Libmonster ID: JP-1493

Author(s) of the publication: S. K. Vasiliev →

Source: Archeology, Ethnography and Anthropology of Eurasia, No. 2.30 June 2008 Pages 34-56 →

UDC 569

Institute of Archeology and Ethnography SB RAS

17 Akademika Lavrentieva Ave., Novosibirsk, 630090, Russia

E-mail: svasiliev@archaeology.nsc.ru

In the late Pleistocene faunal assemblages in the south-east of Western Siberia, the primitive bison (Bison p. priscus) was one of the most widely represented megafauna species. Since the end of the Kazantsev time (about 100-90 thousand years ago), a directed decrease in the size of its body was noted, which reached its minimum at the final stage of the Karginsky interstadial (about 30 thousand years ago). The Sartan cooling again gave a sharp impetus to an increase in the size of Bison p. priscus. In contrast to Europe and Eastern Siberia, in the south of Western Siberia, the size of the body and skull of bison remained virtually unchanged during the late Pleistocene. This conservativeness is obviously explained by the relative stability of the natural and climatic situation, the presence of steppe and forest-steppe spaces favorable for the existence of the species with rich food resources and a small depth of snow cover.

Introduction

In the faunal complexes that successively replaced each other in the south of Western Siberia during the Pleistocene, the primitive bison, along with the horse, was the most massively represented background species. At some stages of the Middle and late Pleistocene, the proportion of bison bones in alluvial taphocenoses could account for up to half of all remains of large mammals. Despite this, this species has remained relatively poorly studied morphometrically and evolutionarily on the territory of Western Siberia. Bison skulls stored in various museums in Western Siberian cities (usually without reliable geological reference) are described and mentioned in a number of works (Gromova, 1935; Flerov, 1977, 1979;

Alekseeva E. V., 1980]. In addition, E. V. Alekseeva's monograph [1980] used measurements of several hundred bones of the postcranial bison skeleton, which she collected mainly on a sand and pebble spit near Krasny Yar in the Tomsk region. The age of this uplift material was presumably estimated by the author within the Middle - Late Pleistocene [Ibid.]. I. V. Foronova (2001), on the contrary, dealt mainly with stratified finds from Pleistocene deposits of the Kuznetsk Basin. However, the total volume of her collection of Bison priscus remains is relatively small, which is why, apparently, it was described very concisely, without specifying in the measurement tables the average values (as in E. V. Alekseeva) and the number of bone specimens. In recent decades, the study of Paleolithic, primarily cave, localities in southern Siberia has accumulated quite significant material on primitive bison. Due to the specific nature of the formation of cave taphocenoses, it is mainly represented by ele-

The study was carried out with the financial support of the Russian Foundation for Basic Research (project N 06 - 04 - 49232) and a grant from the President of the Russian Federation to support leading scientific schools (NSH-6568.2006.6).

page 34
distal limb segments: phalanges, carpal and tarsal bones, and more rarely entire metapodia. Meanwhile, it is precisely for these bison remains (which paleozoologists primarily deal with) that published comparative data are almost completely absent.

The main obstacle that arises when studying the remains of primitive bison is the huge range of individual-sexual variability characteristic of representatives of the genus Bison. Until now, it has remained poorly studied, especially in the case of fossil forms. Often, even within a small sample of Bison priscus skeletal elements of the same name, such extreme variations of variability are observed that make paleoteriologists think about assigning them to different taxonomic or chronological forms [Alekseeva E. V., 1980; Foronova, 2001].

Based on a large amount of stratified material, this article examines the variability and evolutionary development of Bison p. priscus in the south-east of Western Siberia during the late (partly Middle) Pleistocene.

Location Overview

Materials for the study were collected by the author on three large localities of Late Pleistocene fauna in the Upper Ob region. These localities complement each other quite well, chronologically covering all the main stages of the late Pleistocene, which makes it possible to trace the main trends in the evolutionary development of Bison p. priscus at this time in the south-east of Western Siberia.

Krasny Yar is located on the right bank of the Ob River, 17 km below Novosibirsk. In the section of the third erosion-accumulative terrace, two main bone-bearing horizons are revealed here: bluish-gray floodplain loams of layer 4 in the middle part of the outcrop, which belong to the final segment of the Karga interstadial, and diagonally layered sands of layer 6 at the base of the section, which go under the water's edge, dating from the final Kazantsev Interglacial (Volkov and Arkhipov, 1978; Panychev, 1979; Vasiliev, 2002, 2005]. Since 1978, the author has collected 2,703 bone remains dating back to the Kazantsev period and 719 to the Karginsky period (Table 1). All skeletal elements are found in the channel diagonal-layered sands of Layer 6 - from small bones of the wrist and tarsus to entire axial skulls of bison, woolly rhinoceroses and mammoths. The taphocenosis of this layer is dominated by the remains of Bison p. priscus (48.1 %), Equus. ex. gr. gallicus (19.7 %), and Coelodonta antiquitatis (7.3 %).

The bones of deer - giant, noble and elk - make up a total of 17 %. At the end of the Kazantsev Interglacial period, about 100-90 thousand years AGO, forest - steppe landscapes developed on the territory of the Novosibirsk Ob region. In the wide river valleys grew spruce forests with an admixture of larch, cedar and high-stemmed birch. The climate did not differ significantly from the present one and was only slightly drier and cooler (Volkov and Arkhipov, 1978; Panychev, 1979). The predominance of bison and horse bones in the 6th layer, as well as noticeably shortened wolf and elk metapods, compared to modern ones, indicate the relative lack of snow in the Riess-Wurmian winters.

The second half of the Yermakovsky or first half of the Karginsky time is covered by the materials of the grandiose Taradanovsky locality, located on the Ob River below the village of Taradanov (Suzunsky district of the Novosibirsk region). The main bone-bearing horizon of Taradanovsky Yar, from which 99% of all bone remains originate, is located several meters below the low-water edge and is directly eroded by the river. In spring and summer floods, the washed-up material is carried out on a vast sand and pebble bone-bearing shoal located in the lower part of the Yar river. Hydrodynamic conditions in this section of the river are such that mainly small and medium - sized bone remains are heard and deposited on the beach-numerous phalanges, metapods, astragalus, carpal bones and tarsals of large ungulates. Among the bison bones, horn rods, lower jaws and isolated teeth are completely absent, but there are whole shoulder blades that were taken out on the beach due to their"sailability". Of the large limb bones, 11 whole radial and tibial bones (mostly bison females), one whole femur and a single distal half of the humerus were found. During the 2003-2005 seasons, 3,085 bone remains of 20 species of large mammals were collected here (Table 1). 18 radiocarbon dates were obtained from the beach: 13 - in the range of > 45-40 thousand BP, 5-in the range of 35-26 thousand BP [Vasiliev and Orlova, 2006]. Graphical constructions based on the average proportions of more than 180 metacarpal and metatarsal bones of a horse from Taradanov showed its complete identity with Equus ex. gr. gallicus from the Karginsky (ca. 30 Ka BP by 14 S) layer 4 of the Krasny Yar (Vasiliev, 2004, 2005). Thus, the upper age limit of the main bone - bearing horizon of the Taradanovsky Yar most likely exceeds 40-45 thousand years, and the lower age limit does not fall below 90-100 thousand years. Spore-pollen spectra were obtained from the sands lying several meters above the bone layer, at the water's edge, indicating the existence of a forest cover.-

page 35
Table 1. Species composition and number of mammalian bone remains at late Pleistocene localities in the Upper Ob region

Taxon

Krasny Yar, layer 6 (R-W)

Taradanovo (W-1-2)

Krasny Yar, layer 4 (W-2)

R. Orda (W-3)

Number of bones

Lepus sp.

0,07

0,14

Marmota sp.

0,04

Castor fiber L.

0,44

0,10

0,40

Canis lupus L.

0,33

0,13

0,14

Canis sp.

0,19

Ursus arctos L.

0,37

0,20

0,28

0,40

U. (Spelaearctos) rossicus Borissiak

0,26

0,39

Gulo gulo L.

0,04

Meles meles L.

0,04

Crocuta spelaea Goldfuss.

0,04

0,06

Panthera spelaea Goldfuss.

0,41

0,75

0,83

Mammuthus primigenius Blum.

131

4,85

0,71

1,25

12,9

Equus ex. gr. gallicus Prat.

531

19,7

1 210

39,3

446

62,0

105

42,2

Equus (Equus) sp.

0,03

0,14

Equus ex. gr. hydruntinus Regalia

0,44

1,49

Equus cf. hemionus Pallas.

0,14

Coelodonta antiquitatis Blum.

197

7,29

231

7,50

10,7

12,1

Megaloceros giganteus Blum.

187

6,92

2,29

Cervus elaphus L.

128

4,74

138

4,48

3,34

0,80

Alces cf. alces L.

144

5,33

1,85

2,92

0,80

Rangifer tarandus L.

0,03

1,67

0,80

B/son priscus Bojanus

1 300

48,1

1 201

39,0

115

16,0

28,9

Sa/ga borealis Tscherskii

0,11

0,36

0,42

0,80

Ovis ammon L.

0,06

Bovini gen. indet. (large form)

0,04

0,03

Bovini gen. indet. (Soergelia sp.?)

0,65

Total bone remains

2 703

100

3 083

100

719

100

249

100

steppe landscapes with pine-birch and spruce forests (Panychev, 1979). Faunal materials are dominated by the bones of horse (39.3 %), bison (39 %), woolly rhinoceros (7.5 %); there are remains of Pleistocene donkey, saiga antelope, argali, and deer (9.6 %). The composition of the fauna allows us to reconstruct the landscapes of mesophytic steppes or sparse forest-steppe for this time.

Materials from the 4th layer of Krasny Yar indicate that at the end of the Karginsky interstadial (about 30 thousand years ago), the ratio of bison to horse changed in reverse: 62% of the remains here belong to the horse, and bison is relegated to the background (16 %). The number of deer bones is reduced to 7.9 %. The single bone remains of saiga and kulan, with the absolute predominance of bones of the relatively broad-toed Equus ex. gr. gallicus, probably indicate relatively soft, moist soils of the mesophytic steppe. The climate was close to the modern one, but the snow cover was much less thick.

The location on the Orda River, which dates back to the end of the Sartan period (Vasiliev, Nikolaev, and Petrin, 1995), is located between the villages of Rogalevo and Ust-Lukovka in the Ordynsky district of the Novosibirsk Region. From the bones collected in the riverbed (in 1994 and 2006), a series of radiocarbon dates were obtained-approximately from 14-13 to 21-19 thousand years ago.

page 36
There are also more horse remains (42.2 %) than bison (28.9 %); the bones of woolly rhinoceros (12.1%) and mammoth (12.9%) are quite numerous. Red, northern, and elk deer account for a total of 2.4 % (Table 1). The extraordinary shortening and massiveness of the bison and horse metapods indicate a significant decrease in snow cover thickness at this time, when landscapes of cold and low-snow pereglacial steppes developed in the south of Western Siberia (Arkhipov and Volkova, 1994)..

Material and methodology

In total, almost 2.7 thousand Bison p. priscus bone remains were examined, including 1,300 from the Kazantsev layer 6 of Krasny Yar, 115 from the Karginsky layer 4 of this locality, 1,200 from Taradanov (W - 1-2), and 72 from the Sartan deposits of the Orda River. All material, with the exception of specially specified cases, is stored in the IAET SB RAS. For comparison, we used collections of metacarpal bones of bison (132 specimens) from Kurtak (Krasnoyarsk reservoir) and from the Chumysh River in the Altai Territory.

When processing and analyzing the metacarpal bones of bison, V. Eisenmann (1979; Eisenmann and Beckouche, 1986) used a slightly modified technique, which was used by her in the study and description of equine metapodia. The basis of measurements of bison skulls was borrowed, with a number of abbreviations, from the works of V. I. Gromova [1935] and B. S. Rusanov [1975]. Measurements of postcranial skeletal elements were carried out mainly according to the method proposed by A. Driesch (1976). A number of additional measurements were made for the carpal and tarsal bones. The length of carpi radiale, C. intermediate, tarsale IV, width of C. ulnare, carpi 2+3, carpi 4+5 were measured in the projection on the sagittal line of the body. Measurements were made with a caliper with an accuracy of 0.1 mm. A number of difficult, "floating" measurements, such as the diameter of the upper end of the humerus or tibia, were carried out using a special measuring ruler. The distal dimensions of the 1st and 2nd phalanges and the carpi ulnare height were also measured with a three-point ruler.

The author adheres to the concept of K. K. Flerov [1977, 1979], who suggested that the Bison priscus longicornis identified by V. I. Gromova [1935] should be considered identical to the typical race B. p. priscus, which lived in Europe and Western Siberia, and B. p. crassicornis, which occupied the area east of the Yenisei. In the second half of the late Pleistocene, these temporary subspecies were replaced by B. p. mediator in Europe and Western Siberia, and B. p. occidentalis in Eastern Siberia (Flerov, 1977, 1979).

Description and comparison

V. I. Gromova (1935), who studied collections of Bison priscus skulls from various regions of the Soviet Union, noted that the final solution to the question of individual sexual variability of the Pleistocene bison should be expected from large, geologically homogeneous series of skulls from one location. From this point of view, the collection of cranial bison remains from Krasny Yar is an ideal object for research. In the deposits of the Kazantsev layer, 19 frontal-occipital parts of the skull with rods and 19 horn rods with the adjacent part of the skull were collected here. A single axial skull of the male is completely preserved (it is in a private collection), and only intermaxillaria are missing from one female skull. So far, only a single fragment of a female bison skull is known from the Karginsky layer of Krasny Yar. A whole axial skull of a male was pulled out from under the Taradanovsky yar by a net (it is in a private collection). Seven cranial remains of male bison originate from the Orda River (they are kept in the collection of the Horde Museum of Local Lore).

In the Ruslov alluvium of the 6th layer of the Krasny Yar, skulls belonging, no doubt, to large males were found, with the length of the horn rod along the large curvature up to 595 mm and the smallest width of the skull beyond the orbits up to 335 mm. Here, in situ, two skulls of old, judging by the wear of teeth, females were found (Figs. 1, 2) with the smallest forehead width of 242-249 mm and the length of the rod along the large curvature up to 293 mm (N 3188). The left horn shaft of the second skull (N 3263) has a length of 215 mm along the large curvature, and the right one is only 170 mm. The degree of preservation of all the listed material and the conditions of its occurrence (detection) leave no doubt about the synchronicity of the finds to layer 6.

Is such a large scale of sexual dimorphism possible in the size of the Bison priscus skull? Comparison of the range of individual-sexual variability in the skull size of the fossil bison and modern representatives of the genus Bison (Gromova, 1935; Burchak-Abramovich and Naniev, 1954; Nemtsev et al., 2003) showed that it is within the limits of that of the modern bison in most of its main features, i.e. it does not exceed the level of intrapopulation variability. More significant differences are observed in the size of the horn rods: in female Riess-Wurm bison compared to males-

page 37

1. Skull of a female (N 3263) Bison p. priscus Boj, a - top view, b - rear view.

2. Skull of female (N 3188) Bison p. priscus Boj, a -top view, b-rear view.

They are relatively smaller than in the modern bison population. Individual horn variability was also more pronounced in the fossil bison. Thus, according to the average values of the main measurements (Tables 2, N 5-8), the size of female Bison bonansus skulls (according to [Nemtsev et al., 2003]) is 82.8 % of the size of male skulls, whereas for B. p. priscus skulls from the 6th layer of Krasny Yar, this indicator is 80.1 2, N 11-16) - only 70 and 56.1%, respectively.

The analysis of works containing measurement data from the Bison priscus series of skulls (Gromova, 1935; Rusanov, 1975; Alekseeva E. V., 1980; Foronova, 2001; et al.) leads to the conclusion that the basis of the collections studied by the authors was almost exclusively male skulls. What causes this disparity? In the Ruslov alluvium (from which most of the bison skulls that make up museum collections originate), as a rule, the remains of male bison predominate. The example of the 6th layer of Krasny Yar showed that, judging by the metacarpal bones (where the division by sex can be carried out more or less confidently), the remains of female bison in the Ruslov alluvium are less than a third - 31.9 %*. This may be due to some ethological peculiarities of Bison priscus. Fragments of female bison skulls are even rarer in the Ruslov alluvium: 9 out of 38 (23.7 %). The most complete skull of an old female found in situ (N 3188) was buried in riverbed sediments before the connective tissue of the cranial sutures completely decomposed, but it broke apart as soon as it was released from the rock. The second skull of an old female (N 3263), apparently close to the previous one in terms of preservation, was unfortunately already washed out by water at the time of its discovery. From him, it was possible to find frontal bones with horn rods and the left upper jaw with the adjacent lower part of the eye orbit.

Female bison have much less massive and robust skulls than males, with looser and less compact horn rods (Gromova, 1935; Flerov, 1979). Fusion of the sutures of the brain skull in females occurs only after the end of

* It is noteworthy that in the floodplain deposits of layer 4, only four (43 %) male metacarps account for seven metacarpal bones of female bison. On the beach below Taradanovsky Yar, where the lighter bones of females were primarily removed due to hydrodynamic features, 29 metacarpal bones of females account for only 11 (37.9 %) metacarps of males.

page 38
Table 2. Dimensions of Bison priscus skulls

reproductive function [Flerov, 1979], i.e. usually by 13-15, at the latest by 20 years [Baskin, 1979; Sokolov and Tembotov, 1993]. Suture marks on the skull persist even in very old females, especially on the frontal surface. In males, the sutures disappear at 6-7 years of age, and the final fusion of the skull is completed by 8-9 years of age (Flerov, 1979). It is well known that in natural populations, even among such large animals as representatives of the genus Bison, mortality from various causes is very high and a relatively small percentage of individuals survive to old age. Taking into account all of the above, as well as the rigid hydrodynamic sorting of bone remains in the Ruslov alluvium, it becomes obvious that these deposits preserved mainly the frontal-occipital parts of the male skull, and almost all the female skulls disintegrated and collapsed. At best, from them

page 39
the sagittal half of the frontal bone with the horn shaft remained. Such remains, presumably, almost never attracted the attention of not only amateur local historians, but also Quaternary geologists, who took them, perhaps, for fragments of the skull of young individuals. Thus, the majority of museum collections studied by V. I. Gromova [1935], B. S. Rusanov [1975], and other researchers consisted of the skulls of male Bison priscus.

Individual variation in Bison priscus skulls is no less pronounced than sexual dimorphism. The skulls of female bison from Krasny Yar differ relatively little in absolute size (the smallest forehead width beyond the orbits is from 242 to 273 mm), apparently due to the small sample size (9 specimens). The sizes of horn rods vary more widely, with the length of the large curvature varying from 170 to 293 mm, and the girth of the base of the rod - from 180 to 246 mm. Judging from the skeleton part found in the Tomsk Ob region (Shpansky, 1997), the size of the horn rods of female bison could reach 350 mm in large curvature with a base circumference of 230 mm. Measurements of this skull (the largest and smallest width of the forehead, its width between the rods, and the span of the horn rods, respectively, 298, 243, 230, and 700 mm) are within the limits of variability of these characteristics in females of the Red Yar Kazantsevsky bison. The find dates back to the very end of the Pleistocene, possibly the beginning of the Holocene [Ibid.].

Variations in the size and structure of the horn rods of male bison also significantly exceed the variability of the skull itself. So, if the smallest width of the skull beyond the orbits varies from 286 to 335 mm, then the length of the horn rod along the large curvature is from 355 to 595 mm, while the circumference of its base is 278-395 mm. Skull No. 1781, which has the least short horn rods (355 mm), by all criteria, undoubtedly belongs to the old male, and its size in most respects is far from the minimum in the series. The length of the next least short horn rods is 420; 430; 445; 455 mm (2 copies), etc. The setting and shape of the horn rods are also no less variable. The bending index of the rod varies from 64.1 to 81.9, and the index of its massiveness-from 60.4 to 79.3. The collection includes horn rods that are massive from the base to the tip of the horn (with a girth in the middle of their length up to 274 mm) and massive only at the base, and then sharply thinning (girth in the middle of 224 mm) and therefore relatively thin. The position of the rods in relation to the sagittal line of the skull (from 57 to 82°) and the distance of their vertices to the frontal surface (from 50 to 230 mm) differ. Even more impressive is the difference in the span of the horn rods. The most prominent in this respect skull of the old male N 1323 has the longest (595 mm) and relatively thin horny rods, which are significantly straightened (bend index 79.3), spread out to the sides (the angle of divergence of the rods is 82°) and lowered down (the distance from the tops of the rods to the frontal surface is 50 mm), as a result, the distance between the tops of the rods reaches 135 cm, although the absolute size of the skull is not the largest in the series. The next specimen with the most widely spread horns has 112 cm between the ends of the horn rods, while the smallest one has only 80 cm, i.e. the maximum value exceeds the minimum by almost 1.7 times.

So far, only one fragment of a female bison skull (N 3974) has been found in the loam of the Karginsky layer of Krasny Yar. The smallest width of the forehead of this specimen is 252 mm, the largest width along the posterior edge of the eye sockets is 308, the width of the forehead between the rods is 240; the girth of the base of the rod is 204, the actual horn rod is 209, the horizontal and vertical diameters of its base are 65 and 67 mm, respectively; the straight length of the rod is ca 220, the length along the large curvature is ca 270, and the distance between the vertices is approximately 615 mm; the angle of divergence of the rods is 65°. These dimensions fit within the limits of variability of the skulls of female Riess-Wurm bison from Layer 6 (Table 2).

The only complete axial skull of a male from Taradanov (W-1 - 2) has rather large horn rods with relatively small dimensions of the skull itself (Table 2). Among the measurements not included in Table 2, it is necessary to indicate the condylo-basal, basic, total length, and alveolar length of the dentition: 596; 547; 633; 157 mm. The same parameters for the entire axial skull of a male from Krasny Yar are 615; 575; 627; 156 mm. The rest of its measurements (correspond to N 1-19 in Table. 2) following: 279mm; 233; 198; 199; 352; 301,5; 348; 209; 285; 166; 341; 366; 136,5; 108,5; 400; 543,5; 1 090; 130 mm; 76.5°.

As far as can be judged from the seven available cranial remains, the Sartan bison from the Orda River were noticeably larger in skull size, and especially in horn rods, than the Kazantsev Interglacial bison, while retaining all the structural features of the skull of their predecessor (Table 2). Another Sartan bison skull comes from the vicinity of the village of Kudelin Klyuch of the Toguchinsky district.In the Novosibirsk region in 1996, a complete skeleton of an old male was found here, the absolute age of which at 14 S was 23050 ± 255 years (SOAN-3635). This specimen has a span of horn rods of approx. 100 cm, the length of the rod along the base-

page 40
its curvature is 415 mm, its base circumference is 347 mm; the width of the forehead between the rods is 345 mm, and the smallest one in the postorbital constriction is 295 mm.

V. I. Gromova (1935) assumed that the main criterion for separating long - and short - horned (as was assumed in the Middle and Late Pleistocene) Bison forms was such values as the length of the horn rod along the large curvature and the girth of its base, exceeding 550 and 350 mm in B. p. longicornis, respectively.

In terms of skull size, the bison represented by materials from the 6th layer of the Krasny Yar are somewhat smaller than the bulk of the Middle Pleistocene longhorn bison studied by V. I. Gromova [Ibid.]. They are especially inferior to B. p. longicornis (B. p.priscus) in the size of horn rods. Along the length of the horn rod along the large curvature, only 7 specimens are found. out of 27 (25.9 %), and according to the circumference of the base of the horn rod, 17 out of 33 (51.5 %) formally fall into the group of long-horned bison. The horn rods of male bison from Krasny Yar, Taradanov, and the Orda River are curved in a hollow arc, with their ends pointing to the sides, up, and slightly backward, slightly protruding beyond the occiput line and being higher (by an average of 164-169 mm) than the level of the frontal surface. I. V. Foronova (2001) provides data on Middle Pleistocene bison Kuznetsk Basin: the width of the forehead between the rods is 320-410 mm, the length of the horn rod along the large curvature is 620-780 mm, and the girth at the base is 338 - 450 mm, which significantly exceeds the size indicators of the skulls of Late Pleistocene bison from our collection. For a series of B. p. crassicornis skulls from the Sandy Mountain locality on the p. Even shorter and more massive horn rods are characteristic of the Viliu river in Yakutia, dated to the Kazantsev period (Rusanov, 1975), compared to the simultaneous bison of Krasny Yar, with similar absolute skull sizes (Table 2). The same pattern - shortening of horn rods while maintaining the overall large skull size - is observed in simultaneous bison of Western Siberia. Europe (Guenther, 1958; Kahlke, 1958; Flerov, 1977, 1979). Eastern Europe in Riess-Wurm was inhabited by a" very large and long-horned " bison, the span of its horn rods reaching 120 cm (Alekseeva L. I., 1990). Remains of large bison, which in some cases are as large as the Middle Pleistocene B. p. priscus in terms of skull size and horn rods, were found at Late Paleolithic sites in the Angara Valley (Ermolova, 1978). Bison skulls dating back to the Ermakov and Karginsky periods are still practically unknown. Taking into account the decrease in bone size of the postcranial skeleton of B. p. priscus (which had a particularly strong effect at the end of the Karginsky interstadial), it should be assumed that the bison skulls also decreased at this time. The Sartan cold snap again gave some impetus to an increase in the size of the bison, primarily the skull and horn rods. If we take the average values of bison skull measurements from the 6th layer of Krasny Yar as 100 %, then they increased by 3.7% in the Sartan bison from the Orda River. In terms of size and skull structure, the Late Pleistocene bison of the Upper Ob region are comparable to Bison p. priscus (B. p. longicornis), but differed from the latter on average by slightly smaller sizes, especially of horn rods. It is obviously impossible to identify them with Bison priscus mediator (B. p. deminutus).

Thus, the above facts leave no doubt about the gradual (albeit insignificant) decrease in the body size of bison in the south-east of Western Siberia during the middle-late Pleistocene, which affected primarily the skull, and especially the horn rods. This tendency is strongly obscured by the huge individual variability characteristic of members of the genus Bison, and can be detected only in the average values of measurements of sufficiently large, geologically homogeneous series of B. priscus skulls. The fact that the grinding of bison began with the reduction of the horn rods was noticed by V. I. Gromova: "... this process had a particularly strong effect on the shortening of the horns, less on their thinning, and on reducing the size of the skull itself (and probably the entire animal)" [1935, p.131].

The use of bison skulls for biostratigraphic purposes, especially within the Middle - late Pleistocene, is theoretically possible, but, as experience shows, it is hardly feasible in practice. A full-fledged comparison requires a significant, stratigraphically homogeneous series of skulls from one location, which is extremely rare in practice. Interpretation of individual skulls and, especially, horn rods, due to their huge individual-sexual variability (as shown by the materials of Krasny Yar), can lead to erroneous conclusions.

Lower jaw (Mandibula). It is presented only in the materials of Krasny Yar. 57 whole mandibles and their fragments were found in layer 6, and one fragment was found in layer 4. The lower jaws have all the characteristics typical of Bison priscus, an open-space animal (Table 3). Pars dentalis is narrow and high: the height per M3 (on average 82.8 mm) is significantly longer than the length of M l + M2 (60 mm) and slightly longer than the length of M 2 + M3 (79.4 mm). On the molars from the outside, there are additional elements in the form of enamel loops that increase the size of the molars.

page 41
Table 3. Dimensions of the lower and upper jaws of Bison p. priscus Boj.

Dimensions, mm

Krasny Yar, R-W

Mandibula

Maxilla

lim

Jaw length

407 - 492

465,99

P2-M3 alveolar length

156 - 190,2

171,22

135 - 168

153,58

Row length P

56 - 70,5

60,63

51 - 67,3

59,97

Row length M

100 - 124

110,64

78,5 - 106

94,00

Diastema length

121 - 154,5

135,71

Height in the articular process

133 - 169

151,24

Height in the coronal process

216 - 240

229,83

Diastema height, min

33,6 - 46,9

39,78

Diastema thickness, min

16,9 - 26,4

21,86

Height before P 2

42,5 - 62,3

50,41

Height between M 1 and M 2

55 - 77

66,96

Height behind M 3, max

73 - 95

82,81

Thickness under M 3, max

29,5 - 39,3

35,53

Width of the articular process

42 - 62

52,61

P 2, length

11,3 - 14,5

12,98

19,00

width

8,8 - 10,6

9,79

15,30

P 3, length

18 - 23,7

20,22

17,5 - 21,5

19,20

width

11,4 - 14,6

13,00

17 - 20,3

18,93

P 4, length

20,3 - 28

23,54

16 - 21,7

18,49

width

12,7 - 17

15,41

21 - 24,4

22,69

M 1, length

24,5 - 33

27,38

20,5 - 32

25,26

width

16,6 - 26,6

19,26

24,7 - 33

27,50

M 2, length

27,6 - 37

32,57

26,5 - 34,7

31,22

width

17 - 22,4

20,53

23 - 31

28,07

M 3, length

42 - 53,5

46,84

29 - 37

33,06

width

16 - 22,5

19,88

24 - 30,5

28,20

chewing surface and counteracting tooth erasure. Often these loops, especially on M3, have a closed shape.

The postcranial skeleton (Postcranium). Its remains account for over 98 % of the total material studied. In the 6th layer of Krasny Yar and in Taradanov, almost all elements of the postcranial skeleton are represented by significant series-from 20-30 to 100 or more copies of the bones of the same name. This makes it possible to obtain a fairly complete and, in some cases, probably exhaustive morphometric characterization of B. p. priscus skeletal elements. The material from the 4th layer of the Krasny Yar and the Orda River is much less representative. However, it also provides some insight into the morphometric features of bison from the Karginsky and Sartan periods (Table 4). The abundance of phalanges, small carpal bones ,and tarsal bones (mainly from Taradanov) allowed us to give their dimensional characteristics (Table 5). A comparison of the extreme and average values of measurements of elements of the postcranial skeleton of Bison p. priscus from 6The study of the Red Yar and Taradanov layers shows that the bone sizes of these bison, separated, apparently, by a time interval of several tens of thousands of years, practically did not differ from each other. For a number of measurements, the first ones are larger, and for a number of others, the second ones are larger. In general, if we take the bone size of the postcranial skeleton of a bison from the 6th layer of Krasny Yar as 100 %, then the size of the postcranium of a bison from Taradanov decreased by 0.8% in the aggregate of all comparable measurements. A similar comparison of the postcranial remains of B. p. priscus from the 6th and 4th layers of Krasny Yar

page 42
See Table 4. Dimensions of the 1st and 2nd cervical vertebrae and limb bones Bison p. priscus Boj.

Dimensions (mm) and indexes (%)

Krasny Yar, R-W

Taradanovo, W-1-2

Krasny Yar, W-2

Orda River, W-3

lim

Atlas

Width in wings

210 - 256

239,26

214,00

Width of the anterior articular surface

125 - 160

142,24

132,5 - 136

134,83

150 - 166

158,00

Width of the posterior articular surface

114 - 142

132,83

112 - 120

116,67

Highest bone height

100,5 - 129

115,71

102 - 105

103,50

122 - 125

123,50

Axis

Maximum length

109 - 154

139,94

55,40

Vertebral body length

112,7 - 129

120,74

Width of the anterior articular surface

108,3 - 142,3

132,12

117,40

Width of the posterior articular surface

69,5 - 80,3

74,13

57,00

Width of dens ephistrophei

54 - 65,5

59,66

55,40

Vertebral body width, min

70 - 94,8

87,07

74,00

Vertebral height, max

178 - 221,3

196,74

Scapula

Maximum length

571 - 620

594,75

546 - 611

578,79

Width of the upper section

285 - 345

316,25

280 - 320,5

304,31

Neck width

70,5 - 105,5

86,90

70,5 - 98

86,84

74,5 - 76

75,25

Width across the hillock

87,5 - 117

103,22

84,5 - 126

104,94

91 - 98

94,33

Joint surface width

78 - 96,5

88,98

74,5 - 100

88,82

80 - 81,5

80,83

Joint surface diameter

56 - 81,8

70,51

60,5 - 84,2

70,62

57 - 63

60,23

Humerus

1. Bone length

387,5 - 476

432,60

2. Length from head

340 - 414

378,23

341,00

3. Width of the upper end

106 - 153

134,06

4. Upper end diameter

136 - 167

149,70

page 43
Continuation of Table 4

5. Diaphysis width, min

43 - 69

59,68

48,50

62,2 - 64

63,10

53,1 - 65,5

59,30

6. Diameter of the diaphysis, min

46 - 81,2

69,36

57,40

74,5 - 75,5

75,00

63 - 78,7

70,85

7. Width of the lower end

99,3 - 135,5

119,72

100,00

108,5 - 125

115,88

113 - 128

120,50

8. Joint block width

94 - 124,5

108,89

93,00

96 - 109

102,79

100 - 114,5

106,83

9. Medial diameter of the lower end

102,3 - 132

118,10

105,40

111 - 117,5

114,25

104,5 - 125,3

114,90

10. Lateral diameter of the lower end

68,7 - 90

79,55

67,70

69,3 - 81,7

75,50

70,7 - 86,5

78,60

11. Chute diameter, min

46,1 - 63,2

52,45

46,20

46,7 - 51,7

48,42

44 - 55,8

50,10

12. Height of the medial condyle

59 - 76,7

67,98

57,00

53 - 70

63,48

59 - 74

67,13

13. Height of the joint block on the ridge

52,3 - 65,5

58,17

52,00

50,5 - 56,3

53,86

49,3 - 62,7

56,00

3: 1

27,4 - 33,1

30,99

5: 1

13,0 - 15,5

13,80

7: 1

26,7 - 29,9

27,67

3:2

31,2 - 38,8

35,44

5:2

14,8 - 17,6

15,78

15,57

7:2

30,7 - 35,0

31,65

33,14

Radius

1. Bone length

343 - 427

389,87

351,8 - 400,3

374,40

366,00

390 - 391

390,50

2. Width of the upper end

100 - 142

115,93

98,5 - 121,7

106,93

94 - 126,4

107,75

116 - 124

119,00

3. Width of the upper articular surface

94 - 130

107,14

94 - 108,7

99,51

86 - 116,5

100,33

4. Upper end diameter

50 - 71,5

60,32

51,5 - 60,7

54,98

48 - 65

56,56

61 - 68

64,38

5. Width of the diaphysis

51,8 - 86

66,10

49,1 - 71

59,62

50,7 - 58,5

55,58

63 - 67,1

65,05

6. Diameter of the diaphysis

33 - 50

41,33

31,5 - 45,8

37,23

35,5 - 41,5

38,50

7. Width of the lower end

93,2 - 132,5

110,59

82,3 - 111,2

95,79

84 - 98

90,90

111,00

8. Lower end diameter

56,5 - 77,5

67,6

51,7 - 71

60,67

50,3 - 66

56,10

2: 1

27,3 - 33,3

29,74

27,7 - 30,6

28,56

28,85

30,6 - 31,6

30,47

5: 1

14,4 - 20,1

16,95

13,8 - 18,5

15,92

15,44

17,16

7: 1

25,5 - 31,0

28,37

25,3 - 28,1

25,58

26,78

28,39

page 44
Ulna

Maximum length

464 - 515,5

487,89

Olekranon length

135,5 - 173,5

154,87

Diameter of the elbow protuberance

82,7 - 105,5

90,33

Bone body diameter, min

66,2 - 96,6

83,38

72,50

Diameter in the coracoid process

85,5 - 123,3

110,77

115,20

93,5 - 112

102,75

109,5 - 111,5

110,50

Cross-section in a half-moon fillet

53,5 - 79

72,17

56,2 - 77

63,63

60,3 - 73

66,65

69,5 - 72

70,75

Thickness of the elbow protuberance

34,5 - 46

40,77

Joint surface width

52,7 - 77

64,35

57,5 - 64,5

60,03

71,20

63,2 - 65,7

64,45

Metacarpale

1. Bone length

215,2 - 280

240,52

220,3 - 254,5

235,96

226,5 - 246

237,38

218 - 250,7

235,55

2. Width of the upper end

71,5 - 98

85,54

69,8 - 95,4

79,95

73 - 95

81,59

79 - 103

92,63

3. Upper end diameter

40,8 - 56

49,56

38,3 - 55,6

46,41

42,6 - 53

47,41

46 - 59

52,68

4. Width of the diaphysis

42 - 64,5

52,96

40,2 - 61,5

48,11

41 - 60,8

50,16

49 - 62,2

57,48

5. Diameter of the diaphysis

28,5 - 41

34,36

28,5 - 40,7

32,05

29,3 - 36,8

32,48

33 - 43

37,30

6. Width of the lower end

73,7 - 100,2

87,12

72,2 - 93

81,37

72 - 94

83,78

92,7 - 101

97,37

7. Lower end diameter

39,5 - 51,8

46,68

39 - 51

44,47

40 - 51

45,50

47 - 52

50,33

2: 1

31,2 - 42,3

35,56

31,2 - 38,0

33,88

31,6 - 39,6

34,37

39,7 - 44,0

39,32

4: 1

17,2 - 26,9

22,02

17,3 - 25,4

20,39

16,8 - 25,6

21,13

23,6 - 27,3

24,40

6: 1

31,5 - 42,7

36,22

30,6 - 38,8

34,48

32,4 - 39,2

35,29

39,3 - 43,2

41,34

Femur

1. Maximum length

504 - 564

533,89

2. Length from head

450 - 515

488,55

482,00

3. Width of the upper end

163,5 - 199

181,90

4. Upper end diameter

83 - 111,5

98,65

5. Head diameter

61 - 75

68,34

70,00

62,2 - 73

67,60

6. Width of the diaphysis

47 - 60

54,94

55,20

60,00

7. Diameter of the diaphysis

56 - 67

61,51

61,00

61,80

page 45
End of Table 4

8. Width of the lower end

120 - 154

138,17

141,00

9. Medial diameter of the lower end

168,5 - 182,8

174,97

175,70

160,60

10. Lateral diameter of the lower end

128 - 146,5

134,46

134,70

125,30

11. Width of facies patellaris

63 - 80

71,81

69,00

66,00

3: 1

32,5 - 35,7

34,07

6: 1

10,1 - 11,1

10,29

8: 1

25,8 - 28,2

25,88

3:2

34,7 - 38,6

37,23

6:2

10,4 - 12,0

11,25

11,45

8:2

26,5 - 30,5

28,28

29,25

Tibia

1. Bone length

430 - 496

459,44

476 - 489

482,40

413,70

442,50

2. Width of the upper end

122 - 150

136,39

141 - 148,3

144,33

3. Upper end diameter

110,5 - 134,5

123,11

123 - 138

130,80

4. Width of the diaphysis

50 - 74,3

62,50

53,7 - 66,9

60,97

52,3 - 62,5

57,40

60,30

5. Diameter of the diaphysis, min

36,3 - 54,5

43,45

38 - 45,7

42,60

36,1 - 43,1

39,60

6. Width of the lower end

73 - 98,2

85,84

67,3 - 94,6

83,87

79 - 94,3

87,10

76 - 88

80,75

7. Lower end diameter

56,5 - 75,4

65,04

58 - 69

63,83

59 - 72,7

66,40

55 - 63

60,75

2: 1

28,1 - 32,0

29,69

29,5 - 30,3

29,92

4: 1

11,5 - 14,3

13,60

12,5 - 13,7

12,64

13,63

6: 1

17,0 - 19,5

18,68

17,5 - 18,5

17,39

17,97

Astragalus

1. Lateral length

82 - 102,5

91,54

139

76,2 - 104

88,50

82,5 - 95,5

88,80

83,5 - 95

90,16

2. Sagittal length

64,2 - 79,4

71,39

135

70,3 - 92,3

82,03

65,4 - 75,2

69,39

64 - 74,3

70,95

3. Medial length

77,4 - 95,1

84,70

144

58,5 - 80

69,23

77 - 88,9

83,01

76 - 88

84,00

4. Width of the lower end

53,2 - 67,3

60,49

125

47,5 - 63,6

57,00

53,2 - 68,8

58,55

55,5 - 64

60,75

page 46
5. Medial diameter of the lower end

46,5 - 58

52,01

42,5 - 55,3

49,56

46 - 54

49,27

6. Lateral diameter of the lower end

46 - 56,7

51,65

134

41,7 - 57,7

48,94

46,5 - 53,5

49,65

4: 1

61,7 - 74,5

66,08

121

58,2 - 70,7

64,41

58,9 - 72,0

65,93

66,3 - 67,4

67,38

Calcaneus

Bone Length

163 - 201

188,33

170 - 210

186,33

180,50

193,50

Width of the lower end

56 - 75

65,05

51,3 - 74

64,43

62 - 77,5

66,88

74,00

Lower end diameter

66 - 85,7

74,76

65,2 - 82,7

73,70

68 - 77

73,05

77 - 78

77,50

Diaphysis width, min

22,5 - 34,2

28,46

20,7 - 32,7

28,00

23,7 - 36

28,38

Top end width

39,5 - 54,6

49,58

42,7 - 58

50,16

44,70

54,00

Upper end diameter

46 - 57

52,36

44,3 - 59

52,06

50,30

50,00

Centrotarsale

Bone Width

68,2 - 91,4

80,48

63,5 - 88,4

76,16

70,8 - 83,7

75,34

85,50

Width of the upper articular surface

53,2 - 72,5

63,19

51,5 - 68

59,87

56,5 - 67

60,53

70,20

Width of the lower articular surface

57,8 - 76

66,48

57,5 - 75,4

62,97

60 - 68

63,21

Bone diameter

61 - 84

73,32

62 - 85

70,88

66 - 79,5

70,24

82,40

Metatarsale

1. Bone length

278,3 - 314

295,67

264 - 309

292,39

285,5 - 287

286,25

278,5 - 297

288,75

2. Width of the upper end

56,8 - 75,4

66,61

56,2 - 73,2

65,73

57,8 - 61,3

59,60

69,00

3. Upper end diameter

55,5 - 70,5

64,57

56 - 68,5

63,39

55 - 59,2

57,30

4. Width of the diaphysis

34,7 - 50,2

44,43

34,3 - 47,5

41,74

32,8 - 38,8

36,78

35 - 45

40,00

5. The diameter of the diaphysis in the middle

38 - 47

42,02

36,4 - 46,4

42,12

32 - 41

38,25

6. Width of the lower end

68,3 - 86,7

77,66

67,5 - 84

76,57

71,2 - 76,4

73,77

72,5 - 80,7

77,80

7. Lower end diameter

42,1 - 50,9

46,65

40,7 - 50,4

45,99

41,5 - 44

43,10

42 - 47

45,00

2: 1

19,2 - 24,8

22,53

19,8 - 25,5

22,48

21,0 - 21,0

20,82

23,23

4: 1

12,2 - 17,0

15,03

12,2 - 15,5

14,28

13,2 - 13,6

12,85

12,2 - 15,2

13,85

6: 1

24,3 - 29,5

26,27

23,9 - 28,1

26,19

25,7 - 26,8

25,77

26,0 - 28,4

26,94

page 47
See Table 5. Dimensions of the phalanges, kneecaps, wrist and tarsal bones Bison p. priscus Boj.

Dimensions (mm) and indexes (%)

Krasny Yar, R-W

Taradanovo, W-1-2

Krasny Yar, W-2

lim

Phalanx I (front)

1. Maximum length

72,5 - 91,6

79,84

70 - 93

80,35

75 - 82

78,80

2. Sagittal length, min

63,5 - 82

71,43

61 - 82,7

71,85

67 - 74

70,50

3. Width of the upper end

36,5 - 49,5

43,78

34,7 - 52,7

45,63

41,7 - 44,5

43,30

4. Upper end diameter

38 - 56

46,16

38,5 - 55,2

47,18

45,2 - 47

45,80

5. Diaphysis width, min

34 - 47

40,94

33 - 51,2

42,18

38,4 - 41,7

40,22

6. Width of the lower end

35,3 - 49

43,49

35,4 - 56

45,47

40,5 - 44

42,24

7. Lower end diameter

25,5 - 35

30,84

27,2 - 39,2

32,51

29 - 32,3

30,13

3: 1

49,5 - 61,7

54,83

48,3 - 62,6

55,33

50,9 - 56,2

54,95

5: 1

45,2 - 58,3

51,28

46,0 - 59,4

51,15

48,2 - 53,5

51,04

6: 1

48,7 - 61,0

54,47

49,2 - 65,8

55,13

50,0 - 56,4

53,60

Phalanx II (front)

1. Maximum length

46 - 58,8

53,34

101

46 - 62

54,35

51,80

2. Sagittal length, min

39,5 - 49

43,82

104

38,5 - 50,9

44,48

41,00

3. Width of the upper end

39,3 - 54

44,94

34,3 - 52,4

46,58

45,70

4. Upper end diameter

38 - 55

46,41

35 - 54,3

47,90

50,50

5. Width of the diaphysis

31,5 - 44,8

37,21

100

28,5 - 44,2

36,92

37,50

6. Diameter of the diaphysis, min

28,7 - 40

33,42

103

26,4 - 39,5

34,00

32,70

7. Width of the lower end

33,7 - 44,8

38,98

30,7 - 49

41,07

42,70

8. Lower end diameter

35 - 44,3

39,76

32 - 47,5

41,43

41,30

3: 1

74,9 - 94,2

84,25

74,5 - 100,0

85,70

88,22

5: 1

62,4 - 81,5

69,89

61,4 - 85,2

67,93

73,39

7: 1

62,7 - 84,8

73,08

64,4 - 92,1

75,57

82,43

Phalanx III (front + rear)

Maximum length

75 - 104

90,10

76,5 - 108,7

94,54

69 - 106,5

87,92

Dorsal length

58 - 80,8

70,15

61,6 - 85,8

73,29

55 - 79

67,72

Bone Height

45 - 68,3

55,89

48,2 - 71,3

59,23

44,3 - 62,5

53,10

Joint surface height

36 - 51

42,61

37 - 52,6

43,25

38 - 47

41,33

Joint surface width

27 - 41,5

32,01

103

28,2 - 39,7

33,92

26 - 35,5

30,79

Phalanx I (rear)

1. Maximum length

74 - 90

82,01

73,7 - 93

82,35

80,4 - 87,2

83,80

2. Sagittal length, min

66,7 - 88

72,90

64,8 - 82,8

73,69

72 - 76,8

75,13

3. Width of the upper end

33,2 - 47

40,42

35,3 - 48,5

40,96

36 - 43,2

40,10

4. Upper end diameter

39,5 - 52,5

46,44

40,7 - 54

46,26

41,7 - 48,2

45,80

5. Diaphysis width, min

30 - 43

36,46

29 - 43,1

36,64

35,8 - 37,5

36,65

6. Width of the lower end

34,2 - 44,7

39,97

31,2 - 48,1

40,19

40 - 46,8

43,40

7. Lower end diameter

26 - 35

29,73

25 - 36,7

29,63

28 - 29

28,50

3: 1

44,9 - 54,7

49,29

44,5 - 55,1

49,74

48,4 - 48,5

47,85

5: 1

40,5 - 49,8

44,46

35,5 - 50,7

44,49

43,0 - 44,5

43,74

6: 1

45,0 - 52,3

48,74

38,1 - 54,0

48,80

49,8 - 53,7

51,79

page 48
Continuation of Table 5

Phalanx II (rear)

1. Maximum length

50,2 - 59,3

55,07

50 - 63

56,25

60,00

2. Sagittal length, min

43,1 - 49,3

46,97

41,4 - 53,7

47,58

50,50

3. Width of the upper end

35,5 - 45,6

40,00

35,9 - 52

42,10

42,00

4. Upper end diameter

37 - 49

43,53

37 - 54

45,32

45,40

5. Width of the diaphysis

28 - 35,8

31,70

27,5 - 39,3

33,03

30,40

6. Diameter of the diaphysis, min

27,8 - 33,3

29,97

27,3 - 36,4

31,26

27,30

7. Width of the lower end

28,8 - 37

33,59

29,5 - 42,4

35,01

34,70

8. Lower end diameter

32,5 - 42,4

36,61

33 - 45

38,47

36,70

3: 1

68,5 - 78,7

72,63

66,1 - 82,9

74,84

70,00

5: 1

52,8 - 64,8

57,56

50,9 - 66,1

58,72

50,67

7: 1

54,0 - 65,9

61,00

54,3 - 70,5

62,24

57,03

Patella

Length

79 - 95,5

88,13

79,5 - 96,2

88,46

Width

79 - 91,5

83,08

71,5 - 98,3

86,00

Projected height

52 - 60

55,46

43,5 - 60

52,49

Joint surface length

70 - 85

76,89

69 - 84

76,75

Joint surface width

51 - 63

58,22

46 - 66,3

57,54

Carpi radiale

Projection length

50 - 69,5

58,99

50,2 - 72

61,13

Width

31,5 - 44

37,77

32 - 49,5

40,15

Width of the lower joint

surfaces

30 - 42,2

35,99

30 - 44

37,43

Height in the front section, max

35,7 - 45,3

41,38

36,5 - 47,8

41,98

Height in the middle, min

22,5 - 31,8

27,68

24 - 33

28,16

Height in the posterior part (in the process)

34 - 47,5

40,42

36 - 50,3

42,76

Carpi intermedium

Projection length

48,5 - 64

57,14

47,5 - 67

57,24

52 - 58

54,17

Height in the front section, max

37,2 - 42

36,97

32 - 42,5

37,50

35 - 37,3

36,37

Height in the middle, min

25,1 - 33,5

29,72

25,2 - 34

29,48

26 - 29,5

27,57

Front section width, max

33 - 44,8

38,13

33,2 - 44,5

39,25

37,4 - 39

38,20

Carpi ulnare

Width in the projection

28,5 - 40

32,68

30 - 38,5

34,55

30,00

Bone Height

45,3 - 59

50,96

47,3 - 61,6

54,55

49,30

Carpi 2 + 3

Width in the projection

46 - 64,5

55,54

46,5 - 67,5

55,77

53 - 55,5

54,25

Cross section

42 - 53,7

48,52

40,3 - 56,5

48,06

48,5 - 49

48,75

Height, max

23,5 - 35,8

28,70

25,5 - 33,2

28,71

29,80

Carpi 4 + 5

Width

37 - 48,5

43,51

34,9 - 45,5

40,49

Cross section

43,2 - 53,4

47,94

41 - 49,5

45,89

Height, max

30 - 35,5

33,41

28,4 - 34,5

32,11

Carpi accessorium

Maximum length

40 - 44,5

42,45

page 49
End of Table 5

Width of the distal part

38,6 - 45,7

42,03

Distal cross section

27,8 - 32

29,85

Width in the middle, min

32 - 35,2

33,65

Joint surface length

31 - 36,9

34,05

Joint surface diameter

21,7 - 22,6

21,98

Os malleolare

Maximum length

46,2 - 49

47,25

46,1 - 53,3

48,80

51,5 - 52,5

51,75

Maximum diameter

21,7 - 27,7

24,62

21,2 - 30,8

26,33

25,4 - 26,3

25,85

Length of the lower articular surface

35 - 42

37,52

35 - 39

36,48

43,5 - 44

43,75

Diameter of the lower articular surface

17 - 22

19,97

16,4 - 23,4

19,30

20 - 20,5

20,25

Os tarsale IV

Length

48 - 53,5

51,44

50 - 54,2

52,10

Width

30,7 - 34,8

33,01

32 - 33,7

32,85

Height

16,2 - 22

18,60

14,8 - 16

15,40

It shows a significant reduction of the Karginsky interstadial bison-by 4.6 %. In the Sartan period, as shown by materials from the Orda River, the size of skeletal bones increased sharply again (especially against the background of the size depression of bison of the Karginsky period), exceeding those of the Riess-Wurm predecessor by 1.1 %.

In the materials of the Kazantsev period, bones of unusually large bison are sometimes found, which stand out sharply in their size. This is, for example, the radius N 4305: length 427 mm, width of the upper and lower ends and diaphysis, respectively, 142; 132.5 and 86 mm. Prior to its detection, the maximum values of the corresponding radius measurements were 426.5, 129.6, 123.6 and 78 mm.

A comparison of the bone sizes of Bison priscus from the late and earlier stages of the Pleistocene is very interesting. Among the highly mineralized and, as a rule, rounded bone remains of the so-called third preservation group (Q1-2) from the 6th layer of Krasny Yar (Vasiliev, 2002, 2005), a significant number of Bison priscus bones are present. The comparison showed that the sizes (extreme and average values) of the bones of the postcranial skeleton of the Early Middle Pleistocene and Riess-Wurm bison differ little (Table 6). The former are on average somewhat larger and more massive, and among them there are specimens in which the values of a number of characteristics slightly exceed the maximum values for the Kazantsev bison. The smaller size of the astragalus and central cuboid bones of the third group of preservation is mainly explained by their significant roundness. According to A.V. Sher (1997), an increase in the size of the body of Bison priscus was accompanied primarily by an increase in the massiveness of the bones of the postcranial skeleton, without their significant elongation, as well as an increase in the skull and especially in the horn rods.

Palaeotheriologists are usually very interested in the metacarpal bones of Bison. In their works, this topic was also discussed by I. A. Allen [Allen, 1876], I. D. Chersky [1891], and later by E. Schertz [Schertz, 1936], M. F. Skinner and O. K. Kaisen [Skinner, Kaisen, 1947], V. I. Bibikova [1950], B. S. Rusanov [1975Reshetov [1974; Reshetov and Sukhanov, 1979] and a number of other authors. Differentiation of metapods based on gender is of fundamental importance, since the correctness of all further interpretation of the material depends on it. The division of the sample into males and females is usually easy, even visually, without any preliminary measurements or calculations. On average, male bison metacarps are somewhat longer than those of females, and they differ dramatically in their massiveness. The most significant feature in this respect is the width of the diaphysis, where transgression is completely absent. In the measurements and indices of the width of the upper and lower epiphysis, it is observed only to a weak degree (Tables 7, 8). The range of individual metacarpal variability is impressive. Extreme examples of variability are metacarps of males from the 6th layer of Krasny Yar N 3044 and 154 with a length of 280 and 227 mm, respectively, and a relative width of the diaphysis - 21.8 and 26.9.

page 50
Table 6. Limb bone sizes of Early - Middle Pleistocene Bison priscus (Q1 - 2) and Kazantsev time (R-W) from Krasny Yar

Dimensions, mm

Q1 - 2

R-W

lim

Humerus

Diaphysis width

59,8 - 66,8

64,0

43 - 69

59,7

Joint block width

105 - 119

111,5

94 - 124,5

108,9

Radius

Bone Length

371,5 - 413

398,6

343 - 427

389,9

Diaphysis width

52 - 72,3

62,8

51,8 - 86

66,1

Top end width

104,7 - 125,7

115,8

100 - 142

115,9

Metacarpale

Bone Length

221 - 257,2

239,1

215,2 - 280

240,5

Top end width

69,5 - 90,3

88,6

71,5 - 98

85,5

Diaphysis width

44,7 - 61,4

54,5

42 - 64,5

53,0

Diameter of the diaphysis

30,6 - 39,5

34,1

28,5 - 41

34,4

Width of the lower end

72 - 94,5

83,0

73,7 - 100,2

87,1

Astragalus

Lateral length

77,3 - 101,7

89,6

82,1 - 102,5

91,5

Centrotarsale

Bone Width

65 - 88

77,9

68,2 - 91,4

80,5

Metatarsale

Bone Length

275 - 319,5

292,8

278,3 - 314

295,8

Diaphysis width

39,8 - 49,5

44,8

34,7 - 50,2

44,4

Width of the lower end

69,5 - 82,2

77,3

68,3 - 86,7

77,7

Very interesting results were obtained by studying the metacarpal bones of bison according to the method used by V. Eisenmann [Eisenmann, 1979; Eisenmann and Beckouche, 1986] in the study of modern and fossil horses. The value of each of the seven measurements (Tables 7, 8) was correlated with the corresponding parameter of males or females of modern Bison b. bonansus (Reshetov and Sukhanov, 1979); the decimal logarithm of this ratio was calculated. The obtained values were plotted on the coordinate field; the points were connected in a characteristic curve. Comparison of different forms of Bison was made using the average values of measurements and indices. Metacarpal bones of Late Pleistocene B. p. priscus from Kurtak (Krasnoyarsk region) and the Chumysh River of Altai Krai were used as a comparative material. We also used literature data on metacarps B. priscus from the Upper Pleistocene of Belgium (Germonpre, 1993), B. p. crassicornis (Q2) and B. p. occidentalis (Q3) from various localities in Yakutia (Rusanov, 1975), B. p. deminutus (mediator) (Q3) from Amvrosievka (Ukraine), which is considered as a place of death of herds as a result of Paleolithic man's corral hunting (Bibikova, 1950)*. For all the above samples, graphs of average metacarpal proportions were plotted separately for males and females (Tables 7, 8; Figs. 3, 4). The comparison showed extreme similarity, if not identity, of bison metacarps from Krasny Yar, Taradanov, Kurtak, and Chumysh. Bison from Europe and Yakutia differ markedly from the latter in absolute size and proportions. Analysis of the metacarpal bones once again confirms the significant conservativeness in the size of the B. priscus postcranium in the Late Pleistocene of southern Western Siberia observed on other skeletal elements. In this regard, the application of the considered methodology for biostratigraphic purposes in this region is not very promising. At the same time, it allows us to judge the systematic proximity of various representatives of the genus Bison: in closely related forms, the curves on the graphs are similar, and in taxonomically remote ones -

* Unfortunately, this article does not contain measurements of metacarpale cross-sections; the missing data were kindly provided by N. D. Ovodov, who read V. I. Bibikova's dissertation at the time.

page 51
Table 7. Dimensions of metacarpal bones of the genus Bison (males)

Dimensions (mm) and indexes (%)

B. p. priscus

Krasny Yar, R-W

Taradanovo, W-1-2

Krasny Yar, W-2

Kurtak, Q 3

Chumysh River, Q 3

lim

1. Bone length

223 - 280

242,99

222 - 254,5

242,34

229,3 - 246

238,25

231,2 - 251

241,12

218 - 260,6

240,86

2. Width of the upper end

80,5 - 98

89,82

83,7 - 95,4

89,33

84,1 - 95

90,03

82 - 101,5

90,98

79,7 - 99

89,48

3. Upper end diameter

46 - 56

52,11

47,5 - 55,6

51,14

47,6 - 53

50,98

47,3 - 60

52,23

45 - 58,8

51,79

4. Width of the diaphysis

53,5 - 64,5

57,65

53 - 61,5

56,18

54,2 - 60,8

57,10

52,7 - 65,1

58,22

52,4 - 65

57,96

5. Diameter of the diaphysis

31,9 - 41

36,22

33,2 - 40,7

35,94

32,5 - 36,8

35,35

32,5 - 39,5

36,04

31,2 - 39,8

36,23

6. Width of the lower end

81 - 100,2

91,46

86,1 - 93

89,35

89,3 - 94

91,08

78,3 - 104,4

91,21

82 - 99,6

91,59

7. Lower end diameter

43,3 - 51,8

47,94

43,1 - 51

48,19

45,2 - 51

47,10

43 - 55,6

47,39

41,5 - 54,4

48,22

2: 1

33,8 - 42,3

36,96

35,5 - 38,0

36,86

36,7 - 39,6

37,79

33,7 - 41,6

37,73

34,0 - 41,7

37,15

4: 1

21,2 - 26,9

23,73

21,8 - 25,4

23,18

22,6 - 25,6

23,97

21,7 - 26,6

24,15

22,1 - 27,4

24,06

6: 1

33,7 - 42,7

37,64

35,8 - 38,8

36,87

36,6 - 39,2

38,23

33,7 - 42,8

37,83

35,0 - 41,9

38,03

End of Table 7

Dimensions (mm) and indexes (%)

B. p. crassicornis, Yakutia, Q 2 (Rusanov, 1975)

B. p. occidentalis, Yakutia, Q 3 (Rusanov, 1975)

B. P. mediator, Ukraine, Q 3 [Bibikova, 1950]

B. priscus, Belgium, Q 3 [Germonpre, 1993]

B. b. bonansus, modernity (Reshetov and Sukhanov, 1979)

lim

1. Bone length

220 - 249

230,5

214 - 227

217,6

214,5 - 245,4

232,4

225 - 292

246,43

202 - 225

215,0

2. Width of the upper end

85 - 98

89,5

80 - 85

83,0

76,4 - 90,4

84,5

78,1 - 94,8

86,13

70 - 79

74,0

3. Upper end diameter

42 - 59

48,3

42 - 48

45,4

44,2 - 52,6

48,8

46,8 - 59,5

52,02

39,3 - 45,8

42,8

4. Width of the diaphysis

53 - 63

57,1

45 - 56

51,1

46,6 - 57,8

53,1

51,1 - 57,6

55,29

41 - 49

44,0

5. Diameter of the diaphysis

32 - 43

35,57

28 - 36

33,3

30,6 - 36,8

33,7

32,8 - 40,7

36,69

27,6

6. Width of the lower end

84 - 99

90,8

81 - 89

85,6

80 - 98,1

87,0

82,4 - 93,1

86,74

66 - 75

70,0

7. Lower end diameter

43 - 53

46,1

43 - 50

44,7

41 - 49,1

45,4

39 - 57,4

47,53

37,2 - 42,4

39,8

2: 1

36,5 - 41,9

38,8

35,6 - 39,7

38,1

36,36

32,5 - 39,0

34,95

30,6 - 37,6

34,8

4: 1

23,1 - 25,9

24,8

21,1 - 26,1

23,5

22,85

19,6 - 23,8

22,44

18,7 - 22,7

20,6

6: 1

34,8 - 41,8

39,4

37,4 - 41,5

39,3

37,45

30,8 - 37,5

35,20

30,7 - 34,9

33,3

page 52
Table 8. Dimensions of metacarpal bones of the Bison genus (females)

Dimensions (mm) and indexes (%)

B. p. priscus

Krasny Yar, R-W

Taradanovo, W-1-2

Krasny Yar, W-2

Kurta k, Q 3

Chumysh River, Q 3

lim

1. Bone length

215,2 - 258,5

236,65

220,3 - 245,2

233,35

226,5 - 245,5

236,50

213,4 - 247,3

232,55

221 - 253

234,96

2. Width of the upper end

71,5 - 88,8

78,62

69,8 - 82,5

76,57

73 - 80,3

75,97

73,4 - 83,6

78,16

71,3 - 83,6

78,57

3. Upper end diameter

40,8 - 54,6

45,83

38,3 - 50,1

44,69

42,6 - 45

43,85

40 - 50

45,14

40,7 - 48,5

45,76

4. Width of the diaphysis

42 - 50,6

46,10

40,2 - 49,5

45,42

41 - 48,1

44,60

41,2 - 51,5

46,23

41,2 - 51

47,30

5. Diameter of the diaphysis

28,5 - 34,3

31,56

28,5 - 34,3

30,75

29,3 - 31,2

30,57

26,8 - 35

31,43

28,8 - 35,9

31,79

6. Width of the lower end

73,7 - 88,8

80,09

72,2 - 82,4

78,34

72 - 79,5

76,48

74 - 87

79,58

71,2 - 85

79,84

7. Lower end diameter

39,5 - 49,8

44,21

39 - 46,5

42,99

40 - 45,9

43,37

39,6 - 46,9

42,93

38,3 - 47

43,90

2: 1

31,2 - 36,4

33,22

31,2 - 34,9

32,81

31,6 - 32,5

32,12

31,3 - 36,6

33,61

31,3 - 36,3

33,44

4: 1

17,2 - 20,9

19,48

17,3 - 21,6

19,46

16,8 - 20,3

18,86

17,8 - 21,7

19,88

18,6 - 21,9

20,13

6: 1

31,5 - 36,6

33,84

30,6 - 36,3

33,57

32,4 - 32,8

32,34

31,5 - 37,6

34,22

30,2 - 36,9

33,98

End of Table 8

Dimensions (mm) and indexes (%)

B. p. crassicornis, Yakutia, Q 2 (Rusanov, 1975)

B. p. occidentalis, Yakutia, Q 3 (Rusanov, 1975)

B. P. mediator, Ukraine, Q 3 [Bibikova, 1950]

B. priscus, Belgium, Q 3 [Germonpre, 1993]

B. b. bonansus, modernity (Reshetov and Sukhanov, 1979)

lim

1. Bone length

221 - 244

229,9

204 - 227

217,1

206,2 - 237,1

224,69

215 - 230

232,20

201 - 209

205,0

2. Width of the upper end

75 - 82

78,3

70 - 78

73,2

69,5 - 78,6

72,18

62,3 - 79,3

73,16

62 - 72

67,0

3. Upper end diameter

40 - 49

47,1

36 - 44

40,0

40,2 - 47,1

43,10

39,2 - 47,1

43,57

37,3 - 43,1

40,0

4. Width of the diaphysis

42 - 48

45,9

40 - 49

43,9

40,7 - 48,1

42,36

38,6 - 49,9

43,84

32 - 38

35,0

5. Diameter of the diaphysis

29 - 36

31,5

28 - 37

30,3

27,7 - 32,4

29,30

27,6 - 33,3

30,64

24,1

6. Width of the lower end

76 - 86

79,8

74 - 79

75,8

71,5 - 81,6

75,60

68,4 - 81,2

73,77

60 - 67

63,0

7. Lower end diameter

41 - 47

43,7

39 - 42

39,9

39 - 44

41,10

36,9 - 43,7

40,73

34,6 - 37,9

36,5

2: 1

31,9 - 35

34,1

30,2 - 35,5

33,7

32,12

30,8 - 33,1

31,51

30,7 - 34,1

32,3

4: 1

15,6 - 21

20,0

18,6 - 21,4

20,2

18,85

16,9 - 20,8

18,88

15,9 - 18,6

17,3

6: 1

32,3 - 38

34,7

33,8 - 38,7

34,9

33,65

31,2 - 32,7

31,77

29,3 - 32,2

30,6

page 53

3. Diagrams of the average proportions of metacarpal bones Bison p. prisons.

1-Krasny Yar, R-W; 2-Krasnoyarsk reservoir, Kurtak, Q 3; 3-Altai Territory, Chumysh River, Q 3.

4. Diagrams of the average proportions of the metacarpal bones of Bison priscus.

1-B. p. priscus, Krasny Yar, R-W; 2-B. p. crassicornis, Yakutia, Q2 (according to [Rusanov, 1975]); 3-B. p. priscus, Belgium, Q3 (according to [Germonpre, 1993]); 4 - B. p. mediator, Ukraine, Q 3 (according to [Bibikova, 1950]); 5-B. p. occidentalis, Yakutia, Q 3 (according to [Rusanov, 1975]).

they differ significantly. The use of this technique requires compliance with two essential conditions: a geologically equal-aged good series of bones and the correct separation of them by gender.

Discussion

The collection of IAET SB RAS contains excellent materials on Late Pleistocene bison from cave localities in Altai, such as the Hyena Den, Okladnikov Cave, etc. At the Krasnoyarsk Pedagogical University, the author processed large collections of bison remains (mainly from the Karginsky-Sartan period) from the already mentioned Kurtak and a number of other localities along the banks of the Krasnoyarsk reservoir. A special place is occupied by materials from the Larch site near Divnogorsk, dating back to the Sartan period (Akimova et al., 2005). The study of all these collections showed that in the second half of the late Pleistocene Altai and the south of Central Siberia were inhabited by a very large form of Bison priscus, which on average slightly exceeded the size of the bones of the postcranial skeleton of the simultaneous bison of the Upper Ob region.

Thus, an analysis of all available data leads to the conclusion that in the south of Western Siberia, at least since the Middle Pleistocene, there was a directed, but generally insignificant decrease in the body size of Bison priscus. Even among the most recent bison, which lived 20-12 thousand years AGO, there were individuals whose postcranium bones were not inferior in size to the Middle Pleistocene Bison priscus: a slight decrease in size is detected only when comparing the average ones

page 54
measurement values. I. V. Foronova (2001) notes that the line of very large steppe-forest-steppe bison from the priscus group has been traced in the south of Western Siberia since the early Pleistocene, while Western Europe at that time was inhabited by the smaller B. schoetensacki, which was mainly a forest inhabitant [Flerov, 1972; Flerov, David, 1971]. The natural environment in the south of Western Siberia during the Pleistocene did not change as dramatically as in higher latitudes. During the glacial epochs, cold steppe and partly forest - steppe landscapes dominated here, and during the interglacial periods, mainly forest-steppes (Arkhipov, 1971; Arkhipov and Volkova, 1994). The presence of favorable open and semi-open areas with rich food resources probably caused a significant conservativeness in the body size of Bison priscus until its final extinction in this area at the turn of the Holocene. On the other hand, the bison of Europe and northeastern Siberia, by the end of the late Pleistocene, were heavily crushed, transforming into the modern European bison ( Bison b). bonansus and the American forest bison, B. b. athabascae, which disappeared in Yakutia during the Holocene, but has been preserved to this day in northwestern Canada (Flerov and Zablotsky, 1961; Flerov, 1977, 1979; Rusanov, 1975). In the Holocene, small populations of bison continued to exist in a number of places (the most snow-free?). Southern Siberia. Individual fragmentary remains of such bison were found in layers of Middle Holocene monuments of the Priishimskaya forest-steppe (Kosintsev, 1988). In the steppe and forest-steppe zones of Transbaikalia and the Baikal region, bison remains in settlement layers are known up to the 8th-10th centuries AD (Ermolova, 1978).

Conclusion

The study showed that since the end of the Kazantsev period (about 100-90 thousand years AGO), a slight decrease in the body size of Bison p. priscus occurred in the south - east of Western Siberia. It reached its maximum at the final stage of the Karginsky time (about 30 thousand years ago). However, unlike Bison priscus in Europe and Eastern Siberia, bison in the south of Western and Central Siberia did not experience such significant crushing. During the Sartan cold snap, the size of the bison skull and postcranial skeleton (and body, respectively) increased dramatically, again approaching the parameters of Bison p. priscus at the end of the Kazantsev Interglacial.

Conservativeness in the size and morphology of skeletal bones makes the remains of Bison p. priscus unpromising for biostratigraphic studies of Late Pleistocene deposits, which is compounded by the huge scale of their individual-sexual variability.

A series of female bison skulls were collected in situ in the sediments of layer 6 of Krasny Yar, which allowed for the first reliable sex-based separation of cranial remains. For a number of reasons, female skulls are rarely preserved in the fossil state. According to the range of individual and sexual variation in skull size, the Late Pleistocene Bison p. priscus was comparable to the modern representatives of the genus Bison.

The remains of Bison p. priscus in the south of Western Siberia can be used as a kind of indicator of forest-steppe landscapes. The most significant feature here is the ratio of bison and horse bones. The more settled the territories were, the more significantly the number of bison decreased, and the number of horses, on the contrary, increased noticeably; the spread in the watershed areas of forest-steppes caused the opposite process.

List of literature

Akimova E. V., Drozdov N. I., Laukhin S. A., Cheha V. P., Orlova L. A., Koltsova V. G., Sanko A. F., Shpakova E. G. Paleolithic of the Yenisei: Larch. Novosibirsk; Krasnoyarsk: Izd-vo IAET SB RAS, 2005, 180 p. (in Russian)

Alekseeva, L. I., Theriofauna of the Upper Pleistocene of Eastern Europe (large mammals), Nauka Publ., 1990, 109 p. (in Russian).

Alekseeva E. V. Mammals of the Pleistocene of the South-East of Western Siberia, Moscow: Nauka Publ., 1980, 188 p.

Arkhipov S. A. The Quaternary period in Western Siberia. Moscow: Nauka Publ., 1971, 331 p. (in Russian)

Arkhipov S. A., Volkova V. S. Geological history, landscapes and climates of the Pleistocene of Western Siberia. Novosibirsk: SIC OIGSM SB RAS, 1994, 105 p. (in Russian)

Baskin L. M. Ekologiya i povedenie zubr [Ecology and behavior of the bison]. Zubr: Morfologiya, sistematika, evolyutsiya, ekologiya [Bison: Morphology, systematics, evolution, ecology], Moscow: Nauka Publ., 1979, pp. 442-470.

Bibikova, V. I., On some biological features of the primitive bison, Byul. Department of Biol. -1950. - Vol. 55(5). - pp. 35-43.

Burchak-Abramovich N. I., Naniev V. I. Bison in North Ossetia // Tr. Estestv.- istor. G. Zardabi Museum. - Baku, 1954. Issue 8, pp. 134-189.

Vasiliev S. K. Fauna of large mammals of the Kazantsev and Karginsky periods of the Novosibirsk Ob region based on the materials of the Krasny Yar locality / / Fauna of the Urals in the Pleistocene and Holocene. Yekaterinburg: Universitet Publ., 2002, pp. 62-70.

Vasiliev S. K. Taphonomic features of the Taradanovsky secondary alluvial locality //

page 55
Problems of archeology, ethnography, and anthropology of Siberia and adjacent territories. Novosibirsk: Izd-vo IAET SB RAS, 2004, vol. 10, pp. 164-168.

Vasiliev S. K. Large mammals of the Kazantsev and Karginsky time of the Novosibirsk Ob region (based on the materials of the Krasny Yar locality): Author's abstract of the cand. biol. nauk. Novosibirsk, 2005, 26 p. (in Russian)

Vasiliev S. K., Nikolaev S. V., Petrin V. T. On the problem of taphonomy of paleontological remains in the "zaymishchi" of the South of Western Siberia / / Problems of protection, study and use of the cultural heritage of Altai. Barnaul: Alt State University Publ., 1995, pp. 40-42.

Vasiliev S. K., Orlova L. A. K voprosu o vozraste Taradanovskogo mestorozhdeniya fauny krupnykh mammaliyashchikh [On the age of the Taradanovka locality of large mammal fauna]. Novosibirsk: Izd-vo IAET SB RAS, 2006, vol. 12, part 1, pp. 36-42.

Volkov I. A., Arkhipov S. A. Quaternary deposits of the Novosibirsk region. Novosibirsk: IGiG SB AS USSR Publ., 1978, 90 p. (in Russian)

Gromova V. I. Primitive bison (Bison priscus Bojanus) in the USSR // Tr. Zoological Institute of the USSR Academy of Sciences, 1935, vol. 2, vol. 2/3. - p. 77-205.

Ermolova N. M. Teriofauna of the Angara Valley in the late anthropogenic period. Novosibirsk: Nauka Publ., 1978. -220 p.

Kosintsev P. A. Holocene remains of large mammals of Western Siberia / / Current state and history of the animal world of the West Siberian lowlands. Sverdlovsk: Ural Branch of the USSR Academy of Sciences, 1988, pp. 32-51.

Nemtsev A. S., Rautian G. S., Puzachenko A. Yu., Sipko T. P., Kalabushkin B. A., Mironenko I. V. Bison in the Caucasus. - Moscow; Maikop: Kachestvo, 2003. - 292 p.

Panychev V. A. Radiocarbon chronology of alluvial deposits of the Pre-Altai plain. Novosibirsk: Nauka Publ., 1979, 132 p. (in Russian)

Reshetov V. Y. Comparative osteological analysis of the postcranial skeleton of bison in connection with the history of the genus Bison. Novosibirsk: Nauka Publ., 1974, vol. 2, pp. 57-64.

Reshetov V. Yu., Sukhanov V. B. Postcranial skeleton / / Bison: Morphology, systematics, evolution, ecology. - Moscow: Nauka, 1979. - pp. 142-195.

Rusanov B. S. Fossilized bison of Yakutia. Yakutsk: Kn. izd-vo, 1975, 143 p. (in Russian)

Sokolov I. E., Tembotov A. K. Mammals: ungulates. Moscow: Nauka Publ., 1993, 528 p. (Vertebrates of the Caucasus).

Flerov K. K. The oldest representatives and history of the Bison genus. Novosibirsk: Nauka Publ., 1972, vol. 1, pp. 81-86.

Flerov K. K. Bizony Severo-Vostochnoy Sibiri [Bison of North-Eastern Siberia]. Tr. Zool. inta AN SSSR, 1977, vol. 73, pp. 39-58.

Flerov K. K. Systematics and evolution / / Bison: Morphology, systematics, evolution, ecology. - Moscow: Nauka, 1979. - pp. 9-127.

Flerov K. K., David A. I. The genus Bison H. Smith / / Pleistocene of Tiraspol. Chisinau: Stiinca Publ., 1971, pp. 156-165.

Flerov K. K., Zablotsky M. A. O prichinakh izmeneniya arealov bizonov [On the causes of changes in bison ranges]. Byul. MOIP. Ser. biol. - 1961. - Vol. 66 (6). - pp. 99-109.

Foronova I. V. Quaternary mammals of the South-east of Western Siberia (Kuznetsk basin). Novosibirsk: Publishing House of the Siberian Branch of the Russian Academy of Sciences, Phil. "Geo", 2001, 243 p.

Chersky I. D. Description of the collection of post-tertiary mammals collected by the Novosibirsk expedition of 1885-1886-St. Petersburg, 1891-706 p. - (Appl. to vol. 12 " Zap. Imp. Acad. sciences").

Shpansky, A.V., Finding of bison remains in the Tomsk Ob region, Byul. Department of Geology, 1997, Issue 1, pp. 46-52.

Allen I.A. The american bison, living and extinct // Mem. Mus. Compar. Zool. Harvard Coll. - 1876. - Vol. 4, N 10. - P. 1 - 206.

Driesch A., von den. A qiude to the measurement of animal bones from archaeological sites // Peabody Mus. Bull. -1976. - N1. - P. 1 - 136.

Eisenmann V. Les metapodes d'Equus sensu lato (Mammalia, Perissodactula) // Geobios. - 1979. - Vol. 12, N6. - P. 863 - 886.

Eisenmann V., Beckouche S. Identification and discrimination of metapodials from Pleistocene and modern Equus, wild and domestic // Equids in the Ancient World / eds. H. Meadow, H. -P Uerpmann. -Wiesbaden: L. Reichert Verlag, 1986. - P. 116 - 163.

Germonpre M. Osteometric data on Late Pleistocene mammals from the Flemish Valley, Belgium. - Brussels: Department of Paleontology Royal Belgian Ynstitute of Natural scienses, 1993. - 136 p. - (Studiedocumenten vanhet Koninklijk Belgisch Ynstituut voor Natuurwetenschappen; N 72).

Guenther E.W. Feinstratigraphische Unter chung einer Zwischenschicht und der Dechschten der Travertine von Ehringsdorf // Alt-Thuringen: Jahressachriff des Museums fur Ur- und Fruhgeschichte Thuringens. - Weimar, 1958. - Bd. 3. - S. 1 - 9.

Kahlke H.-D. Die jungpleistozanen Saugetierfaunen aus dem Travertingebiet von Taubach - Weimar - Ehringsdorf // Alt-Thuringen: Jahressachriff des Museums fur Ur- und Fruhgeschichte Thuringens. - Weimar, 1958. - Bd. 3. - S. 9 - 130.

Schertz E. Der Geschlechts-Unterschied an Metapodien vonBison // Senkenbergiana. - 1936. - Bd. 18. - S. 357 - 381.

Sher A.V. An Early quaternary bison population from Untermassfeld: Bison menneri sp. nov. // Monogr. Rom. - Mainz: Germ. Zentralmus. - 1997. - Vol. 40, N 1. - P. 101 - 180.

Skinner M.F., Kaisen O.C. The fossil bison of Alaska and preliminary revision of the genus // Bull. Amer. Mus. Natur Hist. - 1947. - Vol. 89, N 3. - P. 123 - 256.

The article was submitted to the Editorial Board on 13.08.07.

page 56

Permanent link to this publication:

https://elib.jp/m/articles/view/BISON-Bison-p-priscus-Bojanus-1827-OF-THE-LATE-PLEISTOCENE-IN-SOUTHEASTERN-WESTERN-SIBERIA

Similar publications: L^Japan L^World Y G

Publisher:

Haruto Masaki → Contacts and other materials (articles, photo, files etc)

Author's official page at Libmonster: https://elib.jp/Masaki

Find other author's materials at: Libmonster (all the World) • Google • Yandex

Permanent link for scientific papers (for citations):

S. K. Vasiliev, BISON (Bison p. priscus Bojanus, 1827) OF THE LATE PLEISTOCENE IN SOUTHEASTERN WESTERN SIBERIA // Tokyo: Japan (ELIB.JP). Updated: 14.12.2024. URL: https://elib.jp/m/articles/view/BISON-Bison-p-priscus-Bojanus-1827-OF-THE-LATE-PLEISTOCENE-IN-SOUTHEASTERN-WESTERN-SIBERIA (date of access: 19.05.2025).

Found source (search robot):

Archeology, Ethnography and Anthropology of Eurasia, No. 2.30 June 2008 Pages 34-56

Publication author(s) - S. K. Vasiliev:

S. K. Vasiliev → other publications, search: Libmonster Japan • Libmonster World • Google • Yandex

Comments:

Reviews of professional authors

Order by: expand all

Per page:

There are no comments yet

New publications:

News from other countries:

ELIB.JP - Japanese Digital Library

Create your author's collection of articles, books, author's works, biographies, photographic documents, files. Save forever your author's legacy in digital form. Click here to register as an author.

Library Partners

	Editorial Contacts Chat for Authors: JP LIVE: We are in social networks:
	About · News · For Advertisers

Permanent link to this publication:

Publisher:

Permanent link for scientific papers (for citations):

Found source (search robot):

Publication author(s) - S. K. Vasiliev:

Comments:

New publications:

Popular with readers:

News from other countries:

ELIB.JP - Japanese Digital Library