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Eastern hunter-gatherer

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Eastern hunter-gatherer
Artifacts and forensic reconstruction of an eastern hunter-gatherer from the site ofYuzhny Oleny island(datedc. 8,100 BP), byM. M. Gerasimov.National Museum of Karelia.[1]
Hunter-gatherers in Europe between 14 ka and 9 ka, with the main area of Eastern Hunter-Gatherers (EHG, ). Individual numbers correspond to calibrated sample dates.[2]

Inarchaeogenetics,eastern hunter-gatherer (EHG),sometimeseast European hunter-gathereroreastern European hunter-gatherer,is a distinct ancestral component that representsMesolithichunter-gatherersofEastern Europe.[3]

The eastern hunter-gatherer genetic profile is mainly derived fromAncient North Eurasian(ANE) ancestry, which was introduced fromSiberia,[4]with a secondary and smaller admixture of Europeanwestern hunter-gatherers(WHG).[5][6]Still, the relationship between the ANE and EHG ancestral components is not yet well understood due to lack of samples that could bridge the spatiotemporal gap.[5]

During the Mesolithic, the EHGs inhabited an area stretching from theBaltic Seato theUralsand downwards to thePontic–Caspian steppe.[7]Along withScandinavian hunter-gatherers(SHG) andwestern hunter-gatherers(WHG), the EHGs constituted one of the three main genetic groups in the postglacial period of earlyHoloceneEurope.[8]The border between WHGs and EHGs ran roughly from the lowerDanube,northward along the western forests of theDniepertowards the westernBaltic Sea.[9]

During theNeolithicand earlyEneolithic,likely during the4th millennium BCEHGs on the Pontic–Caspian steppe mixed withCaucasus hunter-gatherers(CHGs) with the resulting population, almost half-EHG and half-CHG, forming the genetic cluster known asWestern Steppe Herder(WSH).[10][11][12]WSH populations closely related to the people of theYamnaya cultureare supposed to have embarked on amassive migrationleading to the spread ofIndo-European languagesthroughout large parts of Eurasia.

Research

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Schematic formation of the EHGs, through a main ancestry ofAncient North Eurasians(ANE), and a smaller admixture of WHGs
Genetically, the EHG (red) were most closely related to the ANE (pink).

Haak et al. (2015) identified the EHG as a distinct genetic cluster in two males only. The EHG male ofSamara(dated to ca. 5650–5550 BC) carriedY-haplogroupR1b1a1a*andmt-haplogroupU5a1d.The other EHG male, buried inKarelia(dated to ca. 5500-5000 BC) carried Y-haplogroupR1a1and mt-haplogoupC1g.The authors of the study also identified a WHG cluster and an SHG cluster, intermediate between WHG and EHG.[a]They suggested that EHGs harbored mixed ancestry fromAncient North Eurasians(ANEs) and WHGs.[14][15] Researchers have proposed various admixture proportion models for EHGs from WHGs and ANEs.[16][17]Posth et al. (2023) found that most EHG individuals carried c. 70% ANE ancestry and c. 30% WHG ancestry The WHG-like ancestry was most likely not derived from the Oberkassel and Villabruna clusters directly, but from a related and yet unsampledEpigravettianpopulation.[2][18]The high contribution from Ancient North Eurasians is also visible in a subtle affinity of the EHG to the 40,000-year-oldTianyuan manfromNorthern Chinaand other East/Southeast Asians, which can be explained by geneflow from a Tianyuan-related source into the ANE lineage (represented by Malta and Afontova Gora 3), which later substantially contributed to the formation of the EHG.[19]

The formation of the EHG ancestral component is estimated to have happened 13,000–15,000 years BP.[18]EHG associated remains belonged primarily to thehuman Y-chromosomehaplogroupsR1,with a lower frequency ofhaplogroup JandQ.Theirmitochondrial chromosomesbelonged primarily tohaplogroup U2,U4,U5,as well asC1andR1b.[2]Geneflow from an East Asian-like source towards the EHG contributed around 9.4% (4.4%–14.7%).[20]

EHGs may have mixed with "an Armenian-like Near Eastern source", which formed the Yamnaya culture, as early as theEneolithic(5200-4000 BC).[21]The people of theYamnaya culturewere found to be a mix of EHG and a "Near Eastern related population". During the 3rd millennium BC, the Yamnaya people embarked on a massive expansion throughoutEurope,which significantly altered the genetic landscape of the continent. The expansion gave rise to cultures such asCorded Ware,and was possibly the source of the distribution ofIndo-European languagesin Europe.[14]

The people of the MesolithicKunda cultureand theNarva cultureof the easternBalticwere a mix of WHG and EHG, showing the closest affinity with WHG. Samples from theUkrainianMesolithic andNeolithicwere found to cluster tightly together between WHG and EHG, suggesting genetic continuity in theDnieper Rapidsfor a period of 4,000 years. The Ukrainian samples belonged exclusively to the maternal haplogroupU,which is found in around 80% of all European hunter-gatherer samples.[22]

The people of thePit–Comb Ware culture(PCW/CCC) of the eastern Baltic bear 65% EHG ancestry. This is in contrast to earlier hunter-gatherers in the area, who were more closely related to WHG. This was demonstrated using a sample of Y-DNA extracted from a Pit–Comb Ware individual. This belonged toR1a15-YP172.The four samples of mtDNA extracted constituted two samples ofU5b1d1,one sample ofU5a2d,and one sample ofU4a.[23]

Günther et al. (2018) analyzed 13 SHGs and found all of them to be of EHG ancestry. Generally, SHGs from western and northern Scandinavia had more EHG ancestry (ca 49%) than individuals from eastern Scandinavia (ca. 38%). The authors suggested that the SHGs were a mix of WHGs who had migrated into Scandinavia from the south, and EHGs who had later migrated into Scandinavia from the northeast along theNorwegiancoast. SHGs displayed higher frequences of genetic variants that cause light skin (SLC45A2andSLC24A5), andlight eyes(OCA/Herc2), than WHGs and EHGs.[24]

Residual genetic ancestry of European hunter-gatherers during theEuropean Neolithic,between 7.5 ka and 5 ka BP (c. 5,500~3,000 BC)

Members of the Kunda culture and Narva culture were also found to be more closely related with WHG, while the Pit–Comb Ware culture was more closely related to EHG. Northern and eastern areas of the eastern Baltic were found to be more closely related to EHG than southern areas. The study noted that EHGs, like SHGs and Baltic hunter-gatherers, carried high frequencies of the derivedallelesfor SLC24A5 and SLC45A2, which are codings forlight skin.[25]

Mathieson et al. (2018) analyzed the genetics of a large number of skeletons of prehistoric Eastern Europe. Thirty-seven samples were from Mesolithic and Neolithic Ukraine (9500-6000 BC). These were classified as intermediate between EHG and SHG. The males belonged exclusively toRhaplotypes (particularly subclades ofR1b1andR1a) andIhaplotypes (particularly subclades ofI2). Mitochondrial DNA belonged almost exclusively toU(particularly subclades ofU5andU4).[21]

A large number of individuals from theZvejnieki burial ground,which mostly belonged to the Kunda culture and Narva culture in the eastern Baltic, were analyzed. These individuals were mostly of WHG descent in the earlier phases, but over time EHG ancestry became predominant. The Y-DNA of this site belonged almost exclusively to haplotypes ofhaplogroup R1b1a1aandI2a1.The mtDNA belonged exclusively tohaplogroup U(particularly subclades ofU2,U4andU5).[21]

Forty individuals from three sites of theIron Gates Mesolithicin theBalkanswere estimated to be of 85% WHG and 15% EHG descent. The males at these sites carried exclusivelyR1b1aandI(mostly subclades ofI2a) haplotypes.mtDNAbelonged mostly toU(particularly subclades ofU5andU4).[21]

People of theCucuteni–Trypillia culturewere found to harbor about 20% hunter-gatherer ancestry, which was intermediate between EHG and WHG.[21]

Narasimshan et al. (2019) coined a new ancestral component,West Siberian Hunter-Gatherer(WSHG). WSHGs contained about 20% EHG ancestry, 73% ANE ancestry, and 6% East Asian ancestry.[26]

Possible association with Early Indo-European

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The EHG have been argued by some to represent a possible source for thePre-Proto-Indo-European language(see alsoFather Tongue hypothesis). Unlike theYamnaya culture people(or closely related groups), which are associated with speakers of Proto-Indo-European, the EHG-richDnieper–Donets culture peopleshow no evidence ofCaucasus Hunter-Gatherer(CHG) orEarly European Farmer(EEF) ancestry.[27]Both Dnieper-Donets males and Yamnaya males carry the same paternal haplogroups (R1b and I2a), suggesting that the CHG and EEF admixture among the Yamnaya came through EHG males mi xing with EEF and CHG females. Based on this,David W. Anthony,this suggests that theIndo-European languageswere initially spoken by EHGs living in Eastern Europe.[28]

Others have suggested that the Indo-European language family may have originated not in Eastern Europe, but among CHG-rich West Asian populations South of the Caucasus which later absorbed EHG-rich groups North of the Caucasus. It was noted that haplogroups may not correlate with autosomal ancestry components and historical language dispersals.[29]

Physical appearance

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The mutation for blond hair is thought to have originated among theAfontova Gorapopulation of theAncient North Eurasian(ANE) cline of south-central Siberia.[30]

The EHGs are suggested to have had mostlybrown eyesand light skin,[24][31]with "intermediate frequencies of the blue-eye variants" and "high frequencies of the light-skin variants."[32]An EHG from Karelia was determined byGünther (2018)to have high probabilities of being brown-eyed and dark haired, with a predicted intermediate skin tone.[33]Another EHG from Samara was predicted to be light skinned, and was determined to have a high probability of being blue-eyed with a light hair shade, with a 75% calculated probability of being blond-haired.[34][32]

The rs12821256alleleof theKITLGgene that controlsmelanocytedevelopment andmelaninsynthesis,[35]which is associated withblondhair and first found inan individualfrom Siberia dated to around 17,000 BP, is found in three Eastern Hunter-Gatherers from Samara, Motala and Ukrainec. 10,000 BP,suggesting that this allele originated in the Ancient North Eurasian population, before spreading to western Eurasia.[36]

Many remains of East Hunter-Gatherers dated to circa 8,100 BP (6,100 BCE) have also been excavated atYuzhny Olenyisland inLake Onega.[37]TheAncient North Eurasian(ANE) ancestry is by far the main component of the Yuzhny Oleny group, and is among the highest within the rest of the Eastern Hunter-Gatherers (EHG).[4]

  • Reconstruction of burial No. 132 of the Oleneostrovsky burial ground (Yuzhni Oleny island, Lake Onega). Exhibit of the National Museum of the Republic of Karelia.[38]
    Reconstruction of burial No. 132 of the Oleneostrovsky burial ground (Yuzhni Oleny island, Lake Onega). Exhibit of the National Museum of the Republic of Karelia.[38]
  • Artifacts and reconstruction of Eastern Hunter-Gatherers from Yuzhny Oleny island by Gerasimov.[38]
    Artifacts and reconstruction of Eastern Hunter-Gatherers fromYuzhny Olenyisland by Gerasimov.[38]
  • Karelian Petroglyph depicting 5 skiers and a reindeer. These petroglyphs date to 7,000~6,000 years BP.
    Karelian Petroglyphdepicting 5 skiers and a reindeer. These petroglyphs date to 7,000~6,000 years BP.

Material culture

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Adoption of pottery among East European hunter-gatherers, during the 6th millennium BC (from the first adoption circa 5900 BC in the NorthCaspian Sea-or possibly from beyond the Ural area-, to final diffusion circa 5500 BC in theBaltic).[39]

As hunter-gatherers, the EHGs initially relied on stone tools and artifacts derived from ivory, horns or antlers. From circa 5,900 BC, they started to adopt pottery in the area of the northernCaspian Sea,or possibly from beyond the Ural. In barely three or four centuries, pottery spread over a distance of about 3,000 kilometers, reaching as far as theBaltic Sea.This technological spread was much faster than the spread of agriculture itself, and mainly occurred through technology transfer between hunter-gatherer groups, rather than through the demic diffusion of agriculturalists.[40]

See also

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Notes

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  1. ^Lazaridis et al. (2016) found SHGs to be a mix of EHGs and WHGs: "Eastern Hunter Gatherers (EHG) derive 3/4 of their ancestry from the ANE... Scandinavian hunter-gatherers (SHG) are a mix of EHG and WHG; and WHG are a mix of EHG and the Upper Paleolithic Bichon from Switzerland."[13]

References

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  1. ^National Museum of Karelia exhibit
  2. ^abcPosth, Cosimo; Yu, He; Ghalichi, Ayshin (March 2023)."Palaeogenomics of Upper Palaeolithic to Neolithic European hunter-gatherers".Nature.615(7950): 117–126.Bibcode:2023Natur.615..117P.doi:10.1038/s41586-023-05726-0.ISSN1476-4687.PMC9977688.PMID36859578.
  3. ^Haak, Wolfgang; Lazaridis, Iosif; Patterson, Nick; Rohland, Nadin; Mallick, Swapan; Llamas, Bastien; Brandt, Guido; Nordenfelt, Susanne; Harney, Eadaoin; Stewardson, Kristin; Fu, Qiaomei (June 1, 2015). "Massive migration from the steppe was a source for Indo-European languages in Europe". Nature. 522 (7555): 207–211. doi:10.1038/nature14317. ISSN 1476-4687. PMC 5048219. PMID 25731166.
  4. ^abKozintsev, A. G. (January 4, 2022)."Patterns in the Population History of Northern Eurasia from the Mesolithic to the Early Bronze Age, Based on Craniometry and Genetics".Archaeology, Ethnology & Anthropology of Eurasia.49(4): 141.doi:10.17746/1563-0110.2021.49.4.140-151.ANE makes up the principal share of the EHG (Eastern Hunter-Gatherer) autosomal component, whose content is especially high in the genomes of Mesolithic and Early Neolithic inhabitants of northeastern Europe buried at Yuzhny Oleny Ostrov, Popovo, Sidelkino, Lebyazhinka IV, etc. (Haak et al., 2015; Damgaard et al., 2018). They passed EHG on to the Yamnaya people, from whom it was inherited by several filial populations, including Afanasyevans. As early as the Mesolithic, EHG was introduced from northern Russia to Scandinavia, as evidenced by genomes of the Motala people in southern Sweden. Their ancestors had migrated there from the east along the coast of Norway, because the share of EHG in more southern populations, such as the earlier Kunda people of the eastern Baltic, is lower (Haak et al., 2015; Mittnik et al., 2018).
  5. ^abFeldman, Michal; Gnecchi-Ruscone, Guido A.; Lamnidis, Thiseas C.; Posth, Cosimo (2021). "Where Asia meets Europe – recent insights from ancient human genomics".Annals of Human Biology.48(3): 191–202.doi:10.1080/03014460.2021.1949039.PMID34459345.S2CID237348859.
  6. ^Nägele, Kathrin; Rivollat, Maite; Yu, He; Wang, Ke (2022). "Ancient genomic research - From broad strokes to nuanced reconstructions of the past".Journal of Anthropological Sciences.100(100): 193–230.doi:10.4436/jass.10017.PMID36576953.
  7. ^Anthony 2019b,p. 27.
  8. ^Kashuba 2019:"Earlier aDNA studies suggest the presence of three genetic groups in early postglacial Europe: Western hunter–gatherers (WHG), Eastern hunter–gatherers (EHG), and Scandinavian hunter–gatherers (SHG)4. The SHG have been modelled as a mixture of WHG and EHG."
  9. ^Anthony 2019b,p. 28.
  10. ^Haak, Wolfgang; Lazaridis, Iosif; Patterson, Nick; Rohland, Nadin; Mallick, Swapan; Llamas, Bastien; Brandt, Guido; Nordenfelt, Susanne; Harney, Eadaoin; Stewardson, Kristin; Fu, Qiaomei (June 1, 2015)."Massive migration from the steppe was a source for Indo-European languages in Europe".Nature.522(7555): 207–211.arXiv:1502.02783.Bibcode:2015Natur.522..207H.doi:10.1038/nature14317.ISSN1476-4687.PMC5048219.PMID25731166.
  11. ^van de Loosdrecht, Marieke; Bouzouggar, Abdeljalil; Humphrey, Louise; Posth, Cosimo; Barton, Nick; Aximu-Petri, Ayinuer; Nickel, Birgit; Nagel, Sarah; Talbi, El Hassan; El Hajraoui, Mohammed Abdeljalil; Amzazi, Saaïd; Hublin, Jean-Jacques; Pääbo, Svante; Schiffels, Stephan; Meyer, Matthias (May 4, 2018)."Pleistocene North African genomes link Near Eastern and sub-Saharan African human populations".Science.360(6388): 548–552.Bibcode:2018Sci...360..548V.doi:10.1126/science.aar8380.ISSN0036-8075.PMID29545507.
  12. ^Lazaridis, Iosif; Alpaslan-Roodenberg, Songül; Acar, Ayşe; Açıkkol, Ayşen; Agelarakis, Anagnostis; Aghikyan, Levon; Akyüz, Uğur; Andreeva, Desislava; Andrijašević, Gojko; Antonović, Dragana; Armit, Ian; Atmaca, Alper; Avetisyan, Pavel; Aytek, Ahmet İhsan; Bacvarov, Krum (August 26, 2022)."The genetic history of the Southern Arc: A bridge between West Asia and Europe".Science.377(6609): eabm4247.doi:10.1126/science.abm4247.ISSN0036-8075.PMC10064553.PMID36007055.S2CID251843620.
  13. ^Lazaridis 2016.
  14. ^abHaak 2015.
  15. ^Irving-Pease, Evan K.; Refoyo-Martínez, Alba; Barrie, William; Ingason, Andrés; Pearson, Alice; Fischer, Anders; Sjögren, Karl-Göran; Halgren, Alma S.; Macleod, Ruairidh; Demeter, Fabrice; Henriksen, Rasmus A.; Vimala, Tharsika; McColl, Hugh; Vaughn, Andrew H.; Speidel, Leo (January 24, 2024)."The selection landscape and genetic legacy of ancient Eurasians".Nature.625(7994): 312–320.Bibcode:2024Natur.625..312I.doi:10.1038/s41586-023-06705-1.ISSN1476-4687.PMC10781624.PMID38200293.
  16. ^Lazaridis, Iosif (December 1, 2018)."The evolutionary history of human populations in Europe".Current Opinion in Genetics & Development.Genetics of Human Origins.53:21–27.arXiv:1805.01579.doi:10.1016/j.gde.2018.06.007.ISSN0959-437X.PMID29960127.S2CID19158377.
  17. ^Haak, Wolfgang; Lazaridis, Iosif; Patterson, Nick; Rohland, Nadin; Mallick, Swapan; Llamas, Bastien; Brandt, Guido; Nordenfelt, Susanne; Harney, Eadaoin; Stewardson, Kristin; Fu, Qiaomei; Mittnik, Alissa; Bánffy, Eszter; Economou, Christos; Francken, Michael (June 2015)."Massive migration from the steppe was a source for Indo-European languages in Europe".Nature.522(7555): 207–211.arXiv:1502.02783.Bibcode:2015Natur.522..207H.doi:10.1038/nature14317.ISSN1476-4687.PMC5048219.PMID25731166.Haak et al. (2015): 38–40% ANE (MA-1), 60–62% WHG (Fig S8.6). (Alternative topologies where EHG and ANE are unadmixed sister lineages, with WHG being admixed, are not rejected)
  18. ^abAllentoft, Morten E.; Sikora, Martin; Refoyo-Martínez, Alba; Irving-Pease, Evan K.; Fischer, Anders; Barrie, William; Ingason, Andrés; Stenderup, Jesper; Sjögren, Karl-Göran; Pearson, Alice; Sousa da Mota, Bárbara; Schulz Paulsson, Bettina; Halgren, Alma; Macleod, Ruairidh; Jørkov, Marie Louise Schjellerup (January 2024)."Population genomics of post-glacial western Eurasia".Nature.625(7994): 301–311.Bibcode:2024Natur.625..301A.doi:10.1038/s41586-023-06865-0.ISSN1476-4687.PMC10781627.PMID38200295.
  19. ^Villalba-Mouco, Vanessa; van de Loosdrecht, Marieke S.; Rohrlach, Adam B.; Fewlass, Helen; Talamo, Sahra; Yu, He; Aron, Franziska; Lalueza-Fox, Carles; Cabello, Lidia; Cantalejo Duarte, Pedro; Ramos-Muñoz, José; Posth, Cosimo; Krause, Johannes; Weniger, Gerd-Christian; Haak, Wolfgang (April 2023)."A 23,000-year-old southern Iberian individual links human groups that lived in Western Europe before and after the Last Glacial Maximum".Nature Ecology & Evolution.7(4): 597–609.Bibcode:2023NatEE...7..597V.doi:10.1038/s41559-023-01987-0.ISSN2397-334X.PMC10089921.PMID36859553.Currently, the strongest affinity to Tianyuan in Holocene European HGs was reported for Eastern European HGs (EHG). This is because the ancestry found in Mal'ta and Afontova Gora individuals (Ancient North Eurasian ancestry) received ancestry from UP East Asian/Southeast Asian populations54, who then contributed substantially to EHG55.
  20. ^Childebayeva, Ainash; Fricke, Fabian; Rohrlach, Adam Benjamin; Huang, Lei; Schiffels, Stephan; Vesakoski, Outi; Mannermaa, Kristiina; Semerau, Lena; Aron, Franziska; Solodovnikov, Konstantin; Rykun, Marina; Moiseyev, Vyacheslav; Khartanovich, Valery; Kovtun, Igor; Krause, Johannes (June 11, 2024)."Bronze age Northern Eurasian genetics in the context of development of metallurgy and Siberian ancestry".Communications Biology.7(1): 1–12.doi:10.1038/s42003-024-06343-x.ISSN2399-3642.PMC11166947.We then modeled gene flow from the lineage leading to CHB to the EEHG at 9.4% (95% CI 4.4%–14.7%).
  21. ^abcdeMathieson et al. 2018.
  22. ^Jones 2017.
  23. ^Saag 2017.
  24. ^abGünther 2018.
  25. ^Mittnik 2018.
  26. ^Narasimhan 2019.
  27. ^Anthony 2019a,p. 14.
  28. ^Anthony 2019a,pp. 7, 14.
  29. ^Lazaridis, Iosif; Alpaslan-Roodenberg, Songül; Acar, Ayşe; Açıkkol, Ayşen; Agelarakis, Anagnostis; Aghikyan, Levon; Akyüz, Uğur; Andreeva, Desislava; Andrijašević, Gojko; Antonović, Dragana; Armit, Ian; Atmaca, Alper; Avetisyan, Pavel; Aytek, Ahmet İhsan; Bacvarov, Krum (August 26, 2022)."The genetic history of the Southern Arc: A bridge between West Asia and Europe".Science.377(6609): eabm4247.doi:10.1126/science.abm4247.ISSN0036-8075.PMC10064553.PMID36007055.
  30. ^Hanel, Andrea; Carlberg, Carsten (July 3, 2020)."Skin colour and vitamin D: An update".Experimental Dermatology.29(9): 864–875.doi:10.1111/exd.14142.PMID32621306.S2CID220335539.
  31. ^Hanel, Andrea; Carlberg, Carsten (2020)."Skin Colour and Vitamin D: An update".Experimental Dermatology.29(9): 864–875.doi:10.1111/exd.14142.PMID32621306.S2CID220335539."Interestingly, eastern and Scandinavian hunter-gatherers had light skin,[48] in contrast to Baltic hunter-gatherers who kept their dark skin only until 3800 years ago when farming was introduced in this region by the Bronze Age expansion of people of Russian steppe origin.[56, 57]"
  32. ^abPopulation genomics of Mesolithic Scandinavia: Investigating early postglacial migration routes and high-latitude adaptation S8 Text. Functional variation in ancient samples.,doi:10.1371/journal.pbio.2003703.s013
  33. ^Günther 2018,p. 4/28: From Supplementary document S8: "The Karelian individual presents high probabilities of being brown-eyed (0.99), and having a dark hair (0.96). Without speculating about the genetic architecture of skin pigmentation, we suggest an intermediate skin-pigmentation phenotype for the Karelia individual, as it carried the ancestral allele at rs16891982 and the derived allele at rs1426654 (S1 Table). The presence of the rs1426654 light-skin allele, in addition to five additional C11-associated alleles at haplotype defining SNPs (S1 Table) suggests that the Karelian individual carried the C11 light-skin haplotype."
  34. ^Günther 2018,p. 4/28: From Supplementary document S8: "The Samaran individual exhibits high probabilities of being blue-eyed (0.88), light hair shade (0.99); most likely being blond (0.75)."
  35. ^Sulem, Patrick; Gudbjartsson, Daniel F.; Stacey, Simon N.; Helgason, Agnar; Rafnar, Thorunn; Magnusson, Kristinn P.; Manolescu, Andrei; Karason, Ari; Palsson, Arnar; Thorleifsson, Gudmar; et al. (December 2007)."Genetic determinants of hair, eye and skin pigmentation in Europeans".Nature Genetics.39(12): 1443–1452.doi:10.1038/ng.2007.13.ISSN1546-1718.PMID17952075.S2CID19313549.
  36. ^Mathieson et al. 2018"Supplementary Information page 52:" The derived allele of the KITLG SNP rs12821256 that is associated with – and likely causal for blond hair in Europeans is present in one hunter-gatherer from each of Samara, Motala and Ukraine (I0124, I0014 and I1763), as well as several later individuals with Steppe ancestry. Since the allele is found in populations with EHG but not WHG ancestry, it suggests that its origin is in the Ancient North Eurasian (ANE) population. Consistent with this, we observe that the earliest known individual with the derived allele (supported by two reads) is the ANE individual Afontova Gora 3, which is directly dated to 16130-15749 cal BCE (14710±60 BP, MAMS-27186: a previously unpublished date that we newly report here). We cannot determine the status of rs12821256 in Afontova Gora 2 and MA-1 due to lack of sequence coverage at this SNP. "
  37. ^Mittnik, Alissa; Wang, Chuan-Chao; Pfrengle, Saskia (January 30, 2018)."The genetic prehistory of the Baltic Sea region".Nature Communications.9(1): Fig. 1.Bibcode:2018NatCo...9..442M.doi:10.1038/s41467-018-02825-9.ISSN2041-1723.PMC5789860.PMID29382937.
  38. ^abKozintsev, Alexander (January 1, 2021)."Patterns in the population history of Northern Eurasia from the Mesolithic to the Early Bronze Age".Archaeology, Ethnology and Anthropology of Eurasia.49(4): 140–151.doi:10.17746/1563-0110.2021.49.4.140-151.ANE makes up the principal share of the EHG (Eastern Hunter-Gatherer) autosomal component, whose content is especially high in the genomes of Mesolithic and Early Neolithic inhabitants of northeastern Europe buried at Yuzhny Oleny Ostrov, Popovo, Sidelkino, Lebyazhinka IV, etc. (Haak et al., 2015; Damgaard et al., 2018). "," Mesolithic, northern Russian Plain, Yuzhny Oleny Ostrov (Alekseyev, Gokhman, 1984)
  39. ^Dolbunova, Ekaterina; Lucquin, Alexandre (February 2023)."The transmission of pottery technology among prehistoric European hunter-gatherers".Nature Human Behaviour.7(2): 171–183.doi:10.1038/s41562-022-01491-8.ISSN2397-3374.PMC9957732.PMID36550220.
  40. ^Dolbunova, Ekaterina; Lucquin, Alexandre (February 2023)."The transmission of pottery technology among prehistoric European hunter-gatherers".Nature Human Behaviour.7(2): 171–183.doi:10.1038/s41562-022-01491-8.ISSN2397-3374.PMC9957732.PMID36550220.Although demic diffusion may have a role, on the basis of its speed we argue that pottery production was rapidly disseminated through knowledge transfer across established networks between dispersed hunter-gatherer communities

Bibliography

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Further reading

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