to the criteria described in the introduction of this paper. Hence, in taxonomic remarks, we qualify them as candidate species. The taxonomic status of these nominal species should be validated in the future through analyses of multiple unlinked genetic markers and of the contact zone between these lineages and their closest relatives. The relative explanations are included in the species accounts; however, the most important remarks are included below.
1. The entire complex of vipers from the V.kaznakovi group, including V.kaznakovi sensu lato, and V.dinniki, V.tuniyevi, and V.darevskii, requires a thorough study of gene flow across the environmentally associated contact zone, which is currently underway. Genetic differences among these taxa are modest, and whether they are geographically distinct is unclear. The mitochondrial divergence between specimens of these species from Georgia does not exceed 5%; there is no evidence of haplotypes fixed for nominal taxa within this group; and the differences between nominal V.kaznakovi from different parts of the range exceed those between non-conspecific individuals from geographically close areas.
2. The taxonomic status of Darevskiaobscura has been elevated based on both phenotypic and genotypic characteristics of populations from the upper reaches of the Mtkvari (Kura) River. However, the differences between these populations and D.rudis from the neighboring areas are modest, and additional genomic analysis is needed to validate the genetic isolation of these two nominal species.
3. Conversely, stepped phenotypic and genotypic clines exist among brown frogs (Ranamacrocnemis sensu lato) in the Central Lesser Caucasus (Tarkhnishvili et al. 2001); however, the formal taxonomic status of the distinct form R. “camerani” may be revised based on additional genetic studies. The same is true for the eastern population group of Mertensiellacaucasica sensu lato.
4. Presence of hybrid zones is either demonstrated or hypothesized for some closely related species: (a) D.obscura, D.portschinskii, and D.valentini (Tarkhnishvili et al. 2013); (b) Hylaorientalis and H.savignyi – these two species form a broad contact zone in eastern Georgia and may hybridize in this area: this question requires additional study. (c) Two closely related rock lizards, D.caucasica and D.daghestanica, have contiguous ranges in the upper currents of the Tergi (Terek) River, and may hybridize.
5. There is insufficient evidence for separate species status for nominal Bufoverrucosissimus, as mitochondrial haplotypes differ only slightly from those of the nominal B.bufo from Eastern Europe (Sinsch et al. 2008).
6. Profound differences exist between Eremiasvelox from Georgia/Caucasus and the eastern China. Similarly, the long-legged skink’s (Eumecesschneideriiprinceps) sequence differs from that of another nominal subspecies (E.s.pavimentatus). Such profound differences may lead to these two widespread nominal species being revised. The same applies to Ophisopselegans sensu lato, in which sequences from different parts of the range differ by more than 10%.
Comparisons of the sequences from this list with those available in open databases (BOLD Systems and NCBI) revealed multiple knowledge gaps, which would help focus further phylogenetic and taxonomic studies. Those gaps are various.
a) Very high sequence differences among nominally conspecific individuals from different geographic areas. For Mertensiellacaucasica, Eumecesschneiderii, Eremiasvelox, E.arguta, and Ophisopselegans, these infraspecific differences in the structure of the COI mitochondrial gene vary between 8-12%. Specifically, the sequences of E.velox from Georgia and Western China differ by 8.4%; the sequences of E.schneideriprinceps (Georgia) and E.s.pavimentatus (Türkiye) differ by 10.4%; the sequences of O.elegans from Georgia and Türkiye differ by 11.9%. The sequences of M.caucasica from central Georgia and NE Türkiye differ by 11.9% (which aligns with the earlier findings based on the other mitochondrial gene (Tarkhnishvili et al. 2000). The sequences of Platycepsnajadum from Georgia and one sequence from Armenia differ by 6.65%. These cases indicate possible genetic isolation between the mentioned formally conspecific populations and may warrant revision of their taxonomic status.
b) Conversely, there are minor (< 5%) genetic differences between some nominally different species from the same genus. E. g. COI structure between the nominal B.verrucosissimus from Georgia and B.bufo from Central Europe is less than 3%. The sequences of Emysorbicularis from Georgia and E.trinacris from Sicily differ by only 1.6%; the differences between Hylaorientalis from Georgia and H.molleri (Iberian Peninsula), despite a broad geographic gap, reach only 4.3%. Only 1.8% separates the COI structure of Viperarenardieriwanensis from Georgia and V.renardi from Central Asia; therefore, the Georgian form is treated here as a subspecies rather than a separate species. Additionally, genomic data analysis could support validating the non-conspecific status of these geographic forms. Specific stories include species complexes such as Pelophylaxridibundus, Bufotesviridis, and V.kaznakovi. MtDNA differences among species of these complexes are minute, and their separate species status should be validated based on the analysis of multiple genetic markers.
с) The analysis of barcodes revealed uneven coverage of different taxonomic groups by COI barcoding projects. Some are underrepresented in the open databases. For instance, there are 14 species of the genus Telescopus in the World Fauna (Uetz et al. 2025); however, the closest available COI sequence to T.fallaxiberus from our collection is Masticophistaeniatus from the Pacific coast of North America (similarity 87.3%). There are eight described species of geckos in the genus Tenuidactylus, but surprisingly, the closest sequence to that of T.caspius from Georgia was from Crossobamoneversmanni from Pakistan (similarity 81.7%). Although there are sequences belonging to snakes of the genera Zamenis and Eirenis in either BOLD Systems or NCBI databases, the closest non-congeneric sequences to Dolichophisschmidti and Eirenismodestus were non-congeneric representatives of the genera Lytorhynchus and Hemorrhois, respectively. Finally, the COI sequence closest to that of Zamenishohenackeri (similarity ~90%) belonged to a colubrid from Madagascar, Pseudoxyrhopusimerinae.
These facts suggest that many gaps and unresolved phylogenies remain despite the multiple genetic datasets collected over the last few decades; the genomic era might shed more light on these questions. It is important to understand that, from a practical point of view, both under-describing and over-describing species diversity can be counter productive. Indeed, bureaucratic bodies prefer to include species rather than infraspecific units in their conservation lists. However, effective conservation planning requires funds and political decisions, and it is a common illusion that longer lists of species requiring habitat protection force decision-makers to spend more on conservation. Conversely, this might promote ill-founded taxonomic inflation (e.g., Tarkhnishvili 2024; Wuster 2025), which compromises species integrity. Concentration of conservation efforts on populations with very small ranges diverts public attention from more widespread taxa, which can remain unprotected across the largest and most important parts of their range. An opposite example is the elevation of the taxonomic status of regional subspecies of some widespread amphibians and reptiles. Lizard Lacertatrilineata, snakes Dolichophisjugularis, Elaphequatuorlineata, V.ammodytes, V.ursini, amphibians B.viridis and H.arborea are included in Annex II (species requiring special protection) of the Bern Convention on conservation of Wildlife and Natural Habitats (https://www.coe.int/en/web/bern-convention/home). Subspecies of all these species from the Caucasus were re-described as separate species in recent decades. None of those are prioritized in conservation programs supported by the European Commission because they are no longer included in Annex II.
In conclusion, we do not insist that the presented species list of Georgia’s amphibians and reptiles is final. The number of species and nomenclature may (and probably will) change in the future for several reasons. (1) More genomic and population genetic studies of the amphibians and reptiles will either confirm or reject the “true” species status of some formally described taxa, which could elevate the taxonomic status of some geographic populations and, conversely, reject independent evolutionary pathways of others. (2) Formally existing infraspecific taxonomy, based on the analysis of phenotypic characters, rarely reflects actual phylogenetic and phylogeographic patterns and will change in the future. (3) Globalization, landscape change, and growing infrastructure will probably lead to the expansion of some species from the areas bordering Georgia, which can potentially increase the country's species diversity in the future.
Acknowledgements
The authors are thankful to students and researchers of Ilia State University for their assistance in field works. We appreciate comments of two anonymous reviewers for their useful comments. Our special appreciation to Konrad Mebert for throughout revision of the paper draft and multiple useful comments, as well as providing some snake specimens for the collection of ISU.
Additional informationConflict of interest
The authors have declared that no competing interests exist.
Ethical statement
No ethical statement was reported.
Artificial Intelligence (AI) use
Regarding the use of AI in the preparation of this manuscript, the authors declare the following: The authors used an AI-assisted language tool (Chat-GPT) only for grammar correction, spelling, and minor wording improvements during preparation of the first draft of the manuscript. All content was reviewed and approved by the authors, who take full responsibility for the manuscript.
Funding
The study was partially funded by the Shota Rustaveli National Science Foundation of Georgia (award no. FR-23-17324) and by the Federal Ministry of Education and Research of Georgia under grant number 01DK20014A.
All of the data that support the findings of this study are available in the main text or Supplementary Information.
ReferencesAhmadzadeh F, Flecks M, Rödder D, Böhme W, Ilgaz Ç, Harris DJ, Engler JO, Üzüm N, Carretero MA (2013) Multiple dispersal out of Anatolia: biogeography and evolution of oriental green lizards. Biological Journal of the Linnean Society 110(2): 398–408. https://doi.org/10.1111/bij.12129Ananjeva NB (2006) The reptiles of northern Eurasia: taxonomic diversity, distribution, conservation status (No. 47). Pensoft Publishers, 245 pp. https://doi.org/10.1071/9780643093034.reptilesAnanjeva NB, Borkin LY, Darevsky IS, Orlov NL (1988) Dictionary of animal names in five languages. Amphibians and Reptiles. Moscow, Russkij Yazyk, 560 pp.Ananjeva NB, Gabaev VN, Iremashvili GN, Lotiev KY, Petrova TV (2021) The taxonomic status of the vipers of the Pelias (kaznakovi) complex in the middle-flow of the Kura River basin in the East Transcaucasia. Trudy Zoologicheskogo Instituta RAN 325(1): 3–33. https://doi.org/10.31610/trudyzin/2021.325.1.3Arnold EN, Arribas O, Carranza S (2007) Systematics of the Palaearctic and Oriental lizard tribe Lacertini (Squamata: Lacertidae: Lacertinae), with descriptions of eight new genera. Zootaxa 1430(1): 1–86. https://doi.org/10.11646/zootaxa.1430.1.1Arntzen J, Beukema W, Galis F, Ivanović A (2015) Vertebral number is highly evolvable in salamanders and newts (family Salamandridae) and variably associated with climatic parameters. Contributions to Zoology 84(2): 85–113. https://doi.org/10.1163/18759866-08402001Arntzen JW, Wielstra B, Wallis GP (2014) The modality of nine Triturus newt hybrid zones assessed with nuclear, mitochondrial and morphological data. Biological Journal of the Linnean Society 113(2): 604–622. https://doi.org/10.1111/bij.12358Arribas OJ (1999) Phylogeny and relationships of the mountain lizards of Europe and Near East (Archaeolacerta Mertens, 1921, sensu lato) and their relationships among the Eurasian lacertid radiation. Russian Journal of Herpetology 6(1): 1–22.Arribas O, Candan K, Kornilios P, Ayaz D, Kumlutaş Y, Gül S, Yilmaz C, Caynak EY, Ilgaz C (2022b) Revising the taxonomy of Darevskiavalentini (Boettger, 1892) and Darevskiarudis (Bedriaga, 1886) (Squamata, Lacertidae): a Morpho-Phylogenetic integrated study in a complex Anatolian scenario. Zootaxa 5224(1): 1–68. https://doi.org/10.11646/zootaxa.5224.1.1Arribas O, Candan K, Kumlutas Y, Ilgaz Ç (2021) Multivariate analysis of geographic variation in Darevskiaclarkorum (Darevsky & Vedmederja, 1977), correlation with geographic and climatic parameters, and true status of Darevskiadryada (Darevsky & Tuniyev, 1997). Zootaxa 4990(1): 1–22. https://doi.org/10.11646/zootaxa.4990.1.1Arribas O, Candan K, Kurnaz M, Kumlutaş Y, Caynak EY, Ilgaz Ç (2022a) A new cryptic species of the Darevskiaparvula group from NE Anatolia (Squamata, Lacertidae). Organisms Diversity & Evolution 22(2): 475–490. https://doi.org/10.1007/s13127-022-00540-4Astrin JJ, Stuben PE (2008) Phylogeny in cryptic weevils: molecules, morphology and new genera of western Palaearctic Cryptorhynchinae (Coleoptera: Curculionidae). Invertebrate Systematics 22: 503–522. https://doi.org/10.1071/IS07057Audouin JV (1827) Explication sommaire des planches de Reptiles (supplément)... offrant un exposé des caracteres naturelles des genres, avec la distinction des especes: description de l’Egypte, ou Recueil des Observations et des Recherches qui ont été faites en Egypte pendant l’Expedition de l’Armée Française, publie par les Ordres de sa Majesté l’Empereur Napoléon le Grand. Histoire Naturelle 1(4): 161–184.Baig K, Wagner P, Ananjeva N, Boehme W (2012) A morphology-based taxonomic revision of Laudakia Gray, 1845 (Squamata: Agamidae). Vertebrate Zoology 62: 213–260. https://doi.org/10.3897/vz.62.e31388Bakradze MA (1976) New subspecies, Lacertaportschinskiinigrita ssp. n., from Eastern Transcaucasia. Vestnik Zoologii 4: 54–57. [in Russian]Bakradze MA, Darevsky IS (1973) The lidless skink Ablepharuspannonicus (Sauria, Scincidae) in the Caucasus and possible zoogeographical grounds of this finding. Zoologicheskii Zhurnal 52(3): 464–466. [in Russian]Bánki O, Bánki Y, Döring M, Ower G, Hernández Robles DR, Plata Corredor CA, Stjernegaard Jeppesen T, Örn A, Pape T, Hobern D, Garnett S, Little H, DeWalt RE, Miller J, Orrell T, Aalbu R, Abbott J, Abreu C, Acero PA, et al. (2026) Catalogue of Life (2026-01-16 XR). Catalogue of Life Foundation, Amsterdam, Netherlands. https://doi.org/10.48580/dgw64Bannikov AG, Darevsky IS, Ishchenko VG, Rustamov AK, Szczerbak NN (1977) Opredelitel zemnovodnyh i presmykayushchihsya fauny SSSR. Prosveshenie, Moscow, 415 pp. [in Russian]Baran İ, Joger U, Kutrup B, Türkozan O (2001) On new specimens of Viperabarani Böhme & Joger, 1983, from northeastern Anatolia, and implications for the validity of Viperapontica Billing, Nilson & Sattler, 1990 (Reptilia, Viperidae). Zoology in the Middle East. 23(1): 47–53. https://doi.org/10.1080/09397140.2001.10637866Bedriaga J (1886) Beiträge zur Kenntnis der Lacertidenfamilie. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 14: 17–444. https://doi.org/10.5962/bhl.title.10613Bedriaga J (1890) Die Lurchfauna Europas. I. Anura. Froschlurche. Bulletin de la Société Impériale des Naturalistes de Moscou 3: 210–422.Bekoshvili D, Doronin IV (2015) New Data on the Distribution of Snakes in Georgia (Caucasus). Herpetological Review 46(3): 388. https://doi.org/10.3897/herpetozoa.32.e35615Berthold AA (1846) Über das Vorkommen von Tritonen am Kaukasus. Nachrichten von der Georg-Augusts-Universität und der Königl. Gesellschaft der Wissenschaften zu Göttingen, 188–190.Bischoff W (1978) Beiträge zur Kenntnis der Echsen des Kaukasus. Salamandra 14 (4): 178–202.Billing H, Nilson G, Sattler U (1990) Viperapontica sp. n., a new viper species in the kaznakovi group (Reptilia, Viperidae) from northeastern Turkey and adjacent Transcaucasia. Zoologica Scripta 19(2): 227–231. https://doi.org/10.1111/j.1463-6409.1990.tb00257.xBischoff W (1982) Zur Kenntnis der innerartlichen Gliederung der Arwin Eidechse, Lacertaderjugini Nikolskij, 1898. Zoologische Abhandlungen 38(1): 1–52.Bischoff W (1984) Bemerkungen zur innerartlichen Gliederung und zur Verbreitung der Artwiner Eidechse (Lacertaderjugini Nikolskij, 1898) an den Südhängen des Grossen Kaukasus (Sauria: Lacertidae). Salamandra 20(2-3): 101–111.Bischoff W (2007) Auf der Suche nach Gebirgseidechsen in drei Erdteilen. Draco 7(27): 54–73.Böhme W, Joger U (1983) Eine neue Art des Viperaberus-Komplexes aus der Türkei. Amphibia-Reptilia 4(2): 265–271. https://doi.org/10.1163/156853883x00139Böhme W (1985) Zur Nomenklatur der paläarktischen Bogenfingergeckos, Gattung Tenuidactylus Szczerbak & Golubev 1984 (Reptilia: Gekkonidae). Bonner zoologische Beiträge 36: 95–98.Boettger O (1889). Ein neuer Pelobates aus Syrien. Zoologischer Anzeiger 12: 144–147.Boettger O (1892) Wissenschaftliche Ergebnisse der Reise Dr. Jean Valentins im Sommer 1890.Boulenger GA (1881) On the Lizards of the Genera Lacerta and Acanthodactylus. Proceedings of the Zoological Society of London 49(3): 739–747. https://doi.org/10.1111/j.1096-3642.1881.tb01331.xBoulenger GA (1885) Description of a new Species of Frog from Asia Minor. Proceedings of the Zoological Society of London 53(1): 22–23. https://doi.org/10.1111/j.1096-3642.1885.tb02882.xBoulenger GA (1891) Catalogue of the Reptiles and Batrachians of Barbary (Morocco, Alyeria, Tunisia), based chiefly upon the Notes and Collections made in 1880–1884 by M. Fernand Lataste. The Transactions of the Zoological Society of London 13(3): 93–164. https://doi.org/10.1111/j.1096-3642.1891.tb00047.xBoulenger GA (1896) On some little‐known Batrachians from the Caucasus. Proceedings of the Zoological Society of London 64(3): 548–555. https://doi.org/10.1111/j.1096-3642.1896.tb03061.xBoulenger GA (1913) On the geographical races of Viperaammodytes. Annual Magazin of Natural History 8(11): 283–287. https://doi.org/10.1080/00222931308693322Bucci-Innocenti S, Ragghianti M, Mancino G (1983) Investigations of karyology and hybrids in Triturusboscai and T.vittatus, with a reinterpretation of the species groups within Triturus (Caudata: Salamandridae). Copeia, 662–672. https://doi.org/10.2307/1444330Bülbül U, Matsui M, Kutrup B, Eto K (2011) Taxonomic relationships among Turkish water frogs as revealed by phylogenetic analyses using mtDNA gene sequences. Zoological Science 28(12): 930–936. https://doi.org/10.2108/zsj.28.930Carranza S, Arnold EN, Pleguezuelos JM (2006) Phylogeny, biogeography, and evolution of two Mediterranean snakes, Malpolonmonspessulanus and Hemorrhoishippocrepis (Squamata, Colubridae), using mtDNA sequences. Molecular phylogenetics and evolution 40(2): 532–546. https://doi.org/10.1016/j.ympev.2006.03.028Chkhikvadze VM, Tuniyev B (1986) On the taxonomic status of modern land tortoise of the western Transcaucasus. Bulletin of the Academy of Science of Georgian SSR (Soobshch. Akad. Nauk Gruz. SSR) 124(3): 617–620.Conant R, Collins JT (1998) A field guide to reptiles & amphibians: eastern and central North America (Vol. 12). Houghton Mifflin Harcourt, 616 pp.Darevsky IS (1953) On the taxonomic position of the Transcaucasian steppe racerunners Eremiasarguta (Pall.) (Reptilia, Sauria). Doklady AN ArmSSR 16(4): 117–121. [in Russian]Darevsky IS (1957) Systematics and ecology of rock lizards (Lacertasaxicola Eversmann) in Armenia. Zoologicheskie soobshchenija AN Armyanskoj SSR 10: 27–57. [in Russian]Darevsky IS (1967) Rock lizards of the Caucasus: systematics, ecology and phylogenesis of the polymorphic groups of Caucasian rock lizards of the subgenus Archeolacerta. Nauka, Leningrad, 276 pp. [in Russian]Darevsky IS, Eiselt J (1980) Neue felseneidechsen (Reptilia: Lacertidae) aus dem Kaukasus und aus der Türkei. Amphibia-Reptilia 1(1): 29–40. https://doi.org/10.1163/156853880x00042Darevsky IS, Tuniyev BS (1997) A new lizard species from Lacertasaxicola group – Lacertadryada sp. nov. (Sauria, Lacertidae) and some comments relative to Lacertaclarkorum Darevsky et Vedmederja, 1977. Russian Journal of Herpetology 4(1): 1–7. https://doi.org/10.30906/1026-2296-1997-4-1-1-7Darevsky IS, Vedmederja V (1977) A new species of rock lizard Lacertasaxicola Eversmann group from northeastern Turkey and adjoining regions of Adjaria. Trudy Zoologicheskogo Instituta Akademii Nauk SSSR 74: 50–54. [in Russian]Daudin FM (1802) FM 1802–1803. Histoire naturelle, générale et particulière, des reptiles. Imprimerie de F. Dufart, Paris. https://doi.org/10.5962/bhl.title.60678Daudin FM (1803) Histoire Naturelle, Générale et Particulière des Reptiles. vol. 7. Paris: Dufart, pp. 436. https://doi.org/10.5962/bhl.title.60678Delwig W (1928) Eine neue Art der Gattung Pelobates Wagl. aus dem zentralen Transkaukasus. Zoologischer Anzeiger 75: 24–31.DeQueiroz K (2007) Species concepts and species delimitation. Systematic biology 56(6): 879–886. https://doi.org/10.1080/10635150701701083Doronin IV (2011) A description of a new subspecies of rock lizard Darevskiabraunerimyusserica ssp. nov. from the western transcaucasia (Abkhazia), with comments on systematics of Darevskiasaxicola complex. Proceedings of the Zoological Institute RAS 315(3): 242–262. https://doi.org/10.31610/trudyzin/2011.315.3.242 [in Russian]Doronin IV, Dzhelali PA, Lotiev KY, Mazanaeva LF, Mustafaeva GA, Bunyatova SN (2021) Phylogeography of a Darevskia (caucasica) complex (Lacertidae: Sauria) based on the cytochrome b mitochondrial gene analysis. Proceedings of the Zoological Institute RAS 325(1): 49–66. https://doi.org/10.31610/trudyzin/2021.325.1.49 [in Russian]Dubey S, Zwahlen V, Mebert K, Monney JC, Golay P, Ott T, Durand T, Thiery G, Kaiser L, Geser SN, Ursenbacher S (2015) Diversifying selection and color-biased dispersal in the asp viper. BMC Evolutionary Biology, 15: 1–9. https://doi.org/10.1186/s12862-015-0367-4Dubois A, Raffaëlli J (2009) A new ergotaxonomy of the family Salamandridae Goldfuss, 1820 (Amphibia, Urodela). Alytes. Paris 26: 1–85.Dufresnes C, Ghielmi S, Halpern B, Martínez-Freiría F, Mebert K, Jelić D, Crnobrnja-Isailović J, Gippnes S, Jablonski D, Joger U, Laddaga L, Petrovan S, Tomović L, Vörös J, İğci N, Kariş M, Zinenko O, Ursenbacher S (2024b) Phylogenomic insights into the diversity and evolution of Palearctic vipers. Molecular Phylogenetics and Evolution 197: 108095, 1–7. https://doi.org/10.1016/j.ympev.2024.108095Dufresnes C, Mazepa G, Jablonski D, Oliveira RC, Wenseleers T, Shabanov DA, Auer M, Ernst R, Koch C, Ramírez-Chaves HE, Mulder KP, Simonov E, Tiutenko A, Kryvokhyzha D, Wennekes PL, Zinenko OI, Korshunov OV, Al-Johany AM, Peregontsev EA, Masroor R, Litvinchuk S (2019) Fifteen shades of green: the evolution of Bufotes toads revisited. Molecular Phylogenetics and Evolution 141: 106615, 1–25. https://doi.org/10.1016/j.ympev.2019.106615Dufresnes C, Monod‐Broca B, Bellati A, Canestrelli D, Ambu J, Wielstra B, Dubey S, Crochet P-A, Denoël M, Jablonski D (2024a) Piecing the barcoding puzzle of Palearctic water frogs (Pelophylax) sheds light on amphibian biogeography and global invasions. Global Change Biology 30(3): e17180, 1–30. https://doi.org/10.1111/gcb.17180Dufresnes C, Strachinis I, Tzoras E, Litvinchuk SN, Denoël M (2019) Call a spade a spade: taxonomy and distribution of Pelobates, with description of a new Balkan endemic. ZooKeys 859: 131–158. https://doi.org/10.3897/zookeys.859.33634Duméril AMC, Bibron G (1839) Erpétologie Générale on Histoire Naturelle Complète des Reptiles. Vol. 5. Roret/Fain et Thunot, Paris, 871 pp. https://doi.org/10.5962/bhl.title.45973Dunaev EA, Orlova VF (2017) Amphibians and reptiles of Russia. Atlas and determination. 2nd. ed. Moscow: Phyton XXI, 328 pp. https://doi.org/10.1515/9781400885053-006Dwigubsky JA (1832) Essay on the Natural History of Animals from the Russian Empire. 54 pp. [in Russian]Eichwald E (1831) Zoologia specialis, quam expositis animalibus tum vivis, tum fossilibus potissimuni rossiae in universum, et poloniae in specie, in usum lectionum publicarum in Universitate Caesarea Vilnensi. Zawadski, Vilnae: 116–197. https://doi.org/10.5962/bhl.title.51803Eichwald E (1839) De Duabus Novis Amphibiorum Speciebus. Bulletin de la Société impériale des naturalistes de Moscou XII(II): 303–307.Eichwald E (1841) Fauna Caspio-Caucasica nonnullis observationibus novis illustravit. Nouveaux Mémoires de la Société Impériale des Naturalistes de Moscou 7: 1–290. https://doi.org/10.5962/bhl.title.126375Engelmann WE, Fritzsche J, Günther RR, Obst F-J (1993) Lurche und Kriechtiere Europas. Neumann Verlag, 420 pp.Esterbauer H (1985) Die Eidechsennatter, Malpolonmonspessulanus. Syrien. Die Aquarien und Terrarienzeitschrift 38(12): 562–565.Eversmann E (1834) Lizards of Imperial Russia. Nouveaux mémoires de la Société impériale des naturalistes de Moscou: 339–369.Fitzinger LJ (1823) Ueber die im Erzherzogthum Oesterreich vorkommenden Reptilien. Hormayr’sches Archiv, Wien (59).Fleischmann FA (1831) Dalmatiae nova serpentum genera. Acta Academiae Caesareae Leopoldino-Carolinae Germanicae Naturae Curiosorum 16(6): 193–196.Freitas I, Ursenbacher S, Mebert K, Zinenko O, Schweiger S, Wüster W, Brito JC, Crnobrnja-Isailović J, Halpern B, Fahd S, Santos X (2020) Evaluating taxonomic inflation: towards evidence-based species delimitation in Eurasian vipers (Serpentes: Viperinae). Amphibia-Reptilia 41(3): 285–311. https://doi.org/10.1163/15685381-bja10007Fretey J-F (2019) Le nom spécifique lebetinus de la vipère du Levant. Bulletin de la Société Herpétologique de France 2019(1): 1–4.Fritz U (1994) Zur innerartlichen Variabilität von Emysorbicularis (Linnaeus, 1758). 4. Variabilität und Zoogeographie im pontokaspischen Gebiet mit Beschreibung von drei neuen Unterarten. Zoologische Abhandlungen. Museums für Tierkunde Dresden 48(4): 53–93.Fritz U, Baran I, Budak A, Amthauer E (1998) Some notes on the morphology of Emysorbicularis in Anatolia, especially on E.o.luteofusca and E.o.colchica, with the description of a new subspecies from southeastern Turkey. Mertensiella 10: 103–121.Frost DR (2026) Amphibian Species of the World: an Online Reference. Version 6.2 (accessed on 20/02/2026). Electronic Database accessible at https://amphibiansoftheworld.amnh.org/index.php. American Museum of Natural History, New York, USA. https://doi.org/10.5531/db.vz.0001Frost DR, Grant T, Faivovich J, Bain RH, Haas A, Haddad CF, De Sa RO, Channing A, Wilkinson M, Donnellan SC, Raxworthy CJ, Campbell JA, Blotto BL, Moler P, Drewes RC, Nussbaum RA, Lynch JD, Green DM, Wheeler WC (2006) The amphibian tree of life. Bulletin of the American Museum of natural History 297: 1–291. https://doi.org/10.1206/0003-0090(2006)297[0001:tatol]2.0.co;2]2.0.co;2]2.0.CO;2Gabelaia M, Tarkhnishvili D, Adriaens D (2018) Use of three-dimensional geometric morphometrics for the identification of closely related species of Caucasian rock lizards (Lacertidae: Darevskia). Biological Journal of the Linnean Society 125(4): 709–717. https://doi.org/10.1093/biolinnean/bly143Geoffroy Saint-Hilaire E (1827) Description des Reptiles. In: Savigny MJCL de (Ed). Description de l’Égypte ou recueil des observations et des recherches Quintana Roo ont été faites en Égypte pendant l’expedition de l’Armée Française, publié par les ordres de sa Majesté l’Empere Paris, L’imprimerie impériale: 121–160.Gmelin SG (1774) Reise durch Russland zur Untersuchung der drey Natur-Reiche (Vol. 3). Gedruckt bey der Kayserl. Academie der Wissenschaften, 632 pp.Gray JE (1838) Catalogue of the slender-tongued saurians, with descriptions of many new genera and species. Journal of Natural History 1(4): 274–283. https://doi.org/10.1080/00222933809512291Gray JE (1845) Catalogue of the specimens of lizards in the collection of the British Museum. Part I. Catalogue of the specimens of lizards in the collection of the British Museum: 1–80. https://doi.org/10.5962/bhl.title.5499Grechko VV, Bannikova AA, Kosushkin SA, Ryabinina NL, Milto KD, Darevsky IS, Kramerov DA (2007) Molecular genetic diversification of the lizard complex Darevskiaraddei (Sauria: Lacertidae): early stages of speciation. Molecular Biology 41: 764–775. https://doi.org/10.1134/S0026893307050093Griffith H, Ngo A, Murphy RW (2000) A cladistic evaluation of the cosmopolitan genus Eumeces Wiegmann (Reptilia, Squamata, Scincidae). Russian Journal of Herpetology 7(1): 1–16.Gül SE (2020) The most northeastern record of the Turkish endemic viper, Peliasbarani (Böhme and Joger, 1984), from northeastern Anatolia: two viper species in a valley. Amphibian & Reptile Conservation 14(3): 218–225.Gür B, Kalaycı TE (2025) DNA barcoding of some Anura species from Rize. Caucasiana 4 15–24. https://doi.org/10.3897/caucasiana.4.e144670Gvoždík V, Jandzik D, Lymberaki P, Jablonski D, Moravec J (2010) Slow worm, Anguisfragilis (Reptilia: Anguidae) as a species complex: genetic structure reveals deep divergences. Molecular Phylogenetics and Evolution 55(2): 460–472. https://doi.org/10.1016/j.ympev.2010.01.007Hedges SB, Marion AB, Lipp KM, Marin J, Vidal N (2014) A taxonomic framework for typhlopid snakes from the Caribbean and other regions (Reptilia, Squamata). Carribean Herpetology 49: 1–61. https://doi.org/10.31611/ch.49Hijmans RJ, Cameron SE, Parra JL, Jones PG, Jarvis A (2005) Very high resolution interpolated climate surfaces for global land areas. International Journal of Climatology. A Journal of the Royal Meteorological Society 25(15): 1965–1978. https://doi.org/10.1002/joc.1276Hillis DM (2019) Species delimitation in herpetology. Journal of Herpetology 53(1): 3–12. https://doi.org/10.1670/18-123Hofmann S, Mebert K, Schulz KD, Helfenberger N, Göçmen B, Böhme W (2018) A new subspecies of Zamenishohenackeri (Strauch, 1873) (Serpentes: Colubridae) based on morphological and molecular data. Zootaxa 4471(1): 137–153. https://doi.org/10.11646/zootaxa.4471.1.6Hohenacker RF (1831) Notices sur quelques objets d histoire naturelle des provinces méridionales du Caucase que l’on peut se procurer par les roins [sic] de Mr. Hohenaker. Bulletin de la Société impériale des naturalistes de Moscou 3: 363–378.Hohenacker RF (1837) Enumeration animalium quoe in provinciis transcaucasicis Karabach, Schirwan et Taylysh nec non in territorio Elisobetpolensi observavit. Bulletin de la Société Impériale des Naturalistes de Moscou 10: 136–147.Iankoshvili G, Tarkhnishvili D (2021) Distribution of snakes (Reptilia: Serpentes) in Georgia: Social media networks help to improve scientific knowledge. Zoology in the Middle East 67(3): 228–239. https://doi.org/10.1080/09397140.2021.1957208Ishchenko VG (1978) Dynamic Polymorphism of Brown Frogs of the USSR Fauna, Nauka, Moscow, 147 pp. [in Russian]Jablonski D, Kukushkin OV, Avcı A, Bunyatova S, Kumlutaş Y, Ilgaz Ç, Polyakova E, Shiryaev K, Tuniyev B, Jandzik D (2019) The biogeography of Elaphesauromates (Pallas, 1814), with a description of a new rat snake species. PeerJournal 7: e6944, 44 pp. https://doi.org/10.7717/peerj.6944Jablonski D, Mebert K, Masroor R, Simonov E, Kukushkin O, Abduraupov T, Hofmann S (2024) The Silk roads: phylogeography of Central Asian dice snakes (Serpentes: Natricidae) shaped by rivers in deserts and mountain valleys. Current Zoology 70(2): 150–162. https://doi.org/10.1093/cz/zoad008Jimenez-Cazalla F (2012) Malpoloninsignitus (Geoffroy Saint-Hilaire, 1827). In: Martínez G, León R, Jiménez-Robles O, González De la Vega JP, Gabari V, Rebollo B, Sánchez-Tójar A, Fernández-Cardenete JR, Gállego J (Eds). Moroccoherps. Amphibians and Reptiles of Morocco and Western Sahara. Available from www.moroccoherps.com/en/ficha/Malpolon_insignitus/. Version 23/09/2012Joger U (1984) The venomous snakes of the Near and Middle East. Beihefte zum Tübinger Atlas des Vorderen Orients, A 12. Dr. Ludwig Reichert Verlag, Wiesbaden, 115 pp.Kalaentzis K, Koster S, Arntzen JW, Bogaerts S, France J, Franzen M, Kazilas C, Litvinchuk SN, Olgun K, de Visser M, Wielstra B (2025) Phylogenomics resolves the puzzling phylogeny of banded newts (genus Ommatotriton). Molecular Phylogenetics and Evolution 203: 108237, 1–8. https://doi.org/10.1016/j.ympev.2024.108237Kaya U, Agasyan A, Avisi A, Tuniyev B, Crnobrnja Isailovic J, Lymberakis P, Andrén C, Cogalniceanu D, Wilkinson J, Ananjeva N, Üzüm N, Orlov N, Podloucky R, Tuniyev S, Kaya U (2009) Hylaarborea. The IUCN Red List of Threatened Species 2009. https://doi.org/10.2305/IUCN.UK.2009.RLTS.T10351A3197528.enKaya U, Simmons AM (1999) Advertisement calls of the tree frogs, Hylaarborea and Hylasavignyi (Anura: Hylidae) in Turkey. Bioacoustics 10(2-3): 175–190. https://doi.org/10.1080/09524622.1999.9753429Kessler KF (1872) News of the Imperial Society of Naturalists, Anthropology and Ethnography. Volume X, issue 1. Moscow, 168 pp.Kessler KF (1878) A zoological expedition to the Transcaucasian Territory in 1875. Transactions of the St. Petersburg Society of Naturalists 8: 1–200.Kindler C, Böhme W, Corti C, Gvoždík V, Jablonski D, Jandzik D, Metallinou M, Siroky P, Fritz, U (2013) Mitochondrial phylogeography, contact zones and taxonomy of grass snakes (Natrixnatrix, N.megalocephala). Zoologica Scripta 42(5): 458-472. https://doi.org/10.1111/zsc.12018Kindler C, Chèvre M, Ursenbacher S, Böhme W, Hille A, Jablonski D, Fritz U (2017) Hybridization patterns in two contact zones of grass snakes reveal a new Central European snake species. Scientific Reports 7(1): 7378. https://doi.org/10.1038/s41598-017-07847-9Kumar S, Stecher G, Suleski M, Hedges SB (2017) TimeTree: a resource for timelines, timetrees, and divergence times. Molecular biology and evolution 34(7): 1812–1819. https://doi.org/10.1093/molbev/msx116van der Kuyl AC, Ballasina DL, Zorgdrager F (2005) Mitochondrial haplotype diversity in the tortoise species Testudograeca from North Africa and the Middle East. BMC Evolutionary Biology 5(1): 29. https://doi.org/10.1186/1471-2148-5-29Kuzmin SL (2012) The amphibians of the former Soviet Union. KMK, Moscow, 370 pp. [in Russian] https://doi.org/10.30906/1026-2296-1997-4-1-81-82Lantz L (1912) Beitrag zur Biologie von Molge (= Triton) vittata Gray forma ophrytica Berthold. Blatter Aquarien-Terrarienkunde. Magdeburg 22(12): 181–188.Lantz LA, Cyrén O (1918) Nore sur Lacertapraticola Eversm. Proceedings of the Caucasian Museum 11(3-4): 192–195. https://doi.org/10.1080/00222931908673796Lantz LA, Cyrén O (1920) Note sur les Lacertaviridis du Caucase. Bulletin de la Société zoologique de France 45: 33–37.Lantz LA (1928) Les Eremias de l'Asie occidentale. Bulletin du Museum de Géorgie 4: 1–136.Lantz LA, Cyrén O (1936) Contribution à la connaissance de Lacertasaxicola Eversmann. Bulletin de la Société zoologique de France 61: 159–181.Laurenti JN (1768) Specimen medicum, exhibens synopin reptilium emendatam cum experimentis circa venena et antidota reptilium Austriacorum. Trattner, 247 pp. https://doi.org/10.5962/bhl.title.5108Lenk P, Joger U, Wink M (2001) Phylogenetic relationships among European ratsnakes of the genus Elaphe Fitzinger based on mitochondrial DNA sequence comparisons. Amphibia-Reptilia 22(3): 329–339. https://doi.org/10.1163/156853801317050124Lichtenstein MHC (1823) Universität zu Berlin nebst Beschreibung vieler bisher unbekannter Arten von Säugethieren, Vögeln, Amphibien und Fischen. In Verzeichniss der Doubletten des zoologischen Museums der Königl (Vol. 10). Königl. Preuss. Akad. Wiss./T. Trautwein. Berlin. https://doi.org/10.5962/bhl.title.40281Linnaeus CV (1758) Systema naturae. Edition 10: 441–451.Litvinchuk SN, Borkin LJ, Rosanov JM, Skorinov DV (2006) Allozyme and genome size variation in tree frogs from the Caucasus, with description of a new subspecies Hylaarboreagumilevskii, from the Talysh Mountains. Russian Journal of Herpetology 13: 187–206.Litvinchuk SN, Zuiderwijk A, Borkin LJ, Rosanov JM (2005) Taxonomic status of Triturusvittatus (Amphibia: Salamandridae) in western Turkey: trunk vertebrae count, genome size and allozyme data. Amphibia-Reptilia 26(3): 305–323. https://doi.org/10.1163/156853805774408685Martin WCL (1838) A collection of snakes procured by the Euphrates expedition. Proceedings of the Zoological Society of London 81–84.Mayr E (1949) The species concept: semantics versus semantics. Evolution 3(4): 371–372. https://doi.org/10.1111/j.1558-5646.1949.tb00038.xMcDowell SB (1964) Partition of the Genus Clemmys and related problems in the taxonomy of the aquatic Testudinidae. Proceedings of the Zoological Society of London 143(2): 239–279. https://doi.org/10.1111/j.1469-7998.1964.tb03860.xMebert K (2008) Good species despite massive hybridization: genetic research on the contact zone between the watersnakes Nerodiasipedon and N.fasciata in the Carolinas, USA. Molecular Ecology 17(8): 1918-1929. https://doi.org/10.1111/j.1365-294x.2008.03725.xMebert K, Göçmen B, Karış M, İğci N, Ursenbacher S (2016) The valley of four viper species and a highland of dwarfs: fieldwork on threatened vipers in northeastern Turkey. Reptiles & Amphibians 23(1): 1-9. https://doi.org/10.17161/randa.v23i1.14084Mebert K, Iğci N, Kariş M, Yildiz MZ, Schmid PL, Grünewald J, Iankoshvili G, Sanakoeva A, Stümpel N, Tadevosyan T, Çiçek K, Akdag B, Zinenko O, Adakul A, Iremashvili G, Mizsei E, Sos T, Akman B, Elden JM, Jablonski D, Kirschey T (2025a) Critically endangered Darevsky’s Viper: incisive distribution discoveries and taxonomic considerations. Herpetological Conservation and Biology 20(2):438–456.Mebert K, Iskenderov TM, Gasymova GH, Zinenko O (2025c) Does the Iranian steppe viper, Viperaeriwanensisebneri Knoepffler & Sochurek, 1955 occur in Azerbaijan – review and evaluation with new records. Zoology in the Middle East 71(2), 152–162. https://doi.org/10.1080/09397140.2025.2495215Mebert K, Kariş M, Yildiz MZ, Iğci N, Stümpel N, Zinenko O, Iankoshvili G, Sanakoeva A, Schmid PL, Tadevosyan T, Iremashvili G, Mizsei E, Sos T, Iskenderov T, Mammadova A, Çiçek K, Mega B, Avcı A, Rajabizadeh M, Jablonski D, Kirschey T, Elden JM (2025b) Taxonomy, Distribution, and Conservation of Threatened Steppe Vipers of Western Asia: Viperaanatolica and Viperarenardieriwanensis (sensu lato). Herpetological Conservation and Biology 20(2):393–412.Méhely LV (1909) Materialien zu einer Systematik und Phylogenie der muralis-ähnlichen Lacerten. Annales Musei Nationalis Hungarici 7: 409–621.Ménétriés E (1832) Catalogue raisonné des objets de zoologie recueillis dans un voyage au Caucase et jusqu’aux frontières actuelles de la Perse entrepris par ordre de SM l’Empereur. L’Académie Impériale des Sciences, St. Petersbourg, 271 pp. https://doi.org/10.5962/bhl.title.63878Mertens R, Müller L (1928) Liste der Amphibien und Reptilien Europas. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 41(1): 1–62.Mertens R, Müller L (1940) Die Amphibien und Reptilien Europas (Zweite Liste). Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 451.Merrem B (1820) Versuch eines Systems der Amphibien I (Tentamen Systematis Amphibiorum). JC Kriegeri, Marburg, 191 pp. https://doi.org/10.5962/bhl.title.5037Missions L (2020) Landsat normalized difference vegetation index. US Geological Survey. https://www.usgs.gov/landsat-missions/landsat-normalized-difference-vegetation-indexMüller VL (1935) Uber die Smaragdeidechse der Kykladen-Insel Milos. Zoologische Anzeiger 109: 225–236.Murphy RW, Fu J, Macculloch RD, Darevsky IS, Kupriyanova LA (2000) A fine line between sex and unisexuality: the phylogenetic constraints on parthenogenesis in lacertid lizards. Zoological Journal of the Linnean Society 130(4): 527–549. https://doi.org/10.1111/j.1096-3642.2000.tb02200.xMurtskhvaladze M, Tarkhnishvili D, Anderson CL, Kotorashvili A (2020) Phylogeny of caucasian rock lizards (Darevskia) and other true lizards based on mitogenome analysis: Optimisation of the algorithms and gene selection. PLoS One 15(6): e0233680. https://doi.org/10.1371/journal.pone.0233680Musilová R, Zavadil V, Marková S, Kotlík P (2010) Relics of the Europe’s warm past: Phylogeography of the Aesculapian snake. Molecular Phylogenetics and Evolution 57(3): 1245–1252. https://doi.org/10.1016/j.ympev.2010.09.017Muskhelishvili TA (1970) Presmykajushchiesja Vostochnoj Gruzii (The reptiles of the eastern Georgia). Tbilisi, Metsniereba, 241 pp. [in Russian]Nagy ZT, Lawson R, Joger U, Wink M (2004) Molecular systematics of racers, whipsnakes and relatives (Reptilia: Colubridae) using mitochondrial and nuclear markers. Journal of Zoological Systematics and Evolutionary Research 42(3): 223–233. https://doi.org/10.1111/j.1439-0469.2004.00249.xNagy Z, Bellaagh M, Wink M, Paunović A, Korsós Z (2010) Phylogeography of the Caspian whipsnake in Europe with emphasis on the westernmost populations. Amphibia-Reptilia 31(4): 455–461. https://doi.org/10.1163/017353710X518397Nikolskii AM (1898) On two new lizards from Russia. Annuaire du Musée Zoologique de l'Académie Impériale des Sciences de St.-Pétersbourg 3: 284–288.Nikolskii AM (1909a). Novae species reptilium e Caucaso. Izvestija Kavkazskago Muzeja, Tiflis, 4: 301–306.Nikolskii AM (1909b). A new viper species from Caucasus, Viperakaznakovi. Izvestija Kavkazskago Muzeja, Tiflis, 3(4): 173–174.Nikolskii AM (1913) Reptiles and amphibians of the Caucasus. Herpetologia Caucasica. Caucasus Museum Publication, Tiflis, 272 pp. [in Russian]Nilson G, Andrén C, Flärdh B (1988) Die vipern der Türkei. Salamandra (Frankfurt Am Main) 24(4): 215–247.Nilson G, Tuniyev BS, Orlov NL, Höggren M, Andrén C (1995) Systematics of the vipers of the Caucasus: polymorphism or sibling species. Asiatic Herpetological Research 6: 1–26. https://doi.org/10.5962/bhl.part.7982Nilson G, Tuniyev B, Andrén C, Orlov N (1999) Vipers of Caucasus: taxonomic considerations. Kaupia 8: 103–106.Nordmann A von (1840) Observations sur la faune pontique. Ch. F. Schnakenburg, Odessa, 104 pp.Orlov NL, Tuniyev BS (1987) A new species of grass snake from the Caucasus, Natrixmegalocephala sp. nov. (Ophidia: Colubridae). Trudy Zoologicheskogo Instituta Akademii Nauk SSSR–Proceedings of the Zoological Institute, USSR Academy of Sciences 158: 116–139.Orlova VF, Tuniyev BS (1989) On the taxonomy of the Caucasian common toads belonging to the group Bufobufoverrucosissimus (Pallas) (Amphibia, Anura, Bufonidae). Bulleten Moskovskogo Obshchestva Ispytatelej Prirody Serija Biologicheskaya 94(3): 13–24.Pabijan M, Zieliński P, Dudek K, Chloupek M, Sotiropoulos K, Liana M, Babik W (2015) The dissection of a Pleistocene refugium: phylogeography of the smooth newt, Lissotritonvulgaris, in the Balkans. Journal of Biogeography 42(4): 671–683. https://doi.org/10.1111/jbi.12449Pallas PS (1771) Reise durch verschiedene Provinzen des russischen Reichs. Erster Theil. St. Petersburg: Kayserliche Academie der Wissenschaften. [12] + 504 pp.Pallas PS (1773) Reise durch verschiedene Provinzen des Russischen Reiches. Zweyter Theil. St. Petersburg: Kayserliche Academie der Wissenschaften. [8] + 744 pp.Pallas PS (1775) Lacertaapoda descripta. Novi Commentarii Academiae Scientiarum Imperialis Petropolitanae 19: 435–454.Pallas PS (1814) Zoographia Rosso-Asiatica, sistens omnium animalium in extenso Imperio Rossico et adiacentibus maribus observatorum recensionem, domicilia, mores et descriptiones, anatomen atque icones plurimorum. Tomus Tertius. Animalia Monocardia seu Frigidi Sanguinis Imperii Rosso-Asiatici. Petropoli: Ex Officina Caes. Academiae Scientiarum. vii + 428 + cxxv pp. https://doi.org/10.5962/bhl.title.42222Peters G (1958) Die Zauneidechse des Kleinen Kaukasus als besondere Unterart – Lacertaagilisbrevicaudata ssp. n. Zoologischer Jahrbuch 86(1): 127–138.Peters G (1960) Die Grusinische Zauneidechse Lacertaagilisgrusinica nomen novum. Zoologischer Anzeiger, 165(7/8): 279–289.Peters G, Muskheliswili TA (1968) Lacertaagilisioriensis—eine neue Subspecies der kaukasischen Zauneidechsen. Zoologische Jahrbücher Abteilung für Systematik, Geographie und Biologie der Tiere 95: 213-228.Prondzynska KM (2025) First record of the Caspian Bent-toed Gecko, Tenuidactyluscaspius (Eichwald, 1831), from Kutaisi, Georgia. Herpetology Notes 18: 479–482. https://doi.org/10.5281/zenodo.20605527Pyron RA, Burbrink FT, Wiens JJ (2013) A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes. BMC evolutionary biology 13(93): 1–54. https://doi.org/10.1186/1471-2148-13-93QGIS Development Team (2023) QGIS Geographic Information System. Open Source Geospatial Foundation Project. http://qgis.orghttps://doi.org/10.31219/osf.io/nvxcaRadde G (1886) Die Fauna und Flora des südwestlichen Caspi-Gebietes: wissenschaftliche Beiträge zu den Reisen an der persisch-russischen Grenze. FA Brockhaus, 425 pp. https://doi.org/10.5962/bhl.title.36963Raffaëlli J (2022) Salamanders & Newts of the World. Plumelec, France: Penclen Édition, 1100 pp.Ratnasingham S, Hebert PDN (2013) A DNA-based registry for all animal species: The Barcode Index Number (BIN) system. PLoS ONE 8(7): e66213. https://doi.org/10.1371/journal.pone.0066213Reshetnikov AN, Zibrova MG, Ayaz D, Bhattarai S, Borodin OV, Borzée A, Brejcha J, Çiçek K, Dimaki M, Doronin IV, Drobenkov SM, Gichikhanova UA, Gladkova AY, Gordeev DA, Ioannidis Y, Ilyukh MP, Interesova EA, Jadhav TD, Karabanov DP, Khabibullin VF, Khabilov TK, Khan MMH, Kidov AA, Klimov AS, Kochetkov DN, Kolbintsev VG, Kuzmin SL, Lotiev KY, Louppova NE, Lvov VD, Lyapkov SM, Martynenko IM, Maslova IV, Masroor R, Mazanaeva LF, Milko DA, Milto KD, Mozaffari O, Nguyen TQ, Novitsky RV, Petrovskiy AB, Prelovskiy VA, Serbin VV, Shi H-t, Skalon NV, Struijk RPJH, Taniguchi M, Tarkhnishvili D, Tsurkan VF, Tyutenkov OY, Ushakov MV, Vekhov DA, Xiao F, Yakimov AV, Yakovleva TI, Yang P, Zeleev DF, Petrosyan VG (2023) Rarely naturalized, but widespread and even invasive: the paradox of a popular pet terrapin expansion in Eurasia. NeoBiota 81: 91–127. https://doi.org/10.3897/neobiota.81.90473Reuss T (1933) Fortsetzung und Schluss der Originalberichte. Nachr. Bl. Aquarien und Terrarien.-Ver.: 372–373.Schatti B, Monsch P (2004) Systematics and phylogenetic relationships of whip snakes (Hierophis Fitzinger) and Zamenisandreana Werner, 1917 (Reptilia: Squamata: Colubrinae). Revue Suisse de Zoologie 111(2): 239–256. https://doi.org/10.5962/bhl.part.80237Schmidtler JF, Heckes U, Bischoff W, Franzen M (2002) Altitude-dependent character variation in rock lizards of the Darevskiaclarkorum (Darevsky & Vedmederja, 1977)/D.dryada (Darevsky & Tuniyev, 1997) complex: a case of climate parallel variation of pholidosis? (Reptilia: Squamata: Sauria: Lacertidae). Fauniste Abhandlungen Staatliches Museum für Tierkunde Dresden 23: 141–156.Sindaco R, Venchi A, Carpaneto GM, Bologna MA (2000) The reptiles of Anatolia: a checklist and zoogeographical analysis. Biogeographia – The Journal of Integrative Biogeography 21(1): 442–481. https://doi.org/10.21426/b6110017Sinsch U, Tarkhnishvili D, Schneider H (2008) Bufobufo Superspezies – Erdkröten-Artenkreis. In: Handbuch der Reptilien und Amphibien Europas, 191–336.Speybroeck J, Beukema W, Dufresnes C, Fritz U, Jablonski D, Lymberakis P, Martínez-Solano Í, Razzetti E, Vamberger M, Vences M, Vörös J, Crochet P-A (2020) Species list of the European herpetofauna – 2020 update by the Taxonomic Committee of the Societas Europaea Herpetologica. Amphibia-Reptilia 41: 139–189. https://doi.org/10.1163/15685381-bja10010Stejneger L (1907) Herpetology of Japan and adjacent territory. Bulletin of United States National Museum 58: 577 pp. https://doi.org/10.5479/si.03629236.58.iStöck M, Dubey S, Klütsch C, Litvinchuk SN, Scheidt U, Perrin N (2008) Mitochondrial and nuclear phylogeny of circum-Mediterranean tree frogs from the Hylaarborea group. Molecular Phylogenetics and Evolution 49: 1019–1024. https://doi.org/10.1016/j.ympev.2008.08.029Strauch A (1870) Revision der Salamandriden-Gattungen nebst Beschreibung einiger neuen oder weniger bekannten Arten dieser Familie, Commissionnaires de l'Académie impériale des sciences: à St.-Pétersbourg: Eggers 16(4): 109 pp.Strauch A (1873) Die Schlangen des Russischen Reichs, in systematischer und zoogeographischer Beziehung. Mémoires présentés à l'Académie impériale des sciences de St. Petersbourg 11(4): 1–288.Stümpel N, Rajabizadeh M, Avcı A, Wüster W, Joger U (2016) Phylogeny and diversification of mountain vipers (Montivipera, Nilson et al., 2001) triggered by multiple Plio – Pleistocene refugia and high-mountain topography in the Near and Middle East. Molecular Phylogenetics and Evolution 101: 336-351. https://doi.org/10.1016/j.ympev.2016.04.025Suchow GF (1948) Obsor jaschtscheriz podroda Lacerta (Sauria), wstretschaju-schtschychsja w SSSR. Trudy Zoologicheskogo Instituta Akademii Nauk SSSR 7: 101–117.Sztencel-Jabłonka A, Mazgajski TD, Bury S, Najbar B, Rybacki M, Bogdanowicz W, Mazgajska J (2015) Phylogeography of the smooth snake Coronellaaustriaca (Serpentes: Colubridae): evidence for a reduced gene pool and a genetic discontinuity in Central Europe. Biological Journal of the Linnean Society 115(1): 195–210. https://doi.org/10.1111/bij.12496Tarkhnishvili D (2012) Evolutionary history, habitats, diversification, and speciation in Caucasian rock lizards. In: Jenkins OP (Ed). Advances in zoology research, 2: 79–120.Tarkhnishvil DN, Arntzen JW, Thorpe RS (1999) Morphological variation in brown frogs from the Caucasus and the taxonomy of the Ranamacrocnemis group. Herpetologica 55(3): 406–417.Tarkhnishvili D (2014) Ranamacrocnemis (Boulenger, 1885) – Kleinasiatischer Braunfrosch taxon macrocnemis (Boulenger, 1885) – Kaukasischer Braunfrosch taxon camerani (Boulenger, 1896) taxon holtzi (Werner, 1898) taxon pseudodalmatina (Eiselt und Schmidtler, 1971) taxon tavasensis (Baran und Atatür, 1986). In: Handbuch der Reptilien und Amphibien Europas. AULA-Verlag, Wiesbaden, 263–293.Tarkhnishvili D, Gabelaia M, Kandaurov A, Bukhnikashvili A, Iankoshvili G (2017a) Isolated population of the Middle Eastern Phoenicolacertalaevis from the Georgian Black Sea Coast, and its genetic closeness to populations from southern Turkey. Zoology in the Middle East 63(4): 311–315. https://doi.org/10.1080/09397140.2017.1361191Tarkhnishvili D, Gabelaia M, Mumladze L, Murtskhvaladze M (2016) Mitochondrial phylogeny of the Darevskiasaxicola complex: two highly deviant evolutionary lineages from the easternmost part of the range. The Herpetological Journal 26(2): 175–182.Tarkhnishvili DN, Gokhelashvili RK (1999) The amphibians of the Caucasus. Pensoft. Advances in Amphibian Research in the Former Soviet Union 4: 239 pp.Tarkhnishvili D, Gavashelishvili A, Avaliani A, Murtskhvaladze M, Mumladze L (2010) Unisexual rock lizard might be outcompeting its bisexual progenitors in the Caucasus. Biological Journal of the Linnean Society 101(2): 447-60. https://doi.org/10.1111/j.1095-8312.2010.01498.xTarkhnishvili D, Hille A, Böhme W (2001) Humid forest refugia, speciation and secondary introgression between evolutionary lineages: differentiation in a Near Eastern brown frog, Ranamacrocnemis. Biological Journal of the Linnean Society 74(2): 141–156. https://doi.org/10.1006/bijl.2001.0561Tarkhnishvili D, Iankoshvili G (2023) The farther, the closer: geographic proximity and niche overlap versus genetic divergence in Caucasian rock lizards. Biological Journal of the Linnean Society 140(1): 41–57. https://doi.org/10.1093/biolinnean/blad034Tarkhnishvili D, Kandaurov A, Bukhnikashvili A (2002) Declines of amphibians and reptiles in Georgia during the 20th century: virtual vs. actual problems. Zeitschrift fur Feldherpetologie 9: 89–107.Tarkhnishvili D, Kaya U, Gavashelishvili A, Serbinova I (2008) Ecological divergence between two evolutionary lineages of the Caucasian salamander: evidence from GIS analysis. The Herpetological Journal 18(3): 155–163.Tarkhnishvili D, Murtskhvaladze M, Anderson CL (2017b) Coincidence of genotypes at two loci in two parthenogenetic rock lizards: how backcrosses might trigger adaptive speciation. Biological Journal of the Linnean Society 121(2): 365–378. https://doi.org/10.1093/biolinnean/blw046Tarkhnishvili D, Murtskhvaladze M, Gavashelishvili A (2013) Speciation in Caucasian lizards: climatic dissimilarity of the habitats is more important than isolation time. Biological Journal of the Linnean Society 109(4): 876–892. https://doi.org/10.1111/bij.12092Tarkhnishvili D, Thorpe R, Arntzen JW (2000) Pre-pleistocene refugia and differentiation between populations of the caucasian salamander (Mertensiellacaucasica). Molecular phylogenetics and evolution 14(3): 414–422. https://doi.org/10.1006/mpev.1999.0718Tarkhnishvili D, Todua M, Iankoshvili G (2024) Which species of water frogs inhabit ponds of the Caucasus? Taxonomic mess with Pelophylaxridibundus species complex. Caucasiana 3: 31–39. https://doi.org/10.3897/caucasiana.3.e117215Tarkhnishvili D, Todua M, Iankoshvili G, Çetintaş O, Murtskhvaladze M, Yanchukov A (2025) Collapsing landmasses and secondary contacts: introgression and the evolution of adaptive diversity in Caucasian rock lizards. Evolution 79(10): 2193–2207. https://doi.org/10.1093/evolut/qpaf147Tarkhnishvili D, Yanchukov A, Şahin MK, Gabelaia M, Murtskhvaladze M, Candan K, Galoyan E, Arakelyan M, Iankoshvili G, Kumlutaş Y, Ilgaz Ç, Matur F, Çolak F, Erdolu M, Kurdadze S, Barateli N, Anderson CL (2020) Genotypic similarities among the parthenogenetic Darevskia rock lizards with different hybrid origins. BMC Evolutionary Biology 20: 1–25. https://doi.org/10.1186/s12862-020-01690-9Tarroso P, Pereira RJ, Martínez‐Freiría F, Godinho R, Brito JC (2014) Hybridization at an ecotone: ecological and genetic barriers between three Iberian vipers. Molecular ecology 23(5): 1108–1123. https://doi.org/10.1111/mec.12671Tartarashvili RV, Bakradze MA (1989) A new subspecies of salamander from western Georgia. Soobshcheniia Akademii Nauk Gruzinskoi SSR, 133: 177–179. [in Russian]Terentyev PV, Chernov SA (1940) Identification of reptiles and amphibians of the USSR. Leningrad, Uchpedgiz, 184 pp. [in Russian]Thanou E, Jablonski D, Kornilios P (2023) Genome-wide single nucleotide polymorphisms reveal recurrent waves of speciation in niche-pockets, in Europe’s most venomous snake. Molecular Ecology 32: 3624–3640. https://doi.org/10.1111/mec.16944Thunberg CP (1792) Dissertatio entomologica. Sistens insecta Svecica. Quorum partem qvartam. Cons. exp. Facult. Med. Upsal. præside Carol. P. Thunberg, publico examini subjicit Carolus Fredericus Sebaldt, Dalek. in audit. Gust. maj. d. XXIII. Maji MDCCXCII.Tuniyev B, Nilson G, Andrén C (2010) A new species of viper (Reptilia, Viperidae) from the Altay and Saur Mountains, Kazakhstan. Russian Journal of Herpetology 17(2): 110–120.Tuniyev SB, Avcı A, Tuniyev BS, Agasian AL, Agasian LA (2012) Description of a new species of Shield-Head Vipers – Peliasolguni sp. nov. from basin of upper flow of the Kura River in Turkey. Russian Journal of Herpetology 19(4): 314–332.Tuniyev SB, Orlov NL, Tuniyev BS, Kidov AA (2013) On the taxonomical status of steppe viper from foothills of the south macroslope of the East Caucasus. Russian Journal of Herpetology 20(2): 129–146.Tuniyev BS (2016) Rare species of shield-head vipers in the Caucasus. Nature Conservation Research. Zapovednaya Nauka 1(3): 11–25. https://doi.org/10.24189/ncr.2016.023Tuniyev BS, Iremashvili GN, Petrova TV, Kravchenko MV (2018) Rediscovery of the steppe viper in Georgia. Trudy Zoologicheskogo Instituta RAN 322(2): 87–107. https://doi.org/10.31610/trudyzin/2018.322.2.87Tuniyev BS, Petrova TV, Lotiev KY (2023) A New Species of the Genus Darevskia Arribas, 1999 from South Ossetia (Reptilia: Sauria: Lacertidae). Russian Journal of Herpetology 30(4): 237–248. https://doi.org/10.30906/1026-2296-2023-30-4-237-248Tuniyev BS, Ananjeva NB (2024) High herpetological diversity in the Caucasian Ecoregion: an annotated list of species including comments on biogeography and conservation. Zoologicheskij Zhurnal 103(12): 37–76. https://doi.org/10.31857/S0044513424120038Uetz P, Freed P, Aguilar R, Reyes F, Hošek J (eds.) (2025) The Reptile Database, http://www.reptile-database.org. Accessed 31 Jan 2025Utiger U, Helfenberger N, Schätti B, Schmidt C, Ruf M, Ziswiler V (2002) Molecular systematics and phylogeny of Old and New World ratsnakes, Elaphe Auct., and related genera (Reptilia, Squamata, Colubridae). Russian Journal of Herpetology 9(2): 105–124.van Riemsdijk I, Arntzen JW, Babik W, Bogaerts S, Franzen M, Kalaentzis K, Litvinchuk SN, Olgun K, Wijnands JWPM, Wielstra B (2022) Next-generation phylogeography of the banded newts (Ommatotriton): A phylogenetic hypothesis for three ancient species with geographically restricted interspecific gene flow and deep intraspecific genetic structure. Molecular Phylogenetics and Evolution 167: 107361. https://doi.org/10.1016/j.ympev.2021.107361Vedmederja VI, Orlov NL, Tuniyev BS (1986) On the taxonomy of three viper species of the Viperakaznakowi complex. Trudy Zoologicheskogo Instituta AN SSSR 157: 55–61. [in Russian]Veith M, Schmidtler JF, Kosuch J, Baran I, Seitz A (2003) Palaeoclimatic changes explain Anatolian mountain frog evolution: a test for alternating vicariance and dispersal events. Molecular Ecology 12(1): 185–199. https://doi.org/10.1046/j.1365-294X.2003.01714.xVeith M, Fromhage L, Kosuch J, Vences M (2006) Historical biogeography of Western Palaearctic pelobatid and pelodytid frogs: a molecular phylogenetic perspective. Contributions to Zoology 75(3/4): 109–120. https://doi.org/10.1163/18759866-0750304001Waga AS (1876) Nouvelle espèce de Salamandride. Revue et Magasin de Zoologie Pure et Appliquée. Serie 3. Paris 4: 326–328.Werner F (1935) Reptilien der Ägäischen Inseln. Sitzungsber. Akad. Wiss. Wien 144: 81–117.Wied M (1839) Reise in das innere Nord-America in den Jahren 1832 bis 1834 (Vol. 1). J. Hoelscher, 654 pp. https://doi.org/10.5962/bhl.title.51529Wielstra B, McCartney-Melstad E, Arntzen JW, Butlin RK, Shaffer HB (2019) Phylogenomics of the adaptive radiation of Triturus newts supports gradual ecological niche expansion towards an incrementally aquatic lifestyle. Molecular Phylogenetics and Evolution 133: 120–127. https://doi.org/10.1016/j.ympev.2018.12.032Wolterstorff W (1914) Zwei neue Tritonformen der paläarktischen Region. Abhandlungen und Berichte aus dem Museum für Natur-und Heimatkunde zu Magdeburg 2: 371–381.Wolterstorff W (1925) Katalog der Amphibien-Sammlung im Museum für Natur-und Heimatkunde zu Magdeburg: erster Teil: Apoda, Caudata: 155–234.Wüster W (2025) Shedding the mitochondrial blinkers: A long-overdue challenge for species delimitation in herpetology. Vertebrate Zoology 75: 259–275. https://doi.org/10.3897/vz.75.e161536Yablokov AV (ed.) (1976) Sand Lizard: Monographic Description of the Species. Moscow, Nauka. 376 pp. [in Russian]Yanchukov A, Tarkhnishvili D, Erdolu M, Sahin MK, Candan K, Murtskhvaladze M, Gabelaia M, Iankoshvili G, Barateli N, Ilgaz Ç, Kumlutas Y, Matur F, Çolak F, Arakelyan M, Galoyan E (2022) Precise paternal ancestry of hybrid unisexual ZW lizards (genus Darevskia: Lacertidae: Squamata) revealed by Z–linked genomic markers. Biological Journal of the Linnean Society 136(2): 293–305. https://doi.org/10.1093/biolinnean/blac023Zinenko O, Sovic M, Joger U, Gibbs HL (2016) Hybrid origin of European vipers (Viperamagnifica and Viperaorlovi) from the Caucasus determined using genomic scale DNA markers. BMC Evolutionary Biology 16(76): 1–13. https://doi.org/10.1186/s12862-016-0647-7Supplementary materials10.3897/caucasiana.5.e189214.suppl1B2989292-C9AA-5CA2-AF6E-0CF6FFB5113E
Supplementary table contains location name and geographic coordinates, CaBOL identification number, and BOLD BIN for the reference specimens.
https://binary.pensoft.net/file/1711449This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Tarkhnishvili D, Iankoshvili G, Seropian A, Kachlishvili, N, Todua M