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PCA and ADMIXTURE analysis reflecting three time periods in Northern European prehistory. (a) Principal components analysis of 1007 presentday West Eurasians (grey points, modern Baltic populations in dark grey) with 282 projected published ancient and 24 ancient north European samples introduced in this study (marked with a red outline). Population labels of modern West Eurasians are given in Supplementary Information Fig. 2 and a zoomed-in version of the European Late Neolithic and Bronze Age samples is provided in Extended Data Fig. 4. (b) Ancestral components in ancient individuals estimated by ADMIXTURE (k=11). 

PCA and ADMIXTURE analysis reflecting three time periods in Northern European prehistory. (a) Principal components analysis of 1007 presentday West Eurasians (grey points, modern Baltic populations in dark grey) with 282 projected published ancient and 24 ancient north European samples introduced in this study (marked with a red outline). Population labels of modern West Eurasians are given in Supplementary Information Fig. 2 and a zoomed-in version of the European Late Neolithic and Bronze Age samples is provided in Extended Data Fig. 4. (b) Ancestral components in ancient individuals estimated by ADMIXTURE (k=11). 

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Recent ancient DNA studies have revealed that the genetic history of modern Europeans was shaped by a series of migration and admixture events between deeply diverged groups. While these events are well described in Central and Southern Europe, genetic evidence from Northern Europe surrounding the Baltic Sea is still sparse. Here we report genome-w...

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... constructed from 1007 modern individuals from a diverse set of West Eurasian contemporary populations. We also projected previously published ancient samples 4 onto this plot and investigated the relationship of ancient Baltics to other ancient populations in Europe. The two samples from Karelia cluster with previously published Mesolithic EHG ( Fig. 2A), exhibit similar composition of their genetic makeup (Fig. 2b) and share most genetic drift since divergence from Africa with EHG as shown by outgroup f3- statistics (Extended Data Figures 1 and 2). We therefore grouped them together with EHG for all following analyses. Model-based clustering using ADMIXTURE also . CC-BY-NC-ND 4.0 ...
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... West Eurasian contemporary populations. We also projected previously published ancient samples 4 onto this plot and investigated the relationship of ancient Baltics to other ancient populations in Europe. The two samples from Karelia cluster with previously published Mesolithic EHG ( Fig. 2A), exhibit similar composition of their genetic makeup (Fig. 2b) and share most genetic drift since divergence from Africa with EHG as shown by outgroup f3- statistics (Extended Data Figures 1 and 2). We therefore grouped them together with EHG for all following analyses. Model-based clustering using ADMIXTURE also . CC-BY-NC-ND 4.0 International license not peer-reviewed) is the author/funder. It ...
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... copyright holder for this preprint (which was . http://dx.doi.org/10.1101/113241 doi: bioRxiv preprint first posted online Mar. 3, 2017; shows that EHG carry a genetic component (green component in Fig. 2b) that is maximized in hunter-gatherers from the Caucasus (CHG) and shared with Neolithic farmers from Iran and Steppe populations from the Bronze Age, suggesting some common ancestry for these populations, consistent with previous results 21 . Despite its geographically vicinity to EHG, the eastern Baltic individual associated with the ...
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... appear intermediate between WHG/Kunda and EHG in PCA space, and the statistic D(SHG,WHG; EHG, Mbuti) is strongly significant for excess allele sharing of SHG and EHG (Z=12.8). Neither the Kunda individual nor SHG exhibit the major ADMIXTURE component shared between EHG and CHG (green in Figure 2b), bringing into question a direct contribution of EHG into the Mesolithic individuals from Scandinavia and the eastern Baltic. However, using qpWave we cannot reject the previously published result of SHG being formed by admixture of WHG and EHG 6 (58±2% WHG with 42±2% EHG; p=0.55; ...
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... WHG with 42±2% EHG; p=0.55; Supplementary Information Table S7). Both EHG and SHG share a non-negligible component in ADMIXTURE analysis that is maximized in some modern Native American populations which points towards ancient North Eurasian ancestry, as represented by the MA1 and AG3 samples from Palaeolithic Siberia 13 (crimson component in Fig. 2a). Indeed, D-statistics show that EHG and SHG share significantly more alleles with MA1 and AG3 than both Baltic and Western HG (Supplementary Information Table S9). Additionally, mtDNA haplogroups found among EHG point toward an eastern influence: R1b in UzOO77 was previously found in the Palaeolithic Siberian AG3 5 and a haplogroup ...
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... results for the Kunda individual are mirrored in the four later eastern Baltic Neolithic hunter-gatherers of the Narva culture (Fig. 2) and further supported by the lack of significantly positive results for the D-statistic D(Narva, Kunda; X, Mbuti) (Supplementary Information Table S2) demonstrating population continuity at the transition from Mesolithic to Neolithic, which in the eastern Baltic region is signified . CC-BY-NC-ND 4.0 International license not ...
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... contrast to the Eastern Baltic, we see clear evidence of a movement of agriculturalists into Southern Sweden already around 4,000 calBCE. The individuals associated with the Early Neolithic TRB culture (EN TRB) cluster with Middle and Early Neolithic farmers from Europe on the PCA ( Fig. 2A) and in the ADMIXTURE analysis exhibit the component maximized in Levantine early farmers (orange component in Figure 2b). The statistic D(EN_TRB, Middle Neolithic Central Europe; X, Mbuti) does not yield significantly positive results and EN TRB can be modeled as derived from a single source identical with Middle Neolithic (MN) Central ...
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... contrast to the Eastern Baltic, we see clear evidence of a movement of agriculturalists into Southern Sweden already around 4,000 calBCE. The individuals associated with the Early Neolithic TRB culture (EN TRB) cluster with Middle and Early Neolithic farmers from Europe on the PCA ( Fig. 2A) and in the ADMIXTURE analysis exhibit the component maximized in Levantine early farmers (orange component in Figure 2b). The statistic D(EN_TRB, Middle Neolithic Central Europe; X, Mbuti) does not yield significantly positive results and EN TRB can be modeled as derived from a single source identical with Middle Neolithic (MN) Central Europe (p=0.26; ...
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... the EN TRB, with no significant positive results for D(MN_TRB, EN_TRB; X, Mbuti) and qpWave analysis using only EN TRB as source for MN TRB are not rejected (p=0.86). The PWC individuals, who were contemporaneous to the MN TRB but relied mainly on marine resources, appear intermediate between SHG and Middle Neolithic farming cultures on the PCA ( Fig. 2A) and the only models not rejected with qpWave involve a two-way admixture between SHG and EN TRB (91±3% SHG and 9±4% EN TRB Supplementary Information Table S7). By this model, PWC is largely genetically continuous to SHG, which is congruent with their similarities in economy but somewhat at odds with archaeological models indicating ...
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... individual from Sope, Estonia 3 . All Baltic Late Neolithic (LN) individuals (ca. 3,200 to 1,750 calBCE) fall in PCA space in the diffuse European LNBA cluster formed by individuals admixed between Early and Middle Bronze Age (EMBA) pastoralists from the Yamnaya culture of the eastern Pontic Steppe and Middle Neolithic European farmers ( Fig. 2A) and carry the genetic component that was introduced into Europe with this pastoralist migration (green in Fig. 2B). This impact is also reflected . CC-BY-NC-ND 4.0 International license not peer-reviewed) is the author/funder. It is made available under ...
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... in PCA space in the diffuse European LNBA cluster formed by individuals admixed between Early and Middle Bronze Age (EMBA) pastoralists from the Yamnaya culture of the eastern Pontic Steppe and Middle Neolithic European farmers ( Fig. 2A) and carry the genetic component that was introduced into Europe with this pastoralist migration (green in Fig. 2B). This impact is also reflected . CC-BY-NC-ND 4.0 International license not peer-reviewed) is the author/funder. It is made available under ...
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... is dated to the very beginning of the LN, are instead consistent with being derived from the same source as EMBA Steppe pastoralists (p=0.41 and p=0.19, respectively; Supplementary Information Table S4), which corresponds with their ADMIXTURE profiles that lack the early farmer component also missing in EMBA Steppe samples (orange component in Fig. 2b). Coinciding with this steppe- like genetic influx is the first evidence of animal husbandry in the eastern Baltic 15 , suggesting import of this technology by an incoming steppe-like pastoralist population independent of the agricultural societies that were already established to the south and west. Furthermore, the individual ...
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... by an incoming steppe-like pastoralist population independent of the agricultural societies that were already established to the south and west. Furthermore, the individual Spiginas2, which is dated to the very end of the Late Neolithic, has a higher proportion of the hunter-gatherer ancestry, as seen in ADMIXTURE (darker blue component in Fig. 2b), and is estimated to be admixed between 78±4% Central European CWC and 22±4% Narva (Supplementary Information Table S6). A reliance on marine resources persisted especially in the north-eastern Baltic region until the end of the Late Neolithic 29 and in combination with the proposed large population size for Baltic hunter-gatherers a ...
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... Neolithic 2,30 . We observe that the increased affinity to Baltic hunter-gatherers continues in the more recent samples from the Baltic BA (dated between ca. 1,000 and 230 calBCE), that cluster together on the PCA in the same space occupied by modern Lithuanians and Estonians, shifted from other Europeans to WHG and Baltic hunter-gatherers ( Fig. 2A). The statistic D(Baltic_BA, Baltic_LN; Narva, Mbuti) is strongly significant (Z=11.7; Supplementary Information Table S2) demonstrating the increase in allele sharing with local hunter-gatherers in the Baltic populations after the Late Neolithic. Additionally, D-statistics provided significantly positive results for D(Baltic_BA, ...
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... BA gene pool was not completed by admixture between the Baltic CWC population and foragers but involved additional gene-flow from outside the Baltic territory. Direct evidence of this exists in the Baltic BA sample that appears as an outlier on the PCA, falling within the larger European LNBA cluster instead of the tight Baltic BA cluster ( Fig. 2a) and archaeological evidence supports that the site Kivutkalns, which is represented by eight of our individuals, was a large . 3, 2017; bronze-working center located on a trade route that opened to the Baltic Sea on the west and led inland following the Daugava river 31 . The Baltic BA was furthermore the first eastern Baltic ...
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... the regional variant of the CWC, while foraging persisted as an important form of subsistence. The remains were found without any associated artifacts, but in close proximity to a site where the assemblage showed a mix between local hunter-gatherer traditions and CWC influence 32 . On the PCA this sample falls within the European LNBA cluster ( Fig. 2A) and similarly to other individuals from this cluster displays the three components derived from WHG, CHG and Neolithic Levant (Fig. 2b) providing genomic evidence for the influence of both the early Neolithic and LNBA expansions having reached as far as northern Sweden in the third millennium. qpWave rejects both Scandinavia LNBA and ...
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... artifacts, but in close proximity to a site where the assemblage showed a mix between local hunter-gatherer traditions and CWC influence 32 . On the PCA this sample falls within the European LNBA cluster ( Fig. 2A) and similarly to other individuals from this cluster displays the three components derived from WHG, CHG and Neolithic Levant (Fig. 2b) providing genomic evidence for the influence of both the early Neolithic and LNBA expansions having reached as far as northern Sweden in the third millennium. qpWave rejects both Scandinavia LNBA and Baltic LN as single sources for Olsund and all tested models of two-way admixture between Baltic LN or Scandinavia LNBA and other ...
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... the close clustering of modern eastern Baltic populations with Baltic BA on the PCA plot and Lithuanians and Estonians exhibiting the highest allele sharing for ancient Baltic populations with any modern population (Extended Data Figure 2), Baltic BA as a single source for either modern Lithuanians or Estonians is rejected (Supplementary Information Table S4). The statistic D(Lithuanian, Baltic_BA; X, Mbuti) reveals significant positive results for many modern Near Eastern and Southern European populations which can be caused by Lithuanians having received more genetic input from populations with higher farmer ancestry after the Bronze Age (Supplementary Information Table S8). ...
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... analyses involving modern populations we restrict to the intersection of 597,503 SNPs between the 1240k SNP set and the HO dataset. Principle component analysis (PCA) was performed with smartpca in the EIGENSOFT package 54 by constructing the eigenvectors from modern west Eurasian populations (Supplementary Information Figure S2) and projecting ancient individuals on these eigenvectors (Figure 2a, Extended Data Figure 4). Admixture analysis (Figure 2b) was carried out with ADMIXTURE on 3769 modern and 366 ancient individuals for ancestral populations k=2 to k=20. ...
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... analyses involving modern populations we restrict to the intersection of 597,503 SNPs between the 1240k SNP set and the HO dataset. Principle component analysis (PCA) was performed with smartpca in the EIGENSOFT package 54 by constructing the eigenvectors from modern west Eurasian populations (Supplementary Information Figure S2) and projecting ancient individuals on these eigenvectors (Figure 2a, Extended Data Figure 4). Admixture analysis (Figure 2b) was carried out with ADMIXTURE on 3769 modern and 366 ancient individuals for ancestral populations k=2 to k=20. ...
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... component analysis (PCA) was performed with smartpca in the EIGENSOFT package 54 by constructing the eigenvectors from modern west Eurasian populations (Supplementary Information Figure S2) and projecting ancient individuals on these eigenvectors (Figure 2a, Extended Data Figure 4). Admixture analysis (Figure 2b) was carried out with ADMIXTURE on 3769 modern and 366 ancient individuals for ancestral populations k=2 to k=20. The SNP dataset was pruned for linkage disequilibrium with PLINK using the parameters --indep- pairwise 200 25 0.5. ...
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... SNP dataset was pruned for linkage disequilibrium with PLINK using the parameters --indep- pairwise 200 25 0.5. We considered the cross-validation (CV) error and report in Figure 2b the results of k=11 where the CV error levels out at a minimum. To quantify population affinities and admixtures suggested in the PCA and ADMIXTURE analysis, we carried out f-statistics using the programs qp3Pop and qpDstat in the ADMIXTOOLS suite (https://github.com/DReichLab) ...

Citations

... In many ways, the PWC seems to represent a mix of traits from both farmer and hunter-gatherer groups, around the Baltic. A similar mix of farmer and non-farmer is suggested by the genetic analyses of PWC individuals from the eastern Swedish island of Gotland in the Baltic (Skoglund et al. 2014, Mittnik et al. 2017. ...
Article
The Pitted Ware Culture emerged during the Neolithic around 3400 BCE in east-central Sweden and quickly spread south and west to various parts of southern Scandinavia. For the next millennium these groups exhibited less interest in agricultural activity and a general return to the hunting and gathering of wild animals and plants with a focus on seals in some coastal areas. Pitted Ware Culture arrived in northeastern Denmark after 3100 BCE. One of the most important sites of this period was at Kainsbakke, on the peninsula of Djursland. One deep pit at the site contained unusual numbers of bear, red deer, European elk, and boar remains. This study focuses on the isotopic proveniencing of some of these animal remains to evaluate their origins. Confirmation of the import of some of the animals, probably from western Sweden, suggests possible shared ritual activity at the central Kainsbakke site. This evidence also confirms the navigating skills of Neolithic peoples in northern Europe.
... More than a century of research on the Corded Ware Culture (CWC) has resulted in the discovery and study of numerous settlement and burial sites in Europe. With the increased application of biomolecular analyses Frei et al., 2015;Haak et al., 2015;Rasmussen et al., 2015;Sjögren et al., 2016;Valtueña et al., 2017;Mittnik et al., 2017;Saag et al., 2017) within the last few years large scale migrations, the presence of the early plague bacteria Yersina pestis, signs of exogamy, and renewed theories regarding language dispersal (Kristiansen et al., 2017) have been shown among the peoples of the Corded Ware Complexes. These studies provide a larger framework for the CWC research and lay a foundation for more in-depth case studies that further understanding on the complexities of these societies. ...
Article
We present an individual biography of an adult woman from the 3rd millennium BC from the Eastern Baltic. Being a representative of a Corded Ware Culture she is considered one of the first documented cases carrying the early plague bacteria, Yersina pestis. The appearance, life, and death of this individual from Sope, NE Estonia, is provided through an application of a range of osteological and biomolecular analyses. The mortuary practices that accompanied her death are revealed through post-excavation archaeothanatological analysis. The position of bones within the grave indicates that her remains were handled differently from the known contemporary Corded Ware Culture inhumations. The mortuary treatment consisted of at least two phases that resulted in an imitation of a ‘proper’ flexed burial. Probably, the reburial of bones was undertaken during the Corded Ware period.
... The Cap-C58-1 sample contains all defining mutations for H11a1, a descendant subclade of the common western Eurasian H11 haplogroup, as well as a private mutation 16261 T. Similar genetic profiles are currently distributed across most of the northern, central and eastern Europe and in Central Asia. H11a was identified in a Mesolithic hunter-gatherer of the Narva culture from Lithuania (Spiginas1) 26 , in a Bronze Age sample from Hungary (Rise247) 2 and was also found associated to the Unetice culture (BZH1) 27 . H11a1 was found in a medieval sample belonging to an adult female of the Hungarian Conquest period 9 , but it is not an exact match to the C58-1 sample (Fig. 1). ...
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The historical province of Dobruja, located in southeastern Romania, has experienced intense human population movement, invasions, and conflictual episodes during the Middle Ages, being an important intersection point between Asia and Europe. The most informative source of maternal population histories is the complete mitochondrial genome of archaeological specimens, but currently, there is insufficient ancient DNA data available for the medieval period in this geographical region to complement the archaeological findings. In this study, we reconstructed, by using Next Generation Sequencing, the entire mitochondrial genomes (mitogenomes) of six medieval individuals neglectfully buried in a multiple burial from Capidava necropolis (Dobruja), some presenting signs of a violent death. Six distinct maternal lineages (H11a1, U4d2, J1c15, U6a1a1, T2b, and N1a3a) with different phylogenetic background were identified, pointing out the heterogeneous genetic aspect of the analyzed medieval group. Using population genetic analysis based on high-resolution mitochondrial data, we inferred the genetic affinities of the available medieval dataset from Capidava to other ancient Eurasian populations. The genetic data were integrated with the archaeological and anthropological information in order to sketch a small, local piece of the mosaic that is the image of medieval European population history.
... Today, however, this ethnic paradigm (the so-called continuity theory) has been seriously challenged by historical linguistics (e.g. Kallio, 2006;Häkkinen, 2009), archaeo-genetics (Mittnik et al., 2017;Saag et al., 2017;Saag et al., in preparation), and archaeology (Lang, 2015;2018). New data and ideas in all these fields of research cast new light also on the question of fortified settlements. ...
... In the case of the East Baltic Bronze Age, the immigration theory has recently received scientific support from ancient DNA studies. Mittnik et al. 2017 have pointed out that although the Bronze Age population in the East Baltic 6 shows an increased affinity to (western) hunter-gatherers, its genetic composition cannot be explained merely by the admixture between the local Corded Ware people and foragers but involved an additional gene flow from outside the East Baltic territory. On the other hand, although the East-Baltic Bronze-Age population stands genetically closer to the modern population of this region than the Stone Age groups, there have still been important additions afterwards -in the case of the Lithuanians from the south-west and in the case of the Estonians from the east. ...
... At least for the region north of the Daugava river this possible immigration, as suggested by both genetic and linguistic studies, had to come from the east/south-east. 6 The East-Baltic Bronze-Age population -the aDNA of which was analysed by Mittnik et al. 2017 -was represented by ten persons from the Ķivutkalns cemetery (dated to 810-230 BC) and four persons from Turlojiškė (No. 3, dated to 1010-800 BC). ...
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[full article, abstract in English; abstract in Lithuanian] A brief history of research and earlier interpretations of fortified settlements east of the Baltic Sea are provided in the first part of the article. The earlier research has resulted in the identification of the main area of the distribution of fortified settlements, the main chronology in the Late Bronze and Pre-Roman Iron Ages, and their general cultural and economic character. It has been thought that the need for protection – either because of outside danger or social tensions in society – was the main reason for the foundation of fortified sites. The second part of the article adds a new possibility of interpreting the phenomenon of fortified settlements, proceeding from ethnogenesis of the Finnic and Baltic peoples. It is argued that new material culture forms that took shape in the Late Bronze Age – including fortified settlements and find assemblages characteristic of them – derived at least partly from a new population arriving in several waves from the East-European Forest Belt.
... The genetic structure of Europeans today is the result of several layers of migration and subsequent admixture. The incoming source populations no longer exist in unadmixed form, but have been identified using ancient DNA in several studies over the last few years [1][2][3][4][5][6][7][8] . Broadly, present-day Europeans have ancestors in three deeply diverged source populations: European hunter-gatherers who settled the continent in the Upper Paleolithic, Europe's first farmers who expanded from Anatolia across Europe in the early Neolithic starting around 8,000 years ago, and groups from the Pontic Steppe that arrived in Europe during the final Neolithic and early Bronze Age~4,500 years ago. ...
... We merged these 11 individuals with 3,333 published present-day individuals genotyped on the Affymetrix Human Origins platform and 511 ancient individuals sequenced using a mixture of DNA capture and shotgun sequencing. [1][2][3][4]8,22,23 . ...
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European history has been shaped by migrations of people, and their subsequent admixture. Recently, evidence from ancient DNA has brought new insights into migration events that could be linked to the advent of agriculture, and possibly to the spread of Indo-European languages. However, little is known so far about the ancient population history of north-eastern Europe, in particular about populations speaking Uralic languages, such as Finns and Saami. Here we analyse ancient genomic data from 11 individuals from Finland and Northwest Russia. We show that the specific genetic makeup of northern Europe traces back to migrations from Siberia that began at least 3,500 years ago. This ancestry was subsequently admixed into many modern populations in the region, in particular populations speaking Uralic languages today. In addition, we show that ancestors of modern Saami inhabited a larger territory during the Iron Age than today, which adds to historical and linguistic evidence for the population history of Finland.
... Though population genetic analysis indicates some connection of the Conqueror European component to modern Finnish (Fin) and Baltic people (Balt=Latvians+ Lithuanians+Estonians) ( Table 1), but no relation to Saamis (Sam), Mansis and Kanthys (Yug) ( Table 1, S9 Table). The Baltic relation of the European component seems to appear already in the Baltic Bronze Age (BalBA, 1000-230 BCE), [86] measured with the SHD method (Table 1, sequence data are not yet available). BalBA genomes cluster with modern Lithuanians and Estonians, and lack Eastern mtDNA Hg-s and Y-chromosomal haplogroup N-tat, which is typical for Uralic speaking groups, thus Estonians must have received their East Asian-Siberian components after the BalBA period, from a different source [86]. ...
... The Baltic relation of the European component seems to appear already in the Baltic Bronze Age (BalBA, 1000-230 BCE), [86] measured with the SHD method (Table 1, sequence data are not yet available). BalBA genomes cluster with modern Lithuanians and Estonians, and lack Eastern mtDNA Hg-s and Y-chromosomal haplogroup N-tat, which is typical for Uralic speaking groups, thus Estonians must have received their East Asian-Siberian components after the BalBA period, from a different source [86]. According to our data BalBA is best admixed from the closely related Scandinavian Neolith-Bronze Age (NNBA), Afanasevo and European Neolithic populations (S13 Table), so it is unlikely connected to Finno-Ugric groups. ...
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Endre Neparáczki and Zoltán Maróti contributed equally to this study. Abstract It has been widely accepted that the Finno-Ugric Hungarian language, originated from proto Uralic people, was brought into the Carpathian Basin by the Hungarian Conquerors. From the middle of the 19 th century this view prevailed against the deep-rooted Hungarian Hun tradition, maintained in folk memory as well as in Hungarian and foreign written medieval sources, which claimed that Hungarians were kinsfolk of the Huns. In order to shed light on the genetic origin of the Conquerors we sequenced 102 mitogenomes from early Conqueror cemeteries and compared them to sequences of all available databases. We applied novel population genetic algorithms, named Shared Haplogroup Distance and MITOMIX, to reveal past admixture of maternal lineages. Phylogenetic and population genetic analysis indicated that more than one third of the Conqueror maternal lineages were derived from Central-Inner Asia and their most probable ultimate sources were the Asian Huns. The rest of the lineages most likely originated from the Bronze Age Potapovka-Poltavka-Srubnaya cultures of the Pontic-Caspian steppe, which area was part of the later European Hun empire. Our data give support to the Hungarian Hun tradition and provides indirect evidence for the genetic connection between Asian and European Huns.
... However, studies of another lithic technology, the "pressure blade" technique, which first occurred in the northern parts of Scandinavia around 10,200 cal BP, indicates contact with groups in the east and possibly an eastern origin of the early settlers [7,[12][13][14][15] (S1 Text). The first genetic studies of Mesolithic human remains from central and eastern Scandinavian hunter-gatherers (SHGs) revealed similarities to two different Mesolithic European populations, the "western huntergatherers" (WHGs) from western, central, and southern Europe and the "eastern hunter-gatherers" (EHGs) from northeastern and eastern Europe [16][17][18][19][20][21][22][23][24]. Archaeology, climate modeling, and genetics suggest several possibilities for the early postglacial migrations into Scandinavia, including migrations from the south, southeast, northeast, and combinations of these; however, the early postglacial peopling of Scandinavia remains elusive [1,4,[6][7][8][9][10][11][12][13][14][15][16][17][18][19]25,26]. ...
... One individual, SF11, seems to be a slight genetic outlier in the principal component analysis (PCA), which could be due to the lower coverage or driven by nuclear contamination (Table 1, S6 Text). Generally, the pattern of dual ancestry is consistent with a scenario in which SHGs represent a mixed group tracing parts of their ancestry to both the WHGs and the EHGs [17][18][19]22,24,40]. The SHGs from northern and western Scandinavia show a distinct and significantly stronger affinity to the EHGs compared to the central and eastern SHGs (Fig 1). ...
... From about 11,700 cal BP, consistent archaeological evidence of human presence exists in southern Scandinavia following the retreat of the ice sheet [6,41,42] (S1 Text). Artifacts and tools found at these sites show similarities with the Ahrensburgian tradition of northern central Europe [15,43], suggesting that these hunter-gatherers likely had a southern origin from a WHG-like gene pool as no EHG ancestry has been found in central and western Europe [18,21,24,27]. Although this genetic component would have entered from today's northern Germany and Denmark (Fig 2, Scenario a), it remains unclear how and where the EHG component entered Scandinavia (Fig 2, Scenarios b, c and/or d). ...
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Scandinavia was one of the last geographic areas in Europe to become habitable for humans after the Last Glacial Maximum (LGM). However, the routes and genetic composition of these postglacial migrants remain unclear. We sequenced the genomes, up to 57× coverage, of seven hunter-gatherers excavated across Scandinavia and dated from 9,500–6,000 years before present (BP). Surprisingly, among the Scandinavian Mesolithic individuals, the genetic data display an east–west genetic gradient that opposes the pattern seen in other parts of Mesolithic Europe. Our results suggest two different early postglacial migrations into Scandinavia: initially from the south, and later, from the northeast. The latter followed the ice-free Norwegian north Atlantic coast, along which novel and advanced pressure-blade stone-tool techniques may have spread. These two groups met and mixed in Scandinavia, creating a genetically diverse population, which shows patterns of genetic adaptation to high latitude environments. These potential adaptations include high frequencies of low pigmentation variants and a gene region associated with physical performance, which shows strong continuity into modern-day northern Europeans.
... Enamel δ 13 C is relevant to diet, and reflects the entire diet (in this case in early child- hood), in contrast to collagen values which are mainly influenced by protein intake. A comparison with values from Middle Neolithic adult bone samples from the passage grave at Frälsegården (Sjögren/Price 2013;Sjögren 2015;2017) shows that the two early Neolithic women have considerably lower δ 15 N values than most of the Frälsegården population (Tab. 4, Fig. 11). ...
... All three were dated to the EN TRB period. A recent aDNA analysis on individual 3 showed genetic similarity to Central European Early Neolithic individuals, as well as to Middle Neolithic TRB individuals from Sweden (Mittnik et al. 2017). The animal bones were dominated by cattle, sheep/goat and pigs, with minor occurrences of other species. ...
... It was not accompanied by any artefacts, but an association with the Battle Axe culture is possible. Genetic evidence places it on the border between Corded Ware and Late Neolithic individuals (Mittnik et al. 2017). ...
... 25 ). Ancient DNA 26,27 and archaeological evidence 28 suggests that the transition occurred as farming communities to the south responded to the warming climate by expanding north into southern Scandinavia. By around 5,600 cal. ...
Article
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The transition from hunter-gatherer-fisher groups to agrarian societies is arguably the most significant change in human prehistory. In the European plain there is evidence for fully developed agrarian societies by 7,500 cal. yr BP, yet a well-established agrarian society does not appear in the north until 6,000 cal. yr BP for unknown reasons. Here we show a sudden increase in summer temperature at 6,000 cal. yr BP in northern Europe using a well-dated, high resolution record of sea surface temperature (SST) from the Baltic Sea. This temperature rise resulted in hypoxic conditions across the entire Baltic sea as revealed by multiple sedimentary records and supported by marine ecosystem modeling. Comparison with summed probability distributions of radiocarbon dates from archaeological sites indicate that this temperature rise coincided with both the introduction of farming, and a dramatic population increase. The evidence supports the hypothesis that the boundary of farming rapidly extended north at 6,000 cal. yr BP because terrestrial conditions in a previously marginal region improved.
... BP. Ancient DNA evidence has recently shown that it involved an expansion of farming populations from further south [2][3] . In terms of the economy, farming is indicated by the presence of cereal remains and the bones of domestic animals. ...
... Like the TRB this is an agricultural culture, though with probably more emphasis than the latter on domestic animal exploitation. Like the latter, it represents at least in part an expansion of new populations into southern Scandinavia 3,15 . Recent stable isotope and lipid work on ceramic residues from Corded Ware sites in southern Finland 13 dated ca. ...