(Colchis and Talysh) of Southern Caucasus Download

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The results of biogeographical study of the Arcto-Tertiary refugia (Colchis and Talysh) of Southern Caucasus
Arnold M. Gegechkori
I. Javakhishvili Tbilisi State University, Department of Biodiversity, Chavchavadze Ave. 1, 0179
Tbilisi, Georgia
Abstract
South Caucasus two regions Colchis and Talysh are one of the species-rich refugia for many Tertiary relict organisms –
plant and animal species in western Eurasia. Due to two regions peculiar geography, abiotic factors and natural history both
refugial centres of South Caucasus are of particular interest for biogeographical studies. However, present article is a first
attempt to a such interdisciplinary investigaion.
We analysed patterns of floristic and faunistic richness, endemism, including relict endemics, modern distribution of
close relict species in other countries and regions of Northern Hemisphere, specificity of formation of two harbour
territories of Tertiary organisms, palaeo-ecological data of the Black seacoastal and Caspian seacoastal regions, comparative
analyses of the specificity of altitudinal zonation of Colchis and Talysh, at the same time comparing the structure of
surviving in mentioned shelters two ancient elements of native flora and fauna – a) tropical-subtropical, b) organisms of the
Arcto-Tertiary origination. Finally, critical analyse was dedicate to two refugial centres, which by some botanists are
recognized as moist subtropic biocenosis.
Key words: biodiversity, biogeography, biomes, refugia, Colchis, Talysh.
Material and methods
The overview of biomes and comperative analyse of two famous rafugial centres of the Caucasus – Colchis and Talysh is
first and foremost the result of the long-term comprehensive (faunistic, floristic, biogeographical) studies of the wildlife of
the Caucasus by the author of present articles cycle. We focussed first of all on the psyllids (Insecta: Hemiptera,
psylloidea). This group of insects constitude a highly indicative group for environmental
research (Hodkinson, 1974, 2009; Loginova, 1968; Gegechkori, 1984a,b, 1985, 1998). Psyllids
are small phloem feeding insects that are typically monophagous (feed on a single plant
species) or olygophagous (feed on a few relates species). The data were gathered during almost
thirty years (1962-1990) of field work in all biomes and altitudinal zones of the naturalhistorical regions of the Caucasus and in all vegetation seasons (early spring – late autumn).
During the years 1990-2008 similar excursions were made but with lower intensity. Data for
interregional comparison were collected in Middle Asia, Siberia and the Russian
Fig. 1. Leafhopper (psyllid)
Far East (1981-1991), in Asia Minor (1995-1998, 2006), and in other parts of the world
Trichochermes walkeri (Förster)* (1986, 1989, 2000-2003). The results are presented in various monographs and articles
(Gegechkori, 1977, 1978, 1984a,b, 1985, 1991, 2000; Gegechkori & Jibladze, 1976;
Gegechkori & Loginova, 1990).
Map 1. Caucasus Ecoregion with Colchis and Talysh refugia
*All photos belongs to author.
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Colchis – an Important Harbouring Region of Tertiary Organisms
Colchis (Colchida, Kolkheti) part of West Georgia, situated between 41° to 45°N and 40° to 46°E. This political
and administrative perimeter of the Colchis well corresponds with its natural borders ( Kikvidze & Ohsava, 2001).
From the botanical-geographical point of view Colchis belongs to the Eastern Euxinian or Colchian Phytogeographic
Province. This region from West Georgia slightly expands to Russia, in the territory of north-western part of the
Caucasus (Tuapse-Novorosiysk and to Turkey), within north-eastern part of Anatolia (Albov, 1896; Maleev, 1940;
Kolakowsky, 1961; Manjavidze, 1967; Kharadze, Gagnidze, 1970; Gagnidze, 1974, 1999, 2002). Therefore, Colchis
comprises an area, forming so called the ,,Colchic Triangle”(Walter, 1968; Kikvidze & Ohsawa, 2001).
The central place of the most humid area is occupies by South Colchis with West Georgia’s two provinces, Adjaria
and Guria. This is a narrow stripe of coastal lowland between the estuaries of the rivers Tchorokhi and Rioni with
considerably humid and warm-temperate climate. Within this region maximum precipitation known from Mt.Mtirala in
Adjaria (Mtirala from Georgian – ,,cry-one baby”), namely 4500 mm, which is a record data for the Caucasus Isthmus.
In general, the Colchian or Eastern Euxinian Province’s flora consist of about 2500 species, of which approximately
2000 species are distributed in Georgia (Gagnidze, Davitadze, 2000).
On the Eocene-Oligocene boundary the folded structure of the Alpine system began to be formed in the
Mediterrannean orogenic band. It comprises the geosynclinal regions of the Southern slope of the Greater Caucasus and
the Adjara-Trialeti mountain system. The Late Miocene (Sarmatian time) was the turning point in the geological history
of the Caucasus. This time was characterized by huge orogenic movement. The formation of mountain structures and
the division of the territory of Georgian land into two great regions – Western and Eastern ones were continued to this
time. Two regions were separated by Dzirula Masiffe, the same Surami Ridge (Gvozdetky, 1958; Chubukhov, 1966;
Khain, 1975; Milanovski, 1977; Gamkrelidze, 1986, 1997; Adamia et al., 1977, 1991; Volodicheva, 2003).
Approximately in mentioned geological time, the territory of adjasenting to the Black Sea, transformed into the
isolated within the Caucasus region. The warm and humid climate predominated here, promoting to survive the
hygrophilous and thermophilous plants, being died out in the neighbouring territories. Throughout the Late Cenozoic
this, broadly known as Colchic refugium was the most stable from abiotic viewpoint region, where the elements of the
Tertiary flora was survived (Kolakowsky, 1961; Chelidze & Kvavadze, 1987; Dmitrieva, 1990; Shatilova & Ramishvili,
1990; Makharadze, 1998).
Richness of Colchis vegetation as a centre of biological diversity, high ratio of endemism and Tertiary relict species
within borders of western Eurasia, is well known (Grossheim, 1936, 1948; Maleev, 1940, 1941; Kolakowski, 1961;
Ketschoveli, 1960; Gulisashvili et al, 1975; Dolukhanov, 1980; Manjavidze, 1982; Zazanashvili, 1999; Gagnidze &
Davitadze, 2000; Kikvidze & Ohsava, 2001).
Grossheim (1948) indicated that in polydominant forests of Colchis, major forests - building species include 5
species: chestnut (Castanea sativa), an oak (Quercus hartwissiana), beech (Fagus orientalis), hornbeam (Carpinus
betulus), and alder (Alnus barbata).
In some localities of Colchis in the formation of local forests communities participate 2-3 species.
If Talysh phytostructure is favourable with its archaic representatives in the canopy, the Colchis shelter is no less
impressive with its understory.
The botanico-geographical character of Colchic specific evergreen broad-leaved arboreal component of deciduous
forest’s undergrowth, their florogenetic connections attract attention of many botanists (Grossheim, 1936, 1938; Wolfe,
1944; Walter, 1974; Dolukhanov, 1989; Kikvidze, Ohsava, 2001; Nakhutsrishvili, 1999; Zazanashvili, 1999; Schmidt,
2004). To mentioned communities a special monograph was dedicated by A. Dolukhanov (1980), arealogical review
belongs to R. Gagnidze (1999). Phytocoenologically these formations are called “Colchian understorey” (Dolukhanov,
1980), or “Colchian semi-prostrate evergreen underwood” (Gagnidze, 1999), although in composition of Colchis
undergrowth participated deciduous shrubs as well.
The most species-rich relict communities of underforest occur in South Colchis. As it was underlined, this is a
region of maximum accumulation of snow cover and considerably amount of summer precipitations. Therefore, the
present data and corresponding analysis base on mentioned key refugium, or as it is named by botanists, “refugium in
refugium” (Kikvidze & Ohsava, 2001).
The various formations of creeping (semi-prostrate and prostrate) underforest in the Colchis extends mainly from
1700 to 2200m a.s.l. The upper limit of distribution of mentioned thickets coinsides with extreme dispersion of beech
crook-stem forests, and timberline area in general. In maritime zone’s mountains of Adjaria this upper border situated
even about 2250-2370m. The lowest limit of distribution of semi-creeping communities are determined by the types of
relief and competition among this kind of phitocoenosis and straight-trunk beech or spruce forests (Dolukhanov, 1980).
According to this scholar (Dolukhanov, 1974), among all types of vegetation of the Caucasus, semi-prostrate and
prostrate communities are most largely enriched with the ancient relict and relict-endemic species. Mentioned should be
species of following genera: Rhododendron, Ruscus, Ilex, Laurocerasus, Epigaea, Daphne, Hedera and some other
deciduous shrubs (Vaccinium, Rhamnus, etc.). Outside of Colchis, including Hyrcania, these specific formations are in a
state of “dying relicts”, whereas in Colchic ecosystems majority of them perform a role of progressive or not reduce
theirs area relicts, consequently being edificatores within underground stratum.
According to Dolukhanov (1980), there are two ecological groups of Colchic underwood. Representative of
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one group are less shade-resistant, i.e., light-demanding shrubs. They present mainly in subalpine belt and
grow in some local places, where should not be forced out by the tree stands of high trunk forests.
Within underwood of semi-prostrate formations prevail some very ancient relict and endemic species of Colchis:
Betula medwedewii, Quercus pontica, Rhamnus imeretina, Sorbus subfusca, Corylus colchica, Daphne alboviana, and
widely distributed in highmountains of the Caucasus evergreen shrub – Rhododendron caucasicum.
The second ecological groups situated by highlands’ prostrate and semi-prostrate woody plants, known as a
vegetative-migratory shrubs. Frequently they are represented as shade - endurance formations. Some of them in specific
climatic conditions may grow as more or less straight-trunk trees. Among these associations botanists special attention
are paid to species of following genera: Rhododendron, Epigaea, Vaccinium, Laurocerasus, Ilex, Ruscus, Viburnum.
According to Dolukhanov (1980), the relict vegetation of Colchic underforest in Pleistocene’s Ice Ages Periods was
able to survive only in mountain gorges of Colchis and adjacent slopes of the Pontus Ridge near the Black Sea coast but
not in foothills and low forest zone. The winter’s severe frosts of glacial periods, distribution of cold atmospheric
masses, can cause deep temperature depressions of evergreen shrub layer even in coastline zone, hence the latter might
have been survived only under deep snow cover.
Thus Colchis mesophillous semi-prostrate evergreen underwood is a richest formation in western Palaearctic with
so splendid latitudes and altitudinal distribution, florogenetic elements group, which have been formed under conditions
of positive abiotic factors throughout the long-lasting evolution period.
Species connected with centre I. According to Dolukhanov (1980) and Gagnidze (1999) one of the main genera of
Colchic complex is Rhododendron. In Colchis this genera includes 5 species: R. ponticum, R. ungernii, R. smirnowii, R.
caucasicum, R. luteum, and one hybrid form – R. sochadzeae (Rhododendron ponticum mixed with R. luteum).
As it was stressed above the close floristic alliance between eastern
Asia and North America have long been discussed by plant
geographers (Asa Gray, 1846; Wolfe, 1944; Li, 1952, 1972; Graham,
1972; Raven, 1972; Thorne, 1972; Guo, 1999; Guo et al., 1998;
Qian, 2001).
One of the obviously example of the eastern Asia-North
America’s macro-disjunctive area is expressed by genus Epigaea.
From Asia (Japan)
known E. asiatica and E. rosea – in North
America (Takhtajan, 1941, 1997; Wood, 1961; Hersey & van der
Kloet, 1976; Schmidt, 2004).
Mentioned disjunction area become more impressive by the third
representative of this oligotypic genus, E. gaulhteroides, the typical
Colchic – Taurus species (Davis, P.H., 1965-1988). The latter closer
related to the Japaness species. Such configuration of vast disjunct
area confirm the opinion of biogeographers concerning relations of
the Caucasian flora with the centres of development of evergreen
woody plants since pegan time (Wolfe, 1944; Grossheim, 1936;
Dolukhanov 1980).
Fig.2. A pure stand of Rhododendron
ungernii in beech forest, 1800m a.s.l.
(Guria, W. Georgia)
The Epigaea gaultheroides is a classical example of a such relict
vegetation. This kind of organisms – a surviving remnants (,,living
fossils”), as it broadly known, are regarding to ,,regressive relicts”.
In remote past epigea was widely distributed. This suggestion is confirmed by fossil remnants discovered in North
America and Europe. Ilinskaya (1968) in Pliocene’s flora of Transcarpathia found prints of Epigaea baikovskiae, close
species to the Colchian epigea.
Within the South Caucasus two refugia’s animals, according to theirs origin history, it can be outlined two groups:
first one is relict forms. Representatives of this group have been preserved some ancient (plesiomorphic) anatomical and
other features of theirs subtropical-tropical ancestors of the Tertiary Period. This is a geological past, when difference
between the faunas of Europe and tropical Asia were far less significant than it is nowadays.
The second group, which in faunistical literature is frequently mentioned as an “Indian” one, includes species with
contemporary (apomorphic) anatomical features, occur as typical inhabitants of present-day’s tropical-subtropical
latitudes’ countries, although some of them during postglacial period have been extended their areas within some
countries of temperate zone, including the Caucasus. Mentioned should be tiger, porcupine, hyena, etc.
Vertebrate fauna of the Tertiary of Colchis is very poor. Vereschagin (1958) consider that black rat (Rattus
rattus) belongs to the Pliocene relict of Mediterranean proper regions.
As regards invertebrate fauna of Colchis, particular attention should be paid to remarkable moth, Lederer’s
brahmea – Brahmaea ledereri (ordo Lepidoptera) from Colchis (Adjaria) and adjacent north-eastern part of Turkey. It
is endemic and relict to Colchis region, vicarious form of Lenkoran’s brahmea (B. christophi). Large insect (wingspan is
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95-107mm), ornamented with exclusive beautiful, ,,tropical”colour. Habitats presented in lowland’s dark and moist
forests.. Brahmea is living ,,eyewitness” of deep anciency, inhabited in South Colchis since remote past, approximately
from latest Paleogenic (Rijabov, 1958; Gegechkori, 2000; Didmanidze, 2005). Taking into consideration this fact that
entomologists did not discover this species until the last 50-70 years, B. ledereri must be on the range of extinction, or is
already eliminated from Colchis refugium (Gegechkori, 2000; Didmanidze, 2005).
Another species of the genus Brahmaea (B. certhia) occur in the next fomous refugium, in Russian Far East, (r.
Amur Gorge), and then in Korea, China, Japan and India.Thus with so few species (7-8 forms mainly from tropics) is
exhausted the richness of mentioned ancient genus, the remains of which are scattered now within so remote but
climatic viewpoint more or less similar places. On the example of allopatrically dispersed species united in Brahmaea
genus, could provide valuable information for our understanding of global climate change patterns, its influence, of
living organisms eliminations and surviving processes, the role of harbours – refugia – during major Ice Ages in
preservation some archaic thermophilous plants and animal taxons of the Tertiarian Period.
Thus, as we see, like relict woody plants of Colchis, there are two different groups of butterflies, one of them,
resemble to Epigaea gaultheroides, belongs to ,,dying relicts” (species of Brahmaea), the another ones, like Castanea
sativa and particularly Fagus orientalis, characterized by vital capacity, consequently has not only survived till presentday, but found in large areas, renew their ranges in postglacial time.
The most trustworth representative of ancient Tertiary relict fauna of mesophylic ecological type, belongs the
Colchian Isophya (I. redtenbacheri). Chorologically this species is restricted with lowlands forests, feeding on local
woody plants and by its anatomical features has closer relations to another relict species of the same autoecology, the
Caspian isophya (I.caspica) _ one of the obviously element of Hyrcanian centre of antiquity (Bey-bienko, 1958). Thus,
species of Isophya show the same vicarious distribution within Transcaucasia’s two refugia, which is characterized to
above discussed genus Brahmaea.
A typical, the Colchic Tringle distribution attracts to such representative of subphyllum Chelicerata, as is the
Colchis subendemic and relict subspecies of scorpion – Euscorpius mingrelicus mingrelicus. According to V. Fet
(1993), some species and subspecies of genus Euscorpius have many endemics end show remarkabely disjunct
distribution.
Taking into account that scorpions (ordo Scorpiones) are one of the most ancient group not only among class
Arachnida but among terrestrial arthropods in general, their Laurasian and Gondwanian distribution attract particular
interest of many zoogeographers.
Subspecies E. mingrelicus mingrelicus within the Colchic Tringle represented all regions of south-west Russia,
western Georgia and north-eastern Turkey, inhabiting in moist broad-leaved forest of Colchic type.
Allopatric speciacion of E. mingrelicus in subspecies level is good example, which resambles to geographic speciacion of
that Rhododendron. According to Bonacina (1980) and Fet (1993), E. mingrelicus covers an area, including except the
Colchis and adjacent territory of turkey (Taurus district) and southwest Russia (Krasnodar Region), Mediterranean proper
countries as well.
Talysh – Another Refuge of Tertiary Organisms in South Caucasus
The Talysh region is a part of Azerbaijan in the south - eastern part of Transcaucasia. It is located between 38° to 39°
N. The territory of the region covers about 4700 sq. km. The local refuge is known as Hyrcania. It is adjacent to the southern
and south-western (Iranian part of refuge) coast of the Caspian Sea. It consists of two geographical units, the Lenkoran
Lowland and the Talysh mountains. The latter are prolongation of the Lesser Caucasus and form transitional Mountains
chain between the Lesser Caucasus and the Elburz mountains in Iran. In this country they come into contact, but have no
tectonic connections (Antonov, 1955). Iranian part of Talysh region occupies more territories than that in Caucasus’
Azerbaijan (It’s Iranian Azerbaijan as well), (Grossheim, 1926).
Talysh consists of three mountain chains (the highest – peak reaching more than 2400 m a.s.l.) and a number of
ranges. Therefore, unlike of the Greater and the Lesser Caucasus, the Talysh mountains have no highlands with alpine belt.
The mountains system of Talysh is also distinguished with arid climate of steppe type in its upper part - Zuvand region with
Diabar Depression (Hollow), part of Zuvand, embracing Iranian side of the territory. Mid-and lower zones of mountains are
characterized by moist subtropical climate with rich broad - leaved deciduous forests. Therefore, Talysh’s foothills and midmountain slope’s vegetation are situated between two arid landscapes. From lower part this is a Lerik district’s deserts and
semi-deserts, neighbouring Lenkoran Lowland and from upper part - Zuvand Mountain’s desertificated steppes. The
mentioned extraordinary geographical position of the Talysh mountains, directed to the Caspean Sea, turns them into
,,wooded islands”.
Talysh mountains distinctly differ from other orographyc systems of South Caucasus. Hence, all the above - stressed
circumtances are enough to take into consideration the specific history of formation of this region’s flora, consequently,
sharply delimits it from the history of the Caucasus rest regions (Grossheim, 1926).
Like the Black Sea, the Caspian Sea is remnant of the ancient Sarmatian Sea. Till the Mid-Pliocene the Sarmatian Sea
filled large ancient basin, including Ponto-Caspian region, hense among other Caucasian islands, Talysh mountinous island
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emerged above surface of the mentioned ancient sea, and according to palaegeomorphological data, never submerged under
sea waters.This is one more additional fact why so exotic present-day Talysh refuge’s relict and endemic woody plants are
represented here since the Tertiary Period. Grossheim (1960), according to the mentioned circumstance, named local forests
,,Hyrcanian”, indicating Talysh’s phytocenosis age (The pre-Caspian Sea known as ancient Caspium Mare or Hyrcanian
Mare).
The Caspian Sea within its present borders has been formed in the second half of the Pleistocene (Atlas Lithologo –
palaegeographical maps of the USSR, 1967, v.IV, maps 1-55; Mamedov, 1997).
With respect to Talysh refuge, special attention should be paid to the present climatic condition. Azerbaijan is situated
between mountains system of the Greater and the Lesser Caucasus. Therefore, about 80% of its territory are isolated from
the climate influence of neighbouring regions, and only through the Caspian Sea’s basin this country falls into influence of
arid climate of Middle Asia. Despite the Caspian Sea’s (actually lake) site (37.000 sq.km and 78.200 cub.km), this natural
basin is unable to satiate with moisture extremely dry masses, dominating is endless continental, transcaspian extra-arid
deserts. The only exception is Talysh. According to physico-geographical peculiarities and convenient position of this
region and the Elburs mountains’ slopes toward sea, Talysh mid-altitudinal zone receives enough quantity of mean annual
precipitation, which ranges from 1000 to 1500 mm per year.
This fact that Tertiary forests of Talysh have survived owing to internal reasons, namely under influence of the
Caspian Sea’s deep waters, was underlined by Pastuchov (1926), Bush (1935) and other scholars. They considered that large
volume of waters in south-western part of the Caspian, performs role of reservoir to accumulate warm atmospheric masses
in summer, which in winter time supplies with warm and moist air the masses of the coastal zone, therefore improves the
hydrothermical conditions of the Hyrcanian refuge.
Finally, considerable role in formation of harbouring centre in both sides of Hyrcania (Azerbaijan, Iran), is performed
by the following circumstances: this region, unlike Colchis, remained ice – free refugia during all Pleistocene time including
Holocene (Grossheim, 1936; Sagheb-Talebi, 2000; Sagheb-Talebi et al., 2004; Ramezani et al., 2008; Ramezani, 2009b).
All the mentioned phenomena provided conditions for their own exclusive peculiarities to formation of flora and
vegetations of Talysh, and then transformated this region into a powerfull harbouring centre in Western Eurasia (Grossheim,
1936, 1948; Safarov, 1962, 1979; Zohary, 1963; Prilipko, 1980).
According to Grossheim (1926), the ancient Hyrcanian centre of vegetation has been formed and developed in parallel
with the Colchis (Pontic) one, without direct connection between two regions.
This ancient flora of South Europe, including the Talysh Range, was composed by three main groups: Arcto-Tertiary
or Turgai flora, then vegetations of South-Tertiary (occupies in ancient time south regions of Europe and Asia Interior),
and Hyrcanian, i.e. autochtonic flora of Hyrcan Botanical Province (Grossheim, 1926). The penetration of ArctoTertiarian’s origin taxa of the Miocene Age in Caucasus is confirmed by fossil remnants of Parrotia and Zelkova, found in
the Late Miocene deposits of West and Central Europe. For example, an extinct P. fagifolia is very close to the present-day
P. persica.
Despite the above - mentioned, comparing with the Tertiarian forests of some other modern refugia, the relict forests
of Talysh are rather strongly xerophytisized. The latter considerably attract actual lack of an evergreen underforest.
Present-day Talysh flora is represented with about 1750 species of vascular plants (Grossheim, 1926; Safarov, 1965).
The relict forests of Talysh are situate in the Caspian seacoastal lowland and mid-montains zone. These communities,
in general occupy an area from 400 to 600m a.s.l., but some representatives through rivers gorges reach till 1000-1200 m.
a.s.l.
According to Grossheim (1948), the Talysh forests enriched with relict vegetations, resemble another species rich
Colchis forests, but differ not only in quite different composition of woody species and their biology (e.g., double flowering,
cauliflory, two periods of growing of woody plants species, etc), but from phytocoenology point of view. Botanists in
unison emphasize absence of dense evergreen understorey within local forests. Evergreen shrubs sporadically occur only as
intrazonal formations (Grossheim, 1926; Safarov, 1962).
The woody stands of Talysh form two types of forest formations: represented by mixture of some species
(polydominant communities), and communities with predominance of one species (monodominant formations). Safarov
(1962) and Gulisashvili et al. (1975) outline several types of the Talysh relict forests.
Persian ironwood (Parrotia persica) forest. P. persica is single representative of Hamamelidaceae family in
Caucasus, relict of the ancient Arcto-Tertiary flora, as monotypic genera, survived to the present-day.The tree is a native to
the forest of South (Azerbaijan) and South-west (Iran) regions, adjacent to the Caspian Sea.
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Thus, North Iran and Talysh, united in common Hyrcanian Floristic Province, presented a single region of Earth,
where have been preserved forest communities with iron tree of the Tertiary Period (Safarov, 1962, 1972). The closest
relatives of iron tree are distributed in China, the Hymalays and North America.
According to Amdjadi (1958; in Safarov, 1962,1972; Djavanshir, 1976; Sagheb-Talebi, 2000 and Remezani, 2009a),
an iron tree within Iran’s part of the area (south border of distribution of P.persica) appears in mountains till 1000-1400 m
a.s.l., and occurs in formations of beech- hornbeam forests. Thus, as Safarov indicates, from south to northward there takes
place not only quantative decrease of an iron tree within the Hyrcanian forests’ composition, but a considerable decrease of
upper limit of the altitudinal distribution of these species. It can be explained by deterioration of abiotic factors – soil and
climatic conditions in unison of appearing P. persica to the North border of the Hyrcanian Province.
Sometimes P.persica coinsides with native oak - Quercus castaneifolia (ironwood - an oak mixture forest), and with
an oak and Caucasian hornbeam (Carpinus caucasica). The first stratum in such forests is occupied by chestnut leaved oak,
hence only the second stratum of canopy is build up by ironwood.
Arboreal plant layers are composed by several species of Hyrcanian stock. Native hardwood forest consist of the
following species: Quercus castaneifolia, Parrotia persica, Albizzia julibrissin, Zelcova hyrcana, Z.carpinifolia, Fraxinus
coriariafolia, Ficus hyrcana, Diospyros lotus, in lowland communities participate some other trees: Ulmus carpinifolia,
Populus hyrcana and Alnus barbata.
Forests with persian silk acacia (Albizzia julibrissin). A. julibrissin is ancient relict of eastern Asia’s tropicalsubtropical flora. Nowadays silk acacia is characterized by its two ,,islands location”. The first, much largest is JapaneseChinese (Japan, Korea, China), occurs in South-East Asia, the second, particularly smaller – Iranian-Talysh (Hyrcanian) is
situated in western – caspiancoastal region. These two areas are disjunct from each other by huge Afghanistan-Iranian
territory.
A. julibrissin in Elburz covers slopes directed to the Caspian Sea and occupies a narrow stripe in low mountains zone.
In northernward it penetrates within Talysh part of common range (38-40km). In northern part of its areas, silk acacia,
simultaneously with climate deterioration, assumes a new cold-resistance, drought-tolerant features. In Azerbaijan A.
julibrissin extends from the sea level up to 300-400 m a.s.l.
The most vivid example of relict and at the same time of endemic species is manifested by Brahmaea christophi, one
of the most beautiful moths in invertebrate fauna of the Caucasus.
B. christophi resembles much Colchian vicarious species – (B. Ledereri), but is slightly larger (the wing-span reaches
100-130mm). B. christophi covers whole Hyrcanian area, including Iranian Azerbaijan (Gorgan, Mazenderan, Gilian).
Belonging to the relicts of immemorial time, B. christophi is close to extinction (Aliev, Kasparov (eds), 1996).
The following relict and endemic species is presented by Isophya caspica (ordo Othoptera). It is a typical mesic insect
restricted by the Tertiarian forests. The closer relative species, as it was emphasized, endemic of Colchis refuge is I.
redtenbacheri (Bey-Bienko, 1958).
O. Krizhanowski and M. Ter-Minasian (1958) indicate a rather long list of beetles (ordo Coleoptera) species with
restricted area in the Talysh and adjacent territory of Iran, the so - called “Hyrcanian species”. Among probable ancient
species, mentioned scholars considerably stressed Parandra caspia, undoubtedly the Tertiarian relict, all close species of
which are represented out of Palaearctic region including Madagascar. This beetle is one of the most primitive species in its
genus. P.caspia populates various broad-leaved trees. In Talysh this wood-borer beetle is considered as one of the most
serious pest of the wood.
According to Vereshagin (1959), tiger (Panthera tigris) migrated from south-eastern Asia to north-westernward,
probably, only in the late Holocene. East Transcaucasia till very recent time was the extreme western site of the tiger’s
range. In the 60s of the 19th century the Turanian subspecies of tiger (P.t. virgata) was still usual beast in Talysh, where
every year were killed several beasts (Radde, 1899). Then, to the end of the 90s of the same century, tiger was found
frequently enough both in Lenkoran Lowland, and in mountainous forests of Talysh. To the end of the XIX century annually
still was killed even one beast (Satunin, 1915, 1920), but in the beginning of the XX century due to permanent impact of
man (hunting), and hunting of boars – the main food of tigers, anthopogenic influence of forests and poaching caused a
sharp decrease of tiger’s population. From the 20s of the 20th century tigers have come in eastern Transcaucasia from the
forests of south seacoastal zone of the Caspian Sea (Iran’s Azerbaijan) (Firouz, 2005), but yet never settled north part of
Hyrcania. From 1920 and 1950 there were registered two occurrences of tiger’s hunting, one in 1922 near Tbilisi (village
Lelobi) and the second, in 1932, in Talysh. In East Georgia tiger occured through the r. Kura (Mtkvari) riverside forests,
hunting on boars.
With respect to leopard, a small population of beast saxicolor inhabits in the Greater Caucasus Range and some
regions of South Caucasus.
6
Indian crested porcupine (Hystrix indica) is the next “Indian” species. In the beginning of the 20th century animal
occupied only Talysh and Lenkoran Lowland (Satunin, 1915, 1920), in 1923, the animal was found in Armenia (v. Megri,
mid-Aras current). By the end the of 40s, it appeared in Karabakh, in the 60s dwelled in western Azerbaijan, not far from
East Georgia, in 1990 – 2002 animal expanded its area in arid landscapes of East Georgia (vicinity of the rivers Iori and
Alazani), (Bukhnikashvili, Chkhikvadze, 2004). Thus, through the next 80 years it had spread from Lenkoran district of
Azerbaijan to East Georgia (approximately 700-800 km).
Thanks to direct connections with countries, represented in tropical-subtropical latitudes, specific climatic,
orographical and vegetation peculiarities, the Talysh refuge remained in Caucasus troublefree attainable shelter for some
exotic animals till recent time.
The homogenic synecological formations are found not between South Caucasus two refugional centres, from this
viewpoint, as it was emphasized above, Colchis has closer relations with Atlantic Europe and Mediterranean countries, East
and South - eastern Asia, where closely related complex of species occur on disjunctive habitats, than with Hyrcania (Wolfe,
1944; Walter, 1974; Dolukhanov, 1980; Gagnidze, 1999; Kikvidze and Ohsava, 2001, etc.).
Taking into account the above characterized peculiarities of Talysh’s biocoenoses enriched with Arcto-Tertiarian
and tropical-subtropical elements, it should be underlined that all these elements give quite incomporable appearance to the
forest ecosystem of Hyrcania enveloping Talysh as a north outpost of the Elburs’ Range (North Iran).
It’s natural, that so striking features of Iranian part of the Hyrcanian refuge is correspondingly attractive to the animals
biologically bounded with the Hyrcanian Province’s Phytocenoses. That’s why this south part of refuge, for so long time,
performs a role of donor to guaranteeing Talysh region to receive Tertiarian and tropical-subtropical post-glacial elements
during either climatic deterioration within refuge or human’s threat to so exotic for Caucasus animal species.
The Altitudinal Zonation of the Colchian Living Organisms
Colchian type of vertical distribution of vegetation is discussed by some botanists: Ketschoveli (1960), Gagnidze
(1999, 2000), Nakhutsrishvili (1999), Volodicheva (2003).
1. Colchis deciduous, mixed forests (0-500m)
2. Deciduous (chestnut) forests (500-1000m)
3. Deciduous (beech) forests (1000-1500m)
4. Dark coniferous forests (1500-2000m)
5. Subalpine forests–meadow vegetation, (2000-2500m)
6. Alpine (2500-3500; 3800m)
7. Nival zone (3500-3800m and up)
Fig. 3. The scheme of the Colchian or Eastern
Euxinian type of altitudinal zonality
According to the above - mentioned botanists, the Colchian type of altitudinal zonality characterized by development
of hardwood forests (mixed-leaved, deciduous broad-leaved) from the Black seacoast area to 2000-2200 m. From the very
sea level to 500 m a.s.l., large territory is populated by mixed (polydominant) forests. The tree layer of forests is composed
by species of an oak, chestnut, alder, wing-nut, hornbeam, etc. On the vast territory there are extended numerous wetlands
and peat bogs.
Forests, occuping ranges from 500-1000m are composed by chiefly chestnut with admixture of beech, georgian oak,
hornbeam, lime-tree, etc. Beach forests vary mainly in altitudes between 1000 and 1500m. Within the mentioned
communities a considerable part is played by evergreen undergrowth.
The distribution range of dark coniferous forests with participatation of spruce (Picea orientalis) and fir (Abies
nordmanniana) trees, comprises 1500m and 2000m a.s.l., underforest is still presented by mainly evergreen shrubs and low
trees.
Apart from the mentioned type of coniferous communities, in Colchis there occur poor, sometimes well - developed
stands of light coniferous forests with participation of mountains pine tree (Pinus kolchiana).
7
Subalpine belt occupies also rather large areas from 2000m to 2500m. The herbaceous stratum is situated by
subalpine forb meadows and tall herbaceous vegetation; arboreal plants (subalpine forests) are manifested by crook-stem
and park like forests, including beech, birches, native maples, ashes, etc. Participation of the low scrub communities in
highland is insignificant with dekiani (Rhododendron caucasicum).
The next higher altitudes are occupied by an alpine belt (2500-3500m), and sometimes by more elevation with short
grass meadows and forbs, the climax carpet-like alpine meadows, and alpine low and dwarf scrubs (dekiani, species of
bilberries).
To the most striking representatives of fauna belongs: in lowlands Colchian pheasant (Phasianus colchicus), in
highlands – endemic wild goat (Capra caucasica), endemic snowcock (Tetraogallus caucasicus), endemic Caucasian
grouse (Tetrao (Lyrulus) mlokosiewiczi), endemic and relict of the Ice Age Nordmann’s apollo (Parnassius nordmanni) –
one of the most beautiful butteflies of Caucasus.
The specificity of sequence of Colchis
altitudinal zonation
Legend: AT- Arcto-Tertiary; B- Boreal; D Desert and semi-desert;
M - Mediterrannean;
T - Tropical; TS - Tropical-subtropical; ST - SouthTertiary
Fig. 4. Composition of living organisms via colchis vertical
zonation concerning their landscape – genetical belonging
Symbolic designation of various life forms of plants
The Altitudinal Zonation of the Hyrcanian Living Organisms
The Talysh’s vegetational cover with its specific animal communities is characterized by peculiar vertical zonation
The same zonation in Iranian Talysh was studied by Ramezani (2009).
Safarov (1962) outlines that Azerbaijan is the only country in Caucasus, where vertical zonation begins under the sea
level and is represented from 28m below the ocean level in coastal regions to about 4000m a.s.l.
The Talysh type of altitudinal zonality distinguishes by extraordinary structure and is represented by five nature belts
(Grossheim, 1926; Safarov, 1962; Volodicheva, 2003).
Forests of the lowland (- 28m to 50 m a.s.l.). With respect to the forest communities of lowland, these formations
today due to their turning into cultural vegetations, lost their landscape mean (Safarov, 1962). Some fragments of these
forests, as it is mentioned above, can be found in the Hyrcanian lowland’s reserve (covers about 100ha of the area).
According to Grossheim (1926), the primary type of the lowland’s forest is composed by Persian ironwood.
Destruction of Hyrcanian forests is the effect of man’s agricultural activities. Lowland’s communities are distinguished by
presence of marshy forest, not occuring in other region of forested Talysh. In marshy habitats alder occupies a dominant
position among woody plants (Safarov, 1962).
1. Forest of the lowland (- 28m to 50m)
2. Forest of the foothills and lower mountain belt (50 – 600m)
3. Forest of mid-mountain belt(600-1500m)
4. Forest of mountain belt 1500-2200m)
5.The vegetation of mountainous woodless Talysh (Zuvand),
(1800-2500m)
8
Figure 5. The scheme of Hyrcanian
type of altitudinal zonality
Within forests, covering foothills and lower mountain belt (50 – 600m a.s.l.) an edificatory position is occupied by
native oak - Quercus castaneifolia and Persian ironwood -Parrotia persica. These typical for Hyrcanian forest communities
are presented by either monodominant or mixed phytocoenosis. The most striking feature of local forests, except ironwood,
give the grows including Gleditsia caspia and Albizzia julibrissin.
It must be stated that just mentioned two belts are populated by typical Hyrcanian relicts and sometimes relicts and
endemic species of animals (species of Brahmaea, Danais, Isophya, porcupine, in recent time – Turanian tiger, etc.
Forests of mid-mountain belt’s (600-1500m a.s.l.) major forest-forming species on the north slopes occurs beech
(Fagus orientalis), on the south-east slopes - Quercus castaneifolia and Carpinus caucasica, on the south slopes – mainly
Zelkova caprinifolia, accompanied by corresponding herbaceous vegetation. Moist gorges are populated by Pterocaria
pterocarpa.
In upper mountain forest belt’s (1500-2200m a.s.l.), as a leading species, is found Quercus macranthera with
admixture of Carpinus schuschaensis. Within the mentioned belt Arcto-Tertiarian communities of previous belt are replaced
with another type of forest’s formations, which in structural and in floristical viewpoint is closer to montionous forests of
South Europe (Safarov, 1962).
The vegetation of mountainous woodless Talysh (1800-2500m), Diabar depression (part of arid Zuvand), as it was
pointed out in part . . , forest communities completely disappear, instead participations of the mountainous steppe,
mountainous xerophyle (particularly tragacant cushion shrub formations) are insignificant (Grossheim, 1926).
Physical-geographycal location and an extraordinary type of altitudinal zonation of Talysh, unequal by provides
with precipitation, determines some negative features of this refugial centre.
The hardwood forests of Talysh-subcaspian slopes (Elburs in North Iran) are absolutely devoid of broad-leaved
forests composed by chestnut (Castanea sativa), which forms a remarkable natural belt in mountains of Colchis and some
other regions of Transcaucasia.
Vertical zonation of Hyrcanian type is missing needleleaf (Gymnosperm) forests composed by species of spruce, fir
and pine trees (the upper zone manifested only by species of Junipe). As was indicated above, requiring wet climatic
conditions darkconiferous species – Picea orientalis and Abies nordmaniana, seldom light-consumer Pinus kochiana give
some perfect appearance to Colchis vertical zonations.
The most striking feature of so impressive Arcto-Tertiarian forests of Hyrcania is impoverishment with underwood.
To quote Grossheim’s words (1926): ,,There is no evergreen undergrowth, in a proper sense of the term, in the forests of
Talysh”.
The Hyrcanian underforest is characterized by complete absence of genus Rhododendron, where in the western part of
the Caucasus they are manifested by five - six species. In Hyrcan Province there occur Laurocerasus officinalis, Vaccinium
arctostaphylos, Ilex hyrcana, but they, as a rule, are presented as a poor shrub stand.
All above emphasized natural phenomena, as Grossheim (1926) considers, are connected with the history of
Hyrcanian flora and once more underlined a distinctive trend of development of native vegetation cover.
Symbolic designation of woody and semi-woody plants
The specificity of sequence of Talysh altitudinal
zonation
Legend: AT - Arcto-Tertiary; I T (AI) – Interior
Asian – Turanian (Armenian-Iranian); ST – SouthTertiary; T - Tropical; TS - Tropical – subtropical
9
Figure 6. Composition of Living Organisms Through Talysh
Altitudinal Zonation with Respect to theirs Landscape – Genetical Belonging
References
Adamia Sh.A. et al., 1977. Evolution of the ancient continental margin by the example of Alpine history of the Caucasus.
Geotekhtonika, 88-103 (In Russian).
Adamia Sh.A. et al., 1991. Geological background of the Caucasus and adjacent area. Geological events of CretaceousPaleogene boundary. In: Proceed. of Intern. Sympos., Tbilisi. 12, 18-5.
Albov N., 1896. Essay of vegetation of Colchic. In: Zemlevedenie, I: 49-74 (in Russian).
Aliev S.V., Kasparov A.G., (eds.), 1996. Animal world of Azerbaijan. V.I. Invertebrate fauna. Elm,
Baku (in Russian).
Amdjadi, H., 1958. Climat general et types des forests de Iran. Bull. Soc. Roy. Forest Belgique, 65.
Antonov B.A., 1955. Geomorphology of the Lenkoran district (Talysh). Bull. Comission po izuchen. Chetvertichn. Perioda
AN SSSR, 20:71-82.
Atlas of lithologo–palaeogeographical maps of the USSR, 1967, v. IV, Moskva; maps: 1-55 (in Russian).
Bey-Bienko, 1958. Orthoptera and Dermaptera. In: Zhivotnui Mir SSSR, t.V: 435-456. Izdatelstvo AN SSSR, MoskvaLeningrad (in Russian).
Bonacina, A. 1980. Sistematica specifica e sottospecifica del complesso ,,Euscorpius germanus” (Scorpiones, Chalctidae).
Riv. Mus. Scien. Natur. E.Caffi, Bergamo, 2: 47-100.
Bukhnikashvili A, Chkhikvadze V., 2004. Indian crested porcupine (Hystrix indica) in Georgia. Bulleten of the Georgian
Academy of Science, 169, 1: 166-168.
Bush N.A., 1935. Botanical and geographical studies of Caucasus. Izdatelstvo AN SSSR, Moskva-Leningrad (in Russian).
Chelidze L.T., Kvavadze E.V., 1987. Fossil vegetation of Meotical deposits of Abkhazia. Soobsh. AN GSSR,
122, 1: 177-180 (in Russian).
Chubukhov L.A., 1966. The Caucasus. Nauka, Moskva (in Russian).
Davis P.H.,(Ed.)1965-1988. Flora of Turkey and the East Aegean Islands. Vols 1-9. Edinburgh University Press,
Edinburgh.
Didmanidze E.A., 2005. The butterflies of Georgia. Studia 99, Tbilisi, (In English and in Georgian).
Djavanshir K., 1976. Atlas of woody plants of Iran. Teheran (Nation. Soc. for the Conserv. of Natural Res. and Human
Environm.).
Dmitrieva A.A., 1990. The key to the vegetation of Adjaria, v. I,II, Metsniereba, Tbilisi (in Russian).
Dolukhanov A.G., 1974. The subalpine landscapes of Caucasus as a shelter of relict elements of flora. In: Rastitelni mir
visokogorii i ego osvoenie. Probl. Botaniki, t.12: 27-34, Nauka, Leningrad (in Russian).
Dolukhanov A.G., 1980. The Colchian underforest. Metsniereba, Tbilisi (in Russian).
Dolukhanov A.G., 1989. Forest vegetation of Georgia. In vegetation of Georgia, v.1. Metsniereba, Tbilisi (in Russian).
Fet V.,1993. Notes of Euscorpius mingrelicus (Kessler, 1874) (Scorpiones: Chactidae) from the Caucasus. Riv. Mus. Civ.
Sc. Nat. ,,E. Caffi” Bergamo, 16:1-8.
Firouz E., 2005. The complete fauna of Iran. Tauris & Company.
Gagnidze R.I., 1974. Botanical and geographical analysis of the florocoenotic complex of tall herbaceous
vegetation of the Caucasus. Metsniereba, Tbilisi (in Russian).
Gagnidze R.I., 1999. Arealogical review of Colchic evergreen broad-leaved mesophyllous dendroflora species. In: Klötzli
F., Walther G.-R. (Eds.): Conference of recent shifts in vegetation boundaries of deciduous forests, Forests especially
due to general Global warming. Birkhäuser Verlag, Basel-Boston-Berlin. 199-216.
Gagnidze R.I., Davitadze M.D., 2000. Indigenous flora (Vegetation of Georgia): ,,Adjara”. Batumi (in Georgian).
Gagnidze R.I. et al, 2002. Endemic genera of the Caucasian flora. Feddes Repectorium, 113, 7-8: 616-630.
Gajiev G.V., Kakhmatulina I.K. (eds), 2000. Animal World of Azerbaijan. Elm. Baku (in Russian).
Gamkrelidze I.P., 1986. Geodynamic evolution of the Caucasus and adjacent areas in Alpine time. Tectonophysics, 127:
261-277.
Gamkrelidze I.P., 1997. Main trend of tectonic development of Caucasus and adjacent areas of the Mediterranean belt.
Himalayan geology, v. 7: 88-96.
Gegechkori A.M., 1977. Psyllids (Homoptera, Psylloidea) of Talysh. Soobch. AN Gruz. SSR, 86, 1: 197-200 (in Russian).
Gegechkori A.M., 1978. Psyllids of mid-current part of r. Aras. Metsniereba, Tbilisi (In Russian with English
summary).
Gegechkori A.M., 1984a. Psyllids of the Caucasus. Metsniereba, Tbilisi (In Russian).
Gegechkori A.M., 1984b. A species composition and origin of psyllids (Homoptera, Psylloidea) of the Caucasus.
Avtoreferat na soiskanie dissertacii uchenoi stepeni doktora biologicheskich nauk. Leningrad (in Russian).
Gegechkori A.M., 1985. Some aspects of evolution of psyllids. Metsniereba, Tbilisi (in Russian, with English summary and
contents).
10
Gegechkori A.M., 1991. The role of Middle Europe in the formation of psyllid fauna (Homoptera, psylloidea) of
the Caucasus. XII intern. Middle Europe conf., Kiev; 531-534 (in Russian).
Gegechkori A.M., 1997. The ecological-geographical peculiarities of the psyllid (Homoptera, Psylloidea) fauna of the
Caucasus. Meridiani, Tbilisi (in Russian with English summary).
Gegechkori A.M., 2000. Relicts and endemics and Georgia’s biodiversity at the background of Caucasus orogenesis. In:
Biological and Landscapes Biodiversity of Georgia. WWF Georgia, Tbilisi, 83-96.
Gegechkori A.M., Djibladze D.S., 1976. Psyllids (Homoptera, Psylloidea) of Colchis. Metsniereba, Tbilisi (in Russian with
English summary).
Gegechkori A.M., Loginova M.M., 1990. Psyllids (Homoptera, psylloidea) of the USSR (an annotated checklist), Metsniereba, Tbilisi (in Russian with English summary).
Graham A., ed (1972). Floristics and paleofloristics of Asia and Eastern North America. Elsevier, Amsterdam.
Gray A., 1846. Analogy between the flora of Japan and that of the United States. American Journal of Science
and Arts (series 2), 2: 175-176.
Grossheim A.A. 1926. Flora of Talysh. Izdatelstvo NKZ of Azerb. SSR, Baku (in Russian).
Grossheim A.A. 1936. The analysis of Caucasian flora. Trudy Bot. in-ta Azerbaidzh. Fil.AN SSSR. Izd. Az. Fil. AN SSSR,
Baku (in Russian, with English summary).
Grossheim A.A. 1938. Relicts of East Transcaucasia. Problemi reliktov vo flore SSR, I. Izd. Bot. inst-ta AN SSSR: 201-211
(in Russian).
Grossheim A.A. 1948. Vegetational cover of the Caucasus. MOIP, Moskva (in Russian).
Gueguechkori A.M., 1998. Genese de la faune des deserts L’example des psyllides et des desert de l’ancien
monde. In: La Geographie en Georgie, Orstom, Paris: 103-115.
Gulisashvili V.Z., Makhatadze L.B., Prilipko L.I., 1975. Vegetation cover of Caucasus. Nauka, Moskva (in Russian).
Guo Q. –E., 1999. Ecological comparisons between eastern Asia and North America: historical and geographical
perspectives, Journal of Biogeography, 26: 199-206.
Guo Q. –E., Ricklefs R.E., Cody M.L., 1998. Vascular plant diversity in eastern Asia and North America: historical and
ecological explanations. Journal of the Linnean Society, 128: 123-136.
Gvozdetsky N.A., 1958. The physical geography of Caucasus. Izdatelstvo Moskovskogo Universiteta, Moskva (in Russian).
Hersey R.E., van der Kloet S.P., 1967. Taxonomy and distribution of Gaultheria in the Caribbean. Canadian Journal of
Botany, 54, 21: 2465-2472.
Hodkinson I.D., 1974. The biology of the Psylloidea (Homoptera). A review, bull. ent., Res., 64: 325-339.
Hodkinson I.D., 2009. Life cicle variation and adaptation in jumping plant lice (Inseccta: Hemiptera, Psylloidea); a global
syntyhesis, journal of Natural History, vol. 43 (1&2), 65-179.
Ilinskaya I.A., 1959. The Upper Miocene flora of the mountain Povitrule in Transcarpath. In: Botan. Zhurnal,
XLIV, 5: 121-125, Moskva-Leningrad (in Russian).
Ketschoveli N.N., 1960. Vegetation cover of Georgia. Metsniereba. Tbilisi (in Georgian).
Khain V.E., 1975. Structure and main stages in the tectono – magmatic development of the Caucasus: an attempt at
Geodynamic interpretation. Americal Journ. of Sc., 275:131-156.
Kharadze A.L., Gagnidze R.I., 1970. Review of hemixerophilic endemic elements in the flora of the
Novorossiysk sub-province of Caucasus. Not. Syst. Geogr. Inst. Bot. Tbilisi, 28: 56-82 (in Russian).
Kikvidze Z., Ohsava M., 2001. Richness of Colchic vegetation: comparison between refugia of south-western and East
Asia. BMC Ecology, 116: 1-10.
Kolakowsky A.A., 1961. The flora and vegetation of Colchis. MOIP, otd. Bot, 10, XVIII, MGU, Moskva (in Russian).
Kryzhanovski O.L., Ter-Minasian M.E., 1958. Coleoptera of the Caucasus. In: Zhivotni mir SSSR, t.V: 385-431 (in
Russian).
Li H.–L., 1952. Floristic relationships between eastern Asia and eastern North America. Transitions of the
American Philosophical Society (New Series), 42:371-429.
Li H.–L., 1972. Eastern Asia – North America species pairs in wide-ranging genera. Floristics and paleofloristics of Asia
and North America (ed. By A. Graham), Elsevier, New York: 65-78.
Loginova M.M., 1968. New data conserning to fauna and biology of psyllids (Psylloidea, Hemiptera) of Caucasus. Trudy
Vsesoiuzn. Entomol. Obshestva, Leningrad: 275-328 (in Russian).
Makharadze N.V., 1998. The vegetation and climate development of Meotion time of West Georgia. PhD thesis, preprint,
Tbilisi (in Georgian and Russian).
Maleev V.P., 1940. The vegetation of the Black Sea coastal countries in the Euxinean Province of Mediterranean region, its
origin and connection. Trudy Botanicheskogo Instituta Akademii Nauk SSSR, Ser.-Geobotanica,
Maleev V.P., 1941. The Tertiary relicts in the flora of Western Caucasus and main stages of the Quaternary history of its
flora and vegetation. Materialy po istorii Flori i Rastitelnosti SSSR, 1: 61-144 (in Russian).
Mamedov A.V., 1997. The late Pleistocene – Holocene history of the Caspian Sea. Quaternary International. Vols
41/42:161-166.
11
Mandjavidze D.V., 1967. A review of historical development of flora and vegetations of Ajaria in connection of geological
past of Caucasus. In: Trudy Batumskogo Botan. Sada; Flora and Vegetation of Adjaria, 14:16-21 (in Russian).
Mandjavidze D.V., 1982. Relict forests of Adjaria and theirs importance as a timbering materials. Metsniereba, Tbilisi (in
Russian).
Milanovsky E.E., 1977. Neotectonic and modern volcanism of Caucasus. In: Geologia Chetvertichnogo Perioda: 35-43,
Erevan (in Russian).
Nakhutsrishvili G., 1999. The vegetation of Georgia. Braun-Blanquetia, vol. 15 (Camerino).
Pastukhov N.K., 1926. Review of the nature of Talysh. (Trudy Tiflissk. Bot. sada. Ser. 2, 4:33-39 (in Russian).
Prilipko L.I., 1980.: Hyrcanic broad-leaved forests. In: Vegetation of the European part of the USSR. Nauka: 192-196 (in
Russian).
Radde G.I., 1899. Collection of the Caucasian Museum, I. Tiflis (in Russian).
Ramezani E., Mohadjer M.R.M., Knapp H.-D., Ahmadi H., Joosten H., 2008.The Late-Holocene vegetation history of the
Central Caspian (Hyrcanian) forests of Northern Iran. The Holocene, 18, 2:307-321.
Ramezani E., 2009a. Vegetation zones along an altitudinal transect in Central Caspian forest of northern Iran (unpublished
data).
Ramezani E., 2009b. The Holocene development of the Caspian forests. PhD thesis. University of Tehran.
Raven P.H., 1972. Plant species distributions: a summary. Annals of the Missouri Bot. Gard., 59:234-246.
Rijabov M.A., 1958. Lepidoptera of Caucasus. In: Zhivotni mir Kavkaza, t.V: 351-375. Izdatelstvo Akademii Nauk SSSR,
Moskva-Leningrad (in Russian).
Qian H., 2001. A comparison of generic endemism of vascular plants between East Asia and North America. International
Journal of Plant Science, 162:191-199.
Sagheb – Talebi K., 2000. Hyrcanian Forests (North of Iran), the unique ecosystem in Near East Region. XXI IUFRO
World Congress – Forests and Society: the role of research (1-12.08), Kuala Lumpur, Malaysia: 1-8.
Sagheb- Talebi K., Sajedi T., Yazdin F., 2004. Forests of Iran. Research Institute of Forests and Rangelands.
Safarov I.S., 1962. The most important arboreal Tertiarian relicts of Azerbaijan. Izdat. AN Azerb. SSR, Baku, (in Russian).
Safarov I.S., 1972. Ironwood Parrotia persica C.A.M. History, geography, taxonomy and bioecological peculiarities. Bot.
Zh.,57:932-944 (in Russian).
Safarov I.S.,1979. The subtropic forests of Talysh. AN Azerb. SSR., Inst.Bot. im.V. L. Komarova, ELM, Baku.
Satunin K.A., 1915-1920. Mammalians of the Caucasus Region. T. 1-2, Tiflis (in Russian).
Shatilova I.I., Ramishvili I. Sh., 1990. The materials of the history of flora and vegetation of Georgia. Metsniereba, Tbilisi
(in Russian).
Schmidt P., 2004. Bäume und Sträucher Kaukasiens. Teil 3: Laubgehölze der Familien Ebenaceae (Ebenholzgewächse) bis
Frankeniaceae (Frankeniengewächse). Mitt. Dtsch. Dendrol. Ges., 89: 49-71.
Takhtajan A.L., 1941. Botanical-geographical essay of Armenia. Trudy BIN AN Arm. SSR, 4:49-61 (in Russian).
Takhtajan A.L., 1997. Diversity and classification of flowering plants. Columbia University Press. New York,
New York, USA.
Thorne R.F., 1972. Major disjunctions in the geographical ranges of seed plants. Quart. Rev. Biol., 47:365-411.
Vereshagin N.K., 1958. Mammalians of the Caucasus. In: Zhivotni mir SSSR, t. V: 180-219. Izdatelstvo AN SSSR,
Moskva-Leningrad (in Russian).
Vereshagin N.K., 1959. Mammalians of the Caucasus. The history of formation of fauna. In: Zhivotni mir SSSR,
t.V: Izdatelstvo AN SSSR, Moskva-Leningrad (in Russian).
Volodicheva N., 2003. The Caucasus. In: The Physical geography of northern Eurasia. Oxford Regional
Environment. Oxford Univ. Press, Oxford.
Walter H., 1968. Die Vegetation der Erde. V.I. Progress, Moskva (translation in Russian).
Walter H., 1974. Die Vegetation der Erde. V.II. Progress. Moskva (translation in Russian).
Wolfe E.V., 1944. The historical geography of vegetation. Izdatelstvo AN SSSR, Moskva-Leningrad (in Russian).
Wood C.E., 1961. The genera of Ericaceae in the southeastern United States. Journal of the Arnold Arboretum,
42:10-80.
Zazanashvili N., 1999. On the Colkhic vegetation. In: Klötzli F., Walther G. –R. (Eds.): Conference of recent
shifts in vegetation boundaries of deciduous forests, forests especially due to general Global warming.
Birkhäuser Verlag, Basel-Boston-Berlin.
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