Introduction

The population of any region develops to a large extent through its interaction with its closest neighbors, and often with the participation of groups that are significantly geographically distant, but are involved in ethnogenesis by events of different nature: mass migrations due to changes in the ecology of the environment, redistribution of geopolitical influence and the related structure of interethnic relations, transformations in the sphere of material and spiritual culture. societies, etc.

In historical retrospect, we observe a change of cultures and peoples in the same territory. An objective approach to the history of the population of any region requires an analysis of the continuity of all structural elements of multicomponent ethno-cultural formations, including genetic continuity. This aspect of the problem has long been solved by methods of physical anthropology, which allowed us to extract genetic information only indirectly. The current stage of development of biological science provides an opportunity to study the genetic history of populations by means of DNA extracted from paleoanthropological material. Studying the structure of gene pools by analyzing DNA sequences gives a more detailed idea of the origin of populations.

It is now customary for the history of human exploration of any part of the ecumene to be decided by the joint efforts of archaeologists, physical anthropologists and geneticists. Physical anthropology allows us to scan the morphological features of the entire array of remains of carriers of ancient cultures provided by archaeologists and identify key episodes in the reconstructed processes of formation of their anthropological composition. This helps to orient the time-consuming genetic analysis in such a way that sufficient information can be extracted at minimal cost to determine the ethno-cultural space in which different groups of the population are found.

This work was supported by the Russian Foundation for Scientific Research, project No. 06-01-00321a.

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common genetic markers that indicate their relationship, traced when the elements of the cultural complex are dissimilar.

The present study is based on paleoanthropological materials obtained from funerary dents of several archaeological cultures that existed on the territory of Gorny Altai in the chronological range from the Neolithic era (IV millennium BC) to the turn of the new era. It was carried out using the methods of physical anthropology (craniometry) and molecular genetics (analysis of GWS I of the control region of mitochondrial DNA) and aims to find out the main vectors of genetic relationships in the population represented by them.

Earlier, we used the same set of methods to study the carriers of the Pazyryk culture of the Scythian period (IV-III centuries BC), which allowed us to identify their anthropological composition, discuss the formation of the selected components, and outline a promising line of succession of Pazyryk genotypes in the southern Samoyedic groups (Chikisheva, 2003b; Voevoda et al., 2003). From the block of results obtained, we selected for further development the questions of the retrospective plan related to the genetic origins of the Pazyryk groups.

The Pazyryk paleoanthropological material originates from several burial grounds located in the valleys of the main rivers of Gorny Altai. The study of the variability of morphological complex features in craniological series formed in accordance with the location of burials in certain river valleys gives us the right to consider these series as samples from populations (Chikisheva, 2000a, 2003b). Since the paleogenetic analysis was carried out earlier only in one Pazyryk population, which was developing the Ak-Alakha River valley on the Ukok plateau, we considered it necessary to start studying the genetic material in other paleopopulations. In this regard, we studied mitochondrial DNA samples in the paleoanthropological series from the Ulandryk-1 burial ground located in the valley of the river of the same name.

To retrospectively study the genetic relationships of the Pazyryk people, we conducted a study of mitochondrial DNA from bone samples originating from burials of previous cultures: the Neolithic (IV millennium BC) and Karakol period of the developed Bronze Age (first half - mid II millennium BC). It should be noted that the formation of this sample was determined by two tasks. The first is to test at the genetic level the hypothesis about the autochthonous origin of one of the main components of the anthropological composition of the carriers of the Pazyryk culture. It was identified by us at the craniological level [Chikisheva, 1994, 1996, 2000a, 2002, 2003b] and can be characterized as an anthropological type with an intermediate expression of the main features differentiating the Mongoloid and Europoid races. Individuals with this morphology predominate in ordinary burials of the Pazyryk culture (Chikisheva, 2003b). Comparative craniological analysis gave grounds to assume their genetic connection with the Neolithic people and the carriers of the Karakol culture of the Bronze Age of Gorny Altai. The second task of this work is related to the development of a hypothesis proposed by one of the authors of this article about the protomorphy of this morphological complex and the independence of its genesis from the mestizoization of Caucasians and Mongoloids [Chikisheva, 2000b, 2003b].

So, in this study, we studied the haplotype diversity of GWS in the first mtDNA control region from bone samples of the Neolithic era (IV millennium BC) and from burials of the Karakol culture of the Advanced Bronze Age (first half - middle II millennium BC), as well as compared it with that of the remains of the later Pazyryk culture (IV millennium BC). - III centuries BC).

Material and methods

The sample includes 11 samples from burials of various burial grounds located from several tens to several hundred kilometers apart in the Altai Republic (Fig. 1, Table 1).

According to a number of craniological features of high differentiating significance, many skeletons that make up this sample are characterized by features of a kind of intermediacy of their phenotypic parameters in comparison with representatives of the Caucasian and Mongoloid communities (main races). These are signs of general flatness of the facial skeleton and its individual structures: nasomalar and zygomaxillary angles of the horizontal profile, the angle of protrusion of the nasal bones, simotic and dacrial indicators of the protrusion of the nose (Table 2).

Isolation of total DNA from bone. For each sample, the procedure for extracting DNA from bone tissue was repeated several times. The spongy bone fragment was pre-fired in the flame of an alcohol lamp and irradiated with ultraviolet light on each side for 1 h, then ground into a fine powder. To extract total DNA, 4 ml of lysing buffer (4 M guanidine thiocyanate: 0.1 M NaCl, 0.014 M beta-mercaptoethanol, 0.025 M EDTA, 0.5% SDS) was added to 1 g of bone material and incubated for 12 h at 60°C. DNA extraction was performed twice phenol-

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Figure 1. Localization of the sites from which the samples originate.

1-Karakol; 2-Karakol-1; 3-Kaminnaya cave; 4-Ulandryk; 5-Bertek; 6 - Ak-Alakha; 7 - Moynak; 8 - Kuturguntas.

Table 1.

Geographical localization and cultural and chronological characteristics of samples

N n/a

Place of sample collection

Geographical localization

Cultural and chronological characteristics

1

Fireplace Cave

North-western district of Gorny Altai

Neolithic (IV millennium BC)

2

Karakol-1, 1982, mound 3, central pit

The same thing

Karakol culture (mid-2nd millennium BC)

3

Karakol, 1985, mound 3, border 2

Central district of Gorny Altai

The same thing

4

Bertek-56

Ukok Plateau

Mid-2nd millennium BC

5

Ak-Alakha-5, mound 5

The same thing

Pazyryk culture (IV-III centuries BC)

6

Ak-Alakha-1, mound 1, border 2

"

The same thing

7

Ak-Alakha-5, mound 3, border 1

"

"

8

The same, mound 4, border 1

"

"

9

Moynak-2, mound 2, border 1

"

"

10

Kuturguntas-1, kurg. 1

"

"

11

Ulandryk-1, mound 14, border 2

Ulandryk River Valley

"

chloroform and once-chloroform. DNA in 1 M NaCl was precipitated with isopropanol, followed by double washing of the precipitate with 80% ethanol. The resulting precipitate was dried at 56°C and dissolved in 100 µl of deionized water.

Isolation of mitochondrial DNA from the total DNA pool using magnetic particles. Annealing of a biotinylated primer (pr79-B 16024-ttctttcatggggaagcagattt-16046 or pr80-B 16422-attgatttcacggaggatggtg-16401) at a concentration of 1 µm with a complementary fragment of the first hypervariable segment (HVS-1) of the mtDNA control region was performed in a buffer containing 75 mM Tris-HCl (pH 9.0), 20 mM (NH 4)2 SO 4, 0.01% Tween-20. The procedure included 10 minutes of denaturation at 95°C and 60 minutes of annealing at 55°C. The primer-matrix complex (mtDNA) was then re-deposited with PEG to remove primers that were not bound to the target DNA.

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Table 2.

Comparison of parameters of craniological features of remains from a paleosample with their limiting values in the Caucasian and Mongoloid anthropological communities

Individual*

Paul

Nasomalar angle

Zygomaxillary angle

Angle of nasal bone protrusion

Simotic index

Dacrial index

Anthropological type**

1

Wives.

145,1

134,7

25

46,4

50,8

Intermediate

2

Husband.

-

-

-

-

-

"

3

"

136,4

-

-

-

-

"

4

"

144,3

134,9

-

-

-

"

5

"

-

-

-

31,07

-

Mongoloid

6

Wives.

-

124,9

30

56,59

53,59

Caucasian

7

Husband.

143,9

-

-

-

-

Intermediate

8

"

147

128

-

-

-

"

9

"

-

122,2

-

39,22

-

"

10

"

-

-

30

37,86

46,43

Caucasian

11

"

138,3

127,9

20

64

66,67

"

Limit values (min-max) of trait parameters in a Caucasian anthropological community***

135 - 139

125 - 130

25 - 37

39,8 - 49,3

35 - 42

Limit values (min-max) of attribute parameters in the Mongoloid anthropological community***

145 - 148

137 - 142

7,5 - 19

19,5 - 29

51 - 73

-----

* Numbers according to Table 1.

** The anthropological type was diagnosed based on the published description of individuals and craniometric characteristics [Chikisheva, 2000a, 2003b].

*** The data are taken from tables of craniometric constants (Alekseev and Debets, 1964) without taking into account the rarely occurring very small and very large values of signs.

DNA. For this purpose, an equal volume of 20% PEG (MV 6000) in 2.5 M NaCl was added to the resulting solution; the mixture was incubated at 37°C for 15 min; precipitation was carried out by centrifugation at 12000 rpm for 15 min. The precipitate was washed three times with 700 µl of 80% ethanol, dried at 56°C, and dissolved in 35 µl of deionized water.

To reduce non-specific sorption, magnetic particles were incubated in five volumes of Denhard's solution before use. An equal volume of 20 x SSC and 10 µl of paramagnetic particles in 10 x SSC were added to the biotinylated primer - matrix (mtDNA) complex. Covalent binding of streptavidin immobilized on particles with biotinylated primers was carried out at room temperature and stirring constantly for 1 h. Separation of paramagnetic particles with primers immobilized on them in a complex with mtDNA from other components (genomic DNA pool, PCR inhibitor admixtures) It was performed using a magnetic tablet according to the manufacturer's protocol (MERCK for BioBeads Streptavidin).

Elution of the mtDNA fragments required for analysis was performed in 25 µl of deionized water by denaturation of the primer-mtDNA complex at 95°C for 5 minutes. The eluate was separated from the magnetic particles by means of a magnetic plate with visual control of their fixation on the walls of the test tube. Further, the DNA extracted in the solution was used as the initial template in the polymerase chain reaction.

Amplification of ancient DNA. It was performed by the nested-PCR method. For nested-PCR fragment HVS-1

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Two pairs of primers were used for the mtDNA control region: 1) external (fragment 398 bp) - pr79 16024-ttctttcatggggaagcagattt-16046 and pr80 16422-attgatttcacggaggatggtg-16401; 2) internal (fragment 336 bp) - pr_re79 16052-ccacccaagtattgactcaccc-16073 and pr_re80 16388-ctatctgaggggggtcatccat-16367. For amplification of the HVS-1 outer fragment of the mtDNA control region, a PCR mixture with a volume of 12.5 µl was included: 75 mM Tris-HCl (pH 9.0), 20 mM (NH4) 2 SO4, 0.01% Tween-20, 1 µm pr79 and pr80, 0.6 mM of each of the four dNTPs, 5 mM MgCl 2, 3 units. Tag polymerase and 3 µl of eluted DNA. Amplification cycles of 33 included denaturation for 1 min at 95°C, annealing for 1 min at 61°C, and synthesis for 1 min at 72°C.

Amplification of the HVS-1 internal fragment of the mtDNA control region was performed in 50 µl of a reaction mixture containing 75 mM Tris-HCl (pH 9.0), 20 mM (NH4) 2SO4, 0.01% Tween-20, 1 µm pr_re79 and pr_re80, 0.6 mM of each of the four dNTPs, 5 mM MgCl2, 5 units. Tag polymerase and 3 µl amplification of the HVS-1 outer fragment of the mtDNA control region. Amplification cycles of 25 included denaturation for 25 seconds at 95°C, annealing for 25 seconds at 55°C, and synthesis for 25 seconds at 72°C. The PCR product was detected by gel electrophoresis in 4% polyacrylamide gel, followed by staining with ethidium bromide. Next, an equal volume of 20% PEG (MV 6000) was added to the amplification in 2.5 M NaCl, incubated at 37°C for 15 min, precipitated by centrifugation at 12,000 rpm for 15 min, and washed three times with 700 µl of 80% ethanol by centrifugation (12,000 rpm) for 15 min. The resulting precipitate was dried at 56°C and dissolved in 25 µl of deionized water.

Primers pr_re79 and pr_re80 and the BigDye Terminator v 1.1 Cycle Sequencing Kit (Applied Biosystems) were used for the sequencing reaction. The DNA samples were analyzed using an ABI Prism 310 automatic sequencer (Applied Biosystems).

Results

Study of the structure of GWS I nucleotide sequences in the mtDNA control region of 11 ancient DNA samples (Table 1). 3) revealed the presence of nine haplotypes in them, which, according to the generally accepted classification [Wallace, 1995; Richards et al., 1998; Kivisild et al., 2002] belong to five haplogroups: three East Eurasian-A, C, D and two West Eurasian-U5 and H. For two DNA samples from Kaminnaya Cave (N 1) and Ak-Alakha-5, mound 4 (N 8) require additional studies to clarify the haplogroups.

Most of the analyzed mtDNA samples have Western Eurasian haplogroups (63.6%). Among them, haplogroup H prevails (57%), three haplotypes of which are CRS sequences, and one variant has a replacement at position 16304.

When comparing the data of the table. 2 and 3, it was found that only in the case of samples N 5, 10, and 11, we observe a direct correlation between the race-specific craniological features and the belonging of their mtDNA haplotypes to Western or Eastern Eurasian haplogroups. Buried in mound 5, Ak-Alakha-5 (N 5) with the mtDNA genotype of the East Eurasian haplogroup A is characterized by a typical Mongoloid morphotype; the other two (N 10, 11) with the CRS haplotype of the European haplogroup H had European features. Everything else-

Table 3.

Haplotype diversity of GWS I mtDNA of the ancient Altai population in different epochs

N n/a

Haplotype

Haplogroup

Number of repetitions

Burial

1

16224

H/K

4

Fireplace Cave

2

16192 - 16256 - 16270

U5

3

Karakol-1 (1982)

3

16304

K

2

Karakol (1985)

4

CRS

K

3

Bertek-56

5

16223 - 16242 - 16290 - 16319

A

3

Ak-Alakha-5, mound 5

6

16093 - 16129 - 16223 - 16298 - 16327

C

3

Ak-Alakha-1

7

16223 - 16239 - 16319 - 16362

D

3

Ak-Alakha-5, mound 3

8

16129 - 16182C-16183C-16189 - 16362

U5/J

2

The same, kurg. 4

9

16223 - 16311 - 16316 - 16362

D

3

Moynak-2

10

CRS

H

4

Kuturguntas-1

11

CRS

H

3

Ulandryk-1

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Human remains (with the exception of sample N 6) have intermediate anthropometric parameters between Mongoloids and Caucasoids, regardless of the race-specific nature of the haplogroups to which their mtDNA haplotypes belong (see Table 3). In the case of sample N 6 (female), despite the Europeoid morphotype in craniological parameters, the mtDNA haplotype belongs to haplogroup C, which is part of the haplogroup C group. It is part of the superhaplogroup M, variants of which are common among typical Mongoloids.

Analysis of the prevalence of nine haplotypes of ancient remains (see Table. 3) among the modern human populations inhabiting the Altai Mountains and adjacent territories, showed the presence of only four of them in existing databases. Haplotype 16192 - 16256 - 16270 (U5) was detected in Tuvans (0.3%) and Khanty (0.4%); with replacement in position 16304 (H) - in Tuvans (3.5%); CRS (H) - in Tuvans (2.1%), Khakas (1.1%), Altaians (1.1%), Kazakhs (4.1%), Khanty (1.6%) and Mansi (5,3%); 16093 - 16129 - 16223 - 16298 - 16327 (C) - in Tuvans (3.5%), Altaians (1.1%) and Mansi (2.6%).

The haplotype with a replacement at position 16224 is widely represented in populations of Western Europe, Croatia, and Russians. Haplotype 16223 - 16242 - 16290 - 16319 Haplogroup A (N 5) was not detected in populations of Central Asia and Finno-Ugric peoples, but it was detected in Selkups with a frequency of 3.3% (unpublished data by M. A. Gubina, L. P. Osipova). In Central Asia, the other two are common - 16223 - 16290 - 16319 - 16362 (Khakas - 1.1%) and 16242-16290-16293C-16319 (Tuvans-1.2%).

Two haplotypes of haplogroup D - 16223 - 16311 - 16316 - 16362 (N 9) and 16223 - 16239 - 16319 - 16362 (N 7) (see Table 3) - they were not found in modern populations, but their derivatives were registered in the population of Central and South-East Asia. Haplotype 16129-16182C-16183C-16189-16362 (N 8) is not available in databases. 16051 - 16092 - 16129 - 16182 - 16183 - 16189 - 16270 - 16362 and 16069 - 16182 - 16183 - 16189 - 16190 - 16362, belonging to the haplogroups U5b1 and J, respectively, were detected in the British (0.7%) [Helgason et al., 2001].

Discussion of the results

In historical and cultural studies, the physical and geographical Altai-Sayan mountain country (that part of it that lies on the territory of Russia) is sometimes arbitrarily considered as a special region of Southern Siberia, and sometimes Gorny Altai, Kuznetsk Alatau, Kuznetsk Basin and Salair Ridge are included in Western Siberia.

Western and Southern Siberia are located between the areas of the western and eastern circle of the main anthropological communities - Caucasoid and Mongoloid. In the anthropological literature, both regions are also often defined as transition zones, respectively north and south. Transitivity refers to the geographically determined and at certain stages of history realized possibility of contacts and movements of people characterized by contrasting physical features of the Caucasian and Mongoloid racial complexes. This fact was one of the main reasons for the dominance of the concept of mestizoization of Caucasians and Mongoloids in solving the issues of the origin of the anthropological identity of carriers of almost all archaeological cultures discovered in the territory of Gorny Altai. However, the paleoanthropological data obtained in recent years allow us to consider the morphological features of at least a part of this population as a result of the preservation of a certain protomorphy, which consists in an uncoordinated combination of the most important race-differentiating craniological features. The supposed protomorphic morphological complexes in the groups of the ancient population of Gorny Altai are characterized by a large face width, its uniform flatness at the level of the orbits and zygomatic bones, a high bridge of the nose, a slight protrusion of the nasal bones above the general line of the facial profile, an average width of the nasal opening, wide and low orbits. It is difficult to attribute such a complex to the circle of Mongoloid or Europoid anthropological types. It is extremely stable on the territory of Gorny Altai, revealing itself almost unchanged in the anthropological composition of groups of carriers of several cultures from the Neolithic Era to the Early Iron Age inclusive. We analyzed the paleoanthropological materials representing this population from two positions: based on the results of traditional craniometric analysis and the features of variation in the structure of mitochondrial DNA.

Craniometric data have been widely used in the study of the origin and formation of ancient and modern population groups since the mid-19th century, and their interpretative capabilities are well known to any specialist in the field of the genesis of cultures and peoples. A different situation is currently observed around the data of molecular paleogenetics, which makes the first attempts to systematize them in the ethno-cultural space of humanity.

At the beginning of the 1990s, several leading groups of researchers performed a global survey of the-

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2. Diagram of the phylogenetic tree of mitochondrial DNA haplogroups of the peoples of Eurasia (Wallace, 1995; Kivisild et al., 2002; Bermisheva et al., 2002).

Caucasian haplogroups - HV, H, V, J, T, U, K, I, W, X;

Mongoloid-A, B, E, F, Y, M, C, D, G, Z.

a comprehensive screening of mitochondrial genome variability in major human racial groups. As a result, it was possible to identify mutations that are key for determining race-specific mtDNA clusters or haplogroups [Wallace, 1995; Richards et al., 1998].

Data from the sequencing of hypervariable regions of GWS I and GWS II and restriction analysis of the entire mitochondrial genome showed that haplogroups HV, H, V, J, T, U, K, I, W, and X are most characteristic of Caucasian populations [Ibid], while haplogroups A, B, E, F, and Y are most characteristic of Mongoloid populations 2) (Wallace, 1995; Kivisild et al., 2002; Bermisheva et al., 2002).

Let us consider the degree of correlation between the data obtained from craniometric and paleogenetic analyses. The oldest samples of the sample belong to the Neolithic period. There are only two anthropological finds of this era in the Altai Mountains: the skeleton of a woman from the burial in the Kaminnaya Cave (Chikisheva, 2000b) and the skeleton of a man from the burial in the Nizhnetytkeskenskaya-1 cave (Kim and Chikisheva, 1995). Both can be attributed to the same craniological type, the leading features of which are mesobrahicrania and the average height of the cerebral part of the skull, a large width and flatness of the facial region, a relatively high bridge and an average protrusion of the nose above the line of the general profile of the face. Analogies to this combination of craniological features were found in the material from burials in the Krasnoyarsk-Kansk forest-steppe-the Bazaiha burial grounds (Alekseev, 1961) and the Long Lake burial ground (Gerasimova, 1964). Of course, the similarity does not extend to identity, but the described morphological complex is present in all individuals. The genesis of this complex was discussed on the basis of material from Bazaikha and was characterized as a weakened Mongoloid one: V. P. Alekseev (1961) saw the nature of this weakening in the Europoid admixture, and G. V. Shishkin (1961) described it as a weakened Mongoloid one. F. Debetz [1948] equally assumed its protomorphy.

As for the craniological materials of the Neolithic period from the Altai foothill zone, they reveal a complex anthropological composition. The Solontsy-5 series demonstrates a combination of features of the above-mentioned Neolithic population component of the Altai-Sayan Highlands and the Mongoloid component of the Baikal region populations of the same epoch (most closely related to the one presented in the Serov craniological series from the Lena River basin) [Kungurova and Chikisheva, 2002]. The Ust-Isha and Itkul series of burial grounds represent a variant of mestizo Europoids of the ancient hypermorphic Eastern Mediterranean type, whose origin is associated with the southern regions of Central Asia, and Paleosibirian Mongoloids from the Baikal region (Dremov, 1980). Thus, for the Neolithic population of the Altai foothills, a variant of genesis based on a mixture of Mongoloid and Europoid anthropological components is justified.

During the Early Bronze Age (late IV - early II millennia BC), two cultures existed on the territory of Gorny Altai: the Afanasiev culture (represented by a large number of archaeological sites and well - studied in anthropological terms) and the Bolshemyss culture (represented only by ceramic complexes in individual settlements, with a complete lack of any information about people). The Afanasyev culture area also includes the Minusinsk Basin, the northwestern part of the Mongolian Plateau, and the Turfan Basin in Xinjiang. According to the complex of anthropological features, the Afanasyevites are almost identical to the bearers of the synchronous yamnaya culture of the East European Plain. There are several hypotheses about the origin of the Afanasiev population. In the first anthropological studies, the morphological similarity of the Afanasyevites with the population of the far West (carriers of the Yamnaya culture) was explained by the migration of large masses of people from the main area of settlement of Caucasians to Southern Siberia and Central Asia (Debets, 1948; Alekseev, 1961). Later, a hypothesis was put forward about the possibility of existence in the eastern regions

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The steppe zone of Eurasia in the pre-Afanasyev period was an independent focus of formation of one of the local variants of the Caucasian race, which was associated with the genesis of carriers of the Afanasyev culture [Alekseev, 1989, pp. 350-355].

When planning our study, we assumed the study of mtDNA in samples belonging to the Neolithic population of the foothill zone of Altai and to the Afanasyevites of Gorny Altai. Unfortunately, at the moment, it has not been possible to isolate the DNA and genotype these samples in full. We continue to work with them.

During the advanced Bronze Age period between the 18th - 17th and 10th centuries BC, Gorny Altai was home to cultures whose originality and vividness are reflected in the magnificent examples of primitive art - polychrome drawings made on stone slabs of sarcaphagi and on rocks. Archaeologists have not yet established exactly whether all burial and settlement complexes of this chronological period belong to the same or different cultures. The Karakol culture is clearly distinguished (Molodin, 1991, 2002, 2006), whose monuments (Ozernoye, Karakol, Karakol-1, Besh-Ozek, Ust-Kuyum, Kara-Koba-1) are located in the central part of Gorny Altai. They have many-sided parallels with the monuments of the Okunev culture of the Minusinsk basin, as well as a certain similarity with the synchronous monuments of the Krotovo culture of the forest-steppe interfluve of the Ob and Irtysh rivers.

In the paleoanthropological material of the Karakol culture, we found analogies to the craniological type of Neolithic skulls, which gave us reason to assume a genetic connection between both cultural formations. These materials were obtained from two burial grounds excavated by different researchers (A. P. Pogozheva [1984] and V. D. Kubarev [1988]) in the central and north-western regions of Altai (Ongudai and Ust-Kansk by administrative division), but with the same name - Karakol. They were studied and published (Chikisheva 2000b, 2003b). Skulls from both sites have a great morphological similarity. Their characteristic anthropological complex brings them closer to both the Okunevs from the Minusinsk basin, and earlier Neolithic ones from the caves of Gorny Altai, as well as from Bazaikha and Long Lake on the Middle Yenisei.

Based on the materials of two other monuments of approximately the same chronological period discovered in the south of Gorny Altai (the cult complex Kucherla-1 and Bertek-56 mound), V. I. Molodin admits the possibility of the existence of a special archaeological culture, suggesting that the ornamentation of its ceramics is based on the Afanasiev tradition [2002]. However, the study of skeletal remains of a 30 - 3.5-year-old man and a 6.5 - 7-year-old child from Bertek-56 mound (Chikisheva, 2003b) showed that the Caucasian component found in these subjects is not related to the Afanasyev anthropological environment. The morphological specifics of the buried may have been formed as a result of mestizo anthropological type, typical of the carriers of the Karakol culture (this may be evidenced by the structure of the frontal bone, horizontal flatness of the facial skeleton, features of the structure of the teeth), and the Caucasian variant, characterized by general massiveness, a wide and very high face. Among the Caucasian populations, the craniological variant with such features is known from the materials of the Early Tulkhar burial ground, representing pastoral tribes of the Beshkent Valley (Tajikistan) II millennium BC. [Kiyatkina, 1976].

The Late Bronze Age (X-VIII centuries BC) is represented in Gorny Altai by very insignificant materials, among which there are no anthropological finds. These are products made of bronze, ceramics, left by the population of the pre-Altai and West Siberian forest-steppes. No local cultural formations have been identified yet. It is believed that the morphological complex of the Late Bronze Age corresponds to that of the carriers of the Karasuk culture of the Minusinsk basin and the cultures of landless burials in Tuva and Mongolia.

In the Neolithic-Bronze Age cultural - chronological interval, molecular genetic analysis of mtDNA was performed for samples N 1-4 (see Table 3). Haplotype of sample N 1 (female, Neolithic burial in Kaminnaya Cave) contains a substitution at position 16224, which is characteristic of variants of the West Eurasian haplogroups H and K*. Haplotypes N 2, 3 (burials of the Karakol culture in the middle of the second millennium BC) and 4 (burial Bertek-56, dating from the beginning of the second millennium BC) also belong to the West Eurasian haplogroups U5 and H. Of interest is the fact that, according to some studies, haplogroup H originated in the Altai at the turn of the Middle and Upper Paleolithic (approximately 50 thousand years AGO) and then spread widely in Europe after the Ice Age [Loogvali et al., 2004]. The haplotype with a single substitution at position 16304 (sample N 3) is represented all over the world, although in Central Asian populations this variant was detected only in Tuvans (3.5%). Variants of haplogroup U5, to which the haplotype of sample N 2 belongs, are found in many Northern European countries.

* The exact definition of the haplogroup for the analyzed mtDNA variant is under further study.

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Most of them are found in the Sami population (Lahermo et al., 1999; Torroni et al., 1993; Wallace et al., 1999). This haplogroup is part of the U supercluster, which originated on the Eurasian continent and underwent significant phylogenetic differentiation and geographical dispersion. Thus, haplogroup U6 is found on the African continent, U7 is typical for populations of Jordan, Kuwait, Iran, and Saudi Arabia, and U1-U5 are found in Western Europe [Richards et al., 1996, 1998]. It is important to note that the oldest carriers of the analyzed mtDNA samples (N 1-4) belong to the West Eurasian haplo groups H and U5 (see Table. 3), have intermediate craniological parameters (see Table 2).

The Early Iron Age (early nomadic, or Scythian time, VIII-II centuries BC) has been studied most fully in Gorny Altai in comparison with other archaeological periods. There are grounds for cultural and chronological differentiation of monuments of the VIII (or late IX) - early VI century BC and late VI - early II century BC. The former are distinguished by researchers as a special culture [Mogilnikov, 1983; Stepanova, 1986; Kiryushin and Tishkin, 1997]. A term has not yet been chosen for it, and the question of its origin has not yet been resolved. It is assumed that the population of the Early Scythian period of Gorny Altai developed on the same basis as the carriers of the Aldybel culture of Tuva and some groups of Saks of Kazakhstan, and the common origins of the formation of their funeral rite (as one of the leading elements of the cultural complex) are located in Northwestern Mongolia (Kiryushin and Tishkin, 1997; Savinov, 1994). Paleoanthropological material from Early Scythian mounds is relatively sparse and extremely mixed: Mongoloid, Europoid, and also Mongoloid-Europoid complexes of features are distinguished (Tur, 1997).

In relation to the monuments of the next chronological stage (the end of the VI - beginning of the II century BC), the concept of "Pazyryk culture"is widespread in the archaeological literature. In the last three decades, two groups of monuments have also been discovered, similar to Pazyryk, but differing in some features of the funeral rite. Based on one of them, the Kara-Koba culture was identified (Mogilnikov, 1983; Surazakov, 1983). The other group is called Chumysh-Ishinskaya. It is not numerous, but, nevertheless, it is considered as a special culture (Surazakov, 1988). A number of researchers, however, believe that it is illegal to separate monuments of the late VI-early II centuries BC from the Pazyryk culture (Polos'mak, 1994; Shulga, 1986).

The Pazyryk and Kara-Koba cultures are well studied in anthropological terms [Chikisheva, 1994, 1996, 20006, 2002, 2003a]. The autochthonous morphological component, presumably dating back to the Neolithic population of Gorny Altai and the carriers of the Okunevsky circle (Karakol) cultures, was found to be predominant in the composition of the Pazyryk tribes. A comparative analysis of the morphological complex of early nomads of Gorny Altai with data published in the literature showed that this complex dominates a number of ethno-cultural groups of the Scythian period in the southern regions of Eurasia and determines the anthropological similarity of the Pazyryk population of Southern Siberia with part of the tribal associations of the Sako-Usun ethno-cultural community of Central and Central Asia. The Pazyryk people are most closely related to the so-called Early Scythian population that preceded them in the Altai and tend to belong to the Saka and Usun tribes of the Dzungarian Alatau and Tien Shan. The anthropological composition of the Kara-Koba-type burial mounds probably contains a component that connects this group of nomads of the Gorny Altai with the tribes of the forest-steppe Ob-Irtysh interfluve.

Paleogenetic analysis of anthropological finds related to the Pazyryk culture was previously carried out on representative material from mounds in the Ak-Alakha River valley on the Ukok plateau. In this paper, we continued this study with a sample of six samples: N 5-8 from the Ak-Alakha burial ground, N 9 from the Moynak-2 mound, N 10 from the Kuturguntas-1 mound (see Table 3). In addition, a sample from the Ulandryk River valley mound was analyzed (N 11) (see Table 3). In the Pazyryk samples, we observe a higher level of mtDNA genetic diversity than in those belonging to previous cultures (see Table 3). In addition to the West Eurasian haplo groups, we identified three East Eurasian haplo groups-A, D, and C (in samples N 5-7, 9).

It is noteworthy that haplotypes of haplogroup A were not previously found in the anthropological materials of the Pazyryk culture (Molodin et al., 2000). Haplotype 16223 - 16242 - 16290 - 16319 (sample N 5) can undoubtedly be attributed to one of its ancestral variants currently found in Khakass populations (haplotype 16223 - 16290 - 16293 - 16319 - 1,1%) and Tuvans (haplotype 16242-16290 - 16293C-16319-1.2%). All of them are derived from the mtDNA variant with three substitutions in GWS I of the control region (16223 - 16290 - 16319), the highest frequency of which is observed in ethnic groups of Central Asia and China (Wallace et al., 1999). This haplotype is found in the Ainu (3.9%), Evens (3.1%), Chinese (7.7%), Koreans (3.1%), Mongols (2.9%), Uyghurs (7.3%), Kazakhs (9.3%), Kyrgyz (6.3%) and Middle Eastern peoples (2.7%). [Comas et al., 1998; Yao et al., 2002; Derenko,

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Shields, 1997; Derenko et al., 2000]. Intrapopulation pairwise nucleotide differences between similar mtDNA sequences in the populations of East and Central Asia are 2.60 and 3.21, respectively, which indicates to a greater extent the Central Asian origin of the ancestral haplotype of haplogroup A (Molodin et al., 2004). According to some estimates, it originated 48 thousand years ago (Ivanova, 1993). It is possible that the appearance of carriers of haplogroup A variants among the Pazyryk people is associated with the migration (before the IV-III centuries BC) of groups (or groups) of early nomads to the territory of Gorny Altai from the southeast or south. This assumption is supported by the fact that Ak-Alakha-5 buried in mound 5 of the burial ground, whose mtDNA belongs to the East Eurasian haplogroup A (sample N 5), has a well-defined Mongoloid morphotype (see Table 2). Haplotypes of this haplogroup are also found in populations of the Volga-Ural region, but with a low frequency (Bermisheva et al., 2002).

Two Pazyryk samples (N 7 and 9) with intermediate craniological features (see Table. 2) are carriers of two different variants of the Mongoloid haplogroup D, whose haplotypes are relatively widespread in modern indigenous populations of the Altai-Sayan Highlands and Northern Asia (Richards et al., 1996; Kolman, Sambughin, and Bermingham, 1996; Derenko et al., 2001; Ivanova, 1993). For example, haplogroup D is represented in 15% of Tuvans and 7.2% of Khakas (Damba et al., 2003). Haplotype 16223 - 16311 - 16316 - 16362 Sample No. 9 is missing from the mtDNA database, but variants without substitutions at positions 16316 or 16311 were found in Kazakhs (1.4%) and Tuvans (0.3%). Option 16223 - 16239 - 16319 - 16362 Sample N 7 has an additional replacement at position 16239 in comparison with the haplotype common among Tuvans (4.1%), Altaians (3.7%) and Kazakhs (3.4%). 16223 - 16319 - 16362 with three substitutions. Based on the obtained data, it can be stated that intermediate anthropometric features are probably characteristic of carriers of haplotypes of both Caucasian and Mongoloid haplogroups (see Tables 2, 3).

The most impressive result was obtained for sample N 6 with Caucasian craniological features, which was found to be a carrier of the haplotype of the East Eurasian haplogroup C (see Table 3). It should be noted that a similar discrepancy between anthropometric data and genetic parameters was already observed earlier in Pazyryk people buried in the Verkh-Kaldzhin burial ground [Voevoda et al., 1998], with a similar mtDNA haplotype. Variants of haplogroup C are quite widely represented in modern populations of the region. Their frequency reaches 35 and 40%, respectively, among Khakas and Tuvans [Damba et al., 2003], and 13.8% among Kazakhs (unpublished data). In some typical Mongoloid populations (Evenks, Yukagirs) , haplotypes of haplogroup C are more than 50% [Schurr et al., 1999; Starikovskaya et al., 1998]. In the population of Southern and Western Europe, this haplogroup is either absent, or the frequency of occurrence is no more than 1%, which is typical for Finno-Ugric peoples of the Volga-Ural region (from 0 for Komi-Zyryans to 3% for Udmurts), and for Balto-Finn ethnic groups (no more than 2%) [Bermisheva et al. et al., 2002; Meinila, Finilla, and Majamaa, 2001; Villems et al., 1998]. Identical to the haplotype 16093 - 16129 - 16223 - 16298 - 16327 (sample N 6) mtDNA sequences are also found in modern populations of Gorny Altai, but with a low frequency: Tuvans - 3.5%, Altaians-1.1% (Damba et al., 2003).

We have yet to determine to which haplogroup the West Eurasian haplotype 16129-16182C - 16183C - 16189 - 16362 found in sample N 8 belongs (see Table 3). It is not included in the mtDNA database, but there is a variant with an additional substitution at position 16270 in one Englishman [Helgason et al., 2001]. The ancient carrier of the haplotype (sample N 8) has intermediate craniological features, while the other two Pazyrykians (N 10 and 11) with West Eurasian CRS haplotypes are characterized by typical Europoid anthropometric parameters (see Tables 2, 3).

Conclusion

The study of the dynamics of the anthropological composition of the autochthonous population of Gorny Altai leads to the conclusion that the component that occupies an intermediate position in terms of craniological features differentiating the first-order races (Mongoloid and Europoid) prevails. According to the results of molecular genetic analysis of mtDNA samples from the Neolithic and Bronze Ages, only Western Eurasian haplotypes are observed. The uniformity of the structure of the mtDNA gene pool over two to three millennia indicates that there is no significant drift of genetic material in the population. The mixing of western and eastern gene flows in the territory of Gorny Altai by the mtDNA "chronicle" was recorded only in the Iron Age in the carriers of the Pazyryk culture. This is also confirmed by a set of data on ethno-cultural genesis (Molodin, 2003).

An integral assessment of the results obtained allows us to conclude that, apparently, there may not be a strict correlation between the race-specific morphological characteristics of people and the presence of-

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their mtDNA haplotypes belong to the Mongoloid or Caucasian haplogroup. Craniological features, like any other polygenic features, are formed with the participation of many loci of the nuclear genome encoding structural proteins of osteogenesis, hormones, growth cytokines and their receptors, transcription factors, and other regulatory proteins. The genetic content of the optional material inherited from the mother, such as mitochondrial DNA, may only indirectly affect the processes of morphogenesis. The mechanisms of coordinated functioning and evolution of the nuclear and mitochondrial genomes of eukaryotes are currently unclear.

Nevertheless, our results demonstrate the success of parallel craniological and paleogenetic analyses in interpreting cases of intermediate craniometric complexes. According to anthropological criteria, such complexes can be equally attributed to the Europoid-Mongoloid mestizos, and to a specific anthropological community that separated from the Mongoloid or Europoid racial trunk even before the period when pronounced anthropological Europoid and Mongoloid complexes were formed. The structure of the main mtDNA clusters (haplogroups) is clearly associated with the phylogenesis of populations, and their subclusters are geographically vectorized. This quality allows us to find only Western Eurasian haplogroups in the mitochondrial genome of the autochthonous population of Gorny Altai and conclude that it belongs to the gene pool of Caucasian populations. Carriers of the Afanasiev culture did not contribute to the anthropological composition of this population, which does not exclude a certain influence on the external side of life, culture, economy, and beliefs. The absence of biological correlations between the mitochondrial genome and craniological features established by us requires verification of the conclusions obtained on the basis of skeletal morphology by genetic level criteria.

It is highly probable that the Caucasoid component found in the anthropological composition of the Gorny Altai tribes of the second millennium BC indicates their connections with the pastoral population of Near and Central Asia. The existence of such relationships is confirmed by archaeological materials (Molodin et al., 2004; Polos'mak et al., 2006), as well as data from paleogenetic analysis.

It is interesting that the ancient population of the Altai-Sayan Highlands is mainly represented by the Western Eurasian haplotypes of haplogroups H and U5, which are widespread in Western Siberia and Europe. Whereas the modern populations inhabiting this region are dominated by East Eurasian haplogroups (Gubina et al., 2006).

Taking into account the results of anthropological analysis, which extracts genetic information indirectly from an incomparably larger array of ancient samples, it can be concluded that the southern Eurasian anthropological formation identified by us (Chikisheva, 2000a, b; 2003b) is a relic of an ancient protopopulation. These data do not contradict the hypothesis expressed earlier about the formation of the anthropological composition of the modern population of Eurasia on the basis of the original protopopulation that supposedly inhabited the territory of Central and Central Asia [Voevoda et al., 2003].

List of literature

Alekseev V. P. O smeshannom proiskhozhdenii uralskoy rasa [On the mixed origin of the Ural race]. Sverdlovsk: Ural State University Press, 1961, pp. 117-1120.

Alekseev V. P. Istoricheskaya antropologiya i etnogenez [Historical anthropology and ethnogenesis]. Moscow: Nauka Publ., 1989, 445 p.

Alekseev V. P., Debets G. F. Craniometry: Methodology of anthropological research, Moscow: Nauka Publ., 1964, 128 p.

Bermisheva M., Tambets K., Willems R., Khusnutdinova E. Diversity of mitochondrial DNA haplogroups in the peoples of the Volga-Ural region of Russia. - 2002. - Vol. 36, N 6. - pp. 990-1001.

Voevoda M. I., Sitnikova V. V., Chikisheva T. A., Romashchenko A. G., Polosmak P. V., Molodin V. I., Derevyanko A. P., Shumny V. K. Molecular genetic analysis of mitochondrial DNA of representatives of the Pazyryk culture of Gorny Altai (IV-II centuries BC). Dokl. Acad. of Sciences. - 1998. - T. 358, N 4. - pp. 564-566.

Voevoda M. I., Shulgina E. O., Nefedova M. V., Kulikov I. V., Damba L. D., Gubina M. A., Kobzev V. F., Romashchenko A. G. Paleogenetic studies of Early Iron Age culture carriers in Gorny Altai // The population of Gorny Altai in the Early Iron Age as an ethno-cultural phenomenon: origin, genesis, historical destinies (according to archeology, anthropology, and genetics). Novosibirsk: Publishing House of the Siberian Branch of the Russian Academy of Sciences, 2003, pp. 121-147.

Gerasimova M. M. Neolithic burials of the burial ground near the Long Lake. 1964, Issue 18, pp. 135-143.

Gubina, M. A., Damba, L. D., Romashchenko, A. G., Kobzev, V. F., Vilems, R., and Voevoda, M. I., Features of mtDNA haplotype diversity in some populations of Siberia, the Far East, and Central Asia, Sovremennye problemy arkheologii Rossii: Tez. conf. - 2006. - Vol. 2. - pp. 260-263.

Damba L. D., Gub'ina M. A., Konchuk Ch. D., Kobzev V. F., Romashchenko A. G., Voevoda M. P. Features of representation of Mongoloid and Europoid haplogroups of MITO-

page 140

Chondrial DNA in two populations of indigenous residents of the South of Siberia / / Gene Pool of the population of Siberia. Novosibirsk: Izd-vo IAET SB RAS, 2003, pp. 19-24.

Debets G. F. Paleoanthropologiya SSSR. - M.; L.: Izd-vo AN SSSR, 1948. - 392 p. - (TEE; vol. 4).

Derenko, M. V., Denisova, G. A., Malyarchuk, B. A., Dambueva, I. K., Luzina, F. A., Lotosh, E. A., Djorzhu, Ch. M., Karamchakova, O. N., Solovenchuk, L. L., and Zakharov, M. A., Structure of gene pools of ethnic groups in the Altai-Sayan Highlands based on data on mitochondrial DNA polymorphism, Genetika. 2001, vol. 37, No. 10, pp. 1402-1410.

Derenko M. V., Shields D. J. Diversity of mitochondrial DNA nucleotide sequences in three groups of the indigenous population of Northern Asia. - 1997. - Vol. 31, N 5. - pp. 784-789.

Dremov V. A. Anthropological materials from the Ust-Isha and Itkul burial grounds: To the question of the origin of the Neolithic population of the Upper Ob region / / Paleoanthropology of Siberia, Moscow: Nauka, 1980, pp. 19-46.

Ivanova A.V. Polymorphism of mitochondrial DNA in populations of indigenous inhabitants of Chukotka. Institute of Cytology and Genetics SB RAS, Novosibirsk, 1993, 99 p.

Kim A. R., Chikisheva T. A. Pogrebenie iz Nizhnetytkeskenskoi caves I - pervaya doafanas'evskaya mogila na territorii Gornogo Altay [Burial from the Nizhnetytkeskenskaya Cave I-the first pre-Afanas'evskaya grave on the territory of Gorny Altai]. Barnaul: Alt. State University Publ., 1995, Appendix 2, pp. 95-117.

Kiryushin Yu. F., Tishkin A. A. Scythian epoch of the Altai Mountains. Barnaul: Alt. State University Publ., 1997, Part 1: Culture of the population in the Early Scythian period. - 232 p.

Kiyatkina T. P. Materials on paleoanthropology of Tajikistan. Dushanbe: Donish Publ., 1976, 186 p.

Kubarev V. D. Ancient paintings of Karakol. Novosibirsk: Nauka Publ., 1988, 171 p. (in Russian)

Kungurova N. Yu., Chikisheva T. A. Results of the study of the Solontsy-5 Neolithic burial ground on the Biya River // Problems of archeology, ethnography, and anthropology of Siberia and adjacent territories. Novosibirsk: Izd-vo IAET SB RAS, 2002, pp. 121-129.

Mogilnikov V. A. Kurgany Kara-Koba II [Kurgans of Kara-Koba II] / / Archeologicheskie issledovaniya v Gornom Altae v 1980-1982 godakh. Gorno-Altaisk: GANIIIYAL Publ., 1983, pp. 52-89.

Molodin V. I. Developed bronze Mountain Altai // problems of late bronze age and transition to the iron age in the Urals and adjacent territories. Ufa: Publishing House of the Bashkir State University, 1991, pp. 4-25.

Molodin V. I. Gorny Altai in the Bronze Age / / History of the Altai Republic. Gorno-Altaisk: Publishing House of the S. S. Surazakov Institute of Altaistics, 2002, vol. 1, pp. 97-142.

Molodin V. I. Etnogenez, etnicheskaya istoriya i istoricheskie sudby nositelei pazyrykskoi kul'tury Gornogo Altay [Ethnogenesis, ethnic history and historical destinies of the Pazyryk culture bearers of Gorny Altai]. Naselenie Gorny Altay v epokhu rannego zheleznogo veka kak etnokul'turnyi fenomen: proiskhozhdenie, genezis, istoricheskie sudby (po dannym arkheologii, antropologii, genetiki). Novosibirsk: Publishing House of the Siberian Branch of the Russian Academy of Sciences, 2003, pp. 148-178.

Molodin V. I. Karakol culture // Okunevsky collection 2: Culture and its environment. Saint Petersburg: Elexis-Print Publ., 2006, pp. 271-282.

Molodin V. I., Polosmak N. V., Novikov A.V., Bogdanov E. S., Slyusarenko I. Yu., Cheremisin D. V. Archaeological sites of the Ukok plateau (Gorny Altai). Novosibirsk, IAET SB RAS Publ., 2004, 256 p. (in Russian)

Molodin V. I., Romashchenko A. G., Voevoda M. I., Chikisheva T. A. Multidisciplinary analysis of the Pazyryk culture carriers (archeology, anthropology, genetics) / / Scythians and Sarmatians in the VII-III centuries BC: paleoecology, Anthropology and Archeology. 2000. pp. 59-66.

Pogozheva A. I. Kurgany epokhi bronzy na zapad Gornogo Altay [Mounds of the Bronze Age in the West of the Altai Mountains]. AO 1982. - Moscow: Nauka, 1984. - pp. 225-226.

Polosmak N. V. Pazyryk culture // Ancient cultures of the Bertek Valley. Novosibirsk: Nauka Publ., 1994, pp. 137-144.

Polosmak I. V., Kundo L. P., Balakina G. G., Mamatyuk V. I., Vasiliev V. G., Karpova E. V., Malakhov V. V., Vlasov A. A., Kraevskaya I. L., Dovlitova L. S., Korolyuk E. A., Tsareva E. G. Textiles from frozen graves of the Gorny Altai of the IV-III centuries BC (experience in interdisciplinary research). Novosibirsk: Izd-vo IAET SB RAS, 2006, 267 p. (in Russian)

Savinov D. G. Tuva ranneskifskogo vremeni na "perekrestke" kul'turnykh traditsii (aldy-belskaya kul'tura) [Early Scythian Tuva at the "crossroads" of cultural traditions (Aldy-belskaya kul'tura)]. St. Petersburg: Publishing House of St. Petersburg State University, 1994, pp. 76-92.

Stepanova N. F. Kuyumskiy tip pamyatnikov VIII - VI vv. B.C. [The Kuyum type of monuments of the VIII-VI centuries BC]. Barnaul: Alt. State University, 1986, pp. 79-81.

Surazakov A. S. Kurgans of the Early Iron Age in the burial ground of Kyzyk-Telan-1: (On the issue of distinguishing the Kara-Koba culture) // Archaeological research in Gorny Altai in 1980-1982. Gorno-Altaisk: GANIIIYAL Publ., 1983, pp. 42-52.

Surazakov A. S. Gorny Altai and its northern foothills in the Early Iron Age: Problems of chronology and cultural differentiation. Gorno-Altaisk: GANIIIYAL Publ., 1988, 206 p.

Tur S. S. Kraniologicheskie materialy iz ranneskifskikh mogilnikov Altay [Craniological materials from the Early Scythian burial grounds of Altai]. Barnaul: Alt. State University, 1997. Part 1: Culture of the population in the Early Scythian period. - Adj. - p. 136-147.

Chikisheva T. A. Characteristics of paleoanthropological material of Bertek valley monuments // Ancient cultures of the Bertek Valley. Novosibirsk: Nauka Publ., 1994, pp. 167-175.

Chikisheva T. A. On the formation of the anthropological composition of the population of the Pazyryk culture of Gorny Altai // The latest archaeological and ethnographic discoveries in Siberia: Materials of the IV Annual Final Session of the Institute of Archeology and Ethnography SB RAS. December 1996-Novosibirsk: Publishing House of IAET SB RAS, 1996. - pp. 249-252.

Chikisheva T. A. Voprosy proiskhozhdeniya kochevnikov Gornogo Altaya epokhi rannego zheleznogo po dannym antropologii [Questions of the origin of nomads of the Gorny Altai of the Early Iron Age according to anthropology]. - 2000a. - N4 (4). - p. 107-121.

Chikisheva T. A. To the question about the formation of the anthropological composition of the population of Western Siberia in the era

page 141

Late Bronze Age (interpretation of paleoanthropological material from the Old Garden burial ground in Central Baraba) / / Archeology, Ethnography and Anthropology of Eurasia. - 20006. - N 2 (2). - p. 131-147.

Chikisheva T. A. Novye dannye ob antropologicheskom sostav naseleniya Altay v epokhi neolita - bronzy [New data on the anthropological composition of the Altai population in the Neolithic-Bronze Age]. - 2000b. - N 1 (1). - p. 139-148.

Chikisheva T. A. Osobennosti zubnoy sistemy rannykh kochevnikov Gornogo Altay [Features of the dental system of early nomads of the Altai Mountains]. - 2002. - N 1 (9). - p. 149-159.

Chikisheva T. A. K voprosu ob antropologicheskom sostav naseleniya yuzhnykh rayonov Zapadnoy Sibiri v epokhi neolita i bronzy [On the question of the anthropological composition of the population of the southern regions of Western Siberia in the Neolithic and Bronze Ages].

Population of Gorny Altai in the era of early iron according to anthropology // Population of Gorny Altai in the era of early Iron Age as an ethno-cultural phenomenon: origin, genesis, historical destinies (according to archeology, anthropology, genetics). Novosibirsk: Publishing House of the Siberian Branch of the Russian Academy of Sciences, 2003, pp. 63-120.

Shulga P. I. To the question about the culture of the herders of the Altai mountains in VI - II centuries BC. / / Scythian era of Altai. Barnaul: Alt. State University, 1986, pp. 20-23.

Comas D., Calafell E., Mateu E., Pyrez Lezaun A., Bosch E., Martynez Arias R., Clarimon J., Facchini E., Fiori G., Luiselli D., Pettener D., Bertranpetit J. Trading genes along the Silk road: mtDNA sequences and the origin of Central Asian populations // Am. J. Hum. Genet. - 1998. - Vol. 63. - P. 1824 - 1838.

Derenko M. V., Malyarchuk B. A., Dambueva I. K., Shaikhaev C. O., Dorzhu C. M., Nimaev D. D., Zakharov L. A. Mitochondrial DNA variation in two South Siberian aboriginal populations: Implications for the genetic history of Notth Asia // Hum. Biol. - 2000. - Vol. 72. - P. 945 - 973.

Helgason A., Hickey E., Goodacre S., Bosnes V., Stefansson K., Ward R., Sykes B. mtDNA and the Islands of the North Atlantic: Estimating the Proportions of Norse and Gaelic Ancestry // Am. J. Hum. Genet. - 2001. - Vol. 68 (3). - P. 723 - 737.

Kolman C., Sambuughin N., Bermingham E. Mitochondrial DNA analysis of Mongolian populations and implications for the origin of New World founders // Genetics. - 1996. - Vol. 142. - P. 1321 - 1334.

Kivisild T., Tolk H-V., Parik J., Wang Y., Papiha S. S., Bandelt H. J., Villems R. The emerging limbs and twigs the East Asian mtDNA tree // Mol. Biol. Evol. - 2002. - Vol. 19 (10). - P. 1737 - 1751.

Lahermo P., Sajantila A., Sistonen P., Lukka M., Aula P., Peltonen L., Savontaus M. L. The genetic relationship between the Finns and Finnish Saami: analysis of nuclear DNA and mtDNA // Am. J. Hum. Genet. - 1999. - Vol. 64. - P. 232 - 249.

Loogvali E. L., Roostalu U., Malyarchuk B. A., Derenko M. V., Kivisild T., Metspalu E., Tambets K., Reidla M., Tolk H. -V., Parik J., Pennarun E., Laos S., Lunkina A., Golubenko M., Barac L., Pericic M., Balanovsky O. P., Gusar V., Khusnutdinova E. K., Stepanov V., Puzyrev V., Rudan P., Balanovska E. V., Grechanina E., Richard C., Moisan J. P., Cheventre A., Anagnou N. P., Pappa K. I., Michalodimitrakis E. N., Claustres M., Golge M., Mikerezi I., Usanga E., Villems R. Disuniting uniformity: a pied cladistic canvas of mtDNA haplogroup H in Eurasia // Mol. Biol. Evol. - 2004. - Vol. 21. - P. 2012 - 2021.

Meinila M., Finilla S., Majamaa K. Evidence for mtDNA admixture between the Finns and the Saami // Hum. Hered. - 2001. - Vol. 52. - P. 160 - 170.

Richards M., Corte-Real H., Forster P., Macaulay V., Wilkinson-Herbots H., Demaine A., Papiha S., Hedges R., Bandelt H., Sykes B. Paoleolithic and neolithic lineages in the European mitochondrial gene pool // Am. J. Human. Genet. - 1996. - Vol. 59. - P. 185 - 203.

Richards M. B., Macaulay V. A., Bandelt H. -J., Sykes B. C. Phylogeography of mitochondrial DNA in western Europe // Am. Hum. Genet. - 1998. - Vol. 62. - P. 241 - 260.

Schurr T. G., Sukernik R. I., Starikovskaya Y. B., Wallace D. C. Mitochondrial DNA variation in Koryaks and Itel'men: population replacement in the Ochotsk Sea-Bering Sea region during the Neolithic // Am. J. Phys. Anthropol. - 1999. - Vol. 108. - P. 1 - 39.

Starikovskaya Y., Sukernik R., Schurr T., Kogelnik A. M., Wallace D. C, Torroni A. MtDNA diversity in Chukchi and Siberian Eskimos: implications for the genetic history of Ancient Beringia and the peopling of the New Word // Am. J. Hum. Genet. - 1998. - Vol. 63. - P. 1473 - 1491.

Torroni A., Sukernik R. I., Schurr T. G., Starikovskaya Y. B., Cabell M. E., Crawford M. H., Comuzzie A. G., Wallace D. C. Mitochondrial DNA variation of aboriginal Siberians reveals distinct genetic affinities with Native Americans // Am. J. Hum. Genet. - 1993. - Vol. 53. - P. 591 - 608.

Villems R., Adojaan M., Kivisild T., Metspalu E., Parik J., Pielberg G., Rootsi S., Tambets K., Tolk H. -V. Reconstruction of maternal lineages of Finno-Ugric speaking people and some remarks on their paternal inheritance // The roots of peoples and languages of Northern Eurasia I / Eds. K. Wiik, K. Julku. - Turku: Societes Historiae Fenno-Ugricae, 1998. - P. 180 - 200.

Wallace D. C. Mitochondrial DNA variation in human evolution, degenerative diseases, and aging // Am. J. Hum. Genet. - 1995. - Vol. 57. - P. 201 - 223.

Wallace D. C., Brown M. D., Lott M. T. Mitochondrial DNA variation in human evolution and disease // Gene. - 1999. - Vol. 238. - P. 211 - 230.

Yao Y-G., Kong Q-P., Bandelt H-J., Kivisild T., Zhang V. P. Phylogeographic differentiation of mitochondrial DNA in Han Chinese // Am. J. Hum. Genet. - 2002. - Vol. 70. - P. 635 - 651.

The article was submitted to the Editorial Board on 16.04.07.

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