April 1990 Asiatic Herpetological Research Vol. 3, pp. 67-84

On the Independence of the Colchis Center of Amphibian and Reptile Speciation

BORIS S. TUNIYEV 1

^Causasian State Biosphere Reserve, Sochi, USSR

Abstract. -The Colchis region of Western Transcaucasia is characterized by a rather uniform thermal regimen, corresponding to a subtropical climate. The Colchis forests contain an extraordinary abundance and diversity of tree, shrub, and vine species. The herpetofauna of the Colchis forests is surprisingly poor, despite its uniqueness.

Key Words: Amphibia, Reptilia, USSR, Caucasus, biogeography.

Introduction Often they are inhabited by the same species. The question is whether this fauna The herpetofauna of Western has appeared from the west or is it the Transcaucasia is not homogeneous, due to remainder of the fauna that has populated the different age and genesis of the species densely the shores of the Black Sea at one distributions. Along with autochtonous time. It is impossible to answer these and endemic forms, one can find species questions at the present level of our whose main areas of distribution are in the knowledge. But even now I can definitely European part of the USSR and in the say that this fauna by its origin, has nothing Eastern Mediterranean. At the same time, a in common with the faunas in other regions number of species which have main of the Caucasus." distributional centers in the Colchis occur beyond the bounds of Western In 1912, Satunin divided the Caucasian Transcaucasia, in other parts of the Isthmus into five subregions and 11 Caucasian Isthmus. For these reasons, it is districts, including the Colchis in the West- necessary to define the Colchis Transcaucasian district of the Littoral herpetofauna and to determine its place in subregion. the fauna of reptiles and amphibians of the Caucasian Isthmus as a whole. Among other merits of this work by Satunin, one cannot but mention the fact Research on this issue started with the that for the first time, he defined in an exact works of Nordmann (1840), Derjugin way the Colchis region proper. He defined (1899), Silantyev (1903), Brauner (1905), the northern border as the spurs of the Main and Nesterov (1911). However, the first Caucasian Range up to the basin of the well-grounded definition of the fauna in Tuapse River, the southern border as the question from a zoogeographical point of Pontic Range, and the eastern border as the view was presented in the works of Arsijanskij Range. The valley of the Rioni Satunin (1912). Satunin wrote in 1910, River and the adjacent southern slopes of "So far I cannot say much about the genesis the Main Range were defined as the central of the fauna of this region called Western part of the region. Satunin emphasized the Transcaucasia. This country with its depauperate herpetofauna of this region on evergreen plants and scanty fauna one hand, and the presence of endemic resembles a piece of the Mediterranean in species such as Vipera kaznakowi and the narrow sense of the word. True, here Bufo verrucosissimus on the other hand. are endemic species and forms, but not a single genus of vertebrate is unrepresented Nikolsky (1911) assigned the entire in the countries of the Mediterranean. Caucasus, excluding eastern Precaucasia,

© 1990 by Asiatic Herpetological Research Vol. 3, p. 68 Asiatic Herpetological Research April 1990

to the Mediterranean. However, he could Colchis, however, was not distinguished as not differentiate the forest and the alpine an independent center of speciation in this belts of the Greater Caucasus, because of work. the absence of data. Scherbak (1981) included the Colchis in Results and Discussion the Caucasian Region of the Mediterranean Province. He suggested that the typical Investigations of the last decades made it species of the region were Mertensiella possible to add the majority of the species caucasica, Pelodytes caucasicus, Lacerta of the Colchis herpetofauna to an overall saxicola-complex and others. However, picture of Colchis faunal distributions the Colchis proper was again not (Turov 1928; Bartenev and Reznikova distinguished as an independent center of 1935; Khozatsky 1941; Milyanovskiy herpetofaunal formation. 1957; Bannikov et al. 1977; Negmedzyanov and Bakradze 1977; Orlova For the analysis of the herpetofauna of 1973, 1978a, 1978b; Golubev 1980, 1985; the Colchis proper, it is necessary to Tuniyev 1983, 1985). In addition there has exactly define the term "the Colchis been a revision of the taxonomic status of phytolandscapes", and to decide what types such forms as Vipera kaznakowi of vegetation are universally recognized as (Vedmederja et al. 1986; Orlov and "Colchis types". Albov (1885) was the Tuniyev 1986a, 1990 this volume), Lacerta first to clearly depict plant landscapes of agilis (Peters 1960), L. derjugini the Colchis. He singled out a region, (Bartenev and Reznikova 1931; Orlova unique for Russia, of mountain limestone

1978a ; Bischoff 1982, 1984), L. saxicola flora which had been developing mainly (Darevsky 1967; Darevsky and Vedmederja autochthonously in a large refugium with 1977), Anguis fragilis colchicus (Lukina numerous endemic and relict species and 1965; Scherbak and Scherban 1980), and even genera. Kolakovskiy (1980) regarded others. the Colchis flora as basically forest and alpine-meadow, and suggested that its main Accumulation of this information along phytolandscapes had existed since old times with works on fossil amphibians and with changes only in the composition of reptiles of the Caucasus (Vekua et al. 1979; their edificators, except for the extinct Chkhikvadze 1981, 1983, 1984; Bakradze formation of evergreen subtropical forests and Chkhikvadze 1977; Zerova and in the lower mountain belt. The tertiary- Chkhikvadze 1984; Yefimov and relict character of the forest mesophile flora Chkhikvadze 1987) have made it possible and vegetation is fully revealed here due to to revise the zoogeography of the region. slight changes in this region's climatic conditions (Kuznetsov 1891). The most Darevsky (1957) singled out seven characteristic features of the tertiary-relict different groups of species and subspecies Colchis forest are: extraordinary of the herpetofauna in the Caucasus, based abundance and diversity of tree and shrub on their origin. Among the species species, impossibility of singling out the representatives of the region of interest to dominant species (which is also us, it is necessary to pay attention to characteristic of tropical forest with extreme Lacerta strigata (Asia Minor species), density of trees), abundance of vines and Emys orbicularis, Anguis fragilis, epiphytes, and almost total absence of grass Coronella austriaca, Elaphe quatuorlineates cover. All these attributes make the Colchis sauromates, Natrix natrix (European boreal forest similar in many aspects to a tropical species), Testudo graeca, Natrix tessellata rain forest (Pavlov 1984). According to (Mediterranean species), Pseudopus Sinskaya (1933), the Colchis forest apodus, Coluber najadum (east- vegetation underwent three main stages of Mediterranean species), and Lacerta development: the tropical forest; the forest saxicola, L. praticola, L. derjugini, L. of Colchis type, but rich and covering a media (autochthonous species). The wider area; and last, the modern Colchis April 1990 Asiatic Herpetological Research Vol. 3, p. 69

forest. grusinica, Natrix megalocephala, and Vipera kaznakowi are Colchis endemics in The Colchis type of vegetation includes a the broad sense of the word. number of phytocenoses differing in structure, composition, and ecological In addition to the Colchis endemics, peculiarities: it may be mixed there are three more ecological-geographical (polydominant), or may be presented by groups of amphibians and reptiles in the cenosis of one or two species, but the region. They have similar ecological common and obligatory attribute of characteristics (habitat first of all), and phytocenosis of the Colchis type is an overlapping geographic distributions. abundance of tertiary relicts. The area with Colchis type vegetation is characterized by 1. The East-Mediterranean group a rather monotonous thermal regimen, consists of Triturus cristatus karelini, to a corresponding subtropical climate, but Testudo graeca nikolskii (Fig. 1 ), Lacerta with highly diverse soils (Gulisashvili et al. media, L. praticola pontica, L. strigata, 1975). Pseudopus apodus tracius, Natrix tessellata, and Coluber najadum. This The herpetofauna of the Colchis forests group's distribution includes either the is surprisingly poor, despite its uniqueness. Balkans and the Caucasus or the Balkans, The species composition is different in the Crimea, and the Caucasus. According to southeastern and northwestern parts of the ecological characteristics, these are Colchis compared to the other portions of xeromesophiles or hemixerophiles whose its territory. Species such as Mertensiella spreading is related to dry foothills of caucasica, Lacerta clarkorum, L. parvula, Western Transcaucasia up to 200-300 m and L. mixta, whose distributions are above sea level with an annual sum of connected with forests growing on acid temperatures exceeding 5000°C. Thus, soils above volcanic rocks, are found on Testudo graeca, Pseudopus apodus, and the western slopes of the Adzharo- Coluber najadum occur in the Colchis on a Imeretinsky, Shavshetsky and Lazistansky narrow seaside strip of land with enclaves (Pontic) mountain ranges. Similarly, of Mediterranean vegetation from Tuapse to floristic endemics of this part of the Colchis Pitsunda-Sukhumi. A local population of {Rhododendron ungernii, Osmanthus L. strigata occurs in the Pitsunda region, decorus, Betula medwedewii, Epigaea and a local population of L. media occurs gaultheriodes, and others), are Adzharo- in the environs of Pitsunda and Salme. The Lazistan endemics, sometimes with slight majority of localities of L. praticola and radiations to the adjoining regions, but are Natrix tessellata in the Colchis are in the not Colchis endemics in the broad sense of seaside hills up to 400 m above sea level. the word. The same applies to Lacerta It is only along the valleys of large rivers saxicola darevskii and L. saxicola brauneri like the Shakhe River, the Mzymta River, which are in the widespread northwestern the Bzyb River, and others that N . part of the Colchis, but are absent in the tessellata penetrates into the Colchis up to central and southeastern Colchis. These 600 m above sea level. Thus the majority animals, by analogy with floristic endemics of these species are found either in places {Allium candolleanum, Campanula with vegetation of the Mediterranean type, mirabilis, C. bzybica, C. calcarea, C. or in places where the initial Colchis jadvigae, Genista abchasica, Gentiana vegetation has been reduced to zero and the paradoxa, Omphalodes kusnetzovii, and landscapes resemble the Mediterranean others) are northern-Colchis endemics. For ones by their thermo-biotopic conditions example, of the 450 endemic Colchis (substitute shibliaks and tomillares, species of flora, 83 (25%) are endemics of Pitsunda pine groves, foothill post-forest the northern Colchis (Adzinba 1980). glades, and landplots with Erica tetraliz, Triturus vittatus ophryticus, T. vulgaris Juniperus oxicedrus, Pinus pityusa, and lantzi, Bufo verrucosissimus, Pelodytes Arbutus andrachne). caucasicus, Lacerta derjugini, L. agilis Vol. 3, p. 70 Asiatic Herpetological Research April 1990

FIG. 1. Testudo graeca nikolskii is a Mediterranean species, and in the Colchis it is found only in the seaside strip of land with enclaves of Mediterranean vegetation.

2. The Caucasian group includes Hyla The composition of this group is not arborea schelkownikowi (Fig. 2), Rana homogeneous. It includes both species macrocnemis, Lacerta caucasica alpina, L. typical for the steppe areas {Bufo viridis rudis, and Vipera dinniki. Distributions of and Coluber jugularis ) and those that are these species in the Caucasian Isthmus are widely distributed in Europe (all the rest). broader than those of the Colchis group. Only Anguis fragilis and Coronella At the same time, the majority of them are austriaca are widely distributed in the Colchis autochthons. These species are Colchis. This makes it difficult to mesophiles and occur in mesophillous definitely consider them late migrants to the forest and mountain meadow formations. Caucasus. Other species either occur in This group seems to be of Colchis origin, several spots along the Colchis seashore retaining close connections with the main (B. viridis and Natrix natrix ) or populate a center of the formation. Broader ecological narrow strip of land along the sea together tolerance in comparison with typical with the Mediterranean species (C. Colchis species makes it impossible to jugularis ) or a somewhat wider strip include them within the Colchis group. (Emys orbicularis and Rana ridibunda ). Despite the possibility of finding these 3. The European group consists of Bufo species in the typical Colchis forest viridis, Rana ridibunda, Emys orbicularis, formations, the majority of them are still Anguis fragilis, Natrix natrix, Coronella attributed to the Mediterranean type of austriaca, and Coluberjugularis caspius. vegetation. Vol. 71 April 1990 Asiatic Herpetological Research 3, p.

FIG. 2. Hyla arborea schelkownikowi seems to be of a Colchis origin, but because of its broad ecological tolerance, it is found in a large part of the Caucasian Isthmus.

Let us consider the dispersal and Triturus vulgaris lantzi (Fig. 3) occurs distribution of the representatives of the in the same places in the Colchis as T. Colchis group in detail. Triturus vittatus vittatus ophryticus does. Very often both occurs in the territory from the seashore to species are symbiotopic (Tuniyev and the subalpine meadows in all the forest Beregovaya 1986). On the northern slope types. In the place called "the Colchis of the Western Caucasus, its home range is Gates" (lowering of the Main Caucasian wider than that of the previous species, but Range between Mt. Fisht and Mt. it is only through the mesophillous forests Chugush) the species crosses over to the and subalpine meadows that it penetrates northern slope of the Western Calucasus, into the Eastern Transcaucasia up to the reaching the environs of Goriachij Klutch Trialet Ridge. The isolated population in and Krasnodar in the northwest and the Talysh occurs in the Hirkan forests which basin of the Laba River in the northeast. In are ecologically and genetically close to the the eastern part of the area, the species Colchis forests. crosses the Adzharo-Imeretinskij mountain range and reaches the outskirts of Tbilissi- Bufo verrucosissimus (Fig. 4) occurs in Oni. It occurs in the Lagodekhi region as all parts of the Colchis from the sea shore an isolate. Outside the boundaries of the up to the subalpine forests. On the Colchis this species occurs either in the northern slope of the Western Caucasus, it Colchis type forests or in their derivatives. is found on the territory from the environs Vol. 3, p. 72 Asiatic Herpetological Research April 1990

FIG. 3. Triturus vulgaris lantzi. This subspecies is widespread throughout the Colchis.

of Krasnodar in the west to the Psebaj southeastern distributional limits also settlement and the Shakhgirej Canyon in the coincide with that of Bufo verrucosissimus. east, where it is found in the derivatives of Pelodytes caucasicus does not occur east the Colchis forests from 400 m to 1000 m of the Trialet Ridge. There is an isolated above sea level. In the Eastern Caucasus it population in the mesophilous forests in the is found in the Borzhomy Gorge, the Lagodekhi-Zakataly region. Lagodekhi-Zakataly, and the Talysh region, where its distribution is limited to the Lacerta derjugini has a distribution in mesophilous forests, which are abundant in the Colchis similar to P. caucasicus. It the Colchis and Hirkan floral elements. reaches the sub-alpine belt. On the northern slope of the Western Caucasus, it The distribution of Pelodytes caucasicus is found in the Colchis forest derivatives (Fig. 5) in the Colchis is more restricted. It from the Belaja River to the Small Laba does not occur in the coastal belt and oak- River (the Shakhgirej Gorge). The species forests. It is found both in the mesophilous penetrates through the Eastern beech, chestnut, and fir-tree forests, and in Transcaucasia up to the Trialet Ridge. mixed broad-leaved forests with an Separate populations occur in north-eastern evergreen understory. On the northern Georgia up to Lagodekhi-Zakataly. slope of the Western Caucasus, its distribution coincides with that of B. Lacerta agilis grusinica (Fig. 6) is verrucosissimus, but unlike the latter, it is known to occur only on the territory of the not found in deforested places. Its Colchis and the adjoining sea coast up to April 1990 Asiatic Herpetological Research Vol. 3, p. 73 Vol. 3, p. 74 Asiatic Herpetological Research April 1990

FIG. 5. Pelodytes caucasicus occurs in all four Colchis refugia.

with forest and meadow formations of the account the above mentioned peculiarities Colchis type, makes it possible to identify of distribution of herpetofauna within the three more regions in the Caucasian Colchis refugium itself, the differences Isthmus, besides the Western Caucasus become even less significant. In this case, (the Colchis proper), in which the Colchis we deal with three regions smaller in space, herpetofauna occurs. The three regions are: and a wealth of species of the Colchis the Bjelo-Labinskij region on the northern herpetofauna that occur in refugia and have slope of the Western Caucasus, the survived off the main territory of the Kakhetinskij (Lagodekhi-Zakataly) region Colchis. on the southern slope of the Eastern The Belo-Labinskij region is only Caucasus, and the Borzhomskij region in conventionally separated from the Colchis Eastern Transcaucasia (Fig. 10). The by the crest of the Main Caucasian comparative composition of the Mountain Range. All the northern Colchis herpetofauna of these regions is shown in species, except L. a. grusinica, occur on Table 1. the northern slope of the Western Caucasus in the Belaja and the Small Laba river It is evident from Table 1, that the most drainages. It should be stressed that this significant differences are found between unity is based on the fact that the the Colchis and the Kahetinskij regions and characteristic Colchis elements of flora and the least significant differences are between vegetation cross over the Main Caucasian the Colchis and the Borzhomskij and the Range in the place known as "the Colchis Belo-Labinskij regions. Taking into Gates" to its northern slope. The basins of April 1990 Asiatic Herpetological Research Vol. 3, p. 75 Vol. 3, p. 76 Asiatic Herpetological Research April 1990

FIG. 7. Elaphe longissima. The Colchis refugia populations are disjunct from the main distribution in Europe.

the Belaja, Tsitse and Laba rivers abound in The Borzhomskij region is also those elements. To the north of these conventionally separated from the Colchis watersheds their distributions are by the Adzhara-Imeretinskij Mountain continuous up to the Skalistij (Rocky) Ridge. The flora and vegetation of the limestone ridge. Maleyev (1939) has noted Baniskhevskoje Gorge, the Likanskoje that a part of the Maikop district abounds in Gorge, and the upper belt of Mt. Lomis- Colchis elements and is inseparable from Mta, as well as the environs of Bakuriani, the Colchis according to the character of its hardly differ from those of the Colchis. flora and vegetation. Vol. 77 April 1990 Asiatic Herpetological Research 3, p.

FIG. 8. Vipera kaznakowi occurs throughout the territory of the Colchis up to 1000 m above sea level.

In the isolated eastern Kakhetinskij The early Miocene was about the time of region a considerable number of the ancient the formation of the Caucasian Mountains Tertiary vegetation representatives survived (Bogachev 1938). due to the warm and humid climate (Gulisashvili et al. 1975). The warm subtropical climate and vegetation in the Caucasus favored the Modern distributions of eco-geographic evolution and dispersal of heat and groups of amphibians and reptiles mesophilic forms (Triturus vittatus distinguished by our scientists have distinct ophryticus, T. vulgaris lantzi, Pelodytes altitudinal-ecological limits owing to natural caucasicus, Bufo verrucosissimus, Lacerta and historical reasons. saxicola darevskii, L. s. brauneri, L. derjugini, Elaphe longissima, Natrix Migration of ancestoral species of the megalocephala, and Vipera kaznakowi ), as Colchis and the Caucasian groups from the well as species with a broader ecological south apparently took place in the Miocene tolerance (Rana macrocnemis, Hyla arborea when the Caucasian island joined vast schelkownikowi, Lacerta rudis, and L. territories of Asia Minor. Colonization of agilis grusinica ). the Caucasus from the south by different species of mammals during the early Fossil remains of mammals Miocene has been studied by Vereschagin (Mesocricetus, Prometheomys, Sorex, (1958), and that of lizards of the Podarchis- Talpa ) [Vereschagin 1958] and insects Archaeolacerta group by Darevsky (1967). (Orthoptera, Hemiptera, Blattoidea, and Vol. 3, p. 78 Asiatic Herpetological Research April 1990

FIG. 9. Lacerta saxicola darevskii. This lizard is a Colchis endemic whose distribution is restricted to the northwestern part of the Colchis.

Coleoptera) [Rodendorf 1939] suggest a to the broad correlation of the Caucasus and good food supply for amphibians and the Balkans (Vereschagin 1958) and the reptiles during the Miocene. It was also in formation of the steppe landscapes along the Miocene that the majority of these the northern Black Sea coast (Pidoplichko species reached the eastern-most parts of 1954; Scherbak 1966). During that period, the Greater Caucasian Range along its such South-European species as Rana southern slopes and penetrated from there ridibunda, Bufo viridis, Emys orbicularis, into the Talysh across the so-called Anguis fragilis, Coluber jugularis, and "Karabakhi Bridge". Safarov (1966), and Coronella austriaca seem to have other scientists, have studied the former penetrated to the Precaucasia from the west. direct relations between the Colchis and the At the same time, such species as Testudo Hirkan floras. Even at the present time, the graeca, Pseudopus apodus, Triturus floristic composition of the Kakhetinskij cristatus karelini, Lacerta praticola pontica, region and of the Karabakh has many L. media, Coluber najadum, and Natrix common features with that of the Colchis tessellata got into the Colchis from the and the Talysh forests (Arushanyan 1973; west along the Black Sea coast. Sokolov 1977; Takhtadzhan 1978; Gadzhiyev et al. 1985). Early and Middle Pliocene should be considered the beginning of initial The end of the Tertiary period was fragmentation of the Colchis faunal areas characterized by damping of tectonics due when the Greater and the Lesser Caucasian April 1990 Asiatic Herpetological Research Vol. 3, p. 79

FIG. 10. The main refugia of the endemic Colchis herpetofauna.

Mountain ranges underwent substantial 1986). glaciation (Grozdetskiy 1954; Markov et al. 1965). The main center of dispersal of It was in the narrow humid gorges with these species was the Colchis, where a relatively constant thermal regimen that relatively heat-loving vegetation of the the representatives of the Colchis group Caucasian type survived even during the remained intact. At the same time, periods of the extreme Pleistocene cooling independent populations might also have (Vereschagin 1958; Adamyants 1971). been preserved in mid-mountain areas Along with the Colchis, smaller refugia where refugia of the Colchis vegetation sporadically survived on the territory of the exist in the vicinities of the Fisht-Oshtenskij Caucasus Black Sea coast, and also on the Mountain Massive, the Lagonaki Plateau northern slope of the Main Caucasian and even in the Central Caucasus Range between the Pshekha River and the (Kholyavko et al. 1978; Kharadze 1974). Small Laba River. The present distribution Small refugia seem to have also remained of the Tertiary vegetation of the Colchis on the southern slope of the eastern pan of type in the western Caucasus testifies to the Greater Caucasus and in the Kuru River this (Kharadze 1974; Pechorin and Gorge. It is indisputable that the majority Lozovoy 1980; Kholyavko et al. 1978; of the mountainous populations of Colchis Adamyants 1971; Koval and Litvinskaya species perished during the Pleistocene and Vol. 3, p. 80 Asiatic Herpetological Research April 1990

that the ones that survived in refugia have Oshtenskij Mountain Massive (Kholyavko been accumulating unique characteristics et al. 1978; Kharadze 1974; Dolukhanov which led to different geographical forms 1974; Galushko 1974). On the northern (subspecies) on different slopes of the Main slope such vegetation is present in the and the Adzharo-Imeretinskij mountain western parts and changes its character to ranges. The data presented by Takhtadzhan the east, transforming into a steppe type (1946) and Maruashvili (1956) support the (Lavrenko 1980). Modern areas of Vipera hypothesis concerning the preservation of dinniki and Lacerta caucasica alpina have relict Colchis species in the mountains. fixed distributional limits in the subalpine According to their data, the average annual belt influenced by the warm Black Sea. temperature during the glacial periods Final settling of the present-day climate decreased not more than 1.5-2.0°C, while favored fixation of the Colchis species precipitation amounted to not less than distributions, with their distinct populations 1500-2000 mm. exceeding the bounds of the refugia. This was coupled by simultaneous depression Darevsky (1967) considers that this and reduction of the European and argument supports the possibility of foothill especially of the Mediterranean species refugia of reptiles existing on the sea facing distributions. slopes of the Gagrinskij and the Bzhybskij mountain ranges, and in other regions, Concluding this review of the Colchis despite radical reorganization of the herpetofauna and their main refugia, it is distributions of all the species of plants and necessary to enumerate the most important animals in connection with glaciation. characteristics. The Colchis species are characterized by antiquity (conservation During the interglacial and especially the since the Tertiary period), Autochthonity, postglacial periods, reconstruction of all depression—for some species {Vipera vegetation belts took place (Vereschagin kaznakowi; Lacerta clarcorum, L. a. 1958). This favored the isolation of the grusinica), existence of the northern- species of the Colchis and the Caucasian Colchis limestone, and the southern- groups in the above-mentioned refugia, but Colchis volcanic centers of formation of favored wider dispersal of the European narrow-endemic forms. Reptiles have a and the Mediterranean groups in common tendency toward melanism, while Transcaucasia. In the northwestern part of amphibians approach their low temperature the Caucasus Black Sea coast, mesophilic thresholds. These are the adaptive features vegetation gave way to xerophytic acquired during the glacial period. As a vegetation of the Mediterranean type. Plant rule, the modem distribution of the Colchis formations of this type with the prevalence species does not exceed the bounds of the of Juniperetum, Querceto, Pinetum Colchis vegetation refugia or their carpinulosum, Pinetum fruticosum and derivatives. shibliaks are characteristic of the Anpa- Gelendzhik region. Enclaves of The maximal vertical distribution is in Mediterranean vegetation remained still the center of the Colchis up to 1800 m further to the south, up to Pitsunda above sea level, while in the other portions (Takhtadzhan 1978; Kolakovskiy 1961). of the refugia it does not exceed 1000 m above sea level as a rule. The existence of When the xerothermic period ended, the four refugia of the Colchis herpetofauna in climate became more humid again. This the Caucasian Isthmus is determined by favored the reestablishment of the former natural factors of high order- these are the borders of the forest belt (Vereschagin areas with slightly changed climatic 1958). Subalpine meadows and elfin conditions characterized by modern woodlands were expanding throughout the crossing of the January -3°C isotherm and whole subalpine belt of the southern slope 800 mm isohyet. of the Main Caucasian Mountain Range from the Central Caucasus to the Fisht- 1990 April Asiatic Herpetological Research Vol. 3, p. 81

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