N. I. DROZDOV, V. P. CHEKHOV
Institute of Archeology and Ethnography SB RAS Laboratory of Archeology and Paleogeography of Central Siberia

Krasnoyarsk, Akademgorodok, 660036, Russia

E-mail: drozdov@ kspu.ru checha@ kspu.ru

Introduction

One of the phenomena of the Quaternary period was the so-called mammoth fauna. Questions of natural living conditions and the causes of the extinction of some of its representatives, including mammoths, are debatable [Vereshchagin, 1979; Verkhovskaya, 1988; Puchkov, 2001; Sher, 1997; etc.]. The Taimyr Peninsula belongs to the regions of the Russian Subarctic and Arctic that are most saturated with mammoth remains dating in a wide range to the late neo-Pleistocene-Holocene. Thus, the peninsula was one of the places of the most recent residence and subsequent extinction of mammoths.

The Quaternary period in the history of the Taimyr Peninsula has been studied in much more detail than, for example, the northern and central parts of the Central Siberian Plateau. The most significant are the results of research conducted by industry and academic institutions in the late 1930s and 1940s in connection with the development of the Northern Sea Route. These surveys involved:: Geological Institute of the USSR Academy of Sciences, Institute of Arctic Geology (NIIGA), Institute of Arctic and Antarctic Research (AARI, Glavsevmorput). Large-scale geological surveys were carried out here in the 1970s and 1980s by the production geological associations Aerogeologiya and Krasnoyarskgeologiya. Employees of the Biological and Zoological Institutes of the USSR Academy of Sciences conducted paleobotanical and paleozoological research for many years. In the 1990s, the study of the Quaternary period was continued by scientists of the AARI. In parallel with the listed works, studies were conducted on the locations of mammoth remains. Despite many years of studying the nature of Taimyr in the Quaternary period, many issues of paleogeography and paleolandscape studies remain controversial. This determines the diversity of points of view on the evolution and causes of the extinction of individual representatives of the mammoth fauna.

We believe that the habitat conditions and food resources of mammoth fauna in the Late Pleistocene - Holocene periods of Taimyr's history were determined by the following factors:: 1) the scale, form of occurrence and duration of glaciations, transgressions and regressions as powerful factors of paleogeographic changes; 2) climatic and weather indicators, including seasonal ones; 3) the nature and evolution, hydrological features of permafrost; 4) the nature of plant communities, their biological productivity, and the floral composition of phytocenoses. The aim of the work is to review the history of Taimyr nature development in the Late Pleistocene - Holocene, taking into account the peculiarities of mammoth habitat conditions.

Kazantsev time

This stage was characterized by the transgression of the waters of the Arctic basin to the North Siberian lowland with the accumulation of corresponding sediments. Sea on-

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it moved from west to east with decreasing depths in the same direction. The mountains were groups of low-and medium-altitude massifs isolated by the sea. Only in the east of the North Siberian Lowland there were coastal denudation plains, where the relief was leveled and alluvial and lacustrine sediments were formed. Partial degradation of permafrost occurs at this time. At the end of the Kazantsev stage and in the Early Rukta period, accumulative-abrasive terraces with heights of 60-70 and 80-100 m were formed as the sea receded (Strelkov, 1965). The maximum development of sea terraces was achieved in the north of Taimyr. Currently, within the North Siberian Lowland, they are recorded on the tops and slopes of structural denudation ridges. A rich and diverse diatomaceous boreal flora was developed in the lakes, and boreal and arctoboreal foraminiferal complexes were developed in the marine basin.

For the eastern part of the peninsula at this time, the following stages of vegetation cover development are reconstructed in the coastal lowlands: 1) the existence of yernik (with swamp vegetation), willow, and heather vegetation; 2) the appearance of spruce, pine, and birch; and 3) the further advance of forest vegetation northward, approximately to the latitude of Maria Pronchishcheva Bay (75° 30 ' N), i.e., to the area of the modern Arctic tundra [Berdovskaya, Gay, Makeev, 1970]. In the pools of Logaty, Bol. Balakhni (73-73° 30 'N) during the optimum period of the Kazantsev interglacial period, a forest tundra similar to the modern forest tundra of the lower reaches of the Kotui River (71° 30' - 72° N) was developed. Probably, there were areas of sparse northern taiga larch forests with an admixture of spruce, cedar, birch, and pine [Anthropogen Taimyra, 1982; Fischer et al., 1990]. At present, moss-lichen (typical) tundra is developed here. The following climatic parameters were reconstructed for the Kazantsev time: the average annual temperature is -10... -11 °C (-14 °C)* , January temperature -34 °C (-34... -35 °C), July temperature 12 ... 14 °C (6... 8 °C), precipitation 400 mm (250 mm). Thus, the climate in the Kazantsev period was wetter and warmer than today.

Murukta time

The Murukta stage is a time of preglacial regression of the sea, cooling, and ground glaciation. Experts estimate the scale of the latter in different ways. Some believe that ice at the initial, North Siberian stage covered the entire North Siberian lowland from the Yenisei to Popigai. Glaciation centers are located in the North - Siberian (shelf of the Kara Sea, Severnaya Zemlya, Byrranga Mountains), Putoransky and Anabar. The thickness of the ice sheet in the first center was more than 2 km, and the glaciostatic deflection was 300-400 m. The closing of glaciers occurred at the northern edge of the Central Siberian Plateau, in the area of the Khety and Khatanga River valleys. The maximum activity of glaciers was observed in the west of Taimyr. Since the pressure moraine ridges are fragmentary and, in general, the marginal glacial formations are poorly expressed, it can be assumed that the ice quickly lost contact with the glaciation centers and turned into masses of "dead ice". The latter, during deglaciation, provided a wide development of glacial water flows, rocks, and lakes (Strelkov, 1965). Later, at the North Korean stage (Antropogene Taimyr, 1982), ice closure of the first two glaciation centers did not occur in the eastern and central parts of the lowland. Probably, in the Murukta period, the main features of the modern relief were formed, represented by a combination of extensive lowlands, ridges, hills, lake depressions, valleys, and flat areas. From the point of view of other researchers, the Murukta glaciation was so weak and inactive that it did not have any geological impact on the underlying surface. When the ice melted, a hilly ridge relief was formed (Zagorskaya, 1961). D. Y. Bolshiyanov and G. B. Fedorov (Bolshiyanov, Fedorov, and Savelyeva, 2001) agree that ice sheets rather than ice sheets existed in the Arctic and Subarctic in the late Pleistocene. Finally, there is an assumption that the Murukta stage (80-47 KA BP) was associated with the revival of denudation and erosion processes with valley dips up to 40 m, as well as the formation of major relief features - large river valleys and depressions. In the North Siberian lowlands (basins of the Novaya, Balakhnya, Logata, and Verkh rivers). Taimyr, Khatanga) glaciers were absent; the described glacial and water-glacial deposits are traces of mid-Quaternary maximum glaciation (Fischer et al., 1990).

There is very little data on other natural conditions of the Murukta time. According to L. S. Troitsky (1966), representatives of the mammoth fauna in the west of Taimyr appeared only after the retreat of the glacier. Based on the extremely low content of spores and pollen in the corresponding sediments, and the high specific gravity of redeposited Mesozoic-Cenozoic palynoforms, we conclude that the climate is exceptionally harsh (Ukraintseva, 1991). Based on this, we can assume that for the No River basin-

* Current indicators are shown in parentheses here and below.

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The howl (72-73° N) was characterized by a polar-desert type vegetation cover, similar to the present vegetation at m. Although these features, in our opinion, may be a consequence of active processes of denudation and erosion at that time.

Karginsky time

At the Karginsky stage, sea water was again ingressed deep into the lowlands along river valleys. This process is reflected in the ancient coastlines in the lake districts. Taimyr and the Bol River. Balakhni at absolute altitudes of 50-100 m (Antropogen Taimyr, 1982). The sea depth decreased to the east, which indicates the penetration of water from the west. Lake Taimyr was part of an estuary that jutted deeply into the land (Figure 1). The Pyasina River (up to the lake of the same name) was also an estuary. In the Late Karginian period, these two bays were connected by the valleys of the Tareya and Ayatari rivers. Thus, a sharp reduction in sea ingression with the emergence of a multitude of lacustrine basins dates back to the second half of the Karginsky time (32-26 thousand years AGO). It is assumed that the lake basins developed widely outside the ingression boundaries with the formation of a complex of lacustrine, lacustrine-alluvial sediments dating from 14 C 46.3 - 25.7 thousand years ago. L. N. (Fischer et al., 1990). The sediments contain a lot of plant detritus, peat, bone remains, and diatoms, i.e. they are informative in paleogeographic terms. At this stage, the main (third) terrace with a height of 25-30 m was formed. At the end of the Karginsky period, there may have been some rearrangement of the rivers and the formation of the second terrace (15-20 m) began. The Karginsky deposits are permafrost and contain segregated vein ice. During the ingression, some degradation of the permafrost was possible (Strelkov, 1965).

In natural terms, the Karginsky period was heterogeneous. N. V. Kind (1974) and other experts identify three warming periods and two cooling periods in northern Siberia for this period. In the basins of the Zakharova Rassokha and Novaya rivers (72° 30' - 73° N), during the first warming period (50-45 thousand years ago), landscapes similar to the current rare - coniferous northern taiga with larch, spruce, birch, as well as shrubby birch, willow, and alder became widespread. In the lower tier, there are many ferns, mosses, plowshares, and from grasses - aspen, cereals, and various grasses.

The following quantitative paleoclimatic indicators were calculated for the specified time: the average annual temperature is-12 °C (-14°C), the January temperature is -34 °C (-35°C), the July temperature is up to 14 °C (6 ... 8°C), precipitation is 400-450 mm (250 mm). Currently, the area under review is located, according to some data, in the subzone of shrubby (southern) tundra, according to others - in the subzone of northern (or typical) tundra. At the optimum Karginsky time, the climatic parameters were close to the indicated ones; vakhta (Menyanthaceae), urut (Myriophyllum), and rdest (Potamogeton) were found in the water basins. For the first cold snap (45-42 thousand years ago), July temperatures dropped to 10°C and precipitation was about 400 mm (Antropogen Taimyr, 1982).

The stage structure of vegetation development in the Karginsky period in the Novaya River basin is presented by researchers of the Biological Institute of the USSR Academy of Sciences (Ukraintseva, 1991) as follows: 1) grass-mixed and sedge-grass communities (for deposits based on mammoth remains, the date is determined to 14 C: more than 53.2 thousand years ago); 2) shrub tundra (39 thousand years ago), which was later replaced by sedge-larch forests (34.7 thousand years ago, Malokhet warming); 3) larch forests low-growth tundra , shrub and moss tundra (29.8 thousand years ago), which was replaced by spruce - larch forests and by the end of the Karginsky stage (23.2 thousand years ago)-larch rare-coniferous forests (Lipovo-Novoselovo warming). A larch trunk dated to 14 C (ca. 23 KA BP) was discovered south of Maria Pronchishcheva Bay (75° 30 ' N) (Makeev, 1975). Lake basins of the Karginsky period were characterized by increased levels, abundance and diversity of diatoms during the warming epochs-

Fig. 1. Distribution scheme of the marine basin in Taimyr in the Late Rukta-Karga period (according to (Antropogen Taimyr, 1982)). 1 - modern land that was flooded in the Late Tyrian period; 2 - coastline of the Late Tyrian basin; 3-modern land that was flooded by the sea in the Karginsky period; 4-coastlines: a - Early Karginsky (50-33 Ka BP), b - Late Karginsky (32-26 Ka BP) basins.

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howling flora with the presence of South boreal species. During periods of cold weather, the lake level decreased, and the flora was significantly depleted (Cherkasova, 1981).

Thus, Karginsky time is marked by a rhythmic alternation of warmer and wetter days than now, or close to them (50 - 45; 42 - 34; 29 - 24 and colder than today, or close to them (45 - 42; 34 - 29 thousand years ago) periods. The climate situation improved due to higher temperatures in January and July and an increase in annual precipitation. The deterioration of natural conditions was determined by a decrease in July temperatures (Nikolskaya, Klimanov, Borisova et al., 1989). The shift of natural zones (forest tundra, northern taiga) was about 100-200 km; during cold spells, the landscape structure was restored close to the present one (Fischer et al., 1990).

Sartan Time

This is the time of the last significant cooling event in the late Pleistocene. There are polar opinions about the size of the last Late Pleistocene glaciation (Figure 2). According to one, the glaciation was localized within the Putorana Plateau, while the other, it covered the north-west of the Central Siberian Plateau, the North Siberian Lowland, the Byrranga Mountains, and the Kara Sea shelf, and generally recognized a small thickness of ice on the peninsula with a sharp reduction glaciation (as for previous epochs of cooling) from west to east. One of the proofs of the above is the absence of signs of significant glaciostatic deflection and subsequent uplift in the Holocene in Taimyr.

During the Sartan period, lakes existed in the central and eastern parts of the North Siberian Lowland in the relief depressions. Their formation, according to some experts, is associated with the damming of rivers that had a flow to the north and west. Maximum of lake transgression, when lakes could merge to form large basins (for example, the hypothetical lake basin Pra-Labaz, a relic of which today is Lake Baikal). Labaz) was evidently associated with the deglaciation of glaciers (Antropogen Taimyr, 1982). From the point of view of other researchers, the lake cover of the North Siberian Lowland was high and permanent at all stages, which is typical of accumulative lowland plains in permafrost areas.

The disintegration and deglaciation of Sartan ice occurred earlier than 16 thousand years ago. At the beginning of the stage, the level of oz. Taimyr was high due to the sub-damming of the local glacier that blocked the valley of the Niz River by a glacial dam. Taimyrs in the area of the Shrenk River - its left tributary. Later, 16.8 Ka BP, the lake level dropped catastrophically much lower than the present level (Bolshiyanov, Fedorov, Savelyeva, 2001). The second floodplain terrace of the main rivers began to form with the revival of the erosion incision of about 16 thousand years ago. Floodplain deposits of the second terrace of the Mamonty River (Northern Taimyr) with the remains of the "Taimyr mammoth"are dated to 14 from 11.5-11.7 thousand years ago. Permafrost continued to persist throughout the Sartan stage with the formation of syngenetic underground ice, polygonal structures, etc.

The Sartan deposits are characterized by an abundance of mammoth, horse, deer, bison, and musk ox bones, although there are few plant remains compared to the Kargian period. Hence, in Sartan times, the climate conditions were probably much more severe than at present. Numerous Aeolian landforms outside the lake basins also indicate that the climate is very dry (Antropogen Taimyr, 1982; Fischer et al., 1990). Thus, in the valley of the Khety River (below the mouth of the Boyarka River), in the fossil spectra of deposits for which the

2. Scheme of distribution of the Sartan glaciation in Taimyr and adjacent territories (according to [Razvitie landshafts..., 1993]). 1 - the boundary of the maximum area of cover glaciation (the teeth are directed to the extraglacial zone); 2 - the boundary of the minimum area of cover glaciation (the teeth are directed to the extraglacial zone); 3 - the territory of reticulated, semi-covered glaciation with firn fields and low-power, inactive ice caps.

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date to 14 S: 10860 ± 80BP (GIN-674), the leading role is played by pollen and spores of sedges, mosses with an admixture of pollen from cereals, Poland, and isolated shrubs (birch, willow). This makes it possible to consider the vegetation as typical tundra (currently forest-tundra is widespread here) [Nikolskaya et al., 1980], and temperatures are lower than today's [Nikolskaya, Borisov, Kaplyanskaya et al., 1989]. At the same time, for the end of the Sartan stage, plant groups close to modern ones are reconstructed, and late glacial cooling events (Late Dryas or Norilsk stage, 10.2 - 10.9 Ka BP) are not recorded to the south. For example, for the basins of the Bol Lesnaya Rassokha and Zakharova Rassokha rivers (72-73° N), these are sedge - grass, grass-mixed associations, moss tundra, larch forests (about 10.5 thousand years ago), and for the Bol River basin. Balakhnya (73°30' N) - shrubby and shrubby moss tundras (ca. 10.5 thousand years ago), for the Mamont River basin (74° N) - moss and downy-seak tundras with polar willow (ca. 11.5 thousand years ago) [Ukraintseva, 1991]. It should be noted that currently these river basins are located in the subzones of the southern (Bol. Lesnaya Rassokha River), typical (Bol. Lesnaya Rassokha River) and southern (Bol. Lesnaya Rassokha River) subzones. Bol. Balakhnya) and Arctic (Mamont River) tundras. Direct evidence of the existence of woody vegetation in the basin of the Novaya River (Bol Lesnaya Rassokha River) during the Norilsk stage of the Sartan cooling is provided by the larch stumps preserved in the second above-flood terrace in the lifetime state. According to the absolute age of the wood of one of the stumps, larch grew 10500 ± 500 BP (IM-671), i.e. at the very end of the Sargan, in the Late Dryas (Belorusova, Lovelius, Ukraintseva, 1987). There are other data on warming in northern Asia earlier than the Holocene. For example, on Sverdrup Island in the Kara Sea (100 km from the coast of Taimyr), sedge peatlands were formed between 11640 ± 40 and 9770 ± 280 Bp (Tarasov et al., 1995). This allows us to consider the climate for the specified period of time warmer than it is now.

Interesting conclusions were obtained from the study of the isotopic composition of the re-vein ice of m. Saber on the lake. Taimyr (Derevyagin et al., 1999). Ice formation here lasts from the Karginsky period to the present time with small interruptions (about 27 and 12 thousand years ago), fixed by thick peat-mineral interlayers. About 30 thousand years AGO (obviously, during the Konoshchelsky cold snap), the average winter temperatures dropped by 9°C below the current ones. In the Sartan period (18-12 KA BP), the average winter temperatures gradually increased from -29 °C to -25 °C; in the Holocene, they were close to the present (-23 ° C). According to D. Y. Bolshiyanov and G. V. Fedorov (2001), 13.5 - 9.7 Ka and at the optimum Karginsky time (41-35 Ka), the environmental conditions in Taimyr were fundamentally similar and, judging by the abundance of mammoth bones in the sediments of these intervals, the most favorable for the life of these large mammals.

Holocene

The boundary between the Late Pleistocene and Holocene in absolute chronology is usually considered to be 10.2-10.3 Ka BP (Kind, 1974; Khotinsky, 1977). In the Holocene, the first terrace of large rivers of Taimyr (10 - 7 thousand years ago) and their high floodplain (6.5 - 4.5 thousand years ago) were formed, low floodplain, lake and alas deposits were formed, thermokarst, cryogenic and slope processes actively developed. Peat formation continued, especially at the sites of the Karginsky terraces. In Holocene deposits of various ages (alluvial, lacustrine, slope), numerous plant remains are common. Their maximum concentration was recorded in the east of Taimyr in sediments dating from 7 to 5 thousand years ago (Strelkov, 1965; Antropogen Taimyr, 1982). During the subboreal cold snap, glaciers in the Byrranga Mountains became more active. It is assumed that the glaciation at this time was 8-10 times higher than the present one. As already noted, peatlands formed on Sverdrup Island between 11.6 and 9.0 Ka BP. Based on this, it can be assumed that the vegetation on the island was approximately the same as in the zone of typical tundras in the central part of Taimyr, and the coast of the peninsula was occupied by Arctic tundras. Such a violation of geographical zonation corresponded to the transition time from the Sargan to the Holocene period. Restoration of "normal" zoning occurred approx. 8.5-7.5 thousand years AGO, when ice sheets began to form on Severnaya Zemlya. Such a "paleogeographic paradox" cannot yet be explained, but it allows us to speak about the metachronism of climate events in the Arctic, and over the entire circumpolar space (Bolyniyanov, 2000).

In geological sections of pre-Boreal deposits, dated to 14 C. 9.2-9.3 Ka BP, along the Bol rivers. Romanikha, Bol. Balakhnya, Bol. Microfossils were obtained in the Rasocha region (71-74° N, southern tundra, typical tundra, and northern tundra), indicating vegetation close to modern vegetation (Nikolskaya et al., 1980). Progress to the north of Bol is noted. Balakhni of shrubby birch (Nikolskaya and Cherkasova, 1982).

In the Boreal period, the first significant warming of the climate occurred. This is recorded by paleobotanical materials obtained from sediments dated to 14 from 8.8-8.2 thousand years ago. January temperatures increased by 1 - 2 °C, July temperatures-by

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1-6 °C (maximum in the Arctic tundra), and precipitation increased by 50-100 mm (Nikolskaya, Borisova, Kaplyanskaya et al., 1989). At this stage (9 - 8 thousand years AGO) there was intensive peat accumulation and waterlogging of the territory, the southern tundra subzone with larch and alder was spread almost to the latitude of Lake Baikal. Taimyr (74° N) [Belorusova, Lovelius, Ukraintseva, 1987]. According to the results of the peat bog analysis on the Mal River. Kheta in the forest-tundra zone (date to 14 S: 8,500 ± 200 l.n. (GIN-26)), this time corresponds to the forest phase and distribution of spruce. According to N. A. Khotinsky (1977), the Holocene Boreal period in Siberia was the warmest and wettest. The conclusion about warming in the Atlantic period of the Holocene (8-4.5 thousand years AGO) does not raise doubts among researchers, but its scale is estimated in different ways. For the first half of this period, data are scarce; the Holocene climate optimum (5-6 thousand years ago) is well studied. It is believed that it was at this time that Taimyr developed the most favorable conditions for the growth of woody vegetation (Antropogen Taimyr, 1982). In the fossil spore-pollen spectra in the southern tundra subzone, pollen from "thermophilic" tree species such as birch, pine, spruce, larch, and even fir is recorded. Among the shrubs, the more "thermophilic" alder often dominates. In the south of the North Siberian Lowland, rare coniferous northern taiga forests and copses appeared, accompanied by an admixture of birch and spruce. It is believed that in the Holocene optimum, the northern boundary of the larch woodlands (forest tundra) was approximately at the latitude of Lake Baikal. Taimyr or slightly further south (72° 50 ' N)The border of woody vegetation ran along the coast (Belorusova, Lovelius, and Ukraintseva, 1987). 80-100 km south of m. Stumps and trunks of larches were found in Chelyuskin (Miroshnikov, 1958), but their absolute age is not determined. These considerations contradict the opinion of botanists that during the formation of tundra landscapes, woody vegetation did not penetrate into the Arctic tundra. A natural monument of interest is the fossil larch forest in the lower reaches of the Novaya River, described as early as 1937 by L. N. Tyulina and attributed to the Holocene optimum (Belorusova, Lovelius, and Ukraintseva, 1987). The larches here were at that time more powerful than today, growing only in small curtins.

The following climatic parameters were reconstructed for the southern tundra in the range of 6-6.5 thousand years AGO: July temperature was 2-4 °C higher than today, January temperature was 1-2 °C, and precipitation was 50-150 mm higher than now [Nikolskaya, Borisova, Kaplyanskaya et al., 1989]. It was also suggested that abnormally high summer temperatures. Judging by the wood and palynological data, the deviation of the July isotherms from the current values was not less than 8-10 °C. Thus, the average monthly temperature of July at the Chelyuskin m. should have been 10-12 °C (modern 2 °C). The maximum anomalies occurred in the east of Taimyr (Belorusova, Lovelius, and Ukraintseva, 1987).

The results of paleoreconstructions revealed a similarity between the Holocene optimum and the Boreal period (Nikolskaya et al., 1980) and a very large similarity between the Holocene optimum and the Karginsky time optimum (42-43 Ka BP) (Nikolskaya, Klimanov, Borisova et al., 1989). In the Subboreal period (4.5 - 2.5 Ka BP), the climate situation significantly worsened (Nikolskaya et al., 1980), although no significant changes seem to have occurred in the forest-tundra zone. Obviously, forest degradation has begun. In the area of the Ary-Mas tract, instead of spruce-larch woodlands, yernik, and alder forests, only larch woodlands with loss of spruce and tree-like birch began to dominate (Mironenko and Savina, 1975). Peat formation in the region has decreased. Alases were formed on the site of drained thermokarst lakes. Glaciers have been activated in the east of the Byrranga Mountains.

Discussion of the results

According to the most widespread point of view, the most favorable conditions for mammoths were the extreme continental, arid, cryoarid, cold climate with little snow in winter, and solid, solid soils. Under this climate, in the late Pleistocene of northern Eurasia, there probably existed peculiar hyperzonal periglacial landscapes of the cryoxerotic tundra stage type, which were formed during the complete restructuring of the zonal landscape structure during the cold snap epochs (Sher, 1971, 1997; Velichko, 1973; Tomirdiaro, 1980; Shilo, 2001). It is believed that tundra steppes, by analogy with modern grassy steppes, were highly productive ("fodder", according to N. K. Vereshchagin) and had the necessary phytomass for large herbivores. Some scientists even suggest the dominance of the richest high-grass grasslands and other meadows (Tomirdiaro, 1980). In general, it is assumed that the mammoth was adapted to a very narrow range of landscape and climatic changes (Velichko and Zelikson, 2001).

Warming at the Pleistocene-Holocene boundary caused an increase in the water availability of the Subarctic and Arctic regions, the breakdown of hyperzonal vegetation and restoration of the forest zone, the degradation of permafrost, heavy snowfall and increased snow cover, the widespread development of thermokarst, etc.

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swampiness, zaozerennost. Tundras have become "low-fat". All this led to the extinction of individual representatives of the mammoth fauna [Vereshchagin, 1979]. According to the degree of natural changes, the transition to the Holocene is assessed as a "landscape catastrophe". But the nature of North Asia in the late Pleistocene experienced several epochs of warming, which in general did not significantly affect the viability of the mammoth fauna. Therefore, experts suggest that Holocene warming led to the emergence of fundamentally different plant associations that are not similar to those that existed earlier and are not able to feed large herbivores [Sher, 1997]. All this caused the extinction of the latter.

Let us estimate the evolution of the natural environment of Taimyr and mammoth fauna in the late Pleistocene-Holocene, based on the above paleogeographic materials and taking into account the current ecological and geographical conditions of the peninsula.

Glaciations, transgressions, and regressions of the Polar basin. In recent years, new data have been obtained in the course of research, which allows us to speak about low activity, small scales of Sartan glaciation in the Arctic and Subarctic. Such a point of view was justified for Central Siberia as early as in the 1940s by V. N. Sachs (1953, Cheha, 2000). During the maximum Sartan cooling period (20-18 thousand years AGO), the Severnaya Zemlya archipelago joined Taimyr during sea regression. As before, during the relatively warm Late Karginian period, and during the obviously severe climatic period under consideration, mammoths penetrated the Northern Land along the permafrost-bound drained shelf (24-19 Ka BP). Sea level rise from the minimum levels began around 16 KA BP (Bolshiyanov and Makeev, 1995). The separation of the Arctic islands from Taimyr finally occurred, apparently, about 10 thousand years AGO as a result of Holocene transgression. The increase in humidity should have contributed to the growth of glaciers on Severnaya Zemlya, but the scale of glaciation was insignificant and the glaciers were inferior to modern ones. Date to 14 S for a mammoth tusk from October Revolution Island (11500 ± 60 l).N. (LU-610)) [Makeev, Arslanov, and Garutt, 1979] suggests that either mammoths re-penetrated the Northern Land after the ice sheets were reduced, or they survived the Sartan stage of glaciation here. In any case, we can talk about long-distance, multi-hundred-kilometer migrations of these animals and their ecological plasticity, adaptability to sharply different natural conditions on the continent and Arctic islands.

It is noteworthy that there was a significant warming on the Arctic islands (Severnaya Zemlya Archipelago, Sverdrup Island) about 12-10 thousand years AGO, when shrubby and typical tundra appeared there. Additional data on significant warming in the high Arctic latitudes earlier than the Holocene (14-10 Ka BP) and in the Pre-Boreal (10-9 Ka BP) are given above. In modern conditions, mammoths on Severnaya Zemlya would not have been able to survive because of the scanty vegetation [Ibid.]. A special feature of the high-latitude Arctic is the absence of traces of Late Triassic cooling and warming during the Atlantic Holocene period. Obviously, in Taimyr, even in the west of the peninsula, there were no conditions for significant Sartan glaciation either. Normal lacustrine and lacustrine-alluvial accumulation of precipitation occurred.

Permafrost. The presence of permafrost soils was the most important factor determining many paleolandscape characteristics of the region. According to the results of facies studies of frozen strata on the coast of the Laptev Sea, conducted in the 40s-50s of the XX century, it is not necessary to speak about any significant climate fluctuations in the Quaternary (obviously, in the late Quaternary) time, since, firstly, underground ice has a vein origin, and secondly, in the late Quaternary period, it is not possible to there are no traces of epigenetic freezing or signs of freezing after deep thawing (for example, at the end of interglacial periods). A special feature of the history of Siberian glaciation outside the mountains is the extremely weak difference between glacial and interglacial epochs [Vtyurin et al., 1957]. Subsequent studies confirmed that there was no degradation of the permafrost layer north of the Arctic Circle during the Kazantsev, Karginsky, and Holocene periods. During climate warming, the negative average annual temperature of the strata remained within the framework of constant negative average annual air temperatures (Fotiev, Danilova, and Sheveleva, 1974). In fact, the same thing is recorded in the study of vein ice on the Sablera Peninsula, near Lake Baikal. Labaz (Eastern Taimyr).

In general, despite the extremely complex relationships between degradation and agradation processes in permafrost strata, the influence of short-period temperature fluctuations on ice strata is very weak in the northern regions far from the southern permafrost boundary (such as Taimyr). The inertia of permafrost strata is also quite large for large areas of stable cooling under historical climate fluctuations (Popov and Tushinsky, 1973). Apparently, in the past, only within the boundaries of marine transgressions and thick ice sheets could more or less drastic changes occur in the distribution and characteristics of permafrost.

Taking into account the above and taking into account the current temperature parameters of the permafrost zone in Taimyr, we

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We believe that deep regional thawing of frozen strata did not occur either in the Holocene or at earlier stages of the late Pleistocene (possibly excluding the Kazantsevskaya and Karginskaya transgressions). The wide development of thermokarst, lake cover, and mossiness are permanent properties of the tundra and forest-tundra zones of Taimyr with continuous very thick permafrost and icy soils. This can be seen from the wide development of peat, lake and swamp formations in sections of Quaternary Arctic and Subarctic sediments. The scale of these phenomena and the intensity of peat formation, lake and swamp sedimentation processes in the late Pleistocene, of course, changed. But, in our opinion, not so much as to consider it a consequence of a "landscape catastrophe" at the Pleistocene - Holocene boundary, as proponents of the tundra concept believe.

Natural and climatic changes. During the late Pleistocene, Taimyr alternated periods of warming and cooling, which is a reflection of the general planetary evolution of nature. However, the assessment of the depth of natural changes and the correlation of stages with different heat and water availability in this region, as in other regions of Northern Eurasia, is debatable. In general, the different geochronological scales of the late Pleistocene and Holocene are relatively close in terms of the total number of warm and cold epochs. As a result of climate changes, significant shifts in natural zones occurred, reaching 100-200 km. During the warming epochs, the forest-tundra zone expanded to the north, and the typical tundra was replaced by the southern tundra within the zone. Based on the current climatic indicators of these natural zones and subzones, the researchers compiled temperature and humidity characteristics of individual epochs of the late Pleistocene and Holocene. It is obvious that with such high gradients of summer temperatures in Taimyr, which were crucial for vegetation and vegetation growth, the shift of natural zones and subzones should have occurred just as sharply.

Critically assessing the paleogeographic scenarios of the past, we can cite interesting data from geobotanists on the current thermal requirements of trees that make up the northern border of forests in the Subarctic, as well as shrubs and shrubs. If the average daily air temperatures are 7-8 °C, and daytime temperatures exceed 11 °C for three to four hours and such temperatures last for five to six weeks, then Siberian spruce, Siberian larch, dwarf birch and various shrubs (blueberries, lingonberries, etc.) have a normal increase, they bloom and they bear fruit. Soil temperature is of great importance. If at a depth of 15-20 cm (where the main mass of roots is located in the Subarctic) it reaches 5 °C, and in deeper horizons 2-3 °C, then such soils are quite favorable for the growth of the above trees. In summer on Taimyr, in a typical tundra, the soil temperature at the specified depth does not fall below 5-6 °C. Thus, at present, the actual forest boundary does not correspond to the thermal characteristics. In Taimyr, this lag band is 250-350 km (Kryuchkov, 1967). According to the results of paleogeographic reconstructions, the shift of woody vegetation during the warming epochs of the late Pleistocene did not exceed 150-200 km. The conclusion from the above is obvious: the thermal limits for the movement of vegetation zones to the north were not used by vegetation even during periods warmer than today. The reasons for this vary. It can be assumed that the climatic changes in the late Pleistocene were less significant than expected. Even small warming events led to very significant changes in the vegetation cover. Examples of distant penetration of woody and shrubby vegetation to the north are still available today (Pospelov and Pospelova, 2000). As Zh. M. Belorusova, N. V. Lovelius, and V. V. Ukraintseva quite reasonably point out [1987], having once penetrated far to the north, woody vegetation never completely disappeared there, even in the coldest epochs, but quickly recovered during warming periods.

Vegetation, which played an important indicator role of natural changes, was a food base for large mammals, i.e. one of the main environmental factors of the latter's existence. With this in mind, experts have developed a hypothesis about highly productive tundras of Beringia, which have no modern analogues. In recent years, the tundra concept has been severely criticized by geo - and paleobotanists in two ways: 1) the tundra steppe was not a zonal vegetation type; 2) hypothetical tundra forests could not serve as a food base for mammoths in terms of productivity (Kozhevnikov, 1999; Kozhevnikov and Ukraintseva, 1997; Verkhovskaya, 1988).

It should be noted that no researcher mentions the presence of tundra steppe as a zonal landscape type on the peninsula in the past. However, it is obvious that the tundra type of vegetation cover is currently exceptionally widespread here, and the latter is characterized by a pronounced mosaic, complexity, and diversity of plant life forms (psychrophytes, cryophytes, mesophytes, and hygrophytes). Within the tundra and now there is a place for steppe (stepoid, according to Yu. V. Kozhevnikov) vegetation. According to V. G. Mordkovich (1994), parts of the steppe landscape in Central Siberia can be considered as follows:

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from the coast of the Arctic seas to 45° N, "Patches of steppes with a diameter of tens or hundreds of meters are completely isolated from each other and completely surrounded by tundra or taiga landscapes", [Mordkovich, 1994, p. 388]. There are especially many such sites on the slopes of the southern exposure. It is also necessary to take into account the pronounced phenomena of extra - and intrazonality in the tundra, which make it difficult to distinguish zonal vegetation elements.

The tundra's biotopic diversity was also evident earlier: it was more pronounced during periods of warming, and less pronounced during periods of cooling. At the same time, the tundra productivity increased during the warming epochs. Obviously, mineral-trophic meadows and swamps with a wide spatial distribution of shrubby vegetation were the most productive (Kozhevnikov, 1999). In general, phytomass reserves in tundras are always minimal at the tops of interfluvial areas. Even at low altitudes, habitats are being cooled, the wind is increasing, and poor tundras are forming (Bazilevich, Grebenshchikov, and Tishkov, 1986).

These materials do not give grounds to speak about significant differences in the vegetation cover of interglacial epochs and the Holocene. According to the results of paleoecological analysis of carpoids from Upper Quaternary deposits (Anabar estuary, Olenekskaya Channel, Faleevsky Peninsula in the New Siberian Islands archipelago, lower Kolyma River), the natural conditions of the late Pleistocene did not differ in any way from modern ones, i.e. they were not cryoxerotic, as suggested by the proponents of the tundra concept (Solov'ev and Stanishcheva, 1983).

Mammoth fauna, questions of evolution and paleoecology. Mammoth remains in the frozen strata of the Subarctic and Arctic regions, as determined by taphonomic features, are localized in places of natural permanent and spontaneous death of individuals, as well as death in various natural traps - water, swamp, ice, etc. [Vereshchagin and Tomirdiaro, 1995]. It is also obvious that the areas of the richest such localities in the past were also areas of wide distribution of mammoths. It is assumed that these animals have lived in Taimyr since the end of the Kazantsevskaya transgression (65 KA BP). The histogram of radiocarbon dates for the remains of Taimyr mammoths (Sulerzhitsky and Romanenko, 1997) is relatively evenly filled with definitions in the range of 50 - 10 KA BP. The period of 40-38 thousand years AGO (Malokheti optimum) is most saturated with definitions; a few dates correspond to the period of 21-15 thousand years ago (Gydan cold snap). Thus, repeated warming and cooling events during the Karga-Sartan period did not change the overall appearance of the mammoth fauna; along with the remains of the mammoth, horse remains are constantly found, less often bison and musk ox. Individual anomalies may indicate the redistribution of animals in time and space, or a quantitative change in the structure of populations. The map of locations of mammoth bone remains dated to 14 S (Fig. 3) allows us to present the spatial distribution and population density of the Taimyr mammoth at different stages of the late Pleistocene. The most favorable natural conditions for these large animals were found in the central and eastern parts of the North Siberian Lowland (now the typical subzone, southern tundra, and partly forest tundra). Both now and in the past, it was an area with a high continental climate, minimal wind activity, relatively warm summers and harsh winters. This area was characterized by minimal development of marine transgressions at different stages of the late Pleistocene. The western part of Taimyr was characterized by more heavy snowfall, frequent snowstorms, and generally increased cyclonic activity.

The above data, as well as the scheme of mammoth dispersal in Northern Asia (Orlova et al., 2000), as suggested by A. A. Velichko [1973], A. A. Velichko, and E. M. Zelikson [2001], do not allow us to speak about the adaptation of these animals to a narrow range of landscape and climatic conditions. Obviously, mammoths were characterized by a rather high ecological plasticity, which determined different tactics of using the territory-from semi-sedentary existence to long-distance movement during seasonal migrations (Sulerzhitsky and Romanenko, 1997). Eurybionty and polyzonality of proboscis and large ungulates of the Quaternary period were noted (Puchkov, 2001). The polyzonality of mammoths also suggests a certain intraspecific morphophysiological variability of these large mammals with the formation of ecotypes and subspecies, which is rarely observed in the fossil material.

The issue of mammoth food supply is often discussed in the literature. The area of the maximum concentration of the Taimyr mammoth population can be estimated at 10-13 million hectares (central and eastern parts of the North Siberian Lowland). Even if we take as a basis the current minimum annual productivity of the Taimyr tundra (approx. 1 t / ha) and the daily diet of one mammoth (approx. 2 kg of feed), it is not difficult to calculate that this area could feed more than one thousand heads of these large animals during the year. It should also be noted that the Subarctic region is characterized by an increased food value of plants (a lot of protein and lipids), compared to the Arctic.

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3. Map of locations of mammoth fossils in Taimyr, dated by radiocarbon dating. Landscape borders: a - regions; b-districts (po: [Taimyr-Severo-Zemel region. .., 1970]); c-locations of mammoth remains. I - region of the North Taimyr uplands and mountains; II-region of the South Taimyr lowlands (North Siberian Lowland); III-West Siberian Plain; IV-Central Siberian plateau. 1 - Chelyuskin Peninsula area; 2-middle-altitude part of the Byrranga Mountains; 3-hilly plains area; 4 - Byrranga lowlands; 5-hilly hills area; 6 - northern region; 7 - southern region; 8 - Putorana Plateau; 9 - Anabar massif; 10 - Kotui plateau. Radiocarbon dating results (thousand years ago) for individual localities: 1 - 26,7; 2 - 37,0; 3 - 35,8; 4 - 25,1; 5 - 10,7; 6 - 49,7; 7 - 20,4; 8 - 45,0; 9 - 27,5; 10 - 27,3 35,0 36,2 38,3 38,9 40,2; 11 - 41,4; 12 - 38,8; 13 - 23,5 38,5 38,5 39,8 41,2 >52,7; 14 - 14,8; 15 - 42,8 > 53,2; 16 - 22,0 28,9 32,3 36,6 40,3 41,9 >50,0; 17 - 13,3 16,3 32,0 36,8 38,4 38,5 39,1 39,2 47,9 >49,5; 18 - 31,9; 19 - 12,1 22,8 24,9 39,3; 20 - 11,1 43,5 46,1 >49,5; 21 - 40,8; 22 - 23,8; 23 - 29,5 32,0; 24 - 38,8; 25 - 9,7 9,9 10,3; 26 - 12,8; 27 - 10,1 40,5; 28 - 31,8; 29 - 12,3 12,4; 30 - 28,8; 31 - 11,4.

plants of temperate climates (Miroslavov, Voznesenskaya, Bubolo, 1999), and a wide range of mammoth food consumed - grasses, sedges, shrubs, and twig food. It should also be noted that hygro-and mesophilic meadow and swamp vegetation, rather than dry-loving vegetation, as previously assumed, was the main food component of the mammoth and its companions (Verkhovskaya, 1988; Puchkov, 2001). It is unlikely that in the conditions of continuous permafrost with a 10-centimeter seasonally frozen layer, the most forage-rich areas of meadows and swamps could serve as an obstacle to the movement of even such heavy animals.

According to the above radiocarbon dates of mammoth bone remains (see Fig. 3), at certain stages of the Karginsky time (Malokheti, Lipovo-Novoselovo warming), and in the second half of the Sartan time, the Taimyr mammoth moved north to the subzone of the Arctic tundra. Thus, we can talk about the migration of mammoths during periods of warming to the north of the peninsula.

The considered materials describing the Taimyr Peninsula do not indicate a natural catastrophe at the turn of the late Pleistocene - Holocene, but rather the evolutionary development of natural events in the interval 15-9 thousand years ago. Thus, the course of development of the Taimyr nature in the late Pleistocene-Holocene does not correspond to the above hypothesis about the natural cause of the extinction of mammoths at this turn. It seems to us that this problem should also be considered in bioevolutionary terms on the scale of the entire North Asia.

The works of S. S. Shvartz [1961] established that typical Arctic species of mammals and birds are characterized by low metabolism, low fecundity, and slow growth and development. Most of the animals thriving in the tundra zone are at a very low phylogenetic level, characterized by low organization, and a certain primitiveness. The path of passive adaptation (passive ecological and physiological adaptations to the natural environment) in pessimistic conditions for animals is the most profitable. Since the existing gradients of climatic indicators in Taimyr are much more pronounced than in southern latitudes, the composition of fauna and flora, the appearance of communities throughout the tundra zone from its northern to southern borders change extremely sharply. Adaptation to zonal conditions manifests itself in a certain specialization that limits the ability of animals to live in other zones. The harsher the climate, the deeper it is.

The path of active adaptation is fixed by the intensification of growth, the speed of development and maturation, as well as the strengthening of general vital activity and regulatory processes (Chernov, 1980). This pathway is typical for the more southern polyzonal species that moved into the Subarctic and Arctic, living in many less specialized zones. With

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such species inhabit the southern parts of the tundra zone more actively than typical inhabitants of the Arctic.

Judging by the temporal and spatial development frames in Northern Asia, the late - type mammoth Mammuthus primigenius Blum, which migrated to the Subarctic, was also polyzonal and eurybiont. It showed signs of active adaptation to Subarctic conditions and was probably preadapted in the conditions of more southern latitudes of Central Siberia. Here, in some periods of the Pleistocene, forest-steppe and steppe landscapes with tundra-like features were developed (Cheha, 1996). Adaptation to the natural environment was expressed in physiological adaptation processes, as well as in the selection of suitable biotopes, often intrazonal, avoiding adverse natural influences, migrations, etc.

One of the possible scenarios for the collapse of mammoth fauna in the Subarctic and Arctic can be presented in the following form. Over the millennia, it is obvious that the "Arctic typicity" of mammoths has increased due to the change of rather broad natural indicators to tundra ones. Such readaptation processes (reverse development) are usually disastrous for organisms. Readaptation with the intensification of very negative biological processes affected the gradual disintegration of the mammoth fauna. It was not the warming of the climate, but the "pressure" of the Arctic with its specific, highly specialized adaptation requirements that ultimately caused the collapse and extinction of such large animals as mammoths. Under the conditions of readaptation, migration to the south became impossible for them. In search of new favorable habitats, the mammoth population began to shift northward in the late Pleistocene and Holocene. On Wrangel Island, mammoths that had degenerated into a dwarf form survived the Atlantic Holocene period (Vartanyan et al., 1995), but were doomed to extinction.

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The article was submitted to the Editorial Board on 27.04.05.

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