VIII.20. Trans-Urals forest

VIII.20.1. The Seima–Turbino phenomenon

The Seima–Turbino inter-cultural network (main finds ca. 1900–1600 BC) is associated with materials present in the Abashevo, Sintashta–Petrovka, Taskovo–Loginovo (on the Middle and Lower Tobol and Middle Irtysh), Samus (on the Upper Ob), Krotovo (forest-steppe of the Middle Irtysh to the Baraba steppe on the Upper Ob), Elunino, and Okunevo cultures. This expansion through Eurasian forest and forest-steppe societies roughly corresponds to the expansions of the Srubna–Andronovo horizon through the Eurasian steppes (Carpelan and Parpola 2001).

While the Okunevo culture belongs to the Early Bronze Age (ca. 2250–1900 BC), most other cultures date to a later period, during the Pre-Andronovo horizon (ca. 2100–1800 BC). The better quality of tin–bronze proper of Seima–Turbino objects makes the source of both copper and tin probably central Asian ores (e.g. Upper Irtysh–Bukhtarta area of tin, copper, and gold ores), which isapart from the knives with depicted mountain sheep and horses typical of the eastwhy it has been traditionally considered an east–west movement of objects, and potentially of people (Carpelan and Parpola 2001).

Seima–Turbino metalsmiths were the first to regularly use a tin–bronze alloy, and were masters of lost-wax casting (for decorative figures on dagger handles) and thin-walled hollow-mould casting (for socketed spears and hollow axes). Nevertheless, local Okunevo and Afanasevo metallurgy of the Sayan–Altai area is primitive, and it is unlikely that they developed the advanced technology of casting socketed spearheads as one piece around a blank. On the other hand, spearheads of this type appear first in the Caucasus ca. 2000 BC, diffusing early to the Middle Volga–Kama–southern Urals area, where “it was the experienced Abashevo craftsmen who were able to take up the new techniques and develop and distribute new types of spearheads.” The presence of specific animals seems to be a local development, since Seima materials on the Oka river depict European elk types (Carpelan and Parpola 2001).

Sintashta socketed spearheads were made by bending a bronze sheet around a socket form and then forging the seam, while Seima–Turbino types were made by pouring molten metal into a mould that created a seamless cast socket around a suspended core, making a hollow interior, which necessitated tin–bronze rather than arsenical bronze (Anthony 2007). This use of tin–bronzes, of hollow-mould casting method, and of the lost-wax casting technique were probably learned from BMAC, which is probably explained by the exploratory movements of Abashevo, Sintashta, Srubna, and Andronovo settlers into the tin mining sites of the Zeravshan Valley.

The proportion of tin–bronze, arsenical bronze, and pure copper in Seima–Turbino materials from Europe also speaks in favour of a western origin of the material culture and regional adaptations to ore sources. The presence of pure copper in the Altai supports the presence of Abashevo migrants from the Urals, not yet mining arsenical copper. The main ores for Abashevo metal production were on the Volga–Kama–Belaya area sandstone ores of pure copper, and more easterly Urals deposits of arsenical copper. The Abashevo people, expanding from the Don and Middle Volga to the Urals, developed their metallurgy in the Volga–Kama basin (pure copper) and then moved to the east, where they produced harder weapons and tools of arsenical copper. Further to the south, they contributed to the Sintashta society in a territory richest in copper in the whole Urals region (Parpola 2013).

Arsenical copper was probably connected to the Tash–Kazgan deposits situated on the upper reaches of the Ui River in the southern Trans-Urals, which were transported westward over the low-lying Ural range, for about 250–300 km through the mountains. The rather high variation of arsenic concentration suggests that the Tash–Kazgan ore could be smelted on-site or on the Cis-Urals settlements and then transported to Abashevo and Sintashta areas (Koryakova and Epimakhov 2007).

The finding of Abashevo-like pottery in tin miners’ camp at Karnab on the lower Zeravshan, together with contemporary Sintashta-like pottery at Gonur, points to competing exploratory movements including contact and trade from forest-steppe and steppe cultures in Central Asia in look for tin ores near BMAC sites ca. 2100–2000 BC (Anthony 2007).

The Seima–Turbino phenomenon probably shows, therefore, the connection of areas to the west and east of the Urals in a network created by Abashevo settlers expanding into West Siberia through the forest-steppe and forest regions (Figure 48). Supporting this common expansion is the appearance of similar flint projectile points in Seima–Turbino and Sintashta graves, as well as objects of Petrovka origin, and the contemporary Andronovo expansion through the steppes (Anthony 2007).

Figure 48. Formative phase of the “Eurasian Metallurgical Province”, with the distribution of the Abashevo, Sintashta and Petrovka cultures (the map does not show Petrovka’s wider extension to southern Central Asia, Turkmenistan, Tajikistan, Uzbekistan), and the finds of the Seima-Turbino trade network. (After Parpola 2015: 57 Fig. 7.3, based on Chernykh 2007: 77). Image from Parpola (2018).

VIII.20.2. West Siberia

The Middle Bronze Age Pre-Andronovo cultural horizon of the beginning of the 2nd millennium BC in south-western Siberia was constituted by all of the forest-steppe and southern forests from the Trans-Urals to the Yenisei River, which were occupied by several cultures and groups: Tashkovo (Middle Tobol), Loginovo (Middle Ishim), Odino, Krotovo, and Elunino (the Middle Irtysh, Upper Ob’, and Altai areas), as well as the Samus’ culture (Tom–Chulym rivers) (Koryakova and Epimakhov 2007).

They display clear evidence of the diversified economy contributed by the productive and non-productive branches. Their similar pottery morphology and decoration shows alternating holed and combed motifs that cover the entire pot’s surface and reach back to an earlier epoch when they were dominant. These cultures remained rather distinctive and were only slightly touched by steppe influence (Koryakova and Epimakhov 2007).

The Tashkovo and Krotovo are more significant, contributing to later cultural formations in western Siberia, with the latter continuing into the Late Bronze Age. The Tashkovo culture is represented by villages with a circular or semi-circular layout, of ten to twelve houses on the banks of small rivers or lakes, whereas Krotovo shows open dispersed settlements with one- or two-chamber houses of the semisubterranean type and flat burial grounds (Koryakova and Epimakhov 2007).

The eastern Krotovo–Elunino territory (Middle Irtysh and Upper Ob’) shows flat burial grounds situated on elevated riverbanks or terraces, with sprinkled ochre, burials of separated skulls or skeletons without skulls, collective multi-layered burials, and secondary fractured burials. Some graves show rich metal objects, among which are some of Seima–Turbino type (Koryakova and Epimakhov 2007).

The arrival of Andronovo-like cultural horizon in the forest-steppe and southern forest regions replaced these cultures or displaced them to the north and east into the taiga.

VIII.20.3. Andronovo-like cultural horizon

Alakul and Fëdorovo material culture appear in the forest-steppe and forest zones of western Siberia, toward the northern taiga, evidenced by numerous settlement sites and decorated ware. They appear as stockbreeders, metallurgists, hunters and fishers, representing a cultural transformation of the vast area east of the Urals. These syncretic cultures are known as “Andronoid” or “Andronovo-like”, with apparently stronger connections to Fëdorovo. The horizon is composed of several cultures, including Cherkaskul (middle and southern Trans-Urals), Pakhomovo (Middle Irtysh and Tobol), Suzgun (taiga area of the Middle Irtysh), and Elovka (forest area of the Ob’-Irtysh river basin) (Koryakova and Epimakhov 2007).

All these cultures and smaller groups show similar flat-bottomed pottery with high or middle shoulders, short necks, and compact decoration consisting of alternating geometric motives with recognisable forest “images” and “Andronovo spirit”. They share similarities with Cis-, Trans-Urals, and eastern regions of western Siberia. The appearance of Andronovo-type geometric design into the local pottery reduced the area of the local hole and combed ornamental scheme, which still covered the major part of the taiga and Circum-Arctic area. Metallurgy is present, but there is a decrease in the number of metal objects, probably due to the absence of ores. Population density probably remained low, limited by the capacity of the forest ecological niches, with communities concentrated in clusters along rivers (Koryakova and Epimakhov 2007).

The Cherkaskul tradition (ca. 1850–1500 BC) has probably a direct origin in the expansion of Abashevo with the Seima–Turbino phenomenon over the area of the previous Late Garino–Bor and Sayat culture (ca. 2500–1900 BC) of hunter-gatherers, in the middle and south Trans-Urals, with Kazan as its western neighbour. The spread of Cherkaskul materials is thus closely associated with the Seima–Turbino phenomenon, and with the spread of the Fëdorovo tradition of the Andronovo horizon (see above Figure 47), probably as a northern variant directly linked to Abashevo, and not to the Sintashta culture from the steppes (Parpola 2018).

Cherkaskul sites are mainly found concentrated in the southern forest and northern forest-steppe of the Ural Mountains, with some findings as far south as the steppe zone. Cherkaskul materials are also found in thick concentrations on the southern forest and northern forest-steppe of central Asia, including the Upper Irtysh, Upper Ob’, and Upper Yenissei rivers, close to the Altai and Sayan mountains. Cherkaskul pottery appears frequently along with Fëdorovo types, and many sites produced pottery mixing both types. About half of the bone finds in Cherkaskul sites come from hunted animals, attesting to a predominantly hunter-gatherer population (Parpola 2018).

Its pottery includes the frequent use of carpet design in ornamentation, in common with Fëdorovo types, but is denser and more sophisticated than the Fëdorovo tradition, with flat-bottomed pots having a smooth and pleasing profile. In the forest, houses show shallow basements with rectangular pits as grounds for small frame-pillar wooden constructions (ca. 22–50 m2), showing internal hearths. Houses had corridor-like entrances. In the forest-steppe and steppe, houses were larger (ca. 100–200 m2) with deeper basements (Koryakova and Epimakhov 2007).

Settlements show evidence of a stable and settled life, with tolls for hunting, fishing, and bones of domestic animals, including bone dice, and remains of a developed metallurgy. Funerary tradition includes small kurgans with stone fences and mounds, covering individual burials with traces of cremation in the southern area; and inhumation in shallow pits in contracted left-sided position and modest assemblages in the northern and western regions. From the mid–2nd millennium on, different cultures without precise radiocarbon dates evolve from the previous expansion of Cherkaskul (Koryakova and Epimakhov 2007).

The Pakhomovo groups constitute the southern part of the Andronovo-like complex, and are located in the northern forest-steppe and forests of the Tobol-Irtysh basin, with pottery similar to Fëdorovo in morphology, decoration, and manner of surface treatment. Pots covered by monotonous ornaments of the forest style with various holes and figured stamps are common. Settlements are not large (ca. 4,500 m2 in average), with varied large rectangular houses (ca. 100 m2), wooden built with vertical pillars supporting a pyramidal roof, which include fixed hearths and wooden constructions for economic use. Tools related to hunting, fishing, metallurgy, and metal objects evidence their mainly fishing and hunting activity complemented by cattle and horse breeding (Koryakova and Epimakhov 2007).

The Suzgun groups occupy the area to the north of Pakhonovo, partially overlapping it in the northern Isim and Irtysh forest-steppe. Settlements are located on high promontories of the Irtysh River valley and on low fluvial terraces, with the environment conditioning the economy. A wooden walled enclosure with rectangular houses formed with vertical pillars in perpendicular rows and corridor-like entrances is possibly the model settlement. Mass ritual actions connected to sacrifices and common eating of oblational food are found in common sanctuaries (Koryakova and Epimakhov 2007).

Several individual inhumations and collective burials are found in cemeteries, where dead were buried in the extended supine position, with some apparently Pre-Andronovo and some Andronovo-like customs found in the funerary ritual. Bronze metalworking seems to be a part of ritual actions, and the most common artefact found is pottery, with flat and round bottomed pots with well pronounced profiles decorated with geometric motifs. The subsistence economy was diverse, with hunting and fishing being an essential part, but cattle being also dominant, and horse occupying the second position (Koryakova and Epimakhov 2007).

VIII.20.4. Mezhovska–Irmen cultural horizon

The Mezhovska–Irmen cultural horizon (ca. 1500–800 BC) involves a group of cultures with common stylistic similarities and local differences, in the forest-steppe area on both sides of the Ural Mountains, from the middle Kama and Belaya rivers to the Tobol river in western Siberia, with sites reaching up to the Altai (Suppl. Fig. 13). They emerged from the previous Andronovo-like cultural complex of the forest-steppe and southern forest of western Siberia, with influence coming from cross-stamped cultures of the north, and include Mezhovska and Irmen cultures around the Urals, the Sargary–Alekseevka culture to the south, and the Karasuk culture to the east (Koryakova and Epimakhov 2007).

Mezhovska sites were present in the forest and forest-steppe zones on both sides of the Urals, including the forest-steppe region from the Belaya bend to the Middle Kama up to the Chusovaya river (in the Cis-Urals), the Trans-Urals, the Ishim–Irtysh area. The Irmen cultures were distributed in the Ob-Irtysh forest-steppe, with its influence found in Sargary–Alekseevka culture in northern Kazakhstan (Koryakova and Epimakhov 2007).

All settlements are small or middle-sized open settlements (200–300 individuals), mainly seasonal camps, but also stable long-term habitation settlements. They are situated on river terraces or lake or river promontories, possibly with defensive constructions. The largest pillar-frame structures are found to the south, related to the Sargary culture, while smaller houses (ca. 100 m2) are proper of the forest zone. There were vertical pillar-frame constructions, and constructions with deep basement and horizontal frame in its low part. Fireplaces and storage pits are found inside houses, and a corridor-like entrance usually faces the water (Koryakova and Epimakhov 2007).

Subsistence economy in both the Cis-Urals and Trans-Urals depends on the specific ecological niche. It usually includes wild species (up to 15%), with a smaller percentage for ‘fur’ animals; horse and cattle (ca. 30%), and a small percentage of sheep (ca. 13%), as well as fish. Food producing branches were more prevalent in the south, and the high proportion of horses compared to previous periods is probably related to their ability to forge in the winter. Metallurgy was probably not using local ores (as in the succeeding Itkul’ culture), and it was most likely based on domestic needs (Koryakova and Epimakhov 2007).

Characteristic is the decoration of the pottery, usually covering the neck and shoulder, with carved (fretted) elements becoming popular, contrasting with the previous comb–stamped techniques of the region. Ornament patterns are simple, and pottery shapes include pots of globular bodies with short straight or turned up necks, and cans and korchags (large earthenware pots). Bronze objects are represented by massive tools, like celts–axes of the Cimmerian type, gouges of Derbeden type, sickles of Derbeden and Kataisk types, daggers of Kardashinsky and Cimmerian types, as well as double-edged knives with smooth passage to tengue, awls, and needles. Bone and stone arrowheads are also numerous (Koryakova and Epimakhov 2007).

There seems to be a kurgan burial tradition, using stone in the erection of mounds, with one to three inhumations for kurgan, and the dead in extended supine or side position. Secondary burials appear on the ancient surface. Graves are shallow with modest wooden arrangement. Animal bones, mainly cattle, represent remains of the funeral feast, and assemblages include pottery, metal daggers, spearheads, knives, and ornaments (Koryakova and Epimakhov 2007).

The eastern part of the Mezhovska–Irmen horizon is formed by the Irmen culture, with material culture intermediate between Mezhovska and Karasuk. Settlements include traditional large house buildings in variable open villages, small camps and, in a later phase, fortified sites. Houses have several hearths and traces of domestic activities, including animal stabling during the winter, and there are separate houses with structures of economic function. Subsistence economy is also based on livestock breeding, with their bones prevalent over wild animals, and only traces of cultivation. Metallurgy developed depending on the sites and raw material resources (Koryakova and Epimakhov 2007).

Pottery includes large flat-bottomed massive pots, and (proper of burials) smaller pots and jars with flat or round bottoms. Ornamentation includes incised techniques, sometimes accompanied by combed stamps and ‘pearls’, and Andronovo-like motifs appear in funerary pottery. Cemeteries include kurgans with multiple individual burials, with inhumations in crouched right-sided position with southern direction and orientation. Graves are shallow, and wooden frames furnish them. Assemblages consist of metal goods (nail-like pendants, earrings, sewn plaques) and pottery. Collective burials are rare (Koryakova and Epimakhov 2007).

The Sargary culture emerged from Fëdorovo and Cherkaskul traditions in the forest-steppe between Tobol and Irtysh, in northern and central Kazakhstan. It formed part of the “horizon of cultures of the Valikova pottery tradition” (Chernykh 1992), which comprised cultures from the Don–Volga–Ural steppes (late Srubna), southern Urals and Trans-Urals, and Kazakhstan (Sargary, Trushnikovo, Dandybai–Begazy, Amirabad), characterised by poorly ornamented flat-bottomed pottery with clay rollers stuck around the shoulder or neck. Other similarities involved metal artefacts, economic structure, and funerary ritual.

In the Dandybai–Begazy culture, mausoleums were constructed especially for people of high social status. The characteristic pottery of globular form and very small bottoms and cylindrical necks, often with polished surfaces and black, yellow, or red colour, and both Sargary and this culture probably formed a unity with a core area in central Kazakhstan, where the centres of metallurgy lied. In the Trans-Urals region, Sargary material culture is found with Mezhovska, and some Meshovksa finds contain Sargary material, probably from the 13th century BC on (Koryakova and Epimakhov 2007).

Sargary settlements include large territories (ca. 20,000 m2) with semisubterranean buildings with deep basements and up to several dozen large houses, and smaller settlements (1,000–2,500 m2) with up to fifteen houses, yielding a similar number of finds, suggesting that they were inhabited seasonally. Houses are rectangular, frame-pillar constructions with a floor deepened into the ground, and are placed freely along the river bank. Metal tools are numerous (Koryakova and Epimakhov 2007).

Subsistence economy reveals a pattern connected to the steppe area of eastern Europe, with domestic animals predominating, first cattle, then horse and sheep, with likely yearly cycle herding practices. The first trace of agriculture in the region is associated with this culture. Small cemeteries comprising up to three mounds and solitary kurgans of earth and stone are placed on high ground of the initial riverbank and far from river streams, unlike in the preceding period. The dead are placed in contracted position on their side, and assemblages are modest. Funerary rituals are complex and varied, and kurgan ritual and inhumation were reserved for some people, with society tending to atomisation in the forest-steppe, and to concentration in larger settlements in the south (Koryakova and Epimakhov 2007).

The Karasuk culture (ca. 1400–900 BC), genetically at least partly derived from Fëdorovo, flourished around the upper Yenissei, Mongolia and the Ordos region of China. It probably came into being as a result of a migration of different people from the southeast, from the periphery of Shang period China. The beginning of the Karasuk period also marked the return of some Okunevo traditions that did not manifest themselves during the Fëdorovo period (Parpola 2018).

It preceded the transition of the LBA to the EIA Proto-Scythian period, when the use of saddled horse, composite bow, and the ‘animal style’ art became integral parts of the steppe life. Around 1000 BC, the Eurasiatic steppes became uniform culturally from Mongolia to Hungary (see §VIII.18.4. Scythians and Sarmatians), and for a thousand years East Iranian languages were spoken in the region (Parpola 2013). Among the pictorial tradition of petroglyphs in Andronovo-related groups, those of burials of the Karasuk culture in southern Siberia and Kazakhstan represent the latest tradition (Novozhenov 2012).

VIII.20.5. Itkul’–Gamayun and Sargat

The Gamayun and Itkul’ cultures evolved (ca. 800–200 BC) from the Mezhovska cultural region in a narrow band (ca. 150 km wide) on the eastern slope of the Urals. Itkul’ constituted the main metallurgical centre of the Trans-Urals region during the Iron Age. They were in contact with the Ananyino and Akhmylovo cultures, which were the metallurgical centres of the western Urals, and neighboured the Gorokhovo culture (Parpola 2018).

The Gamayun culture is rather archaic in appearance, featuring open or fortified settlements and early fortified house-refuges (of ca. 600 m2), having solid wooden walls with a ditch and strengthened by an earthen bank, and a variable living space (ca. 40–400 m2). Open settlements include small stable villages consisting of several houses, and seasonal short-time hunting-fishing camps. Fortified villages, chiefly concentrated on the periphery of the occupied territory, were variable in size, and occupied the low hills and promontories along riverbanks, with smaller ones representing frontier-guard stations. Houses vary from small hovels and chums to stable pillar-framed wooden houses with one or two sections, and were also used for economic activities (Koryakova and Epimakhov 2007).

Characteristic of the Gamayun culture is the crosslike stamp ornamentation, which was found widespread among the massive of cultures with stamped crosslike ornaments in the southernmost taiga zone and northern forest-steppe, from the Trans-Urals to the Middle Ob’. These cultures emerged from the Late Bronze Age of the Lower Ob’ River forest area, represented by the Lozva–Atlym phase of the Late Bronze Age, when some population groups moved southward in search for game due to the humid conditions at the turn of the 2nd/1st millennium BC. Most migrating communities of the LBA and Gamayun sites show a rather small size (ca. 20–30 people), perhaps extended families or clans (Koryakova and Epimakhov 2007).

The Itkul’ culture represents thus the strict cultural continuation of Mezhovska in the region, and the initial relations with the Gamayun culture seems to be one of conflicts, based on the fortification of houses and on traces of destruction of Gamayun villages. Eventually, both communities formed a symbiotic system based on division of labour and specialisation. Local Itkul’ communities, having harnessed local mineral resources for metal production in a previous period, specialised in metallurgy, while Gamayun groups were hunters, fishers, and most likely miners (Koryakova and Epimakhov 2007).

Itkul’ sites were fortified villages or fortified metallurgical workshops, with the vast majority occupying high ground of rivers or lake terraces. Fortifications were simple, with a wall, moat, ditch, or grove, and bigger settlements took up very high topographic positions usually with an open-order defensive line. Buildings included houses, workshops, and structures connected with special productive functions. Houses were rectangular semisubterranean and surface-based buildings of a pillar-framed construction, and they were rather small, with a simple interior, including one or two hearths. Small working areas served for copper smelting, iron working, and most often metal processing, while mass production of metal was done on larger dwellings or outdoor (Koryakova and Epimakhov 2007).

Artefacts and structures evidence the intensive dedication of the sites to metallurgical activity, with specialisation mainly in bronze production with local mineral deposits. There is operational division involving metal smelting and casting in mountain clusters (and further divisions into full or limited cycle of production), and metalworking in the periphery. Their distinctive pottery includes round-bottomed pots, of chiefly horizontal proportions, and decorated with rather standard comb–stamped patterns, covering the pot’s upper third (Koryakova and Epimakhov 2007).

Itkul’ metallurgists had close relations with the Ananyino populations, which used in part Trans-Uralian ores for manufacturing various objects that diffused westwards. Closer connections existed with neighbouring cultures, with Itkul’ supplying regularly metal products to Gorokhovo and to the southern Siberian Sargat groups. More or less regular contacts existed also with The Upper Ob’ Bolsherechye culture, and with the southern Kazakhstan Saka culture (Koryakova and Epimakhov 2007).

To the east of the Itkul’ culture, up to the Ob river, the Nosilovo, Baitovo, Late Irmen, and Krasnoozero cultures (ca. 900–500 BC) developed, some of them in contact with the Akhmylovo culture of the Middle Volga. All these cultures of the forest steppe were later absorbed into the Sargat culture (Parzinger 2006),

The spread of the Suzgun culture with the Baraba trendmarked by the Baraba–Suzgun pottery, featuring slightly and well-profiled pots with a short throatprobably represents a wide-ranging population expansion in pre-taiga and taiga zones in the Irtysh basin. Later, the Berlik tradition expanded with migrants from the south, interacting in certain sitesas in the local Late Irmen cultural traditionwith Late Irmen people inhabiting the citadel of the settlement, and Berlik immigrants inhabiting the surrounding territory (Molodin, Mylnikova, and Kobeleva 2008).

The Sargat culture emerged from the Late Irmen tradition, succeeding the Sargary culture in its territory, comprising all cultural groups between the Tobol–Irtysh forest-steppe interfluve. Apart from the core settlers of the Mezhovska–Irmen horizon of the Final Bronze Age, intercommunity formation was completed by taiga settlers of the Lozva–Atlym LBA from the north, and southern influence from the Valikova pottery horizon and Arzhan phase of the Scythian and Saka confederation. These groups superimposed each other chronologically and territorially, as evidenced in the Chicha settlement. Open and fortified settlements are characteristic (Molodin, Mylnikova, and Kobeleva 2008).

The Gorokhovo cultural group of the Iset–Tobol area also resulted from a population of Sargary–Mezhovska roots, under the influence of climatic, economic, and social factors. With a clear cultural root in the Trans-Uralian forest-steppe, evidenced by its architecture, fortification system, and pottery, they probably adopted pastoral herding under the cultural and possibly political influence of the Saka confederation, evidenced by the funerary ritual. They formed part of a “forest-steppe–steppe system” that involved the Iktul’ metallurgical centre, South Urals nomads, and the Sargat culture (Parzinger 2006).

Both Gorokhovo and Sargat eventually developed (ca. 5th c. BC on) the “Golden Age” of the western Siberian forest-steppe, under Sargat dominance coming from the east, and increased influence from steppe nomads from the south. Fortified settlements demonstrate a more complex level of architecture, with large elite barrows including very big kurgans, analogous to those found in the steppes, and mainly individual inhumations. Seminomadic stockbreeding became the main subsistence economy. The expansion of Sargat may have caused an initial expansion of Gorokhovo settlers to the west, until their eventual integration under Sargat (Koryakova and Epimakhov 2007).

viii.20.1. Ugrians and Samoyeds

Seima–Turbino-related migrations through the Eurasian forest and forest-steppe zones in western Siberia reflect population movements from west to east, coinciding with the arrival of pastoralists in central Asia (Kılınç et al. 2018), which supports the traditional interpretation of Uralic expanding from west to east, originally with Abashevo-related groups, most likely associated with individuals of Eastern Steppe MLBA ancestry associated with the Andronovo-like cultural horizon (see §viii.17.1. Late Indo-Iranians). While Mezhovska is the best candidate for the original Proto-Ugric-speaking population, Karasuk has been traditionally proposed as the Pre-Samoyedic-speaking community (Parpola 2013). Both Mezhovska and Tagar samples can be modelled as almost completely of Eastern Steppe MLBA ancestry, in contrast with neighbouring groups like Pazyryk or Zevakino-Chilikta—traditionally considered East Iranian cultures—which show likely contributions from neighbouring steppe populations.

Individuals of the Mezhovska culture from the Kapova cave (one dated ca. 1500 BC) are part of the Western Steppe MLBA cluster, with one sample of hg. R1a1a1b1a2-Z280[68], and another of hg. R1b1a1b-M269 (Allentoft et al. 2015). Interesting is the presence of one outlier, with ancestry close to the later South-Eastern Iranian cluster, with increased Near Eastern ancestry, which suggests the potential emergence of this cluster in the Andronovo–Srubna complex, i.e. much earlier than suggested by the available samples. The position of Mezhovska samples is close to later Scythians from Samara, and also close to modern Finns, northern Russians, Early Sarmatians, Estonians, Mordovians, Lithuanians or Belarusians (Unterländer et al. 2017).

The Sargat culture is probably to be identified (at least partly) with the expansion of Proto-Hungarians, who around the 5th century BC “were caught up in a wave of migrations that swept the steppe… Migrating westwards, they settled between the Urals and the Middle Volga region”, staying in Bashkiria until ca. 600 BC, in the so-called Hungaria Magna of medieval sources (Suppl. Fig. 18). That Ugric peoples were horsemen is supported by the number of equestrian terms in Ugric languages, including the word for horse (Parpola 2018), and by the presence of horse riding equipment and horse bones in graves of Early Hungarian frequent riders, evidenced by their skeletal hip changes (Berthon et al. 2018).

Seven individuals of the Sargat culture from the Baraba forest (ca. 500 BC – AD 500) show five hg. N-M231, two hg. R1a-M420 (Bennett and Kaestle 2010), which is expected to be found in a late Ugric community integrating with Palaeosiberian peoples from the LBA cultures of the Lower Ob’ River forest area, likely expanding with hg. N-M231, and becoming integrated both in the Itkul’ culture through its association with Gamayun, and in Sargat through the gradual integration of migrants from the taiga region.

Given the sample of hg. N-M231 from the Mezőcsát Culture (ca. 900 BC) in Hungary (Gamba et al. 2014), belonging to expanding Cimmerians, and the Siberian ancestry expanding with Scythian groups (see §viii.18. Iranians) and later with Altaic peoples (see Suppl. Graph. 15 and Suppl. Graph. 17), it is conceivable that part of the N-M231 lineages integrated among Ugric and part of the Samoyedic peoples accompanied this and similar westward migrations, which is in line with some known shared traits of Uralic and Altaic (Kortlandt 2010).

Modern Ugric populations include the western Mansis, in the immediate Trans-Urals region, with more N-M231(xN1a1-M46) (ca. 60%) than N1a1-M46 (ca. 16%), R1a1a-M198 (ca. 8%), I-M170 (ca. 8%), R1b-M343 (ca. 4%), or J-M304 (ca. 4%). Eastern Khants, on the other hand, show more N1a1-M46 (ca. 49%) than N-M231(xN1a1-M46) (ca. 31%) or N1a1-M46 (ca. 16%), but they also show R1b-M343 (ca. 11%), R1a1a-M198 (ca. 6%), I-M170 (ca. 8%), or J-M304 (ca. 4%) (Tambets, Yunusbayev, et al. 2018). Khants and Nenets share similar ancestry, showing a close affinity with Selkups (in turn intermingled with Yeniseian-speaking Kets), too, which suggests the origin of their admixture in the acculturation of related Siberian peoples (Karafet et al. 2018).

Hungarians, on the other hand, show similar contributions of hg. R1a-M420 (ca. 21-60%), R1b-L23 (ca. 15–20%) and I-M170 (ca. 11-26%), with intermediate frequency of J-M304, E1b-P177 and G2a-P15 (ca. 5–15%), and much lesser N-M231(xN1a1-M46) and N1a1-M46 (ca. 1-6%) (Semino 2000; Csányi et al. 2008; Pamjav et al. 2011; Pamjav et al. 2017). Among Hungarian conquerors, the distribution is similar, with available samples showing among twelve males from Karos-Eperjesszög (AD 900–1000) at least two R1b1a1b-M269, two I2a-L460 (Neparáczki et al. 2017); possibly[69] two N1a1-M46 (Csányi et al. 2008), one from Szabadkigyos-Palliget (ca. AD 950) and another from Ormenykut (ca. AD 975–1000); and among early Hungarians interred in Saint Stephen Basilica, Székesfehérvár (AD 12th c.), three of hg. R1a-M420, including King Béla III of the Árpád dynasty, of hg. R1a1a1b2a2a1d7-YP451 (YP499-), common in modern populations of the northern Caucasus, among Karachays and Balkars; two of hg. R1b-M343, one J1-L255, and one E1b1-P2 (Olasz et al. 2018).

The early mixture found among early Hungarian elites and among modern populations points to the likely domination of a minority of Magyar clansprobably already admixedover a majority of the population of the Carpathian Basin composed of local haplogroups. The finding of haplogroup R1a1a1b1a2-Z280 (ca. 14-15%) and lesser R1a1a1b2-Z93 (ca. 1%), proper of eastern European regions, suggests the arrival of some R1a1a1b1a2-Z280 lineages with Magyar tribes, in contrast to the typical West Slavic subclades of the area, mainly R1a1a1b1a1a-M458 (ca. 7-20% among Hungarians)[70]. Further support of the spread of Hungarian conquerors from the Urals region is found in the shared mitogenomes with the ancient populations (AD 6th – 10th c.) of the Volga–Ural region (Szeifert et al. 2018).

Seven individuals of the Karasuk culture from Arban, Sabinka and Bystrovka (ca. 1530–1260 BC) form a wide cluster reaching from Western to Eastern Steppe MLBA, including contributions of AP ancestry compatible with their admixture with Cis-Baikal populations, with one particular outlier clustering closely with Khövsgöls in northern Mongolia (see §viii.20.3. Turkic peoples and Mongols). Reported haplogroups include two R1a1a1b2a2-Z2124, and one Q1a2a-L712[71] (Allentoft et al. 2015).

The Tagar culture (ca. 1000–200 BC) largely continues the traditions of the Karasuk culture in the Minusinsk basin of the Upper Yenissei. This area is considered the homeland to Proto-Samoyedic, based on the Bulghar Turkic loanwords, and thus the Tagar culture probably represents the expansion of the language (Parpola 2018).  Sampled Tagar individuals (probably ca. 9th c. BC) display increased EHG ancestry compared to other Inner Asian Scythian groups, with unequal ancestry contributions of Steppe MLBA (ca. 83.5%), WSHG (ca. 7.5%), with additional ANE ancestry (ca. 9%), and clear differences of hunter-gatherer ancestry sources with other sampled Sakas, which likely formed a confederation of different peoples (de Barros Damgaard, Marchi, et al. 2018). This is compatible with their origin in the eastern European forest zone, and reported haplogroups include two hg. R1-M173, and one R1a1a1b2a2-Z2124[72].

Modern Samoyedic peoples include the southern Selkups, with a majority of ‘eastern’ haplogroups (ca. 58-66%) like P-P295, Q-M242, R-M207 (xR1a1a-M198, xR1b-M343), or R2-M479, but also more R1a1a-M198 (ca. 14-19%), than other Samoyedic peoples, with lesser R1b-M343 (ca. 6-7%), I-M170 (ca. 0–7%), N-M231(xN1a1-M46) (ca. 7%), N1a1-M46 (ca. 0–2%), or C2-M217 (ca. 2-5%). Among northern groups, Nenets to the west show N-M231(xN1a1-M46) (ca. 57%), N1a1-M46 (ca. 41%), while Enets in central regions show N-M231(xN1a1-M46) (ca. 78%), R1b-M343 (ca. 11%), and N1a1-M46 (ca. 11%). Nganasans to the east, the most recent Palaeosiberian group to adopt Samoyedic languages (Dolgikh 1960, 1962), show N-M231(xN1a1-M46) (ca. 92%) and N1a1-M46 (ca. 3%), and an elevated “Siberian component” which has the highest frequency in three Karasuk samples (Karafet et al. 2018).

Eastern and western Circum-Arctic nomads show a prevalence of N1a1-M46 lineages, which appear in the western area as N1a1a1a2-B211 lineages (formed ca. 5400 BC, TMRCA 1900 BC) among Khanty and Mansi peoples, and in the east among some Nganasans in contact with Yukaghirs (see §viii.20.2. Yukaghirs). Most Nganasans show a deeper N1b-F2905 subclade (formed ca. 16000 BC, TMRCA ca. 13700 BC), though, also found in lesser proportions among Dolgans, Evenks, Evens, as well as in south Siberian Tofalars, Khakassians, Tuvinians, and Shors (Fedorova et al. 2013).

Central Siberian peoples show a majority of N1a2b-P43 lineages (formed ca. 6800 BC, TMRCA ca. 2700 BC), which may suggest an expansion of N1a-F1206 lineages precisely from this central Siberian area. Its western branch N1a2b2-Y3195 (TMRCA ca. 2200 BC) is found in the Cis-Urals region, with Permic peoples and Volga Finns, while its eastern branch N1a2b1-B478/VL64 (formed ca. 2700 BC, TMRCA ca. 1300 BC) is found in central Siberia and East Asia, and N1a2b1b1-B170 in particular coincident with the interaction east of the Urals (Ilumae et al. 2016; Karafet et al. 2018).

viii.20.2. Yukaghirs

Based on proposed Indo-Uralic community (Kortlandt 2002; Kloekhorst 2008; Hyllested 2009), a macro-family formed by Indo-European, Uralic, and Yukaghir is quite likely to have migrated back from the Trans-Urals region with hunter-gatherer pottery (see §ii.3. Indo-Uralians). Remnant populations from this migration probably include two Sintashta outliers of Khvalynsk Eneolithic-like ancestry (ca. 2000–1650 BC), one R1b1a1-P297, the other R1b1a1a-M73 (Narasimhan et al. 2018), both lineages probably related to westward migrations through the Urals. Similarly, there is a possible sample of hg. R1b1a1-P297 in Darra-e Kūr, Afghanistan (ca. 2700 BC), with fully Iran Neolithic-like ancestry. The presence of hg. R1b1a1a-M73 lineages among diverse central Asian populations, in particular among Turkic-speaking groups (see below), also suggest potential remnant groups of hg. R1b1a1-P297 in West Siberia during the Neolithic.

Nevertheless, the continuous population expansions and replacements in Siberia have obscured the potential migration routes for Yukaghirs, whose language has been recorded only recently. Ancient DNA sampled from historical central Siberian peoples show certain discordances with a simplistic model of a macro-Yeniseian community in the taiga zone and southern Siberia up to the Altai–Sayan zone (Kim et al. 2018), and this complexity is in turn compatible with ancient Eurasian and Indo-Uralic population movements through Siberia . Modern Yukaghirs show among ten sampled individualsin a similar distribution to Tungusic Evenks and Evens—four hg. C2-M217, three N1a-L279, two R1a-M420, as well as (not present in Tungusic peoples) one I2a-L460 and one O-M175 (Fedorova et al. 2013).

Four samples of the Bronze Age Glazkovo culture, from the Lake Baikal (ca. 18th–13th c. BC), one of hg R1-M173, show ancestry compatible with modern north-east Siberian populations, compatible with their later described spread to the north (de Barros Damgaard, Marchi, et al. 2018). A Medieval individual near the Yana river (ca. AD 1350) shows hg. N1a1a1a1a4a1-M1993 (formed ca. 1700 BC, TMRCA ca. AD 450), and falls within the widespread Neosiberian cline evidenced by a recent sample from Ust’Belaya (ca. AD 1300) near Lake Baikal (Sikora et al. 2018). The prevalent presence of N1a1a1a1a4-M2019 (formed ca. 4400 BC, TMRCA 1700 BC) in central Siberia and Yakutia, and at lower frequencies in Khants and Mansis (Ilumae et al. 2016), is most likely the result of the expansion of Yukaghir-related languages with acculturated Palaeosiberian lineages.

The expansion of Yukaghir was probably then coincident with Bronze Age migrations, likely continued to the east with N1a1a1a1a4a-M1993 during the Iron Age–Early Middle Ages, as Ugrians and Samoyeds expanded to the north (see §viii.20.1. Ugrians and Samoyeds). This later expansion probably displaced populations of mainly N1a1a1a1a3b-B202 lineages (formed ca. 2800, TMRCA ca. 600 BC) to the extreme north-eastern Siberia, where they retained their Chukotko-Kamchatkan languages, supported by their prevalent N1a1a1a-L708 lineages (ca. 92%). Turkic-speaking Evenks cluster together and overlap with Yukaghirs, and both in turn cluster closely to Nganasans from the Taymyr Peninsula and to Chukchi-speaking Koryaks, revealing a complex acculturation of different East Asian peoples in recent times (Karafet et al. 2018). The wide cluster formed by modern Yukaghirs, including southern Samoyedic speakers in the west and Chukotko-Kamchatkan speakers in the east (see Suppl. Graph. 15) further supports the relatively recent expansion of Yukaghir into the Circum-Arctic region.

viii.20.3. Turkic peoples and Mongols

Cultures succeeding Afanasevo in the Altai region show different lineages and the partial resurgence of WSHG ancestry coupled with different Afanasevo-, Steppe MLBA-, AP-, and AEA-related contributions, suggesting the emergence of different local West Siberian populations (Hollard et al. 2018): the Chemurchek culture (ca. 2300–1800 BC), with one hg. C-M130; the Okunevo culture (ca. 2300–1800 BC), with one hg. R1b1a1b1-L23, three likely N-Z4813, and three likely Q1b1a-L54; the Elunino culture (ca. 2300–1700 BC), with one hg. Q-M242, and one Q1b-M346; the Munkh-Khairkhan culture (ca. 1700–1400 BC), with two N-M231 lineages; and the Sagsai culture (ca. 1400–800 BC), with four hg. Q1b1a-L54, four R1a1a1b2-Z93, and one C-M130.

MBA sites Takhilgat Uzuur and Tsagaan Asga in the Mongolian Altai Mountains also show apparent succeeding replacements from hg. Q1b1a-L54 in the 3rd millennium BC to R1a1a1b2-Z93 in the late 2nd millennium BC, to C-M130 in the early 1st millennium BC  (Hollard et al. 2014).

To the east, near Lake Baikal, a resurgence of AP ancestry (up to ca. 50%) coupled with Afanasevo-related ancestry (ca. 10%) found in EBA individuals is continued during the LBA. Samples of the Deer Stone-Khirigsuur Complex from Khövsgöl in northern Mongolia (ca. 1200–800 BC) show a slightly higher contribution of AP ancestry evidenced by a ‘northern’ shift in the PCA, similar to Karasuk or Okunevo samples (Jeong, Wilkin, et al. 2018). In terms of haplogroups, there are ten probably Q1b-L56, possibly all of Q1b1a-L54 subclade Q1b1a3-L330 (formed ca. 16000 BC, TMRCA ca. 5900 BC), with an estimated successful dispersal of these subclades starting in the Mesolithic ca. 6300 BC from central Asia (Grugni et al. 2019), possibly initially accompanying Dene-Yeniseian-related languages (see §v.8. Palaeosiberians).

A small contribution of Steppe MLBA ancestry (ca. 4-7%) suggests close contacts with Andronovo-related peoples. In particular, the presence of one outlier (ca. 1130–900 BC), of hg. R1a1a1b2a2a1-Z2123, is consistent with the appearance of admixed forest-steppe populations of Eastern Steppe MLBA ancestry like Karasuk in the Altai (ca. 1200–800 BC). Supporting these contacts of Karasuk with eastern Asian steppes is also the Karasuk outlier and the sample of hg. Q1a2a-L712. This points to the spread of pastoralism in the region mediated by acculturation and exogamy more than population replacement (Jeong, Wilkin, et al. 2018). Another sample (ca. 1420-1130 BC) of hg. N1a1a-M178, mtDNA U5a2d1, suggestslike the different groups succeeding Afanasevo in the Altai regionthe acculturation of northern Eurasian communities with post-Neolithic expansions.

To the south-east of Lake Baikal, in the Houtaomuga site from Manchuria, there is genetic continuity from ca. 10000 BC until the Iron Age, but haplogroup N1b1-CTS582, found in the Early Neolithic (ca. 5430–5320 BC), is replaced in the Bronze Age by C2b-L1373, which continues in the Early Iron Age (Ning 2018). Twelve Donghu individuals from the Jinggouzi site (ca. 770–476 BC) show C2a1a-F4032 lineages (formed ca. 12700 BC, TMRCA ca. 12300 BC), as do four samples from Xianbei (ca. AD 4th–10th c.) (Zhang, Wu, et al. 2018; Li et al. 2018). The expansion of the Donghu seems to have caused the expansion of C2-M217 lineages in the Trans-Baikal area.

The first Turkic-speaking community is usually identified with the Xiongnu confederation, with ancient Y-chromosomal data indicating a heterogeneous multi-ethnic cultural organisation, likely emerging initially from local East Asian groups to the east of the Tian Shan Mountains, who showed admixture with central steppe nomads. A more recent West Eurasian ancestry is found among western Xiongnu groups, with Central Sakas being the closest source for their admixture (de Barros Damgaard, Marchi, et al. 2018).

Sampled Xiongnu individuals (ca. 300 BC – AD 200) include hg. R1a1a-M198 and C2-M217 in Duurlig Nars, and hg. N1a1-M46 and Q-M52 and in Egyin Gol Valley (Kim et al. 2010; Petkovski 2006); two likely hg. O2a2b-P164 from Omnogobi, one early sample from the West Xiongnu in Khövsgöl (ca. 330 BC), of hg. R1b2-PH155, and one from an aristocratic burial in Arkhangai (ca. AD 1), likely of hg. R1b2b-PH200[73] (de Barros Damgaard, Marchi, et al. 2018).

The homeland of Turkic peoples is difficult to pinpoint based on scarce samples through wide temporal transects, due to the multiple population replacements in the central and eastern Asian steppes, forest-steppes, and forests, and to the linguistic data and complex ethnogenesis legends pointing to a composite grouping of diverse elements since the reconstructible stage of the language (Golden 1992).

Based on Indo-European (Iranian and Tocharian) and Uralic influences, Proto-Turkic is supposed to have been spoken ca. 1000 BC in some area from the Trans-Urals area to the Altai, with the forest zones of West Siberia being the most likely candidate based on the Indo-Iranian expansions through the steppes, as well as the Uralic expansions through the forest-steppe and forest regions. From there, the ancestors of the Turks migrated east into the Baikal area, where the Xiongnu confederation eventually emerged, and Huns later migrated to West Eurasia (Golden 1992).

The two Sintashta outliers of Khvalynsk Eneolithic-like ancestry, of reported haplogroups R1b1a1-P297 and R1b1a1a-M73, probably correspond to ancient populations of R1b1a1a-M73 lineages widespread from the west in the southern Urals up to the Lake Baikal in the east since the Neolithic, following a WSHG ancestry cline. This is compatible with the Mesolithic expansion of Eurasian through Inner Asia (see §ii.1. Eurasians), and the isolated development of Altaic in the Neolithic (see §v.8. Palaeosiberians).

Haplogroup R1b1a1a-M73 is reported with low frequencies among modern Siberian populations, such as Ugric and Samoyedic peoples, especially southern Selkups (Tambets, Yunusbayev, et al. 2018), and with increasing frequencies among south Siberian populations, in particular among Turkic-speaking Bashkirs (Jeong, Balanovsky, et al. 2018) and Teleuts near the Altai (Karafet et al. 2018), which suggests its presence among Turkic-speaking peoples before their expansion into the Trans-Baikal area and the creation of a community integrating diverse local populations.

The earlier emergence of Turkic-speaking peoples from the forming multi-ethnic groups in the Trans-Baikal area further supports that Turko–Mongolic, and not only Turkic, expanded from the west. The adaptation of horseback riding for more stressful and difficult activities such as warfare—characteristic of these nomadic groups—in the eastern steppes, started probably at the end of the Deer Stone–Khirigsuur Complex or slightly later, evidenced by findings from the Altai to the Baikal region, including the appearance of the bronze snaffle bit and innovations in equine dentistry (Taylor, Bayarsaikhan, et al. 2018).

The Huns, likely representatives of the earliest Turkic-speaking groups in Eurasia, emerged following minor male-driven East Asian gene flow into the preceding Sakas that they invaded, ca. 2,000 years ago. They displaced Iranian-speaking groups (ancestors of the Wusun and Kangju) to the south-east of the Tian Shan mountains, where they became isolated (see §viii.18. Iranians). Sampled nomads from the Kargaly in the Tian Shan region include two early ones (ca. 800–700 BC), one of hg. Q-M242, clustering with Siberian peoples (close to Yeniseians), one intermediate (ca. 425 BC), of hg. R1a1a1b2a2-Z2124, and a later one (ca. 35 BC), of hg. R1b1a1a1b-Y20750 (formed ca. 5300 BC, TMRCA ca. 3300 BC), a subclade of R1b1a1a-M73 (de Barros Damgaard, Marchi, et al. 2018).

This variability is also found later among Huns from Tian Shan (ca. AD 60–600), with three hg. R1a1a1b2a2-Z2124, two R1-M173, one hg. R1b2b-PH200, one N1a1a-M178, and one Q1a2a1-L715[74], in contrast to Tian Shan Sakas, who were all of R1-M173 subclades. Nomads from the central steppe (ca. AD 360), either Huns or Sarmatians, show one sample of hg. R1a1a1b2-Z93. All Huns show an increased shared drift with West Eurasians compared to Xiongnu (de Barros Damgaard, Marchi, et al. 2018).

A sample of the Mongolian Rouran Khaganate from Khermen Tal shows the continuation of haplogroup C2a1a-F4032 in the region (Li et al. 2018), probably expanding with Mongolian-speaking peoples in the eastern steppes. After their defeat (ca. 552 BC) a part of their population migrated to Pannonia where they became known as the Avars and allied with the Longobards to defeat the Gepids, creating the Avar Khaganate (AD 567-805).

Two early female Avars from Szólád (ca. 540–640 BC) show an ancestry similar to Eastern Europeans, with contributions of East Asian ancestry, clustering close to modern West Slavs, which justifies the Central Asian admixture found in a Gepid and a medieval Bavarian individual of the region (Amorim et al. 2018). Twenty-three individuals from a group of elite burials in Hungary (AD 7th–8th c.) show a majority of Inner Asian origin (up to 64%) of their mtDNA (Csáky et al. 2018), although there is high intergroup variation (Šebest et al. 2018).

The Turkic Khaganate assumed military and political organisation of the steppes as the Hunnic Empire broke up and dispersed (AD 6th c.), following the emergence of the Turks, the blacksmiths of the Rourans (Suppl. Fig. 17). Their elite soldiers are genetically closer to East Asians than the preceding Huns of the Tian Shan mountains, with one sample from Berygavoya (ca. 690 BC), of hg. R1-M173. A genetic outlier of the central steppe (ca. AD 270), of hg. R1-M173, shows pronounced European ancestry, and thus ongoing contacts with Europe (de Barros Damgaard, Marchi, et al. 2018).  

The Turkic Khaganate was eventually replaced by short-lived steppe cultures, such as the Kipchak and the Tungusik Kimak poulations, which spread southwards towards the Tian Shan mountains and westward towards the Ural Mountains to form the Kimak Khaganate in the central steppe (ca. 8th–11th c.). One sample from Kimak nomads of the Central Steppes (ca. AD 665), of hg. R1b1a1a1a-Y14051, does not show elevated East Asian ancestry (de Barros Damgaard, Marchi, et al. 2018).

The Kima Khaganate was replaced by local Kipchak groups allied with the Cuman of West Eurasia, hence probably originating near the area of Tuva. Two individuals dating to the Cuman–Kipchak alliance (ca. AD 1050) show one hg. C-M130 and increased East Asian ancestry, and the other one pronounced European ancestry, which is compatible with the incorporation of western and eastern steppe populations. The Karakhanid Khaganate from Turan incorporated some of these groups, with three samples (ca. AD 950–1250) showing further East Asian influx compared to earlier Turks (de Barros Damgaard, Marchi, et al. 2018).

Other unassigned Early Medieval Turkic samples show different proportions of East Asian ancestry, including nine from Tian Shan, among them one early (ca. AD 800–1000), of hg. J2a-M410, and one later (ca. AD 1170) of hg. C2a1a1b1b1-Y12825 (formed ca. 1000 BC, TMRCA ca. 100 BC); one from the Central Steppe (ca. AD 735) of hg. R1b1a1a1a-Y14051; and one from the Caspian Steppe (ca. AD 700) of hg. R1a1a1b2a2-Z2124[75]. Among two samples (ca. AD 1250) of the Medieval Jochi Khan’s Golden Horde in the central steppes, there is one of clearly East Asian ancestry and corresponding PCA cluster, of hg. C2-M217, and one of West Eurasian descent, of hg. R1a1a1b1a2a-Z280, which is further proof of the assimilation of different groups into succeeding Turkic organisations (de Barros Damgaard, Marchi, et al. 2018).

Among modern Tatars, descendants from elite clans of the Golden Horde belong to haplogroup R1b1a1a-M73 (Akchurin et al. 2018). There seems to be a general trend during the Iron Age and Medieval times to a distribution of R1a1a1b2a2-Z2124 lineages in the Pontic–Caspian steppes, of R1b1a1a-M73 lineages in the central steppes, and of R1-M173 (likely R1b2-PH155) and C2a1a1b1b1-Y12825 in the eastern steppes, which may reflect to some extent the different alliances formed by multi-ethnic groups since the time of the formation of the Xiongnu confederacy and the Hunnic expansion, although it may also reflect the initial contacts between peoples of the eastern steppe before the formation of the Xiongnu community. Modern peoples from investigated Xinjiang sites show hg. R1b-M343 including R1b1a1-P297, R1b1a1a-M73 (up to 9%), and R1b2-PH155 subclades, among a majority of typically central Asian lineages, including hg. R1a1a1b2-Z93, especially R1a1a1b2a2-Z2124 (Liu et al. 2018), whose origin cannot be properly interpreted without specific subclades.

Eventually, these khaganates were conquered by the Mongol Empire, which emerged through the unification of East Mongolian and Trans-Baikal tribes, expanding under the rule of Genghis Khan (ca. AD 13th c.). Modern Mongolian tribes show a mixture of East Asian lineages, mainly C2 subclades (ca. 42%), including C2b-F1067 (ca. 29%) and C2a-L1373 (ca. 13%), but also O-M175 (ca. 24%) and N-M231 (ca. 18%). In the PCA, Mongolians cluster in close genetic proximity to a group of North Asian Siberians, including Altaians, Tuvinians, Evenki, and Yakut, with eastern tribes Abaga, Khalkha, Oirat, and Sonid showing the least differentiation, with close interaction between northern Eurasian populations (Bai et al. 2018). Common Mongols likely expanded mainly with a Y-chromosome bottleneck of haplogroup C2a1a1c1-F3796 (TMRCA ca. 500 BC), whose expansion pattern is consistent with the diffusion of most Mongolic-speaking populations (Wei et al. 2018).

Manchu-Tungusic is proposed to have spread either from the Trans-Baikal area or from the Amur River region. Ancient samples from the West Liao River region shows high dynamism, similar to Trans-Baikal areas[76], which may support the emergence of Tungusic-speaking peoples from previous population movements through the eastern steppes. The presence of shared isoglosses with Turkic and Mongolic to the west, forming a likely Altaic family, and with Koreanic (and Japonic) to the east, with less clear links to Altaic, makes the identification of Tungusic still more complicated. The Proto-Tungusic society has been associated either with the Donghu or with an ancestral group from the Amur River region sharing links with eastern peoples. The potential attribution of vowel harmony in Proto-Tungusic to contacts with Mongolic languages (Ko, Joseph, and Whitman 2014) makes the identification of the language with one or the other group still harder.

Present-day Tungusic-speaking peoples, of varied lineages and ancestry, share a similar history to that found among Palaeosiberian peoples integrated among Finnic, Samic, Samoyedic, or Yukaghir-speaking populations, of acculturated Palaeosiberians adopting languages in recent times (Fedorova et al. 2013). This tradition of exogamy notwithstanding, it seems that the recent expansion of C2a1a1b1-M86 lineages among some southern Tungusic groups (as well as other C2-M217 lineages) may have been associated with their expansion from the south-east (Balanovska et al. 2018).