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Thread: Population Genomics of Stone Age Eurasia

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    Default Population Genomics of Stone Age Eurasia

    https://www.biorxiv.org/content/10.1...05.04.490594v1

    Summary:

    The transitions from foraging to farming and later to pastoralism in Stone Age Eurasia (c. 11-3 thousand years before present, BP) represent some of the most dramatic lifestyle changes in human evolution. We sequenced 317 genomes of primarily Mesolithic and Neolithic individuals from across Eurasia combined with radiocarbon dates, stable isotope data, and pollen records. Genome imputation and co-analysis with previously published shotgun sequencing data resulted in >1600 complete ancient genome sequences offering fine-grained resolution into the Stone Age populations. We observe that:

    1) Hunter-gatherer groups were more genetically diverse than previously known, and deeply divergent between western and eastern Eurasia.

    2) We identify hitherto genetically undescribed hunter-gatherers from the Middle Don region that contributed ancestry to the later Yamnaya steppe pastoralists;

    3) The genetic impact of the Neolithic transition was highly distinct, east and west of a boundary zone extending from the Black Sea to the Baltic. Large-scale shifts in genetic ancestry occurred to the west of this "Great Divide", including an almost complete replacement of hunter-gatherers in Denmark, while no substantial ancestry shifts took place during the same period to the east. This difference is also reflected in genetic relatedness within the populations, decreasing substantially in the west but not in the east where it remained high until c. 4,000 BP;

    4) The second major genetic transformation around 5,000 BP happened at a much faster pace with Steppe-related ancestry reaching most parts of Europe within 1,000-years. Local Neolithic farmers admixed with incoming pastoralists in eastern, western, and southern Europe whereas Scandinavia experienced another near-complete population replacement. Similar dramatic turnover-patterns are evident in western Siberia;

    5) Extensive regional differences in the ancestry components involved in these early events remain visible to this day, even within countries. Neolithic farmer ancestry is highest in southern and eastern England while Steppe-related ancestry is highest in the Celtic populations of Scotland, Wales, and Cornwall (this research has been conducted using the UK Biobank resource);

    6) Shifts in diet, lifestyle and environment introduced new selection pressures involving at least 21 genomic regions. Most such variants were not universally selected across populations but were only advantageous in particular ancestral backgrounds. Contrary to previous claims, we find that selection on the FADS regions, associated with fatty acid metabolism, began before the Neolithisation of Europe. Similarly, the lactase persistence allele started increasing in frequency before the expansion of Steppe-related groups into Europe and has continued to increase up to the present. Along the genetic cline separating Mesolithic hunter-gatherers from Neolithic farmers, we find significant correlations with trait associations related to skin disorders, diet and lifestyle and mental health status, suggesting marked phenotypic differences between these groups with very different lifestyles. This work provides new insights into major transformations in recent human evolution, elucidating the complex interplay between selection and admixture that shaped patterns of genetic variation in modern populations.

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    Anatolian farmer ancestry is associated with increased anxiety, guilty feelings and irritability. Explains a lot:

    Interestingly, loci associated with overdispersed mood-related polygenic phenotypes recorded among the UK Biobank individuals (like increased anxiety, guilty feelings, and irritability) showed an overrepresentation of the Anatolian farmer ancestry component; and the WHG component showed a strikingly high contribution to traits related to diabetes
    I1 and probably Proto-Germanic expanded from southern Scandinavia:
    Finally, we investigated the fine-scale genetic structure in southern Scandinavia after the667
    introduction of Steppe-related ancestry using a temporal transect of 38 Late Neolithic and Early668
    Bronze Age Danish and southern Swedish individuals. Although the overall population genomic669
    signatures suggest genetic stability, patterns of pairwise IBD-sharing and Y-chromosome670
    haplogroup distributions indicate at least three distinct ancestry phases during a ~1,000-year time671
    span: i) An early stage between ~4,600 BP and 4,300 BP, where Scandinavians cluster with early672
    CWC individuals from Eastern Europe, rich in Steppe-related ancestry and males with an R1a Y-673
    chromosomal haplotype (Extended Data Fig. 8A, B); ii) an intermediate stage until c. 3,800 BP,674
    where they cluster with central and western Europeans dominated by males with distinct sub-675
    lineages of R1b-L51 (Extended Data Fig. 8C, D; Supplementary Note 3b) and includes Danish676
    individuals from Borreby (NEO735, 737) and Madesř (NEO752) with distinct cranial features677
    (Supplementary Note 6); and iii) a final stage from c. 3,800 BP onwards, where a distinct cluster of678
    Scandinavian individuals dominated by males with I1 Y-haplogroups appears (Extended Data Fig.679
    8E). Using individuals associated with this cluster (Scandinavia_4000BP_3000BP) as sources in680
    supervised ancestry modelling (see “postBA”, Extended Data Fig. 4), we find that it forms the681
    predominant source for later Iron- and Viking Age Scandinavians, as well as ancient European682
    groups outside Scandinavia who have a documented Scandinavian or Germanic association (e.g.,683
    Anglo-Saxons, Goths; Extended Data Fig. 4). Y-chromosome haplogroup I1 is one of the dominant684
    haplogroups in present-day Scandinavians,s, and we document its earliest occurrence in a ~4,000-685
    year-old individual from Falköping in southern Sweden (NEO220). The rapid expansion of this686
    haplogroup and associated genome-wide ancestry in the early Nordic Bronze Age indicates a687
    considerable reproductive advantage of individuals associated with this cluster over the preceding688
    groups across large parts of Scandinavia.
    Yamnaya mostly descended from newly discovered hunter-gatherers from the middle Don intermediate between EHG and CHG:

    From approximately 5,000 BP, an ancestry component appears on the eastern European plains in424
    Early Bronze Age Steppe pastoralists associated with the Yamnaya culture and it rapidly spreads425
    across Europe through the expansion of the Corded Ware complex (CWC) and related cultures 20,21 .426
    We demonstrate that this “steppe” ancestry (Steppe_5000BP_4300BP) can be modelled as a427
    mixture of ~65% ancestry related to herein reported hunter-gatherer genomes from the Middle Don428
    River region (MiddleDon_7500BP) and ~35% ancestry related to hunter-gatherers from Caucasus429
    (Caucasus_13000BP_10000BP) (Extended Data Fig. 4). Thus, Middle Don hunter-gatherers, who430
    already carry ancestry related to Caucasus hunter-gatherers (Fig. 2), serve as a hitherto unknown431
    proximal source for the majority ancestry contribution into Yamnaya genomes. The individuals in432
    question derive from the burial ground Golubaya Krinitsa (Supplementary Note 3). Material culture433
    and burial practices at this site are similar to the Mariupol-type graves, which are widely found in434
    neighbouring regions of Ukraine, for instance along the Dnepr River. They belong to the group of435
    complex pottery-using hunter-gatherers mentioned above, but the genetic composition at Golubaya436
    Krinitsa is different from the remaining Ukrainian sites (Fig 2A, Extended Data Fig. 4). We find437
    that the subsequent transition of the Late Neolithic and Early Bronze Age European gene pool438
    happened at a faster pace than during the Neolithisation, reaching most parts of Europe within a439
    ~1,000-year time period after first appearing in eastern Baltic region ~4,800 BP (Fig. 3). In line440
    with previous reports we observe that beginning c. 4,200 BP, steppe-related ancestry was already441
    dominant in samples from France and the Iberian peninsula, while it reached Britain only 400 years442
    later 11,38,44 . Strikingly, because of the delayed Neolithisation in Southern Scandinavia these443
    dynamics resulted in two episodes of large-scale genetic turnover in Denmark and southern Sweden444
    within a 1,000-year period (Fig. 3).

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    The availability of a large number of modern genomes (n=408,884) from self-identified “white”
    British individuals who share similar PCA backgrounds 107 789 allowed us to further examine the
    790 distribution of ancient ancestries at high resolution in Britain (Supplementary Note 3h). Although
    791 regional ancestry distributions differ by only a few percent, we find clear evidence of geographical
    792 heterogeneity across the United Kingdom as visualised by assigning individuals to their birth
    793 county and averaging ancestry proportions per county (Fig. 5, inset boxes). The proportion of
    794 Neolithic farmer ancestry is highest in southern and eastern England today and lower in Scotland,
    795 Wales, and Cornwall. Steppe-related ancestry is inversely distributed, peaking in the Outer
    Hebrides and Ireland, a pattern only previously described for Scotland 108 796 . This regional pattern was
    797 already evident in the Pre-Roman Iron Age and persists to the present day even though immigrating
    798 Anglo-Saxons had relatively less Neolithic farmer ancestry than the Iron-Age population of
    799 southwest Briton (Extended Data Fig. 4). Although this Neolithic farmer/steppe-related dichotomy
    800 mirrors the modern ‘Anglo-Saxon’/‘Celtic’ ethnic divide, its origins are older, resulting from
    801 continuous migration from a continental population relatively enhanced in Neolithic farmer
    ancestry, starting as early as the Late Bronze Age 109 802 . By measuring haplotypes from these
    803 ancestries in modern individuals, we are able to show that these patterns differentiate Wales and
    804 Cornwall as well as Scotland from England. We also found higher levels of WHG-related ancestry
    805 in central and Northern England. These results demonstrate clear ancestry differences within an
    806 ‘ethnic group’ (white British) traditionally considered relatively homogenous
    , which highlights the
    807 need to account for subtle population structure when using resources such as the UK Biobank
    808 genomes.
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    Group label Sample Site Date Y-DNA mtDNA Latitude and Longitude
    Georgia_UpperPaleolithic NEO283 Kotias Klde -23685 U4'9 42.22, 43.32
    Ukraine_Mesolithic NEO550 Vasilevka-I -9086 H2a2a1 48.2, 35.216
    Ukraine_Mesolithic NEO527 Voloshskoe -8974 U2e1 48.305, 35.136
    Russia_Mesolithic NEO202 Peschanitsa -8903 R1a U4a1 61.272, 38.91
    Ukraine_Mesolithic NEO545 Vasilevka-I -8879 I H2a 48.2, 35.216
    Ukraine_Mesolithic NEO494 Vasilevka-I -8791 R1 U5a1d 48.2, 35.216
    Ukraine_Mesolithic NEO496 Vasilevka-I -8705 GHIJK H2a2a1 48.2, 35.216
    Ukraine_Mesolithic NEO492 Vasilevka-I -8705 I2a1b U2e 48.2, 35.216
    Ukraine_Mesolithic NEO497 Vasilevka-I -8705 R1a U4 48.2, 35.216
    Ukraine_Mesolithic NEO501 Vasilevka-I -8672 Q U5a2 48.2, 35.216
    Ukraine_Mesolithic NEO549 Vasilevka-I -8615 I H2a2a1 48.2, 35.216
    Denmark_Mesolithic NEO254 Koelbjerg -8515 I2a1b U5a2c 55.404, 10.133
    Russia_Mesolithic NEO539 Minino -8107 U4 59.7, 39.5
    Russia_Mesolithic NEO100 Ural River Beach -7977 Q1b H2a2a1 51.566, 53.683
    Sweden_Mesolithic NEO18 Hanaskede -7925 I2a1a U5a1 58.391, 13.558
    Serbia_Mesolithic NEO669 Lepenski Vir -7850 H13 44.5555278, 22.02742
    Georgia_Mesolithic NEO281 Kotias Klde -7773 J2b H13c 42.22, 43.32
    Sweden_Mesolithic NEO27 Bredgĺrden -7743 I2a1a U5a2 57.742, 13.403
    Russia_Mesolithic NEO536 Minino -7618 R1b U4 59.7, 39.5
    Spain_Mesolithic NEO694 Santa Maira -7572 I2a1a U2'3'4'7'8'9 38.73, -0.463
    Denmark_Mesolithic NEO13 Hedegaard (Bislev) -7561 I H2a2a1 56.907, 9.576
    Iran_Neolithic NEO817 Tepe Guran -7442 H2a2a1 33.756, 47.097
    Iran_Neolithic NEO819 Tepe Guran -7341 U7a 33.756, 47.097
    Norway_Mesolithic NEO17 Hummervikholmen -7197 I2 U4 58.064, 7.743
    Denmark_Mesolithic NEO91 Strřby Grřftemark -7184 I2a1a2 K1e 55.393, 12.307
    Denmark_Mesolithic NEO759 Křge Sřnakke -7090 I2 U5b2a+@16192 55.402, 12.367
    Romania_Mesolithic NEO672 Schela Cladovei -6864 K1b2 44.635, 22.66
    Iran_Neolithic NEO816 Tepe Guran -6750 J2a R2 33.756, 47.097
    Serbia_Mesolithic NEO677 Vlasac -6726 R1b1a K1+16362 44.5355833, 22.0444444
    Serbia_Mesolithic NEO655 Vlasac -6726 HV+16311 44.5355833, 22.0444444
    Serbia_Mesolithic NEO657 Vlasac -6726 K1b1b1 44.5355833, 22.0444444
    Romania_Mesolithic NEO671 Schela Cladovei -6709 R1b U4a 44.635, 22.66
    Russia_Siberia_Neolithic_Koshinskaya NEO72 Mergen' 6 -6406 Q1b R1b1 56.019, 69.31
    Russia_Neolithic NEO555 Karavaikha -6327 R1b1a1 T2a1b1 60.414, 38.932
    Spain_Mesolithic NEO646 El Mazo -6324 U5b1b1 43.404, -4.707
    Ukraine_Neolithic NEO521 Igren -6320 U5b2b 48.396, 35.161
    Russia_Neolithic NEO559 Karavaikha -6318 R1b1a1 U5a1 60.414, 38.932
    Russia_Mesolithic NEO87 Zamostje 2 -6254 U5a1 56.666, 38.016
    Russia_Siberia_Neolithic NEO116 Pad’ Tokui -6237 C4a1a1 49.8, 108.8
    Russia_Neolithic NEO561 Karavaikha -6234 U5a1 60.414, 38.932
    Denmark_Mesolithic NEO123 Orehoved Sejlrende -6228 U5b1b 54.96, 11.85
    Denmark_Mesolithic NEO19 Rřntesten -6222 I2a1b U5a2b 56.27, 10.472
    Denmark_Mesolithic NEO122 Orehoved Sejlrende -6206 U5a2b 54.96, 11.85
    Russia_Neolithic NEO558 Karavaikha -6135 R1a K1+16362 60.414, 38.932
    Russia_Neolithic NEO554 Pogostishche I -6115 U5a2d 60.45, 38.633
    Russia_Mesolithic NEO537 Minino -6102 R1b1a U4a 59.7, 39.5
    Serbia_Mesolithic NEO658 Lepenski Vir -6065 N1a1a1a2 44.5355833, 22.0444444
    Russia_Neolithic NEO557 Karavaikha -5962 R1b1a1 U4 60.414, 38.932
    Russia_Neolithic NEO560 Karavaikha -5955 R1b1a1 U5a1 60.414, 38.932
    Spain_Mesolithic NEO938 El Mazo -5928 U5a2a 43.404, -4.707
    Russia_Mesolithic NEO88 Zamostje 2 -5908 R1b1a1 U5a1c 56.666, 38.016
    Denmark_Mesolithic NEO600 Vedbaek boldbaner -5865 I2 K1e 55.848, 12.56
    Hungary_Neolithic_Koros NEO130 Dezsk -5853 V1 46.251, 20.259
    Russia_Siberia_Neolithic_Koshinskaya NEO73 Mergen' 6 -5731 U2e3 56.019, 69.31
    Russia_Siberia_Neolithic NEO117 Zhindo -5706 N1a2 C4a1a+195 49.9, 108
    Armenia_Neolithic NEO110 Aknashen -5644 T1a 40.101, 44.294
    Hungary_Neolithic_Koros NEO137 Hödmezövasarhely Kotac -5642 H5b 46.418, 20.33
    Russia_Siberia_Neolithic NEO115 Zhindo -5600 N1a2 F1b1+@152 49.9, 108
    Russia_Siberia_Neolithic NEO922 Vengerovo-2 -5599 Q1b2 U2e3 55.707, 76.783
    Spain_Mesolithic NEO648 El Toral III -5574 I2a1a U5a2a 43.402, -4.707
    Denmark_Mesolithic NEO683 Tybrind Vig -5571 U5a2b 55.395, 9.826
    Denmark_Mesolithic NEO589 Korsřr Nor -5525 I2a1a2 U5a2b 55.331, 11.145
    Latvia_Mesolithic NEO307 Zvejnieki -5506 I2a1b1a2 U5a2d 57.775, 25.229
    Italy_Neolithic NEO834 Mora Cavorso -5460 G2a2a T2b 41.87, 13.283
    Russia_Neolithic_MiddleDon NEO212 Golubaya Krinitsa -5443 I2a1b1a2 U4a2a 50.072, 39.894
    Denmark_Mesolithic NEO587 Kongemose -5427 I2a1a2 U5b1 55.583, 11.551
    Portugal_Neolithic_Cardial NEO632 Gruta do Caldeirao -5397 I2a1a J2b1a 39.648, -8.415
    Denmark_Mesolithic NEO932 Tudse Hage -5352 K1e 55.249, 11.228
    Russia_Neolithic_MiddleDon NEO113 Golubaya Krinitsa -5348 R1a U2e1a 50.073, 39.878
    Russia_Neolithic_MiddleDon NEO209 Golubaya Krinitsa -5346 R1a U5b2 50.072, 39.894
    Italy_Neolithic NEO695 Maddalena di Muccia -5344 J1c3j 43.075, 13.059
    Ukraine_Neolithic NEO270 Mamaj Gora -5343 I2a1b U4d 47.433, 34.268
    Ukraine_Neolithic NEO551 Vovnigi I -5336 U5a2+16362 48.122, 35.096
    Ukraine_Neolithic NEO268 Mamaj Gora -5295 R1b U5a2+16294 47.433, 34.268
    Russia_Neolithic_MiddleDon NEO204 Golubaya Krinitsa -5278 I2a1b U4 50.072, 39.894
    Russia_Siberia_Neolithic_Kitoi NEO199 Fofonovo -5219 Q1b1 R11 52.2, 107
    Russia_Siberia_Neolithic_Kitoi NEO841 Sosnovyiy Myis -5206 A+152+16362+16189 58.752, 102.634
    Czech_Neolithic_LBK NEO128 Vedrovice -5191 G K1a 49.021, 16.38
    Portugal_Neolithic_Cardial NEO631 Gruta do Caldeirao -5185 I2a1a T2b3+151 39.648, -8.415
    Denmark_Mesolithic NEO748 Henriksholm-Břgebakken (Vedbćk) -5179 I2 U5b1d2 55.853, 12.559
    Denmark_Mesolithic NEO814 Bodal K -5165 U4a2 55.586, 11.538
    Russia_Neolithic_MiddleDon NEO210 Golubaya Krinitsa -5158 R1 U5a2b 50.072, 39.894
    Russia_Neolithic_MiddleDon NEO207 Golubaya Krinitsa -5147 U5a1c 50.072, 39.894
    Ukraine_Neolithic NEO516 Igren -5146 Q1b U5a1i 48.396, 35.161
    Denmark_Mesolithic NEO749 Henriksholm-Břgebakken (Vedbćk) -5122 I2a1a2 U5b1d2 55.853, 12.559
    Russia_Siberia_Neolithic_Kitoi NEO201 Fofonovo -5114 Q1b1 C4 52.2, 107
    Russia_Siberia_Neolithic_Kitoi NEO843 Sosnovyiy Myis -5110 R11 58.752, 102.634
    Russia_Siberia_Neolithic_Sredneirtyshskaya NEO75 Omskaya Stoyanka II -5108 Q1a2 T2d 54.971, 73.366
    Russia_Siberia_Neolithic NEO63 Itkul -5107 Q1a A+152+16362 52.713, 84.692
    Ukraine_Neolithic NEO498 Vovnigi II -5106 R1b1b U4a1 48.132, 35.081
    Russia_Siberia_Neolithic NEO67 Ust'-Isha -5098 C2 U5a2 52.18, 85.965
    Denmark_Mesolithic NEO791 Korsör Nor -5083 I2 U5b1 55.331, 11.145
    Denmark_Mesolithic NEO586 Koed -5080 I2 U5a2c 56.371, 10.573
    Russia_Neolithic NEO556 Karavaikha -5080 U4a1 60.414, 38.932
    Russia_Neolithic_Lyalovo NEO185 Sakhtish IIa -5078 Q1b1 U4a1 56.781, 40.447
    Russia_Siberia_Neolithic_Kitoi NEO200 Fofonovo -5057 Q1b1 C4 52.2, 107
    Ukraine_Neolithic NEO304 Volnensky -5056 I2a1b U5a2d 47.95, 35.39
    Denmark_Mesolithic NEO746 Henriksholm-Břgebakken (Vedbćk) -5036 I U4a2 55.853, 12.559
    Denmark_Mesolithic NEO583 Koed -5030 U5a2b 56.371, 10.573
    Hungary_Neolithic_Tisza NEO149 Gorzsa Cukormajor -5028 I N1a1a1a 46.369, 20.424
    Ukraine_Neolithic NEO553 Vovnigi I -5027 U5a 48.122, 35.096
    Ukraine_Neolithic NEO524 Vovnigi II -5026 I2a1b U5a1j 48.132, 35.081
    Denmark_Mesolithic NEO822 Dragsholm -5023 U5a2b 55.768, 11.388
    Ukraine_Neolithic NEO552 Vovnigi I -5020 I2a1b U2e1 48.122, 35.096
    Russia_Siberia_Neolithic_Sredneirtyshskaya NEO78 Omskaya Stoyanka II -5003 Q1a2 U4a1 54.971, 73.366
    Russia_Neolithic_Lyalovo NEO186 Sakhtish IIa -4969 R1b1a1 U5a1d 56.781, 40.447
    Sweden_Mesolithic NEO679 Skateholm I -4957 U5b2a1a 55.386, 13.481
    Denmark_Mesolithic NEO930 Fannerup F -4942 I2 U5b2b1a 56.4, 10.714
    Ukraine_Neolithic NEO507 Vovnigi II -4937 R1b U5b2 48.132, 35.081
    Russia_Neolithic_Lyalovo NEO192 Sakhtish II -4892 K1b2 56.78, 40.447
    Ukraine_Neolithic NEO528 Vovnigi I -4892 I2a1b T2 48.122, 35.096
    Ukraine_Neolithic NEO302 Volnensky -4889 I2a1b1a2 U4a 47.95, 35.39
    Denmark_Mesolithic NEO733 Dragsholm -4869 U5b2b 55.768, 11.388
    Denmark_Mesolithic NEO732 Dragsholm -4862 U5b2b 55.768, 11.388
    Denmark_Mesolithic NEO745 Henriksholm-Břgebakken (Vedbćk) -4851 I2 U5b1d2 55.853, 12.559
    Russia_Siberia_Neolithic NEO911 Vengerovo-2 -4830 Q1b2 C5c 55.707, 76.783
    Denmark_Mesolithic NEO856 Nederst -4828 U5a2b 56.365, 10.643
    Russia_Neolithic NEO172 Ksizovo 6 -4824 R1b U5a2b2 52.278, 38.955
    Hungary_Neolithic_Tisza NEO145 Gorzsa Cukormajor -4799 T2c1d1 46.369, 20.424
    Ukraine_Neolithic NEO529 Vovnigi I -4794 U5a1 48.122, 35.096
    Denmark_Mesolithic NEO747 Henriksholm-Břgebakken (Vedbćk) -4778 I2a1b R1b1 55.853, 12.559
    Hungary_Neolithic_Tisza NEO147 Gorzsa Cukormajor -4774 J1c6 46.369, 20.424
    Hungary_Neolithic_Tisza NEO140 Gorzsa Cukormajor -4766 H T2b23 46.369, 20.424
    Ukraine_Neolithic NEO503 Vovnigi II -4756 U5a2d 48.132, 35.081
    Russia_Neolithic NEO162 Vasilyevskiy kordon 17 -4731 U5a1d1 52.9, 40.03
    Ukraine_Neolithic NEO300 Volnensky -4728 U5b2 47.95, 35.39
    Hungary_Neolithic_Tisza NEO143 Gorzsa Cukormajor -4706 J2a T2c1d1 46.369, 20.424
    Hungary_Neolithic_Tisza NEO142 Gorzsa Cukormajor -4675 U1a1a 46.369, 20.424
    Ukraine_Neolithic NEO514 Vovnigi II -4673 I2 H2a 48.132, 35.081
    Ukraine_Neolithic NEO502 Vovnigi II -4666 I2a1b U5b1 48.132, 35.081
    Hungary_Neolithic_Tisza NEO148 Gorzsa Cukormajor -4662 U5b2c 46.369, 20.424
    Ukraine_Neolithic NEO305 Vasilevsky -4661 I2a1b1a2 U4a 48.305, 35.139
    Denmark_Mesolithic NEO1 Holmegĺrd-Djursland -4644 I2 U5a'b 56.251, 10.755
    Turkmenistan_Neolithic NEO310 Monjukli-Depe -4600 L1a2 H14 36.848, 60.418
    France_Neolithic_Cardial NEO812 Grotte du Gazel -4595 H1 43.324, 2.42
    France_Neolithic_Cardial NEO813 Grotte du Gazel -4591 H1 43.324, 2.42
    Ukraine_Neolithic NEO262 Lysa Gora -4563 R1b U5a1+@16192 47.451, 35.239
    Denmark_Mesolithic NEO941 Havnř -4424 U5b1b1+@16192 56.709, 10.171
    Denmark_Mesolithic NEO570 Fannerup E -4422 U5b2c 56.401, 10.718
    Russia_Neolithic NEO173 Ksizovo 6 -4393 U5a2b2 52.278, 38.955
    Denmark_Mesolithic NEO751 Bjřrnsholm -4365 I2 U5b2b1a 56.874, 9.215
    Denmark_Mesolithic NEO852 Norsminde -4356 U5b1 56.027, 10.257
    Ukraine_Neolithic NEO508 Vovnigi II -4354 U4a1 48.132, 35.081
    Denmark_Mesolithic NEO855 Fannerup D -4349 I2a1b1a2 U5b1b1+@16192 56.401, 10.715
    Denmark_Mesolithic NEO568 Ertebřlle -4347 I2 U5b1 56.812, 9.179
    Denmark_Mesolithic NEO569 Ertebřlle -4195 I2 U5b2a1a 56.812, 9.179
    Denmark_Mesolithic NEO3 Vćngesř II -4142 I U5a 56.146, 10.552
    Denmark_Mesolithic NEO598 Sřlager -4118 R1b1 55.948, 11.897
    Ukraine_Neolithic NEO522 Voloshskoe -4105 I2a1b U5a1 48.305, 35.136
    Russia_Siberia_Neolithic NEO64 Itkul -4100 C2b R1b1 52.713, 84.692
    Denmark_Mesolithic NEO853 Langř Skaldynge -4095 R1b1 55.545, 10.622
    Russia_Siberia_Neolithic NEO292 Dolgoe Ozero -4086 Q1b1 A+152+16362+16189 56.2, 95.7
    Russia_Neolithic_Volosovo NEO181 Sakhtish IIa -4084 Q1b2a U2e1e 56.781, 40.447
    Denmark_Mesolithic NEO960 Ravnsbjerggĺrd II -4013 I2a1a U5b2a2 55.581, 11.578
    Russia_Neolithic_Volosovo NEO180 Sakhtish IIa -3989 R1b1a1 U5b2c1 56.781, 40.447
    Denmark_Mesolithic NEO645 Rřdhals -3886 I2 K1e 55.914, 11.091
    Russia_Neolithic NEO171 Vasilyevskiy kordon 17 -3885 U5a1+@16192 52.9, 40.03
    Russia_Siberia_Neolithic NEO309 Protoka -3874 Q1b2 A10 56.77, 77.03
    Sweden_Mesolithic NEO260 Evensĺs -3872 I2 R1b1 58.246, 11.492
    Russia_Siberia_Neolithic NEO923 Vengerovo-2 -3864 A10 55.707, 76.783
    Denmark_Neolithic NEO601 Viksř Mose -3857 K1a2b 55.759, 12.238
    Denmark_Mesolithic NEO962 Dragsholm -3836 I U5b2b 55.768, 11.388
    Russia_Siberia_Neolithic NEO102 Afontova Gora -3831 Q U2e1h 56.016, 92.866
    Russia_Neolithic_Volosovo NEO158 Sakhtish II -3804 R1b1a U2e1e 56.78, 40.447
    Russia_Neolithic_Volosovo NEO195 Sakhtish IIa -3797 I2a1a U5a2d 56.781, 40.447
    Russia_Siberia_Neolithic NEO916 Vengerovo-2 -3777 K2b H2a2a1 55.707, 76.783
    Russia_Siberia_Neolithic NEO915 Vengerovo-2 -3777 Q1 C4+152 55.707, 76.783
    Denmark_Neolithic NEO891 Roskilde Fjord (syd for Jyllinge) -3764 H 55.73, 12.1
    Russia_Neolithic NEO164 Vasilyevskiy kordon 17 -3719 R1a U5a1d1 52.9, 40.03
    Denmark_Neolithic NEO790 Tysmose -3718 I2a1a2 HV16 55.711, 12.274
    Russia_Siberia_Neolithic NEO83 Borovjanka XVII -3714 R1b1a1 U4a 55.733, 74.285
    Russia_Neolithic NEO166 Vasilyevskiy kordon 17 -3713 R1b1a1 U5a1f1 52.9, 40.03
    Russia_Neolithic NEO167 Vasilyevskiy kordon 17 -3706 U2e1e 52.9, 40.03
    Russia_Siberia_Neolithic NEO907 Vengerovo-2 -3699 C2 A8a 55.707, 76.783
    Russia_Neolithic NEO163 Vasilyevskiy kordon 17 -3654 J1 U5a2 52.9, 40.03
    Russia_Neolithic_Volosovo NEO189 Sakhtish IIa -3654 R1a U5a2d 56.781, 40.447
    Russia_Neolithic_Volosovo NEO194 Sakhtish IIa -3647 R1b1a1 U4b1b1 56.781, 40.447
    Romania_Neolithic NEO674 Baile Herculane -3640 I2a1b U8b1b1 44.896, 22.428
    Sweden_Neolithic_FBC NEO46 Ängdala -3614 T2b 55.588, 13.113
    Russia_Neolithic NEO170 Vasilyevskiy kordon 17 -3613 U2e1e 52.9, 40.03
    Denmark_Neolithic NEO29 Lohals -3609 H 55.134, 10.903
    Denmark_Neolithic NEO595 Pandebjerg -3595 I2a1a2 J1c 55.789, 11.292
    Denmark_Neolithic NEO23 Jorlřse mose -3586 I2a1a2 U5b1b2 55.601, 11.312
    Russia_Siberia_Neolithic NEO912 Vengerovo-2 -3586 C4+152 55.707, 76.783
    Denmark_Neolithic NEO564 Bygholm Nřrremark -3585 I2a1a J1c3j 55.875, 9.812
    Denmark_Neolithic NEO571 Grřfte -3585 H2a2a 55.404, 11.497
    Ukraine_Neolithic NEO265 Lysa Gora -3585 I2a1b U5a2+16362 47.451, 35.239
    Denmark_Neolithic NEO753 Sigersdal Mose 2 -3583 J1c 55.772, 12.213
    Denmark_Neolithic NEO41 Rude -3581 I J2a1a1a 55.994, 10.248
    Denmark_Neolithic NEO942 Tissře -3545 T2b 55.575, 11.294
    Russia_Neolithic_Volosovo NEO179 Sakhtish II -3515 U5a2d 56.78, 40.447
    Russia_Siberia_Neolithic NEO917 Vengerovo-2 -3512 Q1a2 C4 55.707, 76.783
    Britain_Neolithic NEO847 Hetty Peglers Tump -3511 I2a1b1a1a X2b4 51.697, -2.304
    Denmark_Neolithic NEO28 Salpetermosen syd 8 -3511 I2a1a2 T2b11 55.91, 12.307
    Denmark_Neolithic NEO886 Dalmosegaard -3508 I2a1a2 H26 54.989, 12.419
    Russia_Siberia_Neolithic NEO918 Vengerovo-2 -3508 U2e2a 55.707, 76.783
    Denmark_Neolithic NEO866 Lundby-Falster -3507 R1b1a K1a2b 54.865, 11.841
    Russia_Neolithic_Volosovo NEO184 Sakhtish 8 -3507 U5a2d 56.776, 40.443
    Russia_Neolithic_Volosovo NEO183 Sakhtish IIa -3506 R1b U5b1b1+@16192 56.781, 40.447
    Russia_Neolithic NEO168 Vasilyevskiy kordon 17 -3503 U5a1+@16192 52.9, 40.03
    Denmark_Neolithic NEO757 Sejerby (Sejerř) -3502 T2b 55.902, 11.118
    Russia_Neolithic_Volosovo NEO193 Sakhtish IIa -3501 R1b U2e1e 56.781, 40.447
    Denmark_Neolithic NEO896 Mandemarke -3494 H4a1a1a 54.969, 12.494
    Denmark_Neolithic NEO945 Lćsten Mose -3494 H1b1 56.485, 9.827
    Russia_Neolithic NEO687 Kumyshanskaya Cave -3493 R1a U5a1d2 57.58, 58.2
    Denmark_Neolithic NEO933 Sludegĺrd Sřmose -3488 I2 V 55.253, 10.746
    Italy_Neolithic NEO830 Fontenoce -3443 I1c1a 43.383, 13.55
    Denmark_Neolithic NEO43 Rude -3442 I J2a1a1a 55.994, 10.248
    Russia_Neolithic_Volosovo NEO188 Sakhtish IIa -3440 I2a1b U5a1f1 56.781, 40.447
    Denmark_Neolithic NEO888 Elkenřre -3439 H5 54.732, 11.986
    Russia_Neolithic_Volosovo NEO182 Sakhtish IIa -3436 R1b U5b1b1+@16192 56.781, 40.447
    Denmark_Neolithic NEO744 Vig Femhřve -3413 Q1b2 K1a2a 55.856, 11.59
    Denmark_Neolithic NEO795 Porsmose -3380 I2a1a2 H24 55.276, 11.875
    Russia_Neolithic NEO174 Ksizovo 6 -3357 W6a 52.278, 38.955
    Denmark_Neolithic NEO702 Jřrlundegĺrd -3321 I K1a1a2 55.828, 12.166
    Russia_Neolithic_Volosovo NEO178 Sakhtish II -3318 I2a1a U5a1d 56.78, 40.447
    Russia_Neolithic NEO160 Vasilyevskiy kordon 17 -3313 U5a1d1 52.9, 40.03
    Denmark_Neolithic NEO935 Vibygĺrds Mose -3298 I2a1a2 H 55.564, 12.016
    Russia_Neolithic_Volosovo NEO197 Sakhtish IIa -3295 I2a1a U5a2d 56.781, 40.447
    Denmark_Neolithic NEO7 Sigersdal Mose -3290 CT H 55.761, 12.258
    Denmark_Neolithic NEO597 Storelyng (Řgĺrde boat III) -3264 I2a1a2 U5b1b2 55.587, 11.574
    Denmark_Neolithic NEO865 Lundby-Falster -3234 I2 J1c3g 54.865, 11.841
    Denmark_Neolithic NEO594 Neverkćr mose -3231 I2a1b1a2 K1a+195 55.391, 10.078
    Denmark_Neolithic NEO961 Avlebjerg (Strřby) -3199 K1b1a1 55.368, 12.287
    Denmark_Neolithic NEO599 Vanlřse mose -3191 Q1b K2b1a 55.553, 11.684
    Denmark_Neolithic NEO602 Storelyng (Řstrup Homo II) -3189 I2 U5b2a1b 55.589, 11.576
    Denmark_Neolithic NEO33 Vittrup -3189 I2a1a H2a 57.391, 9.761
    Denmark_Neolithic NEO566 Dřjringe -3185 I2a1a U5b3 55.484, 11.579
    Britain_Neolithic NEO627 Banks tomb -3182 I2a1a2 U5b1c 58.734, -2.937
    Poland_Neolithic_FBC NEO641 Slonowice -3182 H7d 50.267, 20.45
    Sweden_Neolithic NEO36 Hindby mosse -3147 K2b1a 55.565, 13.036
    Britain_Neolithic NEO626 Banks tomb -3132 I2a1a2 H1 58.734, -2.937
    Denmark_Neolithic NEO898 Svinninge Vejle -3131 I2a1a2 K1a3a 55.736, 11.425
    Britain_Neolithic NEO625 Banks tomb -3130 I2a1a2 U5b1c 58.734, -2.937
    Sweden_Neolithic NEO39 Hindby mosse -3124 I2a1a2 H49 55.565, 13.036
    Sweden_Neolithic NEO38 Hindby mosse -3121 K1a3 55.565, 13.036
    Denmark_Neolithic NEO25 Kainsbakke II -3116 K1a+195 56.433, 10.789
    Sweden_Neolithic NEO259 Frälsegĺrden -3010 I2a1a K2a 58.164, 13.454
    Denmark_Neolithic NEO925 Myrebjerg mose -2997 W1 54.771, 10.675
    Russia_Neolithic_Volosovo NEO187 Sakhtish IIa -2990 U2e1e 56.781, 40.447
    Poland_Neolithic_FBC NEO640 Slonowice -2950 G2a2b2a H 50.267, 20.45
    Russia_Siberia_BronzeAge_Okunevo NEO68 Okunevo 5 and 7 -2949 D4j 56.438, 74.907
    Britain_Neolithic NEO630 Banks tomb -2947 I2 H67 58.734, -2.937
    Britain_Neolithic NEO624 Banks tomb -2947 K1b1a1 58.734, -2.937
    Britain_Neolithic NEO717 Banks tomb -2900 I2a1a2 K1a+195 58.734, -2.937
    Russia_Siberia_BronzeAge NEO65 Kostenkova Isbushka -2854 C2 A8a1 52.724, 84.683
    Russia_Siberia_BronzeAge NEO80 Borovjanka XVII -2842 C 55.733, 74.285
    Italy_Neolithic NEO828 Gaudo -2796 J2a N1a1a1 40.416, 15.066
    Russia_Siberia_BronzeAge_Okunevo NEO77 Okunevo 5 and 7 -2782 Q1b C4+152 56.438, 74.907
    Italy_BronzeAge NEO823 Grotta Nisco -2715 G2a2b U1a1a 40.883, 16.733
    Russia_Siberia_BronzeAge_Okunevo NEO79 Okunevo 5 and 7 -2681 U2e1'2'3 56.438, 74.907
    Denmark_Neolithic NEO943 Stenderup Hage -2667 I2a1b1a1b H+152 55.46, 9.694
    Denmark_Neolithic NEO580 Klokkehřj -2662 H2a2a1 55.108, 10.111
    Russia_Neolithic NEO175 Ksizovo 6 -2657 I2a1b U2e1'2'3 52.278, 38.955
    France_Neolithic NEO121 Grotte Mandrin -2577 I2a1a1a1a1 U4a2f 44.466, 4.766
    Estonia_Neolithic_CWC NEO306 Sope -2548 R1b1 59.408, 27.026
    Denmark_Neolithic NEO792 Nćs -2546 I2a1a2 U2e2a1a2 54.87, 12.116
    Kazakhstan_BronzeAge NEO57 Biestamak -2546 A10 52.166, 64.533
    Spain_Neolithic NEO721 Camino de las Yeseras -2545 U5b3 40.439, -3.501
    Sweden_Neolithic NEO44 Dösemarken -2436 J1c2 55.564, 12.946
    Denmark_Neolithic NEO876 Toftum mose -2435 R1a T2a1a 55.895, 9.987
    France_Neolithic NEO119 Grotte Mandrin -2432 U5b3 44.466, 4.766
    Sweden_Neolithic NEO51 Kastanjegĺrden -2391 U4d1 55.555, 13.023
    Portugal_Neolithic NEO603 Sao Paulo 2 -2387 V+@16298 38.676, -9.164
    Portugal_Neolithic NEO609 Sao Paulo 2 -2381 I2a1a K1a1 38.676, -9.164
    Denmark_Neolithic NEO870 Toftum mose -2305 R1b1a1b1a1 T2b2 55.895, 9.987
    Denmark_Neolithic NEO92 Barhřj (Strřby Egede) -2268 R1b1a1b1a1 T2b 55.403, 12.272
    Spain_BronzeAge NEO649 El Toral III -2166 R1b1a1b1a1 H1+16355 43.402, -4.707
    Denmark_Neolithic NEO738 Kolind -2161 I2a1b1a2b J1c3g 56.362, 10.593
    Denmark_Neolithic NEO737 Borreby -2161 R1b1a1b1a1 U4b1a1a1 55.225, 11.303
    Denmark_Neolithic NEO861 Mosede Mose -2161 H6a1a 55.576, 12.275
    Spain_BronzeAge NEO650 El Toral III -2160 R1b1a1b1a1 J1c1 43.402, -4.707
    Russia_Siberia_BronzeAge NEO921 Vengerovo-2 -2148 R1b1 55.707, 76.783
    Russia_Siberia_BronzeAge NEO76 Ostrov 2 -2117 U4a2 56.933, 66.467
    Ukraine_BronzeAge NEO509 Igren -2114 R1b1a1b1b U5a1j 48.396, 35.161
    Denmark_Neolithic NEO878 Kyndelřse -2088 U4a2 55.703, 11.855
    Denmark_Neolithic NEO872 Toftum mose -2066 R1b1a1b1a1 K2a5 55.895, 9.987
    Denmark_Neolithic NEO735 Borreby -2045 R1b1a1b1a1 U4c1 55.225, 11.303
    Denmark_Neolithic NEO875 Toftum mose -2036 I1 U5b3b 55.895, 9.987
    Sweden_Neolithic NEO220 Falköping 5 -2036 I1 H13a1a 58.166, 13.573
    Denmark_Neolithic NEO739 Kolind -2021 I2a1b1a2 N1a1a1a1 56.362, 10.593
    Sweden_Neolithic NEO223 Falköping 5 -2018 I1 W1b1 58.166, 13.573
    Sweden_Neolithic NEO227 Falköping 5 -1974 I1 H1a3 58.166, 13.573
    Sweden_Neolithic NEO224 Falköping 5 -1969 HV6 58.166, 13.573
    Sweden_Neolithic NEO261 Sillvik -1945 I1 T2b 57.736, 11.744
    Sweden_Neolithic NEO228 Falköping 5 -1891 I1 H3h 58.166, 13.573
    Denmark_Neolithic NEO934 Gammellung -1884 W1+119 54.876, 10.784
    Kazakhstan_BronzeAge NEO900 Sjauke -1854 N1 U2e1 52.431, 76.838
    Sweden_Neolithic NEO52 Vattenledningen -1816 R1b1a1b1a1 H3g4 55.503, 13.009
    Sweden_Neolithic NEO226 Falköping 5 -1811 I3d 58.166, 13.573
    Sweden_Neolithic NEO225 Falköping 5 -1798 U5b2a1a1 58.166, 13.573
    Denmark_BronzeAge NEO93 Strřby Ladeplads -1788 I1 K1a1b2b 55.406, 12.285
    Kazakhstan_BronzeAge NEO904 Sjiderti 10 -1750 C4a1 51.651, 74.654
    Sweden_Neolithic NEO221 Falköping 5 -1748 R1a1a1b1a2b H1a3 58.166, 13.573
    Denmark_BronzeAge NEO857 Lollikehuse -1718 I1 H3v+16093 55.758, 12.007
    Denmark_Neolithic NEO860 Mosede Mose -1717 R1b1a1b1a1 K1a1b2a1a 55.576, 12.275
    Denmark_BronzeAge NEO752 Madesř -1699 R1b1a1b1a1a1b U5a1i1 55.621, 11.331
    Kazakhstan_BronzeAge NEO899 Gregorievka 1 -1683 U2e1 52.629, 76.727
    Russia_Siberia_BronzeAge NEO910 Vengerovo-2 -1645 C4+152 55.707, 76.783
    Denmark_BronzeAge NEO815 Vasagĺrd -1524 I1 H3b6 55.055, 14.885
    Spain_BronzeAge NEO653 El Toral III -1468 HV0d 43.402, -4.707
    France_BronzeAge NEO120 Grotte Mandrin -1409 U5a2b3 44.466, 4.766
    Denmark_BronzeAge NEO563 Bybjerg -1405 I1 H6a1a 55.76, 11.798
    Denmark_BronzeAge NEO951 Klćsterupholm Mose -1356 W1 57.29, 10.033
    Russia_BronzeAge NEO61 Bol'shoy Oleni Ostrov -1356 U5a2d 68.966, 33.083
    Russia_BronzeAge NEO60 Bol'shoy Oleni Ostrov -1350 U5a1d2 68.966, 33.083
    Denmark_BronzeAge NEO590 Magleř -1347 I1a2a1 K2a3 55.34, 11.161
    Russia_Siberia_BronzeAge NEO81 Borovjanka XVII -1347 R1a C5c 55.733, 74.285
    Russia_BronzeAge NEO62 Bol'shoy Oleni Ostrov -1336 Z1a1a 68.966, 33.083
    Denmark_BronzeAge NEO946 Hove Ĺ -1195 R1b1a1b1a1a1b U4a1a 55.708, 12.25
    Italy_BronzeAge NEO806 Grotta Delle Mura -1063 J2b2a1a1a1a1a1b N1a1a1 40.957, 17.29
    Russia_Siberia_BronzeAge NEO70 Bazaiha -802 N1 F1b1b 55.956, 92.791
    Ukraine_IronAge NEO278 Kleshnya III -793 R1a U5b1a 48.966, 38.238
    Russia_IronAge NEO538 Minino -300 N1a1a1a1a2 U5a1d2b 59.7, 39.5
    Ukraine_IronAge NEO518 Igren -202 R1a1a1b2 U5a2d 48.396, 35.161
    Kazakhstan_IronAge NEO902 Sjauke 1 -36 Q1b J1b1a1 52.431, 76.838
    Kazakhstan_Medieval NEO901 Sjauke settlement 782 D4e4 52.43, 76.839

  6. #6
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    bump

    Is this the final nail in the coffin for non-European PIE?

    If so, it's been emotional
    Spoiler!

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    I would like to see the autosomal makeup of the Volosovo samples from Russia, they're supposedly one of the main pre-IE cultures in European Russia, as well as that Northern Russian sample from 300 BC with N1a (NEO538), it might be a good pre-Slavic Northern Russian proxy.

    NEO518 from Igren, which is apparently in Dnepropetrovsk oblast, Ukraine, may be some kind of Scythian or Sarmatian.

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    Quote Originally Posted by Creoda View Post
    bump

    Is this the final nail in the coffin for non-European PIE?

    If so, it's been emotional
    I bet steppe hunter-gatherers even more tilted towards CHG will be found further east, around the Volga, which will make Eneolithic steppe pastoralists nearly fully descended from steppe HGs. This CHG stuff is probably very old in the steppe.

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    Thank you for posting. Really interesting paper which I will look at in more detail.

    774 The various hunter-gatherer ancestries are not homogeneously distributed amongst modern
    775 populations (Fig. 5). WHG-related ancestry is highest in present-day individuals from the Baltic
    776 States, Belarus, Poland, and Russia; EHG-related ancestry is highest in Mongolia, Finland, Estonia
    777 and Central Asia; and CHG-related ancestry is maximised in countries east of the Caucasus, in
    Pakistan, India, Afghanistan and Iran, in accordance with previous results 103 778 . The CHG-related
    779 ancestry likely reflects both Caucasus hunter-gatherer and Iranian
    Neolithic signals, explaining the
    relatively high levels in south Asia 104. Consistent with expectations 105,106 780 , Neolithic Anatolian
    781 related farmer ancestry is concentrated around the Mediterranean basin, with high levels in southern
    782 Europe, the Near East, and North Africa, including the Horn of Africa, but is less frequent in
    783 Northern Europe.
    This is in direct contrast to the Steppe-related ancestry, which is found in high
    784 levels in northern Europe, peaking in Ireland, Iceland, Norway, and Sweden, but decreases further
    785 south.
    There is also evidence for its spread into southern Asia. Overall, these results refine global
    786 patterns of spatial distributions of ancient ancestries amongst modern populations.
    The proportion of
    794 Neolithic farmer ancestry is highest in southern and eastern England today and lower in Scotland,
    795 Wales, and Cornwall. Steppe-related ancestry is inversely distributed, peaking in the Outer
    Hebrides and Ireland
    , a pattern only previously described for Scotland
    108 796 . This regional pattern was
    797 already evident in the Pre-Roman Iron Age and persists to the present day even though immigrating
    798 Anglo-Saxons had relatively less Neolithic farmer ancestry than the Iron-Age population of
    799 southwest Briton (Extended Data Fig. 4). Although this Neolithic farmer/steppe-related dichotomy
    800 mirrors the modern ‘Anglo-Saxon’/‘Celtic’ ethnic divide, its origins are older, resulting from
    801 continuous migration from a continental population relatively enhanced in Neolithic farmer
    ancestry, starting as early as the Late Bronze Age 109 802 ... We also found higher levels of WHG-related ancestry
    805 in central and Northern England. These results demonstrate clear ancestry differences within an
    806 ‘ethnic group’ (white British) traditionally considered relatively homogenous, which highlights the
    807 need to account for subtle population structure when using resources such as the UK Biobank
    808 genomes.
    Will look at this paper properly tomorrow as it's late here.

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    The original Borreby guys were R1b and very steppe-rich.
    Borreby, NEO735+737; Magleby 04.04.11-45, Zealand. Passage grave
    Anders Fischer, Morten E. Allentoft and Martin Sikora
    Originally the Borreby passage grave was covered in a tumulus framed by up to 1Ľ m high
    stones that formed a semicircle 16-18 m in diameter. The oval burial chamber, horizontal
    dimensions c. 5˝ by 2 m, was constructed of eleven upright stones and 3 cap stones. The
    ESE facing, 5˝ m long entrance was constructed from 5 pairs of vertical stones and a pair of
    door jambs, topped by capstones (Fig. S6.5). When excavated in 1859 the chamber and
    entrance was full to the roof with human skeletal parts, in between which were scattered
    burial gifts of Middle Neolithic and Late Neolithic date. The excavator judged there were
    remains of at least 60-70 individuals. The Meso-Neo project sampled three of them,
    characterised by unusually coarse facial characteristics (‘Borreby type’ according to previous
    literature) and suspected to represent foreign ancestry (cf. text on the Madesř individual
    below). The two of these skulls that produced DNA of acceptable quality were AMS dated to
    the Late Neolithic period.
    Our genetic analyses have indeed documented shared ancestry for these two individuals,
    and for our third Borreby type individual, from Madesř, since they are all of Y-chromosomal
    haplotype R1b, falling within the earlier cluster of Scandinavian Late Neolithic and Bronze
    Age individuals. Interestingly the non-Scandinavian individuals of this cluster are generally
    from Western Europe, and the R1b haplogroups are also more common there. If this points
    to migration it would likely be from there (cf. main text’s chapter: Fine-scale structure and
    multiproxy analysis of Danish transect).
    Literature: Bröste et al. 1956, p. 32014; Ebbesen 2008, p. 12515; Bennike & Alexandersen
    2002, p. 29716; Hansen 1993, p. 11517.
    Also notice the use of physical antropological terminology in the supplemental files. So much for those that claim physiscal anthropology is dead pseudoscience.

    NEO900 Shauke 1 Burial 2 Burial No. 2 Y-Hg N1 "contained an open, jar-shaped vessel, decorated with a “stepping” comb stamp (made by rolling). A piece of a stone crucible and a tanged arrowhead, and bones of sheep and calf were found nearby. Skeletal remains belonged to a man, 40–50 years old. The skull is dolichocephalic, Europeoid with a narrow face, similar to the type of male population of the Yelunin culture of the Upper Ob region. The sample was dated to 3831±20 uncal BP, OxA39284".

    NEO902 Shauke 1 Burial 3 Y-Hg Q1b "contained two wall sherds of a clay vessel, one of which had a raised cordon decorated with comb stamp impressions. A fragment of an astragalus of a small ruminant covered with green oxides from non-ferrous metal, a fragment of a metal object and 37 animal astragali (cattle – 2 pcs, sheep – 35 pcs), as well as horse and sheep bones were found nearby. Bone remains belonged to an adolescent, presumably male, 14–16 years old. The skull is dolichocephalic, probably Europeoid with somewhat softened Europeoid features. Possibly similar to the type of population of the Yelunin and Krotovo cultures in the interfluve of the Ob and Irtysh. The NEO902 sample was dated to the Iron Age, 2337±18 uncal BP, OxA-39286. This sample therefore most likely comes from another individual, and the bone assemblage may be considered as mixed, probably in connection with the gravel extraction".

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