Supplementary Notes

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Supplementary Notes

Supplementary Note 1. Archaeological and anthropological background

Veretye culture was defined by Svetlana V. Oshibkina based on excavations of Mesolithic sites at the Lacha Lake in the Arkhangelsk region in the Northern part of European Russia (Oshibkina, 1983; 1989; 2006). Excavations of the reference archaeological site of the culture –– Veretye 1 – were conducted by Oshibkina in 1978–1991 (Oshibkina 1997), although Mesolithic contexts in the area were studied already in 1920s30s by Mariya Ya. Foss (Foss, 1941) in the Veretye multilayer site (so-called Nizhneye Veretye) that is situated 80 m from Veretye 1. In the end of the 20th and the beginning of the 21st century other sites with similar archaeological materials were discovered and partly excavated in this territory (Oshibkina, 2006). The distribution of Veretye culture was generally limited by the area of the Vostochnoye Prionezhye region to the east of Lake Onega. This territory consists of glacial lake kettles that are followed now by lake lowlands. Settlements and campsites were located at the river banks often close to the river-mouths at the big lakes. The rather dynamic palaeoenvironmental history of the area (Kvasov 1975; Devyatova 1982; Kosorukova et al. 2018) provided certain conditions as Stone Age archaeological contexts were covered by peat sediments in many cases. Due to the good preservation of organic materials in the archaeological contexts, the material of Veretye culture is diverse, including bone, antler and even wooden artefacts, remains of dwelling and fishing constructions as well as archaeozoological finds etc. Beside settlements and campsites two burial sites of Veretye culture were studied. Remains of two people accompanied by several anthropogenic structures was discovered from the Peschanitsa burial ground, and 10 graves were excavated in the Popovo burial ground (Oshibkina 1982; 1994; 2017). The burial rites display some parallels to the Oleniy Ostrov Late Mesolithic cemetery in Lake Onega, but there are certain differences in the composition of grave goods (Oshibkina 2017). The chronology of Veretye culture (Oshibkina 2004) was originally established based on archaeological typology and results of palaeogeographical studies. A series of conventional radiocarbon dates from samples of organic sediments from cultural, overlapping and underlying layers confirmed the rather long chronology of the culture – from the end of the 9th millennium BC until the second half of the 6th millennium BC. However, radiocarbon dates are still relatively few and their context is not clear. The oldest and youngest radiocarbon dates from charcoal and wood found in Veretye 1 settlements, for which the reservoir effect is excluded, are 9,600±80 (Le-1469) – 9,237–8,766 cal BC with a 95.4% probability – and 7,700±80 (Le-1472) – 6,687-6,422 cal BC with a 95.4% probability (Oshibkina 2006). There are also a number of radiocarbon dates from human bones (Oshibkina 2006), but the potential for a reservoir effect must be considered. Two chronological phases were defined – the early one in the end of the Preboreal until the first half of the Boreal climatic periods, and the late one in the end of the Boreal period until the beginning of Athlanticum. The flint industry of Veretye culture was mainly based on processing local grey moraine flint of rather high quality. Flakes were mainly used as preforms, although blades made up around 20% of the

assemblage. Several blade production technological contexts were defined, including large blades, narrow blades and bladelets for slotted tools. Tools made of blades include arrowheads with and without tang, small points, possible knives and one obvious dagger from Veretye 1. Slate, sandstone and some crystalline rocks were used in the Veretye lithic industry as well. The amount of chopping tools made of both flint and slate is quite significant, over 20%. Sandstone spheroids with holes similar to so-called “mace-heads” were also found, widely presented in Mesolithic contexts in Finland and Karelia. The assemblage of bone and antler tools was presented by a variety of hunting, fishing and household tools, and inset daggers. According to archaeozoological materials, the subsistence of the Veretye culture population was mainly based on exploring forest resources, at least during its early phase. In the later phase, aquatic resources became more sufficient for the economy (Oshibkina 2006). The origin and quaintness of Veretye culture were recently discussed in a larger context of the Early Holocene cultural history of the Eastern European forest belt. Nowadays it is generally accepted that during the Early Mesolithic time the huge territory of the Eastern European Forest belt was a common cultural space due to the developed system of interregional networks (Oshibkina 2004; Zhilin 2003; Gerasimov et al. 2010). From the middle and end of the 8th millennium BC local cultural peculiarities became more pronounceable (Oshibkina 2006; Kriiska & Gerasimov 2014; Kriiska et al. 1916), which resulted in the formation of different cultural traditions. Burial sites included in this article Peschanitsa (Oshibkina 1994; 2006; 2017) burial site is located c 1 km from the eastern shore of the Lacha Lake in the Arkhangelsk region. The site was excavated by S. V. Oshibkina in 1986–87 and 1990– 91. Two graves and several pits surrounding them have been discovered. The pits contained flint adzes, scrapers, bladelets, retouched flakes, remnant cores and some bone artefacts, also animal and bird bones covered in red ochre. In one grave, the remains of ribs and a pelvis were found and a skull found by a local inhabitant during gravel mining can most probably be associated with these. Beside the bones, the grave contained three flint bladelets and separated animal and bird bones. The skull is the object of our study (PES001) and it belongs to a 45–50-year-old male. In the second grave, only leg bones were found. Those are radiocarbon dated to 9,890±120 (GIN-4858), 9,983–8,937 cal BC with a 95.4% probability. The bone collections in the two gaves have been considered to belong to both one or two people. Our dating of the skull (10,728±59 (UBA-41633), 10,785–10,626 with a 95.4% probability) confirms that these are two different individuals. In the case of both dates, however, it must be taken into account that they are affected by the freshwater reservoir effect, the magnitude of which we cannot estimate. Karavaikha 1 multiperiod archaeological site and burial ground is situated on the right bank of the Eloma (a channel of Modlona) river, some 18 km to the south from the Vozhe Lake in the Vologda region. The site was excavated by Alexander Ya. Bryusov during seven field seasons between 1938 and 1955, during which about 580 m2 were unearthed (Bryusov 1941; 1951; 1961; Utkin, Kostyleva 2001). The whole area presents a huge swamp with small patches of ground rising above water some 1–1.5 m. Karavaikha is located on one such small hillwhile the trail of the cultural layer also runs down the hill and was covered by peat and hyttia layers. Archaeological material presents all periods from the Mesolithic to the Bronze Age – there is no stratigraphic sequence, but certain patterns of spatial distribution of finds of different periods has been mentioned (Bryusov 1961; Kosorukova, Piezonka 2014).

A. Bryusov studied 38 ancient graves at the site and four more sites were defined later based on archive data and analysis of the archaeological collection by Alexander V. Utkin and Elena L. Kostyleva (2001), who also suggested an arranged numeration for the graves. The graves were dug in the 30–40 cm thick peat-silt cultural layer or deeper into the underlying clay layer. Burials differ in depth, location on the hill, composition of grave filling and amount of used ochre, which was considered as evidence of their different age. Burial goods are very rare and could instead belong to the cultural layer in general. Five radiocarbon dates associated with the burials (8,200±50 BP (GIN-7173), 7,351–7,066 cal BC with a 95.4% probability; 6,880±90 BP (GIN-7176), 5,978-5,631 cal BC with a 95.4% probability; 5,890±220 BP (GIN-7091), 5,305–4,344 cal BC with a 95.4% probability; 4,760±100 BP (GIN-7090), 3,794–3,127 cal BC with a 95.4% probability; 4,420±50 BP (GN-7172), 3,332–2,915 cal BC with a 95.4% probability) showed a rather wide time range (Utkin, Kostyleva 2001). Our study includes grave 12, which was excavated in 1939. The grave pit was 46 cm deep, reaching the surface of the clay layer. The grave contained a burial of an adult (KAR001), laid on their back in an extended position, head directed to W, hands extended along the body, head turned to the left (Bryusov 1951). The length of the well-preserved skeleton was 175 cm. The grave filling contained a lot of ochre. Grave 12 has been considered to belong to the earlier phase of the burial ground (the Neolithic), although no archaeological finds can be associated with the burial (Utkin, Kostyleva 2001). Mesolithic (mainly Late Mesolithic) archaeological contexts are well-presented in the nearest neighborhood of the site (Kosorukova, Piezonka 2014).

Lyalovo and Volosovo cultures

The Lyalovo culture was defined on the basis of the Lyalovo settlement site near Moscow that was excavated by Boris S. Zhukov (Zhkov 1925). The territory and chronology of the culture has been adjusted multiple times (overview of the research history e.g. Gurina & Krainov 1996; Zaretskaya & Kostyleva 2011). For now, the archaeological sites belonging to the Lyalovo culture have been found between the rivers Volga, Oka and Kostroma in Russia (Gurina & Krainov 1996). Settlement sites belonging to the Lyalovo culture were located on riverbanks and lake shores. Both houses with sunken floors as well as aboveground were built (Gurina & Krainov 1996). Burials can be found at or close to settlement sites (as described by Kostyleva & Utkin 2010). The graves are without inner constructions and typically contain only one inhumation. The deceased were placed in the grave in a supine position, later in a flexed position, and some of them had been either wrapped or tied before the interment. Ochre is known to be found in the graves. The earlier burials can be characterized by a lack of burial goods and embellishments normally found on clothes, except for a few bone artefacts. During the end of the culture period, amber and slate ornaments can be found amongst the artefacts. Lyalovo culture is characterized by large vessels made of clay tempered with mineral admixtures (with ground shell being used as an admixture during the end of the period). The vessels are mainly decorated with pits and comb impressions (Gurina & Krainov 1996; Lozovskiy et al. 2015). In lithic materials, the items of importance include bifacial flint arrow- and spearheads along with polished axes and adzes that were usually made of flint and less often from other rocks (Gurina & Krainov 1996). Bone and wood artefacts, including tools and fishing gear, can be found from settlement sites located in marshlands (Zhilin et al. 2002). The

subsistence of the people of Lyalovo culture was based on fishing, hunting and gathering (Gurina & Krainov 1996). Based on radiocarbon dating, the Lyalovo culture has been placed between ca. 5,200–3,700 cal BC (Zaretskaya & Kostyleva 2011). In the anthropological sense, the Lyalovo culture population is quite heterogenous (Kostyleva & Utkin 2010). The development of the culture (for different opinions on the matter see Gurina & Krainov 1996; Kostyleva & Utkin 2000; Zhilin et al. 2002) has been linked to both the Upper-Volga culture that preceeded the Lyalovo culture and was based in the middle of the European part of Russia (Yu. N. Urban, V. V. Sidorov, A. V. Engovatova), as well as either a widespread or limited migration of small populations from the Northern parts of Russia and Karelia (D. A. Krainov, A. V. Utkin, E. L. Kostyleva). The Volosovo culture succeeds the Lyalovo culture and is based on the area from Lake Ilmen to the Kama river basin district. Research into this culture group was started by Vasily A. Gordotsov who led the excavation at the Vosolovo settlement site in the Vladimir region in Russia (Gorodtsov 1926). The distribution and the chronology of the culture has been concreticised multiple times (research history can be obtained from Nikitin 1974; Krainov 1987a). It is probable that the Volosovo culture consists of multiple archaeological cultures of the same nature, with the most distinct of them based in the Upper/Middle Volga and Central/Lower Oka regions (Krainov 1987a). The settlements of this culture are situated at the shores of rivers and lakes, with a relatively large number of dwellings researched thus far. The houses may have sunken floors or be above ground and can be either quadrangular or, in fewer cases, slightly rounded (Krainov 1987a; Nikitin 2002; Sidorov 2002b). It is likely that more than one of such houses was simultaneously in use in most villages. The burial sites (as described by Kostyleva & Utkin 2010) can be found within the settlements or nearby. More than 30 burial sites with approximately 250 burials are known. The graves are dug into the ground and have no notable inner constructions. As with the Lyalovo culture, in the Vosolovo culture the dead were laid in the grave in a supine position, a practice which later evolved into burying the dead in a flexed position. The graves are usually meant for single burials and can form rows. Some of the graves included ochre. In the later phase of the culture, the practice of single burials lost its importance and collective graves, as well as partial interments, can be detected. The dead are often buried in clothes that have been lavishly decorated (including amber ornaments) and the grave goods include flint and bone anthropo- and zoomorphic figurines, flint arrowheads etc. However, it has been noted that in some cases the burials do not contain any ornaments or grave goods at all. Ritual practice areas and hoards compiled of different artefacts have also been found from the burial sites. The artefacts belonging to the Vosolovo culture are diverse. During the earlier period of the culture, vessels were mainly made of clay tempered with shell admixture, which was later replaced by plant- based admixture. In form, the vessels have a rounded bottom (with flat-bottomed forms found in later stages of the culture) and are decorated mainly with pits and comb impressions (Krainov 1987b). In lithic material, small flint artefacts such as scrapers, knives, arrowheads etc., as well as polished axes and adzes are well known in the Volosovo culture (Krainov 1987b) Different tools and fishing gear (such as harpoons, arrow- and spearheads, fishing hooks etc.) made of bone and wood are known from settlements in marshlands. Anthropo- and zoomorphic figurines and amber ornaments are characteristic to this

culture (Kashina 2002; Kostyleva & Utkin 2010). The subsistence of the Volosovo culture people was based on freshwater fishing, hunting and gathering (Krainov 1987b; Macāne et al. 2019). Based on radiocarbon dating, the Vosolovo culture can be placed between the first half of the 4th millennium cal BC to the first half of the 3rd millennium cal BC (discussion see Macāne et al. 2019). The emergence of the culture has been seen as a gradual development in the area between the rivers Volga and Oka (Krainov 1987b; Nikitin 2011), as well as a result of migrations from the area of the Volga-Kama culture (Khalikov 1969) or from the Easter Baltic region (Zhilin et al. 2002). It has also been suggested that the Vosolovo culture may have developed based on the cultural integration of the populations inhabiting the forest belt and the people migrating from the Valdai upland region (Sidorov & Engovatova 1996; Sidorov 2002a). Burial site included in this article Berendeyevo (Nikitin 1976; Kostyleva & Utkin 2010) burial site is situated on the edge of the Berendeyevo bog in the Yaroslavl region. Graves with three individuals were discovered during the excavation settlement site Berendeyevo I. Our study includes burial 1, which was unearthed by A. L. Nikitin in 1965. Burial 1 (BER001) is a 55–60-year-old male, who was placed in the grave on his left side in a flexed position, head directed to NW. The size of the grave was 1.1 × 0.7 m and the body had been wrapped in birchbark. Grave goods were not found, but the peat had preserved some remnants of clothing on a few bones and some rope, which had been used to wrap the body. The birchbark of burial 1 has been sampled and two radiocarbon datings were made from the sample: 7,730±40 BP (GIN-197) – 6,638–6,478 cal BC with a 95.4% probability – and 4,340±40 BP (GIN-197b) – 3,086–2,890 cal BC with a 95.4% probability. Birchbark was also discovered and radiocarbon dated in other graves: grave 2 4,970±95 (Mo-446) – 3,972–3537 cal BC with a 95.4% probability – and grave 3 5,540±50 BP (GIN- 3886) – 4,487–4271 cal BC with a 95.4% probability. We have dated the individual from burial 1 – 5,487±40 BP (UBA-41612), 4,447–4,259 cal BC with a 95.4% probability. This result is similar to the dating from grave 3, but freshwater reservoir effect cannot be excluded. Burial 1 has been linked to Volosovo culture based on the flexed body position, the birchbark container and the stratigraphy of the site, but according to the radiocarbon date, the Lyalovo culture cannot be ruled out either.

Fatyanovo culture

The antiquities of the Fatyanovo culture are recognized in a wide area of the European part of Russia from the Novgorod oblast to Tatarstan, particularly the Lake Ilmen area to the Vyatka River (e.g. Krainov 1987b; Krenke 2019b). The research history of this culture extends back to about 150 years ago when in 1873 the Fatyanovo burial ground, which also gave its name to the culture, was discovered near the town of Jaroslav, prompting the beginning of excavations on the site the same year (Uvarov 1881). The main features of the culture were defined during the first quarter of the 20th century (Gorodtsov 1915). Research was especially active during the 1950s and 1970s when the topic was handled mainly by Dimitri Krainov in the western and Otto Bader in the eastern part of the culture area. Bader (1950) distinguished a separate Balanovo culture, but the two are still commonly approached as a singular comprehensive culture (e.g. Krainov 1987b; Krenke 2019b). Other subgroups have been recognized in the Fatyanovo culture on the basis of differences in pottery and the shape and material of tools as well as burial methods

(Krainov 1972; 1987b). By now, more than three hundred burial places belonging to the Fatyanovo culture (most of them found by chance during other excavation works) and numerous stray finds (mainly shaft-hole axes) have been found. Information about the settlements of the Fatyanovo culture people is scarce – for a long time, a number of them were only known from the area of the Balanovo group (Krainov 1987b). Found mostly on riverbanks, the settlement sites of the Balanovo group are usually in areas well protected by nature. Both open and fortified settlements were founded by the Balanovo culture group and the remains of dwellings (incl. timber-framed structures), as well as fortifications, have been excavated (Bader & Khalikov 1987). More extensive investigation into the settlement sites of the western part of Fatyonovo culture, including a systematic survey, began in the 2000s (e.g. Krenke 2014; 2019a). The Fatyanovo culture people buried their dead mostly in inhumation cemeteries without visible above ground signs. In a constrained area of the Middle Volga region, they were buried in singular or grouped barrows, located on higher moraine or sandi hills and the shores of watercourses and lakes (Bader & Khalikov 1987; Krainov 1987). The number of graves in a burial ground varies between 2 and 125 (Krainov 1987). Some regularities in the burial customs can be detected, described in more detail below (Krainov 1972; 1987b; Bader & Khalikov 1987; Volkova 2010). The dead are usually buried at a depth of 0.1–2.7 meters and the grave shafts can be relatively large (maximum 3 × 5.6 meters), some with wooden, birch bark or root constructions. The bodies have been laid in the grave mostly on their sides, with elbow, hip and knee joints flexed. In general, the men are laid on their right side and the women on their left. It has been suggested that some of the deceased had been wrapped prior to burial. As an exception, some cremation burials can be found. In some cases, charcoal and ochre have been found in graves. The grave goods usually consist of pottery vessels, battle axes, flint work axes, adzes and knives, as well as metal weapons and jewelry and in special cases, animal bones and shells. Differences between sexes and age groups have been detected in the depth of the graves as well as the grave goods. The stone battle axes are usually found with male burials, rarely with children and women, with the axe often placed near the head. Polished flint work axes and adzes as well as flint knives are usually found in the hip region in male burials and near the feet in female burials. Other items, such as flint arrowheads, scrapers, blades and flakes, bone tools (awls, points, adzes etc.) and ornaments made of animal teeth, bone and amber can also be found. The age-at-death of the deceased is usually between 30–50 years of age, in special cases up to 70 years. Women, however, are rarely older than 40 years (Krainov 1987b). The source of subsistence for the people of the Fatyanovo culture has mostly been thought to be animal husbandry, but also hunting, fishing and gathering, and the possibility of cultivation has not been excluded either (Krainov 1972, 1987b). The bones of domestic animals (pig, goat/sheep, sometimes cattle and horse) and items made from the bones have been found from several burial places (Krainov 1987b). Teeth belonging to horses, cattle and goat/sheep have been found from an extensively excavated RANIS settlement site in the Moscow area (Krenke 2019a) and the bones of domesticated animals have also been recovered from many settlement sites of the Balanovo area (Bader & Khalikov 1987). Both nomadic and sedentary animal husbandry has been assumed. The idea of a nomadic lifestyle has been reasoned from the scarcity of settlement sites (e.g. Tretiakov 1966) while the location of the burial places in the landscape is considered as indirect evidence for agriculture (e.g. Bader 1939; Krainov 1987b). The deforestation on the shores of the Moscow River during the Fatyanovo culture period, visible on pollen

diagrams, is presumed to be related to agriculture (Ershova 2014). However, direct evidence of cereal crop farming has yet to be found. Hunting as a source of subsistence can be inferred from the bone items found from burial places and animal bones from the settlement sites of the Balanovo group. The bones of elk, roe deer, reindeer, beaver, bear, marten and fox as well as other types of wild animals and fish have been differentiated (Bader & Khalikov 1987; Krainov 1987b). In the anthropological sense, the Fatyanovo culture people can be seen as a relatively homogeneous group with the greatest similarities, mostly in cranial features, to other Corded Ware cultures (Denisova 1966; Balueva et al. 1988; Varul et al. 2019). Craniometric studies of skulls from the Upper Volga region reveal that the people of Fatyanovo culture are characterized by dolichocranic skulls (long and narrow neurocranium), medium-height and medium-width face, obvious horizontal profiling and a high protruding noseband. These features are especially clear in the oldest skeletons. Only 14 radiocarbon dates of the Fatyanovo culture sites have been published until this research project and in many cases, the context has been unclear (Krenke 2019b). Based on reliable radiocarbon results, the Fatyanovo culture has been dated to the period between c. 2,750–2,500 (2,300) cal. BC (Krenke 2019b). While our results completely overlap with the beginning of the period, they do shift the end of the culture to be a bit younger, c. 2,150 cal. BC. Different hypotheses have been proposed on the topic of the origin of Fatyanovo culture as well as on its connections to the cultures of the East European Plain that preceded and followed it. In the first half of the 20th century, the idea of an autochthon emergence of the culture was assumed but many of the researchers later renounced the opinion in favour of migration (Krainov 1987b and references therein). The Fatyanovo culture has been quite unanimously treated as a part of European Corded Ware and Battle Axe cultures. The formation and distribution of the culture has on different occurrences been thought to be both prolonged (Krainov 1987b) as well as a relatively fast process (Krenke 2019b). Mostly, the formation of this culture has been described as a migration of settlers to the area that had previously been inhabited by the people of Vosolovo culture whose main form of subsistence was foraging (Krainov 1987a). The most impactful has been Krainov’s (1987) approach where he localized the source of Fatyanovo culture, as well as to all Corded Ware cultures, to the wider area of the basins of rivers Dnepr, Visla and Oder in modern Poland, Ukraine, Belarus, Russia, Lithuania and Latvia. Krainov assumed that the people of the Corded Ware culture settled on the areas of the later Fatyanovo culture in four major migration waves: 1) from Belarus, Lithuania and Eastern Latvia to the Upper-Volga region, 2) from Belarus and Mid- and Upper Dnepr area of Ukraine to Moscow area, 3) from the southernmost areas of Ukraine to Middle-Volga and 4) from Eastern Latvia and Belarus to the areas of Western Dvina and Ilmen. Krainov also did not exclude the idea of some mixing between the settlers and the local populations. From the pottery found from the Moscow river basin during the 2000s, matches have been found in Globular Amphorae, the pottery of the Rzucewo culture in the eastern coast of the Baltic Sea and the epi Corded Ware of the Southern Poland and Ukrainian Volyn areas. It has been assumed that the Fatyanovo culture developed rapidly on the basis of western Globular Amphorae and Corded Ware culture peoples and that the area of its formulation also included the modern Moscow area (Krenke 2019a; 2019b). Burial sites included in this article

Bolshnevo 3 (Krainov 1972; 1975) burial site is on a morainal hillock in the Tver region. It is the only Fatyanovo culture burial place in Bolshnevo that has been excavated – the other two which are known were destroyed by gravel mining. D. A. Krainov excavated the site in 1970. He discovered four graves – two were individual graves, while the other two had four skeletons in each grave. We included individual 1 and 2 from grave 4 and the individual from grave 3. A 40–50-year-old male was buried in grave 3 (BOL003). He was buried on his left side in a flexed position, head directed to NE. Grave goods included a battle axe, flint axe, two ceramic vessels, two flint knives, some bone artifacts and a shell. In grave 4 remains of four adults – two male and two female – and a dog were unearthed. Three skulls had signs of perimortem trauma. Individual 1 (BOL001) was a 45–55-year-old male, who was placed on his left side in a flexed position, head directed to E-NE. He had a perimortem injury. The grave goods included a battle axe and a flint knife, also a bear-tooth pendant and a pig’s bone were near the skeleton. Individual 2 (BOL002) from grave 4 was a 20–25-year-old woman, who was placed on her left side in a flexed position, head directed to SW. Signs of perimortem trauma were also present on her skeleton. AMS radiocarbon dating results from grave 4 are considerably different (burial 1 4,005±39 BP (UBA-41613), burial 2 3,876±36 BP (UBA-41614)) and conventional date from charcoal (3,960±130 BP (GIN-5240), Krenke 2019b), suggesting that possibly the freshwater reservoir effect should be taken into account in when interpreting the data. Goluzinovo (Krainov 1972, Krainov & Gadzyatskaya 1987) burial ground is located on a hillock near Sinyavka river in the Yaroslavl region. Five individual graves were excavated by D. Krainov in 1964– 65, and burial 4 was included into the current study. The individual (GOL001) is a 20–30-year-old male, who was buried on his right side in a flexed position, head directed to NW. Several grave goods were recovered: a battle axe, a flint work axe, three bone items, a ceramic vessel and half of a shell. Ivanovogorsky (Krainov 1963; Krainov 1972) burial ground is situated on the shore of the Ruza river in the Moscow region. The burials of two males and one female were unearthed by K. Ya. Vinogradov in 1928–29. We have included burial 3 to our study. The individual (IVA001) is a 30–40-year-old woman, who was buried in a flexed position on her left side, head directed to NE. Grave goods included two ceramic vessels, a flint arrowhead, two flint knives, a whetstone, several bone items, pendants from teeth and tubular bones, and small shells. Pig bones were recovered near her lower limbs – it has been suggested that these are food offerings. Khaldeevo (Nikitin 1964; Krainov 1972) burial site is on a small morainal hillock in the Yaroslavl region. A. L. Nikitin and D. A. Krainov excavated there in 1959. They discovered one grave, where an 18–25- year-old male (HAL001) was buried. He was placed in a supine position, head directed to SW. Two ceramic vessels, a bone adze and some animal bones were found near the skeleton. Khanevo (Krainov 1972; Kirianova 1976; Sidorov & Engovatova 1992) burial ground is situated on a hillock on the shore of the Iskona river in the Moscow region. Three graves were excavated by A. V. Sidorov in 1969 and three graves by D. A. Krainov in 1970. Our study includes burials 4, 5, 6 and a cranium (HAN001) which most likely was recovered from a grave that was disturbed by human activity. Burial 4 (HAN004) included a male skeleton aged 18–20 years, who was buried on his right side in a flexed position, head directed to SW. Grave goods included a battle axe, a flint spearhead, a flint arrowhead, a stone axe, a ceramic vessel, a flint knife and two bone items. Additionally, pig bones were recovered – most likely remnants of a food offering. Burial 5 (HAN003) included two individuals, a man

and a woman. Both were buried in a flexed position on their right side, heads directed to SW. Several grave goods were found: a battle axe, a stone axe, a flint knife, a flint arrowhead, two ceramic vessels, two bone awls and shells. Near their lower limbs, pig and coat/sheep bones were found – most likely remains from meat that was interred to the grave. Burial 6 (HAN002) is a male aged 18–20 (25) years, who was placed in a supine position, lower limbs flexed from joints, head directed to SW. Grave goods included two work axes, a flint knife, several bone artifacts and a ceramic vessel. Miloslavka (Krainov 1961; Gadzyatskaya 1964; Krainov 1972) burial site is located on a hillock on the bank of a small lake in the Ivanovo region. D. A. Krainov excavated the site in 1958 and 1961, revealing two graves. Also, reports of two additional destroyed graves were gathered from locals. Three graves out of four were single graves, and one was a double grave. The current study included burials 1 and 4. Burial 1 (MIL001) is a 30–35-year-old female, who was placed in the grave on her left side in a flexed position, head directed to E. Her grave goods included a ceramic vessel, a flint flake and several bone artifacts. Burial 4 (MIL002) is a woman, aged approximately 25 years, who was placed in the grave on her left side in a flexed position, head directed to SE. She was inhumed with a 4–5-year-old child. Several grave goods were recovered: a flint work axe, a ceramic vessel, flint flakes, pendants made from shells, and also animal and bird bones. Mytishchi (Bader 1959; Gadzyatskaya 1969; Krainov 1972) burial site is situated on a little hillock in the Ivanovo region. One grave with two individuals was excavated by N. P. Milonova and O. N. Bader in 1928 and D. A. Krainov opened another grave in 1966. Our study included material from burial 3 (MOT001), excavated by Krainov – likely an 18–20-year-old woman, who was placed on her left side in a flexed position, head directed to E. The grave had been partially destroyed. Recovered grave goods included a battle axe, four ceramic vessels, several copper items and a bear-tooth pendant. Naumovskoye (Krainov 1972; Kirianova 1973; Krainov & Gadzyatskaya 1987) burial ground is situated on a hillock next to the Kurbitsa river in the Yaroslavl region. Two graves were unearthed by N. A. Kirianova in 1965. Both individuals are included in the current study. The cranial region of burial 1 was disturbed during gravel mining. The remains of burial 1 (NAU001) derive from a 50–60-year-old male. The individual in burial 2 (NAU002) is an 18–20-year-old male, who was placed into the grave on his left side in a flexed position, head directed to NE. The burial included a battle axe, a ceramic vessel, two flint knives, a bear-tooth pendant, a bone awl, an adze and a circular item made of copper. Nikolo-Perevoz (Raushenbah 1960; Krainov 1963) is located on the right bank of the Dubna river in the Moscow region. In 1958 V. M. Raushenbah excavated a settlement site which had cultural layers from different time periods and unearthed a mass grave with nine individuals. The heads of six of them were directed to SW and three to NE, eight were placed into the grave in a flexed and one in a supine position. Five ceramic vessels, two battle axes, five arrowheads and a spear head were found. In three instances, some arrowheads characteristic to Volosovo culture were recovered amongst the human remains. Most likely the mass grave included individuals who were killed by Volosovo people. Four skeletons were sampled for this study (RDT001, RDT002, RDT003, RDT004). Nikultsino (Krainov 1987; Krainov & Gadzyatskaya 1987) burial site is on a hillock in the Yaroslavl region. 18 graves, some with double burials, were excavated by D. A. Krainov in 1964–65. Our study included burials 3b, 6, 7, 9, 11, 15, 16 and 17. Burial 3b (NIK005) was partially destroyed. The individual is a woman, who was placed in the grave on her left side, head directed to NE. A ceramic vessel and

shards from two additional ceramic vessels were found. Burial 6 (NIK007) is an 18–20-year-old woman, who was placed in the grave on her left side in a flexed position, head directed to SE. A stone work axe, a flint flake and a lump, numerous pendants from animal teeth, shells and copper, also pig bones and three ceramic vessels were recovered. Burial 7 (NIK008) is a 25–30-year-old male, who was placed in the grave on his right side in a flexed position, head directed to SW. His grave goods included a battle axe, a flint blade, two ceramic vessels, a bear-tooth pendant and a bone awl. Burial 9 (NIK004) is a 50– 60-year-old male, who was placed in the grave on his right side in a flexed position, had directed to SSE. His grave goods included a battle axe, a flint work axe, two ceramic vessels, a sheep bone awl and a flint flake. Burial 11 (NIK006) is a 13–16-year-old male, who was placed in the grave on his right side, head directed to SW. A battle axe, a flint knife, two ceramic vessels, flint flakes and quartz flakes were found. Burial 15 (NIK002) is a 40–50-year-old male, who was placed in the grave on his left side in a flexed position, head directed to NE. His grave goods included a battle axe, a flint work axe, a flint knife, a flint arrowhead, a bone pin, two copper items, a grinding stone, a bear-tooth pendant and three ceramic vessels. Burial 16 (NIK003) is a 30–40-year-old male, who was placed in the grave on his left side in a flexed position, head directed to NE. A battle axe was found in the grave. The human remains in burial 17 (NIK001) were poorly preserved and thus age-at-death and sex on the individual have not been assessed. The individual was placed on the left side, most likely in a flexed position. An unidentified flint item, a bone awl, a blank for an awl, a ceramic vessel, animal bones, a piece of a reindeer antler and pieces of red quartzite were found. Skomorokhovo (Krainov 1972; Gadzyatskaya 1976b) is situated on a sandy hillock on the bank of a small stream which flows to the Zmeyka river in the Ivanovo region. D. A. Krainov excavated three graves and discovered two destroyed graves in 1961–62. Our study includes burial 3 (SKO001) – a 20– 30-year-old woman, who was placed in the grave on her right side, head directed to SW. A piece of copper wire and three dog’s teeth were found in the grave. Timofeyevka (Krainov 1964; Krainov 1965; Krainov 1972) burial site is located on a small hillock in the Ivanovo region. In 1959 and 1961, D. A. Krainov excavated 15 single burials and two dog graves. The current study included burials 1, 2, 4, 5, 6, 7, 10, 12, 13, 14 and 15. Burial 1 (TIM007) is a 30–35-year- old male, who was placed on his right side in a flexed position, head directed to SW. The grave goods included a battle axe, a flint knife and a scraper, a ceramic vessel with sheep bones, two flint flakes and some bone artifacts. Burial 2 (TIM006) is a woman with 20–25 (35–40) years of age, who was placed on her left side in a flexed position, head directed to E. Two ceramic vessels, a flint knife, two bone items and sheep bones were found with the individual. Burial 4 (TIM003) is a 35(45)–55-year-old male who was placed in supine position, head directed to SW. He was buried with a flint blade and a flint lump. Burial 5 (TIM004) is a male with 30–40 (45–50) years of age who was placed in a flexed position, head directed to SW. The grave goods included a battle axe, a flint work axe, a flint knife, a fragment of a flint point, a flint scraper, three ceramic vessels, a grinding stone and some bone items. Additionally, five pebbles were placed into the grave. Burial 6 (TIM009) is a (16)18–20-year-old male, who was placed on his right side in a flexed position, head directed to SW. Grave goods included a battle axe, a flint work axe, a boar tusk pendant, a flint point, some bone artifacts, sheep bones and a ceramic vessel. Burial 7 (TIM010) is an 18–20-year-old male, who was placed in the grave on his right side, head directed to SW. A battle axe, flint flakes, a bone dagger, bones from a domestic animal, a round stone and two ceramic vessels were recovered from the grave. Burial 10 (TIM011) is a 12–13-year-old female, who was placed

on her left side in a flexed position, head directed to NE. The grave goods were a copper tube, a flint flake, a bone awl, several animal bones and teeth, and a ceramic vessel. Burial 12 (TIM002) is a 12–13- year-old male, who was placed in the grave on his right side in a flexed position, head directed to NW. Two flint flakes, a bone awl and a ceramic vessel were found. Burial 13 (TIM008) is a male with (30– 35) 50–60 years of age, who was placed on his right side in a flexed position, head directed to NE. His grave goods included a battle axe, a flint work axe, a flint knife, a ceramic vessel with sheep bones and a bone awl. Burial 14 (TIM005) is a female with 18–20 (25–30) years of age, who was placed in the grave on her left side in a flexed position, head directed to SE. A flint knife, two shell pendants, dozens bone tubular and other beads, several animal and bird bones, and two ceramic vessels were found. Burial 15 (TIM001) is an 18–25(30)-year-old male, who was placed on his right side in a flexed position, head directed to SW. A battle axe, a stone slab, a flint flake and a ceramic vessel with pig and dog bones were found in the grave. Volosovo-Danilovsky (Krainov 1972; Krainov & Gadzyatskaya 1987) burial site is located on a morainal hillock on the bank of the Levashevk river in the Yaroslavl region. Stray finds have been collected there since the 1930s, archaeological excavations were held by D. A. Krainov in 1962–64. This is the largest known Fatyanovo culture burial place, where 107 graves with 117 individuals have been discovered. Additionally, a dog’s burial was found. Previously, one radiocarbon dating of the site has been published, but it is unknown what was dated. The dating – 3,650±80 BP (Le-1044) – 2,281–1,775 cal BC with a 95.4% probability – is disputable. Krenke (2019b) has expressed doubt and the previously published dating also does not correlate with our AMS radiocarbon dating of 2570–2299 cal BC. One individual from burial 11 was included in the aDNA study: an adult woman who was placed on her left side in a flexed position, head directed to NE. Her grave goods included three ceramic vessels, a copper circular item, a flint knife and a work axe. Voronkovo (Krainov 1972; Gadzyatskaya 1976; Krainov & Gadzyatskaya 1987) burial site is on a morainal hillock on the bank of the Vondeli river in the Yaroslavl region. O.S. Gadzyatskaya excavated 13 graves from the site in 1965. We have included burials 5, 6, 7, 8 and 13. Burial 5 is a 20–30-year-old male, who was placed to the grave on his right side in a flexed position, head directed to SW. A battle axe, a flint work axe, a flint knife, a flint flake and a ceramic vessel were found from the grave. Burial 6 is a 45–55-year-old male, who was placed on his right side in a flexed position, head directed to SW. A battle axe, a flint work axe, two ceramic vessels, a flint knife and other flint, bone and copper items were in the grave. Burial 7 is an 18–20-year-old male, who was placed on his right side in a flexed position, head directed to SW. His grave goods included a battle axe, a flint work axe, a flint knife, a ceramic vessel, and several bone items and shells. Burial 8 is a 50–60-year-old woman, who was placed on her left side in a flexed position, head directed to SW. A battle axe, a flint work axe, a flint knife, three ceramic vessels, numerous tooth pendants, some bone artifacts and flint flakes were discovered near the remains. Also, bird bones were found from the grave backfill. Burial 13 is a 30–40-year-old male, who was placed on his right side in a flexed position, head directed to W. Only a few grave goods were discovered: a flint work axe, a flint flake and one animal bone.

Estonian regional group of Corded Ware culture

Judging mainly by the characteristic pottery and stone axes, the Estonian group of Corded Ware culture prevailed in present-day Estonia, northern and eastern parts of Latvia, southeastern Finland, as well as in the Karelian Isthmus, Ingria and the Pskov region in northwestern Russia (Jaanits et al. 1982; Nordqvist 2016, Kriiska et al. 2017). Geographically, it is a neighboring group of Fatyanovo culture, but their contact zone is still almost unexplored. Solitary Fatyanovo-type axes in Estonian Corded Ware culture context and Estonian-type axes in the area of Fatyanovo culture indicate some but not very close contacts between these groups (Jaanits 1973; Nordqvist & Häkälä 2014; Kriiska et al. 2015). Also, in time, the Estonian group is almost analogous to Fatyanovo culture, continuing perhaps only a little longer, the averaged range of AMS dates is 2800–2000 cal BC (Kriiska et al. 2015).

Research on the Estonian Corded Ware culture group began in the second half of the 19th century (Grewingk 1871) and the first general presentations were compiled in the first quarter of the 20th century (Ebert 1913; Tallgren 1922). Several important burial sites were studied in the 1920s and 30s, above all Sope and Ardu (Aul 1935; Indreko 1935; 1937). Excavations at settlement sites started more widely in the 1950s (Jaanits 1955; 19669). The Estonian group is characterized by a relatively large number of settlements (a little less than a hundred) located in different natural environments (Kriiska 2000; Kriiska et al. 2015; Paavel et al. 2016). Their burials (more than twenty) are all flat earth graves. The graves are simple pits, less than 1.5 m deep and with varying orientations. The burials are single, sometimes double inhumations, the deceased are in a flexed, rarely in a supine position (Indreko 1935; Varul et al. 2019). The grave goods include battle axes, work axes and adzes of flint and crystalline rock, flint knives as well as different bone tools. The subsistence of the Estonian Corded Ware group was likely based on a mixed economy, combining productive livelihoods with hunting, gathering and fishing (Kriiska 2000; Lõugas et al. 2007; Kriiska 2009). Based on aDNA studies (Saag et al. 2017; Mittnik et al. 2018), Corded Ware culture was brought into this area by newcomers, likely coming from somewhere in eastern or central Europe.

Burial site included in this article

Sope (Jaanits et al. 1982; Kriiska et al. 2007; Varul et al. 2019) is a burial ground situated on a sandy knoll on the bank of the Sope stream in Northeastern Estonia. Several graves were exhumed by locals in the end of the 19th and beginning of the 20th century and the unearthed remains were reburied into an unknown location. The site has been archaeologically studied twice: in 1926 by H. Moora who discovered burial Sope I, and in 1933 by R. Indreko who unearthed burial Sope II. According to reported information the burial site included at least 14 individuals, but only remains of Sope I and II have survived. The current study included a sample from Sope II, an adult woman, who was placed into the grave on her right side, lower limbs flexed from joints, head directed towards NW. The burial included a ceramic vessel, a bone awl, a shell of a freshwater pearl mussel (Margaritifera margaritifera) and some small round stones.

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Iberia_BA Hungary_EN Iberia_Chl Hungary_EN Iberia_Chl Hungary_EN Iberia_Chl Iberia_BA Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_EN Iberia_Chl Iberia_EN Iberia_Chl Iberia_EN Iberia_Chl Iberia_EN Iberia_EN Iberia_MN Iberia_EN Iberia_MN Iberia_EN Iberia_MN Iberia_EN Iberia_MN Iberia_MN LB K_EN Iberia_MN LB K_EN Iberia_MN LB K_EN Iberia_MN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN Remedello LB K_EN Remedello LB K_EN Remedello LB K_EN Sweden_TRB Remedello Trypillia Remedello Trypillia Remedello Trypillia Sweden_TRB Trypillia Trypillia BO L001 Trypillia BO L002 Trypillia BO L003 Trypillia GOL001 BO L001 HAL001 BO L002 HAN002 BO L003 HAN004 GOL001 IVA001 HAL001 MIL001 HAN002 MIL002 HAN004 NAU001 IVA001 NAU002 MIL001 NIK002 MIL002 NIK003 NAU001 NIK004 NAU002 NIK005 NIK002 NIK007 NIK003 NIK008AB NIK004 TIM006 NIK005 TIM008 NIK007 VO D001 NIK008AB VO R003 TIM006 VO R004 TIM008 VO R005 VO D001 SO P002 VO R003 Baltic_CWC VO R004 Baltic_CWC VO R005 Baltic_CWC SO P002 Baltic_CWC Baltic_CWC Baltic_CWC Baltic_CWC Baltic_CWC Baltic_CWC Baltic_CWC Baltic_CWC Baltic_CWC Baltic_CWC Baltic_CWC Baltic_CWC Baltic_CWC Baltic_CWC Central_CWC Baltic_CWC Central_CWC Baltic_CWC Baltic_CWC Central_CWC Central_CWC Central_CWC Central_CWC Central_CWC Central_CWC Central_CWC Central_CWC Central_CWC Central_CWC Central_CWC Central_CWC Central_CWC Central_CWC Central_CWC Central_CWC Central_CWC Central_CWC Central_CWC Central_CWC Central_CWC Central_CWC Central_CWC Central_CWC Central_CWC Central_CWC Central_CWC Central_CWC Central_CWC Central_CWC Central_LNBA Central_CWC Central_LNBA Central_CWC Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA Central_LNBA HallstattBylany Central_LNBA HallstattBylany Central_LNBA Northern_LNBA HallstattBylany Northern_LNBA HallstattBylany Northern_LNBA Northern_LNBA Northern_LNBA Northern_LNBA Northern_LNBA Northern_LNBA Northern_LNBA Northern_LNBA Northern_LNBA Northern_LNBA Northern_LNBA Northern_LNBA Northern_LNBA Northern_LNBA Northern_LNBA Northern_LNBA Sweden_LNBA Northern_LNBA Bell_Beaker Northern_LNBA Bell_Beaker Sweden_LNBA Bell_Beaker Bell_Beaker Bell_Beaker Bell_Beaker Bell_Beaker Bell_Beaker Bell_Beaker Bell_Beaker Bell_Beaker Bell_Beaker Bell_Beaker Bell_Beaker Bell_Beaker Bell_Beaker Bell_Beaker Bell_Beaker Bell_Beaker Bell_Beaker Bell_Beaker Bell_Beaker Bell_Beaker Bell_Beaker Bell_Beaker Bell_Beaker Bell_Beaker Bell_Beaker Bell_Beaker Bell_Beaker Bell_Beaker Bell_Beaker Hungary_BA Bell_Beaker Hungary_BA Bell_Beaker Hungary_BA Hungary_BA Hungary_BA Hungary_BA Hungary_BA Hungary_BA Hungary_BA Hungary_BA Hungary_BA Hungary_BA Hungary_BA Hungary_BA Hungary_BA Hungary_BA Hungary_BA Hungary_BA Hungary_BA Hungary_BA Hungary_BA Hungary_BA Ukraine_Eneolithic Hungary_BA Ukraine_Eneolithic Hungary_BA Ukraine_Eneolithic Ukraine_Eneolithic Ukraine_Eneolithic Ukraine_Eneolithic preScy_Ukr Ukraine_Eneolithic preScy_Ukr Ukraine_Eneolithic preScy_Ukr preScy_Ukr preScy_Ukr preScy_Ukr preScy_Ukr preScy_Ukr preScy_Ukr preScy_Ukr Baltic_BA preScy_Ukr Baltic_BA preScy_Ukr Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA 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Poltavka Poltavka Poltavka Poltavka Poltavka Poltavka Poltavka Poltavka Poltavka Poltavka Poltavka Potapovka Potapovka Poltavka Potapovka Potapovka Potapovka Potapovka Potapovka Potapovka Potapovka Potapovka Potapovka Potapovka Potapovka Potapovka Russia_EBA Russia_EBA Potapovka Yamnaya Yamnaya Russia_EBA Yamnaya_Bulgaria Yamnaya_Bulgaria Yamnaya Yamnaya_Kalmykia Yamnaya_Kalmykia Yamnaya_Bulgaria Yamnaya_Kalmykia Yamnaya_Kalmykia Yamnaya_Kalmykia Yamnaya_Kalmykia Yamnaya_Kalmykia Yamnaya_Kalmykia Yamnaya_Kalmykia Yamnaya_Kalmykia Yamnaya_Kalmykia Yamnaya_Kalmykia Yamnaya_Kalmykia Yamnaya_Kalmykia Yamnaya_Kalmykia Yamnaya_Kalmykia Yamnaya_Kalmykia Yamnaya_Samara Yamnaya_Samara Yamnaya_Kalmykia Yamnaya_Samara Yamnaya_Samara Yamnaya_Samara Yamnaya_Samara Yamnaya_Samara Yamnaya_Samara Yamnaya_Samara Yamnaya_Samara Yamnaya_Samara Yamnaya_Samara Yamnaya_Samara Yamnaya_Samara Yamnaya_Samara Yamnaya_Samara Yamnaya_Samara Yamnaya_Samara Yamnaya_Samara Yamnaya_Samara Yamnaya_Samara 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Karagash_MLBA Kashkarchi_BA Kashkarchi_BA Kashkarchi_BA Kazakh_Mys_MLBA Kazakh_Mys_MLBA Kashkarchi_BA Kazakh_Mys_MLBA Kazakh_Mys_MLBA Kazakh_Mys_MLBA Kazakh_Mys_MLBA Kazakh_Mys_MLBA Kazakh_Mys_MLBA Kazakh_Mys_MLBA Kazakh_Mys_MLBA Kazakh_Mys_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Kazakh_Mys_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Krasnoyarsk_MLBA Lisakovskiy_MLBA_Alakul Lisakovskiy_MLBA_Alakul Krasnoyarsk_MLBA Lisakovskiy_MLBA_Alakul Lisakovskiy_MLBA_Alakul Lisakovskiy_MLBA_Al… Maitan_MLBA_Alakul Maitan_MLBA_Alakul Lisakovskiy_MLBA_Al… Maitan_MLBA_Alakul Maitan_MLBA_Alakul Maitan_MLBA_Alakul Maitan_MLBA_Alakul Maitan_MLBA_Alakul Maitan_MLBA_Alakul Maitan_MLBA_Alakul Maitan_MLBA_Alakul Maitan_MLBA_Alakul Maitan_MLBA_Alakul Maitan_MLBA_Alakul Maitan_MLBA_Alakul Maitan_MLBA_Alakul Maitan_MLBA_Alakul Maitan_MLBA_Alakul Maitan_MLBA_Alakul Maitan_MLBA_Alakul Maitan_MLBA_Alakul Oy_Dzhaylau_MLBA Oy_Dzhaylau_MLBA Maitan_MLBA_Alakul Oy_Dzhaylau_MLBA Oy_Dzhaylau_MLBA Oy_Dzhaylau_MLBA Oy_Dzhaylau_MLBA Oy_Dzhaylau_MLBA Oy_Dzhaylau_MLBA Oy_Dzhaylau_MLBA Oy_Dzhaylau_MLBA Oy_Dzhaylau_MLBA Oy_Dzhaylau_MLBA Oy_Dzhaylau_MLBA Oy_Dzhaylau_MLBA Oy_Dzhaylau_MLBA Oy_Dzhaylau_MLBA Oy_Dzhaylau_MLBA Petrovka Petrovka Oy_Dzhaylau_MLBA Petrovka Petrovka Petrovka Petrovka Petrovka Petrovka Sintashta Sintashta Petrovka Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Sintashta Srubnaya Srubnaya Sintashta Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Srubnaya Zevakinskiy_LBA Zevakinskiy_LBA Srubnaya Zevakinskiy_LBA Zevakinskiy_LBA Zevakinskiy_LBA Zevakinskiy_LBA Zevakinskiy_LBA Zevakinskiy_LBA Zevakinskiy_LBA Zevakinskiy_LBA Zevakinskiy_LBA Zevakinskiy_LBA Zevakinskiy_LBA Zevakinskiy_LBA Zevakinskiy_LBA Zevakinskiy_LBA Zevakinskiy_LBA Kangju Kangju Zevakinskiy_LBA Kangju Kangju Kangju Kangju Kangju Kangju Kangju Kangju Kangju Kangju Kangju Kangju Kangju Kangju Kangju Poprad Poprad Kangju Sarmatian Sarmatian Poprad Sarmatian Sarmatian Sarmatian Sarmatian Sarmatian Sarmatian Sarmatian Sarmatian Sarmatian Sarmatian Sarmatian Sarmatian Sarmatian Sarmatian Sarmatian Sarmatian Sarmatian Sarmatian Sarmatian Sarmatian Sarmatian Sarmatian Sarmatian Sarmatian Sarmatian_SU Sarmatian_SU Sarmatian Sarmatian_SU Sarmatian_SU Sarmatian_SU ScySar_SU ScySar_SU Sarmatian_SU ScySar_SU ScySar_SU ScySar_SU ScySar_SU ScySar_SU ScySar_SU ScySar_SU ScySar_SU ScySar_SU ScySar_SU ScySar_SU ScySar_SU ScySar_SU ScySar_SU ScySar_SU ScySar_SU ScySar_SU ScySar_SU ScySar_SU ScySar_SU ScySar_SU Scy_Kaz Scy_Kaz ScySar_SU Scy_Kaz Scy_Kaz Scy_Kaz Scy_Kaz Scy_Kaz Scy_Kaz Scy_Kaz Scy_Kaz Scy_Kaz Scythian_East Scythian_East Scy_Kaz Scythian_East Scythian_East Scythian_East Scythian_East Scythian_East Scythian_East Scythian_East Scythian_East Scythian_East Scythian_East Scythian_East Scythian_East Scythian_East Scythian_East Scythian_East Scythian_IA Scythian_IA Scythian_East Tagar Tagar Scythian_IA Tagar Tagar Tagar Tagar Tagar Tagar Tagar Tagar Tagar Tagar Tagar Tagar Tagar Tagar Tagar Tagar Tagar Tagar Tagar Tagar Tagar Wusun Wusun Tagar Wusun Wusun Wusun Wusun Wusun Wusun Wusun Wusun Wusun SaltovoMayaki SaltovoMayaki Wusun SaltovoMayaki SaltovoMayaki SaltovoMayaki SaltovoMayaki SaltovoMayaki SaltovoMayaki Kazakh_His Kazakh_His SaltovoMayaki Kazakh_His Kazakh_His Kazakh_His BER001 EHG Kazakh_His KAR001 EHG BER001 PES001 EHG KAR001 EHG EHG PES001 EHG EHG EHG EHG SidelkinoEHG_ML EHG EHG BER001 EHG EHG KAR001 EHG SidelkinoEHG_ML PES001 EHG Baltic_CCC_WHG Baltic_CCC SidelkinoEHG_ML Baltic_CCC_WHG Baltic_CCC Baltic_CCC_WHG Baltic_CCC Baltic_CCC Baltic_CCC_WHG Baltic_CCC Baltic_CCC Baltic_CCC Baltic_CCC Baltic_CCC Baltic_CCC Baltic_CCC Baltic_CCC Baltic_CCC Baltic_CCC Baltic_CCC Baltic_CCC Baltic_CCC Baltic_CCC Baltic_CCC Baltic_CCC Baltic_CCC Baltic_CCC Baltic_CCC Baltic_CCC Baltic_CCC Ukraine_Mesolithic Ukraine_Mesolithic Baltic_CCC Ukraine_Mesolithic Ukraine_Mesolithic Ukraine_Mesolithic Ukraine_Mesolithic Ukraine_Mesolithic Ukraine_Mesolithic Ukraine_Mesolithic Ukraine_Mesolithic Ukraine_Mesolithic Ukraine_Mesolithic Ukraine_Mesolithic Ukraine_Mesolithic Ukraine_Mesolithic Ukraine_Mesolithic Ukraine_Mesolithic Ukraine_Mesolithic Ukraine_Mesolithic Ukraine_Mesolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Mesolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Ukraine_Neolithic Romania_HG Romania_HG Ukraine_Neolithic SHG SHG Romania_HG SHG SHG SHG SHG SHG SHG SHG SHG SHG SHG SHG SHG SHG SHG SHG SHG SHG SHG Baltic_HG Baltic_HG SHG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Baltic_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Iron_Gates_HG Bichon Bichon Bichon WHG WHG WHG WHG WHG WHG WHG WHG WHG WHG WHG WHG WHG WHG WHG WHG WHG WHG WHG WHG WHG WHG WHG WHG WHG WHG WHG WHG WHG WHG Cardial_EN Cardial_EN Central_MN Cardial_EN Central_MN Central_MN Central_MN Central_MN Central_MN Central_MN Central_MN Central_MN Central_MN Central_MN Central_MN Central_MN Central_MN Central_MN Central_MN Central_MN Globular_Amphora Central_MN Globular_Amphora Globular_Amphora Globular_Amphora Globular_Amphora Globular_Amphora Globular_Amphora Globular_Amphora Globular_Amphora Globular_Amphora Globular_Amphora Globular_Amphora Globular_Amphora Globular_Amphora Globular_Amphora Globular_Amphora Globular_Amphora Globular_Amphora Globular_Amphora Globular_Amphora Globular_Amphora Globular_Amphora Globular_Amphora Globular_Amphora Globular_Amphora Globular_Amphora Globular_Amphora Hungary_EN Hungary_EN Hungary_EN Hungary_EN Hungary_EN Hungary_EN Hungary_EN Hungary_EN Hungary_EN Hungary_EN Hungary_EN Hungary_EN Hungary_EN Hungary_EN Hungary_EN Hungary_EN Hungary_EN Hungary_EN Hungary_EN Hungary_EN Hungary_EN Hungary_EN Hungary_EN Hungary_EN Hungary_EN Hungary_EN Hungary_EN Hungary_EN Hungary_EN Hungary_EN Iberia_BA Iberia_BA Iberia_Chl Iberia_BA Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_Chl Iberia_EN Iberia_Chl Iberia_EN Iberia_EN Iberia_EN Iberia_EN Iberia_EN Iberia_EN Iberia_EN Iberia_EN Iberia_EN Iberia_EN Iberia_MN Iberia_EN Iberia_MN Iberia_MN Iberia_MN Iberia_MN Iberia_MN Iberia_MN Iberia_MN Iberia_MN Iberia_MN Iberia_MN LB K_EN Iberia_MN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN LB K_EN Remedello LB K_EN Remedello Remedello Remedello Remedello Remedello Remedello Remedello Sweden_TRB Remedello Sweden_TRB Trypillia Sweden_TRB Trypillia Trypillia Trypillia Trypillia Trypillia Trypillia Trypillia Trypillia Trypillia Trypillia Bell_Beaker Trypillia BO L001 Bell_Beaker BO L001 BO L002 Bell_Beaker BO L002 BO L003 Bell_Beaker BO L003 GOL001 Bell_Beaker GOL001 HAL001 Bell_Beaker HAL001 HAN002 Bell_Beaker HAN002 HAN004 Bell_Beaker HAN004 IVA001 Bell_Beaker IVA001 MIL001 Bell_Beaker MIL001 MIL002 Bell_Beaker MIL002 NAU001 Bell_Beaker NAU001 NAU002 Bell_Beaker NAU002 NIK002 Bell_Beaker NIK002 NIK003 Bell_Beaker NIK003 NIK004 Bell_Beaker NIK004 NIK005 Bell_Beaker NIK005 NIK007 Central_LNBA NIK007 NIK008AB Central_LNBA NIK008AB TIM006 Central_LNBA TIM006 TIM008 Central_LNBA TIM008 VO D001 Central_LNBA VO D001 VO R003 Central_LNBA VO R003 VO R004 Central_LNBA VO R004 VO R005 Central_LNBA VO R005 SO P002 Central_LNBA SO P002 Baltic_CWC Central_LNBA Baltic_CWC Baltic_CWC Central_LNBA Baltic_CWC Baltic_CWC Central_LNBA Baltic_CWC Baltic_CWC Central_LNBA Baltic_CWC Baltic_CWC Central_LNBA Baltic_CWC Baltic_CWC Central_LNBA Baltic_CWC Baltic_CWC Central_LNBA Baltic_CWC Baltic_CWC Central_LNBA Baltic_CWC Baltic_CWC Central_LNBA Baltic_CWC Baltic_CWC Central_LNBA Baltic_CWC Central_CWC Central_LNBA Central_CWC Central_CWC Central_LNBA Central_CWC Central_CWC Central_LNBA Central_CWC Central_CWC Central_LNBA Central_CWC Central_CWC Central_LNBA Central_CWC Central_CWC Central_LNBA Central_CWC Central_CWC Central_LNBA Central_CWC Central_CWC Central_LNBA Central_CWC Central_CWC Central_LNBA Central_CWC Central_CWC Central_LNBA Central_CWC Central_CWC Central_LNBA Central_CWC Central_CWC Central_LNBA Central_CWC Central_CWC Central_LNBA Central_CWC Central_CWC Central_LNBA Central_CWC Central_CWC Central_LNBA Central_CWC Central_CWC Central_LNBA Central_CWC Central_CWC HallstattBylany Central_CWC Central_LNBA HallstattBylany Central_LNBA Central_LNBA Hungary_BA Central_LNBA Central_LNBA Hungary_BA Central_LNBA Central_LNBA Hungary_BA Central_LNBA Central_LNBA Hungary_BA Central_LNBA Central_LNBA Hungary_BA Central_LNBA Central_LNBA Hungary_BA Central_LNBA Central_LNBA Hungary_BA Central_LNBA Central_LNBA Hungary_BA Central_LNBA Central_LNBA Hungary_BA Central_LNBA Central_LNBA Hungary_BA Central_LNBA Central_LNBA Hungary_BA Central_LNBA Central_LNBA Hungary_BA Central_LNBA Central_LNBA Northern_LNBA Central_LNBA Central_LNBA Northern_LNBA Central_LNBA Central_LNBA Northern_LNBA Central_LNBA Central_LNBA Northern_LNBA Central_LNBA Central_LNBA Northern_LNBA Central_LNBA HallstattBylany Northern_LNBA HallstattBylany HallstattBylany Northern_LNBA HallstattBylany Northern_LNBA Northern_LNBA Northern_LNBA Northern_LNBA Northern_LNBA Northern_LNBA Northern_LNBA Northern_LNBA Northern_LNBA Northern_LNBA Sweden_LNBA Northern_LNBA Northern_LNBA SO P002 Northern_LNBA Northern_LNBA Baltic_CWC Northern_LNBA Northern_LNBA Baltic_CWC Northern_LNBA Northern_LNBA Baltic_CWC Northern_LNBA Northern_LNBA Baltic_CWC Northern_LNBA Northern_LNBA Baltic_CWC Northern_LNBA Sweden_LNBA Baltic_CWC Sweden_LNBA Bell_Beaker Baltic_CWC Bell_Beaker Bell_Beaker Baltic_CWC Bell_Beaker Bell_Beaker Baltic_CWC Bell_Beaker Bell_Beaker Baltic_CWC Bell_Beaker Bell_Beaker BO L001 Bell_Beaker Bell_Beaker BO L002 Bell_Beaker Bell_Beaker BO L003 Bell_Beaker Bell_Beaker GOL001 Bell_Beaker Bell_Beaker HAL001 Bell_Beaker Bell_Beaker HAN002 Bell_Beaker Bell_Beaker HAN004 Bell_Beaker Bell_Beaker IVA001 Bell_Beaker Bell_Beaker MIL001 Bell_Beaker Bell_Beaker MIL002 Bell_Beaker Bell_Beaker NAU001 Bell_Beaker Bell_Beaker NAU002 Bell_Beaker Bell_Beaker NIK002 Bell_Beaker Hungary_BA NIK003 Hungary_BA Hungary_BA NIK004 Hungary_BA Hungary_BA NIK005 Hungary_BA Hungary_BA NIK007 Hungary_BA Hungary_BA NIK008AB Hungary_BA Hungary_BA TIM006 Hungary_BA Hungary_BA TIM008 Hungary_BA Hungary_BA VO D001 Hungary_BA Hungary_BA VO R003 Hungary_BA Hungary_BA VO R004 Hungary_BA Hungary_BA VO R005 Hungary_BA Hungary_BA Ukraine_Eneolithic Hungary_BA Ukraine_Eneolithic Ukraine_Eneolithic Ukraine_Eneolithic Ukraine_Eneolithic Ukraine_Eneolithic Ukraine_Eneolithic Ukraine_Eneolithic Ukraine_Eneolithic Ukraine_Eneolithic Ukraine_Eneolithic preScy_Ukr Ukraine_Eneolithic preScy_Ukr preScy_Ukr preScy_Ukr preScy_Ukr preScy_Ukr preScy_Ukr preScy_Ukr preScy_Ukr preScy_Ukr preScy_Ukr preScy_Ukr preScy_Ukr preScy_Ukr preScy_Ukr preScy_Ukr preScy_Ukr Baltic_BA preScy_Ukr Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA Baltic_BA BolshoyOleniOstrov Baltic_BA BolshoyOleniOstrov BolshoyOleniOstrov BolshoyOleniOstrov BolshoyOleniOstrov BolshoyOleniOstrov BolshoyOleniOstrov BolshoyOleniOstrov BolshoyOleniOstrov BolshoyOleniOstrov BolshoyOleniOstrov BolshoyOleniOstrov BolshoyOleniOstrov BolshoyOleniOstrov BolshoyOleniOstrov BolshoyOleniOstrov BolshoyOleniOstrov Baltic_IA BolshoyOleniOstrov Baltic_IA Baltic_IA Baltic_IA Baltic_IA Baltic_IA Baltic_IA Baltic_IA Baltic_IA Baltic_IA Baltic_IA Baltic_IA Baltic_IA Baltic_IA Baltic_IA Baltic_IA Baltic_IA Baltic_IA Baltic_IA Baltic_IA Levänluhta Baltic_IA Levänluhta Levänluhta Levänluhta Levänluhta Levänluhta Levänluhta Levänluhta HungarianScythian Levänluhta HungarianScythian HungarianScythian HungarianScythian HungarianScythian HungarianScythian HungarianScythian HungarianScythian HungarianScythian HungarianScythian HungarianScythian HungarianScythian HungarianScythian HungarianScythian Scy_Ukr HungarianScythian Scy_Ukr Scy_Ukr Scy_Ukr Scy_Ukr Scy_Ukr Scy_Ukr Scy_Ukr Scy_Ukr Scy_Ukr Scy_Ukr Scy_Ukr Scy_Ukr Scy_Ukr Scy_Ukr Scy_Ukr Scy_Ukr Scy_Ukr Scy_Ukr Scy_Ukr Scy_Ukr Scy_Ukr Scy_Ukr Scy_Ukr Scy_Ukr Scy_Ukr Chern Scy_Ukr Chern Chern Chern Chern Chern Chern Chern Hungarian_Med Chern Hungarian_Med Baltic_MA Hungarian_Med Baltic_MA Baltic_MA Baltic_MA Baltic_MA Baltic_MA Baltic_MA Baltic_MA Baltic_MA Baltic_MA Baltic_MA Baltic_MA Baltic_MA Baltic_MA Baltic_MA Baltic_MA Baltic_MA Baltic_MA Baltic_MA Baltic_MA ChalmnyVarre Baltic_MA ChalmnyVarre ChalmnyVarre ChalmnyVarre ChalmnyVarre ChalmnyVarre Bantus Bantus Mandenkas Bantus Mandenkas Yorubas Mandenkas Yorubas Et hiopians Yorubas Et hiopians Mozabites Et hiopians Mozabites Moroccans Mozabites Moroccans Saudis Moroccans Saudis Yemenese Saudis Yemenese Palestinians Yemenese Palestinians Jordanians Palestinians Jordanians Syrians Jordanians Syrians Lebanese Syrians Lebanese Turks Lebanese Turks Kurds Turks Kurds Iranians Kurds Iranians Azeris Iranians Azeris Orcadians Azeris Orcadians Welsh Orcadians Welsh Portugese Welsh Portugese Spaniards Portugese Spaniards French Spaniards French Germans French Germans French_Basques Germans French_Basques Sardinians French_Basques Sardinians North_Italians Sardinians North_Italians Tuscans North_Italians Tuscans Cypriots Tuscans Cypriots Bulgarians Cypriots Bulgarians Hungarians Bulgarians Hungarians Romanians Hungarians Romanians Gagauzes Romanians Gagauzes Slovaks Gagauzes SupplementarySlovaks Figure Slovaks1. ADMIXTURE analysis results with ancient data projected onto the Poles Poles Ukranians Poles Ukranians Belarusians Ukranians Belarusians Russians_Southern Belarusians Russians_Southern Russians_Central Russians_Southern Russians_Central Russians_North Russians_Central Russians_North Lithuanians Russians_North Lithuanians Latvians Lithuanians Latvians Estonians Latvians Estonians Ingrians Estonians Ingrians Finns Ingrians Finns Karelian Finns Karelian Swedes Karelian Swedes Saami_SWE Swedes Saami_SWE Saami_Kola Saami_SWE Saami_Kola Vepsas Saami_Kola Vepsas Maris Vepsas Maris Erza Maris Erza Moksha Erza Moksha Mordva_mix Moksha modernMordva_mix geneticKomis_Permic structureMordva_mix . A ancient individuals at K9, B modern population averages at K9, C ancient Komis_Permic Komis_Zyrian Komis_Permic Komis_Zyrian Udmurts Komis_Zyrian Udmurts Mansis Udmurts Mansis Nenets_Tundra Mansis Nenets_Tundra Nenets_Forest Nenets_Tundra Nenets_Forest Hantis Nenets_Forest Hantis Chuvash Hantis Chuvash Bashkirs Chuvash Bashkirs Tatars Bashkirs Tatars Armenians Tatars Armenians northwest_Mingrelians Armenians northwest_Mingrelians Abkhasians northwest_Mingrelians Abkhasians Balkars Abkhasians Balkars Adygei Balkars Adygei North_Ossetians Adygei North_Ossetians Chechens North_Ossetians Chechens Lezgins Chechens Lezgins Kumyks Lezgins Kumyks Nogais Kumyks Nogais Kalmyks Nogais individualsKalmyks at K3Turkmens to K18, KalmyksD modern population averages at K3 to K18. Turkmens Tajiks Turkmens Tajiks Uzbeks Tajiks Uzbeks Kyrgyzians Uzbeks Kyrgyzians Kazakhs Kyrgyzians Kazakhs Uygurs Kazakhs Uygurs Altaians Uygurs Altaians Khakases Altaians Khakases Hazara Khakases Hazara Pathan Hazara Pathan Burusho Pathan Burusho Balochi Burusho Balochi Brahui Balochi Brahui Makrani Brahui Makrani Sindhi Makrani Sindhi North_Kannadi Sindhi North_Kannadi Malayan North_Kannadi Malayan Paniya Malayan Paniya Sakilli Paniya Sakilli Han Sakilli Han Tu Han Tu Xibo Tu Xibo Daur Xibo Daur Hezhen/Nanai Daur Hezhen/Nanai Oroqens Hezhen/Nanai Oroqens Mongola Oroqens Mongola Mongolians Mongola Mongolians Japanese Mongolians Japanese Buryats Japanese Buryats Tuvinians Buryats Tuvinians Dolgans Tuvinians Dolgans Kets Dolgans Kets Nganasans Kets Nganasans Selkups Nganasans Selkups Shors Selkups Shors Yukaghirs Shors Yukaghirs Nivkhs Yukaghirs Nivkhs Yakuts Nivkhs Yakuts Evenkis Yakuts Evenkis Evens Evenkis Evens Koryaks Evens Koryaks Chukchis Koryaks Chukchis Chukchis

a b 0. 2 0. 19 > > Europe ot her – – Europe Uralic

, Z) Europe Finnic , Z) Siberia other 0. 19 EHG Siberia Uralic Caucasus 0. 18 Near East Central Asia Fatyanovo 0. 18 East Asia

f3(Yorubas; 0. 17 –

0. 17 f3(Yorubas; –

0. 16 0. 16

0. 15 0. 15

– f3(Yorubas; WeRuHG, Z) –> – f3(Yorubas; WeRuHG, Z) –> 0. 14 0. 14 0. 14 0. 15 0. 16 0. 17 0. 18 0. 19 0. 2 0. 14 0. 15 0. 16 0. 17 0. 18 0. 19 0. 2

c d 0. 19 0. 19 > > – – , Z) , Z)

0. 18 0. 18 Fatyanovo Fatyanovo

0. 17 0. 17 f3(Yorubas; f3(Yorubas; – –

0. 16 0. 16

0. 15 0. 15

– f3(Yorubas; YamSam, Z) –> – f3(Yorubas; CeCWC, Z) –> 0. 14 0. 14 0. 14 0. 15 0. 16 0. 17 0. 18 0. 19 0. 14 0. 15 0. 16 0. 17 0. 18 0. 19

Supplementary Figure 2. Outgroup f3 statistics' results of comparisons with modern populations. Outgroup f3 statistics' values of form f3(Yorubas; study population, modern) plotted against each other for two study populations (blue and red axis): A Western Russian hunter-gatherers (WeRuHG) and Eastern hunter-gatherers (EHG), B WeRuHG and Fatyanovo, C Yamnaya Samara (YamSam) and Fatyanovo, D Central Corded Ware culture (CeCWC) and Fatyanovo.

a b 0. 29 Caucasus/Near East 0. 77 > > – – Anatolia/Levant EF Forest/Steppe NCA

0. 76 , Z) , Z) Forest/Steppe EMBA Forest/Steppe MLBA 0. 28 Forest/Steppe IMA 0. 75 Europe HG (EHG–SHG) Europe HG (SHG) Fatyanovo Fatyanovo 0. 74 Europe HG (SHG–WHG) Europe EF Europe LNBA 0. 73 0. 27 Europe IA Europe MA f3(Mbuti; f3(Mbuti; f3(Mbuti; f3(Mbuti; 0. 72

WesternRussiaHG – – Fatyanovo 0. 71 0. 26 0. 7

0. 69 0. 25 0. 68

0. 67 – f3(Mbuti; YamSam, Z) –> – f3(Mbuti; YamSam, Z) – 0. 24 0. 66 > 0. 23 0. 24 0. 25 0. 26 0. 27 0. 28 0. 29 0. 66 0. 67 0. 68 0. 69 0. 7 0. 71 0. 72 0. 73 0. 74 0. 75 0. 76 0. 77

c d 0. 29 > – , Z)

0. 01 0. 28 – D(Yor, YamSam; CeRu, Z) –> Fatyanovo -0.08 -0.06 -0.04 -0.02 0 0. 02 0. 04

-0.01 0. 27 f3_A(Mbuti; f3_A(Mbuti; > – –

-0.03 0. 26 ; CeRu, Z) ; CeRu,

0. 25 -0.05 Fatyanovo

– f3_X(Mbuti; Fatyanovo, Z) –> D(Yor, 0. 24 -0.07 – 0. 66 0. 67 0. 68 0. 69 0. 7 0. 71 0. 72 0. 73 0. 74 0. 75 0. 76 0. 77 Supplementary Figure 3. Outgroup f3 and D statistics' results of comparisons with ancient populations. Outgroup f3 statistics' values of form f3(Mbuti; Yamnaya Samaya/Fatyanovo, ancient) using A autosomal, B X chromosome 1240k capture SNPs, C D statistics’ values of form D(Yorubas, Yamnaya Samaya/Fatyanovo; Central Russians, ancient), D outgroup f3 statistics' values of form f3(Mbuti; chr X/autosomal SNPs, ancient) of Fatyanovo using 1240k capture SNPs. EF – early farmers; EMBA – Early/Middle Bronze Age; MLBA – Middle/Late Bronze Age; IMA – Iron/Middle Ages; HG – hunter-gatherers; LNBA – Late Neolithic/Bronze Age; IA – Iron Age; MA – Middle Ages.

a 120% 120% 110% 110% 100% 1% 100% 2% 9% 7% 90% 12% 14% 90% 12% 80% 80% 70% 70% 60% 60%

50% 99% 50% 101% 98% 91% 88% 93% 40% 88% 86% 40% 30% 30% 20% 20% 10% 10% 0% 0% -1% -10% -10% -20% -20% qpAdm CP qpAdm CP qpAdm CP qpAdm CP Fatyanovo Central CWC Fatyanovo Baltic CWC Baltic CWC Levant N Central CWC Levant N

b 0.40 80

● ●●●●●●●●●●●●●●●●●●●● ●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●● ●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●● 0.35 ●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●● ●●●●●●●●●●●●●●●●●●●●●●●●●●●●●●● ●●●●●●●●●●●●●●●●●●●●●●●● ●●●●●●●●●●●●● ●●●●● 60 ●●●●● ●●●● ●●●●● 2nd degree 0.30 1st degree 40 0.25 Frequency

Identical/Twins 20 0.20 Average pairwise P0 (± 2SE) Average ● Identical/Twins 1st degree 2nd degree 0 0.15

0.20 0.25 0.30 0.35

Average pairwise P0 IVA001NIK005 IVA001NIK002 IVA001NIK003 IVA001NIK007 IVA001MIL002 IVA001NIK004 IVA001MIL001 IVA001TIM008 NIK003NIK005 MIL002NIK005 NIK004NIK005 NIK005NIK007 NIK002NIK005 MIL001NIK005 NIK002NIK003 MIL002NIK003 NIK003NIK004 NIK003NIK007 NIK002NIK007 MIL002NIK002 MIL002NIK007 MIL001NIK003 MIL001MIL002 MIL001NIK004 MIL002NIK004 NIK004NIK007 MIL001NIK007 MIL001NIK002 NIK002NIK004 NIK005TIM008 NIK003TIM008 MIL002TIM008 MIL001TIM008 NIK004TIM008 NIK002TIM008 NIK007TIM008 HAL001IVA001 BOL003IVA001 BOL001IVA001 BOL002IVA001 IVA001NAU001 IVA001NAU002 HAL001NIK005 HAL001NIK003 HAL001NIK007 HAL001MIL001 HAL001MIL002 HAL001NIK002 HAL001NIK004 HAN004IVA001 HAN002IVA001 GOL001IVA001 IVA001VOR005 IVA001VOR003 IVA001VOR004 HAL001TIM008 BOL002NIK005 BOL003NIK005 BOL001NIK005 BOL001NIK003 BOL003NIK003 BOL002NIK002 BOL002NIK003 BOL001MIL002 BOL003MIL002 BOL001MIL001 BOL003NIK002 BOL002NIK007 BOL003NIK007 BOL003NIK004 BOL003MIL001 BOL002MIL002 BOL001NIK002 BOL001NIK007 BOL002MIL001 BOL002NIK004 BOL001NIK004 BOL001TIM008 BOL003TIM008 BOL002TIM008 NAU001NIK005 NAU002NIK005 NAU001NIK003 NAU002NIK003 NAU001NIK007 MIL002NAU001 NAU001NIK002 MIL001NAU001 NAU001NIK004 NAU002NIK007 NAU002NIK002 MIL001NAU002 MIL002NAU002 NAU002NIK004 HAN002NIK005 HAN004NIK005 HAN002NIK003 HAN002MIL002 HAN002NIK007 HAN004NIK003 HAN004NIK007 HAN002NIK002 HAN004MIL002 HAN004NIK002 HAN004MIL001 HAN002MIL001 HAN002NIK004 HAN004NIK004 GOL001NIK005 GOL001NIK003 GOL001MIL002 GOL001NIK002 GOL001NIK007 GOL001NIK004 GOL001MIL001 NAU001TIM008 NAU002TIM008 NIK005VOR005 NIK005VOR003 NIK003VOR005 NIK003VOR003 NIK005VOR004 MIL001VOR005 MIL002VOR005 NIK002VOR005 NIK007VOR005 NIK007VOR003 NIK002VOR003 NIK004VOR005 NIK003VOR004 MIL001VOR003 NIK004VOR004 MIL002VOR003 MIL002VOR004 NIK007VOR004 NIK004VOR003 NIK002VOR004 MIL001VOR004 HAN004TIM008 HAN002TIM008 GOL001TIM008 TIM008VOR005 TIM008VOR003 TIM008VOR004 IVA001NIK008A IVA001NIK008B NIK008ATIM008 BOL001HAL001 BOL002HAL001 BOL003HAL001 NIK005NIK008A NIK003NIK008A NIK005NIK008B MIL002NIK008A MIL001NIK008A NIK002NIK008A NIK007NIK008A NIK004NIK008A NIK003NIK008B BOL001BOL003 BOL001BOL002 BOL002BOL003 NIK007NIK008B MIL001NIK008B NIK002NIK008B NIK004NIK008B MIL002NIK008B HAL001NAU001 HAL001NAU002 GOL001HAL001 NIK008BTIM008 HAL001HAN004 HAL001HAN002 BOL003NAU001 BOL002NAU001 BOL001NAU001 BOL003NAU002 BOL001NAU002 BOL002NAU002 HAL001VOR005 HAL001VOR003 HAL001VOR004 BOL002HAN004 BOL001HAN002 BOL003HAN004 BOL001HAN004 BOL003HAN002 BOL002HAN002 BOL001GOL001 BOL003GOL001 BOL002GOL001 NAU001NAU002 BOL003VOR005 BOL002VOR003 BOL003VOR003 BOL001VOR005 BOL002VOR005 BOL002VOR004 BOL003VOR004 BOL001VOR004 BOL001VOR003 HAN004NAU001 HAN002NAU001 HAN004NAU002 HAN002NAU002 GOL001NAU001 GOL001NAU002 NAU001VOR005 NAU001VOR003 NAU002VOR005 NAU001VOR004 NAU002VOR003 NAU002VOR004 HAN002HAN004 GOL001HAN002 GOL001HAN004 HAN002VOR003 HAN002VOR005 HAN004VOR005 HAN004VOR003 HAN004VOR004 HAN002VOR004 GOL001VOR003 GOL001VOR005 GOL001VOR004 VOR004VOR005 VOR003VOR005 VOR003VOR004 HAL001NIK008A HAL001NIK008B BOL002NIK008A BOL003NIK008A BOL001NIK008A BOL002NIK008B BOL001NIK008B BOL003NIK008B NIK008AVOR005 NIK008AVOR003 NIK008AVOR004 NAU001NIK008A NAU002NIK008A NAU001NIK008B NAU002NIK008B HAN002NIK008A HAN004NIK008A HAN002NIK008B HAN004NIK008B GOL001NIK008A GOL001NIK008B NIK008BVOR005 NIK008BVOR003 NIK008BVOR004 NIK008ANIK008B

Supplementary Figure 4. qpAdm, ChromoPainter/NNLS and kinship test results. A qpAdm and ChromoPainter/NNLS (CP) models with Baltic Corded Ware culture (CWC) or Central CWC and Levant Neolithic (Levant N) as sources, B Fatyanovo kinship test results, 100 most similar pairs out of 253 shown.

H5b

MH605047 Bulgaria E Bronze Age

1630 14007 @146 327 7897 @195 12864 16357 14497 145 14384 16271 1238 228 H5b2 1611 497 SCY300 H5b1 3075 2706 Moldova 146 8854 JQ703241 UK 3952 11914 4-2nd BCE 6710 13734 JQ703921 UK KJ446352 Orcadian NIK008AB 12401 JX128075 Slovak KJ446351 Orcadian Russia 14773 JQ704233 JX128050 FJ794473 English HQ700378 Irish Fatyanovo 14798 UK Czech JQ704238 UK 16224 KF161186 Danish 16278 KF163003 Danish 7058 9948 573.2C 15493 SCY334 JQ704863 EU597568 Moldova 152 13581 5492 9452 6662 16311 Ireland JQ704264 Orcadian 4-2nd BCE 249 UK 1503 JQ703150 MH120483 KF162713 JQ704238 MG952872 573.1C UK Poland Danish UK Hungary JQ704401 KF163051 KF451328 France 10589 Danish Orcadian EU597579 MG646217 Orcadian Poland KF451329 Orcadian b

H6a1a N49 Poland CWC HAN004 Russia Fatyanovo JQ705206 Poland

7202 10237 10936 3548 @16482 @146 5312 3079 93 JQ703161 MF177158 8131 3955 9055 H6a1a2 H6a1a4 H6a1a5 H6a1a10 Poland Poland 16543 8458 10187 JX153182 Finn HM015668 Ashkenazi Jew 0LS09 8834 15226C 9773 41 JX154007 Danish HQ405756 Russian 10250 5048 7706 9055 Estonia VOR003 16168 11662 GU123020 Tatar 16192 10166 11311 16325 Iron Age Russia 16172 H6a1a2a 5302 3523 748 16445 14211 16526 JX153818 Fatyanovo JQ703103 H6a1a2b 9055 16148 93 152 3944 146 4164 7855A 189 2263A JQ703061 16298 2010 JX297423 JQ702606 MH120516 Finn Norway 146 13218 JQ703507 6468 KY670940 JQ703813 13404 5460 12810 1389 14326 Russia KF162603 4947 Swedish Italy Poland JX153173 9145 146 152 KY671015 Norway 13105 Russian Norway KY671061 6611 MH120727 15930 MF177160 Danish GU390313 Finn 16360 @16482 @7202 Russian JQ704472 Russian @16482 Poland MG646120 Poland Jewish KU252600 JQ324568 16189 Poland 3591 MG952811 Poland Swedish France JQ704417 10724 Hungary German 14226 JX307107 Russian c

J1c1b1a

Kociumaka110 Germany BBC N17 Poland E Bronze Age RISE386 Russia Sintashta

14016 16235 2396 16189 210 5972 150 15617 16192 @16189 4204 16093 5987 16319 KF161705 7055 16111 Dane 44.1C 2396 13020 KY670931 J1c1b1a1 JX152910 16145 14214 188 AY714034 Russian Dane 234 15934d 7581 Indian BOL002 VOD001 12192 5078 15934d 11878 12963 Russia FJ502349 MH120721 Russia 5539C 16145 Fatyanovo German Poland Fatyanovo JQ797791 16172 MH120585 Sardinian MH120721 4820 16093 Poland 15047 MH120675 Poland 3075 11150 195 Poland JQ797790 JQ797789 9227 13707 8104 JX153365 Romanian Ukrainian 11348 12534 Dane 14207 JQ703599 Spain JQ705164 NIK005 UK Russia Fatyanovo d

1189 @146 195 @10398 9716 MT079052 Switzerland K1 K2 3330-3216 BC

497 5913 @146 16362 114 @152 2217 @152 6239 709 5231 K1a K1b 498d 11929 10542 4561 14037 por004 Spain 5041-4867 yBP BER001 11962 esp005 Spain Bronze Age 9962 @146 K1c Russia 16093 K2a K2b 10289 @195 WeRuHG KO2 N25 Poland 4600 BCE 114 13117 11485 @195 11722 15946 12738G 9093 Hungary KY782197 Polish 11869A @11914 MT079071 Switzerland 2213-2031 BC 16189 11377 E Neolithic K1a3 K1a4 MT079055 Switzerland 2914-2871 BC MT079041 K1b1 K1b2 13293 324 11969 K2b1 K1a1 MT079026 Switzerland 2914-2872 BC Switzerland K1c1 EU130564 3316 16093 7559 2019 6260 @152 3416 MT079044 Switzerland 3337-2908 BC 3338-3031 BC 16093 12771 5414 MH605028 Bulgaria E Bronze Age K2a3 K2a5 Swedish 151 3106A 13135 5054 3552 15924 11902 13708 RISE179 12612 8029 KX495759 Swedish 12522 MH605042 Bulgaria E Bronze Age JQ703039 321 12771 11016 K1a3a K1a4a K1b2a K2b1a Massy1375 Sweden 13827 12877 152 16189 MH605046 Bulgaria E Bronze Age France 15799 2831 14208 @10398 MT079103 12501 K1a1a K1a1b Germany 152 15884 Nordic LN 16084 150 8433G 13485 310 16309 8400 11923 16438 BR1 Hungary E Bronze Age RISE97 Massy28 13816 Switzerland 16129 K1a4c K2a5b MT079092 Germany 2197-1981 BC E Bronze Age 16327 13590 310 13928 11840 mur002 JQ705901 16093 MT079042 3106A Kociumaka67 8521 15257 13759 12188 520.4C 16320 MG428997 MG646830 Poland 1462 Sweden Germany 16344 2461-2295 BC 16270 4562 K2b1b 11470 3547 151 KY671048 MT079083 Massy44 13740 Russia Swedish HAL25 Switzerland 11914 Germany 1621 16319 629 MG429024 Lithuania 1709 Nordic LN E Bronze Age 16129 MT079038 K1a3a3 K1b2a1 K1b2a3 K2b1a1 310 @152 9617 3745 @114 15799 Massy80 JQ703347 Russian Germany Germany 5264 Bronze Age Germany 3344-3037 BC 13737 E Bronze Age 16354 16463 JQ705467 Lithuania Kunda 310 13708 6492 @152 6734 16343 12130 3199A 195 4883 5786 523.1CA 9903 16368 8473 15031 11151 Massy47 MG646171 Switzerland K1a1b1 K1a4a1 MT079078 MT079065 16172 16129 3336 Germany Norway 1926-1770 BC E Bronze Age 13710 LBK 15924 Kociumaka68SK2 Massy76 Finn CWC/BAC MH605043 RISE563 8621 13866 15746C MG646831 15853 3591 16527 12130 MG952806 JQ702133 HQ658476 Germany Poland KC907699 2912-2877 BC 195 K1a1a2 K1b1a K1c1c K1c1e K1c1h Germany Switzerland BR2 MH120811 11095 E Bronze Age 16093 MT079062 Switzerland 2905 - 2865 BC 16246T 3106A MT079047 Germany Germany 12019 3935 3290 Bulgaria Germany 8834 JQ705481 MG646826 Poland MG646833 8083 KY670849 15853 Hungary Finn Swedish 3079 280 E Bronze Age France 2029-1892 BC 2206-2126 BC Hungary Poland 12103 16184A 16301 593 9932 5583 Massy86 NE6 3513A MT079101 Switzerland 3036-2923 BC MT079049 Switzerland E Bronze Age E Bronze Age 11654 11266 E Bronze Age Bell Beaker TIM008 Poland Poland MG646829 Poland MG646192 Russian MG952859 HG00326 Finn V9 4825 RISE600 Massy67 2581 5460 K1a4c1 K1b2a1a L Bronze Age 12406 6060 1717G Mina3 2483 12007 Germany Hungary MT079027 Switzerland Late Neolithic 12358 13759 13967 JX273244 MH120689 Russia Poland Poland Hungary HG00177 Finn 7270 7158 11914 @152 Estonia VOR005 Russia Germany 16222 16399 MG952788 K1a1b1e JQ702802 8276.1C 16092 Spain E Bronze Age LBK Switzerland 4295 @152 13153 @146 150 @16093 5031 @152 3106A 3100-2928 BC MT079051 RISE586 France Poland Fatyanovo MG646832 JQ703860 Finn MG646131 12630 KY670895 195 Bronze Age Russia Iron Age E Bronze Age KU188280 Hungary K1a1b1b K1a1b1g K1b1a1 K2b1a1a Finn JQ702781 M Neolithic 3011-2893 BC 15884 325 Massy2 297 3796 14180 @16311 10275 9759 114 10101 Switzerland Czech 16524 Poland JX153567 Finn Poland 13497 Russian 16316 Fatyanovo Swedish K1a1a2a Germany 189 10029 Germany 16222 @10398 8994 195 16042 MT079099 @16093 9758 MF362939 3363-2930 BC Unetice Mina6b Spain M Neolithic JX153261 Finn MH120562 JQ702901 12630 K1a4a1a K1a4a1e K1b2a1a1 789 MF362700 114 E Bronze Age JQ704808 15478 JQ703704 Massy99 RISE154 Switzerland JQ703523 11582 Armenian RISE98 Sweden Nordic LN JX153201 Finn Poland Poland KT763050 @152 K1a4a1f 4959 11620 Armenia @12007 MT079054 Swedish JQ704770 France Germany Poland 1693-1609 BC Italy MF362709 JX153541 Finn Swedish 16355 3314 ya 14659 Switzerland 16093 Poland E Bronze Age Unetice 204 Armenia 310 199 10389 3967 5194 JX153615 Finn JQ702131 K1a1a2a1 K1a1b1b1 16093 3330-3216 BC 10370 Massy93 5021 3300 ya 4252 16168 RISE1 16278 KR919601 Finn Finn Massy149 HG00349 Germany 189 @16093 10310 1440 14560 16284 Poland JQ702926 KX079702 undefined @114 @114 Germany 16245 Finn E Bronze Age 1628 16176 ros003 1672 15884 JQ705686 CWC Swedish @16093 5746 E Bronze Age Massy1192SKA 7948 JQ702341 Sweden 3952 JQ705225 Swedish 16362 93 8817 14757 11392 16320 195 K1a4a1a2 MH120771 JQ704983 Massy1380 Germany KC878725 Norway FBC 4825 Dane KF162649 KC170989 EU262720 HG00185 JQ703696 EU753433 @10398 Poland Swedish Germany E Bronze Age Italy TIM001 Dane Russian undefined Finn Finn Finn JX153240 MG646093 E Bronze Age 9377 @146 Russia JX153574 Finn Poland Massy1414 13401 Fatyanovo Finn K1a4a1a2a Germany K1a4a1a2b E Bronze Age JN559854 Swedish HG00331 Finn JQ702797 Swedish

153 @16093 JQ703763 France e

N1a1a

@152

3336 8452 16147A 11476 16344 N1a1a1 RISE555 Russia 16320 @152 16193 5460 E Bronze Age 11884 BAM25a Troc5 N1a1a1a 14259 Hungary Spain N18 Poland 3600 BCE HAL34 E Neolithic E Neolithic Germany @2702 16154 207 15900 LBK 8164 8485 HAL4 9300 N1a1a1a3 16086 Germany HAL2A LBK N1a1a1a1 16300 Germany MH605039 GU123026 Tatar SZEH4b LBK Bulgaria GU290215 Russian Hungary E Bronze Age LBKT 15299 207 13248A 11167 151 2410 15953T 215 15758 16147A 4769 @16172 EF486519 EF486518 5939 15774 GU290206 4200 JX154061 ber2 2392 4838 Czech Russian 7720 EF660944 India 7783 Finn @16189 Sweden N1a1a1a2 MH120439 KM103656 GU290215 Italy 14587 RISE391 BAC NE7 Poland Balkan Russian 15496 Kazakhstan Hungary 16182 Sintashta L Neolithic MH120558 HAL001 Poland Russia Fatyanovo

473 12498 9333 RISE21 12572 Denmark 14207 Nordic BA 15708 TIM002 Russia Fatyanovo f

T1a1

KAR22A Germany L Neolithic JQ797979 Swedish Massy50 Germany E Bronze Age JQ704249 Belarus pcw110 Poland CWC KF057946 Norway pcw250 Poland CWC pcw350 Poland CWC

310 114 4050 1204 5985 16269 5563 8155 1598 151 10143 2380 @@152 12633T 14377 7829 1209 16305T 16290 15529A 9855 3633 14281 4155 15452 JQ797987 12952 5054 1672 MF362862 RISE484 RISE175 KY782164 JQ797985 Italy JQ705335 KT799550 12308 2347 chy002 Armenian Hungary Sweden Polish Italy France Swedish JQ702853 T1a1b 16256 Russia Vatya Nordic BA Finn TIM005 Iron Age X02 Estonia Bronze Age Russia MG429040 Latvia Bronze Age T1a1e Fatyanovo VIII5 Ingeria Iron Age KC782346 Swedish MG646091 Poland 14980 MH120450 Poland KP959734 Polish MG646244 Poland KF161741 Dane JQ798006 MG646160 Poland KF161569 Dane Swedish MG646121 Poland HM184912 Czech

3651T 16179 198 8093 14470 13781 15323 @@195 13711 16293 @16163 9548 9630 14180 4493 @750 5205G 15852 16344 12028 12733 16365 16292 7148 KJ856762 KF161644 KF161331 KY670996 KY782161 KY671018 DQ358975 9025 Buryat AY339576 Dane Dane VII4 319 9438 Russian Polish Russian 8389 4688 MG646139 MH120719 German 12780 Finland Estonia 16263 15110 T1a1b1 Poland Poland 15709 8427 16319 Iron Age KY782246 513 15355 KM047209 16129 JQ797991 Polish 5177 Poland JQ797994 Italy MH120576 JQ797992 Turkey KM213520 JQ702958 Poland MH120770 Greece Poland KF161685 Ukraine Poland Dane 3952 146 204 9128 6185 16244 KC911473 JQ797995 HM852798 Qashqai AY714036 Iran Azeri KC911624 India Persian

13248

JQ797993 Iran g

T2a1b1

TIM010 Russia Fatyanovo ESP26 Germany CWC

9117 6169 8863 9020 T2a1b1a 16114

KAR001 15110 12741 2396 3075 Russia 15452 WeRuHG NAU001 T2a1b1a2-16292 T2a1b1a1 mur003 Russia Russia Fatyanovo Bronze Age 16093 @146 7855 195 3350 5747 16171 X08 15130 V7 Estonia KY670841 Estonia T2a1b1a1b Bronze Age MH120672 Russian Iron Age Poland 3335 13980

KY782216 Polish h

T2b

MT079067 Switzerland 2105 - 2036 BC LBK2155 Germany LBK N28 Poland 4600 BCE KY782206 Polish KY671007 Russian LBK1992 Germany LBK RISE577 Czech Unetice MG429037 Lithuania Bronze Age MH120599 Poland KY670922 Russian pcw260 Poland CWC MG429032 Lithuania Bronze Age MG646118 Poland MG429049 Sweden TRB JK2285 Finland 1200-1700AD KM047204 Poland

10750 3826 12441 11242G 16362 9254 15758 @7521 14016 41 189 16399 11929 14198 6806 @195 16354 4127 9139 7600 16223 151 15865 3483 7226 146 3075 16509 12811 6026 310 14319 16147 8934 319 @16126 16093 7853 5471 11608 14214 3078 3106A 16297 9861 FJ460555 KT763051 KY670930 IIf JK1958 11563 KY671067 JQ705854 15670 6496 MG429034 ESP24 MH605044 T2b3 T2b5 T2b2 T2b+16362 T2b4 T2b36 T2b1 Tunisia Russian 200 5474 KM047191 MH605036 3579 10586 Russian Estonia Finland @16296 Russian 319 5460 Norway KY275503 16178 GOL001 6999 Lithuania Germany T2b30 MT079104 Bulgaria T2b25 FJ460542 13830 12361 Poland Bulgaria 4191C 16261 Medieval 1700-1900AD 10335 6488 Swedish Russia 8743 Bronze Age M Neolithic France E Bronze Age JQ702978 Germany T2b23 JN419195 16304 @195 3157 T2b34 Tunisia 16145 E Bronze Age 4734 16290 KY671046 15379 14341 KY782236 Fatyanovo N26 M Neolithic 183 4907 151 AY495286 undefined @16304 6221 6182 France @152 16172 16368 9067 6530 11899 MF177171 KY782212 @16294 Russian 15511 Polish RDT002 Poland @195 5040 8251 MH120552 @16311 11377 6857 Poland Polish 15697G KX786650 MF362928 MF362808 Russia 4600 BCE 16169 10398 6515 240 4964 9 AY714016 Poland RISE479 KY670904 16272 Russian Armenian KY782223 Armenian Fatyanovo @16189 T2b3+151 3 5231 189 151 195 15937 6254 3496 5201 9587 8158 10310 4167 T2b2b JN035224 India T2b4+152 T2b4a 8591 @11812 HM583750 Hungary RISE276 7521 Russian Polish KY670969 16094 15016 9947 11969 13999 undefined 16111 8292 France Vatya Denmark @16296 SCY311 RISE349 Russian AY495287 JQ798073 JX153367 JX153444 KF161789 KY670918 13563 13440G @16296 AY495299 undefined @16296 6593 JQ702894 TU478 16327 Nordic LBA Moldova Hungary 7150 @146 199 undefind Italy Dane 199 4152 Dane Dane Russian KY670846 T2b5a MH120622 13743 14162 @16296 Finn Finland RISE374 15836 4-2nd BCE MBA 152 14544 KF161624 7337 9948 Russian Poland 16266 JQ798075 KF161818 Dane 600-1200AD Hungary 16136 @1888 195 4486C Dane 11151 16297 JQ703716 German 16274 Italy 12171 10961 4225 @709 152 KY670935 8668 5375 Maros 10398 9096 T2b3c 16192 189 12192 GQ304744 3202 2759 16354 16192 11914 7754 16192 Russian 10398 EF660922 15440 JX153280 T2b3d 8110 152 Italy JX152896 573.XC 8185 @16304 15245 14693 KT897694 6593 11050 7382 2161 1719 7340 6641 16189 6401 9947 13759 8843 4967 16239 @16294 KY670852 Italy TU593 6305 Italy KC286607 KU296183 Dane 8504 9210 JQ798074 EF660978 Sweden 15287 16222 3918 7269 11872 4977 8566 Russian MG952851 JX153064 Finland 7289 MT079079 Austria JQ798072 KF161779 Irish MH120690 Italy T2b2b1 AY495305 GU183768 Italy EU683970 KY671073 KY670997 8718 KF161544 10101 JX153804 KF161497 JQ798067 KC257393 9950 EU494628 KF161778 Dane 15565 Hungary Italy 1550-1650AD 11887 Germany Iran Dane JK1941 Poland undefined France Ireland Russian Russian KY670998 KY670903 10398 Dane Dane JQ702800 Dane Kuwait undefined 11447 undefined 12360 2116-1926 BC T2b5a1 Finland JQ701832 undefined Russian Russian 11563 JQ702058 UK KY670873 13623 1300-1700AD 16181 France JQ702464 14587 9653 Russian JQ703712 UK JK1932 16201 16187 Switzerland JQ703099 Finland 16320 7854 15848 16290 EU492455 HG00356 France 1300-1700AD JF939816 Irish Finn AY339568 @14233 152 3 @16296 JQ705119 JN032298 JQ705208 JF929911 undefined Finn @14766 undefined Belgium undefined undefined JQ702885 JQ703023 JQ702982 JN084060 10649 JX153599 undefined Swedish Switzerland undefined 16193 Finn JQ705432 UK HG00337 Finn i

W6 X12 Estonia Bronze Age VOR004 Russia Fatyanovo

146 @16292 6197 6635 1173 7444 211 2171G 11339 5964 @194 6345 196 3834 44.1C 310 2195 1927 3075 @16192 8610 8002 @16325 8931 KC911604 3350 13278 @8251 3606 JQ245723 16311 @16292 7964 8279 9920 200 6285 15262 2597 TIM006 8658 8020 @1243 9531 Persian 3483 14869 8271 9139 NOssetin KF146286 KF146289 8093 13934 16360 3330 6340 KF161698 3952 Russia 4499 552 W6a 16124 1717 8131 5348 16249 KC911547 16042 8557 12411 Italian Italian MF362807 15301 8939 14560 Danish TIM009 Fatyanovo 16173 TU569 MG429028 Lithuania CWC/BAC W6c 4491 9055 16086 KC911449 Persian 16260 13638 16304 FJ348172 Armenian 16438 15217 @194 10948 KF162879 KF162005 Russia 16352 Finland ESP16 Germany CWC 1119 10217 11016 12373 16298 Persian KC911370 KF146287 JQ245769 JQ245736 STyrol 15769 3766 HQ423397 Danish Danish Fatyanovo JF902025 1200-1500AD 7852 16154 KP763848 @11947 16093 KC911518 Persian Kuwait Turkey NOssetin 215 505 JF286634 6366 Polish English JX152978 4164 13722 11143 214 SVP5 @16192 Indian @12414 KM986576 Qashqai MK491358 MF362927 Armenian 16093 Finn KX949568 Finn KY671014 14094 479 Russia KP763850 12858 Yemen KC911523 Armenian Armenian @16192 9591 JQ705209 undefined Russian KF451323 16311 Yamnaya Indian 15355 Qashqai 16288 9948 16293 KY670866 Russian Orcadian GQ149695 15803 KY367263 13753 TU675 3939 undefined 16148G Armenian MF362895 Finland 8704 @152 @207 16274 2352 8514 16291 16167 15781 HM156681 Armenian 1200-1500AD MH120506 KY670993 JQ703084 KY782195 15088 MH120803 MH120467 KM101979 Indian TU566 Poland Russian 234 @195 Poland Polish Poland Poland USA Finland KP143828 HG00336 12358 6713 MH120820 1200-1500AD Slovak Finn MH120447 MG646162 Poland @195 Poland Poland MF177148 Poland SupplementarySupplementary FigureFigure 5.5. PartialPartial mitochondrial DNA phylogeneticphylogenetic treestrees with with modern modern and and ancient ancient individualsindividuals added.added. Aa H5b, H5b, b B H6a1a, H6a1a, c J1c1b1a,C J1c1b1a, d K, D e K,N1a1a1a, E N1a1a1a, f T1a1, F g T1a1, T2a1b1, G T2a1b1,h T2b, i W6. H T2b, Numbers I W6. Numberson branches on branchesindicate substituted indicate substituted nucleotide nucleotide positions consistent positions withconsiste Reconstructednt with Reconstructed Sapiens Reference Sapiens ReferenceSequence Sequence(RSRS); only (RSRS) transversions; only transversions are further arespecified further with specified capital with letters capital; @ refersletters to; @ a reversionrefers to a reversion(back-mutations) (back-mutations); .XC, .1C; and.XC .2C, .1C are and insertions .2C are where insertions X stands where for Xunknown stands number. for unknown Modern number. data Modernfrom https://www.ncbi.nlm.nih.gov/genbank/ data from https://www.ncbi.nlm.nih.gov/genbank/, https://www.internationalgenome.org/data, https://www.internationalgenome.org/data; ancient from ; ancientAllentoft from et al. Allentoft 2015, Fernandes et al. 2015, et al. Fernandes 2018, Furtwängler et al. 2018, et al.Furtwängler 2020, Gamba et al.et al.2020, 2014, Gamba Haak etet al.al. 2015, 2014, HaakJuras et et al.al. 2015,2017, JurasKnipper et al. et 2017,al. 2017, Knipper Krzewinska et al. 2017, et al. Krzewinska2018, Linderholm et al. 2018,et al. 2020,Linderholm Malmström et al. et2020, al. Malmström2019, Mittnik et etal. al. 2019, 2018, Mittnik Modi etet al. al. 2019, 2018, Saag Modi et alet 2019,al. 2019, Valdiosera Saag et et al al 2019, 2018, Valdiosera Översti et alet 2019.al 2018, Översti et al 2019.

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