On the Mediterranean Influence on the Formation of Herpetofauna of the Caucasian Isthmus and Its Main Xerophylous Refugia
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Russian Journal of Herpetology Vol. 2, No. 2, 1995, pp. 95 – 119 ON THE MEDITERRANEAN INFLUENCE ON THE FORMATION OF HERPETOFAUNA OF THE CAUCASIAN ISTHMUS AND ITS MAIN XEROPHYLOUS REFUGIA B. S. Tuniyev1 Submitted April 25, 1995. The Caucasian Isthmus has common origin within the areas of Ancient Mediterranean and it inherited the united type of mountain belts with Mediterranean mountain systems. On the basing of paleo-areas and re- cent chorology of Caucasian amphibians and reptiles the analysis was given, including the historical changing of the main characteristic of Mediterranean species-complex, its composition and connection with corresponding phyto-landscapes. The recent refugia of Mediterranean species in the Caucasus are de- scribed and made conjecture about stages of colonization the Caucasian Isthmus by these species. The possible ways of preservation of these refugia till now are discussed. Key words: Biogeography, Caucasus, Mediterranean, Herpetofauna, Refugia. Position of Caucasus in the scheme of gin of the Caucasian fauna was made by Satunin biogeographic division of Palaearctic is arguable. Its (1910, 1912), what we have pointed already in the independence in the formation of peculiar flora and previous work (Tuniyev, 1990). Menzbir (1934) in- fauna, or belonging to the adjacent areas of Europe cluded Caucasus and Transcaucasia into the Cauca- and Central Asia are differently interpreted in litera- sian subprovince of East Mediterranean province in ture. It is to the significant degree typical and for the Mediterranean subregion of Palaearctic. In the opin- separate parts of Caucasian Isthmus. Not aiming to ion of Menzbir, Caucasus was under the influence of analyze the whole herpetofauna of the Isthmus and two faunogenetic centers – Mediterranean and Cen- partitioning of its territory, we shall stop on the cer- tral Asian. Puzanov (1938) referred to the Mediterra- tain fauna of Mediterranean area and make an at- nean subregion all Transcaucasia which together with tempt to estimate the influence of this fauna on the the Greece archipelago, Asia Minor, Mountain Cri- formation of the herpetological complexes of mea, enter the East Mediterranean province. The Caucasus. northern slopes of El’brus and Talysh were distin- Zoogeographical works, concerning Caucasus, guished by him into the Hyrkan Province. appeared in the second half of XIX century. Wallace The numerous following works of 1940 – 1960th (1876), as well as later on Haake (1886), divided were based the main on the theriological and ornitho- Caucasian Isthmus along the crest of Great Caucasus, logical partitioning where Caucasus was placed ei- referring Precaucasia to the European subregion and ther to Mediterranean (Kuznetsov, 1949), either to Transcaucasia – to the Mediterranean. Severtsov the province of deciduous forests of Europe (Rusta- (1877) referred to the Mediterranean area the all mov, 1945; Kuznetsov, 1949; Bobrinsky, 1951). Great Caucasus, West Precaucasia and West Trans- caucasia, and the rest Transcaucasia – to the West The work of great expert and investigator of Cau- Asian region and North-East Precaucasia – to the casian fauna Vereshchagin Mammals of Caucasus Middle Asian region. As a matter of fact in this, very (1959) deserves particular attention. Criticizing the close to the recent, scheme of division, Severtsov works of authors, who diminish the significance of have recognized the different of the Caucasus fauna. Mediterranean area (to which Vereshchagin referred The most complete description of heterogeneous ori- Caucasus too), he wrote: “Mediterranean always gave more possibilities for immigration of animals 1 Caucasian State Biosphere Reserve, Sochi, Krasnodar region, and plants than Sahara or Kara Kum, but its signifi- Russia. cance as the independent ancient form genesis center © 1995 Folium Publishing Company 96 B. S. Tuniyev Border of Mediterranean region Border between West- and East-Mediterranean regions Fig. 1. Botanical borders of Mediterranean region (after Rikli, 1946). was not decreasing due to this. The correct interpreta- reshchagin’s interpretation of Mediterranean coin- tion of the concept of “Mediterranean,” and, there- cides as a whole with the boundaries of floristic Med- fore, and Mediterranean type of fauna could be iterranean of Rikli (1946) (Fig. 1), though in bota- achieved only with the complex analysis of natural nists the position of Caucasus also was a subject of a transformation which took place in the Cenozoic in long discussions. Thus, Boissie (1867) in his book the limits of Mediterranean geosynclinal. As it is Flora Orientalis referred the forest Caucasus, occu- known, the Sarmatian sea, which stretched from Gi- pied by deciduous and, partly by the dark coniferous braltar up to Transcaspian, later on in the Pliocene forests north Anatholiya and some regions of Euro- and Pleistocene not once disintegrated on the chain of pean Turkey to the region of Middle Europe. To the isolated basins, being connected sometimes in the ep- “Mediterranean region” he, besides the countries, ad- ochs of transgressions. The Aral coast and the north- joining the Mediterranean seacoast, referred and ern part of Caspian Sea due to the conditions of relief southern Crimea. Not only mountain Crimea, but the and high continental climate were early deserted and whole Caucasus was included in the Mediterranean come out of the united system of zones of Mediterra- subregion (province) on the territory of the former nean type. Nevertheless, the rest plots of this chain of USSR by Alekhin (1938) and Vul’f (1944). Gross- Cenozoic basins, among this and Caucasus, inherited geim (1948) and Maleev (1946) pointed on the affin- a number of single primitive features of landscapes, ity of the Caucasian formations of macquis, garigue, flora and fauna, the roots of which are going into phrygana broad-leaf forests with the same formations Miocene. Just because of this as the Mediterranean, of Mediterranean. Isachenko and Lavrenko (1980) taking in account its paleogeographical essence did not agree with them, referring Caucasus to the should be meant the southern vicinities of Europe (in- two botanical-geographical areas: European broad- cluding the south of Ukraine and Crimea), Northern leaf and Afro-Asian desert. Lavrenko (1958) consid- Africa, Asia Minor, Caucasian Isthmus (excluding ered also earlier that phytogeographically sense the Premanych), Talysh, and El’brus ridge” (p. 462). Ve- forest area of Caucasus in the origin and ecology is Mediterranean Influence on the Formation of Herpetofauna of the Caucasian Isthmus 97 related to the broad-leave forests of Europe, but not ranean group, common for all or the greatest part of to the Mediterranean. Takhtadzhyan (1978) having the East Mediterranean. The latter group in depend- analyzed significant amount of published data, re- ence on the character of species distribution not ferred to the Mediterranean area the foothill part of rarely is divided on the Mediterranean and East Me- West Caucasus between the Anapa, Krasnodar, and diterranean (Darevsky, 1957a). The understanding of Tuapse, the rest parts of Caucasus he referred to the the volume of Mediterranean group varies in differ- Euxine and Caucasian province of the Boreal sub- ent authors (Sobolevsky, 1929; Bodenheimer, 1944; realm and Armenian – Iranian, Hyrkanian, and Tura- Darevsky, 1957a; Anderson, 1968; Alekperov, 1978; nian provinces of Iran – Turanian region. Kireev, 1987; et al.). Alongside with it, the great ma- Stanyukovich (1973) refers the North Caucasus jority of authors include in the number of Mediterra- and Transcaucasia to the type of belting of the moun- nean faunistic elements the next species, distributed tains of the subtropical zone. Vereshchagin consid- on the Caucasus: Triturus cristatus karelinii, Rana ri- ered (1959) that the similarity and close affinity of dibunda, Mauremys caspica, Testudo graeca, Ophi- theriological complexes of the West Mediterranean sops elegans, Pseudopus apodus, Typhlops vermicu- with Caucasus (up to Talysh – El’brus on the east) is laris, Eryx jaculus, Natrix natrix persa, N. tessellata, connected with the common time of origin and for- Coluber najadum, Telescopus fallax iberus, and Mal- mation of high mountain landscape. As reasonably polon monspessulanus. wrote Il’insky (1937) the landscape of Mediterranean Earlier, considering the herpetofauna of the West type biogeographists not rarely mean the contempo- Transcaucasia (Tuniyev, 1990) we included in the rary landscape of Spain, Sicily, and Palestine. As a East-Mediterranean group Triturus cristatus kareli- matter of fact, as in the west so in the east parts of nii, Testudo graeca, Lacerta media, L. strigata, L. pra- Mediterranean alongside with xerophylic landscapes ticola pontica, Pseudopus apodus, Natrix tessellata, of garigue, phrygana and mountain steppes, there are and Coluber najadum. Moreover, in our viewpoint relict landscapes of Tertiary coniferous and decidu- Pelobates syriacus, Mauremys caspica, Ophisops ous forests, mesophytic meadows of alpine and sub- elegans, Ablepharus chernovi, Cyrtopodion kotschyi alpine types. colchicus, Typhlops vermicularis, Eryx jaculus, Nat- Szczerbak (1984) indicated both xerophylous rix natrix persa, Elaphe hohenackeri, E. quatuorli- neata, Coluber caspius, Eirenis modestus, Telesco and mesophylous species of amphibian and reptiles - for the different centers of speciation in the limits of pus fallax, Malpolon monspessulanus, Vipera lebeti- na, and V. ammodytes should be referred to