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Migrations in the south of the West Siberian Plain during the Bronze Age (4th-2nd millennium BC): Archaeological, paleogenetic and anthropological data

Authors:
HUMAN MIGRATIONS IN THE SOUTHERN REGION OF THE WEST SIBERIAN PLAIN DURING THE BRONZE AGE 93
Vyacheslav I. Molodina, Alexander S. Pilipenkob, *, Aida G Romaschenkob,
Anton A. Zhuravlevb, Rostislav O. Trapezovb, Tatiana A. Chikishevaa,
Dmitriy V. Pozdnyakova
Human migrations in the southern region of the West Siberian Plain during
the Bronze Age: Archaeological, palaeogenetic and anthropological data
Introduction
Western Siberia is a vast area of approximately 3 million square kilometers in North Asia. It ranges from
the Ural Mountains in the east to the Yenisei River in the west, and from the Arctic Ocean in the north to
the mountainous regions of Central Asia in the south. The southern part of the region comprises zones
of steppe and forest steppe and has been inhabited by anatomically modern humans since the Stone Age
(Upper Palaeolithic period).
Our work is devoted to the analysis of human migration processes that occurred during the Bronze
Age (4th–early 1st millennium BC) in the forest steppe zone between the Ob and Irtysh rivers (about
800 km from west to east). This area, known as Baraba forest steppe, stretches over 200 km from the
taiga zone in the north to the steppes in the south.
Several factors make the ancient Baraba populations interesting objects of study. Firstly, intensive
archaeological research has been conducted in the Baraba region during the last 30 years. As a result,
a large set of data about all ethno-cultural groups that occupied this territory, from about 14 thousand
years ago to the Late Middle Ages, including the Bronze Age, is currently available. Secondly, the ancient
human groups of Baraba exhibit all general features of the ancient West Siberian population, yet with
some specific features (Molodin, 1985, 2001). Thirdly, we have a unique collection of palaeoanthropo-
logical remains for all Baraba populations from the Bronze Age. The high degree of preservation of
these bones enables their analysis using methods from both physical anthropology and molecular gen-
etics. It is also important that these remains were obtained from burial grounds located within a small
territory (Fig. 1). As a result, the populations occupying this territory were to some extent culturally
homogenous during each of the historical periods considered.
Using this material we have attempted to assess changes in the composition of mtDNA lineages in
the gene pools of ancient Baraba populations during the Bronze Age, comparing the genetic results with
the archaeological and anthropological evidence pointing to the putative migratory events thought to
have occurred during this period.
* Corresponding author: alexpil@mail.ru
a Institute of Archaeology and Ethnography, Siberian
Branch, Russian Academy of Sciences, Novosibirsk,
Russia
b Institute of Cytology and Genetics, Siberian Branch,
Russian Academy of Sciences, Novosibirsk, Russia
Abstract
In this paper we present archaeological and anthropological
data on human migrations in the Western Siberian forest-
steppe region during the Bronze Age (4th–beginning of 1st mil-
lennium BC). These data, accumulated over forty years of in-
tensive research in the region, are compared to new results
showing the diversity of mitochondrial DNA (mtDNA) lin-
eages in this region during that period (92 mtDNA samples
from seven ancient human groups). Preliminary analyses have
demonstrated the usefulness of ancient DNA in tracing and
unravelling patterns of past human migrations.
Keywords
West Siberian forest steppe, Bronze Age cultural groups,
human migrations, ancient mtDNA
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94 VYACHESLAV I. MOLODIN, ALEXANDER S. PILIPENKO ET AL.
Materials and methods of the palaeogenetic study
The ancient West Siberian groups studied are listed in Table 1 and Figure 2. DNA was extracted from the
compact material of long bones (femur, tibia or humerus) and/or from the teeth. Whenever possible,
samples were taken from different parts of the skeleton (eg, postcranial bone and tooth) from each in-
dividual. For a minority of individuals different parts of the same bone had to be used.
DNA was extracted according to the method described by Pilipenko and colleagues (2008, 2010). In
brief, surfaces of bone and tooth samples were cleaned mechanically followed by treatment by bleach
Fig. 1 | Location of the archaeological sites
from which samples for the genetic studies
were obtained. 1 Sopka-2; 2 Tartas-1; 3 Pre-
obrazhenka-6; 4 Stariy Sad; 5 Chicha-1
(including Kurgan Zdvinsk-1)
Table 1 | The Bronze Age West Siberian cultural groups studied
Ancient human group Date
(millennium BC)
Archaeological
sites
Periods
Ust-Tartas Culture 4th – first half of 3rd Sopka-2/3,
Sopka-2/3A
Early Metal Period
Odinovo Culture Beginning of 3rd Sopka-2/4A
Preobrajenka-6
Early Bronze Age
Early Krotovo Culture End of 3rd – beginning of 2nd Sopka-2/4B Middle Bronze Age
Late Krotovo Culture First quarter of 2nd Sopka-2/5, Tartas-1 Middle Bronze Age
Andronovo (Fedorovo) Culture First half of 2nd Tartas-1 Middle Bronze Age
Baraba Late Bronze
Age Culture
End of 2nd Stariy Sad Late Bronze Age
Late Irmen Culture 9th–8th centuries BC Chicha-1 Transition from Bronze
to Iron Age
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HUMAN MIGRATIONS IN THE SOUTHERN REGION OF THE WEST SIBERIAN PLAIN DURING THE BRONZE AGE 95
and UV. Bone powder was drilled from the internal area of the samples. DNA was extracted from the
powder with a 5M guanidinium thiocyanate (GuSCN) buffer (pH 11.0) for 48 hours at 65°C followed by
phenol/chlorophorm extraction. The DNA was precipitated from the aqueous phase with isopropanol in
the presence 1M NaCl. At least two extractions were conducted for each individual under study (in most
cases, three).
Amplification of Hyper-Variable Region (HVR) I of mtDNA was performed using three different
techniques: amplification using four overlapping fragments (Haak et al., 2005), amplification with primer
pairs HA1 and HA3, HB1 and HB3, HC1 and HC3 that produced three overlapping fragments (Adcock et
al., 2001) and nested PCR (two reaction rounds) that produced one long fragment (Pilipenko et al., 2008).
Cloning of PCR products and sequencing of clones have been performed for a half of the samples.
For the remaining samples the cloning procedure is in progress. A set of PCR products was cloned with
pGEM-T Easy Vector Sestem Kit (Promega, USA) and a minimum of 30 clones were sequenced for each
individual.
Sequencing reactions were performed with an ABI Prism BigDye Terminator Cycle Sequencing
Ready Reaction Kit (Applied Biosystems, USA), according to the manufacturer’s instructions. The se-
quences of the products were analyzed on an ABI Prism 3100 Genetic Analyzer (Applied Biosystems,
USA) at the DNA Sequencing Center (Novosibirsk, Russia, www.sequest.niboch.nsc.ru).
The sequencing results were analysed using Sequence Scanner v1.0 (Applied Biosystems, USA)
and DNAStar Lasergene v. 7.1.0 (DNASTAR, USA) software. The phylogenetic tree was constructed
using Network 4.5.1.0 software (http://www.fluxus-engeneering.com).
Precautions against contamination: All stages of work with ancient material were carried out in
specially equipped, isolated clean rooms with positive air pressure, using special cloth, facemasks,
glasses and sterile gloves. All work surfaces were routinely cleaned with a 5% solution of bleach and ir-
radiated by UV light. Blank controls were run in parallel with samples throughout the extraction and
amplification procedures to identify possible contamination. MtDNA HVR I sequences were deter-
mined for all employees working with ancient DNA and (to the extent possible) for participants of exca-
vations and anthropologists who might have had contact with remains.
Results and discussions
The degree of contamination and the authenticity of the aDNA results obtained
Despite the strict experimental conditions we were unable to eliminate the problem of contamination
completely. In particular, contamination was ascertained in a small proportion of the extraction and PCR
blank controls (less than 1% of control number). All extracts or PCR-products were discarded in such
cases. Additionally, we identified singular divergent sequences among clones from several mtDNA
samples studied. These sequences were regarded as the results of contamination as they were not re-
producible in the second PCR or/and extraction. The other necessary conditions for the authenticity of
obtained ancient DNA sequences were as follows: consistency of the results obtained in repeated PCR
from one extract (for the same region or for several overlapping DNA fragments). Consistency between
the results of multiple (at least two) DNA extractions (including extractions from different parts of the
skeleton, such as the femur and teeth). The divergence of obtained aDNA sequences from those of staff
members (geneticists, archaeologists, anthropologists) who may have had contact with the samples be-
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96 VYACHESLAV I. MOLODIN, ALEXANDER S. PILIPENKO ET AL.
fore or during the genetic study. Presence among the clones of a specific pattern of biochemical degra-
dation (cytosine deamination). In our opinion, these criteria meet the main modern requirements for
verification of aDNA data.
In total, we successfully analyzed 92 mtDNA samples from seven ancient human groups from Ba-
raba and adjacent areas (Fig. 3; Table 2–3 see appendix).
The genesis of West Siberian forest-steppe populations
during the Neolithic and Early Metal periods
In order to estimate the impact of the migration waves on ethnogeny it is necessary to characterize the
archaeological, anthropological and genetic contexts preceding the migratory events. For this purpose, it
is also necessary to investigate the early populations of the region.
In contrast to the occupation of the southern region of West Siberia, modern humans arrived in the
Ob-Irtysh interfluve relatively late, at the end of the Pleistocene, about 13–14 thousand years ago (Oklad-
nikov, Molodin, 1983; Petrin, 1986). The absence of burials dating back to this period in the region does
not allow us to conduct a biological investigation of this earliest population. The most ancient anthro-
pological material available is from the Neolithic period (4th–5th millennium BC). More than two dozen
Fig. 2 | Chronological time scale of Bronze Age Cultures from the Baraba region
Fig. 3 | Phylogenetic tree of 92 mtDNA samples obtained from the seven Bronze Age cultural groups
from the Baraba region. Color coding of the groups as in Figure 2
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HUMAN MIGRATIONS IN THE SOUTHERN REGION OF THE WEST SIBERIAN PLAIN DURING THE BRONZE AGE 97
Neolithic burials have been excavated to date. The material from these excavations has allowed the char-
acterization of specific aspects of the material culture and anthropological type of the Neolithic group
(Polos’mak et al., 1989; Molodin, 2001). The anthropological analysis of the material allowed us to de-
tect a specific craniological type in the Baraba population, which was assigned to one of the anthropo-
logical formations discovered by V.V. Bunak in 1956 through the analysis of Neolithic materials from the
northern forest zone of the East European Plain. Bunak called it the “northern Eurasian anthropological
formation” (Bunak, 1956).
This anthropological type developed in a zone that is intermediate to the geographic areas occupied
by the classic Caucasoids and the Mongoloids. The exists substantial anthropological evidence showing
a wide geographic distribution of this anthropological formation: from the Trans-Urals forest and the
Barabian province of Western Siberia in the east to Karelia and the Baltic in the west (Chikisheva, 2010).
It is therefore possible to determine the cultural and anthropological background that existed in the
Baraba region in the beginning of the Bronze Age. The availability of a small number of materials suit-
able for DNA analysis will also allow us to determine the composition of mtDNA lineages of the Baraba
population from the Neolithic (the first results have already been obtained, but are not discussed in this
paper).
The earliest group from the Bronze Age in the Baraba region is represented by the Ust-Tartas popu-
lation (Molodin, 2001). Several Ust-Tartas burial grounds with more than 150 graves have been exca-
vated to date (Fig. 4), some of which revealed the presence of bronze adornments. The results of archaeo-
Fig. 4 | Typical collective burial of Ust-Tartas Culture representatives
(Burial N 655, Sopka-2/3 burial ground)
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98 VYACHESLAV I. MOLODIN, ALEXANDER S. PILIPENKO ET AL.
logical and radiocarbon dating allow us to date this culture to between the 4th millennium BC and the
first half of the 3rd millennium BC.
Another cultural group in the Baraba region existed contemporaneously with the Ust-Tartas cul-
ture, the “Comb-pit Ware culture”. Unlike the Ust-Tartas, the materials from the Comb-pit Ware cul-
ture are associated predominantly with settlements. In addition, several burials have been investigated,
making it possible to reconstruct the burial practices of this culture (Molodin, 1985, 2001).
Representatives of the Ust-Tartas culture belonged to a single craniological type preceding the Neo-
lithic populations of the region (Chikisheva, 2010). Craniometric analyses of materials from burials
with comb-pit ware also show the characteristic features of the northern Eurasian anthropological
formation, but demonstrate greater affinity with the northern periphery of the West Siberian Neolithic
cultures (Chikisheva, 2010), suggesting a predominantly autochthonous development of the Baraba
populations from the Early Metal period.
We have analyzed 18 mtDNA samples from the Ust-Tartas population to date (Fig. 3). The results
obtained thus far allow us to draw several preliminary conclusions about the genetic background in the
region in the beginning of the Bronze Age. By the Early Metal Period the mtDNA pool structure was al-
ready mixed and consisted of both Western and Eastern Eurasian haplogroups in nearly equal propor-
tions. The eastern Eurasian mtDNA cluster was represented by Haplogroups A, C, Z, D, which are most
typical of modern and perhaps ancient populations located in the east of the region studied. Haplo-
groups C and D were predominantly represented by widely distributed root haplotypes. A lineage of
Haplogroup A that was detected in two Ust-Tartas samples represents a subcluster that is apparently
characteristic of West Siberia and the Volga-Ural Region. The observed presence of Haplogroup Z
lineages with a high frequency in the Ust-Tartas group was unexpected, since these lineages are nearly
absent in the gene pool of modern indigenous West Siberian populations.
It is worth noting that the Western Eurasian mtDNA haplogroups in the Ust-Tartas series were rep-
resented only by Haplogroup U lineages, and specifically by the three subgroups – U2e, U4, U5a1. These
results are in agreement with previous data indicating that Haplogroup U lineages (particularly Sub-
groups U5 and U4) predominated in Eastern, Central and Northern European hunter-gatherer groups
from 14000 to 4000 years ago (Bramanti et al., 2009; Malmstrom et al., 2009), and possibly in earlier
periods (Krause et al., 2010). The geographic area within which this genetic feature is observed appears
to be broad (Fig. 5). Apparently, Baraba was near the eastern periphery of this area.
The Early and the beginning of Middle Bronze Age populations: an autochthonous evolution
The local evolution of the cultural groups described above led to the emergence of a new group in the Ba-
raba region in the beginning of the 3rd millennium BC. This group, referred to as Odinovo culture, dif-
fered in terms of their grave goods and funerary practices (Fig. 6) (Molodin, 2008).
The study of Odinovo culture settlements and several burial grounds with characteristic funerary
practices showed that its carriers had well-developed bronze casting and weapons of the Seima-Turbino
type (Molodin, 2008). The material culture of this group combined features of the preceding Comb-pit
ware and Ust-Tartas archaeological cultures. It is possible that both these substrates were involved in the
formation of the Odinovo culture. Anthropological evidence confirms the autochthonous development
of the culture. An anthropological affinity of the group to the Ust-Tartas and the Neolithic craniological
series from the Baraba forest steppe has been established (Chikisheva, 2010).
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HUMAN MIGRATIONS IN THE SOUTHERN REGION OF THE WEST SIBERIAN PLAIN DURING THE BRONZE AGE 99
At the end of the 3rd and beginning of the 2nd millennium BC, a striking and original culture existed in
the West Siberian forest steppe zone. The culture, known as Krotovo, obviously developed autochthon-
ally (Molodin, 1985). Undoubtedly, one of its ancestral components was the preceding Odinovo group.
However, undeniable innovations are observed in the Krotovo funerary practices and inventories. These
new features were probably associated with episodic migrations of representatives of the Petrovo culture
from the modern North Kazakhstan territory (Zdanovich, 1988) to the Baraba forest steppe. Moreover,
present among the Krotovo materials were specific forms of daggers, stalked spearheads and beads of
chalcedony, jaspilite and enstatite (the nearest deposits of these minerals are located in Central Asia and
Kazakhstan) (Fig. 7a,b,c), suggesting the influence of Central Asian cultures in the Baraba region during
the Middle Bronze Age (end of 3rd–beginning of 2nd millennium BC) (Molodin, 1988). This is the most
ancient external influence on the material culture of the Baraba population from the Bronze Age that
might have been accompanied by a migration wave.
The craniological analysis of early Krotovo populations, however, showed the presence of an autoch-
thonous morphological complex that was typical of earlier groups in the region, except for a higher
cranium in Krotovo males (Chikisheva, 2010).
The genetic analysis of the Odinovo and Krotovo groups (10 and 6 samples, respectively) (Fig. 3) did
not reveal any differences between them and the previous Ust-Tartas group, such as the presence of new
mtDNA haplogroups. The mtDNA pool structure was still mixed. The East Eurasian haplogroups were
Fig. 5 | Location of ancient human groups with a high frequency of mtDNA haplogroups U5, U4 and U2e lineages. The area of North-
ern Eurasian anthropological formation is marked by yellow region on the map (References: 1 Bramanti et al., 2009; 2 Malmstrom et
al., 2009; 3 Krause et al., 2010; 4 this study)
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100 VYACHESLAV I. MOLODIN, ALEXANDER S. PILIPENKO ET AL.
Fig. 6 | A burial of Odinovo culture representatives (Sopka-2/
4A burial ground)
Fig. 7 | Specific spearhead (a), daggers (b) and beads (c) from the Early Krotovo Culture sites, indicating the influence
of Central Asian cultures in the Baraba region during the Middle Bronze Age
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represented by the D, C, Z (in both the Odinovo and Krotovo groups) and A (in the Krotovo group) ha-
plogroups. The East Eurasian lineages identified were phylogenetically close (lineages of haplogroups A,
C, Z) or even identical (D haplogroup, 16223–16362 lineages) to the samples from the Ust-Tartas group.
The West Eurasian part of the samples were represented by the U5a1 (Odinovo group) and U2e (Krotovo
group) haplogroup lineages.
Although only a small series of samples have been investigated thus far, the data obtained reveal
continuity between the Odinovo and Krotovo populations and the earlier Ust-Tartas group. These find-
ings are consistent with the autochthonous development of the Baraba populations during the Early and
the beginning of the Middle Bronze Age, as well as with the anthropological evidence.
Our data did not allow us to detect any Central Asian genetic influence on the Krotovo population,
although this conclusion is still preliminary due to the small sample sizes involved.
The Middle and Late Bronze Age: significance of the Andronovo (Fedorovo) population
migration wave
The Krotovo culture continued to develop indigenously for several centuries. At the beginning of the 2nd
millennium BC, the migration of the Andronovo culture population to the south of the West Siberian
Plain started, probably originating in the territory of present-day Central Kazakhstan. Apparently, this
migration wave, which spread from its epicenter to the west, north and east (Kuz’mina, 1994), was a
highly significant phenomenon in Asia.
The newly arrived Caucasian Andronovo population had a great influence on the development of
the indigenous cultures of the West Siberian steppe and forest steppe regions (including the Baraba
forest steppe). According to the archaeological evidence, the newly arrived groups coexisted with the
aborigines for some time, which can be explained by the differing nature of their economic activities
(nomadic and seminomadic pastoralism in the Andronovo group and sedentary or semisedentary pas-
toralism in the aboriginal Late Krotovo group). During this time the migrants had a strong impact on the
material culture of the indigenous group. As a result, the bronze weapons and adornments of the abo-
riginal cultural group changed from Seima-Turbino to Srubno-Andronovo forms (Fig. 8a; 9a). The con-
tact between these two groups is also reflected in their funerary practices and inventories, which com-
bined novel and traditional features of the Baraba region. Archaeological evidence indicates that the
Andronovo impact on the Late Krotovo population was in fact manifold.
The adaptation of the migrant Andronovo group in the West Siberian forest steppe was a long pro-
cess that resulted in the absolute dominance of this population in the region. The Krotovo populations
were partially displaced to the north and partly assimilated. The material and spiritual culture of the An-
dronovo group in West Siberia gradually returned to its classical tradition (Fig. 8b; 9b), although with
some new unique features.
At the late stage of the development of the Krotovo culture, the craniological complex had changed
markedly in comparison to the anthropological type common in all preceding Neolithic and Bronze Age
populations in the Baraba region. The changes observed in males from the Late Krotovo group could be
attributed to the influence of Andronovo representatives. Some Mongoloid features, which were not re-
lated to the previous autochthonous substrate, appeared in females from the Late Krotovo group. The
Europoid component observed in these females does not fall within the generally accepted criteria for
the Andronovo anthropological type.
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102 VYACHESLAV I. MOLODIN, ALEXANDER S. PILIPENKO ET AL.
Fig. 8a | Typical burials of the Late Krotovo Culture; b. typical burial of the Andronovo (Fedorovo) Culture
Fig. 9a | Daggers of Srubno-Andronovo form; b. Ceramic pot with specific ornamentation of the Andronovo (Fedorovo) Culture
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HUMAN MIGRATIONS IN THE SOUTHERN REGION OF THE WEST SIBERIAN PLAIN DURING THE BRONZE AGE 103
Some anthropological data suggest that the Krotovo population probably did not interact directly with
the Andronovo populations, but rather with population groups displaced from the south and west by the
Andronovo migration wave. In these groups, the Andronovo influence was already present (Chikisheva,
2010).
The anthropological analysis of representative groups of the local Andronovo culture in southern
West Siberia revealed a complex population composition. It suggests the existence of different genetic
roots and different degrees of involvement of autochthonous Mongoloid component in their genesis.
The anthropological analysis of the West Siberian Andronovo population shows at least four cran-
iological types. Three types are related to the Palaeocaucasian race and are represented by proto-Euro-
pean anthropological type variants. The fourth, Mongoloid, component is autochthonous. The most
intensive interactions between the Andronovo migrants and the indigenous populations apparently
occurred in the Baraba forest steppe and the right bank of the upper Ob River (Chikisheva and Pozdnya-
kov, 2003).
To investigate the putative impact of Andronovo migrants on the mtDNA pool structure of the
indigenous populations in Baraba, mtDNA samples from the Late Krotovo (n=20) and Andronovo
(n=20) groups in this region were analyzed (Fig. 3) and compared to recently published data (n=10)
(Keyser et al., 2009) and our own unpublished data (n=6) on mtDNA lineages from West Siberian An-
dronovo populations located outside the Baraba forest steppe.
The genetic influence of migrants can be detected by the appearance of a new mtDNA haplogroup
that was absent in the populations preceding the migration wave. This new mtDNA haplogroup, a West
Eurasian T haplogroup, was detected in the Late Krotovo population. The T haplogroup appears simul-
taneously (with a 15% frequency) in the Krotovo and Andronovo groups, but was completely absent in
all preceding Baraba populations. We therefore consider the appearance of the Haplogroup T-lineage as
the most likely genetic marker of the Andronovo migration wave to the region.
This assumption is confirmed by mtDNA studies of Andronovo groups from other West Siberian
areas. Haplogroup T lineages were found, with a frequency of 25%, in the samples (n=16) taken from
two Andronovo groups from the Krasnoyarsk and upper Ob River areas.
We also detected another remarkable feature in the mtDNA pool of the Andronovo group from Ba-
raba. Most mtDNA samples belonged to haplogroups, such as the East Eurasian A and C haplogroups,
that are typical of preceding Baraba indigenous populations. Still, these haplogroups were not found in
the other West Siberian Andronovo groups. Apparently, the Andronovo group from Baraba assimilated
the aboriginal Krotovo population, from which it obtained these East-Eurasian mtDNA haplogroups.
Obviously, there was reciprocal genetic contact between the migrant and indigenous groups in the
region.
The above mentioned ethno-cultural processes had a strong influence on the genesis of the cultural
groups from the Late Bronze Age in this area. The Andronovo group in the south of West Siberia was
replaced by the Irmen culture, which was originally from the West Siberian steppe and forest steppe
zones around the 14th century BC, and occupied the region extending from the Achinsk and Mariinsk
forest steppe in the east to the Irtysh River in the west (Molodin, 1985; Bobrov et al., 1993; Matveev,
1993). The culture existed in the region for more than five hundred years and was characterized by a
powerful economy, which included animal husbandry and agriculture.
A considerable number of Irmen culture settlements and burial grounds have been investigated.
As a result, the characteristic features of the Irmen ceramics, inventory and funerary practices have been
identified. However, the processes underlying the development of this group are not yet completely
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104 VYACHESLAV I. MOLODIN, ALEXANDER S. PILIPENKO ET AL.
understood. Obviously, one major component in the genesis of the group is represented by the preced-
ing Andronovo population, which had already assimilated features from West Siberian aborigines.
Analysis of the anthropological material from the Irmen culture in the Baraba region showed that
their average craniological features are consistent with Caucasoid values, or indicate the presence of a
small Mongoloid admixture. Anthropologically, the local Irmen group from Baraba was relatively homo-
genous. The results of the cross-group statistical analysis confirm the participation of the Andronovo
population in the development of the anthropological type of all local Irmen groups (Chikisheva, 2010).
The study of mtDNA samples from the Irmen population is at an early stage and will not be dis-
cussed in this paper.
The Irmen population engaged in intense contact with neighbouring groups in different parts of its
vast territory. These interactions are reflected in the peculiarities of its material and spiritual culture and
in the structure of the Irmen population as based on the physical anthropology data available. In the
east, the Irmen population interacted with the neighbouring Karasuk culture group (Chlenova, 1972). In
the west, including the Baraba forest steppe region, interactions with neighbours were of a different na-
ture. Several migration waves to Baraba occurred in this period. The first was associated with the mi-
gration of representatives of the Suzgun culture from the north-west. Accordingly, a syncretic culture,
known as the Baraba variant of Suzgun culture, emerged in the northern part of the Baraba forest steppe
(Molodin, Chemyakina, 1984). Two other migratory waves simultaneously occurred in the southern part
of the West Siberian forest steppe. The first was associated with the Roller Ware Culture, which occupied
the vast Eurasian steppes during the Late Bronze Age (Chernykh, 1983). This population also spread
from the West Siberian steppe zone up to Kulunda steppe in the east. Apparently, a small proportion
of this group migrated to the southern forest steppe, as evidenced by the presence of roller ware in the
Irmen settlements (Molodin et al., 2000).
The largest migration wave to the steppe and forest steppe zones of Western Siberia was that of the
Begazy-Dandybay populations during the Final Bronze Age, who came from the territory currently cor-
responding to modern Central Kazakhstan. The adaptation of these populations in the West Siberian
steppe zone resulted in the development of syncretic groups, which then spread farther north and east.
One such group was the Pahomov culture (Korochkova, 2009). This group migrated to the Baraba forest
steppe and interacted intensively with the Irmen and Suzgun populations. A consequence of this pro-
cess is the emergence of archaeological sites, such as the Stariy Sad burial ground in the Baraba region,
where specific burial practices and anthropological types are found (Molodin and Neskorov, 1992). This
cultural group is referred to as Baraba Late Bronze Age culture. The anthropological analysis revealed
differences between males and females in the population that cannot be explained by sexual dimor-
phism. This suggests that the series originated from anthropologically similar but not identical sources.
The main craniological type detected in the population is similar to the Southern Eurasian Anthropo-
logical Formation. The female series shows similarities with the Andronovo populations from Northern
Kazakhstan, which suggests that the latter populations were involved in the development of the Baraba
Late Bronze Age culture. Alternatively, the specific pattern of marriage may have caused the migration
of females from the Begazy-Dandybay population, which possibly brought the southern ceramic tradi-
tion to the Baraba region (Chikisheva, 2010).
A small but informative series of mtDNA samples from the Baraba Late Bronze Age culture popu-
lation (n=5) was analyzed (Fig. 3), revealing the presence of MtDNA lineages (East Eurasian A and C
lineages) that mark the genetic continuity with aboriginal Baraba groups. At the same time, the series
includes the Haplogroup-T lineage, which we believe marks the Andronovo migration wave to West
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HUMAN MIGRATIONS IN THE SOUTHERN REGION OF THE WEST SIBERIAN PLAIN DURING THE BRONZE AGE 105
Siberia. Our data is therefore consistent with the putative origin of the West Siberian Late Bronze Cul-
ture population as the result of interaction between the Baraba indigenous genetic substrate and the
newly arrived group. However, the migration of Begazy-Dandybay culture representatives to the Baraba
region is still questionable.
The transition from the Late Bronze Age to the Early Iron Age: a southern influence
Thus, a complex ethno-cultural structure developed in the end of the Bronze Age in the West Siberian
forest steppe region, becoming even more complex during the transition from the Bronze to the Early
Iron Age (9th–8th century BC). The abrupt cooling of the climate of Western Siberia (at its strongest
in the Holocene [Levina, Orlova, 1993]) led to the deterioration of ecological conditions in the taiga zone.
In consequence, intense migration of taiga populations to the forest steppe zone in the south occurred
in the territory that extends from the Urals in the west to the Yenisei River in the east. As a result, an en-
semble of syncretic archaeological cultures developed in the West Siberian forest steppe region, includ-
ing the Gamajun, Krasnoozersk and Zavyalovo cultures. This powerful migration wave triggered in-
tense movement of various ethnic and cultural groups in the forest, forest steppe and even steppe zones
towards the meridional and latitudinal directions. The study of the Baraba forest steppe area suggests
that such processes occurred. On the one hand, the autochthonous Irmen culture evolved into the Late
Irmen culture, an evolutionary process that was detected in the materials obtained from a large number
of archaeological sites. On the other hand, the discovery and excavation of the Chicha-1 settlement in the
south of the Baraba forest steppe demonstrated both the occasional appearance of migrants from the
north (representatives of the Suzgun and even northern taiga Atlym cultures) and permanent migration
flows of representatives from the Krasnoozersk culture from the north-west and from the Berlik culture
from the Kazakhstan steppes in the south-west (Fig. 10) (Molodin et al., 2009). It is important to note
that the members of the above mentioned cultural groups and the aboriginal Late Irmen group lived in
different parts of the same settlement (Molodin and Parzinger, 2009). Obviously, this led to intense in-
teractions between the ethno-cultural groups, resulting both in the combination of cultural traits and
traditions, as well as in the genetic admixture of populations as a result of intergroup marriages.
The analysis of mtDNA samples from the Chicha-1 population revealed some interesting patterns.
Crucial changes in the composition of mtDNA haplogroups in the gene pool were observed as compared
to the earlier Baraba groups studied (Fig. 3). Dominance of Western Eurasian haplogroups and the near
absence of East Eurasian were observed. Additionally, several new West Eurasian haplogroups appeared
in the region, including Haplogroups U1a, U3, U5b, K, H, J and W.
The phylogeographic analysis suggests that the distribution and diversification centres of several of
these mtDNA haplogroups and specific lineages are located on the west and south west of the Baraba
forest steppe region, on the territory corresponding to modern-day Kazakhstan and Western Central
Asia (Fig. 10). Apparently, the migration wave from the south strongly influenced the gene pool of the
Baraba population in the transitional period from the Bronze to the Early Iron Age. The impact of the
northern human groups was probably less evident in the south of the Baraba forest steppe, at least at the
mtDNA level.
The frequencies of the new haplogroups mentioned above in the mtDNA pools of modern West
Siberian populations show a different pattern however. The haplogroup lineages U1a and U3 are found
only sporadically in the modern populations. Conversely, Haplogroup H is one of the most frequent West
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106 VYACHESLAV I. MOLODIN, ALEXANDER S. PILIPENKO ET AL.
Eurasian haplogroups in the modern indigenous populations of the region. Unexpectedly, the H-haplo-
group lineages appear in the region only during the transition from the Bronze to the Early Iron Age.
Subsequently, in the Scythian-Sarmatian period, a large cultural group, called the Sargat culture,
developed in the region. Its representatives were widespread across the region, from the Ob River to the
Urals. Their development represented one of the most significant cultural events in North Asia.
Conclusions
According to the archaeological and anthropological evidence, the continuity between ethno-cultural
groups from different periods and the inheritance of new cultural and genetic components as a result of
several migration waves markedly affected the genesis of the populations in the West Siberian forest
steppe zone during the Bronze Age.
Ancient mtDNA analyses have confirmed the key role played by autochthonous genetic compo-
nents in composing the gene pool of the populations, especially during the Early and Middle Bronze
Age. These components were represented by the Eastern Eurasian haplogroups A, C and Z, and the
Western Eurasian haplogroup U5a.
On the other hand, the results also reveal some changes in the mtDNA pool structure throughout
the Bronze Age. Some of these changes, which point to migration waves to the West Siberian forest
Fig. 10 | Directions of migration waves in West Siberian forest steppe region during the transition from Bronze to Early Iron Age
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HUMAN MIGRATIONS IN THE SOUTHERN REGION OF THE WEST SIBERIAN PLAIN DURING THE BRONZE AGE 107
steppe zone, are in agreement with the archaeological and anthropological evidence. The most relevant
migration waves occurred during the Middle Bronze Age (represented by the migration of the Andro-
novo culture, probably marked by Haplogroup-T lineages) and the transition from the Bronze to the
Iron Age (represented by the migration from the south, marked by the U1a, U3 and H haplogroup lin-
eages). These preliminary results indicate the usefulness of ancient DNA analysis as an additional tool
for the reconstruction of migratory and ethnogenic processes in the Western Siberian region. To im-
prove the efficacy of genetic methods it will be necessary to analyze other genetic markers in addition to
mitochondrial DNA.
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HUMAN MIGRATIONS IN THE SOUTHERN REGION OF THE WEST SIBERIAN PLAIN DURING THE BRONZE AGE 109
Table 2 | List of samples from Baraba forest-steppe Bronze Age populations studied
Sample
number
Sample
name
Archaeological
site
Burial number
(skeleton number)a
Archaeological culture MtDNA
haplogroup
1 Ut2 Sopka-2/3 600 Ust-Tartas U2e
2 Ut3 Sopka-2/3 655(C) D
3 Ut4 Sopka-2/3 655(D2) C
4 Ut5 Sopka-2/3 655(D4) A
5 Ut7 Sopka-2/3A 627(C) C
6 Ut9 Sopka-2/3A 658(A) U5a1
7 Ut12 Sopka-2/3A 611(K) U2e
8 Ut14 Sopka-2/3A 619 U5a1
9 Ut16 Sopka-2/3A 611(B) C
10 Ut17 Sopka-2/3A 611(C) U2e
11 Ut19 Sopka-2/3A 358 U4
12 Ut31 Sopka-2/3A 341(3) Z
13 Ut32 Sopka-2/3A 628 Z
14 Ut33 Sopka-2/3 656(F) Z
15 Ut34 Sopka-2/3 656(G) Z
16 Ut37 Sopka-2/3 655(A) C
17 Ut38 Sopka-2/3 655(B) A
18 Od1 Preobrazhenka-6 18 Odinovo C
19 Od2 Preobrazhenka-6 2 Z
20 Od3 Preobrazhenka-6 36 D
21 Od12 Sopka-2/4A 184(2) D
22 Od13 Sopka-2/4A 194(1) D
23 Od27 Sopka-2/4A 258(2) D
24 Od29 Sopka-2/4A 523 C
25 Od65 Sopka-2/4A 520(1) D
26 Od66 Sopka-2/4A 594 U5a1
27 Od67 Sopka-2/4A 588(1) D
28 Kr1 Sopka-2/4B 407(1) Early Krotovo C
29 Kr5 Sopka-2/4B 171(1) Z
30 Kr10 Sopka-2/4B 177(1) A
31 Kr79 Sopka-2/4B 305 U2e
32 Kr80 Sopka-2/4B 46 U2e
33 Kr87 Sopka-2/4B 65 D
34 Pkr1 Sopka-2/5 108(1) Late Krotovo T*
35 Pkr2 Sopka-2/5 126 T*
36 Pkr3 Sopka-2/5 123 U5a*
37 Pkr4 Sopka-2/5 141 U5a*
38 Pkr5 Sopka-2/5 117 T*
39 Pkr6 Sopka-2/5 140(2) U5a1
40 Pkr7 Sopka-2/5 349(2) G2a
41 Pkr8 Sopka-2/5 334 A
42 Pkr9 Sopka-2/5 376 U5a1
43 Pkr10 Sopka-2/5 625(1) S
44 Pkr11 Sopka-2/5 132 C
45 TK1 Tartas-1 85 C
46 TK2 Tartas-1 77 C
47 TK3 Tartas-1 76 A
48 TK4 Tartas-1 15 C
49 TK5 Tartas-1 73 U5a1
50 TK6 Tartas-1 84 S
51 TK7 Tartas-1 80(2) S
52 TK8 Tartas-1 72 C
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110 VYACHESLAV I. MOLODIN, ALEXANDER S. PILIPENKO ET AL.
53 TK21 Tartas-1 75 A
54 TA5 Tartas-1 227(1) Andronovo (Fedorovo) T*
55 TA4 Tartas-1 227(2) T*
56 TA7 Tartas-1 218(1) A
57 TA8 Tartas-1 225 S
58 TA9 Tartas-1 217(1) S
59 TA10 Tartas-1 217(2) U5a1
60 TA11 Tartas-1 213 U5a1
61 TA13 Tartas-1 211 C
62 TA14 Tartas-1 180 S
63 TA15 Tartas-1 182(1) T*
64 TA16 Tartas-1 182(2) S
65 TA17 Tartas-1 189 A
66 TA18 Tartas-1 223(1) U5a1
67 TA20 Tartas-1 188 S
68 TA21 Tartas-1 196 S
69 TA22 Tartas-1 208(1) U5a1
70 TA23 Tartas-1 240 C
71 TA25 Tartas-1 124 A
72 TA26 Tartas-1 137 C
73 TA27 Tartas-1 176 C
74 Sts1 Stariy Sad 1 Baraba Late Bronze Age S
75 Sts11 Stariy Sad 49 A
76 Sts7 Stariy Sad 27 C
77 Sts9 Stariy Sad 59 U5a
78 Sts10 Stariy Sad 62 T1
79 Ch1* Chicha-1 D10-B1 Baraba population of
transition from the bronze
to the Early Iron Age
(Chicha-1 Settlement
population)
J
80 Ch2* Chicha-1 D10-B2 H
81 Ch3* Chicha-1 D20-B1 U5b
82 Ch4* Chicha-1 D20-B2 U4
83 Ch5* Chicha-1 D9-B1 U1a
84 Ch6 Chicha-1 D3a-B1 K
85 Ch7* Chicha-1 D3a-B.2 K
86 Ch8 Chicha-1 D8-B.1 D
87 Ch9 Chicha-1 IIIa U3
88 Ch10 Chicha-1 12 W
89 CSh11 Chicha-1 6(2) K
90 Ch12 Zdvinsk-1 1(2) U4
91 Ch13 Zdvinsk-1 2 U4
92 Ch14 Zdvinsk-1 4 H6a1
Sample
number
Sample
name
Archaeological
site
Burial number
(skeleton number)a
Archaeological culture MtDNA
haplogroup
Table 2 | continued
a For the Chicha-1 site burial designations indicate the following: for samples Ch1–Ch8 (infants buried in dwellings of Chicha-1
settlements) – D=dwelling number, B=burial number in that dwelling; Ch9 – individual was buried between dwellings in the
section IIIa of the settlement territory; Ch10, 11 – individuals buried in the ground part of the Chicha-1 necropolis; Ch12–14 in-
dividuals buried in the kurgan part of the Chicha-1 necropolis (previously designated “Zdvinsk-1”)
* These samples were preliminarily published by Pilipenko and colleagues (2008), but were subsequently reanalyzed using more
strict experimental conditions and verification criteria
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HUMAN MIGRATIONS IN THE SOUTHERN REGION OF THE WEST SIBERIAN PLAIN DURING THE BRONZE AGE 111
Table 3 | List of mtDNA HVR polymorphic sites
No of mtDNA
HVR I haplotype
MtDNA
haplogroup
mtDNA HVR I
polymorphic sites
Names of samples
1 A 223T-227C-290T-311C-319A Kr10, TK3, Sts11
2 A 148T-223T-227C-290T-311C-319A Ut5, Ut38
3 A 223T-227C-278T-290T-311C-319A TA17
4 A 189C-223T-290T-319A Pkr8, TA25
5 A 223T-242T-290T-319A TK21
6 A 188T-189C-223T-290T-319A-356C-362C TA7
7 C 223T-298C-327T Ut4, Ut7, Ut16, Ut37, Pkr10, TK1,
TK2, TA9, TA13, TA14, TA16, Sts7
8S129C-223T-298C-327T TK6, TK7, TK8, TA26, TA27
9S223T-297C-298C-327T Od29, Od1
10 S223T-298C-325C-327T Kr1, TA23
11 S223T-298C-325C-327T-362C TA8, Sts1
12 S148T-223T-288C-298C-327T Pkr 11, TK4, TA20, TA21
13 D 223T-362C Ut3, Od12, Od13, Od27, Od65,
Od67, Od3, Kr87
14 D 223T-294T-362C Ch8
15 G2a 223T-227G-278T-362C Pkr7
16 Z 185T-223T-260T-298C-300G Ut31, Ut32
17 Z 185T-223T-260T-263C-298C Ut33, Ut34
18 Z 185T-223T-260T-271C-298C Kr5
19 Z 185T-223T-260T-298C Od2
20 U1a 183S-189C-249C Ch5
21 U2e 129C-189C Ut2
22 U2e 129C-189C-246G Ut12, Ut17
23 U2e 129C-189C-256T Kr79, Kr80
24 U3 343G Ch9
25 U4 356C Ut19, Ch12, Ch13
26 U4 356C-362C Ch4
27 U5a1 192T-256T-270T Ut9, Pkr6, TK5
28 U5a1 192–256–270–318 Ut14
29 U5a1 172–192–256–270 Od66
30 U5a1 192–256–270–304 Pkr9, TA18
31 U5a1 192T-239T-256T-270T TA10,TA11,TA22
32 U5a 256T-270T Sts9
33 U5a 192T-270T Pkr3, Pkr4
34 U5b 189C-260T-270T Ch3
35 K 093C-224C-311C Ch6, Ch7
36 K 213A-224C-311C Ch11
37 T 126C-294T Pkr1, Pkr2, TA4, TA5
38 T2b 126C-189T-292T-294T-296T Pkr5
39 T 086C-126C-140C-294T-296T TA15
40 T1 126C-163G-186T-189C-294T Sts10
41 J 069T-126C Ch1
42 H 366T Ch2
43 H 111T-362C Ch14
44 W 223T-292T Ch10
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112 VYACHESLAV I. MOLODIN, ALEXANDER S. PILIPENKO ET AL.
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... As a result, a large set of data is available concerning the ancient ethno-cultural groups that occupied this territory from the Neolithic period (VII-VI millennium BC or even earlier) to the Late Middle Ages. The most representative materials, including collections of paleoanthropological remains, were obtained for all Baraba populations from the Bronze Age [12,13]. The large burial grounds are associated with the main regional Bronze Age ethno-cultural Table 1. ...
... For this research it is particularly important that early Baraba populations from the Neolithic and Early Bronze Age periods before the arrival of the Andronovo groups in the first half of II millennium BC (namely the Ust-Tartas, Odinovo and Krotovo groups), showing cranial features corresponding to the Uralic (or West Siberian) anthropological type, and belonging to the Northern Eurasian anthropological formation, are used. Using these anthropological materials, we carried out a screening study of the ancient mtDNA gene pool structure and its dynamics in the West Siberian forest-steppe zone, including samples from about 100 individuals to date (some of the preliminary results were published [12,14]). Only 10 samples that turned out to belong to haplogroup A10 after the initial screening were further used in this study. ...
... The following circumstances indicate that ancient samples were not cross-contaminated: experiments with haplogroup A10 samples dated to different periods of the Bronze Age were performed at different times with long time intervals between them; all series of samples include a large variety of lineages belonging to different East and West Eurasian haplogroups, in addition to the lineages of haplogroup A10 described in this paper (these data are being prepared for publication, see [12,14] for some preliminary results); and five lineages of A10 haplogroup with different HVR I haplotypes were present among the ancient mtDNA samples. ...
Article
Full-text available
Background The craniometric specificity of the indigenous West Siberian human populations cannot be completely explained by the genetic interactions of the western and eastern Eurasian groups recorded in the archaeology of the area from the beginning of the 2nd millennium BC. Anthropologists have proposed another probable explanation: contribution to the genetic structure of West Siberian indigenous populations by ancient human groups, which separated from western and eastern Eurasian populations before the final formation of their phenotypic and genetic features and evolved independently in the region over a long period of time. This hypothesis remains untested. From the genetic point of view, it could be confirmed by the presence in the gene pool of indigenous populations of autochthonous components that evolved in the region over long time periods. The detection of such components, particularly in the mtDNA gene pool, is crucial for further clarification of early regional genetic history. Results and Conclusion We present the results of analysis of mtDNA samples (n = 10) belonging to the A10 haplogroup, from Bronze Age populations of West Siberian forest-steppe (V—I millennium BC), that were identified in a screening study of a large diachronic sample (n = 96). A10 lineages, which are very rare in modern Eurasian populations, were found in all the Bronze Age groups under study. Data on the A10 lineages’ phylogeny and phylogeography in ancient West Siberian and modern Eurasian populations suggest that A10 haplogroup underwent a long-term evolution in West Siberia or arose there autochthonously; thus, the presence of A10 lineages indicates the possible contribution of early autochthonous human groups to the genetic specificity of modern populations, in addition to contributions of later interactions of western and eastern Eurasian populations.
... Both anthropological [54] and genetic data [47,55] indicate that until the late Bronze Age Central Asia was populated mainly by Europid Sintashta-Andronovo people while populations with Mongoloid traits and genes were confined East of the Altai. The first Eastern Hg lineages appeared in West Siberia at the beginning of Bronze Age [56] and in the Altai at the Middle Bronze Age [57]. However on the Pontic steppe large scale appearance of Eastern Hg-s is detected just in the Iron Age when European Scythians admixed with Scytho-Siberians, giving rise to 18-26% eastern lineages in European Scythians by the 2 nd century BCE [49]. ...
... As only 7 Estonian mitogenomes are available, they were grouped with other modern Baltic populations (Balt), so the similarity of these to the Conquerors probably derives from BalBA heritage. The connection to modern Finnish population can also be explained from the BalBA and steppe MLBA component which is present in modern Scandinavians, as Finnish sequence matches regularly appear together with Danish ones on our phylogenetic trees (S1 Figure, Networks 14,15,19,25,27,30,35,40,42,43,49,52,56). ...
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Endre Neparáczki and Zoltán Maróti contributed equally to this study. Abstract It has been widely accepted that the Finno-Ugric Hungarian language, originated from proto Uralic people, was brought into the Carpathian Basin by the Hungarian Conquerors. From the middle of the 19 th century this view prevailed against the deep-rooted Hungarian Hun tradition, maintained in folk memory as well as in Hungarian and foreign written medieval sources, which claimed that Hungarians were kinsfolk of the Huns. In order to shed light on the genetic origin of the Conquerors we sequenced 102 mitogenomes from early Conqueror cemeteries and compared them to sequences of all available databases. We applied novel population genetic algorithms, named Shared Haplogroup Distance and MITOMIX, to reveal past admixture of maternal lineages. Phylogenetic and population genetic analysis indicated that more than one third of the Conqueror maternal lineages were derived from Central-Inner Asia and their most probable ultimate sources were the Asian Huns. The rest of the lineages most likely originated from the Bronze Age Potapovka-Poltavka-Srubnaya cultures of the Pontic-Caspian steppe, which area was part of the later European Hun empire. Our data give support to the Hungarian Hun tradition and provides indirect evidence for the genetic connection between Asian and European Huns.
... Both anthropological [61] and genetic data [47,62] indicate that until the Bronze Age Asia was populated mainly by Europid Sintashta-Andronovo people west of the Altai, while popula- tions with Mongoloid traits and genes were confined east of the Altai. The first eastern Hg line- ages appeared in West Siberia at the beginning of Bronze Age [63], in the Altai at the Middle Bronze Age [64], while in Central Asia just around the 6 th century BC corresponding to the Xiongnu invasions [65]. ...
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It has been widely accepted that the Finno-Ugric Hungarian language, originated from proto Uralic people, was brought into the Carpathian Basin by the conquering Hungarians. From the middle of the 19th century this view prevailed against the deep-rooted Hungarian Hun tradition, maintained in folk memory as well as in Hungarian and foreign written medieval sources, which claimed that Hungarians were kinsfolk of the Huns. In order to shed light on the genetic origin of the Conquerors we sequenced 102 mitogenomes from early Conqueror cemeteries and compared them to sequences of all available databases. We applied novel population genetic algorithms, named Shared Haplogroup Distance and MITOMIX, to reveal past admixture of maternal lineages. Our results show that the Conquerors assembled from various nomadic groups of the Eurasian steppe. Population genetic results indicate that they had closest connection to the Onogur-Bulgar ancestors of Volga Tatars. Phylogenetic results reveal that more than one third of the Conqueror maternal lineages were derived from Central-Inner Asia and their most probable ultimate sources were the Asian Scythians and Asian Huns, giving support to the Hungarian Hun tradition. The rest of the lineages most likely originated from the Bronze Age Potapovka-Poltavka-Srubnaya cultures of the Pontic-Caspian steppe. Available data imply that the Conquerors did not have a major contribution to the gene pool of the Carpathian Basin.
... Late Bronze Age (LBA) Srubnaya-Alakulskaya individuals carried mtDNA haplogroups associated with Europeans or West Eurasians (17) including H, J1, K1, T2, U2, U4, and U5 (table S3). In contrast, the Iron Age nomads (Cimmerians, Scythians, and Sarmatians) additionally carried mtDNA haplogroups associated with Central Asia and the Far East (A, C, D, and M) (table S3) (11,18). The ab- sence of East Asian mitochondrial lineages in the more eastern and older Srubnaya-Alakulskaya population suggests that the appear- ance of East Asian haplogroups in the steppe populations might be associated with the Iron Age nomads, starting with the Cimmerians. ...
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For millennia, the Pontic-Caspian steppe was a connector between the Eurasian steppe and Europe. In this scene, multidirectional and sequential movements of different populations may have occurred, including those of the Eurasian steppe nomads. We sequenced 35 genomes (low to medium coverage) of Bronze Age individuals (Srubnaya-Alakulskaya) and Iron Age nomads (Cimmerians, Scythians, and Sarmatians) that represent four distinct cultural entities corresponding to the chronological sequence of cultural complexes in the region. Our results suggest that, despite genetic links among these peoples, no group can be considered a direct ancestor of the subsequent group. The nomadic populations were heterogeneous and carried genetic affinities with populations from several other regions including the Far East and the southern Urals. We found evidence of a stable shared genetic signature, making the eastern Pontic-Caspian steppe a likely source of western nomadic groups.
... Results of multidimensional scaling based on matrix of Slatkin population differentiation (F ST ) based on mtDNA HVRI sequences in the Tagar series and other ancient populations from different regions of Eurasia (details in S3Table). Populations: Tagar-Tagar series (red pentagon) (this study); Iron Age populations related with the 'Scythian world" (red circles): Pazyryk-Pazyryk culture from Altay Mountains (Russia, Kazakhstan, Mongolia)[1,2,4,[22][23][24][25]; Aldy_Bel-series from Aldy Bel culture, Arjan-2 burial complex, Tuva, Russia[4]; Scythians-Classic Scythians from North Pontic region[3,4,26]; Neolithic and Bronze Age populations (black squares): Yamnaya-Yamnaya culture population (Early Bronze Age)[18,[27][28][29]; Catacomb-Catacomb culture population (Bronze Age)[18,29]; Afanasievo-Afanasievo culture population from the Minusinsk Basin (Early Bronze Age)[27,30]; Okunevo-Okunevo culture population from the Minusinsk basin (Bronze Age, pre-Andronovo time); Andronovo_B-Andronovo time population from West-Siberian foreststeppe zone[31]; Andronovo_M-Andronovo culture population from Minusinsk basin[10]; Cisbaikalian_Neo-Serovo and Glazkovo cultures from Cis-Baikal region, Russia (Neolithic and Bronze Age)[32]; Tianshanbeilu-Tianshanbeilu site, eastern Xinjiang, China, Bronze Age (1900-1300 YBC)[33]; Bronze_MA-Middle Bronze Age population from the Mongolian Altai[34]; Lajia_Neo-population from the Lajia site, Qinghai, northwestern China (3800-3400 YBP)[35]; Jiangjialiang_Neo-Neolithic population from the Jiangjialiang site, North China[36]; Iron Age populations not related with the 'Scythian world" (black triangles): Xiongnu-Xiongnu population from Mongolia and Transbaikalia[12,37,38]; Taojiazhai-Taojiazhai site, Qinghai, northwestern China (1900-1700 YBP)[39]; Dondhu-Donghu population from Jinggouzi site, Inner Mongolia, northern China (~2500 YBP)[40].https://doi.org/10.1371/journal.pone.0204062.g003 ...
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Early nomads in the Eurasian steppes since the beginning of the 1st millennium BC played a key role in the formation of the cultural and genetic landscape of populations of a significant part of Eurasia, from Eastern Europe to Eastern Central Asia. Numerous archaeological cultures associated with early nomads have been discovered throughout the Eurasian steppe belt. The Tagar archaeological culture existed in the Minusinsk basin (Sayan Mountains, Southern Siberia, Russia) in the northeastern periphery of the Eurasian steppe belt from the 8th to 1st century BC during the pre-Scythian, Scythian, and Early Xiongnu-Sarmatian periods. In this study, we evaluated mtDNA diversity in the Tagar population based on representative series (N = 79) belonging to all chronological stages of the culture. The Tagar population had a mixed mtDNA pool dominated by Western Eurasian haplogroups and subgroups (H, HV6, HV*, I, K, T, U2e, U4, U5a, and U*) and, to a lesser degree, Eastern Eurasian haplogroups (A*, A8, C*, C5, D, G2a, and F1b). The Tagar population showed a similar mtDNA pool structure to those of other Iron Age populations representing the “Scythian World.” We observed particularly high similarity between the Tagar and Classic Scythians from the North Pontic region. Our results support the assumption that genetic components introduced by Bronze Age migrants from Western Eurasia contributed to the formation of the genetic composition of Scythian period populations in Southern Siberia. Another important component of the Tagar mtDNA pool was autochthonous East Eurasian lineages, some of which (A8 and C4a2a) are potential markers of the westward genetic influence of the eastern populations of the Scythian period. Our results suggest a genetic continuity (at least partial) between the Early, Middle, and Late Tagar populations.
... A variant characterised by substitutions 16223-16298-16327, observed in one Xiaohe individual, is found widely in present-day Eurasia, with the highest frequency in central/ eastern Siberia. It also been detected in a number of ancient individuals, three from Neolithic central Siberia [43], one from northeast Siberia (3600 yBP) [52] , six from northeast Europe (3500yBP) [37], twelve from the Bronze Age West Siberian Plain [53] , one from southern Xinjing(2800- 2011yBP) [54] and four from late Neolithic northwest China [55]. Haplotype 16129-16223-16298-16327 is found mainly in currently northeast, central and south Siberian populations, in Mongolia and central Asia. ...
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The Tarim Basin in western China, known for its amazingly well-preserved mummies, has been for thousands of years an important crossroad between the eastern and western parts of Eurasia. Despite its key position in communications and migration, and highly diverse peoples, languages and cultures, its prehistory is poorly understood. To shed light on the origin of the populations of the Tarim Basin, we analysed mitochondrial DNA polymorphisms in human skeletal remains excavated from the Xiaohe cemetery, used by the local community between 4000 and 3500 years before present, and possibly representing some of the earliest settlers. Xiaohe people carried a wide variety of maternal lineages, including West Eurasian lineages H, K, U5, U7, U2e, T, R*, East Eurasian lineages B, C4, C5, D, G2a and Indian lineage M5. Our results indicate that the people of the Tarim Basin had a diverse maternal ancestry, with origins in Europe, central/eastern Siberia and southern/western Asia. These findings, together with information on the cultural context of the Xiaohe cemetery, can be used to test contrasting hypotheses of route of settlement into the Tarim Basin.
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