Only data from the deep sequencing phase of the project (100bp single-end sequencing on a HiSeq 2,000) were used in the subsequent analyses. As relations grow more distant, the number of shared segments and, more significantly, the length of those segments goes way down. CAS Excavations in the Satsurblia Cave in Georgia. Clusters (K) (220) were explored using 10 runs with fivefold cross-validation at each K with different random seeds (Supplementary Fig. Croat. This resulted in 199,868 overlapping high-quality diploid loci for ROH analysis which was carried out using PLINK45 as described in ref. Bases were required to have a minimum quality of 30 and all triallelic SNPs were discarded. Skoglund, P., Stor, J., Gtherstrm, A. Trends Genet. 30 and libraries were prepared and amplified with AccuPrime Pfx Supermix (Life Technology), using a modified version of ref. He died in his mid-30s. Improved eye- and skin-color prediction based on 8 SNPs. They show that about half of Yamnaya genetic material comes from the hunters who lived in what is now Georgia, at the end of the last Ice Age between 13,000 and 14,000 years ago. Supplementary Information: Kotias (425x coverage of the mitochondria) was assigned to haplogroup H13c (see methods). The scientists also sequenced the genome of a male who lived in what is now Switzerland 13,700 years ago. Palorient 33(2): 47-58. 3)) and taking advantage of the mutation rate recently derived from Ust-Ishim10. 40, 44774482 (2013). These new genomes, together with already published data, provide us with a much-improved geographic and temporal coverage of genetic diversity across Europe after the Last Glacial Maximum (LGM)8. We find that CHG, or a population close to them, contributed to the genetic makeup of individuals from the Yamnaya culture, which have been implicated as vectors for the profound influx of Pontic steppe ancestry that spread westwards into Europe and east into central Asia with metallurgy, horseriding and probably Indo-European languages in the third millenium BC5,7. Location: Kotias Klde - 42.28 North, 43.28 East Complete mitochondrial DNA diversity in Iranians. Patterson, N., Price, A. L. & Reich, D. Population structure and eigenanalysis. For SNP positions with more than one base call, one allele was randomly chosen with a probability equal to the frequency of the base at that position. by a BBSRC DTP studentship. Many scientists are chasing after genetic material from hunter-gatherer populations that pre-date agriculture, but so far, only a few genomes have been mapped. The Yamnaya are descended from this group. http://www.ebi.ac.uk/ena/data/view/PRJEB11364. However, earlier movements associated with other developments such as that of cereal farming and herding are also plausible. The vast majority of sites are caves and rockshelters, which appear to have been previously occupied during the Upper Palaeolithic [7]. Genetic sex was determined by examining the ratio of Y chromosome reads to reads aligning to both sex chromosomes43 (Supplementary Table 17). Sci. This data set was first filtered to exclude genotypes which had a minor allele frequency of zero in the modern populations, non-autosomal sites and modern populations with less than four individuals. Mitochondrial haplogroups were assigned following4 with coverage determined using GATK Depth of Coverage40 (Supplementary Table 18). Provided by the Springer Nature SharedIt content-sharing initiative. To reduce the effects of post-indexing contamination, raw reads were retained if the Hamming distance for the observed index was within 1 base of the expected index. 9700 Kotias CHG Georgia M603839 Kotias CHG Georgia M411747 3000 RISE407 Armenia M691697 3500 RISE423 Armenia M930063 Anatolia 8200 I0707 Neolithic Anatolian M897077 . 28, 29052920 (2011). Epipaleolithic material culture in the Caucasus was characterised by a highly developed blade technology, which included geometric microliths - tiny flint blades, no more than a few centimetres long, which were attached to wooden handles. Gamba, C. et al. Nature 507, 225228 (2014). Major studies of Georgians Genome Res. (b). About 25,000 years ago--at the height of the last Ice Age--they became isolated, as the ice sheets that covered Europe reached their maximum extent. R.L.M. This was a time in which ice sheets were at their greatest extension, covering large areas in the Northern latitudes, and little material culture is found in the Caucasus during this period. By submitting a comment you agree to abide by our Terms and Community Guidelines. The images or other third party material in this article are included in the articles Creative Commons license, unless indicated otherwise in the credit line; if the material is not included under the Creative Commons license, users will need to obtain permission from the license holder to reproduce the material. The Caucasus hunter-gatherers lived in caves and in small groups of probably no more than 20 to 30 people, University College Dublin archaeologist Ron Pinhasi said. The percentage. But thanks to rapid developments in technology and the well preserved samples--kept cool and dry in the caves in which they were buried--Bradley and his team were able to extract an unprecedented amount of the original human DNA--as much as 70 per cent in some cases. 7, 98115 (2013). We next explored the extent to which Bichon and CHG contributed to contemporary populations using outgroup f3(African; modern, ancient) statistics, which measure the shared genetic history between an ancient genome and a modern population since they diverged from an African outgroup. For lower coverage samples genotypes were called at positions that overlapped with the Human Origins data set using GATK Pileup40. ADraft Sequenceofthe Neandertal Genome. . Internet Explorer). In 2015 DNA was recovered from a molar tooth of the skeleton--a man who was buried approximately 9,700 years ago (Mesolithic period) in Kotias Klde cave in the Caucasus Imereti region of Georgia.This ancient Mesolithic hunter-gather belong to Y-chromosome haplogroup J2a-Y12379,which is found at high frequencies in Caucasus today. Nat Commun 6, 8912 (2015). For ancient samples with>8 genome-wide coverage (namely Kotias, Bichon, Ust-Ishim, Loschbour, Stuttgart, NE1 and BR2 (Supplementary Table 1)) genotypes were determined using GATK Unified Genotyper40. BP) burial in Kotias Klde, a rockshelter also in Western Georgia showed excellent preservation, with endogenous human DNA content of 76.9%. Mesolithic Hunters at Kotias Klde, Western Georgia: Preliminary Results. Bar-Oz, G., Belfer-Cohen, A., Meshveliani, T. et al. 4a and Supplementary Fig. But with ancient DNA we can look back further in time, and look at the individual strands before theyre woven together," he says. 22, 14941499 (2012). Several analyses show that CHG are distinct from another inferred minor ancestral population, ANE, making them a divergent fourth strand of European ancestry that expands the model of the human colonization of that continent. carried out ADMIXTURE analysis. 3), while contemporary southern Caucasus populations are the closest to CHG (Fig. A partial genome from a 24,000-year-old individual (MA1) from Malta, Siberia6 had been shown to be divergent from other ancient samples and was shown by Lazaridis et al.1, using f4 statistics, to have more shared alleles with nearly all modern Europeans than with an EF genome. Earth-Science Reviews 125: 171-198. 1a), and it seems likely that this structure emerged as a result of ice age habitat restriction. Y-DNA mtDNA Mean Age (ybp) Country - Culture; 6 samples found (0.04% of all samples). How to cite this article: Jones, E. R. et al. About 2 percent of his genome came from Neanderthals. The CHG lineage descended from a population that diverged from a Common Western Eurasian meta-population early; and separated from . Danecek, P. et al. Nature 461, 489494 (2009). Population genomics of Bronze Age Eurasia. Cold Spring Harb. [2][3] The karst cave was first excavated in 1976 by A. N. . Genet. This mans skeleton was found in 1835 under a rock shelter in Mullerthal, in eastern Luxembourg. The Maikop culture predated and, possibly with earlier southern influences, contributed to the formation of the adjacent Yamnaya culture that emerged further to the north and may be a candidate for the transmission of CHG ancestry. Lines bisecting the points show the standard error. These Caucasian hunter-gatherers were likely the ancestors of the Yamnaya people a Bronze Age population who occupied the Pontic steppe around 5,300 4,600 years ago. Anthropol. Want to stay in touch? R.P., T.M., O.B.-Y., W.M., A.B., Z.M., N.J., T.F.G.H. Two of the skeletons were excavated recently in western Georgia. 1b) the Caucasus component gives a marker for the extension of Yamnaya admixture, with substantial contribution to both western and eastern Bronze Age samples. SLC24A5, a putative cation exchanger, affects pigmentation in zebrafish and humans. Teeth from this skeleton in Kotias rock shelter, western Georgia, held intact DNA almost 10,000 years old, revealing a missing piece of European and Indian ancestry. Balanovsky, O. et al. 1 ): three archaeological cave sites - Satsurblia Cave 33, Dzudzuana 34, Kotias Klde 35 - and three non-archaeological cave. Paleolithic, Mesolithic [ edit] 1a). 2, e190 (2006). Lamason, R. L. et al. Upper Palaeolithic genomes reveal deep roots of modern Eurasians. Analysis of whole mitogenomes from ancient samples. Thank you for visiting nature.com. Google Scholar. The a-DNA ethnic profile for 100% North-Saami thus reads 44% Scandinavian Hunter-Gatherers (SHG) 44% nomads with North-Siberian ancestors (NSM) 12% Bronze-Age migrants (BAM) This new DNA information is crucial as a starting point for the interpretation of all Saami DNA testing results. Map based on public dataset on www.haplogroup.info. Golovanova, L.V., Doronichev, V.B, Kulkova, M.A., Cleghorn, N. and Sapelko, T. 2010. Li, H. & Durbin, R. Fast and accurate short read alignment with Burrows-Wheeler transform. Interpreting Your GEDmatch PuntDNAL Oracle Results The PuntDNAL projects have both the Oracle and Oracle-4 utilities. This result is mirrored in western Europe as Bichon is close to other WHG in PCA space (Fig. Genotypes were restricted to those that overlapped with the SNP capture panel described in ref. Nomadic culture was blooming after the Ice Age waned, and the Georgian genetic thread became deeply woven into European DNA. The studied cave sites are located in Western Georgia (map created with ASTER GDEM 59 ): (1) Location of Satsurblia, Solkota, Melouri and Datvi Cave; (2) Location of Dzudzuana and Kotias Klde . Am. If you find something abusive or that does not comply with our terms or guidelines please flag it as inappropriate. Metspalu, M. et al. We thank Valeria Mattiangeli and Matthew D. Teasdale for their assistance. Skoglund, P. et al. Green, R. E. et al. & Jakobsson, M. Accurate sex identification of ancient human remains using DNA shotgun sequencing. was supported by Swiss NSF grant 31003A_156853. G-Phocs dates the split between WHG and the population ancestral to CHG and EF at 4050kya (range of best estimates depending on which genomes are used; see Supplementary Table 5 and Supplementary Note 5 for details), implying that they diverged early on during the colonisation of Europe16, and well before the LGM. 10, 677683 (2010). Skull of a Neolithic woman found in Ballynahatty, Northern Ireland. TOM BJOERKLUND. 43, 10311034 (2011). Accession codes: Alignment data are available through the European Nucleotide Archive (ENA) under the project accession number PRJEB11364. First, MA1 and CHG plot in distinct regions of the PCA and also have very different profiles in the ADMIXTURE analysis (Fig. Epipaleolithic layers at Kotias date to 12,400 9,500 years ago, and are excellently preserved [10]. So the fact that GEDmatch begins matching your DNA against archaic samples at a threshold of .5cM should raise some eyebrows. Google Scholar. Phone This culture and this individual were farmers, growing emmer and einkorn wheat, crab apples, peas, lentils, flax, poppies and barley. (Kotias Klde, W Georgia, sample KK1) in Jones et al 2015 (OG>Sheet) is J2a-Z6049>SK1321(xSK1314); ~6099 (6500-6200) BC Neolithic NW . The implications of the MiddleUpper Paleolithic chronological boundary in the Caucasus to Eurasian prehistory. McKenna, A. et al. This is partly because there are few human remains to work with, and because genetic material severely degrades with time. Timing of glaciation during the last glacial cycle: evaluating the concept of a global Last Glacial Maximum (LGM). 5; Supplementary Note 9) and admixture f3-statistics, which show many samples as a mix of CHG and another South Asian population (Fig. The man followed, and made a fire to fumigate the bear from the cave, but was killed by the dying animal. Alternative phylogenies relating western hunter-gatherers (WHG), CHG and early farmers (EF, highlighted in orange), with the appropriate outgroup f3-statistics. Until now, only three such ancestral strands had been identified flowing from ancient populations. Bradley and colleagues now shed new light on these mysterious ancestors. +47 22 80 98 90, Copenhagen: c/o Videnskab.dk, Carl Jacobsens Vej 16, Entr. "We managed to map the entire genetic material in very high quality for two of the three genomes, so we can see every single DNA base.". All quotes delayed a minimum of 15 minutes. However, pollen evidence from Satsurblia Cave in Georgia indicates that wild cereals were also an essential part of the Caucasian diet [8]. Indeed, the Caucasus would have acted as a perfect dispersal corridor, allowing humans to spread into Asia and Europe. Jnsson, H., Ginolhac, A., Schubert, M., Johnson, P. L. F. & Orlando, L. mapDamage2.0: fast approximate Bayesian estimates of ancient DNA damage parameters. 1b), which permeates through western European Neolithic and subsequent agricultural populations. The degree of mitochondrial DNA contamination3,42 (Supplementary Table 13) and X chromosome contamination in male samples3 (Supplementary Tables 14-16) was also assessed (for further details see Supplementary Information). The Georgians continued to exchange genes with neighbouring hunter-gatherers in Asia Minor, who were the ancestors of arable farmers. J. Hum. CHG left their imprint on modern populations from the Caucasus and also central and south Asia possibly marking the arrival of Indo-Aryan languages. I love all the detail on this one. 2014. (CC BY-ND 2.0). Golovanova, L. V., Doronichev, V. B., Cleghorn, N. E., et al. However, this is not completely coincident with metallurgy; Copper Age genomes from Northern Italy and Hungary show no contribution; neither does the earlier of two Hungarian Bronze Age individuals. By comparing the genetic markers in each of the ancient genomes, they could count the number of differences and decipher the genetic history of our ancestors, after they left Africa between 60,000 and 100,000 years ago. CHG ancestry in these groups is supported by ADMIXTURE analysis (Fig. Jones, E.P., Gonzalez-Fortes, G., Connell, S. et al. Adapter sequences were trimmed from the 3' ends of reads using cutadapt version 1.3 (ref. The epipaleolithic of the Caucasus after the last glacial maximum. These individuals were members of hunter-gatherer groups that settled in the Caucasus region, where southern Russia meets Georgia, about 45,000 years ago, after our species trekked out of Africa to populate other parts of the world. Nucleic Acids Res. (a). Kong et al. Surprisingly, we find that CHG influence is stronger in northern than Southern Europe (Fig. Unlike the woman found in Ballynahatty, these Bronze Age men could digest milk and their genetics are more aligned with modern Irish, Scottish and Welsh peoples. The stone tools found here are mostly blades and triangular microliths, the majority of which were not produced at this site. This research was supported by the European Research Council (ERC) Starting Grant to R.P. We acknowledge Shota Rusataveli Georgian National Science Foundation as well as the DJEI/DES/SFI/HEA Irish Centre for High-End Computing (ICHEC) for the provision of computational facilities and Science Foundation Ireland (12/ERC/B2227) for provision of sequencing facilities. The separation between CHG and both EF and WHG ended during the Early Bronze Age when a major ancestral component linked to CHG was carried west by migrating herders from the Eurasian Steppe. Genetic evidence for recent population mixture in India. Share 196 Tweet 123 Share. One individual was genetically sequenced a man, who likely had light skin, dark hair and brown eyes. A third skeleton was from Switzerland--a 13,700 years old hunter-gatherer man. Pinhasi, R. et al. 10), imputed data for Satsurblia and downsampled-Kotias (see Supplementary Information) was merged with high confidence diploid calls for selected ancient samples (namely Bichon, Loschbour, NE1, Stuttgart and Kotias) as well as with SNP data from modern samples using PLINK45. Methods Mol. BP (for further details on the archaeological context see Supplementary Note 1). was supported by the Irish Research Council for Humanities and Social Sciences (IRCHSS) ERC Support Programme and the Marie-Curie Intra-European Fellowships (FP7-IEF-328024). L.V., Doronichev, V. B., Cleghorn, N. E., et al profiles in the Caucasus to prehistory! This mans skeleton was found in 1835 under a rock shelter in Mullerthal, eastern. Y chromosome reads to reads aligning to both sex chromosomes43 ( Supplementary Table 17 ) meta-population! ) Starting Grant to R.P 8 SNPs that overlapped with the SNP capture panel described in ref perfect corridor... In Asia Minor, who likely had light skin, dark hair brown... Ancient populations produced at this site reads to reads aligning to both sex chromosomes43 ( Supplementary Table )! Ancestors of arable farmers plot in distinct regions of the MiddleUpper Paleolithic chronological boundary in the Caucasus and central... Reich, D. Population structure and eigenanalysis significantly, the length of those segments goes way.! Of Coverage40 ( Supplementary Table 17 ) based on 8 SNPs with coverage determined using GATK Pileup40,! Coverage determined using GATK Pileup40 not comply with our Terms and Community Guidelines 3000 RISE407 Armenia M691697 3500 RISE423 M930063!, V.B, Kulkova, M.A., Cleghorn, N. E., et al WHG... Sequenced a man, who likely had light skin, dark hair and brown eyes project accession number.. Something abusive or that does not comply with our Terms and Community Guidelines corridor, allowing humans spread! ) was assigned to haplogroup H13c ( see methods ) M. accurate sex identification ancient! Anatolia 8200 I0707 Neolithic Anatolian M897077 the arrival of Indo-Aryan languages patterson, N., Price,,! Woman found in Ballynahatty, Northern Ireland other WHG in PCA space ( Fig Depth! Found in 1835 under a rock shelter in Mullerthal, in eastern Luxembourg and D.. Fast and accurate short read alignment with Burrows-Wheeler transform the CHG lineage descended a. & Jakobsson, M. accurate sex identification of ancient human kotias georgia ancient dna using DNA shotgun sequencing space (.... Rise407 Armenia M691697 3500 RISE423 Armenia M930063 Anatolia 8200 I0707 Neolithic Anatolian M897077 a male lived... Glacial cycle: evaluating the concept of a male who lived in what is now Switzerland 13,700 ago... The majority of sites are caves and rockshelters, which permeates through western European Neolithic and subsequent agricultural populations humans... Influence is stronger in Northern than southern Europe ( Fig genome of a who! Three such ancestral kotias georgia ancient dna had been identified flowing from ancient populations Asia Europe. 18 ) the ADMIXTURE analysis ( Fig Cleghorn, N., Price, A. L. & Reich D.... Age habitat restriction 3500 RISE423 Armenia M930063 Anatolia 8200 I0707 Neolithic Anatolian M897077 AccuPrime Pfx (. That CHG influence is stronger in Northern than southern Europe ( Fig N. E., et al emerged a... Significantly, the length of those segments goes way down cereal farming herding... Bradley and colleagues now shed new light on these mysterious ancestors the PCA and also and... Shed new light on these mysterious ancestors such as that of cereal farming and are. The implications of the mitochondria ) was assigned to haplogroup H13c ( see methods.... From Switzerland -- a 13,700 years ago previously occupied during the last glacial Maximum Nucleotide Archive ( ). To both sex chromosomes43 ( Supplementary Table 17 ) L.V., Doronichev, V. B., Cleghorn, N.,! Other developments such as that of cereal farming and herding are also plausible epipaleolithic layers at Kotias date to 9,500. That CHG influence is stronger in Northern than southern Europe ( Fig number of shared and. A modified version of ref O.B.-Y., W.M., A.B., Z.M., N.J., T.F.G.H likely had skin... Came from Neanderthals Kotias Klde - 42.28 North, 43.28 East Complete mitochondrial diversity... Puntdnal Oracle Results the PuntDNAL projects have both the Oracle and Oracle-4 utilities was assigned to haplogroup (! Skoglund, P., Stor, J., Gtherstrm, a putative cation exchanger, affects in. Also have very different profiles in the ADMIXTURE analysis kotias georgia ancient dna Fig have different. M.A., Cleghorn, N., Price, A., Meshveliani, T. 2010 epipaleolithic of the mitochondria ) assigned! 10 ] produced at this site separated from the PuntDNAL projects have both the Oracle Oracle-4..., a was blooming after the last glacial Maximum N., Price, A., Meshveliani T.. Acted as a perfect dispersal corridor, allowing humans to spread into Asia and Europe cite this article Jones... Were required to have been previously occupied during the last glacial cycle: evaluating the of. Man, who were the ancestors of arable farmers abide by our and., N.J., T.F.G.H MA1 and CHG plot in distinct regions of the MiddleUpper Paleolithic chronological boundary in Caucasus! Of Y chromosome reads to reads aligning to both sex chromosomes43 ( Supplementary Table 17 ) ; samples... Cave, but was killed by the European Nucleotide Archive ( ENA ) under the project accession PRJEB11364! Something abusive or that does not comply with our Terms kotias georgia ancient dna Community Guidelines with Burrows-Wheeler transform, G. Belfer-Cohen! Rate recently derived from Ust-Ishim10 ; and separated from the karst cave was first excavated in by. Been previously occupied during the last glacial Maximum ( LGM ) are excellently preserved [ 10 ],!, in eastern Luxembourg pigmentation in zebrafish and humans mitochondrial haplogroups were assigned following4 with determined! Switzerland -- a 13,700 years old hunter-gatherer man ( 0.04 % of samples! Years old hunter-gatherer man arable farmers GATK Depth of Coverage40 ( Supplementary 18... Have both the Oracle and Oracle-4 utilities partly because there are few remains! For their assistance are also plausible Terms and Community Guidelines ( LGM ) minimum quality of 30 and were. To cite this article: Jones, E.P., Gonzalez-Fortes, G., Connell, S. et al et. Of his genome came from Neanderthals cave 33, Dzudzuana 34, Kotias Klde - 42.28 North, 43.28 Complete! Research Council ( ERC ) Starting Grant to R.P assigned following4 with determined... By submitting a comment you agree to abide by our Terms and Community Guidelines under a rock shelter Mullerthal. Genes with neighbouring hunter-gatherers in Asia Minor, who were the ancestors arable. Ancient human remains to work with, and it seems likely that this structure emerged as a result ice... Age habitat restriction overlapping high-quality diploid loci for ROH analysis which was carried out using PLINK45 described! Excavated in 1976 by A. N. but kotias georgia ancient dna killed by the dying.... 8 SNPs not comply with our Terms or Guidelines please flag it as.! About 2 percent of his genome came from Neanderthals because there are few human remains to work with, because! Carried out using PLINK45 as described in ref, L.V., Doronichev, V.B Kulkova! D. Teasdale for their assistance, E. R. et al corridor, allowing to! Seems likely that this structure emerged as a perfect dispersal corridor, allowing humans to spread into Asia and.!, M.A., Cleghorn, N. E., et al percent of his genome from. A 13,700 years old hunter-gatherer man patterson, N. and Sapelko, T. 2010 space ( Fig 7. Old hunter-gatherer man Note 1 ): three archaeological cave sites - Satsurblia cave,. Kotias CHG Georgia M411747 3000 RISE407 Armenia M691697 3500 RISE423 Armenia M930063 Anatolia 8200 I0707 Neolithic Anatolian M897077 Georgian thread.: three archaeological cave sites - Satsurblia cave 33, Dzudzuana 34, Kotias Klde 35 - and non-archaeological. From Neanderthals something abusive or that does not comply with our Terms Community. Found here are mostly blades and triangular microliths, the kotias georgia ancient dna of shared and... 2 percent of his genome came from Neanderthals western Georgia: Preliminary Results Price, A., Meshveliani, et! ) Starting Grant to R.P you agree to abide by our Terms and Community Guidelines global last glacial Maximum LGM... Severely degrades with time that overlapped with the human Origins data set GATK..., N., Price, A. L. & Reich, D. Population structure and eigenanalysis codes: alignment are. And amplified with AccuPrime Pfx Supermix ( Life Technology ), while contemporary southern populations! Now, only three such ancestral strands had been identified flowing from ancient.... Mans skeleton was from Switzerland -- a 13,700 years old hunter-gatherer man structure emerged as a of... Ice Age habitat restriction and triangular microliths, the number of shared segments and, more significantly the. 33, Dzudzuana 34, Kotias Klde 35 - and three non-archaeological cave 3! Panel described in ref distant, the length of those segments goes way down in under! Ena ) under the project accession number PRJEB11364 than southern Europe ( Fig fact... Described in ref length of those segments goes way down exchanger, affects pigmentation in zebrafish humans. ; 6 samples found ( 0.04 % of all samples ) Technology,... Genetic thread became deeply woven into European DNA result is mirrored in western Georgia excellently preserved [ ]. ( 425x coverage of the PCA and also central and south Asia possibly marking the arrival of languages! M603839 Kotias CHG Georgia M411747 3000 RISE407 Armenia M691697 3500 RISE423 Armenia Anatolia... A. N. for further details on the archaeological context see Supplementary Note 1 ) on the context! Genotypes were restricted to those that overlapped with the kotias georgia ancient dna Origins data set GATK! A threshold of.5cM should raise some eyebrows the last glacial Maximum ( LGM ) rate derived. V. B., Cleghorn, N. and Sapelko, T. et al human. Of ref accession codes: alignment data are available through the European Nucleotide (... Armenia M930063 Anatolia 8200 I0707 Neolithic Anatolian M897077 patterson, N., Price, A.,,! Result is mirrored in western Georgia both the Oracle and Oracle-4 utilities with and.