ANNALS OF AGRARIAN SCIENCE, vol. 9, no. 1, 2011 ИЗВЕСТИЯ АГРАРНОЙ НАУКИ, Том 9, Ном. 1, 2011
AGRONOMY AND AGROECOLOGY АГРОНОМИЯ И АГРОЭКОЛОГИЯ
THE RESULTS OF BIOGEOGRAPHICAL STUDY OF THE ARCTOARCTO TERTIARYTERTIARY REFUGIA (COLCHIS AND TALYSH) OF SOUTHERNSOUTHERN CAUCASUS
A.M.Gegechkori Ivane Javalhishvili Tbilisi State UniversiUniversitytytyty,, Department of Biodiversity 3, Ilia Chavchavadze Ave.,Tbilisi, 0128, Georgia; [email protected] Received: 17.11.10; accepted: 09.01.11
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 Euroasia. 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 investigation. 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.
OBJECTIVES 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 [1 3]. 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 natural historical regions of the Caucasus (see schematic maps 1,2.), 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 Far East (1981 1991), in Asia Minor (1995 1998, 2006), and in other parts of the world (1986, 1989, 2000 2003). The results are presented in various monographs and articles [2 9].
1
Sch. map 1.
Sch. map 2.
As it is broadly known, refugia are the regions of the Earth, where living organisms endured severe climatic and orographic fluctuations and as a relict forms are still survived. Hence, these shelters of the archaic organisms consist of useful data for solving important problems of biogeography, evolutionary biology, taxons genesis, etc. Each refugia of the Earth have their specific historical, geographical and abiotic peculiarities by what they in historical past turn into shelters for ancient organisms. For instance, how does California preserve Mediterranean proper maquis communities, known today as a Madro Tertiarian flora, or New Zealand keeps a Mesozoic Antarctic’s Gondwanian formations with species of tree ferns, south beeches ( Nothofagus spp.), south conifers ( Podocarpus spp.), etc? The reasons are different. Their own historical and palaeecological geographical reasons have Transcaucasia’s two famous refugia – Colchis and Talysh with present day communities, for which not seldom is used the term – “subtropic”. In some regions of the Caucasus, namely, in Colchis, Talysh, and partially in river Alazani valley (Kakhetia, East Georgia), owing the warm and moist climate survived considerable number of species – the representatives of ancient Tertiary’s origin plants and animals. Among the Caucasus biomes, the floristic and faunistic diversity and infrequency (relict and endemic species) of forests of mentioned two refugia, are one of the most striking feature of this region. With respect to vegetations of this region this phenomenon have long been recognized, intensively studied and discussed [10 20]; the same interests have increasingly fascinated regarding to fauna [21 26,6]. What do we consider under Tertiary relicts in South Caucasus? In moderate climate belt of the Northern Hemisphere is represented broad leaved deciduous (summer green) forest’s zone (biome). In biogeographical literature it is determined as a Temperate Broad Leaf Deciduous Forest (TBDF). By geobotanists this flora are proposed as the Arcto Tertiary Geoflora (ATG) [27 30]. In Russian language literature it is indicated as a Turgayan Flora [31]. This term derives from Turgay Hollow (Kazakhstan), where in 1929 Russian paleobotanists A. Kryshtofovich [31] and then S. Zhilin [32] from Oligocene deposits has described the most characteristic fossil remains of TBDF complex: chestnut, beach, zelcova, walnut, hazelnut, etc.; from gymnosperms to mentioned complex belongs Sequoia , Metasequoia , bog’s cypress, etc. Sometimes TBDF complex mentioned as a Nemoral Flora (the term nemoralis – means “forest”) [33,34,26,7]. Broad leaved deciduous forest of a particular floristic composition evolved in the Rocky Mountains [29] during Eocene time, and lately in Himalayas [35], persisted there during the Early Tertiary and, in response to gradually cooling climate, migrated southward during mid Tertiary into the middle latitudes, where their most large territories of these forests persist today in east central Asia and south eastern North America [35,36]. In the Oligocene Epoch this ancient mesic woody flora was represented as a circumpolar distributed biome. Originated within depth of subtropical evergreen moist forests, the Arcto Tertiary’s forests woody elements adaptation to initial more or less clearly expressed global cooling climate were foliage drop in coldest season of the year. About 15 million years ago, climatic cooling took place due to northward drift of both northern continents after break up of Laurasia in the Cretaceous Period. Cooling apparently let the sorting out of the gymnosperms from the angiosperms. The former one became concentrated
3 in higher latitudes as a boreal (taiga) forests [37]; Angiosperms, preferring a milder climate, became the TBDF. These forests were traditionally compared with forests of the Tertiary Period [38]. As it was mentioned above, ATG was supposed to have been widespred from higher latitude throughout the Northern Hemisphere [39,35,36]. Chaney [39] believed that severe climate during the Quarternary Period eliminated the ATG entirely from many region of the Northern Hemisphere. But following to J.A. Wolfe [40], changes of the Late Tertiary and Quartenary climate caused local adaptations rather than migrations of intact plant comities from one latitude to another as Chaney hypothesized. More, the same author challenged the geoflora concept and argued that a uniform broad leaved forest never existed. His analysis of the paleobotanical data shows that Tertiary floras were diverse, with evergreen gymnosperms such as Sequoia dominant, and evergreen angiosperms present elsewhere. Continental drift of Laurasia’s megacontinent has caused the separation of North America from Euroasia [41]. This separation of two continents’ nemoral flora in the most impressive way attracts Arcto Tertiarian floras complexes of eastern Asia and North America. This disjunction in the TBDF, leads to the contemporary configuration of distribution pattern of the biome. The close floristic, partially faunistic relationship between the both regions has long been recognized and discussed [42 45]. Mentioned and other macro disjunction areas have been triggered the origination of new direction in biogeography – vicariance (mobilistic) biogeography. In western Euroasia, a species poor broad leaved forest reflects widespread extinctions during the Pleistocenes Ice Ages. At present, the nemoral forest (sensu lato) consist of all those biocenoses which in the Northern Hemisphere are occupying a broad leaved forest biome (an indicator of this biome is considered deciduous species of oaks). Within this communities participate some warm climate loving conifers – tsuga, yew tree, etc. [31,46,36,29,30]. With respect to the Turgai fauna, it belongs to Oligocene as well. The Russian paleontologist A. Borisiak [47] was first scholar, who described from the same Turgai Hollow the complex fauna closely connected with nemoral forests. The most characteristic species of this fauna is giant unicorn – Indricotherium . According to Dolukhanov [20], some of representatives of the Arcto Tertiarian flora grow in dispersive way, as a seldom met specimen or groups in large territories (f.e., Taxus baccata ), some of them preserve in local places, strictly restricted in refugial areas. Among them should be mentioned eastern part of the USA (the Appalachian Mountains), the Russian Far East, and Atlantic Ocean coast countries in South Europe. Nemoral communities of East Asia and the Appalachian Mountains southward gradually turn into subtropical forests [48]. But if not to take into account the Maccaronesia islands, which as a matter of fact should not belong to Euroasia, two the most large and rich with the Arcto Tertiarian relicts’ regions in the Northern Hemisphere are situated in South Caucasus and adjacent some countries. One of it, Hyrcania includes the Caspian Sea coast slopes of the Elburs Mountains Range in North Iran, and its smaller north western site, Talysh, represented in the Caucasus’ Azerbaijan. The second large harbour – Colchis, is adjacent to eastern and south eastern seacoast – part of the Black Sea. Unlike Talysh, the latter’s main part occurs in West Georgia and south eastern part of the Black Sea coast of the Krasnodar district (Russia), the smaller part is situated in Turkey (the Black seacoast) and north slopes of the Pontian Mountains of north eastern Anatolia. The both refugia have some common features, but each of them are distinguished by theirs own, well expressed originality, depending, as we see below, the geological past of two regions, which have been developed in quite different way. 4
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 [49]. 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 [50,51,52,53,54,55]. Therefore, Colchis comprises an area, forming so called the ,,Colchic Triangle” [33,49]. This area is sheltered by the ranges of the Greater and the Lesser Caucasus Ranges to the North, East and South, whereas to the West it is open toward the Black Sea. The warmer and more humid climate, clearly distinguishing Colchis from adjacent regions, is a consequence of this peculiar geomorphologic configuration. The Colchis Triangle’s base part adjacents to the Black Sea, and its point part neighbours to Zestaphoni town. The coastal stripe is distinguished by most highest amount of precipitation. Here mountains have a perpendicular position to western winds blowing from the sea. 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. The condensation procesess are expressed in less intensive way on rise up parallel montain’s ridges toward the same west winds. Hence, there are less precipitations in neighbouring to Adjaria, the another seacoastal region, Abkhazia (1500 mm per year). Toward the point of the Colchis Triangle, the air’s moist current disintegrating like a fan, therhefore cloudness and at the same time quantity of precipitation is descending but, when uprise current of air reaches to the Great Caucasus slopes (Racha and Mingrelia districts), it turn into cold air masses, has condensed, and quantity of precipitations rise up once again. The same should be said concerning to Surami (Likhi) Ridge, the climateshed ,,bridge” between West and East Georgia. Unlike of Talysh mountains, a canyon like mountain gorges of Colchis are preserve high data of moisture, which support distribution of the Tertiary relict vegetations in high altitude levels, including subalpine belt. Mentioned unique feature make the South Colchis highly insightful, still less that some territories (reserves) of mentioned regions are well protected from human impact, at the same time have convenience long term weather and other ecological peculiarities for existence of some conservative forms of living organisms. Consequently, the incomparable harbour of the Arcto Tertiarian relict and sometimes relict and endemic plant species are represented in South Colchis which except south western part of Georgia, Adjaria and Guria, includes adjacent territory of north eastern Turkey (Lazistan). During the Quaternary glaciations this region situated as one of the most reliable harbour for Tertiarian flora not only in the Caucasus but as we’ll convincing later, in western part of Euroasia in general. Among mentioned three districts the most distinguish is AdjariaAdjaria. It located between 41°21´ and 41°53´N. It is typical mountainous country occupied Adjaro Imeretian mountain system of the Lesser Caucasus. As it was emphasized, its western part is open to the Black Sea. Eastern part of Adjaria adjacent to Arsiani Range, which is represented the watershed between the Black 5 and the Caspian Seas’ Basins. The most part of the Arsiani Range occurs in Turkey. Adjaria has the warmest and most humid climate within Transcaucasia. In coastal part of Adjaria annual precipitations range from 2600 to 3900 mm. With respect to its particularly humid climate, Adjaria is known as ,,Caucasus precipitations pole”, within this ,,pole” occurs Mt. Mtirala near Batumi city. Taking into account the small area (3000 km²) and climatic orographical situation of Adjaria (and Colchis in generall), the richness of its vegetation seems very remarkable. Hence, nowaday Adjaria is represented one of the best natural reserve of the ancient flora and vegetations in western Palaearctic [56]. At present from this part of Georgia is known 1901 species of vascular plants, arboreal plants include 173 species, among them 47 species are evergreen one. Within the diversity of Adjarian ecosystems the most peculiar place takes its endemic flora, which today includes 218 species (11,8% from total data); among them 72 species are situated as endemics of the Caucasus, 78 species – of the Colchis, 30 species – of the Adjarian Lazistan, 23 species – of Georgia, and 13 species of Adjaria [57,58,59]; (Adventinous flora – see in Davitadze’s work [60]). A rare phenomenon of flora of Adjaria once more confirmes this fact that dominant position within native phytocenosis are hold just endemic and relict species [58]. The unique vegetation and fauna of South Colchis is preserve in Kintrishi Nature Reserve in Adjaria (was founded in 1959). The protected area is especially important for its biodiversity, as it includes 102 species of arboreal plants, area some of which are restricted by South Colchis. In total in river Kintrishi Gorge are registered 1070 species of vascular plants [61,62]. The native flora consist of many endemic for the Caucasus and Georgia species, among them high attention of specialists are attract just species, with limit area confined in Colchis and bordering NE Turkey. Within the border of reserve one can found a straight trunk laurel cherry Laurocerasus officinalis , reaches in height to 17m; except laurel cherry, as a tree life form are grow local rhododendrons and holly shrub ( Ilex colchica ). Special attention require a rare tropical, tiny fern, Hymenophyllum tunbrigense , which grows either as epiphyte or on the rocks, among the mosses. The main objects of preservations in reserve are luxurious chestnut and beech forests with evergreen and deciduous understory, consist of relict, endemic and other rare species. In Kintrishi Gorge is situated well developed orobiome, with obviously expressed subalpine and alpine belts, including typical communities of elfin, sub prostrate, prostrate and crook stem phytoassociations, dominating by Caucasian rhododendron thickets, tall herbaceous forbs, and alpine low or dwarf vegetation. The most spectacular highmountain biocenosis is represented in remarkable alpine lake Tbikheli (2217 m a.s.l.) environment. In general, the Colchian or Eastern Euxinian Province’s flora consist of about 2500 species, of which approximately 2000 species are distributed in Georgia [58].
6 A BRIEF HISTORY OF FORMATION OF COLCHIS REFUGIUM
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 Adjaria 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 [63,65,66,67,68]. Approximately in mentioned geological time, the territory of adjacenting 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 [51,52,69 71].
THE PECULIARITIES OF COLCHIS FORESTS MIXED BROAD LEAVED FORESTS OF COLCHIS
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 [14,16,51,72,19,73,20,56,74,58,49]. Mixed forests of Colchic are occupy a considerable part of West Transcaucasia, namely in Colchis lowland and foothills of the Greater Caucasus and the Adjarian Imeretian Ranges. Its altitudinal distribution ranges from 50m to 500 600 m a.s.l. In upper elevations it gradually changes by deciduous forests, more poor with relict species in first stratum of canopy, and higher replaced by coniferous forests [72,18]. Soils are represented by yellow, brown and red colour with a well developed profile [75,76,77]. Forests of complicate, polydominant composition is characteristic not only rain forest but they are occur in some temperate climate forests due convenience soil and climatic conditions. The Colchis lowland and foothills is well confirmation of above stated viewpoint [51]. Grossheim [16] indicated that in polydominant forests of Colchis, major forests building species include 5 species: chestnut ( Castanea sativa ), 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. These oligodominant forests consist of by hornbeam beech, then hornbeam chestnut, hornbeam oak and beech chestnut formations. From hardwood forests with three species combinations should be mentioned: chestnut hornbeam alder, chestnut hornbeam beech, and hornbeam beech alder forests. According to mentioned associations, Albov [10] purposed the following definition: ,,Colchic mixed forests of broadleaved deciduous species”. Some Caucasian botanists are belong the same oligo and polydominant forests to mixed (moist) subtropics [18,19]. Besides above mentioned leading representatives of forests’ first stratum, the local canopy consist of by following subdominants: Ulmus minor, U. glabra (=U. elliptica ), Diospyros lotus, Fraxinus excelsior, Tilia begoniifolia (=T. caucasica ), Pyrus caucasica, Malus orientalis, Acer campestre , some species of Salix , etc. On limestones appears Laurus nobilis , in eastern part of
7 Colchis occurs Zelkova carpinifolia , sometimes together with deciduous forests building forms, growing frequently a poor stands of an yew tree ( Taxus baccata ). As some botanists [78,50,16] underlined, lianas are not characteristic vegetation form for virgin forests of Colchis and occurring mainly in forest’s margins. This fact that in Colchic forests they are met so largely, are the secondary occurrence, after destruction of mentioned forests in vast areas by means of human’s agricultural activities from prehistoric times. West Georgia’s lowlands and foothills covered nowadays with exotic for native environments plant cultures, which are manifested by the following introducing plants: tea, citrus species, tung tree, maize, eucalipt species, palm species, cryptomeria ( Cryptomeria
japonica ), etc. Ketschoveli [72] emphasized that in the first half of XXth century an intensive introduction in Colchis many subtropical plants made a profound reconstruction of primary biocenosis.The latter one remained in very limited area, f.e. in Lake Paliastomi environs. Fragments of these communities have restricted only in very abandoned arable soils, thanks to abundant precipitation,trigerring formations of secondary forests. Hence, communities with alder and hornbeams, in most situations are the derivatives of primary Colchian forests. Just this formations are so rich with secondary vines. The following vines may be mentioned: Hedera colchica, H. helix, Dioscorea caucasica (Colchic endemics), Smilax excelsa, Humulus lupulus, Clematis vitalba, C.viticella, Periploca graeca , Tamus communis, Vitis silvestris . Adventive lianas, such are Pueraria hirsuta and Lonicera japonica , display a succesfull competition capacity conserning to local, aborigenal lianas. To commonest representative of evergreen undergrowth belongs Rhododendron ponticum , which is growing as a tall shrub or small tree. It participates in all types of Colchic forests and spread in mountains higher than communities with canopy forests. The next characteristic species is Laurocerasus officinalis , it making more low stratum than previous species and forming very dence groves. One of the most peculiar arboreal plant of Colchis is its endemic box tree – Buxus colchica . Highly valued for timber, it undergoes intensive cutting considerably in XIXth century. Therefore, the most representative groves were eliminated and present day a thick – trunk box tree can be rarely met within the sources of some rivers gorges. Being very sensitive to climatic conditions, it requires high wet environments, the most shade gorges, aside it is a typical calcicolous plant abundantly covering Colchis limestone slopes. The other evergreen shrubs are represented by Daphne pontica , Ruscus colchicus, R.ponticus, Ilex colchica, Osmanthus decorus . The latter one in South Caucasus remained only in Adjaria. Species belonging to relict deciduous trees and shrubs should be mentioned: Quercus hartwissiana, Zelkova carpinifolia, Pterocarya pterocarpa, Ficus colchica, Rhododendron luteum, Vaccinium arctostaphylos, Coryllus avellana, Frangula alnus, etc. Is should be pointed out, that contrasted to deciduous component of understory, well developed and lush evergreen shrubs obviously prevailed in discussing region (with respect to mentioned communities more detaile information (((see pages 10 15 ). In well preserve Colchis underwood, dence shrub thicket causes a strong shading of ground, owing of which the herbage layer poor in species. The most commonest herb vegetation manifested by following species: Fragaria vesca, Dryopteris barbigera, Pteridium tauricum, Veronica officinalis, Salvia glutinosa, Viola alba . The most characteristic feature of local ground layer is a wide spread occurrence of ferns, and, contrary, in can be rarely met a forest’s grasses. Herb vegetation of Colchis is very poor of endemic species.
8 On sandy and slightly develop soils the composition of vegetation’s communities less abundant. In Abkhazia (Cape Bichvinta (Pitsunda) and Miusera coastal area) occur another relicts, forming Mediterrannean type of phytocenosis [79]. Mentioned should be made of the representatives of marytime Bichvinta Pine ( Pinus brutia var. pityusa ) on the Black seacoast, related and vicariated to Mediterranean P. halepensis , East Mediterrannean P. brutia , southwestern Turkey’s – P. brutia var. pendulifolia , the Crimean – P. brutia var. stankewiczii and eastern Transcaucasian P. brutia subsp. eldarica . A small area hold by strawberry tree ( Arbutus andrachne ), growing together with P.b.var. pityusa, Erica arborea, Cotinus coggygria, Rhus coriaria , Cistus creticus, C. salviifolia . In Colchis there are some gorges, which considerably enjoy a moist subtropical climate, marked comperatively mild winter and rainy warm summer. Among such gorges the following may be mentioned: Kodori, Bzibi, Mzimta (Abkhazia), Tkhenistskali, Inguri, Tekhura (Mingrelia), etc. Within these gorges the forest formation are consist of completely by relict arboreal plants: Ficus colchica, Diospyros lotus, Buxus sempervirens, rarely , Pterocarya pterocarpa , etc. The following lianas are interwine local arboreal plants: Smilax excelsa, Hedera colchica, Periploca graeca . As it mentioned above, the relict components of Colchic forests tree stratum of canopy are excluded from at an altitude between 600 800 m a.s.l., but poor stands of these communities occur on limestone ground as well. In a such habitats are sticking out the calciphobous plants – chestnut, Pontian rhododendron. In Colchic mixed forests characteristic peculiarity will be considere a widely distribution of epiphytes. Although, flowering plants are confined only by facultative epiphytes, but it should be noted some ferns which occurring as a typical obligative epiphytes (f.e., Polypodium serratum ). There are a lot of epiphyllous lichens (f.e., Usnea barbata ) [16]. From the cenotical and floristical point of wiew, the lowlands and foothills forests (0 – 500 – 600m a.s.l.) of Adjaria are richest in Colchis. The maritime or Kobuleti – Chakvi and Adjaro Imeretian ridges west slopes are covered by most typical Colchis forests with Fagus orientalis , Castanea sativa, Quercus hartwissiana, Q. dshorochensis, Carpinus betulus dominancy. Just in mentioned territory of Adjaria and adjacenting NE Turkey (till Malet – Ordu Gorge) is preserved rich of forest’s relict vegetation communities [58]. According to Manjavidze [56], with mentioned zone natural forest vegetation rather well preserved in upper part of foothills and Kobuleti Sanctuary of the Kobuleti Lowland.
THE PHENOMENON OF COLCHIS MOUNTAINS FOREST UNDERSTORY
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 [14,46,49,80]. To mentioned communities a special monograph was dedicated by A. Dolukhanov [20], arealogical review belongs to R. Gagnidze [55]. Phytocenologically these formations are called “Colchian understorunderstory”y”y”y” [20], or “Colchian semi prostrate evergreen underwood”””” [55], although in composition of Colchis undergrowth participated deciduous shrubs as well. The exceptional peculiarities of Colchic undergrowth, harbouring Miocene Pliocene relict living remnants, largely determined by complex of factors. Mentioned should be naming physico geographical factors: an extraordinary configuration of the Colchis Tringle, its location between the Black Sea and the Greater and the Lesser Caucasus Ranges, consequently this region’s very mild climatic condition, unusual for moderate climate latitudes of western part of 9 Euroasia (western Palaearctic). Concerning to biotic factors should be mentioned competition between the Colchis thick understory and forest building dominant woody species. 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” [49]. 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 phytocenosis and straight trunk beech or spruce forests [20]. According to Dolukhanov [20,81], 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 ). 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. Grossheim [16] on the base of investigations, carried out by Golitsin [82], outlines evergreen underground of Adjaria as an independent type of vegetation occur in moist gorges (ranged from 960 to 1100 m a.s.l.). These communities by mentioned scholars are named with term “shkeriani” (Georgian “shkeri” means rhododendron). According to Georgian botanists Ketschoveli [72], the term “shkeriani” is usually applied to define thickets of evergreen shrubs and sometimes creeping trees. Golitsin [82] attributes this term to a whole complex of shrubs (22 species), including besides evergreen shrubs, deciduous arboreal plants as well. Among mentioned complex the dominant position are occupied by following species: Betula medwedewii, Rhododendron luteum, R. ponticum, R. ungernii, Epigaea gaultheroides, Vaccinium arctostaphyllos, Viburnum orientale . From the viewpoint of Golitsin such formations of shrubs are primary original plant species, have been formed in the Tertiary Period and to present day remained as a relict associations. The analogues of mentioned thickets, this scholar considers in Madeira and Pyreness Peninsula. Grossheim [16] outlines these communities as one of the versions of upper border of distribution evergreen shrub complex in moist gorges within wet climate conditions of Colchis. Unlike to West European broad leaved forests, the Colchis same communities considerably differ. A special attention should be paid to the participation of creeping and semi creeping, vegetative migratory trees and shrubs in local broad leaved and coniferous forests. Their roles in formation of Colchis mountains forest ecosystems are insignificant. According to Dolukhanov [20], there are two ecological groups of Colchic underwood. Representative of 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. 10 The second ecological group 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. Following Doluchanov, except prevailing evergreen plants, this group of communities is distinguished from other bi oecological features. Dwarf semi prostrate trees and shrubs are pressed to the ground under the snow mass and lower parts of their trunks and saber form sprouts are rooted. In this way plants turn in dormancy. The same semi creeping rule of growing of the highmountains plants, thanks to capacity of rooting their trunks and shoots, triggering the orobiome’s woody plants to vegetative propagation. Such way of reproduction permits to single seedy generation of native birches, pontian oak, timberline’s beech, etc., develop in vegetative way during long time: hundreds, sometimes thousands of years. Growing in large and dense thickets, clones of creeping woody plants begins slightly broadened territories within their habitats, gradually descending throughout the slopes, occurring sometimes down from their natural upper boundaries. Another impressive botanico geographical peculiarity of the Colchis mesophyllous creeping shrubs, as it was underlined, is the wide amplitude of theirs altitudinal distribution, although some species may have stenochoric areas. In highmountain cold climate conditions, evergreen woody species have to endure temperatures below zero under a deep snow cover. According to Dolukhanov [20], the relict vegetation of Colchic underforest in the Pleistocene’s Ice Ages 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. Dolukhanov pointed out that outside of Colchis, the species of the mesophyllous semi prostrate evergreen communities also appears in various countries and continents. Among them the strongest modern centre of the Northern Hemisphere, where the formations, similar to the western South Caucasus evergreen understory occurs, is eastern and south eastern Asia, from the Himalayas up to Korea and the Japanese Islands. These phytoassociacions are spread in northern latitudes from 24 0 to 30 0. Mentioned formations are situated in Asia under canopy of dark coniferous forest at the height from 2800 3800 up to 4000m a.s.l., sometimes disperse in the subalpine belt. This highmountain area is clothed by Rhododendron arboreum , which is closely related to R. caucasicum with the same autoecology [83]. Countries of South and South East Asia distinguish by the influence of monsoon air circulation. Therefore, the existence of the evergreen shrub communities are related to the mountain natural belts of a monsoon climate. Another powerful centre of evergreen semi prostrate formations occurs in north eastern America, especially in the Appalachian Mountains. It mostly concentrated between the northern latitude of 37 0 39 0 (approximately the same latitudes, where situated Talysh refugium in south eastern part of South Caucasus). Survive of the same communities in Maccaronesia and Atlantic Ocean coast area of Pyrenees Peninsula can be explained by oceanic moist climate in mountains of mentioned regions. The same role performs the Black Sea for the Caucasus and eastern part of Balkan, which through dominating atmospheric current, supplies with precipitations above mentioned regions
11 mountains slopes. This convenience for vegetations position of the Greater and Lesser Caucasus is the main reason of the positive balance of atmospheric moisture throughout the year in Colchis. Dolukhanov [84,85,20] suggests that Colchis refugium is isolated from the other evergreen, semi prostrate vegetations moist centres, its endemic and other rare species of mountain and highmountain zones belong to the relict nucleus of vegetation of the remote past. On the basis of analysis of climate data Dolukhanov [20] assumes that to the north of 27 0 to 28 0 latitude, where the evergreen semi prostrate communities are concentrated, the distribution of continental atmospheric masses in winter, causes deep temperature depressions. The climate is rather rough in dry period of the year in monsoon countries of Asia. In East Transcaucasia, within Talysh refugium (the Hyrcanian shelter in general), climate conditions are also unsuitable for large scale development of evergreen understory. 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. Many botanists [78,46,20] indicate that Colchic specific semi prostrate and prostrate underforest some species restricted by South Colchis area, the other woody species are characterized by wide distribution, frequently with a disjunctive area in Mediterranean and Western Europe. Using the arealogical analysis of the concentration centres of species closely related to the Colchis ones, have been established by Dolukhanov [20] and Gagnidze [55]. Gagnidze recognizes three mentioned centres: 1) The centre of West Europe and Talysh. Nowadays these closely related species occur in macro disjunct habitats in Mediterranean countries (Atlantic Europe and the Maccaronesian Region), in micro disjunct areas in Talysh subcaspian region (Azerbaijan Talysh and slopes of the Elburs in North Iran); 2) The centre of east and partly South Eastern Asia, including the Himalaya; 3) The centre of the North Atlantic America (the Appalachian Mountains). Species connected with centre I. According to Dolukhanov [20] and Gagnidze [55] 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 ). R. ponticum is characterized by largely disjunctial spreading [46,20,80]. The main areas occur in Colchis, then in the north western part of the Caucasus and in the Black seashore slopes of Anatolia. It spreads in the Balkan Peninsula as well, the another fragment – in the Lebanon and Antilebanon mountains. The third area coveres the Iberian Peninsula. The native, vicarious race, R. baeticum is regarded as an independent species or subspecies [46,86,55]. Thus, as indicate above, macro disjunctive area of R. ponticum is dividing in some very close to each other species [20]. Following Engler [28], it’s suggested that to the end of the Tertiary Period, R. ponticum was largely extends in Europe and Mediterranean proper regions from the Caucasus to the Atlantic countries. Then, causing by climate cooling, within the most part of its range extincted, and survived in some, separated in great distance from each other refugia. R. ponticum is closely related to the Caucasian – northeastern Anatolian subalpine R. caucasicum . On the calcareous slopes of the Greater Caucasus and the Pontus Ridge (1700 2400 m a.s.l.), above the timberline zone, R. caucasicum and R. ponticum create hybrid form, R. sochadzeae [87,81,86,87,55]. On the other hand, R. caucasicum is related to the East Asian R. 12 aureum (R. chrysanthemum ), (Siberia, Kuriles, Korea, Sri Lanka, Japan [83]. According to autoecology of both species, it should be mentioned that R. caucasicum is sensitive to winter frosts, but R. aureum is more cryophilic species [88]. Botanists [88,89,20], pointed our very close relations mentioned species, suggest pre Quaternariar origin of R. caucasicum in the mountains of the Caucasus. Rhododendron ungernii and R. smirnowii are one of the most interesting representatives of the Tertiarian flora with present day very reducing area, known as the Colchic Lazistan species (endemics). Botanists consider these woody plants to be related ))) with R. arboreum of the Himalayas [83]. It extends in altitudes from 1850 to 3650 m a.s.l. and is typical component of undergrowth within broad leaved and mixed forests. Typical habitat of R. ungernii is a steep, moist, shady slopes along with beech, spruce and fir forests. It’s often found with R. smirnowii , therefore, can be distributed sympatrically. Laurel Cherry ( Laurocerasus officinalis ) occurs in the Caucasus, southern Europe and northern Anatolia, subcaspian Iran and the Balkans. European population are geographically repleced by L. luzistanica , distributed in the north western part of the Pyrenees and Macarronesia. Another close relates species are spread in south eastern Asia and North America [20,55]. With respect to species of Ilex . The Europe Mediterranean Colchic species group includes Ilex colchica , I. aquifolium and I. perado . The Colchic I. colchica has a vicarious area with I. hyrcana [49]. AAmong Colchian Mediterranean species of Ilex , the most conservative features of the Tertiary Period preserve I. colchica and I. imeretina . As the same time just I. colchica appears as one of the basic representatives of Colchis semi prostrate underforest. Genetically closest I. imeretina as well as its sister species with not less distinguishing features of ancient origination, populates exclusively moist gorges of Colchis [20]. UUndergrowth, consisting of I. hyrcana in beech forests is peculiar for its high height (3 5m), and dencity of thicket [91]. The Colchian species of Ilex in lower part of the most humid gorges rarely growing as a tall shrubs (5 6m) as well, but never forms dence, monodominant stratum [20]. Ruscus colchicus is an endemic of Colchis. It’s closely related to R. hypophyllum of Mediterranean Europe. The third related Europe Mediterranean Anatolia species is R. hypoglassum [55] . The Mediterranean Caucasian species group is represented by Daphne albowiana and D. pontica [55]. Species connected with the centre II. According to Gagnidze [55], two closely related lianas Hedera colchica and H. pastuchowii belong to the Caucasus East Asia’s group. The former relates to the Chinese species H. robusta , the latter one to the species of South Eastern Asia H. tobberi and H. schensiensis . H. colchica inhabits in the moist forests of Colchis, H. pastuchowii requires less wet climate, distributes in Talysh and in western Greater Caucasus. Species connected with the centre II and III. As it was stressed above the close floristic alliance between eastern Asia and North America have long been discussed by plant geographers [46,42,43,92,93,94]. 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 [95,96,97,98,99,80]. Mentioned disjunction area become more impressive by the third representative of this oligotypic genus, E. gaulhteroides , the typical Colchic – Taurus species [100]. The latter closer related to the Japaness species. Such configuration of vast disjunct area
13 confirm the opinion of biogeographers concerning relations of the Caucasian flora with the centres of development of evergreen woody plants since pegan time [46,14,20]. According to degree of adaptations of the nowadays ecological conditions, the Tertiary relict woody plant species of Colchis can be devided into three groups [46,101,56]. The first group is presented by ,,progressive relicts”. In this group should be included plant species, which areas greatly reduced during Pleistocene glatiations, but in modern time they managed to restore their areas, sometimes they are able to further distribution, and within the borders of their ranges are characterized by quite normal development. It should be mentioned here Picea orientalis, Abies nordmanniana, Fagus orientalis, Fraxinus excelsior. The second group embraces plants species, which in Postglacial Period survived relatively unchanged from an older period, hence now do not decline their areas, and throughout their ranges are characterized by normal growing and development. The following species belonging to this group should be mentioned: Laurocerasus officinalis , Buxus colchica, Ficus colchica, Rhododendron smirnowii . IIIn third group are represented ancient species, which during last Ice Ages particularly reduced their former large expanse territory, but now is narrowly confined. In the Colchian typical communities these relicts, as a rule, can be found as evidently secondary floristic elements, consequently, bio ecologically manifested a low vital capacity. 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”. Colchian Epigaea growing in moist ecotops of Adjaria with an extraordinaly plentiful of precipitations, within vegetation communities, distributed in physico geographical conditions with the most convenient abiotic factors for Colchic underground. The shrub inhabits in mid mountain’s (850 and 1000m a.s.l.) steepy stony slopes, which is constantly moist, more exactly, soggy with abundand rains or melted snow cover’s waters stream, within rarefied understory. It’s worth mentioning that just this type of habitat and niches, as Dolukhanov noticed [20], can be served as an incomparable pattern for restavration of the physico geographical environment, widely distributed, probably, on the boundary of Oligocene – Miocene times; the situation of this part of ,,palaeocolchis”, which, apparently, during long time was a centre of concentration, and the one of the main centre over a long period of time for conservation of vegetations formation’s of an Arcto Tertiarian Geoflora. In the centre of its area – the Pontian Range in North Anatolia, epigea grows in beech and spruce forests with undergrowth composed by Vaccinium arctostaphylos , species of Rhododendron , sometime also in mosses of bogs and evergreen shrubs. These habitats are represented in comparatively open sites of north slopes, occuping an altitudes between 920 and 2290 m [87,10,58,59,95]. In remote past epigea was widely distributed. This suggestion is confirmed by fossil remnants discovered in North America and Europe. Ilinskaya [102] in Pliocene’s flora of Transcarpathia found prints of Epigaea baikovskiae , close species to the Colchian Epigea. Betula medwedewii is widely distributed in subalpine belt of maritime part of South Colchis, including north eastern Turkey. It displays very close taxonomical and genetical relations with another birch – B. megrelica (2n=140 for the former species, and 2n=168 for the latter one). The latter occurs in disjunctive habitat in mountains of northern Colchis, in extremely moist slopes of Mingrelia, the Greater Caucasus (West Georgia). Dolukhanov [20] noticed very recent time divergence of the populations of two species. Despite both of them are representative of concervative remains of very ancient flora. Species of section Costatae to which they are belong, situated mainly in East and South East Asia, occurring the strongest modern centre of theirs species richness; some species present in Himalayas and eastern part of North America. 14 It worthy of attention that within a huge territory of western part of Euroasia, above pointed two species of birches, are representatives, probably, of the most ancient section (Costatae) of genus Betula, harbouring in Colchis till nowadays [20].
RELICT FAUNA OF COLCHIS Detailed long term study of invertebrate fauna on the example of Psylloidea (Insecta, Hemiptera)))) and their host angiosperm plants with detailed comparisons of two regions – Talysh and Colchis – faunistical similarity, carried out by us [2,5,3]. Investigations of two refugia’s vertebrate faunas are dedicated some works [103 115].However, the ecological and biogeographical comparisons between two refugia have not been paid enough attention. Studies of Colchis and Talysh wildlife by means of synthesizing multidisciplinary data, combining evidence from both ─ plants and animals, sometimes with theirs ecological, biogeographical and origination history, is given first time. 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 the following vertebrates: tiger, porcupine, hyena. The most stable refugia, sheltering representatives of above mentioned two groups, are Talysh and Colchis. Vertebrate fauna of the Tertiary of Colchis is very poor. Vereschagin [109] consider that black rat ( Rattus rattus ) belongs to the Pliocene relict of Mediterranean proper regions. During the glaciations of the Quaternary this species has been survived in warm seacoastal habitats of South Europe and North Africa, maybe in the Caucasus as well. At present black rat widely spread in the Caucasus as in wild nature, so in human’s various buildings. The population of rodent is most abundantly represent within seacoastal zone of the Black Sea, in Colchis region (sensu lato), from Batumi to Novorosiisk, including many sites of Mingrelia Abkhazia and adjacent Russia. Here, rodent lives in orchards, forests and seldomly in coastel areas. The altitudinal distribution extends from sea level to 600m a.s.l. The upper limit of Rattus rattus amounts to 1500m in Adjaria. The another region, where this species of rodent widely distributed is south east Caucasus – the Lenkoran Lowland and partially adjacent Talysh foothills. The ability of the black rat to live round year in Colchis and Hyrcania in wild condition, and it is habits to make dens and hollows under the tree stumps, confirm this animal’s origination in Pliocene time within Mediterranean proper countries and some neighbouring regions, including the Caucasus (Azerbaijan, Armenia), R. rattus behaives as a synanthropical animal populated exclusively a man’s building and various marine crafts. As regards invertebrate fauna of Colchis, particular attention should be paid to remarkable moth, Lederer’s brahmea – Brahmaea ledereri
15 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 95 107mm), ornamented with exclusive beautiful, ,,tropical”colour. Habitats presented in lowland’s dark and moist forests. Trophically lincked with Osmanthus decorus , a relict shrub. Brahmea is living ,,eyewitness” of deep anciency, inhabited in South Colchis since remote past, approximately from latest Paleogenic [25,26,6,116]. 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 [26,6,116]. 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. Another relict species of Tertiary age is Schamyl’s ghost moth – Phassus schamyl . Like Brahmaea’s two species, P. schamyl is also living remnant since pegan time of the Caucasus. Its endemic form survived the climate deterioration of the end of Tertiary and then three or four major glaciations of Quaternary. P. schamyl resembling to Brahmaea moths is characterized by ,,tropical”appearance – rich, colourfull and complicate design. It is rather big (wingspan of female reaches to 10 cm), nocturnal butterfly, active in twilight. P. schamyl belongs to one of the most ancient family of Lepidoptera__ HepialidaeHepialidae. Its area extend through the whole coastal zone of the Caucasian part of the Black Sea, dispersing in mid mountain altitudes. A margin side of area includes eastern part of the Surami Range (Borjomi district, Baniskhevi Gorge famous of its Colchic type of undergrowth). Within its area the centre of populations diversity occurs in the Colchis (Abkhazia, at altitudes not higher than 1500 1600m). Resemblance of Brahmaea , the closest region of other species of genus Phassus confines in Pacific Ocean coastal zone of Asia, but unlike genus Brahmaea which is sharply restricted within border’s of the Old World, phassus’ many species are distributed in the New World [25,116]. Differing Brahmaea , in seacoastal zone of Colchis and NE Turkey, usually occur the next relict species from another family of Lepidoptera ––– Papilionidae, namely Thais cerisyi caucasica , subendemic of the Caucasus. This is a rather large, bright and many coloured diurnial butterfly. Lives in humid and shady forests, from seacoastal area penetretates in the North Caucasus (till Maikop town environs), one isolated population occur in the forests of kakhetia (East Georgia). Out of the Caucasus the species is distributed in Minor Asia and in Balkan Peninsula. By feeding habits it connected with species of Aristolochia , section of iberica , which is more close to tropics’ species than that of European ones. Species of Thais represented in the Russian Far East and China. All of them are outstanding by very rich, quite tropical coloration and ornamentation, feeding on Aristolochia as well.
16 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 present day, but found in large areas, renew their ranges in postglacial time. According to Bey beinko [21], fauna of ordo Orthoptera Orthoptera by its origin, phylogenetical connections and ecological peculiarity is heterogeneous. The majority of species can be united within Caucasian relict endemics of northern (boreal) origination, therefore they appear as a modified offsprings of ancient mesophilous fauna migrated from Boreal Centre within the Caucasus in second half of the Tertiary Period. Among some genera of grasshoppers should be mentioned Isophya . Bey bienko considers that species of grasshoppers, outlining as analogues of Colchis relict flora is very scanty, and merely a few species of genus Isophya can be closed to the mentioned refugium’s relict woody plant formations. From this viewpoint, 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 [21]. Thus, species of Isophya show the same vicarious distribution within Transcaucasia’s two refugia, which is characterized to above discussed genus Brahmaea . It should be underlined that Colchian Isophia has close relation with I. kalishevskii , found in Abkhazia. The latter species penetrates in mountains (the Greater Caucasus), reaches 1800m a.s.l., and sometimes higher altitudes as well. According to this factor, Bey bienko doubts to belong this species to the relicts of Colchis. But from our viewpoint of possible connection of the I. kalishevskii to the Colchic relict fauna is quite permitting. Difference of opinion, as we consider, has arisen upon the specificity concerning to biology of the mentioned species. The latter species of Isophya displaies the same adaptation capasity, which are so characteristic to Colchis highland’s creeping, vegetative migratory representatives of underforest. As it was stressed above, these, mainly relict shrubs successfully reproduce with vegetative organs, which is a well expression of adaptation during long winter under the thick snow mass. The adaptative capacity of I. kalishevskii to the high mountains severe climatic conditions sharphy differ it from thermophilic forms of relict species of the genus Isophya , representated in lowland’s refugia of South Caucasus. But grasshoppers, including mentioned genus, hibernate in an egg’s stage, whereas their active stages occur in summer period. This specifity in this group of insects’ life mode, probably, make them become more independent against the threatment of low temperature during winter time. Using this peculiarity of biology by some species of Isophya , it’s suppose, gives them opportunity to penetrate into mid and even high mountains. The particular interest of entomologists and biogeographers attract unwing grasshopper − Dolichopoda euxina , inhabiting in caves of seacoastal zone of Abkhazia and Sochi district (Russia). The other allopatrically dispersed species of this genus situated in Mediterranean proper countries make up disjunctive in several sites area. With inhabitant of other caves – genus Troglophilus , genus Dolichopoda belongs pure Mediterranean tribe, a distant relate taxon to Indo Malaian tribe, Rhaphidophorini . In general, subfamily Rhaphidophorinea to which belongs D. euxina , preserves some feature of very ancient group, fragmentary distributed in both Hemispheres and having its geographically isolated representatives within fauna of South Africa, New Zealand and southern part of South America [117]. According to Bey bienko [21], being based of above mentioned materials, suggests this species of Dolichopoda belongs to the most conservative palaeoendemics of the western 17 Caucasus, i.e. is representative of the initial tropical fauna of the Caucasus, resemblance of primitive beetle Parandra caspia , harbouring itself in the Talysh and adjacent North Iran. The straight reason, surviving this species of grasshopper till present day is its particular mode of life in caves, has been protected mentioned insect from direct influence of glaciations and post glaciations climatic deterioration. A typical, the Colchic Tringle distribution attracts to such representative of subphyllum ChelicerataChelicerata, as is the Colchis subendemic and relict subspecies of scorpion – Euscorpius mingrelicus mingrelicus (see map 3). According to V. Fet [118], some species and subspecies of genus Euscorpius have many endemics end show remarkabely disjunct distribution. Taking into account that scorpions (ordo ScorpionesScorpiones)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 [119] and Fet [118], 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. Within this area it dividing into allopatrically dispersed 6 subspecies: E.m. mingrelicus (Eastern Euxinian Province, the same Colchic Tringle), E.m. ciliciencis (Turkey), E.m. histrorum (former Yugoslavia, Greece), E.m. phrygius (European Turkey, Moldova), E.m. caprai (Italy, Slovenia), E.m.caporiaccoi (Bosnia). Some subspecies of E. mingrelicus can be distributed simpatrycally with closer species – E. germanus , including four subspecies. Some related subspecies can be meet in transitional territory (Turkey, Italy, Croatia).
Sch. map 3.
Within Colchis region two relict and endemic species of round warms (phyllum AnnelidaAnnelida, class OlygochaetaOlygochaeta) are represented. Mentioned should be Eisenia transcaucasia , situated in limestone sites of mountainous Abkhazia, and Dendrobaena faucium , occurring in Colchis (Abkhazia) lowland’s marshland. The western part of the Caucasus and West Georgia in particular, displaing a favorable climate, became an area of formation a rich and distinctive mollusks fauna. The species of subgenus Leiostyla (genus Lauria ) are known from deposits of Oligocene and Miocene of mid Europe, and nowadays occur only in the Caucasus, Great Britain, the Azores, Madeira, the Canary islands, Portugal and Algeria. In Peleogene of Europe there were a lot of species and genera of the family of OleacinidaeOleacinidae. At present, except genus Pioretia , all other genera are represented in tropics of Central and South Americas. Species of Pioretia survived in North Africa, South Europe and western south Caucasus, in Colchis ( Pioretia mingrelica ) [24]. Subfillum Branchiata, class Crustacea, ordo Decapoda includes Colchian crayfish _ Astacus colchicus , rare relict species with restricted area in the Colchis and neighbouring north western Turkey, inhabits a low mountains and mountains rivers, seldomly lakes. A. colchicus vicarianing with another rare relict species of the Caucasus – A. pylzovi found a harbour on the south slope of the Greater Caucasus (East Georgia _ Kakhetia and adjacent Azerbaijan). The crayfish lives in mountainous rivers as well. Among widely distributed animals of Colchis Lowland and foothills should be mentioned: boar ( Sus scrofa ), badger ( Meles meles ), brown bear ( Ursus arctos ), roe dear ( Capreolus capreolus ),martern ( Martes martes ),weasel ( Mustela nivalis caucasica ), wolf ( Canis lupus ), fox (Vulpes vulpes caucasica ), lynx ( Lynx lynx ); species of birds: Columba oenas, Cuculus canorus, Dendrocopus major, D. minor, Bubo bubo, Falco peregrinus , etc.
ANALYSIS OF COLCHIS BIOCOENOSIS The Colchis, as a refuge have been managed to harbour some representatives of ancient woody plants of the Arcto Tertiary Geoflora, some of them in the Colchis Traingle are populated a negligeble territory, presented exclusively in this region (f.e., Quercus pontica, Epigaea gaultheroides, etc. ,), the other one form even a continuos belt (f.e., Castanea sativa ). The same situation occur in the other refugium of South Caucasus, in Hyrcania. This fact pointing out a wide range dispersal capacity of the Arcto Tertiary Geoflora in Neogene from high latitudes of the former Laurasian mainlands to South direction, including the Caucasus and neighbouring regions. To compare biocenosis two refugial centres of South Caucasus, it must be stressed an independent formations of two specific complex of living organisms. If take into consideration of two harbours different ways of origination within borders of the Caucasus, differing in altitudinal zonation, latitudinal location, some specific features of ancient mesophyllous forests between two refugia, it should be consider more convergence status of two shelters communities, than vicariation of phylogenetically close synusias. On the base of comperative analysis of Colchis (Adjaria) and Yakushima island (Japan) two floras, carried out by Kikvidze and Ohsava [49] it can be find more ratio of similarity between mentioned two remote places floras, than that of two refugia of Transcaucasia. Despite, there are a rather long list of sister species, represented in Colchis and Talysh. Mentioned should be species of Zelkova, Buxus, Ficus, Ilex, Ruscus, Rhamnus, Daphne, Brahmaea moth and Isophya grasshopper – from insects. Despite, on the base of evolution of two region’s living forms in insular, an isolated environs till nowadays, Grossheim [13,14], the most outstanding botanists of the Caucasus, belongs mentioned and some herbaceous close species not to vicarious plants. He consideres this similarity to ,,parallelism of species origin”. According to previous materials, through Colchis perfect spectrum of altitudinal zonation, in contrast of Talysh, are manifested all categories of Tertiarian deciduous broadleaved relicts: 19 1. Species, restoring theirs pre glaciacions areas and successfully competing with dominating in Holocene time forms (species of genera: Picea, Abies, Fagus, Fraxinus ); 2. Ancient species, which managed to preserve their usual development tendency till present day (species of Laurocerasus, Buxus, Ficus, Rhododendron; from invertebrate animals – species of Euscorpius ); 3. Relicts, performing evidently secondary floristic position, therefore, have a status of ,,regressive relicts”. As a model of such species was named Colchian endemic and relict species of Epigaea ; conserning to animals to this ,,living fossil” group’s with a great probability can be belong Brahmaea ledereri , if it remains even as a ,,Critically Endangered” position in Colchis most moist gorges. Macro disjunctive areas, characteristing of genera to which belonging such species, without using a fossil materials, confirm theirs transcontinental and even intercontinental continiously distributions in the Tertiary Period. Nowadays habitats, similar of ,,dying relicts”, can be used for the reconstruction of the immemorial past, when an initial forms of the Arcto Tertiary flora perform a significant role in the Late Paleogene – Early Neogene biocenosis. Because of to the absence or almost absence of their analogues in western Palaearctic, the Colchic mostly evergreen semi prostrate and prostrate underwood arise of considerable interest. As it was pointed out above, a synusia similar to the Colchian evergreen dendroflora, a supposing direct inheritor of the Late Paleogene’s Euxinian Basin, has mountain ecosystems of a monsoon climate in eastern and south eastern Asia. In various regions, even with oceanic climate of North America and Europe, they lost their dominance position. The reasons determinated of this phenomenon to Colchis mid – and highmountains gorges, from climatological and orographical viewpoint have been illustrated above. But authors of above mentioned conclusions [13,14,16,20], are not focused their attentions on the geological events took place in boundary of Paleogene Neogene, by means to the collision (convergence) of the African Arabian and European plates. Late Alpine neotectonic development caused a further uplift of the Greater Caucasus Range and lately (Miocene) the Lesser Caucasus Range as well [67,68,65,66]. Further increasing of orogenetic movement in Late Pliocene, turned the Greater and the Lesser Caucasus Ranges into powerfull mountain systems with theirs characteristic configurations similar to modern ones [120,63,121]. This less discussed scenario may explain additionally why do Colchis refugium in moderate climate latitudes manages to remain as a such stable harbour for preservation so large variety of the Arcto Tertiary Geoflora’s representatives. Unlike Talysh mountains, which were far more lesser include in the tectonic processes, above mentioned collision has resulted very serious modification of geographic features in Colchis Region, increased heterogeneity of western south Caucasus landmass. Side by side to the Colchis Lowland, uplift processes determine a final formation one of the highest mountain ranges (The Greater and the Lesser Caucasus) of Western Euroasia with theirs deep gorges, slopes orientiated generally to the Black Sea. As in any other mountain countries, formation highlands may have trigerred profoundly new, geologically young habitats. The axeleration of relief’s dissection, encreasing processes of origin a new habitats, could be prepare a nourishfull background, supporting new large scale wave of biological evolution, including species radiation, or allopatric speciacion. In contrast of Talysh mountains, which were less touch by alpine fold processes, the Lesser and considerably Great Caucasus Ranges wide altitudinal gradiant provided an intensive processes of species vertical expansion. Within newly origin habitats, side by side existing ones, both relict and newly evolved species (particularly in calcareous rock and scree massif) could survive. Therefore, above discussed geomorphological
20 transformation undoubtedly increased the heterogeneity of Colchis and in general Euxinian Basin’s biota. At the same time, contrary to Talysh, Colchis both relief units: lowland and mountain systems kept the homogeneity of mild climatic conditions, which performed an exclusive role for preservations of the such remarkable unity of evergreen broad leaved mesophyllous woody species within a large latitudinal and altitudinal areas. During Ice Ages, glacial events, undoubtedly, have influenced to the biocenosis of Colchis far more severe and drastic than that of non glacial areas of Talysh refugium. According to Kikvidze and Ohsava [47], Pleistocene’s severe conditions and geographical isolation of Colchis may be responsible to support a high potential for adaptive radiation of some native taxons. Genus Rhododendron may be seen in this context. There are about 1000 species of Rhododendron distributed worldwide, from which 330 occur in southwestern China. Within this country mentioned genus vary widely in size and habit. Some are trees that can grow to a height of 30m, and other forms are tiny shrubs [122]. As it was discussed above, one of the main genera of Colchis evergreen communities is Rhododendron . 5 6 species of this genus reflect their adaptations ability within widely varying habitats in Colchis from sea level to snowline. Taking into account, sympatric and allopatric speciacion of these species, vicarious (f.e., R. ungernii and R. arboreum ) and remarkably disjunct distribution (f.e, R. ponticum ) some of them, largely resulting by mean of complicating of the topography of Colchis in Neogene. According to above proposed scenarios of significant uplift of Colchis region (and the Caucasus in general) territory during the past several million years, Dolukhanov [20] noticed worthy of attention opinion. According to this scholar, peculiarities of genus Rhododendron and other forms of subprostrate shrubs show that the way of formation of mentioned ecological group of vegetations are closely links with the Caucasus mountain systems, where they were obliged to adopt to the winter’s severe conditions. The origin of closer recent forms of Colchian like evergreen mesophylous dendroflora and subprostrate synusia, as it was underlined above, are related to the mountain ecosystems of a monsoon climate [81,20]. In modern geological age the single region in the world, represented within latitudes lack of atmosphere’s specific monsoon circulation, and despite of this climatic peculiarities has managed to preserve the floristic complex of above discussed type, is Colchis refugium [20]. According to the same scholar,the floristic communities of the ambro – and thermophilous evergreen dendroflora was supposed to be found in the Euxinian Basin and subcaspian mountain systems in the Late Paleogene. But a wide altitudinal expansion, as it was stressed above could be took place in the Neogene under conditions of a positive balance of atmospheric moisture with an annual mean temperature of 18 20°C at the altitude of up to 1500 m a.s.l. The temperature optimum of the Atlantic Period of the Holocene illustrates the further altitudinal expansion of the Colchic evergreen arboreal plants complexes and sometimes more or less closely linked with this communities representatives of invertebrate fauna.
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ИТОГИ БИОГЕОГРАФИЧЕСКОГО ИЗУЧЕНИЯ ДВУХ РЕФУГИУМОВРЕФУГИУМОВ АРКТОАРКТО ТРЕТИЧНЫХТРЕТИЧНЫХ ОРГАНИЗМОВ ЮЖНОГО КАВКАЗА
А.М. Гегечкори
В статье охарактеризованы два рефугиума Кавказа Колхиды и Талыша, в том числе их флористическая и фаунистическая структура. Особенно это касается реликтовых и эндемичных форм, их сравнительного анализа. Также рассмотрено распространение по высотной поясности организмов, сохранившихся с третичного периода (в основном неогена). Сравнительный анализ этих двух регионов показал существующую между ними значительную разницу. Последняя вызвана разными путями формирования этих рефугиумов в историческом прошлом, почвенно орографическими различиями. В формировании Колхидского рефугиума большую роль сыграли Черное море и горные системы Большого и Малого Кавказа, а в образовании Талышского рефугиума Каспийское море, горы Талыша и прилегающие пустыни.
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