Высокое герпетологическое разнообразие Кавказского экорегиона: аннотированный список видов, включая комментарии по биогеографии и охране

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The Caucasus has always been one of the most alluring places for herpetologists. In spite of a long-term history of herpetological research in this unique region, many issues remain unclear ranging from taxonomic diversity, different views on taxonomic position and intraspecific structure of herps inhabiting the Caucasus.

Due to its geographical location, properties of landscapes and climates, and geological history of the formation the Caucasus Isthmus fauna has been characterized by exceptional high diversity of its flora and fauna. Recognition of this fact and obvious evidences of biodiversity loss have driven the Critical Ecosystem Partnership Fund (CEPF), an organization dedicated to promotion of global conservation in biodiversity-rich areas, to designate the Caucasus ecoregion in 2004. According to World Wide Fund for Nature (WWF), this region is among the world’s 200 ecoregions containing biodiversity of global importance. As to CEPF (www.panda.org/caucasus/cepf), the Caucasus belongs to 25 biodiversity hotspots that are biologically most diverse, but also most vulnerable to environmental degradation (Zazanashvili et al., 2004). These 25 hotspots are selected on a basis of species diversity per unit of area: covering only 1.4% of the Earth’s land, they accommodate about 44% of all species of vascular plants and 35% of all species in four terrestrial groups (Myers et al., 2000). Hence, the Caucasus has been globally perceived as a problematic ecoregion with unique flora and fauna which is inhabited by rare, relict and endemic species and which has a global importance for preservation of our planet’s biological diversity.

Fig. 1. The map of the Caucasus ecoregion.

The Caucasus ecoregion, historically considered as the isthmus between the Black Sea, Sea of Azov and the Caspian Sea, covers an area of 580,000 km² and comprises Abkhazia, Armenia, Azerbaijan, Georgia, Ciscaucasian part of the Russian Federation, South Ossetia, northeastern Turkey and northwestern Iran (Fig. 1). To reflect the fullness of biodiversity, five priority areas (corridors) were identified. They serve as home to the majority of threatened species and their relatively preserved distribution areas (Fig. 2). These are the corridors of the Greater Caucasus Ridge, western and eastern corridors of the Lesser Caucasus, Caspian and Hyrcanian corridors. The governments of all countries of the Caucasus ecoregion have joined important international conventions to reinforce their efforts in biodiversity conservation.

Fig. 2. The regions of conservation priority and their corridors in the Caucasus ecoregion with our modifications highlighted in red (from Tuniyev et al., 2019).

Research and progress of environmental strategies and priorities in the Caucasus require current knowledge of taxonomic and ecological diversity of animal species and understanding of trends in range structure of these species.

In the first ecoregion conservation plan for the Caucasus (Williams et al., 2006), there were mentioned 77 species of reptiles in the Caucasus with 28 endemics among them and 14 species of amphibians with 4 endemic species. In 2010, the Caucasus Biodiversity Council (CBC) and WWF proposed revising the ecoregion conservation plan. Since then, the herpetofauna of Ecoregion has undergone a number of major changes due new finds of species, description of new species and subspecies, progress of molecular-research and observations in poorly visited areas (mainly high-mountains). That is why, the main task is to support and contribute to the concluding steps of the ECP 2012 revision process through compilation and structuring updated information on Reptiles and Amphibians of the Caucasus ecoregion. The Caucasus has been characterized by an extreme high diversity of vipers Vipera sensu lato (19 of approximately 40 species in the world fauna), the Pelias species (14 of 22 known species) as well as the rock lizards (Darevskia spp.) (27 of 41 species). Many of these species occupy total ranges of only a few thousand square kilometres. Discovering of the high level of diversity of shield-head vipers in the Caucasus ecoregion is essential to establish the conservation status of rare narrow spread species, the vast majority of whom are included in the IUCN Red List of Threatened species (2024). The lizards of the genus Darevskia from the Lacertidae family demonstrate particular diversity. In other words, this area plays a key role in the conservation of the variety of species of shield-head vipers and lacertid lizards (Tuniyev, 2016).

Fig. 3. Endemic Caudata Amphibians: 1 – Lissotriton lantzi, 2 – Paradactylodon persicus, 3 – Mertensiella caucasica, 4 – Mertensiella djanaschvilii.

Therefore, we provide the update full list of herpetofauna of the Caucasian ecoregion on the base of literature sources, databases and own results of study of the Caucasian herpetofauna. The taxonomic and nomenclature changes and additions follow recent literature sources and databases (Frost, 2024: Uetz et al., 2024; IUCN Red List of Threatened species, 2024) with some modifications. Knowledge about biodiversity remains inadequate because most species living on Earth were still not formally described (the Linnean shortfall) and because geographical distributions of most species are poorly understood and usually contain many gaps (the Wallacean shortfall) (Bini et al., 2006; Hortal et al., 2015).

Fig. 4. Distribution of Ommatotriton ophtyticus.

We did not set the goal of listing all the synonymy in the taxonomy of amphibians and reptiles of the Caucasus ecoregion, as well as a detailed discussion of the existing different and sometimes contraversal views on the status of different species/subspecies. At the same time, on the base of current data and own research of this region for many decades, we offer our opinion on the composition and distribution of herpetofauna in the Caucasus ecoregion with comments on taxonomic and conservation status of included taxa. Type territories indicated for endemics of the Caucasus only given in the tables 1–4. In these tables, we marked with an asterisk those species which status is under discussion and a revision of their status is possible. For the conservation purposes many forms under consideration are included in the Red Data lists including the IUCN Red List of Threatened species (2024), and accessed a certain conservation status.

AMPHIBIANS AND REPTILES DIVERSITY

Amphibians. During the last decades, several forms of amphibians were described from the Caucasus ecoregion, such as Bufo verrucosissimus tertyshnikovi Kidov, 2009 from Stavropol Highland (Russia); Bufo eichwaldi Litvinchuk, Borkin, Skorinov et Rosanov, 2008 and Hyla orientalis gumilevskii Litvinchuk, Borkin, Skorinov et Rosanov, 2008 from Talysh Mountains (Azerbaijan). Genus’s names’ changes took place in salamanders (from Triturus Rafinesque, 1815 (part) to Lissotriton Bell, 1839 and Ommatotriton Gray, 1850, from Batrachuperus Boulenger, 1878 and Iranodon Dubois and Raffaëlli, 2012 to Paradactylodon Risch, 1984, Green Frogs (from Rana Linnaeus, 1758 to Pelophylax Fitzinger, 1843) and in Green Toads (from Bufo Garsault, 1764 to Bufotes Rafinesque, 1815). Taxonomic status was changed for Shelkownikow’s treefrog to Hyla orientalis Bedriaga, 1890; taxonomic position rised up with changes of genus names at Banded Newt – Ommatotriton ophryticus (Berthold, 1846) and Lantz’s Smooth Newt – Lissotriton lantzi (Wolterstorff, 1914). Taxonomic status of Karelin’s Crested Newt – Triturus karelinii (Strauch, 1870) was changed to species level. Beside, Persian Mountain Salamander – Iranodon persicus (Eiselt et Steiner, 1970) was absent in List of the Caucasus ecoregion species (Zazanashvili, 2009). Only 40 years after the changes in status to the species level of Bufo verrucosissimus (Pallas, 1814) (Orlova, Tuniyev, 1989), the species status was recognized as valid based on the results of molecular genetic studies (Özdemir et al., 2020). The species status of Mertensiella djanaschvilii (Tartarashvili et Bakradze, 1989) (Frost, 2024), was not immediately recognized. The problems of species diversity in the genera Bufotes Rafinesque, 1815 and Pelophylax Fitzinger, 1843 remains open.

Table 1. List of amphibian species of the Caucasian ecoregion

As for 1 July 2024 there are known 18 amphibian species belonging to the two orders. The order Caudata in the Caucasian ecoregion includes two families: Hynobiidae with one genus Paradactylodon (one species) and Salamandridae with four genera: Mertensiella (two species), Triturus (one species), Ommatotriton (one species), and Lissotriton (one species). The order Anura is represented with five families: Bombinatoridae with one genus Bombina (one species); Bufonidae with two genera Bufo (two species) and Bufotes (one species); Hylidae with one genus Hyla (two species); Pelobatidae with one genus Pelobates (two species); Pelodytidae with one genus Pelodytes (one species); Ranidae with two genera Pelophylax (one species) and Rana (two species). The half (nine species) of these amphibians are endemics (Table 1, Fig. 3–6).

Fig. 5. Endemic Toads of the Caucasus ecoregion: 1a – Bufo verrucosissimus circassicus, 1b – B. v. turowi, 1c – B. v. tertyshnikovi, 1d – B. v. verrucosissimus, 2 – B. eichwaldi.

Fig. 6. Distribution of Pelodytes caucasicus.

The Caucasian Salamander (Mertensiella caucasica) and Djanashvili’s Salamander (Mertensiella djanaschvilii) are found in the western part of Lesser Caucasus in Georgia and in Pontic Ridge in Turkey only with slightly irradianions beyond of ecoregion. Persian Salamander (Paradactylodon persicus) occurs in Hyrcanian forests of the Alborz Mountains in Iran. Banded Newt (Ommatotriton ophtyticus) and Lantz’s Smooth Newt (Lissotriton lantzi) are distributed mainly in western half of ecoregion. Caucasian Parsley Frog (Pelodytes caucasicus) and Colchis Toad (Bufo verrucosissimus) live in mountain forests of the Western Caucasus with a limited reach to Central Trancaucasus and separately on uppermost part of Stavropol Upland. Hyrcanian Toad (Bufo eichwaldi) is endemic of Hyrcanian forests in the Talysh and Alborz Mountains. Five species were included in the IUCN Red List in categories NT (Table 1); four amphibian species still not assesed.

Reptiles. Previously 77 species of reptiles were registered in the Caucasus (Williams et al., 2006). The update list includes 110 species: 5 species of the turtles and tortoises, 59 species of lizards and 46 species of snakes (Table 2–4). Among them 32 species (44 subspecies) are endemic to the ecoregion.

Table 2. List of turtles and tortoises species of the Caucasian ecoregion

The members of four families present the Caucasian fauna of the turtles and tortoises. They are Emydidae with two genera Emys Dumeril, 1806 (one species with four subspecies); Trachemys Agassiz, 1857 (one introduced species); Geoemydidae with one genus Mauremys Gray, 1869 (one species); Cheloniidae with one genus Chelonia Brongniart, 1800 (one species) and Testudinidae with one genus Testudo Linnaeus, 1758 (one species with four subspecies). Three species were included in the IUCN Red List in categories from NT to EN; one species (Mauremys caspica) still not assessed (Table 2). A confusing situation arose around subspecies structure of Emys orbicularis and Testudo graeca. Eichwald described Emys orbicularis iberica from Astrakhan. There, according to modern concepts, lives the nominative subspecies. Animals from southern Dagestan and Transcaucasia were described as subspecies E. o. kurae Fritz, 1994. Later E. o. kurae was transferred to the list of synonyms (Uetz et al., 2024). However, we considered it possible to apply this name to turtles of this sector of the Caucasus ecoregion. It is important to note that the intraspecific taxonomy of Emys orbicularis and Testudo graeca remains controversial, also to the point of denying the validity of Testudo graeca nikolskii Ckhikvadze et Tuniyev, 1986 and synonymizing it with Testudo graeca ibera Pallas, 1814 (Türkozan et al., 2021).

The Caucasian saurofauna (Table 3) is characterized by high taxonomic diversity and is presented by the members of five families. They are: Agamidae with three genera, Paralaudakia Baig, Wagner, Ananjeva et Bohme, 2012 (one species), Trapelus Cuvier, 1817 (two species) and Phrynocephalus Kaup, 1825 (four species); Gekkonidae with three genera Mediodactylus Szczerbak et Golubev, 1977 (one species), Hemidactylus Goldfuss, 1820 (one introduced species) and Tenuidactylus Szczerbak et Golubev, 1984 (one species); Anguidae with two genera Anguis Linnaeus, 1758 and Pseudopus Merrem, 1820, with one species each; Scincidae with three genera:.Ablepharus Fitzinger in Eversmann, 1823 (three species), Eumeces Wiegmann, 1834 (one species) and Heremites Gray, 1845 (one species). The most taxonomically (42 species) and phylogenetically diverse is Lacertidae (Bonaparte, 1831) family. It is presented by eight genera: Darevskia Arribas, 1997 (29 species), Iranolacerta Arnold, Arribas et Carranza, 2007 (one species), Eremias Wiegmann, 1834 (five species), Lacerta Linnaeus, 1758 (three species), Ophisops Menetries, 1832 (one species), Parvilacerta Harris, Arnold et Thomas, 1998 (one species), Phoenicolacerta Arnold, Arribas et Carranza, 2007 (one introduced species) and Podarcis Wagler, 1830 (one introduced species) (Table 3). According to the First Ecoregion Conservation Plan for the Caucasus (Willams et al., 2006), that time from the known 26 species of genus Darevskia in the world, 25 species were noted in the Caucasus, with 23 endemics of ecoregion. The next Ecoregion Conservation Plan for the Caucasus mentioned 21 besexual species (36 subspecies) and four parthenogenetic species (Tuniyev et al., 2020). The Caucasian Ecoregion is one of the main centre of taxonomical diversity of rock lizards from genus Darevskia. The present upgrated list includes 24 bisexual species (40 subspecies) within Caucasus ecoregion from the known 31 bisexual species (54 subspecies) in the world and 5 parthenogenetic species of seven in the world fauna (Table 3). In recent decades, several species and subspecies have been described in the Caucasus ecoregion, including Darevskia caucasica vedenica (Darevsky et Roitberg, 1999), D. brauneri myusserica Doronin, 2011, D. praticola hyrcanica Tuniyev, Doronin, Kidov et Tuniyev, 2011, D. praticola loriensis Tuniyev, Doronin, Tuniyev, Aghasyan, Kidov et Aghasyan, 2013, D. rudis mirabilis Arribas, Ilgaz, Kumlutaş, Durmuş, Avci et Üzüm, 2013, D. aghasyani Tuniyev et Petrova, 2019, D. tuniyevi Arribas, Candan, Kurnaz, Kumultas, Caynak et Ilgaz, 2022 and D. arribasi Tuniyev, Lotiev et Petrova, 2023. D. raddei and D. defilippi is now regarded as a complex of cryptic species as well as D. chlorogaster (Ahmadzadeh et al. 2013). A number of subspecies have been elevated to species level, such as Darevskia brauneri (Méhely, 1909), D. szczerbaki (Lukina, 1963), D. pontica (Lantz et Cyrén, 1918), D. nairensis (Darevsky, 1967), D. alpina (Darevsky, 1967), D. daghestanica (Darevsky, 1967), D. adjarica (Darevsky et Eiselt, 1980). At the same time, many open questions remain with the systematics of the Caucasian rock lizards. The allocation of subspecies to species such as Darevskia rudis (Bedriaga, 1886) and Darevskia valentini (Boettger, 1892) has been questioned (Candan et al., 2021). The status of taxa (species/subspecies) of the “Darevskia praticola – pontica” complex is controversial (Tuniyev et al., 2011; Freitas et al., 2016; Speybroeck et al., 2020). In addition, the subspecific rank of D. p. hyrcanica and D. p. loriensis is recognized by some authors (Freitas et al., 2016) and rejected by others (Saberi-Pirooz et al., 2018). The validity of the name Darevskia dryada (Darevsky et Tuniyev, 1997) is disputed (Schmidtler et al., 2002); Arribas et al. (2021) argue the ambiguity of such a conclusion. It was suggested that the continuous range and clinal variability revealed in Darevskia derjugini (Nikolsky, 1898) indicate monotypy and conspecificity of the described subspecies (Tuniyev, Ostrovskikh, 2006), which is also confirmed in the Turkish part of the species range (Kurnaz et al., 2017). The known parallels in the abdominal coloration and osteology characters of Darevskia parvula (Lantz et Cyrén, 1913) and Darevskia defilippii (Camerano, 1877) in two mesophilic tertiary refugia of the Caucasus ecoregion (Colchis and Hyrcanica) require further more detailed study of these species and their relationships (Arribas, 2012). In our opinion, the species status for Darevskia portschinskii nigrita (Bakradze, 1976), D. caucasica vedenica (Darevsky et Roitberg, 1999) and Lacerta agilis boemica Suchow, 1929 cannot be ruled out. Forteen species were included in the IUCN Red List in categories from NT to CR, one species has category DD, and eight species are still not assessed (Table 3).

Table 3. List of lizard species of the Caucasian ecoregion

The analysis of taxonomic diversity of the Caucasian snakes shows that new records, phylogenetic studies and taxonomic revisions have led to the most significant changes in the numbers of genera and species. Recent taxonomic views were made on the base of the new results of study of the phylogeny of advanced snakes (Colubroidea) (Pyron et al., 2010) with resurrection of the family Psammophiidae Boie in Fitzinger, 1826 in particular with genera Malpolon Fitzinger, 1826 and Psammophis Boie in Fitzinger, 1826 for the Caucasian herpetofauna. Tuniyev et al., (2019) carried out a detailed analysis of this fauna according to its status as of 2019. This monograph provides a comprehensive bibliography on all aspects of the diversity of the ophidiofauna of the Caucasus up to 2019. In present paper, due to its limited volume, we use and review more recent works in this field, as well as new descriptions. All snake species occurring in the Caucasus are related to the five families. They are Typhlopidae with one genus Xerotyphlops Hedges, Marion, Lipp, Merin et Vidal, 2014 (one species), Boidae with genus Eryx Daudin, 1803 (two species) and taxonomically and phylogetically diverse family Colubridae (ten genera with 21 species) with two subfamilies: Natricinae with one genus Natrix Laurenti, 1768 (three species), and Colubrinae with nine genera: Dolichophis Gistel 1868 (two species), Hemorrhois Boie, 1826 (two species), Platyceps Blyth, 1860 (one species), Coronella Laurenti, 1768 (one species), Eirenis Jan, 1863 (four species), Elaphe Fitzinger, 1833 (three species), Zamenis Wagler, 1830 (three species), Rhynchocalamus Günther, 1864 (one species) and Telescopus Wagler, 1830 (one species). The family Psammophiidae is represented in the Caucasian ecoregion by two genera: Malpolon (one species) and Psammophis (one species).

Fig. 7. Endemic сolubrids of the Caucasus ecoregion: 1 – Natrix megalocephala, 2 – Rhynchocalamus satunini, 3 – Zamenis persicus.

The assessment of taxonomic diversity of the groups Elaphe sensu lato and Coluber sensu lato is problematic and undergoing radical changes, stimulated by remarkable discoveries of new taxa. The system of colubrid systematics hinges on the results of intensive phylogenetic and taxonomic studies, research of nomenclature and colubrid descriptions (Salvi et al., 2018; Tuniyev et al., 2019; Uetz et al., 2024) (Fig. 7). Seventeen species were included in the IUCN Red List in categories from NT to CR, one species has category DD, and three species still not assessed (Table 4).

Table 4. List of snake species of the Caucasian ecoregion

The diverse family (20 species) of Viperidae family includes the members of two subfamilies: Crotalinae with one genus Gloydius Hoge et Romano-Hoge, 1981 (one species) and Viperinae with four genera: Montivipera Nilson, Tuniyev, Andrén, Orlov, Joger et Herrmann, 1999 (three species), Macrovipera Reus, 1927 (one species), Pelias Merrem, 1820 (14 species) and Vipera Laurenti, 1768 (one species). The Caucasus has been characterized by an extreme diversity of vipers Vipera sensu lato (19 out of approximately 40 species of this genus occur here) and represents the node of taxonomic diversity of the Pelias species (14 of 22 known species).

Fig. 8. Endemic vipers of kaznakovi-complex of the Caucasus ecoregion: 1 – Pelias orlovi, 2 – P. magnifica, 3 – P. dinniki, 4 – P. kaznakovi, 5 – P pontica, 6 – P. olguni, 7 – P. tuniyevi, 8a – P. darevskii darevskii, 8b – P. d. kumlutasi, 8c – P. d. uzumorum.

Cryptic speciation plays a significant role in the formation of this group. Tracking the history of research of the Pelias vipers in the Caucasus in the past 100 years one can find a trend in description of cryptic species from the Caucasus vipers of P. kaznakovi – P. renardii species groups (Tuniyev, 2016; Tuniyev et al., 2019; Freitas et al., 2020) (Fig. 8, 9).

Fig. 9. Endemic vipers of ursinii-renardi-complex of the Caucasus ecoregion: 1 – Pelias lotievi, 2a – P. shemakhensis kakhetiensis, 2b – P. sh. shemakhensis, 3 – P. eriwanensis.

The results of recent phylogenetic research of the Caucasian vipers have demonstrated the polyphyly of the evolutionary lineages of P. ursinii, P. orlovi, P. renardi and P. dinniki, making the existence of complexes of cryptic species and/or the hybrid origin of some species quite possible (Zinenko et al., 2015, 2016; Thanou et al., 2023). The latest genome-wide data show the nuclear diversity of Palearctic vipers and supports the results obtained by ddRAD-seq analyses on particular species sets (Zinenko et al., 2016; Joger, Zinenko, 2021; Thanou et al., 2023; Dufresnes et al., 2024). They broadly corresponds to the mitochondrial phylogeny (Freitas et al., 2020) and underlines the importance of the hybridization both within and between the major clades. As many species living in the Caucasus ecoregion have disjunctive ranges, their distribution patterns can be considered as the result of relict insularization.

The discovery of a high level of diversity of Pelias vipers in the Caucasus has important implications for the assessment of conservation status of rare and narrowly distributed species many of which are recorded on the IUCN Red List of Threatened Species (Table 4). In the future, we should expect changes in the taxonomic composition of the ophidiofauna because of new records, discovery of new cryptic taxa of vipers and new revisions of taxonomically complicated genera of colubrids and viperids.

A number of species mentioned for the Caucasus ecoregion were not confirmed: (Pelophylax lessonae (Camerano, 1882), Chamaeleo chamaeleon (Linnaeus, 1758), either provided by local finds of the alien species, in particular Phoenicolacerta laevis (Gray, 1838) from western Georgia, Trachemys scripta elegans (Wied-Neuwied, 1839) in Krasnodar and Stavropol Territories and Dagestan, Hemidactylus turcicus (Linnaeus, 1758) in Sochi (Kukushkin et al., 2017; Dunaev, Imshenitzky, 2018; Tuniyev et al., 2023).

Fig. 10. Similarity of the batrachofauna of the countries of the Caucasus ecoregion according to the Jaccard coefficient.

Fig. 11. Similarity of the batrachofauna of the countries of the Caucasus ecoregion according to the Sörensen-Chekanovsky coefficient.

Comparative analysis of the herpetofauna of the countries of the Caucasus ecoregion, using cluster analysis of the values of the Sorensen–Czekanovsky and Jaccard coefficients by the method of Ward (1963), showed identical results on the dendrograms (Fig. 10– 13).

Fig. 12. Similarity of the herpetofauna of the countries of the Caucasus ecoregion according to the Jaccard coefficient.

Fig. 13. Similarity of the herpetofauna of the countries of the Caucasus ecoregion according to the Sörensen- Chekanovsky coefficient.

At the same time, the results of the analysis for amphibians and reptiles were somewhat different. Stably, common clusters for amphibians and reptiles were formed by Azerbaijan and Iran. In terms of the level of similarity of the batrachofauna, the Colchis refugium contributes to the formation of a common cluster of Georgia and Turkey, with sufficient proximity to Russia, while the dominant mountainous xerophilic landscapes of the Armenian Highlands affect the high similarity of the reptile fauna of Armenia with Turkey. The presence of both mesophilic and xerophilic landscapes in Russia and Georgia unites the reptilian fauna of these countries into a common cluster, which is closest to the single cluster of small areas of the southern macroslope of the Greater Caucasus in Abkhazia and South Ossetia.

ENDEMIC SPECIES: RICHNESS AND CONSERVATION STATUS AND THREATS

Endemic species richness is high in the Caucasian ecoregion (Russia, states of Transcaucasia, northeast Turkey and Caspian Iran). They are concentrated in a number of refugia, both mesophyllic (Caucasian Black Sea coast of Russia, Georgia, Turkey; southeast Azerbaijan and Caspian coast of Iran) and xerophylous (Turkey eastward from Anatolian Diagonal; Armenia, Azerbaijan (Nakhichevan, Zuvand), Georgia (Vashlovani), Russia (Dagestan, semiarid Hollows in East Caucasus) (Tuniyev et al., 2019). The updated list includes 18 amphibians’ species with endemic 13 subspecies of 10 species, nine species of which occur in Caucasian ecoregion only (Table 1). Among 110 reptile species in the ecoregion, 32 are endemic species and 44 are endemic subspecies (Tables 2–4). The distribution of the number of endemic species across the countries of the ecoregion is uneven and depends on both the area occupied and the landscape diversity. The maximum number of endemics inhabit the territories of Georgia (28 species), Turkey (24 species) and Russia (19 species) (Fig. 14).

Fig. 14. Representation of endemic species in the countries of the Caucasus ecoregion (see Chapter Endemic Species: Richness and Conservation Status and Threats and Table 5).

Taking into account the artificiality of the administrative approach to assessing the originality of faunas, we note the importance of understanding the volume of biodiversity of endemic species both for the countries themselves and for the adoption of transboundary conservation strategies.

Globally threatened species: CR, EN, VU redlisting categories according to IUCN. We compiled a dataset of 18 extant species of amphibians and 110 species reptiles occurring in the Caucasus ecoregion from the Reptile Database and IUCN Red List of Threatened Species. Of these, 112 have published assessments of extinction risk on the IUCN Red List. Twenty-two species (28 subspecies, 19.6%) were assessed as threatened with extinction (categories Vulnerable, Endangered and Critically Endangered). Two species has category Data Deficient (DD) and 17 species – Near Threatened (NT). Therefore, 71 species (63.4%) belong to category Least Concern (LC) in Caucasus ecoregion. Sixteen species (9.6%) are still not assessed, due to late descriptions or changes in taxonomy. In addition, the assessments are absent for almost all subspecies, with exceptions of Testudo graeca nikolskii. The declining populations present majority of species.

Regionally threatened species. In the Caucasus ecoregion, as in many other regions of the world, the conservation of herpetofauna is actually effective only in nature reserves and national parks, although it is formally organized in protected areas of a lower level. Therefore, the analysis of compliance of the extant Econet (network of protected areas) to the needs of herpetofauna preservation becomes a profoundly important task. .It should be emphasized that despite the inclusion of many species in regional Red Books (Table 5), the normative form of protection, as experience shows, is not very productive and for the effective conservation of the herpetofauna of the Caucasian ecoregion. The recent international experience shows that that a territorial form of protection is optimal and should be applied as an effective conservation measure for the Caucasian ecoregion.

Table 5. Representation of amphibian and reptile species in the regional red data books of the Caucasian ecoregion

Notes: In horizontal: 1 – Russian Federation (RF); 2 – Azerbaijan; 3 – Armenia; 4 – Georgia; 5 – Adygey, RF; 6 – Dagestan, RF; 7 – Ingushetia, RF; 8 – Kabardino-Balkaria, RF; 9 – Kalmykia, RF; 10 – Karachay-Cherkessia, RF; 11 – Krasnodar Territory, RF; 12 – Rostov Region, RF; 13 – North Ossetia – Alania, RF; 14 – Stavropol Territory, RF; 15 – Chechen Republic, RF; 16 – South Ossetia.

In columns in uppercase: 1 – subspecies Emys orbicularis colchica Fritz, 1994; 2 – populations from Crimea Peninsular and Caucasian Black Sea coast; 3 – populations from Crimea Peninsular and Krasnod Territory; 4 – subspecies Lacerta agilis grusinica Peters 1960 and L. a. mzymtensis (Tuniyev S. et Tuniyev B., 2008); 5 – populations from Caucasian Black Sea coast; 6 – populations from Caucasian Black Sea coast; 7 – within Elaphe sauromatmes; 9 – populations from Caucasian Black Sea coast; 10 – populations of Crimea Peninsular and Precaucasia; 11 – named as Phrynocephalus helioscopus; 12 – named as Phrynocephalus helioscopus; 13 – within Elaphe sauromatmes; 14 – within Pelias renardi; 15 – mentioned as Pelias kaznakovi; 16 – Black Sea coast population; 17 – mentioned as Pelobates vespertinus; 18 – subspecies Lacerta agilis exigua; 19 – subspecies Eremias arguta transcaucasica; 20 – subspecies Darevskia caucasica vedenica; 21 – mentioned as Anguis fragilis; 22 – mentioned as Triturus vulgaris; 23 – mentioned as Hyla arborea; 24 – within Mertensiella caucasica; 25 – mentioned as Malpolon monspessulanus Mertens et Mueller 1928.

The Priority Conservation Areas (PCA) and the corridors between them designated by CEPF in the Caucasus (Fig. 2) almost fully represent the key areas of the most important ecosystems in Armenia, Georgia, Azerbaijan and the neighbouring areas of Turkey and Iran. In these countries, the regional Econets are delineated as based mainly on the existing or planned protected areas. The Caucasian parts of Turkey and Iran are nearly fully set aside as priority areas. The situation in Russia and Abkhazia is different. The central axial part of highlands of the Greater Caucasus Ridge is defined quite artificially and almost deprived of herpetofauna. The vast tracts of the Black Sea coast, remnants of pristine steppes and meadow-steppes in the West and Central Ciscaucasus, unique dry steppes and sandy lands in the East Ciscaucasus, foothills and mid-elevations in the northern Jurassic depression between the Skalistyi and Bokovoy ridges, foothills and maritime Dagestan regions remain greatly underrepresented in protected areas (Tuniyev et al., 2019).

Biodiversity should be preserved in its intrinsic diversity and complexity, but the foremost regional goal and responsibility is set on saving endemic species from extinction. The status of such endemic species as Natrix megalocephala, Zamenis persicus, Pelias dinniki, P. lotievi and Montivipera raddei is quite safe because a substantial part of their ranges is covered by protected areas. Contrary to this, a number of other species are represented in protected areas insignificantly (Phrynocephalus horvathi, Darevskia szczerbaki, Eremias pleskei, Pelias kaznakovi, P. magnifica, P. eriwanensis, P. ebneri, P. tuniyevi) or not at all (Trapelus sanguinolentus sanguinolentus, Darevskia aghasyani, Darevskia arribasi, Darevskia caucasica vedenica, Darevskia dryada, Pelias orlovi, P. pontica, Eryx miliaris nogajorum). Moreover, the habitats of Pelias orlovi and P. magnifica were not even embraced by CEPF’s PCAs.

To preserve the Szczerbak’s Rock Lizard and Orlov’s viper, it is essential to establish a reserve or a national park between Gelenjik and Jubga, incorporating the peaks Oblego and Papai, and to expand the territory of Sochi National Park so that to include the Mt. Bolshoi Pseushkho (Tuniyev, 2016). Some of these areas are in establishing now. Conservation of the Relict viper also implies adding the Malyi Bambak Ridge to Caucasian Biosphere Reserve. The Black Sea viper can be saved solely by creating a national park near Borçka. The Lake Arpi National Park was established in Armenia in 2009 largely for the protection of the Darevsky’s viper. The Steppe Agama, Desert Sand boa and associated psammophilic communities can be protected only through the establishment of Nogaysky Steppe Reserve. The Shemakha and Kakhety Steppe Vipers need special protected areas near Shamakhi in Azerbaijan and on Shirak Plateau in Georgia.

The next regional goal is focused on conservation of globally threatened or endangered species of wide distribution, in the Caucasus – Pelias renardi, Elaphe sauromates and Dolichophis caspius. The first species is left beyond the PCAs and the last two are almost neglected. No large protected areas can be set aside for Pelias renardi for the strong patchiness of its remnant habitats. However, it is feasible to establish zoological natural monuments or cluster sites of existing protected areas, e. g. in the East Priazovie (Yasenskaya Spit and others), foothills of the West and Central Caucasus (vicinities of the Raevskaya village, Maikop and Kislovodsk towns, Gerpegem Ridge), along the Kuma-Manych Depression (Lake Manych-Gudilo shoreline). These areas are also important for preservation of the Caspian whip snake and the Eastern four-lined ratsnake, but it also demands for the establishment of new protected areas between the Cape Bolshoi Utrish and the Kabardinka Settlment on the Black Sea coast, in Primanychie (area around the Lake Manych-Gudilo) and the Kudriavaya (Curly) Ravine in the Stavropol Territory. Caspian whip snakes will also benefit from the above-mentioned Nogaysky Steppe Reserve.

Particular attention should be paid to the hotspots of amphibians and reptiles species diversity in the Caucasian ecoregion. Such areas are the Black Sea coast (excluding the Rioni Lowland), lower çoruh basin, semi-arid depressions in the East Ciscaucasus, maritime Dagestan, Artwin Hollow, Kura-Arax Lowland (aside from its deserts), Arax riverside in Armenia and Azerbaijan, and the Talysh–Alborz massif.

The analysis of biological diversity, spatial and altitudinal distribution of existing protected areas, including water bodies and wetlands, demonstrates the inability of protected areas to fulfill their goals and to ensure sustainable management of natural ecosystems in their complexity (Krokhmal, Tuniyev, 2003). First, the Econet in Russian Caucasus and Abkhazia needs much improvement (Fig. 2). In these two countries, protected areas are concentrated in mid-elevation and highland areas of the West and Central Caucasus to protect their natural heritage. In Chechen and Dagestan Republics (East Caucasus), highlands and mid-elevations are not protected, including the unique limestone region of Inner Dagestan saturated with xerophilic endemics and relicts. In nearly all the North Caucasus, protected areas with stringent protection regime (reserves) are lacking at low elevations 800–1200 m a. s. l. The two areas retaining the unique for Russia Mediterranean semi-arid juniper and pistachio-juniper forests (Anapa – Gelenjik and foothills of Dagestan) are out of state protection as well. The northernmost enclave of the Hyrcano-Alborz biota in the Samur mouth is in Samur National Park in Dagestan and Samur-Yalama National park is located in Azerbajan.

Much effort in international cooperation is required to establish such transboundary protected areas as East Caucasus Transboundary Area (Russia, Azerbaijan, and Georgia), North-West Caucasus (Kavkazsky, Pskhu-Gumista reserves, Teberdinsky, Ritsa Relict and Sochi National Parks in Russia and Abkhazia) and Kolkhida International Biosphere Polygon (Sochi National Park, Psou – Bzyb interfluve and Pitsunda-Mussera Reserve in Russia and Abkhazia). Implementation of this Econet development programme, along with other conservation measures described above for preservation of endemic and threatened species, will allow approaching the long-term goal of sustainable conservation of herpetofauna and other biodiversity in the Caucasian ecoregion.

MAIN THREATS AND PROPOSED CONSERVATION ACTIONS

The main threats to ECA reptiles, according to the IUCN Red List of Threatened Species (2024) are agriculture, residential/commercial development, and biological resource use. These threats primarily cause habitat fragmentation and loss (Visconti et al., 2018).

The major threat of habitat loss affects in particular relic forest species, and species of the steppe and semi-desert ecosystems, which are often not able to persist on agricultural and other transformed lands. Eremias pleskei (Armenia, Azerbaijan, Turkey and Iran) is listed as Critically Endangered, based on a population decline of more than 80% over ten years; its natural sandy habitat has virtually disappeared due to human disturbance. For habitat specialists, such as Phrynocephalus horvathi (Critically Endangered) which is primarily limited to patches of saltwort, wormwood semi-desert, and highly specific soils, habitat conversion can have a major impact. The disappearance of steppe vipers of the “ursinii-renardi” complex throughout most of the previously occupied habitats in the ECA is associated with ploughing of steppes for agriculture (Tuniyev, 2016).

Significant threats include the illegal capture of commercially valuable species for the pet trade (all representatives of the vipers, turtles, and some species of lizards) in Turkey, the Caucasus and Central Asia. Prosecution of snake species continues in the area, especially in Turkey, the Caucasus, southern regions of Russia, which is associated with low levels of environmental education (Visconti et al., 2018).

Invasive and other predatory species play a particularly important role for endemic and relic species. For example, Procyon impact is very high to amphibians in Western Caucasus and Lenkoran-Talysh region.

Climate change is likely to play a major role in the region in the future. Climate change has led to an increase in summer temperatures and length of dry summer period in the Western Caucasus, resulting in a reduction of habitats of mesophylic Colchis reptile species (Darevskia derjugini) and the increase in the number of Eastern Mediterranean species of snakes (Dolycophis caspius, Platiceps najadum) on the Black Sea Coast.

Other threats are less prominent in the IUCN Red List of Threatened species (2024) such as pollution; however, a recent risk evaluation of pesticide use to protected European reptiles suggests that ten species, including all six Habitat Directive Annex II turtles, are at above-average pesticide risk (Visconti et al., 2018).

Some examples for conservation strategy:

Aghasyan’s Lizard – Darevskia aghasyani. Conservation status of taxa assessed as CR B2ac(iv) “Critically Endangered”, because on the basis of expert estimates found that the habitat area is less than 10 km², it may have fragmented, but at the moment consists of only one locality and marked by extreme fluctuations in number of mature individuals (Tuniyev, Petrova, 2019). Previously Urts Ridge was part of “Khosrov Forest” State Reserve, as a cluster area. Given the exceptional uniqueness of flora and fauna of this ridge, description of about ten species of plants from Urts Ridge and the breeding for Mouflon, together with our species of relic described lizards, it is advisable to consider returning Urts Ridge to the Khosrov Reserve.

Darevsky’s Viper – Pelias darevskii. The new finds of species in Armenia (Agasian, Agasian, 2008) were located in direct closeness (12 km) from type locality and actually did not extend the known distributional area of species. In July 2014 the species actually was found by us in Ninotsminda, Akhalkalaki, Aspidindza, Akhaltsihe districts of Somkheto-Dzhavakheti Region of Georgia at Dzhavakheti Highland (Mt. Madatapa) and Erusheti Mountains (Mt. Gumbati, Mt. Airilanbashi) (Tuniyev et al., 2014). Established guidance on finding the Darevsky’s viper in Turkey at a considerable distance from the known habitat were of great interest. Our research in Turkey showed that vicinity of Posof inhabits a distinct species – Pelias olguni (Tuniyev S. et al., 2012). Endemic subspecies live in humid (Pelias darevskii uzumorum) and semidry (Pelias darevskii kumlutasi) segments of Yalnizçam Dağlari (Arsiyansky) Ridge. The taxonomic status of other finds is under discussion (Tuniyev, 2016).

The density of populations of Pelias darevskii in Armenia is extremely low, as the number of vipers in the identified locality in Georgia. The main threats are the killing of animals, overgrazing, trapping for the illegal zoo trade. The expansion of Darevsky’s viper distribution range cannot substantially influence on its conservation status. The Darevsky’s viper still remains narrow distributed sporadically spread species, not forming high-dense populations. Category rarities on the global population estimated as CR B2ab (ii, iii). Species is listed in the IUCN Red List of Threatened species (2024) and Red Data Book of the Republic of Armenia (2010) (Table 5).

Species effectively guarded exclusively in Arpilich National Park (Armenia). It is necessary to include it to recently creating Dzhavakheti National Park in Georgia area on the Dzhavakheti Highland and Erusheti Mountains and optimization borders transboundary with Arpilich National Park Mt. Madatapa in Georgia, with the participation of Darevsky’s viper biotopes (Tuniyev, 2016).

Dinnik’s Viper – Pelias dinniki. It is endemic to the Highlands of the Great Caucasus, sporadically distributed within the Russian Federation, republics of Abkhazia, Georgia and South Ossetia (Tuniyev et al., 2009). In Russian Federation, it is known from the Krasnodar Territory and republics of Adygei, Karachay-Cherkessia, Chechen and Dagestan (Tuniyev, Tuniyev, 2008; Red Data Book of Ingushetia Republic, 2007). During the last decade significantly expanded understanding of the range of the species. We have found isolated populations on Mt. Zhitnaya of Lagonaksky Ridge, Adygei (north-western limit), the tops of Hakudzh, Lysaya of Chernomorsky Chain, Krasnodar region (western limit) (Tuniyev, Tuniyev, 2007, 2013); in vicinity of pass Genukh and upper basin of Dzhurmut River, Dagestan (eastern limit). In addition, the new localities we have discovered on Aibga Ridge (Krasnodar Territory), Kutakheku Ridge (Abkhazia), upper basin of Bolshaya Liakhvi River (South Ossetia), etc. There are no reliable findings in the territory of the republics of Kabardino-Balkaria and Ingushetia yet, but it is likely theylivethere too (Tuniyev, 2016). Probably represents a complex of cryptic forms in the east of range.

It is listed in the IUCN Red List of Threatened species (2024), the Red Books of the Russian Federation (2021), Krasnodar Territory (2017), the republics of Adygei (2021), Karachay-Cherkessia (2013), Chechen (2020), Dagestan (2020), Republic of North Ossetia-Alania (2022), Kabardino-Balkaria (2018) and Republic of South Ossetia (2017) (Table 5). In Chechen Republic and Dagestan species is extremely rare, sporadically distributed on most mesophylous parts of subalpine belt. The main threats for western populations is a recreational development, construction of ski facilities on the territory of Sochi National Park, the planned mastering of the Caucasian reserve, development of resorts in the North Caucasus on existing protected territories; for eastern populations is overgrazing and absence of high rank protected areas (reserves and national parks). Category rarities on the global population estimated as VU B1ab (iii, v), in a new edition of the Red Data Book of the Russian Federation (2021) mind category VU C1 + 2a.

To save the polymorphic populations of species, it is necessary to abandon ambitious plans for transformation of natural landscapes in the Highlands of the Sochi National Park (Aibga Ridge) and in Caucasian reserve (basins of the upper flowing of rivers Mzymta and Malaya Laba); create a new Federal protected areas in the Itum-Kalinskaya Hollow of Chanty-Argun River (based on former Soviet sanctuary) in Chechen Republic and Dagestan reserve plots clustered Tokhor River, Mt. Guton and vicinity of Genukh pass (Tuniyev, 2016).

Armenian Steppe Viper – Pelias eriwanensis. Disjunctive distributed on the southern slopes of the Lesser Caucasus and the Armenian Highland endemic relict species. The species range includes Armenia, Azerbaijan, Turkey and Northwestern Iran (Rajabizadeh et al., 2011). Recently, the species has been found on the shores of Lake Kartsakhi in South Georgia (Tuniyev S. et al., 2014) and on Bargushat Ridge in Armenia (Tuniyev, 2016).

Isolated populations of Pelias eriwanensis inhabit rocky mountain meadow-steppe, juniper bushes on the slopes of quite volcanoes and arid ridges (Urc, Karabakh, etc.), rarely rise down along the canyons of the rivers to the middle-mountain belts, where the steppe outliers in the canyons (Kasakh River, Armenia). Sporadic animals, rarely up to 5 specimens at 1 km of the route. The main threats are the killing of animals and destruction of their habitats, overgrazing both in Transcaucasian Republics and in Turkey. Category rarities on the global population estimated as VU B1ab (iii, v). Species is listed in the Red Book of Armenia (2010) (Table 5). It is necessary to list species in Red Books of Republics Azerbaijan and Georgia. .

Erevan mountain-steppe viper currently protected in Armenia on Khosrov reserve territory, formally defined in Sevan National Park and Zangezur Sanctuary. Species habitats on Urc (former Sarajbulag) Ridge has dropped out from under protection in connection with the deletion of this site from the Khosrov reserve. To save the species requires the creation of transboundary protected area around Kartsakhi Lake in South Georgia and Turkey, creating local reserves on Coast of Childyr Lake in Turkey.

Kaznakov’s Viper – Pelias kaznakovi. The oppressed relict endemic to Colchis with slightly irradiation, sporadically distributed in foothill areas of Krasnodar Territory, Republics of Abkhazia, Adygei, in western Georgia and Adzharia within the former Soviet Union and on the Lazistan coast of Turkey (Orlov, Tuniev, 1986). In the Russian Federation, area covers foothill areas along the Black Sea coast from vicinity of Tuapse to the border with Abkhazia, on the northern slope of the Western Caucasus from the town Goryachy Kluch to the foothills of Apsheronsky District of Krasnodar Territory and Tulsky District of Adygei Republic. It is found in a number of new localities in Turkey (Afsar, Afsar, 2009; Tuniyev et al., 2019).

Pelias kaznakovi has never been numerous and now progressively disappears throughout the area (Ostrovskikh, 1991). Currently, most biotypes, or destroyed or heavily transformed in connection with the construction of Olympic venues and involving them in the infrastructure of the Mzymta – Psou Rivers area. Apparently, the total number of species in the Russian Federation is not more than 2000 individuals.

The main threats are the killing of animals and destruction of their habitats, intensive trapping for the illegal zoo trade, recreational mastering the Black Sea coast of the Caucasus, to change the zoning of the Sochi National Park, with the transfer of plots of vipers and the protected habitat of protected zones in the lower category of zoning. Category rarities on the global population estimated as EN B2ab (ii, iii, v), in a new edition of the Red Data Book of the Russian Federation (2021) mind category EN A4abcd; B2ab (i, ii, iii, iv, v). Pelias kaznakovi is listed in the IUCN Red List of Threatened species (2024), was listed the Red Book of the USSR (1984), the Red Data Book of Georgian SSR of the former Georgian Soviet Socialist Republic (1982). It is included into the Red List of Georgia (2024), Red Books of Russian Federation (2021), Krasnodar Territory (2017), Republic of Adygei (2021) (Table 5).

The bulk of the range in Russia remains on the territory of Sochi National Park, small populations are found in the Caucasian reserve. Formally, species is protected in Ritza relic National Park, Pskhu-Gumista reserve (Abkhazia), in the Kintrish reserve and Mtirala National Park (Georgia), Kamili National Park (Turkey).

To save Pelias kaznakovi in Sochi National Park it should be included in the functional zoning in especially protected or reserved zone. Important measure is breeding in captivity with subsequent reintroduction (Tuniyev, 2016).

Lotiev’s Viper – Pelias lotievi. Endemic species probably represents a complex of cryptic forms. The total area covers mainly middle mountain area of Great Caucasus within Russia, with little penetration in Georgia and Azerbaijan. In the Russian Federation, this snake was found in semi-arid hollows in Eastern and Central Caucasus, where it inhabits shibliaks, frigana, mountain-steppe areas from 1200 to 1800 m a. s. l. In the Krasnodar Territory found in the midlands band at an altitude of 1500–1700 m a. s. l., where Pelias lotievi inhabits xerophylous subalpine meadows, scrub and rocky associations and talus slopes. For species description material was available only from 8 localities; but later after the numerous expeditions, covering the whole of the Northern Caucasus the detailed distribution data (about 70 localities) were collected. In the East, the range of Pelias lotievi reaches coastal Dagestan in vicinity of settlement Izerbash (Kolichi River), covering almost the entire foothill and mountainous Dagestan, except in the highland sections of the Main Ridge. To the west, Pelias lotievi inhabits semiarid hollows of Chechen Republic, Ingushetia, North Ossetia-Alania, xerophylous slopes of Mt. Elbrus and Skalisty (Rocky) Ridge in Kabardino-Balkaria. Further west it is a chain of isolates in hemixerophylous landscapes of Bokovoy (Forward) and Skalisty (Rocky) ridges in the Karachay-Cherkessia Republic up to the extreme western finds at the watershed of Bolshaya Laba and Malaya Laba Rivers in the Krasnodar Territory.

The main threats are overgrazing, reducing the protective properties of biotopes, direct destruction of man, and weak coverage of the habitats of the species by existing network of high rank protected areas, i. g. reserves and national parks. Category rarities on the global population estimated as NT, in relation to the total area of the species, not exceeding 20000 km². Pelias lotievi is listed in the Red Books of Krasnodar Territory (2017), the Chechen (2020), North Ossetia-Alania (2022), Kabardino-Balkaria (2018) and Karachay-Cherkessia Republics (2013) (Table 5). An isolated population is protected in the territory of the Caucasian reserve; part of the range is included in Teberdinsky, Prielbrusye, Alania National Parks, Erzi, Kabardino-Balkarsky Vysokogorny and Severo-Ossetinsky reserves. Formally preserved on the territory of Charodinsky sanctuary in Dagestan. It is necessary to create new protected areas in Sadono-Unal′skaya Hollow of North-Ossetia-Alania, Itum-Kalinskaya Hollow of Chechen Republic, and in several places of inner-mountain Dagestan (Tuniyev, 2016).

Relic Viper – Pelias magnifica. The oppressed endemic relict species, has a limited amount of habitat, conservation depends even on short anthropogenic influences. The entire area is located in the Russian Federation and covers Skalysty (Rocky) Ridge within Krasnodar Territory, Republic of Adygei, possibly being in the Karachay-Cherkessia Republic (Tuniyev, Ostrovskikh, 2006). Species range covers so-called Belo-Labinsky Refugio of Colchis biota (Tuniyev, 1990) and represents a complex option derivates of Colchis vegetation enriched with elements of Mediterranean and even the steppe vegetation. Habitats represented by light oak groves, dry stretches of grassy meadows and shrubs on rocky limestone cornices of arrays in the range of heights from 700 to 1000 m a. s. l. (Tuniyev, 2016; Tuniyev et al., 2016).

Population’s density is extremely low for a day excursion numbered no more than three specimens. The main threats are naturally archaic area, which constantly highlights the grassroots fires, increasing recreational press, trapping animals for illegal zoo trade.

Pelias magnifica included in the IUCN Red List of Threatened species (2024) category EN A1cd + 2 CD, Red Data Book of the Russian Federation (2021) with category EN A2abc; B2ac(iv), Red Book of Krasnodar Territory (2017) with category EN A1cd+2cd and Red Book of Adygei Republic (2021) with category EN B1ab(i, ii, iii, v); C2a(i) (Table 5). Thus, a rapid assessment of the year 2014 allowed to state the existence of stable small, exposed to anthropogenic limiting factors population of Relic Viper in the type locality and agree to the terms in the IUCN Red List of Threatened species (2024) category of threats the existence of species within Krasnodar Territory – EN A1cd+2cd (Tuniev S. et al., 2016). Insignificant part of the population is protected in the Caucasian reserve, to which it is necessary to include the Maly Bambak Ridge.

Orlov’s Viper – Pelias orlovi. Endangered relict endemic species with number declining progressively. The entire area is located in the Krasnodar Territory of the Russian Federation and covers both slopes of the lowest Northwestern part of the Greater Caucasus from Markhot Ridge in the West to the peak Bolshoy Pseushkho in the East. Distribution area lies entirely within the area of influence of the Mediterranean climate and development of xero-mesophyt habitats. Species occurs in different versions of submediterranean landscapes: from intrazonal riparian glades to steppe-meadows and ecotones of Juniper forests. Range of high-altitude distribution ranges from 450 to 950 m a. s. l. (Tuniyev, 2016).

The number of species is very low, at the end of the last century for a day excursion numbered no more than 3 specimens. Express score 2014–2016 years testified to the disappearance of a species in a number of localities (northern foothill of Mt. Papay, vicinity of Gelendzhik City, etc.) and a catastrophic populations’ density decreasing in the surviving habitats. The main threat was and it remains illegal trapping of animals for zoo trade. Category rarities on the global population estimated as CR B1ab (i, v); C2A(i), in a new edition of the Red Data Book of the Russian Federation (2021) has category EN A2abcd. It is listed in the IUCN Red List of Threatened species (2024) and Red Book of Krasnodar Territory (2017).

This species formally defined at recent organizing regional Markhot Nature Park. It is necessary to organize cluster plot of Utrish reserve at Mt. Papay, or creation of a full-fledged Novorossiysk reserve. It is in accordance with the earlier recommendations as well as implementing the decision of UNEP within the framework of the compensatory actions Sochi-2014 Winter Olympics to include in the composition of the Sochi National Park Mt. Bolshoy Pseushkho (Tuniyev, Timukhin, 2013).

Eastern Steppe Viper – Pelias renardi. Species is reduced and continuing to reduce population size and habitat caused by human activities. The total area covers the steppe and semi-desert area of the South-Eastern European countries, Kazakhstan and Central Asia. In the Russian Federation, it is found from the Volga-Kama region north to the Precaucasia region South and Central Altai Mountains in the East (Ananjeva et al., 2006). In the Caucasus ecoregion, area covers the lowland and the foothills of the Northern Caucasus. According to our observations, in the Central Caucasus Pelias renardi maximum penetrates mountain country.

Found on the plains of various types (loess, alluvial-loess, terraced), on hills, in lower-mountain belts. It inhabits forest edge, shrub associations, shibliaks, steppe slopes (Ostrovskikh, Plotnikov, 2003). In the Southeast of the range rises up in Krasnodar Territory to 1000 m a. s. l., is able to live on the sandy sea spits. In conditions of anthropogenic landscape forms narrow stripe-like settlements in badlands and bargain sites, tree planting, etc. (Ostrovskikh, Plotnikov, 2003а). In the Central Caucasus, in the area of the Elbrus gap forest vegetation, Pelias renardi together with other steppe species of plants and animals penetrate mountains up to 1500 m a. s. l. (Tuniyev, 2016).

At the end of the last century, the density in the Caucasus had up to 30 animals per 1 hectare. Currently the number of habitats and the number of snakes they declined sharply. Major threats remain steppe and forest-steppe landscape transformation and physical extermination by man. Category rarities on the global population estimated as VU A1c+2c, in a new edition of the Red Data Book of the Russian Federation (2021) has category VU A4cd; B1ab (i, ii, iii, iv). Pelias renardi is listed in the IUCN Red List of Threatened species (2024), in the Caucasus ecoregion – in the Red Books of Stavropol (2013) and Krasnodar (2017) territories, republics of Dagestan (2020), Chechen (2020), Karachay-Cherkessia (2013), Adygei (2021), North Ossetia-Alania (2022), Kabardino-Balkaria (2018).

In the Caucasian part of the distributional area species it is not protected, a tiny population exists on the northern periphery of the reserve Utrish and Kislovodsk National Park. The cluster is required to organize sections of the existing reserves in areas of dense habitation vipers, an extension of the recently established Kislovodsk National Park, with including of Pelias renardi’s biotopes in the Region of Caucasian Mineral Waters (Tuniyev, 2016).

Shemakha Viper – Pelias shemakhensis. Narrow-endemic, oppressed relic species, whose area covers the Shamakhi District of Azerbaijan. Until recently, no records was known from the northeastern steppe regions of Eastern Georgia. In 2016, it was rediscovered here (Tuniyev et al., 2018). Alekperov (1978) wrote that in vicinity of Shemakha a viper occurs on the altitude of a 700 m a. s. l. on the stony areas of mountains, covering by a xerophylous grasses and shrub vegetation, also in the small agricultural sowing, in the heaps of stone on the plough-lands and along the roads. According to our observations, the biotopes of species (in both Azerbaijan and Georgia) are presented by shibliaks of Paliurus spina-christy and ecotones of dry Andropogon steppes and derivates of the broad-leaved forests on the stony slopes of foothills. Alekperov (1978) recorded the numerous specimens originated from Shemakha: only for feeding study, 22 specimens were dissected. At present, the species became very rare in Northwest Azerbaijan; in Northeast Georgia is on the verge of extinction. As a rare taxa, limited in range and abundance, Pelias shemakhensis named as Pelias renardi is included in the Red Book of Azerbaijan (2013). Pelias shemakhensis should be included in the IUCN Red List of Threatened species (2024) and Red Book of Georgia with the status category CR B1ab (i, ii, iii, iv, v).

To save the species it is necessary to organize protected areas in vicinity of Shemakha in Azerbaijan and cluster plots of Vashlovani reserve on the Shirak Plateau in Georgia.

Tuniyev’s Viper – Pelias tuniyevi. Endangered narrow distributed relict endemic species with declining progressively number. All known finds are located in the left-bank basin of the middle flow of the Kura River from the Borzhom Gorge (east slope of Meskheti Ridge) in Georgia to middle-mountain districts of South Ossetia (east foothills of Likhsky Ridge and south spurs of Central Caucasus) (Ananjeva et al., 2021). It inhabits mid-altitude broad-leaved forests, Carpinus orientalis shibliak and post-forest glades in the range of elevations from 700–850 m in the Borzhom Gorge of Georgia to more than 1300 m a. s. l. in the ruins of Atsriskhevi of South- Ossetian Reserve. Currently, biotopes of P. tuniyevi are characterized by more xero-mesophilic traits and moderately warm mezo-climate in comparison with typical mesophilic biotopes of North-Colchian (Krasnodar Territory – Abkhazia) and, especially, Adzharo-Lazistanian for P. kaznakovi, living in the humid subtropical climate.

Pelias tuniyevi is listed in the Red Books of Republic of South Ossetia (2017) (named P. kaznakovi) and partly under name P. kaznakovi in Red Data Book of Georgian SSR (1982). Small parts of area protected in South-Ossetian Reserve and Borjomi-Kharagauli National Park.

As can be seen from the foregoing material, relatively positive protection present with such endemics as Pelias darevskii, P. dinniki, P. lotievi, a significant part of the habitats of these vipers is covered by the territories of the nature reserves and national parks. At the same time, it is not a case for Pelias orlovi, P. shemakhensis; Pelias kaznakovi, P. magnifica, P. ebneri, P. eriwanensis and P. tuniyevi which are poorly represented within the protected areas. Moreover, Pelias orlovi and P. magnifica habitats were not even included in the priority regions of the CEPF.

We provide the recommendations on the organization of new protected areas, or the expansion of existing protected areas within the Caucasian Isthmus. Analysis of biological diversity and spatio-altitudinal characteristics of protected natural territories and species in the Caucasus showed that they not fully ensure the main goals for the protection and rational use of natural resources, including long-term preservation of shield-head vipers. In some cases, international cooperation is required to create protected areas. They are East-Caucasian Transboundary Territory, uniting Mt. Guton, Tlyaratinsky Sanctuary, vicinity of Genukh pass, Lagodekhi and Zakataly State Reserves, Shakhdag National Park (Russia, Azerbaijan, Georgia); North Colchis Transboundary Territory, uniting Caucasian, Pskhu-Gumista, Pitsunda-Myussera Reserves, Sochi and Ritza National Parks (Abkhazia, Russia) and Ashotsk-Javakheti Transboundary Territory, uniting the Arpilich National Park, Javakheti National Park and Erusheti Mountains (Armenia, Georgia, Turkey).

Of course, this is not a complete list of protected areas which should be considered for the transboundary cooperation between Georgia and Turkey, between Armenia, Iran and Azerbaijan.This important goal should be the subject of a separate discussion. The implementation of this programme, along with suggested above other protected areas for the protection of all endemic and endangered species, will come as close to a sustainable long-term reliable completeness of viperidae snakes biodiversity conservation (as an umbrella-species) in the Caucasian ecoregion.

ACKNOWLEDGEMENTS

The authors would like to thank Nikolai Orlov and Igor Doronin for valuable comments an earlier version of the manuscript and all the colleagues and friends who shared field work in the Caucasus with us. We are grateful to Alexey Egoshin, who performed cluster analysis of our data, Sofia Tuniyeva and Mikhail Kravchenko for processing photographs and drawings. We are sincerely grateful to anonymous reviewers for their kind help and useful comments, which helped us to improve the previous version of the manuscript. Thanks a lot for Ted Papenfiss for his kind help with English translation.

FUNDING

This study wascompleted with partial financial support from under the state theme 122031100282-2. No additional grants to carry out or direct this particular research were obtained.

ETHICS APPROVAL AND CONSENT TO PARTICIPATE

The present work is based on the literature sources and own long-term study of the Caucasian ecoregion whoch does not contain any sources on human, or animals that meet the criteria of Directive 2010/63/EU.

CONFLICT OF INTEREST

The authors declare no conflicts of interest.

Об авторах

Б. С. Туниев

Автор, ответственный за переписку.
Email: btuniyev@mail.ru
Россия, Московская ул., 21, Сочи, 354000

Н. Б. Ананьева

Email: Natalia.Ananjeva@zin.ru
Россия, Университетская наб., 1, Санкт-Петербург, 199034

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1. JATS XML

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2. Рис. 1. Карта Кавказского экорегиона.

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3. Рис. 2. Регионы природоохранного приоритета и их коридоры в Кавказском экорегионе с нашими изменениями, выделены красным цветом (из Туниева и др., 2019).

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4. Рис. 3. Эндемичные хвостатые амфибии: 1 – Lissotriton lantzi, 2 – Paradactylodon persicus, 3 – Mertensiella caucasica, 4 – Mertensiella djanaschvilii.

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5. Рис. 4. Распространение Ommatotriton ophtyticus.

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6. Рис. 5. Эндемичные жабы Кавказского экорегиона: 1а – Bufo verrucosissimus circassicus, 1б – B. v. turowi, 1в – B. v. tertyshnikovi, 1г – B. v. verrucosissimus, 2 – B. eichwaldi.

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7. Рис. 6. Распространение Pelodytes caucasicus.

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8. Инжир. 7. Эндемичные ужеобразные Кавказского экорегиона: 1 – Natrix megalocephala, 2 – Rhynchocalamus satunini, 3 – Zamenis persicus.

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9. Инжир. 8. Эндемичные гадюки комплекса Kaznakovi Кавказского экорегиона: 1 – Pelias orlovi, 2 – P. magnifica, 3 – P. dinniki, 4 – P. kaznakovi, 5 – P pontica, 6 – P. olguni, 7 – P. tuniyevi, 8а – P. darevskii darevskii, 8б – P. d. роскошный, 8в – П. д. узуморум.

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10. Инжир. 9. Эндемичные гадюки комплекса ursinii-renardi Кавказского экорегиона: 1 – Pelias lotievi, 2а – P. shemakhensis kakhetiensis, 2б – P. sh. шемахензис, 3 – P. eriwanensis.

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11. Рис. 10. Сходство батрахофауны стран Кавказского экорегиона по коэффициенту Жаккара.

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12. Рис. 11. Сходство батрахофауны стран Кавказского экорегиона по коэффициенту Серенсена-Чекановского.

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13. Рис. 12. Сходство герпетофауны стран Кавказского экорегиона по коэффициенту Жаккара.

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14. Рис. 13. Сходство герпетофауны стран Кавказского экорегиона по коэффициенту Серенсена-Чекановского.

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15. Рис. 14. Представленность эндемичных видов в странах Кавказского экорегиона (см. главу Эндемичные виды: богатство, статус сохранения и угрозы и таблицу 5).

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