VIII.15. Eastern European Forest Zone

VIII.15.1. Balanovo

During the Eneolithic, subsistence economy on the Cis-Urals forest zone was based on the effective hunting of big hoofed animals (reindeer, elk, antelope, wild pig, bear, and beaver), gathering, and productive fishing (sturgeon, grayling, pike, chub, idus, tench, etc.). Large wooden rectangular houses were arranged in rows along the riverbank, and connected to each other and with farmyards by roofed passages. Small settlements were concentrated at the confluence of rivers (Koryakova and Epimakhov 2007).

Groups of the Cis-Urals regiondescended from sub-Neolithic Volga–Kama and Kama cultures, and probably related to Volosovolike Novolin, Garino–Bor, and Yurtik cultures showed late and simple metalwork, limited in forms, made from poor copper of the western Urals region. The eastward diffusion of cattle breeders and farmers of Corded Ware cultures brought a change in pottery design, stone tools, subsistence economy, and funerary ritualincluding flat and kurgan burials containing crouched skeletons, accompanied mostly by globular short-necked vessels and stone battle–axeswhich can only be explained by the sudden irruption of western settlers (Koryakova and Epimakhov 2007).

The north-east province of the Balanovo culture, a variant of Fatyanovo, was the easternmost group of the Corded Ware culture, occupying the Kama–Vyatka–Vetluga interfluve. Hundreds of sites, including villages, cemeteries, and numerous stone axes found by chance, represent the expansion of the culture in the region, with sites usually located on the high hills of riverbanks, and villages consisting of several above-ground houses (ca. 16–28 m2) built from wooden logs and saddle roofs, joined by passages (Koryakova and Epimakhov 2007).

Cemeteries contain both individual and collective graves, with men buried on the right side, women on the left side, both in contracted position, and dead wrapped in animal skins or birch bark, and placed into wooden constructions. Grave assemblages depend on sex, age, and social position, with copper axes accompanying elites, stone axe–hammers with men and teenagers, and flint axes with everyone, except chiefs. Balanovo layers are basically connected or overlap the late Volosovo and Garino layers, which indicates that Balanovo settlers occupied previous Eneolithic sites (Koryakova and Epimakhov 2007).

Balanovo brought a more advanced economic and cultural tradition than that of their neighbours. Their subsistence economy was based on animal husbandry, primarily pigs and sheep, but also later including cattle and horses depending on the local ecological conditions. They used draught cattle and wagons, exploited the local copper sandstone depositsbringing thus metallurgical tradition to the regionand pioneered the swidden method of farming (Koryakova and Epimakhov 2007).

VIII.15.2. Netted Ware, Chirkovo, Kazan

The Netted Ware or Textile Ware culture (Figure 76) appeared ca. 1900 BC in the Upper Volga–Oka region, derived from Fatyanovo–Balanovo settlers that inhabited previous Volosovo area and interacted with the Seima–Turbino network. It spread ca. 1900–1800 BC to the north into inner Finland, at the same time as the Pozdnyakovo branch of early Srubna exerted its influence on it.

Netted Ware expanded into regions previously occupied by cultures producing asbestos- and organic-tempered wares, reaching the Narva River on the eastern border of Estonia to the west, the Oulu River to the north, and Karelia but did not settle in the coastal zones (Figure 77). The early period of Netted Ware in Finland and Karelia is represented by the Sarsa–Tomitsa ceramics, starting ca. 1700 BC, with a later subgroup of Kalmistonmáki ceramics (Parpola 2018).

Figure 76. Vessels of the Netted Ware culture from Karelia. Modified from Kosmenko (1996).

It is not clear yet what the relationships between this immigrant tradition and the local populations of inner Finland were, but it is evident that changes took place in all fields of life: settlements, material culture, means of subsistence, and thus Weltanschauung. This does not imply a synchronous or abrupt change, or a complete population turnover, because traditional forms of subsistence held their ground alongside slash–and–burn agriculture for centuries and even millennia (Nordqvist 2018).

Figure 77. Distribution of eastern Textile Ceramics in northern Europe. 1 ‘Wafer’ ceramics; 2 Area of eastern Textile Ceramics; 3 Area in the early stage. Modified from Kosmenko (1996).

To the east of the textile ceramics area, in the Mid–Volga region, Abashevo settlers fought for the possession of the area between the mouths of the Kama and Vyatka, rich in copper deposits. The Chirkovo culture (ca. 1800–700 BC) formed eventually by the fusion of Abashevo and Balanovo elites over previous Volosovo territory, with Balanovo remains found well into the second half of the 2nd millennium BC. In its early phase, it included the Seima siteone of the type sites for the Seima–Turbino phenomenon, with evidence of materials from as far away as the Krotovo culture—and participated in the Seima–Turbino network, from the forest-steppe of the middle Irtysh to the Baraba steppe on the upper Ob (Parpola 2018).

The Kazan culture (ca. 1900–800 BC) developed in the Vetluga–Volga–Vyatka interfluve based on Balanovo settlers over previous Garino–Bor territory, from which it retained local traditions. Bronzes of the first phase are of the Seima–Turbino type, and it eventually comprised the Kama and Belaya basins, as well as areas of the Middle Volga almost up to Samara to the south, interacting with early Srubna. Its final Maklasheevka phase (ca. 1000–800 BC) immediately preceded its replacement by the Ananyino culture (Parpola 2018).

Similarly, Late Bronze Age groups of the Cis-Urals Prikamsky subarea (Prikazanska, Erzovska, Lugovska, Kurmantau, Buiska), which continued Eneolithic traditions in pottery and house designs, had adopted animal husbandry based on cattle, horse, and to a lesser extent pigs, and sheep, with evidence of cultivation and continuing influence from Andronovo–Cherkaskul, which points to the settlement of Corded Ware-related peoples and partial acculturation of the region (Koryakova and Epimakhov 2007).

VIII.15.3. Ananyino and Akozino

At the turn of the 2nd–1st millennium BC, the Early Iron Age begins in the steppes with the expansion of ‘Pre-Scythian’ horse-riding nomads. In the Middle Volga area, the Ananyino culture appears in the Kama River and its tributaries Vyatka, Chusovaya, and Belaya, famed for its metallurgy, with revived features of the Seima–Turbino and Eurasian types. It represented the main Cis-Urals metallurgical centre, and features tin, tin–antimony–arsenic, and pure copper alloys, widespread in the Bronze Age. It produced a large number of socketed axes, spearheads, arrowheads, battle hammers, knives-daggers, and plenty of ornaments (Parpola 2013).

It expanded to the north and north-east into the basins of the Pechora, Vychegda, and Mesen’ rivers in the first half of the 1st millennium BC. To the south, a buffer zone existed between Ananyino territory and the area of the South Uralian and Volga Sauromatian nomads, although they were in close contact with the north Pontic area, the Caucasus, and Kazakhstan, by this time developing Pre-Sauromatian traditions. Their metallurgical techniques were based in part on technology from the Caucasus, and from the Pontic–Caspian steppes were imported Cimmerian bronzes, such as daggers with cross-linked handles, Koban bronze axes, or two-ring horse bits. Only later (ca. 4th c. BC on) would the local production change its orientation to that of Sauromatian needs (Vasilyev 2002).

Up to eleven territorial variants of the culture can be distinguished, although ceramics are quite homogeneous. Two main interrelated groups can be distinguished: the Post-Maklasheyevo group, at the core of Ananyino expansion to the east; and the Textile group (see below Akozino), both occupying the broad-leafed forests of the Volga–Kama area and being more advanced economically, possessing a quite developed bronze metallurgy. Northern cultural groups occupying the forests of the Kama, Vyatka, and Vetluga rivers under the influence of Ananyino were orientated mainly to hunting and fishing, with an archaic stone industry rooted in the preceding time (Koryakova and Epimakhov 2007).

There are open and fortified settlements, the latter appearing first in the south as a reaction to the nomadic threat and spreading “the Age of Fortresses” in the forest zone, characteristic of the Iron Age throughout eastern Europe and western Siberia. The expansion of a chiefdom-based system is marked thus by the appearance of fortified settlements in the core area of the culture. The Ananyino society shows the greatest degree of militarisation compared to other societies of the Eurasian temperate forest zones, including a great quantity of advanced forms of weaponry. Settlements were divided according to function into a clear hierarchy, with large fortified settlements as administrative centres, smaller sites as watch posts, and open villages of various sizes. Male chiefs probably related by kinship are distinguished by rich and distinguished goods, including imported objects (Koryakova and Epimakhov 2007).

Fortresses are therefore proper of middle and late stages, with earlier settlements being smaller and simpler, in close proximity to water. Fortified settlements were usually located on narrow promontories of high river banks, separated some 20–40 km from each other. They were variable in size, from large administrative and ceremonial centres (up to 30,000 m2) to small fortified subordinated settlements (up to ca. 4,000 m2), and both had usually two sides defended by steep slopes, and a third size limited by a big earthen moat and ditch. There were also seasonal, temporary hunting camps (Koryakova and Epimakhov 2007).

Settlements had numerous rectangular houses with a trend to develop large, above-ground pillar or log wooden constructions, from the early small semisubterranean dwellings. Hearths were located in the central part of a house floor, with one or two entrances to the houses, which were aligned in rows. There could have been a division according to function, with some central structures serving for ritual ceremonies, including sacrifices (e.g. horses, cows), and some exterior zones used for economic activities (Koryakova and Epimakhov 2007).

Ananyino graveyards were located near rivers, on high banks and terraces connected to certain villages. Graves with likely burnt wooden logs above them (in a sort of “mortuary house” above the burial) appear in the dozens or hundreds, organised in rows parallel to the river. The dead were placed in shallow oval or rectangular pits in an extended position, with their legs orientated to the river. Apparently, the Volga and Kama rivers were of great importance for the local population, which could have considered these running waters to be pathways for the dead (Koryakova and Epimakhov 2007).

The Ananyino cosmological model is connected in its upper level with the sun, an image that is repeated on various objects: round discs with the depiction of the face, round plaques with a concentric design, concentrically decorated spindle-whorls, etc. The middle level is associated with animals like elk, bear, wolf, and horse, while the underwater and underground creatures represent the lower level. Communal and tribal ritual centres show remains of fire surrounded by posts with various offerings, great accumulation of ash, charcoal, crushed bones, votive objects, thousands of arrowheads, hundreds of anthropomorphic and zoomorphic figurines, dogs, and bees (Koryakova and Epimakhov 2007).

Individual primary burials were the norm, but there were some collective and double burials, and some secondary burials with cleaned bones or separate skulls only. All graves contained pottery, and there were some practices involving fire cult, based on the charcoal in the grave infilling, possibly from pyres or sacrificial places with cremation. Male graves contained horse bones, weapons (spearheads, arrowheads, battle hammer-axes, socketed axes), tools (knives), decorations (belts with pendants, torques, bracelets). Female burials contained cattle bones, decorations, needles, and spindle whorls. Tombs of chiefs were more complex, with a circle of stone and a wooden roof cover, as well as rich assemblages (Koryakova and Epimakhov 2007).

Pottery is round bottomed, with profiled neck and smooth, sometimes polished surface. Corded and comb decoration combined with holes covers the upper part. Subsistence economy is based on stalled animal husbandry, which make up the majority of animal bones recovered, anddepending on the local environmentincluded cattle (30–40%), horses (ca. 30%), big pigs (ca. 20–40%) and sheep (ca. 10%), also used a source of wool. Dogs were commonly used, and hunting was common for fur and as an essential dietary complement, together with fishing and gathering. The advanced bronze metallurgy was also complemented by bone- and woodworking and iron production, which eventually replaced bronze production. Its main demographic growth due to the use of natural resources must have happened around the 5th–4th c. BC (Koryakova and Epimakhov 2007).

At the same time as the Ananyino culture begins to expand ca. 1000 BC, the Netted Ware tradition from the middle Oka expanded eastwards into the Oka–Vyatka interfluve of the Middle Volga region, until then occupied by the Chirkovo culture. Eventually, the Akozino and Akhmylovo groups (ca. 800–300 BC) emerged from the area, showing a strong cultural influence from the Ananyino culture (and often considered the Textile variant of the culture), by that time already expanding into the north-east forest of the Cis-Urals region (Parpola 2013).

The Akozino culture remained nevertheless linked to the western Forest Zone traditions, showing (Kuzminykh and Chizhevskij 2009):

·         Netted Ware ceramics;

·         socketed axes of “Akozino–Mälar” type, produced in a special metallurgical furnace distinct from that of the Ananyino culture, related to influences from the Lusatian culture from Poland (Parpola 2013); and

·         funeral customs involving the inclusion of the Akozino–Mälar axes and other specific weapons among the grave goods.

An extensive interregional trade system developed during this time in the eastern Baltic area. Celts of Ananyino type and shape are found widespread from ca. 10th to the 5th centuries BC in the eastern Baltic, including Sweden, Finland, and Karelia. Findings with Ananyino origins also include bimetallic, single-edged daggers with iron blade and bronze hilts, characteristic of the second stage of Ananyino development, ca. 8th – 7th centuries BC. On the other hand, Mälar-type celtsso called because of the original belief (as with axes) that they were produced in Mälaren, Swedenappear with an even wider geographical distribution, including Norway, southern Scandinavia, the Northern European Plains (Paavel et al. 2019), showing in the east a higher concentration on the western area of the Volga–Kama region, which connects it to the Akozino culture (Yushkova 2010).

The Gorodets culture (ca. 800 BC – AD 800) from the forest-steppe zone north and west of the Volga, shows fortified settlements during the Iron Age. Incursions of Gorodets iron makers into the Samara valley are seen by deposits of their typical pottery and a bloom or iron in the region (Kuznetsov and Mochalov 2016). This attests to continued contact between forest and steppe areas in the Cis-Ural region.

In the western region of the Forest Steppe, the Dnieper–Dvina and the Hatched Ware cultures develop with slight differences. Both of them feature jar-shaped and slightly-profiled forms, but while the Dnieper–Dvina culture shows smooth vessels, Early Hatched Ware culture shows mainly hatched vessels. Haching is also seen in Dnieper–Dvina, whose slightly-profiled pots show inverted or vertical rims, in contrast to the elongated and inverted rims of the Hatched Ware. In later stages, during the first centuries AD, carinated pottery features in the Late Hatched Ware, while profiled vessels with elongated rims prevail in the Dnieper–Dvina culture.

The Ananyino culture evolution in the Iron Age can be followed through its zoomorphic styles into Iron Age Pyanobor and Glyadenovo cultures, which partially reject the previous style and ‘barbarise’ (simplify) the ornamentations, under the influence from Scythian and related styles. The Glyadenov style of flat anthropomorphic and zoomorphic figurines can be followed in morphology and styles to the analogous Ural-Siberian Middle Age culturesItkuska, Ust’-Poluiska, Kulaiska cultureswhich can in turn be considered as prototypes of Permian styles (Vasilyev 2002).

viii.15. Mordvins and Mari-Permians

The Cis-Uralian forest-steppe zone north of the Pontic–Caspian steppes represents a key prehistorical linguistic and cultural frontier, a north–south ecotone between the pastoral steppes to the south and the forest zone to the north. Pastoral steppe subsistence is associated with dynamic organisations represented by Turkic, before that Iranian, and before that eastern Proto-Indo-European. The Forest Zone is clearly represented pre- and proto-historically by Uralic languages, before they were displaced by Russian (Anthony 2016).

The synchronous appearance of closely related cultures in the forest-steppe and steppe regions, coupled with the emergence of fortified settlements (and thus a chiefdom-based system) in the whole Middle Volga area, should be identified as one of the few cases known where this stable prehistoric linguistic and cultural frontier has been crossed, allowing for the infiltration of Abashevo peoples from the forest into the steppe, probably boosted by the expansion of late Corded Ware groups in the region. This and subsequent interactions between northern and southern Abashevo settlers for centuries left a strong genetic and linguistic impact in both forest and steppe cultures, although each territory eventually retained their own ancestral culture and subsistence economies, which in turn determined their ethnolinguistic identification.

Uralic is widely supported to be spread through the east European forest zone already during the Fatyanovo-Balanovo and Abashevo expansion (Koivulehto 2001, 2003; Kallio 2002, 2017; Parpola 2013). The presence of Proto-Indo-Iranian and Proto-Iranian loanwords in Finno-Ugric (Kallio 2002; Koivulehto 2003; Koivulehto 1991), as well as the Uralic influence in Proto-Indo-Iranian (Kallio 2001), prove that these communities bordered each other in long-lasting contacts. Non-Uralic toponyms in the Volga–Oka and Volga–Kama areas support the admixture of incoming Uralic speakers with non-Uralic-speaking local hunter-gatherers of the Pit–Comb Ware culture (Kallio 2015; Zhivlov 2015), some of which may have survived in northern Russia up to the Common Era, based on topo-hydronyms of the area (Helimski 2001).

The synchronous appearance of Sintashta–Potapovka–Filatovka complex in the steppes, derived in great part from the migration of Abashevo settlers and transformation of southern Abashevo territory (see §VIII.18.1. Sintashta–Potapovka–Filatovka), should be identified in part with this interaction area in the forest-steppe and steppe areas. Southern Abashevo and later Pozdnyakovo (from Srubna), as well as the steppes, would then correspond to the Pre- and Proto-Indo-Iranian period, while northern Abashevo and related Fatyanovo and Battle Axe cultures to the north would correspond to the evolving Finno-Ugric community.

While Abashevo has not been sampled yet, the Poltavka outlier from the Sok River in Samara (ca. 2900–2500 BC), of hg. R1a1a1b2a-Z94, clustering closely with Corded Ware, probably represents an Abashevo-related immigrant. Similarly, early Sintashta and Srubna samples, of intrusive R1a1a1b2-Z93 lineages (and some of hg. R1a1a1b1a2-Z280 in Srubna) suggest the mixture of both haplogroups in Abashevo, probably with a gradient of increasing R1a1a1b1a-Z282 to the north in Fatyanovo–Balanovo.

The expansion of Balanovo- and Abashevo-related groups to the north-east, and the eventual formation of the Ananyino culture, are probably to be associated with the evolution of Proto-Permic. “It is commonly accepted by archaeology, ethnography, and linguistics that the ancestors of the Permian peoples (the Udmurts, Komi-Permians, and Komi-Zyryans) left the sites of Ananyino cultural intercommunity. In the west, in the Middle Volga basin, the Ananyino groups were neighbors of the groups that produced textile ceramics, these were ancestors of the Volga Finns (Goldina 1999; Napolskikh 1997)” (Koryakova and Epimakhov 2007).

The sudden emergence of the Ananyino and Akhmylovo cultures ca. 1000–800 BC is linked to the expansion of Iranian-speaking cultures of the southern steppe, creating a ‘push’ event that caused the emergence of a chiefdom-based social system in eastern Europe. Ananyino later split into two cultures, the Pyanobor culture (ca. 300 BC – AD 200) on the Vyatka, middle and lower Kama, and lower Balaya river, associated with Proto-Udmurt; and the Glyadenovo culture (ca. 200 BC – AD 500) on the upper Kama, connected with Proto-Komi (Parpola 2013).

The expansion of Ananyino culture into the north-east Cis-Urals region, with the formation of the Northern cultural groups of the Kama, Vyatka, and Vetluga rivers, is probably to be associated with the integration of some N1a1a1a1a-L392 lineages (formed ca. 4400 BC, TMRCA ca. 2900 BC)by this time probably widely distributed in the Circum-Arctic region and in central and western Siberia (see §§viii.16. Saami and Baltic Finns, §viii.21.1. Yukaghirs, and §viii.21.2. Turkic peoples and Mongols)into the Permic genetic stock. The best fit for northern Komi including AP ancestry (Sikora et al. 2018), instead of Baikal-related ancestry (as is common for some eastern Uralic groups), further supports the integration of distinct Circum-Arctic and Palaeosiberian groups among different expanding Uralic-speaking populations.

The expansion of Netted Ware from Abashevo near the Middle Volga, and its expansion into inner southern Finland, including the Sarsa and Tomitsa groups, possibly represented an expansion of West Uralic peoples, although Textile ceramics in the north are most likely also associated with West Uralic Balto-Finnic and Samic dialects. The emergence of Akozino in Netted Ware territory probably represents the spread of Proto-Mordvinic and closely related Volga-Finnic dialects, while the Gorodets culture can be more specifically correlated with the development of Mordvinians (Parpola 2013).

The expansion of Akozino warrior-traders under influence of Ananyino is probably to be related to a Y-chromosome bottleneck involving N1a1a1a1a1a-VL29 lineages (formed ca. 2100 BC, TMRCA ca. 1600 BC), accompanying migrants to the west into cultures around the Baltic area, likely through alliances and exogamy practices, in an integration facilitated by the emergence of chiefdoms in eastern Europe and the cultural influence and demographic growth of Ananyino and Akozino (see §viii.16.2. Baltic Finns). The more intense contacts with forest populations, and later expansion westwards by Akozino may justify a common, recent origin of certain non-Uralic loanwords shared by Balto-Finnic and Mordvinic (Häkkinen 2009).

An origin of haplogroup N1a1a1a1a1a-VL29 in western Siberia, between the Urals and Lake Baikal, can be proposed based on modern (Ilumäe et al. 2016) and ancient DNA (Cui et al. 2013; de Barros Damgaard, Martiniano, et al. 2018). The connection of its upper clade N1a1a1a1a1-Y6058/CTS10760 (formed ca. 2900 BC) with populations of Northern and Eastern Eurasia, and in particular the expansion of sister clade N1a1a1a1a3a-F4205 with Avars (see §viii.21.2. Turkic peoples and Mongols), and N1a1a1a1a-L392 with Northern Asian populations (see §viii.17.1. Ugrians), further supports its ancestral connection with Palaeosiberian populations (see §v.8. Palaeosiberians) and its lack of relationship with Uralic peoples, until their gradual incorporation during the expansion of Uralic dialects to the east and north (Figure 78).

Figure 78. Map of archaeological cultures in north-eastern Europe ca. 8th–3rd centuries BC. Modified from (Vasilyev 2002). Shaded area represents the Ananyino cultural-historical society. In pink and purple, Circum-Arctic and Siberian populations. Solid arrows depict the expansion of Uralic languages to the north, and steppe populations to the west, while dotted arrows represent the direction of “Siberian” ancestry and haplogroups from Akozino into the Baltic. Labelled are regions where N1a1-Tat is probably going to be found, with pink color representing non-N1a1a1a1a-L392 lineages. N1a1a1a1a2-Z1936 shows an ancestral connection between Palaeo-Laplandic and forest peoples from the Trans-Urals, while N1a1a1a1a1a-CTS10760 connects the Volga region with Altaic peoples. Image modified from Vasilyev (2002).

In fact, some available ancient DNA samples attributed to steppe nomads since the LBA–Iron Age transition show likely N1a1-Tat subcladessuch as an LBA individual from Afontova Gora (ca. 1000–850 BC), a Cimmerian from Hungary (ca. 980–930BC), and Scytho-Siberians (see below §viii.19. Iranians) which suggests a potential expansion of lineages ancestral to N1a1a1a1a1a-VL29 with Pre-Scythian and Scythian populations through a southern route across the Urals, and their integration into the Middle Volga area because of the intense contacts of Ananyino and Akozino–Akhmylovo with steppe peoples.

Mordovians seem to be less affected by the admixture with Palaeosiberian populations from the Circum-Arctic region, showing an ancestry closer to Corded Ware populations of the forest zone, formed by CWC (ca. 79%), WHG (ca. 9%) and Nganasan-like (ca. 12%), compatible with a lesser admixture with Palaeo-Arctic-like populations (Jeong et al. 2019). Mordovians also show more hg. R1a1a-M198 (ca. 27%), with less I-M170 (ca. 20%), N1a1-Tat (ca. 16%), R1b-M343 (ca. 13%), and J-M304 (ca. 12%).

Maris show a north-south cline of N1a2b-P43 (ca. 42% in Bashkirian Maris) against N1a1-Tat (ca. 13% among Bashkirian Maris, with up to 46% in northern Maris), with R1a1a-M198 lineages represented in both populations (ca. 28% in the south, 22% in the north), with approximately half R1a1a1b1a2-Z280 (16%), and half R1a1a1b2-Z93 (16%). None of the N1a1-Tat subclades among Bashkirian Maris investigated belonged to common subclades N1a1a1a1a1a-VL29 or N1a1a1a1a2-Z1936 (Dudás et al. 2019).

Among Permic peoples, Udmurts and Besermyans  show a similar ancestry, formed by CWC (ca. 71–74%) and Nganasan-like (ca. 25–29%), which support closer contacts with Palaeo-Arctic and Trans-Uralian populations (Jeong et al. 2019). Southern Udmurts show more hg. N1a1-Tat (ca. 54%) than R1a1a-M198 (ca. 19%) or N-M231(xN1a1-Tat) (ca. 17%); while northern Komis show less differences between N1a1-Tat (ca. 37%), R1a1a-M198 (ca. 27%) or N-M231(xN1a1-Tat) (ca. 19%).

The cluster formed by modern Mordovians, Estonians, as well as some Finns, Karelians, and Mansis probably represents the original Uralic cluster near Abashevo (Tambets et al. 2018). The presence of N-M231(xN1a1-Tat) lineages suggests contacts with Palaeosiberian peoples to the north, possibly involving migrations from the east (see §viii.17. Ugrians and Samoyeds).