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Post by Admin on Jan 22, 2022 21:27:11 GMT
Figure 2. Principal component and ADMIXTURE analyses’ results. A principal component analysis results of modern West Eurasians with ancient individuals projected onto the first two components (PC1 and PC2), B ADMIXTURE analysis results for a selection of ancient population averages at K9 with ancient individuals projected onto the modern genetic structure. EF – early farmers; HG – hunter-gatherers; LNBA – Late Neolithic/Bronze Age; IA – Iron Age; MA – Middle Ages. Next, we used outgroup f3 and D statistics to compare the genetic affinity of WeRuHG to those of other relevant populations. We found that WeRuHG and EHG are similar in their genetic affinities both to other ancient and to modern populations (Fig. 3A, Supplementary Fig. 2A). On the other hand, when comparing WeRuHG to the later Fatyanovo, we found that WeRuHG shares more with EHG-like populations, Western Siberian HG, ancient Iranians and modern populations from East Asia and Siberia, while Fatyanovo shares more with most ancient European and Steppe populations and modern populations from the Near East, the Caucasus and Europe (Fig. 3B, Supplementary Fig. 2B). Figure 3. Outgroup f3 statistics’ results of comparisons with ancient populations. Outgroup f3 statistics’ values of form f3(Yorubas; study population, ancient) plotted against each other for two study populations (blue and red axis): A Western Russian hunter-gatherers (WeRuHG) and Eastern huntergatherers (EHG), B WeRuHG and Fatyanovo, C Yamnaya Samara (YamSam) and Fatyanovo, D Central Corded Ware Culture (CeCWC) and Fatyanovo. 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. Early farmer ancestry in Fatyanovo Culture individuals Then, we turned to the Bronze Age Fatyanovo Culture individuals and determined that their maternal (subclades of mtDNA hg U5, U4, U2e, H, T, W, J, K, I and N1a) and paternal (chrY hg R1a-M417) lineages (Table 1, Supplementary Fig. 5, Supplementary Data 2) were ones characteristic of CWC individuals elsewhere in Europe13,14,29,32,28. Interestingly, in all individuals for which the chrY hg could be determined with more depth (n=6), it was R1a2-Z93 (Table 1, Supplementary Data 2), a lineage now spread in Central and South Asia, rather than the R1a1-Z283 lineage that is common in Europe38,39. On the PCA, the Fatyanovo individuals (and the Estonian CWC individual) group together with many European Late Neolithic/Bronze Age (LNBA) and Steppe Middle/Late Bronze Age (MLBA) individuals on top of modern Northern and Eastern Europeans (Fig. 2A). This ancient cluster is shifted towards Anatolian and European EF compared to Steppe Early/Middle Bronze Age (EMBA) populations, including the Yamnaya. The same could be seen in ADMIXTURE analysis where the Fatyanovo individuals are most similar to LNBA Steppe ancestry populations from Central Europe, Scandinavia and the Eastern Baltic (Fig. 2B, Supplementary Fig. 1). These populations are composed of the blue “WHG” and yellow “Khanty” component and two brown components maximized in HG from the Caucasus and Iran, similarly to Yamnaya populations. However, the European LNBA populations (including Fatyanovo) also display a green component most frequent in Anatolian and European EF populations, which is not present in the Yamnaya from Russia. We compared the affinities of the Fatyanovo individuals to those of other populations using f3 and D statistics and found that Fatyanovo shares more with European EF populations and modern Near Easterners than Yamnaya_Samara does (Fig. 3C, Supplementary Fig. 2C, Supplementary Fig. 3C). Importantly, this signal can also be seen when using either autosomal or X chromosome positions from the Lazaridis et al. 201631 ancient dataset instead of the autosomal positions of the EBC-chipDB (Supplementary Fig. 3A–B). However, when comparing Fatyanovo to Central_CWC, there were no clear differences in their affinities to different ancient or modern population groups (Fig. 3D, Supplementary Fig. 2D). Furthermore, we confirmed the presence of sex-biased admixture previously seen in CWC individuals from Estonia, Poland and Germany40,41,32,42,43 in the Fatyanovo (Supplementary Fig. 3D). This is also supported by the presence of mtDNA hg N1a in two Fatyanovo individuals – a hg frequent in Linear Pottery Culture (LBK) early farmers, but not found in Yamnaya individuals44,13,29. Since the previous analyses suggested that the genetic makeup of the Fatyanovo Culture individuals was a result of admixture between migrating Yamnaya individuals and contemporary European populations, we used two complementary methods (qpAdm and ChromoPainter/NNLS) to determine the most suitable proxies for the admixing populations and the mixing proportions. The model with the highest p-value and the lowest standard errors for qpAdm had Yamnaya and Levant Neolithic (N) as sources while the model with the smallest residuals for ChromoPainter/NNLS included also WHG, so we are presenting both models for both analyses (Fig. 4A–B). We ran the analyses for Fatyanovo, Central CWC and Baltic CWC and saw that in the results of the more complex model, some of the Yamnaya ancestry from the simpler model got reassigned as WHG but the proportions of Levant N ancestry remained similar (within 4%). We found that although the results from qpAdm and ChromoPainter/NNLS differed somewhat, both showed more Levant N ancestry in Fatyanovo (~22% and ~17%) than in Central/Baltic CWC (~11% and ~10%). What is more, this result is supported by models where Fatyanovo and Central/Baltic CWC are modeled as a mixture of Levant N and Baltic/Central CWC, respectively (Supplementary Fig. 4A).
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Post by Admin on Jan 23, 2022 3:30:36 GMT
Figure 4. qpAdm and ChromoPainter/NNLS results. A models with Yamnaya and Levant Neolithic (Levant N) as sources, B models with Western hunter-gatherers (WHG), Yamnaya and Levant N as sources. CP – ChromoPainer/NNLS; CWC – Corded Ware Culture. Lastly, we looked for closely related individuals in the Fatyanovo Culture sample-set using READ45. There were no confirmed cases of 2nd degree or closer relatives (Supplementary Fig. 4B). Phenotype informative allele frequency changes in Western Russia We imputed the genotypes of 115 phenotype-informative positions connected to diet (carbohydrate, lipid and vitamin metabolism), immunity (response to pathogens, autoimmune and other diseases) and pigmentation (eye, hair and skin). We used previously published Eastern Baltic individuals for comparison27,32,33. Here, we focus on variants associated with pigmentation (39 SNPs of the HIrisPlex-S system), lactase persistence (rs4988235, rs182549; MCM6) and fatty acid metabolism (rs174570C; FADS2-3) (Table 2), the latter three alleles showing a significant increase in frequency through time. Although the results should be interpreted with due caution because of the small sample size, we inferred that the examined WeRuHG individuals carried alleles connected to brown eyes, dark brown to black hair and intermediate or dark skin pigmentation while around a third of the Fatyanovo individuals had blue eyes and/or blond hair. Furthermore, we infer that the frequency of the two alleles associated with lactase persistence (rs4988235, rs182549; MCM6) is 0% in WeRuHG and ~10% in Fatyanovo Culture individuals (similar to Eastern Baltic populations from the same time periods), but has a significant increase to over 40% by the Late Bronze Age in the Eastern Baltic33. On the other hand, an allele connected to an increase in cholesterol and a decrease in triglyceride levels in serum (rs174570C; FADS2–3) significantly rises in frequency by the CWC period – from ~20% in Eastern Baltic and Western Russian HG to ~50% in CWC individuals from the two regions. Interestingly, the frequency of the allele increases further in the subsequent time periods in the Eastern Baltic. Table 2. Phenotype prediction results. Phenotype proportions per period.
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Post by Admin on Jan 23, 2022 19:44:02 GMT
Discussion After the Last Glacial Period, at the end of the 10th and the beginning of the 9th millennium BC, vast areas in the Eastern Baltic, Finland and northern parts of Russia were populated relatively quickly by hunter-gatherer groups’46–49. Flint originating from several places in the Eastern Baltic and the European part of Russia and the similarities in lithic and bone technologies and artefacts suggest the existence of extensive social networks in the forest belt of Eastern and Northern Europe after it was populated50,51,48. This has led to the hypothesis that the descendants of Paleolithic HG from Eastern Poland to the central areas of European Russia took part in the process52,7,53 and remained connected to their origin, creating a somewhat stretched social network8. However, these connections ceased after a few centuries, as can be witnessed from the production of stone tools from mostly local materials, and new geographically smaller social units appeared in the middle of the 9th millennium BC8. Ancient DNA studies of this area have included Mesolithic individuals that are from the 8th millennium BC or younger and reveal two genetic groups: WHG in the Eastern Baltic and EHG in Northwestern Russia14,29,27,32,28,30. However, no human genomes from the settlement period have been published so far, leaving the genetic ancestry/ancestries of the settlers up for discussion. The individual PES001 from around 10,700 cal BC (probably somewhat affected by the freshwater reservoir effect) presented here with 4x coverage provides evidence for EHG ancestry in Northwestern Russia close to the time it was populated. This, in turn, raises the question of the ancestry of the first people of the Eastern Baltic, keeping in mind the shared social network of the two areas at the time of settlement next to the presence of genetically different groups later in time – a question hopefully answered by future studies.
The formation of Fatyanovo Culture is one of the main factors that affected the population, culture and lifestyle of the previously hunter/fisher-gatherer culture of the Eastern European forest belt. The Fatyanovo Culture people were the first farmers in the area and the arrival of the culture has been associated with migration17,21. This is supported by our results as the Stone Age HG and the Bronze Age Fatyanovo individuals are genetically clearly distinguishable. The sample size of our HG is admittedly low, but the three individuals form a genetically homogenous group with previously reported EHG individuals23–28 and the newly reported PES001 is the highest coverage whole-genome sequenced EHG individual so far, providing a valuable resource for future studies. What is more, the Fatyanovo Culture individuals (similarly to other CWC people) have mostly Steppe ancestry, but also some EF ancestry which was not present in the area before and thus excludes the northward migration of Yamnaya Culture people with only Steppe ancestry as the source of Fatyanovo Culture population. The strongest connections for Fatyanovo Culture in archaeological material can be seen with the Middle Dnieper Culture20,54 spread in present-day Belarus and Ukraine55,56. Importantly, the territory of what is now Ukraine is where the most eastern individuals with European EF ancestry and the most western Yamnaya Culture individuals are from based on published genomic data26,57 (Fig. 1, Supplementary Data 1). Furthermore, archaeological finds show that LBK reached Western Ukraine around 5,300 BC58 and the Yamnaya Culture (burial mounds) arrived in Southeastern Europe around 3,000 BC and spread further as far as Romania, Bulgaria, Serbia and Hungary59. These findings suggest present-day Ukraine as the possible origin of the migration leading to the formation of the Fatyanovo Culture and of the Corded Ware cultures in general.
The exact timing of and processes involved in the emergence of the Fatyanovo Culture in European Russia and the local processes following it have also remained unclear. Until recently, the Fatyanovo Culture was thought to have developed later than other CWC groups and over a longer period of time17,20. However, radiocarbon dates published last year21 and the 25 new dates presented here point towards a fast process, similar in time to CWC people reaching the Eastern Baltic and southern Fennoscandia60–62. Importantly, the archaeological cultures are clearly differentiated between the areas. What is more, it has been suggested that the Fatyanovo Culture people admixed with the local Volosovo Culture HG after their arrival in European Russia6,17,36. Our results do not support this as they do not reveal more HG ancestry in the Fatyanovo people compared to other CWC groups or any visible change in ancestry proportions during the period covered by our samples (2,900–2,050 BC).
Finally, allele frequency changes in Western Russia and the Eastern Baltic revealed similar patterns in both areas. Interestingly, the frequencies of alleles in the MCM6 and FADS2–3 genes, which have been hypothesized to have increased due to dietary changes from the Neolithic onwards63,29,34, had significant changes in frequency at different times – the MCM6 alleles by the CWC period and the FADS2–3 allele after it. This points toward a more complex scenario for the onset of strong selection on these alleles than just the arrival of farming, as has already been suggested previously13,29,34,64,28,33.
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Post by Admin on Jan 23, 2022 21:38:35 GMT
Supplementary Notes Supplementary Note 1. Archaeological and anthropological background Veretye culture 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.
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Post by Admin on Jan 24, 2022 2:01:30 GMT
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 plantbased 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 5 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)
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