Eastern European mammoth distribution and environments during the Middle Valdai Briansk Interstade (33,000-24,000 BP)

A.K. Markova1, A.N. Simakova2, A.Yu. Puzachenko1, L.M. Kitaev1 1Institute of Geography of the Russian Academy of Sciences, Moscow, Russia [email protected], [email protected], [email protected] 2Geological Institute of the Russian Academy of Sciences, Moscow, Russia [email protected]

SUMMARY: One of the most remarkable intervals within the Valdai Glaciation was a time of significant warming – the Briansk Interstade (33,000-24,000 yr. BP); there are abundant palaeotheriologic and palaeob- otanic data, including mammoth finds, available for this interval. The principal goal of this study was an inte- grated analysis of the extensive database with respect to main features of mammoth environments during this Interstade. For this purpose we used mathematical as well as traditional research methods. Information on and plant species composition, geology and geographical position of the sections as well as absolute and relative ages of localities has been included into databases. All data were organized in PARADOX software and then moved to ARC/VIEW GIS; the latter formed the basis for construction of electronic maps of indicator mammal and plant ranges and recognition of the principal biomes.

1. MATERIAL drainage basin (~60°N) in the north to the lower Dniester drainage basin (~ 48°N). They 1.1 Mammal data were found in the middle courses of the Dnieper and Don (~ 50 – 52°N), in the Kama 45 Briansk mammal sites have been included basin (~ 55 –60°N) and in others regions (Fig. into this study; most of them belong to cultural 1). It is possible that mammoth lived farther layers of Palaeolithic sites and are dated by north, but we have not enough information for radiocarbon. One site corresponds to mole- and Subarctic zones at present. No site courses of the Briansk soil, another to fluvial of this age is known in the Crimea either. In the deposits. from five sites have been early Valdai time the range of mammoth was studied by one of the authors (Markova 1982; wider and included the Crimean Peninsula. Markova et al. 1995). A few primary literature sources were also used, such as papers by 1.3 Paleobotanical data Alexeeva (1990); Vereshchagin and Baryshnikov (1980), Guslitser and Kanivets During the last decades, a number of (1965), Kalinovski (1983); Rekovets (1985), researchers, among them. Artushenko (1973), Rogachev et al. (1981), Tatarinov (1977) and Bolikhovskaya (1995), Gurtovaya (1985), others. Spiridonova (1991) and others, published detailed palynological materials for the 1.2 Mammoth finds Briansk Interstade on the Russian Plain; they are supported by radiocarbon dates and by Briansk mammoth remains are widely spread geological data. over the East European Plain, from the Kama 52 plant remain localities of the Briansk time 299 The World of Elephants - International Congress, Rome 2001

Fig.1 - The sites with Mammuthus primigenius remains (33 – 24 ka).

have been studied (Simakova & Kozharinov lists of mammals have been established for 1995). The palynological data reveal a compli- every class of localities. The results of this clas- cated vegetation and climatic history during the sification were analyzed both in artificial MDS Briansk Interstade. dimension space, and in geographical physical space according to the coordinates (latitude, 2. METHODS OF THE ENVIRONMENT longitude). The palaeofloristic data were classi- RECONSTRUCTION fied analogously, but on the basis of Euclidian distance matrix. The distances between the The principal goal of this study was to recon- sites were calculated for logarithm spectrum. struct the mammoth environment using all The distinguished classes were characterized available data. The integrated analysis of mam- by mean values of pollen spectra. The results of mal and plant data permitted to elucidate the pollen analysis reflect their relative appearance biogeographical situation on the Russian Plain (maximum value for a sample, in %) for 20 during the Briansk Interstade warming. The most widespread species, genera and also taxa lots of theriological and botanical data of the of higher order. At the second phase the limits Briansk Interstade were jointly analyzed. of biomes were defined using results of inte- Mathematical methods have been used for grated analyses of these classifications, these aims (Markova et al. in press). At the first obtained for mammals and plants, and with stage of analysis the localities were classified particular reference to geographic position of by mammal and plant composition. The initial all distinguished groups of sites. theriological materials were presented as matrix of “presence - absence” of 46 genera (58 3. THE RESULTS OF CLASSIFICATION OF species) of mammals which have been recov- THERIOLOGICAL AND FLORISTIC DATA ered from the Briansk deposits. The squared dissimilarity matrix was calculated (using the Two groups of localities with palynological Jakar distance) for 45 mammalian sites located materials have been distinguished as a result of in 19 geographical points. The cases in which classification (I, II). Each of these groups both species were absent have been ignored includes two sub-groups. Their ordination in during the matrix calculation. Then we ana- the space of MDS axis could be interpreted as lyzed this matrix for reproducing the distances, follows: sites of the first group (I, sub-groups 1 based on several underlying dimensions. The and 2) feature relatively high proportion of 300 Eastern European mammoth distribution and environments during Middle Valdai Briansk Interstade... cold-loving and moisture-loving forest taxa; 4.1 Forest-tundra sites of the second group (II, sub-groups 3 and 4) include high amount of xerophytic steppe A combination of tundra and steppe plant and semi-desert taxa and also some broad- communities with “islands” of Pinus-Betula leaved trees (sub-group 4). open forest and with the assemblage of sub-arc- Five groups of mammal localities have been tic mammals (, polar fox, pied and true distinguished as a result of classification. The lemmings, and narrow-sculled voles). first group includes the most isolated sites. Mammoth was very rare here. Possibly, the bio- The second and forth groups are the most mass in the north of Eastern Europe was not suf- remote from the first one. The third group lies ficient for subsistence of this large herbivore. closely to the forth one but very far from the second group. The fifth group occupies an 4.2. Periglacial forest-tundra-steppe intermediate position in relation to all the rest groups. A mosaic of plant communities, including Betula-Pinus and Picea forested “islands”, meadow and halophytes steppes, and tundra 4. PRINCIPAL BIOMES DURING BRIANSK vegetation patches. Mammal assemblages con- INTERSTADE sisted of tundra and forest-steppe species (, primitive bison, wild ox, The following biomes could be reconstructed reindeer, red deer, roe deer, horse, saiga, from north to south based on the described wolverine, polar fox, ground squirrel, common above procedures (Fig. 2). Mammoth Mam- hamster, pied and true lemmings, narrow- muthus primigenius lived in most of them: sculled, red-backed and field voles and others).

Fig.2 - Biogeographical provinces on the Russian Plain and the Crimea during the Briansk Interstade after botanical and mammal data. I. Forest-tundra, II. Periglacial forest-tundra-steppe, III. Periglacial forest-steppe, IV. Periglacial steppe, V. Forest-steppe of plains (A), and highlands and mountains (B). 301 The World of Elephants - International Congress, Rome 2001

The environments with sufficient quantity of 5. CONCLUSIONS biomass favored the mammoth and other large herbivore existence in this zone. The reconstructed biomes reflect specific environmental and climatic conditions of the 4.3 Periglacial forest-steppe Briansk Interstade where mammoth found favorable habitats. The analogues of modern Meadow-steppes with Pinus-Betula forest- natural zones did not exist in this time on the ed “islands”, with scarce broadleaved trees, Russian Plain. The reconstructed landscapes and with forest-steppe assemblage of mam- reflect moderate-cool climate of this period, mals (woolly mammoth, woolly , when the majority of northern subarctic plant primitive bison, wild ox, reindeer, red deer, and mammal species extended their ranges roe deer, horse, saiga, cave carnivores, bobac considerably to the south and was distributed marmot, ground squirrels, great jerboa, not only in the Northern, but also in the Central steppe lemming, narrow-sculled and field Russian Plain. At the same time, steppe species voles, and others). The environment of this penetrated farther north and west, which sug- biome was beneficent for Mammuthus primi- gests a destruction of continuous forest zone genius. Mostly open landscapes with high during the Briansk Interstade. Fragments of biomass were favorable for mammoth, primi- forests persisted in topographically diversified tive bison, woolly rhinoceros and other large areas with many local habitats (in highlands, herbivores. The density of mammoth sites is mountains, and river valleys). The Briansk bio- the highest in this area. The mammal species mes were similar to modern ecotones in their richness is also highest among the plain high diversity. Earlier Guthrie, Graham, regions of Eastern Europe. Lundelius, Semken and others have noted sim- ilar features of periglacial biota in North 4.4 Periglacial steppe America. The differences between East European bio- Forb steppes with the steppe mammal assem- geographical provinces were smoothed during blage (woolly mammoth, woolly rhinoceros, the Briansk time: subarctic plants and mam- horse, saiga, bobac marmot, ground squirrels, mals were present practically in every of them. great jerboa, greater mole rat, northern mole- Forest species, more sensitive to warmth and vole, steppe and yellow lemmings, and others humidity, were not abundant on the Russian and others including reindeer). Drier conditions Plain. However, the periglacial forest-steppe of this biome determined lower biomass with and forest-tundra steppe were more extensive respect to the previous one. This accounts for during the Briansk time than at the Late Valdai lower quantity of mammoth localities in these Maximum. That indicates rather temperate cli- regions. mate of the Briansk Interstade. The Briansk climate undoubtedly was more 4.5 Plain (A) and mountain (B) forest-steppe severe than recent one, with lower winter tem- peratures and low precipitation (Frenzel 1992). Betula-Pinus forests areas with few Several factors had the major effect: the exis- broadleaved trees and mammal assemblage of tence of small ice sheet in Scandinavia; a sig- south periglacial forest-steppe (woolly mam- nificant marine glaciation (pack ice), which moth, woolly rhinoceros, primitive bison, wild affected the moisture supply to the continent; ox, horse, saiga, giant and red deers, reindeer, wide distribution of ; and also the wild boar, steppe and yellow lemmings, yel- decrease of air masses transfer from the West to low-necked mouse, field and common voles; in the East (van Andel & Tzedakis 1996; Velichko the Crimea the assemblage lacks mammoth, but 1973). Such climatic conditions influenced the the wild ass and Microtus obscurus appearance and wide distribution of different are present. types of periglacial vegetation and “mixed” 302 Eastern European mammoth distribution and environments during Middle Valdai Briansk Interstade... mammal assemblages over the Russian Plain; 7. REFERENCES many of them have no analogues in modern biota. Various types of open periglacial envi- Alexeeva, L.I. 1990. Theriofauna of Late ronments with great diversity of grass and bush Pleistocene of Eastern Europe. Moscow: vegetation were comfortable for large herbi- Nauka (In Russian). vores, including woolly mammoth. The paly- Andel van, T.H. & Tzedakis, P.C. 1996. nological studies have been performed on Palaeolithic landscapes of Europe and envi- stomach and bowels contents from the frozen rons, 150,000 – 25,000 ago: an mammoth bodies, found mostly in . It overview. 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