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Am. 4, but show both models in Supplementary Data4. For downstream analyses, we used bamutils (version 1.0.13, https://github.com/statgen/bamUtil.git) TrimBam to trim two bases at the start and end of all reads. Torroni, A. et al. Li, H. et al. 23, 553559 (2013). K.M. Department of Archaeogenetics, Max Planck Institute for the Science of Human History, 07745, Jena, Germany, Thiseas C. Lamnidis,Kerttu Majander,Choongwon Jeong,Elina Salmela,Wolfgang Haak,Johannes Krause&Stephan Schiffels, Institute for Archaeological Sciences, Archaeo- and Palaeogenetics, University of Tbingen, 72070, Tbingen, Germany, Department of Biosciences, University of Helsinki, PL 56 (Viikinkaari 9), 00014, Helsinki, Finland, Kerttu Majander,Elina Salmela&Pivi Onkamo, The Eurasia3angle Project, Max Planck Institute for the Science of Human History, 07745, Jena, Germany, Department of Cultures, Archaeology, University of Helsinki, PL 59 (Unioninkatu 38), 00014, Helsinki, Finland, Peter the Great Museum of Anthropology and Ethnography (Kunstkamera), Russian Academy of Sciences, University Embankment, 3, Saint Petersburg, 199034, Russia, Vyacheslav Moiseyev&Valery Khartanovich, Vavilov Institute of General Genetics, Ulitsa Gubkina, 3, Moscow, 117971, Russia, Research Centre for Medical Genetics, Moskvorechye Ulitsa, 1, Moscow, 115478, Russia, Biobank of North Eurasia, Kotlyakovskaya Ulitsa, 3 12, Moscow, 115201, Russia, Max Planck Institute for Evolutionary Anthropology, Deutscher Pl. All qpWave and qpAdm models were run using the option allsnps: YES. These results suggest that the geographic range of the Saami extended further south in the past, and points to a genetic shift at least in the western Finnish region since the Iron Age. 19, 16551664 (2009). AllSaami corresponds to a population consisting of the two genomes from the SGDP and the genome from this study (ModernSaami). The Native-American-related ancestry seen in the EHG and Bolshoy corresponds to a previously reported affinity towards Ancient North Eurasians . B Biol. Additionally, we gain further insights into the genetic history of the Saami in Finland, by showing that during the Iron Age, close genetic relatives of modern Saami lived in an area much further south than their current geographic range. A common variation in EDAR is a genetic determinant of shovel-shaped incisors. Patterson, N., Price, A. L. & Reich, D. Population structure and eigenanalysis. Nat. We analysed low coverage genomes from four additional individuals of the Levnluhta site using PCA (Supplementary Figure3), confirming the exclusive position of Levnluhta_B compared to all other six individuals (including the four low-coverage individuals) from that site, as is consistent with the ADMIXTURE and qpAdm results. This Siberian ancestry was subsequently admixed into many modern populations in the region, particularly into populations speaking Uralic languages today. Chuan-Chao Wang, Hui-Yuan Yeh, David Reich, Eirini Skourtanioti, Harald Ringbauer, Philipp W. Stockhammer, Choongwon Jeong, Oleg Balanovsky, Johannes Krause, Daniel M. Fernandes, Alissa Mittnik, David Reich, Simone Andrea Biagini, Neus Sol-Morata, Francesc Calafell, Perle Guarino-Vignon, Mal Lefeuvre, Cline Bon, Ludovica Molinaro, Francesco Montinaro, Luca Pagani, Martin Sikora, Vladimir V. Pitulko, Eske Willerslev, Nature Communications Am. ALDER provides a relative date estimate for a single-pulse admixture event in generations. In short, this program provides an automated search through the Y haplogroup tree (as provided within yHaplo, as accessed from ISOGG on 04 Jan 2016) from the root to the downstream branch based on the presence of derived alleles and assigns the most downstream haplogroup with derived alleles. The novel genome-wide data presented here from ancient individuals from Finland opens new insights into Finnish population history. Hear this story. Y-chromosome analysis of ancient Hungarian and two modern Hungarian-speaking populations from the Carpathian Basin. The Yamnaya-related steppe ancestry has been described as a mixture of Eastern- and Caucasus hunter-gatherers from the Pontic-Caspian steppes, dating to 3,300-2,600 BCE, which eventually spread further to the Altai region in the East in the form of people associated to the Afanasievo Culture. Uralic-speaking populations are highlighted in dark purple. The steppe ancestry component within modern Europeans (green), which is associated with the Yamnaya population, is maximised in ancient Iranian populations and to a lesser extent Caucasus hunter-gatherers (CHG). Nat Commun 9, 5018 (2018). Results from the least complex model for each test population/individual are shown. Rohland, N., Harney, E., Mallick, S., Nordenfelt, S. & Reich, D. Partial uracil-DNA-glycosylase treatment for screening of ancient DNA. . Katoh, K., Misawa, K., Kuma, K.-I. Native Americansparticularly the Hopi Tribehave long received credit for cultivating blue . This is also consistent with the increased proportion of early European farmer ancestry related to Neolithic Europeans (Fig. Mixture proportions from five sources estimated using qpAdm. Skoglund, P. et al. 4). Finally, all mitochondrial reads were aligned to their respective consensus sequence, using bwa aln (version 0.7.12-r1039)64 with a maximum number of differences in the seed (-k) set to 5 and the maximum number of differences (-n) to 10, and bwa samse (version 0.7.12-r1039)64. Of the 100l extract, 20l was used to immortalize the sample DNA as a double-stranded library. For samples from the sites of Bolshoy and Chalmny Varre, we used leftover tooth powder that was originally processed at the Institute of Anthropology at the University of Mainz for replication purposes as described in ref. We imported the trimmed mitochondrial reads for each individual with mapping quality >30 into Geneious (version 10.0.9, https://www.geneious.com)68 and reassembled these reads to the reference genome RSRS78, using the Geneious mapper, with medium sensitivity and 5 iterations. Bioinformatics 25, 17541760 (2009). 26, 12441253 (2013). 3(ed. Nature 528, 499503 (2015). Our data suggest that this fourth genetic component found in modern-day north-eastern Europeans arrived in the area before 3500 yBP. A.Wes., V.M., V.K., A.S., P.O., O.B., W.H. Markers used correspond to the ones used in Fig. The supernatant was separated from the pellet of bone powder by centrifugation (14,000rpm). PLINK: a tool set for whole-genome association and population-based linkage analyses. The variant calls were filtered for variants with a quality score above 30, and a custom script was used to convert the variants into EigenStrat format. For each library, a unique pair of eight-bp-long indexes was incorporated using a Pfu Turbo Cx Hotstart DNA Polymerase and a thermocycling program with the temperature profile as follows: initial denaturation (98C for 30sec), cycle of denaturation/annealing/elongation (98C for 10sec/ 60C for 20sec/ 72C for 20sec) and final extension at 72C for 10min61. USA110, 1575815763 (2013). Ancient individuals from this study (black box)are represented by thicker bars and shown in bold. Saag, L. et al. Kimura, R. et al. Der Sarkissian, C. et al. and S.P. Whole-genome sequencing of 175 Mongolians uncovers population-specific genetic architecture and gene flow throughout North and East Asia, Genomic insights into the formation of human populations in East Asia, Ancient DNA reveals admixture history and endogamy in the prehistoric Aegean, The genetic history of admixture across inner Eurasia, The spread of steppe and Iranian-related ancestry in the islands of the western Mediterranean, People from Ibiza: an unexpected isolate in the Western Mediterranean, Genome-wide analysis of a collective grave from Mentesh Tepe provides insight into the population structure of early neolithic population in the South Caucasus, West Asian sources of the Eurasian component in Ethiopians: a reassessment, The population history of northeastern Siberia since the Pleistocene, https://sourceforge.net/projects/bio-bwa/files, https://github.com/stschiff/sequenceTools.git, https://github.com/TCLamnidis/Sex.DetERRmine, https://www.genetics.ucla.edu/software/admixture/download.html, https://github.com/TCLamnidis/ContaminateGenotypes, https://www.biorxiv.org/content/early/2016/11/19/088716, Description of Additional Supplementary Files, http://creativecommons.org/licenses/by/4.0/, Inferring biological kinship in ancient datasets: comparing the response of ancient DNA-specific software packages to low coverage data, Admixture has obscured signals of historical hard sweeps in humans, The Anglo-Saxon migration and the formation of the early English gene pool, Dairying, diseases and the evolution of lactase persistence in Europe, Phylogenetic history of patrilineages rare in northern and eastern Europe from large-scale re-sequencing of human Y-chromosomes, Nobel Prize in Physiology or Medicine 2022. The sequenced DNA fragments were mapped to the human reference genome, and pseudohaploid genotypes were called based on a random read covering each targeted SNP (see Methods). Genet. We show that the genetic makeup of northern Europe was shaped by migrations from Siberia that began at least 3500 years ago. Out of the 13 originally processed Iron-Age samples from Finland, seven proved to be of an adequate quality to be used in downstream analyses. performed laboratory work. 7, 447458 (1999). Lavento, M.)3036 (Finnish Antiquarian Society & Archaeological Society of Finland, 2004). AllSaami refers to a grouping including the 2 individuals from the SGDP (Saami(SGDP)) and the high-coverage modern Saami shotgun genome in this study (Modern Saami). CAS Ameur, A. et al. All of these were below 1.6% contamination (Table1). Csnyi, B. et al. Cite this article. The Yamnaya culture or the Yamna culture ( Russian: , Ukrainian: lit. Taavitsainen, J.-P. in The Oxford Handbook of the Archaeology and Anthropology of Hunter-Gatherers (eds Cummings, V., Jordan, P. & Zvelebil, M.) Ch. The study accession is PRJEB29360. 72, 519534 (2008). XXVI, 81105 (2009). Genetics 192, 10651093 (2012). b LD decay curve for Bolshoy, using Nganasan and EHG as sources. Anthropol. The genetic prehistory of the Baltic Sea region. CAS Specifically, AdapterRemoval was used to trim the sequencing adapters from our reads, with a minimum overlap of 1bp, and using a minimum base quality of 20 and minimum sequence length of 30bp. In this study we extend the available information from this area considerably, and present the first ancient genome-wide data from Finland. One of the individuals from Levnluhta (JK2065/Levnluhta_B) rejects a cladal position with modern Saami to the exclusion of most modern Eurasian populations. Females are expected to have an x-rate of 1 and a y-rate of 0, while males are expected to have both x- and y-rate of 0.5 (ref. Biol. Historical sources note Lapps living in the parishes of central Finland still in the 1500s21. Kearse, M. et al. 5b). The Finno-Ugric branch of the Uralic language family, to which both Saami and Finnish languages belong, has diverged from other Uralic languages no earlier than 40005000 years ago, when Finland was already inhabited by speakers of a language today unknown. These horse-riding pastoralists from the western steppe, known as the Yamnaya, may not have been responsible for bringing horse . Among the European sources, Lithuanians gave the most negative results for Estonians, Russians and Mordovians. Thank you for visiting nature.com. Ten of the eleven ancient individuals from this study fall on this Uralic cline, with the exception of one individual from Levnluhta (ID JK2065, here named Levnluhta_B), who instead is projected closer to modern Lithuanian, Norwegian and Icelandic populations. Allentoft, M. E. et al. Living tribespeople do descend in part from three ancient Native Americans who lived in the region 2500 to 6000 years ago . 2010, db.prot5448 (2010). 2a, Supplementary Figures3a, b for a version focusing on West Eurasia). Calculated f4 (Mbuti, Nganasan; Lithuanian, Test). We calibrated the radiocarbon date of Bolshoy, reported in refs 24,55 as 347342 years BP, using Intcal1356 as the calibration curve, using OxCal 4.357. Green, R. E. et al. The teeth were then sand-blasted using aluminium oxide abrasive (Harnisch & Rieth) and ground to fine powder using a mixer-mill (Retsch). A Copernican reassessment of the human mitochondrial DNA tree from its root. 17, 60 (2016). The sampling and subsequent processing of the ancient human remains was done in dedicated clean-room facilities (Methods). Genome Biol. The origin and timing of this East Asian-related contribution is unknown. By T101 in forum Paleogenetics Cold Spring Harb. This ancestry component is also present in modern Armenians from the Caucasus, Bedouins from the Arabian Peninsula and South Asian populations. Int. One Levnluhta individual (JK1970) showed slightly lower affinity to central Europe than modern-day Saami do, while still rejecting a cladal position with modern-day Finns. Am. Written informed consent was obtained from the Saami individual whose genome was analysed in this study, which was approved of by The Hospital District of Helsinki and Uusimaa Ethical Committee (Decision 329/13/03/00/2013) and the ethics committee of the University of Leipzig (Approval reference number 398-13-16122013). Greenlandic Inuit show genetic signatures of diet and climate adaptation. When multiple admixture events have occurred, such a single estimate should be interpreted as a (non-arithmetic) average of those events46,48. G3 7, 34593468 (2017). Archaeol. Upper Palaeolithic Siberian genome reveals dual ancestry of Native Americans. A majority of Yamnaya ancestry came from Caucasus-based hunter-gatherers and a minority . Seven individuals stem from excavations in Levnluhta, a lake burial in Isokyr, Finland. For the Eurasian PCA (Fig. Also known as the Yamnaya Culture, Pit Grave Culture or Ochre Grave Culture. Second, as shown in our analyses, the admixture patterns found in historic and modern Uralic speakers are complex and in fact inconsistent with a single admixture event. Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Ancient Fennoscandian genomes reveal origin and spread of Siberian ancestry in Europe, https://doi.org/10.1038/s41467-018-07483-5. Yamnaya Native Americans African East Asian South Asian 35% 25% 15% 10% 8% 4% 3% 25% Agriculture in Europe has origins in the Near East, back to the first farmers who cultivated barley and wheat in the hillocks of the Fertile Crescent. Google Scholar. In the meantime, to ensure continued support, we are displaying the site without styles Curr. This indicates that the people inhabiting Levnluhta during the Iron Age, and possibly other areas in the region as well, were more closely related to modern-day Saami than to present-day Finns; however, their difference from the modern Saami may reflect internal structure within the Saami population or additional admixture into the modern population. Hkkinen, J. Jatkuvuusperustelut ja saamelaisen kielen leviminen (OSA 2). Am. Duplicate removal was carried out using DeDup v0.12.1. J. Hum. According to research published since 2013, MA-1 belonged to the population of Ancient North Eurasians, who were genetically "intermediate between modern western Eurasians and Native Americans, but distant from east Asians", and partial genetic ancestors of Siberians, American Indians, and Bronze Age Yamnaya and Botai people of the Eurasian steppe. Today, the inhabitants of the area speak Finnish and Swedish. carried out the processing of the sequenced reads and generating the genotypes of the modern Saami genome. For each Test population, if outgroup set OG1 did not produce a working full model (p<0.05), we tried alternative outgroup sets with one right population removed. The genetic structure of Europeans today is the result of several layers of migration and subsequent admixture. Eur. Suomalais-Ugrilaisen Seuran Aikakauskirja96, 287316 (2017). The fitted trendline considers a minimum distance of 1cM. We preferred models with OG1-4 over OG5 in general, because OG5 contains more ancient genomes with potential biases in the right populations, which more often causes failing models for modern Test populations. Natl Acad. Horses were domesticated some time before 3,000 BC in central Asia. T.C.L., K.M., E.S., C.J. Contacts between Siberia and Europe are also recognised in linguistics. umkin, V. J. Ramsey, C. B. Bayesian analysis of radiocarbon dates. Our results show that all of the test populations are indeed admixed, with the most negative values arising when Nganasan are used as the Siberian source (Supplementary Data3). Following a brief ceremony April 17 that included the burning of sage and the singing of Native American prayers, five bison from the Laramie Foothills . Significantly negative f3 values correspond to the Test population being admixed between populations related to the two source populations42. Broushaki, F. et al. EAGER: efficient ancient genome reconstruction. Preprint at https://www.biorxiv.org/content/early/2016/11/19/088716 (2016). We used samtools mpileup (version 1.3)65, filtering for map- (-Q) and base- (-q) quality of 30, deactivating per-Base Alignment Quality (-B), on the trimmed bam files, to generate a pileup of reads mapping to a set of 43 phenotypic SNPs4,40,41,79 that are part of our genome capture panel. Google Scholar. The Ristola Site In Lahti And The Earliest Postglacial Settlement Of South Finland(Lahti City Museum, 2004). Therefore the admixture date estimate for Bolshoy does not preclude earlier admixture events bringing Siberian Nganasan-related ancestry into the population, in multiple waves. This was possible for all samples with UDG-half treatment, except for the individuals from Levnluhta, which represented too little damage for the PMD-filtering to be applied. Blankholm, H. P) 139167 (Equinox, 2018). Sampling and extracting ancient DNA requires a strict procedure in order to avoid contamination introduced by contemporary genetic material. The preliminary workflow included documenting, photographing and storing the samples in individual, ID-coded plastic tubes and plastic bags. et al. Forensic Sci. Huyghe, J. R. et al. IntCal13 and Marine13 radiocarbon age calibration curves 050,000 years cal BP. We first confirmed that the deamination pattern at the terminal bases of DNA reads were characteristic of ancient DNA (1.044.5% for non-UDG libraries, and 4.79.5% for non-UDG libraries, see Supplementary Table1), using mapDamage (version 2.0.6)66. and T.C.L. Suom.-Ugr. Linguistic evidence shows that Saami languages were spoken in Finland prior to the arrival of the early Finnish language and have dominated the whole of the Finnish region before 1000 CE16,17,18. In the case of PWC from Sweden where none of the outgroup sets OG1-4 produced a working model, a revised set of right populations was used (OG5) which includes Samara_HG to provide more power to distinguish hunter-gatherer ancestries. 266, 63117 (2012). For the West Eurasian PCA (Supplementary Figure3a, b), the following populations were used to construct principal components: Abkhasian, Adygei, Albanian, Armenian, Balkar, Basque, BedouinA, BedouinB, Belarusian, Bulgarian, Canary_Islander, Chechen, Chuvash, Croatian, Cypriot, Czech, Druze, English, Estonian, Finnish, French, Georgian, Greek, Hungarian, Icelandic, Iranian, Italian_North, Italian_South, Jew_Ashkenazi, Jew_Georgian, Jew_Iranian, Jew_Iraqi, Jew_Libyan, Jew_Moroccan, Jew_Tunisian, Jew_Turkish, Jew_Yemenite, Jordanian, Kumyk, Lebanese, Lezgin, Lithuanian, Maltese, Mordovian, North_Ossetian, Norwegian, Orcadian, Palestinian, Polish, Russian, Sardinian, Saudi, Scottish, Sicilian, Spanish, Spanish_North, Syrian, Turkish, Ukrainian. PLoS Genet. Nature 551, 368372 (2017). performed the laboratory work of the modern Saami genome. Finally, 5000 to 4800 years ago, nomadic herders known as the Yamnaya swept into Europe. Use the Previous and Next buttons to navigate the slides or the slide controller buttons at the end to navigate through each slide. They established that the Yamnaya pastoralists of the Pontic steppe contributed a substantial proportion of ancestry to modern Europeans (later, the same was found to be the case in Indians and other Asians). The 3500-year-old ancient individuals from Bolshoy represent the highest proportion of Siberian Nganasan-related ancestry seen in this region so far, and possibly evidence its earliest presence in the western end of the trans-Siberian expanse (Fig. Fort Collins Coloradoan. In addition, we sequenced the whole genome of a modern Saami individual to 17.5-fold coverage, for whom genotyping data has previously been published1. Salmela, E. et al. You can view a sample below. A multiple alignment of the consensus sequence and a reference set of 311 mitochondrial genomes69 was generated, using mafft (version v7.305b)70,71,72 with the --auto parameter. Terminal base deamination damage calculation was done using mapDamage, specifying a length (-l) of 100bp (Supplementary Table1). Mallick, S. et al. For the fill-in step, a mixture of 0.4U/l Bst-polymerase and 125M dNTP mix was added and the mixture then incubated in a thermocycler (30min 37C, 10min 80C). This was followed by two subsequent washing steps of 450l of wash buffer (High Pure Viral Nucleic Acid Large Volume Kit) and two dry spin steps of 1min centrifugation at 14,000rpm. On the ethnogenesis of the Sami: an archaeological view. For example, North America had a big demographic shift starting around 1500 AD. 128, 415423 (2005). Haplogroup N1c, to which this haplotype belongs, is the major Y-chromosomal lineage in modern north-east Europe and European Russia. 2, e190 (2006). Finally, the Imiyakhtakhskaya culture from Yakutia spread to the Kola Peninsula during the same period24,53. We used EAGER63 (version 1.92.50) to process the sequenced reads, using default parameters (see below) for human-originated, UDG-half treated, single-end sequencing data, when processing the UDG-half libraries for all individuals. The merged reads were extracted from the resulting bam files, and merged with the bam file containing reads from the single end sequence run using samtools merge (version 1.3)65. By Sciacchitano in forum DNA Testing & General Genetics Replies: 0 Last Post: 24-05-18, 07:45. The fact that the Nganasan-related genetic component is consistently shared among Uralic-speaking populations, with the exceptions of absence in Hungarians and presence in the non-Uralic speaking Russians, makes it tempting to equate this genetic component with the spread of Uralic languages in the area. Bone powder from a cave bear was processed in parallel serving as a positive control. The Simons Genome Diversity Project: 300 genomes from 142 diverse populations. Hidden and remote: new perspectives on the people in the Levnluhta Water Burial, Western Finland (c.ad 300800). If either Yamnaya or EHG could be dropped . Behar, D. M. et al. Fu, Q. et al. Phenotypes related to both Estonians (about half of Yamnaya ancestry comes from EHGs with slight native american) and Circassians/Georgians (the caucasus/west asian/"hindu kush" ancestry that is slightly less than half of Yamnaya, and also has ANE) Less balticized and less gracilized than those modern populations, though. For three Levnluhta individuals exceeding the threshold coverage of 1% in the preliminary screening, we used the non-UDG treated libraries to confirm the authenticity of the ancient data. We note that a low but significant amount of Neolithic European ancestry is also present in the Bolshoy population. The third Levnluhta individual (Levnluhta_B), however, lacks this Siberian component. Anyone you share the following link with will be able to read this content: Sorry, a shareable link is not currently available for this article. Genet. 28(University of Tartu Publisher, 2018). supervised sequencing of the modern Saami and post-sequencing bioinformatics. Cell 134, 416426 (2008). Fennosc. Parallel palaeogenomic transects reveal complex genetic history of early European farmers. Toim.=Mmoires De. We used a custom script (https://github.com/TCLamnidis/Sex.DetERRmine) for the calculation of each relative coverage as well as their associated error bars (Supplementary Figure1, Supplementary Note3 for more information on error calculation). The genetic history of Ice Age Europe. Populations containing individuals from this study are shown in bold. The molecular copy number in pre-indexed libraries ranged from ~10E8 to ~10E9 copies/l, indicating a successful library preparation, whereas the indexed libraries ranged from ~10E10 to ~10E12 copies/l, stating an admissible indexing efficiency. Science editor, BBC News website. COLUMBUS, Ohio (AP) Thousands of Native American remains in Ohio could finally be laid to rest under a provision that has passed the state House, the start of a process that tribal members have waited on for decades. Fourth, we calculated mitochondrial contamination for all ancient individuals using ContamMix29. For the female individuals from these two sites, we note that they are projected close to the males in PCA space (Fig. Here, we report five Yamnaya individuals well-dated to 3021 to 2501 calibrated BCE from kurgans in Romania, Bulgaria, and Hungary, displaying changes in bone morphology and distinct pathologies associated with horseback riding. Nature Communications (Nat Commun) Biol. Second, we estimated potential contamination rates through heterozygosity on the single-copy X chromosome for male individuals (all below 1.6%)27 (Table1, see Supplementary Figure1 for sex determination). We found that Bolshoy works as source in some ancient individuals, but not for modern Uralic speakers (see Supplementary Data4). Scandinavian hunter-gatherers from Motala in Sweden have also been found to carry haplotypes associated with this allele4. 90, 809820 (2012). Ancestors of present-day Finnish speakers possibly migrated from northern Estonia, to which Finns still remain linguistically close, and displaced but also admixed with the local population of Finland, the likely ancestors of todays Saami speakers23. J. Hum. Error bars represent one standard error and are plotted to the right of their associated mixture proportion. 370, 20130624 (2015). Lahermo, P. et al. We would like to also thank the sequencing team at Max Planck Institute of Evolutionary Anthropology for the sequencing of the modern Saami genome. For modern Hungarians, the European source giving most negative results was French, while both Bolshoy and Nganasan gave equally negative results when used as the Siberian source. We tested the power of this method to detect contamination and find that it can detect contamination that is distantly related to the ancestries present within the test individuals already at rates of 58%, but lacks the power to identify contamination closely related to the test individuals (see Supplementary Note2). For the 13 Iron-Age individuals from Finland available to us, the sampling took place in a clean-room facility dedicated to ancient DNA work, at the Institute for Archaeological Sciences in Tbingen. We used smartpca (version #16000)73 (https://github.com/DReichLab/EIG) to carry out Principal Component Analysis (PCA), using the lsqproject: YES and shrinkmode: YES parameters. Bioinformatics 28, 16471649 (2012). First, we confirmed the deamination patterns at the terminal bases of DNA reads being characteristic of ancient DNA (Supplementary Table1). The remaining dentine was collected by carefully separating it from the enamel with a dentist drill and cooled-down diamond drill heads, rotated at a speed below 15rpm, to avoid possible heat-caused damaging to the ancient DNA. For the modern Saami individual, total DNA was phenol-chloroform extracted and physically sheared using COVARIS fragmentation. Finally, for individuals with sufficient SNP coverage, we carried out principal component analysis (PCA), projecting damage-filtered and non-filtered versions of each individual, to show that the different datasets cluster together regardless of the damage-filtering30 (see Supplementary Figure3 and Methods). Our results suggest that a new genetic component with strong Siberian affinity first arrived in Europe at least 3500 years ago, as observed in our oldest analysed individuals from northern Russia. Google Scholar. This model, however, does not fit well for present-day populations from north-eastern Europe such as Saami, Russians, Mordovians, Chuvash, Estonians, Hungarians, and Finns: they carry additional ancestry seen as increased allele sharing with modern East Asian populations1,3,9,10. Artefacts from the site have been dated to the Finnish Iron Age (300800 CE)25,26. This genocide in Spain by the Yamnaya culture took place during the bronze age. Mittnik, A. et al. Hausen, R. Registrum Ecclesiae Aboensis Eller bo Domkyrkas Svartbok Med Tillgg Ur Skoklosters Codex Aboensis (Kansallisarkisto, 1890). The hunter-gatherer genetic ancestry in Europeans (blue) is maximized in European Upper Palaeolithic and Mesolithic hunter-gatherers, including the 8000-year-old Western European hunter-gatherers from Hungary and Spain (WHG), the 8000-year-old Scandinavian hunter-gatherers from Motala (SHG) and the Narva and Kunda individuals from the Baltics. The procedure included a blunt-end repair, adapter ligation and adapter fill-in steps, as described by Meyer and Kircher59. The Native-American-related ancestry seen in the EHG and Bolshoy corresponds to a previously reported affinity towards Ancient North Eurasians (ANE)2,33 contributing genes to both Native Americans and West Eurasians.

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