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HERPETOZOA 19 (3/4): 99 - 110 99 Wien, 30. Jänner 2007
On the Quaternary reptilian fauna of Bashkortostan (South Urals, Russia)
Zur Quartären Reptilienfauna von Baschkortostan (Südural, Rußland)
VlNER KHABIBULLIN
KURZFASSUNG Die Geschichte der quartären Reptilienfauna von Baschkortostan (Südural, Rußland) wird im Zusammen- hang mit einem kurzen Überblick zur Entwicklung des Klimas, der Landschaften und der Vegetation dargestellt. Fünf Reptilienarten {Lacerta agilis, Anguis fragilis, Lacerta vivipara, Natrix natrix, Vipera berus) sind für das Spätquartär des Untersuchungsgebietes anhand von Fossilbelegen nachgewiesen. Die neuzeitliche Herpetofauna des Urals begann sich im späteren Pliozän herauszubilden. Die gegenwärtige Reptilfauna entwickelte sich im mitt- leren Holozän, mit der Rückkehr der Reptilien aus den eiszeitlichen Refugialräumen. Die Gebirgsketten des Urals sowie die Tätigkeit des Menschen hatten auf die Ausbildung der gegenwärtigen Kriechtierfauna der Region keinen bedeutenden Einfluß.
ABSTRACT
The history of the Quaternary reptilian fauna of Bashkortostan (South Urals, Russia) is described and the development of climate, landscapes and vegetation are briefly reviewed. Five late Quaternary fossil reptile species {Lacerta agilis, Anguis fragilis, Lacerta vivipara, Natrix natrix, Vipera berus) had been identified for the region. The modern herpetofauna of the Urals started to form approximately in the Late Pliocene. In the South Urals the complete present day herpetofauna had established in the Middle Holocene due to reptiles dispersion from glacial réfugia. The Ural Mountains as well as anthropogenic factors had little influence on the evolutionary history of the recent reptile fauna in the region.
KEY WORDS
Reptilia, paleontology, evolutionary history, Quaternary, Pliocene, Pleistocene, Holocene, post-glacial dis- persion, glacial réfugia, Republic of Bashkortostan, South Urals, Russia
INTRODUCTION
Origin and evolutionary history of that issue (i.e. GARANIN 1983; KHABIBULLIN present-day faunas are among the most 2001a, 2002). interesting and intricate topics in zoology. The modern reptilian fauna in temper- They were poorly studied and challenged as ate North Eurasia (and therefore in Bash- far as the North Eurasian temperate zone kortostan) is primarily a result of the Late reptilian fauna is concerned, the evolution- Cenozoic post-glacial dispersion from gla- ary history of which, especially in the Plei- cial réfugia in South Europe and South- stocene, was rather dramatic. This paper West Asia (NIKOLSKY 1947; LENK et al. presents a synthesis of our knowledge of 1999). Therefore the evolutionary history the Quaternary herpetofauna of Bashkorto- of the modern herpetofauna of Bash- stan (South Urals, Russia) urged by the kortostan is restricted to the history of post- paucity and fragmentation of literature on glacial colonization.
MATERIALS AND METHODS
Published data is summarized from scheme proposed for the former USSR by the fields of paleontology, paleogeography YAKCHIMOVICH (1965, 1992) was used. A and biogeography and the stratigraphie general picture was created to analyze not ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at
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Fig. 1: Map of locations of late Cenozoic reptile remains in the territory of the Republic of Bashkortostan (RB) and adjacent regions, a - Krasny Bor; b - Akkulaevo; c - Kyzyrbak; d - Ziganskaya; e - Zapovednaya I, II; f- Lemeza I, II, IV; g - Nukatskaya. The administrative border of RB is represented by a dash-dotted line. Abb. 1 : Karte der Fundorte spätkänozoischer Reptilienreste auf dem Territorium der Republik Baschkortostan und angrenzender Gebiete, a - Krasny Bor; b - Akkulaevo; c - Kyzyrbak; d - Ziganskaya; e - Zapovednaya I, II; f- Lemeza I, II, IV; g - Nukatskaya. Die administrativen Grenzen der Republik Baschkortostan sind durch die strich-punktierte Linie bezeichnet.
only data from the precise area within the Brief characteristics of Bashkortostan administrative borders of Bashkortostan, but also from adjacent territories. Bashkortostan (Republic of Bashkor- Fossil reptile remnants from six excav- tostan, RB) is one of 89 administrative units ation sites were analyzed (fig. 1). Quater- of the Russian Federation with over four nary reptilian fossil materials were found million inhabitants and a total area of mainly in caves (eight out often sites). Only 143,000 square kilometers. Bashkortostan data on Late Cenozoic fossil materials is pre- is situated in the South Urals at the "meeting sented. point" of Europe and Asia between 51 °31 ' - All interpretations below if not stated 56°25' N and 53°10' - 60°00' E. The water- otherwise, primarily refer to Bashkortostan shed of the Uraltau mountain ridge divides and the South Urals (within the administra- Bashkortostan into the following natural tive borders of Bashkortostan, Orenburg and regions: the western part of RB (Fore-Urals) Chelyabinsk regions). But as far as the evo- belongs to the Volga river basin while the lutionary history of the modern reptilian eastern part (Trans-Urals) belongs to the fauna of Bashkortostan is concerned, it is Ural and Ob river basins. In between these tightly associated with that of the adjacent two parts are the Ural Mountain ridges that Ural region and the whole Eastern Europe, form the mountainous part of RB. The ele- and most generalizations can be extrapolated vations of Bashkortostan range from 58.7 m to the temperate zone of North Eurasia. to 1,640 m a.s.l., with the average altitude ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at
On the Quaternary reptilian fauna of Bashkortostan (South Urals, Russia) 101
Table 1 : The recent reptile fauna of Bashkortostan (South Urals, Russia) according to KHABIBULLIN (2001a). Tab. 1: Die rezente Reptilienfauna von Bashkortostan (Süd-Ural, Rußland) nach KHABIBULLIN (2001a).
Order Suborder Family Species Ordnung Unterordnung Familie Art Testudines Cryptodira Emydidae Emys orbicularis (LINNAEUS, 1758) Squamata Sauria Anguidae Anguis fragilis LINNAEUS, 1758 Lacertidae Lacerta agilis LINNAEUS, 1758 Zootoca vivipara (JACQUIN, 1787) Serpentes Colubridae Coronella austriaca LAURENTI, 1768 Elaphe dione (PALLAS, 1773) Natrix natrix LINNAEUS, 1758 Natrix tessellata (LAURENTI, 1768) Viperidae Vipera berus (LINNAEUS, 1758) Vipera ursinii (BONAPARTE, 1835)
being 326 m a.s.l. (KJ-HSMATOV & AICHME- The mosaic of landscapes in the South TOv 1984). Climate in RB is continental with Urals is very diverse. The region is charac- substantial differences between seasons and terized primarily by its transitional character regions (mountainous parts, Fore- and from low grasslands to mountain forests. Trans-Urals). RB is one of the most conti- But a variety of habitats including swamps nental regions in Europe: for example, the (both lowland and mountain), grassland, annual range of temperatures is up to 87 °C woodland thicket and colonizing scrub is (from minimal -50°C in January to maximal also represented. +37°C in July). The Ural Mountains great- The Ural Mountains are of Paleozoic ly influence climate, landscape, vegetation origin. Until the Late Oligocene, the South and other environmental parameters and Ural Mountains were relatively low with cause great differences between mountain- average heights around 300-400 m a.s.l. ous regions and their western and eastern Their modern appearance and the present slopes. Humid air masses from the Atlantic day heights were obtained only in the Ocean cannot cross the Ural Mountains and Cenozoic Era after Neogene (Miocene) tec- thus, precipitate over the western slopes. tonic activity (YAKCHIMOVICH 1992). Thus, the climate of eastern slopes and the Trans-Ural region is much more arid and Modern reptilian fauna of Bashkortostan continental with very hot summers and rather cold winters, however with very little The modern reptilian fauna of Bash- snow coverage. Cold air masses from the kortostan consists of 10 widespread species Arctic Ocean and dry air masses from including one turtle, three lizard and six Kazakhstan also influence the climate con- snake species (KHABIBULLIN 2001a) (table ditions of the region. 1); species endemic to RB are not known.
RESULTS
A summary of bones and tiny unidentified fossil particles" paleontological data (SUKHOV 1972a: 137). Remnants of five reptile species were identified; all five are Late Cenozoic reptile remnants from widespread common recent species. Due to ten excavation sites were analyzed (fig. 1). lack of quantitative analyses in the relevant Quaternary reptilian fossil materials were publications a species list is all that could be found mainly in caves (eight out often sites). compiled (table 2). The main source of bone concentra- Fossil records of Emys orbicularis tions were owl fecal pellets which were have not been described from the territory "practically teemed with lizard scutes, frog of RB so far (KHABIBULLIN 2001b) although ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at
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Table 2: Fossil reptile remains of the late Cenozoic from the area of the Republic of Bashkortostan (South Urals, Russia) Tab. 2: Fossile Reptilienreste des späten Känozoikums aus dem Gebiet der Republik Baschkortostan (Südural, Rußland). Cave - Höhle.
Epoch Locality Fossil reptiles found Literature Epoche Fundort Fossilfunde von Reptilien Schriften Holocene Zapovednaya (cave) A. fragilis, Z. vivipara, V. berus SATAEV & MAKAROVA (1997) Zapovednaya II (cave) A.fragilis, Z.vivipara, N.natrix, V.berus, YAKOVLEVA(2002) Unidentified snakes Ziganskaya (cave) Unidentified lizards, snakes / SUKHOV(1978) unbestimmte Eidechsen, Schlangen Lemeza (1) (cave) A.fragilis, N. natrix, V. berus YAKOVLEVA(2002) Lemeza (li) (cave) N. natrix, V.berus YAKOVLEVA(2002) Lemeza IV (cave) A.fragilis, N. natrix, V. berus YAKOVLEVA (2002) Nukatskaya (cave) A. fragilis, Z. vivipara, N. natrix, V. berus YAKOVLEV et al. (2000) Pleistocene Krasny Bor A.fragilis, L. agilis, N. natrix, V. berus(?) SUKHOV (1972b); CHKHIKVADZE & SUKHOV (1977) Kyzyrbak (cave) Unidentified lizards, snakes / SUKHOV (1978) unbestimmte Eidechsen, Schlangen Eopleistocene Akkulaevo Lacerta sp. SUKHOV 1972a
they are abundant in numerous excavations VADZE et al. 1983; ZERO VA & CHKHIKVADZE of Western and Central Europe, Ukraine, Cau- 1984). casus, and Central European Russia (ROZH- In the South Urals, fossil Lacerta DESTVENSKY & TATARINOV 1964), i.e. within agilis were found in Krasny Bor (Late Plei- the limits of the turtle's present day geo- stocene). Fossils of Zootoca vivipara were graphical range. found in the Holocene sediments of the Za- In the South Urals Anguis fragilis is povednaya cave (SATAYEV & MAKAROVA known from Late Pleistocene (near Krasny 1997), Nukatskaya cave (YAKOVLEV et al. Bor: SUKHOV 1972b [age re-determined by 2000) and Lemeza IV cave (YAKOVLEVA A. G. YAKOVLEV 1996]), Holocene (Zapo- 2002). Lacerta agilis evolved in the Early vednaya cave: SATAYEV & MAKAROVA 1997; Pliocene (KALYABINA et al. 2001) within the Nukatskaya cave: YAKOVLEV et al. 2000; Za- Caucasus region (YABLOKOV 1976). Earlier povednaya II, Lemeza II caves: YAKOVLEVA information on fossil Lacerta found in Kaz- 2002) and Late Holocene (Lemeza I, IV akhstan remained unconfirmed (CHKHIK- caves: YAKOVLEVA 2002) excavations. Fos- VADZE et al. 1983). Possible explanations sil remnants of A. fragilis are abundant in could be (KHABIBULLIN 2002): the recent L. Quaternary excavations in the European part agilis and Z. vivipara, currently distributed of the former Soviet Union (ROZHDEST- widely across Northern Kazakhstan, did not VENSKY & TATARINOV 1964). colonize Kazakhstan before the post glacial In RB unidentified fossil Lacerta sp. period; or both these species were not abun- remnants were found (SUKHOV 1972a) in dant and just not represented in examined the Akkulaevo excavation site (Eopleisto- fossil materials. cene). In the former USSR, fossils of the From RB, fossil bones of Natrix natrix lizard genus Lacerta are known from the are known from the Late Pleistocene (Kras- Pliocene of the Ukraine (ROZHDESTVENSKY ny Bor excavation: SUKHOV 1972b; CHKHIK- & TATARINOV 1964), the Miocene of the VADZE & SUKHOV 1977) and from the Middle Caucasus, the Eopleistocene of the Perm (Nukatskaya cave: YAKOVLEV et al. 2000; Za- region (SUKHOV 1975), the Low Pleistocene povednaya II and Lemeza II caves: YAKOV- of the Nizhny Novgorod region (RATNIKOV LEVA 2002) and Late (Lemeza I and Lemeza 1998) and the Pleistocene of the Belgorod IV caves: YAKOVLEVA 2002) Holocene. region (RATNIKOV 1988). In the Pliocene, Fossil Colubridae are known from lacertid lizards had a wide distribution in Quaternary excavations of Moldavia, Ukra- the region north to the Black Sea (CHKHIK- ine and European Russia (ZEROVA & CHKHIK- ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at
On the Quaternary reptilian fauna of Bashkortostan (South Urals, Russia) 103
VADZE 1984). Fossils of Eopleistocene Ela- the most important biological structures to phe dione are known from Poland (KHO- evolve in the Miocene was grass. ZATSKY 1982), from the Pleistocene of the In the Pliocene, continentality and Crimea (ZEROVA & CHKHIKVADZE 1984), aridity increased greatly in the southeastern and from the Low Pleistocene of the Nizhny part of the Russian (East-European) plain. Novgorod region (RATNIKOV 1998), far to Coniferous mountain forests with patches of the north of the present geographic range of open grass and water/marsh vegetation the species. Fossil Natrix tessellata is dominated. The Lower Pliocene sediments known from the Pleistocene of the Crimea were very rich in grass and bush pollen (ZEROVA & CHKHIKVADZE 1984), and the (NEMKOVA 1981), which indicates the pres- Pleistocene of the Belgorod region (RATNI- ence of open grasslands. The basis of mod- KOV 1998), fossil records of N. natrix from ern vegetation zones in the Fore-Urals was Middle Pleistocene sediments of Georgia established by the Early Pliocene. (BARYSHNIKOV & NESOV 1999), Ukraine As a result of increased water levels, (ZEROVA & CHKHIKVADZE 1984), and the the Caspian Sea ingressed twice into the Low Pleistocene of the Nizhny Novgorod South Urals region in Late Eopleistocene region (RATNIKOV 1998). (YAKCHIMOVICH 1970). The first, or Early Fossil V. berus is known from the Akchagyl ingression was chronologically Anthropogene of the Ukraine (ZEROVA & shorter but caused deeper submergence of CHKHIKVADZE 1984), from the Middle Holo- the land than the second. Simultaneously, cene (Zapovednaya cave: SATAYEV & MAKA- the boreal waters from the Arctic Ocean ROVA 1997; Nukatskaya cave: YAKOVLEV et expanded to the South Urals along the west- al. 2000; Zapovednaya II, Lemeza II caves: ern slopes of the Ural Mountains. Thus, in YAKOVLEVA 2002) and Late Holocene Early Eopleistocene, the Ural region was (Lemeza I, IV caves: YAKOVLEVA 2002) of cut off from the Russian Plain for a short South Urals.. Fossil Upper Pleistocene rem- period. The maximal sea ingression into the nants supposedly belonging to V. berus were South Urals occurred in the Middle Akcha- found in Krasny Bor (SUKHOV 1972b; gyl. Along the valleys of the ancient river CHKHIKVADZE & SUKHOV 1977). systems waters invaded the South Fore- Fossil Viperidae (no species indicated) Urals: one arm from the north-west along are known from the Low Pleistocene of the the Paleo-Belaya river basin, the other from Nizhny Novgorod region (RATNIKOV 1998). the south along the Paleo-Sakmara and its Vipera ursinii is geochronologically older tributary river valleys. These two arms than V. berus (GARANIN 1983), which had were demarcated by the hills of the Obschiy possibly evolved only during the Ice Age in Syrt (YAKCHIMOVICH 1965). Thus, the Central Europe in Romania (NIKOLSKY South Ural was cut off from the Russian 1947; CARLSSON 2003; CARLSSON et al. Plain in the Northwest, Southwest and 2004). South. However, the land connection was soon reestablished. A review of Late Cenozoic landscapes, During the Pleistocene when much of climate and vegetation the world's temperate zones were alternate- ly covered by glaciers during cool periods The landscapes of modern appearance and ice-free during the warmer interglacial in the South Urals started development by periods, climate fluctuations had a striking the end of the Miocene (YAKCHIMOVICH impact on the floras and faunas. MONIN & 1992). At the same time (Late Miocene) the SHISHKOV (1979) found that there were four present-day two-rivered orography net consecutive glaciations in East Europe (fig. established: waterflows from western and 2). According to these authors the Dniper southern mountain slopes were part of the glaciation, maximal for the Russian Plain, Paleo-Volga and Paleo-Ural river basins was not so for the Ural: in this region the whose waters flowed into the Caspian Sea, first Ice period, Oka - about 18,000 years while the waters from the eastern slopes ago, had the biggest southward expansion were part of the Paleo-Ob river basin whose when the Oka ice sheet had spread as far as waters flowed into the Arctic Ocean. One of 55° N to the south, i.e., all northern part of ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at
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30
50
Fig. 2: Scheme of the southern limits of peistocene glaciations in the area of the Russian plain (according to MONIN & SHISHKOV 1979). 1 - Oka; 2 - Dniper; 3 - Moscow; 4 - Valday. Abb. 2: Schematische Darstellung der Südgrenzen pleistozäner Vereisungen auf dem Gebiet der Russischen Tiefebene (nach MONIN & SHISHKOV 1979). 1 - Oka; 2 - Dnjepr; 3 - Moskau; 4 - Valdai.
the Republic of Bashkortostan was covered During the following epoch, the Holo- by ice. cene, the climate in the South Urals was The south Ural Mountains and foot- slightly warmer than during the Pleistocene. hills were not glaciated even at the time of In the Boreal, the coniferous forests, espe- maximal (Dniper? Oka?) glaciation; when cially pine forests, became dominant. Global there were small ice caps only on the high- warming and decline of continentality in the est mountain peaks during the Middle and Early Holocene favored forests develop- Late Pleistocene (YAKCHIMOVICH 1965). ment, although in the Fore-Urals for the most Very typical for the Pleistocene were of the Holocene as well as for interglacials, widespread birch forests with patches of coni- the notable role of broad-leaved woods as ferous and broad-leaved woods; to the south, compared with the Russian Plain has been the area was dominated by open grass vege- reported (NEMKOVA 1981; 1992). During the tation with patches of gallery forest along the climate optimum of the Atlantic (with the cli- streams. Vegetation in the South Urals and mate conditions slightly warmer than in adjacent territories was very scarce with modern times), in medium height Ural fungi dominating, as is typical for periglacial mountains the broad-leaved forests became steppes (YAKCHIMOVICH 1965). more abundant. For the Sub-boreal of the Pleistocene biota were extremely sim- south, lime forests and increased portions of ilar to modern ones - many genera and even birch and broad-leaved forests have been species of Pleistocene reptilians existed in reported. In the Sub-Atlantic, the vegetation that period. patterns resembled those of today; the fir- ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at
On the Quaternary reptilian fauna of Bashkortostan (South Urals, Russia) 105
pine woods with lime, oak and elm patches broad-leaved forests, the latter transformed and the mainly woodless spaces emerged at to the modern type flora (YAKCHIMOVICH the very end of the Sub-Atlantic. In the 1992). The entirely modern flora in the Holocene there was a general vegetation suc- South Urals has been formed by Apsheron cession from cold to warm steppe-forests. (i.e. in Pre-Ice Age!) though the hypothesis So, during the Cenozoic Era in the of its Late Akchagyl origin was not fully re- South Urals, a double flora succession took jected (NEMKOVA 1981; YAKCHIMOVICH place: in the Oligocene the ancient Poltava 1992). evergreen flora was replaced by the Turgai
DISCUSSION
All recent families of the reptilian on the reptilian fauna of the Urals, as well as fauna of the former Soviet Union had on that of the whole Palearctic. Even already evolved by the end of the Paleogene though the ice sheets did not cover the terri- (BAKRADZE & CHKHIKVADZE 1988). The tory of the South Urals, the periglacial cold substantial Middle Cenozoic cooling caused zone and ice-related events spread over the shrinkage of the geographic ranges of the whole Ural Mountain region (VELICHKO thermo- and hygrophilous reptiles. It is be- 1975). By the end of the Valday glaciation lieved that, during the Late Eocene - Early the southernmost edge of permafrost ex- Oligocene the most significant changes tended southward as far as 46° N. More- occurred in the reptilian fauna of Kazakh- over, the regions of formerly only seasonal stan, Caucasus and Europe (CHKHIKVADZE et Siberian-like congelation now reached the al. 1983). Caucasus and the North of the Caspian Sea Unlike modern herpetofaunas of Cent- (GERASIMOV & VELICHKO 1982). This ral Asia and Kazakhstan, which most likely means that the whole East European Plain started to form no later than in the Middle and a major part of the West Siberian Plain Oligocene - Early Miocene (CHKHIKVADZE were frozen lands. et al. 1983), the modern herpetofauna of the In the territory of European Russia, a Urals must have begun formation in the sophisticated complex of surface structures Late Pliocene (KHABIBULLIN 2002). All of cryogen origin with dominating large 20- genera and most recent reptile species 30 m long and 4-5 m broad crack systems which today represent the North Eurasian had evolved at this time. To the east of the herpetofauna, had already evolved by the Ural Mountains, in the territory of the South Early Pliocene. Trans-Urals and West Siberia "...cryogen Despite the fact that Lacertidae are the transformations like involutions and ice only fossil reptiles known from the Pliocene clines occurred frequently and served .... as of Bashkortostan (KHABIBULLIN 2001b), an indicator of long-term frozen ground and fossil materials of other widespread and severely cold climate" (STEPHANOVSKY & numerous present-day families (Anguidae, MALEEVA 1977). On the other hand, in some Colubridae and Viperidae) can be expected microhabitats on the western slopes of the to be found. Until now there are no records South Ural Mountains the nemoral forest of Pliocene-Holocene reptilian fossils vegetation and broad-leafed floras managed which differ from recent taxa and also no to survive even during the very cold peri- records on other fossil reptiles that inhabit- ods. Despite this fact as well as the evi- ed the territory of Bashkortostan in the past. dence of fossil findings of underground The paucity of paleontological materials on dwellers like the mole-rat Myospalax the South Urals Quaternary reptiles does not myospalax LAXMANN, 1773, the poikilother- allow for more comprehensive conclusions. mic reptiles did not survive these cold cli- The gradual cooling in the Pliocene - mate conditions, and disappeared from most Eopleistocene and particularly the Pleisto- parts of Siberia, the Urals and the Russian cene Ice Age (with the Oka Ice Age as a Plain (NIKOLSKY 1947), as well as North probable exception) had a striking impact Europe. ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at
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The impact of Pleistocene climatic mate became warmer some 12,000 to 8,000 changes on the fauna of Europe was sub- years ago in the Pre-Boreal period of the stantial: The low temperatures during the Holocene (KRIVENKO 1991). This expan- glacials displaced most thermophilic groups: sion significantly accelerated during the Reptile life was heavily affected; many Holocene climate optimum. The main species became extinct or lost large parts of migratory flow of reptiles that now inhabit their former ranges (LENK et al. 1998). The Central and North Europe and Siberia, was situation was aggravated by the existence of directed from réfugia in the Caucasus, and natural barriers, preventing the animals from South and South-Eastern Europe towards escaping to the South. Such barriers were North and Northeast through the East- the Mediterranean, Black and Caspian Seas, European Plain and Ural Mountains as far the Balkans, Caucasus and Central Asian as Siberia. Analysis of the clinal variability mountain chains as well as arid areas in of several morphological parameters in rep- Central Eurasia. Only species that could tilian species of the temperate zone support- survive in one of various restricted however ed this hypothesis (YABLOKOV 1976; ANISI- climatically favoured areas ("glacial réfu- MOVA 1981). gia") in the southern extremities of Europe In the Sub-Boreal and Sub-Atlantic and adjacent regions were saved from ex- Holocene periods, after slight cooling of the tinction (LATTIN 1949; HEWITT 1996). After climate and subsequent retreat of most ani- analyzing data on birds' paleontology and mals' geographical ranges, the climate con- biogeography, ornithologists proposed the ditions became relatively stable (KRIVENKO existence of several such réfugia in Central 1991). and Eastern Asia (KRIVENKO 1991). As one can see from atlases of reptil- Hence, the recent presence of reptiles ian geographical ranges, all reptile species in Eurasia north to a line which is roughly inhabiting the western slopes of the Urals, represented by the 42nd parallel must have can also be found on their eastern slopes. been caused by postglacial migration/colo- So the mountainous landscapes of the Urals nization events (STUGREN & KOHL 1980; as a physical phenomenon did not globally COOPER et al. 1995; TABERLET et al. 1998). influence the reptilian post-glacial disper- The sequence of glaciations and intergla- sion and colonization process in North cials made the animal's geographic ranges Eurasia, though mountain landscapes do repeatedly shrink and expand. However, play a significant role on the microhabitat the fact that fossil reptilian's remnants have level. been found in temperate Eurasia sediments Great forest complexes remained un- support the idea of relatively warm local cli- changed only in the mountainous part of the mate during Pleistocene interglacial periods South Urals, unlike the steppe regions that (RATNIKOV 1996). are mainly agriculture lands now. The For the Eastern European herpetofau- mountains of the South Urals and, conse- na the most crucial réfugia must have been quently, their forests extend southwards into those in the Balkans, Caucasus, Asia Minor the steppe zone. This fact favors the south- and probably in Central Asia (KHABIBULLIN ward dispersion of boreal species like Zoo- 2001a, 2002). From these réfugia started toca vivipara and Vipera berus. However, the inter- and postglacial dispersion into the because of human activities, these condi- present day distribution areas. tions favorable for forest species evolved As to originally widespread species, only in the most recent past. Intensive agri- practically each refugium gave origin to its cultural, farming and forestry activities own subspecies. The diverse subspecies mainly affected the plain regions, not the structure of several modern European rep- mountains. On the other hand, the Ural tile species with vast geographic range like Mountains form the northern border of the Lacerta agilis (see KALYABINA et al. 2001) geographic range of several reptile species or Emys orbicularis (see FRJTZ 1998) sup- like Emys orbicularis, Elaphe dione and ports this idea. Vipera ursinii. For these species montane The dispersion from glacial réfugia temperate zone environmental conditions was induced when glaciers melted and cli- are unfavorable. The Ural Mountains are ©Österreichische Gesellschaft für Herpetologie e.V., Wien, Austria, download unter www.biologiezentrum.at
On the Quaternary reptilian fauna of Bashkortostan (South Urals, Russia) 107
barriers between the water basins of Volga South Urals. The whole sum of anthro- River, Ob River and Ural River, and from pogenic impacts on environment and local this point of view they influenced the distri- biota is very diverse, aprogressively bution patterns and played a particular role increasing and by nowadays very intense. in the post-glacial colonization process of This primarily results in natural landscape hydrophilous reptiles (i.e. N. natrix disper- transformations, habitat fragmentation, sion from the Caspian sea northward along diminished biodiversity and reduction of the Ural river). number of suitable environments. The first humans arrived at the South At least in some reptile species, the Urals not before the Late Holocene. In recent range regression has been caused Bashkortostan the earliest known archeo- and, possibly, post-glacial colonization logical camp is Mysovaya near Lake Kara- process in Central and North Europe has balykty in the Trans-Urals that dates back to been influenced not only by suboptimal the Early Paleolithic (SHAKUROV 1996). conditions of their natural environment but During the Paleolithic and Mesolithic the also, more important, direct and indirect dis- so-called "consuming" economy (i. e. hunt- turbances by man (e. g. Emys orbicularis - ing, fishing and foraging) dominated. In the see FRITZ 1998). As a consequence the Neolithic (6,000-4,000 years BC) in the northern distribution limit of reptiles in South Urals the transition from "consum- Europe is not a natural one. Instead, it re- ing" to "productive" economy (i. e. agricul- flects impacts over the course of thousands ture, cattle breeding and fish farming) took of years in a heavily disturbed environment. place. The intensive colonization of the The patterns of larger-scale Holocene rep- South Urals territories dates back to the tile colonization (for example, migratory Bronze Age when the first stationary settle- routes) were less affected by man in Eastern ments were established. Until the 17th-18th Europe because it is much less populated as century the main activities of semi-nomadic compared to Western Europe. However, the local Bashkir people remained the cattle local distributional "mosaic" is to a consider- breeding and early apiculture. Only by the able degree the result of human, mostly agri- end of nineteenth century the thoroughly cultural, activities. settled way of life spread over the whole
ACKNOWLEDGMENTS
I thank A. G. YAKOVLEV (Laboratory of Regional Scientific Centre of the Russian Academy of Science) Geology and Geophysics, Institute of Geology, Ufa for consultations in the paleontological part of the paper.
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DATE OF SUBMISSION: November 19, 2003 Corresponding editor: Heinz Grillitsch
AUTHOR: Viner KHABIBULLJN, Faculty of Biology, Bashkir State University, Ufa, 450074, Russia [email protected] >