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      Megalithic tombs in western and northern Neolithic Europe were linked to a kindred society

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          Significance

          A new phenomenon of constructing distinctive funerary monuments, collectively known as megalithic tombs, emerged around 4500 BCE along the Atlantic façade. The megalithic phenomenon has attracted interest and speculation since medieval times. In particular, the origin, dispersal dynamics, and the role of these constructions within the societies that built them have been debated. We generate genome sequence data from 24 individuals buried in five megaliths and investigate the population history and social dynamics of the groups that buried their dead in megalithic monuments across northwestern Europe in the fourth millennium BCE. Our results show kin relations among the buried individuals and an overrepresentation of males, suggesting that at least some of these funerary monuments were used by patrilineal societies.

          Abstract

          Paleogenomic and archaeological studies show that Neolithic lifeways spread from the Fertile Crescent into Europe around 9000 BCE, reaching northwestern Europe by 4000 BCE. Starting around 4500 BCE, a new phenomenon of constructing megalithic monuments, particularly for funerary practices, emerged along the Atlantic façade. While it has been suggested that the emergence of megaliths was associated with the territories of farming communities, the origin and social structure of the groups that erected them has remained largely unknown. We generated genome sequence data from human remains, corresponding to 24 individuals from five megalithic burial sites, encompassing the widespread tradition of megalithic construction in northern and western Europe, and analyzed our results in relation to the existing European paleogenomic data. The various individuals buried in megaliths show genetic affinities with local farming groups within their different chronological contexts. Individuals buried in megaliths display (past) admixture with local hunter-gatherers, similar to that seen in other Neolithic individuals in Europe. In relation to the tomb populations, we find significantly more males than females buried in the megaliths of the British Isles. The genetic data show close kin relationships among the individuals buried within the megaliths, and for the Irish megaliths, we found a kin relation between individuals buried in different megaliths. We also see paternal continuity through time, including the same Y-chromosome haplotypes reoccurring. These observations suggest that the investigated funerary monuments were associated with patrilineal kindred groups. Our genomic investigation provides insight into the people associated with this long-standing megalith funerary tradition, including their social dynamics.

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          Ancient human genomes suggest three ancestral populations for present-day Europeans

          We sequenced genomes from a $\sim$7,000 year old early farmer from Stuttgart in Germany, an $\sim$8,000 year old hunter-gatherer from Luxembourg, and seven $\sim$8,000 year old hunter-gatherers from southern Sweden. We analyzed these data together with other ancient genomes and 2,345 contemporary humans to show that the great majority of present-day Europeans derive from at least three highly differentiated populations: West European Hunter-Gatherers (WHG), who contributed ancestry to all Europeans but not to Near Easterners; Ancient North Eurasians (ANE), who were most closely related to Upper Paleolithic Siberians and contributed to both Europeans and Near Easterners; and Early European Farmers (EEF), who were mainly of Near Eastern origin but also harbored WHG-related ancestry. We model these populations' deep relationships and show that EEF had $\sim$44% ancestry from a "Basal Eurasian" lineage that split prior to the diversification of all other non-African lineages.
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            Massive migration from the steppe was a source for Indo-European languages in Europe

            We generated genome-wide data from 69 Europeans who lived between 8,000-3,000 years ago by enriching ancient DNA libraries for a target set of almost 400,000 polymorphisms. Enrichment of these positions decreases the sequencing required for genome-wide ancient DNA analysis by a median of around 250-fold, allowing us to study an order of magnitude more individuals than previous studies and to obtain new insights about the past. We show that the populations of Western and Far Eastern Europe followed opposite trajectories between 8,000-5,000 years ago. At the beginning of the Neolithic period in Europe, ∼8,000-7,000 years ago, closely related groups of early farmers appeared in Germany, Hungary and Spain, different from indigenous hunter-gatherers, whereas Russia was inhabited by a distinctive population of hunter-gatherers with high affinity to a ∼24,000-year-old Siberian. By ∼6,000-5,000 years ago, farmers throughout much of Europe had more hunter-gatherer ancestry than their predecessors, but in Russia, the Yamnaya steppe herders of this time were descended not only from the preceding eastern European hunter-gatherers, but also from a population of Near Eastern ancestry. Western and Eastern Europe came into contact ∼4,500 years ago, as the Late Neolithic Corded Ware people from Germany traced ∼75% of their ancestry to the Yamnaya, documenting a massive migration into the heartland of Europe from its eastern periphery. This steppe ancestry persisted in all sampled central Europeans until at least ∼3,000 years ago, and is ubiquitous in present-day Europeans. These results provide support for a steppe origin of at least some of the Indo-European languages of Europe.
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              Genomic insights into the origin of farming in the ancient Near East

              We report genome-wide ancient DNA from 44 ancient Near Easterners ranging in time between ~12,000-1,400 BCE, from Natufian hunter-gatherers to Bronze Age farmers. We show that the earliest populations of the Near East derived around half their ancestry from a ‘Basal Eurasian’ lineage that had little if any Neanderthal admixture and that separated from other non-African lineages prior to their separation from each other. The first farmers of the southern Levant (Israel and Jordan) and Zagros Mountains (Iran) were strongly genetically differentiated, and each descended from local hunter-gatherers. By the time of the Bronze Age, these two populations and Anatolian-related farmers had mixed with each other and with the hunter-gatherers of Europe to drastically reduce genetic differentiation. The impact of the Near Eastern farmers extended beyond the Near East: farmers related to those of Anatolia spread westward into Europe; farmers related to those of the Levant spread southward into East Africa; farmers related to those from Iran spread northward into the Eurasian steppe; and people related to both the early farmers of Iran and to the pastoralists of the Eurasian steppe spread eastward into South Asia.
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                Author and article information

                Journal
                Proc Natl Acad Sci U S A
                Proc. Natl. Acad. Sci. U.S.A
                pnas
                pnas
                PNAS
                Proceedings of the National Academy of Sciences of the United States of America
                National Academy of Sciences
                0027-8424
                1091-6490
                7 May 2019
                15 April 2019
                15 April 2019
                : 116
                : 19
                : 9469-9474
                Affiliations
                [1] aHuman Evolution, Department of Organismal Biology, Uppsala University , SE-752 36 Uppsala, Sweden;
                [2] bCentre for Anthropological Research, Department of Anthropology and Development Studies, University of Johannesburg , 2006 Auckland Park, South Africa;
                [3] cDepartment of Archaeology and Ancient History, Uppsala University–Campus Gotland , SE-621 67 Visby, Sweden;
                [4] dResearch Centre in Evolutionary Anthropology and Paleoecology, School of Natural Sciences and Psychology, Liverpool John Moores University , L3 3AF Liverpool, United Kingdom;
                [5] eDepartment of Surgery, Royal Prince Alfred Hospital , Sydney, NSW 2050, Australia;
                [6] fOsteoarchaeological Research Laboratory, Department of Archaeology and Classical Studies, Stockholm University , SE-106 91 Stockholm, Sweden;
                [7] gMuseums and Special Collections, Sir Duncan Rice Library, University of Aberdeen , AB24 3AA Aberdeen, Scotland;
                [8] hDepartment of Human Evolution, Max Planck Institute for Evolutionary Anthropology , 04103 Leipzig, Germany;
                [9] iDepartment of Prehistorical Archaeology, Institute of Archaeology of Czech Academy of Sciences , CZ-11801 Prague, Czech Republic;
                [10] jArchaeological Research Laboratory, Department of Archaeology and Classical Studies, Stockholm University , SE-106 91 Stockholm, Sweden
                Author notes
                2To whom correspondence may be addressed. Email: jan.stora@ 123456ofl.su.se or mattias.jakobsson@ 123456ebc.uu.se .

                Edited by Anne C. Stone, Arizona State University, Tempe, AZ, and approved March 15, 2019 (received for review October 19, 2018)

                Author contributions: A.G., J.S., and M.J. designed research; F.S.-Q., H.M., M.F., L.G.-F., E.M.S., L.G.S., R.G., N.H., A.G., J.S., and M.J. performed research; H.M., M.F., L.G.-F., G.B., G.N., K.B., S.T., N.C., H.B., R.S., and J.S. contributed samples and conducted archaeological analyses; F.S.-Q., H.M., and M.F. analyzed data; and F.S.-Q., H.M., M.F., J.S., and M.J. wrote the paper with input from all authors.

                1F.S.-Q., H.M., and M.F. contributed equally to this work.

                Author information
                http://orcid.org/0000-0002-0201-6204
                http://orcid.org/0000-0003-4714-088X
                http://orcid.org/0000-0002-4609-6402
                http://orcid.org/0000-0001-7840-7853
                Article
                201818037
                10.1073/pnas.1818037116
                6511028
                30988179
                c0d75a09-8047-488e-a146-1138e63ae967
                Copyright © 2019 the Author(s). Published by PNAS.

                This open access article is distributed under Creative Commons Attribution-NonCommercial-NoDerivatives License 4.0 (CC BY-NC-ND).

                History
                Page count
                Pages: 6
                Funding
                Funded by: Riksbankens Jubileumsfond (RJ) 501100004472
                Award ID: M13-0904:1
                Award Recipient : Anders Götherström Award Recipient : Jan Storå Award Recipient : Mattias Jakobsson
                Funded by: Knut och Alice Wallenbergs Stiftelse (Knut and Alice Wallenberg Foundation) 501100004063
                Award ID: N/A
                Award Recipient : Anders Götherström Award Recipient : Jan Storå Award Recipient : Mattias Jakobsson
                Funded by: Berit Wallenberg foundation
                Award ID: N/A
                Award Recipient : Magdalena Fraser
                Categories
                Biological Sciences
                Genetics
                Social Sciences
                Anthropology
                From the Cover

                paleogenomics,population genomics,migration,megalithic tombs

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