Asiatic Herpetological Research

April 1990

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 abundanceand 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.

IntroductionThe

herpetofaunaofWestern

Transcaucasia is not homogeneous, due tothe different age and genesis of the species
distributions.
Along with autochtonousand endemic forms, one can find species
whose main areas of distribution are in theEuropean part of the USSR and in the
Eastern Mediterranean. At the same time, anumber of species which have main
distributional centers in the Colchis occur
beyond the bounds of Western
Transcaucasia, in other parts of the
Caucasian Isthmus. For these reasons, it isnecessary to define the Colchis
herpetofauna and to determine its place inthe fauna of reptiles and amphibians of the
Caucasian Isthmus as a whole.

Research on this issue started with theworks of Nordmann (1840), Derjugin(1899), Silantyev (1903), Brauner (1905),
and Nesterov (1911). However, the firstwell-grounded definition of the fauna inquestion from a zoogeographical point ofview was presented in the works of

Satunin (1912). Satunin wrote in 1910,"So far I cannot say much about the genesisof the fauna of this region called Western
Transcaucasia.
This country with its
plants and scanty fauna
resembles a piece of the Mediterranean in
the narrow sense of the word. True, here
are endemic species and forms, but not asingle genus of vertebrate is unrepresented
in the countries of the Mediterranean.

evergreen© 1990 by Asiatic Herpetological Research

Often they are inhabited by the same
species. The question is whether this fauna
has appeared from the west or is it theremainder of the fauna that has populateddensely the shores of the Black Sea at one
impossible to answer these
the present level of ourquestionsButeven now I can definitely
knowledge.
fauna
that
this
say
by its origin, has nothing
time.

It

is

at

in common with the faunas in other regionsof the Caucasus."

In 1912, Satunin divided the Caucasian
Isthmus into five subregions and 11
districts, including the Colchis in the WestTranscaucasian district of the Littoral

subregion.

Among other merits of this work by
Satunin, one cannot but mention the factthat for the first time, he defined in an exact
way the Colchis region proper. He defined
the northern border as the spurs of the Main
Caucasian Range up to the basin of theTuapse River, the southern border as the
Pontic Range, and the eastern border as the
Arsijanskij Range. The valley of the Rioni
River and the adjacent southern slopes ofthe Main Range were defined as the central
part of the region. Satunin emphasized thedepauperate herpetofauna of this region on
one hand, and the presence of endemic
species such as Vipera kaznakowi and
Bufo verrucosissimus on the other hand.

Nikolsky (1911) assigned the entire
Caucasus, excluding eastern Precaucasia,

Asiatic Herpetological ResearchVol. 3, p. 68

to the Mediterranean.

However, he could

not differentiate the forest and the alpine
belts of the Greater Caucasus, because of
the absence of data.

April 1990

Colchis, however, was not distinguished as
an independent center of speciation in thiswork.

Scherbak (1981) included the Colchis in

Results and Discussion
Investigations of the last decades made it
possible to add the majority of the speciesof the Colchis herpetofauna to an overall
picture of Colchis faunal distributions(Turov 1928; Bartenev and Reznikova
1935; Khozatsky 1941; Milyanovskiy
Bannikov
et
al.
1957;
1977;
Negmedzyanov and Bakradze 1977; Orlova
1973, 1978a, 1978b; Golubev 1980, 1985;
Tuniyev 1983, 1985). In addition there has
been a revision of the taxonomic status ofsuch forms as Vipera
kaznakowi
etal.
Orlov and
1986;
(Vedmederja
1990
this
1986a,
volume), Lacerta
Tuniyev
1960), L.
derjugini
andReznikova
1931; Orlova(Bartenev
1978a Bischoff 1982, 1984), L. saxicola(Darevsky 1967; Darevsky and Vedmederja
1977), Anguis fragilis colchicus (Lukina
1965; Scherbak and Scherban 1980), and(Peters

agilis;others.

Accumulation of this information alongwith works on fossil amphibians andreptiles of the Caucasus (Vekua et al. 1979;
Chkhikvadze 1981, 1983, 1984; Bakradze
and Chkhikvadze 1977; Zerova andthe Caucasian Region of the Mediterranean

Province. He suggested that the typical
species of the region were Mertensiellacaucasica, Pelodytes caucasicus, Lacerta

saxicola-complex and others. However,
the Colchis proper was again notdistinguished as an independent center ofherpetofaunal formation.

For the analysis of the herpetofauna of
the Colchis proper, it is necessary toexactly define the term "the Colchis
phytolandscapes", and to decide what typesof vegetation are universally recognized as"Colchis types". Albov (1885) was thefirst to clearly depict plant landscapes of
the Colchis.
He singled out a region,
for
Russia, of mountain limestone
uniqueflora

which had been developing mainly

autochthonously in a large refugium withnumerous endemic and relict species andeven genera. Kolakovskiy (1980) regarded
the Colchis flora as basically forest andalpine-meadow, and suggested that its main
phytolandscapes had existed since old times
with changes only in the composition of
their edificators, except for the extinct
formation of evergreen subtropical forestsin the lower mountain belt.

The tertiary-

1984; Yefimov andChkhikvadze 1987) have made it possible

relict character of the forest mesophile flora

to revise the zoogeography of the region.

slight changes in this region's climaticconditions (Kuznetsov 1891). The mostcharacteristic features of the tertiary-relict
Colchis forest are:extraordinary
abundance and diversity of tree and shrub
species, impossibility of singling out thedominant species (which is alsocharacteristic of tropical forest with extreme
density of trees), abundance of vines andepiphytes, and almost total absence of grasscover. All these attributes make the Colchis
forest similar in many aspects to a tropicalChkhikvadze

Darevsky(1957) singled out sevendifferent groups of species and subspeciesof the herpetofauna in the Caucasus, basedon their origin.
Among the species
representatives of the region of interest tous, it is necessary to pay attention toLacerta strigata (Asia Minor species),
Emys orbicularis, Anguis fragilis,
Coronella austriaca, Elaphe quatuorlineatessauromates, Natrix natrix (European boreal
species), Testudo graeca, Natrix tessellata(Mediterranean species), Pseudopus
apodus, Coluber najadum (eastMediterranean species), and Lacerta
saxicola, L. praticola, L. derjugini, L.
media (autochthonous species). The

and vegetation is fully revealed here due torain forest (Pavlov 1984).

According toforestColchisSinskaya
underwent
three
main
vegetation
stages of(1933),

thedevelopment: the tropical forest; the forestof Colchis type, but rich and covering awider area; and last, the modern Colchis

Asiatic

April 1990

Herpetological Research

forest.

The Colchis type of vegetation includes anumber of phytocenoses

differingVol. 3, p. 69

grusinica, Natrix megalocephala, andVipera kaznakowi are Colchis endemics inthe broad sense of the word.

incomposition, and ecologicalitpeculiarities:
may be mixed(polydominant), or may be presented bycenosis of one or two species, but thecommon and obligatory attribute ofphytocenosis of the Colchis type is anabundance of tertiary relicts. The area with

In addition to the Colchis endemics,

structure,

there are three more ecological-geographicalgroups of amphibians and reptiles in theregion.
They have similar ecologicalcharacteristics (habitat first of all), andoverlapping geographic distributions.

The

Colchis type vegetation is characterized bya rather monotonous thermal regimen,

consists of Triturus

corresponding to a subtropical climate, butwith highly diverse soils (Gulisashvili et al.

Testudo graeca nikolskii (Fig. 1 ), Lacerta
media, L. praticola pontica, L. strigata,1975).

Pseudopus

1.

tessellata,

The herpetofauna of the Colchis forests
is surprisingly poor, despite its uniqueness.

The species composition is different in thesoutheastern and northwestern parts of the
Colchis compared to the other portions ofSpecies such as Mertensiella
Lacertacaucasica,
clarkorum, L. parvula,and L. mixta, whose distributions areconnected with forests growing on acidsoils above volcanic rocks, are found on
the western slopes of the AdzharoImeretinsky, Shavshetsky and Lazistansky
its territory.

(Pontic)

mountain ranges.

Similarly,

floristic endemics of this part of the Colchis{Rhododendron ungernii,

Osmanthus

decorus, Betula medwedewii, Epigaea
gaultheriodes, and others), are AdzharoLazistan endemics, sometimes with slightradiations to the adjoining regions, but are
not Colchis endemics in the broad sense of

the word. The same applies to Lacerta
saxicola darevskii and L. saxicola brauneri

East-Mediterranean
cristatus

group

karelini,

apodus tracius, Natrix
and Coluber najadum. This

group's distribution includes either theBalkans and the Caucasus or the Balkans,
Crimea, and the Caucasus. According toecological characteristics, these arexeromesophiles or hemixerophiles whose
spreading is related to dry foothills ofWestern Transcaucasia up to 200-300 m
above sea level with an annual sum of
temperatures exceeding 5000°C. Thus,
Testudo graeca, Pseudopus apodus, andColuber najadum occur in the Colchis on a
narrow seaside strip of land with enclavesof Mediterranean vegetation from Tuapse to

Pitsunda-Sukhumi. A local population of
occurs in the Pitsunda region,
and a local population of L. media occurs
in the environs of Pitsunda and Salme. The
majority of localities of L. praticola andNatrix tessellata in the Colchis are in theseaside hills up to 400 m above sea level.
It is only
along the valleys of large rivers
L. strigata

like the Shakhe River, the Mzymta River,N

which are widespread in the northwesternthepart of the Colchis, but are absent in thecentral and southeastern Colchis. These
animals, by analogy with floristic endemics

tessellata penetrates into the Colchis up to{Allium

candolleanum,

Campanulamirabilis, C. bzybica, C. calcarea, C.

jadvigae, Genista abchasica, Gentiana

Bzyb

River,and others that

.

m

600
above sea level. Thus the majorityof these species are found either in places
with vegetation of the Mediterranean type,or in places where the initial Colchis
vegetation has been reduced to zero and theparadoxa, Omphalodes kusnetzovii, andothers) are northern-Colchis endemics. For
example, of the 450 endemic Colchis
species of flora, 83 (25%) are endemics ofthe northern Colchis (Adzinba 1980).

landscapes resemble the Mediterranean
ones by their thermo-biotopic conditions

Triturus vittatus ophryticus, T. vulgaris
lantzi, Bufo verrucosissimus, Pelodytes
caucasicus, Lacerta derjugini, L. agilis

Juniperus oxicedrus, Pinus pityusa, andArbutus andrachne).

(substitute shibliaks and tomillares,
Pitsunda pine groves, foothill post-forest
glades, and landplots with Erica tetraliz,Vol. 3, p. 70

Asiatic Herpetological Research

FIG. 1. Testudo graeca nikolskii

is aApril 1990

Mediterranean species, and in the Colchis it is found only in theseaside strip of land with enclaves of Mediterranean vegetation.

2. The Caucasian group includes Hylaarborea schelkownikowi (Fig. 2), Rana
macrocnemis, Lacerta caucasica alpina, L.
rudis, and Vipera dinniki. Distributions ofthese species in the Caucasian Isthmus arebroader than those of the Colchis group.
At the same time, the majority of them are
Colchis autochthons. These species aremesophiles and occur in mesophillous
forest and mountain meadow formations.

The composition of this group is nothomogeneous. It includes both species
typical for the steppe areas {Bufo viridis
and Coluber jugularis ) and those that are

widely distributed in Europe (all the rest).

Only Anguis fragilis
austriaca
Colchis.

and Coronella

are widely distributed in theThis makes it difficult todefinitely consider them late migrants to the
Caucasus. Other species either occur inThis group seems to be of Colchis origin,
retaining close connections with the main
center of the formation. Broader ecologicaltolerance in comparison with typicalColchis species makes it impossible toinclude them within the Colchis group.

several spots along the Colchis seashore(B. viridis and Natrix natrix ) or populate anarrow strip of land along the sea together
with the Mediterranean species (C.
jugularis ) or a somewhat wider strip(Emys orbicularis and Rana ridibunda ).
Despite the possibility of finding these

The European group consists of Bufo

species in the typical Colchis forest
formations, the majority of them are still
attributed to the Mediterranean type of3.

viridis, Rana ridibunda, Emys orbicularis,

Anguis fragilis, Natrix natrix, Coronella
austriaca, and Coluber jugularis caspius.

vegetation.

April 1990

Asiatic Herpetological ResearchVol. 3, p. 71

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
same places in the Colchis as T.

distribution of the representatives of thein theColchis group in detail. Triturus vittatus
occurs in the territory from the seashore tovittatus ophryticus

the subalpine meadows in all the forest
In the place called "the Colchis
types.

Gates" (lowering of the Main CaucasianRange between Mt. Fisht and Mt.
Chugush) the species crosses over to thenorthern slope of the Western Calucasus,
reaching the environs of Goriachij Klutch
and Krasnodar in the northwest and thebasin of the Laba River in the northeast. In
the eastern part of the area, the species
crosses the Adzharo-Imeretinskij mountain
range and reaches the outskirts of TbilissiOni. It occurs in the Lagodekhi region as
an isolate. Outside the boundaries of the
Colchis this species occurs either in the
Colchis type forests or in their derivatives.

does. Very often botharesymbiotopic (Tuniyev andspeciesOn the northern slope1986).
Beregovayaof the Western Caucasus, its home range iswider than that of the previous species, but
is only through the mesophillous forestsand subalpine meadows that it penetrates
into the Eastern Transcaucasia up to theit

The isolated population inin
the Hirkan forests which
occursTalyshare ecologically and genetically close to the
Colchis forests.
Trialet Ridge.

Bufo verrucosissimus (Fig. 4) occurs in
Colchis from the sea shore

all parts of theOn theto the subalpine forests.
northern slope of the Western Caucasus, it
is found on the territory from the environsup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
settlement and the Shakhgirej Canyon in theeast, where it is found in the derivatives of
the Colchis forests from 400
to 1000
above sea level. In the Eastern Caucasus it
is found in the Borzhomy Gorge, themm

Lagodekhi-Zakataly, and the Talysh region,
where its distribution is limited to themesophilous forests, which are abundant in
the Colchis and Hirkan floral elements.

The distribution of Pelodytes caucasicus(Fig. 5) in the Colchis is more restricted. It
does not occur in the coastal belt and oakforests. It is found both in the mesophilous

beech, chestnut, and fir-tree forests, and in
mixed broad-leaved forests with anevergreen understory. On the northernslope of the Western Caucasus, its
distribution coincides with that of B.
verrucosissimus, but unlike the latter, it is
not found in deforested places.
Itssoutheastern distributional limits also
coincide with that of Bufo verrucosissimus.
Pelodytes caucasicus does not occur eastof the Trialet Ridge. There is an isolated
population in the mesophilous forests in the

Lagodekhi-Zakataly region.

Lacerta derjugini has a distribution in
the Colchis similar to P. caucasicus.

It

reaches the sub-alpine belt.
On thenorthern slope of the Western Caucasus, it
is found in the Colchis forest derivatives
from the Belaja River to the Small Laba
River (the Shakhgirej Gorge). The species
theEasternpenetrates
through
Transcaucasia up to the Trialet Ridge.
Separate populations occur in north-eastern
Georgia up to Lagodekhi-Zakataly.

Lacerta agilis grusinica(Fig. 6) is

known to occur only on the territory of the
Colchis and the adjoining sea coast up toApril 1990

Asiatic Herpetological ResearchVol. 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
Colchis type, makes it possible to identify
three more regions in the Caucasian
Isthmus, besides the Western Caucasus(the Colchis proper), in which the Colchis

account the above mentioned peculiarities
of distribution of herpetofauna within the
Colchis refugium itself, the differences

herpetofauna occurs. The three regions are:the Bjelo-Labinskij region on the northernslope of the Western Caucasus, theKakhetinskij (Lagodekhi-Zakataly) region
on the southern slope of the Eastern
Caucasus, and the Borzhomskij region in
Eastern Transcaucasia (Fig. 10).
The
of
the
comparative compositionofthese regions is shown inherpetofauna
Table 1.

and a wealth of species of the Colchis
herpetofauna that occur in refugia and haveevident from Table 1, that the mostsignificant differences are found between
the Colchis and the Kahetinskij regions andIt isthe least significant differences are between
the Colchis and the Borzhomskij and theBelo-Labinskij regions.

Taking into

become even less significant.

In this case,

we deal with three regions smaller in space,survived off the main territory of the
Colchis.

The Belo-Labinskij regionis only
fromtheColchisconventionally separated
Main
Caucasian
of
the
the
crest
by
Mountain Range. All the northern Colchis
species, except L. a. grusinica, occur on
the northern slope of the Western Caucasus
in the Belaja and the Small Laba riverdrainages.

unity

is

It

should be stressed that thisbased on

the

fact

that

the

characteristic Colchis elements of flora andvegetation cross over the Main CaucasianRange in the place known as "the Colchis
Gates" to its northern slope. The basins ofApril 1990

Asiatic Herpetological ResearchVol. 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

those elements.
To the north of thesewatersheds their distributions arecontinuous up to the Skalistij (Rocky)
limestone ridge. Maleyev (1939) has notedthat a part of the Maikop district abounds in
Colchis elements and is inseparable fromthe Colchis according to the character of its
flora and vegetation.

The

Borzhomskij

regionisalsoconventionally separated from the Colchis
by the Adzhara-Imeretinskij Mountain

Ridge.

The flora and vegetation of theBaniskhevskoje Gorge, the Likanskoje
Gorge, and the upper belt of Mt. LomisMta, as well as the environs of Bakuriani,
hardly differ from those of the Colchis.

Asiatic Herpetological Research

April 1990

Vol. 3, p. 77

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

the isolated eastern Kakhetinskij
region a considerable number of the ancient
Tertiary vegetation representatives survived
In

due

to

the

The early Miocene was about the time of
the formation of the Caucasian Mountains(Bogachev 1938).

warm and humid climate
The warm subtropical climate and(Gulisashvili et al. 1975).

Modern distributions of eco-geographic
groups of amphibians and reptiles
distinguished by our scientists have distinct
altitudinal-ecological limits owing to naturaland historical reasons.

vegetation in the Caucasus favored theevolution and dispersal of heat andmesophilic forms (Triturus vittatus
ophryticus, T. vulgaris lantzi, Pelodytes
caucasicus, Bufo verrucosissimus, Lacerta
saxicola darevskii, L. s. brauneri, L.

derjugini,

Elaphe longissima, Natrix

when the Caucasian island joined vast

megalocephala, and Vipera kaznakowi ), as
well as species with a broader ecologicaltolerance (Rana macrocnemis, Hyla arborea
schelkownikowi, Lacerta rudis, and L.

territories of Asia Minor.

agilis grusinica ).

Migration of ancestoral species of the
Colchis and the Caucasian groups from thesouth apparently took place in the Miocene
Colonization of

the Caucasus from the south by different

species of mammals during the early
Miocene has been studied by Vereschagin(1958), and that of lizards of the PodarchisArchaeolacerta group by Darevsky (1967).

remainsofmammals
Fossil
(Mesocricetus, Prometheomys, Sorex,
Talpa ) [Vereschagin 1958] and insects(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 thenorthwestern part of the Colchis.

Coleoptera) [Rodendorf 1939] suggest agood food supply for amphibians andreptiles during the Miocene. It was also in
the Miocene that the majority of these
species reached the eastern-most parts of
the Greater Caucasian Range along its
southern slopes and penetrated from there
into the Talysh across the so-called"Karabakhi Bridge". Safarov (1966), andother scientists, have studied the former
direct relations between the Colchis and the
Hirkan floras. Even at the present time, thefloristic composition of the Kakhetinskij
region and of the Karabakh has many
common features with that of the Colchis
and the Talysh forests (Arushanyan 1973;
Sokolov 1977; Takhtadzhan 1978;

to the broad correlation of the Caucasus andthe Balkans (Vereschagin 1958) and theformation of the steppe landscapes along
the northern Black Sea coast (Pidoplichko

1954; Scherbak 1966). During that period,
such South-European species as Ranaridibunda, Bufo viridis, Emys orbicularis,
Anguis fragilis, Coluber jugularis, andCoronella austriaca
seem to havepenetrated to the Precaucasia from the west.
At the same time, such species as Testudo
graeca, Pseudopus apodus, Trituruscristatus karelini, Lacerta praticola pontica,
L. media, Coluber najadum, and Natrix
tessellata got into the Colchis from thewest along the Black Sea coast.

Gadzhiyev et al. 1985).

Early and Middle Pliocene should beconsidered the beginning of initial

The end of the Tertiary period wascharacterized by damping of tectonics due

fragmentation of the Colchis faunal areas
when the Greater and the Lesser CaucasianApril 1990

Asiatic Herpetological ResearchVol. 3, p. 79

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

Mountain ranges underwent substantial
glaciation (Grozdetskiy 1954; Markov et al.
1965). The main center of dispersal ofthese species was the Colchis, whererelatively heat-loving vegetation of the
Caucasian type survived even during theperiods of the extreme Pleistocene cooling(Vereschagin 1958; Adamyants 1971).
Along with the Colchis, smaller refugia
sporadically survived on the territory of the
Caucasus Black Sea coast, and also on thenorthern slope of the Main CaucasianRange between the Pshekha River and the
Small Laba River. The present distributionof the Tertiary vegetation of the Colchis
type in the western Caucasus testifies to(Kharadze 1974; Pechorin andLozovoy 1980; Kholyavko et al. 1978;
Adamyants 1971; Koval and Litvinskaya
this1986).
It

was in the narrow humid gorges witha relatively constant thermal regimen thatthe representatives of the Colchis groupintact.
At the same time,remained

independent populations might also havebeen preserved in mid-mountain areas
where refugia of the Colchis vegetation
exist in the vicinities of the Fisht-Oshtenskij

Mountain Massive, the Lagonaki Plateau
and even in the Central Caucasus(Kholyavko et al. 1978; Kharadze 1974).
Small refugia seem to have also remainedon the southern slope of the eastern pan of
the Greater Caucasus and in the Kuru River
Gorge. It is indisputable that the majorityof the mountainous populations of Colchis
species perished during the Pleistocene andAsiatic Herpetological ResearchVol. 3, p. 80that the ones that survived in refugia havebeen accumulating unique characteristics
which led to different geographical forms(subspecies) on different slopes of the Main
and the Adzharo-Imeretinskij mountain

The data presented by Takhtadzhan(1946) and Maruashvili (1956) support the
hypothesis concerning the preservation ofrelict Colchis species in the mountains.
According to their data, the average annual
ranges.

temperature during the glacial periods
decreased not more than 1.5-2.0°C, while
precipitation

amounted to not less than

1500-2000 mm.

Darevsky (1967) considers

that thisargument supports the possibility of foothill

April 1990

Oshtenskij Mountain Massive (Kholyavko
et al. 1978; Kharadze 1974; Dolukhanov
1974; Galushko 1974). On the northernslope such vegetation is present in thewestern parts and changes its character tothe east, transforming into a steppe type
(Lavrenko 1980). Modern areas of Vipera
dinniki and Lacerta caucasica alpina have
fixed distributional limits in the subalpine
belt influenced by the warm Black Sea.
Final settling of the present-day climate
favored fixation of the Colchis species
distributions, with their distinct populations

exceeding the bounds of the refugia. Thiswas coupled by simultaneous depression
and reduction of the European andespecially of the Mediterranean species

refugia of reptiles existing on the sea facing
slopes of the Gagrinskij and the Bzhybskij

distributions.

mountain ranges, andConcluding this review of the Colchis
herpetofauna and their main refugia, it isnecessary to enumerate the most important

despiteradicalin other regions,reorganizationof thedistributions of all the species of plants andanimals in connection with glaciation.

During the interglacial and especially the
postglacial periods, reconstruction of allvegetation belts took place (Vereschagin1958). This favored the isolation of thespecies of the Colchis and the Caucasiangroups in the above-mentioned refugia, butfavored wider dispersal of the European
and the Mediterranean groups inTranscaucasia. In the northwestern part of
the Caucasus Black Sea coast, mesophilicvegetation

gave

way

to

xerophytic

vegetation of the Mediterranean type. Plant
formations of this type with the prevalenceof Juniperetum, Querceto, Pinetum

characteristics. The Colchis species arecharacterized by antiquity (conservation
since the Tertiary period), Autochthonity,
depression for some species {Vipera—

kaznakowi; Lacerta clarcorum, L. a.
grusinica), existence of the northernColchis limestone, and the southernColchis volcanic centers of formation of
narrow-endemic forms. Reptiles have acommon tendency toward melanism, whileamphibians approach their low temperaturethresholds. These are the adaptive featuresacquired during the glacial period. As arule, the modem distribution of the Colchis
species does not exceed the bounds of the
Colchis vegetation refugia or theircarpinulosum, Pinetum fruticosum andshibliaks are characteristic of the AnpaGelendzhik region.
Enclaves ofMediterranean vegetation remained still
further to the south, up to Pitsunda(Takhtadzhan 1978; Kolakovskiy 1961).

derivatives.

When the xerothermic period ended, theclimate became more humid again. Thisfavored the reestablishment of the former
borders of the forest belt (Vereschagin

four refugia of the Colchis herpetofauna in
the Caucasian Isthmus is determined bynatural factors of high order- these are theclimatic
areas with slightly changed1958).
Subalpine meadows and elfin
woodlands were expanding throughout thewhole subalpine belt of the southern slopeof the Main Caucasian Mountain Range
from the Central Caucasus to the Fisht-

conditions

The maximal vertical distribution is in
up to 1800 m
above sea level, while in the other portionsof the refugia it does not exceed 1000 m
above sea level as a rule. The existence ofthe center of the Colchis

characterized by modernofthecrossing
January -3°C isotherm and

800 mm isohyet.

Asiatic Herpetological Research

April 1990

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